Topics in Biodiversity and Conservation 20

Florencia Montagnini Editor

Biodiversity Islands:
Strategies for
Conservation in
Human-Dominated
Environments
Topics in Biodiversity and Conservation

Volume 20

Series Editors
Professor David L. Hawksworth CBE, The Natural History Museum, London, UK
Dr. Anurag Chaurasia, Indian Institute of Vegetable Research, Indian Council of
Agricultural Research, Varanasi, Uttar Pradesh, India
Springer’s book series, Topics in Biodiversity and Conservation, brings together
some of the most exciting and topical papers in biodiversity and conservation
research. The result is a series of useful themed collections covering issues such as
the diversity and conservation of specific habitats or groups of organisms, and the
key dilemma of resource use versus conservation.
Based on Springer’s popular journal, Biodiversity and Conservation, the series
provides access to selected peer-reviewed papers which represent the cutting edge of
current research to provide a valuable overview of progress in each topic addressed.
With their diversity of case studies and depth of investigation, these collections will
be of particular interest for courses including biodiversity and/or conservation issues,
and to advanced students and researchers working in related fields.

More information about this series at [Link]

Florencia Montagnini
Editor

Biodiversity Islands:
Strategies for Conservation
in Human-Dominated
Environments
Editor
Florencia Montagnini
School of the Environment,
The Forest School
Yale University
New Haven, CT, USA

ISSN 1875-1288 ISSN 1875-1296 (electronic)

Topics in Biodiversity and Conservation
ISBN 978-3-030-92233-7 ISBN 978-3-030-92234-4 (eBook)
[Link]

© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland
AG 2022
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether
the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and
transmission or information storage and retrieval, electronic adaptation, computer software, or by
similar or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or
the editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
This book is dedicated to my father.
Foreword

It is unfortunate that a book on this topic needed to be written, but it is on a highly

relevant subject given the worldwide destruction of natural habitats and the loss of so
many biological species. As we, at present, face the sixth great species extinction, the
remaining biodiversity survive in small patches or islands, which have become
extremely important for the survival of the species they contain or the migrant
birds they host. As someone who has worked a lifetime in botanical gardens, I am
very aware of the small patches of original vegetation that they often contain and
their value for the pollinators that visit the flowers or the occasional visit of
migratory birds as they pause on their journey. At the Royal Botanical Gardens at
Kew in London, there is rare-listed hoverfly and an endangered species of lichen
among other species preserved in this biodiversity island. In my field work in the
highly fragmented Atlantic rainforest of Brazil and Argentina, we are still finding
new and undescribed species of plants in the small remnants of the original forest. It
is fortunate that still many species of animals and plants survive in these often small
islands, making them extremely important, and a book drawing attention to them is
most welcome.
Until relatively recently, much more attention was given to marine islands
following the work of MacArthur and Wilson in 1967 and because of their much-
threatened biodiversity, but now there is a growing realisation of the importance of
human-made islands on the mainland. The creation of biodiversity islands has been
the topic of important research in the Biological Dynamics of Forest Fragments
Project near Manaus, Brazil. I have spent many hours identifying plant species for
this project. The original name of the project “Minimum Critical Size of Ecosystems
Project” indicates its original research purpose to provide data about the minimum
area needed to preserve a functioning area of rainforest to assist in the establishment
of reserves and conservation areas. This book clearly demonstrates that today there
are biodiversity islands of many different sizes, shapes and purposes.
This book treats a great variety of different types of biodiversity islands, all of
which are areas of high biodiversity surrounded by highly degraded or intensely

vii
viii Foreword

used landscapes that act as refuges for the surviving species of the original ecosys-
tem. The many examples given here clearly show the critical importance of biodi-
versity islands for conservation, restoration and sustainable management of several
productive agroforestry systems. It is good to be taken around the world with
examples of biodiversity islands in both the tropical and the temperate regions. I
like the fact that these examples include not only areas of pristine natural habitats
such as the Monteverde Cloud Forest in Costa Rica or the forest islands in the
Paraguayan Chaco but also several examples from highly managed islands in
agroforestry and regenerative agriculture systems. Some of the examples of the
policies and political motivations given in various chapters should be helpful to
anyone involved in the creation or management of a biodiversity island. Several
chapters here show examples of harmonising food production with conservation.
This unity of purpose is important and is far more likely to be of long-term success
than placing conservation and agriculture in separate camps. Several chapters show
the importance of alternative ways to produce food from more integrated manage-
ment systems that also preserve biodiversity. The social, ecological, ethical and
economic benefits of such systems are clearly outlined in several of the chapters.
I congratulate the editor of this book for gathering together such a varied and
useful compilation of the ongoing work on biodiversity islands. This will be of
considerable use to people involved in the design of future biodiversity islands
because it has much to say about the motivations and politics and also about their
size and spatial distribution whether from fragments of the original vegetation or
from restoration of degraded and intensely used areas. It will be a most useful tool for
both conservation and restoration. My hope is that this will be used by conservation
organisations, local communities and indigenous peoples to create effective islands
of biodiversity in many different ecosystems of the world and for many more
creative types of management.

Lyme Regis, United Kingdom Professor Sir Ghillean T. Prance FRS, VMH
e‐mail: siriain01@[Link]
Preface

A biodiversity island is an area of high biodiversity located within ecologically

degraded, human-dominated landscapes. Biodiversity in the “islands” exceeds the
surrounding landscape biodiversity baseline. These biodiversity islands thereby act
as ecological refuges, promoting restoration and conservation in altered ecosystems
prevalent today throughout the globe.
Biodiversity islands can provide food, water, fuels, and fibers, as well as genetic,
medicinal, biochemical, and ornamental resources, pollination services, biological
pest control, and maintenance of life cycles of migratory species. These landscapes
hold promise for protecting a multitude of plant and animal species for present and
future generations. The presence of biodiversity islands spread over a large area can
decrease the chances of habitat loss from fire, disease, and other disturbances.
Biodiversity islands can exist within a wide range of human-dominated land-
scapes, including forest, agricultural, and urban settings, and can vary in scale from
square meters to thousands of square kilometers. Design strategies for biodiversity
islands depend on the spatial distribution of reserves throughout the landscape, the
degree of site degradation, the species present, and their locations within the urban to
rural spectrum.
This book is intended to provide an overview for the identification and establish-
ment of biodiversity islands, presenting examples and case studies where the biodi-
versity islands approach is being used in a variety of locations and contexts
worldwide. This book will contribute to design parameters on appropriate sizing
and spatial distribution of biodiversity islands to be effective in conservation and
regeneration across the landscape, using integrated landscape management
approaches.
The chapters discuss current challenges faced today by biodiversity conservation
researchers, practitioners, and policy makers and propose innovative approaches to
tackle them. Contributors are an assemblage of researchers, academicians, and
practitioners from biodiversity conservation, environmental management, forestry,
agroecology, agroforestry, and related fields who approach the issues from unique
perspectives.

ix
x Preface

This book comprises five parts: Part I, Introduction, establishes the framework
for understanding the complexities of biodiversity islands and the variety of strate-
gies that can be used to establish them. The Introduction defines the term “biodiver-
sity islands” and their size, location, and distribution in the landscape; stresses their
many ecological, social, and economic benefits; and discusses potential limitations
of the use of this framework along with ways to overcome them. Part II, Biodiver-
sity Islands Establishment and Management: Challenges and Alternatives,
shows how design strategies may depend on landscape use within the matrix of
habitat fragmentation, with integrated landscape management (ILM), including
sustainable agriculture, agroforestry, and community-led action, providing a frame-
work for implementation. Part III, Biodiversity Islands Across the Globe: Case
Studies, shows how varied agroecological strategies were applied in the formation
or conservation of biodiversity islands in human-dominated landscapes in Paraguay,
Peru, Costa Rica, Colombia, Great Britain, Argentina, Panama, and the USA. The
variety of case studies from different types of landscapes from several regions of the
world reveals the role biodiversity islands play in conserving local flora and fauna
that have been largely diminished by anthropogenic activities, while providing
cultural connections to nature and supplying ecosystem services that make biodi-
versity islands advantageous to farmers and nearby communities. Part IV,
Safeguarding the Environmental, Economic, and Social Benefits of Biodiversity
Islands, further details the economic, social, political, and cultural aspects of the
establishment and persistence of biodiversity islands in anthropogenic landscapes,
emphasizing how community-led action contributes to their development and sub-
sequent management, with examples from Puerto Rico, Ecuador, Brazil, India, the
USA, Panama, and Ethiopia. Part V, Conclusions, summarizes the lessons learned
while compiling this volume and lays out the pending challenges and potential
solutions ahead.
One late summer afternoon, about 2 years ago, while relaxing in the porch of a
house in suburban/rural Northford, Connecticut, a fox ran across the garden, appar-
ently not feeling too threatened by our presence. When wondering where this small
animal was coming from, and where did it go when it finally ran away, Kjell E Berg
suggested that the water reservoir located about 100 meters from the house was a
nice undisturbed forest that perhaps was functioning as a biodiversity island. Soon
the idea of digging more into the concept grew in all directions; the next day, Brett
Levin at Yale enthusiastically took it as his own project, and soon we wrote the
introductory chapter of this book among the three of us.
Other ideas followed as we developed a website: [Link]
and led a meeting session called “Biodiversity Islands: Pockets of Protected Land in
Human Dominated Environments” at a IUFRO (International Union of Forest
Research Organizations) conference in Posadas, Misiones, Argentina, in October
2018. The structure and contents of this book further developed as we met and held
conversations with students, colleagues, and friends whose enthusiasm, energy, and
joyful attitude made this book possible from start to end. The more than a 100 authors
who contributed chapters for this book drove the rest of the way with their
Preface xi

dynamism, dedication, and persistence. Numerous colleagues and friends also

helped with their intellectual input and moral support.
There was a total of 105 contributors from 11 countries (32 Argentina; 2 Brazil; 1
Canada; 14 Colombia; 2 Costa Rica; 10 Mexico; 4 Panama; 11 Paraguay; 3 Peru;
23 USA; 2 UK). Different chapters report research, case studies, and experiences
from 14 countries: Argentina, Brazil, Colombia, Costa Rica, Ecuador, Ethiopia,
India, Mexico, Panama, Paraguay, Peru, Puerto Rico, the UK, and the USA. Thus,
the book includes examples of biodiversity islands from tropical as well as temperate
regions, ranging from natural habitats to agroforestry and regenerative agriculture
systems, and from relatively small to large geographic areas of the world.
A holistic, multidisciplinary perspective was taken in approaching each theme,
encompassing factors and variables from multiple disciplines. The contributing
authors present views from the academic, practitioner, and policy-making perspec-
tives, offering alternatives and suggestions for promoting strategies that support
biodiversity conservation through intentionally designed frameworks for sustainable
forest landscapes. With the current worldwide trend of habitat destruction and the
need to preserve biodiversity and its values, this book is an essential tool as it
provides suggestions and concrete examples that can be used by a variety of
stakeholders in various settings throughout the world. This book is useful to
researchers, farmers, foresters, landowners, land managers, city planners, and policy
makers alike.

New Haven, CT, USA Florencia Montagnini

Acknowledgments

Many chapter contributors acted as independent reviewers of other colleagues’

chapters. In addition, other external reviewers generously gave their time to read
and offer useful suggestions to improve the chapters. There was a total of
55 reviewers from the academic as well as from the practitioner’s realms. The
following is a list of chapter reviewers: Oscar J Abelleira, Dara Albrecht, Victor
Arroyo-Rodriguez, Gary Bentrup, Kjell E Berg, Robert Bushbacher, Jonathan
Cornelius, Sara del Fierro, Beatriz Eibl, Alberto Esquivel, Ben Everett-Lane,
Glenn Galloway, Sergius Gandolfi, Eva Garen, Libertario González, Heather
Griscom, David Hawksworth, Karen Kainer, Keith Kirby, B. Mohan Kumar,
Rafaela Laino Guanes, Ariel Lugo, Brett Levin, Philip Marshall, Paula Meli,
Zoyla Mireya Clavo Peralta, Irene Montes-Londoño, Gabriela Morales-Nieves,
Mathew Moran, Carlos Navarro, Quint Newcomer, Fernando Niella, Joseph Orefice,
Alison Ormbsy, Nahuel Pachas, Pablo Peri, Daniel Piotto, Julio Prieto, Neptali
Ramírez-Marcial, Juan Rivero de Aguilar, Carmen María Rojas González, Ricardo
Rozzi, Rocío Santos-Gally, John Schelhas, Sara Scherr, Emily Sigman, Jacob
Slusser, Ryan Smith, Rosina Soler, Eric Toensmeier, Mateo Vega, Zoe Volenec,
Sheila Ward, Catherine Watson, and Gustavo Zuleta. Many thanks to them for their
generosity and dedication.
Sara del Fierro and Ryan Smith, both from Yale University’s School of the
Environment (YSE), performed multiple roles as most efficient, dynamic, and
enthusiastic book editors, assistants, reviewers, and co-authors. Dara Albrecht and
Ben Everett-Lane, both at Yale College, majoring in environmental science, gener-
ously volunteered their time as dedicated, energetic, and passionate editors, assis-
tants, reviewers, and co-authors. They all made the task feel more important and
appreciated, and their contributions and collegiality are immensely appreciated.
Financial, logistical, and administrative support from Yale School of the Envi-
ronment (YSE) made this book possible.

xiii
xiv Acknowledgments

Finally, this book was written to soothe the grief of losing Sunset, constant and
faithful companion whose energy, strength, and perseverance were always conta-
gious and made the ride through life smooth and enjoyable for so many years.

Northford and New Haven, CT, USA Florencia Montagnini

June 12, 2021
Contents

Part I Introduction
1 Introduction. Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments . . . . . . . . . . . . . . . . . . . . . . . . 3
Florencia Montagnini, Brett Levin, and Kjell E. Berg

Part II Biodiversity Islands Establishment and Management:

Challenges and Alternatives
2 The Importance of Small Rainforest Patches for Biodiversity
Conservation: A Multi-taxonomic Assessment . . . . . . . . . . . . . . . . . 41
Víctor Arroyo-Rodríguez, Ricard Arasa-Gisbert,
Norma P. Arce-Peña, Martín J. Cervantes-López,
Sabine J. Cudney-Valenzuela, Carmen Galán-Acedo,
Manuel A. Hernández-Ruedas, Miriam San-José, and Lenore Fahrig
3 Regenerative Agriculture as Biodiversity Islands . . . . . . . . . . . . . . 61
Brett Levin
4 Functions of Agroforestry Systems as Biodiversity Islands
in Productive Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Florencia Montagnini and Sara del Fierro
5 Biodiversity Islands: The Role of Native Tree Islands Within
Silvopastoral Systems in a Neotropical Region . . . . . . . . . . . . . . . . 117
Rocio Santos-Gally and Karina Boege
6 Riparian Forests: Longitudinal Biodiversity Islands
in Agricultural Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Lina Paola Giraldo, Julián Chará, Zoraida Calle,
and Ana M. Chará-Serna

xv
xvi Contents

7 Conservation and Registration of Seed Sources in Reserve

Remnants in the Province of Misiones, Argentina . . . . . . . . . . . . . . 157
Beatriz I. Eibl, Florencia Montagnini, Lucas N. López,
Héctor F. Romero, Claudio J. Dummel, Guillermo Küppers,
Miguel A. López, Pedro Lavignolle, Jorge Cortes,
and Marcelo A. De La Vega

Part III Biodiversity Islands Across the Globe: Case Studies

8 Islands of Forests Among Savannas: Key Elements
for Conservation and Production in the Paraguayan
Humid Chaco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Rafaela Laino, Karim Musalem, Luis Domingo Laino,
Andrea Caballero-Gini, Diego Bueno-Villafñae, Lidia Aranda,
Alberto Esquivel, Marcela Ferreira Riveros, Lía Romero Nardelli,
Nicolás Cantero, and Rebeca Irala
9 Biodiversity Islands and Dominant Species in Agricultural
Landscapes of the South Western Amazon, Perú . . . . . . . . . . . . . . 207
Zoyla Mirella Clavo Peralta, Jorge Washinton Vela Alvarado,
and Carlos Mariano Alvez-Valles
10 The Monteverde Cloud Forest: Evolution of a Biodiversity
Island in Costa Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Quint Newcomer, Fabricio Camacho Céspedes, and Lindsay Stallcup
11 A Highly Productive Biodiversity Island Within
a Monoculture Landscape: El Hatico Nature Reserve
(Valle del Cauca, Colombia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
Zoraida Calle D, Carlos Hernán Molina C,
Carlos Hernando Molina D, Enrique José Molina D,
Juan José Molina E, Bernardo Murgueitio C,
Amalia Murgueitio C, and Enrique Murgueitio R
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture
Amidst a Transformed Landscape in the Colombian Andes . . . . . . 305
Irene Montes-Londoño, Alicia Calle, Olimpo Montes,
and Arturo Montes
13 Islands of Trees in Long-Fragmented Landscapes
in Great Britain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
Keith J. Kirby
14 Natural Landscape of the Pampa Region in Santa Fe Province,
Argentina: Environmental Resilience and Opportunity
for Changing the Agri-Food Paradigm . . . . . . . . . . . . . . . . . . . . . . 353
Libertario Hugo González, Germán Neffen, Victoria Benedetto,
Marta Sánchez Miñarro, Andrea García, Ricardo Biasatti,
Pablo Rimoldi, Cristian Alesio, and Daniel Paiz
Contents xvii

15 Residential Garden Design for Urban Biodiversity Conservation:

Experience from Panama City, Panama . . . . . . . . . . . . . . . . . . . . . 387
Helen R. C. Negret, Rafael Negret, and Irene Montes-Londoño
16 Biodiversity Islands at the World’s Southernmost City:
Plant, Bird and Insect Conservation in Urban Forests
and Peatlands of Ushuaia, Argentina . . . . . . . . . . . . . . . . . . . . . . . . 419
Rosina Soler, Julieta Benítez, Francisco Sola,
and María Vanessa Lencinas
17 Paradise Lot: A Temperate-Climate Urban Agroforestry
Biodiversity Island . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439
Eric Toensmeier
18 Contribution to the Domestication and Conservation
of the Genetic Diversity of Two Native Multipurpose Species
in the Yabotí Biosphere Reserve, Misiones, Argentina . . . . . . . . . . . 461
Fernando Niella, Patricia Rocha, Ariel M. Tuzinkievicz,
Ricardo Buchweis, Christian Bulman Hartkopf, Peggy Thalmayr,
José González, Florencia Montagnini, and Sandra Sharry

Part IV Safeguarding the Environmental, Economic,

and Social Benefits of Biodiversity Islands
19 How Community-Led Action Can Advance the Development
of Biodiversity Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
Brett Levin
20 Priorities, Perspectives, and Use of a Community Forest
by Surrounding Residents in Mayagüez, Puerto Rico:
Protecting the Forest for Its Services . . . . . . . . . . . . . . . . . . . . . . . . 505
Gabriela M. Morales-Nieves
21 Sacred Church Forests in Northern Ethiopia: Biodiversity
and Cultural Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 531
Mabel Baez Schon, Carrie L. Woods, and Catherine L. Cardelús
22 Beyond the Island: Integrated Approaches to Conserving
Biodiversity Islands with Local Communities . . . . . . . . . . . . . . . . . 551
Michael S. Esbach, Mahi Puri, Robinson Botero-Arias,
and Bette A. Loiselle
23 Agroecology and Forest Conservation in Three Types
of Land Reform Communities in the Cacao Region
of Bahia, Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 569
Kathleen R. Painter, Robert Buschbacher, Luiz Carlos Souto Silva,
and Emerentina Costa e Silva
xviii Contents

24 Preserving Biodiversity in Appalachian Mixed Mesophytic

Forests Through the Permit-Based Harvest of American
Ginseng and Other Forest Botanicals . . . . . . . . . . . . . . . . . . . . . . . 601
Karam Sheban
25 Farmer Perceptions of Tropical Dry Forest Restoration
Practices on the Azuero Peninsula of Panama – Implications
for Increasing Biodiversity in a Human-Dominated Landscape . . . . 629
Vicente Vásquez, Cristina Barber, Yassine Dguidegue,
T. Trevor Caughlin, Roxana García, and Ruth Metzel
26 Safeguarding Biodiversity Islands in Northern Ethiopia
Amidst Political Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647
Emily Sigman

Part V Conclusions
27 Conclusions: Challenges and Opportunities in Implementing
Biodiversity Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 677
Florencia Montagnini, Ryan T. Smith, Benjamin Everett-Lane,
Sara del Fierro, and Dara Albrecht

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 705
About the Editor

Florencia Montagnini has over 30 years of experience researching and teaching in

topics on sustainability of managed ecosystems in the tropics, such as forest, tree
plantations, and agroforestry systems, with a special emphasis on Latin America.
Her work as a scientific advisor and consultant has also taken her to Africa and South
East Asia. Her research encompasses sustainable land-use systems that integrate
ecological principles with economic, social, and political factors; the principles and
applications of forest landscape restoration; the reforestation of degraded lands with
native species; identification and quantification of ecological services (biodiversity,
carbon sequestration, and watershed protection); organic farming using indigenous
resources; biodiversity conservation in human-dominated landscapes; and biodiver-
sity islands. She received her BS in agronomy from the National University of
Rosario, Argentina; her master’s degree in ecology from the Venezuelan Institute
of Scientific Research (IVIC), Caracas, Venezuela; and her PhD in ecology from the
University of Georgia. Since 1989, she has worked as a professor and researcher at
the Yale School of the Environment, as well as the Tropical Agriculture Research
and Higher Education Center (CATIE). She has written 11 books and over 250 sci-
entific articles about the ecology of tropical forests, agroforestry systems, native
species reforestation, and forest landscape restoration.

xix
 Part I
Introduction
Chapter 1
Introduction. Biodiversity Islands:
Strategies for Conservation in
Human-Dominated Environments

Florencia Montagnini, Brett Levin, and Kjell E. Berg

Abstract This chapter serves to conceptualize, identify and promote implementa-

tion of the framing tool we term Biodiversity Islands: ecological refuges where
plants and animals can thrive without major interference from human activity,
thereby providing ecological, economic, and social benefits at the ecosystem, land-
scape, and global levels. Design strategies for these biodiversity islands depend on
their purpose, as well as on the spatial distribution of reserves throughout the
landscape, degree of landscape degradation, species present, and location within
the urban-rural spectrum. Biodiversity islands can exist in a range of human-
dominated landscapes (e.g., agricultural, wetland, urban) and can range from square
meters to many square kilometers in size. In promoting biodiversity islands, we
encourage the application of integrated landscape management and inclusive com-
munity led action, and discuss possible financial and non-monetary incentives for
their implementation and protection. This book presents examples of biodiversity
islands from throughout the world and discusses potential difficulties with their use.
We expect this book to be useful to researchers, farmers, foresters, landowners, land
managers, city planners, and policy makers.

Keywords Agroforestry · Biodiversity conservation · Community action ·

Degraded landscapes · Ecosystem services · Landscape restoration

F. Montagnini (*)
School of the Environment, The Forest School, Yale University, New Haven, CT, USA
e-mail: fl[Link]@[Link]
B. Levin
Dietrick Institute for Applied Insect Ecology, Ventura, CA, USA
K. E. Berg
Bioren Systems, Hampden, MA, USA

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 3

F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
4 F. Montagnini et al.

1.1 Introduction

The concept of biodiversity islands helps focus and frame the importance of saving
intact sections of land where plants and animals can thrive without major degener-
ative interference from human activity. A biodiversity island is an area of high
biodiversity within ecologically degraded or threatened, human-dominated land-
scapes. Building upon the foundations of island biogeography theory (MacArthur
and Wilson 1967), small and large biodiversity islands act as ecological refugia,
protected areas, or reserves within the landscape, with biodiversity in the “island”
greatly exceeding the surrounding landscape’s biodiversity baseline.
Historically, the refugia concept was used to describe climatically stable areas in
which species survived past Quaternary glacial–interglacial oscillations. The frame-
work of “Anthropocene refugia” extends this concept in recognition of the so-called
“Anthropocene” era in which human activities are the dominant driving force on
ecosystems and are major factors limiting species distributions. “Anthropocene
refugia” then refer to areas that provide spatial and temporal protection from
human activities and that will remain suitable for a given taxonomic unit in the
long-term. This framework focuses on developing appropriate conservation strate-
gies for wildlife taxa around the world (Monsarrat et al. 2019). As we refer to them
here, biodiversity islands go a step further in proposing a framework to contribute to
the protection of a multitude of plant and animal species and to the provision of
ecological, social, and economic benefits for present and future generations.
The size, configuration, and position of biodiversity islands in the landscape may
be mostly opportunistic, determined by their purpose, as well as by patterns of
human settlement, development, and utilization or extraction of natural resources.
When newly established, planned and designed, their characteristics, i.e. size and
configuration may vary according to various scientific guidelines (Laurance 2008;
Laurance et al. 2018). Matrix type and quality are also important determinants of
taxonomic diversity (Boesing et al. 2018). Biodiversity islands may act as buffer
zones between areas of greater human impact or degradation. Multiple biodiversity
islands spread over a large area in an optimal configuration can decrease chances of
biodiversity loss through creation of repopulation reserves and biological corridors
(Harvey et al. 2008).
Biodiversity islands can also be designed and established to serve restoration
purposes, as in applied nucleation/island tree planting strategies used to accelerate
recovery of degraded tropical forests and pastures (Holl 2002; Holl et al. 2017;
Santos-Gally et al. 2019). Depending on their design and specific considerations for
their management and preservation, biodiversity islands can effectively serve to
protect all levels of biological diversity, including genetic variability, species diver-
sity, functional diversity, as well as ecosystems and landscape diversity (Eibl et al.
2022; Niella et al. 2022; Santos-Gally and Boege 2022).
A key challenge in designing and managing biodiversity islands, as explained in
chapters of this book authored by Arroyo-Rodríguez et al., Clavo Peralta et al.,
Kirby, and a few others, is the question of priorities and tradeoffs inherent in most
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 5

conservation approaches. Designing a protected area with emphasis on one or few

target species may not favor other species with different habitat requirements.
Deciding on a specific type of biodiversity island design without considering the
broader landscape, or the value of alternative land uses can undermine wider
conservation objectives. Biodiversity islands must be always designed and managed
with careful attention to both broad and local contexts and objectives.
Biodiversity islands are particularly valuable in areas of biodiversity hotspots.
Biodiversity hotspots are regions that contain significantly high concentrations of
plant or animal endemism and experience high rates of habitat loss. Because of those
features, attention is needed to protect their biodiversity. The 36 global hotspots
currently identified represent just 2.3% of Earth’s land surface, but they contain
around 50% of the world’s endemic plant species and 42% of all terrestrial vertebrate
species (Mittermeier et al. 2011; Conservation International 2014). Integrating
biodiversity islands within these biodiversity hotspots would allow multiple small
sites on a relatively small area of land to contribute to significant beneficial biodi-
versity outcomes.
Although we emphasize tropical forest landscapes, biodiversity is also highly
threatened in other types of ecosystems. Temperate grasslands ecosystems, for
instance, are especially threatened because they have been long preferred for eco-
nomic uses and human settlement. Another good example from temperate regions
are ancient woodland islands in the British countryside, as described by Kirby in this
volume. Wetlands and drylands are at increasing risk as well. In drylands, there are
fewer species adapted to the harsh environmental conditions, and these are especially
important and difficult to substitute, in contrast with humid forests where many
ecological functions are provided by several different species. Other non-forest
ecosystems, such as well-designed and managed agricultural systems, grazing
areas, non-native tree cover, ‘green’ infrastructure and eco-friendly built structures,
can also improve ecosystem services such as watershed functions, sedimentation
control, and carbon sequestration and storage. Therefore biodiversity islands are just
as important in non-forest ecosystems, as shown in respective chapters in this book
(example Laino et al., Vásquez et al.).
Recognizing, developing, and protecting biodiversity correlates with an increase
in economic benefit due to a greater presence of natural capital. Natural capital is the
stock of natural assets that make human life possible. These include the environ-
mentally derived provisions necessary for fresh air, food, clean water, medicine,
warmth, and shelter. Many of the benefits that may arise through the development of
biodiversity islands could be captured by natural capital accounting by placing a
monetary value on the environmental or ecosystem services provided. The estimated
total value of ecosystem services worldwide has been estimated to be between $125
and $145 trillion USD per year, offering an enormous contribution to human
wellbeing (Costanza et al. 2014). Biodiversity plays an important role within this
valuation by supplying provisions of food and water, fuels and fibers, genetic
resources, medicinal and other biochemical supplies, ornamental products, pollina-
tion services, biological control, and habitat for the maintenance of life cycles of
migratory species (Kumar 2010). Assessing economic value for biodiversity can
6 F. Montagnini et al.

guide strategies and policies for preserving ecosystem integrity over time (Pimentel
et al. 1997).
While it is important to recognize these economic benefits as a tool for evaluating
environmental gain through the development of biodiversity islands, it is also
important to recognize the non-monetary benefits. These can take the form of
cultural, religious, spiritual, educational, and experiential value within the landscape,
in addition to the inherent value of the biodiversity itself. One may view biodiversity
islands as natural capital “banks” within the landscape, which preserve and store
these important resources not only for their present value but also for the future. For
example, a natural area or piece of fallow land that may otherwise be disregarded or
developed can be protected as a biodiversity island to provide value to a farm,
community, or region for generations to come.
Although the threats to biodiversity are dramatic and broad-ranging, binding
international agreements for biodiversity conservation have thus far been largely
unsuccessful. The Convention on Biological Diversity (CBD) established a strategic
action plan for 2002 to 2010 which ran short of reaching its goals. The 2010 Aichi
Biodiversity Targets which aimed to reduce loss of natural habitats and expand
nature reserves and other effective area-based conservation reserves from 10% to
17% of the world’s land by 2020, have also fallen behind (Watts 2018). Following
the 14th Meeting of the Conference of the Parties to the Convention on Biological
Diversity (CBD COP 14 Egypt) in 2018 there was a unanimous decision to accel-
erate action to achieve the Aichi Biodiversity Targets by 2020 at the global, regional,
national and subnational levels. The meeting also developed a comprehensive and
participatory process for the post-2020 global biodiversity framework to be agreed
upon in Beijing at the next Conference of Parties (COP 15) in 2020 (State Informa-
tion Service 2018).
The post-2020 global biodiversity framework applies a “theory of change”
approach, a strategic planning framework used to help plan, implement and evaluate
the impacts of the actions taken. It provides a powerful tool for organizing measur-
able goals and solutions, and for evaluating both short-and long-term impacts in a
consistent, meaningful and transparent structure ([Link]
f84/a892b98d2982a829962b6371/[Link]). In this context, it
becomes imperative to increase efforts to promote biodiversity conservation strate-
gies, such as biodiversity islands, that can be effective at the local, regional and
global levels.
The interaction between climate change and habitat isolation (mainly caused by
habitat loss; Fahrig 2003) adds another significant threat to biodiversity conserva-
tion. Restoration of degraded ecosystems across the world can work towards
addressing climate change and biodiversity loss (Lovejoy and Hannah 2019). For
example, favoring natural regeneration and reforestation of degraded lands can
contribute to carbon sequestration, as well as to recovery of habitats and biodiver-
sity. The work of recognizing and developing biodiversity islands provides one step
in the restoration process, which can contribute to reversing isolation and increasing
biodiversity worldwide.
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 7

Natural resource managers urgently need a synthesis of our rapidly growing

understanding of these issues in order to promote strategies for biodiversity conser-
vation in human-dominated landscapes. Building on existing literature on relevant
conceptual aspects, as well as on case studies and experiences with biodiversity
conservation throughout the world, this book serves to identify and promote strat-
egies for framing and planning biodiversity islands within a landscape. We advocate
for the adoption of biodiversity islands in human modified landscapes, whether
private, public, rural, or urban. We present examples of biodiversity islands at
several different scales throughout the world and discuss potential difficulties with
the use of this framework. We expect this book to be useful to researchers, farmers,
foresters, landowners, land managers, city planners, and policymakers.

1.2 Design Considerations: Configuration, Size,

and Location in the Landscape

Design strategies for biodiversity islands depend on many factors including their
purpose, patch dynamics and spatial distribution of reserves throughout the land-
scape, the degree of site degradation, species present, and their locations within the
urban to rural spectrum. Our understanding of the relationship between spatial
distribution of reserves throughout fragmented landscapes and biodiversity conser-
vation has evolved significantly in recent years. The Theory of Island Biogeography,
developed by MacArthur and Wilson (1967), first framed the dynamics of popula-
tion ecology and speciation across fragmented landscapes of marine islands through
time around the factors of island size, distance from other islands, and methods of
species dispersal and immigration. This framework has since been applied numerous
times to terrestrial ecosystems worldwide to understand population dynamics and
diversity in areas where reserves act as “islands” throughout fragmented landscapes.
For example, Diamond (1975) recognized patch dynamics as an essential com-
ponent in reserve design. A minimum dynamic area must provide recolonization
sources for relevant species diversity to prevent extinction (Pickett and Thompson
1978). Open areas, such as pastures and annual croplands, decrease landscape
connectivity and resource availability for forest species and recolonization (Dunning
et al. 1992).
More recently however, habitat fragmentation research has gone beyond island
biogeographic theory (Laurance 2008). Island biogeography provides few predic-
tions about how community composition in fragments should change over time, and
which species would be most vulnerable. In addition, edge effects can be an
important driver of local species extinctions and ecosystem change. The matrix of
modified vegetation surrounding fragments can strongly influence fragment connec-
tivity, affecting the demography, genetics, and survival of local populations. Most
fragmented landscapes are also altered by other anthropogenic changes, such as
8 F. Montagnini et al.

hunting, logging, fires, and pollution, which can interact synergistically with habitat
fragmentation (Laurance 2008).
The “island” metaphor has been long used because of its simplicity in commu-
nicating the concept to a wide range of stakeholders. However, the concept is
complex and multifaceted (Haila 2002), and varies by location and habitat, as
different contributors of this book demonstrate in their respective chapters (Kirby
2022; Santos-Gally and Boege 2022; Montes-Londoño et al. 2022; Negret et al.
2022; Soler et al. 2022; and others).
The scale of biodiversity islands can range from square meters to thousands of
square kilometers (Table 1.1). Examples presented as case studies in this book range
from small persisting biodiverse land patches in the British countryside as described
by Kirby, to relatively larger protected areas in tropical forest locations in Costa
Rica, Peru, Ecuador as shown in contributions by Newcomer et al., Clavo Peralta
et al., Esbach et al. respectively, and several in between. There has been significant
debate around promoting increased biodiversity through few large reserves versus
more numerous small reserves (Simberloff and Abele 1982; Tjørve 2010). Research
as to whether single large or several smaller reserves (SLOSS) are superior, also
known as “the SLOSS debate,” led to the development of the Biological Dynamics
of Forest Fragments Project (formerly the Minimum Critical Size of Ecosystem
Project), which monitors features of multiple size fragmentation regimes in Amazon
forests over time (Lovejoy and Bierregaard 1990; Laurance et al. 2018).
This unresolved debate is critical for prioritizing conservation actions, particu-
larly for increasingly disturbed biodiversity hotspots. Recent research results suggest
that small patches have greater importance for biodiversity conservation than previ-
ously anticipated (Arroyo-Rodríguez et al. 2009; Hernández-Ruedas et al. 2014;
Volenec and Dobson 2019; Wintle et al. 2019). Results from the Lacandona
rainforest of Mexico, where the effect of forest patch size on species density of
different taxonomic groups was examined, support this assessment (Arroyo-
Rodríguez et al. 2022).
Recent study further supports the “habitat amount hypothesis,” which emphasizes
total habitat area within a landscape (Watling et al. 2020). This theory argues that the
species density (i.e., number of species in plots of fixed size) is more strongly and
positively related to habitat amount (e.g., forest) in the landscape than to the size of
the patch in which the plot is located (Fahrig 2013). This suggests that the greater the
total area of reserves within a fragmented landscape, the greater the potential
biodiversity, whether it be numerous small reserves, fewer large reserves, or any
mixture of these sizing parameters (Fahrig 2013, 2017).
In addition, many remaining forest patches are non-randomly distributed and
strongly correlated with topography, and soil types and quality due to human
disturbances. Many fragments are located at inaccessible or less desirable areas at
higher elevations and on steeper slopes, with varying slope aspects. The spatial
distribution of forest fragments influences tropical tree conservation, as fragment
location is the main driver of tree species maintenance within landscapes (Liu and
Slik 2014).
Table 1.1 Examples of different types of existing biodiversity islands throughout the world
Name Scale Location Landscape Species Objective Source
Islands of Several Worldwide Indigenous Native for- Sustainable food produc- United Nations IPBES
nature square territories est species tion and biodiversity (2019)
kilometers
Private For- Hectares Organic yerba mate farms, Agriculture and Native for- Sources of pollinators, bio- Montagnini et al. (2011),
est reserves Misiones, Argentina, and other, Forest est species logical pest control, water Montagnini et al. (2022)
in many locations worldwide protection, ecotourism
Islands of Hectares Santa Fe Province, Argentina Extensive indus- Exotic and Restoration of biodiversity Libertario González, pers.
resilience trial agriculture native comm. (June 2019) and
species González et al. (2022)
Dayak Hectares Indonesia Managed forest Native Long term production of Peters (2018)
gardens species crops, fruits, and timber
Forest Hectares Several locations worldwide Agricultural Native and Windbreak, connectivity Montagnini and del Fierro
windbreaks exotic (2022)
species
Hutan Desa Hectares Indonesia Native managed Native and Protection of community Moeliono et al. (2015)
forests and production resources
agricultural species
Church 3–300 ha Ethiopia Deforested land Native Fertile oasis Abbott (2019) and Baez
forests species Schon et al. (2022)
1 Introduction. Biodiversity Islands: Strategies for Conservation. . .

Religious Vary Several locations worldwide (see Native forests, Native Spiritual, conservation Several authors, see text
sites and text) lakes, rivers species
sacred
groves
Native tree Square Los Tuxtlas, Veracruz, Mexico Degraded pastures Native tree Pasture restoration and Santos-Gally et al. (2019)
Species meters restored to species productivity and Santos-Gally and
Islands Silvopastoral Boege (2022)
systems
(continued)
9
Table 1.1 (continued)
10

Name Scale Location Landscape Species Objective Source

Patches of Square Latin America / tropics Degraded agricul- Native tree Landscape restoration Holl (2002) and Holl et al.
trees meters tural or pasture species (2017)
(nucleation)
Isolated Square Valle del Cauca, Colombia, and Degraded pastures Pioneer Catalyze (jump start) Esquivel and Calle (2002),
trees in meters many other locations native tree succession others
pastures species
Edible For- Square New England, USA Urban and Native and Sustainable food produc- Toensmeier and Bates
est garden meters suburban exotic tion and biodiversity (2013) and Toensmeier
species (2022)
F. Montagnini et al.
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 11

The edge effect within the matrix influences a set of environmental conditions
that may be favorable to different species and highly detrimental to others (Tuff et al.
2016; Laurance et al. 2018). For example, Pfeifer et al. (2017) demonstrates that
edge effects can be both positive and negative, with highly endangered species
tending to show stronger negative responses to edges. Overall, increasing vegetation
cover types in the matrix is an efficient conservation strategy for maintaining higher
biodiversity levels in fragmented landscapes (Boesing et al. 2018). Also, a certain
amount of disturbance may provide additional niches for greater biodiversity to be
housed (Fahrig 2017).
In human managed landscapes, an ideal biodiversity friendly landscape could be
an area with significant continuous forest cover in a single large reserve, along with
sections of fragmented landscapes of different sized patches (Melo et al. 2013).
Land-use intensification can be mitigated by maintaining isolated trees, living
fences, and a limited number of open areas within the heterogeneous anthropogenic
matrix. Local efforts to avoid chronic disturbances and decrease further intensifica-
tion may include protection against excessive hunting, unsustainable logging, and
mismanaged firewood extraction (Fig. 1.1).

Fig. 1.1 Fragmented landscape on the road to Meru District, near Nairobi, Kenya, where the World
Agroforestry Center promotes tree planting and agroforestry systems among small farmers and
farmers’ associations to improve livelihoods, enhancing soil fertility, biodiversity and other eco-
system services ([Link]). (Photo: F. Montagnini)
12 F. Montagnini et al.

Computational methods have been developed for understanding patterns of

biodiversity throughout reserve networks. Site-selection algorithms provide compu-
tational methods to identify reserves containing as many species as possible to
maximize the representation of species (Cabeza and Moilanen 2001). Part of the
method utilizes irreplaceability indices and vulnerability indices for determining
species’ conservation value. Such innovations may prove valuable with greater
technological advancement through time.
Other factors to take into consideration in the design of biodiversity islands are
the degree of degradation and the species present in the targeted landscape. Moni-
toring of key plant and animal species is important to understanding how such
population dynamics reflect the baseline of biodiversity within the area prior to
degradation. After determining the degradation status, it is possible to decide the
most feasible and appropriate restoration pathway (Clewell et al. 2004). Strategies
should reflect past land use history and presence of damaging invasive species, when
applicable.
When biodiversity islands are designed and established to serve restoration
purposes, as in applied nucleation or tree island planting strategies, several factors
can influence their design and effectiveness. This approach is based on nucleation
theory (Yarranton and Morrison 1974), a natural recovery process where pioneer
shrubs and trees establish patchily and facilitate recruitment via enhanced seed
dispersal and improved establishment conditions. Patches spread outward clonally
and/or by facilitating the colonization of later-successional species. The use of “tree
islands” or applied nucleation has shown a great deal of promise as a restoration
approach, given that it simulates a natural recovery pattern and reduces tree planting
costs. Recently this strategy has been tested to discern to what extent restored plant
communities are similar to reference forests (Holl 2002; Holl et al. 2017). It has also
been successfully applied to restore and add biodiversity to tropical pastures (Santos-
Gally et al. 2019; Santos-Gally and Boege 2022), as well as to restore degraded
riparian and open forests in temperate regions. These nucleation strategies provide
clusters or islands of habitat, which can help to restore forests and increase their
connectivity.

1.3 Biodiversity Islands in Rural and Urban Settings

Biodiversity islands may exist anywhere within the rural to urban spectrum. In
human-dominated landscapes, the degree and matrix of fragmentation is context
specific, determined by various social and ecological dimensions. Whether urban or
rural, the protection of natural areas and restoration efforts are both needed for
further creation of biodiversity islands. The first and most crucial step in doing so
is recognizing and protecting these biodiversity islands within the landscape as
valuable assets.
Creation of government administered parks, private reserves, and voluntarily
conserved lands allow for large biodiversity islands to emerge within rural,
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 13

cultivated contexts. In addition, many farmers deliberately leave portions of original

forest cover in their properties to serve as a refuge for pollinators, to provide a source
of beneficial insects and birds that can contribute to pest control, and to protect water
sources. For example, in Misiones, Argentina, several organic farms feature these
relatively large areas that sometimes also serve ecotourism purposes (Montagnini
et al. 2011) (Table 1.1). Farmers may also have motivations such as personal
conviction along with ethical, cultural, or spiritual drivers that lead them to create
and protect reserves in their private lands.
In urban and built environments, including buildings, housing, bridges, roads,
airports and other, biodiversity islands such as urban green infrastructure, green-
ways, habitat networks, parks, and planted trees, can provide important habitat for
wildlife and other species and offer other services, such as enhanced climate
resilience. Communities can play a leading role in these efforts to ensure that they
are initiated and managed sustainably and locally.
In both rural and urban settings, relict areas that have since been engulfed by
development can function as biodiversity islands. For example, the old Governor’s
House in the center of Chennai (Madras), India has ancient forest remnants where
even a new fungus on a new insect species has been collected (David Hawksworth,
personal communication, October 2019). Similarly, the old farm houses that were
abandoned as they were surrounded by the extensive monoculture agriculture
systems that now dominate the landscape in the “pampas” or Argentinian plains
are now covered by diverse vegetation that has repopulated the area and harbors rich
fauna. Currently, researchers from INTA1 are studying the biodiversity of these
“resilience islands”, as they call them, and are proposing ways to integrate them in
the productive landscapes of the region (González et al. 2022).

1.3.1 The Contribution of Sustainable Agriculture

to Biodiversity Islands

Sustainable agricultural management techniques geared toward harmonizing eco-

system productivity and biodiversity conservation can contribute to mitigating or
reversing detrimental effects of human impacts on landscapes (Montagnini and Berg
2019). Incorporation of biodiversity friendly land uses into actively managed buffer
zones or biological corridors can contribute to the long-term conservation value of
protected areas (DeFries et al. 2007; Harvey et al. 2008; Chazdon et al. 2009; Clavo
Peralta et al. 2022; Giraldo et al. 2022; Laino et al. 2022, among others). In
landscapes lacking protected areas or intact forests, agriculture, agroforestry,

1
The National Institute for Agricultural Technology (Instituto Nacional de Tecnología
Agropecuaria, INTA) is an Argentine federal extension agency in charge of the generation,
adaptation and diffusion of technologies, knowledge and learning procedures for agriculture, forest
and agro-industrial activities within an ecologically clean environment.
14 F. Montagnini et al.

remnant vegetation, plantations, wetlands and managed forest patches provide

critical habitats and refugia for biodiversity (Harvey et al. 2006, 2008; Harvey and
González 2007; Bhagwat et al. 2008; Chazdon et al. 2009; Levin 2022a).
For example, the planting of diverse flowering insectary hedgerows may serve as
an optimal buffer zone between land sparing areas and agricultural areas on a farm.
Insectary hedgerows can offer diverse sources of pollen and nectar, creating a
balanced on-farm insect ecology inclusive of beneficial predatory insects that help
to manage pests (Chandler et al. 1998; Landis et al. 2000). Other interventions may
include agroforestry systems (AFS), which may contribute to higher species diver-
sity within a relatively homogeneous landscape (Galluzzi et al. 2010; Udawatta et al.
2019; Montagnini 2020; Montagnini and del Fierro 2022).
In recent years, the general consensus on the escalating rates of biodiversity loss
in agricultural systems has given rise to an increase in the number of international,
national and local actions related to biodiversity management in agricultural sys-
tems. A few notable initiatives among these include the World Project for Farm
Conservation at Bioversity International (formerly IPGRI, The International Plant
Genetic Resources Institute); the People, Land Management and Environmental
Change Project (PLEC); the network for Development and Conservation of Farm
Biodiversity (CBDC); the International Center for Tropical Agriculture (CIAT); the
Institute for Soil Biology and Fertility (TSBF); the World Project for Pollinators
supported by the FAO; and other projects supported by the Global Environment
Facility (GEF), among several others (Jarvis et al. 2011; Montagnini et al. 2022).
Many national and regional rural development programs also recognize the
importance of biodiversity and thus include biodiversity conservation goals, for
example, SANBIO in South Africa ([Link] which is work-
ing in several African countries on projects to improve local livelihoods. In addition,
some large World Bank-funded projects, and the Green Climate Fund pipeline
project for landscape restoration ([Link] that currently
focus more on carbon, could be (and in some cases are) incorporating explicit
biodiversity objectives.

1.3.2 The Role of Agroforestry in Biodiversity Islands

Agroforestry systems (AFS), which combine trees and crops on the same land, can
increase productivity in the short and long term while promoting biodiversity and
bringing social, environmental and economic benefits to the farmer and society
(Montagnini and Metzel 2017; Montagnini 2020; Montagnini and del Fierro
2022). AFS are often important components of buffer zones of protected areas.
Thus, they can be a great tool for biodiversity islands. The most common AFS are
shaded annual and perennial crops, silvopastoral systems, live fences, and wind-
breaks. The characteristics of the systems vary greatly according to their design,
objectives, species involved, and regions. Caution should therefore be taken when
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 15

deciding on design and management of AFS to be included as component parts of

biodiversity islands.
Several examples of indigenous multistrata AFS that are closely integrated within
the forest landscape show remarkable biodiversity (Redford and Padoch 1992).
These types of multistrata AFS may constitute a kind of biodiversity island on
their own. Alternatively, they may be integrated within biodiversity islands or may
be used to create buffer zones. Other types of multistrata AFS, including
homegardens and shaded perennial crops such as cacao, coffee and yerba mate,
can also contribute significantly to biodiversity conservation (Fig. 1.2) (Montagnini
and Berg 2019; Montagnini 2020; Montagnini and del Fierro 2022).
Likewise, silvopastoral systems (SPS) can be easily integrated with other land-
scape level strategies, such as connectivity corridors, in order to conserve biodiver-
sity and enhance other environmental services within agricultural landscapes
(Fig. 1.3). In many regions where cattle production predominates, several unpro-
tected forest remnants of high conservation value are often embedded within a
matrix of cattle grazing areas made up of pasture monocultures with few trees. In

Fig. 1.2 Homegarden and nursery in Nkenlikok, Cameroon. The World Agroforestry Centre
introduced agroforestry methods to rural farmers in the central African country some 20 years
ago as part of projects intended to ensure smallholder households increase their use of trees in
agricultural landscapes to improve food security, nutrition, income, health, shelter, social cohesion,
energy resources and environmental sustainability (Montagnini and Metzel 2017). Several tradi-
tionally used tree and other plant species are reproduced in farmers’ nurseries aiding to in-situ
conservation of local genetic resources. (Photo: F. Montagnini)
16 F. Montagnini et al.

Fig. 1.3 A traditional silvopastoral system in Indonesia: buffaloes grazing under rubber trees near
Lubuk Beringin Village, Bungo Regency, Jambi Province, Sumatra. The first village forest (Hutan
Desa) with 2356 ha was inaugurated and designated in 2009 under the management of Lubuk
Beringin village administration. Under the Forestry Minister Regulation No. p49/Menhut-II/2008,
village communities can be granted legal right to manage state forests for their own prosperity
(World Agroforestry Centre (ICRAF), Bogor (Indonesia) 2010). Livestock in small farms in the
village include goats, sheep, chickens and buffalo. (Photo: F. Montagnini)

such areas, SPS with complex vegetation structures can support important levels of
biodiversity (Harvey et al. 2005, 2006; Sáenz et al. 2007) and provide ecosystem
services such as natural pest management, carbon sequestration, water and soil
conservation, nutrient cycling, hydrological protection, and crop pollination
(Chazdon et al. 2009; Calle et al. 2010). Hence it is possible to enhance biodiversity
by strategically placing elements such as live fences, scattered trees, riparian buffers,
and connectivity corridors within the landscape (Esquivel and Calle 2002;
Murgueitio et al. 2011; Calle and Holl 2019; Santos-Gally et al. 2019; Calle et al.
2022; Giraldo et al. 2022; Santos-Gally and Boege 2022) (Table 1.1).
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 17

Windbreaks and living fences are examples of AFS that connect portions of
remnant forests in agricultural landscapes (Francesconi et al. 2011). Windbreaks
tend to be favored by farmers and can be instrumental in biodiversity conservation
and landscape connectivity in fragmented areas. Windbreaks therefore have the
potential to play similar roles in the connectivity of biodiversity islands across a
landscape (Montagnini 2020; Montagnini and del Fierro 2022).
The low intensive management multistrata AFS with native species have the
greatest potential to harbor high amounts of biodiversity. Biodiversity decreases as
management intensity increases in more homogeneous tree crop/animal combina-
tions. However, even the less heterogeneous types of AFS provide greater biodiver-
sity than would otherwise be realized in conventional monoculture agriculture or in
degraded landscapes. In addition, farmers value and protect AFS because of their
contributions to their livelihoods, thus ensuring the conservation of services they
provide, including biodiversity (Montagnini 2020; Montagnini and del Fierro 2022).

1.3.3 Biodiversity Islands in Urban Environments

Urban areas are highly modified and complex landscapes, within which green or
open areas are valuable for human well-being as well as wildlife (Pickett et al. 2004).
Urban Green Infrastructure (UGI) provides important contributions to the sustain-
ability of urban systems and offers a nature-based solution to reduce the impact of
rapid urbanization, a consequence of the increased rates of rural to urban migrations
happening today worldwide. Urban species diversity and richness and the use of
green spaces for biodiversity conservation are of paramount importance for devel-
oping climate-resilient cities (Biazen 2015). The UGI may include several types of
vegetation assemblages in parks, playgrounds, sidewalk tree plantings and even
cemeteries when specific efforts are made to increase their aesthetic value to visitors
([Link]
diverse-plants-animals/).
Indicators of the size of the UGI, such as the amount of green space per person,
are used by funding institutions such as the Interamerican Development Bank
(IADB, [Link] as criteria to decide on financing specific urban pro-
jects. For example, the Emergent and Sustainable Cities IADB initiative has desig-
nated the city of Rosario, Argentina as the greenest city of the country, with an
average of 12 m2 of green space per inhabitant. In contrast, in the capital city of
Buenos Aires, there are only 3.5 green m2 per person ([Link]
mundo/destinos/argentina/rosario-fue-declarada-la-ciudad-mas-ecologica-de-argen
tina/). The city of Rosario, which has a population of about 1.2 million covers a total
of 178 Km2, with a total of 2070 streets, avenues and short alleys (Fig. 1.4). There
are a total of 250 green municipal areas, with the Independence Park (Parque de la
Independencia) of a total of 126 ha standing out for its floristic richness. The growth
of urban vegetation has increased over the last three decades thanks to tree planting
as well as the creation of new green areas ([Link]
18 F. Montagnini et al.

Fig. 1.4 View of downtown Rosario, Argentina showing flowering trees and other vegetation of
the abundant urban green infrastructure. (Photo: “Carácter” by Sebastian Infante; www.
[Link]). This picture won the first prize in a contest organized in November 2019
by Aehgar, Asociación Empresaria Hotelera Gastronómica y Afines Rosario (Hotels, Gastronomy
and Related Empresarial Association of Rosario)

Cities provide habitats for a rich and diverse range of plants and animals.
Enhancement of biodiversity in urban ecosystems can have a positive impact on
the quality of life and education of urban dwellers and thus facilitate the preservation
of biodiversity in natural ecosystems. An essential first step to managing biodiversity
in urban environments more effectively is to gain a full understanding of the
interplay between landscape (matrix effects) and local factors (patch effects) that
affect urban biodiversity (Savard et al. 2000).
Many cities have a network of habitat fragments or urban greenways comprising
areas of semi-natural habitats, secondary succession, ruderal and pioneer environ-
ments and open areas. These habitats may be important features for biodiversity both
as stable and as transient habitats (McIntyre et al. 2001), and may also be valuable
for their possible function as corridors and stepping stones to facilitate species
dispersal.
In urban landscape planning, urban greenways and wildlife corridors are increas-
ingly advocated for in order to encourage animals and plants to move around urban
areas, thereby preserving or enhancing urban biodiversity. Urban planning needs to
consider the design and establishment of greenways to serve as dispersal route ways
as well as habitat, providing a chain of different habitats that permeate the urban
environment. City planners can have a positive impact on urban biodiversity by
slowing the pace of redevelopment (Angold et al. 2006).
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 19

The establishment of parks and the integration of ecological landscape design

bring increased biodiversity to suburban and urban settings. Actively designing or
setting aside areas within an urban park to serve as biodiversity islands can contrib-
ute significantly to biodiversity conservation. Some community led efforts, sustained
and managed locally, can also integrate biodiversity islands and make substantial
contributions to biodiversity conservation. For example, in Philadelphia, the long-
established Fairmont Park contains a wide range of species and habitats such as
pollinator and butterfly gardens within the extensive urban forests following the
Schuylkill River and Wissahickon Creek ([Link] Similarly, the
Urban Oases in the New Haven Watershed program in New Haven, CT bring
together a variety of community partners to establish a network of green spaces
for wildlife and community members at parks throughout the city (http://
[Link]/).
Planting trees and increasing vegetation in cities can create small scale biodiver-
sity islands. These efforts also have social and educational value. For example, the
Urban Resources Initiative, a university non-profit collaboration based in New
Haven, CT, USA, is one of many likeminded organizations that plants trees while
working with the local community, thereby increasing habitat for biodiversity
([Link]
Similarly, in Philadelphia, PA, USA, Philly Peace Park is an example of a
neighborhood-managed ecology campus championing sustainable production of
food while fostering education and community involvement (www.
[Link]). As another example, the small town of Salliqueló in the
Buenos Aires Province in Argentina has carried out a tree-planting program called
“Nace un niño, nace un árbol” (“A child is born, a tree is born”) for 24 years, with
over 3000 trees planted so far ([Link]. Salliqueló, una ciudad que piensa
en verde).
In urban settings, gardens can play a variety of roles as biodiversity islands at
different scales, including at the landscape level, contributing to conservation of
forest habitat and creating new habitats for other species. In many cases this is
attained by improving connectivity between remaining natural forest fragments
within city limits, thereby creating “stepping stones” across the urban landscape
and reducing the edge-effect at forest boundaries (Negret et al. 2022; Toensmeier
2022). Urban green areas potentially serve as venues for promoting not only local
biodiversity conservation but also novel species assemblages that are different from
those found in pristine areas (Soler et al. 2022).

1.4 Landscape Management Approaches

for the Implementation of Biodiversity Islands

Biodiversity islands can exist in a wide range of human-dominated landscapes. They

may be actively implemented as part of a complex landscape design that may include
agriculture, forests, plantations, wetlands or other uses, or they may be the result of a
20 F. Montagnini et al.

passive management practice, i.e., as part of the landscapes that were left untouched
for practical or other reasons. The method of implementation is important, as it will
determine the characteristics, position in the landscape, and management of the
biodiversity island.
Land management decision-making may take many different forms. Public,
private, and communal land each involve different stakeholders as key decision
makers who may choose to implement biodiversity islands. An integrated landscape
approach is needed to understand the effects of landscape structure and dynamics on
conservation of biodiversity, provision of ecosystem services, and sustainability of
rural livelihoods (Tscharntke et al. 2005; Chazdon et al. 2009). Integrated landscape
management (ILM) is an approach inclusive of various stakeholders that can be
instrumental in furthering the creation of biodiversity islands. ILM supports engage-
ment across sectors and scales, increases broad multilateral coordination, and
ensures harmonization of planning, implementation and monitoring processes at
the landscape, regional and national levels (Thaxton et al. 2016).
With the ILM approach, landscapes can be designed and managed to secure a
future for both biodiversity and rural livelihoods. This rationale can be used in
planning strategies to promote the adoption and development of biodiversity islands.
Landowners can integrate small scale land sparing to set aside pieces of the property
as untouched natural settings that can act as biodiversity islands. On the other hand,
when coexistence is identified as an end goal, land sharing may allow wildlife to
thrive in human-dominated landscapes (Crespin and Simonetti 2019). While land
sparing (e.g., protected areas) is needed to protect biodiversity, land sharing (e.g.,
agroforestry systems) may contribute to the maintenance of ecological services. The
debate between land sparing and sharing continues, with several authors advocating
for the land sparing alternative for the sake of conservation (Phalan 2018; Phalan
et al. 2011). For a given area, the question could be resolved with consideration of
site-specific conditions of habitat disturbance and development, e.g., conventional
pastures versus degraded lands versus improved sustainable cattle raising versus
reserves. Both land sparing and land sharing as complementary strategies can
provide valuable protection of species diversity over time (Grass et al. 2019).
Strategies to promote biodiversity islands at the landscape level may include
mitigating threats to biodiversity loss in agricultural landscapes; protecting, diversi-
fying, and sustainably managing tree cover within an agricultural matrix, including
all types of agroforestry systems (AFS); promoting and conserving indigenous,
traditional, and ecologically based agricultural practices; and restoring degraded
lands (Harvey et al. 2006, 2008; Chazdon et al. 2009).
The notion of a “landscape approach” is not new, but in recent years it has gained
in importance and is a major topic of national and international policy discourse
(Denier et al. 2015). The landscape may be the most appropriate scale for action
between national and local scales. A landscape approach, using ILM, can allow
stakeholders to decide on land and water use in such a manner that community,
commercial and conservation interests are more balanced and sustainable. ILM arose
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 21

from diverse, innovative strategies – from indigenous territorial development, to

integrated watershed management, to landcare. It involves new levels of collabora-
tion through place-based partnerships that are inclusive of communities, govern-
ments, businesses, and land managers (Montagnini et al. 2022).
Many examples of applications of ILM with a strong biodiversity focus have been
documented recently (Denier et al. 2015). For example, in northwest Vietnam, the
Australian Centre for International Agricultural Research (AGIAR) and the Consor-
tium of International Agricultural Research Centers (CGIAR) have introduced
agroforestry techniques into monocropped landscapes at the farm level, while also
providing techniques for the expansion of AFS to the landscape level. The tech-
niques include “training of trainers,” championing farmers, organizing farmer field
days and setting up community tree nurseries. Agroforestry was scaled up in
collaboration with provincial governments and local farmers. The project not only
diversifies income at the farm level, but also provides important environmental
services at the landscape scale, such as reduced pressure on forests for timber,
reduced soil erosion and protection against storms ([Link]
org/project/agroforestry-livelihoods-smallholder-farmers-northwest-viet-nam).
The landscape approach to development and conservation is proliferating world-
wide, particularly in the past five years (Sara Scherr, EcoAgriculture Partners,
personal communication, January 2020). ILM initiatives in Africa, Latin America/
Caribbean and South/Southeast Asia have inspired the creation of new partnerships
that emphasize biodiversity (Denier et al. 2015). International conservation-oriented
organizations working in this field include, among others, the African Wildlife
Foundation ([Link] which uses a landscape approach to conserva-
tion to improve the livelihoods of local people, and the International Union for
Conservation of Nature (IUCN)‘s Livelihoods and Landscapes Strategies, which
address human and environmental needs in large areas of land with a special
emphasis on the sustainable use of forests ([Link]
countries/thailand/livelihoods-and-landscapes-strategies).
The new alliances inspired by the ILM approach promote collaboration in
research, in the co-design of conservation programs and policies, and in management
of human-modified landscapes. This can be accomplished using participatory and
multidisciplinary approaches in research and management (Chazdon et al. 2009;
Esbach et al. 2022; Newcomer et al. 2022; Painter et al. 2022; Vásquez et al. 2022).
In addition, socioeconomic, legal, and political actions can be used to promote
biodiversity islands at the landscape level. These may include, for example, the use
of economic instruments, such as payments for ecosystem services, as discussed in
Sect. 1.6 of this chapter, or improving environmental laws and enforcement to
reduce deforestation, regulate logging, conserve on-farm tree cover, and reduce
agrochemical use (Sheban 2022; Sigman 2022). Action can also be taken to promote
ecologically sustainable production systems such as agroforestry or to leverage local
and regional political support for existing initiatives for biodiversity protection
(Newcomer et al. 2022).
22 F. Montagnini et al.

1.5 How Community Led Action Can Advance

the Development of Biodiversity Islands

Community led action can contribute to the development of biodiversity islands.

Motivations for such action may be inherent within cultures or learned and applied
with ethical, philosophical, or scientific motivations. Land access and land tenure
provide the basis for community led efforts, while other powerful instruments and
legal tools to be used in service of conservation include education, political engage-
ment, and conservation easements. Community developed biodiversity islands may
be governed as public lands, commons, cooperatives, or through tactics of commu-
nity based conservation, in which social complexities are appreciated and demo-
cratic decision-making processes are used. Examples of grassroots community
action for the advancement of conservation practices are numerous, diverse, and
worldwide (Otto et al. 2013; Levin 2022b; Morales-Nieves 2022). Both non-profit
and for-profit organizations are capable of further expanding community led action
for the development of biodiversity islands.
The types of legal frameworks, methods of enforcement, modes of implementa-
tion, and levels of community engagement that can be used to protect and manage
biodiversity islands are site and context dependent (Levin 2022b). Conservation
easements are one such tool that may be advocated for by a community for the
development of biodiversity islands. Conservation easements are voluntary legal
agreements between landowners and a land trust or government agency that protect
conservation values on a property by permanently limiting uses of the land and
offering tax incentives to the land owner. A land trust is a non-profit organization
that acquires land or conservation easements through support from donations or
government funding. In many instances, a land trust can also act as a conservation
organization to help draft, implement, and ensure compliance with the easement.
Conservation easements may include land uses such as recreation, forestry,
agroforestry, or agriculture. These easements, which provide conservation value in
addition to opportunities for revenue generation, are known as working land con-
servation easements. Throughout the world, communities can support and develop
local land trusts to assist in the development of biodiversity islands by supporting
landowners in putting their land into conservation. One of many such examples is
the Harrison Family of North Branford, CT, who recently transferred ownership of
their 8.5 ha property to the North Branford Land Conservation Trust (NBLCT)
([Link]). The family donated the land, although it was prime for
real estate development, so that it could be protected and preserved in its natural state
in perpetuity (Figs. 1.5 and 1.6) (Totoket Times 2018). As such, the Harrison Farm,
now under the care and maintenance of NBLCT, serves as a true biodiversity island
in an otherwise suburban landscape.
Religious sites and sacred groves are other examples where community led action
protects biodiversity rich sites on a landscape in ways that can be considered
biodiversity islands. Examples include the Mizorom sacred groves in Northeastern
India, sacred pools protected by Tchabè communities in Central Benin, sacred cacao
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 23

Fig. 1.5 A section of forest recently donated to the North Branford Land Conservation Trust
(NBLCT) by the Harrison Family and protected in perpetuity by a conservation easement. (Photo:
F. Montagnini)

groves of the Maya, and other sacred groves in Zimbabwe, Ghana, Thailand, China,
and Nepal (Gómez-Pompa et al. 1990; Gadgil et al. 1993; Bhagwat and Rutte 2006;
Ceperley et al. 2010). One such successful example is church forests in Ethiopia
(Table 1.1), which are small pockets of protected forest surrounding orthodox
churches. They number over 35,000 and are spread all over the country. These
small but fertile oases, which range from 3 to 300 ha, are some of the last remnants of
the tall, lush native forests that once covered Ethiopia, and which, along with their
biodiversity, have all disappeared (Abbott 2019; Baez Schon et al. 2022).
Other community protected sites include the village forests in Indonesia, known
as Hutan Desa, which are legally recognized for the ecosystem services and benefits
to society they provide. Their management and protection are guided by traditional
communal governance as well (Moeliono et al. 2015). Although the original purpose
in most cases was not to create biodiversity islands, they may be developed as a
consequence of such community actions.
There are numerous examples of organized community efforts to support biodi-
versity outcomes that align with the creation of biodiversity islands (UNDP 2006;
Levin 2022b). From the Zapatistas farmers movement in southern Mexico (Esteva
1999), to community restoration projects in Northern Ethiopia (Ethiopian Biodiver-
sity Institute 2014; Sigman 2022), to the newly emerging global concept of Ecosys-
tem Restoration Camps ([Link] rural
24 F. Montagnini et al.

Fig. 1.6 The Harrison Family during a ribbon cutting ceremony at the newly created Harrison
Farm Preserve, celebrating the transfer of ownership from the family to North Branford Land
Conservation Trust (NBLCT). (Photo: F. Montagnini)

community action provides alternative methods for enhancing biodiversity out-

comes in rural settings (Nazarea et al. 2014).
Non-profit organizations such as EcoAgriculture Partners ([Link]
org/) and The Forests Dialogue ([Link] aim to help engage
communities for biodiversity conservation in rural settings through the ILM
approach previously described. For example, in their new initiative, “1000 Land-
scapes for 1 Billion People. A Radical Collaboration for Resilient Communities and
Restored Nature”, EcoAgriculture supports locally led initiatives, for example, via
their Landscape Action Platform co-designed with communities (Concept Note for
Discussion. EcoAgriculture Partners and Rainforest Alliance, September 2019).
Numerous other international non-profit organizations, such as Commonland
([Link] Conservation International ([Link]
[Link]/), Solidaridad ([Link] World Wildlife
Fund ([Link] and Wetlands International ([Link]
[Link]/), just to name a few, promote practical strategies for local use of
natural resources in harmony with biodiversity conservation in many of their pro-
jects around the world. Likewise, networks of landscape initiatives, for example, the
International Model Forest Network ([Link] and the Satoyama Initiative
([Link] work with and connect member organizations
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 25

worldwide to promote ways for local communities to manage their natural environ-
ment for production activities, to nurture their traditions and culture, and to maintain
ecosystems and biodiversity.
In suburban settings, parks, home gardens, community garden projects, and
ecovillages also act as biodiversity islands. In the urban context, the rise of urban
forestry, parks, urban community gardens, and educational centers which support
these ends continue to grow in popularity. Along this wide spectrum of population
density, there is great potential for the continuation of community led action to
promote and sustain biodiversity islands.

1.6 The Way Forward: Valuing and Financing Biodiversity

Conservation

Biodiversity conservation is in great peril due to significant gaps in what is available

and what is needed for financing conservation in a feasible and tangible way.
According to current estimates, for example, conserving biodiversity to meet the
Aichi targets and other conservation efforts worldwide would require between $150
and $440 billion (CBD High-Level Panel 2014; Credit Suisse et al. 2014); with some
estimates as high as $977 billion (Talberth and Gray 2012). Meanwhile, only about
$30 billion has been spent on biodiversity in the 2010 decade (i.e., only 3+ billion
dollars/year) via different mechanisms, such as biodiversity-relevant taxes, PES
programs, biodiversity offsets, and bilateral biodiversity-relevant overseas develop-
ment assistance (OECD 2018). Other studies including other instruments estimate
spending between $39 and $49 billion annually (OECD 2019), with some as high as
$50 billion (Parker et al. 2012).
There are many specific examples throughout the world that serve to illustrate the
urgent need to find alternative feasible ways to finance conservation efforts such as
biodiversity islands. Biodiversity alone is not easy to finance; however, it can be
financed through strategies that incorporate other valued resources or ecosystem
services such as water or carbon (Sheban 2022). When businesses and communities
recognize these integrated benefits as essential to their own supply chains, resource
bases, and human health, financing becomes more tangible.
The valuation of ecosystem services is a first step in financing biodiversity
conservation. Assessment of ecosystem services varies among spatial scales, land-
scapes, and interactions with stakeholders. Proper valuation is critical for developing
programs that seek to compensate farmers and other actors for the environmental
services and benefits they provide, including biodiversity (Pagiola et al. 2017). Such
programs may include Payments for Environmental Services (PES), which encour-
age projects that enhance conservation, restoration, production, and rural develop-
ment via compensation (Montagnini and Finney 2011).
Recent decades have witnessed a considerable increase in PES programs that pay
or compensate land users financially for land management practices intended to
26 F. Montagnini et al.

provide or ensure ecosystem services, with over 550 active programs around the
globe and an estimated US$36–42 billion in annual transactions (Salzman et al.
2018). The biodiversity and habitat PES sector uses offsets directed to ensure habitat
and biodiversity protection. This sector remains the least developed in terms of
geographical scope and is most challenging for countries to put in place. Mitigation
credit banks are growing but primarily in developed countries, with a few examples
also in developing countries. With transactions estimated at US$3.6 billion per year,
compensatory mitigation banking continues to grow. In developing countries, mit-
igation carried out directly by the party producing the impact or by a subcontractor,
known as ‘permittee-responsible mitigation’, is the most commonly found option for
compliance, although many allow developers to pay a compensation fee in lieu of an
offset, which is generally used to fund conservation projects carried out by the public
sector or an NGO. Voluntary biodiversity offsets are a recent policy development
and remain small (Salzman et al. 2018).
Unlike in water PES for which the beneficiaries are straightforward and local, the
beneficiaries of biodiversity are often widespread, and the specific benefits are
indirect or non-material. Institutions comparable to water utilities that can collect
fees on behalf of many beneficiaries do not exist, and common metrics are difficult to
determine. As a result, biodiversity PES programs remain limited (Salzman et al.
2018).
Compared to PES systems that include only one environmental service, systems
that incorporate bundling or layering of multiple services can make sustainable land
uses more attractive to farmers and reduce perverse incentives (Montagnini and
Finney 2011). These valuation systems that incorporate bundling of multiple ser-
vices may protect biodiversity even when PES are given for the provision of some
other ecosystem service such as water or carbon. For example, various land use types
can be assigned an “Environmental Service Index” based on tree cover and habitat
type, bundling biodiversity and carbon (Montagnini and Finney 2011). Under the
provision of external funding, a payment can be estimated for each land use type to
compensate farmer participation and thus encourage establishment of biodiversity
islands on farms. Bundling services can protect biodiversity islands much like the
market mechanisms supporting the protection of the Panama Canal watershed
(Adamowicz et al. 2019).
Perhaps one of the greatest opportunities for biodiversity conservation is to
incorporate biodiversity objectives into the new generation of climate change action
programs, for which funds are increasingly available. While the language currently
places the emphasis on climate change mitigation and adaptation with terms such as
“climate-smart agriculture” or “climate-smart landscapes” many of these programs
can and do incorporate multiple ecological objectives, beyond just climate.
In many jurisdictions, taxes levied on activities detrimental to ecosystems world-
wide can be used to share the costs of environmental services such as carbon storage
and biodiversity. As an example, Sweden recently introduced emissions taxes on
airline travel to discourage the use of fossil fuels (Nordic Business Insider 2018). A
fair and equitable distribution could divide the proceeds proportionately between
countries that are exposed to the air travel emissions. These funds could be used to
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 27

finance biodiversity islands, preserves and carbon sequestration in affected

countries.
In contrast to the taxed approach, industries may recognize the importance of self
regulation to address environmental degradation of their practices. Continuing with
the example of the aviation industry, a global climate agreement has been reached,
CORSIA (Carbon Offsetting and Reduction Scheme for International Aviation),
which focuses on investing in emission reductions in unrelated fields to compensate
for airline emissions that exceed the 2020 Aichi Targets ([Link]
policy/environment/Pages/[Link]). World ecosystem services could be greatly
enhanced if these types of funds and efforts to offset excessive emissions focused on
supporting carbon sequestration and biodiversity protection through biodiversity
islands and preserves.
Likewise, worldwide investments aligned with positive environmental outcomes
are gaining popularity. For example, Norway’s $1 Trillion pension fund has com-
mitted to zero deforestation in its public procurements (Rainforest Foundation
Norway 2016). Similar initiatives could be applied in other parts of the world,
perhaps with a focus on a more equitable allocation of emissions tax revenues
towards protecting natural resources.
The integration of biodiversity goals in agricultural and other supply chains
(including even products for high fashion, sophisticated consumer markets, govern-
ment procurement, etc.) is gaining momentum as well (see examples in Montagnini
and Metzel 2017; Montes-Londoño et al. 2022; Newcomer et al. 2022; Sheban
2022). In this context, the Coalition for Private Investment in Conservation (CPIC)
works to support and scale biodiversity-friendly private investment (http://
cpicfi[Link]/). In its Statement of Intent, CPIC affirms the need for “investment
opportunities that provide measurable, science-based conservation benefits and
social impact to participating communities and to biodiversity, while delivering
at-scale financial returns for investors.” To address the current gap in low invest-
ment, inadequate returns, and lack of coordinated knowledge around scalable
investment, CPIC calls for “a concerted, systematic effort focused on creating
investment products that provide a conservation and financial bottom line. . .”
(cpicfi[Link]/wp-content/uploads/2017/05/05_10_17_CPIC-Statement-of-Intent-
[Link]). The Statement is endorsed by numerous NGOs and public and private
entities who have agreed to be part of this collaborative effort.
Because it takes advantage of smaller parcels or fragments of land, the concept of
biodiversity islands may be more amenable to funding allocations by governments
and policymakers than more conventional sparing approaches that dedicate large
areas of land to biodiversity conservation. In situations when governments feel
pressured to develop lands to satisfy economic needs of the population, dedicating
relatively smaller areas such as biodiversity islands to conservation, instead of
designating them for strictly economic uses, may seem more feasible (Montagnini
et al. 2022).
28 F. Montagnini et al.

1.7 Conclusions: Potential for Recognizing and Integrating

Biodiversity Islands in Human-Dominated Landscapes

Based on the evidence and experiences presented in this book, there is great potential
for recognizing and further integrating biodiversity islands in human-dominated
landscapes to achieve positive economic, ecological, and social outcomes. Design
strategies for biodiversity islands are dependent on landscape use within the matrix
of habitat fragmentation. Integrated landscape management (ILM), including ele-
ments of sustainable agriculture, agroforestry and community led action, may
provide the guiding framework for implementation of biodiversity islands in com-
plex landscape matrices involving a variety of stakeholders. Land use systems made
up of complex assemblages embedded in a forestry matrix, as is the case in many
traditional indigenous sacred sites and agroforestry systems, can be considered
biodiversity islands as well.
In more degraded landscapes, mixed production systems can act as buffer zones
surrounding preserved sites, protecting biodiversity islands or serving as biodiversity
islands themselves. Restructuring financial systems to value strategies that protect
ecosystem services can enable individuals, businesses, and communities to carry out
actions to support the establishment and preservation of biodiversity islands. While
financial incentives are crucial, still, other motivations such as personal conviction
along with ethical, cultural, or spiritual drivers may prevail in fostering actions
towards the creation and protection of biodiversity islands over time.
As with other conservation and restoration initiatives, there are some inherent
potential difficulties in the use of this framework. The configuration, design, and
other factors previously discussed influence the effectiveness of biodiversity islands.
For example, sometimes biodiversity islands may be created to protect target
species, disregarding other species with less mobility or specific resource
requirements.
In addition, not all biodiversity islands have the same conservation value. It
would be useful to prioritize to some degree the conservation value of different
biodiversity islands. For instance, large reserves may not have the same value as
isolated trees in urban areas or living fences in agricultural lands. Conservation plans
can be misguided without this prioritization, e.g. a landowner may cut his or her
forest, maintaining some isolated trees or living fences because the latter are also
classified as biodiversity islands. In fact, biodiversity islands should be prioritized
within each context where they may be applied, whether that be urban, rural,
wetlands, etc. A forest in a city cannot be compared to a forest in a rural setting.
This prioritization should be done by the respective local planners in each specific
case, e.g., municipal authorities, local or international NGOs, park and recreation
agencies, etc., with consideration of the location’s specific biodiversity goals and
constraints.
Moreover, depending on the social and economic context, emphasizing biodiver-
sity conservation may be prone to criticism by those who place higher value on the
provision of certain services for society. On the other hand, biodiversity friendly
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 29

land uses, such as biodiversity islands themselves or agroforestry systems, may be

seen by many as enough of a contribution to biodiversity conservation, thereby
excluding or replacing other necessary efforts.
Another challenge of biodiversity islands is that because of their tendency to be
small and few within a larger landscape, climate change may pose a serious problem
to their viability as biodiversity habitat. Parks and protected areas are already facing
this challenge as the habitat for species of critical interest shifts with climate change.
Finally, many may argue in favor of the growing trend of recognizing the rights of
nature independent of the services it provides, for which biodiversity islands may be
used to protect nature regardless of its purpose or potential uses (Prieto Méndez
2013). The growing international movement around the rights of nature provides an
alternative justification for biodiversity islands based on the inalienable right of
nature to exist, which goes beyond the more anthropocentric rationale based on the
ecosystem services they provide ([Link]
nature/).
The purpose of this book is to present examples and case studies where the
biodiversity islands approach is being used in a variety of locations and contexts
worldwide. This book contributes to informing design parameters on appropriate
sizing and spatial distribution of biodiversity islands in order to be effective in
conservation and regeneration across the landscape, using integrated landscape
management approaches. The book is organized in five parts: I. Introduction;
II. Biodiversity Islands Establishment and Management: Challenges and Alter-
natives; III. Biodiversity Islands across the Globe: Case Studies;
IV. Safeguarding the Environmental, Economic, and Social Benefits of Biodi-
versity Islands; and V. Conclusions.
The contributing authors present views from the academic, the practitioner and
the policymaker perspectives, offering alternatives and suggestions for promoting
strategies that support biodiversity conservation through intentionally designed
frameworks for sustainable human-dominated landscapes. This book provides sug-
gestions and concrete examples that can be used by a variety of stakeholders in
various settings throughout the world. In all, this book covers examples and case
studies from 16 countries, including 10 from Latin America (Argentina, Brazil,
Colombia, Costa Rica, Ecuador, Mexico, Panama, Paraguay, Peru and Puerto
Rico), as well as from the USA, Sweden, Finland, Germany, France, India, Indone-
sia, Cameroon, Kenya, Ethiopia and others. This book is useful to researchers,
farmers, foresters, landowners, land managers, city planners, and policy makers
alike.

Acknowledgements The authors thank Rafael Herrera, Joseph Orefice, Victor Arroyo-Rodríguez,
Sara Scherr and two anonymous reviewers who offered constructive and useful criticism on this
chapter. A Conference Session entitled: Biodiversity Islands: Pockets of Protected Land in Human
Dominated Environments, that the authors led as part of the IUFRO Conference “Adaptive
Management for Forested Landscapes in Transformation”, Posadas, Misiones, Argentina, October
1, 2018, inspired and initiated this chapter as well as other contributions contained in this book.
30 F. Montagnini et al.

References

Abbott A (2019) Biodiversity thrives in Ethiopia’s church forests. Nature 565:548–549

Adamowicz W, Calderon-Etter L, Entem A, Fenichel EP, Hall JS, Lloyd-Smith P, Ogden FL,
Regina JA, Rad MR, Stallard RF (2019) Assessing ecological infrastructure investments. Proc
Nat Acad Sci. [Link]
Angold PG, Sadler JP, Hill MO, Pullin A, Rushton S, Austin K, Small E, Wood B, Wadsworth R,
Sanderson R, Thompson K (2006) Biodiversity in urban habitat patches. Sci Total Environ 360:
196–204
Arroyo-Rodríguez V, Pineda E, Escobar F, Benítez-Malvido J (2009) Value of small patches in the
conservation of plant-species diversity in highly fragmented rainforest. Conserv Bio 23:729–
739
Arroyo-Rodríguez V, Arasa-Gisbert R, Arce-Peña N, Cervantes-López MJ, Cudney-Valenzuela SJ,
Galán-Acedo C, Hernández-Ruedas MA, San-José M (2022) The importance of small rainforest
patches for biodiversity conservation: a multi-taxonomic assessment in the Lacandona region,
Mexico. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in biodiversity and conservation. Springer, Cham,
pp 41–60
Baez Schon M, Woods CL, Cardelús CL (2022) Sacred church forests in northern Ethiopia:
biodiversity and Cultural Islands. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments, Series: Topics in Biodiversity and Conserva-
tion. Springer, Cham, pp 531–549
Bhagwat SA, Rutte C (2006) Sacred groves: potential for biodiversity management. Frontiers Ecol
Environ 4(10):519–524
Bhagwat SA, Willis KJ, Birks HJB, Whittaker RJ (2008) Agroforestry: a refuge for tropical
biodiversity? Trends Ecol Evol 23:261–267
Biazen M (2015) The value of urban green infrastructure and its environmental response in urban
ecosystem: a systematic review. Inter J Environ Sci 4(2):89–101
Boesing AL, Nichols E, Metzger JP (2018) Biodiversity extinction thresholds are modulated by
matrix type. Ecography 41(9):1520–1533
Cabeza M, Moilanen A (2001) Design of reserve networks and the persistence of biodiversity.
Trends Ecol Evol 16(5):242–248
Calle A, Holl KD (2019) Riparian forest recovery following a decade of cattle exclusion in the
Colombian Andes. For Ecol Manag 452. [Link]
Calle Z, Guariguata MR, Giraldo E, Chará J (2010) La producción de maracuyá (Passiflora edulis)
en Colombia: perspectivas para la conservación del hábitat a través del servicio de polinización.
Interciencia 35(3):207–212
Calle Z, Molina CCH, Molina DCH, Molina DEJ, Molina EJJ, Murgueitio B, Murgueitio A,
Murgueitio E (2022) A highly productive Biodiversity Island within a monoculture landscape:
El Hatico nature reserve (Valle del Cauca, Colombia). In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in biodi-
versity and conservation. Springer, Cham, pp 279–304
CBD High-Level Panel (2014) Resourcing the Aichi Biodiversity Targets: an assessment of
benefits, investments and resource needs for implementing the strategic plan for biodiversity
2011–2020. Second report of the high-level panel on global assessment of resources for
implementing the strategic plan for biodiversity 2011–2020. Montreal, Canada [Link]
[Link]/financial/hlp/doc/[Link]
Ceperley N, Montagnini F, Natta A (2010) Significance of sacred sites for riparian forest conser-
vation in Central Benin. Bois Forêts Tropiques 303:5–23
Chandler J, Corbett A, Lamb C, Long R, Reberg-Horton C, Stimmann M (1998) Beneficial insects
move from flowering plants to nearby crops. Calif Agric 52:23–26
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 31

Chazdon RL, Harvey C, Komar O, Griffith DM, Ferguson BG, Martínez-Ramos M, Morales H,
Nigh R, Soto-Pinto L, van Bruegel M, Philpott S (2009) Beyond reserves: a research agenda for
conserving biodiversity in human-modified tropical landscapes. Biotropica 41(2):142–153
Clavo Peralta ZM, Vela Alvarado JW, Alvez-Valles CM (2022) Biodiversity islands and dominant
species in agricultural landscapes of the Western South Amazon, Peru. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in biodiversity and conservation. Springer, Cham, pp 207–235
Clewell A, Aronson J, Winterhalder K (2004) The SER international primer on ecological resto-
ration. Society for Ecological Restoration International Science & Policy Working Group
(Version 2: October, 2004) [Link]
Conservation International (2014) Hotspots: targeted investment in nature’s most important places.
[Link] Retrieved 1 Sep 2018
Costanza R, de Groot R, Sutton P, van der Ploeg S, Anderson SJ, Kubiszewski I, Farber S, Turner
RK (2014) Changes in the global value of ecosystem services. Glob Environ Chang 26:152–158
Credit Suisse, World Wildlife Fund, McKinsey & Company (2014) Conservation finance: moving
beyond donor funding toward an investor-driven approach. [Link]
media/cc/docs/responsibility/conservation-fi[Link]. Retrieved 1 Nov 2018
Crespin SJ, Simonetti JA (2019) Reconciling farming and wild nature: integrating human–wildlife
coexistence into the land-sharing and land-sparing framework. Ambio 48:131–138. [Link]
org/10.1007/s13280-018-1059-2
DeFries R, Hansen A, Turner BL, Reid R, Liu JG (2007) Land use change around protected areas:
management to balance human needs and ecological function. Ecol Appl 17:1031–1038
Denier L, Scherr S, Shames S, Chatterton P, Hovani L, Stam N (2015) The little sustainable
landscapes book. Global Canopy Programme, Oxford
Diamond JM (1975) The island dilemma: lessons of modern biogeographic studies for the design of
natural reserves. Biol Conserv 7(2):129–146
Dunning JB, Danielson BJ, Pulliam HR (1992) Ecological processes that affect populations in
complex landscapes. Oikos 65:169–175
Eibl B, Montagnini F, López MA, Lavignolle P, Cortes J, De La Vega M, López LN, Dummel C,
Küppers G (2022) Conservation, connectivity and registration of seed areas in remnants of
natural reserves in the province of Misiones, Argentina. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in biodi-
versity and conservation. Springer, Cham, pp 157–181
Esbach MS, Puri M, Botero-Arias R, Loiselle BA (2022) Beyond the island: safeguarding biodi-
versity requires integrated approaches. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in biodiversity and conserva-
tion. Springer, Cham, pp 551–568
Esquivel MJ, Calle Z (2002) Árboles aislados en potreros como catalizadores de la sucesión en la
Cordillera Occidental Colombiana. Agrofor Amér 9(33–34):43–47
Esteva G (1999) The Zapatistas and people’s power. Cap Class 23(2):153–182
Ethiopian Biodiversity Institute (2014) Government of the Federal Democratic Republic of Ethio-
pia: Ethiopia’s fifth national report to the convention on biological diversity, Ethiopian Biodi-
versity Institute Addis Ababa, Ethiopia. 86 pp
Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Ann Rev Ecol Evol Syst 34:487–
515. [Link]
Fahrig L (2013) Rethinking patch size and isolation effects: the habitat amount hypothesis. J
Biogeogr 40(9):1649–1663
Fahrig L (2017) Ecological responses to habitat fragmentation per se. Ann Rev Ecol Evol Syst 48:
1–23
Francesconi W, Montagnini F, Ibrahim M (2011) Living fences as linear extensions of forest
remnants: a strategy for restoration of connectivity in agricultural landscapes. In:
Montagnini F, Finney C (eds) Restoring degraded landscapes with native species in Latin
America. Nova Science Publishers, New York, pp 115–126
32 F. Montagnini et al.

Gadgil M, Berkes F, Folke C (1993) Indigenous knowledge for biodiversity conservation. Ambio
22:151–156
Galluzzi G, Eyzaguirre P, Negri V (2010) Home gardens: neglected hotspots of agro-biodiversity
and cultural diversity. Biodivers Conserv 19(13):3635–3654
Giraldo LP, Chará J, Calle Z, Blanco JF, Chará-Serna AM (2022) Riparian corridors: longitudinal
biodiversity islands in farming landscapes. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human-dominated environments. Series: Topics in biodiversity and conser-
vation. Springer, Cham, pp 139–156
Gómez-Pompa A, Flores JS, Fernández MA (1990) The sacred cacao groves of the Maya. Lat Amer
Antiq 1(3):247–257
González LH, Neffeng G, Benedetto V, Sánchez Miñarro M, García A, Biasatti R, Rimoldi P,
Alesio C, Paiz D (2022) The anthropization of the natural landscape in the Pampa region in
Santa Fe province, Argentina. Resilience of the environment, opportunity for a change of the
Agri-food paradigm. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Series: Topics in biodiversity and conservation. Springer,
Cham, pp 353–386
Grass I, Loos J, Baensch S, Batáry P, Librán-Embid F, Ficiciyan A, Klaus F, Riechers M, Rosa J,
Tiede J et al (2019) Land-sharing/sparing connectivity landscapes for ecosystem services and
biodiversity conservation. Peo and Nat 1:262–272
Haila Y (2002) A conceptual genealogy of fragmentation research: from island biogeography to
landscape ecology. Eco App 12(2):321–334
Harvey CA, González J (2007) Agroforestry systems conserve species rich but modified assem-
blages of tropical birds and bats. Biodiv Conserv 16:2257–2292
Harvey CA, Villanueva C, Villacís J, Chacón M, Muñoz D, López M, Ibrahim M, Taylor R,
Martínez JL, Navas A, Sáenz J, Sánchez D, Medina A, Vílchez S, Hernández B, Pérez A,
Ruiz F, López F, Lang I, Kunth S, Sinclair FL (2005) Contribution of live fences to the
ecological integrity of agricultural landscapes in Central America. Agric Ecosyst Environ
111:200–230
Harvey CA, Medina A, Sánchez D, Vílchez S, Hernández B, Saenz J, Maes J, Casanoves F, Sinclair
FL (2006) Patterns of animal diversity associated with different forms of tree cover retained in
agricultural landscapes. Ecol Appl 16:1986–1999
Harvey CA, Komar O, Chazdon R, Ferguson BJ, Finegan B, Griffith DM, Martínez-Ramos M,
Morales H, Nigh R, SotoPinto L, Van Breugel M, Wishnie M (2008) Integrating agricultural
landscapes with biodiversity conservation in the Mesoamerican hotspot. Conserv Bio 22(1):
8–15
Hernández-Ruedas MA, Arroyo-Rodríguez V, Meave JA, Martínez-Ramos M, Ibarra-
Manríquez G, Martínez E, Jamangapé G, Melo FPL, Santos BA (2014) Conserving tropical
tree diversity and forest structure: the value of small rainforest patches in moderately-managed
landscapes. PLoS One 9:e98931
Holl KD (2002) Tropical moist forest restoration. In: Perrow MR, Davy AJ (eds) Handbook of
ecological restoration, vol 2. Cambridge University Press, Cambridge, pp 539–558
Holl KD, Leighton Reid J, Chaves-Fallas JM, Oviedo-Brenes F, Zahawi RA (2017) Local tropical
forest restoration strategies affect tree recruitment more strongly than does landscape forest
cover. J Appl Ecol 54:1091–1099
Jarvis DI, Padoch C, Cooper HD (2011) Manejo de la agrobiodiversidad en los ecosistemas
agrícolas. Bioversity International, Rome
Kirby K (2022) Islands of trees in Long-fragmented landscapes in Great Britain. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in biodiversity and conservation. Springer, Cham, pp 337–352
Kumar P (ed) (2010) The economics of ecosystems and biodiversity ecological and economic
foundations. Earthscan, London/Washington. 422 pp
Laino R, Musalem K, Laino L, Caballero A, Bueno D, Aranda L, Esquivel A, Ferreira Riveros M,
Romero Nardelli L, Cantero C, Irala R (2022) Island forests among savannahs: conservation of
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 33

biodiversity in rangelands of the Paraguayan humid Chaco. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in Bio-
diversity and Conservation. Springer, Cham, pp 185–205
Landis DA, Wratten SD, Gurr GM (2000) Habitat management to conserve natural enemies of
arthropod pests in agriculture. Ann Rev Entomo 45(1):175–201
Laurance WF (2008) Theory meets reality: how habitat fragmentation research has transcended
island biogeographic theory. Biol Conserv 141(7):1731–1744
Laurance WF, Camargo JL, Fearnside PM, Lovejoy TE, Williamson GB, Mesquita RC, Meyer CF,
Bobrowiec PE, Laurance SG (2018) An Amazonian rainforest and its fragments as a laboratory
of global change. Biol Rev 93(1):223–247
Levin B (2022a) Regenerative agricultural systems as biodiversity islands. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in biodiversity and conservation. Springer, Cham, pp 61–88
Levin B (2022b) The role of community led action for the creation of biodiversity islands. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human-dominated envi-
ronments. Series: Topics in biodiversity and conservation. Springer, Cham, pp 487–504
Liu J-J, Slik JWF (2014) Forest fragment spatial distribution matters for tropical tree conservation.
Biolog Cons 171:99–106
Lovejoy TE, Bierregaard RO (1990) Central Amazonian forests and the minimum critical size of
ecosystems project. In: Gentry AH (ed) Four neotropical rainforests. Yale University Press, New
Haven, pp 60–71
Lovejoy TE, Hannah L (eds) (2019) Biodiversity and climate change transforming the biosphere.
Yale University Press, New Haven. 416 pp
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press,
Princeton, p 203
McIntyre NE, Rango J, Fagan WF, Faeth SH (2001) Ground arthropod community structure in a
heterogeneous urban environment. Landsc Urban Plan 52:257–274
Melo FP, Arroyo-Rodríguez V, Fahrig L, Martínez-Ramos M, Tabarelli M (2013) On the hope for
biodiversity-friendly tropical landscapes. Trends Ecol Evol 28(8):462–468
Mittermeier RA, Turner WR, Larsen FW, Brooks TM, Gascon C (2011) Global biodiversity
conservation: the critical role of hotspots. In: Zachos FE, Habel JC (eds) Biodiversity hotspots.
Springer, Verlag Berlin Heidelberg, pp 3–22
Moeliono M, Mulyana A, Adnan H, Manalu P, Yuliani L (2015) Village forests (hutan desa):
empowerment, business or burden? World Agroforestry Centre-ICRAF, Southeast Asia
Regional Office, Bogor
Monsarrat S, Jarvi S, Svenning J-C (2019) Anthropocene refugia: integrating history and predictive
modelling to assess the space available for biodiversity in a human-dominated world. Phil Trans
of the R Soc B 374:20190219
Montagnini F (2020) The contribution of agroforestry to restoration and conservation: biodiversity
islands in degraded landscapes. In: Gupta SR, Dagar JC, Teketay D (eds) Agroforestry for
degraded landscapes: recent advances and emerging challenges, vol 1. Springer Nature, Singa-
pore, pp 445–479. [Link]
Montagnini F, Berg KE (2019) Feeding the world. In: Carver F, Gossington H, Manuel C (eds)
Sustainable development goals. Transforming our world. Witan Media Ltd/United Nations
Association, Painswick/London, pp 98–100. [Link]
world/
Montagnini F, del Fierro S (2022) Functions of agroforestry systems as biodiversity islands in
productive landscapes. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Series: Topics in biodiversity and conservation. Springer,
Cham, pp 89–116
Montagnini F, Finney C (2011) Payments for environmental services in Latin America as a tool for
restoration and rural development. Ambio 40:285–297
34 F. Montagnini et al.

Montagnini F, Metzel R (2017) The contribution of agroforestry to sustainable development goal 2:

end hunger, achieve food security and improved nutrition, and promote sustainable
agriculture. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity conser-
vation and food sovereignty, Advances in Agroforestry 12. Springer, Cham, pp 11–45
Montagnini F, Eibl B, Barth SR (2011) Organic yerba mate: an environmentally, socially and
financially suitable agroforestry system. Bois For Tropiq 308:59–74. https://
drfl[Link]/organic-yerba-mate-an-environmentally-socially-and-
financially-suitable-agroforestry-system/
Montagnini F, Smith RTW, Everett-Lane B, del Fierro S, Albrecht D (2022) Conclusions: lessons
learned and pending challenges. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in biodiversity and conserva-
tion. Springer, Cham, pp 677–703
Montes-Londoño I, Calle A, Montes-Calderon A, Montes-Botero O (2022) Hacienda Pinzacuá: an
island of regenerative agriculture in the piedmont of the Colombian Central Andes. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human-dominated envi-
ronments. Series: Topics in biodiversity and conservation. Springer, Cham, pp 305–335
Morales-Nieves GM (2022) Priorities, perspectives and use of a community forest by surrounding
residents in Mayagüez, Puerto Rico: protecting the forest for its services. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in biodiversity and conservation. Springer, Cham, pp 505–530
Murgueitio RE, Calle Z, Uribe F, Calle A, Solorio B (2011) Native trees and shrubs for the
productive rehabilitation of cattle ranching lands. For Ecol Manag 261:1654–1663
Nazarea VD, Rhoades RE, Andrews-Swann JE (eds) (2014) Seeds of resistance, seeds of hope:
place and agency in the conservation of biodiversity. University of Arizona Press, Tucson,
AZ. 288 pp
Negret HRC, Negret R, Montes-Londoño I (2022) Residential garden Design for Urban Biodiver-
sity Conservation: Experience from Panama City, Panama. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in biodi-
versity and conservation. Springer, Cham, pp 387–417
Newcomer Q, Camacho Céspedes F, Stallcup L (2022) The Monteverde Cloud Forest: evolution of
a biodiversity island in Costa Rica. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in biodiversity and conserva-
tion. Springer, Cham, pp 237–278
Niella F, Rocha P, Tuzinkievicz A, Buchweis R, Bulman C, Thalmayr P, González J, Montagnini F,
Sharry S (2022) Native multipurpose species in the Yaboti Biosphere Reserve, Misiones-
Argentina. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in biodiversity and conservation. Springer, Cham,
pp 461–483
Nordic Business Insider (2018) Norwegian Air axes two Stockholm-US direct routes. https://
[Link]. Retrieved 1 Dec 2018
OECD (2018) Tracking economic instruments and finance for biodiversity. OECD Publishing,
Paris. [Link]
[Link]
OECD (2019) Biodiversity: finance and the economic and business case for action. Report prepared
for the G7 Environment Ministers’ meeting, 5–6 May 2019. Available at: [Link]
environment/resources/biodiversity/G7-report-Biodiversity-Finance-and-the-Economic-and-
[Link]
Otto J, Zerner C, Robinson J, Donovan R, Lavelle M, Villarreal R, Salafsky N, Alcorn J,
Seymour F, Kleyneyer C, Pearl M (2013) Natural connections: perspectives in community-
based conservation. Island Press, Washington DC
Pagiola S, Honey-Rosés J, Freire-González J (2017) Assessing the permanence of land use change
induced by payments for environmental services: evidence from Nicaragua. World Bank
eLibrary. [Link]
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 35

Painter K, Buschbacher R, Souto Silva LC, Costa e Silva E (2022) Agroecology and forest
conservation in three types of land reform community in the Cacao Region of Bahia,
Brazil. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in biodiversity and conservation. Springer, Cham,
pp 569–599
Parker C, Cranford M, Oakes N, Leggett M (2012) The little biodiversity finance book, 3rd edn. The
Global Canopy Programme, Oxford. 101 pp
Peters CM (2018) Managing the wild. Stories of people and plants and tropical forests. Yale
University Press, New Haven. 208 pp
Pfeifer M, Lefebvre V, Peres CA, Banks-Leite C, Wearn OR, Marsh CJ, Butchart SHM, Arroyo-
Rodríguez V, Barlow J, Cerezo A et al (2017) Creation of forest edges has a global impact on
forest vertebrates. Nature 551:187–191
Phalan BT (2018) What have we learned from the land sparing-sharing model? Sustainability
10(1760):doi:10.3390/su10061760
Phalan B, Onial M, Balmford A, Green RE (2011) Reconciling food production and biodiversity
conservation: land sharing and land sparing compared. Science 333(6047):1289–1291
Pickett ST, Thompson JN (1978) Patch dynamics and the design of nature reserves. Biol Conserv
13(1):27–37
Pickett STA, Cadenasso ML, Grove JM (2004) Resilient cities: meaning, models, and metaphor for
integrating the ecological, socio-economic, and planning realms. Landsc Urban Plan 69:369–
384
Pimentel D, Wilson C, McCullum C, Huang R, Dwen P, Flack J, Tran Q, Saltman T, Cliff B (1997)
Economic and environmental benefits of biodiversity. BioSci 47(11):747–757
Prieto Méndez JM (2013) Derechos de la naturaleza. Fundamento, contenido y exigibilidad
jurisdiccional. Corte Constitucional del Ecuador, CEDEC, Quito. 280 pp
Rainforest Foundation Norway (2016) Norwegian state commits to zero deforestation. [Link]
[Link]/en/news/norwegian-state-commits-to-zero-deforestation-1. Retrieved 7 Feb 2018
Redford KH, Padoch C (eds) (1992) Conservation of Neotropical forests. Working from traditional
resource use. Columbia University Press, New York, p 475
Sáenz JC, Villatoro F, Ibrahim M, Fajardo D, Pérez M (2007) Relación entre las comunidades de
aves y la vegetación en agropaisajes dominados por la ganadería en Costa Rica, Nicaragua y
Colombia. Agrofor Amér 45:37–48
Salzman J, Bennett G, Carroll N, Goldstein A, Jenkins M (2018) The global status and trends of
payments for ecosystem services. Nat Sustain 1:136–144
Santos-Gally R, Boege K (2022) Biodiversity islands. Native tree islands within silvopastoral
systems in a neotropical region. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in biodiversity and conserva-
tion. Springer, Cham, pp 117–138
Santos-Gally R, Boege K, Fornoni J, Domínguez CA (2019) Cómo transitar hacia una ganadería
tropical sostenible. Oikos. [Link]
Savard JPL, Clergeau P, Mennechez G (2000) Biodiversity concepts and urban ecosystems. Land
Urb Plan 48:131–142
Sheban K (2022) Preserving biodiversity in Appalachian mixed mesophytic forest through permit-
based harvest of ginseng and other forest botanicals. In: Montagnini F (ed) Biodiversity islands:
strategies for conservation in human-dominated environments. Series: Topics in biodiversity
and conservation. Springer, Cham, pp 601–628
Sigman E (2022) Safeguarding the benefits of biodiversity islands in northern Ethiopia in the midst
of political change. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Series: Topics in biodiversity and conservation. Springer,
Cham, pp 647–674
Simberloff D, Abele LG (1982) Refuge design and island biogeographic theory: effects of frag-
mentation. Amer Nat 120(1):41–50
36 F. Montagnini et al.

Soler R, Benítez J, Solá F, Lencinas MV (2022) Biodiversity islands at the world’s

southernmost city: plant, bird and insect conservation in urban forests and peatlands of Ushuaia,
Argentina. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in biodiversity and conservation. Springer, Cham,
pp 419–437
State Information Service (2018) COP14 decisions, political declaration unanimously adopted-
Minister. [Link]
mously-adopted%2D%2Dminister?lang¼en-us. Retrieved 7 Feb 2019
Talberth J, Gray E (2012) Global costs of achieving the Aichi biodiversity targets a scoping
assessment of anticipated costs of achieving targets 5, 8, and 14. Center for Sustainable
Economy, Washington, DC [Link]
[Link]
Thaxton M, Forster T, Hazlewood P, Mercado L, Neely C, Scherr SJ, Wertz L, Wood S, Zandri E
(2016) Landscape partnerships for sustainable development: achieving the SDGs through
integrated landscape management. White paper produced by The landscapes for people, food,
and nature initiative task force on the sustainable development goals. www.
[Link]. 28 pp
Tjørve E (2010) How to resolve the SLOSS debate: lessons from species-diversity models. J Theor
Biol 264(2):604–612
Toensmeier E (2022) Paradise lot: a temperate-climate urban agroforestry biodiversity Island. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human dominated-
environments. Series: Topics in biodiversity and conservation. Springer, Cham, pp 439–459
Toensmeier E, Bates J (2013) Paradise lot; two plant geeks, one tenth of an acre, and the making of
an edible garden oasis in the city. Chelsea Green Publishing, White River Junction, VT
Totoket Times (2018) Press release from the North Branford conservation trust. celebrating a new
chapter and leaving a legacy. Totoket. Times 25(17):14
Tscharntke T, Klein AM, Kruess M, Steffan-Dewenter I, Thies C (2005) Landscape perspectives on
agricultural intensification and biodiversity—ecosystem service management. Ecol Lett 8:857–
874
Tuff KT, Tuff T, Davies KF (2016) A framework for integrating thermal biology into fragmentation
research. Ecol Lett 19(4):361–374
Udawatta RP, Rankoth LM, Jose S (2019) Agroforestry and biodiversity. Sustainability 11:2879.
[Link]
UNDP (2006) Community action to conserve biodiversity. linking biodiversity conservation with
poverty reduction. Case studies from Latin America and the Caribbean. UNDP, New York.
[Link]
and_biodiversity/[Link]. Retrieved 7 Feb 2019
United Nations IPBES (2019) Global assessment report on biodiversity and ecosystem services.
[Link]
Vásquez V, Barber C, Dguidegue Y, Caughlin TT, García R, Metzel R (2022) Farmer perceptions
of tropical dry forest restoration practices on the Azuero Peninsula of Panama – implications for
increasing biodiversity in a human-dominated landscape. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in biodi-
versity and conservation. Springer, Cham, pp 629–646
Volenec ZM, Dobson AP (2019) Conservation value of small reserves. Conserv Bio. [Link]
org/10.1111/cobi.13308
Watling J, Arroyo-Rodríguez V, Pfeifer M, Baeten L, Banks-Leite C, Cisneros L, Fang R, Hamel-
Leigue C, Lachat T, Leal I, Lens L, Possingham H, Raheem D, Ribeiro D, Slade E, Urbina-
Cardona N, Wood E, Fahrig L (2020) Support for the habitat amount hypothesis from a global
synthesis of species density studies. Ecol Lett 23(4):674–681. [Link]
13471. (Advances in Agroforestry 12. Cham: Springer) pp 179–209
1 Introduction. Biodiversity Islands: Strategies for Conservation. . . 37

Watts J (2018) Stop biodiversity loss or we could face our own extinction, warns UN. [Link]
[Link]/environment/2018/nov/03/stop-biodiversity-loss-or-we-could-face-our-own-
extinction-warns-un. Retrieved 27 Nov 2018
Wintle BA, Kujala H, Whitehead A, Cameron A, Veloz A, Kukkala A, Moilanen A, Gordon A,
Lentini PE, Cadenhead NCR, Bekessy SA (2019) Global synthesis of conservation studies
reveals the importance of small habitat patches for biodiversity. PNAS 116:909–914
World Agroforestry Centre (ICRAF), Bogor (Indonesia) (2010) Lubuk Beringin village forest. The
first in Indonesia. SEA Regional Office. Contributor(s): Rewards for Use of and Shared
Investment in Pro-Poor Environmental Services (RUPES). [Link]
publication/lubuk-beringin-village-forest-first-indonesia
Yarranton GA, Morrison RG (1974) Spatial dynamics of a primary succession: nucleation. J Ecol
62:417–428
 Part II
Biodiversity Islands Establishment
and Management: Challenges
and Alternatives
Chapter 2
The Importance of Small Rainforest Patches
for Biodiversity Conservation:
A Multi-taxonomic Assessment

Víctor Arroyo-Rodríguez, Ricard Arasa-Gisbert, Norma P. Arce-Peña,

Martín J. Cervantes-López, Sabine J. Cudney-Valenzuela,
Carmen Galán-Acedo, Manuel A. Hernández-Ruedas, Miriam San-José,
and Lenore Fahrig

Abstract Tropical forests are being rapidly deforested worldwide. The remaining
forest is distributed in different-sized forest patches, but the species preservation
value of small patches remains debated. Some studies suggest that edge effect can
decrease forest-specialist species diversity, particularly in small patches, which are
expected to be mainly occupied by a few disturbance-adapted species. We tested this
hypothesis by sampling plants, dung beetles, amphibians, reptiles, birds, and mam-
mals in the fragmented Lacandona rainforest in Mexico. We separately evaluated
forest-specialist and habitat generalist species. As positive patch size effects on
species richness can be simply related to the sample area effect (i.e. larger samples
have a higher chance of holding more species), we assessed the effect of patch size
on the number of species of each group in samples of constant size (species density).
We also evaluated whether species density is lower in forest patches than in
continuous forest sites. We found that patch area was generally a poor predictor of

V. Arroyo-Rodríguez (*)
Laboratorio de Ecología de Paisajes Fragmentados, Instituto de Investigaciones en Ecosistemas
y Sustentabilidad, Universidad Nacional Autónoma de México, Morelia, Michoacán, Mexico
Escuela Nacional de Estudios Superiores, Universidad Nacional Autónoma de México, Mérida,
Yucatán, Mexico
R. Arasa-Gisbert · N. P. Arce-Peña · M. J. Cervantes-López · S. J. Cudney-Valenzuela ·
M. A. Hernández-Ruedas · M. San-José
Laboratorio de Ecología de Paisajes Fragmentados, Instituto de Investigaciones en Ecosistemas
y Sustentabilidad, Universidad Nacional Autónoma de México, Morelia, Michoacán, Mexico
C. Galán-Acedo
Escuela Nacional de Estudios Superiores, Universidad Nacional Autónoma de México, Morelia,
Michoacán, Mexico
L. Fahrig
Geomatics and Landscape Ecology Laboratory, Department of Biology, Carleton University,
Ottawa, Canada

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 41

F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
42 V. Arroyo-Rodríguez et al.

species density, and that in most study groups the density of species did not differ
between continuous forest and forest patches. Remarkably, most results were inde-
pendent of habitat specialization. These findings add to the increasing evidence that,
on a per-sample area basis, small patches are valuable biodiversity islands for
conservation of forest-specialist species and are not the near-exclusive habitat of
generalist species. Our results indicate a need to redress the neglect of small patches
in conservation plans, even for forest-specialist species in fragmented rainforests, in
order to help maintain species diversity.

Keywords Biodiversity-friendly landscapes · Habitat fragmentation · Habitat loss ·

Landscape configuration · Landscape restoration

2.1 Introduction

Deforestation is causing the net annual loss of more than 12 million hectares of forests
globally (Hansen et al. 2013). The remaining forest is distributed in millions of
different-sized forest patches, most of them very small, with the global meanpatch
size ranging between 13 and 17 ha (Taubert et al. 2018). This situation threatens
biodiversity maintenance since forest loss is a major cause of species loss (Newbold
et al. 2016; Watling et al. 2020). Because forest area can have an effect on species loss
at a local scale, some studies suggest that small forest patches can be of relatively low
conservation value because: (1) they cannot maintain viable populations of forest-
specialist species (i.e. those that use forest interior as their primary habitat);
(2) populations in small patches are more susceptible to human-caused disturbances
(e.g. hunting, logging, livestock incursion, wildfire); and (3) most forest-specialist
species are negatively impacted by edge effects (i.e. biotic and abiotic changes at
forest edges) and should therefore be more prone to extinction in small patches, which
have a high edge-to-area ratio (Diamond 1975, 1976; Willis 1984; Laurance et al.
2002; Banks-Leite et al. 2010; Fletcher et al. 2018; Phalan 2018). Thus, smaller
patches are expected to have a lower number of species than large ones.
The lower number of species in small patches can, however, be caused by the
sample area effect, i.e. small samples (patches) can contain a lower number of
individuals, so they have a higher chance of containing a lower species richness as
well (Fahrig 2013; Chase et al. 2018). Thus, to better understand the impact of patch
size on species diversity, the number of species needs to be measured in same-sized
samples (i.e. species density; Fahrig 2013). Small patches may be edge-affected
habitats of ideal conditions for a few generalists and/or invasive species (i.e. those
that use resources from forest interior, forest edges, regenerating forest stands, and
agricultural lands), but of less favorable conditions for most forest-specialist species
(Laurance et al. 2002; Banks-Leite et al. 2010; Tabarelli et al. 2012; Fletcher et al.
2018). Therefore, the species density should be positively related to patch size.
Similarly, compared to samples in continuous forest sites, species density should be
lower in forest patches, especially when considering patches of <100 ha, as edge
effects can penetrate 500 m or more into forest patches (Laurance et al. 2002; Harper
et al. 2005; Pfeifer et al. 2017).
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 43

In contrast to this hypothesis, evidence indicates that on a per-sample area basis,

small patches can have as high or higher value for conservation of forest species as
large patches (Arroyo-Rodríguez et al. 2020; Fahrig 2020; Watling et al. 2020).
Among other factors (reviewed by Fahrig 2020), this may be related to the fact that
most species are able to use resources from multiple patches in the landscape rather
than being constrained by the resources available in a single patch. This is plausible
if forest edges and the surrounding anthropogenic matrix do not represent a barrier to
the movement of individuals in fragmented landscapes, as demonstrated in several
studies (Mendenhall et al. 2013; Ferreira et al. 2018; Galán-Acedo et al. 2019).
Therefore, the number of species in a given site (species density) may depend more
on the forest cover available in the local landscape surrounding the site than on the
size of the patch in which the site is located (Fahrig 2013). A recent global meta-
analysis supports this hypothesis (Watling et al. 2020). However, considerable
skepticism remains about the value of small patches in biodiversity conservation,
especially in tropical regions, so this controversial topic requires further evaluation
(Fletcher et al. 2018).
Here, we report results of studies in which we compiled data on the seed rain,
saplings, trees, dung beetles, amphibians, reptiles, birds, bats, medium- and large-
bodied terrestrial mammals, and arboreal mammals recorded in the interior of
old-growth forest sites (forest sites, hereafter). The area of study included
different-sized forest patches and continuous forest sites in the Lacandona rainforest,
Mexico (Fig. 2.1). These data were collected as part of several of our previous
studies on the same subject (Table 2.1). In particular, we assessed, for the first time:
(1) the effect of patch size on species density (i.e. species richness in samples of fixed
size); and (2) the difference in species density between forest patches and continuous
forest sites. These analyses were done by separately assessing forest-specialist and
forest-generalist species. Such an evaluation is important in the context of biodiver-
sity island design, as our findings will indicate if forest patches of any size can be
beneficial to the protection of forest biodiversity.

2.2 Methods

2.2.1 Study Area

The Mexican portion of the Lacandona rainforest (16
100 5600 N - 90
530 2100 W) is
located in southeastern Chiapas State, Mexico (Fig. 2.1). The climate is warm with a
mean annual temperature of 24
C and humid with a mean annual rainfall of
2143 mm. This region preserves one of the largest rainforest tracts in Mexico and
represents a priority area for biodiversity conservation in Mesoamerica (Arriaga
et al. 2000). Within this region, the Montes Azules Biosphere Reserve preserves
331,200 ha of contiguous, largely undisturbed forest. Nevertheless, deforestation
outside this reserve has resulted in the loss of more than 45% of original forest cover,
due to its conversion to cattle pastures, annual crops, and oil-palm plantations
44 V. Arroyo-Rodríguez et al.

Fig. 2.1 Location of the Lacandona rainforest in southeastern Mexico. We sampled continuous
forest sites within the Montes Azules Biosphere Reserve, and forest patches in the Marqués de
Comillas municipality (black stars). Forest cover is indicated with dark green, and the anthropo-
genic matrix with brown. The blue line is the Lacantún River. Source: Digitized by C. Galán-Acedo
using Landsat images and the CLASlite software

(Carabias et al. 2015; Fig. 2.2a and b). Such forest loss has been particularly notable
in the Marqués de Comillas region, which is separated from the Biosphere Reserve
by the Lacantún River (Figs. 2.1 and 2.2a).

2.2.2 Study Forest Sites

As part of previous independent studies of different taxonomic groups, we have data

from 12 to 26 forest sites depending on the taxa: 0 to 4 sites within the Montes
Azules Biosphere Reserve, and 5 to 24 forest patches within the Marqués de
Comillas municipality (Table 2.1; Fig. 2.1). Continuous forest sites refer to circular
areas of 100 ha each within the Montes Azules Biosphere Reserve (continuous forest
sites hereafter), which were spaced 4 km apart on average and at least 1 km from the
nearest edge of the Lacantún river (Fig. 2.1). Forest patches within the Marqués de
Comillas municipality were spaced at least 1500 m apart, and forest patch size ranges
differed among studies (Table 2.1). All sites were located in lowland areas
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 45

Table 2.1 Summary of study designs. Rows refer to independent studies of different biological
groups in forest patches (FP) and continuous forest sites (CF) in the Lacandona rainforest, Mexico
Biological Response Sampling # forest Patch Sampling method in
group variablea period sites size (ha) each site Refsb
Seeds Species Feb. 19 2.8–129 9 seed traps (0.5 m2 1
density 2015 – FP + 1 each) in a grid of
Feb. 2016 CF 88m
Saplings Species Jan. – 19 2.8–157 25 plots (8 m2 each) in a 2
density May 2018 FP + 1 grid of 120  120 m
CF
Trees Species Apr. – 19 2.8–129 10 parallel 50  2-m 3
density Jun. 2015 FP + 1 plots (plot
CF distance ¼ 10 m)
Dung Species Jul. – Sep. 21 2.8–129 4 baited pitfall traps per 4
beetles density* 2012 FP + 3 transect. Number of
CF transects proportional to
patch area
Amphibians Species May – 9 2.8–91.9 Visual encounters in 5
& reptiles density Sep. 2012 FP + 3 6 non-fixed transects
CF
Amphibians Species May – 5 FP 1.7–69.8 Visual encounters in 6
& reptiles density Nov. 6 non-fixed transects
2018
Birds Species May – 17 2.8–129 Point counts (number 7
density* Aug. FP + 3 proportional to patch
2012 CF size)
Bats Species May – 12 2.8–91.9 5 mist nests located 50 m 8
density Sep. 2012 FP + 3 between each other
CF
Terrestrial Species Apr. – 24 2.8–129 Camera traps (150 cam- 9
mammalsc density Aug. FP + 4 era trap nights per site)
2011 CF
Species Apr. – 24 2.8–129 Camera traps (150 cam- 10
density Aug. FP + 4 era trap nights per site)
2017 CF
Arboreal Species May 19 2.8–157 Camera traps on 5 focal 11
mammals density 2018 – FP + 1 trees
May 2019 CF
a
In all cases but two (indicated with *), we calculated the number of species (i.e. count response
variable) recorded in samples of fixed size (species density). The responses marked with asterisk
were recorded with a sampling effort that increased with patch size. In those cases, species density
was calculated as the mean number of species per transect (dung beetles) or point count (birds), so
they are continuous response variables
b
References: 1. San-José et al. (2020); 2. R. Arasa-Gisbert (unpubl. data); 3. M.A. Hernández-
Ruedas (unpubl. data); 4. Sánchez-de-Jesús et al. (2015); 5. Russildi et al. (2016); 6. Cervantes-
López (unpubl. data); 7. Carrara et al. (2015); 8. Arroyo-Rodríguez et al. (2016); 9. Garmendia et al.
(2013); 10. N. Arce-Peña (unpubl. data); 11. S. Cudney-Valenzuela (unpubl. data)
c
Terrestrial mammals were sampled in the same sites but in different time periods. We calculated
the accumulated number of species sampled at each site across time periods
46 V. Arroyo-Rodríguez et al.

Fig. 2.2 Pictures from the study landscape showing the Lacantun river (panel a), which separates
the continuous forest of the Montes Azules Biosphere Reserve (left side) and the Marqués de
Comillas region (right side). This latter region is highly heterogeneous, as the remaining forest
patches are surrounded by a matrix of cattle pastures (panels a and b) and crops (e.g. oil-palm
plantations in panel c), with different treed elements (e.g. living fences and isolated standing trees in
panels a and b). (Photos: Víctor Arroyo-Rodríguez)
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 47

(100–200 meters above sea level (m.a.s.l.)) with similar soil and weather conditions
to avoid the potential confounding effects of these variables. Also, because most of
the region was rapidly deforested during the 1980s, we assume that most patches
have a similar age (30–40 years).

2.2.3 Study Taxa

We surveyed seeds, saplings, trees, dung beetles, amphibians, reptiles, birds, bats,
medium- to large-bodied terrestrial mammals, and arboreal mammals (Table 2.1).
All surveys were done within forest interior (>50 m from the nearest forest edge).
The sampling methods and efforts are detailed in their original studies and summa-
rized and referenced in Table 2.1, but a brief overview is given here. Note that all
taxa but two (i.e. birds and dung beetles) were sampled in sample areas of
constant size.

[Link] Seeds, Saplings and Trees

We sampled seeds, saplings and trees in the center of each forest site (n ¼ 20 sites in
all cases; Table 2.1). To sample seeds, we placed 9 seed traps (trap area ¼ 0.5 m2,
1-mm nylon mesh) in a grid of 3  3 traps separated by 4 m (total sampling
effort ¼ 20  9 ¼ 180 traps). Traps were hung approximately 90 cm above the
ground, and trap contents were recovered every 15 days for one year (see San-José
et al. 2020). Saplings were recorded in 25 circular plots (1.60-m radius, 8-m2 each;
total sampling area per site ¼ 200 m2) in a grid of 5  5 plots (inter-plot
distance ¼ 30 m). We counted and identified all tree saplings (excluding palms
and lianas) with 30 cm height and < 1 cm of diameter at breast height (DBH). Trees
were sampled using a modification of Gentry’s (1982) protocol; at each site, we
recorded only tree species (including palms) with DBH  1 cm in ten parallel
50  2-m transects (i.e. 0.1 ha per patch), separated by 10 m.

[Link] Dung Beetles

We collected dung beetles in 24 forest sites (Table 2.1) using baited pitfall traps
(i.e. 1 L plastic containers buried with the top edge leveled with the soil surface). We
baited traps with 25 g of a mixture of human and pig excrement (7:3). We placed
four traps separated by 50 m along transects. Sampling effort increased with patch
area (i.e. one transect every 20 ha, and five transects in continuous forest sites).
When sampling 2 transects in a site, we separated them by 150 m. In total, we
used 196 pitfall traps, and we collected, identified, and counted all individuals and
species (Sánchez-de-Jesús et al. 2016).
48 V. Arroyo-Rodríguez et al.

[Link] Amphibians and Reptiles

We sampled amphibians and reptiles as part of two studies in 12 and 5 forest sites
(Table 2.1) but using the same protocol. In particular, we used visual encounter
surveys (Crump and Scott 1994). We surveyed all forest sites (continuous forest sites
and forest patches) using the same methods and a similar sampling effort in all
sample sites independently of their size. In particular, we divided each sample site in
six sections of similar size, and then sampled all amphibians and reptiles using
non-fixed transects (one transect per section). During each visit we sampled a
different section, totaling 6 days per forest site. Each visit included a day (10:00 to
13:00 h) and night (19:00 to 22:00 h) sampling period (3 h each). All individuals
were identified in situ using the field guides of Lee (2000) and Campbell (1998) and
released in the same place where they were captured (see Russildi et al. 2016).

[Link] Birds

We sampled birds in 20 forest sites (Table 2.1) using unlimited radius point counts
(Bibby et al. 2000). We sampled each site three times (once per month) following a
randomly selected order. We distributed point counts by dividing each site in three
sections of similar size, and during each visit, we sampled a different section from 5:
30 to 10:30 h. Point counts were separated 200 m from each other. The number of
point counts increased with the size of the patches and sections (i.e. 1 to 5 points per
section depending on its size). In total, we surveyed 130 point counts in patches and
72 in continuous forest sites. In each point count we recorded all birds seen or heard
during a 15 min period, considering only those perched on trees, on the floor, feeding
or using other resources of the study forest (see Carrara et al. 2015).

[Link] Bats

We sampled bats in 15 forest sites placing 5 mist nests (12  2.5 m) 50 m apart in
natural linear openings (e.g. animal trails, streams) at the center of each site
(Table 2.1). Mist nets were kept open for 5 h after sunset and checked every
30 min (Kunz 1982). Each site was sampled for six non-consecutive nights
(n ¼ 90 nights). We did not sample bats during the full moon because of their
reduced activity. We placed captured bats in cloth bags (30  40 cm) for later
identification using a field key (Medellín et al. 2008), and we marked them on the
right tibia to avoid repeated counts. We released them within 2 h of capture at the
capture site (see Arroyo-Rodríguez et al. 2016).
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 49

[Link] Medium- and Large-Bodied Terrestrial Mammals

Terrestrial mammals were sampled as part of two studies in the same 28 forest sites
and using exactly the same sampling protocol (Table 2.1). In each forest site, we
placed a camera trap in an apparently ‘high quality’ location, i.e. with evident signs
of use by mammals (e.g. area with several mammal footprints). The camera was left
for 30 days, with a recovery time of 30 seconds per picture. Cameras were serviced
once a month (e.g. change of batteries, download pictures). After 30 days, we moved
the camera to a different ‘high quality’ location within the same forest site, but 100 to
1000 m apart, as done in previous studies (Saito and Koike 2013). In total, we
registered 150 camera trap nights per site, with five sample locations in each site
(Garmendia et al. 2013). For the analyses described below, we combined the
information from the two studies by accumulating the species sampled in each site
after the two sampling periods.

[Link] Arboreal Mammals

We also used camera traps to record arboreal mammals in 20 forest sites (Table 2.1).
At the geographical center of each forest site, we selected five trees (four reached the
canopy and one the understory) with good climbing conditions (i.e. branches
20 cm wide, preferably hardwood species, with adequate architecture to install a
camera trap facing other main branches). We placed one camera trap on one of the
five trees per patch and we changed the location of the cameras, rotating them once a
month among the five trees, except from October to December when they remained
on the same tree (7387 total camera trap nights). We serviced cameras (e.g. change
of batteries, download pictures) every time we changed their location. We used bait
(tuna, peanut butter with oatmeal, and a banana) on one tree per site to increase the
probability of photo-capturing arboreal mammals.

2.2.4 Data Analyses

We separately assessed responses of forest-specialist and generalist species, which

were classified based on scientific literature (Garmendia et al. 2013; Hernández-
Ruedas et al. 2014; Carrara et al. 2015; Sánchez-de-Jesús et al. 2016; Arroyo-
Rodríguez et al. 2016; Russildi et al. 2016; IUCN 2019), and the assistance of
local experts. All analyses were done with R 3.0.1 (R Core Team 2013).
50 V. Arroyo-Rodríguez et al.

[Link] Effect of Patch Size on Response Variables

Our response variables were estimated on a per-sample area basis: i.e. species
density. This was already the case when sampling effort was the same in each
patch (i.e. seed rain, saplings, trees, amphibians, reptiles, bats, medium- and large-
bodied terrestrial mammals, and arboreal mammals). When sampling effort
increased with patch size (i.e. dung beetles and birds, Table 2.1), we calculated the
mean number of species per transect (dung beetles) or per point count (birds).
After a visual inspection of the scatter plots, we discovered that species density
was not related to patch size in most species groups (i.e. null patch size effect), and
when related, patch size seemed to be linearly related to species density (Table 2.2).
Therefore, to statistically test these relationships we used generalized linear models
(GLM) and compared those models to the null model (including only the intercept).
Such a comparison was done following an information-theoretic criteria. In partic-
ular, we estimated the Akaike Information Criterion corrected for small samples
(AICc) and selected the model with lower AICc. When the linear and null models
showed similar empirical support (i.e. ΔAICc <2), we selected the simplest (null)
model. We tested GLMs with a Poisson error distribution for count response vari-
ables (see column of response variables in Table 2.1), and we corrected for over-
dispersion by including a quasi-poisson error distribution when the ratio between
residual deviance to degrees of freedom was higher than 1. For responses variables
that were means (dung beetles and birds; Table 2.1), we used a Gaussian error
distribution (Crawley 2007).

[Link] Differences Between Continuous Forest and Forest Patches

Differences in all response variables per unit area (as in Sect. [Link]) between
continuous and fragmented forests were also tested with GLMs, but only for those
taxa for which we had enough replicates of continuous forest (n  3 sites). As
described in Sect. [Link], we fitted a Poisson error distribution to count response
variables (number of species) and a Gaussian error distribution to continuous
response variables (mean number of species). Here, we also corrected the models
for over-dispersion, when needed.

2.3 Results

2.3.1 Effect of Patch Size on the Number of Species

In total, we sampled 111 species of tree seeds, 161 tree saplings, 152 trees, 43 dung
beetles, 26 amphibians, 42 reptiles, 84 birds, 34 bats, 21 terrestrial mammals, and
15 arboreal mammals. Patch size was generally a poor predictor of species density,
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 51

Table 2.2 Effect of forest patch size on the species density of different taxa in the Lacandona
rainforest, Mexico
Taxon Habitat Model Coef AICc Δi wi
Seeds FS Null 92.54 0.0 0.75
GLM 0.0015 94.74 2.2 0.25
FG Null 87.17 0.0 0.74
GLM 0.0015 89.37 2.2 0.25
Saplings FS Null 111.61 0.0 0.77
GLM 0.0004 114.01 2.4 0.23
FG Null 143.32 0.0 0.60
GLM 0.0014 144.12 0.8 0.40
Trees FS Null 120.95 0.0 0.73
GLM 0.0010 122.95 2.0 0.27
FG Null 132.09 0.0 0.74
GLM 0.0007 134.19 2.1 0.26
Dung beetles FS GLM 20.062 100.95 0.0 0.92
Null 105.95 5 0.08
FG GLM 0.040 102.69 0.0 0.55
Null 103.09 0.4 0.45
Amphibians FS Null 58.5 0.0 0.64
GLM 0.006 59.6 1.1 0.36
FG GLM 0.008 61.1 0.0 0.53
Null 61.3 0.2 0.47
Reptiles FS Null 52.56 0.0 0.70
GLM 0.005 54.26 1.7 0.30
FG Null 64.16 0.0 0.53
GLM 0.006 64.36 0.2 0.47
Birds FS Null 11.27 0.0 0.74
GLM 0.003 13.37 2.1 0.26
FG GLM 20.028 57.80 0.0 0.85
Null 61.3 3.5 0.15
Bats FS Null 48.56 0.0 0.81
GLM 0.0002 51.46 2.9 0.19
FG Null 51.95 0.0 0.81
GLM 0.0011 54.85 2.9 0.19
Terrestrial mammals FG Null 110.48 0.0 0.67
GLM 0.0026 111.88 1.4 0.33
Arboreal mammals FG Null 87.55 0.0 0.77
GLM 0.0006 89.95 2.4 0.23
We separately assessed forest-specialist (FS) and forest-generalist (FG) species. We compared the
generalized linear model (GLM) with the null model (which included only the intercept) using
information-theoretic criteria (AICc: Akaike Information Criterion corrected for small samples; Δi:
difference in AICc between the best model and model i; wi: Akaike weights). The selected model in
each case is in boldface. Note that when the linear and null models showed similar empirical support
(i.e. ΔAICc <2), we selected the simplest (i.e. null) model
52 V. Arroyo-Rodríguez et al.

Fig. 2.3 The most

important effects of patch
size on species density of
forest-specialist dung
beetles (black dots) and
generalist birds (gray dots)
in the Lacandona rainforest,
Mexico. Only models with
empirical support are shown
(see Table 2.2). The red line
shows the predicted
estimates from the best
fitting models and the dotted
lines indicate 95%
confidence intervals

with the null model being the best fitting model in 16 of 18 assessments (89%;
Table 2.2). Only the mean number of specialist dung beetles and generalist birds
were linearly and negatively related to patch size (Fig. 2.3).

2.3.2 Continuous Forest vs Forest Patches

In 8 out of 11 cases (72%), the species density was similar in forest patches and
continuous forest sites (Table 2.3). The density of terrestrial mammals (all generalist
species) and the densities of forest-specialist amphibians and reptiles were 1.3 to 1.7
times higher in continuous forest than in forest patches (Table 2.3).

2.4 Discussion

This study demonstrates the high conservation value of small forest patches for
biodiversity conservation in the Lacandona rainforest – a species-rich but vanishing
tropical region in southeastern Mexico. We predicted that if small patches are of low
quality for forest-specialist species, the density of forest-specialist species should be
positively related to patch size and be higher in continuous forest than forest patches.
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 53

Table 2.3 Differences in the mean number of forest-specialist and generalist species between
continuous forest sites (CF) and forest patches (FP) in the Lacandona rainforest, Mexico, for
biological groups sampled in at least three continuous forest sites (see Table 2.1)
Study taxa CF FP t p
Forest-specialist species
Dung beetles 3.9  0.1 6.5  2.8 1.69 0.09
Amphibians 6.0  4.2 4.4  1.6 2.5 0.01
Reptiles 7.0  2.2 4.0  1.3 2.56 0.01
Birds 0.5  0.1 0.8  0.3 0.6 0.55
Bats 8.0  3.0 5.3  1.4 1.69 0.09
Generalist species
Dung beetles 2.7  0.2 5.8  2.6 2.02 0.06
Amphibians 3.3  1.5 5.5  2.0 1.49 0.14
Reptiles 5.0  1.7 7.0  2.2 1.21 0.23
Birds 1.6  0.2 3.6  1.3 1.69 0.09
Bats 9.7  3.2 8.2  1.3 0.8 0.43
Terrestrial mammals 15.0  1.6 11.6  2.8 2.26 0.03
Mean (SD) values are indicated in each forest type. Significant differences are highlighted with
boldface (t ¼ t value in GLM tests, p ¼ probability)

Conversely, if generalist species can use resources from forest interiors, forest edges,
and even from some matrix types, we expected no effect of patch size on generalists,
and a similar density of generalist species in both forest types.
Our findings support our predictions for generalist species, but not for specialist
ones. Patch area is generally a poor predictor of species density. In fact, where there
was a difference (specialist dung beetles and generalist birds), species density was
higher in small than large patches. Also, in all but three study groups (generalist
terrestrial mammals, and specialist amphibians and reptiles), the density of species
did not differ between continuous forest sites and forest patches. Importantly, most
findings were similar for forest-specialist and habitat generalist species. Therefore,
our findings add to an increasing number of studies (e.g. Fahrig 2020; Watling et al.
2020) indicating that, on a per-area basis, a unit of habitat in a small forest patch may
not only have a conservation value similar to that of large ones, but may be even
greater.
The fact that species density is rarely related to patch area and that it does not
usually differ between continuous forest and forest patches can be related to several
non-exclusive factors. First, in this region, forest patches are embedded in a highly
heterogeneous matrix with many treed elements (i.e. living fences, isolated trees, tree
crops; Fig. 2.2a–c), and this spatial context can reduce edge influence on forest
patches (Harper et al. 2005; Arroyo-Rodríguez et al. 2017). Second, these treed
elements in the matrix can also provide important supplementary resources to forest
species (Asensio et al. 2009; Mendenhall et al. 2013; Hernández-Ordóñez et al.
2015; Ferreira et al. 2018; Galán-Acedo et al. 2019), and can therefore prevent the
loss of species in small patches (reviewed by Dunning et al. 1992, Arroyo-Rodríguez
et al. 2020). In fact, there is evidence that species-area relationships are shallower
54 V. Arroyo-Rodríguez et al.

(less extinction driven) where matrix quality is higher (Reider et al. 2018). Third,
forest-specialist and generalist species can use these treed elements to move across
the landscape (Mendenhall et al. 2013; Galán-Acedo et al. 2019), allowing individ-
uals to use resources from multiple patches rather than being constrained to the
resources available in a single patch (Fahrig 2013, 2020). This is particularly likely
in Lacandona’s moderately-deforested region, where mean (SD) inter-patch isola-
tion distance (edge-to-edge) in 1000-m radius landscapes is relatively short
(99.8  104.9 m; San-José et al. 2020). Therefore, as predicted by the ‘habitat
amount hypothesis’ (Fahrig 2013) and supported by empirical evidence (Watling
et al. 2020), species density in this region may depend more on forest cover in the
local landscape surrounding the site than on the size of the patch in which the sample
site is located. Although we did not test the effect of forest cover on species density,
our database does not show any relation between forest cover and patch size
(Arroyo-Rodríguez et al. 2013). Thus, our findings are, to some extent, consistent
with this hypothesis, which predicts that if habitat amount (i.e. forest cover) remains
constant, species density should also be independent of patch size (see Fig. 7b in
Fahrig 2013). These results provide potential for a new approach to future research
in the region.
Interestingly, our findings were independent of the degree of habitat specializa-
tion of the species. This contradicts the idea that small patches are mainly valuable
for conserving generalist species, but not for habitat specialist species (Diamond
1976; Willis 1984; Tabarelli et al. 2012; Fletcher et al. 2018). In her recent review,
Fahrig (2020) also found no support for this assumption, possibly because, “as long
as patches are close enough together, persistence across multiple patches could occur
by frequent immigration and/or by individual space use that incorporates multiple
patches” (Fahrig 2020). In fact, as stated above, individuals from many forest
specialist species have been recorded moving through the anthropogenic matrix,
using several small patches as part of their home ranges (Asensio et al. 2009;
Mendenhall et al. 2013; Ferreira et al. 2018; Galán-Acedo et al. 2019). Therefore,
the high conservation value of small forest patches is not limited to generalist
species, as these small patches can also contain forest specialists.
However, it is important to consider that the density of forest-specialist amphib-
ians and reptiles is lower in forest patches than continuous forest. This finding
supports previous studies in the region showing that some amphibian and reptile
species from continuous forest sites are very rare or absent in forest patches
(Hernández-Ordóñez et al. 2014, Russildi et al. 2016). These included amphibians
such as Craugastor alfredi, Smilisca cyanosticta, Smilisca cyanosticta, and
Bolitoglossa mexicana, and reptiles such as Celestus rozellae, Anolis capito and
Porthidium nasutum (Fig. 2.4). Currently there is scarce information on the life
history traits that limit the persistence of these forest-specialist species in forest
patches, but they could have high population variability and restricted habitat needs
(Gibbs 1998). Thus, in agreement with recent proposals to design biodiversity-
friendly landscapes (Arroyo-Rodríguez et al. 2020), a mixed strategy of preserving
many small patches and a few large patches would maximize biodiversity conser-
vation in the region.
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 55

Fig. 2.4 Examples of generalist terrestrial mammals (a–c), and forest-specialist reptiles (d–e) and
amphibians (f–g) that are very rare or absent in all studied forest patches, despite the fact our
sampling effort was higher at forest patches than continuous forest (see Table 2.1). Species
names ¼ Panthera onca (a), Puma concolor (b), Tapirus bairdii (c), Anolis capito (d), Porthidium
nasutum (e), Smilisca cyanosticta (f) and Bolitoglossa mexicana (g). Photographers: Norma Arce-
Peña (pictures a–c), and Martín J. Cervantes-López (pictures d–g)

The density of medium- and large-sized terrestrial mammals was also lower in
forest patches than continuous forest. This is surprising because they are all gener-
alist species. This decrease of medium- and large-sized mammal density might be
related to higher hunting pressures in forest patches than in the continuous forest of
the reserve. Hunting is a well-known driver of terrestrial mammal elimination in
human-modified landscapes (Lamb et al. 2017; Benítez-López et al. 2019; Osuri
et al. 2020; Deere et al. 2020). In fact, our database, which includes large-bodied
vertebrates such as the Jaguar (Panthera onca), Cougar (Puma concolor), Baird’s
tapir (Tapirus bairdii), Collared peccary (Pecari tajacu), and White-tailed deer
(Odocoileus virginianus) (Fig. 2.4; see the complete list of species in Garmendia
56 V. Arroyo-Rodríguez et al.

et al. 2013), are all frequently hunted in human-modified tropical forests, resulting in
“empty” or “half-empty” forests (e.g. Peres 2001; Peres and Palacios 2007; Deere
et al. 2020). Therefore, we speculate that hunting is a major driver of mammal
“defaunation” in forest patches, although this remains to be tested. If this hypothesis
is correct, in addition to preventing forest loss in the region, the conservation of
terrestrial mammals will likely require enforcing strict controls on hunting, as has
already been highlighted for other fragmented regions (e.g. Deere et al. 2020).

2.5 Final Remarks and Conservation Implications

Given the accelerated deforestation and fragmentation of natural forests worldwide,

understanding the importance of forest patch size in preserving biodiversity has
never been so urgent. Our findings suggest that patch size is a poor predictor of
species density in fragmented tropical forests. This does not mean that forest loss
does not have negative effects on biodiversity. Forest loss is a well-known driver of
population decline and species extinction (Fahrig 2003; Watling et al. 2020), and
thus, preventing deforestation in all landscapes, including those where little or no
deforestation has yet occurred, should be a top priority, especially in the tropics
(Peres 2005; Gibson et al. 2011; Phalan 2018; Edwards et al. 2019; Walker et al.
2019; Arroyo-Rodríguez et al. 2020).
In tropical regions that have been already deforested and fragmented, an impor-
tant question is whether preservation and restoration strategies should be focused on
large (Diamond 1975) or small patches (Fahrig 2020). The present study supports
recent conclusions that preservation of all forests is important to biodiversity,
especially preservation of small patches in human-dominated areas (Deane and He
2018; Belder et al. 2019; Wintle et al. 2019; Fahrig 2020). This is important since
conservation strategies usually prioritize large patches. In the Lacandona rainforest,
for instance, payments for ecosystem services are only given to landowners that
preserve forest areas of >100 ha (CONAFOR 2020), leaving smaller patches open to
future destruction. This threatens biodiversity maintenance in the region, because
based on our and previous findings, small patches have, on a per-sample area basis,
as high or higher value for biodiversity conservation as large patches and are not the
near-exclusive habitat of generalist species. Thus, an urgent priority for biodiversity
conservation is to redress the neglect of small forest patches in conservation plans
within this and other biodiversity hotspots.

2.6 Conclusion

Several previous studies have suggested that small forest patches are less suitable to
forest-specialist species due to their high edge-to-area ratio, which amplifies detri-
mental edge effects. This has led to a neglect of small forest patches as key
biodiversity islands that contribute to the preservation of species diversity. Our
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 57

results show that, independently of the taxa and habitat specialization, there are
equivalent levels of species density across different-sized forest patches, and species
density does not differ between continuous forest and forest patches. This contra-
dicts the notion that forest-specialist species are unable to survive in small patches,
likely because they are able to move across the landscape to use resources from
several forest patches and even from the anthropogenic matrix. These findings have
critical applied implications in the context of biodiversity islands. As deforestation
expands, forest patches are becoming increasingly prevalent and smaller, particu-
larly in rainforest landscapes. While prior work has been done to encourage land-
owners to preserve large forest patches, a greater attempt should be made to conserve
small land patches. These small patches not only increase the amount of habitat for
forest species, but they can also be a keystone of species movement, which is why
their preservation should be considered a priority towards the conservation of the
region’s habitat. In examining the different factors that contribute to biodiversity
island design, small forest patches should be given as much consideration as larger
patches because they can often have the same or greater benefit to the wellbeing of
the regional environment and the species within it.

Acknowledgments We are indebted to A. Garmendia, E. Carrara, H. A. Sánchez-de-Jesús,

C. Rojas, M. Solà, I. M. González-Perez, D. Jiménez-González and G. Russildi for collecting
some of the data. This research was supported by Rufford small grants (projects 16237-1, 22049-1,
23706-1, 24642-1), SEP-CONACyT (project 2015-253946), Programa de Apoyo a Proyectos de
Investigación e Innovación Tecnológica (PAPIIT), DGAPA-UNAM (projects IA-203111,
IB-200812, RR-280812, and IN-204215), and Idea Wild. CG-A thanks DGAPA-UNAM for her
postdoctoral scholarship. We thank the landowners in the Marqués de Comillas region for allowing
us to collect data on their properties, as well as the Montes Azules Biosphere Reserve and the
National Commission of Natural Protected Areas. We especially thank Audon Jamangapé, Adolfo
Jamangapé, Gilberto Jamangapé and Marta Aguilar for their invaluable field assistance and
accommodation in the Marqués de Comillas region. Zoe Volenec, Ben Everett-Lane and an
anonymous reviewer made valuable comments on the manuscript that helped us to improve it.

References

Arriaga L, Espinoza J, Aguilar C, Martínez E, Gómez L, Loa E (2000) Regiones terrestres

prioritarias de México. CONABIO, Mexico City
Arroyo-Rodríguez V, González-Perez IM, Garmendia A, Solà M, Estrada A (2013) The relative
impact of forest patch and landscape attributes on black howler monkey populations in the
fragmented Lacandona rainforest, Mexico. Landsc Ecol 28:1717–1727
Arroyo-Rodríguez V, Rojas C, Saldaña-Vázquez RA, Stoner KE (2016) Landscape composition is
more important than landscape configuration for phyllostomid bat assemblages in a fragmented
biodiversity hotspot. Biol Conserv 198:84–92
Arroyo-Rodríguez V, Saldaña-Vázquez RA, Fahrig L, Santos BA (2017) Does forest fragmentation
cause an increase in forest temperature? Ecol Res 32:81–88
Arroyo-Rodríguez V, Fahrig L, Tabarelli M, Watling JI, Tischendorf L, Benchimol M, Cazetta E,
Faria D, Leal IR, Melo FPL, Morante-Filho JC, Santos BA, Arasa-Gisbert R, Arce-Peña N,
Cervantes-López MJ, Cudney-Valenzuela S, Galán-Acedo C, San-José M, Vieira ICG, Slik
58 V. Arroyo-Rodríguez et al.

JWF, Nowakowski J, Tscharntke T (2020) Designing optimal human-modified landscapes for

forest biodiversity conservation. Ecol Lett 23:1404–1420
Asensio N, Arroyo-Rodríguez V, Dunn JC, Cristóbal-Azkarate J (2009) Conservation value of
landscape supplementation for howler monkeys living in forest patches. Biotropica 41:768–773
Banks-Leite C, Ewers RM, Metzger JP (2010) Edge effects as the principal cause of area effects on
birds in fragmented secondary forest. Oikos 119:918–926
Belder DJ, Pierson JC, Ikin K, Blanchard W, Westgate MJ, Crane M, Lindenmayer DB (2019) Is
bigger always better? Influence of patch attributes on breeding activity of birds in box-gum
grassy woodland restoration planting. Biol Conserv 236:134–152
Benítez-López A, Santini L, Schipper AM, Busana M, Huijbregts MAJ (2019) Intact but empty
forests? Patterns of hunting-induced mammal defaunation in the tropics. PLoS Biol 17:
e3000247
Bibby CJ, Burgess ND, Hill DA, Mustoe S (2000) Bird census techniques. Academic, London
Campbell J (1998) Amphibians and reptiles of Northern Guatemala, the Yucatan, and Belize,
Animal Natural History series. University of Oklahoma Press, Norman
Carabias J, de la Maza J, Cadena R (2015) Conservación y desarrollo sustentable en la Selva
Lacandona. 25 años de actividades y experiencias. Natura y Ecosistemas Mexicanos A.C,
Mexico City
Carrara E, Arroyo-Rodríguez V, Vega-Rivera JH, Schondube JE, de Freitas SM, Fahrig L (2015)
Impact of landscape composition and configuration on forest specialist and generalist bird
species in the fragmented Lacandona rainforest, Mexico. Biol Conserv 184:117–126
Chase JM, McGill BJ, McGlinn DJ, May F, Blowes SA, Xiao X, Knight TM, Purschke O, Gotelli
NJ (2018) Embracing scale-dependence to achieve a deeper understanding of biodiversity and
its change across communities. Ecol Lett 21:1737–1751
CONAFOR (2020) Apoyos CONAFOR. Comisión Nacional Forestal, Mexico. [Link]
mx/conafor/acciones-y-programas/apoyos-conafor
Crawley MJ (2007) Statistical modelling in the R book. Wiley, Chichester
Crump ML, Scott NJ Jr (1994) Visual encounter surveys. In: Heyer WR, Donnelly MA, McDiarmid
RW, Hayek LC, Foster MC (eds) Measuring and monitoring biological diversity: standard
methods for amphibians. Smithsonian Institution Press, Washington, pp 84–92
Deane DC, He F (2018) Loss of only the smallest patches will reduce species diversity in most
discrete habitat networks. Glob Change Biol 24:5802–5814
Deere NJ, Guillera-Arroita G, Platts PJ, Mitchell SL, Baking E, Bernard H, Haysom JK,
Reynolds G, Seaman DJI, Davies ZG, Struebig MJ (2020) Implications of zero-deforestation
commitments: forest quality and hunting pressure limit mammal persistence in fragmented
tropical landscapes. Conserv Lett 13:e12701
Diamond JM (1975) The island dilemma: lessons of modern biogeographic studies for the design of
natural reserves. Biol Conserv 7:129–146
Diamond JM (1976) Island biogeography and conservation: strategy and limitations. Science 193:
1027–1029
Dunning JB, Danielson BJ, Pulliam HR (1992) Ecological processes that affect populations in
complex landscapes. Oikos 65:169–175
Edwards DP, Socolar JB, Mills SC, Burivalova Z, Pin Koh L, Wilcove DS (2019) Conservation of
tropical forests in the Anthropocene. Curr Biol 29:R1008–R1020
Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst 34:487–
515
Fahrig L (2013) Rethinking patch size and isolation effects: the habitat amount hypothesis. J
Biogeogr 40:1649–1663
Fahrig L (2020) Why do several small patches hold more species than few large patches? Glob Ecol
Biogeogr 29:615–628
Ferreira AS, Peres CA, Bogoni JA, Cassano CR (2018) Use of agroecosystem matrix habitats by
mammalian carnivores (Carnivora): a global-scale analysis. Mammal Rev 48:312–327
2 The Importance of Small Rainforest Patches for Biodiversity Conservation:. . . 59

Fletcher RJ, Didham RK, Banks-Leite C, Barlow J, Ewers RM, Rosindell J, Holt RD, Gonzalez A,
Pardini R, Damschen EI, Melo FPL, Ries L, Prevedello JA, Tscharntke T, Laurance WF,
Lovejoy T, Haddad NM (2018) Is habitat fragmentation good for biodiversity? Biol Conserv
226:9–15
Galán-Acedo C, Arroyo-Rodríguez V, Andresen E, Arregoitia LV, Vega E, Peres CA, Ewers RM
(2019) The conservation value of human-modified landscapes for the world’s primates. Nat
Commun 10:152
Garmendia A, Arroyo-Rodríguez V, Estrada A, Naranjo E, Stoner KE (2013) Landscape and patch
attributes impacting medium- and large-sized terrestrial mammals in a fragmented rain forest. J
Trop Ecol 29:331–344
Gentry AH (1982) Patterns of Neotropical plant species diversity. In: Hecht MK, Wallace B, Prance
GT (eds) Evolutionary biology. Plenum Press, New York, pp 1–84
Gibbs JP (1998) Distribution of woodland amphibians along a forest fragmentation gradient.
Landsc Ecol 13:263–268
Gibson L, Lee TM, Koh LP, Brook BW, Gardner TA, Barlow J, Peres CA, Bradshaw CJA,
Laurance WF, Lovejoy TE, Sodhi NS (2011) Primary forests are irreplaceable for sustaining
tropical biodiversity. Nature 478:378–383
Hansen MC, Potapov PV, Moore R, Hancher M, Turubanova SA, Tyukavina A, Thau D, Stehman
SV, Goetz SJ, Loveland TR, Kommareddy A, Egorov A, Chini L, Justice CO, Townshend JRG
(2013) High-resolution global maps of 21st-century forest cover change. Science 342:850–853
Harper KA, MacDonald SE, Burton PJ, Chen J, Brosofske KD, Saunders SC, Euskirchen ES,
Roberts D, Jaiteh MS, Esseen P (2005) Edge influence on forest structure and composition in
fragmented landscapes. Conserv Biol 19:768–782
Hernández-Ordóñez O, Martínez-Ramos M, Arroyo-Rodríguez V, González-Hernández A,
González-Zamora A, Zárate DA, Reynoso VH (2014) Distribution and conservation status of
amphibian and reptile species in the Lacandona rainforest, Mexico: an update after 20 years of
research. Trop Conserv Sci 7:1–25
Hernández-Ordóñez O, Urbina-Cardona N, Martínez-Ramos M (2015) Recovery of amphibian and
reptile assemblages during old-field succession of tropical rainforests. Biotropica 47:377–388
Hernández-Ruedas MA, Arroyo-Rodríguez V, Meave JA, Martínez-Ramos M, Ibarra-Manríquez G,
Martínez E, Jamangapé G, Melo FPL, Santos BA (2014) Conserving tropical tree diversity and
forest structure: the value of small rainforest patches in moderately-managed landscapes. PLoS
One 9(6):e98931
IUCN (2019) The IUCN red list of threatened species. Version 2019-3. [Link]
Downloaded on 10 December 2019
Kunz TH (1982) Ecology of bats. Plenum Publishing Corporation, New York
Lamb CT, Mowat G, McLellan BN, Nielsen SE, Boutin S (2017) Forbidden fruit: human settlement
and abundant fruit create an ecological trap for an apex omnivore. J Anim Ecol 86:55–65
Laurance WF, Lovejoy TE, Vasconcelos HL, Bruna EM, Didham RK, Stouffer PC, Gascon C,
Bierregaard RO, Laurance SG, Sampaio E (2002) Ecosystem decay of Amazonian forest
fragments: a 22-year investigation. Conserv Biol 16:605–618
Lee JC (2000) A field guide to the amphibians and reptiles of the Maya world: the lowlands of
Mexico, Northern Guatemala, and Belize. Cornell University Press, Ithaca
Medellín R, Arita H, Sánchez O (2008) Identificación de los murciélagos de México. Clave de
Campo. Asociación Mexicana de Mastozoología, México D.F
Mendenhall CD, Kappel CV, Ehrlich PR (2013) Countryside biogeography. In: Levin SA
(ed) Encyclopedia of biodiversity, 2nd edn. Academic, Waltham, pp 347–360
Newbold T, Hudson LN, Arnell AP, Contu S, De Palma A, Ferrier S, Hill SLL, Hoskins I, Lysenko
AJ, HRP P, Burton VJ, CWT C, Emerson S, Gao D, Hale GP, Hutton J, Jung M, Sanchez-
Ortiz K, Simmons BI, Whitmee S, Zhang H, Scharlemann JPW, Purvis A (2016) Has land use
pushed terrestrial biodiversity beyond the planetary boundary? A global assessment. Science
353:288–291
60 V. Arroyo-Rodríguez et al.

Osuri AM, Mendiratta U, Naniwadekar R, Varma V, Naeem S (2020) Hunting and forest modifi-
cation have distinct defaunation impacts on tropical mammals and birds. Front For Glob Change
2:87
Peres CA (2001) Synergistic effects of subsistence hunting and habitat fragmentation on Amazo-
nian forest vertebrates. Conserv Biol 15:1490–1505
Peres CA (2005) Why we need megareserves in Amazonia. Conserv Biol 19:728–733
Peres CA, Palacios E (2007) Basin-wide effects of game harvest on vertebrate population densities
in Amazonian forests: implications for animal-mediated seed dispersal. Biotropica 39:304–315
Pfeifer M, Lefebvre V, Peres CA, Banks-Leite C, Wearn OR, Marsh CJ, Butchart SHM, Arroyo-
Rodríguez V, Barlow J, Cerezo A, Cisneros L, D’Cruze N, Faria D, Hadley A, Harris SM,
Klingbeil BT, Kormann U, Lens L, Medina-Rangel GF, Morante-Filho JC, Olivier P, Peters SL,
Pidgeon A, Ribeiro DB, Scherber C, Schneider-Maunoury L, Struebig M, Urbina-Cardona N,
Watling JI, Willig MR, Wood EM, Ewers RM (2017) Creation of forest edges has a global
impact on forest vertebrates. Nature 551:187–191
Phalan BT (2018) What have we learned from the land sparing-sharing model? Sustainability 10:
1760
R Core Team (2013) R version 3.0. 1. The R core team, 2001e2013. [Link]
Reider IJ, Donnelly MA, Watling JI (2018) The influence of matrix quality on species richness in
remnant forest. Landsc Ecol 33:1147–1157
Russildi G, Arroyo-Rodríguez V, Hernández-Ordóñez O, Pineda E, Reynoso VH (2016) Species-
and community-level responses to habitat spatial changes in fragmented rainforests: assessing
compensatory dynamics in amphibians and reptiles. Biodivers Conserv 25:375–392
Saito M, Koike F (2013) Distribution of wild mammal assemblages along an urban-rural-forest
landscape gradient in warm temperate East Asia. PLoS One 8(5):e65464
Sánchez-de-Jesús H, Arroyo-Rodríguez V, Andresen E, Escobar F (2016) Forest loss and matrix
composition are the major drivers shaping dung beetle assemblages in a fragmented rainforest.
Landsc Ecol 31:843–854
San-José M, Arroyo-Rodríguez V, Meave JA (2020) Regional context and dispersal mode drive the
impact of landscape structure on seed dispersal. Ecol Appl 30:e02033
Tabarelli M, Peres CA, Melo FP (2012) The ‘few winners and many losers’ paradigm revisited:
emerging prospects for tropical forest biodiversity. Biol Conserv 155:136–140
Taubert F, Fischer R, Groeneveld J, Lehmann S, Müller MS, Rödig E, Wiegand T, Huth A (2018)
Global patterns of tropical forest fragmentation. Nature 554:519–522
Walker RT, Simmons C, Arima E, Galvan-Miyoshi Y, Antunes A, Waylen M, Irigaray M (2019)
Avoiding Amazonian catastrophes: prospects for conservation in the 21st century. One Earth 1:
202–215
Watling JI, Arroyo-Rodríguez V, Pfeifer M, Baeten L, Banks-Leite C, Cisneros LM, Fang R,
Hamel-Leigue AC, Lachat T, Leal IR, Lens L, Possingham HP, Raheem DC, Ribeiro DB, Slade
EM, Urbina-Cardona JN, Wood EM, Fahrig L (2020) Support for the habitat amount hypothesis
from a global synthesis of species density studies. Ecol Lett 23:674–681
Willis EO (1984) Conservation, subdivision of reserves, and the antidismemberment hypothesis.
Oikos 42:396–398
Wintle BA, Kujala H, Whitehead A, Cameron A, Veloz S, Kukkala A, Moilanen A, Gordon A,
Lentini PE, Cadenhead NCR, Bekessy SA (2019) Global synthesis of conservation studies
reveals the importance of small habitat patches for biodiversity. Proc Natl Acad Sci U S A 116:
909–914
Chapter 3
Regenerative Agriculture as Biodiversity
Islands

Brett Levin

Abstract When the amount of biological diversity in an agricultural system is

significantly higher than the baseline biodiversity of the surrounding area, the
agricultural system itself may be recognized as a biodiversity island. Regenerative
agricultural systems, which build and maintain fertility through time, may increase
and maintain biodiversity as an integrated component of food production. Increases
in biodiversity within an agricultural system can span all biological taxonomic
kingdoms and vast numbers of classes and species within each. As such, regenera-
tive agricultural management techniques geared toward harmonizing agricultural
productivity and biodiversity conservation can contribute to mitigating or reversing
detrimental effects of human impacts on landscapes. Greater diversity through
intercropping, companion planting, combinations of perennial and annuals crops,
cover cropping, hedgerows and diverse edge plantings, reduced agrochemical use,
silvopasture with rotational grazing, and selection of rare, heirloom, underutilized, or
diverse genetics allows for biodiversity to harmonize with agricultural production. In
landscapes lacking protected areas or intact ecosystems, habitat restoration and
preservation within agricultural systems can enable both farm productivity and
biodiversity to increase. An integration of restoration and agriculture through farmer
managed natural regeneration, rewilding, and incorporation of traditional ecological
knowledge as operational management approaches within a regenerative agricultural
framework may also achieve such ends. Much of the origins of regenerative agri-
culture emerged from indigenous practice of food production and traditional eco-
logical knowledge that maintains biodiversity. Examples of regenerative agriculture
as biodiversity islands, where farm productivity and improved biodiversity are
achieved, span a multitude of crops, regions, and cultures throughout the world.

Keywords Agroforestry · Cover cropping · Intercropping · Habitat restoration ·

Hedgerows · Reduced agrochemical use · Silvopasture · Traditional ecological
knowledge

B. Levin (*)
Dietrick Institute for Applied Insect Ecology, Ventura, CA, USA
e-mail: [Link]@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 61

F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
62 B. Levin

3.1 Introduction

Historically, expansion of agriculture has contributed to biodiversity loss globally.

Conversion of native ecosystems to less diverse agriculturally productive landscapes
tends to follow trends in global human population (Crist et al. 2017). Generally, a
positive feedback loop exists related to population growth and ecosystem loss; as
human populations increase, more land is converted to agriculture, providing the
calories necessary for further population growth and greater land conversion (Hen-
derson and Loreau 2019, Gustafson et al. 2020). While the technological and
operational achievements of modern industrial agriculture provide relatively inex-
pensive calories to significant populations on a daily basis, the negative social costs
in pollution, farmworker exploitation and health, and biodiversity loss are extensive.
Even with considerable advancements and gains in yield per hectare through genetic
selection and operational improvements, global biodiversity loss trends continue in
response to agricultural expansion (Newbold et al. 2016). An inverse relationship
typically emerges between increased crop specialization with production to achieve
economies of scale and decreased landscape biodiversity and ecological function
(Klasen et al. 2016). From deforestation for cattle ranching in the Amazon to
commodity crop expansion in sub-Saharan Africa and corn and soybean production
throughout the Midwestern United States, the global trends of rising global popula-
tion linked with agricultural expansion and biodiversity loss continue (Fearnside
2017).
With the increase of such landscape degradation, ecosystem loss, and loss of
species diversity throughout the globe, an urgency of design and communication
frameworks is needed to protect biodiversity within the agricultural matrix (Kidd
et al. 2019). Biodiversity islands, which may be defined as areas of high biodiversity
nested within ecologically degraded, human-dominated landscapes, are one such
instrument (Montagnini et al. 2022). The number of species in a biodiversity island
is greater than the biodiversity of the surrounding human managed landscape. As
such, biodiversity islands can take many forms, such as parklands surrounded by
urban sprawl, conserved forestland surrounded by degraded and overgrazed pasture,
or riparian corridors in a monocropped agricultural matrix (Montagnini et al. 2022).
These ecological refugia can provide social, economic, and environmental value
through time. Agricultural systems which utilize the practices of regenerative agri-
culture may harmonize ecosystem and agricultural productivity with biodiversity
conservation, thus operating as biodiversity islands within the landscape.
Regenerative agriculture is an emerging term with a variety of definitions stem-
ming from a diversity of land use approaches, ecological and social contexts, and
lineages of agricultural practice. Consensus regarding an exact scientific definition
for regenerative agriculture poses a challenge and frameworks for socio-economic
and social implementation are sparse within the academic literature (Schreefel et al.
2020). In recent years, the term regenerative agriculture has gained popularity,
differentiating itself from organic, conventional, conservation, or sustainable agri-
culture. Explicit practice-based definitions of regenerative agriculture may by
3 Regenerative Agriculture as Biodiversity Islands 63

limiting, given the system’s approach and broad range of contexts towards which
regenerative agricultural principles may be applied (Soloviev and Landua 2016).
Regenerative agriculture draws from centuries of indigenous and traditional agricul-
tural practices and decades of scientific study and applied research on organic
farming, soil health, agroecology, permaculture, holistic management, and agrofor-
estry around the globe. Generally, regenerative agriculture is a system of farming
principles and practices that increase biodiversity, enrich soils, improve watersheds,
and enhance ecosystem services (White 2020). They aim to capture carbon in soil
and aboveground biomass, reversing current global trends of atmospheric carbon
dioxide accumulation while offering increased yields, resilience to climate instabil-
ity, and higher health and vitality for farming and ranching communities (www.
regenerativeagriculturedefi[Link]). The social aspects of agricultural production
are also addressed by regenerative agriculture, in which production supports just and
reciprocal relationships amongst all stakeholders. While a sustainable system main-
tains itself through time, a regenerative system builds and enhances ecological and
social functioning, recognizing whole systems rather than reductionist viewpoints
(Gibbons 2020). The definition and practice of regenerative agriculture continues to
evolve.
This chapter focuses on regenerative agricultural systems which support wildlife,
biodiversity conservation, and a diversity of genetic resources harmonized with farm
productivity. Such agricultural methods may take many forms, from land sparing
and land sharing, through traditional cultivation methods, and various other
working-lands, agroecological management, and operational techniques (Perfecto
et al. 2009; Gliessman 2016; Altieri 2018; Wagner 2020). Through time, as agricul-
tural practices enhance fertility, sequester carbon, improve soil structure and water
holding capacity, and reduce agrochemical inputs, farm biodiversity may increase as
well (Toensmeier 2016; Rhodes 2017; Meena et al. 2020). When such biodiverse
agricultural areas are within ecologically degraded human dominated surroundings,
they act as biodiversity islands within the landscape. While general practices are
described in this chapter, frameworks for implementation must consider social,
economic, environmental, and cultural circumstances of each location. The follow-
ing techniques and considerations described are useful for farmers, policy makers,
researchers, and decision makers in landscape management.

3.2 Regenerative Management Increasing Biodiversity

Regenerative agricultural systems may be designed and managed to increase the

on-farm presence of cultivated and wild species from numerous taxonomic king-
doms. This can include systems for water catchment, roads and pathway placement,
and crop selection as well as more specific practices such as intercropping,
polycultures, agroforestry, insectary hedgerows, reduced agrochemical use, and
habitat restoration. These practices may also be interwoven with cultural and social
restoration and the use of traditional ecological knowledge to fully foster
64 B. Levin

regeneratively managed agricultural systems as part of the development of biodi-

versity islands within a landscape.

3.2.1 System Design and Management Plans

System design and management plans for increasing biodiversity through regener-
ative agriculture are highly specific to the region, social context, diversity of crop
selection, and particular biophysical attributes of the site, such as geology, soil, and
hydrology (Cabeza and Moilanen 2001; Mendenhall et al. 2014). The degree of
existing ecological degradation, surrounding patch dynamics, and associated popu-
lation ecology as it relates to island biogeography are also important factors deter-
mining species migration and baseline site biodiversity (Tavares et al. 2019). In
severely degraded sites or existing monoculture industrialized agricultural systems,
significant changes in cultural and management practices may be necessary to
increase on-farm biodiversity. Yield of singular specialized crops may need to
decrease to achieve greater on-farm biodiversity, while greater diversified crop
yields, species abundance, and provision of ecosystem services can result (Altieri
2015). In many indigenous, traditional, and agroecological agricultural systems,
management and design integrate biodiversity into production. A key principle of
such agroecological management is designing agricultural ecosystems to mimic the
function of local ecosystems through productive and diverse native species or
agronomic crop analogs (Gliessman 2016; Altieri 2018). Integrating such practice
through improved agricultural methods promotes habitat for a broad range of
microbial, animal, plant, and fungi communities (Altieri 1999; Benayas and Bullock
2012). Sustainable intensification of agriculture through the application of agroeco-
logical principles can also increase trophic complexity, niche formation, and the
biodiversity potential of the agroecosystem (Liere et al. 2017; Atkinson and Watson
2019).
Accordingly, system design and associated management for improved soil health
can greatly increase biodiversity potential (Wagg et al. 2014). Terrestrial ecosystem
functioning and biodiversity are controlled largely by soil microbial dynamics and
soil health, whereas soil health is the capacity of soil to function as an essential living
system, within ecosystem and land-use boundaries, to sustain plant and animal
productivity, maintain or enhance water and air quality, and promote plant and
animal life (Giller et al. 1997; Lehmann et al. 2020). Management practices that
enhance soil health are key indicators of ecosystem productivity and associated
biodiversity (Brussaard et al. 2007).
From the physical and biophysical perspective, proper system design is essential
in developing biodiverse and productive agricultural systems. Well-designed sys-
tems can be more productive, pest resistant, and water efficient, and they conserve
and cycle nutrients more effectively (Doré et al. 2011; Ching 2018). Darren
Dougherty’s Regrarian’s Platform®, built upon P.A. Yeoman’s Keyline Scale of
Permanence, offers one framework for considering factors according to the amount
3 Regenerative Agriculture as Biodiversity Islands 65

of time and effort required for farm modification by humans (Yeomans 1958). The
consideration of these factors, including climate, geography, water, access, forestry,
buildings, fences, soil, economy, and energy, can lead to the lasting commercial and
ecological viability of the agricultural system (Chabay et al. 2015, [Link].
org).
Long-term improved biodiversity can also correlate directly with other farm
system design considerations that utilize an ecosystem approach (Dominati et al.
2019). For example, effective context-specific water management is extremely
important for landscape productivity and improved biodiversity potential. When
culturally, ecologically, and financially appropriate, earthworks designed and
implemented to optimize on-farm water retention can increase the capacity for onsite
cultivation and associated biodiversity (Socci et al. 2019). Keyline design, devel-
oped by agricultural innovator P.A. Yeomans, involves processes such as bringing
water from valleys to ridgelines, to capture, slow, spread, store, and integrate water
on the farm (Yeomans 1958). Utilizing such water management approaches, rain-
water can become a tool for soil building, increased biomass accumulation, and
increased biodiversity, rather than an erosive and potentially polluting force.
Planting arrangement and species selection are also important design consider-
ations. For example, several vegetation strata, including low-lying groundcovers,
understory herbs, low to mid story shrubs, and trees, some of which may be nitrogen-
fixing, reduce dependency on agricultural inputs, enhancing synergisms among both
biological and biophysical system functioning (Nair 2017). As these strategically
designed biodiverse agricultural systems develop, in-situ mulching, improved nutri-
ent cycling, increased water retention, more buffered temperatures, reduced soil
evaporative loss, and increased biological control may result (Schroth et al. 2001;
Lichtfouse 2018). Use of plants and animal breeds adapted to local conditions can
also reduce dependence on foreign inputs and further increase nutrient cycling, soil
fertility, and add biodiversity to the system (Enri et al. 2017; Jose et al. 2019).
Regenerative agriculture may harness such principles, design strategies, and man-
agement techniques to achieve improved biodiversity outcomes for the farm system.
If located within a degraded landscape with low biodiversity, such practices may
allow for a biodiversity island to emerge.

3.2.2 Intercropping and Polycultures

Polycultures and intercropping involve the cultivation of multiple plant species in

mixtures, typically in two or more parallel rows, though they may be planted in
complex assemblages. These practices increase biodiversity and are utilized
throughout the world for several agricultural benefits including diversified yields,
improved biological pest control, weed control, reduced wind erosion, and improved
water infiltration (McLaughlin and Mineau 1995; Corrado et al. 2019). Compared to
a monocrop system, an intercropped system is inherently more biodiverse, given the
increased number of plant species and varieties in cultivation, either planted at the
66 B. Levin

same time or successionally on the same land. Food web interactions and habitat
complexity also increase (Moss et al. 2020). Synergistic relationships of vertebrate,
invertebrate, and microbial communities support and harbor more complex, resilient,
and biodiverse farm ecosystems (Jackson et al. 2007; Zhao et al. 2019).
Intercropping can consist of annuals, perennials, or combinations of the two.
The selection of intercropped species requires knowledge of which species grow
well together. The degree of biodiversity improvements will depend on the species
selected and cultivation strategy. In some instances, weeds are left to grow as a trap
crop for insect pests, increasing invertebrate diversity (Reddy 2017). In other
instances, diversified polycultures are intentionally planted from nursery stock or
seeded in situ. Studies indicate that intercropping increases invertebrate abundance
compared to monocrop systems (Cárcamo and Spence 1994; Tilman 2020). Simi-
larly, in terms of microbial biodiversity, intercropping of diverse landscapes and tree
species results in greater soil microbiological diversity (Lacombe et al. 2009; Chen
et al. 2020). Overall, through intercropping, biodiversity of plants, soil, and animal
life can increase and contribute to the creation of biodiversity islands in otherwise
degraded landscapes.

3.2.3 Agroforestry

Agroforestry systems (AFS), an intensive land management system that optimizes

benefits from the biological interactions created when trees and/or shrubs are
deliberately combined with crops and/or livestock, can increase farm productivity
while supporting biodiversity and providing social and economic benefits for
farmers (Leakey 1999; Jose 2009, 2012; Montagnini and Metzel 2017; Udawatta
et al. 2019; Montagnini 2020, [Link] AFS are heterogeneous in
their design, management, and species composition, and therefore have diverse
values in terms of restoration, productivity, and conservation.
Within AFS, the highest amounts of biodiversity are typically found within
successional AFS, home gardens, forest gardens, and other complex multi-strata
systems (Huang et al. 2002). Other simpler AFS with fewer species of plants or
animals will typically foster less biodiversity, though they may still be considered a
biodiversity island if the surrounding landscape is degraded. In most instances, the
AFS will be the most biodiverse cultivated system of the human dominated land-
scape (Schroth et al. 2004; McNeely and Schroth 2006). Riparian corridors, living
fences, windbreaks, and perimeter hedges may also provide connectivity in
fragmented agricultural landscapes and help bring greater biodiversity to a farm
(Jose 2012). Perennial crops under shade or silvopastoral systems (SPS) with
scattered trees for shade can also provide greater biodiversity and ecological
benefits than monocultures or degraded fallow lands, though providing less biodi-
versity than complex multistrata AFS (Leakey 1999). To favor biodiversity restora-
tion and conservation, AFS should increase their structural complexity in terms
of strata and number of species (Montagnini and del Fierro 2022). Management
3 Regenerative Agriculture as Biodiversity Islands 67

and regenerative agricultural practices that employ agroforestry systems can increase
biodiversity and aid in the development of agricultural lands as biodiversity islands
within the landscape.
Correspondingly, successional agroforestry systems established through enrich-
ment planting can mimic natural regeneration to produce biodiverse and productive
food systems (Young 2017). Successional agroforestry systems consist of multi-
strata multifunctional species assemblages that collectively appear to have a similar
structure to native forests. They can include both introduced and native species.
Native species may emerge from the existing seedbank where seeds are otherwise
unavailable. Tree-growth and crop productivity are achieved by management that
promotes functional characteristics of key natural successional stages of the native
landscape. As stands mature, unique habitats emerge, creating the conditions for
greater biodiversity and opportunities for the establishment of a greater variety of
successional productive species.
Many native or imported species cannot be planted in open plantations because
seedlings are shade tolerant and otherwise will not germinate. As such, enrichment
plantings of food bearing species within degraded landscapes may produce highly
diverse agroforestry systems (Montagnini et al. 1997). Such practices are not new.
For centuries, numerous indigenous cultures recognized the resiliency and food
bearing potential of biodiverse successional agroforestry systems as forest analogues
(Bertsch 2017).

3.2.4 Cover Cropping

Cover cropping, an agricultural technique where pure or mixed stands of perennial or

annual herbaceous plants are grown to cover soil and improve fertility, has also been
shown to increase farm biodiversity. As a tool for regenerative agriculture, cover
cropping legumes, cereals, and other plant mixtures can improve soil structure, soil
fertility, pest management, and biodiversity. Moreover, cover cropping improves
water holding capacity and infiltration, reduces soil erosion, adds organic matter to
soil horizons, cycles nutrients, nourishes the soil food web, reduces weed competi-
tion, and aids in the regulation of soil temperatures (Altieri 2015).
Cover crop management practices vary significantly depending on regional
climate, species selection, tillage and clipping frequency, and time of seeding
(Finch and Sharp 1976). Rye (Secale cereale L.), clovers (Trifolium spp.), vetches
(Vicia spp.), alfalfa (Medicago sativa), and leguminous Pueraria, Stylosanthes, and
Centrosema are commonly planted as cover crops (Altieri 1995). One study indi-
cated the planting of leguminous cover crops (Mucuna pruriens var. utilis) increases
soil macrofauna and nematofauna in maize cultivation (Blanchart et al. 2006). In
rubber plantations, Kuzdu, Pueraria phaseoloides is a nitrogen fixing legume
commonly used as a cover crop. Wild peanuts (Arachis pintoi) are commonly used
as cover crops for coffee and silvopastural systems in Central and South America
(De La Cruz et al. 1994).
68 B. Levin

A practice used from tropical to temperate systems, cover crops also promote
invertebrate diversity, increase populations of beneficial parasitoids, and can
improve biological pest control (Altieri and Schmidt 1985). As a tool in regenerative
agriculture, cover cropping provides multiple benefits and increases belowground
and aboveground biodiversity of the farm system. As such, cover cropping can
enhance the capacity of regenerative agricultural systems as biodiversity islands.

3.2.5 Insectary Hedgerows

Planting a broad range of flowering perennial and annual species in hedgerows

throughout a farm can harbor a diverse and balanced insect ecology while greatly
reducing pest pressure on crops and maintaining on-farm biodiversity in regenerative
agricultural systems (Long et al. 1998; Landis et al. 2000). These insectary hedge-
rows include plants that attract both pests and associated beneficial predatory insects,
providing a breeding ground for beneficial insect populations to increase and expand
into cultivated spaces. Flowering species which bloom in succession throughout the
growing season should be included to ensure that nectar is available to support
beneficial insects throughout the growing season (Holland 2019).
Within insectary hedgerows, plants can perform different functions related to bio-
logical pest control and on-farm biodiversity. Plants that are more attractive to a pest
than the crop plant may be monitored as indicators of pest populations and developing
pest pressures. For example, pole beans are more attractive to spider mites than tomato,
pepper, cucumber, or strawberry. As such, indication of spider mites on pole beans can
allow farmers to control outbreaks before significant crop damage occurs. Similarly,
trap crops are plants which are more attractive to pests than the commercial crop, taking
most of the pest damage and sparing the desired crop (Parker et al. 2016). Used in
conjunction with one another, a monitoring plant can also act as a trap plant.
When beneficial insects become established by feeding on the pests, these trap
crops can become banker crops providing a food source for the increase of beneficial
insect populations (Miller et al. 2017). Banker plants attract and host pests and are
used as an insectary to grow more beneficial insects (Balzan 2017). For example, fast
growing cereal grasses such as ryegrass can be used to attract aphids that become a
food source for aphid predators and parasites. In each of these instances, the simple
increase of plant and arthropod diversity in the system, through the planting of
insectary hedgerows, promotes great biodiversity and establishment of the agricultural
system as a biodiversity island in an otherwise degraded, human dominated landscape.

3.2.6 Reduced Agrochemical Use

Another important aspect of regenerative agriculture as it relates to biodiversity is the

reduction of agrochemical use. Use of pesticides, herbicides, and conventional
3 Regenerative Agriculture as Biodiversity Islands 69

fertilization all may contribute towards decreases in biodiversity (Benton et al. 2003;
Mandal et al. 2020). Agrochemical use may also negatively affect nutrient cycling,
the soil food web, decomposition of soil organic matter, beneficial insects and
natural predator populations. Excessive use of agrochemicals may also increase
NO2 and other greenhouse gas emissions, thus affecting air quality and farmworker
health, which are antithetical to the outcomes and principles of regenerative agri-
culture (Kimbrell 2002).
Insect populations have significantly decreased in recent years, largely attribut-
able to increased pesticide use (Sánchez-Bayo and Wyckhuys 2019). In many
instances, the effects of these chemical applications go beyond their point of use
and can be associated with decreased biodiversity in the broader ecosystem. Runoff
of excess nitrogen and phosphorus in fertilizers enters waterways and reduces
aquatic biodiversity (Ali et al. 2011).
Fortunately, as described in this section, biological practices for pest manage-
ment, weed abatement, and fertility are feasible and can increase on farm biodiver-
sity without agrochemical use (Jørgensen and Kudsk 2006). Such regenerative
practices can also decrease the costs of inputs through time by improving in situ
nutrient cycling (Coleman et al. 1983). Agrochemical applications contradict the
biological practices of regenerative agriculture particularly related to biodiversity.
Therefore, their use should be minimized in the establishment and maintenance of
agricultural systems as biodiversity islands.

3.2.7 Habitat Restoration Within Regenerative Agriculture

Another method to increase on-farm biodiversity is through the restoration of habitat

and ecosystems within low diversity farm systems. Establishing areas of natural
vegetation on farms allows the landscape to fulfill greater ecological function and
provides additional ecosystem services simultaneously with agricultural production.
In degraded lands, restoration of habitat towards these ends directly relates to the
ecological objectives and goals of regenerative agriculture. The co-benefits of
on-farm habitat restoration include production of nonagricultural products, habitat
for various life forms, prevention of soil erosion through runoff and wind, increased
carbon sequestration, and increased water infiltration and watershed health (Benton
et al. 2003). The intentional integration of habitat restoration within the landscape is
therefore a strategy a farmer may choose to implement as part of a regenerative
agricultural system. Examples of biodiversity enhancing on-farm habitat restoration
include farmer managed natural regeneration, successional agroforestry systems
which integrate native species, and rewilding of farmlands.
Farmer managed natural regeneration increases biodiversity and farm productiv-
ity by allowing the existing on-farm sources of regeneration to germinate, grow, and
compete with other vegetation. Through observation and selection of which useful
species emerge, one can manage, tend, and harvest from more diversified farm
ecosystems. This can be achieved by allowing natural regeneration to take place in
70 B. Levin

fields or on selected patches within the farm (Wintle et al. 2019) Valuable species are
selected to persist, thus creating a low-cost and biodiverse foundation from which
productive agricultural systems may emerge. For example, in the Sahel region of
Niger, rather than weeding all species, farmers may select specific species to remain
in the fields. By caring for these naturally regenerating drought tolerant species,
greater diversity and yields result. These practices have been a contributing factor in
a low-cost option for increased diversity and indigenous genetics of gardens and
agriculture throughout the region (Reij and Garrity 2016).
Allowing succession to occur in a slightly more hands-off approach may be
known as rewilding or natural regeneration. Allocating land for rewilding, some
areas of crop cultivation may be lost, but the trade-off results in greater diversity,
pollination, and other ecosystem services (Navarro and Pereira 2015). These areas
may also be seeded with a diversity of desired annual and perennial species, with
minimal continued management.
The beneficial outcomes of natural regeneration on sections of farmland are
particularly clear in certain grazing systems. For example, The Knepp Wildland
Project in the United Kingdom originally utilized a traditional pastureland. As cattle
were removed from sections of the farm, those areas underwent rapid natural
regeneration. In some areas, existing seed banks were able to emerge and other
areas were seeded with desirable species. After tree establishment, cattle were
reintroduced to sections of the farm, where they had access to increased forage
and greater shade, functioning as a silvopastoral system (Tree 2019). This integra-
tion of rewilding and natural regeneration provided habitat for a vastly greater
number of local species, while still providing farm yields. The farm was transformed
into a biodiversity island within the landscape.

3.2.8 Silvopasture and Rotational Grazing

Silvopasture with rotational grazing is another management strategy which can

improve agricultural biodiversity as part of a regenerative agricultural system (Jose
et al. 2019). Silvopasture is a type of agroforestry system consisting of the inten-
tional combination of trees, forage plants and livestock together as an integrated,
intensively-managed production system ([Link] Silvopasture
can provide profitable opportunities for tree growers, forest landowners, and live-
stock producers through the integration of what are typically separate production of
tree crops and livestock. The benefits of rotational grazing are site and context
specific but can include improved forage production, soil health, fertility, soil carbon
storage, drought resistance, weed control, human and animal relationships, animal
welfare, an extended grazing season, reduced forage waste, and reduced parasite
problems (Orefice and Carroll 2017; Jose et al. 2019). Combined with silvopasture,
additional economic benefits of tree production may emerge such as reduced fertil-
izer requirements, improved yields, increased weight gain, and reduced fodder needs
(Gabriel 2018). Compared to monoculture tree cultivation or continuous grazing,
3 Regenerative Agriculture as Biodiversity Islands 71

silvopasture with rotational grazing can greatly increase biodiversity (McDermott

and Rodewald 2014). Silvopastoral systems can harbor a high diversity of cultivated
species in addition to a wide range of arboreal wildlife habitat.
As for rotational grazing, the diversity, quality, and longevity of forage species
can increase when adequate rest is given to the grazed area, when compared to a
continuous grazing system. Such outcomes are dependent on stocking rate, paddock
size, longevity of grazing, and regional climatic and biophysical factors of the farm
(Gabriel 2018). If managed optimally for the appropriate context of the farm,
rotational grazing provides opportunities for a greater variety of forage species to
persist and for greater profitability (Orefice et al. 2019). Additionally, greater farm
biodiversity may be present in rare, native, or unique livestock breeds as well as the
incorporation of different species including goats, chickens, ducks, pigs, cattle,
buffalo, and others (Gabriel 2018). With proper management, silvopasture and
rotational grazing can allow for greater biodiversity to emerge (McAdam and
McEvoy 2009).
With both silvopasture and rotational grazing, one should learn the benefits as
well as the risks before adopting the practices. Integration of multi-species grazing
schemes may increase parasite loads if not managed properly. In certain areas,
legislation prevents grazing on lands used for the cultivation of food crops within
a specified time period preceding harvest in order to prevent contamination risks.
Transition of land into silvopasture or grazing areas without proper management can
damage soil, cause erosion, and eliminate opportunities for natural regeneration
when appropriate or desired (Orefice et al. 2019). The complexity and diversity of
approaches for integrating silvopasture and rotational grazing depends on farm
location and larger holistic framework of farm context (Savory and Butterfield
1998). If properly applied and managed, silvopasture with rotation grazing is an
agricultural practice which may increase on-farm biodiversity and allow for a farm to
become a biodiversity island within a degraded landscape.

3.2.9 The Use of Rare, Heirloom, and Underutilized Species

and Cultivars

When unique, rare, and diverse species of plants and livestock are cultivated in
regenerative agricultural systems, these farm systems can serve as biodiversity
islands within a human dominated and degraded landscape. Greater crop diversity
of cultivated species increases the overall biodiversity of the agricultural system and
can allow for increased food security, decreased pest pressures, more resilience to
climate change, and enhanced connection between cultures and locally produced
foods (Smith et al. 2008; Chateil et al. 2013; Gaudin et al. 2015).
Rare, heirloom, regional and family cultivars of fruit and vegetable crops were
once commonplace globally, though an inverse relationship tends to exist between
industrialized agriculture expansion and landrace presence and diversity (Nazarea
72 B. Levin

2005). Fortunately, farms, organizations, and community groups are working to

continue to keep such species alive while increasing genetic diversity through time
(Abebe 2005). Regenerative agriculture systems such as urban community gardens
and homegardens commonly cultivate genetically diverse and heirloom crops
(Bardhan et al. 2012; Redondo-Brenes and Montagnini 2010). Conservation of
on-farm crop diversity is extremely important to both biodiversity and the cultures
from which these crops arose (Brush 2000). In situ and ex situ methods of conser-
vation allow for genetic resources to be preserved through time while expanding
crop diversity (Swanson and Goeschl 2000). More diverse crops have the potential
to support greater soil life diversity and insect diversity, with differing phenologies,
nutrient requirements, decomposition rates, and structure (Redlich et al. 2018).
When in urban environments, these systems can be important refugia for biodiver-
sity, as well as places where people can connect with nature (Toensmeier 2022).
When rare, heirloom, or underutilized species are cultivated, seeds saved and passed
on through time, greater genetic diversity can persist in the human dominated
landscape.
Maintaining the diversity of plants in cultivation can take many forms. Rowen
White, through her work as an indigenous seed breeder, cofounder of the Sierra
Seeds Cooperative, and chair of the board of The Seed Savers Exchange, emphasizes
the genetic, cultural, and historical importance of seed saving (White et al. 2018).
The Felix Gillet Institute, founded by the late Amigo Bob Cantisano, explores
neglected homesteads and agricultural sites seeking surviving heirloom varieties of
fruit and nut trees throughout California ([Link] Plant explorers
such as David Fair-Child traveled the globe bringing diverse and underutilized
species into cultivation (Fair-Child 1939). Plant breeders such as the famed Luther
Burbank utilize innovative breeding techniques to greatly expand favorable charac-
teristics and useful varieties and diversity of plants in cultivation (Burbank 1915).
Through seed saving, recovery of heirloom varieties, exploration of underutilized
species, and innovative breeding for new genetics, the diversity of cultivated plants
can expand, even when challenged by economic forces spurring an opposite trend
(Nazarea 2005). Regenerative agriculture can utilize the broad range of cultivar
diversity to keep the genetics, stories, and species diversity alive. In doing so,
regenerative agricultural sites can continue to develop as biodiversity islands within
the landscape.

3.3 Social Dimensions of Regenerative Agriculture

as Biodiversity Islands

In attaining greater biodiversity in regenerative agricultural systems, social and

cultural factors should not be overlooked. Restorative action can go beyond practices
of cultivation and ecological management to address restoration and regeneration of
community and human relationships. Respect for the cultural origins of regenerative
3 Regenerative Agriculture as Biodiversity Islands 73

agriculture and the historical ecology of cultivated lands, farmworker health,

empowerment, right-livelihoods, and the affordability and access to regeneratively
cultivated foods by consumers must integrate with enhanced biodiversity for the
agricultural system to be truly regenerative. The potential to address these social
considerations is a key task for regenerative agriculture into the future.

3.3.1 Traditional Ecological Knowledge: The Roots

of Regenerative Agriculture

Traditional Ecological Knowledge (TEK) or Ecological Indigenous Knowledge

(EIK) and the historical ecology of cultivated lands may contribute significantly
towards the development and continuation of food production systems which act as
biodiversity islands within a landscape. Much of regenerative agriculture is built
upon this knowledge. When not already in place, indigenous, knowledge-holding
representatives should hold positions of authority and decision-making within agri-
cultural organizations. Land return to indigenously managed lands is another
approach towards empowerment and social restoration in regenerative agriculture.
Many indigenous communities inhabit areas where the diversity of plant and animal
species have been utilized for thousands of years (Rocha et al. 2017). This can be
seen in many terraced landscapes throughout the globe (Fig. 3.1).
Most inhabited places of the earth have an associated historical ecology, although
in many instances this knowledge has been deeply eroded due to various social and
economic factors (Balée 1998). Where traditional cultures remain, often certain
members of the indigenous culture still hold knowledge of traditions and practices
related to cultivating, managing, processing, and consuming diverse, native species
(Berkes et al. 1994). For example, there is extensive knowledge of Native American
management and cultivation of California’s pre-colonial landscape (Anderson
2013). In areas where such knowledge is nearly lost from collective memory or
culture, there is significant opportunity in the rediscovery and re-empowerment of
such traditional knowledge as a fundamental component of productive and biodi-
verse regenerative agricultural systems of the future.
Technical and scientific knowledge of such traditional ecological knowledge may
provide medicinal, nutritional, and otherwise valuable products. Among many
others, guayusa, yerba mate, and cacao have well-documented examples of indige-
nous knowledge being integrated with scientific techniques to develop modern
cultivation practices. In instances where local knowledge is used to gain insights
into cultivation, harvesting, and processing, and bringing new products to market,
one must be cautious to avoid exploitation and acknowledge the social responsibility
within regenerative agriculture. For example, Yoco (Paullinia yoco), a vine that is
wild-harvested for its caffeinated bark by the Secoya people, has had populations
greatly reduced in recent years. A Yale researcher began working to restore
populations of Yoco by designing systems with the community to enrich the forests
74 B. Levin

Fig. 3.1 A village outside Muktinath, Nepal. Irrigated terrace agriculture with incorporation of
various annual grains and tree crops, adding to the biodiversity of the landscape. Techniques of
traditional and regenerative agriculture allow for subsistence farming, organic nutrient cycling, and
efficient use of water to transform otherwise inhospitable terrain into a biodiverse landscape.
(Photo: Brett Levin)

with more yoco while monitoring key environmental outcomes of these systems.
(Fig. 3.2) The project looks very carefully under which ecological conditions the
yoco can be established, such as light, soils, and species assemblages. This connec-
tion between traditional, technical, and community empowerment is an example of
how regenerative agriculture can enhance biodiversity through the incorporation of
traditional ecological knowledge. These productive sites emerge as biodiversity
islands within landscapes that are rapidly being cleared due to logging and conver-
sion of forest to pasture.
Correspondingly, ethnobiology and ethnophenology address the human and
cultural component of how species and genetics are selected through time (Nabhan
2016). Ethnobiology explores the complex interactions among cultures, their lan-
guages and resource management practices with genes, foods, medicines, habitats,
and landscapes for addressing critical links between culture, cultivation, and eco-
logical diversity. Ethnophenology refers to the cultural perception of the timing of
recurrent natural history events and environmental conditions in the selection and
managing of specific species. For example, records from the early 1900s provide
anecdotal evidence that for the Hidatsa people of the Missouri River, the sunflower
seed was always the first seed planted in the spring based on observations of the
melting of ice along the banks of the Missouri River around April. This was followed
by planting of corn in May based on the observation of the emergence of leaves of
3 Regenerative Agriculture as Biodiversity Islands 75

Fig. 3.2 Expanding the traditional Secoya people cultivation through forest enrichment with Yoco
in a multilayered, biodiverse forest system in the Amazon of Ecuador. (Photo: Luke Weiss)

the wild gooseberry bushes. These strong observations of environmental cues are
learned by cultures with longstanding connections to land.
Through the incorporation of greater biodiversity into farm systems, regenerative
agriculture can learn and build upon these traditions to provide a more perceptive,
inclusive, and harmonious approach to cultivation, rather than mandate by strict
agronomic management procedures (Nabhan 2014; Albuquerque and de Sousa
2016). Regenerative agriculture farms may become locations of applied practice
towards cultural restoration and biodiversity enhancement in addition to agronomic
cultivation.

3.3.2 Farmworker Health, Empowerment,

and Right-Livelihood

Farmworker health, empowerment, and right-livelihood are other important social

matters that must be considered as part of biodiversity-enhancing regenerative
agricultural systems. Is an agricultural system that enhances biodiversity but harms
and exploits its workers truly regenerative? Farmworker health issues caused by
exposure to toxic agrochemicals is rampant in industrialized agriculture (Salzman
76 B. Levin

2018; Saxton 2021). By implementing biodiversity-enhancing agricultural systems,

farmworker health can improve (Afshari et al. 2021). Opportunities in farmworker
training and empowerment may also be integrated into biodiversity-friendly agri-
cultural systems (Braun and Duveskog 2011). One such opportunity may be the
training of farmworkers to recognize, monitor, and collect data on soil health and
pest pressures, which can allow for more targeted biological approaches towards
cultivation and pest management. A survey of Farmer Field School (FFS) programs,
which employ local knowledge sharing and training in pest management, not only
reduced pesticide use and improved associated farmworker health, but proved
economically advantageous (Rejesus and Jones 2020).
Additionally, workers should be paid a living wage for their time and energy if the
agricultural system is to be recognized as regenerative. Worker-ownership and
cooperative business structures can provide long-term equity, wealth building and
authority to all levels of the organization while biodiversity practices are
implemented (Alkon and Guthman 2017). Such approaches are utilized in coffee
plantations throughout El Salvador and farmworker cooperatives of the United
States (Bacon et al. 2008; Gray 2013). Sylvanaqua farms is a unique example in
the Washington DC area focused on both the social and ecological components of
regenerative agriculture ([Link] Overall, for a farm system
that enhances biodiversity to be recognized as regenerative, it must also consider the
health and wellbeing of the farmers who work the land.

3.3.3 Affordability and Access to Regenerative Agriculture

The lack of affordability and access of regeneratively grown crops and associated
products is another often overlooked social dimension of regenerative agriculture. If
only the wealthy can afford regenerative agriculture, is it truly regenerative? When
on-farm biodiversity improvements cause prices to increase, low-income consumers
become excluded from the market. Similarly, when the negative social costs of
degradative agriculture are not factored into pricing, prices remain artificially low
(Pascual and Perrings 2007).
Such issues can be addressed through numerous progressive and grassroots
strategies. Removal of existing governmental subsidies that support degradative
practices is essential. Support of new subsidies for regenerative practices advance
affordability. Negative social costs such as biodiversity loss, soil loss, greenhouse
gas emissions, water quality degradation, and effects of industrial agriculture on
human health must be factored into pricing (Mouysset et al. 2015). Payments for
ecosystem services can also reduce prices of food and products to consumers that are
grown regeneratively (Lankoski 2016). Additionally, with grassroots efforts to
develop local regenerative agricultural systems and community supported agricul-
ture projects as biodiversity islands, low-income access and affordability can
improve. Examples are numerous and worldwide, spanning urban to rural areas
3 Regenerative Agriculture as Biodiversity Islands 77

lacking access to regeneratively grown produce (Adam 2006; Duchemin et al. 2008;
Lovell 2010).
Overall, a transition from degenerative agricultural practices to regenerative
practices requires a cultural shift to one that sees natural systems as essential,
valuable, and inherently interconnected. Regenerative agricultural practices that
increase biodiversity should also improve social and human wellbeing. Much of
the origins of regenerative agriculture emerged from indigenous practices of food
production and traditional ecological knowledge that maintains biodiversity. Rec-
ognizing, appreciating, and empowering this history is an essential component of the
story of regenerative agriculture that is commonly appropriated, dismissed, or
ignored. For agriculture to be truly regenerative, it must use a systems approach
and consider impacts to the interrelated human systems that make cultivation,
distribution, and food access possible.

3.4 Regenerative Agriculture in the Modern Age

As regenerative agriculture continues to expand, it is important to highlight some of

the key organizations currently shaping the conversation. Future considerations
regarding funding, monitoring of environmental outcomes, and education are essen-
tial for the continued widespread adoption of regenerative agriculture which may act
as biodiversity islands.

3.4.1 Organizations Supporting Regenerative Agriculture

for Biodiversity Conservation

The promotion and adoption of regenerative agriculture continues to expand in

various sectors. Organizations and projects supporting the advancement of biodi-
verse regenerative agriculture are diverse and worldwide. Investment entities, farms,
service organizations, consumer packaged goods manufacturers, and nonprofit
organizations are rapidly expanding their language and practices surrounding regen-
erative agriculture ([Link]
try-map/). The International Federation of Organic Agriculture Movements
(IFOAM), the National Center for Appropriate Technology (NCAT) and their
ATTRA program, Ecological Farming Association (Ecofarm), The Tropical Agri-
cultural Research and Higher Education Center (CATIE), The Society of Ethnobi-
ology, and countless other organizations support research and communication of
these ideas. Some additional examples are reviewed in greater detail below to
highlight the diversity of geography and scope of work from which supporting
organizations exist.
78 B. Levin

SOCLA (Sociedad Cientifica de Agricultura Latino Americana de Agroecologia,

Latin America Scientific Society of Agroecology) is a network of researchers, pro-
fessors, extensionists, and other professionals which promotes agroecological and
regenerative practices to confront the crisis of industrial agriculture. SOCLA pro-
motes agroecology as a scientifically justifiable and sustainable rural development
strategy in Latin America. To accomplish its objectives SOCLA organizes scientific
congresses, holds short training courses in various countries, produces publications
on key issues, and maintains working groups that provide information, analysis and
technical advice to numerous civil and farmers organizations involved in agroecol-
ogy in the region. Recently, the North American SOCLA chapter was launched. This
work promotes the integration of greater biodiversity within regenerative agricul-
tural systems ([Link]
Rodale Institute has been a leader in organic and regenerative agriculture since its
founding in 1947 in Kutztown, Pennsylvania. As a pioneer in this field, they support
new farmers, contribute valuable research, and educate consumers regarding the
benefits of organic products. A key component of their work encourages biodiversity
through regenerative agriculture. Rodale recognizes that a rich mix of microorgan-
isms, plants, and animals on the farm creates healthy soil, strong crops, and resilient
natural systems that don’t require chemical intervention to manage pests and dis-
eases. This knowledge is shared broadly through their public, outreach and educa-
tion, in addition to being applied on their own agricultural land. In 2018, Rodale
Institute helped spearhead a new, holistic, high-bar standard for agriculture certifi-
cation. Regenerative Organic Certification, or ROC, is overseen by the Regenerative
Organic Alliance, a non-profit made up of experts in farming, ranching, soil health,
animal welfare, as well as farmer and worker fairness ([Link]
The certification consists of three pillars: soil health, animal welfare, and social
fairness. Attaining certification supports approaches to land management and asso-
ciated processes that contribute to the health of ecosystems and human communities.
The Land Institute, founded by Wes Jackson in the 1970s, has been working to
develop perennial grain crops and support biodiverse polycultures. Located in
Salinas, Kansas, they have recently had success with the development of a new
species of perennial grain called Kernza, which has the potential to transform much
of the world’s grain production into perennial agriculture, thereby contributing to
soil protection and the preservation of waterways. The Land Institute is also devel-
oping a crop protection program that relies on biological control using natural
enemies ([Link]
The Al Bahaya project in Saudi Arabia is transforming a barren desert into
productive savanna grasslands and agroforestry systems using regenerative agricul-
ture techniques and extensive stone terracing to capture water. Choosing the appro-
priate species has been essential. When the system was first established, irrigation
was utilized and later it was cut off. For 31 months there was no precipitation. After it
finally rained, the species were able to recover and begin a biological cascade
towards rejuvenation. ([Link] Here,
once the system was established, regenerative agricultural processes utilize the
resources available, no matter how scarce, to build biodiversity and productive
3 Regenerative Agriculture as Biodiversity Islands 79

agricultural systems. Such considerations are the basis of the future of regenerative
agriculture as biodiversity islands within degraded landscapes.
The Savanna Institute, a nonprofit organization located in Wisconsin, is a leader
in temperate agroforestry research, laying the groundwork for widespread agrofor-
estry in the Midwest US. Working in collaboration with farmers and scientists, the
Savanna Institute is developing perennial food and fodder crops within
multifunctional polyculture systems, grounded in ecology, and inspired by the
savanna biome, with an emphasis on tree crops. Chestnuts and hazelnuts tend to
be the backbone of The Savanna Institute’s diverse agroforestry systems and they
strategically enact their mission via research, education, and outreach (http://
[Link]/).
Numerous family farms with goals of integrating biodiversity and food produc-
tion also continue to emerge. New Forest Farm is a diverse restoration agriculture
research site in southwestern Wisconsin, USA. Located on a former cornfield,
through the efforts of Mark Shephard, the land has been transformed into a biodi-
verse perennial agriculture ecosystem. Utilizing innovative water management tech-
niques, various trees, shrubs, vines, canes, grasses, forbs and fungi have been
planted, organized to optimize yield and efficiency in harvesting and management.
Woody crops include hazelnuts, chestnuts, walnuts, and apples (Shepard 2013). The
diverse plantings and biology present within New Forest Farm make it a biodiversity
island within the surrounding vast expanse of monoculture corn and soy production.
Polyface farm is another example of a biodiverse regenerative agriculture family
farm. Spearheaded by the Salatin family, the operation produces pastured poultry
and a broad range of crops focusing on soil health, community health, and the
continued improvement of the land base. Through time, measured improvements
in biodiversity have resulted (Salatin 2010). Such operations as biodiversity islands
within the landscape integrate old farm knowledge with new innovations, paving the
future of a new, regenerative, and biodiverse agricultural paradigm.
The Savory Institute and Holistic Management International both promote, advo-
cate, and teach about regenerative agriculture through holistic rangeland manage-
ment and holistic decision making ([Link] [Link]
global/). Holistic management was born from the work of Allan Savory, a Zimba-
bwean ecologist. Properly managed livestock are the ecological foundation of the
holistic context. The general objectives are to help ranchers and land stewards
strengthen local economies, improve local food quality, heal the environment,
improve wildlife habitats, and enhance community. The teachings train farmers to
recognize their goals, plan appropriately based on specific contexts, and manage
livestock to mimic natural ecological patterning of mob grazing while improving soil
carbon sequestration and overall rangeland biodiversity as compared to conventional
grazing and cattle raising operations.
These organizations are a small sampling of many more groups focused on
advancing biodiversity through regenerative agriculture. It is also important to
recognize the millions of smallholders practicing similar techniques and sharing
traditional knowledge throughout the world. As awareness and interest continues to
grow for increasing biodiversity in degraded landscapes while producing food, one
80 B. Levin

can expect the influence of these bodies to continue to expand and new organizations
to continue to emerge.

3.4.2 Considering the Future: Funding, Monitoring,

and Education

Alongside private sector approaches, governments can continue to support and grow
programs for agricultural practices that encourage farmers to increase farm biodi-
versity. Governments can work towards goals of increased agricultural biodiversity
in the same way successful widespread adoption of organic programs in Europe took
place. This was achieved through increased funding of training programs, offsetting
certification costs, and improving the quality of government advisory services, all of
which have proven highly effective (Mills et al. 2020). In the United States, the
Department of Agriculture and the Natural Resources Conservation Service cur-
rently have several financial incentives for farmers to adopt practices such as riparian
corridors, windbreaks, and hedgerows (Duru et al. 2015). The Environmental Qual-
ity Incentives Program (EQIP) aids agricultural producers through technical and
financial support through public funding to address natural resource degradation and
to improve the environment through increased water and air quality, conserved
ground and surface water, increased soil health, reduced soil erosion, improved or
created wildlife habitat, and mitigation against increasing weather volatility through
public funding. Of these conservation practices, many contribute to the development
of agricultural biodiversity islands within a landscape ([Link]
wps/portal/nrcs/main/national/programs/financial/eqip/). Though well-funded with a
budget of $1.75 billion in fiscal years 2019 and 2020, $1.8 billion in fiscal year 2021,
$1.85 billion in fiscal year 2022 and $2.025 billion in fiscal year 2023, there remains
opportunity for greater financing of biodiversity enhancing conservation practices
([Link]
farm-bill). This type of financial assistance can be greatly expanded upon, and
include all the previously mentioned practices, which can increase farm resilience,
yields, and on-farm biodiversity. Within the United States, this can be addressed
through a revision of funding priorities federally in the Farm Bill, and locally
through state action and cooperative extensions.
Additionally, as private funding and markets for payments for ecosystem services
and carbon sequestration in agriculture continue to advance, it is important to
consider the potential to integrate biodiversity within such projects. Many carbon
offset projects focus solely on biomass production and carbon sequestration. Focus-
ing on the maximum biomass growth possible to obtain as many carbon credits as
possible may place higher value on fast growing species than native, bio-regionally
appropriate food-bearing species. In these instances, where projects focus on bio-
mass generation for either carbon or bioenergy, biodiversity can decrease through
time rather than improve (Abreu et al. 2017). By incorporating some of the practices
3 Regenerative Agriculture as Biodiversity Islands 81

mentioned above, these projects can have mutually beneficial outcomes of biomass
production, sequestration, and improved biodiversity outcomes.
Advancements in monitoring of biodiversity coupled with carbon sequestration
and other ecosystem services may provide another significant increase in the adop-
tion of regenerative agriculture. When the benefits and positive impacts of these
practices are measurable with greater certainty, value can be associated with such
practices, and the positive social benefits can be attributed to individual farms and
farmers. The externalities of any farm, positive or negative, influence the rest of the
landscape. When such externalities are properly monitored and valued, society is
more able to perceive those benefits, which in turn makes regenerative agriculture
more attractive. This opens further opportunity for community engagement, invest-
ment, funding, and more widespread adoption of biodiverse regenerative farming,
sparking the development of biodiversity islands throughout degraded landscapes
([Link] [Link]
Online educational opportunities for learning regenerative agricultural practices
that enhance biodiversity outcomes have grown significantly. Reports, podcasts,
webinars, workshops, conferences, virtual university extension programs which are
now widely available for free, provide information that is both conceptual and
specific for bioregional applied practice. Many examples of such media can be
found on websites and platforms such as, [Link] https://
[Link]/, [Link] [Link] [Link]
[Link]/en-us/, [Link] [Link]
[Link] and others mentioned in Chap. 1 Sect. 1.5 of this
volume.
Additionally, there are a growing number of technical and scientific publications
accessible to a broad audience, such as Working with Nature: Resource Management
for Sustainability (Jordan 1998), Tomorrow’s Biodiversity (Shiva 2000), Call of the
Reed Warbler (Massy 2017), Growing a Revolution (Montgomery 2018), and
Reclaiming the Commons (Shiva 2020). Such resources and writings are inspiring
a new generation of educators, policy makers, and farmers to engage in the work of
developing biodiverse regenerative agricultural systems which may act as biodiver-
sity islands within the landscape.
Regenerative agriculture emerged from traditional knowledge and ecological
observations through time. While conducted mostly by indigenous people and
smallholders throughout the world, over the past century, writers and practitioners
worldwide have continued to advance the science and practice of regenerative
agriculture in the western paradigm. Such notable proponents include Amigo Bob
Cantisano, Bill Mollison, Christine Jones, Cyril G Hopkins, Darren Dougherty,
David Montgomery, Edward Faulkner, Eric Toesnmeier, Ethan Soloviev, Eve Bal-
four, Everette “Deke” Dietrick, F.H. King, Gabe Brown, J. Russell Smith,
J.I. Rodale, Joel Salatin, John Jeavons, John Kempf, John Lundgren, Judith
Schwartz, Miguel Altieri, Leah Penniman, Mark Shepard, Masanobu Fukuoka,
Newman Turner, P.A. Yeomans, Reginaldo Haslett-Marroquin, Richard Perkins,
Rudolph Steiner, Sir Albert Howard, Thomas Barrett, Vandana Shiva, William
Albrecht, Wendell Berry, and many others. Through an ever-growing application
82 B. Levin

of scientific, philosophical, ethical, and on-the-ground practice, the role and impact
of biodiverse regenerative agriculture continues to expand, increasing the develop-
ment of biodiversity islands in degraded lands.

3.5 Conclusions

Biodiversity enhancing regenerative agricultural practices can be applied wherever

agriculture is practiced. Throughout the twentieth century, a shift in agricultural
production to large scale, industrial, monoculture production with the use of agro-
chemicals became commonplace. While providing cheap calories, this agriculture
has a myriad of negative consequences for both humans and the environment. This
chapter has described successful examples and techniques for the advancement of a
different approach towards agriculture, where agrochemical use is limited or elim-
inated, diverse genetics are utilized, and design and agricultural techniques are
examined to enhance biodiversity within agricultural systems.
Much of regenerative agriculture emerged from indigenous land use systems.
Building from this knowledge as science, diverse agroecosystems can continue to
spread throughout the globe with great success. A diversity of organizations and case
studies were presented to highlight the scope, scale, and diversity of regenerative
agriculture as it contributes to biodiversity islands in a landscape. If managed
following the approaches described throughout this chapter, farms can become
biodiversity islands within the matrix of degraded landscapes.
Financial, cultural, and ecological opportunities abound in the transition of
degraded lands into agriculturally based biodiversity islands. As adoption of biodi-
versity into agricultural lands becomes more commonplace and more farms through-
out the landscape harbor greater levels of biodiversity, one can envision an
agricultural future in which biodiversity islands are the norm. As such, speciation
and species preservation may continue to increase through time in harmony with
human habitation and agricultural production, and the biodiversity island which is
planet earth may flourish towards a greater bounty and beauty that is evident in the
diversity of life.

References

Abebe T (2005) Diversity in homegarden agroforestry systems of southern Ethiopia. PhD thesis,
Wageningen University, Wageningen
Abreu RC, Hoffmann WA, Vasconcelos HL, Pilo NA, Rossatto DR, Durigan G (2017) The
biodiversity cost of carbon sequestration in tropical savanna. Sci Adv 3(8):e1701284. https://
[Link]/10.1126/sciadv.1701284
Adam KL (2006) Community supported agriculture. ATTRA-National Sustainable Agriculture
Information Service, Butte. [Link]
3 Regenerative Agriculture as Biodiversity Islands 83

Afshari M, Karimi-Shahanjarini A, Khoshravesh S, Besharati F (2021) Effectiveness of interven-

tions to promote pesticide safety and reduce pesticide exposure in agricultural health studies: a
systematic review. PLoS One 16(1):e0245766
Albuquerque UP, de Sousa DCP (2016) Ethnobiology and biodiversity conservation. In: Albuquer-
que UP, de Sousa DCP (eds) Introduction to ethnobiology. Springer, Cham, pp 227–232
Ali M, Khan SJ, Aslam I, Khan Z (2011) Simulation of the impacts of land-use change on surface
runoff of Lai Nullah Basin in Islamabad, Pakistan. Landsc Urban Plan 102(4):271–279
Alkon A, Guthman J (eds) (2017) The new food activism: opposition, cooperation, and collective
action. University of California Press, Berkeley
Altieri MA (1995) Biodiversity and biocontrol: Lessons from insect pest management. Academic
Press Adv in Plant Path 11(10):191–209
Altieri MA (1999) The ecological role of biodiversity in agroecosystems. In: Altieri MA
(ed) Invertebrate biodiversity as bioindicators of sustainable landscapes. Elsevier, Amsterdam,
pp 19–31
Altieri MA (2015) Agroecology, key concepts, principles and practices. Penang, Third World
Network (TWN)-SOCLA, Jutaprint
Altieri MA (2018) Agroecology: the science of sustainable agriculture. CRC Press, Boca Raton
Altieri MA, Schmidt LL (1985) Cover crop manipulation in northern California orchards and
vineyards: effects on arthropod communities. Biol Agric Hortic 3(1):1–24
Anderson MK (2013) Tending the wild: native American knowledge and the management of
California’s natural resources. University of California Press, Berkeley
Atkinson D, Watson CA (eds) (2019) The science beneath organic production. Wiley, New York
Bacon C, Mendez VE, Gliessman S, Goodman D, Fox J (eds) (2008) Confronting the coffee crisis:
fair trade, sustainable livelihoods and ecosystems in Mexico and Central America. MIT Press,
Cambridge
Balée W (1998) Historical ecology: premises and postulates. In: Balée W (ed) Advances in
historical ecology. Columbia University Press, New York, pp 13–29
Balzan MV (2017) Flowering banker plants for the delivery of multiple agroecosystem services.
Arthropod Plant Interact 11(6):743–754
Bardhan S, Jose S, Biswas S, Kabir K, Rogers W (2012) Homegarden agroforestry systems: an
intermediary for biodiversity conservation in Bangladesh. Agrofor Syst 85(1):29–34
Benayas JMR, Bullock JM (2012) Restoration of biodiversity and ecosystem services on agricul-
tural land. Ecosystems 15(6):883–899
Benton TG, Vickery JA, Wilson JD (2003) Farmland biodiversity: is habitat heterogeneity the key?
Trends Ecol Evol 18(4):182–188
Berkes F, Folke C, Gadgil M (1994) Traditional ecological knowledge, biodiversity, resilience and
sustainability. In: Berkes F, Folke C, Gadgil M (eds) Biodiversity conservation. Springer,
Dordrecht, pp 269–287
Bertsch A (2017) Indigenous successional agroforestry: integrating the old and new to address food
insecurity and deforestation. In: Montagnini F (ed) Integrating landscapes: agroforestry for
biodiversity conservation and food sovereignty, Advances in agroforestry 12. Springer, Cham,
pp 65–178
Blanchart E, Villenave C, Viallatoux A, Barthès B, Girardin C, Azontonde A, Feller C (2006)
Long-term effect of a legume cover crop (Mucuna pruriens var. utilis) on the communities of
soil macrofauna and nematofauna, under maize cultivation, in southern Benin. Eur J Soil Biol
42:S136–S144
Braun A, Duveskog D (2011) The farmer field school approach – history, global assessment and
success stories, Background paper for the IFAD rural poverty report. International Fund for
Agricultural Development, Rome
Brush SB (ed) (2000) Genes in the field: on-farm conservation of crop diversity. Boca Raton,
International Development Research Centre and International Plant Genetic Resources Institute/
Lewis Publishers
84 B. Levin

Brussaard L, De Ruiter PC, Brown GG (2007) Soil biodiversity for agricultural sustainability. Agric
Ecosyst Environ 121(3):233–244
Burbank L (1915) Luther Burbank: his methods and discoveries and their practical application, vol
12. Luther Burbank Press, New York and London
Cabeza M, Moilanen A (2001) Design of reserve networks and the persistence of biodiversity.
Trends Ecol Evol 16(5):242–248
Cárcamo HA, Spence JR (1994) Crop type effects on the activity and distribution of ground beetles
(Coleoptera: Carabidae). Environ Entomol 23(3):684–692
Chabay I, Frick M, Helgeson J (eds) (2015) Land restoration: reclaiming landscapes for a sustain-
able future. Academic, New York
Chateil C, Goldringer I, Tarallo L, Kerbiriou C, Le Viol I, Ponge JF, Salmon S, Gachet S, Porcher E
(2013) Crop genetic diversity benefits farmland biodiversity in cultivated fields. Agric Ecosyst
Environ 171:25–32
Chen XD, Dunfield KE, Fraser TD, Wakeli SA, Richardson AE, Condron LM (2020) Soil
biodiversity and biogeochemical function in managed ecosystems. Soil Res 58(1):1–20
Ching LL (2018) Agroecology for sustainable food systems, Environment and development series,
19. Third World Network, Penang
Coleman D, Reid C, Cole C (1983) Biological strategies of nutrient cycling in soil systems. Adv
Ecol Res 13:1–55
Corrado C, Elena T, Giancarlo R, Stefano C (2019) The role of agrobiodiversity in sustainable food
systems design and management. In: Nandwani D (ed) Genetic diversity in horticultural plants.
Springer, Cham, pp 245–271
Crist E, Mora C, Engelman R (2017) The interaction of human population, food production, and
biodiversity protection. Science 356(6335):260–264
De La Cruz R, Suárez S, Ferguson JE (1994) The contribution of Arachis pintoi as a ground cover in
some farming systems of Tropical America. CIAT Publication, Cali
Dominati EJ, Maseyk FJ, Mackay AD, Rendel JM (2019) Farming in a changing environment:
increasing biodiversity on farm for the supply of multiple ecosystem services. Sci Total Environ
662:703–713
Doré T, Makowski D, Malézieux E, Munier-Jolain N, Tchamitchian M, Tittonell P (2011) Facing
up to the paradigm of ecological intensification in agronomy: revisiting methods, concepts and
knowledge. Eur J Agron 34(4):197–210
Duchemin E, Wegmuller F, Legault AM (2008) Urban agriculture: multi-dimensional tools for
social development in poor neighbourhoods. Field Actions Sci Rep 2(1):1–8. [Link]
5194/facts-2-1-2009
Duru M, Therond O, Martin G, Martin-Clouaire R, Magne MA, Justes E, Journet EP, Aubertot JN,
Savary S, Bergez JE, Sarthou JP (2015) How to implement biodiversity-based agriculture to
enhance ecosystem services: a review. Agron Sustain Dev 35(4):1259–1281
Enri SR, Probo M, Farruggia A, Lanore L, Blanchetete A, Dumont B (2017) A biodiversity-friendly
rotational grazing system enhancing flower-visiting insect assemblages while maintaining
animal and grassland productivity. Agric Ecosyst Environ 241:1–10
Fair-Child D (1939) The world was my garden: travels of a plant explorer. Soil Sci 48(4):356
Fearnside P (2017) Business as usual: a resurgence of deforestation in the Brazilian Amazon. Yale
Environ 360:1–6
Finch CU, Sharp WC (1976) Cover crops in California orchards and vineyards. US Department of
Agriculture, Washington, DC
Gabriel S (2018) Silvopasture: a guide to managing grazing animals, forage crops, and trees in a
temperate farm ecosystem. Chelsea Green Publishing, White River Junction
Gaudin AC, Tolhurst TN, Ker AP, Janovicek K, Tortora C, Martin RC, Deen W (2015) Increasing
crop diversity mitigates weather variations and improves yield stability. PLoS One 10(2):
e0113261
Gibbons LV (2020) Regenerative—the new sustainable? Sustainability 12(13):5483
3 Regenerative Agriculture as Biodiversity Islands 85

Giller KE, Beare MH, Lavelle P, Izac AM, Swift MJ (1997) Agricultural intensification, soil
biodiversity and agroecosystem function. Appl Soil Ecol 6(1):3–16
Gliessman S (2016) Transforming food systems with agroecology. Agroecol Sustain Food Syst
40(3):187–189. [Link]
Gray M (2013) Labor and the locavore: the making of a comprehensive food ethic. University of
California Press, Berkeley
Gustafson JP, Raven PH, Ehrlich PR (2020) Population, agriculture, and biodiversity: problems and
prospects. University of Missouri Press, Columbia
Henderson K, Loreau M (2019) An ecological theory of changing human population dynamics.
People Nat 1(1):31–43
Holland JM (2019) Contribution of hedgerows to biological control. In: Dover JW (ed) The ecology
of hedgerows and field margins. Routledge, Abingdon, pp 123–146
Huang W, Luukkanen O, Johanson S, Kaarakka V, Räisänen S, Vihemäki H (2002) Agroforestry
for biodiversity conservation of nature reserves: functional group identification and analysis.
Agrofor Syst 55(1):65–72
Jackson LE, Pascual U, Brussaard L, de Ruiter P, Bawa KS (2007) Biodiversity in agricultural
landscapes: investing without losing interest. Agric Ecosyst Environ 121(3):193–195
Jordan CF (1998) Working with nature: resource management for sustainability. Taylor & Francis,
Amsterdam
Jørgensen LN, Kudsk P (2006) Twenty years’ experience with reduced agrochemical inputs: effect
on farm economics, water quality, biodiversity and environment. In: Home-Grown Cereals
Authority (HGCA) agronomist conference, Grantham, UK
Jose S (2009) Agroforestry for ecosystem services and environmental benefits: an overview.
Agrofor Syst 76(1):1–10
Jose S (2012) Agroforestry for conserving and enhancing biodiversity. Agrofor Syst 85(1):1–8
Jose S, Walter D, Kumar BM (2019) Ecological considerations in sustainable silvopasture design
and management. Agrofor Syst 93(1):317–331
Kidd LR, Garrard GE, Bekessy SA, Mills M, Camilleri AR, Fidler F, Fielding KS, Gordon A,
Gregg EA, Kusmanoff AM, Louis W (2019) Messaging matters: a systematic review of the
conservation messaging literature. Biol Conserv 236:92–99
Kimbrell A (ed) (2002) The fatal harvest reader: the tragedy of industrial agriculture. Island Press,
Washington, DC
Klasen S, Meyer KM, Dislich C, Euler M, Faust H, Gatto M, Hettig E, Melati DN, Jaya INS,
Otten F, Pérez-Cruzado C (2016) Economic and ecological trade-offs of agricultural speciali-
zation at different spatial scales. Ecol Econ 122:111–120
Lacombe S, Bradley RL, Hamel C, Beaulieu C (2009) Do tree-based intercropping systems increase
the diversity and stability of soil microbial communities? Agric Ecosyst Environ 131(1–2):
25–31
Landis DA, Wratten SD, Gurr GM (2000) Habitat management to conserve natural enemies of
arthropod pests in agriculture. Annu Rev Entomol 45(1):175–201
Lankoski J (2016) Alternative payment approaches for biodiversity conservation in agriculture.
[Link]
Leakey RR (1999) Agroforestry for biodiversity in farming systems. In: Collins WW, Calvin OQ
(eds) Biodiversity in agroecosystems. Lewis Publishers, New York, pp 127–145
Lehmann J, Bossio DA, Kögel-Knabner I, Rillig MC (2020) The concept and future prospects of
soil health. Nat Rev Earth Environ. [Link]
Lichtfouse E (ed) (2018) Sustainable agriculture reviews 31: biocontrol, vol 31. Springer,
New York
Liere H, Jha S, Philpott SM (2017) Intersection between biodiversity conservation, agroecology,
and ecosystem services. Agroecol Sustain Food Syst 41(7):723–760
Long R, Corbett A, Lamb C, Reberg-Horton C, Chandler J, Stimmann M (1998) Beneficial insects
move from flowering plants to nearby crops. Calif Agric 52(5):23–26
86 B. Levin

Lovell ST (2010) Multifunctional urban agriculture for sustainable land use planning in the United
States. Sustainability 2(8):2499–2522
Mandal A, Sarkar B, Mandal S, Vithanage M, Patra AK, Manna MC (2020) Impact of agrochem-
icals on soil health. In: Prasad MNV (ed) Agrochemicals detection, treatment and remediation.
Butterworth-Heinemann, Oxford, pp 161–187
Massy C (2017) Call of the reed warbler: a new agriculture—a new earth. University of Queensland
Press, St. Lucia, p 569
McAdam JH, McEvoy PM (2009) The potential for silvopastoralism to enhance biodiversity on
grassland farms in Ireland. In: McAdam JH, McEvoy PM (eds) Agroforestry in Europe.
Springer, Dordrecht, pp 343–356
McDermott ME, Rodewald AD (2014) Conservation value of silvopastures to Neotropical migrants
in Andean forest flocks. Biol Conserv 175:140–147
McLaughlin A, Mineau P (1995) The impact of agricultural practices on biodiversity. Agric Ecosyst
Environ 55(3):201–212
McNeely JA, Schroth G (2006) Agroforestry and biodiversity conservation–traditional practices,
present dynamics, and lessons for the future. Biodivers Conserv 15(2):549–554
Meena RS, Kumar S, Datta R, Lal R, Vijayakumar V, Brtnicky M, Sharma MP, Yadav GS, Jhariya
MK, Jangir CK, Pathan SI (2020) Impact of agrochemicals on soil microbiota and management:
a review. Land 9(2):34
Mendenhall CD, Karp DS, Meyer CF, Hadly EA, Daily GC (2014) Predicting biodiversity change
and averting collapse in agricultural landscapes. Nature 509(7499):213–217
Miller TP, Rebek EJ, Schnelle MA (2017) Banker plants for control of greenhouse pests. Oklahoma
Cooperative Extension Service, Oklahoma State University, Stillwater. [Link]
[Link]/
Mills J, Ingram J, Dibari C, Merante P, Karaczun Z, Molnar A, Sánchez B, Iglesias A, Ghaley BB
(2020) Barriers to and opportunities for the uptake of soil carbon management practices in
European sustainable agricultural production. Agroecol Sustain Food Syst 44(9):1185–1211
Montagnini F (2020) The contribution of agroforestry to restoration and conservation: biodiversity
islands in degraded landscapes. In: Dagar JC, Gupta SR, Teketay D (eds) Agroforestry for
degraded landscapes: recent advances and emerging challenges. Springer, New York, pp
445–479. [Link]
Montagnini F, del Fierro S (2022) Functions of agroforestry systems as biodiversity islands in
productive landscapes. In: Montagnini F (ed) Biodiversity Islands: strategies for conservation in
human-dominated environments. Topics in Biodiversity and Conservation, Springer, Cham,
pp 89–116
Montagnini F, Metzel R (2017) The contribution of agroforestry to sustainable development goal 2:
end hunger, achieve food security and improved nutrition, and promote sustainable
agriculture. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity conser-
vation and food sovereignty, Advances in agroforestry 12. Springer, Cham, pp 11–45
Montagnini F, Eibl B, Grance L, Maiocco D, Nozzi D (1997) Enrichment planting in overexploited
subtropical forests of the Paranaense region of Misiones. Argent For Ecol Manage 99(1–2):
237–246
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity Islands: strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity Islands: strategies
for conservation in human-dominated environments, Topics in Biodiversity and Conservation.
Springer, Cham, pp 1–35
Montgomery D (2018) Growing a revolution: bringing our soil back to life. W.W. Norton and
Company Inc., New York
Moss C, Lukac M, Harris F, Outhwaite CL, Scheelbeek PF, Green R, Berstein FM, Dangour AD
(2020) The effects of crop diversity and crop type on biological diversity in agricultural
landscapes: a systematic review protocol [version 2; peer review: 1 approved, 2 approved
with reservations]. Wellcome Open Res 4(101). [Link]
15343.2
3 Regenerative Agriculture as Biodiversity Islands 87

Mouysset L, Doyen L, Pereau JC, Jiguet F (2015) Benefits and costs of biodiversity in agricultural
public policies. Eur Rev Agric Econ 42(1):51–76
Nabhan GP (2014) Food security, biodiversity and human health: ethnobiology as a predictive
science. J Ethnobiol 34(1):7–11
Nabhan GP (2016) Enduring seeds: native American agriculture and wild plant conservation.
University of Arizona Press, Tucson
Nair PK (2017) Managed multi-strata tree+ crop systems: an agroecological marvel. Front Environ
Sci. [Link]
Navarro LM, Pereira HM (eds) (2015) Rewilding European landscapes. Springer, Cham
Nazarea VD (2005) Heirloom seeds and their keepers: marginality and memory in the conservation
of biological diversity. University of Arizona Press, Tucson
Newbold T, Hudson LN, Arnell AP, Contu S, De Palma A, Ferrier S, Hill SL, Hoskins AJ,
Lysenko I, Phillips HR, Burton VJ (2016) Has land use pushed terrestrial biodiversity beyond
the planetary boundary? A global assessment. Science 353(6296):288–291
Orefice JN, Carroll J (2017) Silvopasture—it’s not a load of manure: differentiating between
silvopasture and wooded livestock paddocks in the northeastern United States. J For 115(1):
71–72
Orefice J, Smith RG, Carroll J, Asbjornsen H, Howard T (2019) Forage productivity and profit-
ability in newly-established open pasture, silvopasture, and thinned forest production systems.
Agrofor Syst 93:51–65
Parker JE, Crowder DW, Eigenbrode SD, Snyder WE (2016) Trap crop diversity enhances crop
yield. Agric Ecosyst Environ 232:254–262
Pascual U, Perrings C (2007) Developing incentives and economic mechanisms for in situ biodi-
versity conservation in agricultural landscapes. Agric Ecosyst Environ 121(3):256–268
Perfecto I, Vandermeer JH, Wright AL (2009) Nature’s matrix: linking agriculture, conservation
and food sovereignty. Routledge, London
Reddy P (2017) Intercropping. In: Reddy P (ed) Agro-ecological approaches to pest management
for sustainable agriculture. Springer, Singapore, pp 109–131
Redlich S, Martin EA, Steffan-Dewenter I (2018) Landscape-level crop diversity benefits biological
pest control. J Appl Ecol 55(5):2419–2428
Redondo-Brenes A, Montagnini F (2010) Contribution of homegardens, agrosilvopastoral systems,
and other human dominated land-use types to the avian diversity of a biological corridor in
Costa Rica. Adv Environ Res 2(2):111–148
Reij C, Garrity D (2016) Scaling up farmer-managed natural regeneration in Africa to restore
degraded landscapes. Biotropica 48(6):834–843
Rejesus RM, Jones MS (2020) Perspective: enhancing economic evaluations and impacts of
integrated pest management Farmer Field Schools (IPM-FFS) in low-income countries. Pest
Manag Sci 76(11):3527–3536
Rhodes CJ (2017) The imperative for regenerative agriculture. Sci Prog 100(1):80–129
Rocha P, Niella F, Keller H, Montagnini F, Metzel R, Eibl B, Kornel J, Romero F, López L,
Araujo J, Barquinero J (2017) Ecological indigenous (EIK) and scientific (ESK) knowledge
integration as a tool for sustainable development in indigenous communities. Experience in
Misiones, Argentina. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity
conservation and food sovereignty. Springer, Cham, pp 235–260
Salatin J (2010) Sheer ecstasy of being a lunatic farmer. Chelsea Green Publishing, White River
Junction
Salzman T (2018) The farmworker health landscape: barriers to improving health and safety in US
agriculture. Doctoral dissertation, Tufts University, Medford/Somerville
Sánchez-Bayo F, Wyckhuys KA (2019) Worldwide decline of the entomofauna: a review of its
drivers. Biol Conserv 232:8–27
Savory A, Butterfield J (1998) Holistic management: a new framework for decision making. Island
press, Washington, DC
88 B. Levin

Saxton DI (2021) The Devil’s fruit: farmworkers, health, and environmental justice. Rutgers
University Press, New Brunswick
Schreefel L, Schulte RPO, de Boer IJM, Schrijver AP, van Zanten HHE (2020) Regenerative
agriculture – the soil is the base. Glob Food Sec 26:100404. [Link]
100404
Schroth G, Lehmann J, Rodrigues MRL, Barros E, Macêdo JL (2001) Plant-soil interactions in
multistrata agroforestry in the humid tropics. Agrofor Syst 53(2):85–102
Schroth G, Izac AMN, Vasconcelos HL, Gascon C, da Fonseca GA, Harvey CA (eds) (2004)
Agroforestry and biodiversity conservation in tropical landscapes. Island Press, New York
Shepard M (2013) Restoration agriculture. Acres U.S.A., Austin
Shiva V (2000) Tomorrow’s biodiversity. Thames & Hudson, London
Shiva V (2020) Reclaiming the commons: biodiversity, indigenous knowledge, and the rights of
mother earth. Synergetic Press, Santa Fe
Smith RG, Gross KL, Robertson GP (2008) Effects of crop diversity on agroecosystem function:
crop yield response. Ecosystems 11(3):355–366
Socci P, Errico A, Castelli G, Penna D, Preti F (2019) Terracing: from agriculture to multiple
ecosystem services. In: Oxford research encyclopedia of environmental science. Oxford Uni-
versity Press, Oxford
Soloviev ER, Landua G (2016) Levels of regenerative agriculture. [Link]
wp-content/uploads/2017/03/Levels-of-Regenerative-Agriculture-1. Accessed Online March
22 2021
Swanson T, Goeschl T (2000) Optimal genetic resource conservation: in situ and ex situ. In: Genes
in the field: on-farm conservation of crop diversity. International Development Research Centre
and International Plant Genetic Resources Institute/Lewis Publishers, Boca Raton, pp 165–191
Tavares PD, Uzêda MC, Pires ADS (2019) Biodiversity conservation in agricultural landscapes: the
importance of the matrix. Floresta e Ambiente. [Link]
Tilman D (2020) Resource competition and community structure (vol. 17). Princeton University
Press. Princeton, New Jersey
Toensmeier E (2016) The carbon farming solution: a global toolkit of perennial crops and
regenerative agriculture practices for climate change mitigation and food security. Chelsea
Green Publishing, White River Junction
Toensmeier E (2022) Paradise lot: a temperate-climate urban agroforestry Biodiversity Island. In:
Montagnini F (ed) Biodiversity Islands: strategies for conservation in human-dominated envi-
ronments. Topics in Biodiversity and Conservation. Springer, Cham, pp 439–459
Tree I (2019) Wilding: returning nature to our farm. New York Review of Books, London
Udawatta R, Rankoth L, Jose S (2019) Agroforestry and biodiversity. Sustainability 11(10):2879
Wagg C, Bender SF, Widmer F, Van Der Heijden MG (2014) Soil biodiversity and soil community
composition determine ecosystem multifunctionality. Proc Nat Acad Sci 111(14):5266–5270
Wagner DL (2020) Insect declines in the anthropocene. Annu Rev Entomol 65:457–480
White C (2020) Why regenerative agriculture? Am J Econ Soc 79(3):799–812
White R, Charlebois E, Martin S (2018) Seed “rematriation”. Panel discussion at the intertribal food
sovereignty summit. Pequot Museum and Research Center, 21–23 August 2018, Ledyard,
CT, USA
Wintle BA, Kujala H, Whitehead A, Cameron A, Veloz S, Kukkala A, Moilanen A, Gordon A,
Lentini PE, Cadenhead NC, Bekessy SA (2019) Global synthesis of conservation studies reveals
the importance of small habitat patches for biodiversity. Proc Nat Acad Sci 116(3):909–914
Yeomans PA (1958) The challenge of landscape: the development and practice of Keyline. Keyline
Pub. Pty, Sydney
Young KJ (2017) Mimicking nature: a review of successional agroforestry systems as an analogue
to natural regeneration of secondary forest stands. In: Montagnini F (ed) Integrating landscapes:
agroforestry for biodiversity conservation and food sovereignty. Springer, Cham, pp 179–209
Zhao ZH, Hui C, Reddy GV, Ouyang F, Men XY, Ge F (2019) Plant species richness controls
arthropod food web: evidence from an experimental model system. Ann Entomol Soc Am
112(1):27–32
Chapter 4
Functions of Agroforestry Systems
as Biodiversity Islands in Productive
Landscapes

Florencia Montagnini and Sara del Fierro

Abstract Given their ability to harmonize productivity with environmental func-

tions, agroforestry systems (AFS) are an important strategy for conservation within
human managed landscapes. AFS are heterogeneous in their design, management,
and species composition, with consequences for their restoration, conservation, and
productivity functions. AFS can function as biodiversity islands or can be incorpo-
rated into existing biodiversity islands as buffer zones because they can be integrated
into already productive landscapes. This chapter provides an overview of the various
ecological, social, and economic benefits of the main types of AFS systems and their
applications as and within biodiversity islands. It also discusses the use of incentives
to support and promote AFS in order to safeguard the contributions they provide to
landscape biodiversity and rural communities.

Keywords Buffer zones · Certification · Connectivity · Markets · Organic farming ·

Payments for ecosystem services (PES)

4.1 Introduction

Sustainable agricultural management that seeks to balance ecosystem productivity

and conservation can play an important role in mitigating or reversing detrimental
human induced effects on landscapes. Agroforestry systems (AFS), which combine
trees and crops or pasture on the same land, can increase productivity in the short and
long term, while also supporting biodiversity and providing social and economic
benefits for farmers (Jose 2009, 2012; Montagnini et al. 2015; Montagnini and
Metzel 2017; Montagnini 2020; Udawatta et al. 2019).

F. Montagnini (*)
School of the Environment, The Forest School, Yale University, New Haven, CT, USA
e-mail: fl[Link]@[Link]
S. del Fierro
United States Department of Agriculture (USDA), Washington, DC, USA

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 89

F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
90 F. Montagnini and S. del Fierro

A growing body of evidence supports the overall assertion that floral, faunal, and
soil microbial diversity are significantly greater in AFS as compared to monocul-
tures, adjacent crop lands, and even some forests (Bhagwat et al. 2008; Udawatta
et al. 2019). Among the soil organisms, arbuscular mycorrhizal fungi, bacteria, and
enzyme activities have been found to be significantly greater in AF than in conven-
tional agriculture (Udawatta et al. 2019). Agroforestry can also create spatially
concentrated high biodiversity near trees due to favorable soil-plant-water-microcli-
mate conditions. The greater biodiversity in AFS has been attributed to their more
favorable microclimate and soil conditions, and to their heterogeneity in comparison
with monocrops.
AFS are used in a variety of contexts and systems, serving various needs and
functions and incorporating different crops and local practices to meet those needs.
The specific characteristics of each AFS vary strongly according to system design,
objectives and species involved, with strong differences both within and between
ecological regions. Their locations, and within them the context-specific economic,
social, and political factors, can influence the practices used, along with their
productivity, sustainability, and environmental services (Montagnini and Metzel
2017; Montagnini 2020). Therefore, functions of AFS, including their role in
biodiversity conservation and restoration, can vary widely, however, they tend to
enhance conservation of biodiversity when compared to monoculture systems.
As the agricultural frontier continues to advance and landscapes become further
fragmented, driven by the need to supply food and resources to a growing (in number
and area) human population, measures to preserve biodiversity are urgently needed
(Montagnini and Berg 2019). One such measure is the promotion of biodiversity
islands, protected and managed areas of high biological diversity, within otherwise
degraded human-dominated landscapes, where plants and animals can thrive without
major degenerative interference from human activity (Montagnini and Berg 2019;
Montagnini et al. 2022). Since AFS are often important components among land
uses in buffer zones of protected areas, they work well as parts of biodiversity
islands, creating a smoother transition to areas of greater human impact. AFS can
also serve as biodiversity islands themselves within cultivated landscapes, consisting
of higher species compositions and providing greater species habitat than otherwise
conventional systems.
This chapter provides an overview of AFS systems, drawing from studies around
the world. Since biodiversity in AFS varies strongly according to system character-
istics, the following section discusses features and biodiversity contributions of the
most frequent AFS: multistrata systems including homegardens and successional
AFS, perennial crops under shade, silvopastoral systems, and living fences and
windbreaks. For each type, the role of local and indigenous knowledge and several
of their ecological, social, and economic benefits are highlighted. The chapter then
offers ways to consider AFS as components of biodiversity islands in human-
dominated landscapes. The chapter ends by discussing incentive systems that can
be incorporated to promote and sustain AFS in order to safeguard the contributions
they provide to both biodiversity and rural communities.
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 91

4.2 Predominant Types of Agroforestry Systems and Their

Benefits

The predominant types of AFS used globally are multistrata AFS, perennial crops
under shade, silvopastoral systems, living fences, windbreaks and riparian buffers.
The main features of these major types, as well as of sub-types within them, along
with their ecological, economic, social, and cultural benefits, are summarized in
Table 4.1.

4.2.1 Multistrata Agroforestry Systems

[Link] Homegardens

Homegardens, also called ‘forest gardens,’ are defined as intimate, multistory

combinations of various trees and crops, sometimes in association with domestic
animals, around homesteads (Nair and Kumar 2006). In homegardens, plants are
generally categorized into three main groups: cultivated, protected or spared. Culti-
vated plants are those that are sown or planted by the owner, while protected plants
are transplanted or spontaneous plants that are encouraged by the farmer. Spared
plants are those that spontaneously grow in the garden and are not removed
(Blanckaert et al. 2004). Each design and management strategy can contribute to
the presence and preservation of species of interest to the farmer and to landscape
biodiversity (Fig. 4.1).
Food plants, including crops and trees, are often the most abundant species found
in homegardens. Homegardens are also valuable sources of income via cash crops
(Nair and Kumar 2006). Species complexity in homegardens is a result of deliberate
and meticulous selection and management by farmers to grow and steward the
products they consider important for their subsistence and livelihood. Different
factors, such as remoteness from urban centers, management, and modernization,
influence biodiversity in homegardens throughout the world, as seen from examples
from Kerala, India (Peyre et al. 2006), southern Ethiopia (Abete et al. 2006), the
Peruvian Amazon (Wezel and Ohl 2006), and Meso America (Kumar and Nair
2006; Montagnini 2006).
Homegardens span a wide range of climatic and geographic conditions, including
areas of typically lower plant diversity. Table 4.1 shows cases from Puebla, Mexico,
eastern Cuba, and the Mexican Plateau where homegardens in particularly semi-arid
contexts provide culturally important and sustenance species in areas that would
otherwise have lower biodiversity and fewer economic options.
Homegardens also serve as sites for domestication of useful species in several
regions. Cases from Pará, Brazil, Copán, Honduras, and the Tehuacán-Cuicatlán
valley in central Mexico demonstrate the use of homegardens in these regions to
serve as “banks” for ancient and potential future crop species. They also serve as
92 F. Montagnini and S. del Fierro

Table 4.1 Summary of types of AFS, their ecological and economic/cultural/social benefits, and
case studies cited in the text.
Type of Economic/cultural/social
system Ecological benefits benefits Case studies cited
Homegardens Increased plant diversity Production of culturally Puebla, Mexico
in arid to semi-arid areas important food crops (Blanckaert et al. 2004)
where low diversity is (chilli (Capsicum spp.),
expected tomato (Lycopersicum
esculentum), and several
Cactaceae and
Solanaceae species) in
arid to semi-arid areas
Reduced climatological Production of medicinal Mexican plateau
impacts; increased soil and food plants (Terrones Rincón et al.
fertility 2011)
Increased species diver- Production of fruit trees Eastern Cuba (Wezel
sity, higher ecosystem in semi-arid area where and Bender 2003)
productivity other options are more
difficult and less viable
Conservation of crop Serve as crop species Pará, Brazil (Callo-
species “banks”, validation facil- Concha and Denich
ities for farmer decision- 2011)
making, crop improve-
ment and propagation
Serve as gene bank, pre- Serve as research fields El Camalote, Honduras
serving species not found for new varieties and (House and Ochoa
elsewhere, including edi- cultivars and gene banks 1998)
ble vegetable species and for edible vegetable and
fruits, such as the chayo fruit species
(Cnidoscolus
chaymansa)
Increased species diver- Larger, better quality Tehuacán-Cuicatlán,
sity and conservation fruits, such as highly Mexico (González
consumed fruit, Soberanis and Casas
tempesquistle 2004)
(Sideroxylon palmei)
Serve as refuge for wild- Production of food Petén, Guatemala
life during disturbances sources for wildlife; food (Griffith 2000)
like fire and for critical and cash income for
food source post-fire farmers
Successional Improved soil fertility; Preservation of indige- Chiapas, Mexico
AFS bees attracted for nous system using dis- (Diemont et al. 2006;
pollination tinct phases: Milpa Diemont et al. 2011)
(cultivated maize field),
arbusto (shrub with
planting), and acahual
(fallow shrub), then
return to Selva Alta (high
forest); readily harvest-
able crops; honey bees
(continued)
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 93

Table 4.1 (continued)

Type of Economic/cultural/social
system Ecological benefits benefits Case studies cited
Managed gardens mirror Production of fruit trees Dayak people of Indo-
natural succession stages nesia (Peters 2018)
and functions
Cultivated crops serve Increase in agricultural NE Brazil (Schulz
ecological functions of production; reduced risk 2011).
natural plants; restoration of drought-related har-
of degraded lands vest loss
Attract wildlife for seed Attract wildlife for Worldwide (Bertsch
dispersal, increased soil hunting 2017)
fertility
Perennial Display high biodiversity Provision of coffee crop, Mexico (Davidson
crops under and high structural com- product diversification, 2005)
shade: Coffee plexity; provide habitats; decreased economic
serve as perches/nesting risks, decreased need of
sites; provide food external inputs
sources; improve local
microclimates; increase
sustainability
Seed deposition; serve as Provision of coffee crop, Mexico (Moguel and
buffer zones and biologi- product diversification, Toledo 1999)
cal corridors decreased economic
risks, decreased need of
external inputs
Increased butterfly spe- Provision of coffee crop, Chiapas, Mexico (Mas
cies richness product diversification, and Dietsch 2003)
decreased economic
risks, decreased need of
external inputs
Increased plant diversity, Competitive coffee yields Costa Rica (Rossi et al.
better soil cover, soil in intermediate intensity 2011).
conservation managed systems
Provide habitat for Provision of coffee crop, Neotropics (Somarriba
migrating birds higher sustainability, et al. 2004).
product diversification,
decreased need of exter-
nal inputs
Increased bird diversity Provision of coffee crop, Colombia, worldwide
higher sustainability, (Chait 2015; several
product diversification, others).
decreased need of exter-
nal inputs
Increased diversity of Provision of coffee crop, Neotropics, worldwide
arthropods, mammals, higher sustainability, (Mas and Dietsch 2003;
amphibians and reptiles product diversification, Teodoro et al. 2011;
decreased need of exter- Rossi et al. 2011; Chait
nal inputs 2015).
(continued)
94 F. Montagnini and S. del Fierro

Table 4.1 (continued)

Type of Economic/cultural/social
system Ecological benefits benefits Case studies cited
Increased diversity of Increases in yield, weight Colombia, worldwide
pollinators of the bean, and quality (Chait 2015)
of the coffee
Perennial Conservation of Atlantic Provision of cacao crop, Cabruca systems in
crops under forest biodiversity higher sustainability, Bahia and Espírito
shade: Cacao product diversification, Santo in eastern Brazil
decreased need of exter- (Rolim and Chiarello
nal inputs 2004).
Maintained species rich- Increased yield, higher Indonesia (Clough et al.
ness of trees, fungi, sustainability, product 2011)
invertebrates, and diversification, decreased
vertebrates need of external inputs
Landscape restoration; Increased resilience of Cacao Alliance partici-
increased water infiltra- productive systems in the pants in El Salvador
tion capacity; soil and face of climate change; (Montagnini and
water conservation; greater food sovereignty Metzel 2017)
expanded areas with bio- and employment
logical significance; opportunities
enhanced biological
corridors
Increased plant diversity, Improved livelihoods via Indonesia (Roshetko
better soil cover, soil stronger resilience to et al. 2016)
conservation pests and diseases
Perennial Maintained bird species Provision of yerba mate Paraguay (Cockle et al.
crops under abundance (relative to crop 2005)
shade: Yerba adjacent forest reserve),
mate including globally threat-
ened species
SPS Increased bird fauna Provision of cattle prod- Cordoba, Colombia
habitat ucts, higher sustainabil- (Múnera et al. 2009)
ity, product
diversification, improved
cattle comfort and
productivity
Preservation of native Production of timber, Neotropics (Calle et al.
species; nitrogen fixation, edible fruits for cattle 2017)
soil conservation, and
natural biological control
of pests
Provide resources for Provision of cattle prod- Neotropics (Rivera
native wildlife; reduce ucts, higher sustainabil- et al. 2013; Montoya-
risk of local extinction ity, product Molina et al. 2016)
diversification, improved
cattle comfort and
productivity
(continued)
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 95

Table 4.1 (continued)

Type of Economic/cultural/social
system Ecological benefits benefits Case studies cited
Intensive SPS Greatest richness in bird Increased cattle produc- El Hatico, Colombia
species of land uses; tivity, higher sustainabil- (Fajardo et al. 2009).
three times as many bird ity, product
species as pasture sys- diversification, improved
tems without trees cattle comfort and
productivity
Increase in abundance Increased cattle produc- La Vieja, Colombia
and diversity of birds; tivity, higher sustainabil- (Calle et al. 2009).
general plant and animal ity, product
diversity; improved natu- diversification, improved
ral biological control; cattle comfort and
more rare/endangered productivity
species sightings
Living fences Promote biodiversity, Protect agricultural plots Neotropics (Harvey
increase connectivity and cattle; provide fuel- et al. 2005, 2008;
among forest patches, wood, fruits, fodder, and Francesconi et al.
serve as biological shade for cattle 2011a, b; Ibrahim et al.
corridors 2011)
Higher species richness Divide, separate, and Esparza, Costa Rica
of butterflies than in pas- protect agricultural plots (Tobar et al. 2007)
tures with high or low
tree densities
Preferred habitat and Divide, separate, and Esparza, Costa Rica
corridors for mantled protect agricultural plots (Rosales and Sáenz
howler monkeys 2007)
Windbreaks Maintain spatial organi- Serve as windbreaks to Chaco, Argentina
zation of landscape; con- protect crops, provide (Tamashiro 2018)
tribute to fauna species tree products
mobility, serve as part of
biological corridors

research sites for validating and testing new varieties and methods, and as venues for
developing products, such as fruits, that are larger or have better quality than in the
wild (Table 4.1).
Homegardens are human impacted areas due to their proximity to homes. This
makes them unique in that their biodiversity is often linked to human habitation—
more so than other agroforestry systems that are practiced in remote locations where
biodiversity can be more “wild” and less novel. Through the vast potential of species
diversity of food plants and the associated habitat generated from such species,
homegardens may harbour greater amounts of biological diversity than otherwise
conventional agricultural systems or degraded landscapes, allowing for
homegardens to function as biodiversity islands in the landscape.
96 F. Montagnini and S. del Fierro

Fig. 4.1 Homegarden in Embu district, Kenya, featuring coffee, manioc, plantain, sugar cane, and
a number of fruit, fuelwood, and timber species in the upper strata. The World Agroforestry Center
has been studying and promoting homegardens and other agroforestry systems since the 1990s in
collaboration with local organizations, based on ethnobotanical surveys and small farmer’s needs,
using local as well as exotic tree species for fodder, soil fertility and soil erosion control. ([Link]
[Link]/Units/Library/Books/Book%2006/html/14.5_kari_kefri_icraf_agrpfore.
htm?n¼135). Photo: F. Montagnini

[Link] Successional Agroforestry Systems

In restoration ecology, successional processes are often manipulated to restore

devastated landscapes (Hobbs et al. 2007). Successional AFS (SAFS) attempt to
replicate the spatial and temporal dynamics of forest succession, with species
assemblages that are planted, maintained, and modified over time to mirror the
successional stages of secondary forest development, for achieving both ecosystem
conservation or restoration as well as for production of subsistence and marketable
goods (Fig. 4.2). Successional agroforests can be similar to natural forest ecosystems
as they are characterized by high plant diversity and managed in accordance with the
natural succession of species (Peneireiro 1999; Micollis et al. 2016).
Such systems have been practiced by indigenous peoples for centuries. The
Lacandon Mayan of Chiapas, Mexico and the Kenyah Dayak of Kalimantan,
Indonesia have long managed homegardens as SAFS according to traditional indig-
enous practices (Table 4.1). In many indigenous SAFS, plant species are selected not
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 97

Fig. 4.2 In this system,

indigenous people from the
Napo Province in the
Ecuadorian Amazon
introduce useful species
such as fruit, medicinal, and
food species, which occupy
consecutive successional
stages, thereby restoring and
enriching the biodiversity of
the disturbed forests. This
management practice can be
considered a type of
successional agroforestry
system. Photo:
F. Montagnini

only to provide fruit and products, including honey, for personal consumption and
soil fertility, but also to attract wildlife, which deposit waste thus increasing soil
fertility, bring new seed sources, and can be hunted for food (Bertsch 2017).
Many of these SAFS practices from traditional, indigenous knowledge have been
adapted to modern agroecosystems (Schulz 2011; Young 2017). In northeastern
Brazil, Nicaragua, and Belize, farmers integrate and enhance natural succession
within their AFS, for instance by planting locally adapted edible plants with similar
functional characteristics as plants of the same successional level within the native
ecosystem, beginning with plants that augment organic material, and then slowly
integrating plants of higher successional levels. Highly degraded areas have been
regenerated with this method, resulting in an approximately four-fold increase of
agricultural production, while also reducing the risk of drought-related harvest loss
via crop diversification and the use of perennial plants (Schulz 2011).
98 F. Montagnini and S. del Fierro

4.2.2 Perennial Crops Under Shade in Agroforestry Systems

AFS of perennial crops under shade are broadly used throughout the tropics, with
cacao and coffee being the most frequent worldwide (Montagnini et al. 1992; Beer
et al. 1998; Chait 2015). The environmental value of shade trees is provided by their
forest-like structure (Perfecto et al. 2005). Shade trees also have social and economic
value in reducing the vulnerability of households to climatic stress, pest outbreaks,
falling prices and food insecurity (Tscharntke et al. 2011). Enriching the diversity of
natural shade trees, for instance by planting leguminous species, can also provide
additional positive impacts such as increased soil fertility (Montagnini et al. 1992;
Beer et al. 1998). Reducing pesticide spraying protects the functional
agrobiodiversity of the system, including organisms that provide biological control
against pests and diseases and pollinators that enhance cacao and coffee yield. From
a landscape perspective, natural forest maintained alongside agroforestry increases
the diversity of functionally important organisms (Tscharntke et al. 2011).
In AFS of perennial crops under shade, the amount of crop shading cover is a
proxy of agricultural intensification (Beer et al. 1998). Shade trees provide long-term
resistance and resilience in the presence of unmanageable pest pressure, vulnerabil-
ity to changing climate and difficulties in rejuvenating the perennial crops. Shade
removal, although it may increase short-term yield gains, may compromise this long-
term resilience.

[Link] Coffee

Coffee (Coffea arabica) is of paramount economic importance in more than 50 coun-

tries worldwide, and although it has potential to influence biodiversity conservation
over large areas, its cultivation can also be a cause of deforestation if done poorly
(Somarriba et al. 2004). Many regions where coffee is cultivated fall within areas
identified as mega-diversity sites (Somarriba et al. 2004). In some areas, the land-
scape has been so severely degraded that the only remaining tree cover is in coffee
plantations. For example, in El Salvador, the most densely populated country in
Latin America with less than 10% of its natural forests remaining, the vast majority
of remaining tree cover is associated with shade coffee (F. Montagnini, personal
observations 2002; Blackman et al. 2012).
Traditional, indigenous shaded coffee AFS harbor relatively high biological
diversity and provide high structural complexity (Moguel and Toledo 1999).
These systems illustrate valuable features of AFS: providing habitat for a variety
of species, serving as perches and nesting sites, providing food resources, and
improving local microclimates that are amenable for a wide variety of species
(Davidson 2005). Shaded coffee AFS can also serve as sites for seed deposition
and germination, act as buffer zones and biological corridors, and serve as a refuge
for wildlife from surrounding areas that have been deforested (Moguel and Toledo
1999).
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 99

Coffee AFS play an important role in providing habitat for–and increasing the
local diversity of–birds, arthropods, mammals, and, to a lesser extent, amphibians
and reptiles. In Chiapas, Mexico, Costa Rica, and elsewhere in the Neotropics,
several studies have found an inverse relationship between the diversity of certain
species and the intensity of the management of the coffee AFS (Table 4.1). In
experimental coffee AFS at CATIE,1 Costa Rica, intermediate management intensity
produced competitive coffee yields, and organically managed plots had high herba-
ceous diversity and were as productive as chemically managed plots, suggesting that
AFS can balance agricultural productivity while maintaining a significant number of
herbaceous species (Rossi et al. 2011).

[Link] Cacao

Cacao (Theobroma cacao) AFS can conserve natural resources and improve small
farmers’ livelihoods and self-sufficiency, by offering a varied production of food and
cash crops (Cerda et al. 2014; Gross et al. 2016). Studies from Bahia and Espírito
Santo, Brazil, and Indonesia, have demonstrated that in addition to the economic
benefit they provide, cacao AFS conserve local biodiversity. In Indonesia, the
species richness of trees, fungi, invertebrates, and vertebrates did not decrease
with increased cacao yield, indicating that moderate shade and adequate labor can
be combined with a complex habitat structure to provide both high biodiversity and
high yields (Clough et al. 2011).
As with coffee and other crops, cacao’s benefits to biodiversity greatly depend on
design and management of the AFS. Even though moderate shade levels rarely
reduce cacao or coffee yield, farmers in many parts of the world are converting
shaded cacao and coffee systems into unshaded monocultures to increase short-term
income (Tscharntke et al. 2011). However, benefiting from the long-term advantages
of shaded cacao agroforestry does not necessarily exclude intermediate levels of
intensification. For example, in Sulawesi, Indonesia, it has been shown that reducing
canopy cover from 80% to 40% can double the income of local farmers with only
minor changes in biodiversity and associated ecosystem services (Tscharntke et al.
2011). In another study in southern Cameroon of a project where increased use of
fungicides and the expansion of cultivated area aimed to reduce rural poverty,
overall plant diversity decreased only slightly with management intensification
(Gockowski et al. 2010). In another example in Sulawesi, Indonesia, the transfor-
mation of the conventional cacao cultivation systems to cacao AFS has improved
livelihoods for small farmers by increasing diversity in the cacao AFS, which
enhanced yields that were previously poor due to pests and diseases (Roshetko
et al. 2016).

1
Centro Agronómico Tropical de Investigación y Enseñanza, Tropical Agriculture Research and
Higher Education Center, Turrialba, Costa Rica.
100 F. Montagnini and S. del Fierro

Fig. 4.3 Agroforestry system of organic yerba mate (Ilex paraguariensis) and timber trees in
Misiones, Argentina, subtropical Atlantic Forest region. Organic yerba mate, grown generally under
several species of native trees and shrubs of timber, fruit or other uses, can get price surpluses,
which has led to an increased interest in organic farming and in yerba mate cultivation under shade
in recent years (Montagnini et al. 2011; Eibl et al. 2017). Photo: F. Montagnini

[Link] Yerba Mate

Yerba mate, Ilex paraguariensis, is a native tree from South America whose leaves
are used to prepare an infusion or tea of popular local consumption, with a market
expanding internationally due to its nutritious and energizing properties (Montagnini
et al. 2011; Eibl et al. 2015, 2017). Yerba mate trees are usually grown in mono-
cultures with conventional management, resulting in decreased plant productivity
and soil erosion in the long term. Since the yerba mate tree grows naturally in
subtropical forest and is shade tolerant, however, it is adequate for growing under the
canopy of other tree species in AFS (Fig. 4.3).
Yerba mate grows naturally in the Atlantic forest of southeastern Brazil, north-
eastern Argentina, and eastern Paraguay, a region that is one of the world’s biodi-
versity hotspots, with about 1–8% of all species worldwide, and high rates of plant,
insect and mammal endemism (Myers et al. 2000; Calmon et al. 2011). The Atlantic
forest is one of the most highly impacted rainforest areas in the world, where over
five centuries of deforestation resulted in a ~ 84% loss of area, with deforestation
continuing currently at a rate of 20,000 ha per year (Ribeiro et al. 2009; SOS Mata
Atlântica and INPE 2014). Agriculture, cattle-ranching, and industry have replaced
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 101

much of the Atlantic forest, and its diverse fauna is threatened by high grade logging,
hunting, habitat loss, and habitat fragmentation (Cockle et al. 2005; Brewer 2011).
Yerba mate is grown by small or medium to large farmers as family businesses,
farmers’ cooperatives, or large-scale enterprises, both for local consumption and for
export. Yerba mate cultivation thus expands a whole range of systems, from
extensive monocultures, to AFS with 1–2 tree species for shade, to more complex,
multistrata systems in the case of most of the organic yerba mate AFS (Ilany et al.
2010; Montagnini et al. 2011; Eibl et al. 2015, 2017) (Fig. 4.3).
In a study in Paraguay, bird species’ presence and abundance were compared
between a forest reserve and an adjacent plantation of shade-grown yerba mate,
where some of the forest understory and trees were removed and yerba mate was
planted below the tree canopy. Of the 145 species that were regularly recorded in the
forest, 66%, including five globally threatened species, were also recorded in the
yerba mate AFS. Within the yerba mate AFS, higher tree density did not lead to a
greater abundance of birds. Yerba mate AFS under native trees could therefore be
used to rehabilitate cleared land and allow recolonization by Atlantic forest bird
species (Cockle et al. 2005).
In another study in the ‘San Rafael’ Reserve and its buffer area (Itapua Depart-
ment, Paraguay), researchers analyzed the potential benefits of forests with shade-
grown yerba mate for birds, amphibian and reptiles, comparing species richness and
composition between three environments: forest with shade-grown yerba mate,
forest edge and monoculture crop plantations. Their results suggest that forests
with yerba mate plantations maintain high bird species richness, with its species
composition differing significantly from edges and croplands (Cabral et al. 2020).
Yerba mate AFS can also boost the biodiversity conservation capabilities of
forest fragments through increasing connectivity. Currently, with the expansion of
the yerba mate cultivation area and an increased interest in growing it in AFS due to
potential price surpluses, it would be interesting to further ascertain the role of yerba
mate AFS on restoring and conserving biodiversity, as compared to other cultivating
systems (Montagnini et al. 2011; Montagnini 2020).

4.2.3 Silvopastoral Systems

Silvopastoral systems (SPS) can incorporate more sustainable and biodiversity

friendly approaches into livestock rearing operations. Silvopastoral systems, which
involve the combination of trees with pastures and livestock, are classified by the
functions and configuration or structure of trees within the system. SPS may include
dispersed trees in pastures, live fences in pastures, fodder banks, tree alley pasture
systems, and pastures with windbreaks (Pezo and Ibrahim 1999). Because they are
more structurally complex than grass monocultures, SPS have important benefits for
biodiversity. Many types of SPS (e.g., high density trees in pastures and live multi-
strata fences) have levels of species richness comparable to those of early secondary
forest, and networks of live fences in pastures are important for landscape
102 F. Montagnini and S. del Fierro

Fig. 4.4 Silvopastoral systems using planted hybrid pines (Pinus taeda x Pinus caribaea), Braford
cattle (hybrids of Brahman x Hereford), with Brachiaria bryzantha grass in Misiones, Argentina.
This highly technified type of system based on exotic species of animals, trees and grasses is less
biodiverse than more traditional systems based on local species of plants and animals, however, it is
more structurally complex and therefore more biodiverse than grass monocultures. Photo:
F. Montagnini

connectivity (Harvey et al. 2005; Francesconi et al. 2011a, b; Ibrahim et al. 2011). For
example, in a comparison of bird diversity among different land uses in Cordoba,
Colombia, the SPS had the greatest total number of bird species, followed by old
fallows, forest fragments, and pastures with low tree density (Múnera et al. 2009).
The biodiversity benefits of SPS depend on the system components and manage-
ment, with larger biodiversity present in the more complex systems, such as SPS
with natural regenerating trees in pastures, than in the fodder banks or planted timber
trees in pastures (Fig. 4.4). In particular, using native species as part of SPS confers
several advantages to biodiversity conservation (Montagnini and Finney 2011;
Murgueitio et al. 2011; Montagnini et al. 2013; Santos-Gally and Boege 2022).
Native trees and palms play important roles in tropical livestock systems by provid-
ing direct benefits through production of timber and edible fruits for the cattle, and
indirect benefits through nitrogen fixation, soil conservation, and natural biological
control of pests, as well as by providing resources for wildlife (Rivera et al. 2013;
Montoya-Molina et al. 2016; Calle et al. 2017). Endangered or vulnerable tree and
palm species that are deliberately added to cattle ranching systems may have a lower
risk of local extinction.
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 103

[Link] Intensive Silvopastoral Systems

Intensive Silvopastoral Systems (ISPS) are agroforestry arrangements that combine

high-density cultivation of fodder shrubs (4000–40,000 plants per ha) with improved
tropical grasses, and trees or palms at densities of 100–600 individuals per ha. ISPS
were initially developed in Colombia and have expanded to Mexico and Brazil,
among other countries (Murgueitio et al. 2009, 2011; Chará et al. 2017). Several
agroecological principles and strategies are applied in managing ISPS, including:
(i) use of several layers of vegetation (herbs, shrubs, trees, and palms) to maximize
energy transformation; (ii) reduced dependency on agrochemical inputs and energy,
emphasizing interactions and synergisms among biological components to enhance
recycling and biological control; and (iii) incorporation of biodiversity into the
system components and its surroundings (Chará et al. 2017). Under these conditions,
biodiversity restoration and conservation are enhanced in ISPS relative to other SPS,
and even more so to conventional pastures.
In ISPS, the canopy cover, tree diversity, and structural complexity of vegetation
contribute to their functions as habitat, refuge and food resources (Chará et al. 2015).
In various studies from Colombia, examples of ISPS had greater richness of birds
and other animal and plant species, provided natural biological control, and resulted
in increased sightings of rare and endangered species (Table 4.1; Fajardo et al. 2009;
Calle et al. 2009).
At the landscape level, ISPS can contribute to connectivity among patches of
forest as well as to the recovery of strategic sites for the provision of environmental
services (Calle et al. 2012, 2022). Several examples have been documented where
the movement of organisms has been facilitated by ISPS. A matrix permeable to bird
movement can avoid the collapse of small populations of wildlife that are isolated in
forest fragments (Chará et al. 2015).
Because ISPS intensify production in high yielding systems on less land, they can
help to avoid deforestation. At the same time, ISPS are a form of production that
utilize multi-functional landscapes of high value for biodiversity without sacrificing
productivity and economic feasibility (Chará et al. 2015). In this way, they incor-
porate the benefits of both land sparing and land sharing in a single productive and
biodiversity friendly system.

4.2.4 Living Fences and Windbreaks

[Link] Living Fences

The principal role of living fences is to divide, separate, and protect agricultural plots
or cattle. They also provide several services and products—including fuelwood,
fruits, fodder, and shade for cattle—and a major environmental function: promotion
of biodiversity (Harvey et al. 2005, 2008; Francesconi et al. 2011a, b; Ibrahim et al.
2011). In Esparza, Costa Rica, species richness of butterflies was found to be higher
104 F. Montagnini and S. del Fierro

in multi-strata live fences than in pastures with high and low tree densities (Tobar
et al. 2007; De Tobar and Ibrahim 2010). In this same area, Rosales and Sáenz
(2007), found that Mantled Howler Monkeys (Alouatta palliata) preferred riparian
forest and forest fragments for daily activities and also used living fences to move
within and across pastured areas.
Living fences promote bird abundance and diversity by providing habitat and can
be used by generalist and savanna specialist species. At the landscape level, living
fences can provide effective connectivity among forest patches (Francesconi et al.
2011a, b; Francesconi and Montagnini 2015).
Like other AFS, their structure and composition are important factors influencing
their usage by bird species. The presence of birds in living fences could be improved
by increasing tree diversity and allowing trees to grow to mature stages or to develop
broad crowns (Francesconi et al. 2011a, b). However, some of these features, such as
having larger trees in the fences, may not be as practical for the farmer as it may be
harder to manage the wire, and the trees maybe more difficult to prune. Thus living
fence design should recognize the tradeoffs between their productive and conserva-
tion functions.

[Link] Windbreaks

As they are often the only arboreal component of an agricultural landscape, wind-
breaks and hedges play important roles in providing habitats and resources for
animals and other plants. Like living fences, windbreaks and hedges also function
as natural corridors for animal movements across landscapes (Harvey et al. 2005,
2008). Windbreaks tend to be favored by farmers and can be instrumental in
biodiversity conservation and landscape connectivity in fragmented areas. For
example, forest windbreaks in the perimeter of agricultural fields are frequently
used in the Chaco region of Argentina, where about 32% of the original forest
remains on average. These windbreaks range from 30 to 50 m wide and 1700 m long,
representing just 5% of the forest area but providing up to 40% connectivity among
forest fragments. Windbreaks therefore help to maintain the spatial organization of
the landscape and can contribute to the mobility of different species among forest
fragments (Tamashiro 2018). Like living fences, windbreaks have a variety of
agricultural functions and their design and management for environmental functions
must be compatible with their agricultural use.
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 105

Higher/more intensive human impact Lower/less intensive human impact

Human-managed,
g , biodiverse landscapes
& biodiversity islands
Convenonal,
monoculture
Agroforestry systems
agriculture Intact forests
Silvopastoral
Degraded lands Living fences
systems Mulstrata
Windbreaks
High impact human Perennial crops agroforestry
Riparian
landscapes under shade systems
buffers

Less biodiversity More biodiversity

Fig. 4.5 Schematic representation of landscapes along a spectrum of high to low human impact
and low to high biodiversity

4.3 Applying Agroforestry Systems as and Within

Biodiversity Islands

As demonstrated by the different types of AFS and examples of their applications in

a variety of contexts, the contributions of AFS to biodiversity in managed landscapes
depend on the type of AFS, its management, component species, and position within
the landscape matrix (Montagnini 2020). The low-intensive management,
multistrata AFS with native species has the greatest potential to harbor the largest
biodiversity (Figs. 4.1, 4.2, and 4.3). As management intensity increases, biodiver-
sity decreases accordingly (Fig. 4.4). However, even the less heterogeneous AFS
provide greater biodiversity than would otherwise be realized in conventional
monoculture agriculture or degraded landscapes. In addition, farmers value and
protect AFS for their contributions to their livelihoods, thereby also ensuring the
maintenance of ecosystem services they provide, including biodiversity. Figure 4.5
illustrates the location of AFS along this spectrum of high to low human impact and
low to high biodiversity. The diagram points to intact forests as, generally, the land
use supporting the highest biodiversity. However, certain types of AFS can some-
times have higher diversity of species than intact forests nearby, because the AFS
can have open areas that attract species that would not use the forest, for instance,
some species of birds and butterflies (Múnera et al. 2009).
Because AFS can be incorporated into a range of managed landscapes, they have
great potential to function as biodiversity islands or as component parts of biodiver-
sity islands. Building upon the foundations of island biogeography, biodiversity
islands act as ecological refugia, protected areas, or reserves within a landscape and
106 F. Montagnini and S. del Fierro

can exist in a wide range of human dominated landscapes (MacArthur and Wilson
1967; Tjørve 2010). They may be actively implemented as part of a complex
landscape that includes several land uses (e.g., agriculture, forest plantations, etc.)
or they may be part of a passive management practice (i.e., areas left untouched for
practical or economic reasons). Biodiversity islands size, configuration, and position
within the landscape may vary according to different features, including patterns of
human settlement, development, and utilization of natural resources (Laurance
2008). The method of implementation is important, as it will determine the charac-
teristics, position in the landscape, and management of the biodiversity island and
surrounding landscapes.
Living fences and windbreaks function as biodiversity islands within cultivated
areas, providing, as described earlier, habitat and resources for wildlife and other
plant species. Like other biodiversity islands, they can serve as biological corridors,
providing landscape connectivity within fragmented areas. Even relatively small
patches of land, when strategically located around perimeters throughout a cultivated
landscape, can provide important services to biodiversity.
Similarly, homegardens and successional AFS can also serve as important bio-
diversity islands within suburban or urban landscapes (Negret et al. 2022; Soler et al.
2022; Toensmeier 2022). Though their relative area may be small, the presence of
these AFS within developed landscapes can provide otherwise missing ecological
and biological functions.
AFS are also important components among land uses in buffer zones of protected
areas. Biodiversity islands may therefore include a buffer zone that incorporates AFS
to transition from the preserved “island” to areas of greater human impact or
degradation. AFS may uniquely occupy these buffer zones since they are a source
of productivity, but still provide ecological benefits, offering a more gradual transi-
tion for wildlife from the more protected areas. Multiple biodiversity islands spread
over a large area in an optimal configuration can decrease chances of biodiversity
loss through creation of repopulation reserves and biological corridors.
To ensure the effectiveness of different types of AFS as part of biodiversity
islands, land users should take a landscape approach, considering both the prevalent
land uses and the natural ecosystems in the region of study. The prevalent land uses
in a landscape can be arranged along a continuum of successional stages, from the
earliest stages of succession (degraded lands) to more mature stages (forests), with
AFS lying in between these two extremes. Landowners can integrate small scale land
sparing to set aside pieces of the property as untouched natural settings to act as
biodiversity islands. Alternatively, or in addition, they can explore land sharing by
using AFS to incorporate biodiversity conservation and food production on the same
land. Both land sparing and land sharing can provide valuable protection of species
diversity through time (Phalan et al. 2011). In either instance, outcomes result in a
greater biodiversity within the biodiversity islands as compared to otherwise
degraded landscapes.
Deciding on the design and management of AFS within a landscape must,
however, be done appropriately and cautiously. For example, AFS promotion may
be inappropriately used to justify forest cutting and to advance the agricultural
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 107

frontier, citing the role that AFS plays in restoring or preserving biodiversity. AFS
should instead be used to compensate for biodiversity loss by restoring and preserv-
ing biodiversity in regions where the landscape has already been converted to
agriculture or otherwise degraded. In addition, it must be noted that the type of
biodiversity provided by AFS might not be the desired or natural biodiversity of the
region. Increasing overall biodiversity (through novel habitats in AFS) could be in
fact detrimental to some types of native biodiversity. The type of AFS, its intensity,
and use of native versus new crop species all impact the type of biodiversity to be
promoted or restored.
The same judiciousness should be taken when planning and designing AFS as
component parts of biodiversity islands. When compared to a completely unnatural
system (like crop monoculture), AFS stand out for their biodiversity. However,
compared to an intact native forest, AFS biodiversity may have less benefits,
although it may still be better than the alternative monocultures or degraded land-
scapes. While AFS should not be a replacement for intact forests in terms of
biodiversity value, they are a useful tool for conservation in increasingly human
impacted landscapes.

4.4 Supporting and Promoting Agroforestry Systems

as Biodiversity Islands with Incentive Programs

Given the ecological and social benefits of the various types of AFS presented in this
chapter, several studies have looked at programs or designs that have helped to
incentivize and expand their applications. In several examples worldwide, certifica-
tion schemes that guarantee higher quality and ecologically-sound management have
helped to facilitate the sale of AFS products to specialty export markets (Montagnini
and Metzel 2017; Rocha et al. 2017). In combination with direct sale made possible
by aggregating the harvest through producer cooperatives, certification holds the
potential to help sustain the livelihoods of family farmers confronting an evolving
market. By compensating farmers for the extra labor required to produce certified
organic, biodiversity friendly products, certification can bridge the gap between
financial and biodiversity benefits. Organic AFS are also more biodiversity friendly
since the lack of pesticide and herbicide use favors both plant and animal diversity
(Montagnini et al. 2011; Rossi et al. 2011).
Several commodities grown as perennial crops in AFS such as coffee, cacao,
yerba mate, guayusa, and açaí have been able to achieve price surpluses that can
serve as an incentive for the farmer to turn to certified organic or biodiversity
friendly products (Montagnini and Metzel 2017; Rocha et al. 2017). In Pará, Brazil,
açaí (Euterpe oleraceae) is harvested from forests as well as from homegardens in
the estuary of the Amazon River. When marketed well, local producer associations
and cooperatives facilitate the collective sale of açaí, offering alternative points of
sale that may recognize the higher quality product, in response to the demand from
108 F. Montagnini and S. del Fierro

export markets that value traditional production (Pepper and de Freitas Navegantes
Alves 2017). In Argentina, although some times yerba mate production may not be
very attractive due to price instabilities, organic yerba mate producers can get
substantial price surpluses on their product, thereby increasing interest in organic
farming and in yerba mate cultivation under shade in recent years (Montagnini et al.
2011; Eibl et al. 2017).
Using a similar model that promotes a product that has a favorable niche in the
market, biodiverse cacao AFS have often been incorporated into restoration and rural
development projects in Latin America and beyond (Cerda et al. 2014; Gross et al.
2016). This is especially true where the resulting biodiversity friendly product can
obtain higher market prices, as is the case with wild and cultivated cacao in the
Amazon region of Bolivia (Rocha et al. 2017). In El Salvador, the Cacao Alliance
seeks to position the country as an exclusive origin for high quality fine aromatic
cacao in the profitable gourmet segments in international markets (Montagnini and
Metzel 2017). Cacao AFS generate social and environmental benefits such as:
(a) restoring productive landscapes through increased vegetative cover;
(b) increasing water infiltration capacity; (c) increasing size and quality of areas
with restored biological significance where there has been reduction of ecological
niches due to habitat fragmentation; and d) improving connections between biolog-
ical corridors (Frank Sullyvan Cardoza Ruiz, Cacao Alliance, El Salvador, personal
communication, September 2016).
As cattle ranching is expected to continue being an important land use in many
regions, the use of payments for ecosystem services (PES) mechanisms can be one
way to provide incentives for farmers to make their cattle ranching activities more
environmentally friendly. A recent project in Latin America has examined whether
PES has increased the adoption of SPS on cattle farms in Esparza (Costa Rica),
Matiguás (Nicaragua), and Quindío (Colombia) (Ibrahim et al. 2011). An environ-
mental service index (ESI) was developed to determine the level of PES, with birds
as the primary indicator of biodiversity. The number of bird species observed in
pastures with high tree densities or multistrata live fences was higher than that in
degraded pastures and grass monocultures and was comparable to the number of
species in riparian and secondary forest (Sáenz et al. 2007). The percentage of tree
cover and the number of tree species were the two most important parameters that
explained variation in bird species on different land uses.
Before the PES project began, farmers managed the pastures with the use of
herbicides to control weeds, which was associated with high mortality of saplings
and juvenile stages of native tree species (Ibrahim and Camargo 2001). With the
implementation of PES, the use of herbicides was reduced significantly. In addition
to managing natural regeneration to increase tree cover in pastures, private farmers in
Costa Rica and Colombia were trained and supported to produce plants of focal tree
species of interest for conservation. These plants were sold to many cattle farmers
receiving PES and were planted along live fence lines and riparian forest that were
fenced off to keep cattle away from the water sources.
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 109

Similarly, the Mainstreaming Biodiversity into Sustainable Cattle Ranching

project (MBSCR) promotes the planting of 50 focal species of native trees and
palms of global conservation concern in cattle farms in five regions in Colombia
(Calle et al. 2015, 2017). In this context, focal species are native trees and palms that
can be incorporated directly into SPS, live fences, or riparian buffers to enhance
biodiversity and environmental services in cattle dominated landscapes. To achieve
this goal, the MBSCR project uses a short-term PES to partially offset investment
costs in land uses that are compatible with biodiversity. In recognition of the special
effort that must be made to adopt focal species, farmers eligible for PES receive an
additional bonus for planting and caring for these native species on their farms.
These programs and systems have been used in various contexts to provide
support and incentives to farmers. As agricultural activities and human settlements
continue to expand, such incentives may play an increasing role in aiding farmers
and other land-users in incorporating AFS and other biodiversity friendly practices
onto their productive lands or into their neighborhoods, in order to create biodiver-
sity islands that foster connectivity and biodiversity. Figure 4.6 provides a concep-
tual diagram that summarizes the elements contributing to AFS, the predominant
types of AFS, and their primary ecological and economic or social benefits.

4.5 Conclusions

With increasing threats to natural ecosystems worldwide due to human population

pressures, and consequent biodiversity losses and fragmentation, new strategies are
needed to restore, conserve, and connect ecosystems and landscapes. AFS present a
compromise that meet both the needs of biodiversity conservation and sustainable
productivity. Multistrata systems, including homegardens and successional AFS,
hold the highest biodiversity, when compared to more simple AFS. The more
simplified AFS designs, such as some types of perennial crops under shade or SPS
with only few trees species for shade, still can provide more ecological benefits than
monocultures or degraded lands, though they may offer lower biodiversity than the
more complex multistrata systems. Living fences, windbreaks and hedges along the
perimeters of cultivated land can provide connectivity in fragmented agricultural
landscapes. To favor biodiversity restoration and conservation, AFS should increase
their structural complexity in terms of number of species and strata.
Biodiversity islands can contribute to the protection of plants and wildlife in
human managed landscapes and AFS have great potential to be incorporated into
their designs, because of their intermediate role between intensively managed land
and intact, protected land. AFS can be integrated as components of biodiversity
islands, especially in buffer zones of protected areas, to smooth transitions from
higher to lower impact areas. Alternatively, more complex AFS can also constitute
biodiversity islands themselves, for instance among more intensively cultivated land
or within human settled landscapes, where they can provide both economic and
social functions along with their biological and ecological roles. Promotion of AFS
110 F. Montagnini and S. del Fierro

should not be used to justify conversion of forests to agricultural systems, but rather,
can be planned within a broader strategy that seeks to maintain or restore areas of
natural forest within cultivated landscapes.
Like other conservation endeavors, financial incentives are often needed to
promote and sustain AFS in order to provide a competitive advantage over more
intensive agriculture. AFS inherently provide economic benefits because of the
agricultural products they provide, but additional incentives may help to bolster
their biodiversity and ecological contributions. Payments for Ecosystem Services
(PES), certification schemes such as organic certification, and niche marketing, have
been used successfully in this regard and may be integrated into AFS to promote its
usage and to encourage more favorable environmental practices, including pesticide-
free management and/or the use of native species. These types of incentives may
support the increased use of AFS as and within biodiversity islands, in turn provid-
ing services for biodiversity and rural communities alike.

Potenal contribung elements

Indigenous and tradional knowledge

Incenve systems: PES, Cerficaon schemes

Agroforestry systems

Complementary/
Mulstrata Perennial
Silvopastoral linear systems
systems crops under
systems
x Homegardens shade x Living fences
x Successional x Intensive SPS x Windbreaks
x Cacao
agroforestry
x Coffee x Riparian buffer strips
x Yerba mate

Main benefits

Ecological Economic/social
x Increased biodiversity relave to x Increased overall system
monocultures and degraded land producvity and sustainability
x Increased connecvity x Cultural preservaon
x Restoraon of degraded lands x Increased income
x Preservaon of rare species x Diversified income sources

Fig. 4.6 Schematic representation of elements contributing to AFS, the predominant types of AFS,
and their primary ecological and economic/social benefits
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 111

Acknowledgements Thanks to Brett Levin (Yale University School of Forestry and Environmen-
tal Studies) who provided valuable assistance in the preparation of this chapter, and to Kjell Berg for
his continued input and encouragement. Brett Levin and Joseph Orefice provided useful comments
and insights as reviewers.

References

Abete T, Wiersum KF, Bongers F, Sterck F (2006) Diversity and dynamics in homegardens of
southern Ethiopia. In: Kumar BM, Nair PKR (eds) Tropical homegardens: a time-tested
example of sustainable agroforestry. Advances in agroforestry 3. Springer, Dordrecht, 377 pp,
pp 123–142
Beer J, Muschler R, Kass D, Somarriba E (1998) Shade management in coffee and cacao
plantations. Agrofor Syst 38:134–164
Bertsch A (2017) Indigenous successional agroforestry: integrating the old and new to address food
insecurity and deforestation. In: Montagnini F (ed) Integrating landscapes: agroforestry for
biodiversity conservation and food sovereignty, Advances in agroforestry 12. Springer, Cham,
pp 165–178
Bhagwat SA, Willis KJ, Birks HJB, Whittaker RJ (2008) Agroforestry: a refuge for tropical
biodiversity? Trends Ecol Evol 23(5):261–267
Blackman A, Ávalos-Sartorio B, Chow F (2012) Land cover change in agroforestry: shade coffee in
El Salvador. Land Econ 88(1):75–101
Blanckaert I, Swennen RL, Paredes Flores M, Rosas López R, Lira Saade R (2004) Floristic
composition, plant uses and management practices in homegardens of San Rafael Coxcatlán,
Valley of Tehuacán, Mexico. J Arid Environ 57:39–62
Brewer MD (2011) Strategies for cost-effective native species restoration in the sub-tropical
Atlantic Forest of southern Brazil. In: Montagnini F, Finney C (eds) Restoring degraded
landscapes with native species in Latin America. Nova Science Publishers, New York, pp
173–195
Cabral H, Coronel-Bejarano D, Ruiz R, Canete L, Britez E, Rojas V (2020) Living in the shadows:
diversity of amphibians, reptiles and birds in shade-grown yerba mate (Ilex Paraguariensis)
plantations. Austral Ecol:1–12. [Link]
Calle A, Montagnini F, Zuluaga AF (2009) Farmer’s perceptions of Silvopastoral system promotion
in Quindío, Colombia. Bois For Trop 300(2):79–94
Calle Z, Murgueitio E, Chará J (2012) Integrating forestry, sustainable cattle ranching, and
landscape restoration. Unasylva 63:31–40
Calle Z, Hernández M, Murgueitio E, Giraldo AM, Uribe F, Zuluaga AF (2015) Especies focales
del Proyecto Ganadería Colombiana Sostenible. Carta Fedegán 148:54–60
Calle Z, Giraldo AM, Cardozo A, Galindo A, Murgueitio E (2017) Enhancing biodiversity in
neotropical silvopastoral systems: use of indigenous trees and palms. In: Montagnini F
(ed) Integrating landscapes: agroforestry for biodiversity conservation and food sovereignty,
Advances in agroforestry 12. Springer, Cham, pp 417–438
Calle Z, Hernando C, José E, José J, Hernán C, Murgueitio B, Murgueitio A, Murgueitio E (2022) A
highly productive biodiversity island: El Hatico Nature Reserve (Valle Del Cauca,
Colombia). In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Topics in Biodiversity and Conservation. Springer, Cham,
pp 279–304
Callo-Concha D, Denich M (2011) Operationalizing environmental services of agroforestry sys-
tems: functional biodiversity assessment in Tomé-Açú, Pará, Brazil. In: Montagnini F,
Francesconi W, Rossi E (eds) Agroforestry as a tool for landscape restoration. Nova Science
Publishers, New York, pp 143–156
112 F. Montagnini and S. del Fierro

Calmon M, Brancalion PH, Paese A, Aronson J, Castro P, da Silva SC, Rodrigues RR (2011)
Emerging threats and opportunities for large-scale ecological restoration in the Atlantic Forest of
Brazil. Restor Ecol 19(2):154–158
Cerda R, Deheuvels O, Calvache R, Niehaus L, Saenz Y, Kent J, Vilchez S, Villota A, Martinez C,
Somarriba E (2014) Contribution of cocoa agroforestry systems to family income and domestic
consumption: looking toward intensification. Agrofor Syst. [Link]
014-9691-8
Chait G (2015) Café en Colombia: servicios ecosistémicos, conservación de la biodiversidad. In:
Montagnini F, Somarriba E, Murgueitio E, Fassola H, Eibl B (eds) Sistemas Agroforestales.
Funciones productivas, socioeconómicas y ambientales, Serie Técnica Informe Técnico 402.
CATIE/Fundación CIPAV, Turrialba/Cali, pp 349–361
Chará J, Camargo JC, Calle Z, Bueno L, Murgueitio E, Arias L, Dossman M, Molina EJ (2015)
Servicios ambientales de sistemas silvopastoriles intensivos: mejoramiento del suelo y
restauración ecológica. In: Montagnini F, Somarriba E, Murgueitio E, Fassola H, Eibl B (eds)
Sistemas agroforestales: funciones productivas, socioeconómicas y ambientales, Serie técnica
informe técnico No. 402. CATIE/Editorial CIPAV, Turrialba/Cali, pp 331–347
Chará JD, Rivera J, Barahona R, Murgueitio E, Deblitz C, Reyes E, Martins Mauricio R, MolinaJJ
FM, Zuluaga AF (2017) Intensive silvopastoral systems: economics and contribution to climate
change mitigation and public policies. In: Montagnini F (ed) Integrating landscapes: agrofor-
estry for biodiversity conservation and food sovereignty, Advances in agroforestry 12. Springer,
Cham, pp 395–416
Clough Y, Barkmann J, Juhrbandt J, Kessler M, Cherico Wangera T, Anshary A, Buchorig D,
Cicuzza D, Darrasi K, Dwi Putrak D, Erasmi S, Pitopang M, Schmidt C, Schulze CH, Seidell D,
Steffan-Dewenter I, Stenchly K, Vidal S, Weista M, Wielgoss AW, Tscharntke T (2011)
Combining high biodiversity with high yields in tropical agroforests. Proc Natl Acad Sci
USA 108(20):8311–8316
Cockle KL, Leonard ML, Bodrati AA (2005) Presence and abundance of birds in an Atlantic forest
reserve and adjacent plantation of shade-grown yerba mate, in Paraguay. Biodivers Conserv 14:
3265–3288
Davidson S (2005) Shade coffee agro-ecosystems in Mexico: a synopsis of the environmental
services and socio-economic considerations. J Sustain For 21(1):81–95
De Tobar L, Ibrahim M (2010) ¿Las cercas vivas ayudan a la conservación de la diversidad de
mariposas en paisajes agropecuarios? Rev Biol Trop 58(1):447–463
De Tobar L, Ibrahim M, Casasola F (2007) Diversidad de mariposas en un paisaje agropecuario del
Pacífico central de Costa Rica. Agroforestería en las Américas 45:58–65
Diemont SAW, Martin JF, Levy-Tacher SI (2006) Lacandon Maya forest management: restoration
of soil fertility using native tree species. Ecol Eng 28:205–212
Diemont SAW, Bohn JL, Rayome DD (2011) Comparisons of Mayan forest management, resto-
ration, and conservation. For Ecol Manag 261:1696–1705
Eibl B, Montagnini F, López M, Montechiesi R, Barth S, Esterche E (2015) Ilex paraguariensis
A. St.-Hil., yerba mate orgánica bajo dosel de especies nativas maderables, una propuesta de
producción sustentable. In: Montagnini F, Somarriba E, Murgueitio E, Fassola H, Eibl B (eds)
Sistemas agroforestales: funciones productivas, socioeconómicas y ambientales, Serie técnica
informe técnico No. 402. CATIE/Editorial CIPAV, Turrialba/Cali, pp 153–177
Eibl B, Montagnini F, López M, López LN, Montechiesi R, Esterche E (2017) Organic yerba mate,
Ilex paraguariensis, in association with native tree species: a sustainable production
alternative. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity conser-
vation and food sovereignty, Advances in agroforestry 12. Springer, Cham, pp 261–281
Fajardo ND, González RJ, Neira LA (2009) Sistemas ganaderos amigos de las aves. In:
Murgueitio E, Cuartas C, Naranjo J (eds) Ganadería del futuro: Investigación para el desarrollo,
2da edn. Fundación CIPAV, Cali, pp 171–203
Francesconi W, Montagnini F (2015) Los SAF como estrategia para favorecer la conectividad
funcional del paisaje fragmentado. In: Montagnini F, Somarriba E, Murgueitio E, Fassola H,
Eibl B (eds) Sistemas agroforestales: funciones productivas, socioeconómicas y ambientales,
Serie técnica informe técnico No. 402. CATIE/Editorial CIPAV, Turrialba/Cali, pp 363–379
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 113

Francesconi W, Montagnini F, Ibrahim M (2011a) Living fences as linear extensions of forest

remnants: a strategy for restoration of connectivity in agricultural landscapes. In: Montagnini F,
Finney C (eds) Restoring degraded landscapes with native species in Latin America. Nova
Science Publishers, New York, pp 115–126
Francesconi W, Montagnini F, Ibrahim M (2011b) Using bird distribution to evaluate the potential
of living fences to restore landscape connectivity in pasturelands. In: Montagnini F,
Francesconi W, Rossi E (eds) Agroforestry as a tool for landscape restoration. Nova Science
Publishers, New York, pp 133–142
Gockowski J, Tchatat M, Dondjang JP, Hietet G, Fouda T (2010) An empirical analysis of the
biodiversity and economic returns to cocoa agroforests in Southern Cameroon. J Sustain For
29(6):638–670. [Link]
González-Soberanis C, Casas A (2004) Traditional management and domestication of
tempesquistle, Sideroxylon palmeri (Sapotaceae) in the Tehuacan-Cuicatlán Valley, Central
México. J Arid Environ 59:245–258
Griffith DM (2000) Agroforestry: a refuge for tropical biodiversity after fire. Conserv Biol 14:325–
326
Gross L, Castro-Tanzi S, Scherr SJ (2016) Connecting farms to landscapes for biodiversity
conservation. The CAMBio project in Central America. Ecoagriculture discussion paper
no. 15, Ecoagriculture Partners, Washington DC, 31 pp
Harvey C, Villanueva C, Villacís J, Chacón M, Muñoz D, López M, Ibrahim M, Gómez R,
Taylor R, Martínez J, Navas A, Sáenz J, Sánchez D, Medina A, Vílches S, Hernández B,
Pérez A, Ruíz F, López F, Lang I, Kunth S, Sinclair F (2005) Contribution of live fences to the
ecological integrity of agricultural landscapes. Agric Ecosyst Environ 111:200–230
Harvey CA, Komar O, Chazdon R, Ferguson BG, Finegan B, Griffith DM, Martínez-Ramos M,
Morales H, Nigh R, Soto-Pinto L, Van Breugel M, Wishnie M (2008) Integrating agricultural
landscapes with biodiversity conservation in the Mesoamerican hotspot. Cons Biol 22(1):8–15
Hobbs RJ, Walker LR, Walker J (2007) Integrating restoration and succession. In: Walker LR,
Walker J, Hobbs RJ (eds) Linking restoration and ecological succession. Springer, New York,
pp 168–179
House PH, Ochoa L (1998) La diversidad de especies útiles en diez huertos en la aldea de Camalote,
Honduras. In: Lok R (ed) Huertos caseros tradicionales de América Central: características,
beneficios e importancia, desde un enfoque multidisciplinario. Centro Agronómico Tropical de
Investigación y Enseñanza (CATIE), Turrialba, pp 61–84
Ibrahim M, Camargo JC (2001) ¿Cómo aumentar la regeneración de árboles maderable en potreros?
Agroforestería en las Américas 8(32):35–41
Ibrahim M, Casasola F, Villanueva C, Murgueitio E, Ramírez E, Sáenz S, Sepúlveda C (2011)
Payment for environmental services as a tool to encourage the adoption of silvo-pastoral
systems and restoration of agricultural landscapes dominated by cattle in Latin America. In:
Montagnini F, Finney C (eds) Restoring degraded landscapes with native species in Latin
America. Nova Science Publishers, New York, pp 197–219
Ilany T, Ashton M, Montagnini F, Martinez C (2010) Using agroforestry to improve soil fertility:
effects of intercropping on Ilex paraguariensis (yerba mate) plantations with Araucaria
angustifolia. Agrofor Syst 80(3):399–409
Jose S (2009) Agroforestry for ecosystem services and environmental benefits: an overview.
Agrofor Syst 76:1–10
Jose S (2012) Agroforestry for conserving and enhancing biodiversity. Agrofor Syst 85(1):1–8
Kumar BM, Nair PKR (2006) (eds) Tropical Homegardens: a time-tested example of sustainable
agroforestry, Advances in agroforestry 3. Springer, Dordrecht, 377 pp
Laurance WF (2008) Theory meets reality: how habitat fragmentation research has transcended
island biogeographic theory. Biol Conserv 141(7):1731–1744
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press,
Princeton, p 203
114 F. Montagnini and S. del Fierro

Mas AH, Dietsch T (2003) An index of management intensity for coffee agroecosystems to evaluate
butterfly species richness. Ecol Appl 13(5):1491–1501
Miccolis A, Mongeli Peneireiro F, Rodrigues Marques H, Mascia Vieira DL, Francia Arco-Verde-
M, Rigon Hoffmann M, Rehder T, Barbosa Pereira AV (2016) Agroforestry Systems for
agroecological restoration: how to reconcile conversation with production, options for the
Cerrado and the Caatinga. Brasília: Instituto Sociedade, População e Natureza – ISPN/World
Agroforestry Centre – ICRAF, 240p. [Link]
PDFS/[Link]
Moguel P, Toledo V (1999) Biodiversity conservation in traditional coffee systems of Mexico.
Cons Bio 13(1):11–21
Montagnini F (2006) Homegardens of Mesoamerica: biodiversity, food security, and nutrient
management. In: Kumar BM, Nair PKR (eds) Tropical homegardens: a time-tested example
of sustainable agroforestry, Advances in agroforestry, vol 3. Springer, Dordrecht, pp 61–84
Montagnini F (2020) The contribution of agroforestry to restoration and conservation: biodiversity
islands in degraded landscapes. In: Dagar JC, Gupta SR, Teketay D (eds) Agroforestry for
degraded landscapes: recent advances and emerging challenges, vol 1. Springer, Singapore, pp
445–479. [Link]
Montagnini F, Berg KE (2019) Feeding the world. In: Carver F, Gossington H, Manuel C (eds)
Sustainable development goals. Transforming our world. Witan Media/United Nations
Association – UK, Painswick/London, pp 98–100. [Link]
feeding-the-world/
Montagnini F, Finney C (2011) Payments for environmental services in Latin America as a tool for
restoration and rural development. Ambio 40:285–297
Montagnini F, Metzel R (2017) The contribution of agroforestry to sustainable development goal 2:
end hunger, achieve food security and improved nutrition, and promote sustainable
agriculture. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity conser-
vation and food sovereignty, Advances in agroforestry 12. Springer, Cham, pp 11–45
Montagnini F et al (1992) Sistemas agroforestales. Principios y aplicaciones en los trópicos, 2d.
ed. Organización para Estudios Tropicales (OTS). San José, 622 pp. [Link]
images/downloads/information-resources/library/[Link]
Montagnini F, Eibl B, Barth SR (2011) Organic yerba mate: an environmentally, socially and
financially suitable agroforestry system. Bois For Tropiques 308:59–74. https://
drfl[Link]/organic-yerba-mate-an-environmentally-socially-and-
financially-suitable-agroforestry-system/
Montagnini F, Ibrahim M, Murgueitio Restrepo E (2013) Silvopastoral systems and mitigation of
climate change in Latin America. Bois For Tropiques 316(2):3–16
Montagnini F, Somarriba E, Murgueitio E, Fassola H, Eibl B (eds) (2015) Sistemas agroforestales:
funciones productivas, socioeconómicas y ambientales. Serie técnica informe técnico No. 402.
CATIE/Editorial CIPAV, Turrialba/Cali, 454pp
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity islands: strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human-dominated environments, Topics in Biodiversity and Conservation.
Springer, Cham, pp 1–35
Montoya-Molina S, Giraldo-Echeverri C, Montoya-Lerma J, Chará J, Escobar F, Calle Z (2016)
Land sharing vs land sparing in the dry Caribbean lowlands: a dung beetles’ perspective. Appl
Soil Ecol 98:204–212
Múnera E, Bock BC, Bolívar Vergara DM, Botero Botero JA (2009) Composición y estructura de la
avifauna en diferentes hábitats en el Departamento de Córdoba, Colombia. In: Murgueitio E,
Cuartas C, Naranjo J (eds) Ganadería del futuro: Investigación para el desarrollo, 2da
Ed. Fundación CIPAV, Cali, pp 206–225
Murgueitio E, Cuartas C, Naranjo J (eds) (2009) Ganadería del futuro: Investigación para el
desarrollo, 2da Ed. Fundación CIPAV (Centro para la Investigación en Sistemas Silvopastoriles
de Producción Agropecuaria), Cali, p 490
4 Functions of Agroforestry Systems as Biodiversity Islands in Productive. . . 115

Murgueitio E, Calle Z, Uribe F, Calle A, Solorio B (2011) Native trees and shrubs for the productive
rehabilitation of tropical cattle ranching lands. For Ecol Manag 261(10):1654–1663
Myers N, Mittermeier RA, Mittermeier CG, Fonseca G, Kent J (2000) Biodiversity hotspots for
conservation priorities. Nature 403:858–863
Nair PKR, Kumar BM (2006) Introduction. In: Kumar BM, Nair PKR (eds) Tropical homegardens:
a time-tested example of sustainable agroforestry, Advances in agroforestry, vol 3. Springer,
Dordrecht, pp 1–10
Negret HRC, Negret R, Montes-Londoño I (2022) Residential garden design for urban biodiversity
conservation: experience from Panama City, Panama. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Topics in Biodiversity
and Conservation. Springer, Cham, pp 387–417
Peneireiro FM (1999) Sistemas Agroflorestais Dirigidos pela Sucessão Natural: um Estudo de Caso.
(Agroforestry systems driven by natural succession: a case study). Universidade de São Paulo,
Escola Superior de Agricultura “Luiz de Queiroz”. Master’s dissertation, 138 pp
Pepper LG, De Freitas Navegantes Alves L (2017) Small-scale Açaí in the global market: adding
value to ensure sustained income for forest farmers in the Amazon Estuary. In: Montagnini F
(ed) Integrating landscapes: agroforestry for biodiversity conservation and food sovereignty,
Advances in agroforestry 12. Springer, Cham, pp 211–234
Perfecto I, Vandermeer J, Mas AA, Soto Pinto L (2005) Biodiversity, yield, and shade coffee
certification. Ecol Econ 54:435–446
Peters CM (2018) Managing the wild. Stories of people and plants and tropical forests. Yale
University Press, New Haven, p 208
Peyre A, Guidal A, Wiersum KF, Bongers F (2006) Homegardens dynamics in Kerala, India. In:
Kumar BM, Nair PKR (eds) Tropical homegardens: a time-tested example of sustainable
agroforestry, Advances in agroforestry, vol 3. Springer, Dordrecht, pp 87–104
Pezo D, Ibrahim M (1999) Sistemas Silvopastoriles, 2nd ed. Proyecto Agroforestal CATIE/GTZ,
Módulo de Enseñanza Agroforestal No. 2. CATIE, Turrialba, Costa Rica, 275pp
Phalan B, Onial M, Balmford A, Green RE (2011) Reconciling food production and biodiversity
conservation: land sharing and land sparing compared. Science 333(6047):1289–1291
Ribeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, Hirota MM (2009) The Brazilian Atlantic
Forest: how much is left, and how is the remaining forest distributed? Implications for conser-
vation. Biol Conserv 142:1141–1153
Rivera L, Armbrecht I, Calle Z (2013) Silvopastoral systems and ant diversity conservation in a
cattle-dominated landscape of the Colombian Andes. Agric Ecosyst Environ 181:188–194
Rocha P, Niella F, Keller H, Montagnini F, Metzel R, Eibl B, Kornel J, Romero F, López L,
Araujo J, Barquinero J (2017) Ecological indigenous (EIK) and scientific (ESK) knowledge
integration as tool for sustainable development in indigenous communities. experience in
Misiones, Argentina. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity
conservation and food sovereignty, Advances in agroforestry 12. Springer, Cham, pp 235–260
Rolim SG, Chiarello AG (2004) Slow death of Atlantic forest trees in cocoa agroforestry in
southeastern Brazil. Biodivers Conserv 13:2679–2694
Rosales M, Sáenz JC (2007) Uso de coberturas de vegetación por tropas de monos congos (Alouatta
palliata) y carablanca (Cebus capucinus) en un agropaisaje y percepciones de finqueros con
respecto a la conservación de los primates en Esparza, Costa Rica. In: III Congreso
Iberoamericano Sobre Desarrollo y Ambiente (CISDA), Heredia, Costa Rica
Roshetko JM, Purwanto E, Moeliono M, Widayati A, Purnomosidhi P, Mahrizal, Wau D,
Perdana A, Martini E, Gaol A, Paramita E, Dahlia L, Suyanto KN, Manurung G, Yuliani L,
Rohadi D, Manalu P, Umar A, Millang S (2016) Agroforestry and Forestry in Sulawesi: linking
knowledge with action. Annual progress report year 5 (April 2015–March 2016). World
Agroforestry Centre (ICRAF) Southeast Asia Regional Program, Center for International
Forestry Research; Bogor, Indonesia, Operation Wallacea Trust, Bogor, Indonesia, Faculty of
Forestry, Hasanuddin University, Makassar, Indonesia, 137 pp
Rossi E, Montagnini F, de Melo E (2011) Effects of management practices on coffee productivity
and herbaceous species diversity in agroforestry systems in Costa Rica. In: Montagnini F,
116 F. Montagnini and S. del Fierro

Francesconi W, Rossi E (eds) Agroforestry as a tool for landscape restoration. Nova Science
Publishers, New York, pp 115–132
Sáenz JC, Villatoro F, Ibrahim M, Fajardo D, Pérez M (2007) Relación entre las comunidades de
aves y la vegetación en agropaisajes dominados por la ganadería en Costa Rica, Nicaragua y
Colombia. Agroforestería en las Américas 45:37–48
Santos-Gally R, Boege K (2022) Biodiversity islands. Native tree islands within silvopastoral
systems in a neotropical region. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments, Topics in Biodiversity and Conservation.
Springer, Cham, pp 117–138
Schulz J (2011) Imitating natural ecosystems through successional agroforestry for the regeneration
of degraded lands – a case study of smallholder agriculture in northeastern Brazil. In:
Montagnini F, Francesconi W, Rossi E (eds) Agroforestry as a tool for landscape restoration.
Nova Science Publishers, New York, pp 3–17
Soler R, Benítez J, Solá F, Lencinas MV (2022) Biodiversity islands at the world’s
southernmost city: plant, bird and insect conservation in urban forests and peatlands of Ushuaia,
Argentina. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments, Topics in Biodiversity and Conservation. Springer, Cham, pp 419–437
Somarriba E, Harvey CA, Samper M, Anthony F, Gonzalez J, Staver C, Rice RA (2004) Biodi-
versity conservation in neotropical coffee (Coffea Arabica) plantations. In: Schroth G, da
Fonseca GAB, Harvey CA, Gascon C, Vasconcelos HL, Izac MN (eds) Agroforestry and
biodiversity conservation in tropical landscapes. Island Press, Washington, D.C., pp 198–226
SOS Mata Atlântica, INPE (2014) Atlas dos remanescentes florestais da Mata Atlântica: período
2012–2013. Instituto Nacional de Pesquisas Espaciais [Link] [Link]
[Link]/
Tamashiro S (2018) Corredores verdes reducen impactos del desmonte. Retrieved 17 October 2018,
from [Link]
keting&utm_admin¼8097&utm_medium¼email&utm_campaign¼#.[Link]
Teodoro AV, Munoz A, Tscharntke T (2011) Early succession arthropod community changes on
experimental passion fruit plant patches along a land-use gradient in Ecuador. Agric Ecos
Environ 140(1–2):14–19
Terrones Rincón TDRL, Hernández Martínez MA, Ríos Ruiz SA, Martínez Ayala C (2011)
Non-wood products from native multipurpose trees from agroforestry homegardens in the
semiarid Mexican Plateau. In: Montagnini F, Francesconi W, Rossi E (eds) Agroforestry as a
tool for landscape restoration. Nova Science Publishers, New York, pp 85–97
Tjørve E (2010) How to resolve the SLOSS debate: lessons from species-diversity models. J Theor
Biol 264(2):604–661
Toensmeier E (2022) Paradise lot: a temperate urban multistrata agroforestry island of
biodiversity. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human
dominated environments. Topics in Biodiversity and Conservation, Springer, Cham, pp 439–459
Tscharntke T, Clough Y, Bhagwat SA, Buchori D, Faust H, Hertel D, Holscher D, Juhrbandt J,
Kessler M, Perfecto I, Scherber C, Schroth G, Veldkamp E, Wanger TC (2011) Multifunctional
shade-tree management in tropicalagroforestry landscapes – a review. J Appl Ecol 48:619–629
Udawatta RP, Rankoth LM, Jose S (2019) Agroforestry and biodiversity. Sustainability 11:2879.
[Link]
Wezel A, Bender S (2003) Plant species diversity of homegardens of Cuba and its significance for
household food supply. Agrofor Syst 57:39–49
Wezel A, Ohl J (2006) Homegarden plant diversity in relation to remoteness from urban centers: a
case study from the Peruvian Amazon. In: Kumar BM, Nair PKR (eds) Tropical homegardens: a
time-tested example of sustainable agroforestry, Advances in agroforestry 3. Springer, Dor-
drecht, pp 159–184
Young K (2017) Mimicking nature: a review of successional agroforestry systems as an analogue to
natural regeneration of secondary forest stands. In: Montagnini F (ed) Integrating landscapes:
agroforestry for biodiversity conservation and food sovereignty, Advances in agroforestry 12.
Springer, Cham, pp 179–209
Chapter 5
Biodiversity Islands: The Role of Native
Tree Islands Within Silvopastoral Systems
in a Neotropical Region

Rocio Santos-Gally and Karina Boege

Abstract Neotropical rainforests have lost about 35% of their native vegetation due
to deforestation for agricultural purposes and/or livestock grazing. In addition, many
forest remnants are immersed in a grassland matrix with low productivity, due to soil
degradation and erosion, high temperature and low humidity. Under such a scenario,
silvopastoral systems (SPS) can be a sustainable alternative to increase the profit-
ability of livestock production, allowing areas not suitable for livestock to be
restored for biodiversity conservation. In this chapter we review the advantages of
SPS from their prehistoric appearance to the more innovative intensive SPS (iSPS),
and propose that the inclusion of native tree islands (as ecological restoration plots)
can further increase the recovery of biodiversity in tropical regions. For the imple-
mentation of these native tree islands, we review different approaches for ecological
restoration, from passive ones, such as exclusion of cattle in pastures, to the most
labor-intensive ones, such as planting seedlings within cattle farms. We also discuss
the relevance of considering the different components of biological diversity
(genetic, functional and phylogenetic diversity) for species selection during these
ecological restoration efforts and highlight the use of phylogenetic diversity as a
useful predictor of functional diversity and ecological dynamics. Finally, we present
an example of a recent intervention in a tropical region of Mexico including the
implementation of islands of native trees taking phylogenetic diversity into account.
Our proposal is to conserve and restore, not only the species but the unique
evolutionary history of the great biodiversity accumulated in the tropics.

Keywords Agroforestry · Biodiversity conservation · Ecosystem functioning ·

Functional traits · Livestock · Phylogenetic diversity

R. Santos-Gally (*)
CONACYT-Instituto de Ecología, Universidad Nacional Autónoma de México, México,
CDMX, Mexico
e-mail: rsantos@[Link]
K. Boege
Instituto de Ecología, Universidad Nacional Autónoma de México, México, CDMX, Mexico

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 117
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
118 R. Santos-Gally and K. Boege

5.1 Introduction

We are currently facing a global biodiversity loss of a magnitude not seen before
during human history. To date, 35% of the original vegetation cover around the
world has been lost, and of the remaining forested areas about 82% have been
degraded by human activities (Watson et al. 2018). Furthermore, it is estimated
that every year between 11,000 and 58,000 plant species go extinct (Ceballos et al.
2017). In the case of vertebrate animals, more than 50% of the species are endan-
gered (Hoffmann et al. 2010; Dirzo et al. 2014; Ceballos et al. 2017), and
populations of most representative insect orders are decreasing in numbers
(Hoffmann et al. 2010; Dirzo et al. 2014). For plants, the situation is critical as
well, particularly in the tropics, as most of the world’s tropical tree species now are
considered to be threatened (Steege et al. 2015). Last, but not least important, our
reduced knowledge regarding microorganism diversity does not even allow us to
estimate how many species are being lost at this taxonomic level.
A generalized decrease in biodiversity across the world has led to a striking
reduction of ecosystem functionality and different ecological processes providing
climate regulation, water availability, nutrient cycling, soil fertility and disease
control (Haines-Young and Potschin 2010; Segan et al. 2016). The loss of these
ecosystem services has affected human socioeconomic resilience, compromising
natural resources and livelihood for future generations. Food and water provisioning,
resilience to extreme climatic events, incidence of pest and zoonotic diseases, are
just some processes already affected by anthropogenic activities in the last century
(Haines-Young and Potschin 2010; Watson et al. 2018).
In both terrestrial and aquatic ecosystems, the main drivers of species extinction
afecting ecosystem processes are habitat loss, the spread of invasive species, acute
climate change, widespread environmental pollution, and the over-exploitation of
species (Dirzo et al. 2014; Bellard et al. 2016; Román-Palacios and Wiens 2020). In
particular, the expansion of farmland has led to the loss of ~100 million ha of tropical
forests around the world in just over three decades (AGAL 2008; Lewis et al. 2015).
Tropical regions in Latin America have been severely affected in the last decades,
due to the expansion of agricultural and livestock activities, with the loss of between
3.8 (Achard et al. 2014) and 4.88 (Baccini et al. 2012) million ha/year of wet and dry
forests, with a mean annual deforestation rate of 0.49% (Achard et al. 2014). For
example, as a result of an aggressive policy promoting the introduction of African
improved grasses and cattle species for extensive livestock production, Mexico has
lost up to 80% of the original tropical forest cover in the last 50 years (Challenger
and Soberón 2008).
Extensive cattle ranching requires large extensions of pastures, and a heavy
investment in herbicides and pesticides to maintain high productivity, which
makes it economically challenging and, in the long run, inefficient (Arellano et al.
2018). Without any tree cover, open pastures promote a significant increase in local
temperatures, the loss of water sources, soil compaction, a reduction of available
nutrients in the system and an overall biodiversity loss at all taxonomic levels, from
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 119

soil microbes to large vertebrates at the top of food chains. Hence, this productive
activity causes a continuous deforestation of tropical forests (Steinfeld et al. 2006).
The maintenance of healthy and sustainable socio-ecological systems requires a
generalized transformation process and reconsideration of the current ways of
carrying out agricultural, forestry and livestock activities. Different alternatives
should aim for sustainable food production systems encompassing the conservation
of biodiversity, ecosystem functioning and ecological processes, incorporating the
values, knowledge and interests of local producers and the enhancement of local
markets. In this sense, agroforestry and silvopastoral systems (SPS) play an impor-
tant role in advancing the interactive trajectories of socio-economic and environ-
mental changes (Altieri 1999; Montagnini 2017; Chará et al. 2019; Calle 2020).
Through so-called land sharing, these systems optimize ecological processes by
increasing functional trait diversity (e.g. incorporating nitrogen fixing species) that
in turn delivers ecosystem services to replace external inputs (e.g. fertilizers), turning
fragmented and less-productive landscapes into a biodiverse production matrix
(Perfecto and Vandermeer 2010; Cardinale et al. 2012; Calle et al. 2013; Kremen
and Merenlender 2018). For example, the combination of different vegetation strata
promotes the use of solar energy in the conversion of biomass, which can be used in
animal feed, and can also add nutrients to the soil through leaf decomposition, and
water filtration by roots (Cardinale et al. 2012). In this sense, agroecology and its
associated systems can be considered an inextricable component of the biodiversity
conservation agenda.
In this chapter we review how SPS in the neotropics can be a sustainable
alternative for livestock production. We highlight the productive and economic
benefits derived from SPS, especially those where the use of high protein forage
species is intensified, and how, ideally, such systems can liberate fragile areas within
farms for ecological restoration. We discuss different approaches to ecological
restoration (from cattle exclusion and natural regeneration to plantations), both in
un-productive abandoned pastures, and within cattle ranches. As SPS prove their
productive efficiency and producers become more sensitive to increasing biodiver-
sity in their land, the discussion of which species are the most suitable for the proper
functioning of the ecosystem becomes crucial. Therefore, we emphasize the impor-
tance of considering genetic, functional and phylogenetic diversity when selecting
plant species for ecological restoration.
For example, previous studies suggest that an increase in genetic diversity is
associated with a higher ability to adapt and respond to environmental changes
(Kettenring et al. 2014). Phylogenetic diversity – a measure of the amount of
evolutionary history represented within a community – has also been suggested to
increase ecosystem resilience and productivity due to a broader diversity of traits that
represent the functional diversity of the ecosystem (Cadotte et al. 2008, 2013).
Community diversity is quantified through different measures of phylogenetic and
functional diversity, which are based on the distance that separates the species from
their most recent common ancestor. Conserving and/or restoring biodiversity, taking
into account these evolutionary relationships, means that we are protecting not only
ecological processes but also their evolutionary history.
120 R. Santos-Gally and K. Boege

5.2 Silvopastoral Systems

5.2.1 SPS as a Sustainable Alternative for Livestock

Production

Silvopastoral systems are characterized by intentional and/or spontaneous arrange-

ments that combine trees or shrubs, herbs and grasses for nutrition, fiber, medicine,
and energy for people and animals. This alternative way of feeding livestock has a
long history related to human use of available resources in particular ecosystems
(Jose et al. 2019). Among the oldest examples is the Dehesa landscape from the
Iberian Peninsula (also called Montado in Portugal), Italy, France and Morocco,
which includes the combination of different trees (Quercus ilex, Q. suber, Q. rubur,
Argania spinosa, Platanus orientalis, etc.) and grasses with a variety of livestock
types (cows, sheep, pigs, horses, goats) (Fig. 5.1). On a historic timescale, use of
Dehesas likely began in the mid-late Neolithic (ca. 4500–3300 BC), when humans
used management interventions such as thinning to establish pastures underneath
Mediterranean tree canopies to feed livestock (López-Sáez et al. 2007, 2014).
Paleoecological records show a transition from a Mediterranean vegetation, domi-
nated by oaks and junipers in the tree layer (Q. ilex-type, Juniperus-type), and olives
(Olea-type), lentisk (Pistaccia lentiscus-type) and gum rockrose (Cystus ladanifer-
type) in the shrub strata, towards an increase of heliophilous taxa, such as ribwort
plantain (Plantago lanceolata-type), nettle (Urtica dioica type), Amaranthaceas and

Fig. 5.1 Silvopastoral systems in the South of Europe and North Africa, (a) SPS in France with
Quercus robur, grasses and Aubrac cattle, (b) Quercus suber, herbs and grasses with black Iberian
pigs in the Sierra Norte of Seville, Spain; (c) Moroccan mountain goats with Argania spinosa; d)
France, Platanus orientalis, herbs and grasses with horses. (Photos: R. Santos-Gally)
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 121

other nitrophilic species (those adapted to growth in nitrogen-rich soils), such as

Rumex-type and Aster-type, associated with the presence of domestic animals and
agriculture. In addition, archaeological records suggest the use of acorns to produce
flour, which could have been used to supplement the human diet (Cerrillo et al.
2005) providing an additional benefit. The perpetuation of the Dehesa landscape
since prehistoric times is an example showing that anthropic intervention in ecosys-
tems can be sustainable, ensuring productive benefits while conserving biodiversity
and climate regulation (Garrido et al. 2017; Ferraz-de-Oliveira et al. 2016). In fact,
the Dehesa has been considered a traditional cultural ecosystem or semi-natural
ecosystem (according to the European Union legal context) and serves as a model
ecosystem in ecological restoration (Gann et al. 2019).
Neotropical livestock farming, since its inception in the sixteenth century, also
originally occurred under the shelter of trees (Murgueitio and Ibrahim 2008;
Guevara et al. 2018; Jose et al. 2019), ensuring a sustainable productive system
preserving local biodiversity. It was not until the last century that a mistaken
conception of incompatibility between trees and grasses permeated among ranchers,
basically due to the introduction of heliophilous grasses of the genera Cynodon,
Megathyrsus, Brachiaria, Urochloa, Pennisetum, Dichanthium, Cenchrus,
Bothriochloa and others which were well adapted to browsing by large herbivores
and to fire. Their use promoted what we know today as extensive cattle ranches
without any tree cover.
Nevertheless, the presence of trees within the productive landscape is still com-
mon in tropical farms (Kú et al. 1999; Guevara et al. 2018). These trees have
multiple uses as a supply of fruit, medicine, forage, shade, wood, live fences,
windbreaks and river corridors. The presence of trees is also evidence of the
importance of traditional ecological knowledge and that their presence is of utmost
importance for the management and conservation of natural resources (Raymond
et al. 2010; Guevara et al. 2018; Tarbox et al. 2020). This knowledge should be an
ally in the adoption and expansion of silvopastoral practices. Furthermore, synergies
between local and scientific ecological knowledge can promote co-produced inno-
vations for the transition towards more sustainable practices (Reed et al. 2008).

5.2.2 Innovations of Intensive Silvopastoral Systems

for Livestock Sustainability in Latin America

Intensive silvopastoral systems (iSPS) are a novel agroecological alternative

consisting of a well-designed combination of different vegetation strata such as
grasses, herbs, shrubs, trees and/or palms. This species arrangement increases
biodiversity at different levels and therefore, functional traits that underpin different
functions in ecosystemic processes and their benefits for productive systems such as
nutrient recycling, soil fertility enhancement, increased primary productivity, and cli-
mate buffering. In particular, highly proteinic forage plants, with functional traits
122 R. Santos-Gally and K. Boege

such as high leaf N and P content for direct browsing by livestock are a key element
in iSPS. Species currently used for the shrub strata within Latin American iSPS are
Leucaena leucocephala, Tithonia diversifolia and Guazuma ulmifolia (Murgueitio
et al. 2015). Planted in high densities (more than 10,000 ha
1) within the paddocks,
L. leucocephala facilitates high fixation and transfer of nitrogen, while T. diversifolia
favors the solubilization of phosphorous in acid soils thus benefitting the associated
pastures (Ojeniyi et al. 2012; González 2013; Bacab 2013). These species also
provide high protein content for cattle, which translates into higher productivity.
The result of this arrangement is a more productive and diverse system, with greater
species richness and more diverse functional traits, although biodiversity is not
necessarily maximized.
Intensive rotation of livestock among paddocks practiced with permanent or
mobile electric fences is essential to allow forage plant and grass recovery. It is
also fundamental to ensure a supply of good quality water through fixed or mobile
drinkers and to provide mineral salt. Live fences often divide paddocks and are used
to limit the area where the cows graze (Murgueitio et al. 2019). In addition,
arrangements in the tree stratum may include timber, fruit and forage species
which serve as food and shade for livestock, as well as isolated or small groups
(i.e. “vegetation islands”) of native trees providing ecological services such as
pollination, nutrient recycling, water filtration, fruit and seed dispersion, and carbon
fixation. Depending on the ecoregion and productive interest, the number of adult
trees can range from 100 to 600 ha
1 (Chará et al. 2019). The rotation of livestock
and the retention of forest cover, through living fences and trees in paddocks, play a
key role for ecological restoration (Martínez-Ramos et al. 2016). The first helps
reduce soil compaction and improves nutrient recycling, thus facilitating the appear-
ance of new propagules. The latter act as biological corridors, to facilitate the
movement of pollinators, seed dispersers, herbivores and other animals (Omeja
et al. 2016).
The different components of the iSPS (Fig. 5.2) increase biodiversity and its
associated functional traits. Ideally, these systems allow a transformation of live-
stock farming landscapes of homogeneous vegetation to complex networks of
vegetation, where the areas adjacent to the pastures are restored to form biological
corridors, and the points where the productive zones join can form native tree islands
(see green squares in Fig. 5.2). Native tree islands become a source of propagules,
nesting sites, biological pest control and a sink for soil nutrients, and the increase in
these ecological processes is linked to environmental, productive and economic
improvements. Cattle is fed better thanks to efficient and quality grazing resulting
from high-protein forage. The animals suffer less heat stress, as the temperature in
wooded paddocks can decrease between 4–8  C in reference to open pasture areas,
and distances to search for water or food are reduced through access to mobile
drinkers and more forage biomass. All this can result in an increase of 5 to up to
10 times the amount of meat production and up to an additional 80% in the volume
of milk produced in comparison with conventional pastures (Murgueitio and Giraldo
2009; Navas 2010; González 2013; Bacab et al. 2013; Chará et al. 2019; Jose et al.
2019; Murgueitio et al. 2019). In addition, intensive rotation of livestock results in an
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 123

Fig. 5.2 Scheme of a cattle ranch with intensive silvopastoral systems, native-tree islands (green
squares), a riparian vegetation corridor, a water system for drinking fountains and living fences.
(Modified from Santos-Gally et al. 2019)

increase of stocking rate per ha of four to five times higher than what is achieved in
extensive cattle ranching (Murgueitio et al. 2019). Expenses on external inputs, such
as fertilizers, can be reduced to zero due to the increased nitrogen fixation and other
nutrients contributed by forage shrubs (González 2013, Murgueitio et al. 2019).
Forage biomass is also increased by up to 47% compared to that within a monocul-
ture of pastures, therefore reducing the need for food supplementation by more than
half (González 2013; Calle et al. 2013).
An iSPS, when well designed and managed, is a habitat with a more complex
structural vegetation than the treeless grass monocultures of extensive pastures, and
therefore, becomes a refuge enabling increased diversity of arthropofauna, local and
migratory birds, reptiles and small mammals (Harvey and Haber 1998; Giraldo et al.
2010; Rivera et al. 2013; Omeja et al. 2016). The increased biodiversity of iSPS
allows some of the ecological processes of natural habitats to be restored, such as the
acceleration of manure recycling by different types of dung beetles. Some iSPS have
been shown to conserve 60.7% of forest beetle species, thus reducing laying sites of
blood-eating cattle flies which can negatively impact weight gain by cattle
(Murgueitio and Giraldo 2009; Giraldo et al. 2018). Additional ecological processes
are restored such as fruit and seed dispersal, pollination and nutrient recycling. In
this way, iSPS could also present a greater diversity of functional traits, although not
necessarily maximizing phylogenetic diversity.
At an interspecific level, functional traits determine the adaptive attributes by
which an organism can survive in a given environment (response traits) and their
124 R. Santos-Gally and K. Boege

contribution to ecosystem functioning and services (effect traits) (Violle et al. 2007;
Cardinale et al. 2012; Srivastava et al. 2012). For example, in tropical forests,
whether roots are deep or superficial has implications for resource competition, as
species with deep roots can survive better during long periods of water stress (Paz
et al. 2015). Species with pivotal deep roots contribute to soil development, ground-
water and streamflow regulation, soil carbon storage and moisture content in the
atmosphere (Pierret et al. 2016). Other functional traits, such as leaves and roots with
lower C:N ratios, influence organic matter decomposition rates (these being faster
when the C:N ratios are lower) and thus soil microbiota and nutrient cycling. In this
context, the inclusion of native-tree islands within SPS could significantly increase
phylogenetic diversity and hence the number of functional groups in a particular
landscape.
Furthermore, the change of diet resulting from the increased diversity of tannin-
rich species modifies the digestive processes of cows, reducing the amount of
methane and nitrous oxide emitted into the atmosphere, and thus their contribution
of greenhouse gas emissions (Rivera et al. 2015). Compared with conventional
extensive livestock, the iSPS generates an increase in the environmental services
that result from the greater diversity in functional traits of the different species that
are found within it. Therefore, an ecological restoration program that prioritizes
phylogenetic diversity also prevents the erosion of evolutionary history, when
species with distant recent common ancestors are purposely chosen, as in the
hypothesized case of islands of native vegetation within an iSPS (as shown in
Fig. 5.2).
In general, iSPS allow intensifying production within the parts of the property
that are most suitable for livestock, generating greater productivity and profitability,
ideally freeing up areas that have a vocation other than that of livestock production.
These areas are ideal to fulfill ecological restoration processes, becoming forests
fragments used for conservation purposes and the recovery of ecosystem services in
the long term (Perfecto and Vandermeer 2010; Giraldo et al. 2018). In the following
sections we discuss different implementation experiences and challenges related
with the establishment of native tree species either within the degraded pastures
and iSPS, or in adjacent areas released from cattle ranching and designated for
conservation. We emphasize the importance of considering plant phylogenetic
diversity as a novel tool for ecological restoration in these areas.

5.3 Experiences of Pasture Restoration in Dry and Humid

Neotropical Forests

Different restoration experiences of tropical forests in abandoned cattle pastures

have shown the complex and context-dependent factors influencing the success of
such efforts (Martínez-Garza et al. 2016). Many empirical studies provide different
approaches to using tree islands or forest patches of native vegetation within cattle
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 125

ranches to recover biodiversity and ecosystem services (Aide et al. 2000; Griscom
et al. 2005; Martínez-Ramos et al. 2012; González-Tokman et al. 2018). Whereas
most studies have been performed in abandoned pastures (Aide et al. 1994, 2000;
Martínez-Ramos et al. 2012), a few experiences show that even in active livestock
pastures, vegetation restoration is possible and is a viable way of achieving more
sustainable cattle ranching practices (Martínez-Garza and Howe 2003; de la Peña-
Domene et al. 2013; Calle et al. 2017). Next, we describe some key lessons from
experiences with ecosystem restoration in Latin America and Mexico in particular,
providing important details and challenges that must be considered when
establishing native tree islands within livestock ranches.

5.3.1 Intensity and Costs of Interventions

Depending on the site conditions, the intensity of soil degradation present in resto-
ration areas (Chazdon 2003), and on the available economic resources, restoration
interventions in degraded pastures can range from minimal, low-cost approaches to
very intensive and costly plantations (Chazdon 2003; Griscom et al. 2005; Martínez-
Garza et al. 2016). Minimal intensity interventions consist of the exclusion of
particular areas from cattle, allowing natural vegetative succession to occur (Hobbs
and Norton 1996; Aide et al. 2000). Studies of natural succession in abandoned
pastures in neotropical dry and humid forests show that, if propagule species are
available nearby, cattle exclusion and fire protection allow the establishment of early
and intermediate stages of forest succession in a range of 25–40 years, with species
richness levels equivalent to mature forests, although species enrichment can be
required to achieve the same species composition (Aide et al. 1994, 2000).
Similarly, in a tropical dry forest of Panama, Griscom et al. (2005) report that,
when excluding cattle from abandoned pastures, up to 67 species (mostly pioneer
species) were established in experimental plots, with significantly greater stem and
basal areas than plants established where cattle was not excluded. In Mexico,
chronosequence studies of abandoned pastures in both tropical dry and humid forests
show that after 20 years of cattle exclusion, different biotic communities including
plants, amphibians, reptiles, bats and birds can reach the same diversity and structure
as mature forests, although functional traits may take longer to recover (Martínez-
Ramos et al. 2012). However, these forest succession trends are strongly influenced
by the level of land degradation and distance to propagule sources (Chazdon 2003;
Martínez-Ramos et al. 2012). Although these studies suggest that restoration of
tropical forests is possible by excluding cattle and protecting target areas from
fires, recovering ecosystem services often takes much longer. Hence, incorporating
vegetation islands into sustainable cattle ranching initiatives may require some
degree of additional intervention.
An intermediate level of intervention involves the exclusion of cattle from the
areas to be restored, together with actions accelerating secondary succession. These
actions include the enrichment of restoration areas with pioneer species (Martínez-
126 R. Santos-Gally and K. Boege

Ramos and García-Orth 2007), the removal of competitor species such as grasses,
vines or ferns (Martínez-Ramos et al. 2012, González-Tokman et al. 2018), and the
protection of seeds to reduce seed predation (Martínez- Ramos 2012). For example,
in Southeastern Mexico, the enrichment of areas with early successional stages
(“acahuales”) with the pioneer species Ochroma pyramidale was reported to accel-
erate the re-establishment of different late-successional species, increasing the avail-
ability of organic matter in the soils, and the eradication of fern species arresting
natural succession (Levy et al. 2016). Other experiences have also included the
protection of isolated trees within pastures or even the introduction of artificial bird
perches to stimulate seed dispersal from forest fragments (Shiels and Walker 2003;
Laborde et al. 2008; Martínez-Garza et al. 2016). The presence of isolated fig trees in
livestock pastures in Los Tuxtlas, México, for instance, has been shown to promote
the establishment up to 73 native tree and shrub species (Laborde et al. 2008).
The most intensive level of intervention to restore ecosystem functioning is the
establishment of tree plantations of a diverse array of native plant species within
areas excluded from cattle (Martínez-Gaza et al. 2016). This strategy, although more
costly, may accelerate the processes of natural secondary succession, the recovery of
soil fertility, nutrient cycling and availability of different niches for higher trophic
levels (del-Val et al. 2016), facilitating the recovery of species interactions and
ecological processes. The establishment of plantations requires adequate species
selection, depending on the purposes of the desired ecological restoration and the
availability of native propagules in the surrounding areas. For example, if the
objective of the plantation is to accelerate the process of natural succession, planting
a combination of pioneer and late successional tree species, in particular those
dispersed by animals, may further enhance the recruitment of new species (Martí-
nez-Garza et al. 2013; de la Peña-Domene et al. 2013). If long-term carbon fixation is
of interest, species with high wood density and greater water conduction efficiency
are recommended (Martínez-Cabrera et al. 2009). Furthermore, species selection
may determine the success of these efforts, as not all tree species are adapted to the
same soil characteristics and climatic conditions. Which species to include in a
native vegetation island should consider not only the local availability of seeds or
propagules, but also previous experience and local knowledge on their propagation,
phenology, germination techniques or ability to be propagated via stakes.

5.3.2 Technical and Biological Factors Influencing

the Establishment of Vegetation Islands

The technical and biological solutions are complex and need to be adjusted for each
case, considering the costs, suitable species and the design of the areas to be
recovered. Equally important to consider are the primary objectives for which
those vegetation islands or ecological restoration plots are created. Are landholders
interested in recovering native biodiversity for ethical, cultural and/or economic
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 127

reasons? Are they aware of the abilities of vegetation islands to improve ecosystem
services, thereby enhancing the productivity of their cattle ranches? Are they
interested in including timber or fruit tree species to diversify the products coming
from their ranches? Addressing this diversity of interests is necessary for species
selection and may require the inclusion of timber or fruit trees in vegetation islands
within livestock pastures (Minnemeyer et al. 2011; Calle et al. 2012). For example,
in Colombia, the successful improvement of pastures combines the establishment of
high densities of forage species for cattle, such as Guazuma ulmifolia, with strips
(instead of vegetation islands) of native timber species such as Cordia gerascanthus
and Tabebuia rosea and the endangered Pachira quinata (Calle et al. 2012).
Once species are selected, choosing the right propagation techniques is also
relevant. Some species can be established vegetatively through stakes, whereas
others perform better when planted as seeds or saplings. For example, in Chiapas,
Mexico, a restoration experiment demonstrated that plant survivorship and perfor-
mance was increased when plants were planted as seeds or saplings, rather than
propagated from stakes (Douterlungne and Ferguson 2016). This was probably due
to greater survival of plants with pivotal roots, in contrast with stakes that need to
develop their roots.
Overall, these different ecological restoration efforts in tropical forests can
provide guidelines for the implementation of native-tree islands within cattle pas-
tures and conservation areas, in which the costs and work implicit at different level
of intensity described here must be taken into account. Most of these experiences
however, have been performed in abandoned pastures, without the challenge of
livestock exerting a continuous pressure on the excluded plots. Some strategies to
deal with this pressure are a) excluding livestock from plots with long-lasting living
fences, and b) planting buffers of forage plants to prevent livestock from entering the
plantations.

5.4 Relevance of Incorporating Phylogenetic Diversity

in Ecological Restoration

Intensive silvopastoral systems that contain vegetation islands can protect natural
ecosystems, recover biodiversity, and restore ecological functioning in landscapes
modified and fragmented by productive activities. Therefore, these systems are
useful for reducing biodiversity loss, which can lead to ecosystem degradation and
loss of ecological benefits (Perfecto and Vandermeer 2010). In addition, when
deforestation is followed by agriculture and livestock grazing, the potential for forest
regeneration is decreased. Ecological restoration projects with native tree species in
SPS in Colombia have used the focal species approach for plant species selection
(Calle et al. 2017). This includes the selection of those plant species most threatened
by degrading processes such as habitat loss and fragmentation, climate change,
alteration of biochemical cycles, or species introductions, in turn facilitating the
128 R. Santos-Gally and K. Boege

conservation of less vulnerable species (Lambeck 1997). It has been proposed that
the specific management and landscape restoration requirements for certain species
could be representative of the requirements of other species of the same or different
taxonomic groups (Lindenmayer et al. 2002). The use of vulnerable or endangered
species in SPS, as for example, Mimosa trianae and Swietenia macrophylla in
Colombia (Calle et al. 2017) has contributed to their conservation, with parallel
benefits such as the provision of habitat for birds and insects or improved micro-
habitats for the establishment of other species. Other restoration efforts have shown
that the introduction of greater plant species diversity or greater plant functional trait
diversity can enhance particular ecosystem processes and the associated ecosystem
services for human well-being (Benayas et al. 2009; Doherty et al. 2011; Montoya
et al. 2012).
There is sufficient evidence that biodiversity and the diverse functional traits
within a given environment determine the stability of ecosystem functions through
time (Cardinale et al. 2012). Furthermore, a congruence between phylogenetic
diversity and functional trait diversity has been reported in different ecological
communities (Cadotte 2019), indicating that those communities with species dis-
persed along the branches of the phylogenetic tree of life, i.e. the tree-like represen-
tation of the phylogenetic relationships that describes the evolutionary history of
Earth’s species, tend to be more functionally diverse.
Phylogenetic diversity can be viewed as a proportion of the diversity of a group of
interest that is represented within a specific ecological community. If the composi-
tion of these species includes lineages with distant common ancestors, then there is a
greater probability of finding different functional traits between the species due to a
greater degree of independent evolutionary histories. Phylogenetically distant spe-
cies tend to have dissimilar traits, whereas closely related species tend to have
similar traits and ecological niches, competing for the same resources and
responding similarly to natural selection agents (Felsenstein 1985; Díaz et al.
2013). For example, under current climate change conditions, tropical dry forests
face an increase in temperature and drought (IPCC 2007; Aguirre-Gutiérrez et al.
2020). In some tropical forest communities in West Africa, an increase in the
composition of drought tolerant species with low ratios of leaf mass to sapwood
mass (LM:SM) and higher photosynthetic rates has been observed (Fauset et al.
2012). A shift towards greater species homogeneity in landscapes due to changing
environmental conditions has been demonstrated at functional and phylogenetic
levels (Aguirre-Gutiérrez et al. 2020), highlighting the need to cover the different
components of diversity for more effective ecosystem conservation. Such homoge-
nization of species composition at different levels, including genetic variation,
phylogenetic diversity, and in particular, functional traits, can reduce the diversity
of species responses to environmental fluctuations, overall reducing community
resilience to changing climates.
Hence, species diversity and species functional traits are important to biotic
interactions and dynamics of communities (Navarro-Cano et al. 2014). Given the
intra and interspecific variation in functional traits (morphological, physiological,
structural, behavioral, biochemical, etc.) their quantification to assess their value to
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 129

ecosystem processes is not a simple task. In addition, measuring certain traits such as
the structure and size of tree roots is also challenging. Therefore, phylogenetic
diversity has the added practical advantage of being a useful proxy for functional
diversity.
Recently, Cadotte et al. (2019) found that the degree of correlation between
phylogenetic and functional diversity reported in 36 studies was positive with values
ranging from 0.4 to 1.0. The tendency for phylogenetically distant species to be
functionally disparate allows for their coexistence, improving their efficiency in the
use of limiting resources and buffering harsh abiotic conditions (Navarro-Cano et al.
2014). Studies on plant assemblages of high phylogenetic diversity have shown
higher productivity in terms of plant biomass (Cadotte 2013), higher soil microbial
productivity (Navarro-Cano et al. 2014), or decreased plant-pathogen infection range
(Gilbert and Webb 2007). Phylogenetic and functional diversity are thus two metrics
that can be used to restore diverse, functional and resilient communities (Forest et al.
2007; Verdú et al. 2012; Hipp et al. 2015; Thévenin et al. 2018; Cadotte et al. 2019).
Intraspecific genetic variation in plants is another form of diversity that can
modify ecosystem functioning. This may be relevant to ecological restoration for
its ability to modify soil properties, microbiota composition, and ultimately the role
of plant-soil feedbacks in determining plant performance (Schweitzer 2018). Indeed,
a comparison of different genotypes of Pinus pinaster from three different geo-
graphic regions showed that genetic variation determines the phylogenetic structure
of rhizosphere microbial communities. This, in turn, modulated the enzymatic
processes related to the C, N and P cycles (Pérez-Izquierdo et al. 2019). Thus,
plant genotypes affect ecosystem properties and services through the different
impacts that they can have on nutrient availability and soil fertility.

5.5 Phylogenetic Diversity in Native Tree Islands Within

Silvopastoral Systems

Genetic, functional and phylogenetic diversity are associated with increases in the
abilities of communities to adapt and respond to changes in the environment, as well
as the resilience and productivity of the ecosystem (Díaz et al. 2013). To test our
hypothesis that high phylogenetic diversity in native-tree islands within iSPS can
enhance the ecological outcomes of restoration efforts, thereby promoting a high
diversity of functional traits, we implemented an iSPS with native-tree islands within
the pastures. These vegetation islands had different degrees of phylogenetic diver-
sity, allowing for future studies to test how different plant communities influence
ecological processes and the recovery of ecosystem services. Because they were
only recently established (2018–2019) and data collection is in process, no results
regarding the influence of phylogenetic diversity on ecosystem processes and ser-
vices has yet been obtained, but here it is described how native tree islands were
implemented to inspire further restoration efforts within iSPS elsewhere.
130 R. Santos-Gally and K. Boege

We implemented a native-tree island experiment in the Mexican humid tropics

(18.552469 N,
94.998846 W) (Santos-Gally et al. 2019) within extensive cattle
pastures in a region where a high evergreen rainforest was gradually deforested
starting in the 1940s. As a result, by 1990 the region consisted mainly of a patchwork
of extensive cattle pastures with different species of native grasses (Andropogon sp.,
Digitaria sp., Lasiacis divaricata, Oplismenus burmannii, Panicum and Setaria sp.),
some introduced species (Cynodon plectostachyus. C. dactylon, Brachiaria sp.), and
few scattered native trees. The soil has a clay to clay-loam texture and is acidic to
moderately acidic (pH 5.2–5.7), probably due to the high Fe values. The land use is
mostly semi-extensive livestock farming with 0.9 cows/ha, mainly dual-purpose
zebu and zebu-swiss cow breeds. Within a 100 ha livestock farm, we chose an
area of 10 ha to develop an iSPS including ecological restoration with scattered
native-tree islands. Because soils are acidic to moderately acidic, we chose Tithonia
diversifolia, Leucaena diversifolia and Guazuma ulmifolia as the species for the
high-density shrub stratum. In the design of the selection of forage species, we
considered the rancher’s preferences, the presence of the species in natural habitats,
preference as a fodder species by the cows, high protein content (20–28%), as well as
the species’ ability to fix nitrogen and solubilize phosphorus in the soil
(L. diversifolia and T. diversifolia, respectively).
Within the cattle pasture, we established native-tree islands which had either high
or low phylogenetic diversity (HPD and LPD, measured as the sum of all the
branches that comprise each group in a time-calibrated phylogenetic tree of all
angiosperm families), with 30 plant species per island, belonging to 25 and 10 dif-
ferent plant families, respectively, according to the Angiosperm Phylogeny Group
(APG III) (Stevens 2001) classification (N ¼ 11 islands per treatment). With the
intention of accelerating plant succession in the plant community assemblage, a
greater number of tolerant species than pioneer species was selected. Within each
treatment, 20 species were shade-tolerant while 10 species were pioneers. Twelve
species were shared between treatments (six tolerant and six pioneers) and 36 species
were not shared between treatments (28 shade-tolerant and eight pioneers). In total
we selected 48 native tree species. To measure the phylogenetic diversity of each
group, we used a time-calibrated phylogenetic tree reconstruction of all angiosperms
families and some gymnosperm families as an outgroup (Fiz-Palacios et al. 2011).
Two phylogenetic trees were reconstructed according to the families included in
each treatment. Phylogenetic diversity was measured for each treatment using the
package picante (Kembel et al. 2010) in R (R Team 2018) and calculated as the sum
of all branches spanning the group of species of interest (Faith 1992). The HPD
treatment, composed of species from 25 angiosperm families, had a phylogenetic
diversity that was three times greater than the LPD treatment, with species from
10 families.
Seeds were collected from at least 3 to 5 mother trees for both phylogenetic
groups, for the purpose of increasing intraspecific genetic variation. Seeds were
germinated in a mixture of black soil and soil from around the parent tree in an effort
to incorporate parent tree’s soil microbial community. Seedlings were planted in the
experimental islands (plots) of the two phylogenetic groups between August 2018
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 131

15 m
1m

15 m
1m

Native-tree island

Fig. 5.3 (a) Map of Mexico showing the location of the study site in Sontecomapan, Veracruz; (b)
distribution of native-tree islands in Rancho Los Amigos, black symbols represent islands with high
phylogenetic diversity, white symbols those with low phylogenetic diversity. (c) Sketch of a native-
tree island representing how saplings where planted

and January 2019. The seedlings were planted once they had reached an appropriate
size for planting, determined by the phenology of the species, their germination and
growth rate. Grass was controlled several times by clearing around the saplings. The
location and type (HPD or LPD) of each 15 m2 biodiversity island within the
10-hectare paddock was assigned at random (Fig. 5.3). The islands were excluded
from cattle with two barbed wire lines.
Physicochemical and biological soil parameters including measurements of
arthropods and microbes, as well as the diversity of pollinator and invertebrate
species in the plant stratum, were recorded. These measurements were taken prior
to the establishment of the vegetation islands to determine responses to high and low
phylogenetic diversity treatments in biodiversity islands over the 10-year study.
These measurements will be compared 3, 6 and 9 years after the establishment of
the vegetation islands. In addition, the survival of seedlings, competition, litter
decomposition, growth rate, plant palatability to herbivores, seed dispersion and
pollination diversity will be compared between treatments.
Most of the results of this ecological restoration experiment within a silvopastoral
system will be obtained in coming years. Nonetheless, we present some early results
from the implementation stage. First, we found that the seeds of the species that were
132 R. Santos-Gally and K. Boege

used to implement the biodiversity islands had an average germination success of

32% (Santos-Gally et al. 2019). Survival of saplings for both HPD and LPD plots
was greater after the rainy season (96%) than after the dry season (69%). Because we
expect HPD islands will contain a greater diversity of functional traits, allowing
greater complementarity in the use of resource when compared to LPD islands, we
predict that HPD islands will present higher productivity, better decomposition of
organic matter and greater availability of nutrients, which in turn will be facilitated
by a greater diversity of microorganisms in the soil (Cadotte et al. 2008; Cadotte
2013). If the diversity of reproductive and dispersal traits increases with phyloge-
netic diversity, we expect that the diversity of pollinator and disperser species will
increase in islands with high phylogenetic diversity. Furthermore, if structural
diversity of vegetation increases in islands with HPD, then we expect greater
diversity in the recruitment of seedlings on these islands.
Previous studies have shown that seedling survival is frequently affected by
environmental variability such as that which occurs during a prolonged dry season
(Martínez-Garza et al. 2013), as well as by competition with grasses (González-
Tokman et al. 2018) or, in the case of restoration attempts within production
paddocks, livestock pressure. To avoid this pressure, we recommend isolating
restoration areas with an electric fence and connecting plots with corridors. Such
corridors could have the additional advantage of serving to separate paddocks and
provide shade for cattle (e.g. paddock division in Fig. 5.2), as well as potentially
functioning as corridors for some small vertebrates. In our case, the control of the
exotic grass Brachiaria brizantha CIAT 6780, was an important two-pronged
problem. First, because it is known that its root system competes efficiently for
water resources (Guenni et al. 2002), and second, once established as a pasture, its
eradication requires continuous clearing. Sometimes if the exotic grasses turn out to
be a very aggressive competitor with seedlings, the use of graminicides is also
suggested at this stage.

5.6 Conclusions

Intensive silvopastoral systems play a crucial role as part of the biodiversity conser-
vation agenda, especially in Neotropical regions such as Colombia, Brazil and
Mexico. Native tree islands within iSPS increase native biodiversity, serving as a
source of propagules, nesting sites, biological pest control and contributing to soil
nutrient recycling. The iSPS enriched with tree islands and/or corridors can be
envisioned as a mosaic of forest fragments and grasslands that allow for increased
vegetation cover, reestablishing ecological and evolutionary processes, and improv-
ing ecosystem services in Neotropical regions. Hence, establishing iSPS with islands
of phylogenetically and functionally diverse native vegetation can contribute to the
stability, resilience, and functioning of tropical ecosystems.
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 133

In our experience, the implementation of native tree islands within productive

pasturelands as a form of active restoration has posed several challenges. For
example, the costs related to seed collection, germination and transplanting seed-
lings, as well as the establishment of a nursery, are significant and may be
unaffordable to producers. In addition, native tree islands need to be protected
from cattle and competition from grasses such as Brachiaria. Once these challenges
are overcome, native tree islands can present an opportunity for the integration of
livestock production and biodiversity conservation in human-modified landscapes.

Acknowledgements J.R. Pérez-Ishiwara, S. Sinaca-Colin assisted with different aspects of the

work. Special thanks for F. Serrano of LANCIS- IE-UNAM for the development of tools for plant
mapping within the vegetation islands, and to M. Baltazar with computing assistance. This work
was supported by funding from the Universidad Nacional Autónoma de México (PAPIIT
IN-202419).

References

Achard F, Beuchle R, Mayaux P, Stibig H-J, Bodart C, Brink A, Donnay F, Lupi A, Carboni S,
Desclee B, Donnay F, Eva HD, Lupi A, Rasi R, Seliher R, Simonetti D (2014) Determination of
tropical deforestation rates and related carbon losses from 1990 to 2010. Glob Chang Biol 20:
2540–2554
AGAL (2008) Livestock policy briefs 03: cattle ranching and deforestation. FAO-ONU, Rome.
[Link]
Aguirre-Gutiérrez J, Malhi Y, Lewis SL, Fauset S, Adu-Bredu S, Affum-Baffoe K, Baker TR,
Gvozdevaite A, Hubau W, Moore S, Peprah T, Ziemińska K, Phillips OL, Oliveras (2020)
Long-term droughts may drive drier tropical forests towards increased functional, taxonomic
and phylogenetic homogeneity. Nat Commun 11:3346. [Link]
16973-4
Aide TM, Cavelier J (1994) Barriers to lowland tropical forest restoration in the Sierra Nevada de
Santa Marta, Colombia. Restor Ecol 2:219–229
Aide TM, Zimerman JK, Pascarella JB, Rivera LW, Humfredo M-V (2000) Forest regeneration in a
chronosequence of tropical abandoned pastures: implications for restoration ecology. Restor
Ecol 8:328–338
Altieri MA (1999) Applying agroecology to enhance productivity of peasant farming systems in
Latin America. Env Dev Sust 1:197–217
Arellano L, López-Ortiz S, Pérez-Hernández J, Alonso G, Torres-Rivera JA, Huerta C, Cruz-
Rosales M (2018) La Ganaderia Bovina Veracruzana. Hacia la sutentabilidad. In: Halffter G,
Cruz M, Huerta C. (Comps.) Ganadería sustentable en el Golfo de México. Instituto de
Ecología, A.C., México pp. 189–212
Bacab HM, Madera NB, Solorio FJ, Vera F, Marrufo DF (2013) Los sistemas silvopastoriles
intensivos con Leucaena leucocephala: una opción para la ganadería tropical. Avances Inv
Agrop 17:67–81
Baccini A et al (2012) Estimated carbon dioxide emissions from tropical deforestation improved by
carbon-density maps. Nature Clim Change 2:182–185
Bellard C, Cassey P, Blackburn TM (2016) Alien species as a driver of recent extinctions. Biol Lett
12:20150623
Benayas JMR, Newton AC, Diaz A, Bullock JM (2009) Enhancement of biodiversity and ecosys-
tem services by ecological restoration: a meta-analysis. Science 325:1121–1124
134 R. Santos-Gally and K. Boege

Cadotte MW, Cardinale BJ, Oakley TH (2008) Evolutionary history and the effect of biodiversity
on plant productivity. Proc Natl Acad Sci U S A 105:17012–17017. [Link]
pnas.0805962105
Cadotte MW (2013) Experimental evidence that evolutionarily diverse assemblages result in higher
productivity. Proc Natl Acad Sci U S A 110:8996–9000
Cadotte MW, Carboni M, Si XF, Tatsumi S (2019) Do traits and phylogeny support congruent
community diversity patterns and assembly inferences? J Ecol 107:2065–2077
Calle A (2020) Can short-term payments for ecosystem services deliver long-term tree cover
change? Ecosyst Serv 42:101084
Calle Z, Murgueitio E, Chará J (2012) Integrating forestry, sustainable cattle ranching and land-
scape restoration. Unasylva 239:31–40
Calle Z, Murgueitio E, Chará J, Molina CH, Zuluaga AF, Calle A (2013) A strategy for scaling up
intensive silvopastoral systems in Colombia. J Sustain Forest 32:677–693
Calle Z, Giraldo AM, Cardozo A, Galindo A, Murgueitio E (2017) Enhancing biodiversity in
Neotropical silvopastoral systems: use of indigenous trees and palms. In: Montagnini F
(ed) Integrating landscapes: agroforestry for biodiversity conservation and food sovereignty.
Advances agroforestry, vol 12. Springer, Dordrecht, pp 417–438
Cardinale B, Duffy JE, Gonzalez A, Hooper DU, Perrings C, Venail P (2012) Biodiversity loss and
its impact on humanity. Nature 486:59–67
Ceballos G, Ehrlich PR, Dirzo R (2017) Biological annihilation via the ongoing sixth mass
extinction signaled by vertebrate population losses and declines. Proc Natl Acad Sci U S A
114:E6089–E6096
Cerrillo E, Pastor Villegas J, Prada Gallardo A, González Cordero A, Morales Muñiz A, López Sáez
JA, López García P, Arnanz A, Juan-Treserras J, Matamala JC, Garrido García JA (2005) Bases
económicas y ambientales para el estudio de las comunidades neolíticas del centro-oeste
peninsular: perspectivas desde el yacimiento de Los Barruecos. In: Arias Cabal P., Ontañón
Peredo, R. García-Moncó, C. (Eds.), III Congreso del Neolítico en la Península Ibérica.
Monografías del Instituto Internacional de Investigaciones Prehistóricas de Cantabria
1, Universidad de Cantabria, Santander. 1056 pp
Challenger A, Soberón J (2008) Los ecosistemas terrestres. In Capital natural de México. Vol. I:
Conocimiento actual de la biodiversidad. Conabio, México, pp 87–108
Chará J, Reyes E, Peri P, Otte J, Arce E, Schneider F (2019) Silvopastoral systems and their
contribution to improved resource use and sustainable development goals: evidence from Latin
America. FAO, CIPAV and Agri Benchmark, Cali, p 60
Chazdon RL (2003) Tropical forest recovery: legacies of human impact and natural disturbances.
Perspect Plant Ecol Evol Syst 6:51–71
de la Peña-Domene M, Martínez-Garza C, Howe HF (2013) Early recruitment dynamics in tropical
restoration. Ecol Appl 23:1124–1134
del-Val E, Martínez-Garza C, Boege K, Juan-Baeza I, Solis L (2016) Restauración de poblaciones
de invertebrados e interacciones bióticas en selvas estacionales de Jalisco y Morelos. In Ceccon
E. and Martínez-Garza C. (coord.) Experiencias mexicanas en la restauración de los
ecosistemas. Universidad Nacional Autónoma De México. ISBN: 978–607–02-8157-0
Díaz S, Purvis A, Cornelissen JHC, Mace GM, Donoghue MJ, Ewers RM, Jordano P, Pearse WD
(2013) Functional traits, the phylogenyof function, and ecosystem service vulnerability. Ecol
Evol 3:2958–2975
Dirzo R, Young HS, Galetti M, Ceballos G, Isaac NJB, Collen B (2014) Defaunation in the
Anthropocene. Science 345:401–406
Douterlungne D, Ferguson BG (2016) Estrategias para el establecimiento de árboles en pastizales
para la restauración de la selva húmeda en Chiapas. In Ceccon E. and Martínez-Garza (coord.)
Experiencias mexicanas en la restauración de los ecosistemas. Universidad Nacional Autónoma
de México. ISBN: 978–607–02-8157-0
Doherty J, Callaway JC, Zedler JB (2011) Diversity–function relationships changed in a long-term
restoration experiment. Ecol Appl 21:2143–2155
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 135

Faith DP (1992) Conservation evaluation and phylogenetic diversity. Biol Conserv 61:1–10
Fauset S, Baker TR, Lewis SL, Feldpausch TR, Affum-Baffoe K, Foli EG, Hamer KC, Swaine MD
(2012) Drought-induced shifts in the floristic and functional composition of tropical forests in
Ghana. Ecol Lett 15:1120–1129
Felsenstein J (1985) Phylogenies and the comparative method. Am Nat 125:1–15
Ferraz-de-Oliveira MI, Azeda C, Pinto-Correia T (2016) Management of Montados and Dehesas for
high nature value: an interdisciplinary pathway. Agr Syst 90:1–6
Fiz-Palacios O, Schneider H, Heinrichs J, Savolainen V (2011) Diversification of land plants:
insights from a family-level phylogenetic analysis. BMC Evol Biol 11:341
Forest F, Grenyer R, Rouget M, Davies TJ, Cowling RM, Faith DP, Balmford A, Manning JC,
Procheş Ş, van der Bank M, Reeves G, Hedderson TAJ, Savolainen V (2007) Preserving the
evolutionary potential of floras in biodiversity hotspots. Nature 445:757–760
Gann G, McDonald T, Walder B, Aronson J, Nelson CR, Eisenberg C et al (2019) International
principles and standards for the practice of ecological restoration. Restor Ecol 27:S3–S46
Garrido P, Elbakidze M, Angelstam P, Plieninger T, Pulido F, Moreno G (2017) Stakeholder
perspectives of wood-pasture ecosystem services: a case study from Iberian dehesas. Land Use
Policy 60:324–333
Gilbert GS, Webb CO (2007) Phylogenetic signal in plant pathogen-host range. Proc Natl Acad Sci
U S A 104:4979–4983
Giraldo C, Escobar F, Montoya S (2018) Escarabajos del estiércol en paisajes ganaderos de
Colombia. Fundación CIPAV, Cali
Giraldo C, Escobar F, Chará J, Calle Z (2010) The adoption of silvopastoral systems promotes the
recovery of ecological processes regulated by dung beetles in the Colombian Andes. Insect
Conserv Divers 4:115–122. [Link]
González JM (2013) Costos y beneficios de un sistema silvopastoril intensivo (SSPI), con base en
Leucaena leucocephala. Estudio de caso en el municipio de Tepalcatepec, Michoacán. México
Avances Inv Agrop 17:35–50
González-Tokman D, Barradas VL, Boege K, Domínguez CA, del-Val E, Saucedo E, Martínez-
Garza C (2018) Mulching maximizes growth rates but not survival of 11 tree species in
restoration plantings in a Mexican dry forest. Restor Ecol 26:642–649
Griscom HP, Ashton PMS, Berlyn GP (2005) Seedling survival and growth of native tree species in
pastures: implications for dry tropical forest rehabilitation in Central Panama. For Ecol Manage
218:306–318
Guenni O, Marín D, Baruch Z (2002) Responses to drought of five Brachiaria species. I. Biomass
production, leaf growth, root distribution, water use and forage quality. Plant and Soil 243:229–
241
Guevara S, Laborde J, Sánchez-Ríos G (2018) El paisaje de la ganadería: una experiencia
tropical. in: Halffter G., Cruz M., Huerta C. (Comps.) Ganadería sustentable en el Golfo de
México. Instituto de Ecología, A.C., México, 432 pp
Haines-Young R, Potschin M (2010) The links between biodiversity, ecosystem services and
human Well-being. In: Raffaelli D, Frid C (eds) Ecosystem ecology: a new synthesis. Ecological
Reviews Series, Cambridge University Press, Cambridge
Harvey CA, Haber WA (1998) Remnant trees and the conservation of biodiversity in costa Rican
pastures. Agr Syst 44:37–68. [Link]
Hipp AL, Larkin DJ, Barak RS, Bowles ML, Cadotte MW, Jacobi SK, Lonsdorf E, Scharenbroch
BC, Williams E, Weiher E (2015) Phylogeny in the service of ecological restoration. Am J Bot
102:647–648
Hobbs, RJ Norton, DA (1996) Towards a conceptual framework for restoration ecology. Restor
Ecol 4: 93–110
Hoffmann M, Hilton-Taylor C, Angulo A, Bohm M, Brooks TM, Butchart SH, Carpenter KE,
Chanson J, Collen B, Cox NA et al (2010) The impact of conservation on the status of the
world’s vertebrates. Science 630:1503–1509
136 R. Santos-Gally and K. Boege

IPCC (2007) Climate change 2007: impacts, adaptation and vulnerability. Fourth Assessment
Report, Geneva
Jose S, Walter D, Mohan Kumar B (2019) Ecological considerations in sustainable silvopasture
design and management. Agr Syst 93:317–331
Kettenring KM, Mercer KL, Reinhardt Adams C, Hines J (2014) Application of genetic diversity-
ecosystem function research to ecological restoration. J Appl Ecol 51:339–348
Kembel SW, Cowan PD, Helmus MR, Cornwell WK, Morlon H, Ackerly DD, Blomberg SP, Webb
CO (2010) Picante: tools for integrating phylogenies and ecology. Bioinformatics 26:1463–
1464
Kremen C, Merenlender AM (2018) Landscapes that work for biodiversity and people. Science
362:6020
Kú VJ, Ramírez AL, Jiménez FG, Alayón AJ, Ramírez CL (1999) Árboles y arbustos para la
producción animal en el trópico mexicano. En: Sánchez MD, Rosales MM editores.
Agroforestería para la producción animal en América Latina. Roma, Italia. 231–258 pp
Laborde J, Guevara S, Sánchez-Ríos G (2008) Tree and shrub seed dispersal in pastures: the
importance of rainforest trees outside forest fragments. Écoscience 15:6–16
Lambeck RJ (1997) Focal species: a multi-species umbrella for nature conservation. Conserv Biol
11:849–856
Levy-Tacher SI, Aguirre-Rivera JR, Vleut I, Román-Dañobeytia F, Perales-Rivera H, Zúñiga-
Morales, J, et al. (2016) Experiencias y perspectivas para la rehabilitación ecológica en zonas
de amortiguamiento de las áreas naturales protegidas Montes Azules (Chiapas) y Calakmul
(Campeche). In Ceccon E. and Martínez-Garza C. (coord.) Experiencias mexicanas en la
restauración de los ecosistemas. Universidad Nacional Autónoma de México. ISBN:
978–607–02-8157-0
Lewis SL, Edwards DP, Galbraith D (2015) Increasing human dominance of tropical forests.
Science 349:827–832
Lindenmayer DB, Manning AD, Smith PL, Possingham HP, Fischer J, Oliver I, McCarthy MA
(2002) The focal-species approach and landscape restoration: a critique. Conserv Biol 16:33–
345
López-Sáez JA, López-García P, López-Merino L, Cerrillo E, González A, Prada A (2007) Origen
prehistórico de la dehesa en Extremadura: Una perspectiva paleoambiental. Rev Estudios
Extremeños 63:493–509
López-Sáez JA, Abel-Schaad D, Pérez-Díaz S, Blanco-González A, Alba-Sánchez F, Dorado M,
Ruiz-Zapata B, Gil-García MJ, Gómez-González C, Franco-Múgica F (2014) Vegetation
history, climate and human impact in the Spanish central system over the last 9000 years.
Quat Int 353:98–122
Martínez-Cabrera HI, Jones CS, Espino S, Schenk HJ (2009) Wood anatomy and wood density in
shrubs: responses to varying aridity among transcontinental transects. Am J Bot 96:1388–1398
Martínez-Garza C, Howe HF (2003) Restoring tropical diversity: beating the time tax on species
loss. J Appl Ecol 40:423–429
Martínez-Garza C, Bongers F, Poorter L (2013) Are functional traits good predictors of species
performance in restoration plantings in tropical abandoned pastures? Forest Ecol Manag 303:
35–45
Martínez-Garza C, Campo J, Ricker M, Tobón W (2016) Effect of initial soil properties on six year
growth of 15 tree species in tropical restoration plantings. Ecol Evol 6:8686–8694
Martínez-Ramos M, García-Orth X (2007) Sucesión ecológica y restauración de las selvas
húmedas. B Soc Bot Mex 80:69–84
Martínez-Ramos M, Barraza L, Balvanera P, Benítez Malvid J, Bongers F et al (2012) Manejo de
bosques tropicales: bases científicas para la conservación, restauración y aprovechamiento de
ecosistemas en paisajes rurales. Invest Ambiental 4:107–126
Martínez-Ramos M, Pingarroni A, Rodríguez-Velázquez R, Toledo-Chelala L, Zermenño-
Hernaández I, Bongers F (2016) Natural forest regeneration and ecological restoration in
human-modified tropical landscapes. Biotropica 48:745–757
5 Biodiversity Islands: The Role of Native Tree Islands Within Silvopastoral. . . 137

Minnemeyer S, Laestadius L, Sizer N, Saint-Laurent C, Potapov P (2011) A world of opportunity.

Washington, DC, World Resources Institute. Available at: [Link]/restoringforests
Montagnini F (Ed.) (2017) Integrating landscapes: agroforestry for biodiversity conservation and
food sovereignty. Advances in Agroforestry,12. Springer, Dordrecht
Montoya D, Rogers L, Memmott J (2012) Emerging perspectives in the restoration of biodiversity-
based ecosystem services. Trends Ecol Evol 27:666–672
Murgueitio E, Ibrahim M (2008) Ganadería y Medio Ambiente en américa Latina. In: Murgueitio E,
Cuartas C, Naranjo J (eds) Ganadería del futuro: Investigación para el desarrollo. Fundación
CIPAV, Cali, Colombia, p 490
Murgueitio E, Giraldo C (2009) Sistemas silvopastoriles y control de parásitos. Rev Carta Fedegan
115:60–63
Murgueitio E, Chará J, Barahona R, Rivera JE (2019) Development of sustainable cattle rearing in
silvopastoral systems in Latin America. Cuban J Agric Sci 53:1–7
Murgueitio E, Barahona R, Chará J, Flores MX, Mauricio RM, Molina JJ (2015) Los Sistemas
silvopastoriles intensivos en América Latina alternativa sostenible para enfrentar el cambio
climático en la ganadería. Cuban J Agric Sci 49:541–554
Navas A (2010) Importance of silvopastoral systems on caloric stress reduction in tropical livestock
productions. Ver Med Vet 19:113–122
Navarro-Cano JA, Goberna M, Valiente-Banuet A, Montesinos-Navarro A, García C, Verdú M
(2014) Plant phylodiversity enhances soil microbial productivity in facilitation-driven commu-
nities. Oecologia 174:909–920
Ojeniyi SO, Odedina SA, Agbede TM (2012) Soil productivity improving attributes of Mexican
sunflower (Tithonia diversifolia) and siam weed (Chromolaena odorata). Emirates J Food Agr
24:243–247
Omeja PAO, Lawes MJ, Corriveau A, Valenta KV, Sarkar D, Paim FP, Chapman CAC (2016)
Recovery of tree and mammal communities during large-scale forest regeneration in Kibale
National Park, Uganda. Biotropica 48:770–779
Paz H, Pineda-García F, Pinzón-Pérez L (2015) Root depth and morphology in response to soil
drought: comparing ecological groups along secondary succession in a tropical dry forest.
Oecologia 179:551–561
Pérez-Izquierdo L, Zabal-Aguirre M, González-Martínez SC, Buée M, Verdú M, Rincón A,
Goberna M (2019) Plant intraspecific variation modulates nutrient cycling through its below-
ground rhizo-spheric microbiome. J Ecol 107:1594–1605
Perfecto I, Vandermeer J (2010) The agroecological matrix as alternative to the landsparing/
agriculture intensification model. Proc Natl Acad Sci U S A 107:5786–5791
Pierret A, Maeght JL, Clément C, Montoroi JP, Hartmann C, Gonkhamdee S (2016) Understanding
deep roots and their functions in ecosystems: an advocacy for more unconventional research.
Ann Bot 118:621–635
R Team (2018) R: a language and environment for statistical computing. R Foundation for
Statistical Computing, Vienna, Austria
Raymond C, Fazey I, Reed M, Stringer L, Robinson G, Evely A (2010) Integrating local and
scientific knowledge for environmental management. J Environ Manage 91:1766–1777
Reed MS, Dougill AJ, Baker T (2008) Participatory indicator development: what can ecologists and
local communities learn from each other? Ecol Appl 18:1253–1269
Rivera LF, Armbrecht I, Calle Z (2013) Silvopastoral systems and ant diversity conservation in a
cattle-dominated landscape of the Colombian Andes. Agric Ecosyst Environ 181:188–194
Rivera JE, Molina IC, Donney’s G, Villegas G, Chará J, Barahona R (2015) Dinámica de
fermentación y producción de metano en dietas de sistemas silvopastoriles intensivos con
Leucaena leucocephala y sistemas convencionales orientados a la producción de leche. Livest.
Res. Rural Dev. 27: Article #76. Available: [Link]
Román-Palacios C, Wiens JJ (2020) Recent responses to climate change reveal the drivers of
species extinction and survival. Proc Natl Acad Sci U S A 117:4211–4217
138 R. Santos-Gally and K. Boege

Santos-Gally, R., Boege, K., Fornoni, F. & Domínguez, C. 2019. Ganadería sostenible en la región
de Los Tuxtlas, Veracruz, México: el equilibrio entre la producción y la conservación de la
biodiversidad. Chará, J., Peri, P. & Rivera, J., (eds). ActasIX Congreso Sistemas Silvopastoriles.
Aporte a los objetivos de desarrollo sostenible. Cali, Colombia, p. 318.
Schweitzer JA, Van Nuland M, Bailey JK (2018) Intraspecific plant–soil feedbacks link ecosystem
ecology and evolutionary biology. In: Ohgushi T, Wurst S, Johnson S (eds) Aboveground–
belowground community ecology. Springer Nature Switzerland AG, Cham, Suiza, pp 69–84
Segan DB, Murray KA, Watson JEM (2016) A global assessment of current and future biodiversity
vulnerability to habitat loss–climate change interactions. Global Ecol Conserv 5:12–21
Shiels AB, Walker LR (2003) Bird perches increase forest seeds on Puerto Rican landslides. Restor
Ecol 11:457–465
Srivastava DS, Cadotte MW, MacDonald AAM, Marushia RG, Mirotchnick N (2012) Phylogenetic
diversity and the functioning of ecosystems. Ecol Lett 15:637–648
Steege NC, Pitman A, Killeen TJ, Laurance WF, Peres CA et al (2015) Estimating the global
conservation status of more than 15,000 Amazonian tree species. Science 1:1–10
Steinfeld H, Gerber P, Wassenaar T, Castel V, Rosales M, De Haasn C (2006) Livestock’s long
shadow. Environmental issues and options. Food and Agriculture Organization of the United
Nations, Rome, p 390
Stevens PF (2001 onwards) Angiosperm Phylogeny Website. Version 14, July 2017 [and more or
less continuously updated since] [Link]
Tarbox BC, Swisher M, Calle Z, Wilson CH, Flory SL (2020) Decline in local ecological
knowledge in the Colombian Andes may constrain silvopastoral tree diversity. Restor Ecol.
[Link]
Thévenin C, Mouchet M, Robert A, Kerbiriou C, Sarrazin F (2018) Reintroductions of birds and
mammals involve evolutionarily distinct species at the regional scale. Proc Natl Acad Sci U S A
115:3404–3409
Verdú M, Gómez-Aparicio L, Valiente-Banuet V (2012) Phylogenetic relatedness as a tool in
restoration ecology: a meta-analysis. Proc Biol Sci 279:176–1767
Violle C, Navas ML, Vile D, Kazakou E, Fortunel C, Hummel I, Garnier E (2007) Let the concept
of trait be functional! Oikos 116:882–892
Watson JEM, Evans T, Venter O et al (2018) The exceptional value of intact forest ecosystems. Nat
Ecol Evol 2:599–610. [Link]
Chapter 6
Riparian Forests: Longitudinal Biodiversity
Islands in Agricultural Landscapes

Lina Paola Giraldo, Julián Chará, Zoraida Calle D,

and Ana M. Chará-Serna

Abstract Riparian buffers safeguard the only remaining forest fragments in many
agricultural landscapes of the Colombian Andean region. These linear landscape
elements contribute to the conservation of terrestrial biodiversity in agricultural
landscapes by providing shelter, reproduction sites, food, and connectivity for
arthropods, amphibians, mammals and birds. Thus, riparian buffers play a critical
role as biodiversity islands. In addition, forested riparian buffers protect aquatic
environments and water quality by reducing the input of pollutants from catchment
areas, improving physical habitat with shade, and adding allochthonous materials
that provide the main source of energy for stream ecosystems. This chapter summa-
rizes the results of research conducted during the past two decades by the Center for
Research in Sustainable Agricultural Systems (CIPAV) in the Central Andes coffee-
growing region of Colombia. These studies highlight the critical role of forested
riparian buffers for conservation and ecosystem services. We provide a synthesis of
lessons learned on the effects of both cattle grazing and riparian forest cover on
stream ecosystems. This body of research also demonstrates that streams protected
by riparian forests support complex and biodiverse macroinvertebrate assemblages
and may respond positively to the ecological restoration of riparian strips. The
chapter concludes with recommendations for restoring and protecting riparian
buffers from agricultural practices, partially through incentives to landowners.
These insights have emerged from decades of research and institutional experience
on riparian restoration initiatives.

L. P. Giraldo (*)
Centro para la Investigación en Sistemas Sostenibles de Producción Agropecuaria (CIPAV),
Cali, Colombia
Instituto de Biología, Universidad de Antioquia, Medellín, Colombia
e-mail: lina@[Link]
J. Chará · Zoraida Calle D · A. M. Chará-Serna
Centro para la Investigación en Sistemas Sostenibles de Producción Agropecuaria (CIPAV),
Cali, Colombia
e-mail: julian@[Link]; zoraida@[Link]; ana@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 139
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
140 L. P. Giraldo et al.

Keywords Andean streams · Colombian Andes · Ecological restoration · Riparian

biodiversity · Riparian buffers · La Vieja River

6.1 Introduction

Forested riparian buffers (also known as riparian forests) are strips of vegetation that
grow along rivers and streams, and around springs and wetlands. These longitudinal
elements, distributed along the water network, act as biodiversity islands, connecting
forest fragments, conserving water sources, and providing environmental benefits to
adjacent agroecosystems (Ericsson and Stevens 1996; Naiman et al. 2000; Schroth
et al. 2004; Lees and Peres 2007; Palmer et al. 2014; Luke et al. 2019).
In Colombia, forested riparian buffers, which have an average width of 24 m, are
often the only patches of woody vegetation that remain in many agricultural land-
scapes. Although forested riparian buffers often occupy small land areas, they make
a disproportionate contribution to the landscape-scale conservation of birds, arthro-
pods, and other organisms that provide essential services such as biological pest
control, seed dispersal, pollination and carbon sequestration (Schroth et al. 2004;
Marczak et al. 2010).
However, despite their importance, these forest strips are being destroyed and
replaced with pastures or cropland, which has had negative effects on aquatic
environments, water quality, and terrestrial biodiversity (Braccia and Voshell
2007; Chará et al. 2007; Riseng et al. 2011; Skłodowski et al. 2014). This is the
case of the Central Andes coffee-growing region of Colombia, where many forests,
including riparian corridors, were replaced with coffee or banana plantations and
pastures during the second half of the twentieth century (Sadeghian et al. 1999). In
some areas of this region, land cover transformation has been successfully reversed
through restoration projects carried out by local farmers and the Center for Research
in Sustainable Agricultural Systems (CIPAV, an autonomous Colombian organiza-
tion with 35 years of experience in research, training and outreach on sustainable
agricultural production systems; Calle 2020). Several of the restoration projects
developed by CIPAV in this coffee-growing region have focused on the implemen-
tation of environmentally friendly agroforestry and silvopastoral systems and the
release of riparian areas for forest restoration (Calle 2020).
This chapter synthesizes the findings of research conducted for two decades along
with restoration projects in this coffee-growing region. These studies evaluated the
role of riparian forests in the protection of terrestrial biodiversity and aquatic
environments by monitoring the results of several restoration initiatives focused on
these key landscape elements. The chapter ends with a synthesis of the lessons
learned from these studies, together with recommendations that can be applied to
riparian restoration.
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 141

6.1.1 Study Area

The region of central-western Colombia known as the “Eje Cafetero” (coffee-

growing region) includes a variety of ecosystems of the Central Andes (Cordillera
Central), from lowland rainforests to snow-capped mountains. Its high biodiversity
and unique ecosystems are threatened by landscape transformation to establish
pastures or crops. For this reason, it has been recognized internationally as a
conservation priority (CARDER-FONADE 2002; Uribe-Gómez 2008). In the
1990s, many coffee growers in the region eliminated their plantations as a result of
the economic instability triggered by the fall of international coffee prices, making
pastures for bovine livestock become the dominant land use at elevations between
1200 and 1800 m (Sadeghian et al. 1999). The adoption and management of
livestock grazing systems have changed the species composition and vegetation
structure of riparian buffers in this region (Chará-Serna et al. 2015; Fig. 6.1).
CIPAV studies were done at La Vieja river basin in the Central Andes coffee-
growing region (Fig. 6.2). These watersheds are located on rolling hills and valleys,
at altitudes between 900 and 2400 m, with average annual rainfall of 1900–2600 mm
and mean temperature between 12 and 24
C, varying with elevation. Rainfall
exhibits a bimodal seasonality, with two annual periods of high precipitation
(April–May and October–November). The studies focused on the effects of agricul-
tural systems (mainly cattle ranching) on headwater micro-basins (<100 ha), the role
of forested riparian buffers in mitigating these effects on aquatic environments, and
the conservation of terrestrial biodiversity.

Fig. 6.1 Riparian forest in a cattle ranching landscape, coffee-growing region of Colombia. (Photo:
Julián Chará)
142 L. P. Giraldo et al.

Fig. 6.2 Study area in the Central Andes coffee-growing region, Colombia. (Map: Julián Mendivil)

Close to 20% of the land in La Vieja river basin is covered by secondary, mature
and riparian forests. The most species-rich botanical families in this area are
Lauraceae, Rubiaceae, Moraceae, Euphorbiaceae and Fabaceae. Abundant species
in riparian forests include Ocotea sp., Calliandra pittieri, Miconia sp., Cordia
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 143

alliodora, Guadua angustifolia, Cupania americana, Sorocea trophoides,

Oreopanax cecropifolius, Piper crassinervium, Anacardium excelsum, Cecropia
angustifolia, Croton magdalenensis, Heliconia platystachys, Brosimum alicastrum,
Aiphanes horrida, Cinnamomum triplinerve, Guarea guidonia and Urera
caracasana (Méndez and Calle 2010). A large portion of this species richness is
confined to riparian forests, in contrast with the agricultural matrix, where the
diversity of woody plants is very low (Méndez and Calle 2010).

6.2 Riparian Forests and Biodiversity

Riparian forests link terrestrial and aquatic ecosystems through processes that occur
at different spatial scales. At the watershed scale, these forests contribute to ecolog-
ical functions that depend on species movement or landscape connectivity. On a
local scale, riparian forests provide organic matter and shade to streams, maintain
slope stability, and protect stream beds, thereby determining habitat quality and
aquatic biodiversity.

6.2.1 Terrestrial Biodiversity (Regional or Watershed Scale)

Riparian forests protect some of the most biodiverse and dynamic ecosystems on the
planet (Naiman et al. 2005). Their vegetation may include unique species assem-
blages and sustain animal populations that depend on these ecosystems for shelter,
reproduction, food and passage (Moore and Richardson 2003; Sabo et al. 2005).
Studies on mammals, birds and dung beetles have shown that forested riparian
buffers support more terrestrial biodiversity than the surrounding agricultural matrix
(Fajardo et al. 2009; Gray et al. 2014; Zimbres et al. 2017; Luke et al. 2019). These
ecosystems are biodiversity islands because they are often the last forest remnants in
agricultural landscapes. In this context, their conservation and restoration help
mitigate biodiversity loss and habitat fragmentation (Lees and Peres 2007). Due to
their linear configuration, riparian forests act as biological corridors, connecting
forest patches in fragmented landscapes, and facilitating migration and dispersal of
birds, mammals, reptiles, amphibians, insects and other organisms that provide key
ecosystem services (Naiman et al. 2005; Medina et al. 2007; Gray et al. 2017).
Studies conducted in agricultural landscapes of the Colombian coffee-growing
region have found that compared to all other types of ecosystems, forested riparian
buffers and forest fragments have the most structurally complex and species-rich
vegetation (Table 6.1). Out of 390 woody plant species known to exist in the
agricultural basin of La Vieja river, 278 species (71%) were found in riparian forests
(Calle and Méndez 2009), where the dominant species include Guadua angustifolia
(bamboo), Cupania americana, Sorocea trophoides, Oreopanax cecropifolius,
Piper crassinervium, Anacardium excelsum and Cecropia angustifolia.
144 L. P. Giraldo et al.

Table 6.1 Bird, woody plant and dung beetle species richness in riparian forests and other land
uses in cattle farms of the Central Andes coffee-growing region (CSCRP: Colombian Sustainable
Cattle Ranching Project) ([Link]
Woody plant species Bird species
richness richness Woody plant species
(Calle and Méndez (Fajardo et al. richness
Land use 2009) 2009) (CSCRP)
Riparian forest 278 103 183
Secondary and mature forest 264 92 199
Secondary growth areas – 88 –
Bamboo forest (Guadua 89 – –
angustifolia)
Agriculture 86 – –
Scattered trees in paddocks – – 15
Live fences – – 25
Intensive silvopastoral – – 18
systems
Enhanced treeless pasture – 45 1
Natural treeless pasture 102 38 –

More recent studies done in this coffee-growing region by the Colombian Sus-
tainable Cattle Ranching Project ([Link] found a
high diversity of plants in riparian forests (unpublished data; Table 6.1). Some
threatened and scarce tree species were found in riparian forests, including Cedrela
odorata, Swietenia macrophylla, Podocarpus oleifolius, Anacardium excelsum and
Astronium graveolens.
Additionally, compared to other landscape elements and land uses in the region,
riparian forests in this basin showed the highest bird species richness (103 of the
229 species in the landscape, 45%), and contained most of the 41 bird species of
global conservation concern recorded in the region (Fajardo et al. 2009) (Table 6.1).
Endemic and nearly endemic bird species observed in this study include the ‘grayish
piculet’ (Picumnus granadensis), ‘flamerumped tanager’ (Ramphocelus
flammigerus), ‘apical flycatcher’ (Myiarchus apicalis), ‘crested ant-tanager’
(Habia cristata), ‘bar-crested antshrike’ (Thamnophilus multistriatus), ‘scrub tana-
ger’ (Tangara vitriolina) and ‘grasshopper sparrow’ (Ammodramus savannarum).

6.2.2 Aquatic Biodiversity (Local Scale)

Forested riparian buffers mitigate the impact of agricultural activities on aquatic

ecosystems through different mechanisms. Riparian vegetation filters and retains
sediments, organic matter, nutrients, chemical substances and pathogens released
from the catchment area, preventing them from entering aquatic ecosystems. Fur-
thermore, tree shade reduces fluctuations in water temperature, and the roots of dense
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 145

Fig. 6.3 Stream protected by a riparian forest on a cattle farm in the coffee-growing region,
Colombia. (Photo: Carlos Pineda)

vegetation stabilize riverbanks, protecting them from erosion (Osborne and Kovacic
1993; Mingoti and Vettorazzi 2011; Schilling and Jacobson 2014; Tanaka et al.
2016) (Fig. 6.3). Together, these mechanisms enhance hydrological regulation,
improve water quality and contribute to the conservation of aquatic biodiversity.
Studies done in agricultural landscapes of the Central Andes coffee-growing
region have shown that headwater streams protected with riparian forests often
contain a considerable diversity of aquatic macroinvertebrates (Chará et al. 2007;
Giraldo et al. 2014; Villada et al. 2017; Ramírez et al. 2018). This biodiversity is also
related to water quality and characteristics of the streambed such as the abundance of
stones (Table 6.2). Macroinvertebrate orders like Ephemeroptera, Plecoptera and
Trichoptera (also known as EPT taxa) play an important role in processing leaf litter
contributed by the riparian vegetation to the aquatic environment and are particularly
sensitive to habitat alteration. Therefore, they are considered bioindicators of con-
served ecosystems. Up to 77% of the families and 42% of the genera of Trichoptera
reported for Colombia were found to be associated with forested riparian buffers
(Ascúntar et al. 2014).
Recent studies of small streams protected by riparian forests within agricultural
landscapes have expanded the known distributions of several species of the orders
Trichoptera, Plecoptera and Coleoptera in Colombia (Zúñiga et al. 2014, 2015;
González-Córdoba et al. 2015, 2016). Additionally, research in these small ecosys-
tems has resulted in the discovery and description of new aquatic insect species for
the country (Molineri et al. 2016). These findings support the value of small streams
as unexpected reservoirs of biodiversity in agricultural landscapes.
146 L. P. Giraldo et al.

Table 6.2 Mean values of physical and biological variables in watersheds of the Central Andes
coffee-growing region of Colombia. The impact estimate is the arithmetic difference between
watersheds with forested and pasture-dominated riparian buffers. Based on Chará et al. (2007),
Giraldo et al. (2014), Villada et al. (2017), Ramírez et al. (2018) and summarized in Giraldo (2019)
Watersheds with
forested riparian Watersheds with pasture- Impact
buffers dominated riparian buffers estimate
Variable n ¼ 24 n ¼ 30 (%)
Width of streambed 2.3 4.2 82.6 (+)
(bank to bank) (m)
Depth (cm) 17.5 13.1 25.1 ()
% of rocks 67 13 80.5 ()
% of mud 18 61 238.8 (+)
Macroinvertebrates
Mean abundance 751.8 2811.2 274 (+)
Richness 83 72 13.2 ()
% EPTa 36.1 4.2 88.3 ()
% Diptera 25.5 42.5 66.6 (+)
% Mollusca 9.7 42.2 335 (+)
Water quality
Temperature (
C) 18.4 21.8 18.4 (+)
Total solids (mg L1) 85.5 146.8 71.6 (+)
Total suspended 9.7 139 1332.9 (+)
solids (mg L1)
BOD 5-20
C (mg. 2.3 6.2 169.5 (+)
L1O2)
Ammonia nitrogen 0.47 0.67 42.5 (+)
(mg.L1 N-NH3)
Dissolved oxygen 6.1 4.3 29.5 ()
(mg.L1)
Fecal coliforms 1596.3 36200.6 2167.7 (+)
(MPN. 100 mL1)
a
Ephemeroptera, Plecoptera, Trichoptera
BOD biochemical oxygen demand, MPN most probable number

Similar studies have shown that the elimination of riparian forests often triggers
severe changes in the composition of the aquatic fauna, such as a loss of diversity of
sensitive EPT taxa, and an increase in the abundance of groups that are tolerant to
organic pollution, such as Diptera and Mollusca (Chará et al. 2007; Giraldo et al.
2014; Ramírez et al. 2018) (Table 6.2). Agricultural practices have also been shown
to affect aquatic macroinvertebrates indirectly, by increasing nitrogen concentrations
and reducing the width of forested riparian strips. These alterations reduce habitat
quality for aquatic fauna by limiting the availability of coarse substrates within
stream channels (Chará-Serna et al. 2015).
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 147

Riparian forests provide large quantities of coarse particulate organic material

(leaves, flowers, fruits, branches) to the aquatic environment. These organic inputs
are the primary source of energy for food webs in forested headwater streams, where
the closed canopy limits light availability and primary productivity (Hynes 1975;
Vannote et al. 1980; Wallace et al. 1997). The processing of allochthonous inputs of
organic matter initiates the transfer of energy that then flows through the aquatic
food web in these ecosystems (Jones 1997; Wallace et al. 1997; Lamberti and
Gregory 2006; Aldridge et al. 2009; Yoshimura 2012). Coarse plant material also
maintains aquatic biodiversity by providing habitats and physical structures that are
used by fauna (Suurkuukka et al. 2014). In a comparative study of protected and
unprotected streams, Giraldo (2019) found a greater abundance (584 vs. 40 individ-
uals), richness (10 vs. 5 genera) and biomass (3.6 vs. 0.35 g) of leaf-processing
macroinvertebrates per grasp in streams with riparian forests.
In addition, riparian vegetation offers shelter for emerging adult stages of aquatic
insects, which may not be able to fly far and constitute an important food source for
birds, bats, amphibians and other insectivorous species. Riparian forests provide
habitat for other species of arthropods, mollusks, crustaceans, and small fish that are
consumed by terrestrial organisms, contributing to the transfer of energy between
terrestrial and aquatic ecosystems (Paetzold et al. 2005).

6.3 Impacts of the Loss of Riparian Forests on Streams

Even though riparian forests perform essential functions related to the protection of
water quality and biodiversity, they have been highly impacted around the world
(Kuglerová et al. 2014). Farmers often remove riparian forests to establish pastures
and crops because the riparian forest soils are richer in nutrients than the surrounding
areas (Naiman et al. 2005). Replacing native forests with pastures or crops leads to
several negative effects, including increased inputs of sediments and pollutants to
water sources, reduced water regulation capacity, and biodiversity loss (Duehr and
Siepker 2006; Chará et al. 2007; Lorion and Kennedy 2009; Turunen et al. 2019).
Table 6.2 summarizes several differences in physical, biological and water quality
variables between water sources with and without the protection of the riparian
vegetation. Unprotected sites tend to have shallower water, lower proportions of
coarse substrates, lower species richness, higher abundance of organisms and higher
values in parameters such as water temperature, solids, biochemical oxygen demand
(BOD), nitrogen, and fecal coliforms.
The following list presents some lessons learned about the effects of livestock and
riparian forests on headwaters of the Central Andes coffee-growing region
(described in detail in Giraldo (2019), based on studies of water quality, habitat
quality, aquatic macroinvertebrates and the flow of coarse particulate organic matter
(Chará et al. 2007, 2011; Camargo et al. 2011; Giraldo et al. 2014; Chará-Serna et al.
2015; Galindo et al. 2017; Ramírez et al. 2018; Giraldo 2019).
148 L. P. Giraldo et al.

• Cattle grazing in catchment areas causes undesirable effects such as soil com-
paction, reduced infiltration capacity, varying degrees of erosion, and the loss of
forests that protect streams. Comparative studies of soils under two types of
riparian vegetation carried out in the area have found that soils had lower apparent
density (0.7 vs. 0.9 g/cm3), higher total porosity (70% vs. 60%) and lower
susceptibility to compaction (85% vs. 88.3%) in bamboo (Guadua angustifolia)
riparian forests than in pastures (Camargo et al. 2011).
• The degradation or removal of riparian forests reduces canopy cover and shade.
Vegetation structure and composition become simplified as plant covers domi-
nated by grasses and pioneer shrubs (mostly Piperaceae and Melastomataceae)
replace more diverse woody vegetation.
• Although woody plants from nearby areas continue to disperse their seeds to
degraded riparian strips, the vigorous growth of grasses may temporarily inhibit
the establishment of trees and shrubs. In sites without restoration treatments,
pastures can cover up to 52% of the area (Galindo et al. 2017).
• The loss of riparian forests and their buffering services amplifies the negative
effects of grazing on watersheds. Without shade, water temperature, organic
matter, nutrients and pathogens increase while dissolved oxygen decreases
(Chará et al. 2007; Giraldo et al. 2014). Each of these changes implies a loss of
water quality with negative consequences for nearby human populations in
addition to local species.
• The removal of riparian woody vegetation facilitates the direct access of cattle to
streambeds. Without the strong roots that stabilize stream margins, cattle tram-
pling rapidly deteriorates banks and slopes.
• Damage to the banks accelerates erosion and sedimentation of the streambed and
changes channel morphology. The average width of the bed in unprotected
streams is 5.4 m, compared to 2.2 m in sites protected by riparian forests
(Chará et al. 2007).
• Streams where riparian forests have been eliminated and cattle have direct access
to the channel tend to be shallower than protected streams, with a significant
fraction of coarse substrates being replaced by fine sediments such as silt and
sand. In cattle areas, up to 100% of the riverbed of unprotected streams can
become covered by very fine substrates (Giraldo et al. 2014).
• The loss of riparian forest reduces the inputs of wood, litter, and other coarse
materials in streams. Fallen organic matter forms important microhabitats such as
pools and small turbulences, provides colonization substrates for organisms, and
is an essential source of energy for macroinvertebrates. Pools occupy a smaller
proportion of the area in streams impacted by livestock activities than in those
protected by riparian forests (13% vs. 46%, respectively; Chará et al. 2007).
• Lower water quality and modified physical conditions of streams cause changes
in macroinvertebrate communities. In these circumstances, groups that tolerate
habitat degradation, such as mollusks (Physidae) and dipterans (mainly of the
Chironomidae and Simuliidae families) tend to increase in abundance and dom-
inance, but the overall richness of species, families, and orders tends to decrease.
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 149

• On average, streams protected by riparian forests receive 7.3 times more leaf litter
per year than unprotected streams (9150 kg vs. 1255 kg per hectare of woody
vegetation, respectively; Giraldo 2019).
• Leaves form the largest proportion of litter that enters forested streams. However,
in buffer strips covered by grasses, a qualitative change occurs in the composition
of accumulated material when the inputs of wood, flowers and fruits are lost. Due
to the lack of logs that form pools and structures that retain materials on the
stream bed, the rate of storage can be four times lower in streams with grasses
(Giraldo 2019).
• Although the studied watersheds are immersed in agricultural landscapes and
occupy relatively small areas (<100 ha), their aquatic ecosystems harbor impres-
sive biodiversity, represented mainly by macroinvertebrates. The conservation of
these watersheds is essential to protect this biota. Small watersheds are also the
main sources of water for rural communities, so their conservation is also critical
from a public health perspective.

6.4 Restoration of Riparian Forests in Agricultural

Landscapes

Riparian restoration should start by guaranteeing the protection of existing forest

remnants and improving connectivity between the upper and lower sections of
watersheds. When restoring riparian forests, it is useful to define a set of clear
objectives or reference conditions in terms of ideal forest structure and composition.
The selection of restoration techniques will depend on the specific conditions of each
site, including the characteristics of the remnant vegetation, soil conditions, and the
proximity to seed sources (e.g., other forest fragments).
The restoration of riparian areas in agricultural landscapes should be prioritized,
facilitating the gradual reestablishment of woody vegetation and its associated
ecological functions in terrestrial and aquatic ecosystems (Meli et al. 2019). Certain
characteristics of restored riparian forests, such as their width, length, and vegetation
structure, determine the magnitude of the environmental benefits of restoration, such
as reductions in nutrient cycling rates, the protection of aquatic environments, and
landscape-scale biodiversity conservation.
Fencing riparian strips is one of the most popular methods to initiate the restora-
tion of riparian forests in agricultural landscapes. Fencing has been shown to
enhance the natural regeneration of riparian vegetation by preventing the access of
cattle to riparian areas. For example, riparian strips that had been protected from
grazing during the last 10 to 14 years in La Vieja river basin had recovered a similar
assemblage of dominant species as reference forest ecosystems in the area, including
common species of trees such as Cupania americana, Inga edulis, Cecropia
angustifolia and Croton magdalenensis, and some locally threatened species such
as Anacardium excelsum, Oreopanax cecropifolius, Trichilia pallida, Aiphanes
horrida, Nectandra turbacensis, Ocotea macropoda and Machaerium capote
150 L. P. Giraldo et al.

Table 6.3 Vegetation structure in reference sites and riparian buffers undergoing restoration at La
Vieja river basin (Calle and Holl 2019)
Reference Restored
Average tree species density 8.5 species ha1 19 species ha1
Density of tree stems 300 stems ha1 750 stems ha1
Basal area 8 m2 ha1 14.6 m2 ha1
Average canopy cover Not available 89%
Grass cover Not available < 5%

(Calle and Holl 2019). Additionally, restored areas had more species, a higher
density of tree stems, higher canopy cover and lower grass cover than reference
forests (Table 6.3).
Different factors may slow down or prevent the spontaneous regeneration of
woody species in fenced riparian buffers. A frequent issue in deforested cattle
ranching watersheds is the uncontrolled growth of grasses on the riparian strips.
Dense grass growth may inhibit the regeneration of shrubs and trees, even after the
removal of grazing, temporarily halting secondary succession. Techniques of
assisted natural regeneration, such as the periodic control of competing plants and
the enrichment planting of pioneer trees, can be used to accelerate forest recovery.
Fast-growing shrubs can be planted to shade out grasses, slow their growth and
facilitate the regeneration of woody plants, offsetting the inhibitory effects of grass
growth. For example, Galindo et al. (2017) studied the effect of Tithonia diversifolia
and Piper auritum planted at high-density to shade the grasses and facilitate the
establishment of native trees. After 15 months, T. diversifolia was able to reduce
grass cover by 81% and enhanced the survivorship of native trees planted
underneath.
Riparian restoration efforts can also have beneficial effects on aquatic environ-
ments. A recent study of several cattle ranching watersheds in the Andean region of
Colombia showed that the early growth of native vegetation in riparian strips
enhances the chemical and biological properties of aquatic ecosystems (Giraldo
et al. 2020). The biochemical oxygen demand (BOD), which measures organic
water pollution, was significantly lower in the studied streams 36 months after the
beginning of riparian restoration activities. Similarly, restored streams showed an
increase in dissolved oxygen, as well as a decrease in turbidity and fecal coliforms.
Regarding the composition of biological communities, the relative abundance of
tolerant aquatic insects of the family Chironomidae (Diptera) significantly decreased
through the 3 years of sampling, whereas the abundance of the family
Hydropsychidae (Trichoptera) showed moderate increases. Together these
bio-indicators suggest that the stream ecosystem is being restored.
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 151

6.5 Conclusions and Recommendations

Given the strategic value of riparian forests, all initiatives to protect and restore these
landscape elements and their ecosystems should be designed with the complete basin
in mind. The following recommendations are based on lessons learned through
various studies carried out by CIPAV in Colombian cattle ranching landscapes
(Murgueitio and Ibrahim 2009; Chará and Giraldo 2011; Calle et al. 2012; Chará
et al. 2018; Giraldo 2019).

6.5.1 Watershed Scale

• Conservation initiatives for riparian areas and aquatic environments should

prioritize landscape elements that can gradually reestablish some ecological
functions of forests. Tree coverage should be increased not only in stream
banks, but in the entire catchment areas.
• The adoption of silvopastoral systems, agroforestry systems, strategically placed
live fences, windbreaks, and other tree-based elements promote the recovery of
soil, biodiversity, hydrologic regulation, microclimate, natural biological pest
control, and carbon sequestration.
• Agrochemicals should be reduced gradually and finally eliminated in pastures
and cropland. Veterinary medicines should be used more sparingly to reduce
water pollution.
• Solid wastes and urine from livestock facilities should be treated with agroeco-
logical practices that enhance nutrient recycling in farming systems, such as water
decontamination with biodigesters and the production of compost.

6.5.2 Stream Segment Scale

• A necessary first step when restoring forest cover in heavily eroded or degraded
riparian strips is the restriction of livestock access to the streams.
• Drinking stations must be provided at each paddock to prevent cattle from
entering into streams.
• One key action when conserving riparian forests within agricultural landscapes is
fencing of existing forest fragments and guaranteeing their effective protection.
• After fencing the riparian strip, the process of assisted natural regeneration
involves controlling competing plants and enrichment planting with native spe-
cies. Together, these actions promote the recovery of aquatic environments.
• Riparian forest restoration accelerates the recovery of aquatic biodiversity and
key ecological functions, such as the processing and transfer of leaf litter within
the ecosystem.
152 L. P. Giraldo et al.

• An important principle that should guide the restoration of riparian forests is the
provision of heterogeneous leaf litter to the aquatic ecosystem. Planted and
regenerating species should provide thick and thin, small and large leaves, and
both fast and slow decaying litter. Since the organisms that contribute to the
fragmentation of leaf litter require some palatable, nutrient dense food resources,
tree species that provide such attractive resources for aquatic organisms should be
included in the restoration treatments.
• Trees that contribute significant amounts of litter to the soil of the riparian areas
will be attractive to edaphic macrofauna and will accelerate the recovery of the
stream segment’s hydrological properties.
• The richness and singularity of entomofauna in Andean stream ecosystems prove
that forested riparian buffers function as biodiversity islands and support efforts
to restore these landscape elements.

6.5.3 Social Issues

• The main environmental service provided by riparian forests in headwaters is the

maintenance of water quality for rural populations, urban areas, and agriculture.
Clear measures are required to achieve their conservation and recovery in order to
ensure their productive use.
• The vulnerability of riparian forests is a direct result of conservation and man-
agement decisions made by landowners. Preserving these landscape elements,
their ecosystems, and their myriad services requires an increasing willingness of
farmers to adopt environmentally friendly and sustainable agricultural practices
and undertake the restoration of areas critical for biodiversity. Colombia and
other Latin American countries need clear policies for headwater conservation.
These policies must guarantee the sustainable use of resources and the continuity
of environmental services.
• Incentives such as payment for environmental services and discounts in property
taxes motivate farmers to conserve and restore forest remnants adjacent to aquatic
environments.

Acknowledgements The authors gratefully acknowledge the support of Minciencias and the
Colombian Autonomous Fund for Science, Technology and Innovation Francisco José de Caldas
through contract 80740-006-2020, and the support to L.P. Giraldo through the scholarship for
doctoral studies, call 567.
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 153

References

Aldridge KT, Brookes JD, Ganf GG (2009) Rehabilitation of stream ecosystem functions through
the reintroduction of coarse particulate organic matter. Restor Ecol 17:97–106
Ascúntar O, Zuñiga MDC, Giraldo LP, Ramírez Y, Chará J, Ramos B (2014) Contribución al
conocimiento de los tricópteros adultos (Insecta: Trichoptera) de los andes orientales de
Colombia. In: La biodiversidad sensible: patrimonio natural irreemplazable. IV Congreso
Colombiano de Zoología. Diciembre 1–5 de 2014. Libro de resúmenes. Asociación Colombiana
de Zoología. Disponible en línea: [Link]; www.
[Link]
Braccia A, Voshell R (2007) Benthic macroinvertebrate responses to increasing levels of cattle
grazing in blue Ridge Mountain streams, Virginia, USA. Environ Monit Assess 131:185–200
Calle A (2020) Can short-term payments for ecosystem services deliver long-term tree cover
change? Ecosyst Serv 42:101084
Calle A, Holl KD (2019) Riparian forest recovery following a decade of cattle exclusion in the
Colombian Andes. For Ecol Manag 452:117563
Calle Z, Méndez LE (2009) Estructura y Composición de la Vegetación Arbórea en el Agropaisaje
del Río La Vieja. In: Rodriguez JM, Camargo JC, Niño J, Pineda AM, Arias LM, Echeverry
MA, Miranda CL (eds) Valoración de la Biodiversidad en la Ecorregión del Eje Cafetero.
CIEBEREG, Pereira, pp 185–192
Calle Z, Murgueitio E, Chará JD (2012) Integrating forestry, sustainable cattle ranching and
landscape restoration. Unasylva 239:31–40
Camargo JC, Chará JD, Giraldo LP, Chará-Serna AM, Pedraza GX (2011) Beneficios de los
corredores ribereños de Guadua angustifolia en la protección de ambientes acuáticos en la
Ecorregión Cafetera de Colombia. 1. Efectos sobre las propiedades del suelo. Recursos
Naturales y Ambiente 61:53–59
CARDER-FONADE (Ministerio del Medio Ambiente) (2002) Ecorregión Eje cafetero: un territorio
de oportunidades. Corporación Alma Mater, Pereira
Chará JD, Giraldo C (2011) Servicios ambientales de la biodiversidad en paisajes agropecuarios.
Fundación CIPAV, Cali, 76p
Chará JD, Pedraza G, Giraldo LP, Hincapié D (2007) Efecto de corredores ribereños sobre el estado
de quebradas en la zona ganadera del río La Vieja, Colombia. RAFA 45:72–78
Chará JD, Giraldo LP, Chará-Serna AM, Pedraza GX (2011) Beneficios de los corredores ribereños
de Guadua angustifolia en la protección de ambientes acuáticos en la Ecorregión Cafetera de
Colombia. 2. Efectos sobre la escorrentía y captura de nutrientes. Revista Recursos Naturales y
Ambiente 61:60–66
Chará J, Reyes E, Peri P, Otte J, Arce E, Schneider F (2018) Silvopastoral systems and their
contribution to improved resource use and sustainable development goals: evidence from Latin
America. FAO, CIPAV. Editorial CIPAV, Cali, 58p. Licence: CC BY-NC-SA 3.0 IGO. isbn:
978-958-9386-84-2
Chará-Serna AM, Chará JD, Giraldo LP, Zúñiga MDC, Allan JD (2015) Understanding the impacts
of agriculture on Andean stream ecosystems of Colombia: a causal analysis using aquatic
macroinvertebrates as indicators of biological integrity. Freshwater Sci 34:727–740
Duehr JP, Siepker MJ (2006) Relation of riparian buffer strips to in-stream habitat,
macroinvertebrates and fish in a small Iowa stream. J Iowa Acad Sci 113:49–55
Ericsson A, Stevens V (1996) Fringe benefits: a land owner’s guide to the value and stewardship of
riparian habitat. Forest Renewal BC. Fraser River Action Plan. Environment Canada, 28pp.
[Link]
Fajardo D, Johnston-González R, Neira L, Chará J, Murgueitio E (2009) Influencia de sistemas
silvopastoriles en la diversidad de aves en la cuenca del río La Vieja, Colombia. Recursos
Naturales y Ambiente 58:9–16
Galindo V, Calle Z, Chará J, Armbrecht I (2017) Facilitation by pioneer shrubs for the ecological
restoration of riparian forests in the Central Andes of Colombia. Restor Ecol 25:731–737
154 L. P. Giraldo et al.

Giraldo LP (2019) Efecto de la vegetación ribereña sobre la dinámica de la materia orgánica

particulada gruesa e insectos fragmentadores en quebradas de paisajes ganaderos (Eje Cafetero
de Colombia). PhD thesis, Universidad de Antioquia, Medellín, Colombia
Giraldo LP, Chará J, Zúñiga MDC, Chará-Serna AM, Pedraza G (2014) Impacto del uso del suelo
agropecuario sobre macroinvertebrados acuáticos en pequeñas quebradas de la cuenca del río La
Vieja (Valle del Cauca, Colombia). Rev Biol Trop 62:203–219
Giraldo LP, Chará J, Chará-Serna AM, Ramírez YP (2020) Restauración de corredores ribereños en
paisajes ganaderos de la zona andina colombiana: Efectos tempranos sobre el ambiente acuático.
Revista Acad Colomb Ciencias Exactas 44:652–664
González-Córdoba M, Zúñiga MDC, Manzo V (2015) Riqueza genérica y distribución de Elmidae
(Insecta: Coleoptera, Byrrhoidea) en el departamento del Valle del Cauca, Colombia. Biota
Colombiana 16:51–74
González-Córdoba M, Zúñiga MDC, Manzo V, Giraldo LP, Chará J (2016) Notelmis Hinton y
Onychelmis Hinton (Coleoptera: Elmidae: Elminae) de Colombia: Claves taxonómicas
ilustradas. Boletín del Museo de Entomología de la Universidad del Valle 16:1–17
Gray CL, Slade EM, Mann DJ, Lewis OT (2014) Do riparian reserves support dung beetle
biodiversity and ecosystem services in oil palm-dominated tropical landscapes? Ecol Evol 4:
1049–1060
Gray REJ, Slade EM, Chung AYC, Lewis OT (2017) Riparian reserves in oil palm plantations may
provide movement corridors for invertebrates. bioRxiv:204990. [Link]
Hynes H (1975) The stream and its valley. Verh Int Ver Theor Angew Limnol 19:1–15
Jones JB (1997) Benthic organic matter storage in streams: influence of detrital import and export,
retention mechanisms, and climate. In: Webster JR, Meyer JL (eds) Stream organic matter
budgets. J North Am Benthol Soc 16:109–119
Kuglerová L, Agren A, Jansson R, Laudon H (2014) Towards optimizing riparian buffer zones:
ecological and biogeochemical implications for forest management. For Ecol Manag 334:74–84
Lamberti GA, Gregory SV (2006) CPOM transport, retention, and measurement. In: Hauer FR,
Lamberti GA (eds) Methods in stream ecology, 2nd edn. Academic, San Diego, pp 273–292
Lees AC, Peres CA (2007) Conservation value of remnant riparian forest corridors of varying
quality for Amazonian birds and mammals. Conserv Biol 22:439–449
Lorion C, Kennedy B (2009) Relationships between deforestation, riparian forest buffers and
benthic macroinvertebrates in neotropical headwater streams. Freshw Biol 54:165–180
Luke SH, Slade EM, Gray CL, Annammala KV, Drewer J, Williamson J, Agama AL, Ationg M,
Mitchell S, Vairappan CS, Struebig MJ (2019) Riparian buffers in tropical agriculture: scientific
support, effectiveness and directions for policy. J Appl Ecol 56:85–92
Marczak LB, Sakamaki T, Turvey SL, Deguise I, Wood SLR, Richardson JS (2010) Are forested
buffers an effective conservation strategy for riparian fauna? An assessment using meta-
analysis. Ecol Appl 20:126–134
Medina A, Harvey CA, Merlo DS, Vílchez S, Hernández B (2007) Bat diversity and movement in
an agricultural landscape in Matiguás, Nicaragua. Biotropica 39:120–128
Meli P, Calle A, Calle Z, Ortiz-Arrona CI, Sirombra M, Brancalion PHS (2019) Riparian-forest
buffers: bridging the gap between top-down and bottom-up restoration approaches in Latin
America. Land Use Policy 87:104085. [Link]
Méndez LE, Calle Z (2010) Plantas de la Cuenca media del río La Vieja. CIPAV y
CIEBEREG, Cali
Mingoti R, Vettorazzi CA (2011) Relative reduction in annual soil loss in microwatersheds due to
the relief and forest cover. Eng Agric 31:1202–1211
Molineri C, Zúñiga MDC, Ramos BC, Giraldo LP, Cardona W (2016) Three new species of
Leptohyphes Eaton (Ephemeroptera: Leptohyphidae) from Colombia. Iheringia, Série Zoologia
106:e2016024
6 Riparian Forests: Longitudinal Biodiversity Islands in Agricultural Landscapes 155

Moore RD, Richardson J (2003) Progress towards understanding the structure, function and
ecological significance of small stream channels and their riparian zone. Can J For Res 33:
1349–1351
Murgueitio E, Ibrahim M (2009) Ganadería y Medio Ambiente en América Latina. In:
Murgueitio E, Cuartas C, Naranjo J (eds) Ganadería del futuro: Investigación para el desarrollo.
Fundación CIPAV, Cali, pp 19–39
Naiman RJ, Bilby RE, Bisson PA (2000) Riparian ecology and management in the Pacific coastal
rain forest. Bioscience 50:996–1011
Naiman RJ, Décamps H, McClain ME (2005) Riparia: ecology, conservation and management of
streamside communities. Elsevier Academic Press, San Diego
Osborne LL, Kovacic DA (1993) Riparian vegetated buffer strips in water-quality restoration and
stream management. Freshw Biol 29:243–258
Paetzold A, Schubert CJ, Tockner K (2005) Aquatic-terrestrial linkages along braided-river:
riparian arthropods feeding on aquatic insects. Ecosystems 8:748–759
Palmer MA, Hondula KL, Koch BJ (2014) Ecological restoration of streams and rivers: shifting
strategies and shifting goals. Annu Rev Ecol Evol Syst 45:247–269
Ramírez YP, Giraldo LP, Zúñiga MDC, Ramos BC, Chará J (2018) Influencia de la ganadería en los
macroinvertebrados acuáticos en microcuencas de los Andes centrales de Colombia. Rev Biol
Trop 66:1244–1257
Riseng CM, Wiley MJ, Black RW, Munn MD (2011) Impacts of agricultural land use on biological
integrity: a causal analysis. Ecol Appl 21:3128–3146
Sabo JL, Sponseller R, Dixon M, Gade D, Harms T, Heffernan J, Jani A, Katz G, Soykan C,
Watts J, Welter J (2005) Riparian zones increase regional species diversity by harboring
different, not more species. Ecology 86:56–62
Sadeghian S, Rivera J, Gómez M (1999) Impacto de sistemas de ganadería sobre las características
físicas, químicas y biológicas de suelos en los Andes de Colombia. In: Sánchez M, Rosales M
(eds) Agroforestería para la Producción Animal en América Latina, Producción y Sanidad
Animal No. 143. FAO, Roma, pp 123–142
Schilling KE, Jacobson P (2014) Effectiveness of natural riparian buffers to reduce subsurface
nutrient losses to incised streams. Catena 114:140–148
Schroth G, da Fonseca GAB, Harvey CA, Gascon C, Vasconcelos HL, Izac AM (2004) Agrofor-
estry and biodiversity conservation in tropical landscapes. Island Press, Washington, DC
Skłodowski M, Kiedrzynska E, Kiedrzynski M, Urbaniak M, Zielinska KM, Kurowski JK,
Zalewski M (2014) The role of riparian willows in phosphorus accumulation and PCB control
for lotic water quality improvement. Ecol Eng 70:1–10
Suurkuukka H, Virtanen R, Suorsa V, Soininen J, Paasivirta L, Muotka T (2014) Woodland key
habitats and stream biodiversity: does small-scale terrestrial conservation enhance the protection
of stream biota? Biol Conserv 170:10–19
Tanaka MO, Souza ALT, Moschini LE, Oliveira AK (2016) Influence ofwatershed land use and
riparian characteristics on biological indicators of stream water quality in southeastern Brazil.
Agric Ecosyst Environ 216:33–339
Turunen J, Markkula J, Rajakallio M, Aroviita J (2019) Riparian forests mitigate harmful ecological
effects of agricultural diffuse pollution in medium-sized streams. Sci Total Environ 649:495–
503
Uribe-Gómez F (2008) La Dimensión Ambiental en la Construcción de la Ecorregión Eje
Cafetero. In: Rodriguez JM, Camargo JC, Niño J, Pineda AM, Arias LM, Echeverry MA,
Miranda CL (eds) Valoración de la Biodiversidad en la Ecorregión del Eje Cafetero.
CIEBEREG, Pereira, pp 21–38
Vannote RL, Minshall GW, Cummins KW, Sedell JR, Cushing CE (1980) The river continuum
concept. Can J Fish Aquat Sci 37:130–137
Villada S, Triana LA, Días L (2017) Grupos funcionales alimentarios de insectos acuáticos en
quebradas andinas afectadas por agricultura y minería. Caldasia 39:370–387
156 L. P. Giraldo et al.

Wallace JB, Eggert SL, Meyer JL, Webster JR (1997) Multiple trophic levels of a forest stream
linked to terrestrial litter inputs. Science 277:102–104
Yoshimura M (2012) Effects of forest disturbances on aquatic insect assemblages. J Entomol Sci
15:145–154
Zimbres B, Peres CA, Machado RB (2017) Terrestrial mammal responses to habitat structure and
quality of remnant riparian forests in an Amazonian cattle-ranching landscape. Biol Conserv
206:283–292
Zúñiga MDC, Giraldo LP, Calero H, Ramírez YP, Chará JD (2014) Anacroneuria caraca stark y
A. jewetti stark (Insecta: Plecoptera: Perlidae): Primeros registros para los Andes Orientales y el
pie de monte de la orinoquía Colombiana. Boletín del Museo de Entomología de la Universidad
del Valle 15:12–19
Zúñiga MDC, Giraldo LP, Ramírez YP, Chará JD, Ramos BC (2015) Neoatriplectides (Trichoptera:
Atriplectididae) en Colombia:Notas sobre su taxonomía, ecología y distribución en el
Neotrópico. Rev Colomb Entomol 41:149–152
Chapter 7
Conservation and Registration of Seed
Sources in Reserve Remnants
in the Province of Misiones, Argentina

Beatriz I. Eibl, Florencia Montagnini, Lucas N. López, Héctor F. Romero,

Claudio J. Dummel, Guillermo Küppers, Miguel A. López,
Pedro Lavignolle, Jorge Cortes, and Marcelo A. De La Vega

Abstract The Upper Parana Atlantic Forest, a biodiversity hotspot of highly threat-
ened biodiversity, extends to northeastern Argentina in the province of Misiones. In
Argentina, national and provincial laws promote the restoration of degraded forest
areas. This is a crucial process necessary to the establishment of biodiversity islands,
which requires plant propagation material of native species to ensure their viability
and diversity. Since 2018, a registry of forest areas and seed trees has been
implemented nationwide to serve as a repository of biodiversity and plant propaga-
tion material, controlled by the National Seed Institute (INASE) Regulation n 318/
18, elaborated in scientific/technical cooperation with the School of Forestry of the
National University of Misiones. In these registered areas, which can be in private,
public, or protected lands, propagation material is certified, with priority given to
arboreal species and rare, endemic, threatened and/or vulnerable species. Each
species is registered in every site where it is documented in order to record as
much genetic variability as possible. Propagation material certification establishes
a protocol that allows native species to be included, under certified control, in
sustainable, productive, and multiple-purpose land use systems. To date, more
than 1000 registered seed trees have been recorded in more than 54 hectares of

B. I. Eibl (*) · L. N. López · H. F. Romero · C. J. Dummel · G. Küppers · M. A. López

Facultad de Ciencias Forestales, Universidad Nacional de Misiones, Eldorado, Misiones,
Argentina
e-mail: hcr_lopez@[Link]
F. Montagnini
School of the Environment, The Forest School, Yale University, New Haven, CT, USA
e-mail: fl[Link]@[Link]
P. Lavignolle · J. Cortes · M. A. De La Vega
Instituto Nacional de Semillas (INASE), Dirección de Certificación y Control, Buenos Aires,
Argentina
e-mail: plavig@[Link]; jcortes@[Link]; madelavega@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 157
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
158 B. I. Eibl et al.

forest remnants. Registered seed trees facilitate restoration of degraded areas,

increasing site diversity via dispersion by birds, small animals and wind. The
certification of native species for multiple use contributes to their conservation and
to their use for restoration, while also generating resources for owners, helping to
conserve biodiversity.

Keywords Atlantic forest · Diversity · Germination · Natural reserves · Production ·

Propagation material · Restoration

7.1 Introduction

The eco-region of the Atlantic Forest in South America is an internationally recog-

nized biodiversity hotspot due to its biologically rich terrestrial ecosystems
(Galindo-Leal and Câmara 2003; Mittermeier et al. 2011). In Argentina, the Atlantic
Forest, locally known as the Upper Parana Atlantic Forest, emerges from the coast
and stretches into the interior, sharing portions with Brazil and Paraguay (Di Bitetti
et al. 2003). The part of the Upper Parana Atlantic Forest in the Province of
Misiones, known as the Selva Misionera or Selva Paranaense, contains the greatest
biodiversity in Argentina, with dense multi-stratified vegetation in the forests and
isolated trees in open fields. Many previously unfound species continue to be
discovered in the forest and added to the local biodiversity lists, proving it has the
ideal habitat conditions to support a variety of life (Keller 2007).
Restoration and conservation practices in this region of the Atlantic Forest can
help protect one of the last relicts of native forest and therefore conserve native
species. In this region, restored areas of protective forests on the borders of streams
and slopes provide connectivity between different areas of restored forests in the
basins of the Uruguay, Iguazú and Paraná rivers, which contributes to the biodiver-
sity of the region. Additionally, because the Upper Parana Atlantic Forest shares its
atmosphere, biodiversity, and surface and underground water with Brazil and Par-
aguay, its biodiversity is connected to those nations, so protecting the Argentinian
section of forest will contribute to the protection of land in those nations as well.
However, the advancement of the agricultural frontier has caused changes in land
use, population growth, timber extraction from natural forests and increased isola-
tion of protected areas, which has compromised the integrity of the genetic and
species diversity of these regions and the durability of these natural formations
(Young 2003; Gibbs et al. 2010; Mittermeier et al. 2011; Armenteras et al. 2017).
The protection of the Atlantic Forest is crucial to the environmental health of the
region, which is why the Argentinian government has made previous efforts to
conserve the land.
The government of Argentina has passed laws promoting restoration and conser-
vation efforts, and restoration efforts are ongoing. Strategies for biodiversity con-
servation in this region must include a series of actions applied by public institutions
that complement each other in order to provide an enforced, comprehensive envi-
ronmental protection plan. For instance, the Provincial Laws XVI n 60 “Green
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 159

Corridor” (Ley Provincial XVI nº60 2010) and n 29 (Ley Provincial nº29 2010) of
Protected Natural Areas present an opportunity to maintain natural forest connec-
tivity in the Province of Misiones by conserving forest in public areas, private parks
and nature reserves.
Conserving biodiversity can also be a profitable renewable resource through the
use of local species for economic purposes. Therefore, following the principles of its
conservation and rational use, biodiversity can be considered as an asset for eco-
nomic and social development purposes, providing additional benefits toward con-
servation practices (Ugarte-Guerra et al. 2010).
However, several factors must be considered when conserving land and
establishing biodiversity islands. The degree of degradation and the species present
in the targeted landscape are important factors to evaluate when designing biodiver-
sity islands. After determining the degradation status, it is possible to determine the
most feasible and appropriate restoration pathway that will maximize the efficiency
and impact of conservation efforts (Montagnini et al. 2022). In Argentina, the
national and provincial laws establish and promote the restoration of degraded
areas, which requires propagation material of native species including seeds, fruits,
cuttings, etc. As part of our previous work on restoration of degraded lands with
native tree species that can be useful for productive purposes, a list of plant species
was compiled. The creation of this database facilitated the selection of species most
compatible with the productive systems in the region (including plantations, agro-
forestry, and silvopastoral systems) (Eibl et al. 2015b). These specifically-chosen
plant species provide long-term products and store atmospheric carbon, in addition
to the advantages of diversity conservation via native forests strips and patches.
In this chapter, we describe the methodology, procedures and contributions of a
successful registry and certification system of seed sources on land areas in need of
restoration and conservation of species diversity in the region. The registry allows
for the propagation of genetic material used in restoration and/or enrichment plans to
be traced and monitored to evaluate its success and potential use in domestication
programs.

7.2 Legal Framework Supporting Tree Planting

in Argentina

In Argentina, the National Law on Minimum Budgets for the Environmental Protec-
tion of Native Forests, n 26,331 (Ley Nacional nº26331 2007). classifies native forests
into three management categories: conservation, sustainable management and poten-
tial conversion to other uses. Forests that fall in the first two categories must be
restored with native species, which may be required by the governing agency in each
province. In the case of Misiones, the governing agency is the Ministry of Ecology and
Renewable Natural Resources (MEyRNR). In addition to government regulation,
National Law n 27,487 (Ley Nacional nº27487 2008), establishes incentives for
160 B. I. Eibl et al.

restoration projects or productive projects which involve planting and harvesting. In

order to receive the incentive, scientific procedures for planting propagation material
need to be available and used.
Misiones has enacted a series of provincial laws in order to promote restoration
efforts of threatened forest and protect regional biodiversity. Provincial Law XVI
n 53 (Ley Provincial XVI nº53 2010), utilizes protective forests and riparian corri-
dors to aid in the recovery of stream and slope borders. The law establishes that
restoration of border areas is mandatory, and that tree species may be used to
improve restoration efforts based on a management plan previously approved by
the relevant authorities. The Misiones Provincial Law of Renewable Energies XVI
n 97 (Ley Provincial XVI nº97 2010), promotes the planting of species for biomass
purposes. By means of the database of plant species, ideal plant material can be
selected for restoration of a habitat. For example, plantation of potential native
species for timber purposes can have benefits to forest restoration (Eibl et al.
2015a). Law XVI n 98 (Ley Provincial XVI nº 98 2010), of the Provincial Bank
of Germplasm of Misiones, which aims to preserve the genetic base of native plants,
facilitates the availability of seeds for conservation, research and productive use.
Likewise, the Provincial Law XVI n 47 (Ley Provincial XVI nº 47 2010), regulates
the conservation and sustainable use of biological diversity and its components, in
accordance with the Global Strategy for the Conservation of Plant Species (GSPC
2020), of which Argentina is a signatory.
The implementation of these restoration efforts requires appropriate propagation
material. In order to ensure plant material has the greatest benefit to a region while
avoiding potential invasive or diseased species requires the application of specific
certification protocols. These certifications ensure that restoration projects use prop-
agation material that will maximize their contribution to the biodiversity of the
region. In addition, successful restoration efforts require proper monitoring and
verification in order for the desired functions and objectives of propagation material
to be properly fulfilled.
The important restoration effort that Argentina is making by protecting forests
and providing renewable energy fulfills its commitments made alongside a total of
195 countries at the United Nations Climate Change Conference in COP21 (2015).
In COP25 (2019) Argentina was one of the Latin American countries that signed the
“Declaration for Restoration,” committing 30 million hectares within the region to
be restored by 2030. This goal aims to bring back vital ecosystems and promote
social and economic development of the region.
Forests are expected to make key contributions to reduce greenhouse gases
(GHG) emissions and to ensure the long-term stability of global climate. Restoration
and management of degraded forests can contribute to resilient landscapes by
absorbing carbon and maintaining other ecosystem services, such as the provision
of water for sustainable consumption and productive use. Conservation of forests
and reforestation schemes contribute to achieving sustainable development goal
(SDG) 15 on Life on Land by maintaining and increasing the proportion of the
world’s land area that is forested (IUCN 2020b). In fact, FAO’s 2018 State of the
World’s Forests describes how forests and trees contribute to 28 targets relating to
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 161

ten SDGs (FAO 2018; FAO 2020), showing the importance of degraded land
restoration.

7.3 Strategies and Methods for Certification of Seed

Sources of Native Species

The current strategies for conservation of native species for environmental and
productive purposes in Argentina were tested in private reserves by technicians
and professionals, and subsequently disseminated in workshops and participatory
courses led by the School of Forestry of the National University of Misiones
(FCF/UNaM). For each stage of the certification process, beginning by harvesting
seeds and other specimens, there are a minimum of criteria to be considered before
progressing to the next stage in order to ensure that species are properly conserved
and successfully used in restoration practices.
Current conservation strategies include the establishment and upkeep of a registry
of tree seed species, the storage of seeds of native species, and the certification of the
characteristics of the seeds in their botanical, physical and physiological aspects as is
typically done in a seed analysis laboratory. In addition, plants that are produced in
nurseries receive the Certification of Nurseries of Native Species through specific
protocols, which are then registered in the National Registry of Seed Trade and
Control (RNCyFS) of the National Seed Institute (Instituto Nacional de Semillas,
INASE). Certification protocols for trees follow the Forest Species Certification
System, regulated by INASE Resolution n 256/99 (INASE 1999), which establishes
a system of classes and categories for basic propagation material. The National
Network of Germplasm Banks of Native Plant Species (RNBGEVN) contributes to
the exchange of information and training on germplasm conservation of native
species (De Viana et al. 2011).

7.3.1 Conservation Areas That Contain Propagation Material

National parks and public nature reserves can serve as germplasm reservoirs for
conservation or research. Private nature reserves can also provide germplasm for
propagation and productive uses if the material is properly certified.
In the Province of Misiones, some private landowners participate in the country’s
private nature reserve network with their forest remnants (Schiaffino and Bertolini
2016). Most of the small traditional producers in the province maintain their own
nature reserves because of their commitment to the environment, in response to legal
requirements, and/or for the possibility of participating in certification systems for
their products. It is important to generate resources for the owners who are currently
162 B. I. Eibl et al.

conserving biodiversity to ensure the persistence of their private reserves and their
willingness to protect them through time (Eibl 2006; Eibl and López 2017).
The existing national and provincial legal framework does not provide a basis for
specifically certifying propagation material harvested from nature reserves, which
could be a potential profitable resource as well as beneficial to the restoration
process. Our project aims to provide specific actions to conserve remaining natural
forests areas, to create new productive forests through natural succession, and to
generate a productive process with native species that are part of restoration activ-
ities. Finally, our project attempts not only to provide specific conservation and
restoration actions, but also to provide technical background for designing appro-
priate legal tools to certify areas that house these native seeds. These tools would
establish a restoration process that is more efficient and easier, which would provide
greater incentives for landowners to protect their land.

7.3.2 Seed Tree Network

The Faculty of Forest Sciences who works on identification and registration of seed
trees in areas of private reserves, developed their techniques in joint cooperation
with private companies, using information that had been gathered in 20–30 years of
projects where they had been participants. These projects were run or funded by
organizations such as the Mellon Foundation (1990/1998), Wildlife Foundation
(1998), Perez Companc (2002), Tajy Project of Tabaco Norte (2007/2010),
Biofábrica Misiones SA (2010/2012), the Registry of Seed Areas in projects with
yerba mate (Ilex paraguariensis), small farmers with the UCAR (Unidad de Cambio
Rural), Ministry of Agriculture, and INTA (Instituto Nacional de Tecnología
Agropecuaria) (2014/2017) (Eibl and López 2017; Niella et al. 2016, 2017). Using
results of those projects, a list of potential native tree species was taken as a basis,
with a total of 336 tree species represented in 67 botanical families that occupy the
middle and upper stratum of the Misiones forest (Paranaense forest) (Gartland and
Bohren 2008).
Based on the interest expressed by individual landowners, who were identified
through local information and references, technical visits to each site were made. In
each reserve area, species are selected from this list according to the preferences of
the owners as well as the classification of chosen species as valuable, rare, vulner-
able, and/or threatened species (IUCN 2020a). Specimens of greater size, with good
health, and those whose tree form is the most representative for each of the species
are given priority when choosing. These criteria are used for all species, with greater
emphasis on species of ecological importance. If individual specimens are no longer
found in desired conditions, they are registered in the state in which they appear, in
order to guarantee natural regeneration of that species. When possible, each species
is recorded at every site of their natural distribution in order to ensure accuracy in
documentation, always considering the recommendations of Thomas et al. (2014) on
genetic considerations in ecosystem restoration when using native tree species.
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 163

Seed trees are registered by their identity, geographical position (GPS point),
health status, location on the site, and description of their surrounding environment.
A data sheet is prepared for each individual, containing information on size and
photographs. Isolated trees of interest are also considered for registration. A list of all
selected individuals is prepared, using a code with acronyms for identifying the
region, species, number of individuals and date (Eibl et al. 2001, 2002; Eibl and
Báez 2004; Eibl and López 2017). This allows leaders of restoration efforts to have
ample data available to them in order to make informed decisions on the proper
propagation material to use to have the most effective impact on the intended area of
conservation.

7.3.3 Harvest, Identification, and Conditioning of Fruits

and Seeds

Fruit harvesting is done when there is a specific demand of propagation material.

Desired seed plants are located based on a database of species already registered by
the School of Forestry (FCF/UNaM). The time of end of maturation and beginning
of dispersion is identified from the phenological observation of the individuals to be
harvested. This is the stage in which fruits and seeds have the lowest dry weight and
the greatest percentages of germination and vigor (Eibl et al. 2012). These location
data and observations allow harvesters to easily find and collect fruit when needed.
In general, the species of dehiscent fruits which split at maturity releasing their
contents must be harvested on the tree, while those of indehiscent fruits, such as
legumes and berries, can be harvested from the soil, as well as on the tree when they
are in the last stage of dispersion. Treetop harvesting is done by specially trained
staff who climb up the trees. A net is previously placed on the ground to collect the
fruits and seeds of both, fruits that fall naturally and those that have to be harvested
(Gold et al. 2004).
The harvested fruits corresponding to each individual specimen represent the
propagation material of that tree. These materials are labeled in the field and entered
in the Seed Bank notebook, with a number that guarantees their traceability. This
information includes owner name, GPS point, harvest date, tree number, species, and
location. In addition to fruits and seeds of trees (Eibl 2013), collection of propagation
material can also come from bushes, herbs, grasses, epiphytes, orchids, ferns, and
others. After being harvested, propagation materials are classified and certified by
origin, provenances and identity. Their identity is verified using the website darwin.
[Link] which is based on information from the Darwinion Botanical Institute
(Instituto de Botánica Darwinion) (Flora del Conosur 2020).
After harvest, the seeds are conditioned by separating them from their fruits and
cleaning them of impurities such as dust and woody sticks. The seeds are dried until
they reach equilibrium with the environment and then they are prepared for storage
and/or for nurseries. The seeds that are entered into active seed banks or germplasm
164 B. I. Eibl et al.

banks are conditioned for their purpose using drying and/or ultra-drying techniques
and are packed in suitable containers (to avoid excess moisture), in order to maintain
their viability for a longer time (Eibl et al. 2012, 2013a).

7.3.4 Certification of Physical and Physiological Seed Quality

The germination to produce a normal seedling (percent germination, PG), the

moisture of the seed at the time of harvest and during storage (H%), purity (whole
seeds and/or portion of germinable seeds), and the weight of seeds (weight of 1000
seeds in grams) are determined in the laboratory, following recommendations from
the International Seed Test Standards (ISTA 2019) and Ministério da Agricultura
(2009). Germination tests are used to determine the best pre-germination treatments,
while the germination conditions, times appropriate for each species, the minimum
requirements for germination and purity percentages are established for each species
following procedures described by Eibl et al. (2012).

7.3.5 Nursery and Field Planting of Native Species

For production of plants in the nursery, different technical specifications are required
for each species in terms of pre-germination treatments, nursery times, substrates,
containers and nutrients. Likewise, technical specifications are used in determining
quality indicators for height and diameter at the collard or base (dac), slenderness
(ratio of stem/dac), length, lignification of seedlings (highest dry weight) and the
consistency of the roots with the substrate used for growing the seedlings (Eibl et al.
2013b).
Field planting of native species requires knowledge of the silvicultural system
and certification requirements. The selection of the species to be planted depends on
the owner’s preferences and conditions of the site. Individual planting holes of at
least 60–80 cm in depth promote the development of the trees, ensuring their support
and improving their source of nutrition.
The natural vegetation of the site can protect the planted seedlings from direct
sunlight, from high and low temperatures, and from frost, drought, wind and insects.
If the natural vegetation is not present, pioneer or species tolerant to direct exposure
should be planted first to ensure greatest survival and good initial growth. Prior
evidence indicates that agroforestry systems (AFS) or trees planted in areas
supporting early natural succession are likely to guarantee the survival of the
seedlings (Montagnini et al. 2005, 2006; Barth et al. 2008; López et al. 2013; Eibl
et al. 2015b), as well as to contribute to the recovery of degraded soils (Day et al.
2011; Eibl et al. 2019b).
For example, as part of our own research on restoration of degraded lands using
native trees, we planted 11 native tree species in sites with extreme degradation due
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 165

to intensive agricultural use and invasive grasses. The initial trees planted were
heliophytes of rapid growth. In a second stage, we added valuable shade tolerant
woody species, in order to recover the potential of the site to produce wood and
fuelwood and fix atmospheric carbon (Montagnini et al. 2006; Eibl et al. 2019b). The
evidence from this research showed that tree plantations can facilitate the establish-
ment of natural regeneration, adding diversity of species to the site and allowing the
generation of new multi-diverse forests by tending to the species of interest
(Montagnini et al. 2005; Eibl et al. 2019c).

7.4 Advances in the Conservation and Certification of Seed

Sources of Native Tree Species
7.4.1 Conservation Areas That Contain Propagation Material

In a joint work with the National Seed Institute of Argentina (INASE), a new
regulation for the Registration of Conservation Areas and the certification of prop-
agation material was recently achieved. The Resolution N 318/18 (INASE
2018) refers to the registration of the seed production area of native species and is
being disseminated by technical personnel from the National Seed Institute and the
Forestry School of the National University of Misiones (FCF-UNaM). In addition,
through training and communication workshops, the Resolution is being distributed
among interested provinces that must adhere to the proposal at the national level.
The Province of Misiones followed the national proposal with an agreement
subscribed on September 24, 2019 (n 81), between INASE and MEyRNR (Ministry
of Ecology and Renewable Natural Resources of Misiones). Simultaneously, an
agreement was reached between the FCF-UNaM and MEyRNR for the implemen-
tation and the scientific and technical support of the proposal in the field. The
identification and documentation of tree diversity that is contained in an area or
private reserve that the owner agrees to protect are registered under a number
assigned by the RNCyFS (Fig. 7.1) in order to keep accurate record of species
counts and types present. Examples of lists of identified trees and seedlings in private
reserves are shown in Tables 7.1 and 7.2.
The propagation material of all species (fruits, seeds, pollen, buds, branches,
roots, others) that is harvested for use is certified. In designating this certification,
several factors are considered, including the sustainability of the source. Certifica-
tion allows for more informed decisions to be made regarding what propagation
material will have the greatest beneficial impact to the land and not result in any
invasive or diseased species. Therefore, a management plan designed by a qualified
professional is critical and has to be appropriate to the area and approved by the
MEyRNR of the Province of Misiones (Ley Provincial XVI nº105 2010).
The FCF-UNaM had previously worked in more than 30 natural reserves with a
total area of 54 hectares of remnants of small and medium private forests, registering
166 B. I. Eibl et al.

Fig. 7.1 Forest reserve with selected native tree species, in productive areas in a family farm in
Andresito, Province of Misiones. (Photo: B. Eibl)

the trees and monitoring the biodiversity, providing valuable information prior to the
creation of the new national regulation. A field demonstration of certification at the
level of trees, shrubs, epiphytes, and herbs of the propagation material collected from
the origin was carried out in these reserves. We also evaluated the opportunity to sell
fruit, seeds and/or plant seedlings in the market, as a way to generate income to the
owners to compensate them for their conservation efforts (Fig. 7.2).

7.4.2 Seed Tree Network

In accordance with the existing seed tree network protocol, all the species living in
each area are included in the registry, from the crown strata with epiphytes and ferns
to the undergrowth with herbs and shrubs as well as mushrooms and mosses. Trees
are registered in every landscape where they are present, whether they are part of
forests, riverine strips or growing in fields with grasses (Table 7.3). For example,
some hardwood species, such as Anadenanthera colubrina, Astronium balansae,
Handroanthus heptaphyllus, and palms such as Butia yatay, and Acrocomia aculeate
tend to predominate in open fields in the region.
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 167

Table 7.1 Adult tree species in the forest reserve of the Stadler family farm, in General Alvear,
Province of Misiones
Common Appearance/
N Scientific name name Botanical family frequency
1 Casearia sylvestris Sw. var. Burro caá FLACOURTIACEAE 1
sylvestris
2 Cordia trichotoma(Vell.) Arráb. Loro negro BORAGINACEAE 1
exSteud.
3 Cupania vernalis Cambess. Camboata SAPINDACEAE 1
Colorado
4 Matayba eleagnoides Radlk. Camboata SAPINDACEAE 1
Blanco
5 Helietta apiculata Benth. Canela de RUTACEAE 1
venado
6 Casearia decandra Jacq. Guazatumba FLACOURTIACEAE 1
chica
7 Rauvolfia sellowii Müll. Arg. Quina APOCYNACEAE 1
8 Machaerium stipitatum (DC.) Isapu’ymoroti FABACEAE 1
Vogel
9 Chrysophyllum gonocarpum Aguaí SAPOTACEAE 1
(Mart. &Eichler) Engl.
10 Diatenopteryx sorbifolia Radlk. María preta SAPINDACEAE 1
11 Myrocarpus frondosus Allemão Incienso FABACEAE 1
12 Aralia warmingiana (Marchal) Sabuguero ARALIACEAE 1
J. Wen
13 Zanthoxylum petiolareA. St.- Fagara RUTACEAE 1
Hil. &Tul. naranjillo
14 Chrysophyllum marginatum Vasuriña SAPOTACEAE 1
(Hook. &Arn.) Radlk. ssp.
Marginatum
15 Erythrina falcataBenth. Ceibo FABACEAE 1
16 Syagrus romanzoffiana (Cham.) Pindo PALMAE 1
Glassman
17 Lonchocarpus campestris Mart. Rabo ita FABACEAE 1
exBenth.
18 Pisonia zapallo var. Zapallo Pisonia NYCTAGINACEAE 1
Griseb. zapallo
19 Hovenia dulcis Thunb. Hovenia RHAMNACEAE 1
20 Picrasma crenata (Vell.) Engl. Palo amargo SIMAROUBACEAE 1
21 Ocotea puberula (Rich.) Nees Laurel guaíca LAURACEAE 2
22 Dendropanax cuneatus (DC.) Ombu ra ARALIACEAE 2
Decne. &Planch.
23 Nectandra lanceolata Nees Laurel LAURACEAE 2
amarillo
24 Myrsine balansae (Mez) Otegui Pororoca MYRSINACEAE 2
25 Sorocea bonplandii (Baill.) Ñandipa MORACEAE 2
W.C. Burger, Lanj. &Wess. Boer
(continued)
168 B. I. Eibl et al.

Table 7.1 (continued)

Common Appearance/
N Scientific name name Botanical family frequency
26 Prunus brasiliensis (Cham. Persiguero ROSACEAE 2
&Schltdl.) D. Dietr.
27 Jacaratia spinosa(Aubl.) A. DC. Yacaratia CARICACEAE 2
28 Nectandra megapotamica Laurel negro LAURACEAE 2
(Spreng.) Mez
29 Parapiptadenia rigida (Benth.) Anchico FABACEAE 2
Brenan Colorado
30 Balfourodendron riedelianum Guatambú RUTACEAE 3
(Engl.) Engl. Blanco
31 Cordia americana (L.) Guayubira BORAGINACEAE 3
Gottschling& J.S. Mill.
32 Holocalyx balansae Micheli Alecrín FABACEAE 3
33 Apuleia leiocarpa (Vogel) Grapia FABACEAE 4
J.F. Macbr.
34 Alchornea triplinervia (Spreng.) Mora blanca EUPHORBIACEAE 4
Müll. Arg. grande
35 Maclura tinctoria ssp. Tinctoria Mora amarilla MORACEAE 4
(L.) Steud.
36 Trichilia claussenii C. DC. Catigua MELIACEAE 4
guazú
37 Cedrela fissilis Vell. Cedro MELIACEAE 5
misionero
38 Aspidosperma austral Müll. Guatambú APOCYNACEAE 5
Arg. Amarillo
39 Inga marginataWilld. Inga’i FABACEAE 6
40 Cecropia pachystachya Trécul Ambay CECROPIACEAE 7
41 Cabralea canjerana Cancharana MELIACEAE 12
(Vell.) Mart.
42 Ficus luschnathiana (Miq.) Higuera de MORACEAE 16
Miq. Monte
Source: Eibl et al. (2016a). Project: Productive experiences with multipurpose native species in
environmental restoration areas and strategies for the conservation of forest remnants. Rural Change
Unit (UDI) n 01/2013. Sustainable Forest Plantations (SFPC)

The Registry of Seed Trees currently contains more than 1000 individuals
registered in different areas of the province since 1990, whose propagation materials
are currently available to be harvested for a nursery supply. Endangered, vulnerable
and rare species such as Araucaria angustifolia, Aspidosperma polyneuron,
Myrocarpus frondosus, Maclura tinctoria, Aralia warmingiana, Aspidosperma aus-
trale, and others are the priority species in the records, even if the individuals are old
or deteriorated, in order to prevent species extinction and preserve valuable genetic
information (Fig. 7.3).
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 169

Table 7.2 Regenerating tree seedlings in the forest reserve of Stadler family, in General Alvear,
Province of Misiones
Common Appearance/
N Scientific name name Botanical family frequency
1 Casearia sylvestris Sw. var. Burro caá FLACOURTIACEAE 1
sylvestris
2 Chrysophyllum gonocarpum Aguaí SAPOTACEAE 1
(Mart. &Eichler) Engl.
3 Trichilia catigua A. Juss. Catiguá MELIACEAE 1
Colorado
4 Matayba eleagnoides Radlk. Camboata SAPINDACEAE 1
Blanco
5 Eugenia burkartiana MYRTACEAE 1
(D. Legrand) D. Legrand
6 Jacaranda micrantha Cham. Caroba BIGNONIACEAE 1
blanca
7 Ocotea puberula (Rich.) Nees Laurel LAURACEAE 1
guaíca
8 Trichilia elegansA. Juss. Catigua MELIACEAE 1
Chico
9 Carica papaya L. Mamon CARICACEAE 1
10 Ilex brevicuspis Reissek Caona AQUIFOLIACEAE 1
11 Alchornea triplinervia (Spreng.) Mora blanca EUPHORBIACEAE 1
Müll. Arg. grande
12 Diatenopteryx sorbifolia Radlk. María preta SAPINDACEAE 1
13 Ficus luschnathiana (Miq.) Miq. Higuera de MORACEAE 1
Monte
14 Daphnopsis racemosa Griseb. Daphnopsis THYMELAEACEAE 1
15 Trichilia catigua A. Juss. Catiguá MELIACEAE 1
Colorado
16 Citronella paniculata (Mart.) Pasto de ICACINACEAE 1
R.A. Howard anta
17 Cecropia pachystachya Trécul Ambay CECROPIACEAE 1
18 Tabernaemontana catharinensis Horquetero APOCYNACEAE 1
A. DC.
19 Solanum granulosum-leprosum Fumo bravo SOLANACEAE 1
Dunal
20 Celtis iguanaea (Jacq.) Sarg. Tala CELTIDACEAE 1
21 Schefflera morototoni (Aubl.) Cacheta ARALIACEAE 1
Maguire, Steyerm. &Frodin
22 Cordia americana (L.) Guayubira BORAGINACEAE 2
Gottschling& J.S. Mill.
23 Guarea macrophylla ssp. Cedrillo MELIACEAE 2
Spicaeflora (A. Juss.) T.D. Penn.
24 Hennecartia omphalandra Ñandipa’ra MONIMIACEAE 2
J. Poiss.
25 Trema micrantha (L.) Blume Palo polvorá CELTIDACEAE 2
(continued)
170 B. I. Eibl et al.

Table 7.2 (continued)

Common Appearance/
N Scientific name name Botanical family frequency
26 Balfourodendron riedelianum Guatambú RUTACEAE 2
(Engl.) Engl. Blanco
27 Nectandra lanceolata Nees Laurel LAURACEAE 3
amarillo
28 Nectandra megapotamica Laurel negro LAURACEAE 3
(Spreng.) Mez
29 Pilocarpus pennatifolius Lem. Jaborandi RUTACEAE 3
30 Cupania vernalisCambess. Camboata SAPINDACEAE 3
Colorado
31 Maclura tinctoria ssp. Tinctoria Mora MORACEAE 3
(L.) Steud. amarilla
32 Myrsine balansae (Mez) Otegui Caápororocá MYRSINACEAE 4
33 Trichilia claussenii C. DC. Catigua MELIACEAE 4
guazú
34 Cedrela fissilis Vell. Cedro MELIACEAE 5
misionero
35 Aspidosperma australe Müll. Arg. Guatambú APOCYNACEAE 5
Amarillo
36 Cabralea canjerana (Vell.) Mart. Cancharana MELIACEAE 8
37 Inga marginata Willd. Inga’i FABACEAE 8
38 Sorocea bonplandii (Baill.) Ñandipa MORACEAE 23
W.C. Burger, Lanj. &Wess. Boer
39 Actinostemon concolor (Spreng.) Laranjeira EUPHORBIACEAE 31
Müll. Arg.
Ref. Eibl et al. (2016a). Project: Productive experiences with multipurpose native species in
environmental restoration areas and strategies for the conservation of forest remnants. Rural Change
Unit (UDI) n 01/2013. Sustainable Forest Plantations (SFPC)

7.4.3 Harvest, Identification and Conditioning of Fruits

and Seeds

As described in the methods, at the time of harvest, the fruits and seeds must be ripe,
with the maximum dry weight and minimum humidity on the mother plant (a time
that usually corresponds to the greatest germination and vigor). This moment
coincides with the initial and maximum phase of fruit maturity and dispersion, as
is the case for Cabralea canjerana, for example (Fig. 7.4).
Throughout the year, different species are found in their dispersal stage. When
harvested, the containers are labeled with data on site, species, and date of harvest.
The seeds of the recently collected ripe fruits have around 10–18% humidity for dry
fruits (orthodox seeds) and over 40% for fresh fruits (recalcitrant seeds). Condition-
ing the fruits, which consists of separating the seeds to achieve maximum purity, is
done manually or using sieves and fans to blow them into the seed containers.
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 171

Fig. 7.2 Diversity

monitoring guided by the
owners in the natural reserve
of the Stadler family farm in
General Alvear, Province of
Misiones. (Photo: B. Eibl)

7.4.4 Certification of the Physical and Physiological Seed

Quality

Based on our experiences following the protocols described, we have learned and
refined the methodology in a number of ways. For instance, the storage unit or the
nursery unit depends on the characteristic of the species. In general terms the unit is a
seed; however, where the separation of the seed is difficult the unit is a fruit
(i.e. Balfourodendron riedelianum and Helietta apiculata). The manual separation
of the seeds allows purity levels to exceed 90% (Eibl et al. 2013a). For the purpose of
certification, collected seed samples are expected to achieve a germination percent-
age higher than 70%. Exceptions are taken only in specific cases where germination
is naturally low (under 50%), as is the case for Bastardiopsis densiflora, Aralia
warmingiana, Scheflera morototoni, among others.
The germination rate is an expression of vigor and is linked to the storage
potential of the specimen. As the germination rate increases, the chance of
maintaining viability in storage also increases. For example, Cedrela fissilis seeds
germinate faster and more evenly when the dry weight of the seeds is greater
172 B. I. Eibl et al.

Table 7.3 Registered seed trees in the forest reserve of the Stadler family farm, in General Alvear,
Province of Misiones
Common
Scientific name name Botanical family Code
Alchornea triplinervia (Spreng.) Müll. Mora blanca MORACEAE 13-ALCTRI-
Arg. 002-15
Apuleia leiocarpa (Vogel) J.F. Macbr. Grapia FABACEAE 13-APULEI-
010-15
Aspidosperma australe Müll. Arg. Guatambú APOCYNACEAE 13-ASPAUS-
Amarillo 003-15
Cabralea canjerana (Vell.) Mart. Cancharana MELIACEAE 13-
CABCAN-
009-15
Cabralea canjerana (Vell.) Mart. Cancharana MELIACEAE 13-
CABCAN-
010-15
Cedrela fissilis Vell. Cedro MELIACEAE 13-CEDFIS-
Misionero 012-15
Cordia trichotoma (Vell.) Arráb. ex Peteribí BORAGINACEAE 13-CORTRI-
Steud. 022-15
Cordia trichotoma (Vell.) Arráb. Peteribí BORAGINACEAE 13-CORTRI-
exSteud. 023-15
Holocalyx balansae Micheli Alecrín FABACEAE 13-HOLBAL-
008-15
Maclura tinctoria (L.) [Link]. Mora MORACEAE 13-MACTIN-
tinctoria amarilla 006-15
Maclura tinctoria (L.) [Link]. Mora MORACEAE 13-MACTIN-
tinctoria amarilla 007-15
Rauvolfia sellowii Müll. Arg. Quina de APOCYNACEAE 13-RAUSEL-
Monte 001-15
Schefflera morototoni (Aubl.) Maguire, Cacheta ARALIACEAE 13-
Steyerm. &Frodin SCHMOR-
004-15
Source: Niella F, Eibl B, Keller H, Maiocco D, Rocha P, Vega V (2017) Network of trees and seed
areas for the conservation, domestication and rescue of native plant genetic resources in the
Province of Misiones. SILVA N 17/2013. Project Biodiversity Conservation Component in Forest
Productive Landscapes. GEF 090118. Ministerio de Agricultura Ganaderia y Pesca (MAGyP)

(Ocampo et al. 2015). Araucaria angustifolia seeds also maintain their initial vigor
in growth, two years after planted in the field (Eibl et al. 2017).
For seeds that are less likely to survive drying and freezing conditions during
ex-situ conservation, known as recalcitrant seeds, the storage humidity should be
generally maintained above 35%, and they should be preserved in moist sand under
cold temperatures (Rodríguez et al. 2019). Such is the case of seeds of Eugenia
involucrata of the Myrtaceae family, among others. The seeds of Cabralea
canjerana are very recalcitrant. Even when stored in wet sand and a cold environ-
ment, they can only survive for 4 months (González et al. 2015).
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 173

Fig. 7.3 Selected and

registered tree of
Aspidosperma polyneuron
in productive areas in a
family farm, in Andresito,
Province of Misiones.
(Photo: B. Eibl)

Fig. 7.4 Ripe fruits and dispersion of Cabralea canjerana indicate the optimal moment for harvest.
(Photo: B. Eibl)
174 B. I. Eibl et al.

Seeds of orthodox species should be dried until they reach equilibrium with the
environment and subsequently placed in containers with silica-gel as a desiccator to
reduce their humidity to 5% or less. They need to be stored in airtight containers and
in a cold environment to maintain seeds long term in germplasm banks. In storage
for conservation, the humidity must stay below 5% and germination must not fall
below 70% (Eibl et al. 2013a). Ultra-dried seeds of Aspidosperma polyneuron were
reported as viable after 5 years of storage (Otegui et al. 2016). This species’ seeds
were dehydrated to 5% and expressed 90% viability two (2) years after storage in dry
and cold conditions (Eibl et al. 2019a).
The certification of the physical and physiological quality of the seeds is regis-
tered in a label that contains identity data, seed origin, purity, germination power,
harvest date and weight, following procedures stated in resolution INASE n 42/2000
(INASE 2000) of seed marketing, and specifically for tree seeds, resolution INASE
n 256/99.

7.4.5 Nurseries and Field Planting of Native Species

As part of our research, silvicultural knowledge is already available for more than
forty native species of multiple use (firewood, wood, landscape, restoration, honey,
food, medicines, others) (Eibl et al. 2015b, 2016a, 2019c). The nurseries distributed
in different regions have received training and advice from personnel working at the
Experimental Nursery of Native Species linked to the Seed Bank of the Forestry
School of the University of Misiones in Eldorado, Misiones, Argentina. Seedlings of
30–40 cm in height of more than 40 different species, placed in rigid polypropylene
containers of 100 to 230 cm3, using composted pine bark substrates and slow-release
fertilizer at a rate of 1.5 to 3 kg/m3 were produced for field planting, attending to the
development of proper root formation. The nursery time to achieve adequate quality
plants is from 6 to 24 months, depending on the species.
Regarding the availability of propagation material, the selected trees present
abundant fruiting every 3 years if weather conditions are adequate. Recalcitrant
seeds and/or those that are difficult to store, as well as those of rare, vulnerable
and/or valuable species such as Handroanthus sp., Aspidosperma polyneuron,
Myrocarpus frondosus, Araucaria angustifolia and others, are not usually available
for nurseries. Under those circumstances these species can be stored in nurseries in
the form of living plants grown in rigid containers, for more than 3 years, as a Bank
of Plants (Eibl et al. 2017).
Demonstration trials with native tree species in open plantations, enrichment
planting in degraded forest, and species associated with agroforestry systems
(AFS) form a network of demonstration productive experiences for the region that
include more than 40 species of interest. Survival exceeding 70% and 200–300
plants achieved per ha are expected, given an initial implementation of 400 individ-
ual plants or more (Eibl et al. 2015a).
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 175

The technical certifications at 18 months after plantation allow the producer to

access the benefits of forestation offered by National Law n 26,432. These payments
can be approximately 500 USA $/ha, depending on whether the trees are planted for
enrichment, restoration and/or in pure plantations with native species. According to
National Law n 27,487, incentives also vary for medium (20–40 years) and/or long
term (40–60 years) restoration projects, with time of tree harvest depending on the
species and quality of the desired wood. In addition to the main goals of restoration
practices towards conservation of species, these projects also provide ecological and
productive benefits including mitigation of climate change and wood harvesting,
even at the start of the project (Eibl et al. 2015b).
Results of our recent research attest to the positive impacts of some of these
species on the restoration of degraded land. For example, in areas of degraded
physical and chemical soil conditions, occupied by invasive pastures, 29 years
after the intervention with an initial plantation of 11 native species, there were
62 different tree species represented by 28 families (Eibl et al. 2019b). The propa-
gules that appeared on the site were mainly dispersed by birds, small animals, and
wind (Dohman et al. 2019).
The higher growth and survival of species of economic interest, such as
Anadenanthera colubrina, Bastardiopsis densiflora, Nectandra lanceolata,
Balfourodendron riedelianum, Peltophorum dubium and Machaerium stipitatum,
underscore their importance as tolerant to degraded soil conditions (Eibl et al.
2019c). These findings suggest the possibility of creating new productive islands
of biodiversity from natural regeneration after an initial intervention with a planta-
tion. This can be considered as an alternative in degraded ecosystems where no
natural regeneration occurs. This also points to the need for creating an area of
propagation material using the trees that grew successfully in the restored land.
Meetings were held at the Selva Misionera Botanical Garden (JBSM) as part of
the Argentine Botanic Gardens Network (RAJB 2006), with local farmers, techni-
cians, students, and staff from local institutions. The main purpose was to raise
awareness about the environmental benefits of forests and knowledge of species and
their multiple uses. These actions resulted in higher local commitment with envi-
ronmental actions (Eibl et al. 2016a,b). Based on this initiative, the creation of
botanical gardens in all the municipalities of the province is encouraged in order
to display plants and facilitate the knowledge of species and their current and
potential uses for society, in addition to their inherent role in ex-situ conservation
(Fig. 7.5).
Several field experiences including native species have been established, with
selected seed trees marked at their original site, which will annually provide part of
the needed seeds. Several nurseries are already in production, thus ensuring the
availability of quality plants for plantations that, in combination with several species
of multiple uses, will allow future uses of wood and firewood, which may be
marketed under the certification of sustainable management.
176 B. I. Eibl et al.

Fig. 7.5 Traceability for

certification, from seed
origin and plants of
Araucaria angustifolia to
field plantations. (Photo:
B. Eibl)

7.5 Conclusion

The certification of propagation material obtained from conservation areas, seed

banks, nursery plants and tree plantations grown in the field follows a certification
protocol for quality control that, under Resolution INASE n 318/18, allows native
species to be included in productive systems with a purpose of sustainable manage-
ment in productive plantations, restoration and/or conservation and to provide
income to the owners.
This certification facilitates the care of the sources of seeds (propagation mate-
rial), enable the traceability of the material used in the form of an eco-label, promotes
dissemination and generates information for the work that currently includes pro-
ductive uses (such as firewood, wood, medicinal, ornamental, honey, food, essences,
landscape, restoration, others) for species from local plant diversity.
The establishment of biodiversity islands often necessitates restoration of
degraded lands for which it is crucial to have the appropriate plant propagation
material that can ensure their viability and diversity. The massive use of native
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 177

species for multiple purposes is a contribution to the production, restoration, and

conservation of their diversity.

References

Armenteras D, Espelta JM, Rodríguez N, Retana J (2017) Deforestation dynamics and drivers in
different forest types in Latin America: three decades of studies (1980–2010). Glob Environ
Chang 46:139–147
Barth SR, Eibl BI, Montagnini F (2008) Adaptabilidad y crecimiento de especies nativas en áreas en
recuperación del noroeste de la provincia de Misiones. 13as. Jornadas Técnicas Forestales y
Ambientales. Facultad de Ciencias Forestales, Universidad Nacional de Misiones, y Estación
Experimental Montecarlo del INTA, 5 al 7 de junio Eldorado, Misiones, Argentina. [Link]
[Link]/la-facultad/secretarias/ciencia-tecnica-y-posgrado/jornadas-tecnicas-
forestales-y-ambientales/
COP21 (2015) Convención Marco sobre el Cambio Climático. Acuerdo de las Partes15 de
diciembre de 2015, Paris Francia. [Link]
Accessed 10/2016
COP25 (2019) Argentina entre los países latinoamericanos que se comprometen a restaurar
250 millones de hectáreas al 2050. [Link]
argentina-entre-los-paises-latinoamericanos-que-se-comprometen-a-restaurar-250-millones-de-
hectareas-al-2050/. Accessed Jan 2020
Day S, Montagnini F, Eibl BI (2011) Effects of native trees in agroforestry systems on the soils and
yerba mate in Misiones, Argentina. In: Montagnini F, Francesconi W, Rossi E (eds) Agrofor-
estry as a tool for landscape restoration. Nova Science Publishers, New York, pp 99–112
De Viana ML, Morandini MN, Giamminola EM, Diaz RC (2011) Conservación ex situ, un banco
de germoplasma de especies nativas. Lhavet 1(1):35–41
Di Bitetti MS, Placci G, Dietz LA (2003) Una Visión de Biodiversidad para la Ecorregión del
Bosque Atlántico del Alto Paraná: Diseño de un Paisaje para la Conservación de la
Biodiversidad y prioridades para las acciones de conservación. World Wildlife Fund,
Washington DC. [Link]
Ecorregin-del-Bosque-Atlntico-del-Alto-Paran
Dohmann R, Suárez S, Eibl B, Dummel C, Robledo L, Montagnini F, López M (2019)
Caracterización de grupos ecológicos y formas de dispersión en un área de restauración luego
de 29 años de plantación de especies nativas en la Provincia de Misiones, Argentina. Actas de
las XVIII Jornadas Técnicas Forestales y Ambientales. 17 al 19 de octubre, Eldorado, Misiones,
pp 382–384. [Link]
jornadas-tecnicas-forestales-y-ambientales/
Eibl BI (2006) La estrategia global para la conservación de la biodiversidad al 2010. www.
[Link] edición n 2510, del 21 de mayo del 2006
Eibl BI (2013) Biodiversidad. Semillas y Frutos de la Selva Misionera. Fundación Bosques Nativos
Argentinos para la Biodiversidad. Año 3 N 6. 20 pp, [Link]
Eibl BI, Báez MN (2004) Áreas productoras de semillas de especies nativas de la Selva Misionera.
Actas de las Jornadas de Mejoramiento Genético. Facultad de Ciencias Forestales. Instituto
Nacional de Tecnología Agropecuaria. 14 y 15 de Julio de 2004. Centro de Convenciones.
Posadas, Misiones, pp 22–28
Eibl BI, López MA (2017) Estrategias para la conservación y el uso productivo de especies nativas
del Bosque Atlántico Interior (Selva Paranaense). Conferencia. Actas del IV Congreso Nacional
de Ciencias Agrarias, Facultad de Ciencias Agrarias, Universidad Nacional de Asunción. 19 al
21 de abril, Campus San Lorenzo, Asunción, Paraguay, pp 135–141
178 B. I. Eibl et al.

Eibl BI, Paredes D, Gómez J (2001) Gigantes de Misiones. Comunicación Revista Forestal
Yvyrareta 10:87–88
Eibl BI, Méndez RM, Di Stassi M, Bohren AV, Sosa G, Robledo F (2002) Selección de árboles
semilleros en propiedades y reservas de la selva misionera. Trabajo n 60 en formato CD. Actas
de las IX Jornadas Técnicas Forestales. 15 al 17 de Mayo de 2002. Facultad de Ciencias
Forestales, Universidad Nacional de Misiones, Eldorado, Misiones
Eibl BI, González C, Otegui M, Dreyer N (2012) Protocolos tentativos para la propagación de
20 especies nativas de interés productivo de la selva misionera.15as Jornadas Técnicas
Forestales y Ambientales. Facultad de Ciencias Forestales, UNaM – EEA Montecarlo, INTA.
7, 8 y 9 de Junio de 2012, Eldorado, Misiones, Argentina. [Link]
facultad/secretarias/ciencia-tecnica-y-posgrado/jornadas-tecnicas-forestales-y-ambientales/
Eibl BI, González C, Otegui M, Dreyer N, Mallozi G, Weinberger V (2013a) Conservación de
semillas de especies nativas en bancos de germoplasma. Resumen Actas, p. 45. Trabajo
completo n 13CA en CD. Jornadas Científico Tecnológicas 40 Aniversario UNAM, 15–17
mayo 2013, Posadas. Misiones
Eibl BI, González C, Dreyer N, Branco F, Bohren AV (2013b) Indicadores de calidad de plantas en
vivero para el establecimiento de ensayos de producción de biomasa para energía con especies
nativas. Resumen Actas, pp. 46. Trabajo completo n 14CA en CD. Jornadas Científico
Tecnológicas 40 Aniversario UNAM.15 al 17 de mayo de 2013. Posadas. Misiones
Eibl BI, López MA, Bohren AV, Montagnini F, López LN, Barbaro L, Sforza O (2015a)
Restauración de áreas mediante plantaciones de especies nativas con fines de biomasa leña y
maderables en la Provincia de Misiones. IV Congreso Iberoamericano y del Caribe de
Restauración Ecológica. Libro Actas M231, pp 131. SIACRE. 12–16 de Abril 2015, Buenos
Aires, Argentina
Eibl BI, Montagnini F, López MA, Montechiesi R, Barth SR, Esterche E (2015b) Ilex
paraguariensis A. St.-Hil., yerba mate orgánica bajo dosel de especies nativas maderables,
una propuesta de producción sustentable. In: Montagnini F, Somarriba E, Murgueitio E,
Fassola H, Eibl B (eds) Sistemas Agroforestales. Funciones productivas, socioeconómicas y
ambientales. Serie Técnica Informe Técnico 402, CATIE, Turrialba, Costa Rica. Fundación
CIPAV. Cali, Colombia, pp 158–177
Eibl BI, López MA, Barbaro L, Sforza O, López J, Branco F, López M, Stadler N, Méndez R,
Steitzer N (2016a) Experiencias productivas con especies nativas multipropósito en áreas de
restauración ambiental y estrategias para la conservación de remanentes de bosques. Actas XVII
Jornadas Tecnicas Forestales y Ambientales. 17 al 19 de agosto 2018. Posadas, Misiones. http://
[Link]/la-facultad/secretarias/ciencia-tecnica-y-posgrado/jornadas-tecnicas-
forestales-y-ambientales/
Eibl BI, Rocha SP, González C, González CA, Silveira T, Krauczuk E, Suárez S, Hancherek FA
(2016b) El Jardín Botánico Selva Misionera, una herramienta para la concientización ambiental.
Actas de las XVII Jornadas Técnicas Forestales y Ambientales. 17 al 19 de octubre. Posadas,
Misiones, [Link]
jornadas-tecnicas-forestales-y-ambientales/
Eibl BI, López MA, Stadler N, López M, González C (2017) Plantaciones productivas de Araucaria
angustifolia (Bertol.) Kuntze utilizando progênies de dos orígenes de la Provincia de Misiones.
Actas del IV Congreso Nacional de Ciencias Agrarias, Facultad de Ciencias Agrarias
Universidad Nacional de Asunción. Formato CD. 19 al 21 de abril, Campus San Lorenzo,
Asunción, Paraguay, pp 469–472
Eibl BI, González C, Lecanda Prado A, Ortega MN, Rodríguez GA, Kiefl JM (2019a) Germinación
de Aspidosperma polyneuron Müll. Arg., luego del secado y el almacenamiento de las semillas.
Actas de las XVIII Jornadas Técnicas Forestales y Ambientales. 17 al 19 de octubre, Eldorado,
Misiones, pp 319–321. [Link]
posgrado/jornadas-tecnicas-forestales-y-ambientales/
Eibl BI, López MA, Montagnini F, Suárez S, Dohman R, Dummel CJ, Robledo L (2019b)
Diversidad de especies arbóreas luego de 29 años de la plantación de especies nativas en un
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 179

área de restauración en la Provincia de Misiones, Argentina. Actas de las XVIII Jornadas

Técnicas Forestales y Ambientales. 17 al 19 de octubre, Eldorado, Misiones, pp 307–309,
[Link]
tecnicas-forestales-y-ambientales/
Eibl BI, López MA, Montagnini F, Suárez S, Dohman R, Dummel C, Robledo L (2019c)
Crecimiento de especies arbóreas luego de 29 años de la plantación de especies nativas en un
área de restauración en la Provincia de Misiones, Argentina. Actas de las XVIII Jornadas
Técnicas Forestales y Ambientales. 17 al 19 de octubre, Eldorado, Misiones, pp 310–312.
[Link]
tecnicas-forestales-y-ambientales/
FAO (2018) The state of the World´ s forests-forest pathways to sustainable development. http://
[Link]/documents/card/en/c/I9535EN/
FAO (2020) Integrating the SDGs: showing contributions by forests and trees to the 2030 Agenda.
Sustainable Forest Management (SFM) Toolbox. [Link]
management/toolbox/sfm-home/detail-events/en/c/1145494/
Flora del Conosur (2020) Instituto de Botánica Darwinion. [Link]
FloraArgentina/[Link]. Accessed on Jan 2020
Galindo-Leal C, Câmara IDG (2003) Atlantic forest hotspot status: an overview. In: Galindo-
Leal C, IDG C (eds) Conservation International/Center for Applied Biodiversity Science. The
Atlantic Forest of South America: biodiversity status, threats, and outlook. Island Press,
Washington DC, pp 3–11
Gartland HM, Bohren AV (2008) Consideraciones sobre la biodiversidad forestal a nivel de
especies arbóreas, en la selva paranaense de Misiones Argentina. Revista Forestal Yvyrareta
15:39–49
Gibbs HK, Ruesch AS, Achard F, Clayton MK, Holmgren P, Ramankutty N, Foley JA (2010)
Tropical forests were the primary sources of new agricultural land in the 1980s and 1990s. Proc
Natl Acad Sci U S A 107:16732–16737
Gold K, León-Lobos P, Way M (2004) Manual de recolección de semillas de plantas silvestres para
conservación a largo plazo y restauración ecológica. Instituto de Investigaciones Agropecuarias,
Centro Regional de Investigación Intihuasi, vol 110. Boletín INIA, La Serena, Chile, p 62
González C, Eibl BI, Dreyer NB (2015) Almacenamiento de semillas de Cabralea canjerana
(Vell.) Mart. In: Actas versión Digital. Grupo forestales. Congreso Nacional de viveros cítricos,
forestales y ornamentales. INASE, 4 al 6 de agosto de 2015, Posadas, Misiones
GSPC (2020). Global Strategy for Plant Conservation. [Link]
INASE (1999) Resolución n 256/99. Normas para la certificación, producción e importación de
semillas de especies forestales. [Link]
56-1999
INASE (2000) Resolución n 42/2000. Inscripción en el Registro Nacional de Comercio y
Fiscalización de semillas y proceso de rotulación de semillas. [Link]
normativa/nacional/resolucion42-2000
INASE (2018) Resolución n 318/18. Registro de Área Productora de Semillas de Especies Nativas
(APSEN) [Link]
ISTA (2019) International Rules for Seed Testing. International Seed Testing Association (ISTA).
Zürichstr. 50, CH-8303. Bassersdorf, Switzerland. 300 pp
IUCN (2020a) Red list of threatened species. [Link]
iucn-red-list-threatened-pecies#RL_categories. Accessed Feb 2020
IUCN (2020b) Soluciones basada en naturaleza. UICN Sur. [Link]
commission-ecosystem-management/soluciones-basadas-en-la-naturaleza. Accessed Jan 2020
Keller HA (2007) Solanum vaillantii (Solanaceae), nueva cita para la flora Argentina. Bonplandia
16(3–4):275–278
Ley Nacional n 26331 (2007) Bosques Nativos. Presupuestos Mínimos de Protección Ambiental de
los Bosques Nativos. Boletín Oficial Diciembre 26/2007. SAIJ, Sistema Argentino de
Información Jurídica. Ministerio de Justicia, Buenos Aires, Argentina
180 B. I. Eibl et al.

Ley Nacional n 27487 (2008) Prorroga y Reforma de la Ley n 25080. Decreto Argentina, Boletín
Oficial. 29/12/2008. Resolución134/2019-APN-SECAGYP. SAIJ Sistema Argentino de
Información Jurídica. Ministerio de Justicia. Buenos Aires, Argentina
Ley Provincial de Energías Renovables XVI n 97 (ex Ley n 4439) (2010) Marco regulatorio de los
recursos energéticos renovables. Provincia de Misiones. [Link]/
digesto_juridico
Ley Provincial XVI N 105 (ex ley n 3260) (2010) Provincia de Misiones el ordenamiento de los
Bosques Nativos y los mecanismos a implementar en la para la Conservación, de los Bosques
Nativos y el Régimen de Promoción de Manejo Sostenible. [Link]/
digesto_juridico
Ley Provincial XVI n 29 (ex Ley n 2932) (2010) Áreas Naturales Protegidas. Provincia de
Misiones. [Link]/digesto_juridico
Ley Provincial XVI N 47 (ex ley n 3337) (2010). Conservación y aprovechamiento sostenible de la
diversidad biológica y sus componentes. [Link]/digesto_juridico
Ley Provincial XVI n 53 (ex Ley n 3426) (2010) Bosques protectores de cursos de agua y
pendientes. [Link]/digesto_juridico
Ley Provincial XVI n 60 (ex Ley n 3631) (2010) Área Integral de Conservación y Desarrollo
Sustentable Corredor Verde. Provincia de Misiones. [Link]
juridico
Ley Provincial XVI N 98 (ex ley n 4464) (2010) Banco Provincial de Germoplasma de Misiones.
[Link]
López MA, Eibl BI, Palavecino J (2013) Valoración de especies nativas en áreas de restauración
ambiental, Provincia de Misiones, República Argentina. En: Educacao Ambiental: Premisa
inafastavel ao desenvolvimento económico sustentavel. Editores Joao Almeida e Da Silva
Almeida. Ed. Lumen Juris, Rio de Janeiro pp 371–391
Ministério da Agricultura (2009) Regras para análise de sementes. Ministério da Agricultura,
Pecuária e Abastecimento, Secretaria de Defesa Agropecuária, Mapa/ACS, Brasília, Brasil.
399 pp
Mittermeier RA, Turner WR, Larsen FW, Brooks TM, Gascon C (2011) Global biodiversity
conservation: the critical role of hotspots. In: Zachos FE, Habel JC (eds) Biodiversity hotspots.
Distribution and protection of conservation priority areas. Springer, Berlin, Heidelberg, pp 3–22
Montagnini F, Cusack D, Petit B, Kanninen M (2005) Environmental services of native tree
plantations and agroforestry systems in Central America. J Sustain For 21(1):51–67
Montagnini F, Eibl BI, Fernandez R (2006) Rehabilitation of degraded lands in Misiones, Argen-
tina. Bois et Forests des Tropiques 288:51–65
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity Islands. Strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human-dominated environments. Topics in Biodiversity and Conservation,
Springer, Cham, pp 1–35
Niella F, Eibl B, Rocha P, Dummel C, Keller HA (2016) Estrategia para la conservación,
domesticación y rescate de recursos fitogenéticos nativos a la selva paranaense. XVII Jornadas
Técnicas Forestales y Ambientales, Posadas, Misiones. [Link]
modules/uploads/2017/03/Acta_de_Resumenes_XVII_Jotefa_2016.pdf
Niella F, Eibl B, Keller H, Maiocco D, Rocha P, Vega V (2017) Red de árboles y áreas semilleras
para la conservación, domesticación y rescate de recursos fitogenéticos nativos de la Selva
Paranaense. SILVA N 17/2013. Conservación de la Biodiversidad en Paisajes Productivos.
GEF 090118. Ministerio de Agricultura, Ganadería y Pesca. [Link]
areas/proyectos_forestales/biosilva/SILVA%20017%[Link]
Ocampo T, Eibl BI, González C (2015) Indicadores de calidad a partir de variables morfo-
fisiológicas en frutos, semillas y plantas de Cedrela fissillis Vell. En: Actas versión Digital.
Grupo forestales. Congreso Nacional de viveros cítricos, forestales y ornamentales. INASE, 4 al
6 de agosto de 2015, Posadas, Misiones
7 Conservation and Registration of Seed Sources in Reserve Remnants in the. . . 181

Otegui MB, Ojeda DS, Totaro ME, Eibl BI, Valdés MB, Mallozzi GY (2016) Comportamiento y
Longevidad de Aspidosperma polyneuron Müll. Arg. frente al Ultrasecado. Seed conservation
of palo-rosa. RECyT 18(25):15–19
RAJB (2006) Red Argentina de Jardines Botánicos. Boletín Año 2(3) [Link]
documentos/red-argentina-de-jardines-botanicos-rajb
Rodríguez G, González C, Eibl BI, González G, Grance L (2019) Viabilidad de semillas de Plinia
rivularis (Cambess.) Rotman durante el almacenamiento. Actas de las XVIII Jornadas Técnicas
Forestales y Ambientales, 17 al 19 de octubre, Eldorado, Misiones, pp 328–330. [Link]
[Link]/la-facultad/secretarias/ciencia-tecnica-y-posgrado/jornadas-tecnicas-
forestales-y-ambientales/
Schiaffino K, Bertolini MP (2016) Reservas Naturales Privadas, avances de un trabajo en Red.
Conferencia. Actas de las XVII Jornadas Técnicas Forestales y Ambientales, 17–19 de agosto,
Posadas, Misiones. [Link] [Link]
facultad/secretarias/ciencia-tecnica-y-posgrado/jornadas-tecnicas-forestales-y-ambientales/
Thomas E, Jalonen R, Loo J, Boshier D, Gallo L, Cavers S, Bordacs S, Smith P, Bozzano M (2014)
Genetic considerations in ecosystem restoration using native tree species. For Ecol Manage 333:
66–75
Ugarte-Guerra J, Alemán F, Nieto V, Prado L, Galvan O, Silva I, Ohashi S, García W (2010)
Estudio de las cadenas de abastecimiento de germoplasma forestal en cinco países amazónicos.
World Agroforestry Centre (ICRAF), Lima, Perú
Young CF (2003) Socieconomic causes of deforestation in the Atlantic forest of Brazil. In:
Conservation International/Center for Applied Biodiversity Science (author). In: Galindo-
Leal C, de Gusmao CI (eds) The Atlantic Forest of South America: biodiversity status, threats,
and outlook. Island Press, Washintong/Covelo/London, pp 103–117
 Part III
Biodiversity Islands Across the Globe:
Case Studies
Chapter 8
Islands of Forests Among Savannas: Key
Elements for Conservation and Production
in the Paraguayan Humid Chaco

Rafaela Laino, Karim Musalem, Luis Domingo Laino,

Andrea Caballero-Gini, Diego Bueno-Villafñae, Lidia Aranda,
Alberto Esquivel, Marcela Ferreira Riveros, Lía Romero Nardelli,
Nicolás Cantero, and Rebeca Irala

R. Laino (*) · L. D. Laino · L. R. Nardelli

American Chaco Research Center, Benjamín Aceval, Paraguay
e-mail: info@[Link]; [Link]
K. Musalem
American Chaco Research Center, Benjamín Aceval, Paraguay
WWF Paraguay, Asunción, Paraguay
[Link]
A. Caballero-Gini · D. Bueno-Villafñae
American Chaco Research Center, Benjamín Aceval, Paraguay
Instituto de Investigación Biológica del Paraguay, IIBP, Asunción, Paraguay
L. Aranda
American Chaco Research Center, Benjamín Aceval, Paraguay
Universidad Nacional de Asunción, San Lorenzo, Paraguay
A. Esquivel
WWF Paraguay, Asunción, Paraguay
Wildlife Paraguay, Luque, Paraguay
M. F. Riveros
American Chaco Research Center, Benjamín Aceval, Paraguay
Instituto de Investigación Biológica del Paraguay, IIBP, Asunción, Paraguay
Guyra Paraguay, Asunción, Paraguay
[Link]
N. Cantero
Wildlife Paraguay, Luque, Paraguay
Universidad Nacional de Córdoba, Córdoba, Argentina
R. Irala
Wildlife Paraguay, Luque, Paraguay
[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 185
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
186 R. Laino et al.

Abstract This chapter provides information on islands of forests immersed in

flood-prone savannah-dominated landscapes for biodiversity conservation and pro-
ductive activities. Our study area in the Paraguayan Chaco is in a region dedicated to
low intensity cattle rearing, with native grasslands as forage within a mosaic of
natural plant formations. The region is highly diverse due to the variety of plant
formations and the great amount of wild fauna. We present results of research
projects conducted by the American Chaco Research Center, a Paraguayan research
initiative. These results suggest that extensive or semi-extensive ecosystem-based
livestock production could be an opportunity to maintain ecosystem services,
including biodiversity, in the region via a production system that is highly compat-
ible with the conservation of forest species. Adverse environmental conditions for
production (such as floods from rain and periodic floods of the Paraguay River)
underscore the importance of maintaining the regulating functions of natural eco-
systems, which leads to an opportunity for the conservation of threatened biodiver-
sity. The mosaic of forest islands, flood-prone palm savannas, and wetlands is
fundamental for water quality and hydrological regulation as it reduces the intensity
of the effects of flooding on neighboring areas, as well as for carbon sequestration.
Inclusion of live fencing and the maintenance of continuous areas of intact or lightly
disturbed vegetation can increase habitat connectivity, provide a barrier to manage
fires, and serve as a refuge for animals when fires occur. These native ecosystems are
at high risk of disappearing due to the intensification of agriculture and livestock
production, as well as the expansion of urban areas, so strategies for their conserva-
tion, including tailor-made incentives are needed.

Keywords Biodiversity conservation · Ecosystem services · Floodable forest ·

Habitat connectivity · Livestock farming · Wildlife

8.1 Introduction

8.1.1 Forests as Islands Immersed in Flood-Prone

Savannas

This chapter describes the relevance of forest islands as part of a mosaic of savannas,
palm groves and wetlands (an image of the study area is shown in Fig. 8.1).
The presence of any of these formations is determined mainly by the geomorphology
of its terrain. The landscape includes plains and depressions where water settles for
different periods of time according to weather conditions (Mereles et al. 2020a).
Wetlands occupy the lowest areas, generally with permanent waters where species
linked to water develop with different life forms: floating, submerged (free or not)
and rooted in the mud of the bottom. Some of the plant species recorded in the
wetlands of the study area are Cyperus giganteus, Typha domingensis and Sagittaria
montevidensis, with Cyperus giganteus being the most abundant and frequent
(Macedo 2018). Palm groves are monotypical formations that can flood frequently.
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 187

Fig. 8.1 Image of the study site in the Humid Chaco ecoregion. Dense forest island immersed in
extensive savannas. (Photo: Gianfranco Mancusi)

The characteristic species is the native palm locally known as Karanday (Copernicia
alba), accompanied by a rich herbaceous stratum, whose density varies according to
the presence of water (Mereles and Rodas 2014; Mereles et al. 2013).
The landscape consists of dense forest islands immersed in savannas, palm
groves and wetlands, resulting in different forest types: Dense Subtropical Forest,
which occurs naturally on islands associated with palm groves in the floodplain of
the Paraguay River (Pérez de Molas 2015); Dense Mesoxerophytic Forest, with Red
Quebracho (Schinopsis balansae) being the dominant tree species (Mereles et al.
2020a); and Riparian Forest, which develops adjacent to the riverbeds forming strips
50–100 meters wide (Maturo et al. 2005; Peña-Chocarro et al. 2006). Main arboreal
tree species found in the area are shown in Table 8.1.
This chapter is based on field studies that were conducted in rangelands located
on the right bank of the Paraguay River (Fig. 8.2), where average rainfall is
1200 mm/year and the average temperature is 24
C (Mereles et al. 2013). During
some years, with the occurrence of the El Niño phenomenon, intense rainfall causes
generalized flooding. In other years, drought is accentuated and prolonged
(Ginzburg and Adámoli 2006; Junk et al. 2013). The study site itself has experienced
both of these extremes (most recently in 2015 and 2019): over half of its surface area
remained flooded during one part of the year and then suffered an extreme drought in
the same year. The pulsing water-level in the rainy season and the pronounced dry
and wet periods create an aquatic-terrestrial transition zone where important ecolog-
ical processes occur (Mereles et al. 2020b). Most of the studies were conducted in
nearby ranches, which are located within the Key Biodiversity Area 22 La Rafaela
(Cartes and Clay 2009), and which have been identified as potential birding tour
areas (Mamede et al. 2019) and as important corridors for connectivity of the Great
American Chaco (Mereles et al. 2020b).
188 R. Laino et al.

Table 8.1 Forest types and tree species recorded in dense forest islands of the study area
Forest type Tree species Source
Dense subtropical Peltophorum dubium, Enterolobium contortisiliquum, Ficus El Raiss
forest enormis, Ocotea diospyrifolia, Sapium haematospermum, (2014)
Gleditzia amorphoides, Guazuma ulmifolia, Chloroleucon
tenuiflorum, Handroanthus heptaphyllus, Syagrus
romanzoffiana, Copernicia alba.
Dense Schinopsis balansae, Rollinia emarginata, Aspidosperma Lubián
mesoxerophytic quebracho blanco, Forsteronia sp., Tabernaemontana (2014)
forest catharinensis, Syagrus romanzoffiana, Acrocomia aculeata,
Copernicia alba, Tabebuia nodosa, Handroanthus
heptaphyllus, Tabebuia aurea, Cordia americana, Carica
papaya, Cecropia pachystachya, Celtis sp., Terminalia
triflora, Sapium longifolium, Gleditsia amorphoides,
Parapiptadenia rigida, Albizia niopoides, Enterolobium
contortosiliquum, Prosopis affinis, Inga uraguensis,
Peltophorum dubium, Copaifera langsdorfii, Pterogyne nitens,
Vitex megapotamica, Ocotea diospyrifolia, Trichilia catigua,
Trichilia pallida, Maclura tinctoria, Sorocea sprucei, Psidium
guajava, Myrcianthes pungens, Genipa americana,
Calycophyllum multiflorum, Zanthoxylum petiolare,
Zanthoxylum riedelianum, Casearia sylvestris, Diplokeleba
floribunda, Sapindus saponaria, Chrysophyllum marginatum,
Guazuma ulmifolia, Seguieria paraguayensis, Ruprechtia
laxiflora, Phyllostylon rhamnoides.
Riparian forest Ocotea diospyrifolia, Lonchocarpus sp., Terminalia triflora, Macedo
Celtis sp., Peltophorum dubium, Nectandra angustifolia, (2018)
Copernicia alba, Guazuma ulmifolia, Zanthoxylum petiolare,
Machaonia spinosa, Xylosma venosa, Inga uraguensis,Vitex
sp., Genipa americana, Pouteria glomerata, Albizia inundata,
Chrysophyllum gonogocarpum, Zigia sp., Vitex
megapotamica, Chrysophyllum marginatum, Enterolobium
contortisiliquum, Sorocea sprucei, Myrsine sp.

The chapter begins with a description of the study site explaining why these
forests exist as islands immersed in flood-prone savannas. The next part focuses on
ecosystem services such as carbon sequestration and water quality and describes the
native fauna housed within the forest islands and surroundings. In the following
sections, we present information about the role that these biodiversity islands play in
functional connectivity and as fire-breaks in the savanna. Finally, we emphasize the
benefits of forest islands to cattle ranching in native grasslands and the challenges for
their conservation.
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 189

Fig. 8.2 Study site in the Humid Chaco ecoregion. Landscape is dominated by naturally occurring
savannas, palm groves, wetlands and dense forests

8.2 Ecosystem Services Evaluated in the Study Site

8.2.1 Carbon Sequestration and Water Quality

Measurements of carbon storage were made in the dense forest islands, in the
surrounding savannas and palm groves, and in the wetland soils. Lubián (2014)
determined 254 tons of CO2e/ha in the forest and 24.5 tons of CO2e/ha in the palm
groves and grasslands. Another evaluation focusing on the forest islands (Boródn
2015) calculated 175.8 tons CO2e/ha (following IPCC 2005 report) and 291 tons
CO2e/ha (following Sato et al. 2014 equation) for the aboveground biomass, and
103.9 tons/ha for forest carbon soil. For wetland soil, Brun (2013) reported average
values of 3.15 tons of CO2/ha.
Wetlands are among the most productive primary ecosystems on the planet,
depending on the hydrological regime, i.e., how often they are flooded and how
long they remain flooded (Kandus et al. 2010). Water originates in the lowest areas,
between savannas, marshlands and swamps with high productivity of the herbaceous
stratum (Benzaquen et al. 2017). The herbaceous cover of the wetlands decreases
water speed and facilitates the sedimentation and retention of suspended materials,
thus improving water quality (Kandus et al. 2010). Wetlands of the study area were
190 R. Laino et al.

of ‘good’ qualitative conservation status according to the ECELS index1 evaluated

by Ferreira (2018) and Brun (2013). In addition, exploratory analysis of water
quality (including pH, Electrical Conductivity, Suspended Solids, Nitrate, Total
Phosphorus, Total Nitrogen, Chloride, Sulphate, Biochemical Oxygen Demand,
Chemical Oxygen Demand, Coliforms and water temperature) found no relevant
pollution in wetlands and rivers of the study area, except for specific sites where the
flow decreased in the dry season (Chaparro 2014, PINV 15-143 2018).
In addition to maintaining water quality, the mosaic of forest islands, flood-prone
palm savannas and wetlands is fundamental for hydrological regulation as it reduces
the intensity of the effects of flooding on neighboring areas. Although these ecosys-
tems do not prevent flooding, they reduce the river’s flood peaks, retain excess
runoff after rainfall and release it slowly afterwards, and encourage aquifer recharge
(Kandus et al. 2010).

8.2.2 Wildlife Housed in the Forest Islands and Surroundings

The study of amphibians, reptiles, mammals, and birds was carried out inside of the
different forest islands (FI) and surrounding savannas (S), palm groves (PG) and
wetlands (W) through different sampling methods.
Amphibian surveys were made using three different sampling techniques, as
these vary in their effectiveness depending on intrinsic species characteristics (Ali
et al. 2018). The three sampling methods used from August 2017 to June 2018 were
visual encounter surveys (Crump and Scott 1994), pitfall traps with drift fences
(Corn 1994) and PVC pipes of 40 mm of diameter as refuge for tree frog species
(Boughton et al. 2000).
All together we registered 2449 individuals corresponding to 29 species included
in the families: Bufonidae, Hylidae, Leptodactylidae, Phyllomedusidae, Microhylidae
and Odontophrynidae, which represent 48% of all the species registered in the
Humid Chaco ecoregion (Brusquetti and Lavilla 2006; Frost 2020). Some of these
species are shown in Fig. 8.3. Regarding their conservation status, all species found
are categorized as Least Concern (LC) at international (IUCN 2019) and national
levels (Motte et al. 2019), except for the Rio Grande Dwarf Frog (Physalaemus
riograndensis) (Motte et al. 2019). This species is considered to be in the data
deficient category (DD) at the national level due to its scarce records (Table 8.2),
which are mostly from the southern region of the country (Brusquetti and Lavilla
2006; Motte et al. 2019; Frost 2020). In addition, we obtained the first depart-
mental record for the White spotted Humming frog (Chiasmocleis albopunctata),

1
ECELS (Estado de Conservación de Ecosistemas Lénticos Someros Index) is a methodological
tool used to determine the ecological status of wetlands, which was developed by Agencia Catalana
del Agua (ACA 2004).
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 191

Fig. 8.3 Amphibian species registered at the study area. A. Monkey frog (Pithecopus azureus).
B. Polka-dot Tree Frog (Boana punctata). C. Male (on the top) and female of Dwarf tree-frog
(Dendropsophus nanus). D. Uruguay Arlequin Frog (Lysapsus limellum). E. White lipped-tree frog
(Trachycephalus typhonius). F. Two colored Oval frogs (Elachistocleis bicolor). G. Common
192 R. Laino et al.

a rare species mainly due to its fossorial habits,2 size and mimetic coloration. Only
one individual was registered during the study time frame through the pitfall trap
method. Nevertheless, the record is interesting since it extends the distribution range
of the species 57 km northwest from the nearest locality at the Surubi’i Urbanization,
Central Department (Aquino et al. 2004; Brusquetti and Netto 2008).
Through pitfall traps, visual encounter surveys and camera traps, we registered a
total of 12 reptile species including snakes, lizards of the families Tupinambidae and
Teiidae, and the Jacare caiman (Caiman yacare). Individuals of Jacare caiman were
actively searched for at night, and we found them mostly isolated and using different
water body types, including cattle ponds. When isolated sections of the wetlands
were searched, we found juvenile aggregations along with adults, which indicates a
preference in habitat use and reproductive success. Further research will improve our
knowledge about the species habitat use, reproductive sites and its estimated density
in the area, which harbors great potential for reproduction and conservation of Jacare
caiman. Another great lizard found was the Black-and-white Tegu (Salvator
merianae), which was recorded frequently with camera traps.
Fifteen mammal species were registered using camera traps that were placed in
different types of environments in the area for three years. We obtained records of
species categorized as vulnerable at the national level (APM and SEAM 2017),
mainly as a consequence of fragmentation and habitat conversion associated with
agricultural and livestock activities, illegal hunting, and road run overs, among
others (Table 8.2). We found the Giant anteater (Myrmecophaga tridactyla) and
even the Maned wolf (Chrysocyon brachyurus), which is typically an elusive species
(Mujica 2014). Furthermore, we obtained records of Puma (Puma concolor) through
camera traps. Although this species is categorized as least concern (APM and SEAM
2017), it still has conservation problems and its populations are decreasing (Nielsen
et al. 2015). The Azara’s night monkey (Aotus azarae), the Black and Gold howler
monkey (Alouatta caraya) and the four-eyed gray opossum (Philander opossum)
were also registered through night sightings. Presence of Jaguar (Panthera onca)
was confirmed with camera traps on November 2020, after this chapter closed
edition. Some of the registered mammal species are shown in Fig. 8.4.
The Humid Chaco harbors more than 430 species of birds, which represents
~60% of the avifauna of Paraguay (Del Castillo 2019). Specifically, in the Key
Biodiversity Area 22 La Rafaela and nearby ranches, numerous census and surveys
have been carried out by several ornithologists and researchers throughout the last
decades. We compiled records obtained during the 2000–2020 period from available

Fig. 8.3 (continued) Lesser Escuerzo (Odontoprhynus americanus). H. Klappenbachmmmmms

Red-bellied Toad (Melanophryniscus klappenbachi) in amplexus. (Photographs: A - D, F:
A. Caballero-Gini. E, G, H: M. Ferreira)

2
Species that are adapted to digging and living in burrows.
Table 8.2 Fauna species of national and global conservation concern, with details of habitat and breeding status
Habitat and Conservation status Conservation status Breeding
Class Common name Scientific name ecology Paraguay IUCN status
Amphibia Rio Grande dwarf frog Physalaemus S DD LC –
riograndensis
Birds Greater Rhea Rhea americana FI, S, PG – NT BR
Bare-faced curassow Crax fasciolata FI Threatened VU BR
Turquoise-fronted Amazona aestiva FI, S, PG – NT BR
parrot
Bearded Tachuri Polystictus pectoralis S, PG Threatened NT BR
Sharp-tailed tyrant Culicivora caudacuta S Endangered VU BR
Dinellimmmmms Pseudocolopteryx W – NT AM
Doradito dinelliana
Strange-tailed tyrant Alectrurus risora S Endangered VU BR
Dark-throated Sporophila ruficollis S – NT BN, AM
seedeater
Rufous-rumped Sporophila hypochroma S – NT BN, AM
seedeater
Mammalia Giant anteater Myrmecophaga FI, W Threatened VU –
tridactyla
Maned wolf Chrysocyon brachyurus S, PG Threatened NT –
8 Islands of Forests Among Savannas: Key Elements for Conservation. . .

Neotropical otter Lontra longicaudis W Least concern NT –

Azarammmmms agouti Dasyprocta azarae FI Least concern DD –
Habitat and ecology: S, Savannas; FI, Forest islands; PG, Palm groves; W, Wetlands
Conservation status Paraguay (APM and SEAM 2017; MADES 2019; Motte et al. 2019): Threatened, “Amenazada de extinción”; Endangered, “En peligro
de extinción”; DD, Data Deficient
Conservation status IUCN (IUCN 2019): LC, Least Concern; NT, Near Threatened; VU, Vulnerable; DD, Data Deficient
Breeding status (Guyra Paraguay 2004): BR, Breeding permanent resident; BN, Breeding resident but northern austral migrant; AM, Austral migrant
193
194 R. Laino et al.

Fig. 8.4 Mammals registered in the Humid Chaco (Paraguay) through camera traps and visual
encounters. A. Giant anteater (Myrmecophaga tridactyla) carrying its cub. B. Gray Brocket
(Mazama gouazoubira). C. Puma (Puma concolor). D. Lesser Capybara (Hydrochoerus
hydrochaeris). E. Collared peccary (Pecari tajacu). F. Neotropical otter (Lontra longicaudis).
G. South American Coati (Nasua nasua). H. Gold howler monkey (Alouatta caraya). (Photographs:
A - G: camera traps installed during 2017 to 2018. H: K. Musalem)
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 195

data in the eBird online platform (eBird 2020a, b), Guyra Paraguay’s Biodiversity
Database ([Link]) and personal checklists, obtaining a total of 249 spe-
cies included in 51 families, which represents 35% of the species of Paraguay and
58% of the species registered at the Humid Chaco ecoregion.
Habitat preferences of these species, based on our records in the area, have been
identified for four main ecosystems: forest islands (FI), savannas (S), palm groves
(PG) and wetlands (W). A total of 125 species (51% of the total species of the area)
use FI as one of their habitats, and 82 species (33%) depend mainly on this habitat.
Nine bird species are globally threatened or near threatened (IUCN 2019), four of
which are also nationally threatened or endangered (MADES 2019). Bare-faced
curassow (Crax fasciolata), Sharp-tailed tyrant (Culicivora caudacuta) and Strange-
tailed tyrant (Alectrurus risora) are categorized as Vulnerable (IUCN 2019), and the
former depends strictly on forest island habitats in the area (Table 8.2). At the
national level, the Bare-faced curassow and the Bearded tachuri (Polystictus
pectoralis) are categorized as Threatened species, while the Sharp-tailed and
Strange-tailed tyrant are Endangered species. Moreover, four species are endemic
of the Chaco (Guyra Paraguay 2004): Chaco chachalaca (Ortalis canicollis), Cream-
backed woodpecker (Campephilus leucopogon), Dinelli’s doradito
(Pseudocolapterix dinelliana) and Black-capped warbling-finch (Microspingus
melanoleucus). Forty-one species are migratory. Six of those are austral migrants,
found mainly during austral winter. Twenty-two are northern austral migrants, which
breed in Paraguay but are less abundant or absent during the winter. Seven are
southern austral migrants, breeding also in Paraguay, but increasing in numbers
during the winter. Some bird species observed in the study area are shown in
Fig. 8.5.

8.3 Role of Biodiversity Islands in Functional Connectivity

at Local and Regional Levels

At a regional level, a recent study identified our study area as part of priority
biological corridors relevant for connectivity of the Great American Chaco, an
ecoregion that extends through Paraguay, Argentina and Bolivia (Mereles et al.
2020b). The study highlights the importance of biological corridors to maintain a
long-term vision of biodiversity, which maintains the connection between key areas
for conservation. Loss of continuous areas can lead to changes in the structure and
function of the remaining fragments (Lindenmayer and Fischer 2006). One of the
problems caused by habitat reduction and fragmentation is a contraction in popula-
tion size, along with increases in inbreeding and the consequent decrease of genetic
diversity (Shaffer 1990).
Although there is considerable uncertainty regarding how fast species respond to
habitat loss, and how time-delayed responses vary in space (Semper-Pascual et al.
2018), we found evidence linked to population reduction sizes and fragmentation of
196 R. Laino et al.

Fig. 8.5 Birds registered in the Humid Chaco, Paraguay. A. Scarlet-headed Blackbird
(Amblyramphus holosericeus). B. Strange-tailed Tyrant (Alectrurus risora). C. Crimson-crested
Woodpecker (Campephilus melanoleucos). D. Black-collared Hawk (Busarellus nigricollis).
E. Red-billed Scythebill (Campylorhamphus trochilirostris). F. Toco toucan (Ramphastos toco).
G. Blue-crowned Trogon (Trogon curucui). (Photographs: A - B: N. Cantero. C - E, G:
A. Esquivel. F: A. Merenciano)
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 197

habitat at Chaco and other regions (e.g., Gómez Fernández et al. 2016; Crooks et al.
2017; Zastavniouk et al. 2017; Pereyra et al. 2019). For example, according to
Rodrigues et al. (2008), the main causes of decline of the Giant anteater populations
are the reduction, deterioration and fragmentation of habitats. Habitat loss may lead
to a decrease in population size and isolation among remnant populations. In this
sense, Collevatti et al. (2007) warned that the population of Myrmecophaga
tridactyla in Emas National Park (Brazil) has a low level of genetic diversity and a
high level of inbreeding.
The Paraguayan Chaco is undoubtedly highly diverse due to the variety of plant
formations and the great amount of wild fauna that remains (Mereles et al. 2020b).
At a local level, the Humid Chaco ecoregion borders the Dry Chaco and the Pantanal
to the North, and the Cerrado and the Alto Paraná Atlantic Forest to the Southeast
(Rumbo 2010), therefore, the conservation of native ecosystems could benefit the
fauna exchange with the other four ecoregions present in Paraguay. In both Humid
Chaco and Pantanal ecoregions, the wooded formations of Schinopsis balansae, an
emblematic tree species with a high content of natural tannins, are also found. These
vegetation types constitute one of the most diverse ecotonal formations in the Great
Chaco, where plant species converge from the Atlantic Forest, the Cerrado, the
Amazon and the Dry Chaco, among others (Mereles et al. 2020b).
In the study site, El Raiss (2014) examined the functional connectivity of forest
islands for the Black-and-gold Howler Monkey (Alouatta caraya) within the native
silvopastoral system (i.e., the cow-calf system where animals graze on savannas and
are interspersed with mixed native forests). The author’s findings show an Equiva-
lent Connected Area (ECA) of 79%, indicating a good connectivity condition of
forest islands in the landscape. The author also points out that 10 to 15% removal of
forest islets would drastically affect this connectivity due to this species’ dependence
on forest islets. Thus, the study concluded that the current production system is
highly compatible with the conservation of this species.
In addition, the inclusion of live fencing, which is not a common practice in the
region, can increase habitat connectivity. The maintenance of continuous areas of
intact or lightly disturbed vegetation is a priority issue to consider in conservation
policies.

8.4 Forest Islands as Fire-Breaks in the Savanna

Fires are a normal and frequent event that occur in flooded savannas around the
world (Whelan 2006). For example, in South Africa, fire is considered a natural
factor in the development and maintenance of the vegetation of the Kruger National
Park (Govender et al. 2012). At the regional scale, Silveira et al. (1999) recommend
a fire management program to minimize the danger of uncontrolled fires, using
controlled burns on a rotational basis in different sections of the Emas National Park
in Brazil. The program was also meant to improve the availability of food for
herbivores and control the spread of alien grass species.
198 R. Laino et al.

In addition to flooding, fire is a main agent of disturbance in the Chaco.

According to Morello et al. (2009), floods put pressure on ecosystems with equal
intensity as the fires. Fire is an ecological component in the vegetation distribution of
the Humid Chaco, due to the high productivity of the herbaceous stratum during the
wet season and the insufficient number of herbivores to assimilate all the production
(Herrera et al. 2003). Fire consumes plant production that is not consumed by
herbivores and other foraging species (such as ants) and re-opens space for growth.
Extensive fires are provoked by natural causes such as electric storms or by anthro-
pogenic burning during the management of pastures and forests (Ginzburg and
Adámoli 2006; Benzaquen et al. 2017). In almost all of the world’s natural savannas,
the frequency of spontaneous fires is increased by human action and its effects can
influence the existing balance in the natural vegetation (Whelan 2006).
Fire is used as a management tool in livestock production in savannas. After
burning, the herbaceous stratum reaches a higher concentration of nitrogen and
protein in the regrowth (Ginzburg and Adámoli 2006), and lignified grasses are
eliminated, thereby stimulating the growth of native grasses, which are more palat-
able for cattle (López-Hernández and Hernández Valencia 2009). Inadequate fire
management during pasture burning and the lack of regulation of the livestock load
(such as intensity and frequency of grazing), lead to significant incidence of forest
fires (Galindo et al. 2009). Inappropriate fire regimes threaten biodiversity conser-
vation because high-intensity fires kill plants and animals and change the landscape
for years, decades, or even centuries in some natural communities (Whelan 2006).
Therefore, the conservation of forest islands in landscapes such as the Humid Chaco
is of utmost importance as a barrier to manage fires that may get out of control. The
islands of vegetation serve as a refuge for animals when fires occur (Silveira et al.
1999). In addition to the maintenance of forest islands immersed in savannas,
controlled grazing is also proposed as a fire prevention measure (Ruiz-Mirazo
et al. 2007).

8.5 Benefits of Forest Islands to Cattle Ranching in Native

Grasslands

The farms located in the study area are usually dedicated to cattle raising, with a
calving percentage3 of 50% in cows and 80% in heifers, and weaned calves between
six and eight months of age having a live weight of 130 to 150 kg. The stocking rate
is 0.5 animal unit (of 400 kg) per ha, which is relatively low because it considers the
possibility of flooding or drought, though it may increase during spring and summer
in the crucial period of the birth of calves (Laino et al. 2017). Some ranches spare
land purposely (as reserves, with no grazing animals) to avoid losing control of the

3
Calving percentage (porcentaje de parición) is the percentage of females that give birth to calves
from the total of cows/heifers serviced by bulls.
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 199

herd in difficult terrain or as a buffer area to avoid thefts, however no estimation is

available of this practice.
Lubián (2014) determined the existence of a variety of native grasses that produce
between 973 and 3612 kg of dry grass ha1 year1 in the study site, the most
palatable being Clavel grass (Hemarthria altissima) and Kapií-pytá (Andropogon
lateralis). Extensive cattle production is based mainly on three factors: the natural
supply of grasses for cattle feed, the availability of water and its accumulation in
savannas, and the presence of dense forest islands immersed in savannas, palm
groves and wetlands in the study area. The forest islands play a role in livestock
breeding since they provide shelter for animals during floods and in extreme
conditions of heat (up to 50
C) in summer and cold (3
C) weather in winter. In
addition, management of forests, with occasional extractions of wood for corrals,
cattle ponds,4 fences and bridges, allows savings of up to 40% of infrastructure costs
in a productive unit. These savings would not be possible if forests were converted to
exotic pastures (Laino et al. 2017).
The combination of these three factors - natural grasses, water and forest islands -
has allowed extensive livestock farming in the context of climate change adaptation.
At the same time, frequent floods and droughts lead the farmer to seek economic
stability at the producer level due to low and unstable profits (Laino et al. 2017). At
this intersection of climate adaptation and the consequent search for economic
viability, land use change happens through the clearing of forests for planting exotic
pastures of high yield, which maximizes economic production, but constitutes a risk
for conservation in the region.

8.6 Challenges for Conservation of Forests Islands

in the Chaco Region of Paraguay

Ecosystem degradation in the Chaco is occurring at the regional level (WWF 2016).
From 2012 to 2018, a total of 2,925,030 ha experienced land use change in the South
American Great Chaco, according to data from the land use change monitoring
carried out by the NGO Guyra Paraguay. The work demonstrates the gradual degree
of modification of the Great Chaco ecoregion, which until recently comprised one of
the largest natural areas in the world (Guyra Paraguay 2018). These studies are
almost entirely based on monitoring of deforestation of dense forests, but no studies
focus on savannas or open landscapes. In particular, the farms included in this
chapter are at higher risk of deforestation because of their proximity to urban areas
and also due to illegal extraction of trees by intruders during high floods when farms
can be accessed by boat.
The South American Great Chaco is not only being affected by a very intense
process of deforestation, but is also suffering a loss of natural grasslands, both on

4
These are “tajamares” or ponds that have been dug out for rain water collection.
200 R. Laino et al.

higher land and in wetlands, with a rate of disappearance even higher than that of the
forests (Bucher 2016). In most tropical and subtropical biomes, conservation strat-
egies are mainly focused on the preservation of forests, but savanna ecoregions and
open habitats deserve conservation attention as well (Grau et al. 2015). Many South
American countries have no specific wetland management programs. In areas with
low population density and without agro-industrial activities, wetlands are less
impacted (Junk et al. 2013). The complex interactions between biophysical and
socioeconomic processes that drive the trends of Chaco natural grasslands represent
a major scientific challenge to preserve this shrinking environment and its valuable
biodiversity (Grau et al. 2015).
Although many variables can affect the impact of livestock practices on ecosys-
tems, low intensity cattle ranching, with low densities of cattle on native pastures
(savannas) and conserved forest islands, could be beneficial for the survival of the
wild fauna that still remains. The presence of cattle consuming the high herbaceous
productivity could be a key factor in avoiding extensive fires. This in turn also
contributes to maintaining wetlands and associated riparian forests in the region,
which serve as natural refuges for wildlife.
The high relative abundance of aquatic animal species increases the conservation
value of the Paraguayan Humid Chaco, even though there are no endemic fauna
species (Mereles et al. 2013). Despite its high biodiversity, the Humid Chaco does
not have enough protected areas (Caballero-Gini et al. 2020). Thus, cattle ranchers of
the study area play an important role in the conservation of species at the local level,
since its livestock activities allow the coexistence of wild native and domesticated
exotic species.
The vast majority of wild protected areas of the Paraguayan Chaco are located in
the Dry Chaco, Continental Sand Dunes and Cerrado (Mereles et al. 2020b),
therefore it is crucial to highlight the importance of conserving the ecosystems of
the Humid Chaco as well. In addition, as described, the ecosystems of the Humid
Chaco are also important in flood mitigation, aquifer recharge and water quality
improvement (Benzaquen et al. 2017). Future economic development, combining
production with biodiversity conservation in a sustainable way, may be possible in
the region (Mereles et al. 2020b).
This chapter has described general patterns of the richness of species of flora and
fauna of the region. The aim is to highlight the value of these areas for conservation,
despite being intended predominantly for economic productive activities. However,
it is not clear yet if there is a deliberate intention of farmers to conserve certain
natural elements, or if it is simply an unintended consequence of their management
(or lack thereof). While this production-based conservation model may be
interpreted as a low income generating activity by some (or perhaps even inefficient
in economic terms), it may alternatively be interpreted as an opportunity for conser-
vation motivated by non-economic reasons such as cultural, family ties, research, or
appreciation of nature, or as a combination of both. Further research is needed to
understand drivers for intensification and also motivations to preserve natural eco-
systems in the area. However, the key message is that the type of management
discussed here allows for conservation of natural elements that more intensive
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 201

economic activities may not allow. The chapter is not intended to present this
production-conservation model as a substitute for the need to spare areas exclusively
for biological conservation, but as a complement in the landscape.
The lack of a previous baseline of biodiversity present in the area before cattle
ranching activities began (approximately 100 to 150 years ago), also limits our
understanding of how cattle ranching activities have affected biodiversity in the
past decades. Thus, this chapter provides only information of the current presence of
fauna and flora under the existing management and makes no assumptions about the
trends of the presence of the species. Comparisons through time, and especially with
less intervened areas, are needed to understand the impact of the productive activity
on biodiversity in the region over the long term.

8.7 Conclusion

Extensive or semi-extensive ecosystem-based livestock production could help to

maintain ecosystem services in the region. The need to address adverse environ-
mental conditions from the productive point of view, in this case floods from rain
and periodic floods of the Paraguay River, could represent an opportunity for the
conservation of threatened biodiversity. These native ecosystems are at high risk of
disappearing due to the intensification of agriculture and livestock production, as
well as the expansion of urban areas, and so strategies for their conservation,
including tailored incentives, are needed.
Currently, an increase of higher yielding crops for the region are being proposed
or piloted, and major roads and infrastructure are being planned or discussed locally
for production of rice and other commodity crops. Dropping prices of meat in
Paraguay and the poor recognition of these systems as a land sharing strategy,
could in the future lead to other more profitable production systems, which could
jeopardize relevant local ecosystem services such as wildlife, water quality, flood
regulation, as well as global services such as carbon sequestration. Implementation
of better management practices for cattle rearing, reducing taxes for sustainable
farming, providing national and international incentives, increasing prices for meat
produced under these environmentally friendly systems, and promoting ecotourism,
among other strategies, are some of the possible solutions to promote conservation of
the Humid Chaco forest islands, savannas and wetlands along with sustainable
production.

Acknowledgments The American Chaco Research Center is a local initiative promoting biodi-
versity conservation, environmental research and sustainable cattle production in native grasslands
of the Humid Chaco in Paraguay ([Link]). Institutionally, it is supported by the
Manuel Gondra Foundation as well as the Estancia Playada associates.
This chapter is based on works carried out in the framework of several research projects: Project
PINV 15-143 “Multidisciplinary contribution to the eco-hydrology of the wetlands of the Humid
Chaco” and PINV 14-350 “State of the caiman populations in six localities of Paraguay” both
financed by Consejo Nacional de Ciencia y Tecnología (CONACYT) of Paraguay, and the Project
202 R. Laino et al.

“Initial evaluation of the richness of medium and large sized mammals using camera traps in a
silvopastoral system in the American Chaco Research Center” carried out with the support of IDEA
WILD. We also presented data obtained in field trips made by researchers and the results of theses
of undergraduate and graduate students from national and foreign universities. We also thank
Silvina Chaparro, Humberto Sánchez, Paloma Moreno, Lucas Cñaete and people in charge at
Estancia Playada who helped during the fieldwork and for making our stay more enjoyable. We
dedicate this chapter to the memory of Aldo Florentín, who worked with us in the field during the
projects.

References

Agencia Catalana del Agua (ACA) (2004) Caracterización, regionalización, elaboración de

herramientas para la determinación del estado ecológico de los humedales de Catalunya.
Catalunya, Espñaa. [Link]/aca/documents/ca/directiva_marc/zones_humides.pdf.
Accessed 2 Mar 2020
Ali W, Javid A, Bhukhari SM, Hussain A, Hussain SM, Rafique H (2018) Comparison of different
trapping techniques used in herpetofaunal monitoring: a review. Punjab Univ J Zool 33(1):
57–68
Aquino L, Colli G, Reichle S, Silvano D, Scott N (2004) Chiasmocleis albopunctata. The IUCN red
list of threatened species 2004, p e.T57748A11680057. [Link]
[Link]. Accessed 13 Jan 2020
Asociación Paraguaya de Mastozoología (APM), Secretaría del Ambiente (SEAM) (2017) Libro
Rojo de los Mamíferos del Paraguay: especies amenazadas de extinción. Editorial CREATIO,
Asunción, Paraguay, p 137
Benzaquen L, Blanco DE, Bo R, Kandus P, Lingua G, Minotti P, Quintana R (eds) (2017) Regiones
de Humedales de la Argentina. Ministerio de Ambiente y Desarrollo Sustentable, Fundación
Humedales/Wetlands International, Universidad Nacional de San Martín y Universidad de
Buenos Aires, Buenos Aires
Boródn D (2015) Estimación del contenido de carbono en la cobertura forestal y en el suelo de un
bosque mesoxerofítico, Chaco Paraguayo. Tesis de Ingeniería Forestal, Universidad Nacional
de Asunción, Asunción, Paraguay
Boughton RG, Staiger J, Franz R (2000) Use of PVC pipe refugia as a sampling technique for hylid
treefrogs. Am Midl Nat 144:168–177
Brun V (2013) Servicio ecosistémico de almacenamiento de carbono en el suelo del humedal del
Río Negro, departamento de Presidente Hayes, Paraguay. Tesis de Ingeniería Ambiental,
Universidad Nacional de Asunción, Asunción, Paraguay
Brusquetti F, Lavilla EO (2006) Lista comentada de los anfibios del Paraguay. Cuad Herpetol 20(2):
3–79
Brusquetti F, Netto F (2008) Geographic distribution: Chiasmocleis albopunctata. Herpetol Rev 39:
362
Bucher E (2016) El futuro incierto de los humedales del Chaco: el caso de los bñaados del río Dulce.
Paraquaria Natural 4(2):11–17
Caballero-Gini A, Bueno-Villafñae D, Laino R, Musálem K (2020) Diversity of mammals and birds
recorded with camera-traps in the Paraguayan humid Chaco. Bol Mus Nac Hist Nat Parag 24(1):
5–14
Cartes JL, Clay RP (2009) Paraguay. In: Devenish C, Díaz Fernández DF, Clay RP, Davidson I,
Yépez Zabala I (eds) Important bird areas of the Americas – priority sites for biodiversity
conservation. BirdLife international (BirdLife conservation series no. 16), Quito, Ecuador, pp
297–306
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 203

Chaparro P (2014) Análisis de la relación cobertura vegetal-calidad del agua en el humedal del Río
Negro, Presidente Hayes, Paraguay. Tesis de Ingeniería Ambiental, Universidad Nacional de
Asunción, Asunción, Paraguay
Collevatti R, Leite K, de Miranda G, Rodrigues F (2007) Evidence of high inbreeding in a
population of the endangered giant anteater, Myrmecophaga tridactyla (Myrmecophagidae),
from Emas National Park, Brazil. Genet Mol Biol 30(1):112–120
Corn P (1994) Straight-line drift fences and pitfall traps. In: Heyer WR, Donnelly MA, McDiarmid
RW, Hayek LC, Foster MS (eds) Measuring and monitoring biological diversity. Standard
methods for amphibians. Smithsonian Institution Press, Washington DC
Crooks K, Burdett C, Theobald D, King S, Moreno Di Marco C, Boitani L (2017) Quantification of
habitat fragmentation reveals extinction risk in terrestrial mammals. PNAS 114(29):7635–7640.
[Link]
Crump ML, Scott NJ (1994) Visual encounter surveys. In: Heyer WR, Donnelly MA, McDiarmid
RW, Hayek LC, Foster MS (eds) Measuring and monitoring biological diversity. Standard
methods for amphibians. Smithsonian Institution Press, Washington DC
Del Castillo H (2019) Segunda Actualización de la Lista Comentada de Aves del Paraguay (Guyra
Paraguay 2004). Paraquaria Natural 7:8–18
eBird (2020a) EBird Basic Dataset. Cornell Lab of Ornithology, Ithaca, New York. [Link]
org/paraguay/hotspot/L2762215?yr¼all &m¼&rank¼mrec. Accessed 30 Jan 2020
eBird (2020b) EBird Basic Dataset. Cornell Lab of Ornithology, Ithaca, New York. [Link]
org/paraguay/hotspot/L6126375?yr¼all&m¼&rank¼mrec. Accessed 30 Jan 2020
El Raiss Z (2014) Análisis de la conectividad para el mono carayá (Alouatta caraya) en un sistema
silvopastoril del chaco úhmedo sudamericano. Tesis para el Máster de Restauración de
Ecosistemas, Universidad de Alcalá, Espñaa
Ferreira E (2018) Análisis del estado de conservación de tres humedales bajo distintos impactos
ambientales antrópicos, Chaco Húmedo, Paraguay. Tesis de Ingeniería Ambiental, Universidad
Nacional de Asunción, Asunción, Paraguay
Frost DR (2020) Amphibian Species of the World: an Online Reference. Version 6.0 (Date of
access). Electronic Database accessible at [Link]
[Link]. American Museum of Natural History, New York, USA. [Link]
vz.0001
Galindo I, Barrón J, Padilla JI (2009) Relación entre ganadería extensiva e incendios en zonas
forestales del estado de Colima. Avances en Investigación Agropecuaria 13(3):17–33
Ginzburg R, Adámoli J (2006) Situación Ambiental en el Chaco Húmedo. In: Brown A, Martínez
Ortiz U, Acerbi M, Corcuera J (eds) La Situación Ambiental Argentina 2005. Fundación Vida
Silvestre Argentina, Buenos Aires, pp 103–113
Gómez Fernández A, Alcocer I, Matesanz S (2016) Does higher connectivity lead to higher genetic
diversity? Effects of habitat fragmentation on genetic variation and population structure in a
gypsophile. Conserv Genet 17:631–641
Govender N, Mutanga O, Ntsala D (2012) Veld fire reporting and mapping techniques in the Kruger
National Park, South Africa, from 1941 to 2011. African J Range Forage Sci 29(2):63–73
Grau HR, Torres R, Gasparri NI, Blendinger PG, Marinaro S, Macchi L (2015) Natural grasslands
in the Chaco. A neglected ecosystem under threat by agriculture expansion and forest-oriented
conservation policies. J Arid Environ 123:40–46
Guyra Paraguay (2004) Lista comentada de las aves de Paraguay. Annotated checklist of the Birds
of Paraguay. Guyra Paraguay, Asunción, Paraguay. 200 pp. [Link]
Guyra Paraguay (2018) Monitoreo Mensual del Cambio de Uso y Cobertura de la Tierra, Incendios
y Variación de la Cubierta de Aguas en el Gran Chaco Americano. Informe Técnico. Asunción,
Paraguay. [Link]
Herrera P, Torrella S, Adámoli J (2003) Los incendios forestales como modeladores del paisaje en
la región chaqueña. In: Kunst C, Bravo S, Panigatti J (eds) Fuego en los ecosistemas argentinos.
INTA, Santiago del Estero
204 R. Laino et al.

IPCC (2005) Carbon dioxide capture and storage. Metz B, Davidson O, de Coninck H, Loos M,
Meyer L (eds.) [Link]
IUCN (2019) The IUCN Red List of Threatened Species. Version 2019–3. [Link]
org. Accessed 30 Jan 2020
Junk WJ, An S, Finlayson CM, Gopal B, Kveˇt J, Mitchell SA, Mitsch WJ, Robarts RD (2013)
Current state of knowledge regarding the world’s wetlands and their future under global climate
change: a synthesis. Aquat Sci 75:151–167
Kandus P, Morandeira N, Schivo F (eds) (2010) Bienes y servicios ecosistémicos de los humedales
del Delta del Paraná. Fundación Humedales/Wetlands International, Buenos Aires, Argentina
Laino L, Musálem K, Laino R (2017) Perspectivas para un desarrollo sustentable: un estudio de
caso de producción ganadera en la región del Chaco paraguayo. Población y Desarrollo 45:95–
106
Lindenmayer DB, Fischer J (2006) Habitat fragmentation and landscape change: An ecological and
conservation synthesis. Island Press, Washington, DC
López-Hernández D, Hernández Valencia I (2009) Nutritional aspects in Trachypogon savannas as
related to nitrogen and phosphorus cycling. In: Del Claro K, Oliveira P, Rico-Gray V (eds)
Tropical biology and conservation management, volume X, savanna ecosystems. UNESCO/
EOLSS, Oxford
Lubián A (2014) Valoración económica de sistemas de producción con inclusión de servicios
ecosistémicos en el Bajo Chaco, departamento de Presidente Hayes, Paraguay. Tesis de
Ingeniería Ambiental, Universidad Nacional de Asunción, Asunción, Paraguay
Macedo R (2018) Análisis de la estructura vegetal humedal-bosque ripario en el humedal del río
Negro, Chaco Húmedo, Paraguay. Tesis de Ingeniería Ambiental, Universidad Nacional de
Asunción. Asunción, Paraguay
MADES (2019) Resolución N
254/19 “Por la cual se actualiza el listado de las especies protegidas
de la vida silvestre de la clase aves”. Ministerio del Ambiente y Desarrollo Sostenible.
Asunción, Paraguay. 10 p
Mamede S, Benites M, Esquivel A, Clay R, de Lima MG, Rodrigues Alho CJ (2019) Turismo de
observação de aves no Chaco: oportunidades e desafios ao Corredor Bioceânico, segmento
Brasil/Paraguai. Interacções 20:159–177
Maturo HM, Oakley LJ, Prado DE (2005) Vegetación y posición fitogeográfica de la Reserva El
Bagual. In: Di Giacomo AG, Krapovickas SF (eds) Historia natural y paisaje de la Reserva El
Bagual, Formosa, Argentina. Asociación Ornitológica de La Plata, Buenos Aires. 592pp
Mereles F, Rodas O (2014) Assessment of rates of deforestation classes in the Paraguayan Chaco
(great south American Chaco) with comments on the vulnerability of forests fragments to
climate change. Clim Change 127:55–71
Mereles F, Cartes JL, Clay R, Cacciali P, Paradeda C, Rodas O, Yanosky A (2013) Análisis
cualitativo para la definición de las ecorregiones de Paraguay occidental. Paraquaria Nat 1(2):
12–20
Mereles F, Céspedes G, De Egea-Elsam J, Spichiger R (2020a) Estudios fitosociológicos en el Gran
Chaco: estructura, composición florística y variabilidad del bosque de Schinopsis balansae en el
Chaco Húmedo Boreal, Paraguay. Bonplandia 29(1):39–55
Mereles MF, Cespedes G, Cartes JL, Goerzen R, De Egea-Elsam J, Rodriguez L, Yanosky A,
Villalba L, Weiler A, Cacciali P (2020b) Biological corridors as a connectivity tool in the region
of the great American Chaco: identification of biodiversity hotspots in the ecoregions of the
Paraguayan Chaco. Res Ecology 2(1):27–36
Morello J, Rodríguez A, Silva M (2009) Clasificación de ambientes en áreas protegidas de las
ecorregiones del Chaco Húmedo y Chaco Seco. In: Morello J, Rodriguez A (eds) El Chaco sin
bosques: la pampa o el desierto del futuro. Orientación Gráfica Editora, Buenos Aires, pp 53–91
Motte M, Zaracho V, Caballero-Gini A, Ferreira-Riveros M, Romero Nardelli L, Coronel-Bejarano-
D, Netto F, Carosini A, Rojas V, Bueno D, Cabral H, Martínez N (2019) Estado de conservación
y lista roja de los anfibios del Paraguay. Bol Mus Nac Hist Nat Parag 23(1):1–62
8 Islands of Forests Among Savannas: Key Elements for Conservation. . . 205

Mujica N (2014) Habitat factors affecting occupancy and detection of mammals in the Paraguayan
Chaco. Master’s Thesis, Southern Illinois University, Carbondale, Illinois, USA
Nielsen C, Thompson D, Kelly M, Lopez-Gonzalez CA (2015) Puma concolor (errata version
published in 2016). The IUCN Red List of Threatened Species 2015: e.T18868A97216466.
[Link] Accessed 8 Jan 2020
Peña-Chocarro M, De Egea J, Vera M, Maturo H, Knapp S (2006) Guía de árboles y arbustos del
Chaco úhmedo. Guyra Paraguay, Asunción, Paraguay, 291pp
Pereyra M, Pol RG, Galetto L (2019) Ant community patterns in highly fragmented Chaco forests
of Central Argentina. Austral Ecol 44(4):668–679
Pérez de Molas L (2015) Manual de familias y géneros de árboles del Paraguay. FAO, San Lorenzo,
p 216
PINV 15-143 (2018) Proyecto Aporte multidisciplinario a la eco-hidrología de los humedales del
Chaco Húmedo. Centro de Investigación del Chaco Americano y Consejo Nacional de Ciencia y
Tecnología (CONACYT). del Paraguay, Asunción, Paraguay
Rodrigues FHG, Medri IM, De Miranda GHB, Camilo-Alves C, Mourao G (2008) Anteater
behavior and ecology. In: Vizcaíno SF, Loughry WJ (eds) The biology of the Xenarthra. Florida
University Press, Gainesville, pp 257–268
Ruiz-Mirazo J, Robles Cruz AB, Jiménez Piano R, Martínez Moya JL, López Quintanilla J,
González Rebollar JL (2007) La prevención de incendios forestales mediante pastoreo
controlado: el estado del arte en Andalucía. Sesión 3, Wildfire, Sevilla, Espñaa
Rumbo M (2010) Análisis biogeográfico de los mamíferos de Paraguay. Bol Mus Nac Hist Parag
16(1):18–29
Sato JH, Figueiredo CC, Marchão RL, Madari BE, Benedito LEC, Busato JG, Souza DM (2014)
Methods of soil organic carbon determination in Brazilian savannah soils. Sci Agric 71(4):
302–308. [Link]
Semper-Pascual A, Macchi L, Sabatini F, Decarre J, Baumann M, Blendinger P, Gomez-Valencia B,
Mastrangelo M, Kuemmerle T (2018) Mapping extinction debt highlights conservation oppor-
tunities for birds and mammals in the south American Chaco. J Appl Ecol 55(3):1–12
Shaffer M (1990) Population Viability Analysis. Conserv Biol 4:39–40
Silveira L, Rodrigues FHG, de Almeida Jácomo AT, Diniz Filho JAF (1999) Impact of wildfires on
the megafauna of Emas National Park. Central Brazil Oryx 33(2):108–114
Whelan RJ (2006) The ecology of fire - developments since 1995 and outstanding questions,
proceeding of bushfire 2006 - life in a fire-prone environment: translating science into practice
conference. Griffith University, Brisbane
World Wildlife Fund (WWF) (2016) Análisis social, económico y ambiental de la producción de
soja y carne en Paraguay. WWF-Paraguay, Asunción, Paraguay, Asunción, Paraguay
Zastavniouk C, Weir LK, Fraser D (2017) The evolutionary consequences of habitat fragmentation:
body morphology and coloration differentiation among brook trout populations of varying size.
Ecol Evol 7(17):6850–6862
Chapter 9
Biodiversity Islands and Dominant Species
in Agricultural Landscapes of the South
Western Amazon, Perú

Zoyla Mirella Clavo Peralta, Jorge Washinton Vela Alvarado,

and Carlos Mariano Alvez-Valles

Abstract The Ucayali region, in the Peruvian Amazon, is characterized by forests

that host a mega diversity of species. These forests have been affected by land use
changes that were in some cases supported by public policies, such as in the 1960s,
that encouraged the inhabitants of the Sierra (highlands) and Costa (coastal) regions
of Peru with incentives to populate the Amazon. The majority of settlers in Ucayali
were initially from the Sierras and had livestock rearing backgrounds. They became
established along the road that connects the Ucayali region with the city of Tingo
María in the Huánuco region. Most of them established their pastures by felling
nearly all the trees, though fragments of different sizes of primary and secondary
forest of different regeneration ages were conserved and used for various subsistence
activities. Many farmers value the trees and use live fences as limits of their
properties or their paddocks. Over the years, and due to the boom in agro-industrial
crops, most farmers reduced their grazing fields to plant oil, palm, and cocoa. The
objective of this chapter is to describe these islands of biodiversity and the dominant
species in the agricultural landscapes of this region. We describe the characteristics
of the Peruvian Amazon with an emphasis on the Ucayali region, its predominant
land uses, and the deforestation and degradation therein; the characteristics of the
biodiversity islands and their uses; the dominant flora species and their conservation
value and adaptation ranges; biological connectivity; and conservation strategies for
designing public policies. Effective design of land use or conservation programs in
accordance with current needs will only succeed with knowledge and dissemination
of the current state of our Amazon.

Z. M. Clavo Peralta (*) · C. M. Alvez-Valles

Facultad de Medicina Veterinaria, Instituto Veterinario de Investigaciones Tropicales y Altura
(IVITA), Universidad Nacional Mayor de San Marcos (UNMSM), Ucayali, Peru
e-mail: zclavop@[Link]; calvezv@[Link]
J. W. Vela Alvarado
Facultad de Ciencias Agropecuarias, Centro de Investigación de Fronteras Amazónicas (CIFA),
Universidad Nacional de Ucayali (UNU), Ucayali, Peru

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 207
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
208 Z. M. Clavo Peralta et al.

Keywords Conservation and development · Deforestation · Flora · Rainforest ·

Ucayali region

9.1 Introduction

The Peruvian Amazon is a megadiverse region that provides important ecosystem

services, such as carbon storage and biodiversity. The region holds diverse flora and
fauna, as well as high cultural diversity. It also supplies natural resources such as
hydrocarbons, and minerals, and its lands are used for the execution of development
projects in agriculture and livestock. For many political administrations, it represents
the largest area of expansion of the agricultural frontier.
At present times, legal and illegal productive and extractive anthropic activities
directly and indirectly cause deforestation and habitat loss. This directly contributes
to the present danger of a loss of balance in forest ecosystems and biodiversity. At
this crucial time, as researchers from two academic institutions invited by the author
of this book, we have reviewed and systematized the rich existing secondary
information, in addition to experiences from our own research on the forest resources
of the Peruvian Amazon in the Ucayali region, focusing on biodiversity and its
conservation potential through current strategies such as the framing tool of biodi-
versity islands.
Forest fragmentation and habitat loss are the result of environmental expansion
and degradation due to human activities (e. g. deforestation), all leading to a gradual
deterioration of the quality of remaining ecosystems (Morera et al. 2007). This also
results in changes in the forest microclimate (higher temperatures, lower humidity),
and in the dynamics of biological communities (increased predation and decreased
productivity) (Bierregaard et al. 1992). Likewise, the decrease in the size of forest
remnants and geographic isolation of natural ecosystems reduces the potential for
dispersion and colonization (Bennett 1998), restricting the natural movement of
species, reducing the possibility of genetic exchange, and consequently decreasing
biodiversity (Bennett 1998; Kattan 2002). The risk of extinction, especially for
species that depend solely on certain habitats to survive, also increases as a conse-
quence of forest fragmentation and habitat loss (McIntyre 1995).
The problem with decreasing the size and quality of habitat for species is that it
leads to a reduction of their populations (García 2002). Plant populations and the
populations of organisms with which they interact (pollinators, dispersers, herbi-
vores) are reduced. Therefore, the species richness of a forest remnant depends on
the suitability of the remnant as a habitat for several species (Begon et al. 1996). The
larger the remnant, the greater the possibility of keeping interior habitat or the greater
the probability of existence of a variety of habitats, which leads to a greater
possibility of species survival (Begon et al. 1996). The smaller the population, the
greater its probability of extinction. Consequently, when the population has been
greatly reduced, genetic deterioration is a big threat (Kattan 2002).
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 209

Considering the current high rates of deforestation and fragmentation due to the
increase in anthropic pressure on natural cover, which on a large scale can radically
alter the physical environment and climate, there is a need to implement conserva-
tion strategies in altered and fragmented environments. One of these strategies is the
establishment of connectivity between fragments of isolated or poorly connected
forests through the use of ecological corridors, which can facilitate the structural and
functional connectivity of biotic elements (Colorado Zuluaga et al. 2017). This
strategy should not only focus on the diversity attributes of biological communities,
but also on the maintenance of their natural dynamics, including the conservation of
their habitats and ecological processes at different spatial and temporal scales
(i.e. ecosystem functionality) that they require for their sustainability and biodiver-
sity conservation (Nott and Pimm 1997; Armenteras and Vargas 2016). Habitat
connectivity facilitates the dispersion and migration of species (their entry and exit
flow) through the landscape to meet basic habitat requirements (Bergoeing 1998).
In the southwestern Amazon, specifically the Ucayali Department of the Peruvian
Amazon, several immigrants from the coast and highlands (“Sierra”) of Peru settled
over time. These settlers, many of who have experience with livestock, were
encouraged by public policies over time with the goal of populating and utilizing
the Amazonian lands. For this reason, cattle raising was the main activity of these
families for their income. The cutting and burning of forests for pasture cultivation
resulted in the loss of large areas of vegetation cover. Nowadays, it is notable to see
these large areas without forests, which later were transformed into monocultures
such as oil palm and cocoa. Following this activity, an agricultural landscape can be
observed where large areas of pastures are dominant and the few areas with
vegetation cover are small patches of different sizes and areas of forests, called
“biodiversity islands.” These islands play a fundamental role in the conservation of
biodiversity because they act as shelters for species of flora and fauna. In addition,
these forest fragments promote connectivity while maintaining ecological integrity
and increasing landscape diversification.
The content of this chapter is based on secondary information on the Peruvian
Amazon region, as well as on research carried out by the authors over many years of
work on the subject as university professors as well as in research institutions.
Researchers working at the Veterinary Institute for Tropic and Altitude Research
(IVITA), Pucallpa, and of the Faculty of Veterinary Medicine of the Universidad
Nacional Mayor de San Marcos, have been conducting research since the 1980s on
the species of trees that predominate in the pastures as products of natural regener-
ation, on tree species that are suited for agricultural production, on the characteriza-
tion of plant succession after pastures or agricultural crops, on tree species resistant
to fire, on species with adequate vegetative reproduction suited for live fences
(“cercos vivos”), and on the use of the various species by the rural population.
Since the 1990s, faculty from the National University of Ucayali, Faculty of
Agricultural Sciences (UNU) and the Experimental Station of the National Institute
of Agrarian Research (INIA), Pucallpa, have generated knowledge on animal pro-
duction systems, the management of tropical pastures, silvopastoral systems from
natural regeneration of tree species, and tree planting.
210 Z. M. Clavo Peralta et al.

The chapter has four parts: it begins with the characterization of the Ucayali
region of the Peruvian Amazon and the classification of its vegetation; the second
part describes land use, conservation, deforestation and forest fragmentation, and
drivers of deforestation; the third part describes the dominant flora species, their
conservation value and key species; and the fourth section characterizes the ranges of
adaptation and biological connectivity. The information presented can be used to
guide strategies for conserving the biodiversity of the region.

9.2 Characterization of the Peruvian Amazon

and the Ucayali Region

The Peruvian Amazon is located at the eastern Andean flank and covers an area of
78.5 million ha, of which 23.11% correspond to the so-called High Forest which is
located between the Andean mountains at 500–2000 m above sea level, and 76.89%
correspond to the Low Forest located at less than 500 m above sea level (Gazzo
1982). The Ucayali region is located in the southwestern Amazon and in the central
and eastern part of the Peruvian territory, between the following coordinates: by the
North 7 0 200 0000 S Latitude and 74 0 320 0500 West Longitude; by the East, 9 0 250
0900 of S and 70 0 290 4600 W; by the South, 11 0 270 3500 S and 72 0 340 5500 W; and
from the West, to 08 0 400 1900 S and 75 0 580 0800 W (Vivanco 2004) (Fig. 9.1).
The average temperature is higher than 24  C in the tropical area and around
22  C in the sub-tropical parts. The precipitation varies according to the life zone: in
the subtropical area, the precipitation exceeds 3000 mm per year, while in the
tropical humid area it approaches 2000 mm per year and in the tropical dry area it
approaches 1200 mm per year.
The total area of the Ucayali region is 102,410.55 km2, corresponding to 13.15%
of the total area of the Peruvian Amazon and to 7.96% of Peru, being the second
largest region of the country, after the Loreto region (INEI, cited by Vivanco 2004).
The population of the Ucayali region according to the 2017 census was 496,459
inhabitants, of which approximately 75% live in urban areas, with a population
density of 5 inhabitants/km2 (INEI 2018).
Three morphologically different altitudinal floors are found in the Ucayali region:
Ceja de Selva (“Forest Brow”), Selva Alta (“High Forest”), and Selva Baja (“Low
Forest”). The Ceja de Selva begins at the headwaters of the Sepa, Unini, and
Catsingari rivers in the Atayala province at 1000 m above sea level, and in the
headwaters of the Aguaytía and Yurac rivers in the Padre Abad province, which
reaches as high as 3000 m above sea level. The Selva Alta is found in valley bottoms
of high altitude and narrow width, with terraces staggered in four levels, the lowest
having agricultural aptitude. Finally, the Selva Baja occupies the largest area in the
region and has acidic soils with low content of organic matter and phosphorous, low
base saturation, and high levels of aluminum (Table 9.1) (MINAGRI 2012).
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 211

Fig. 9.1 Geographical location of the Ucayali region in the southwestern Amazon. (Source:
Sonaira Silva. Federal University of Acre. Floresta Campus. Cruzeiro Do Sul)

Table 9.1 Morphological characteristics of the forests of the Ucayali region

Area Altitude
Altitudinal floors (km2) (masl) Characteristics
Ceja de Selva (“forest 1135.00 1000–3000 It begins in the provinces of Atalaya and
brow”) Padre Abad
Selva Alta (“high 12,948.00 500–1000 Neshuya districts, Curimaná, Campo
forest”) Verde
Selva Baja (low forest”) 88,434.00 0–500 Callería districts, Masisea
Source: Adapted from MINAN (2020), GeoBosques Program
212 Z. M. Clavo Peralta et al.

The Ucayali region is crossed from South to North by the Ucayali River, which
constitutes the middle course of the Amazon River. The Ucayali river is born at the
junction of the Tambo and Urubamba rivers and culminates when its waters pour
into the Marañón River. From that confluence it begins to be called Amazon River,
the largest river in Peru, which is approximately 3000 km long with a width that
varies from 400 to 2000 m.
The Ucayali river has a winding riverbed that forms many meanders along its
way. Some of them become “cochas” or “tipishcas,” large areas which are very rich
in fish, and some of them have also become touristic zones.
The river level rises between the months of September and October and reaches
its maximum in February. The water levels start to recede in May, reaching their
minimum level in July and August; according to bathymetric studies, there is a
difference of 12 m between the highest and lowest water levels, each cycle lasting an
average of 6 months. This cycle generates a distinct type of coastal tropical or
subtropical moist broadleaf forest area called “restinga” in the river shores, as well
as beaches and mud areas that form a complex of banks, in which large amounts of
silt and fine sediments accumulate. These sediments originate from materials that are
dragged along the rivers of the slopes of the Eastern Andes (Kalliola and Puhakka
1993). The sediments originate soils of high fertility, used for planting yellow corn,
plantain, grain legumes, rice, cassava, peanuts, and soybeans. The agriculture of
these crops plays an important role in food security of the riverine population as well
as a high percentage of the urban population.

9.2.1 Vegetation Classification of the Peruvian Amazon

At first look, Amazonian vegetation gives an impression of homogeneity, but this

cannot be farther from reality. The large number of species makes it difficult to
identify the different existing vegetation units. Different geological and geomorpho-
logical formations create habitats with different drainage conditions and very diverse
soils, which cause differences in the structure and floristic composition of the
vegetation.
In tropical America, numerous studies have been carried out to classify the
vegetation, for example, those of Holdridge (1967), Encarnación (1985), Malleux
(1971, 1982), all cited by Tuomisto (1993). According to the Holdridge system, the
life zones of the Ucayali region are classified mostly as Tropical Dry Forest and
Tropical Humid Forest. The Tropical Dry Forest begins in the southern Contamana
district, follows the Ucayali river upstream, and ends at the confluence of the
Urubamba and Tambo rivers. The Tropical Humid Forest begins at the border
with the Loreto region and follows the border with Brazil to the province of Purús
in the Ucayali region. The wettest natural life zone in Peru is located between Tingo
María and Pucallpa, where the sub-tropical rainforest and the very humid tropical
forests are located (Figs. 9.2 and 9.3) (Cochrane and Sánchez 1982).
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 213

Fig. 9.2 Tropical dry forest, Abujao River Basin, Pucallpa, Perú. (Photo: Jorge Vela, National
University of Ucayali. Pucallpa, Perú)

Malleux (1971, 1982) and Encarnación (1985), both cited by Tuomisto (1993)
have defined different types of vegetation for the Peruvian Amazon. Malleux based
his definitions on visible characteristics as seen in photographs, especially the
topography of the land and the texture of the vegetation cover, while Encarnación
based his on deep knowledge of the vegetation of the Loreto region, and used the
same vernacular nomenclature used by the inhabitants of the region. Both authors
classify the vegetation as: (a) Swamp forest, such as “aguajales”; (b) Temporarily
flood forests, such as riparian forests and restinga forests; and (c) mainland forest,
such as terrace forests, hill forests, secondary and degraded forests, and protection
forest (Figs. 9.4 and 9.5).
214 Z. M. Clavo Peralta et al.

Fig. 9.3 Tropical humid forest, Yurac river basin, border of Pucallpa and Tingo María, Padre Abad
Province, Ucayali region. (Photo: Mirella Clavo, IVITA, Pucallpa, Peru)

9.3 Land Use, Conservation, Deforestation,

and Fragmentation in the Ucayali Region, Southwestern
Amazon

9.3.1 Land Use

Land use capacity is an important variable given by the soil and its territorial
distribution, posing real challenges in the exploitation of this resource. The Peruvian
Amazon has an approximate extension of 78,308,801 ha, of which 69,380,729
(92.7%) are of mature forests (primary forest and secondary forests older than
12 years). The Ucayali region has aptitude and vocation for forestry, and a great
richness of natural resources, with high availability of water and diversity of flora
and fauna. Of the 10,515,536 ha of territorial extension of the Ucayali region,
88.41% corresponds to forests, and the rest to agriculture, grasslands, wetlands
and rural settlements as shown in Table 9.2.
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 215

Fig. 9.4 Swamp forest “aguajal”. Abujao river basin, Coronel Portillo Province, Ucayali region.
(Photo: Jorge Vela, National University of Ucayali, Pucallpa, Peru)

9.3.2 Conservation of Natural Resources

Peru is one of the ten megadiverse countries in the world, housing much of the
biological diversity of the planet. It has 28 climates, 84 out of the total 104 life zones,
and eight biogeographic provinces. It has three major river basins that contain
12,201 lakes and ponds, 1007 rivers, and 3044 glaciers. Much of this natural wealth
is conserved in the country’s 77 Natural Protected Areas, 17 Regional Conservation
Areas and 108 Private Conservation Areas, which in total conserve
22,584,586.19 ha (MINAM 2020). Most of the protected area belongs to the
Amazon region, specifically to the Ucayali region; however, it represents only a
small fraction of its territory.
The Ucayali region harbors a wealth of natural resources that combines well with
the cultural diversity of its people, wealth that must be valued in order to achieve a
harmonious balance between humans and nature to support the life of this planet.
Although the Ucayali region is one of the most biodiverse in the Peruvian Amazon,
only 12% of its territory is protected by different conservation categories. It only has
three National Parks, two Communal Reserves and one Regional Reserve, covering
a total of 1,261,864.32 ha. Furthermore, these areas, despite being protected by the
government, are undergoing a deforestation process, with 18,335 ha deforested so
far (MINAM 2020).
216 Z. M. Clavo Peralta et al.

Fig. 9.5 Forest remnants in cattle ranches, in areas of the Pucallpa to Tingo María road. (Photo:
Jorge Vela, National University of Ucayali, Pucallpa, Peru)

Table 9.2 Land use in the Ucayali region

Greater use capacity Aptitude or vocation Area (ha) %
Forest land Forest 9,296,795 88.41
Flooded areas in forests 95,601 0.91
Agricultural land Agriculture 297,378 2.83
Prairies Secondary vegetation 165,650 1.58
Pastures/grasslands 320,210 3.05
Hydromorphic Savannah 83 0.00
Wetlands Water body 285,354 2.71
Flooded areas in non-forest sites 36,503 0.35
Settlement Artificial areas 17,731 0.17
Mining areas 221 0.001
Other lands Bare land 10 0.001
Total 10,515,536 100.00
Source: Adapted from MINAN (2020) GeoBosques program
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 217

In Peru, Private Conservation Areas are the conservation category with the
highest number of protected areas, and yet, none of them are in the Ucayali region
(MINAM 2020). In addition to conservation areas promoted by the government, the
government of Peru also grants land to native communities, currently covering an
area of 1793.2 km2 and representing 17.06% of the territory of the Ucayali region
(MINAM 2020).

9.3.3 Deforestation and Forest Fragmentation in the Amazon

Currently, the balance of biological diversity worldwide is threatened. The crisis is

generated by anthropic activities of extractive and productive nature, such as illegal
and indiscriminate logging, the extraction of non-timber forest products, exploitation
of hydrocarbons, mining, hunting, illegal trade in native species, drug trafficking,
migratory agriculture, and extensive cattle ranching among others. These activities
have resulted in the disappearance of invaluable habitats and ecosystems. Likewise,
ethnodiversity, whose security and permanence has been threatened since ancient
times, is also currently in crisis because of the intrusion of entities or ideologies that
undermine ancestral customs, issues of territoriality, and a lack of resources. These
cause the decline of native human groups and the progressive excision or cultural
disappearance of these populations due to the loss of traditional knowledge.
Peru, as part of the Amazon basin, is no stranger to this deforestation process that
progressively removes a high percentage of its forests year after year and conse-
quently its biodiversity. MINAM (2020) has monitored deforestation in the Ucayali
region, calculating that from 2001 to 2018, 384,474 ha have been lost. In 2013,
36,793 ha were lost, representing the year of highest deforestation. In the following
years, that number has gradually decreased. For example, in 2018, deforestation
decreased to 25,991 ha (Fig. 9.6).

Fig. 9.6 Forest loss in the Ucayali region 2001–2018 (hectares). (Source: Adapted from MINAN,
Programa GeoBosques 2020)
218 Z. M. Clavo Peralta et al.

9.3.4 Drivers of Deforestation and Fragmentation

The occupation of the Amazonian territory, and consequently, the impacts of

deforestation have been occurring for many years, which continues to be both a
cause for concern and a subject of studies by many researchers and scientists. Toledo
and Serrao (1982) were of the opinion that the wisest decision to conserve the
Amazon would be to completely preserve it without modifying the ecosystem.
They also however raised the question of how long it would be possible to prevent
human occupation of this region. At that point in time, it was already too late. The
Amazon was already being invaded by people driven by socioeconomic pressures,
as well as demographic and regional integration policies in countries such as
Colombia, Ecuador, Peru, Bolivia and Brazil.
Rodríguez (2003) indicated that the Amazon, in the context of several develop-
ment models of the Peruvian economy, had only fulfilled roles as a provider of
natural resources. At the time, this market had a national and international heyday,
increasing demand for products like rubber cases, rosewood, Lonchocarpus nicou
(“barbasco”), wood, oil, and Theobroma cacao (“cocoa”). As promoted by the
governments of the day, infrastructure changed through the construction of access
roads and development projects for easier access to these profitable resources. Thus,
the Amazon became a sort of “escape valve” to mitigate the poverty of the Andean
populations. It was also subject to border colonization projects, implemented as a
security measure due to its status as a border region with countries such as Colombia,
Ecuador, Bolivia and Brazil.
According to Dourojeanni (1987), Robiglio et al. (2015), and Porro et al. (2015),
in the 1980s, agriculture and livestock activities were the main direct causes of
deforestation in Peru. Urban development, building of communication infrastruc-
ture, mining and oil exploitation were also other direct causes of deforestation
(Dourojeanni 1987). This author also stated that logging was not a direct cause of
deforestation at the time. While it is true that those activities were not major direct
causes of deforestation at the time due to their relatively small area, they were the
spearhead for the entry of large projects for agricultural and livestock development,
which take advantage of the facilities of the communication infrastructure and
internal roads and highways generated by logging. These large agriculture and
livestock projects included the sowing of large areas of oil palm, cocoa, coffee,
plantain, “camu camu” (Myrciaria dubia), and cattle raising for meat and milk.
Furthermore, the planting of illicit crops such as coca, which has been an important
economic engine in the Ucayali region since the 1980s, is not mentioned in official
statistics. The province of Padre Abad in Ucayali has the highest levels of coca
expansion in the country (Salisbury and Fagan 2011; Porro et al. 2015).
Deforestation and fragmentation of the forest of the Peruvian Amazon continues
rapidly today. The Monitoring of the Andean Amazon Project (MAAP), an initiative
of Amazon conservation (ACCA), uses satellite technology to monitor deforestation
in real time on the megadiverse Andean Amazon (Amazon basin of Peru, Colombia,
Ecuador and Bolivia) (ACCA 2020). In the last 17 years (2001–2017),
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 219

approximately 2.1 million ha of Peruvian forests have been lost, representing 50% of
the total Andean Amazon forests (ACCA 2020). Some of the main drivers of
deforestation are: (a) agriculture, which includes crops such as oil palm, cocoa and
other crops of small and large scale; (b) livestock, (c) gold mining, (d) forest roads
and (e) highways; with agriculture and livestock the drivers that most threaten the
entire Amazon region.
Most of the deforested areas (74%) are of small scale, with an average of less than
5 ha, which makes forest fragmentation very marked. Satellite imagery reveals many
small and large patches, as well as isolated forest fragments, known as biodiversity
islands (Finer and Mamani 2018).
ACCA (2020), MINAN (2020), and AIDER (2020) report that the greatest
amount of forest loss in the Ucayali region occurs in the province of Coronel Portillo,
in the Campo Verde and Nueva Requena districts, with 138,111 ha, representing 4%
of its area; and in the province of Padre Abad, in the districts of Curimaná, Neshuya,
Von Humboldt, Irazola and Padre Abad, with 162,581 ha, representing 25% of its
area. For Robiglio et al. (2015), the loss of forest is influenced by the opening of land
communication routes, expansion of the agricultural frontier, and development of
permanent crop plantations motivated by legal financial institutional incentives, as
well as the existence of perverse incentives that promote the shift from forest use to
crops.
Such is the case of the processes for granting proof of possession and titling
promoted by the National Commission for Development and Life without Drugs
(DEVIDA) that seek to eradicate the cultivation of coca. DEVIDA promotes uses in
a variety of agricultural activities, without necessarily considering or valuing the
importance of maintaining forests to ensure the quality of life and food security of
the population, as well as the important ecosystem services they provide (Robiglio
et al. 2015).
In addition, the study carried out by Porro et al. (2015) in the provinces of
Coronel Portillo and Padre Abad with samples of mestizo populations and indige-
nous peoples with diverse ethnic composition and different environments shows
that, on average, almost 40% of annual income comes from forests and environ-
mental products (including fish), followed by agriculture (25%), wages (17%),
livestock activity and animal products (11%), demonstrating the importance of
multiple uses of forest products for a large part of the local population. Public
policies for sustainable use could therefore be expected to promote a balance
between forest conservation and use.
220 Z. M. Clavo Peralta et al.

9.4 Dominant Flora Species, Conservation Value, Key

Species

9.4.1 Characteristics of the Secondary Forests

of the Peruvian Amazon

Several studies on the floristic composition of vegetation succession after the use of
the primary forest were initiated in the 1980s. Secondary forests known as “purmas”,
or fallows, are a consequence of the type of shifting agriculture practiced in the
region. After cutting and burning the forest, maize and rice are grown in the first year
and cassava or plantains in the second year. Then, the farmer makes the decision to
either plant grass or to leave the area abandoned to start the successional process.
They usually wait for a minimum of 7 years, a period considered to be enough for the
soil to regain its fertility, and then they can begin the cycle again. The period of time
that the land is left to recover its fertility is currently shorter than in the previous
generations, when farmers waited for periods of 10–15 years before starting a new
agricultural cycle (Toledo and Serrao 1982; Tuomisto 1993; Clavo et al. 2006).
Observations and studies indicate that within the flora of these secondary or
fallow forests there are species with diverse use potential (wood, firewood, fruits,
fibers, medicinal, among others), including fast-growing species such as Cecropia
sp, Ochroma piramidale and Guazuma sp. These fast-growing species are relatively
easy to manage because they do not require good soil quality and can withstand
variations in environmental conditions (Sabogal et al. 2001; Porro et al. 2015). The
more traditional use of the land, including shifting cultivation in large areas of land
and allowing fields to remain fallow for at least 10 years, gave rise to extensive areas
of abandoned fields with a population density of two or three people per square
kilometer. This migratory cultivation system was sustainable and self-sufficient and
also allowed for the conservation of biodiversity (Chávez 1991).
In recent years, the population has increased to five inhabitants per square
kilometer and new crop initiatives have been started (INEI 2018). New inhabitants
with large capitals have established monocultures of crops such as oil palm and rice
on large areas of fallows or “purmas” at different stages of succession. There are also
grasslands with scattered trees and wetlands. Forest fragmentation has increased,
especially along the road that connects the department of Ucayali with Lima.
According to Colan, cited by Sabogal et al. (2001), in the Peruvian Amazon
region, secondary vegetation is distributed in small patchy areas within the agricul-
tural units, usually located near the residual forests that have some seed trees of
commercial species. Small producers dedicated to the production of crops or live-
stock generally establish at least 2 ha of oil palm, taking advantage of the monetary
incentives provided by the government, and leave, according to the extension of land
that they own, small areas of primary or secondary forest of about a hectare or more.
Farmers also leave vegetation around water sources, which they use as watering
holes for cattle, indirectly conserving riparian vegetation and water. These constitute
the so-called islands of biodiversity that not only conserve the plant and animal
species but also the ecological processes and the landscape.
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 221

The presence of scattered trees in the pastures or fields also contributes to the
conservation of biodiversity and to biological connectivity. For example, a total of
87 tree species were identified in studies carried out in the department of Ucayali in
cultivated pastures of 40 farmers (Clavo and Fernández-Baca 1999; Riesco et al.
1995). The most frequent and economically valuable species that can be considered
as key species were Cordia ucayalensis, Ochroma pyramidale, Handroanthus
serratifolius, Terminalia spp., Trema micranta and Trichilia sp. These key species
are fast growing, have high regeneration capacity with good ability for regrowth and
good seed production, and some are fire resistant. In recent evaluations, we found an
increase in individuals per tree species that were growing in the boundaries of the
farms or as live fences of the pastures, providing biological connectivity and acting
as biological corridors (Vela et al. 2019).

9.4.2 Vegetation Studies in Different Stages of Succession

IVITA researchers, as part of a project on Sustainable Amazon Systems (SAS),

characterized the composition of secondary forest vegetation in different succes-
sional stages following abandoned agriculture or pastures (Riesco et al. 1995). A
total of 16 tree and shrub species were found in a 2-year old secondary forest
developed after the abandonment of agricultural use. In this forest, 98% of the
shrub and arboreal stratum was made up of individuals over 2 m high and less
than 5 cm in diameter at breast height (DBH). Among these, the species
“llausaquiro” (Heliocarpus popayenensis) was the most frequent, followed by the
“sachahuaca negra” (Baccharis floribunda). In this successional stage, the prolifer-
ation of individuals is large, due to the availability of light and the large number of
seeds produced, but few individuals manage to establish and develop due to com-
petition (Table 9.3). We must emphasize that “sachahuaca negra” is always present
in the first successional stages of the vegetation in the region, along with other
species of the Asteraceae family, constituting one of the botanical families that is
characteristic of the vegetation recovery of the Peruvian Amazon forests (Riesco
et al. 1995; Clavo and Fernández-Baca 1999).
In the 5-year old secondary forests of agricultural origin, 34 shrub and tree species
were identified, with the most frequent being “black ocuera” (Vernonia
baccharoides) with a frequency of 12.36%, followed by “black shimbillo” and
“alcanfor moena” (Table 9.4). In this successional stage, it is also common to find
trees of Piper hispidum, Jacaranda copaia, Cecropia membranacea and Cordia
ucayalensis. The abundance and density of herbaceous vegetation begins to decrease
at this stage (Table 9.4) (Riesco et al. 1995; Clavo and Fernández-Baca 1999).
In the 10-year old secondary forests of agricultural origin, 61 shrubs/trees and
8 liana species were identified. The most frequent tree species was the “white
tahuari” (Handroanthus sp.), followed by “black cordoncillo” (Piper laevigatum)
and “shimbillo bean” (Inga pundata). At this stage, the amount of herbaceous
species decreases, the number of trees increases, and liana species such as “cat’s
 222

Table 9.3 Frequency and density of tree species by diameter classes in 2-year old secondary forest of agricultural origin
Diametric classes Total
No. Scientific name Common name >2 to <5 cm 5 to 10 cm >10 cm Number of trees/ha Frequency (%)
1 Heliocarpus popayanensis H.B.K Llausaquiro 10,133.00 979.78 40.0 11,152.78 47.16
2 Ochroma pyramidale Urbam. Topa negra 1333.33 162.96 24.0 1520.2.09 6.43
3 Solanum grandiflorum R&P. Shucahuito 2133.33 133.33 0 2266.66 9.58
4 Cordia ucayalensis Jonhst Añallu caspi 133.33 29.63 0 162.96 0.69
5 Inga thiboudiana D.C Shimbillo negro 533.33 14.81 0 548.14 2.32
6 Baccharis floribunda H.B.K Sachahuaca negra 3733.33 0 0 3733.33 15.79
7 Cecropia membranacea Trecul Cetico blanco 533.33 0 0 533.33 2.26
8 Vernonia baccharoides R. et P. Ocuera negra 1333.33 0 0 1333.33 5.64
9 Hura crepitans L. Catahua 133.33 0 0 133.33 0.56
10 Hymatanthus Sucuuba (Spruce) Wood Bellaco caspi 400.0 0 0 400.0 1.69
11 Trema micrantha (L.) Blume Atadijo 666.67 0 0 666.67 2.82
12 Acacia sp Pashaquilla colorada 400.0 0 0 400.0 1.69
13 Vitex sp Cormiñon colorado 266.67 0 0 266.67 1.13
14 Banara Guianensis Aubl. Varilla blanca 133.33 0 0 133.33 0.56
15 Bahuinia sp. Pashaquilla pata de vaca 133.33 0 0 133.33 0.56
16 Lisianthus alatus (Aubl) Tabaquillo 266.62 0 0 266.62 1.13
Total 22,266.2 1320.5 64.0 23,650.7 100.0
Source: Riesco et al. (1995)
Z. M. Clavo Peralta et al.
Table 9.4 Frequency and density of tree species by diameter classes in 5-year old secondary forest of agricultural origin
Diametric classes Total
>2 to 5 to Number of trees/ Frequency
No. Scientific name Common name <5 cm 10 cm >10 cm ha (%)
1 Apeiba tibourbou Aubl. Maquisapa ñacha 0 14.81 4.0 18.82 0.21
negra
2 Casearia sp Sanipanga 0 0 4–0 4.0 0.04
3 Cecropia membranacea Trecul Cetico blanco 533.33 162.96 12.0 708.3 7.81
4 Cecropia engleriana Snethlage Cetico shiari 133.33 103.70 24.0 261.04 2.88
5 Cordia ucayalensis Jonhst Añallu caspi 0 148.14 4.0 152.15 1.68
6 Dictyoloma peruviana Plach. Huamanzamana negra 0 29.63 4.0 33.63 0.37
7 Rollinia insignis R.E. Frus Anonilla blanca 133.33 0 4.0 137.33 1.51
8 Inga Thiboudiana D.C Shimbillo negro 666.66 385.18 48.0 1099.85 12.13
9 Jacaranda copaia (Aubl.) [Link] Huamanzamana blanca 0 29.63 16.0 45.63 0.5
10 Miconia sp Rifari 0 0 4.0 4.0 0.04
11 Leonia glycicarpa R. & P. Tamara negra 0 0 4.0 4.0 0.04
12 Rollinia ulei Diels Sacha anona, anonilla 0 44.44 8.0 52.44 0.58
13 Sclerolobium sp Ucshaquiro blanco 133.33 133.33 32.0 298.67 3.29
14 Solanum grandiflorum R&P. Shucahuito 0 0 4.0 4.0 0.04
15 Banara sp Varilla 0 14.81 0 14.82 0.16
16 Alchornea triplinervia Uchumullaca 0 14.81 0 14.82 0.16
17 Banara guianensis Aubl. Varilla blanca 266.66 44.44 0 311.11 3.43
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . .

18 Vismia baccifera sub sp. ferruginea Pichirina colorada 0 14.81 0 14.82 0.16
(Kunth)Ewam
19 Trattinickia sp Tanque negro 533.33 29.63 0 562.96 6.21
20 Cordia nodosa Lam. Pucaruro caspi negro 0 14.81 0 14.82 0.16
21 Hymatanthus sucunba (Spruce) Woodson Bellaco caspi 0 14.815 0 14.82 0.16
22 Inga att. Alba (Swartz) Willo Shimbillo blanco 666.66 14.81 0 681.48 7.51
223

(continued)
Table 9.4 (continued)
224

Diametric classes Total

>2 to 5 to Number of trees/ Frequency
No. Scientific name Common name <5 cm 10 cm >10 cm ha (%)
23 Micropholis venulosa Mart. & Eichl Tanque blanco 0 14.81 0 14.82 0.16
24 Theobroma subincanum Mart. Cacahuillo 0 14.81 0 14.82 0.16
25 Vernonia baccharoides R. et P. Ocuera negra 913.33 207.40 0 1120.74 12.36
26 Banara arguta Briq. Varilla negra 266.66 0 0 266.67 2.94
27 Baccharis floribunda H.B.K Sachahuaca negra 533.33 0 0 533.33 5.88
28 Bactris sp Ñejilla 533.33 0 0 533.33 5.88
29 Naucleopsis glabra Sprice ex Beill Tamamurillo 133.33 0 0 133.33 1.47
30 Bahuinia sp. Pashaquilla pata de va 266.66 0 0 266.67 2.94
31 Capirona decorticans Spruce Capirona negra de 266.66 0 0 266.67 2.94
altura
32 Ocotea killipii A.C. Smith Alcanfor moena 800 0 0 800.00 8.82
33 Piper hispidum SW Cordoncillo 533.33 0 0 533.33 5.88
34 Cassia ruiziana Benth. Mataru amarillo 133.33 0 0 133.33 1.47
Total 7446.6 1451.8 172.0 9070.5 100
Source: Riesco et al. (1995)
Z. M. Clavo Peralta et al.
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 225

claw” (Uncaria guianensis) and “clavo huasca” (Tinanthus palyanthus), both of

medicinal and commercial value, appear more frequently (Table 9.5) (Riesco et al.
1995; Clavo and Fernández-Baca 1999).
When comparing forests of the same successional ages but of different origin,
differences in composition of species can be found in the early stages of secondary
succession. Secondary forests that originated from abandoned grassland have a
greater amount of herbaceous species than those from abandoned agricultural use,
and they are all similar (Riesco et al. 1995; Clavo and Fernández-Baca 1999).
In a study of the composition of seed banks of secondary forests (fallow) of
different successional stages and different origin, López et al. (2005) found no
significant differences in seed diversity, with a similar value of the Shannon index.
Fallows of agricultural origin, in different stages, had a smaller value than those of
pasture origin, probably due to the greater quantity of herbaceous species. The
Shannon indices were as follows: 1-year fallow of agricultural origin 2.58, 1-year
of pasture origin 3.09, fallow of pasture origin of 5 years 2.96 and fallow of
agricultural origin of 5 years 2.50. Herbaceous species that are more abundant in
fallows of pasture origin may contribute to the higher values of Shannon index
diversity. The species of greatest commercial value such as cedar, mahogany, kapok
and hardwoods such as “shihuahuaco” (Dipteryx sp.) were not recorded in any
successional stage probably due to their seed characteristics, which are more affected
by fire and lack of seed dispersers (Riesco et al. 1995; Clavo and Fernández-Baca
1999).

9.4.3 Use of Species from Secondary Forests and Islands

of Biodiversity

The secondary forests of the Peruvian Amazon are generally underutilized. Lack of
knowledge of the species is a constraint on their use and affects farm productivity.
Farmers in the region prefer species that provide them with products to increase their
income throughout the year and are diversifying their farms with this objective. The
secondary forest is part of the farm as a component for the future, waiting for the
recovery of soils and as a provider of some resources. However, secondary forests
constitute part of the biodiversity islands of the region, so it is essential to improve
knowledge of their species composition, and to quantify the products that they can
provide so that they can be managed properly to enhance their biodiversity conser-
vation (Riesco et al. 1995; Clavo and Fernández-Baca 1999).
Studies conducted in Pucallpa, department of Ucayali, show that the settlers
(producers) know the name of desirable and undesirable plants, but they have less
knowledge of their classification than other native peoples elsewhere in Peru or in
other parts of the world. Farmers in Pucallpa do not use plants as indicators of soil
fertility, they know few uses, such as building materials and edible fruits, and they
have very little knowledge on medicinal plants. They do, however, have good
Table 9.5 Frequency and density of tree species by diameter classes in 10-year old secondary forest of agricultural origin
226

Diametric classes Total

>2 to Number of trees/ Frequency
No. Scientific name Common name <5 cm 5 to 10 cm >10 cm ha (%)
1 Cassia spectabilis D.C. Retama negra 0 0 20.0 20.0 0.32
2 Cecropia membranacea Trecul Cetico blanco 0 0 36.0 36.0 0.58
3 Trichilia rubra C.D,C. Tushmo blanco 0 14.81 28.0 42.81 0.69
4 Inga edulis Mart. Guabilla 0 29.63 32.0 61.63 1.0
5 SD Chupo huayo 133.33 0 0 133.33 2.16
6 Ceiba samauma (Mart.) K. Schum. Huimba negra 0 0 12.0 12.0 0.19
7 Cassia lucens Vagel Retama colorada 0 59.25 48.0 107.25 1.74
8 Heliocarpus popayanensis H.B.K. Llausaquiro 0 14.81 8.0 22.81 0.37
9 Guatteria phanerocampta Diels Carahuasca negra 0 44.44 8.0 52.44 0.85
10 Vismia cayennensis (Jack) Pers. Pichirina blanca 0 0 12.0 12.0 0.19
11 Inga brachyrhachis Harms Charapa shimbillo 0 59.25 16.0 75.25 1.22
12 Terminalia oblonga (R&P) Steud. Yacushapana blanca 0 0 8.0 8.0 0.13
13 Guazuma crinita C. Mart Bolaina blanca 0 0 28.0 28.0 0.45
14 Cochlospermum orinocense (Kunth) Steud. Topa negra 0 0 16.0 16.0 0.26
15 Dictyoloma peruviana Plach. Huamanzamana negra 0 14.81 4.0 18.81 0.31
16 Cordia ucayalensis Jonhst Añallu caspi 0 0 12.0 12.0 0.19
17 Eschweilera coriácea (DC)[Link] Machimango blanco 0 0 4.0 4.0 0.06
18 Bactris gassipaes Kunth Pijuayo 0 0 4.0 4.0 0.06
19 Schizolobium amazonicum Huber ex Ducke Pashaco colorado 0 0 32.0 32.0 0.52
20 Talisia cf. cerasina (Benth.) Radlk. Caspi ubos 0 0 4.0 4.0 0.06
21 Guazuma ulmifolia Lam. Bolaina negra 0 0 8.0 8.0 0.13
22 Vitex klugii Mold Cormiñon 0 0 12.0 12.0 0.19
23 Croton matourensis Aubl Aucatadijo 0 0 4.0 4.0 0.06
24 Lonchocarpus spiciflorus Mart. ex Benth Maria buena 0 0 4.0 4.0 0.06
Z. M. Clavo Peralta et al.
25 Piptadenia pteroclada Benth Pashaco blanco 0 0 4.0 4.0 0.06
26 Hura crepitans L. Catahua 0 0 4.0 4.0 0.06
27 SD Desconocido cortezaun 133.33 0 0 133.33 2.16
28 Acacia loretensis J.F. Macbr. Pashaco negro con 0 14.81 8.0 22.81 0.37
espina
29 Uncaria guianensis (Aubl.) Gmel. Uña de gato 933.33 0 0 933.33 15.14
30 Trema micrantha (L.) Blume Atadijo 0 0 12.0 12.0 0.19
31 Inga calantha Ducke Rosca pacaya 0 14.81 4.0 18.81 0.31
32 Rollinia insignis R.E. Frus Anonilla 0 0 4.0 4.0 0.06
33 Sapium marmieri Huber Caucho masha 0 0 8.0 8–0 0.13
34 Astrocaryum sp Huicungo 0 0 4–0 4.0 0.06
35 Cordia nodosa Lam. Pucaruro caspi negro 0 14.81 0 14.81 0.24
36 Hymatanthus sucuuba (Spruc.) Wood Bellaco caspi 0 14.81 0 14.82 0.24
37 Erythroxylum cf. k Penth Charichuela blanca 0 44.44 0 44.44 0.72
38 Eschweilera sp Machimango colorado 0 14.81 0 14.81 0.24
39 Ichnosiphon cf. rotundifolius Trompetero sacha 573.33 0 0 573.33 9.3
40 Siparuna guianensis Aubl. Pishi huayo 0 29.63 0 29.63 0.48
41 Terminalia amazonia (J.F. Gmel) Exel Yacushapana negra 0 14.81 0 14.81 0.24
42 Aegiphila sp Sacha tol blanco 133.33 59.25 0 192.59 3.12
43 Banara guianensis Aubl. Varilla blanca 0 14.81 0 14.81 0.24
44 Cassia macrophilla Kunth. Maturu negro 0 14.81 0 14.81 0.24
45 Tovomita umbellata Benth. Chullachaqui caspi rojo 0 14.81 0 14.81 0.24
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . .

46 Inga pundata Willd. Poroto shimbillo 133.33 14.81 0 148.14 2.4

47 Lacmella arborescens (Muell. Arg.) Monach. Sachavaca huayo 0 14.81 0 14.81 0.24
48 Tococa guianensis Aubl. Pucaruro sacha 0 14.81 0 14.81 0.24
49 Morisonia oblongifolia Britt. Tamara blanca 0 14.81 0 14.81 0.24
50 Adenocalymna impressum (Rusby) Sandw. Huangana huasca blanca 0 14.81 0 14.81 0.24
51 Naucleopsis glabra Sprice ex Beill Tamamurillo 133.33 0 0 133.33 2.16
227

(continued)
Table 9.5 (continued)
228

Diametric classes Total

>2 to Number of trees/ Frequency
No. Scientific name Common name <5 cm 5 to 10 cm >10 cm ha (%)
52 Casearia aculeata Jacq. Limon casha 0 14.81 0 14.81 0.24
53 Perebea guianensis Aubl. Pama negra 133.33 14.81 0 148.15 2.4
54 Handroanthus sp Tahuari blanco 266.66 14.81 0 281.48 4.56
55 Garcinia gardneriana (Planch & Triana) Charichuela negra 0 14.81 0 14.81 0.24
Zappi
56 Ruizodendron ovale (R&P) [Link]. Paujil ruro 0 14.81 0 14.81 0.24
57 Tinanthus polyanthus Clavo huasca 666.66 0 0 666.66 10.81
58 Doliocarpus dentatus (Aubl)Standl Paujil chaqui 133.33 0 0 133.33 2.16
59 Ocotea killipii A.C. Smith Alcanfor moena 133.33 0 0 133.33 2.16
60 Simarouba amara Aubl. Marupa 133.33 0 0 133.33 2.16
61 Solanum grandiflorum R&P. Shucahuito 133.33 0 0 133.33 2.16
62 Calicobolus sericeus (HBX) House. Camote huasca 266.67 0 0 266.66 4.32
63 Banara arguta Briq. Varilla negra 133.33 0 0 133.33 2.16
64 Costus sp Sacha huiro blanco 133.33 0 0 133.33 2.16
65 Piper laevigatum H.B.K Cordoncillo negro 266.67 0 0 266.66 4.32
66 Trattinickia aspera (Standl) Swcrtz Sacha uvilla 133.33 0 0 133.33 2.16
67 Couepia sp Parinari 133.33 0 0 133.33 2.16
68 Dioscorea decorticans Presl. Soga alambre 133.33 0 0 133.33 2.16
69 Trichilia sp Requia blanca 133.33 0 0 133.33 2.16
Total 5106.6 651.8 408 6166.5 100
Source: Riesco et al. (1995). SD: unidentified
Z. M. Clavo Peralta et al.
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 229

knowledge of timber species that can be commercialized (Fujisaka et al. 2000). It is

important to note that most of the population in the productive area of Pucallpa are
migrants from different regions of Peru, and they try to use and manage the local
species in the same way as they did in their region of origin. Some farmers are not
even interested in knowing the uses of the plants they find in their plots. Thus, it is
important to disseminate the various uses of the species so that farmers may value
them better and so that they manage properly those species that are part of the islands
of biodiversity (Riesco et al. 1995; Clavo and Fernández-Baca 1999).
As previously mentioned, the islands of biodiversity in the area have different
sizes and shapes: some are fragments of secondary forests, others have originated
from primary forests, and many are riparian forests whose species contribute to
hydrological regulation through processes such as water storage and decreasing the
impact of floods. The riparian plant species give stability to the water channels and
provide refuge for the fauna, as well as climate regulation. Given the high value of
the ecosystem services that they provide for human societies, these biodiversity
islands need strategies to become areas of conservation with recognized value by the
government and with incentive mechanisms that ensure their permanence over time
(Riesco et al. 1995; Clavo and Fernández-Baca 1999).
These islands of biodiversity also house medicinal and food plants whose use
must be promoted so that the population can value them. In this manner, the
biodiversity islands can become Areas of Conservation Value, taking into consider-
ation the definitions of High Conservation Value Resource1 Network and the criteria
used by the Forest Stewardship council (FSC) to establish Conservation Value
Areas.

9.5 Landscape Fragmentation and Biological Connectivity

The fragmentation processes cause a decrease in the vegetation covers, leaving the
original vegetation of a given area as small fragments isolated from each other.
Increasing the number of fragments, reducing their area, and increasing the distance
between them, are limitations for ecological processes such as seed dispersal,
colonization, migration and interaction between species (Bennett 1998). Therefore,
landscape fragmentation is a process that severely affects biodiversity (Bennett
1998; Kattan 2002; Morera et al. 2007), and as habitat loss increases, connectivity
decreases and the edge effect increases (Fig. 9.7).
The Ucayali region presents alteration in its landscapes due to the different land
uses carried out there, mainly agricultural activities with crops of oil palm, coca,
banana, pineapple and cacao; and cultivation of pastures for livestock. Taking into

1
High conservation value of natural resources in an area is established according to the criteria and
principles of responsible forest management. Sustainable forest Deforestation hotspots in the
Peruvian Amazon (FSC 2020).
230 Z. M. Clavo Peralta et al.

Fig. 9.7 Landscape alteration process in relation to habitat loss and fragmentation. (Source:
Modified from Hobbs and Wilson 1998)

account the existence of altered and fragmented landscapes, it is necessary to

implement conservation strategies in these types of landscapes. Ecological corridors
are a strategy that facilitates biological connectivity between isolated or poorly
connected forest fragments (Colorado Zuluaga et al. 2017). This connectivity facil-
itates the dispersal and migration of species across the landscape to meet basic
habitat requirements (Bergoeing 1998). For this reason, connectivity between forests
in a fragmented landscape is important for the conservation and maintenance of their
integrity and biodiversity. Within these fragmented forests in the Ucayali region,
there are native species of various uses such as timber, medicinal, fruit or other,
which places greater emphasis on their conservation.
One of the alternatives that is being promoted as a biological corridor in the
Ucayali region is the implementation of agroforestry systems (AFS), mainly with
silvopastoral systems (SPS) as land use alternatives in agricultural landscapes. These
systems can serve to recover degraded areas, mitigate climate change by reducing
deforestation pressure in forests, increase biodiversity, and reestablish ecological
interactions through planting trees and perennial species. Despite the fact that this
research is still in progress, some areas have already been established with
silvopastoral systems, including one in the research areas of the IVITA station –
Pucallpa (Fig. 9.8). Positive results are being obtained regarding the increase in
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 231

Fig. 9.8 Left. Silvopastoral system with Simarouba amara (“marupa”) plantations. Right. Live
fences with Erythrina sp. (“Amasisa”). Both established at the IVITA – Pucallpa station. (Photos:
Carlos Mariano Alvez-Valles and Zoyla Mirella Clavo Peralta, IVITA, Pucallpa, Peru)

biodiversity, mainly with records of bird and amphibian species (Vela et al. ongoing
research project). Additionally, Redondo-Brenes and Montagnini (2010) mention
that agrosilvopastoral systems present a greater ecological complexity than that of
monocultures and, in turn, are found within land uses that harbor intermediate values
of bird species richness (143) within the Paso de la Danta Biological Corridor, south
of Costa Rica.
Live fences (“cercas vivas”) are also part of the AFS that provide land cover and
can serve as agroecological strategies to promote landscape connectivity (Schelhas
2007; Francesconi and Montagnini 2015). Such is the study by Gabriel and Pizo
(2005) in southwestern Brazil, which documents the frequent use of live fences by
birds during their movement and vocalizations, whether for rest or foraging. In the
Ucayali region, these systems are widely used in agricultural landscapes, where
planted trees and/or shrubs delimit the divisions of the field and the boundaries
between farms (Fig. 9.8), but further studies are still lacking in this part of the
Amazon confirming this system as an ecological corridor.
Riparian forests, apart from functioning as habitats, also function as ecological
corridors. The interaction between vegetation and river dynamism causes a hetero-
geneous mosaic of very productive habitats, due to the combination of fertility and
water availability of their soils (Martín et al. 2013). The most relevant factors that
affect the viability and functionality of riparian forest as biological corridors are the
existence of forest cover and the continuous or temporary presence of water. This
connective function is not only restricted to the circulation of living beings, since
these types of forest also act as conduits for runoff, sediment, organic matter and
other material (Martín et al. 2013). Therefore, the restoration of riparian forests
promotes their role as ecological corridors, facilitating displacement and providing
extensive resources as food and refuge for wildlife in fragmented landscapes
(Estrada and Coates-Estrada 2001; Gurrutxaga 2004).
Finally, Francesconi and Montagnini (2015) mention that agroforestry systems
(AFS) increase the vegetation cover of the landscape and therefore, can act as
232 Z. M. Clavo Peralta et al.

ecological strategies to restore functional connectivity in fragmented landscapes, but

still need further studies to confirm this function. Likewise, it is necessary to
consider and prioritize conservation at the landscape level in areas where the native
habitat is almost always fragmented (Vandermeer et al. 2008). In promoting con-
servation strategies of plant diversity in the fragments, their use by local human
populations should not be ignored, since in most cases forest fragments positively
favor the local economy (Brokaw 2002).

9.6 Conclusions

The southwestern Amazon is a very complex region, rich in life formations,

megadiverse and of great importance for the conservation and maintenance of
the ecological balance of the planet. There is wide and varied information on the
ecological characterization, types of vegetation, and ecosystem services of the
southwestern Amazon. The research carried out in the southwestern Amazon dates
back a long time, with a strong initial onset that has since apparently decreased in its
intensity, when compared with studies of the Amazon forest conducted in other
countries such as Brazil, Colombia, and Ecuador. The approaches were also differ-
ent, initially using aerial photography, field work and local knowledge, and then
varying as new technologies became available. Currently, with the advancement of
satellite technology, remote sensing, specialized software, and drones, among others,
deforestation and its causes can be detected in real time. It is very important to have
early warning of land use changes so that the competent authorities can take
immediate action.
In the 1980s, due to the expansion of agriculture and livestock in the southwestern
Amazon, large areas of forest were lost. After growing one or two crops, these lands
were abandoned to recover their soil fertility. As a result of this anthropic activity,
large tracts of “purmas”, or fallows, appeared, many of them forming fragments of
forests, patches or islands of biodiversity. The majority of unbridled areas are on
average less than 5 ha.
The diverse species of plants present in the islands of biodiversity in the south-
western Amazon fulfill different functions that contribute to ecological processes,
with many key species of large size that ensure their own continuity as well as the
conservation of several other species of flora and fauna. These areas of conservation
value are often used by the local population and communities, who know their
various uses that facilitate their daily lives, and maintain this knowledge from
generation to generation. However, the more recent settlers who may not be aware
of these varied uses could benefit from greater dissemination of knowledge around
these species to enhance their conservation value. Fragments of forests or relics,
known as islands of biodiversity, can be an alternative for the conservation of
biodiversity and all its ecological processes in areas close to cities or agricultural
landscapes. With the adequate management of the species they contain, the biolog-
ical connectivity necessary for the survival of the species can be achieved.
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 233

References

AIDER (2020) Promoviendo y concertando el desarrollo rural bajo en emisiones en la Región

Ucayali bajo el enfoque de producción, protección e inclusión (GCF-TF/PNUD Ucayali).
Análisis integrado de causas y mecanismos causales de deforestación y cambio de uso. Informe
Final de Análisis. Equipo técnico de Apoyo Regional Ucayali. Pucallpa, Perú, 71p
Armenteras D, Vargas O (2016) Patrones del paisaje y escenarios de restauración: acercando
escalas. Acta Biol Colomb 21((1) Supl):S229–S239. [Link]
50848
Begon M, Harper JL, Townsend CR (1996) Ecology individuals, populations and communities.
Blackwell Science, London, p 1068
Bennett A (1998) Enlazando el paisaje: el papel de los corredores y la conectividad en la
conservación de la vida silvestre. IUCN, Cambridge, 276pp
Bergoeing J (1998) Geomorfología de Costa Rica. Instituto Geográfico Nacional, San José, 408pp
Bierregaard RO, Lovejoy TE, Kapos V, dos Santos AA, Hutchings RW (1992) The biological
dynamics of the tropical rainforest fragments: a prospective comparison of fragments and
continuous forest. Bioscience 42:859–866
Brokaw N (2002) La persistencia de las especies vegetales en los fragmentos de bosque neotropical.
Source: Kattan GH, Fragmentación: patrones y mecanismos de extinción de especies. In:
Guariguata MR, Katan GH (eds) Ecología y Conservación de Bosques Neotropicales.
EULAC – GTZ, 1st edn. Ediciones LUR, Cartago, pp 564–565
Chávez F (1991) Composición florística de los bosques secundarios en tres etapas sucesionales en la
zona de Pucallpa. Tesis para optar el grado de Bachiller en Ciencias Biológicas. Universidad
Nacional San Antonio Abad del Cusco, Perú
Clavo M, Fernández-Baca J (1999) Regeneración natural de especies arbóreas para el
establecimiento de sistemas silvopastoriles. Revista de Investigaciones Veterinarias del Perú
10(1):71–81
Clavo Z, Roncal S, Ricse A, Sabogal C (2006) Composición florística post-quema en áreas
degradadas por la agricultura en la Región Ucayali, Amazonia peruana. Revista Forestal del
Perú 29(1–2):78–91
Cochrane T, Sánchez P (1982) Recursos de tierras, suelos y su manejo en la región Amazónica:
Informe acerca del estado de conocimientos. In: Hecht SB (ed) Amazonía, Investigación sobre
agricultura y uso de tierras. Centro Internacional de Agricultura Tropical, Cali, pp 141–218
Colorado Zuluaga GJ, Vásquez Muñoz JL, Mazo Zuluaga IN (2017) Modelo de conectividad
ecológica de fragmentos de bosque andino en Santa Elena (Medellín, Colombia). Acta Biol
Colomb 22(3):379–393
Conservación Amazónica–ACCA (2020) Hotspots y tendencias de deforestación en la Amazonía.
Monitoring of the Andean Amazon Project. [Link]
Consulted 18 May 2020
Dourojeanni M (1987) Aprovechamiento del barbecho forestal en áreas de Agricultura migratoria
en la Amazonía peruana. Revista Forestal del Perú CEDINFOR14(2):1–33
Estrada A, Coates-Estrada R (2001) Bat species richness in live fences and in corridors of residual
rain forest vegetation at Los Tuxtlas, Mexico. Ecography 24:94–102
Finer M, Mamani N (2018) Deforestation hotspots in the Peruvian Amazon. MAAP (Mapping the
Andean Amazon Project). Síntesis #3. [Link] Consulted
26 Jan 2020
Forest Stewardship Council (2020) Principios y criterios del FSC para el manejo forestal
responsable. FSC-STD-01-001 V5-2 E. Secretariado Internacional Charles-de-Gaulle-Str.5.
53113 Bonn, Germany, 32p. [Link] Consultado el 17 May 2020
Francesconi W, Montagnini F (2015) Los SAFs como estrategias para favorecer la conectividad
funcional del paisaje fragmentado. In: Montagnini F, Somarriba E, Murgueitio E, Fassola H,
Eibl B (eds) Sistemas Agroforestales. Funciones Productivas, Socioeconómicas y Ambientales.
Serie técnica. Informe técnico 402. CATIE, Turrialba. Editorial CIPAV, Cali, 454p
234 Z. M. Clavo Peralta et al.

Fujisaka S, Escobar G, Veneklaas E (2000) Weedy fields and forests: interactions between land use
and the composition of plant communities in the Peruvian Amazon. Agric Ecos Environ 78:
175–186
Gabriel VA, Pizo MA (2005) The use of fencerows by birds in a fragmented landscape of the
Atlantic rainforest. Natureza e Conservacão 3:79–89(portugués), 182–192(inglés)
García R (2002) Biología de la conservación: conceptos y prácticas. INBio, Santo Domingo de
Heredia, 166pp
Gazzo J (1982) Políticas y planes de desarrollo para la región amazónica del Perú. Amazonía,
Investigación sobre agricultura y uso de tierras. In: Hecht SB (ed) Centro Internacional de
Agricultura Tropical, Cali, pp 87–108
Gurrutxaga M (2004) Conectividad Ecológica del Territorio y Conservación de la Biodiversidad.
Nuevas Perspectivas en Ecología del Paisaje y Ordenación del Territorio. Departamento de
Agricultura y Pesca, Gobierno Vasco
Hobbs RJ, Wilson AM (1998) Corridors: theory, practice and achievement of conservation
objetives. In: Dover W, Bunce RGH (eds) Key concepts in landscape ecology. International
Association for Landscape Ecology, Preston, pp 265–279
Instituto Nacional de Estadística e Informática (INEI) (2018) Perú, Perfil Sociodemográfico. Censos
Nacionales 2017: XII de Población, VII de Vivienda y III de Comunidades Indígenas. Lima,
Perú, 641p
Kalliola R, Puhakka M (1993) Geografía de la Selva Baja Peruana. In: Kalliola R, Puhakka M,
Danjoy W (eds) Amazonía Peruana. Vegetación húmeda tropical en el llano Sub Andino.
Publicado por Proyecto Amazonía, Universidad de Turko y la Oficina Nacional de Evaluación
de Recursos Naturales, Lima, pp 9–21
Kattan G (2002) Fragmentación de patrones y mecanismos de extinción de especies. In: Guariguata
MR, Katan GH (eds) Ecología y Conservación de Bosques Neotropicales. EULAC – GTZ, 1st
edn. Ediciones LUR, Cartago, pp 561–590
López E, Baldoceda R, Clavo ZM, Veneklaas E (2005) Estudio del banco de semillas presentes en
suelos de barbechos originados por efecto de agricultura y pastura. Revista Forestal de Ucayali
2(2):1–15
Martín J, Fernández L, Urios G (2013) Los Bosques isla en Andalucía. Consejería de Medio
Ambiente y Ordenación del Territorio, Junta de Andalucía, Sevilla, 192pp
McIntyre NE (1995) Effects of forest patch size on avian diversity. Landsc Ecol 10(2):85–99
Ministerio de Agricultura (MINAGRI) (2012) Plan Estratégico Regional del sector agrario de
Ucayali. Gobierno Regional de Ucayali. Dirección Regional Agraria de Ucayali. Periodo:
2008–2012. Pucallpa, Perú
Ministerio del Ambiente (MINAN). (2020). Programa GeoBosques. Plataforma de monitoreo de
cambios sobre la cobertura de los bosques. Bosque y pérdida de bosque en la región Ucayali.
Lima. Perú. [Link] Accesed on line
28 Jan 2020
Ministerio del Ambiente (MINAM) (2020) Servicio Nacional de Áreas Naturales Protegidas
(SERNANP). Sistema de Áreas Naturales Protegidas del Perú. Áreas naturales protegidas de
administración nacional con categoría definitiva. Lima Perú. [Link]
documents/10181/165150/Listado+ANP+[Link]/5e1a7af5-ab65-455c-9094-
f86bf9b994d4. Consulted 28 Jan 2020
Morera C, Pintó J, Romero M (2007) Paisaje, procesos de fragmentación y redes ecológicas:
aproximación conceptual. In: Chassot O, Morera C (eds) Corredores Biológicos: acercamiento
conceptual y experiencias en América. Centro Científico Tropical/Universidad Nacional de
Costa Rica, San José, 128pp
Nott MP, Pimm SL (1997) The evaluation of biodiversity as a target for conservation. In: Pickett
STA, Ostfeld RS, Shachak M, Likens GE (eds) The ecological basis of conservation. Springer,
Boston
Porro R, López A, Vela J (2015) Forest use and agriculture in Ucayali, Peru: livelihood strategies,
poverty and wealth in an Amazon frontier. Forest Policy Econ 51:47–56
9 Biodiversity Islands and Dominant Species in Agricultural Landscapes. . . 235

Redondo-Brenes A, Montagnini F (2010) Contribution of homegardens, agrosilvopastoral systems,

and other human-dominated land-use types to the avian diversity of a biological corridor in
Costa Rica. Adv Environ Res 2(2):111–148
Riesco A, Ara M, De La Torre M (1995) Informe final Proyecto Sistemas Amazónicos Sostenibles.
SAS. Financiado por CIID Canadá, Ejecutado por IVITA-Pucallpa
Robiglio V, Reyes M, Castro E (2015) Diagnóstico de los productores familiares en la Amazonia
Peruana. ICRAF, Oficina Regional para América Latina, Lima, Perú. Por encargo de
GGGI&DIE
Rodríguez F (2003) Perú: Ordenamiento Territorial con base en la Zonificación Ecológica
Económica en la Amazonia. Programa de Ordenamiento Ambiental. Instituto de Investigaciones
de la Amazonía Peruana. Iquitos, Perú
Sabogal C, Colán V, Galván O, Current D, Dominguez G, Yepes F (2001) Aspectos metodológicos
para determinar opciones de manejo de bosques secundarios (purmas) en la Amazonia peruana,
con especial referencia a la zona de Pucallpa. Revista Forestal del Perú 24(1–2):85–109
Salisbury DS, Fagan C (2011) Coca and conservation: cultivation, eradication, and trafficking in the
Amazon borderlands. GeoJournal 11:1–20
Toledo J, Serrao A (1982) Producción de pastos y ganado en la Amazonía. In: Hecht SB
(ed) Amazonía, Investigación sobre agricultura y uso de tierras. Centro Internacional de
Agricultura Tropical, Cali, pp 295–324
Tuomisto H (1993) Clasificación de vegetación en la Selva Baja Peruana. In: Kalliola R,
Puhakka M, Danjoy W (eds) Amazonía Peruana. Vegetación húmeda tropical en el llano Sub
Andino. Publicado por Proyecto Amazonía, Universidad de Turko y la Oficina Nacional de
Evaluación de Recursos Naturales, Lima, pp 103–112
Vandermeer J, Perfecto I, Philpott S, Chappell MJ (2008) Reenfocando la conservación en el
paisaje: la importancia de la matriz. In: Harvey CA, Sáenz JC (eds) Evaluación y conservación
de biodiversidad en paisajes fragmentados de Mesoamérica. INBio, Santo Domingo de Heredia,
pp 75–104
Vela J, Clavo M, Alvez-Valles C, Mori J, Panduro N, Ramírez N, Bautista J, Tavera I, Fernández V
(2019) Construcción de un modelo de ganadería sostenible con pequeños productores de leche
como una opción de mitigación y adaptación al calentamiento global. Amazonía Sur Occidental,
Pucallpa, Perú. Informe de Avance 2019. Facultad de Ciencias Agropecuarias, Vicerrectorado
de investigación, Universidad Nacional de Ucayali, Pucallpa, Perú, 40p
Vivanco L (2004) Gran enciclopedia de la región Ucayali. Amazonía Peruana: Identidad Regional
Pucallpa. Perú, 303p
Chapter 10
The Monteverde Cloud Forest: Evolution
of a Biodiversity Island in Costa Rica

Quint Newcomer, Fabricio Camacho Céspedes, and Lindsay Stallcup

Abstract Monteverde, Costa Rica represents an example of dynamic processes

shaping an ever-changing, landscape-scale biodiversity island. Monteverde is inter-
nationally renowned for biodiversity conservation efforts initiated by
non-governmental organizations and private citizens that led to the creation of the
Monteverde Reserve Complex—a network of reserves spanning the region. Located
in the Tilarán Mountain Range, an area of high endemism in the Central American
isthmus, Monteverde’s reserves provide habitat for over half of the species found in
the entire country of Costa Rica, including 55 species of birds, mammals, amphib-
ians and reptiles with some degree of threatened status on the IUCN Red List. One
characteristic that makes Monteverde unusual is the number of research scientists
that have settled in the area and studied the region over multiple decades. Some of
these scientists helped secure international funding to purchase land for the
Monteverde Cloud Forest Biological Preserve and the Children’s Eternal Rainforest
and participated in the creation of local non-governmental organizations to promote
conservation, education, and sustainable community development. Recognizing that
Monteverde’s biodiversity island requires habitat connectivity across a larger land-
scape to support seasonal migratory species, Monteverde’s organizations established
the Bellbird Biological Corridor. The impacts of changing climate conditions—in
particular, the increase of daily minimum temperatures and the increase in number of
consecutive dry day periods—are being observed in Monteverde’s cloud forests and
further threaten the conservation of habitat and species. Holistic policies and pro-
grams spanning tourism, agriculture, transportation, energy, and environmental
sectors are needed for continued conservation successes.

Q. Newcomer (*)
University of Georgia, Warnell School of Forestry and Natural Resources, Athens, GA, USA
F. Camacho Céspedes
Tropical Science Center, Monteverde, Costa Rica
L. Stallcup
Monteverde Conservation League, Monteverde, Costa Rica

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 237
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
238 Q. Newcomer et al.

Keywords Biodiversity conservation · Biological corridors · Endangered species ·

Grassroots conservation · Private land conservation · Tropical Forest conservation

10.1 Introduction

The arc of the Central American isthmus spans approximately 1,835 km (1,140 mi)
from the southeast in Panama to the northwest in Guatemala. At its narrowest point,
the isthmus is only 50 km (30 mi) wide. The region serves as a land bridge between
North and South America, a critical path and melting pot for species from both
hemispheres that colonized the isthmus during the past three million years. Coastal
plains on both Atlantic and Pacific slopes are connected by a mountainous spine. Yet
the mountains also form a barrier of separation between Atlantic and Pacific slopes,
which have distinct ecological conditions shaped by both geology and climate
patterns. As a result, in a relatively small area, Central America has great biodiversity
with many micro-climates. In Costa Rica alone, Holdridge (1967) described 13 life
zones.
Monteverde, Costa Rica is a small town (population less than 1,000) at the end of
a meandering mountain road. The town of Monteverde lends its name to the
surrounding region, which includes the population center of Santa Elena, and several
other communities including San Luis, Cerro Plano, Cañitas, and La Lindora.
Monteverde also lends its name, particularly within the international tourism indus-
try, to the region’s natural environment—the Monteverde Cloud Forest.
In this chapter, we describe the broader region of Monteverde, including both the
town and the region’s cloud forests, as a case study example of the processes shaping
a dynamic, large-scale biodiversity island.1 Monteverde is internationally renowned
for privately-owned biodiversity conservation efforts that led to the creation of the
greater Monteverde Reserve Complex—a network of reserves spanning the region.
Its exceptional natural environment is complemented by a distinctive socio-cultural
environment which was fundamental to the establishment of the network of reserves
and more recently the expansion of the biological corridor extending outward from
Monteverde through the region (see Sect. 3.3). Monteverde’s outstanding conserva-
tion successes are tempered by many challenges. Based on our personal experience
as administrators of local organizations and active participants in community-based
conservation initiatives during the past three decades, it the authors’ perspective that
the ability of Monteverde’s citizenry—including governmental offices,

1
For background and general description of the region’s ecology and conservation history, we rely
heavily on the seminal work of Nalini Nadkarni and Nathaniel Wheelwright, Monteverde: Ecology
and Conservation of a Tropical Cloud Forest, first published in 2000 with updated chapters
published in 2014. We encourage readers interested in a more in-depth understanding of
Monteverde to reference this outstanding resource. Our reflections on conservation initiatives in
Monteverde, including the Bellbird Biological Corridor, the Children’s Eternal Rainforest, Enlace
Verde, and local reforestation projects are largely derived from our personal involvement in these
initiatives and organizations.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 239

non-governmental organizations, private-sector businesses, and engaged individual

community members—to self-reflect, adapt to local issues and to external, global-
scale threats, and continue to lead biodiversity conservation efforts makes for an
interesting case study.
Nestled high in the Tilarán Mountain Range in northwestern Costa Rica, the
Monteverde region is home to the Monteverde Reserve Complex, a 27,500-hectare
block of protected forest comprised of three privately owned reserves (the
Monteverde Cloud Forest Biological Preserve, Children’s Eternal Rainforest, and
Bosqueterno) and the state-owned Santa Elena Cloud Forest Reserve (see Fig. 10.1).
The Reserve Complex is the centerpiece of a larger protected area, which we will
refer to as the Monteverde Arenal Bioregion (MAB), that includes the state-owned
Arenal Volcano National Park and Alberto Manuel Brenes Biological Reserve (see
Fig. 10.2) and other smaller reserves. The MAB is connected to other protected areas
via biological corridors, including the Bellbird Biological Corridor, which aims to
connect Monteverde with the mangrove forests on Costa Rica’s Pacific gulf coast.

Fig. 10.1 The Monteverde Reserve Complex is comprised of three privately-owned reserves—the
Children’s Eternal Rainforest, owned by the Monteverde Conservation League; the Monteverde
Cloud Forest Biological Preserve, owned by the Tropical Science Center; and Bosqueterno S.A., a
corporation whose land assets are managed by the Tropical Science Center and which is owned by
Quaker families who settled in Monteverde in the 1950s. Map Source: Yuber Rodríguez,
Monteverde Conservation League 2020
240 Q. Newcomer et al.

Fig. 10.2 The Monteverde Arenal Bioregion surrounding the Monteverde Reserve Complex
includes two large government-owned protected areas—Arenal Volcano National Park and the
Alberto Manuel Brenes Biological Reserve, several smaller reserves and protected areas, and the
Bellbird Biological Corridor which connects Monteverde with the Gulf of Nicoya. Map Source:
Yuber Rodríguez, Monteverde Conservation League 2020

The properties within the Monteverde Reserve Complex span seven Holdridge
life zones (Haber 2000a; Holdridge 1967; Tosi et al. 1969). The elevational gradient
of the Monteverde Reserve Complex ranges from 700 meters above sea level (masl)
on the Pacific slope up to the Continental Divide—1,850 masl at its highest point—
and down the Atlantic slope to 466 masl. The species on the Pacific slope have
greater affinity with the tropical dry forest ecosystem, while organisms on the
Atlantic slope tend to be more adapted to the humid conditions of the tropical rain
forest. The species found at higher elevations are also tolerant to cool, humid
conditions and are adapted to resist strong easterly trade winds, heaviest between
December – March with gusts up to 80 kmph (Tropical Science Center unpublished
data). Much of the Monteverde region falls within the tropical montane cloud forest
vegetation type (see Fig. 10.3), which, according to Nadkarni and Wheelwright
(2000: 9) is “one of the world’s most threatened ecosystems.”
As traditionally understood, islands undergo geological processes of formation—
the eruption of volcanoes creating igneous landforms, and the collision and subduc-
tion/abduction of continental plates lifting up land masses. While geological forces
created the underlying conditions for the establishment of the cloud forest and the
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 241

Fig. 10.3 A landscape view of the Monteverde Cloud Forest Preserve and the Children’s Eternal
Rainforest from the Continental Divide looking down the Atlantic slope. The Monteverde Reserve
Complex represents the largest privately-owned protected area in Central America. Image Source:
Fabricio Camacho Céspedes

evolution of the ecological niches that established along the Continental Divide in
the Tilarán Mountain range in Costa Rica, the formation of Monteverde’s biodiver-
sity island was driven by strong socio-political, economic, cultural components over
decades, rather than millennia. Expansionist development policies encouraged
human population incursion into forested regions. Frontiersmen’s axes felled seem-
ingly inexhaustible tropical forests, fires “cleaned” the land, and cattle farms
expanded across the landscape. As Harvey and Haber (1999) describe, this process
resulted in the creation of small, sometimes single remnant tree biodiversity islands
throughout the Monteverde region.
Agricultural expansion in the surrounding landscape completed the transforma-
tion of the remaining forests in the Monteverde region into a biodiversity island.
Forested highland hillsides were replaced by coffee plantations and dairy cattle
operations. Rivers flowing to the Caribbean were dammed and diverted to the
Pacific, providing hydropower and supplying irrigation to the dry northwestern
province of Guanacaste. The arid tropical dry forest and drained coastal wetlands
were replaced with extensive fields of cattle, rice, sugar cane, and eventually
pineapple. To the north of the Monteverde region, extensive tourism development
242 Q. Newcomer et al.

modified the landscape near the Arenal Volcano; ornamental plants, pineapple, and
family farms dominated areas east of the Monteverde Reserve Complex.
Yet in the midst of this landscape transformation, there were also concerted
efforts to protect the region’s natural resources. In the 1950s, Quaker settlers in
Monteverde recognized the need to maintain forested lands that protected highland
springs, the community’s primary water source. This area, known as Bosqueterno,
would eventually become the nucleus of the Monteverde Cloud Forest Biological
Preserve, which was founded in 1972. In 1977, the Costa Rican government
declared protected areas in the region (the Arenal Forest Reserve, later to become
the Zona Protectora Arenal Monteverde, or ZPAM). The government was unable to
purchase or effectively manage the lands it had declared as a reserve, however, and
private landowners continued agricultural expansion into their farms in this area.
Private conservation initiatives, led by the Tropical Science Center and Monteverde
Conservation League, sprang up in an effort to purchase primary rainforest before it
was felled (see Sect. 3.1). In the 1980s and 1990s, Costa Rica’s international debt
would be swapped for investments into the expansion of the Monteverde biodiver-
sity island, eventually—in combination with funds raised by youth around the
world—creating the Children’s Eternal Rainforest.
The focus has shifted over time from protecting water to protecting specific
endangered species, then to ecosystem conservation (Burlingame 2000). Most
recently, the Monteverde community is engaged in an evolving initiative integrating
farm-scale to watershed-scale ecosystem management and sustainable economic
development with the goal of climate change resiliency for the region’s biodiversity
and the growing human population (Brenes et al. 2019). Simultaneously, research
scientists and conservation organizations are working to develop a more unified
approach to share data, collaborate across disciplines, and perhaps develop new
methods and approaches for understanding and disseminating information about the
region’s ecology (Allen and Hoekstra 1992; Zamsow et al. 2018).
The following sections highlight natural history characteristics as well as some of
the organizations and initiatives that make Monteverde’s biodiversity island unique
and of critical importance as a large node in Costa Rica’s national landscape matrix
of conservation. We then describe some of the challenges for ongoing biodiversity
conservation, many of which are not unique to the Monteverde region. We remind
the reader that the complete story of biodiversity conservation in Monteverde, Costa
Rica is far more complex than we are able to present in this snapshot.

10.2 Species Accounts

“Biodiversity in the broadest sense is a characteristic feature of Monteverde”. Wheelwright

2000:420.

The diversity of micro-climates (or life zones) in the Monteverde region positively
impacts the area’s overall biodiversity. The Atlantic slope of Monteverde has the
greatest bird diversity in Costa Rica—40% of Central America’s mammal species
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 243

have been identified in Monteverde. The area is home to one-third of Costa Rica’s
plant species, including as many families of epiphytes as the entire country of
Mexico. Ten percent of Monteverde’s plants are endemic to Costa Rica’s Tilarán
mountain range (Wheelwright 2000).In this section, we offer a brief glimpse into
Monteverde’s natural history. For a much more complete and detailed description
refer to Nadkarni and Wheelwright (2000, 2014).

10.2.1 Epiphytes, Orchids

Characteristic to Monteverde’s vegetation are its diverse epiphyte communities

which represent the most diverse plant group in Monteverde with more than 800 spe-
cies including 471 species of monocots, 230 species of dicots, and 177 species of
ferns (Haber 2000a, b). These communities not only add to the overall biodiversity
in Monteverde, but also play a significant role in cloud forest ecosystem dynamics.
For example, many of these plants decompose in the forest canopy, producing an
abundant accumulation of canopy soil that is rich in organic matter and which
contributes to nutrient cycling and conservation (Nadkarni 1981; Nadkarni et al.
2002). The canopy epiphyte root-humus mat exhibit high potential water storage
capacity (Köhler et al. 2007). However, the actual water availability in this canopy-
mat is conditioned to rainfall and evaporation patterns (Köhler et al. 2007), which is
one of the main contributing factors that suggest that this may be one of the most
vulnerable communities to the effects of climate change (Clark et al. 2014).
Orchids represent the most diverse epiphyte family in Monteverde with more than
450 species (Haber 2000a, b), of which more than 30 species are new to science
(Atwood and Dressler 1995). This richness makes Monteverde one of the most, if
not the most, orchid-diverse locations on the planet (Haber 2000a, b). The most
important aspects that add up to this diversity are the abundance of life zones,
pollinators and the intensity of studies in the area (Atwood and Dressler 1995).
Orchid diversity in Monteverde represents about one third of the estimated number
of orchid species in Costa Rica (Atwood and Dressler 1995). Although it is one of
the most prominent families in terms of species diversity, many species known as
miniature orchids are inconspicuous and difficult to recognize among other epi-
phytes by the untrained eye.

10.2.2 Vascular Plants

Within the seven Holdridge life zones (Holdridge 1967) included in the Monteverde
Flora Project study area, there are three forest types—Pacific slope seasonal forest,
cloud forest, and Atlantic slope rain forest (Haber 2000a). The Monteverde Flora
Project was an initiative of the Missouri Botanical Garden and the Manual to the
Plants of Costa Rica Project. Through the Monteverde Flora Project, the flora of the
244 Q. Newcomer et al.

Monteverde area, from the Continental Divide down both Atlantic and Pacific slopes
to 700 masl, was collected and identified (Haber 1991, 2000a, b).
In addition to the Monteverde Flora Project, multiple studies since the mid-1970s
have contributed to the identification and classification of Monteverde’s flora (e.g.,
Dyer 1979; Hartshorn 1983). By 2000, the list of vascular plants for Monteverde
included 3,021 species with 755 species of trees (Haber 2000a, b). This represents
about one-third of all vascular plant species in Costa Rica (Haber 2000a, b). About
10% of Monteverde’s flora species are endemic to the Tilarán Mountains (Haber
2000a, b), and over 10% of the 216 of the common or characteristic tree species of
the life zones of Monteverde face some degree of threat as per the IUCN Red List
(see Table 10.1).
The diversity of wild avocados—66 species of Lauraceae representing eight
genera (Haber 2000a, b)—in Monteverde is notably greater than in adjacent areas.
Lauraceae play a key role as a food source for many frugivorous birds, including
Three-wattled Bellbirds (Procnias tricarunculata) and Resplendent Quetzals
(Pharomacros moccino) (see Fig. 10.4), which synchronize their migration routes
to follow the fruiting patters of these and other tree species (Powell and Bjork 1994).
The lipids contained in the wild avocados serve as one of the main sources of energy
to support the reproduction of these iconic birds.

10.2.3 Avifauna

Many early scientific publications (e.g., Snow 1977; Wheelwright 1983) focused on
Monteverde’s spectacular avifauna, capturing the attention of the international
birding community. Natural history films (e.g., BBC’s Forest in the Clouds),
books (e.g., National Geographic’s Mountain Worlds), and magazine articles (e.g.,
International Wildlife’s “Is This the Garden of Eden?”) based on the region’s
impressive biodiversity also brought international attention to the area (Burlingame
2000).
To date, more than 400 of Costa Rica’s 850 avian species have been reported in
the area (Fogden 2000), including a number of IUCN Red Listed species such as the
Great Curassow (Crax rubra), Keel-billed Motmot (Electron carinatum), Blue-and-
gold Tanager (Bangsia arcaei), the above-mentioned Three-wattled Bellbird and
Resplendent Quetzal, and the Bare-necked Umbrellabird (Cephalopterus
glabricollis) (see Table 10.1). Avian diversity is amplified by the combination of
resident and migratory communities, which include latitudinal and altitudinal
migrants. Among the latter, the most economically important species is the Resplen-
dent Quetzal, due to its importance to Monteverde’s ecotourism industry.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 245

Table 10.1 Species found in the Monteverde area which are listed as Near Threatened, Vulnerable,
Endangered, Critically Endangered, Extinct in the Wild, or Extinct on the IUCN Red List
Common
Species name IUCN Red List Status Citation
Plantsa
Agonandra Vulnerable Nelson (1998a)
macrocarpa
Bombacopsis Vulnerable Sandiford (1998)
quinata
Capparis Near Threatened Mitré (1998a)
discolor
Chrysophyllum Near Threatened World Conservation
hirsutum Monitoring Centre
(1998a)
Costus nitidus Endangered Skinner (2014)
Dichapetalum Vulnerable World Conservation
costarricense Monitoring Centre
(1998b)
Elaeagia Endangered World Conservation
uxpanapensis Monitoring Centre
(1998c)
Eugenia Endangered Nelson (1998b)
salamensis
Ficus lateriflora Critically Endangered Page (1998)
Ilex Vulnerable Mitré (1998b)
costaricensis
Magnolia Near Threatened Khela (2014)
poasana
Ocotea Quizzará Critically endangered Joslin et al. (2018)
monteverdensis Blanco
Ocotea Vulnerable World Conservation
viridiflora Monitoring Centre
(1998d)
Oreomunnea Endangered Americas Regional
pterocarpa Workshop (1998)
Persea Coyo Endangered Wegier et al. (2017)
schiedeana Avocado
Pouteria austin- Vulnerable World Conservation
smithii Monitoring Centre
(1998e)
Pouteria Vulnerable World Conservation
congestifolia Monitoring Centre
(1998f)
Pouteria Vulnerable World Conservation
fossicola Monitoring Centre
(1998g)
Sideroxylon Near Threatened World Conservation
capiri Monitoring Centre
(1998h)
(continued)
246 Q. Newcomer et al.

Table 10.1 (continued)

Common
Species name IUCN Red List Status Citation
Sideroxylon Near Threatened World Conservation
persimile Monitoring Centre
(1998i)
Terminalia Endangered Nelson (1998c)
bucidoides
Ticodendron Duranzo de Near Threatened Rivers et al. (2019)
incognitum Ardilla
Zinowiewia Near Threatened Mitré (1998c)
costaricensis
Amphibians2
Agalychnis Blue-sided Endangered IUCN SSC Amphib-
annae Treefrog ian Specialist Group
and NatureServe
(2014)
Agalychnis Lemur Leaf Critically Endangered Solís et al. (2008d)
lemur Frog
Atelopus varius Variable Har- Critically Endangered Pounds et al. (2010)
lequin Frog
Bolitoglossa La Palma Endangered Pounds et al. (2008c)
subpalmata Salamander
Craugastor Atlantic Rob- Critically Endangered Pounds et al. (2008a)
andi ber Frog
Craugastor Angel Robber Critically Endangered Pounds et al. (2008b)
angelicus Frog
Craugastor Cerro Utyum Near Threatened Solís et al. (2010a)
podiciferus Robber Frog
Duellmanohyla Costa Rica Endangered NatureServe and
uranochroa Brook Frog IUCN SSC Amphib-
ian Specialist Group
(2013)
Ecnomiohyla Heredia Endangered Solís et al. (2008a)
fimbrimembra Treefrog
Ecnomiohyla Cope’s Brown Vulnerable Solís et al. (2010b)
miliaria Treefrog
Incilius Golden Toad Extinct Savage et al. (2008)
periglenes
Isthmohyla Narrow-lined Critically Endangered Solís et al. (2008c)
angustilineata Treefrog
Isthmohyla American Critically Endangered Solís et al. (2010c)
rivularis Cinchona
Plantation
Treefrog
Isthmohyla tica Starrett’s Critically Endangered IUCN SSC Amphib-
Treefrog ian Specialist Group
and NatureServe
(2013a)
(continued)
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 247

Table 10.1 (continued)

Common
Species name IUCN Red List Status Citation
Isthmohyla Zetek’s Near Threatened Solís et al. (2008b)
zeteki Treefrog
Lithobates Rancho Vulnerable IUCN SSC Amphib-
vibicarius Redondo Frog ian Specialist Group
and NatureServe
(2013b)
Nototriton Monteverde Vulnerable Pounds et al. (2008f)
gamezi Moss
Salamander
Nototriton La Estrella Near Threatened Bolaños et al.
picadoi Salamander (2008b)
Oedipina poelzi Quarry Worm Endangered Bolaños et al.
Salamander (2008a)
Oedipina Cienega Colo- Near Threatened Wake et al. (2008)
uniformis rado Worm
Salamander
Pristimantis Mountain Near Threatened Pounds et al. (2008d)
altae Robber Frog
Pristimantis La Loma Near Threatened Pounds et al. (2008e)
caryophyllaceus Robber Frog
Reptilesb
Celestus hylaius Rain Forest Near Threatened Solórzano et al.
Caiman (2013)
Lizard
Trimetopon Dunn’s Tropi- Endangered Porras et al. (2013)
simile cal Ground
Snake
Insects (Odonata)c
Libellula Near Threatened Paulson (2009b)
mariae
Palaemnema Endangered Paulson and von
baltodanoi Ellenrieder (2006a)
Perigomphus Vulnerable Paulson and von
pallidistylus Ellenrieder (2006b)
Birdsd
Ara ambiguus Great Green Endangered (once abundant in BirdLife Interna-
Macaw Atlantic Slope lower elevations, but tional (2016a)
has not been recently sighted)
Antrostomus Chuck-will’s Near Threatened BirdLife Interna-
carolinensis Widow tional (2018a)
Bangsia arcaei Blue-and-gold Near Threatened BirdLife Interna-
Tanager tional (2018b)
Buteogallus Black Solitary Near Threatened BirdLife Interna-
solitarius Eagle tional (2016b)
(continued)
248 Q. Newcomer et al.

Table 10.1 (continued)

Common
Species name IUCN Red List Status Citation
Cephalopterus Bare-necked Endangered Birdlife International
glabricollis Umbrellabird (2016c)
Contopus Olive-sided Near Threatened BirdLife Interna-
cooperi Flycatcher tional (2017a)
Crax rubra Great Vulnerable BirdLife Interna-
Curassow tional (2016d)
Electron Keel-billed Vulnerable BirdLife Interna-
carinatum Motmot tional (2016e)
Grallaricula Ochre-breas- Near Threatened BirdLife Interna-
flavirostris ted Antpitta tional (2017b)
Hylocichla Wood Thrush Near Threatened BirdLife Interna-
mustelina tional (2017c)
Morphnus Crested Eagle Near Threatened BirdLife Interna-
guianensis tional (2017d)
Patagioenas Ruddy Pigeon Vulnerable BirdLife Interna-
subvinacea tional (2016f)
Pharomachrus Resplendent Near Threatened Birdlife International
mocinno Quetzal (2016g)
Procnias Three-wattled Vulnerable Birdlife International
tricarunculatus Bellbird (2016h)
Sclerurus Gray-throated Near Threatened BirdLife Interna-
albigularis Leaftosser tional (2016i)
Setophaga Cerulean Near Threatened BirdLife Interna-
cerulea Warbler tional (2019a)
Spizaetus Ornate Hawk- Near Threatened BirdLife Interna-
ornatus Eagle tional (2016j)
Sturnella Eastern Near Threatened BirdLife Interna-
magna Meadowlark tional (2019b)
Tinamus major Great Near Threatened BirdLife Interna-
Tinamou tional (2017e)
Touit Red-fronted Vulnerable BirdLife Interna-
costaricensis Parrotlet tional (2018c)
Vermivora Golden- Near Threatened BirdLife Interna-
chrysoptera winged tional (2018d)
Warbler
Mammals5
Ateles geoffroyi Black-handed Endangered Cuarón et al. (2008)
Spider
Monkey
Ectophylla alba Caribbean Near Threatened Rodriguez and
White Bat Pineda (2015)
Leopardus Northern Vulnerable Payan and de
tigrinus Tiger Cat Oliveira (2016)
Leopardus Margay Near Threatened de Oliveira et al.
wiedii (2015)
(continued)
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 249

Table 10.1 (continued)

Common
Species name IUCN Red List Status Citation
Myrmecophaga Giant Anteater Vulnerable (once present in Miranda et al. (2014)
tridactyla Monteverde, but not recently
sighted)
Panthera onca Jaguar Near Threatened Quigley et al. (2017)
Sylvilagus Tapeti Endangered Ruedas and Smith
brasiliensis (2019)
Tapirus bairdii Baird’s Tapir Endangered García et al. (2016)
Tayassu pecari White-lipped Vulnerable (once present in Keuroghlian et al.
Peccary Monteverde, but not recently (2013)
sighted)
Vampyrum Spectral Bat Near Threatened Solari (2018)
spectrum
a
Based on database search of IUCN Red List ([Link]) using the 216 species listed in
Table 3.2. Common or characteristic tree species of the life zones in the Monteverde area in the
chapter Plants and Vegetation (Haber 2000a, b) in Nadkarni and Wheelwright (2000). An exhaus-
tive list of the status of all flora in the Monteverde area is beyond the scope of this chapter.
Furthermore, many of the 3021 species of vascular plants known to the Monteverde area have not
been evaluated for their IUCN Red List endangered status. Therefore, a representative sample of the
most common or characteristic tree species (216 of the 755 known tree species in Monteverde) was
deemed appropriate for the purposes of this chapter
b
Based on database search of IUCN Red List ([Link]) using the complete species list of
Amphibians and Reptiles of Monteverde provided in Pounds and Fogden (2000). This was also
cross-referenced with the most recent unpublished Amphibians and Reptiles species lists for the
Monteverde Cloud Forest Biological Preserve compiled by the Tropical Science Center (2019
unpublished data) to ensure any new species added to the lists of observed species in the
Monteverde area were included
c
An exhaustive study of all insect orders in the Monteverde area has yet to be completed. We have
chosen to use the order Odonata as an example subset of insects representing the Monteverde area.
Based on database search of IUCN Red List ([Link]) using the complete species list
(n ¼ 102) of Odonata collected at 700 m or higher on both Pacific and Atlantic slopes within the
Monteverde area provided by Haber (personal communication, 2020 unpublished data). An addi-
tional species of Odonata, Epigomphus subsimilis, found in the lower Pacific slopes (355 m) is
IUCN Red Listed as Endangered (Paulson 2009a, b). 51 of the 102 species identified by Haber in
the Monteverde area have not been evaluated for inclusion in the IUCN Red List Database, and five
of those 51 species were not included in the Catalogue of Life database ([Link]
org/col/search/all). One Odonata species was included in the IUCN Red List database, however its
status could not be determined due to deficient data
d
Based on database search of IUCN Red List ([Link]) using the complete species list of
Birds of the Monteverde Area provided in Fogden (2000). This was also cross-referenced with the
most recent unpublished Bird species list for the Monteverde Cloud Forest Biological Preserve
compiled by the Tropical Science Center (2019 unpublished data) to ensure any new species added
to the lists of observed species in the Monteverde area were included
e
Based on database search of IUCN Red List ([Link]) using the complete species list of
Mammals of Monteverde provided in Timm and LaVal (2000). This was also cross-referenced with
the most recent unpublished Mammals species lists for the Monteverde Cloud Forest Biological
Preserve compiled by the Tropical Science Center (2019 unpublished data) to ensure any new
species added to the lists of observed species in the Monteverde area were included
250 Q. Newcomer et al.

Fig. 10.4 The Resplendent Quetzal (Pharomacros moccino) and the Three-wattled Bellbird
(Procnias tricarunculata) represent two of Monteverde’s iconic, IUCN Red Listed bird species.
Both species have attracted scientific researchers to the region and have played an important role in
attracting bird watchers and nature-based tourism in general to Monteverde. (Image Sources:
Alvaro Cubero (Resplendent Quetzal) and Orlando Calvo (Three-wattled Bellbird))

Even this flagship species has not escaped environmental threats; the Tropical
Science Center2 estimates a current population of 55 individual quetzals (male,
female and juveniles) in the Monteverde Cloud Forest Biological Preserve (TSC
unpublished data) in contrast to the 50 reproductive couples estimated by Wheel-
wright (1983). Hamilton et al. (2003) report habitat loss and fragmentation on the
Pacific Slope as the main causes of population declines of Three-wattled Bellbirds in
Monteverde. This species declined from 135 individuals in 1997, to just 90 in 2002.
Current monitoring efforts by the Tropical Science Center in the Monteverde region
revealed that the population of Bellbirds may be increasing; 102 individuals were
identified in 2018 and 120 in 2019 (TSC unpublished data).
Over one-third of Costa Rica’s hummingbird species have been identified in
Monteverde (Feinsinger 1977). It is believed that cooler air temperatures at higher
elevations may result in a lower abundance of pollinating insects, thus providing
conditions that support a higher diversity of hummingbirds due to lower resource
competition. Several species of vascular plants, including Justicia sp. (Acanthacea)

2
The Tropical Science Center (TSC) is Costa Rica’s oldest environmental non-profit,
non-governmental organization. TSC was established in 1962 by research scientists Leslie
Holdridge, Joseph Tosi, and Robert Hunter together with several local businessmen. See Sect.
3.1.1 for further description of TSC’s role in the Monteverde Reserve Complex.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 251

and Satyria sp. (Ericaceae), have coevolved pollination systems to capitalize on

diverse hummingbird phenotypes (Deliso 2008).

10.2.4 Insects

There has been no comprehensive study of insects in Monteverde, by far the most
diverse and abundant group of organisms in the region. The region is home to
102 species of damselflies and dragonflies (W Haber personal communication).
Monteverde’s butterflies have been studied by Haber (1993) and Stevenson
(W Haber and R Stevenson, unpublished data), who indicate a total of 658 species
in the Monteverde zone (Stevenson and Haber 2000, W Haber personal communi-
cation). More than half of the area’s butterfly species are altitudinal migrants,
moving up and down the Pacific and Caribbean slopes with the wet and dry seasons
(Stevenson and Haber 2000). Habitat availability is critically important, and chang-
ing climate conditions are driving up-slope migration of Pacific lowland butterfly
species—in addition to lowland birds and mammals—into the Monteverde area
(J Porter personal communication; Wheelwright 2000; Timm and LaVal 2000).
Ant faunas of cloud forests are strongly specialized for cloud forest habitat and
sharply differentiated from the adjacent lowlands. The ant fauna of Monteverde has
been extensively studied by Longino, who estimated that about 70 species are cloud
forest specialists (J Longino personal communication). About a third of these are
widespread, known from other cloud forest sites in Central America. Most of the rest
are restricted to the cloud forests of Costa Rica and adjacent parts of Panama. A few
are known only from Monteverde, but it is difficult to know if this is true local
endemism or a result of under-sampling elsewhere. Recent DNA studies are showing
that some of the species shared among different Costa Rican mountain ranges
actually have deep genetic divergences and have been isolated from each other for
millions of years (J Longino personal communication).

10.2.5 Larger Mammals

Three species of monkeys—Mantled Howler Monkeys (Alouatta palliata), White-

faced Capuchins (Cebus capucinus), and the IUCN Red List endangered Black-
handed Spider Monkeys (Ateles geoffroyi); agoutis (Dasyprocta punctate), pacas
(Cuniculus paca), and prehensile-tailed porcupines (Coendou mexicanus) among
other rodents; coatis (Nasua narica), peccaries (Tayassu pecari and Pecari tajacu),
sloths (Bradypus variegatus and Choloepus hoffmanni), kinkajous (Potos flavus),
olingos (Bassaricyon gabbii), and opossums (Didelphis marsupialis and Marmosa
mexicana) may be observed during visits to one of Monteverde’s reserves. The area
is home to all six of Costa Rica’s wild felines (Herpailurus yagouaroundi,
Leopardus pardalis, Leopardus tigrinus, Leopardus wiedii, Panthera onca, Puma
252 Q. Newcomer et al.

concolor). Camera trap research shows regular and widespread presence of pumas
and ocelots, and occasional presence of jaguars (Zamzow et al. 2018). Baird’s Tapir
(Tapirus bairdii), Costa Rica’s largest land mammal and IUCN Red Listed as
endangered (see Table 10.1), is also present in lower abundance. As is the case for
other taxonomic groups in the Monteverde region, comprehensive monitoring pro-
grams to estimate populations and conservation status of mammals have not been
established in Monteverde.
By night, insectivorous, nectar-feeding, and frugivorous bats parallel the daytime
roles of birds—pollinating, dispersing seeds, and preying on invertebrates in
Monteverde’s forests (Timm and LaVal 1998; Muchhala 2003). These ecosystem
services are essential in maintaining the functional integrity of the cloud forest
ecosystem. Montverde’s bat diversity includes 58 species, which represents nearly
half of the total bat diversity in Costa Rica (Timm and LaVal 2000; Wainwright
2007). Migration outside of Monteverde’s protected areas on the Pacific Slope for
foraging and pollination make bats especially important in the process of natural
habitat restoration, as they help disperse fruits and seeds of pioneer species such as
Piperaceae and Solanaceae. More research on bat migration is needed, however, in
order to more fully understand their contribution to habitat restoration, which may in
turn promote greater bat conservation (Caughlin et al. 2007).

10.2.6 Amphibians

Historically, Monteverde was home to 60 species of amphibians, including 2 caeci-

lians, 5 salamanders, and 53 anurans, many of which are part of a “distinctive upland
assemblage rich in endemic species” (Pounds and Fogden 2000). As is the case
globally, Monteverde’s amphibians have experienced dramatic and alarming declines
over the past four decades due to habitat loss as well as the amphibian chytrid fungus,
Batrachochytrium dendrobatidis (Bd) (James et al. 2015; Whitfield et al. 2017;
Whitfield et al. 2016). In 1990, researchers found only 25 of the 50 anuran species
expected to be present in the Monteverde area; notably absent was the iconic and
endemic Golden Toad (Incilius periglenes) (Pounds and Fogden 2000).
However, Monteverde’s extensive protected areas apparently function as a refuge
and foster recovery for some species. For example, Forrer’s Leopard Frog
(Lithobates forreri) was later found (Pounds and Fogden 2000), the Green-eyed
Frog (L. vibicarius) (see Fig. 10.5) and Starrett’s tree frog (Isthmohyla tica) also
reappeared in remote areas of the Children’s Eternal Rainforest, and other species
have since been rediscovered as well (Garcia-Rodriguez et al. 2012; Whitfield et al.
2017; M. Wainwright personal communication). In the case of L. vibicarius,
research indicates that these remnant populations can survive and persist even with
Bd infection (Whitfield et al. 2017).
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 253

Fig. 10.5 The Green-eyed Frog (Lithobates vibicarius) was thought to be extinct but later
reappeared in the Children’s Eternal Rainforest. (Image Source: Mark Wainwright)

10.2.7 Soils and Microbial Communities

The taxonomy and ecology of soil fauna are largely missing from biodiversity
inventories in Monteverde, although two studies have examined the biology of
cloud forest canopy soils (Nadkarni et al. 2002; Rains et al. 2003). This could be
the next frontier of biological research in the area. The implementation of new DNA
sequencing technologies available in Costa Rica provides a significant opportunity
to study the composition of microorganisms in cloud forest soil ecosystems.
254 Q. Newcomer et al.

10.2.8 Endemism, Climate Change, and Species Decline

The Monteverde region exhibits high endemism rates in comparison to areas at

lower elevations. For example, Haber (2000a, b) reports approximately 10% of
Monteverde’s plant species to be endemic to the Tilarán mountain range. A com-
prehensive study of endemic species, invasive species, and the vulnerability of
endemic species to invasive species and changes in climate is needed for the region.
In the 50 years between 1940 and 1990, deforestation and agricultural expansion
were the primary forces creating the island-effect around Monteverde; however, the
primary current threat to Monteverde’s biodiversity is climate change. Strikingly, the
number of dry days and the number of dry days in runs—five or more consecutive
days with no precipitation, which have an impact on mist frequency in the cloud
forest—have shown a steadily increasing trend over time (Pounds et al. 2006).
Through the 1970s, there were less than 10 dry days in runs per year, with some
years having no such runs (Pounds et al. 2006); 108 total dry days were recorded in
2019, with most of the zero precipitation days during January–April (Monteverde
Institute, unpublished data). The shift of environmental variables regulating ecosys-
tem function may produce a rapid transformation in life zone distribution and species
composition across the region. Should this change happen more quickly than
ecosystems and species are able to adapt, species declines—and extinctions—will
likely result.
New disease vectors invading the Monteverde region may become a leading
cause of species declines (Pounds et al. 2006). This is perhaps most vividly
highlighted by the example of amphibian species declines and extinctions at the
hands of the chytrid fungus B. dendrobatidis, mentioned above. This phenomenon
may be exacerbated by changing climate conditions.

10.3 Biodiversity Conservation in Monteverde

“Within Costa Rica, the Monteverde Zone is atypical, with its multi-cultural population, the
large number of people within high education al levels, sharp awareness of conservation
and sustainable development, relative prosperity based on dairy farming and ecotourism,
and ability to create grassroots organizations to deal with local issues”. (Burlingame
2000: 374)

Given the Monteverde region’s biodiversity, as highlighted in Sect. 2, and in light of

past and current threats to the region’s forests, biodiversity conservation initiatives
have played a central role in maintaining the region’s biodiversity and in the
emergence of Monteverde as a world-renowned nature-based tourism destination.
The Monteverde Reserve Complex—the amalgamation of the various protected
areas described in the Introduction (see Sect. 1, Fig. 10.1)—is the accomplishment
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 255

of private citizens who established organizational structures and carried out

fundraising to acquire and maintain large swaths of land in private ownership for
conservation purposes. Initiatives from two Costa Rican NGOs, the Tropical Science
Center (TSC, owner of the Monteverde Cloud Forest Biological Preserve) and the
Monteverde Conservation League (MCL, owner of the Children’s Eternal
Rainforest) and their international fundraising counterparts grew out of the recogni-
tion that the Costa Rican government did not have the funds to effectively purchase
or steward this [Link] requiring Herculean effort to successfully carry out
fundraising for land purchase, Monteverde had a compelling story about immediate
threats to species such as the charismatic Resplendent Quetzal and the endemic
Golden Toad. Furthermore, the concept of a Children’s Eternal Rainforest—a
privately-owned protected area established by many small contributions from chil-
dren all over the world—began with Swedish elementary school students, whose
fundraising efforts subsequently spread around the globe thanks to several interna-
tional NGOs that were formed to collect funds raised by the students. The interna-
tional fundraising efforts were able to raise hundreds of thousands of dollars from
tens of thousands of small donations. In doing so, they brought further attention to
and grew the mystique of Monteverde as a special place on the planet where
biodiversity protection is paramount.
One of the great challenges of large-scale land conservation, whether public or
private, is the cost of management. Purchase is a one-time event; maintenance and
stewardship are forever. Neither the TSC nor the MCL was able to raise sufficient
funding to establish long-term endowment funds to support the staff, materials, and
other costs related to forest protection and management. They depend in part on
revenue-generating use of the reserves to support annual budgets, which requires
ongoing investment in infrastructure, and larger annual budgets for operations and
maintenance. Both organizations have experienced periods of institutional growth to
ramp up operations and generate more revenue, followed by periods of contraction
and re-organization as revenue cycles ebbed.
How these organizations managed reserve borders and encroachments has been
an important factor in the conservation successes of the Monteverde Reserve
Complex. The MCL and TSC realized early on that effective conservation was
better achieved through non-confrontational means, and, in contrast to government
park rangers, their park guards do not carry weapons (Burlingame 2000). When park
guards come across poachers and others encroaching on reserve properties, the
guards take the approach of dialog and explain conservation goals. Many of the
early forest guards were former hunters and farmers themselves, and they understood
the mindset of those living around the reserves. While this approach did not always
work in terms of stopping encroachment, it did succeed in building trust and respect
between reserve personnel and neighbors. Anecdotal conversations with park guards
reveal that while poaching continues to be a problem in the private reserves, there is
much less hunting today than in the past; some hunters also simply move to
neighboring properties (including state owned reserves) in order to respect the
boundaries of the Monteverde Cloud Forest Biological Preserve and Children’s
Eternal Rainforest (H Chacón personal communication). Some neighboring
256 Q. Newcomer et al.

landowners have seen the economic opportunities associated with growing ecotour-
ism, particularly from birdwatching (e.g., the 83 ha Curi Cancha Reserve), and have
adopted conservation measures, leaving remnant forest trees on their properties and
allowing pastures to return to forest.

10.3.1 The Monteverde Reserve Complex

[Link] Tropical Science Center, the Monteverde Cloud Forest
Biological Preserve, and Bosqueterno, S.A.

The Monteverde Cloud Forest Biological Preserve (MCFBP), owned and operated
by the Tropical Science Center, was founded in 1972. Through strategic land
purchases, the MCFBP has grown from its initial 328 ha to 4,125 ha in 2019
(Burlingame 2000; C Hernandez personal communication).
Biologists who came to Monteverde to study the cloud forest were instrumental in
fundraising and land purchase efforts, and helped generate broad interest about
conservation of this unique ecosystem. In particular, George Powell played a crucial
role in raising awareness—and funds—on an international scale in support of
conservation of the Golden Toad, Resplendent Quetzal, Bare-necked Umbrellabird,
and wild feline species. Powell was also the person who first approached the TSC,
Costa Rica’s first nonprofit conservation organization, about taking ownership of the
newly acquired lands (Burlingame 2000).
As part of an effort to expand the MCFBP, in 1974 the TSC secured a 90-year
lease on 554 ha of land belonging to Bosqueterno S.A., a Costa Rican corporation
established by the Quaker community.3 When Quakers originally settled in
Monteverde in the 1950s, they set aside this land—high on the mountain, shrouded
in dense cloud forest, and unsuitable for farming—in order to protect
the community’s principal water source (Burlingame 2000). A small portion of the
Bosqueterno property forms part of the main trail system open to visitors in the
MCFBP.
Visitation at the MCFBP has grown from 471 visitors in 1973–1974 (Burlingame
2000) to more than 100,000 visitors in 2019 (Y Méndez personal communication).
As the Monteverde area’s most visited reserve, the MCFBP has established a
maximum limit of 250 persons at any given time on its 13 km of trails; the maximum
limit for daily visitation is 450 people. Of the MCFBP’s 4,125 ha, slightly more than
80 ha (or about 2% of the total reserve area) are open to the public ([Link]
[Link]/contenido/our-protected-areas).
The MCFBP’s environmental education program, founded in 1992, brings local
students to the cloud forest for talks, workshops, and hikes. Environmental educators

3
The 554 ha owned by Bosqueterno, S. A. and managed by the Tropical Science Center are not
included in the 4,215 ha owned by the Tropical Science Center.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 257

also visit local classrooms and participate in environmental education activities in

conjunction with the Monteverde Environmental Education Commission. The
MCFBP’s research and monitoring program is important for carrying out censuses
of birds, amphibians, mammals, and other groups, as well as monitoring Resplen-
dent Quetzals and other species of special interest. Most of the MCFBP’s operations
and programs—including environmental education, research and monitoring, forest
protection, and maintenance—are funded via tourist visitation to the reserve, interest
on investments, and participation in the Costa Rican government’s Payments for
Environmental Services (PES) program.4
Also located in the Monteverde area, the TSC’s 250 ha San Luis Biological
Reserve protects a critical tract of habitat connecting the cloud forest to the lower dry
forests on the Pacific slope within the Bellbird Biological Corridor. This property is
not open for tourism and is primarily used for conservation and research.

[Link] Monteverde Conservation League and the Children’s Eternal

Rainforest

One of the first objectives of the Monteverde Conservation League, a Costa Rican
nonprofit organization founded in 1986 by a group of Monteverde residents, was to
raise funds for the purchase and preservation of forest that would otherwise be lost or
severely degraded by agricultural expansion. The land purchased would soon
become known as the Children’s Eternal Rainforest (CER), today Costa Rica’s
largest privately-owned reserve. The 22,600-hectare forested expanse traverses
seven geopolitical districts and three provinces, bridges an elevational range of
>1,200 m, and spans the Continental Divide that separates Atlantic and Pacific
watersheds. As the centerpiece of the Monteverde Reserve Complex, the CER is a
vital nexus for natural habitats and populations.
The CER (known locally as “Bosque Eterno de los Niños”) was purchased and
protected thanks to donations from children, adults, and organizations in more than
40 countries around the globe. In addition to its key role in biodiversity conservation,
the CER also benefits Costa Ricans through the conservation of five major water-
sheds that provide a continuous supply of clean water for human consumption,
agriculture, and hydroelectric production; opportunities in the ecotourism sector; and
via innovative outreach services such as facilitating participation by neighboring
landowners in national PES programs. The MCL was also instrumental in early
reforestation efforts on private farms in the Monteverde region (see Bosques en
Fincas, Section 3.3.1, below), and has maintained an active environmental education
program since 1986. In this way, the reach of the CER extends beyond the borders of

4
Mechanisms for the Costa Rican Payment for Environmental Services (PES) program and the
National Forestry Financing Fund (FONAFIFO) were established in Costa Rica’s Forestry Law
7575 in 1996 (Asamblea Legislativa de la República de Costa Rica 1996). FONAFIFO is the
administrative entity for the national program. See Sect. 4 for further discussion of PES as a tool for
conservation in Monteverde.
258 Q. Newcomer et al.

the Monteverde Reserve Complex, creating the human and natural connections
needed to ensure the welfare of human and nonhuman communities in the future.
As is the case for many conservation organizations, the quest for economic
stability over time has been one of the MCL’s main challenges. Currently, about
half of the MCL’s total gross income comes from PES, including participation in
Costa Rica’s national program with FONAFIFO, and contracts with two private
hydroelectric companies in recognition of the ecosystem services (abundant, clean
water throughout the year) provided by thousands of hectares of protected forest
upstream of their dams. Unfortunately, the cooperative spirit that led to the original
signing of the private PES contracts did not last, and the MCL has had to fight to
defend both agreements—including one that is still in the appeals process in the
Costa Rican court system.
The MCL also receives important income from visitation at its field stations and
trails. Though most of the CER is not open to the public, there are four visitation
centers, including the Bajo del Tigre Reserve, which receives the most visitation
(approximately 9,000 visitors in 2019). Two field stations, San Gerardo and Pocosol,
offer rustic lodging, meals and trails to student groups and ecotourism visitors. Finca
Steller, on the eastern border of the CER, is home to the MCL’s environmental
education program and native tree nursery, and also has a small trail system.
Donations and grants continue to provide crucial funding as well, although the
increase in environmental crises on a global scale—NOAA (the United States
Department of Commerce’s National Oceanic and Atmospheric Administration;
(2020)) reports that droughts, flooding, freezing, severe storms, tropical cyclones,
wildfires, and winter storms caused $531.7 billion in damage between 2015 and
20195—combined with changes in US tax law increasing the threshold for deduc-
tions for charitable giving (many of the CER’s donors are US-based), have brought
new challenges to the nonprofit financing landscape.

[Link] Santa Elena Cloud Forest Reserve

The Santa Elena Reserve (SER) is another example of a unique idea for biodiversity
conservation piloted in Monteverde. The Santa Elena Technical-Professional High
School (CTPSE, Colegio Técnico Profesional de Santa Elena)—the public high
school which serves students from communities throughout the Monteverde
region—provides technical skill training in areas relevant to the local economy. As
tourism’s influence grew to a substantial portion of the local economy, preparing
local youth for careers in the tourism sector became a priority. The CTPSE had

5
These NOAA (2020) figures do not account for other global environmental crises, including
habitat loss due to large-scale Amazonian fires, the current Australian wildfires, world-wide coral
reef decline, impacts of increasing plastic contamination in the world’s oceans, melting glaciers and
ice caps, and localized disruptions due to coastal zone flooding. Dollar values are based on losses
which would not have been incurred had the event not taken place and include both insured and
non-insured losses.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 259

previously signed a long-term lease with the Costa Rican government for a 310-ha
farm; however, using this farm for agricultural purposes did not prove successful
(Burlingame 2000). In 1992, the Santa Elena Reserve was established as a training
ground for ecotourism and gave students the opportunity to learn about park
management and natural history guiding. The SER borders MCL’s CER (see
Fig. 10.1) and includes about 80% primary forest on the upper Atlantic slope
(Burlingame 2000). In 2019, the SER received 51,164 visitors (Y Arias personal
communication). Given its more remote location from the towns of Monteverde and
Santa Elena, the SER receives fewer visitors than the MCFBP; however, this has
made the SER an attractive alternative for birdwatchers and tourists who prefer less
crowded environments. For researchers and ecology students, the SER also provides
a good site for comparative research and study with respect to other reserves in
the area.

[Link] Alberto Manuel Brenes Biological Reserve, Arenal Volcano

National Park, and Other Surrounding Protected Areas

The Alberto Manuel Brenes Biological Reserve (RBAMB, Reserva Biológica

Alberto Manuel Brenes), located contiguous to the CER and the north-eastern
edge of the Bellbird Biological Corridor, spans 7,800 ha and is managed by the
University of Costa Rica. Although very remote in terms of access via poorly-
maintained roads from the nearest town, the RBAMB includes basic lab and
dormitory space to support field research and field study courses.
The Arenal Volcano National Park (PNVA, Parque Nacional Volcán Arenal)
spans 29,692 ha and includes its namesake, the Arenal Volcano, which was one of
the hemisphere’s most active volcanoes from the late 1960s through the first decade
of the 2000s. Established in 1994, PNVA is an important buffer region on the
Atlantic slope of the Monteverde Reserve Complex, extending forest connectivity
to the north and east through lower elevations and protecting critical habitat and
connectivity for migratory birds and large mammals requiring expansive ranges.
The Zona Protectora Arenal Monteverde6 (Arenal-Monteverde Protected Zone)
includes the MCFBP, CER, SER, Bosqueterno, RBAMB, PNVA, and a number of
private farms, as well as a sizeable property to the north of Arenal Volcano National
Park belonging to ICE, Costa Rica’s electric utility (see Fig. 10.2). These are
primarily located on the region’s Atlantic slope. All of these additional protected
areas have helped to expand the region’s biodiversity island across the landscape.

6
In Costa Rica, Zona Protectora is one of seven management designations for “protected wildlife
areas.” These also include National Parks, Forest Reserves, Biological Reserves, National Wildlife
Refuges, Wetlands, and National Monuments. These are defined in Chapter VII, Article 32 of the
Ley Orgánica del Ambiente, No. 7554, of October 4, 1995.
260 Q. Newcomer et al.

10.3.2 Sustainable Agriculture and Agroecology

Agricultural production in the earlier years of Monteverde’s settlement was largely

focused on subsistence because of the region’s remote location and poor road access
to external markets. Partly due to lack of access and partly because of personal
values, many local farmers did not use agrochemicals, thereby contributing to more
sustainable small-scale production (Griffith et al. 2000); nonetheless, agricultural
expansion was a leading cause of deforestation in the Monteverde region through the
1970s.
Monteverde’s Quaker settlers developed the nucleus of a large dairy production
region, the “Monteverde milkshed,” supplying the Productores de Monteverde
S.A. dairy which, at its peak, grew into a large-scale producer of cheeses distributed
throughout Central America (Griffith et al. 2000; Burlingame 2000). Due to low
prices for milk, some of the pastures previously dedicated to dairy and mixed dairy-
beef production have been abandoned and have returned to secondary growth forests
(Stuckey et al. 2014).
Agriculture continues to be an important component of the local economy, but it
does not currently contribute to continued deforestation. Instead, agriculture has
moved toward a more sustainable model, particularly at higher elevations (Stuckey
et al. 2014). Lack of support from extension agents is an ongoing challenge for local
producers who have adopted organic and other sustainable farming methodologies
(O Salazar personal communication).
While coffee has long played an important role in the local agricultural sector, the
focus of Monteverde’s coffee sector has shifted toward organic/sustainable produc-
tion and is marketed primarily toward tourism (Stuckey et al. 2014). This includes
direct sales to tourists as well as agroecology tours of sustainable farms, which
diversifies the income stream for the farmers and their families. Integrated
sustainably-produced coffee and agrotourism projects represent a rapidly-growing
segment of Monteverde’s agricultural economy.
A recent study by the TSC found a growing local market for sustainably produced
agricultural products, but insufficient organization to connect producers to consumers
and promote this emerging market for locally and sustainably produced foods (Tropical
Science Center 2018). Consequently, TSC has initiated a new program,
Encadenamientos Productivos (or Agricultural Products Network), to link local
farmers with restaurants and other consumers, particularly within the Bellbird Biolog-
ical Corridor. Some local businesses have independently developed similar initiatives.
For example, the University of Georgia Costa Rica Campus (UGA CR) implemented
the policy of purchasing the majority of food from within a 400 km radius of the
campus, with an emphasis on on-site organic production and purchasing from local
farmers in the Monteverde area and in the nearby lowlands who also followed
sustainable agricultural practices. The Hotel Belmar and several other Monteverde-
based hotels and restaurants have established on-site food production and local food
purchasing components to their operations. Weekly farmer’s markets in Santa Elena
and Guacimal offer outlets for connecting local farmers with local consumers.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 261

10.3.3 Biological Corridors

“The main focus of conservation biology at Monteverde since 2000 has been on the role of
landscape features in preserving biodiversity, particularly connectedness between habitats
at different spatial scales”. (Wheelwright 2014: 2).

This section describes some of the ways in which Monteverde has integrated
landscape-level conservation into the surrounding communities, functionally
extending the limits of the Monteverde biodiversity island beyond the physical
limits of the core protected areas.

[Link] Bosques en Fincas

In the mid-1990s, the MCL administered the program Bosques en Fincas (or Forests
in Farms), which encouraged local farmers to maintain existing forest fragments and
connect them by planting windbreaks. Dairy farmers realized the effectiveness of
windbreaks at maintaining pasture productivity during the extremely windy season.
Many of these windbreaks are now 20–30 years old. The Bosques en Fincas program
set the stage for expanding forest connectivity across the privately-owned landscape
surrounding the Monteverde Reserve Complex and spawned research suggesting the
critical importance of remnant trees for biodiversity conservation (Guindon 1996;
Harvey and Haber 1999). Brownson et al. (2019) describe Bosques en Fincas as an
example of a successful local PES program, in which farmers were engaged by the
MCL’s outreach team and given the species of trees they were interested in planting
on their farms. This program triggered a cultural shift toward reforestation in the
region (K Brownson personal communication), which set the stage for other refor-
estation programs focusing on native species reforestation (see Sect. 3.3.4, below).

[Link] Enlace Verde

In the mid-1990s, the Monteverde Institute promoted a corridor program, Enlace

Verde, aimed at protecting existing forests on individual farms within the town of
Monteverde using conservation easements. This program intended to connect the
Children’s Eternal Rainforest with the Bajo del Tigre Reserve, both owned by the
MCL. At the time, the San José, Costa Rica-based Environmental and Natural
Resource Law Center (CEDARENA, Centro de Derecho Ambiental y de los
Recursos Naturales) was introducing the concept of conservation easements within
Costa Rica and partnered with the Monteverde Institute to support the Enlace Verde
program. Three easements were signed in 1998, including unique reciprocal ease-
ment contracts. Despite multiple charrette exercises and individual meetings with
landowners, including preparation of drafts of easement contracts and accompanying
maps, the overall program did not expand as hoped across some fifty properties
located in six neighborhood clusters (Scrimshaw et al. 2000:382).
262 Q. Newcomer et al.

[Link] Reforestation Initiatives

Following in the footsteps of the MCL’s Bosques en Fincas program, the Fundación
Conservacionista Costarricense (FCC, or Costa Rican Conservation Foundation)
was established in 2002 in response to the observed need to expand Pacific slope
habitat for the Three-wattled Bellbird and other migratory species in this range. The
FCC’s reforestation program was started in 1998 as the “Bellbird Project” with the
support of local organizations and funds provided by the British Embassy in San
José, Costa Rica (D Hamilton personal communication). Following several years of
project implementation, the formal non-profit foundation was established.
The FCC’s tree nursery produces native forest species that provided free of
charge to local landowners. The FCC has studied multiple areas where trees have
been planted to understand and develop best practices that yield the greatest benefit-
to-cost related to minimizing mortality rates and maximizing growth rates during
establishment (D Hamilton personal communication). The FCC does not use formal
agreements with recipient landowners to maintain the trees. To date, FCC has
planted over 250,000 trees of more than 143 species and 42 families (D Hamilton
personal communication).
The former University of Georgia Costa Rica Campus (UGA CR)7 reforestation
program was established in 2008 to offset University of Georgia students’ emissions
related to study abroad in Costa Rica, and ran for 11 years until 2019. In 2010 the
FCC partnered with UGA CR, providing funding to expand the tree nursery. UGA
CR’s native species nursery primarily focused on species native to the San Luis
Valley, where the campus was located. Planting was done both on-campus and on
neighboring farms in San Luis, with saplings given to local landowners. UGA CR
asked participant landowners to sign agreements which noted the numbers and
species of saplings provided and stated that landowners agreed to maintain the
saplings through establishment, however there was no enforcement. Beginning in
2011, UGA CR student groups and interns monitored planting sites to determine
mortality and measure growth rates.
Between 2008 and 2016, the UGA CR program planted more than 35,000 trees
representing over 90 species from the local forests (UGA CR unpublished data). For
carbon offset estimates, mortality was estimated at 25%; however, at several sites,
monitoring revealed high mortality due to cattle encroachment on one property and
development of a housing lot on another. Other sites with effective protection for
saplings during establishment had less than 5% mortality (UGA CR
unpublished data).
Reforestation projects with non-binding agreements for landowner care of the
trees planted are only as effective as the quality of the management provided until
tree establishment. In their study of formal and informal payment for ecosystem
services programs (see Sects. 3.4 and 4 for further discussion), Brownson et al.

7
The University of Georgia sold this property in 2019. Long-term research and reforestation
programs were not maintained by the current owner.
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 263

(2019) found that the more informal local reforestation programs in the Monteverde
region promoted greater tree species diversity and more effectively engaged with
less economically prosperous landowners than the formal national ecosystem ser-
vices program.

[Link] Corredor Biológico Pájaro Campana (Bellbird Biological

Corridor)

At the same time the Monteverde Institute was working on Enlace Verde, the TSC
proposed the concept of a biological corridor following the Guacimal River water-
shed from Monteverde down the Pacific slope to the mangrove forests along the Gulf
of Nicoya. As a step toward implementing this project, TSC acquired a 240 ha
(593 acre) farm in the lower San Luis Valley, securing the largest remaining Pacific
slope forest patch in the upper elevation of this watershed (Burlingame 2000).
Further expansion of the corridor was not immediately put into place; yet the idea
for a corridor initiative, like the seed of an Ocotea monteverdensis tree buried in the
forest floor, was firmly planted and waiting for the right conditions to germinate.
The corridor initiative was revived in 2006, and in 2007 a group of local
conservation and education organizations established the Local Advisory Commit-
tee for the Corredor Biológico Pájaro Campana (CBPC, or Bellbird Biological
Corridor), including representatives from TSC, MCL, UGA CR, FCC, and the
Monteverde Institute. The CBPC has been successful in large part due to common
interests, shared vision, cooperative efforts, and commitment of time and funding by
the organizations involved. The Local Advisory Committee successfully addressed
differences in opinion, listened to and incorporated concerns from a wide body of
stakeholder groups, and implemented the initiative in a professional manner.
The CBPC spans 88,738 hectares within three adjacent watersheds which flow
from the Monteverde cloud forest to the Gulf of Nicoya—the Aranjuez, Guacimal,
and Lagarto River watersheds (see Fig. 10.2). Eleven of Costa Rica’s 13 Holdridge
Life Zones (Holdridge 1967) are represented within the corridor, and approximately
50% of Costa Rica’s terrestrial vertebrate species are found within this region
(Welch et al. 2011).
The CBPC was recognized by the Costa Rican government as part of the formal
national network of biological corridors connecting Costa Rica’s larger parks and
reserves. The CBPC’s Local Advisory Committee secured funding from the UNDP
Small Grants Program to develop a strategic plan for the corridor (Welch et al. 2011).
This strategic plan defined key areas of common interest, and sub-committees then
developed sector-specific work plans from these broadly defined goals (J Welch
personal communication). Over the course of the next decade, the Local Advisory
Committee was able to secure additional grant funding from the UNDP Small Grants
Program to develop promotional and educational materials, hold workshops in
communities throughout the corridor region, and establish multiple community-
based sub-committees throughout the three watersheds. The Local Advisory Com-
mittee was sensitive to the appearance of conservation initiatives being driven by the
264 Q. Newcomer et al.

organizations—both literally and figuratively—“at the top of the mountain” (and in

the communities which received the most direct economic benefits from ecotour-
ism), and worked to build broad common interest in the concept of maintaining and
expanding habitat connectivity. Local concerns were incorporated into the strategic
plan and, subsequently, continued UNDP Small Grant Program funding helped to
establish sub-regional committees to generate locally driven initiatives (J Welch
personal communication).
A growing number of PhD dissertations, master’s theses, and undergraduate
thesis projects have been completed related to the Bellbird Corridor (e.g., Brownson
2019; Camacho Céspedes 2019; Padgett-Vasquez 2019; O’Halloran et al. 2018;
Powlen 2018; Allen 2016; Chinchilla Ramos 2015; Piedrahíta López 2013). The
University of Georgia and Lynchburg College carried out a long-term water quality
monitoring program in the region for 8 years, documenting the macroinvertebrate
communities in the three watersheds (T Shahady personal communication).
The CBPC also is the focus of current land purchase priorities for the Monteverde
Conservation League, whose goal is to increase ecological connectivity between the
Children’s Eternal Rainforest and forested areas at lower elevations on the Pacific
slope. The MCL has purchased more than 100 hectares in the CBPC since 2014 and
continues to raise funds for land purchase and conservation in the area.

10.3.4 Payment for Environmental Services (PES)

Costa Rica’s national Payment for Environmental Services (PES, or Pago por Servicios
Ambientales) program administered by the National Forestry Financing Fund
(FONAFIFO, Fondo Nacional de Financiamiento Forestal) has been extensively
studied by academics and practitioners alike. The early years of Costa Rica’s PES
program coincided with a noticeable trend of afforestation across the country, follow-
ing decades where Costa Rica experienced some of the highest deforestation rates in the
world (FONAFIFO 2001). The successes of PES in helping to protect privately-owned
forest land in Costa Rica is highlighted by figures such as over 10% of the country
having been protected via PES forest conservation contracts (Ringhofer et al. 2013).
Within the CBPC, 11 Holdridge life zones were represented by PES contracts issued
between 2008 and 2012, with most of these PES contracts located in areas not
represented by the Monteverde Reserve Complex (Padgett-Vasquez 2019). During
this time period, 51 properties throughout the CPBC region were inscribed by
FONAFIFO in the PES program (Padgett-Vasquez 2019). These properties ranged in
size from 11 to 300 hectares (Padgett-Vasquez 2019). The majority of the 51 properties
were located along or in very close proximity to rivers and streams, which suggests that
the protection of these forested areas through PES contributes to maintaining the
functional integrity of the region’s ecological systems (Padgett-Vasquez 2019).
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 265

10.3.5 The Monteverde-Arenal Bioregion Initiative (MABI)

One of the characteristics that makes Monteverde unique is the number of research
scientists that have settled in the area and continued to study the region over multiple
decades. The Monteverde-Arenal Bioregion Initiative (MABI) was initiated in 2014
by several of Monteverde’s long-term research scientists and educators to coordinate
and expand research, teaching, and outreach collaborations in support of biodiversity
conservation. Given the ongoing observations and growing collective understanding
about changing climate conditions in the Monteverde region and the likely impacts
for biodiversity conservation (e.g., Pounds et al. 1999; Pounds et al. 2006; Nadkarni
and Solano 2002; Eaton et al. 2012), MABI was created to develop shared database
resources and coordinate regular conferences which serve to share information,
define research priorities for the region, develop coordinated research proposals,
and support each other’s research, teaching, and outreach. To date, funding limita-
tions have limited new region-wide collective research efforts, however the multiple
MABI conferences held during 2014–2016 served to share information and build
new relationships among the many scientists and educators working in the region
(N Nadkarni personal communication).

10.3.6 The Monteverde Commission for Resilience to Climate

Change (CORCLIMA)

The same capacity for community mobilization and organization and deeply-rooted
environmental values that led to the creation of the Monteverde Reserve Complex
and more recently the Bellbird Biological Corridor has also spawned a new initia-
tive, The Monteverde Commission for Resilience to Climate Change (CORCLIMA).
Promoting Costa Rica’s National Strategy for Climate Change, local residents and
institutions have joined forces and are coordinating across sectors and scales with the
goal of leading the Monteverde Zone to become carbon neutral and develop climate
resilience strategies for the region (Brenes et al. 2019, [Link]). While the
carbon balance benefits are global in scale, the adjustments Monteverde seeks to
achieve in meeting carbon neutral goals and implementing climate change resilience
will enhance the quality of life for local residents, help further protect habitat to
support the region’s biodiversity, and promote Monteverde’s reputation as an
authentic, sustainable tourism destination. CORCLIMA’s actions serve as a demon-
stration for other communities seeking to implement similar plans.
266 Q. Newcomer et al.

10.4 Challenges for the Future

How to sustain the local economy, improve quality of life for a growing local
population, and continue to protect the region’s natural capital while adapting to
changing climate conditions is the biggest overarching challenge facing Monteverde
today. Should the projected changes in climate impact the cloud forest habitat to the
extent that it further threatens the populations of quetzals, bellbirds and other iconic
species which draw many tourists to the region, what impact will that have on
tourism and thus the local economy? Long-term biodiversity conservation is tied
to economic stability, and economic downturn can lead to increased pressure on
forests and forest resources. How can Monteverde diversify the local economy and
also find alternative funding mechanisms for biodiversity conservation to balance
fluctuations in tourism?
Addressing the challenges requires both top-down and continued grass-roots
leadership. Coordinated strategic visioning with an alignment of national policies
integrating environmental conservation, agricultural production, and continued
development for sustainable tourism, sustainable manufacturing, and service sectors
are also needed. Costa Rica’s national law recognizing biological corridors as
regions where such integrated conservation and development takes place provides
a broad legal framework and national administrative structure to support local
initiatives. Yet as Padgett-Vasquez (2019) notes, one of the challenges in
landscape-level conservation is to establish functional wildlife corridors within
these broadly declared conservation regions spanning privately-owned property.
The accomplishments of the Bellbird Biological Corridor reflect how local leader-
ship capacity and coordination is critical to implement such broad national conser-
vation plans. The challenge of funding staff and administrative overhead to support
the ongoing coordination of the CBPC reflects the common economic struggle faced
by grass-roots conservation initiatives around the world. Volunteer boards com-
prised of dedicated representatives of local NGOs can only carry conservation
initiatives so far, particularly in small communities.
Planning and zoning is of critical importance as populations grow and small rural
towns like Santa Elena undergo rapid urbanization. Of the 82 cantons in Costa Rica,
40 have developed zoning plans, 21 have comprehensive environmental planning
regulations, and only four have completed studies of hydrological vulnerability and
established guidelines regarding water resource protection (INVU 2019). While the
springs emanating from the protected reserves supply clean water for the region,
wastewater management and contamination of local waterways continues to threaten
public health as well as Monteverde’s aquatic diversity outside of the protected
areas.
A growing body of literature through the first two decades of the twenty-first
Century describes the shortcomings of public land conservation programs to provide
sufficient habitat to address long-term biodiversity conservation concerns (Norton
2000; Chacón 2005; Mayer and Tikka 2006; Pasquini et al. 2011; Kamal et al. 2015).
In particular, there is growing recognition that connectivity across a wider landscape
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 267

is critical to provide sufficient habitat to support migratory species through an

increasingly more densely human-dominated landscape, and to support long-term
ecological resiliency and both floral and faunal adaptation to changing global climate
conditions (Zamzow et al. 2018). Yet for privately-owned projects and conservation
initiatives led by non-governmental organizations to be successful, there must be
supportive public policy in place to encourage and support conservation practices
(Pasquini et al. 2011). Private nature reserves and those owned by non-governmental
organizations have some advantages compared to public protected areas—manage-
ment flexibility, ability to make decisions more rapidly, and broader economic
opportunities (Pasquini et al. 2011). Indeed, in the case of Monteverde, initial
government designation of the region as a Zona Protectora was ineffective and led
to the local establishment of NGOs to fill the conservation gap. Private land
conservation—including NGO-owned conservation reserves—cannot exist in a
vacuum, however, and requires lock-step coordination with the public sector and
the long-term public sector commitment to establish and maintain such policy.
Comprehensive conservation policy supporting private land conservation must
address ecological, economic, and social aspects, including public education
required to embed the value and importance of biodiversity conservation as core to
the human condition. From the local to global-scale, there must be coordination and
commitments from the public sector reflecting that biodiversity conservation is a
priority beyond partisan politics.
For private land conservation programs to successfully meet conservation goals,
there is a need for thoughtfully designed, contextually-appropriate incentive pro-
grams to encourage private land conservation that address both ecosystem conser-
vation goals as well as economic productivity which is considered a right of private
land ownership (Mayer and Tikka 2006). The ability of farmers to adopt more
sustainable agricultural practices is largely influenced by broader socio-economic
trends, national-level rural development policies, and institutional capacity of agen-
cies, such as the Ministry of Agriculture, to provide technical support for organic and
other forms of sustainable agriculture (Wegner 2016). Conversations with coffee
farmers in Monteverde reveal the complete absence of Ministry of Agriculture
extension support for farmers practicing organic agriculture (O Salazar personal
communication), despite the growing international demand for certified organic
products.
As a tool in the biodiversity conservation toolbox, payment for ecosystem
services has been employed in Costa Rica since the mid-1990s. While studies of
the earlier years of the PES program found that PES was effective at increasing forest
cover within biological corridors in Costa Rica (Morse et al. 2009; Newcomer 2007),
most evidence in Costa Rica, however, points toward PES not being particularly
helpful in changing landowner decision-making, and there is mixed evidence at best
about whether PES payments broadly contribute to avoided deforestation (Allen
2018, K Allen personal communication). In the case of private landowners in the
Monteverde area, PES subsidizes land abandonment and/or protection of forests
which are not necessarily in imminent threat (K Allen personal communication) and,
in general across Costa Rica, has favored larger landowners (Zbinden and Lee 2005;
268 Q. Newcomer et al.

Newcomer 2007). Furthermore, individuals have expressed a lack of trust with the
government (Newcomer 2007; Allen and Padget-Vasquez 2017), in part driven by
restrictions placed on properties in PES during and following the termination of PES
contracts (S Padgett-Vasquez personal communication). Thus, local PES programs,
such as the reforestation programs described in Sect. 3.3.3, are effective ways to
build trust relationships with smaller local landowners, increase tree species diver-
sity across the landscape, and provision ecosystem services (Brownson et al. 2019).
Despite these noted issues, however, the national PES program is an important
mechanism that provides critical funding for ongoing habitat protection and biodi-
versity conservation of large protected areas owned by NGOs such as MCL.
Fundraising for the purchase of land for conservation purposes is challenging;
fundraising for ongoing maintenance and protection of these properties is far more
difficult. While Costa Rica’s PES program promotes biodiversity conservation as
one of the four primary ecosystem services recognized in all PES contracts, carbon
sequestration and hydrological services are the primary services connecting buyers
(e.g., those purchasing gasoline, those using water in manufacturing) to sellers (i.e.,
forest land owners). In the case of NGOs, including MCL, which own large tracts of
land that serve as regional biodiversity islands, the service provision of biodiversity
conservation is a high value-added contribution to the bundle of ecosystem services.
Given FONAFIFO’s ongoing struggles to fund the national PES program via gas
taxes, water fees, and subsidies from international agencies (including the World
Bank and the Global Environment Fund), a more coordinated, holistic national
policy strategy might incorporate a small fee for the provision of biodiversity
services as a mechanism to help support local NGOs’ land conservation initiatives,
such as the Monteverde Cloud Forest Preserve and the Children’s Eternal Rainforest.
Efforts to integrate natural capital accounting into the national accounting struc-
ture can help Costa Rica better value and manage its natural resources. Providing
such an overarching framework allows private industry, non-governmental organi-
zations (NGOs), and individuals to then pursue innovative approaches toward
achieving a more sustainable economic and ecological future for Costa Rica. Such
a holistic approach toward national development policy would provide the frame-
work for sustaining and expanding biodiversity islands such as Monteverde.

10.5 Conclusion

Monteverde reflects a unique conservation situation where the national government

of Costa Rica recognized the importance of the region for biodiversity conservation
but was not able to dedicate resources to establish a public protected area. Interna-
tional research scientists partnered with existing Costa Rican NGOs and established
new local NGOs, filling the federal conservation gap by securing funds to purchase
and manage large swaths of forest across the region. At the heart of the region’s
privately-held protected lands are 554 hectares of forest that Quaker settlers set aside
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 269

to protect their community’s water source some 20 years prior to the establishment
of larger protected areas.
Many of the international scientists studying the area’s ecology decided to make
Monteverde their home and/or have spent significant amounts of time in the region
over decades. They helped to establish local NGOs that not only protect land and
study the region’s ecology, but also help to share this knowledge with the local
community and teach local people about what they are studying. Over time, they
have created a vast body of knowledge about Monteverde’s biodiversity, some of
which we have highlighted in Sect. 2 as representative of the Monteverde region. In
turn, many local residents have subsequently committed their professional careers to
support conservation initiatives, working as naturalist guides, park guards, research
technicians, administrators, and educators. Together, this blended community
established local schools that incorporate experiential learning and environmental
stewardship into their curriculum, spreading conservation values among the younger
generations. As described, the NGOs translate knowledge into outreach programs
supporting local farmers and broader community interests, all the while maintaining
a core mission of habitat protection. The local NGOs capitalized on the increased
popularity of Costa Rica as an ecotourism destination and the region’s charismatic
species and were able to move more nimbly than many of the public parks, setting up
infrastructure, administration, and support services to promote nature-based tourism.
This grew the local economy and increased general prosperity in the region. As
described in previous sections, the diversity of different types of initiatives—uni-
versity partnerships, sustainable tourism, international fundraising campaigns, pur-
chasing policies for sustainable products, carbon offsets, farmer and landholder
outreach, technical school partnerships, etc.—have formed a patchwork quilt of
biodiversity conservation that has endured for nearly 50 years. As some pieces of
this quilt fray, the Monteverde community has remained dedicated to the long-term
biodiversity conservation goals and has developed new initiatives to fill the gaps.
From the initial successes of establishing the MVCFBP, the CER, and the SER, the
scale of conservation initiatives has expanded from bounded protected areas focused
on Monteverde’s cloud forests to landscape-level protection spanning both Atlantic
and Pacific slopes.
Changing climate conditions now make Monteverde an interesting place to study
the impacts of these changes on the cloud forest ecosystem. How this biodiversity
island will evolve, how biological corridors will support species migration, and how
climate impacts and projected resource constraints will affect the region’s socioeco-
nomic conditions and how that, in turn, impacts biodiversity conservation remains to
be seen. If there is a big-picture takeaway message from the case study of
Monteverde, it is that biodiversity conservation—through a portfolio of private
land conservation initiatives, public parks and reserves, and public-private partner-
ships—is a continuously evolving process made ever more challenging by changing
climate conditions, yet made possible and successful by highly engaged, coordi-
nated, cooperative private-sector, public-sector, and NGO-led initiatives.

Acknowledgements We are grateful to our many colleagues in Monteverde for the knowledge and
insights they have shared with us through the years living and working in Monteverde. In particular,
270 Q. Newcomer et al.

thanks to Orlando Calvo, Alvaro Cubero, Bill Haber, Deb Hamilton, Russ Kumai, Jack Longino,
Yoryineth Méndez, Nalini Nadkarni, Alan Pounds, Yúber Rodríguez, Katy VanDusen, Mark
Wainwright and Justin Welch for their contributions to this chapter.

References

Allen K (2016) Conservation planning in a global Era: the role of market-based mechanisms in
promoting sustainable development in Costa Rica. PhD dissertation, The University of Georgia,
Athens, GA
Allen K (2018) Why exchange values are not environmental values: explaining the problem with
neoliberal conservation. Conserv Soc 16(3):243–256
Allen TFH, Hoekstra TW (1992) Toward a unified ecology. Columbia University Press, New York
Allen KE, Padgett-Vasquez S (2017) Forest cover, development, and sustainability in Costa Rica:
can one policy fit all? Land Use Policy 67:212–221. [Link]
05.008
Americas Regional Workshop (Conservation & Sustainable Management of Trees, Costa Rica,
November 1996) (1998) Oreomunnea pterocarpan. The IUCN Red List of Threatened Species
1998: e.T30663A9570512. [Link]
[Link]. Downloaded on 24 January 2020
Asamblea Legislativa de la República de Costa Rica (1996) No 7575 Ley Forestal. Alcance No 21 a
La Gaceta No 72, La Uruca, San José, Costa Rica, 16 April 1996, 8 pp
Atwood JT, Dressler RL (1995) Another new species of Stellilabium from Monteverde, Costa Rica.
Selbyana 16:239–241
BirdLife International (2016a) Ara ambiguous. The IUCN Red List of Threatened Species 2016: e.
T22685553A93079606. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2016b) Buteogallus solitarius. The IUCN Red List of Threatened Species
2016: e.T22695849A93530532. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2016c) Cephalopterus glabricollis. The IUCN Red List of Threatened
Species 2016: e.T22700933A93805184. [Link]
[Link]. Downloaded on 19 January 2020
BirdLife International (2016d) Crax rubra. The IUCN Red List of Threatened Species 2016: e.
T22678521A92776389. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2016e) Electron carinatum. The IUCN Red List of Threatened Species 2016:
e.T22682989A92971734. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2016f) Patagioenas subvinacea. The IUCN Red List of Threatened Species
2016: e.T22690328A93269904. [Link]
[Link]. Downloaded on 27 January 2020
BirdLife International (2016g) Pharomachrus mocinno. The IUCN Red List of Threatened Species
2016: e.T22682727A92958465. [Link]
[Link]. Downloaded on 19 January 2020
BirdLife International (2016h) Procnias tricarunculatus. The IUCN Red List of Threatened Species
2016: e.T22700946A93806341. [Link]
[Link]. Downloaded on 19 January 2020
BirdLife International (2016i) Sclerurus albigularis. The IUCN Red List of Threatened Species
2016: e.T22702968A93897178. [Link]
[Link]. Downloaded on 25 January 2020
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 271

BirdLife International (2016j) Spizaetus ornatus. The IUCN Red List of Threatened Species 2016:
e.T22696197A93548774. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2017a) Contopus cooperi (amended version of 2016 assessment). The IUCN
Red List of Threatened Species 2017: e.T22699787A110734937. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2017b) Grallaricula flavirostris (amended version of 2016 assessment). The
IUCN Red List of Threatened Species 2017: e.T22703362A110974550. [Link]
2305/[Link]. Downloaded on 25 January 2020
BirdLife International (2017c) Hylocichla mustelina (amended version of 2016 assessment). The
IUCN Red List of Threatened Species 2017: e.T22708670A111170926. [Link]
2305/[Link]. Downloaded on 25 January 2020
BirdLife International (2017d) Morphnus guianensis. The IUCN Red List of Threatened Species
2017: e.T22695991A118209977. [Link]
[Link]. Downloaded on 27 January 2020
BirdLife International (2017e) Tinamus major (amended version of 2016 assessment). The IUCN
Red List of Threatened Species 2017: e.T22678148A110915916. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2018a) Antrostomus carolinensis. The IUCN Red List of Threatened
Species 2018: e.T22689778A131764040. [Link]
[Link]. Downloaded on 27 January 2020
BirdLife International (2018b) Bangsia arcaei. The IUCN Red List of Threatened Species 2018: e.
T22722558A132016322. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2018c) Touit costaricensis. The IUCN Red List of Threatened Species 2018:
e . T 2 2 6 8 6 0 1 7 A 1 3 1 5 1 7 2 7 4 . h t t p s : / / d o i . o rg / 1 0 . 2 3 0 5 / I U C N . U K . 2 0 1 8 - 2 . R L T S.
[Link]. Downloaded on 25 January 2020
BirdLife International (2018d) Vermivora chrysoptera. The IUCN Red List of Threatened Species
2018: e.T22721618A132145282. [Link]
[Link]. Downloaded on 25 January 2020
BirdLife International (2019a) Setophaga cerulea. The IUCN Red List of Threatened Species 2019:
e . T 2 2 7 2 1 7 4 0 A 1 5 3 6 9 1 3 2 0 . h t t p s : / / d o i . o rg / 1 0 . 2 3 0 5 / I U C N . U K . 2 0 1 9 - 3 . R L T S.
[Link]. Downloaded on 25 January 2020
BirdLife International (2019b) Sturnella magna (amended version of 2018 assessment). The IUCN
Red List of Threatened Species 2019: e.T22735434A155622113. [Link]
[Link]. Downloaded on 25 January 2020
Bolaños F, Chaves G, Wake D, Savage J (2008a) Oedipina poelzi. The IUCN Red List of
Threatened Species 2008: e.T59321A11916146. [Link]
[Link]. Downloaded on 19 January 2020
Bolaños F, Wake D, Savage J (2008b) Nototriton picadoi. The IUCN Red List of Threatened
Species 2008: e.T59299A11902609. [Link]
[Link]. Downloaded on 20 January 2020
Brenes J, VanDusen K, Welch J (2019) Pathways toward Climate Change Resilience in
Monteverde, Costa Rica. Report prepared for the Monteverde Community Fund, Monteverde,
Costa Rica. [Link]
[Link]
Brownson K (2019) From incentives to impact: an evaluation of the socioecological impacts of
payments for ecosystem services. PhD dissertation, The University of Georgia, Athens, GA
Brownson K, Guinessey E, Carranza M, Esquivel M, Hesselbach H, Madrid Ramirez L, Villa L
(2019) Community-Based Payments for Ecosystem Services (CB-PES): implications of com-
munity involvement for program outcomes. Ecosyst Serv 39:Article 100974. [Link]
1016/[Link].2019.100974
272 Q. Newcomer et al.

Burlingame L (2000) Conservation in the Monteverde zone: contributions of conservation

organizations. In: Nadkarni NM, Wheelwright NT (eds) Monteverde: ecology and conservation
of a tropical cloud Forest. Oxford University Press, New York, pp 351–388
Camacho Céspedes F (2019) Biofertilización orgánica de almácigos de café (Coffea arabica L.) con
compost producido a partir de residuos biomásicos locales, microorganismos de montaña y
lodos digeridos de biodigestor en la región de Monteverde, Costa Rica. PhD dissertation,
Universidad Estatal a Distancia, Programa DOCINADE, San José, Costa Rica
Caughlin TT, Ganesh T, Lowman MD (2007) COS 159-8: the role of frugivorous birds and giant
fruit bats as agents for reforestation in South India. Ecological Society of America/Society of
Restoration Ecology Joint Meeting, August 5–August 10, 2007, San Jose, California. https://
[Link]/eco/2007/techprogram/[Link]
Chacón CM (2005) Fostering conservation of key priority sites and rural development in Central
America: the role of private protected areas. Parks Prot Areas Program Priv Protected Areas
15(2):39–47
Chinchilla Ramos R (2015) Conservación y manejo integral a través del análisis del uso de la tierra
y la fragmentación boscosa en el Corredor Biológico Pájaro Campana, Pacífico Central,
Puntarenas. Thesis for Licenciatura, University of Costa Rica, San José, Costa Rica
Clark KL, Lawton RO, Butler PR (2014) The physical environment – update 2014. In: Nadkarni
NM, Wheelwright NT (eds) Monteverde: ecology and conservation of a tropical cloud forest –
2014 updated chapters. Oxford University Press, New York
Cuarón AD, Morales A, Shedden A, Rodriguez-Luna E, de Grammont PC, Cortés-Ortiz L (2008)
Ateles geoffroyi. The IUCN Red List of Threatened Species 2008: e.T2279A9387270. https://
[Link]/10.2305/[Link]. Downloaded on 19 January 2020
de Oliveira T, Paviolo A, Schipper J, Bianchi R, Payan E, Carvajal SV (2015) Leopardus wiedii.
The IUCN Red List of Threatened Species 2015: e.T11511A50654216. [Link]
[Link]. Downloaded on 19 January 2020
Deliso E (2008) Climate Change and the Hummingbirds of the Monteverde Cloud Forest, Costa
Rica. Centro Científico Tropical 1 ed. San José, Costa Rica 42 pp:22 X 28 cms
Dyer V (1979) List of plants collected. Unpublished report, Tropical Science Center, San José,
Costa Rica
Eaton WD, Roed M, Chassot O, Barry D (2012) Differences in soil moisture, nutrients and the
microbial community between forests on the upper Pacific and Caribbean slopes at Monteverde,
Cordillera de Tilaran: implications for responses to climate change. Trop Ecol 53(2):235–240
Feinsinger P (1977) Notes on the hummingbirds of Monteverde, Cordillera de Tilarán, Costa Rica.
Wilson Bull 89(1):159–164
Fogden MP (2000) Appendix 9: birds of the Monteverde area. In: Nadkarni NM, Wheelwright NT
(eds) Monteverde: ecology and conservation of a tropical cloud Forest. Oxford University Press,
New York, pp 541–552
FONAFIFO (2001) Reactivación de la Reforestación Comercial en Costa Rica. Proyecto
REFORESTA. Propuesta de Asistencia Técnica y Financiera. MINAE-FONAFIFO. San José,
Costa Rica
García M, Jordan C, O’Farril G, Poot C, Meyer N, Estrada N, Leonardo R, Naranjo E, Simons Á,
Herrera A, Urgilés C, Schank C, Boshoff L, Ruiz-Galeano M (2016) Tapirus bairdii. The IUCN
Red List of Threatened Species 2016: e.T21471A45173340. [Link]
[Link]. Downloaded on 19 January 2020
García-Rodríguez A, Chaves G, Benavides-Varela C, Puschendorf R (2012) Where are the survi-
vors? Tracking relictual populations of endangered frogs in Costa Rica. Divers Distrib 18:204–
212. [Link]
Griffith K, Peck DC, Stuckey J (2000) Agriculture in Monteverde: moving toward sustainability. In:
Nadkarni NM, Wheelwright NT (eds) Monteverde: ecology and conservation of a tropical cloud
Forest. Oxford University Press, New York, pp 389–417
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 273

Guindon CF (1996) The importance of Forest fragments to the maintenance of regional biodiversity
in Costa Rica. In: Schelhas J, Greenberg RS (eds) Forest patches in tropical landscapes. Island
Press, Washington, DC, pp 168–186
Haber WA (1991) Lista provisional de las plantas de Monteverde, Costa Rica. Brenesia 34:63–120
Haber WA (1993) Seasonal migration of monarchs and other butterflies in Costa Rica. In: Malcolm
SB, Zalucki M (eds) Biology and conservation of the monarch butterfly. Los Angeles County
Museum of Natural History, Los Angeles, pp 201–207
Haber WA (2000a) Plants and vegetation. In: Nadkarni NM, Wheelwright NT (eds) Monteverde:
ecology and conservation of a tropical cloud forest. Oxford University Press, New York, pp
39–94
Haber WA (2000b) Appendix 1: vascular plants of Monteverde. In: Nadkarni NM, Wheelwright
NT (eds) Monteverde: ecology and conservation of a tropical cloud forest. Oxford University
Press, New York, pp 457–518
Hamilton D, Molina V, Powell G (2003) El Estatus del Pájaro Campana (Procnias tricarunculata):
Un Ave en Peligro de Extinción. Zeledonia 7:15–24
Hartshorn GS (1983) Plants. In: Janzen DH (ed) Costa Rican natural history. University of Chicago
Press, Chicago, pp 118–157
Harvey CA, Haber WA (1999) Remnant trees and the conservation of biodiversity in Costa Rican
pastures. Agrofor Syst 44:37–68. [Link]
Holdridge LR (1967) Life zone ecology, Rev. edn. Tropical Science Center, San Jose
Instituto Nacional de Vivienda y Urbanismo (INVU) (2019) Planes Reguladores. [Link]
[Link]/planes-reguladores
IUCN SSC Amphibian Specialist Group and NatureServe (2013a) Isthmohyla tica. The IUCN Red
List of Threatened Species 2013: e.T55675A3031693. [Link]
[Link]. Downloaded on 19 January 2020
IUCN SSC Amphibian Specialist Group and NatureServe (2013b) Lithobates vibicarius. The IUCN
Red List of Threatened Species 2013: e.T58746A3072875. [Link]
[Link]. Downloaded on 19 January 2020
IUCN SSC Amphibian Specialist Group and NatureServe (2014) Agalychnis annae. The IUCN Red
List of Threatened Species 2014: e.T55288A3027705. [Link]
[Link]. Downloaded on 20 January 2020
James TY, Toledo LF, Rödder D, da Silva LD, Belasen A, Betancourt-Román C, Jenkinson TS,
Soto-Azat Z, Lambertini C, Longo AV, Ruggeri J, Collins JP, Burrowes PA, Lips KR, Zamudio
KR, Longcore JE (2015) Disentangling host, pathogen, and environmental determinants of a
recently emerged wildlife disease: lessons from the first 15 years of amphibian chytridiomycosis
research. Ecol Evol 5:4079–4097
Joslin JD, Haber WA, Hamilton D (2018) Ocotea monteverdensis. The IUCN Red List of Threat-
ened Species 2018: e.T48724260A117762662. [Link]
[Link]. Downloaded on 19 January 2020
Kamal S, Grodzińska-Jurczak M, Brown G (2015) Conservation on private land: a review of global
strategies with a proposed classification system. J Environ Plan Manag 58(4):576–597. https://
[Link]/10.1080/09640568.2013.875463
Keuroghlian A, Desbiez A, Reyna-Hurtado R, Altrichter M, Beck H, Taber A, Fragoso JMV (2013)
Tayassu pecari. The IUCN Red List of Threatened Species 2013: e.T41778A44051115. https://
[Link]/10.2305/[Link]. Downloaded on
19 January 2020
Khela S (2014) Magnolia poasana. The IUCN Red List of Threatened Species 2014: e.
T193992A2293588. [Link]
Downloaded on 24 January 2020
Köhler L, Tobón C, Frumau KA, Bruijnzeel LS (2007) Biomass and water storage dynamics of
epiphytes in old-growth and secondary montane cloud forest stands in Costa Rica. Plant Ecol
193(2):171–184
274 Q. Newcomer et al.

Mayer AL, Tikka PM (2006) Biodiversity conservation incentive programs for privately owned
forests. Environ Sci Pol 9(7–8):614–625. [Link]
Miranda F, Bertassoni A, Abba AM (2014) Myrmecophaga tridactyla. The IUCN Red List of
Threatened Species 2014: e.T14224A47441961. [Link]
[Link]. Downloaded on 19 January 2020
Mitré M (1998a) Capparis discolor. The IUCN Red List of Threatened Species 1998: e.
T37802A10077032. [Link]
Downloaded on 24 January 2020
Mitré M (1998b) Ilex costaricensis. The IUCN Red List of Threatened Species 1998: e.
T30657A9570425. [Link]
Downloaded on 24 January 2020
Mitré M (1998c) Zinowiewia costaricensis. The IUCN Red List of Threatened Species 1998: e.
T30585A9558303. [Link]
Downloaded on 24 January 2020
Morse WC, Schedlbauer JL, Sesnie SE, Finegan B, Harvey CA, Hollenhorst SJ, Kavanagh KL,
Stoian D, Wulfhorst JD (2009) Consequences of environmental service payments for Forest
retention and recruitment in a Costa Rican biological corridor. Ecol Soc 14(1):23
Muchhala N (2003) Exploring the boundary between pollination syndromes: bats and humming-
birds as pollinators of Burmeistera cyclostigmata and B. tenuiflora (Campanulaceae).
Ocecologia 134:373–380
Nadkarni NM (1981) Canopy roots: convergent evolution in rainforest nutrient cycles. Science
214(4524):1023–1024
Nadkarni NM, Solano R (2002) Potential effects of climate change on canopy communities in a
tropical cloud forest: an experimental approach. Oceologia 131:580–586
Nadkarni NM, Schaeferz D, Matelson TJ, Solano R (2002) Comparison of arboreal and terrestrial
soil characteristics in a lower montane forest, Monteverde, Costa Rica. Pedobiologia 46:24–33
Nadkarni NM, Wheelwright NT (eds) (2000) Monteverde: Ecology and conservation of a tropical
cloud forest. Oxford University Press
Nadkarni NM, Wheelwright NT (eds) (revised 2014) Monteverde: Ecology and conservation of a
tropical cloud forest. Oxford University Press
NatureServe and IUCN SSC Amphibian Specialist Group (2013) Duellmanohyla uranochroa. The
IUCN Red List of Threatened Species 2013: e.T55314A3028798. [Link]
[Link]. Downloaded on 19 January 2020
Nelson C (1998a) Agonandra macrocarpa. The IUCN Red List of Threatened Species 1998: e.
T36132A9984004. [Link]
Downloaded on 24 January 2020
Nelson C (1998b) Eugenia salamensis. The IUCN Red List of Threatened Species 1998: e.
T36135A9984195. [Link]
Downloaded on 24 January 2020
Nelson C (1998c) Terminalia bucidoides. The IUCN Red List of Threatened Species 1998: e.
T32502A9710189. [Link]
Downloaded on 24 January 2020
Newcomer Q (2007) Innovations in Private Land Conservation: An Integrated Evaluation of
Payment for Environmental Services in the Path of the Tapir Biological Corridor in Costa
Rica. PhD dissertation Yale University School of Forestry and Environmental Studies, New
Haven, CT
NOAA National Centers for Environmental Information (NCEI) (2020) U.S. Billion-Dollar weather
and climate disasters. [Link]
Norton DA (2000) Conservation biology and private land: shifting the focus. Conserv Biol 14(5):
1221–1223
O’Halloran DT, Mowatt E, Leclerc ED, Bhatia M (2018) Designing a marketing plan for rural
Tourism in Costa Rica. Worcester Polytechnic Institute Interactive Qualifying Projects.
Retrieved from [Link]
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 275

Padgett-Vasquez S (2019) Forest connectivity from a Bird’s eye view: linking Cloud Forests to
Mangroves in the Bellbird Biological Corridor of Costa Rica. PhD dissertation, The University
of Georgia, Athens, GA
Page W (1998) Ficus lateriflora. The IUCN Red List of Threatened Species 1998: e.
T30556A9552527. [Link]
Downloaded on 24 January 2020
Pasquini L, Fitzsimons J, Cowell S, Brandon K, Wescott G (2011) The establishment of large
private nature reserves by conservation NGOs: key factors for successful implementation. Oryx
45(3):373–380. [Link]
Paulson DR (2009a) Epigomphus subsimilis. The IUCN Red List of Threatened Species 2009: e.
T7836A12854505. [Link]
Downloaded on 19 January 2020
Paulson DR (2009b) Libellula mariae. The IUCN Red List of Threatened Species 2009: e.
T165077A5960306. [Link]
Downloaded on 30 January 2020
Paulson D, von Ellenrieder N (2006a) Palaemnema baltodanoi. The IUCN Red List of Threatened
Species 2006: e.T60261A12317735. [Link]
[Link]. Downloaded on 30 January 2020
Paulson D, von Ellenrieder N (2006b) Perigomphus pallidistylus. The IUCN Red List of Threatened
Species 2006: e.T29603A9503838. [Link]
[Link]. Downloaded on 30 January 2020
Payan E, de Oliveira T (2016) Leopardus tigrinus. The IUCN Red List of Threatened Species 2016:
e.T54012637A50653881. [Link]
[Link]. Downloaded on 19 January 2020
Piedrahíta López C (2013) Lineamientos Técnicos para la Planificación de la Adaptación al Cambio
Climático en Áreas Funcionales para la Conservación de la Biodiversidad. MSc thesis Centro
Agronómico Tropical de Investigación y Enseñanza (CATIE), Turrialba, Costa Rica
Porras LW, Chaves G, Solórzano A (2013) Trimetopon simile. The IUCN Red List of Threatened
Species 2013: e.T203605A2769074. [Link]
[Link]. Downloaded on 20 January 2020
Pounds JA, Fogden MP (2000) Appendix 8: amphibians and reptiles of Monteverde. In: Nadkarni
NM, Wheelwright NT (eds) Monteverde: ecology and conservation of a tropical cloud forest.
Oxford University Press, New York, pp 537–538
Pounds J, Fogden M, Campbell J (1999) Biological response to climate change on a tropical
mountain. Nature 398:611–615. [Link]
Pounds JA, Bustamante M, Coloma L, Consuegra JA, Fogden MPL, Foster PN, La Marca E,
Masters KL, Merino-Viteri A, Puschendorf R, Ron SR, Sánchez-Azofeifa GA, Still CJ, Young
B (2006) Widespread amphibian extinctions from epidemic disease driven by global warming.
Nature 439:161–167. [Link]
Pounds J, Bolaños F, Chaves G (2008a) Craugastor andi. The IUCN Red List of Threatened Species
2 0 08 : e . T 5 6 41 4 A 11 4 7 3 1 54 . h t t p s : / / d oi . or g / 1 0 .2 3 0 5 / I U C N . U K .2 0 0 8 . R L T S.
[Link]. Downloaded on 20 January 2020
Pounds J, Bolaños F, Chaves G (2008b) Craugastor angelicus. The IUCN Red List of Threatened
Species 2008: e.T56418A11473990. [Link]
[Link]. Downloaded on 20 January 2020
Pounds J, Bolaños F, Chaves G, Wake D, Savage J (2008c) Bolitoglossa subpalmata. The IUCN
Red List of Threatened Species 2008: e.T59212A11897656. [Link]
[Link]. Downloaded on 20 January 2020
Pounds J, Bolaños F, Solís F, Ibáñez R, Chaves G, Savage J, Jaramillo C, Fuenmayor Q (2008d)
Pristimantis altae. The IUCN Red List of Threatened Species 2008: e.T56406A11470632.
[Link] Downloaded on
19 January 2020
276 Q. Newcomer et al.

Pounds J, Bolaños F, Solís F, Ibáñez R, Chaves G, Savage J, Jaramillo C, Fuenmayor Q, Castro F,

Grant T, Lynch J, Lips K (2008e) Pristimantis caryophyllaceus. The IUCN Red List of
Threatened Species 2008: e.T56497A11475825. [Link]
[Link]. Downloaded on 20 January 2020
Pounds J, Bolaños F, Wake D, Savage J (2008f) Nototriton gamezi. The IUCN Red List of
Threatened Species 2008: e.T59294A11900971. [Link]
[Link]. Downloaded on 19 January 2020
Pounds J, Puschendorf R, Bolaños F, Chaves G, Crump M, Solís F, Ibáñez R, Savage J, Jaramillo C,
Fuenmayor Q, Lips K (2010) Atelopus varius. The IUCN Red List of Threatened Species 2010:
e.T54560A11167883. [Link]
Downloaded on 19 January 2020
Powell G, Bjork R (1994) Implications of altitudinal migration for conservation strategies to protect
tropical biodiversity: a case study of the Resplendent Quetzal Pharomacrus mocinno at
Monteverde, Costa Rica. Bird Conserv Int 4(2–3):161–174. [Link]
S0959270900002744
Powlen K (2018) The motivations and barriers of landowner participation in reforestation in the
bellbird biological corridor in Costa Rica. MSc Thesis Colorado State University, Fort
Collins, CO
Quigley H, Foster R, Petracca L, Payan E, Salom R, Harmsen B (2017) Panthera once (errata
version published in 2018). The IUCN Red List of Threatened Species 2017: e.
T15953A123791436. [Link]
Downloaded on 19 January 2020
Rains KC, Nadkarni NM, Bledsoe CS (2003) Epiphytic and terrestrial mycorrhizas in a lower
montane costa Rican cloud forest. Mycorrhiza 13:257–264
Ringhofer L, Singh SJ, Smetschka B (2013) Climate change mitigation in Latin America: a
mapping of current policies, plans and programs. Institute for social ecology working paper 143
Rivers MC, Barstow M, Fuentes ACD (2019) Ticodendron incognitum. The IUCN Red List of
Threatened Species 2019: e.T37468A128258819. [Link]
[Link]. Downloaded on 24 January 2020
Rodriguez B, Pineda W (2015) Ectophylla alba. The IUCN Red List of Threatened Species 2015: e.
T7030A22027138. [Link]
Downloaded on 19 January 2020
Ruedas L, Smith AT (2019) Sylvilagus brasiliensis. The IUCN Red List of Threatened Species
2019: e.T87491102A45191186. [Link]
[Link]. Downloaded on 19 January 2020
Sandiford M (1998) Bombacopsis quinata. The IUCN Red List of Threatened Species 1998: e.
T32905A9738742. [Link]
Downloaded on 24 January 2020
Savage J, Pounds J, Bolaños F (2008) Incilius periglenes. The IUCN Red List of Threatened
Species 2008: e.T3172A9654595. [Link]
[Link]. Downloaded on 19 January 2020
Scrimshaw N, Gibbons W, Burlingame LJ (2000) Conservation Easements in Monteverde: the
Enlace Verde project. In: Burlingame L Conservation in the Monteverde Zone in: Nadkarni NM
and Wheelwright NT (eds) Monteverde: ecology and conservation of a tropical cloud forest.
Oxford University Press, New York, pp. 381–382
Skinner D (2014) Costus nitidus. The IUCN Red List of Threatened Species 2014: e.
T56347808A56353108. [Link]
[Link]. Downloaded on 19 January 2020
Snow BK (1977) Territory behavior and courtship of the male three-wattled bellbird. Auk 94:623–
645
Solari S (2018) Vampyrum spectrum. The IUCN Red List of Threatened Species 2018: e.
T22843A22059426. [Link]
Downloaded on 19 January 2020
10 The Monteverde Cloud Forest: Evolution of a Biodiversity Island in Costa Rica 277

Solís F, Ibáñez R, Chaves G, Savage J, Jaramillo C, Fuenmayor Q (2008a) Ecnomiohyla

fimbrimembra. The IUCN Red List of Threatened Species 2008: e.T55482A11304367.
[Link] Downloaded on
20 January 2020
Solís F, Ibáñez R, Chaves G, Savage J, Jaramillo C, Fuenmayor Q, Bolaños F, Lips K(2008b)
Isthmohyla zeteki. The IUCN Red List of Threatened Species 2008: e.T55701A11351517.
[Link] Downloaded on
27 January 2020
Solís F, Ibáñez R, Pounds J, Bolaños F, Chaves G (2008c) Isthmohyla angustilineata. The IUCN
Red List of Threatened Species 2008: e.T55390A11290945. [Link]
[Link]. Downloaded on 27 January 2020
Solís F, Ibáñez R, Savage J, Jaramillo C, Fuenmayor Q, Kubicki B, Pounds J, Chaves G, Jungfer K,
Lips K (2008d) Agalychnis lemur. The IUCN Red List of Threatened Species 2008: e.
T55855A11381418. [Link]
Downloaded on 20 January 2020
Solís F, Ibáñez R, Chaves G, Savage J, Jaramillo C, Fuenmayor Q, Bolaños F (2010a) Craugastor
podiciferus. The IUCN Red List of Threatened Species 2010: e.T56870A11547021. [Link]
org/10.2305/[Link]. Downloaded on 20 January 2020
Solís F, Ibáñez R, Chaves G, Savage J, Jaramillo C, Fuenmayor Q, Kubicki B, Bolaños F (2010b)
Ecnomiohyla miliaria. The IUCN Red List of Threatened Species 2010: e.T55561A11319134.
[Link] Downloaded on
20 January 2020
Solís F, Ibáñez R, Pounds JA, Bolaños F, Chaves G, and Lips K (2010c) Isthmohyla rivularis. The
IUCN Red List of Threatened Species 2010: e.T55627A11343375. [Link]
[Link]. Downloaded on 20 January 2020
Solórzano A, Acosta Chaves V, Chaves G (2013) Celestus hylaius. The IUCN Red List of
Threatened Species 2013: e.T203032A2758756. [Link]
[Link]. Downloaded on 20 January 2020
Stevenson R, Haber WA (2000) Section 4.5.2: Migration of butterflies through Monteverde. In:
Hanson P (ed) Insects and spiders. In: Nadkarni NM, Wheelwright NT (eds) Monteverde:
ecology and conservation of a tropical cloud Forest. Oxford University Press, New York, pp
118–119
Stuckey J, Camacho Céspedes F, Vargas Leitón G, Stuckey SA, Vargas Leitón JL (2014) Agricul-
ture in Monteverde, moving toward sustainability—update 2014. In: In Nadkarni NM, Wheel-
wright NT (eds) Monteverde: ecology and conservation of a tropical cloud forest – 2014
updated chapters. Oxford University Press, New York
Timm RM, LaVal RK (1998) A field key to the bats of Costa Rica, Occasional publication series 22:
1–30. Center of Latin American Studies, The University of Kansas, Lawrence
Timm RM, LaVal RK (2000) Appendix 10: Mammals of Monteverde. In: Nadkarni NM, Wheel-
wright NT (eds) Monteverde: ecology and conservation of a tropical cloud Forest. Oxford
University Press, New York, pp 553–557
Tosi JA, Holdridge LR, Centro Científico Tropical (1969) República de Costa Rica, mapa ecológico
San José. Centro Científico Tropical, Costa Rica. [Link] Web.
18 December 2021
Tropical Science Center (2018) Promoción de Encadenamientos Productivos Sostenibles en el
Corredor Biológico Pájaro Campana. Programa de Corredores Biológicos, Gerencia de
Desarrollo, Centro Científico Tropical, San José, Costa Rica
Wainwright M (2007) The mammals of Costa Rica: a natural history and field guide,
No. 599 W142m. Cornell University Press, Ithaca
Wake D, Savage J, Bolaños F (2008) Oedipina uniformis. The IUCN Red List of Threatened
Species 2008: e.T59327A11918239. [Link]
[Link]. Downloaded on 20 January 2020
278 Q. Newcomer et al.

Wegier A, Lorea Hernández F, Contreras A, Mastretta-Yanes A (2017) Persea schiedeana. The

IUCN Red List of Threatened Species 2017: e.T34402A61528670. [Link]
[Link]. Downloaded on 24 January 2020
Wegner GI (2016) Payments for ecosystem services (PES): a flexible, participatory, and integrated
approach for improved conservation and equity outcomes. Environ Dev Sustain 18:617. https://
[Link]/10.1007/s10668-015-9673-7
Welch J et al (2011) Plan Estratégico del Corredor Biológico Pájaro Campana (CBPC) 2011–2016.
Monteverde, Costa Rica
Wheelwright NT (1983) Fruits and the ecology of resplendent quetzals. Auk 100:286–301
Wheelwright N (2000) Conservation biology. In: Nadkarni NM, Wheelwright NT (eds)
Monteverde: ecology and conservation of a tropical cloud forest. Oxford University Press,
New York, pp 419–456
Wheelwright N (2014) Conservation biology – update 2014. In: Nadkarni NM, Wheelwright NT
(eds) Monteverde: ecology and conservation of a tropical cloud Forest – 2014 updated chapters.
Oxford University Press, New York
Whitfield SM, Lips KR, Donnelly MA (2016) Amphibian decline and conservation in Central
America. Copeia 104(2):351–379
Whitfield SM, Alvarado G, Abarca J, Zumbado H, Zuñiga I, Wainwright M, Kerby J (2017)
Differential patterns of Batrachochytrium dendrobatidis infection in relict amphibian
populations following severe disease-associated declines. Dis Aquat Org 126:33–41. https://
[Link]/10.3354/dao03154
World Conservation Monitoring Centre (1998a) Chrysophyllum hirsutum. The IUCN Red List of
Threatened Species 1998: e.T37729A10074123. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998b) Dichapetalum costaricense. The IUCN Red List of
Threatened Species 1998: e.T36492A9999314. [Link]
[Link]. Downloaded on 19 January 2020
World Conservation Monitoring Centre (1998c) Elaeagia uxpanapensis. The IUCN Red List of
Threatened Species 1998: e.T31855A9665319. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998d) Ocotea viridiflora. The IUCN Red List of Threat-
ened Species 1998: e.T30597A9560236. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998e) Pouteria austin-smithii. The IUCN Red List of
Threatened Species 1998: e.T34409A9864731. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998f) Pouteria congestifolia. The IUCN Red List of
Threatened Species 1998: e.T32686A9717700. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998g) Pouteria fossicola. The IUCN Red List of Threat-
ened Species 1998: e.T37716A10072704. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998h) Sideroxylon capiri subsp. capiri. The IUCN Red
List of Threatened Species 1998: e.T36041A9977046. [Link]
[Link]. Downloaded on 24 January 2020
World Conservation Monitoring Centre (1998i) Sideroxylon persimile subsp. subsessiliflorum. The
IUCN Red List of Threatened Species 1998: e.T36057A9971292. [Link]
[Link]. Downloaded on 24 January 2020
Zamzow BK, Nieman SJ, Davis CN, Garro Cruz M, Monroe A, Stallcup L, Moran MD (2018)
Status of large terrestrial vertebrates in the Monteverde-Arenal bioregion, northwestern Costa
Rica. Tropical Conservation Science. [Link]
Zbinden S, Lee DR (2005) Paying for environmental services: an analysis of participation in Costa
Rica’s PSA program. World Dev 33:255–272
Chapter 11
A Highly Productive Biodiversity Island
Within a Monoculture Landscape: El
Hatico Nature Reserve (Valle del Cauca,
Colombia)

Zoraida Calle D, Carlos Hernán Molina C, Carlos Hernando Molina D,

Enrique José Molina D, Juan José Molina E, Bernardo Murgueitio C,
Amalia Murgueitio C, and Enrique Murgueitio R

Abstract This chapter describes the landscape-scale, national and regional influ-
ence of a rural property that forms a biodiversity island within a monoculture
landscape. Managed continuously by nine generations of the Molina family, El
Hatico Nature Reserve embodies a set of values grounded in a deep connection to
the land. Between 1960 and 1990, the fertile flatlands of the Cauca River valley lost
almost all dry forest remnants, wetlands, traditional annual crops, and agroforestry
systems, adopting a uniform method of sugarcane production that modified stream
banks, eliminated the small-scale topographic heterogeneity, and integrated periodic
burning and herbicide applications as part of the management protocols. Meanwhile,
El Hatico gained tree cover, enhanced its soil quality, conserved its forest fragments,
transformed its conventional pastures into biodiverse silvopastoral systems and
transitioned to agroecological sugarcane production. El Hatico’s long tradition of
agricultural and livestock research and detailed production records helped develop

Coauthor Carlos Hernán Molina passed away during the final stage of editing this book. None of
the innovations described in this chapter would exist were it not for his leadership, moral courage
and commitment to generational exchange and agroecology. With admirable clarity and wisdom, he
guided his family through decades of dramatic land use change and stood up for a diversified
agricultural production against the ravaging advance of monoculture. Thanks to Carlos Hernán,
El Hatico persists as a biodiversity island.

Zoraida Calle D (*) · Bernardo Murgueitio C · Amalia Murgueitio C · Enrique Murgueitio R

Centro para la Investigación en Sistemas Sostenibles de Producción Agropecuaria (CIPAV),
Cali, Colombia
e-mail: zoraida@[Link]
Carlos Hernando Molina D · Enrique José Molina D · Juan José Molina E
Centro para la Investigación en Sistemas Sostenibles de Producción Agropecuaria (CIPAV),
Cali, Colombia
El Hatico Nature Reserve, El Cerrito, Colombia

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 279
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
280 Zoraida Calle D et al.

the highly efficient intensive silvopastoral systems in which cattle graze on nitrogen-
fixing fodder shrubs interspersed with grasses and under the shade of native trees.
These silvopastures have inspired thousands of technical assistants, extension
workers and farmers in Colombia and other Latin American countries to undertake
the transformation of conventional cattle ranching systems. Simultaneously, El
Hatico developed organic sugarcane by applying the principles of agroecology to
produce sugar while storing carbon, enhancing the soil biota and making an efficient
use of water. El Hatico’s forest fragment is surrounded by a wildlife-friendly
silvopastoral matrix that is permeable to the movements of birds and arthropods.
This property’s unique combination of land uses provides a model for the integration
of agroecology, agroforestry and ecological restoration.

Keywords Agroecology · Ecological restoration · Functional biodiversity ·

Intensive silvopastures · Silvopastoral systems · Sugarcane

11.1 Introduction

Can a single rural property make a difference for biodiversity and sustainability? If
so, on which scales can that occur? This chapter explores the spheres of influence of
a family estate that stands out as a biodiversity island within an intensive monocul-
ture landscape. Seen from the air, El Hatico Nature Reserve, located in the fertile
flatlands of the geographic Cauca river valley in Colombia, is the only woodland in a
landscape dominated by sugarcane plantations. On closer inspection, what appears
to be a secondary forest is in fact an old growth dry forest remnant, partially
surrounded by silvopastures with a high density and diversity of trees. On one
edge, this forest fragment borders a sugarcane plantation that has been managed
agroecologically for two decades, produces certified organic sugar, and has lines of
native palms and trees between rows of sugarcane plants.
The first section of this chapter provides a brief historical context of land use
change in the geographic Cauca river valley and the lower Amaime river basin,
where El Hatico is located. The following sections describe two land-use changes
that were occurring at El Hatico while the surrounding landscape was following the
opposite trends: the transition from conventional cattle ranching to silvopastoral
systems and the adoption of agroecological practices in sugarcane production. Then,
we summarize the results of research projects done at El Hatico on the spatial
distribution of ants, parasitic wasps, spiders and birds, and the functional biodiver-
sity in silvopastures and sugarcane. In the final section, we explore El Hatico’s
influence at larger spatial scales through research, training and inspiring farmers,
extension workers, and decision makers. We also discuss El Hatico’s impacts on
land use policy related to livestock, sustainable agriculture, and private conservation
initiatives.
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 281

This chapter focuses on why El Hatico is an outstanding leader in tropical

sustainable agriculture, with broad impacts from local to global scales. The farm
provides critical habitat for biodiversity within its agricultural matrix, modeling the
alignment of increased productivity with enhanced ecological functioning.

11.2 Historical Context: Land Use Change

in the Geographic Cauca River Valley

The geographic Cauca river valley comprises 421,000 hectares of flatlands in the
high Cauca river basin, surrounded by the central and western Andean ranges
(Cordillera Central and Cordillera Occidental) in the Colombian departments of
Valle del Cauca, Cauca and Risaralda. Between 1957 and 1986, 66% of the forest
cover in the upper Cauca river basin was transformed into agricultural land (CVC
1990). Today this landscape retains only 1.76% of its original forest, represented by
scattered fragments with a mean area of 6 ha (Arcila et al. 2012). Wetlands once
occupied almost one quarter of the geographic Cauca river valley; the few remaining
wetlands are besieged by urban and agricultural expansion (Rivera et al. 2007).
As a result of fragmentation, the extreme isolation of forest remnants and the
opportunity cost of the land, the agricultural landscape of the geographic Cauca
River valley has very limited opportunities for conservation based on protected areas
and restored biological corridors alone. Close to 75% of the remaining forest
fragments are separated from their nearest neighbor by distances of 500 m or more
(Arcila et al. 2012). Approximately 50% of these fragments lack a real forest interior
area when assuming a 50 m edge effect. The fact that the isolated forest patches are
surrounded by hostile matrices that barely promote the movement of organisms
heralds an uncertain future for the regional biodiversity. Small populations of
animals and plants in isolated remnants tend to have a high probability of local
extinction, related to low genetic variability combined with demographic and sto-
chastic effects (Eibl et al. 2022; Niella et al. 2022). All conservation or restoration
initiatives in the geographic Cauca river valley should involve redesigning and
managing the agricultural matrix to enhance the movement of plants and animals
in a landscape dominated by sugarcane (Calle et al. 2012, 2013).
El Hatico Nature Reserve is located at the lower Amaime river basin, close to the
site where the Amaime river joins the larger Cauca river (162 km south-west of
Pinzacuá farm; see Fig. 12.1 in Montes-Londoño et al. 2022). The lower Amaime
basin retained an extensive forest cover until the 1950s. Between 1950 and 1970,
agriculture expanded in this area with public and private investment at the expense of
forests, wetlands and even cattle ranching (Murgueitio 2019). Large areas were
planted with cotton, millet, corn, soy, beans, rice and sugarcane. Landscape trans-
formation in the 1960s was driven mainly by the economic blockade of Cuba and the
increasing demand for sugar in the United States. The growing dominance of
intensive sugarcane production has continued until the present.
282 Zoraida Calle D et al.

Fig. 11.1 Sugarcane, silvopastures and forest at El Hatico Nature Reserve. (Photo: Juan Diego
Vanegas)

Murgueitio (2019) analyzed the changes in land cover that took place between
1986 and 2018 in the lower Amaime river basin. Land covered by perennial crops
(mainly sugarcane) increased from 62.2 to 74%, while annual crops declined from
15.3 to 2.2%. Areas occupied by infrastructure increased by 1.9%, areas covered by
natural vegetation increased by 0.8%, and pastures declined by 1.6%.
El Hatico Nature Reserve occupies 0.64% of the lower Amaime river basin but
conserves 4% of its forested areas (Murgueitio 2019). This 288-ha property com-
bines silvopastoral systems (140 ha), sugarcane (100 ha), forest (14 ha), Guadua
angustifolia forest (26 ha), restored biological corridors (2 ha) and mixed fruit trees
(5 ha); Figs. 11.1 and 11.2). El Hatico is located at 1000 m of altitude and has
average temperature of 24
C, average annual rainfall of 750 mm and 75% relative
humidity (Molina-Castro et al. 2012). Due to its location at the center of the
geographic Cauca river valley, evapotranspiration at El Hatico (1600 mm year1)
far exceeds rainfall, creating a significant moisture deficit. The Cauca river valley is
considered a dry tropical forest according to the Holdridge life zone system,
although most trees retain their foliage even during the driest periods. This mild
deciduousness is explained by the bimodal distribution of rainfall and the superficial
water table.
In 1942, land use in El Hatico showed a pattern similar to other rural properties in
the area, with one forest fragment, giant bamboo (Guadua angustifolia) stands,
pastures with a low density of trees, and sugarcane plantations. However, an aerial
photograph from 1986 shows a radically different landscape context; El Hatico’s
forest had become the only remaining fragment in the area, and the surrounding
matrix had been transformed into a simplified grid of sugarcane plantations with few
strips of giant bamboo forest. Satellite images from 2007 and 2018 show another
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 283

Fig. 11.2 Land use distribution at El Hatico Nature Reserve

reality, in which tree density and canopy cover had increased in El Hatico’s
silvopastures, creating an open woodland physiognomy in which the edges of the
forest fragment had become blurred (Molina-Castro et al. 2012). In two decades,
vegetation structure was enhanced and the once simplified agricultural matrix
became suitable habitat for diverse groups of organisms.

11.3 From Conventional Cattle Ranching to Silvopastoral

Systems

El Hatico has a long tradition of cattle farming. For several generations, livestock
production focused on beef, horses, mules and bovine breeding stock. Dairy farming
played a marginal role until the development of local road infrastructure in the
1950s.
The foundations of El Hatico’s silvopastoral systems go back to the first decades
of the twentieth century. The first Colombian publication on the role of trees in cattle
pastures, written by a member of the Molina family, mentions the failure of the initial
trials with temperate legumes planted as feed for cattle in the tropics and highlights
the nutritional value of fruits from native leguminous trees such as Samanea saman
and Enterolobium cyclocarpum (Molina-Garcés 1938).
284 Zoraida Calle D et al.

Trees of a high-yield ecotype of Prosopis juliflora were planted in paddocks of El

Hatico in the 1940s to provide nutritious fruits for the cattle. These trees offer a high
energy supplement in the form of sweet pods during the driest months of the year,
when grass biomass drops to its lowest level. In the 1980s, Gliricidia sepium trees
were planted at high density to provide cut-and-carry fodder; this Gliricidia fodder
bank was used for more than two decades. However, the manual harvest of forage is
labor-intensive and most tree and shrub species with highly nutritional forage do not
tolerate cattle browsing. This motivated El Hatico’s team to integrate leguminous
shrubs that could be browsed directly in the paddocks. An empirical observation of
cattle browsing Leucaena leucocephala trees led to the initial idea of planting this
species in an intensive silvopastoral system (Murgueitio et al. 2015).

11.3.1 Intensive Silvopastoral Systems

El Hatico pioneered the adoption of silvopastoral systems in the 1970s and the
development of intensive silvopastoral systems (ISS) in the 1990s. ISS are charac-
terized by the high-density cultivation of fodder shrubs (5000–80,000 plants ha1)
interspersed with improved tropical grasses, legumes, trees and palms (Murgueitio
et al. 2015; Chará et al. 2017; Santos-Gally and Boege 2022).
Today, El Hatico’s ISS form a complex and wildlife friendly agricultural matrix
that combines grass species such as Cynodon plectostachyus, Megathyrsus maximus,
Cynodon dactylon and Paspalum notatum; a high density of the nitrogen-fixing tree
Leucaena leucocephala (up to 30,000 plants ha1 managed as fodder shrubs); 30–50
medium sized planted and regenerating trees ha1; a lower density of large shade and
timber trees and palms (Murgueitio et al. 2011; Calle et al. 2013; Chará et al. 2015).
Paddocks are separated by 40 km of live fences of trees such as Gliricidia sepium
(some of which are more than 100 years old), broad leaf mahogany (Swietenia
macrophylla), Guazuma ulmifolia, Maclura tinctoria, the large bromeliad Bromelia
plumieri and mixed live fences formed by naturally regenerating trees, shrubs and
herbs (Molina-Castro et al. 2012). Table 11.1 presents the most common tree and
palm species found in silvopastures; Figure 11.3 shows a live fence of broad-leaf
mahogany.
The grazing method applied in El Hatico’s ISS combines high animal loads
(50 Lucerna dairy cows with an average weight of 450 kg in 4000 m2 paddocks)
with brief rotations (2 days), followed by long periods of recovery (45 days).
Therefore, throughout the year, each individual paddock is grazed intensively for a
total of 16 days and recovers during the remaining 349 days.
El Hatico has access to two sources of irrigation: a concession agreement
providing a fixed volume of water from the Amaime river and groundwater that is
pumped from a deep well (100 m) at a high energetic cost. Both sources were used
for several decades to irrigate pastures. The gradual adoption of complex
silvopastoral systems allowed El Hatico to increase its per-hectare productivity
from 7436 l of milk in 1996 to 18,299 l in 2004 (El Hatico, unpublished data).
After reaching that historical peak yield, the owners decided to suspend irrigation,
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 285

Table 11.1 Common trees and palms in silvopastures and live fences at El Hatico
Scientific name Local name Family Origin Uses
Attalea butyracea (Mutis ex L. f.) Palma corozo de Arecaceae P, NR FW
Wess. Boer puerco
Roystonea regia (Kunth) O.F. Cook Palma real Arecaceae P, NR A
Syagrus sancona H. Karst. Palma zancona Arecaceae P A, FW
Bromelia plumieri (E. Morren) Piñuela Bromeliaceae P LF, FW
L.B. Sm.
Enterolobium cyclocarpum (Jacq.) Orejero Fabaceae NR T, FC
Griseb.
Gliricidia sepium (Jacq.) Kunth Matarratón Fabaceae P LF, T
ex Walp.
Prosopis juliflora (Sw.) DC. Mezquite, Fabaceae P, NR T, FC
Algarrobo
Samanea saman (Jacq.) Merr. Samán Fabaceae NR T, FC
Senna spectabilis (DC.) H.S. Irwin & Vainillo Fabaceae NR FC
Barneby
Ceiba pentandra (L.) Gaertn. Ceiba Malvaceae NR A
Guazuma ulmifolia Lam. Guácimo Malvaceae NR FC, LF
Guarea guidonia (L.) Sleumer Cedro macho Meliaceae NR T, FW
Swietenia macrophylla King Caoba Meliaceae P T, LF
Cedrela odorata L. Cedro rosado Meliaceae P, NR T
Maclura tinctoria (L.) D. Don ex Dinde Moraceae P, NR LF, T,
Steud. FW
Zanthoxylum rhoifolium lam. Tachuelo Rutaceae NR T, FW
Origin: Planted (P), Natural Regeneration (NR); Uses: Aesthetics (A), Fruits for cattle (FC), Fruits
for wildlife (FW), Live fence (LF), Timber (T)

making their milk production depend on the water stored in the soil, known as “green
water”. Since then, ISS became 100% rainfed and milk yield stabilized around
15,000 l ha1 year1 (Molina-Castro et al. 2012).
Organic dairy products from El Hatico (and the closely related Lucerna farm1)
have been a logical consequence of the superior quality and safety of this agroeco-
logical milk, rather than being an explicit goal of the adoption of silvopastoral
systems. By replacing its conventional monocultures of African star grass (Cynodon
plectostachyus) with ISS, El Hatico cut down fixed costs, added value to the milk
through organic certification and increased the profitability of dairy farming. The
cost of mineral salt was reduced by 42% because ISS provide abundant minerals and
the animals limit their salt intake. The costs of irrigation and fertilizer were
completely eliminated. Without ISS, the current per hectare profit would be US$ -
27 ha1 month1 for a milk price of US $0.35. With ISS, the monthly per hectare
profit is $206 for a market price of US $0.47 l1 for certified organic milk (El Hatico,
unpublished records).

1
Lucerna is another rural property dedicated to organic milk and sugarcane production, located in
Bugalagrande, Valle del Cauca.
286 Zoraida Calle D et al.

Fig. 11.3 Live fence of broad-leaf mahogany (Swietenia macrophylla). (Photo: Carlos Pineda)

11.3.2 Soil Recovery in ISS

Several studies on soil conditions and other environmental factors that influence crop
and animal productivity have been conducted at El Hatico. For example, Vallejo
et al. (2010) studied the microbiological, physical and chemical properties of the soil
in a chronosequence of silvopastoral systems (3–6, 8–10 and 12–15 years old), and
compared them to a conventional pasture outside of El Hatico that had been grazed
intensively for 35 years and illustrates the baseline condition for the farm’s
silvopastures. In the conventional pasture, lower microbial responses (hydrolytic
enzyme activities) from bacteria involved in decomposition and nutrient minerali-
zation were explained by the high bulk density and penetration resistance that create
a less favorable environment for root exploration of nutrients and water (Vallejo
et al. 2010). In contrast, lower penetration resistance values in silvopastoral soils
indicate improved soil aggregation and greater pore space, both of which enhance
microbial habitats and activity.
The oldest silvopastures at El Hatico (12–15 years) showed the highest microbial
biomass and enzyme activity levels when normalized for carbon or clay contents
(Vallejo et al. 2010). The microbial responses summarized in Table 11.2 indicate
that silvopastures are improving soils; however, the results of this research suggest
that it takes at least 8 years (though likely >12 years) to fully appreciate this effect
under the conditions of El Hatico (Vallejo et al. 2010).
The diverse and multi-layered plant community (as shown in Fig. 11.4) is one of
the factors that explains greater microbial responses in ISS. With the adoption of
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 287

Table 11.2 Total carbon (per clay unit), bulk density, soil penetration resistance and enzyme
activity in silvopastures, conventional pasture and forest at El Hatico Nature Reserve
Relative enzyme
activity per unit
of C:
(B-glucosidase,
alkaline
phosphatase and
Land Use Age (years) C clay1 (x100%) BD (g cm3) SPR (MPa) urease)
SS12 12–15 10.0a 1.39b 3.30b *** *** ***
SS8 8–10 8.5a 1.40b 2.47b ** ** **
SS3 3–6 8.6a 1.44b 2.85b ** * **
CP >30 7.9a 1.52a 3.98a ** * **
F >100 8.3a 1.21c 1.49c *** * ***
Source: Vallejo et al. (2010, 2012)
C clay1 Total carbon per clay unit basis, BD Bulk density, SPR Soil penetration resistance.
Asterisks indicate the relative magnitude of enzyme activity levels; land uses sharing the same
number of asterisks are not significantly different at P < 0.05

silvopastures, fodder biomass production at El Hatico increased from 23 to

39 Mg ha1 year1 (Mahecha 2003). Higher biomass production and fodder yield
in these systems imply a higher return of carbon to the soil. Animal manure and urine
enhance soil microbial activity by providing readily usable substrates (Clegg et al.
2006). The root systems of multi-layered silvopastures likely have a higher biomass
that fully explores the soil profile. Increased above and belowground carbon inputs
seem to drive the microbial responses in older silvopastures. The results summarized
in Table 11.2 suggest that silvopastoral systems promote agroecosystem sustainabil-
ity by improving the ability of soils to perform decomposition and nutrient miner-
alization, as reflected by hydrolytic enzyme activities (Vallejo et al. 2010).
The enhanced soil moisture and water use efficiency of ISS is probably related to
the recovery of soil organic matter (Vallejo et al. 2010, 2012). The baseline condi-
tion for soil organic matter (SOM) in El Hatico’s paddocks was 2.9% in 1994, while
the reference condition for the forest is 4.3% (Arias 1994). The chronosequence of
ISS illustrates two contrasting situations: 3.4 to 3.7% of SOM outside tree crowns
and 4.4 to 4.9% under trees (Vallejo et al. 2012). These results show that the soil
under ISS as the one shown in Fig. 11.3 can store significant amounts of carbon and
that trees form high-carbon “islands” within the paddocks.
An additional benefit of El Hatico’s ISS is the reduction of greenhouse gas
emissions. Methane is a potent greenhouse gas with a global warming potential
equivalent to 25 times that of carbon dioxide (CO2). In cattle ranching, this gas
results mostly from the activity of anaerobic microorganisms of the Archaea domain,
that hydrolyze proteins, starches and cell wall components in the rumen. Animals
with high fiber and dry matter intakes (DMI) release more methane. However, diet
composition affects enteric methane production; when forages have high digestibil-
ity, less methane is produced per unit of DMI. Also, the presence of condensed
tannins in leucaena is known to reduce methane production by inhibiting the growth
288 Zoraida Calle D et al.

Fig. 11.4 Intensive silvopasture with Leucaena leucocephala and native trees. (Photo: Zoraida
Calle)

of cellulolytic and proteolytic bacteria. Methane released by the cattle per unit of
degraded dry matter is 30% lower in El Hatico’s ISS with leucaena than in conven-
tional star grass monocultures (Molina et al. 2015, 2016 and references therein).

11.4 Agroecological Sugarcane

For several decades, sugarcane in the geographic Cauca river valley has been planted
in intensive monocultures that use chemical fertilizers, herbicides and insecticides.
Pre-harvest herbicide is applied to induce stress and increase the concentration of
saccharose in the sugarcane. Pre- and post-harvest burning are done to facilitate the
manual harvest and to eliminate crop residues, respectively. Conventional practices
result in significant water pollution, soil compaction, greenhouse gas emissions and
an increasing vulnerability of sugarcane to pests. Currently, the sector is
transitioning from manual to mechanical harvest with heavy machinery in the
region; this makes burning unnecessary but will bring new challenges related to
soil physical degradation.
El Hatico joined the sugarcane industry in 1960 and implemented pre- and post-
harvest burning in 1972. In 1994, they realized that 50% of the soil organic matter
had been lost after two decades of conventional sugarcane cultivation (Arias 1994).
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 289

Since 1993 they had been monitoring parameters such as per hectare monthly
sugarcane biomass production with the Colombian Sugarcane Research Center
(Cenicaña, [Link]).
The transition to organic sugarcane production included the adoption of agroeco-
logical practices such as minimum tillage, mulching crop residues instead of burning
them, and integrating nitrogen-fixing legumes (Vigna unguiculata and Crotalaria
juncea) between cane rows to provide green manure. Herbicides were replaced with
African hair sheep (breed of Ovis aries that does not grow wool) that consume all
grasses and weeds, complemented with selective manual weed control. These
practices have promoted functional biodiversity and water economy (Sadeghian
and Madriñan 2000; Hincapié et al. 2019).
Stem borers (Diatraea spp., Lepidoptera: Crambidae) are considered the main
pests of sugarcane in the Americas and cause severe economic losses (Vargas et al.
2015; Solis and Metz 2016). Conventional biological control involves releasing
parasitoids such as Lydella minense, Billaea claripalpis, Trichograma demanun
and Cotesia flavipes, which have low survival rates in conventional sugarcane
plantations, where they lack vital resources such as pollen and nectar. Natural control
of Diatraea spp. by ants such as Solenopsis achinid, Wasmannia auropunctata,
Ectatomma ruidum, Pheidole spp., Solenopsis spp., Camponotus sp., Nylanderia
spp., and Pachycondyla ferruginea has been observed at El Hatico and other organic
farms (Gómez and Vargas 2014; Rivera et al. 2019). The Tachinid fly Genea jaynesi
is a native parasitoid that controls this pest in the Cauca river valley. Its mass-rearing
has not been successful in the laboratory, but it thrives in native vegetation strips,
where it feeds on flowers of common weeds (Vargas et al. 2006, 2015; Cenicaña
2017; Rivera et al. 2019).
In 1996, El Hatico obtained its organic certification for sugarcane and livestock
production. Shortly after, they started to integrate lines of native trees such as the
endangered Caesalpinia ebano and palms (Sabal mauritiiformis and Syagrus
sancona) within the sugarcane plots as shown in Fig. 11.5. Palm and tree lines
were established between the sugar cane lines, spaced 36 m apart, which is equiv-
alent to 24 lines of sugarcane, since the distance between sugarcane lines is 1.5 m.
Planting distance between palms was 2 m to facilitate the manual harvest of the palm
leaves for thatching, and there were 5 m between trees. Additional Syagrus sancona
palms are being planted between each pair of trees.

11.4.1 Yields of Agroecological Sugarcane at El Hatico

In the geographic Cauca river valley, the yield of sugarcane per kilogram of nitrogen
applied to the crop has declined steadily in conventional sugarcane, from 1.03 ton in
1985 to 0.58 ton in 2015 (Cenicaña, unpublished results). This waning response to
chemical nitrogen fertilization has triggered a soil degradation alert in the region. In
contrast to conventional sugarcane, the yield of El Hatico’s agroecological sugar-
cane varied between 1.3 and 1.87 ton per kilogram of nitrogen (from organic
290 Zoraida Calle D et al.

Fig. 11.5 Line of Sabal mauritiiformis palms in agroecological sugarcane plantation at El Hatico.
(Photo: Carlos Pineda)

fertilizer and green manure) between 2002 and 2018 (El Hatico, unpublished yield
records). Conventional producers currently use 180–220 kg of synthetic N
ha1 year1, compared to 80 kg ha1 year1 from poultry manure used at El Hatico.
Crop residues, green manure crops and free-living N-fixers such as soil bacteria
provide approximately 80 kg of additional nitrogen at El Hatico (Cenicaña,
unpublished data); however, this nitrogen is more difficult to quantify.
Between 2001 and 2018, El Hatico’s agroecological sugarcane consistently
outperformed its conventional counterpart (average yield: 9.99 vs. 8.58 tons of
sugarcane ha1 month1, respectively). Even more important than the higher yield
is the fact that agroecological sugarcane behaves as a long-lived perennial crop.
Several plots at El Hatico have been harvested 18 times between 2001 and 2020
without a decline in yield (El Hatico, unpublished yield records); in contrast, the
conventional counterparts are being replaced after only five harvests due to decreas-
ing productivity (Cenicaña 2001–2018, annual reports). This frequent replanting of
sugarcane has high financial and environmental costs.
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 291

11.4.2 Environmental Impacts of Conventional

and Agroecological Sugarcane

In conventional sugarcane, herbicides are used periodically to eliminate grasses and

other weeds in plots and alleys between plots (which occupy 10% of the land in
plantations). El Hatico solved the problem of weed control by integrating African
hair sheep. This makes sense, both financially and environmentally. Sheep use
marginal areas that produce a high biomass of grasses, legumes and other species.
Grazing replaces costly and unsustainable management practices such as the appli-
cation of herbicides and mechanic weed control in sugarcane alleys and plantations.
In addition to making marginal areas productive, sheep reduce the cost of weed
control by 35% (Molina et al. 2013, 2014). Each hectare planted with sugarcane
supports two to three adult hair sheep, transforming weeds into high-quality meat
and manure that can be used as organic fertilizer (Molina et al. 2013, 2014). Some
manure remains in the crop and another part is collected in enclosures (750–1000 g
are collected daily from each 30–40 kg animal) (Fig. 11.6).
Agroecological management of sugarcane enhances soil biological activity
(Sadeghian and Madriñan 2000). Data summarized in Table 11.3 show that
populations of beneficial soil fungi and bacteria differ significantly between con-
ventional and agroecological sugarcane plantations at different depths in the soil
profile (Manrique et al. 2006).
Pardo (2009) found dramatic differences in macroinvertebrate biomass related to
the type of management (agroecological at El Hatico vs. conventional in a nearby
property) and between the moist and dry seasons. Macroinvertebrate biomass was
15.3 times higher in agroecological than in conventional sugarcane during the moist
season (26.44 vs. 1.73 g per 0.75 m3 of soil), and 59% higher during the dry season
(6.56 vs. 4.11 g per 0.75 m3 of soil). These differences were largely driven by
variations in the biomass of earthworms and millipedes during the moist season, and
earthworms during the dry season (Pardo 2009; Pardo et al. 2017).
Since 1994 soil organic matter (SOM) has increased from 2% to 4% at El Hatico’s
agroecological sugarcane, which means that it has almost recovered the forest
reference value (4.2%). With the increase in SOM, the use of irrigation water has
declined from 10,000 to 6000 m3 ha1 cycle1 at El Hatico (a cycle at El Hatico is
12 months). This water economy has huge economic and environmental implica-
tions. The doubling of SOM in less than three decades highlights the climate change
mitigation potential of agroecological sugarcane. If the whole region transitions to
sustainable practices, sugarcane producers will be able to sequester large amounts of
carbon, save precious water resources and cut down production costs while increas-
ing yields significantly. Scaling-up agroecological practices in sugarcane would
enhance ecosystem services and revitalize the regional economy.
292 Zoraida Calle D et al.

Fig. 11.6 African hair sheep under a line of endangered Caesalpinia ebano trees. (Photo: Zoraida
Calle)

11.5 Biodiversity Studies at El Hatico: Arthropods

and Birds
11.5.1 Ants

Ants are considered good indicators of biodiversity, disturbance, ecological succes-

sion and ecosystem rehabilitation (Dominguez-Haydar and Armbrecht 2011).
Armbrecht and Chacón (1997, 1999) studied ant species richness and diversity in
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 293

Table 11.3 Populations of Beneficial fungia Conventional Agroecological

beneficial fungi and soil bac-
0–5 cm 37 88
teria at different depths in the
soil profile in Hatico’s agro- 5–10 cm 30 60
ecological sugarcane planta- 10–20 cm 23 37
tions and reference 20–40 cm 10 30
conventional plantations >40 cm 7 25
Beneficial bacteriab Conventional Agroecological
0–5 cm 400 580
5–10 cm 280 500
10–20 cm 250 480
20–40 cm 155 400
>40 cm 50 100
Source: Manrique et al. (2006)
a
Colony forming units g1  104
b
Colony forming units g1  106

the seven largest forest fragments of the Cauca river valley and their surrounding
agricultural matrixes, including El Hatico as one of their study sites. They found a
total of 137 morphospecies, grouped into 37 genera and 6 subfamilies; 90% of the
ant species were captured in forest fragments and 54% in their surrounding matrixes.
They also found a significant correlation between ant species richness in the forest
fragments and their matrixes (Armbrecht and Chacón 1999).
In these studies, the highest species richness and diversity values were found at El
Hatico: a total of 81 ant species (66 in the forest and 35 in the surrounding
silvopastoral matrix). El Hatico also had the highest number of exclusive ant species
(ants that were not found in any other study site) in the agricultural matrix (6).
Interestingly, the diversity index for El Hatico’s silvopastoral systems (H0 ¼ 3.07)
was higher than the diversity indexes of four of the seven forest fragments (H0
between 1.96 and 2.87) (Armbrecht and Chacón 1999). The studied agricultural
matrixes were on average 7.7
C warmer than the forest fragments; however, El
Hatico was only 3.7
C warmer. El Hatico’s silvopastures formed the coolest matrix,
with an average temperature of 30.1
C (Armbrecht 1995).
In another study conducted in dry forest fragments in this same region Armbrecht
et al. (2001) found that the assemblages of ant species in dry forest fragments were
not simple subsets of the regional pool of species, but rather, that each fragment
preserved an assembly with unique elements. The loss of any one of these small
forest patches would cause the disappearance of a fraction of the regional ant
diversity. This study highlights the importance of every remaining forest fragment
for the long-term viability of the populations of several ant species (Armbrecht et al.
2001).
294 Zoraida Calle D et al.

11.5.2 Spiders

Spiders (Aranae order) are considered as appropriate models for studies of commu-
nity structure, composition and dynamics because they are diverse and abundant in
terrestrial ecosystems, and their communities are affected by habitat, land use,
vegetation structure and plant species composition (Pearce and Venier 2006). In
tropical ecosystems, structurally complex vegetation tends to support diverse spider
assemblages (Baldissera et al. 2012).
Delgado et al. (2014) studied the species composition and diversity of spiders at
El Hatico in 1-ha plots located in a silvopastoral system, a forest fragment and
agroecological sugarcane. They collected 3635 adult spiders, belonging to
156 morphospecies and 30 families; these species represent approximately 75% of
El Hatico’s total estimated spider fauna (Delgado et al. 2014). The silvopastoral
system plot had the highest number of spider species (74), followed by the forest
(71 species), and the agroecological sugarcane plot (46 species). Average similarity
between land uses in the composition of spider assemblages was 45.4%; this
suggests a high beta diversity or species turnover between land uses at El Hatico.
The forest fragment and the silvopastoral system shared 54.4% of spider species
(Delgado et al. 2014).
In the study conducted at El Hatico, the silvopastoral system was the most diverse
habitat type, while the forest fragment was the least diverse one, given that half of the
collected spiders belonged to a single species (Leucauge sp.). El Hatico’s spider
fauna found in this research included eight different guilds; this ecological diversity
shows that different land uses provide a variety of microhabitats and resources for
spiders with different hunting strategies. A high diversity of spiders suggests the
presence of diverse prey species at El Hatico.

11.5.3 Parasitoid Wasps

Parasitoid wasps form a large group of hymenopterans that lay their eggs on
(or inside) the bodies of other arthropods, causing the slow death of their hosts.
Different parasitoid groups specialize in hosts from different insect orders. López
et al. (2013) studied the diversity of parasitoid wasps in four silvopasture plots and
the forest fragment at El Hatico. They collected 1376 parasitoids belonging to
7 super-families, 18 families and 42 morphospecies.
Vegetation structure and plant species composition in different land uses at El
Hatico were clearly related to the abundance of parasitoids. These small wasps were
more abundant in the silvopastures (319–364 individuals captured in traps through-
out the study) than in the forest fragment (11 captured individuals). Species richness
was also higher in silvopastures (22–26 species) compared to the forest fragment
(11 species). However, the diversity index (H0 ) was higher at the forest (H0 ¼ 2)
compared to the silvopastures (H0 values from 1 to 1.22), as a result of the higher
equitability index (0.572 in the forest and 0.281–0.397 in the silvopastures).
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 295

The abundance of parasitoid wasps was correlated with plant species richness.
The diversity of parasitoids in silvopastures is related to shade and the diversity of
complementary food resources (nectar and pollen), including common weeds such
as Lantana camara (Verbenaceae), Parthenium hysterophorus (Asteraceae) and
Sida acuta (Malvaceae); parasitoids require nectar and pollen apart from insect prey.
The parasitioid wasp fauna identified at El Hatico includes species that have been
released in the farm since the 1960s as biological control agents for pest species such
Diatraea saccharalis (Pyralidae) in sugarcane and Spodoptera sp. (Noctuidae) in
corn and sorghum (López et al. 2013). Apparently, these commercial biological
control agents have established populations at El Hatico’s silvopastures, which
probably enhances biological pest control.

11.5.4 Birds

El Hatico’s bird fauna is outstanding. In a one-year study based on periodic bird

censuses, Cárdenas (1998) observed a total of 135 species, belonging to 39 families
and 17 orders. Two thirds of the birds (89 species) used agroecosystem habitats
during the censuses. Tyrannidae was the richest bird family with 19 species (14%),
followed by Fringillidae (11 species, 8%) and Thraupidae (8 species, 6%); 30 fam-
ilies were represented by 4 species or less. Waterfowl richness and the presence of
species belonging to specialized taxonomic groups were also noteworthy. A total of
51 bird species showed evidence of reproductive activity. The bird list compiled by
Cárdenas (1998) includes nine species that had not been reported previously in the
Valle del Cauca and that expand the latitudinal distribution described by Hilty and
Brown (1986) for the birds of Colombia.
This study found a considerable bird species richness in silvopastoral systems:
57 species in a system with fruit trees, 46 in an ISS with Leucaena leucocephala
shrubs and 43 in a star grass silvopasture. The forest and the agroecological
sugarcane had 33 bird species each. The lowest bird species richness was observed
in the bamboo forest with 29 species and in a conventional sugarcane (outside of El
Hatico) with 19 species (Cárdenas 1998).
Diversity indices (H0 ) were consistent with those of species richness: 3.21 for the
silvopasture with fruit trees, 3.07 for ISS with L. leucocephala, 2.98 for the
silvopasture with star grass, 2.86 for the forest, 2.73 for the bamboo forest, 2.43 for
the agroecological sugarcane and 1.53 for the conventional sugarcane (Cárdenas
1998). Bird diversity and richness in the agroecological sugarcane are likely related
to the proximity of the forest fragment and the bamboo forest, as well as the presence
of live fences and tree-lined alleys that enhance structural and floristic diversity.
In a more recent but shorter study of El Hatico’s bird fauna, Hurtado-G et al.
(2016) observed a total of 109 bird species belonging to 37 families and 16 orders.
They observed the highest species richness in the agroecological sugarcane system
(57), followed by the fruit orchards (50), forest and silvopastoral systems (43 each).
Conventional sugarcane plantations outside of El Hatico had 40 bird species
(Hurtado-G et al. 2016).
296 Zoraida Calle D et al.

Of the bird species observed by Hurtado-G et al. (2016), 9.2% were migratory.
The forest and agroecological sugarcane had the highest number of migratory birds
(4 species each), followed by the silvopastoral system (3 species), conventional
sugarcane (2) and fruit trees (1).
Migratory birds are not restricted to pristine areas in the tropics; instead, they use
agroecosystems during their winter residence. Trees in crops or paddocks favor the
arrival and permanence of Nearctic-Neotropical migratory species by providing
perches, shelter, foraging substrates and corridors (Rice and Greenberg 2004).
Hurtado-G et al. (2016) also studied the diet of migratory bird species at El
Hatico. Food fragments obtained from fecal samples of migratory birds included
mostly arthropods (99%) from the orders Coleoptera (64%), Hymenoptera (18%),
Araneae (9%), Hemiptera (5%), Diptera (1.6%), Lepidoptera (1.6%), Acari (0.4%)
and Psocoptera (0.4%). Most migratory bird species fed on similar items, although in
different proportions related to their foraging habits. Beetles were important com-
ponents of the diet of most migratory birds, varying from 28% for Catharus
ustulatus to 72% for Setophaga petechia. Consumption of Hemiptera and spiders
was common among all species except for Hirundo rustica; this bird consumed a
high proportion of Hymenoptera (40%). Hurtado’s results confirm that migratory
birds that visit El Hatico each year are mostly insectivorous.

11.6 Functional Biodiversity: Natural Enemies of Pests

The term functional biodiversity includes the value and range of species and
organismal traits that influence ecosystem functioning (Tilman 2001). Here we use
the term in the narrower agroecological sense, referring to species with positive
effects on agroecosystems such as natural enemies of pests.
Many organisms that behave as pests in conventional cattle ranching and agri-
culture are controlled naturally by different species at El Hatico without external
inputs or energy. Some examples include:
• Birds such as the cattle egret (Bubulcus ibis), the yellow-headed caracara
(Milvago chimachima) and the smooth-billed ani (Crotophaga ani) contribute
to the integrated management of ticks (Rhipicephalus microplus) and other
ectoparasites of cattle.
• The ant Ectatomma ruidum, the entomopathogenic fungus Nomuraea rileyi and
minute wasps (Trichogramma sp.) that are endoparasitoids of insect eggs control
periodic outbreaks of the lepidopteran Azeta versicolor, which completely defo-
liates Gliricidia sepium trees (Gómez et al. 2002).
• The cattle egret, together with spiders and entomopathogenic fungi, control two
important pests of sugarcane, the fall armyworm (Spodoptera frugiperda) and the
small mocis moth (Mocis lapites), rendering insecticides unnecessary in agro-
ecological sugarcane (CH Molina, personal observations).
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 297

• Bacillus thuringiensis, Trichogramma sp. and paper wasps (Polistes

erythrocephalus, Vespidae) controlled an outbreak of a defoliator worm
(Noctuidae) that reduced the available biomass of Leucaena leucocephala by
30% in El Hatico’s ISS during the El Niño drought in 2008–2010 (Montoya et al.
2010).
• Lacewings (Chrysopidae) and at least 10 parasitoid wasps from 8 different
families control the microlepidopteran Eccopsis galapagana, which defoliates
Prosopis juliflora trees in El Hatico’s ISS (Reyes et al. 2012).
• The smooth-billed ani (Crotophaga ani), a common large bird in the cuckoo
family, controls outbreaks of Calligo illioneus butterflies, which lay their eggs on
the leaves of sugarcane (Gómez and Lastra 1998).
• Dung beetles such as Dichotomius belus, Ontophagus marginicollis and
Coprophanaeus jasius disrupt the biological cycle of flies by burying manure,
thus controlling populations of the stable fly Stomoxys calcitrans and the horn fly
Haematobia irritans, both of which suck blood from cattle (Giraldo et al. 2018b).
• Ants such as Ectatomma ruidum, Crematogaster sp., Nylanderia fulva, Azteca
sp. and Dolichoderus sp., wasps (Polistes erytrociphalus), together with spiders
and birds such as Theristicus caudatus, Vanellus chilensis, Crotophaga ani,
Milvago chimachima and Bubulcus ibis are predators of leafcutter ants (consid-
ered the most problematic herbivores in the Neotropics) at El Hatico. Arthropods
and birds that control winged leafcutter ants prevent the proliferation of new
colonies (Castaño et al. 2019).
Diverse birds, arthropods (including parasitoids) and fungi play important produc-
tive roles at El Hatico as they help avoid the economic and environmental costs of
pest control in animal and plant-based systems. As seen, predation by birds, together
with entomopathogenic fungi and management practices facilitate the production of
organic milk by helping avoid the use of toxic substances for tick control (Giraldo
and Uribe 2007). It is remarkable that judicious management and associated biodi-
versity control pests and weeds in agroecological sugarcane, thus making the use of
toxic agrochemicals unnecessary. Hopefully, farmers in the region and beyond will
try to emulate these examples, as discussed in the following sections.

11.7 El Hatico’s Influence on Society at Different Scales

El Hatico has strong a commitment to the generation of knowledge on sustainable

agriculture and livestock production. Technical knowledge is shared openly with
research centers, networks, universities, farmers and decision makers; a total of
13,732 people from 815 institutions and 59 countries visited El Hatico between
2005 and 2016 (El Hatico, unpublished records). This section describes the scales of
influence of El Hatico, demonstrating how a single rural property can make a
difference for biodiversity and sustainability at a regional scale and beyond.
298 Zoraida Calle D et al.

11.7.1 Promoting Sustainable Cattle Ranching

The local or landscape-scale impacts of sustainable ranching practices developed at

El Hatico have been modest because cattle grazing is no longer an important
economic activity in the geographic Cauca river valley. However, El Hatico has
become a national and international reference for sustainable cattle ranching. Field
visits of ranchers, extensionists and decision makers have been instrumental to
overcome skepticism about tree-based ranching practices (Calle et al. 2012, 2013).
El Hatico played a key role in convincing the Colombian government and the
national cattle ranchers association (FEDEGAN) about the strategic value of
silvopastoral practices. Extensionists and other members of the technical team of
the Colombian Sustainable Cattle Ranching Project2 were trained at El Hatico and
later influenced 4200 farms (95,000 ha) in five regions of Colombia (Giraldo et al.
2018a). In 2011, El Hatico received FEDEGAN’s national sustainable livestock
award.
Ten Colombian universities have integrated field visits and scientific knowledge
developed at El Hatico into their academic programs of agronomy, environmental
studies, animal science, veterinary medicine and biology. Different research groups
have studied the productivity and ecosystem services of silvopastoral systems at El
Hatico (Chará et al. 2015).
Through a strategic alliance with the Celema dairy plant in Manizales, El Hatico
and Lucerna farm developed Colombia’s first organic ultra-pasteurized or extended
shelf-life milk and a variety of certified organic dairy products that have penetrated
national markets.

11.7.2 The Expansion of Agroecological Sugarcane

Agroecological sugarcane developed by El Hatico’s team has impacted land use

change at the local and landscape scales, and has inspired a large number of
international producers. The agroecological protocol developed for the farm’s
100 ha of sugarcane has influenced the regional sugar sector by guiding producers
in the adoption of sustainable practices. More than 20,000 ha of sugarcane that
supply eight sugar mills have shifted to organic production in the Valle del Cauca
and more than 20% of the sugarcane in the lower Amaime river basin (surrounding

2
The Colombian Sustainable Cattle Ranching Project (CSCRP) was designed by an alliance
between the Global Environment Fund (GEF), the UK government, FEDEGAN, The Nature
Conservancy (TNC), CIPAV and Fondo Acción, under the supervision of The World Bank. It
took place from 2010 to 2020 in five ecoregions where cattle ranching exists close to protected
areas, and aimed to overcome the main barriers to the adoption of sustainable practices.
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 299

El Hatico), is now organic (Murgueitio 2019). Agroecological and biodiversity-

friendly farming practices have entered the vocabulary of the regional sugar industry
and have influenced Cenicaña’s research agenda (Torres 2006; Hincapié et al. 2019).
El Hatico and its closely related farm Lucerna inspired the adoption of more
efficient practices for brown sugar production in Mexico. Sugar mills from El
Salvador, Brazil and Argentina began the transition to organic sugarcane after
their teams visited El Hatico. The Better Sugarcane Initiative of Bonsucro (global
sugarcane platform, [Link]) integrates technical knowledge developed
at El Hatico. Organic sugar from El Hatico and Lucerna is exported to Europe, Asia
and North America.

11.7.3 Rural Sustainability Values and Generational

Exchange

El Hatico is a founding member of Resnatur,3 the Colombian Network of Private

Nature Reserves, which currently joins 168 farm-scale conservation initiatives that
preserve 57,296 ha of natural ecosystems in the five regions of the country. El Hatico
influenced Resnatur’s vision of biodiversity conservation integrated to sustainable
agriculture and livestock production. Currently, the Colombian System of Protected
Areas (SINAP, [Link]
de-areas-protegidas-sinap/) recognizes this conservation strategy and its philosophy
of sustainable production.
El Hatico’s vision of integrated livestock production, agriculture and forestry,
rooted in the principles of agroecology and ecological restoration, has reached
national and regional audiences through ELTI,4 CIPAV5 and SOCLA’s6 field
courses on Agroecology and Restoration. Case studies of El Hatico are available
for global audiences through ELTI’s Online Program. Since 1986, training events
offered by CIPAV and El Hatico have inspired thousands of alumni from 40 coun-
tries to undertake sustainable livestock production and agriculture projects.

3
[Link]
4
Environmental Leadership & Training Initiative at Yale University’s School of the Environment, a
program dedicated to capacity building for forest and landscape restoration in the tropics.
5
Center for Research on Sustainable Agricultural Production Systems ([Link]), a
Colombian organization dedicated to research, training and outreach on sustainable agriculture
and livestock production, ecosystem services, water-based systems and ecological restoration.
6
The Latin American Scientific Society for Agroecology ([Link])
300 Zoraida Calle D et al.

11.8 Influencing Policy

Knowledge on sustainable cattle ranching generated at El Hatico has influenced

projects developed by The World Bank, FAO, IDB, The Nature Conservancy and
CIAT. Silvopastoral systems were included in Colombia’s public policy and the
current government’s development plan (DNP 2019). In addition, tree-based sus-
tainable ranching is part of Colombia’s strategy to meet the 20% reduction goal for
greenhouse gas emissions committed by the country for 2030 (DNP 2019).
In 2014, El Hatico hosted the meeting of the institutional members of the Global
Agenda for Sustainable Livestock.7 This was a unique opportunity for livestock
specialists from around the world to closely analyze El Hatico’s silvopastoral
systems. One effect of that meeting was the creation of the Global Network on
Silvopastoral Systems that brings together 107 members from 50 institutions and
29 countries. This multi-stakeholder partnership works to strengthen and scale-up
silvopastoral systems worldwide, by generating, exchanging and disseminating
knowledge, documenting public policy and facilitating dialogue to address the
challenges related to the sustainable development goals (https://
[Link]/). A case study of silvopastoral systems developed
by FAO, CIPAV and Agri Benchmark (Germany) analyzes natural resource use
efficiency, land productivity, economic performance and environmental benefits of
ten silvopastoral production models in Colombia (including El Hatico’s ISS), Mex-
ico and Argentina (Chará et al. 2019). This document provides policy recommen-
dations for promoting and scaling-up silvopastoral systems in Latin America and
other regions.

11.9 Conclusions

El Hatico has played a pivotal role in spreading two ideas that are key for
implementing sustainable farming systems. First, farm-scale biodiversity conserva-
tion should go beyond the protection of natural ecosystem remnants to include
productive areas. Second, the redesign of farming systems based on the principles
of agroecology can result in higher yields, better quality of agricultural products,
lower production costs and multiple ecosystem services, without known trade-offs.
Such changes can take place when landowners have a strong intergenerational

7
The Global Agenda for Sustainable Livestock, established in 2011, is a multi-stakeholder part-
nership with the aim of fostering and guiding the sustainable development of the global livestock
sector in alignment with the SDG framework of the UN Agenda 2030. It provides a platform to
address comprehensively the sector’s multiple challenges towards sustainable development by
facilitating global dialogue and encouraging local practice and policy change, focusing on innova-
tion, capacity building, incentive systems and enabling environments ([Link].
com)
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 301

commitment to the land, motivation to innovate and improve their farming systems,
and a desire to share knowledge with peers, scientists and decision makers.
Biodiversity conservation in fertile productive landscapes, such as the geographic
Cauca river valley, should include the agricultural matrix with actions that enhance
habitat for native species and facilitate wildlife movements. Even small forest
fragments such as the one protected at El Hatico can contribute to regional conser-
vation if surrounded by wildlife-friendly agroecosystems.
Synergies between sustainable livestock production, agroecology and ecological
restoration can allow rural properties to increase their profitability and productivity,
while protecting forests and integrating native trees into cattle grazing areas and
agriculture. Innovative and efficient farming systems such as those found at El
Hatico can inspire positive change at local, regional and global scales by motivating
producers to enhance thousands of hectares. Mahatma Gandhi wrote that “a small
body of determined spirits fired by an unquenchable faith in their mission can alter
the course of history”. El Hatico inspires people to believe that the sum of individual
decisions in the right direction has the power to transform whole regions.

References

Arcila AM, Valderrama C, Chacón P (2012) Estado de fragmentación del bosque seco en la cuenca
alta del río Cauca, Colombia. Biota Colombiana 13(2):86–101
Arias J (1994) Evaluación exploratoria del comportamiento de la materia orgánica por prácticas de
quema en molisoles del Valle del Cauca. Tesis de Posgrado en Suelos y Aguas. Facultad de
Ciencias Agropecuarias, Universidad Nacional de Colombia, Palmira
Armbrecht I (1995) Comparación de la mirmecofauna en fragmentos boscosos del Valle Geográfico
del río Cauca, Colombia. Bol Mus Ent Univ Valle 3(2):1–14
Armbrecht I, Chacón P (1997) Composición y diversidad de hormigas en bosques secos relictuales
y sus alrededores en el Valle del Cauca, Colombia. Revista Colombiana de Entomología
23(1–2):45–50
Armbrecht I, Chacón P (1999) Rareza y diversidad de hormigas en fragmentos de bosque seco
colombianos y sus matrices. Biotropica 31(4):646–653
Armbrecht I, Tischer I, Chacón P (2001) Nested subsets and partition patterns in ant assemblages
(Hymenoptera, Formicidae) of Colombian dry forest fragments. Pan Pac Entomol 77(3):
196–209
Baldissera R, Rodriguez E, Hartz S (2012) Metacommunity composition of web spiders in a
fragmented neotropical forest: relative importance of environmental and special effects. PLoS
One 7(10):1–9
Calle Z, Murgueitio E, Chará J (2012) Integrating forestry, sustainable cattle-ranching and land-
scape restoration. Unasylva 239(63):31–40
Calle Z, Murgueitio E, Chará E, Molina CH, Zuluaga AF, Calle A (2013) A strategy for scaling-up
intensive silvopastoral systems in Colombia. J Sustain For 32(7):677–693. [Link]
1080/10549811.2013.817338
Cárdenas G (1998) Comparación de la composición y estructura de la avifauna en diferentes
sistemas de producción. BS Thesis, Universidad del Valle, Cali
Castaño K, Giraldo C, Rodríguez J, Giraldo NV, Calle Z, Reyes K, Molina CH, Molina EJ, Molina
JJ, Armbrecht I, Montoya J (2019) Aspectos ecológicos, enemigos naturales y métodos de
control de la hormiga arriera Atta cephalotes (Hymenoptera: Myrmicinae) en la Reserva Natural
302 Zoraida Calle D et al.

El Hatico. In: Castaño K, Chará J, Giraldo C, Calle Z (eds) Manejo integrado de insectos
herbívoros en sistemas ganaderos sostenibles. CIPAV, Cali, pp 150–179
Cenicaña (2001–2018) Informes anuales. Cenicaña, Cali. [Link]
de-cenicana/
Cenicaña (2017) Manual de reconocimiento de arvenses en el cultivo de la caña de azúcar.
Cenicaña. 168 p [Link]
arvenses/manual_reconocimiento_arvenses.pdf
Chará J, Camargo JC, Calle Z, Bueno L, Murgueitio E, Arias L, Dossman M, Molina EJ (2015)
Servicios ambientales de sistemas silvopastoriles intensivos: mejoramiento del suelo y
restauración ecológica. In: Montagnini F, Somarriba E, Murgueitio E, Fassola H, Eibl B (eds)
Sistemas agroforestales: funciones productivas, socioeconómicas y ambientales, Serie Técnica,
Informe Técnico 402. CATIE/Editorial CIPAV, Turrialba/Cali, 454 p
Chará JD, Rivera J, Barahona R, Murgueitio E, Deblitz C, Reyes E, Martins Mauricio R, Molina JJ,
Flores M, Zuluaga AF (2017) Intensive silvopastoral systems: economics and contribution to
climate change mitigation and public policies. In: Montagnini F (ed) Integrating landscapes:
agroforestry for biodiversity conservation and food sovereignty, advances in agroforestry 12.
Springer, Cham, pp 395–416
Chará J, Reyes E, Peri P, Otte J, Arce E, Schneider F (2019) Silvopastoral systems and their
contribution to improved resource use and sustainable development goals: evidence from Latin
America. FAO, CIPAV and Agri Benchmark. Cali, 58 p. License CC BY-NC-SA 3.0
IGO. ISBN: 978-958-9386-84-2 [Link]
Clegg CD, Attard E, Degrange V, Klumpp K, Richaume A, Soussana JF, Le Roux X (2006) Impact
of cattle grazing and inorganic fertiliser additions to managed grasslands on the microbial
community composition of soils. Soil Biol Biochem 31:73–82
Corporación Autónoma Regional del Valle del Cauca (CVC) (1990) Comparación de cobertura de
bosques y humedales entre 1957 y 1986 con delimitación de las comunidades naturales críticas
en el valle geográfico del río Cauca. CVC Report 90–7. Cali, 84 p
Delgado J, Armbrech I, Flórez E, Molina CH (2014) Arañas (Arachnida: Araneae) asociadas a
cuatro tipos de manejo del hábitat en la Reserva Natural El Hatico (Colombia). Revista Ibérica
d e A r a c n o l o g í a 2 5 : 5 9 –6 9 . h t t p : / / g i a . s e a - s oc i o s .c o m / PD F/ 2 5/ 0 59 - 0 6 9 RI A 2
[Link]
Departamento Nacional de Planeación (2019) Plan Nacional de Desarrollo 2018–2022: Pacto por
Colombia, pacto por la equidad. Bogotá, Colombia, 477 p
Dominguez-Haydar Y, Armbrecht I (2011) Response of ants and their seed removal in rehabilita-
tion areas and forests at El Cerrejón coal mine in Colombia. Restor Ecol 19:178–184. [Link]
org/10.1111/j.1526-100X.2010.00735.x
Eibl B, Montagnini F, López MA, Lavignolle P, Cortes J, De La Vega M, López LN, Dummel C,
Küppers G (2022) Conservation, connectivity and registration of seed areas in remnants of
natural reserves in the province of Misiones, Argentina. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human- dominated environments. Topics in Biodiversity
and Conservation, Springer, Cham, pp 157–181
Giraldo C, Uribe F (2007) Manejo integrado de garrapatas en sistemas sostenibles de producción
ganadera. Red Electrónica de Garrapatas y Enfermedades Transmitidas por Garrapatas en
América Latina y El Caribe. FAO, Rome
Giraldo C, Chará J, Uribe F, Gómez JC, Gómez M, Calle Z, Valencia LM, Modesto M, Murgueitio
E (2018a) Ganadería Colombiana Sostenible: entre la productividad y la conservación de la
biodiversidad. In: Halffter G, Cruz M, Huerta C (eds) Ganadería sustentable en el Golfo de
México. Instituto de Ecología, A.C., Xalapa, pp 31–61
Giraldo C, Montoya S, Escobar F (2018b) Escarabajos del estiércol en paisajes ganaderos de
Colombia. CIPAV, Cali
Gómez LA, Lastra LA (1998) Caligo illioneus: el gusano cabrito. Carta Trimestral Cenicaña 20(1):
8–13. [Link]
11 A Highly Productive Biodiversity Island Within a Monoculture Landscape:. . . 303

Gómez LA, Vargas G (2014) Los barrenadores de la caña de azúcar, Diatraea spp., en el valle del
río Cauca: investigación participativa con énfasis en control biológico. Documento de Trabajo
734, Cenicaña, Cali
Gómez ME, Rodríguez L, Murgueitio E, Ríos CI, Rosales M, Molina CH, Molina CH, Molina EJ,
Molina JP (2002) Árboles y arbustos utilizados en alimentación animal como fuente proteica.
Editorial CIPAV, Cali
Hilty SL, Brown WL (1986) A guide to the birds of Colombia. Princeton University Press,
Princeton
Hincapié E, Sánchez J, Carbonell JA (2019) IoT network applied to agriculture: monitoring stations
for irrigation management in soils cultivated with sugarcane. In: Corrales J, Angelov P, Iglesias
J (eds) Advances in information for communication technologies for adapting agriculture to
climate change II, AACC 2018. Advances in intelligent systems and computing, vol 893.
Springer, pp 249–259. [Link]
Hurtado-G A, Cruz-B L, Molina-D EJ (2016) Dieta de aves migratorias en un sistema
agroecológico del Valle del Cauca, Colombia. Bol Cient Mus Hist Nat U de Caldas 20(2):
151–163. [Link]
López L, Armbrecht I, Montoya-Lerma J, Molina EJ (2013) Diversidad de avispas parasitoides en
un sistema silvopastoril orgánico de producción ganadera de Colombia. Revista Avances en
Investigación Agropecuaria 17(1):65–78. [Link]
Mahecha L (2003) Importancia de los sistemas silvopastoriles y principales limitantes para su
implementación en la ganadería colombiana. Rev Col Cienc Pec 16:11–18
Manrique GM, Guzmán ML, Victoria JI (2006) Flora microbiana en suelos cultivados con caña de
azúcar y otros sistemas de exploración. In: Torres JS (ed) Manejo del cultivo en condiciones de
caña verde, Serie Técnica 35. Cenicaña, pp 54–63. [Link]
serie_tecnica/st_35/st_35.pdf
Molina EJ, Molina CH, Molina JJ, Suárez JF, Uribe P, Valencia LM, Murgueitio E, Uribe F (2013)
Ovinos y caña de azúcar. Revista DeCarne 7:55–59
Molina EJ, Molina CH, Molina JJ, Suárez JF, Uribe P, Valencia LM, Murgueitio E, Uribe F (2014)
Integración con beneficios. Revista DeCarne 8:48–51
Molina IC, Donney G, Montoya S, Rivera JE, Villegas G, Chará J, Barahona R (2015) La inclusión
de Leucaena leucocephala reduce la producción de metano de terneras Lucerna alimentadas con
Cynodon plectostachyus y Megathyrsus maximus. Livestock Research for Rural Development
27, Article 96. Retrieved June 12, 2020, [Link]
Molina IC, Angarita EA, Mayorga OL, Chará J, Barahona R (2016) Effect of Leucaena
leucocephala on methane production of Lucerna heifers fed a diet based on Cynodon
plectostachyus. Livest Sci 185:24–29
Molina-Castro CH, Molina-D CH, Molina-D EJ, Molina-E JJ (2012) Manejo agroecológico de caña
de azúcar y sistemas silvopastoriles intensivos. Revista Tecnicaña 29:28–35. [Link]
revistatecnicana/docs/tec_no29_2012/37
Molina-Garcés C (1938). Árboles para sombrío y forraje. Revista de la Academia Colombiana de
las Ciencias Exactas, Físicas y Naturales 2(6): 273–278
Montes-Londoño I, Calle A, Montes-Calderon A, Montes-Botero O (2022) Hacienda Pinzacuá: an
island of regenerative agriculture in the piedmont of the Colombian Central Andes. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human-dominated envi-
ronments. Topics in Biodiversity and Conservation, Springer, Cham, pp 305–355
Montoya J, Calle Z, Murgueitio E (2010) Biodiversidad funcional como reguladora de insectos
plaga en los sistemas silvopastoriles durante la sequía del fenómeno del niño 2009–2010. In
Memorias del VI Congreso Latonoamericano de Agroforestería para la Producción Pecuaria
Sostenible, Panama, Sept 28–30, 2010. CATIE/CIPAV. Turrialba/Costa Rica, pp 29–30
Murgueitio B (2019) Análisis del cambio de la cobertura del suelo en la zona baja de la cuenca del
río Amaime, periodo 1986–2018. Colegio La Arboleda, Cali, 138 p
Murgueitio E, Calle Z, Uribe F, Calle A, Solorio B (2011) Native trees and shrubs for the productive
rehabilitation of tropical cattle ranching lands. For Ecol Manag 261:1654–1663
304 Zoraida Calle D et al.

Murgueitio E, Xóchitl M, Calle Z, Chará J, Barahona R, Molina CH, Uribe F (2015) Productividad
en sistemas silvopastoriles intensivos en América Latina. In: Montagnini F, Somarriba E,
Murgueitio E, Fassola H, Eibl B (eds) Sistemas agroforestales: funciones productivas,
socioeconómicas y ambientales, Serie Técnica, Informe Técnico 402. CATIE/Editorial
CIPAV, Turrialba/Cali, pp 59–101. 454 p
Niella F, Rocha P, Tuzinkievicz A, Buchwies R, Bulman C, Thalmayr P, González J, Montagnini F
(2022) Native multipurpose species in the Yaboti Biosphere Reserve, Misiones-Argentina. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human-dominated envi-
ronments. Topics in Biodiversity and Conservation, Springer, Cham, pp 461–483
Pardo LC (2009) Macroinvertebrados edafícolas en sistemas agrícolas del municipio de El Cerrito
(Valle del Cauca), con énfasis en la comunidad de escarabajos Melolonthidae (Coleoptera:
Scarabaeoidea). Tesis de doctorado. Universidad del Valle, Cali
Pardo LC, Sevilla F, Amézquita E (2017) Macroinvertebrados edáficos en cultivos de caña de
azúcar, ganadería silvopastoril y un relicto forestal en el Valle del Río Cauca (Colombia).
Boletín de la Sociedad Entomológica Aragonesa 60:301–312
Pearce JL, Venier LA (2006) The use of ground beetles (Coleoptera: Carabidae) and spiders
(Araneae) as bioindicators of sustainable forest management: a review. Ecol Indic 6:780–793
Reyes K, Montoya J, Giraldo C, Brown JW (2012) Description of the early stages of Eccopsis
galapagana Razowski & Landry (Tortricidae), a defoliator of Prosopis juliflora (SW.)
DC. (Fabaceae) in Colombia. J Lepidopt Soc 66(3):156–164
Rice RA, Greenberg R (2004) Silvopastoral systems: ecological and socioeconomic benefits and
migratory bird conservation. In: Schroth G, Fonseca GAB, Harvey CA, Gascon C, Vasconcelos
A (eds) Agroforestry and biodiversity conservation in tropical landscapes. Island Press,
Washington, DC, pp 453–472
Rivera C, Naranjo L, Duque AM (2007) De María a un mar de caña: Imaginarios de naturaleza en la
transformación del paisaje vallecaucano entre 1950 y 1970. Universidad Autónoma de
Occidente, Cali
Rivera LF, Escobar F, Philpott SM, Armbrecht I (2019) The role of natural vegetation strips in
sugarcane monocultures: ant and bird functional diversity responses. Agric Ecosyst Environ
284:106603. [Link]
Sadeghian KH, Madriñan R (2000) La actividad microbiana-CO2 en suelos cultivados en caña de
azucar con y sin quema. Universidad Nacional de Colombia (Colombia)
Santos-Gally R, Boege K (2022) Biodiversity islands: the role of native tree islands within
silvopastoral systems in a neotropical region. In: Montagnini F (ed) Biodiversity islands:
strategies for conservation in human-dominated environments. Topics in Biodiversity and
Conservation, Springer, Cham, pp 117–138
Solis MA, Metz MA (2016) An illustrated guide to the identification of the known species of
Diatraea Guilding (Lepidoptera, Crambidae, Crambinae) based on genitalia. ZooKeys 565:73–
121
Tilman D (2001) Functional diversity. In: Levin SA (ed) Encyclopedia of biodiversity, vol
3. Academic Press, NewYork, pp 109–120
Torres JS (ed) (2006) Manejo del cultivo en condiciones de caña verde. Cenicaña, Serie Técnica
35 [Link]
Vallejo V, Roldán F, Dick RP (2010) Soil enzymatic activities and microbial biomass in an
integrated agroforestry chronosequence compared to monoculture and a native forest of Colom-
bia. Biol Fertil Soils 46(6):577–587
Vallejo V, Arbelia Z, Teránb W, Lorenzc N, Dickc RP, Roldan F (2012) Effect of land management
and Prosopis juliflora (Sw.) DC trees on soil microbial community and enzymatic activities in
intensive silvopastoral systems of Colombia. Agric Ecosyst Environ 150:139–148
Vargas GA, Obando V, Gómez LA (2006) Jaynesleskia jaynesi: otra alternativa para el manejo de
Diatrea spp. Carta Trimestral Cenicaña (Colombia) 28(2):3–5
Vargas G, Gómez LA, Michaud JP (2015) Sugarcane stem borers of the Colombian Cauca River
valley: pest status, biology and control. Fla Entomol 98:728–735
Chapter 12
Hacienda Pinzacuá: An Example
of Regenerative Agriculture Amidst
a Transformed Landscape in the Colombian
Andes

Irene Montes-Londoño, Alicia Calle, Olimpo Montes, and Arturo Montes

Abstract The central Andes of Colombia is a region of high biodiversity that has
been intensely transformed, mostly by unsustainable agricultural practices, and
especially by extensive cattle grazing. Whereas cattle farms are often considered
ecological deserts, cattle production can be approached in a different way: by
integrating more trees into the pastures, introducing better animal management
practices, and restoring protective forests, so that productivity, biodiversity, and
the flow of ecosystem services can be positively impacted. In this case study, we
describe the agroecological transformation of Hacienda Pinzacuá, a 45-hectare farm
that has become an island of regenerative agriculture amidst a highly fragmented
landscape. We explain how the farm’s land use history led to the severe degradation
of its once fertile soils; how key land management decisions were made and
gradually implemented through trial and error; and the significant land cover
changes that occurred over 20 years of transformation. We also provide data on
how these changes have impacted productivity, biodiversity, and ecosystem services
to illustrate how conservation and production can work synergistically to transform
the land and the people. Today Pinzacuá stands out as an island of vegetation in an
otherwise treeless landscape and has become a high-quality matrix that serves as
habitat or refuge for a variety of taxa striving to persist in this fragmented landscape.
Finally, we reflect on the challenges faced along the process, and the prospects for
maintaining Pinzacuá as both an island of biodiversity and an example for other
farmers seeking more resilient productive alternatives.

I. Montes-Londoño (*) · O. Montes · A. Montes

Hacienda Pinzacuá, Alcalá, Valle del Cauca, Colombia
Miami, FL, USA
A. Calle
Environmental Leadership & Training Initiative, Yale School of the Environment, New Haven,
CT, USA

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 305
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
306 I. Montes-Londoño et al.

Keywords Agroforestry · Biodiversity conservation · Degraded landscapes ·

Ecosystem services · Landscape restoration · Silvopastoral systems

12.1 Introduction

The Andean region of Colombia is part of the Northern Andes biodiversity hotspot,
an area known for its ecosystem diversity and high levels of endemism, and its
significance for global biodiversity (Mittermeier et al. 2011). The region is home to
34% of the country’s population—almost 17 million people—and generates 32% of
the national Gross Domestic Product (GDP) (Delgado and Pérez 2018). Human
activities such as agriculture, cattle ranching, urbanization, infrastructure develop-
ment, and species introductions have rapidly transformed this landscape, creating
enormous pressure on the remaining natural ecosystems, their biodiversity, and the
vital services they provide (Kattan and Alvarez-López 1996). Today, only 18–25%
of the original forest cover remains, mostly as fragmented remnants embedded in a
matrix of agricultural land uses (Etter 1998). As a result, Colombia’s Andean
ecosystems are among the most threatened in the world (Mittermeier et al. 2011),
and the human populations who inhabit them face increasing risks from the impacts
of land degradation.
Across Latin America, this pattern of ecosystem transformation has led to a
widespread biodiversity crisis that is further compounded by climate change. A
variety of nature-based solutions—actions to protect, restore and sustainably manage
both natural and transformed landscapes—have been identified that can contribute to
preserve biodiversity, combat climate change, and improve human well-being
(IUCN 2020). For example, improving the management of agricultural lands to
achieve a balance between a sustainably managed matrix, protected ecosystem
remnants, and high connectivity is critical to support biodiversity, sustain the flow
of ecosystem services, and enhance climate change resilience (Chazdon et al. 2009;
Perfecto and Vandermeer 2010; Vílchez et al. 2013; Griscom et al. 2017; Kremen
and Merenlender 2018).
Increasing tree cover through practices such as adding live fences, planting,
establishing or retaining trees in pastures, and using complex agroforestry and
silvopastoral systems can enhance the conservation value of agricultural landscapes
without compromising production (Perfecto and Vandermeer 2010; Harvey et al.
2011; Tscharntke et al. 2012; Mendenhall et al. 2014; Prevedello et al. 2018).
Although lack of access to technical assistance and financial incentives still limit
the widespread adoption of many such practices (Calle et al. 2013; Calle 2020), a
growing number of landowners are realizing the need to re-evaluate their conven-
tional practices and test alternative methods to work more closely with nature.
This chapter presents a case study examining the transformation of Pinzacuá, a
farm located in the central Andean region of Colombia and whose landowners,
having witnessed the degradation of their once fertile lands, decided to change
course. We describe the efforts made over 20 years to restore productivity in this
45-hectare farm by increasing tree cover and recovering soil health. We begin by
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 307

Fig. 12.1 Location of Pinzacuá in the La Vieja River watershed in Colombia’s Andean
region. Map: Alicia Calle

explaining the landscape context and land use history, then describe how gradual
changes were implemented, and the impacts of the process on biodiversity, farm
productivity, and ecosystem services. Finally, we reflect on the future of the farm
and the challenges and opportunities for others who may decide to tackle such an
effort.

12.2 Landscape and Social Context

Hacienda Pinzacuá is located on the western slope of Colombia’s Central Cordillera,

in the La Vieja river central watershed which marks the limits between the depart-
ments of Quindío and Valle del Cauca (Fig. 12.1). The farm is adjacent to Alcalá, a
town of 20,000 people located at 1290 m.a.s.l. Annual precipitation averages
1900 mm distributed bimodally from March to May and September to November;
average annual temperature is 23.4
C and relative humidity is 85% (Comisión
Conjunta 2008). The area’s original vegetation is typical of the transition between
premontane and low montane humid forest (Espinal 1977). However, only 11% of
the original forest cover remains, mostly in bamboo forests and unconnected frag-
ments <5 hectares (Camargo and Cardona 2005). Soils are Inceptisols and Andisols
of quaternary alluvial and volcanic origins respectively, which are young and deep
(20–30 m), ranging from sandy to clayey loam in texture, free of rock and acidic
(pH 4.5–5.5), and highly productive (Montes Londoño et al. 2017).
While the region has a strong agricultural tradition, the type of agriculture
practiced has changed over the past century in response to external drivers. The
original forests were first replaced by traditional coffee agroforests, and eventually
converted to single-crop coffee plantations of the more productive sun-loving
308 I. Montes-Londoño et al.

varieties. The collapse of coffee prices in the 1980s and 90s resulted in further
conversion, mostly to highly fertilized pastures for cattle ranching (Calle and
Piedrahíta 2007), disregarding the fact that the fragile soils and steep slopes are
unsuitable for grazing (Sadeghian et al. 2001). Today, the area is an agricultural
mosaic of cash crops (e.g., coffee, plantain, citrus) and cattle pastures, which occupy
33% of the land (DANE 2014). Pinzacuá is surrounded by undulating hills covered
by a mix of croplands with some isolated trees, monocultures of African grasses with
sparse tree cover, some live fences, small fragments of disturbed secondary forests
and riparian areas, and small bamboo forest patches.
Historically, this region enjoyed a strong economy and a high quality of life for
two main reasons. First, it is located at the center of the so-called “golden triangle”
formed by Bogotá, Medellín and Cali, Colombia’s main cities. Second, for most of
the twentieth century, this was the country’s prime coffee-growing region, and the
majority of its nearly four million inhabitants were dedicated to this activity. High
productivity and a strong international coffee market resulted in an economic boom
that allowed for the development of excellent infrastructure and services, and the
highest quality of life in the country (Toro Zuluaga 2005; Mejía Cubillos 2013). The
crash of the coffee markets in the 1990s led to a general economic contraction and
the emergence of new problems: unemployment, loss of income, unequal access to
essential public services, and increased levels of drug-related violence, all
compounded by the arrival of people displaced by the armed conflict that was taking
place elsewhere in the country (Toro Zuluaga 2005). Ultimately, the human devel-
opment index (HDI) stagnated for almost a decade and its gap with respect to the
national average widened (UNDP 2004).
The decline of coffee production gave way to new sectors such as telecommuni-
cations, services, tourism, construction and commerce that replaced agriculture.
Urban centers now concentrate most economic activities and the majority of the
population, while less than 15% of the people remain in rural areas (Mejía Cubillos
2013). Moreover, the farmer population is aging; more than 40% are between 40 and
54 years old (DANE 2014). The remaining farms are mostly privately owned, small
and medium properties between 5 and 200 hectares (UPRA 2019a, b, c). Smaller
farms are usually family-owned and subsistence-oriented, with a mix of cash crops
and livestock. Meanwhile, larger farms are largely monoculture-based commercial
operations. As farming declines, the price of land in this densely populated region
has skyrocketed: the price of an average hectare of cropland in 2014 was $4000
(Becerra et al. 2017a, b, c), but near urban or touristic areas it can be as high as
$40,000 (López Murillo 2015). High prices are forcing a land ownership transfor-
mation in this landscape where only large developers or consolidated farm owners
can afford the land, and the latter tend to stick to the conventional high-input-high-
output monoculture model.
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 309

12.3 Transforming Pinzacuá: An Incremental Process

12.3.1 History of a Transformation

Pinzacuá is a 45-hectare family-owned and operated business dedicated to raising

high-quality female Brangus cattle for breeding and beef production. The farm was
acquired in 1985, and its land use history since then can be divided in two distinct
periods: before and after 1997. A summary of the main events in the farm’s history is
provided in Table 12.1.
Before 1997, the farm followed the prevailing high-input, treeless pasture model
dictated by the dominant agricultural intensification trend of the 1960s, 70s and 80s.
Under this model, a succession of different crops was planted: first coffee, then
tobacco, and finally pasture. Despite the high yields, the farm was never profitable
because the prices of these commodities could not compensate for the excessive
costs of the imported agrochemicals on which production depended. Moreover, as
the soils were gradually depleted, higher doses of fertilizers were required every year
simply to maintain production. After a decade of struggling to keep the farm afloat,
the need to reduce input dependency and keep production costs under control
became evident.
In 1997 the decision was made to stop chemical fertilization altogether. This not
only required cutting animal load by half, but it also revealed the severity of soil
degradation. Finding a strategy to restore soil fertility without applying external
inputs thus became the priority. Experience and self-reflection about traditional
production systems eventually led to a question: if the soils of coffee systems shaded
with nitrogen-fixing Inga trees were so rich that they required little fertilization and
no pesticides, could the same principles apply to pastures? So, defying the wide-
spread idea that trees and pastures do not mix, in 2002 a 2-hectare trial plot was
established with Inga spp. trees planted at a density of 100 trees ha1.
The first 2 years of this trial were difficult: establishing trees in active pastures
turned out to be more challenging and expensive than anticipated. Damage by cattle,
leaf-cutter ants and humans all contributed to the high mortality rate of the planted
seedlings. The idea that trees do not belong in pastures was deeply engrained, and
farm workers were reluctant to protect and care for the seedlings which resulted in
the need to replant constantly. Protecting the young trees from cattle browsing and
trampling was also problematic and required creative solutions (Fig. 12.2). Trees
were initially protected with individual bamboo corrals, but eventually a better
method was developed by planting lines of trees protected with electrical fence.
Fellow cattle ranchers mocked these efforts arguing that intentionally increasing tree
cover in the paddocks was nonsense as it would reduce pasture productivity.
In 2002, as the landowner began to question the introduction of Inga trees in the
pastures, news about the launching of a sustainable ranching project in the region
reached him. The Regional Integrated Silvopastoral Ecosystem Management
(RISEM) project was providing technical assistance and payments for ecosystem
services to promote the adoption of silvopastoral systems, previously unknown in
310 I. Montes-Londoño et al.

Table 12.1 Main events in Pinzacuá’s land-use history

Year Event
1985 Pinzacuá is acquired. The farm is a traditional
agroforestry plantation with coffee growing
under diverse assemblages of shade trees,
mainly Inga.
1986 (Inspired by the Green Revolution, the Shade coffee is replaced with sun-coffee man-
National Coffee Federation promoted intensifi- aged with intensive application of synthetic
cation efforts during these years, paying farmers fertilizers and pesticides, and shade trees are
to eliminate tree cover and replace traditional completely eliminated.
shade coffee for more productive sun-loving
varieties. Tree cover was eliminated in 50% of
this territory as a result of both coffee intensi-
fication campaigns and a push to replace coffee
with pastures in the 1990s (Rodríguez et al.
2004). These changes led to landscape-level
degradation and fragmentation of the remaining
natural habitats, and in some cases triggered
local or regional extinctions (Kattan et al.
2004). Although yields did increase, approxi-
mately half of the increased production was lost
due to a labor shortage)
1993 Sun coffee is eliminated due to low market
prices, and replaced by a monoculture of the
African grass Cynodon plectostachium without
trees, and a monocrop plantation of tobacco.
1994 Tobacco is eliminated and the entire farm is
converted to pasture. Pasture management
based on intensive application of fertilizers and
amendments (1 ton of urea plus 2 tons of
dolomite lime ha1 year1) continues in order
to sustain a high animal load (10 heads ha1).
1998 Use of agrochemicals becomes economically
unviable and is completely eliminated. As a
result, animal loads are reduced and soil deg-
radation becomes evident. Shifting away from
grass monoculture to a regenerative system
becomes the priority.
2000 A trial plot with Inga trees is established with
the hope of restoring soil fertility.
2002 Using a farm planning approach, plots are
classified according to their potential and lim-
itations. Streams are isolated and riparian areas
reforested with native trees and bamboo
(Guadua angustifolia).
2015 Fodder hedgerows are established within some
paddocks.
2016 A small agroforestry plot with shade-loving
arabica coffee is established.
2018 A forage bank to feed goats is established.
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 311

Fig. 12.2 Methods to plant and protect trees in established active pastures. Individual bamboo and
shade cloth corral method (left), and 2 m-wide strips protected with electrical fence (right), an
innovation devised in Pinzacuá and later replicated in other farms. Photos: Álvaro Zapata

the region (Pagiola and Ríos 2013). Pinzacuá, along with other 74 farms, joined the
project and started working with assistance from the Center for Research on Sus-
tainable Agriculture Production Systems (CIPAV, [Link]), whose technical
staff advocated increasing tree cover throughout the farm and encouraged the Inga
trial. Although the project’s economic incentive was capped at US $6500 per farm
and only covered approximately 30% of the implementation costs incurred, the
motivation and support provided by CIPAV were instrumental for Pinzacuá to
persevere in its tree-planting efforts.
From that point on, tree planting became the centerpiece of Pinzacuá’s transfor-
mation. A land use plan was devised to utilize each plot according to its potential.
Trials planting a variety of trees in different spacing arrangements were expanded
throughout the farm to identify the optimal combinations. Paddocks were divided
with live fences and cattle were rotated in short periods. All streams and water
courses were fenced off, and riparian buffer areas were reforested with native tree
species and Guadua angustifolia, a native giant bamboo. The steepest areas of the
farm, which were unsuitable for cattle, were planted with mixed native species. More
recently, an agroforestry system with arabica coffee was established and other
non-timber forest products such as vanilla and pepper were tested. Fodder hedge-
rows were planted in some paddocks and a forage bank to feed goats was
established.
312 I. Montes-Londoño et al.

The changes implemented in Pinzacuá described in the following sections were

inspired by a variety of sources. As a former coffee grower, the landowner drew
initial ideas for tree planting densities and pruning regimes from technical manuals
for shade coffee published by Colombia’s Coffee Grower’s Federation
([Link]). Drawing on decades of experience developing
silvopastoral systems and management, CIPAV staff offered guidance on tree and
fodder species and pasture management, and helped adjust the systems to Pinzacuá’s
conditions. Visits to demonstration farms, especially to the flagship silvopastoral
farm El Hatico Nature Reserve (Calle et al. 2022), enabled farmer-to-farmer learning
and inspired many ideas for small scale trials. Ultimately, the landowner’s own
curiosity and innovative spirit, observation skills, and proclivity to old-fashioned
empirical learning were critical for the refinement of techniques that worked for
Pinzacuá, some of which went on to become standard silvopastoral practice (Calle
2008).

12.3.2 Trees and Living Soils, the Pillars of Change

Ever since coffee was introduced in Colombia, trees have played a key role in
traditional coffee agroforestry systems. Pinzacuá’s transformation and its current
land use practices are rooted in this traditional knowledge and in the realization that
the mechanisms that sustained high natural productivity in shaded coffee systems are
largely based on the benefits provided by trees. Today, the entire farm is planted with
trees and all land uses—silvopastures, riparian areas, and the homegarden—have
some form of tree cover (Fig. 12.3).
The impact of trees on productivity is not direct, but rather mediated by their
effect on different components of the production system, from the livestock to the
soils. The strength and quality of high-performing soils rest on the interactions
between a multitude of beneficial macro and microorganisms that constitute the
soil food web. Earthworms, arthropods, nematodes, fungi, protozoa and bacteria
consume energy-rich materials (e.g., leaves, roots, root exudates) all of which are
directly or indirectly sourced from plants, especially trees, via litterfall and roots.
Without plentiful and diverse plant-derived organic inputs, the soil food web cannot
thrive (Moebius-Clune et al. 2016). For this reason, and despite the inherent com-
petition between trees and pastures, the main objective of all actions implemented in
Pinzacuá since 1997 has been to adequately feed and provide the best conditions for
the life belowground. Plants, and especially trees, have been the farm’s best allies in
restoring soils and making them alive, dynamic, and productive.
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 313

Fig. 12.3 Aerial views of Pinzacuá in 2003 (top) and 2016 (bottom). Main land use in 2003:
treeless pasture; in 2016: silvopastoral systems, riparian forests, and homegarden. Land uses in
neighboring farms can also be observed. Photos: RISEM Project
314 I. Montes-Londoño et al.

12.3.3 Land Use Planning

Today, the level and type of tree cover (i.e., tree density and arrangement) are
intentionally managed throughout the farm. As a general rule, soil suitability (e.g.,
fertility, depth, slope, vulnerability to erosion and compaction) determines the type
of land use, which in turn determines tree cover and tree species (Table 12.2). For
example, steep hillsides, which are vulnerable to erosion and compaction, are not
used for cattle but instead are planted with trees at a high density using a mixed
native species forestry model described below. By contrast, the flat areas atop hills,
which are more fertile and less prone to erosion, are reserved for cattle production
and planted with trees at a lower density to ensure that pastures receive sufficient
sunlight.

12.3.4 Tree Species Selection

Pinzacuá’s planned tree diversity includes more than 60 different species that have
been intentionally incorporated into the different land uses, including some of
conservation concern. With the exception of the homegarden, native species are
preferred over exotics because they are better adapted to the local climatic and
edaphic conditions and provide food and shelter for the organisms above and
below ground.
Trees for the silvopastoral systems were selected for specific characteristics
including: (i) nitrogen fixation or high production of rapidly decomposing litter
(e.g., Inga edulis); (ii) hardiness to withstand herbivory, both from cattle and leaf
cutter ants (e.g., Maclura tinctoria); (iii) rapid growth and vigor to outcompete
grasses (e.g., Gliricidia sepium); (iv) seeds, fruits and forage production to supple-
ment cattle and horse nutrition (e.g., Senna spectabilis); and (v) timber production
for farm use or local markets (e.g., Anacardium excelsum). Meanwhile, riparian area
reforestation favored two fast-growing native species: Guadua angustifolia, a giant
bamboo with high local economic and cultural value; and Colubrina sp., a tree with
good timber properties.
The homegarden includes a mix of native and exotic species of cash crops (e.g.,
Vanilla planifolia), medicinal plants (e.g., Sechium edule), and ornamentals (e.g.,
Anthurium sp.), as well as timber trees (e.g., Swietenia macrophylla), fruit trees (e.g.,
Garcinia madruno), multipurpose trees (e.g., Inga edulis) and palms (e.g., Aiphanes
caryotifolia).
For the forestry plantations, dinde (Maclura tinctoria (L.) D. Don ex Steud) was
the main species selected because of its timber quality, high survivorship relative to
other native species, and ability to provide fruit and habitat for birds and other
wildlife (Martins and Setz 2000; Chízmar-Fernández 2009; Suárez et al. 2012;
Montes-Londoño et al. 2017). Also known as old fustic or Argentine osage orange,
dinde is valued for heavy construction, flooring, furniture, turnery, fence posts and
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 315

Table 12.2 Current land uses and their extension, relief, and species planted
Tree
density
Land use and (trees
area Relief type ha1) Plant species
Silvopastoral Hilltops 100–150 Albizia guachapele,
system Anacardium excelsum,
15 ha Anadenanthera peregrina,
Brachiaria sp., Cassia
grandis, Cynodon
plectostachyus, Enterolobium
cyclocarpum, Gliricidia
sepium, Hymenaea courbaril,
Inga edulis, Leucaena
leucocephala, Pennisetum sp.,
Psidium guajava, Samanea
saman, Senna spectabilis,
Syagrus sancona
Mixed planted Hillsides Initial den-
forest sity: 1100;
15 ha
Final density: Anacardium excelsum, Aniba
250. perutilis, Cedrela odorata,
Lafoensia acuminata, Maclura
tinctoria, Magnolia
hernandezii, Ocotea
helicterifolia, Swietenia
macrophylla, Vachellia
macracantha
Riparian areas Depressions >900 Anacardium excelsum,
5 ha Guadua angustifolia
Shaded coffee Hillside 500 shade Albizia carbonaria,
1 ha trees, 2000 Anacardium excelsum,
coffee Caesalpinia ebano (threat-
shrubs ened in Colombia), Citrus sp.,
Inga edulis, Jacaranda
mimosifolia, Pourouma
cecropiifolia, Pouteria
caimito, Quararibea cordata,
Swietenia macrophylla
Natural regen- Hillside/ depression 500–900 Cedrela odorata, Colubrina
eration and sp., Erythina fusca, Erythrina
enriched sec- poeppigiana, Juglans
ondary forest neotropica, Magnolia
1 ha hernandezzi, Montanoa
quadrangularis
Homegarden Hilltop 500 Acca sellowiana, Aiphanes
0.24 ha caryotifolia, Anthurium sp.,
Arachis pintoi, Bactris
gasipaes, Bismarckia nobilis,
Brownea ariza, Bougainvillea
(continued)
316 I. Montes-Londoño et al.

Table 12.2 (continued)

Tree
density
Land use and (trees
area Relief type ha1) Plant species
sp., Cassia grandis, Ceiba
pentandra, Citrus sp.,
Codiaeum variegatum,
Crescentia cujete, Diospyros
sp., Eugenia stipitata,
Euphorbia pulcherrima,
Garcinia madruno, Garcinia
mangostana, Inga edulis,
Juglans neotropica,
Myrciaria cauliflora,
Passiflora edulis, Passiflora
ligularis, Persea americana,
Piper nigrum, Sechium edule,
Selenicereus megalanthus,
Senna spectabilis, Swietenia
macrophylla, Tamarindus
indica, Vanilla planifolia
Forage/protein Hillside 3000–4000 Alocasia macrorrhiza,
bank Boehmeria nívea, Guazuma
0.5 ha ulmifolia, Pennisetum sp.,
Tithonia diversifolia,
Trichanthera gigantea.

railroad crossties, and specialty wood items, and is culturally and economically
important throughout Latin America. Dinde is the source of fustic, a yellow pigment
used as dye for khaki and other color textiles (Rangel 1949; Roig 1974). The leaves,
sap and wood have been used in traditional medicine, and have potential for
extraction of a non-toxic broad-spectrum antioxidant (Cioffi et al. 2003). Once
common in La Vieja river watershed, dinde populations have been decimated;
today it is rarely found in the forests or in the markets, although it is considered a
species of least concern (LC) for conservation (Rivers et al. 2017).

12.3.5 Tree Density and Shade Management

[Link] Silvopastures

The main factor affecting pasture productivity in silvopastures is shade, which in turn
depends on the level of canopy cover and the characteristics of the tree canopy. Thus,
identifying the level of shade that maximizes pasture productivity is key to successful
silvopasture management. In Pinzacuá, informal trials manipulating shade with
different planting densities and pruning intensities have shown that the pasture of
choice, star grass (Cynodon plestostachys), performs best with <50% shade.
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 317

Fig. 12.4 Final tree spacing arrangement in the silvopastoral system. Trees are planted in an east-
west direction to allow maximum light penetration to the understory. Photo: Fernando Uribe, 2018

Three different spatial arrangements identified through empirical observation

were tested to determine the optimal tree density to both restore the soils and
allow enough light into the understory: (i) a 10  10 m grid (100 trees ha1);
(ii) rows planted every 20 m with 5 m between trees (100 trees ha1); and (iii) rows
planted every 15 m with 5 m between trees (133 trees ha1). The latter design has
worked best as the initial planting arrangement, and is then gradually reduced to a
final density of approximately 100 trees ha1 to avoid canopy closure (Montes
Londoño unpublished data) (Fig. 12.4).
The optimal level of shade at this tree density is maintained by pruning three
fourths of the tree crown every 2 years. Branches growing in east-west direction are
retained while those growing north-south and directly shading the pasture corridors
are removed (Fig. 12.5). While pruning is labor intensive, the residue is used to make
artisanal charcoal and sold to cover the labor costs and to get some additional
revenue (Fig. 12.5). Furthermore, the residue from the charcoal heaps is used as a
high-quality biochar amendment to improve soil fertility throughout the farm.

[Link] Forestry Plantations

The forestry plantations were established with a mix of native timber trees planted in
a 3  3 m spacing arrangement. The grid pattern spreads trees out to minimize
competition, and the high initial density prevents early branching and promotes
self-thinning, reducing the need for frequent pruning. Because dinde is good at
self-thinning, plantations require less intensive management than silvopastures.
318 I. Montes-Londoño et al.

Fig. 12.5 Pruning of Inga trees (left) and artisanal charcoal production from the pruning material
(right). Photos: Álvaro Zapata

Thinning is performed according to a specific guideline developed for dinde

(Montes-Londoño et al. 2017).1

12.4 Impact on Land Cover and Biodiversity

12.4.1 Land Cover Change

Although Pinzacuá’s transformation has been slow and gradual, 20 years into the
process the accumulated changes in land use and tree cover are substantial. A spatio-
temporal analysis based on high resolution satellite images revealed that between
2003 and 2016 land uses with high tree cover (i.e., forests, silvopastures and live
fences) increased by 74.9% at the expense of uses with low tree cover (i.e., treeless
pastures and croplands) (Table 12.3, Fig. 12.6) (Alicia Calle 2018, unpublished
data). Forests included both reforestation and forestry plantations with a closed
canopy and natural regeneration, and silvopastures were all areas of pasture with
intermediate or high tree density. The most important changes during this period
occurred in forests, which increased by 14.4% (6.3 ha); pastures with tree cover,
which increased by 62.9% (27.6 ha); and pastures without trees, which initially

1
Given the lack of information on growth and silvicultural management for dinde, research was
conducted on 12 farms of the region to develop preliminary spacing and thinning guidelines (see
Montes et al. 2017). Dinde performed similar to other trees used for reforestation in the humid
American tropics, with a Mean Annual Increment in Diameter at Breast Height (MAIDBH) of
2.56 cm year1, a growth rate which is higher than it has been reported for sites in Honduras and
Cuba (Cordero and Boshier 2003) but lies on the lower end of fast-growing native and exotic
species in Central America. This growth rate stands overall within the range that has been reported
for other native species in tropical humid regions elsewhere in Latin America (Piotto et al. 2004;
Wishnie et al. 2007; van Breugel et al. 2011).
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 319

Table 12.3 Land cover change in Pinzacuá and the surrounding landscape from 2003 to 2016
(Alicia Calle, 2018, unpublished data)
Surrounding
Pinzacuá landscape
2003 Area 2016 Area Change %
(ha) (ha) (ha) Change % Change
Forests 2.85 9.18 6.33 14.44 2.82
Silvopastures 1.75 29.31 27.56 62.86 0.80
Live fences 1.23 0.17 1.06 2.43 0.62
Pasture no tree 34.74 2.82 31.92 72.81 15.85
cover
Cropland (no trees) 1.21 0.05 1.16 2.64 10.92
Other/No 2.06 2.32 0.25 0.58 1.38
information
Total 43.84 43.84

Fig. 12.6 Land cover in Pinzacuá, 2003 and 2016. Maps: Alicia Calle

comprised most of the farm but decreased by 72.8% (31.9 ha) (Calle unpublished
data). Given the small size of the property, the decision to release areas for forest
restoration and convert all treeless pastures to silvopastures underscores the land-
owners’ appreciation for both the direct and indirect benefits of trees.
320 I. Montes-Londoño et al.

The magnitude of changes implemented in Pinzacuá, however, is perhaps best

appreciated in the context of the surrounding landscape. During the same 13-year
period, land cover change in a total of 557.4 hectares covered by farms surrounding
Pinzacuá followed a different trend. Forest cover increased modestly (2.8%) and
treeless pastures decreased by 15.9%, mostly to accommodate cropland expansion
(10.9%) with only marginal improvements in pasture tree cover (1.4%) (Calle 2020)
(Table 12.3, Fig. 12.6). The neighboring farms, which were not part of the RISEM
project, had no access to payments for ecosystem services or technical assistance to
improve their pasture areas. Instead, they appear to have planted cash crops in
response to short-term spikes in market prices. Although spillover of silvopastoral
systems onto neighboring farms did happen in other project farms, Pinzacuá’s
neighbors were not interested. Location adjacent to a large town and a main road
connecting to markets may partly explain why other farmers opted for high-value
cash crops.

12.4.2 Biodiversity

Since the transformation began, Pinzacuá has been the site of numerous biodiversity
studies. While the lack of baseline assessments does not allow for direct compari-
sons, the minimal vegetation cover and severe land degradation evident in the 2003
satellite image (Fig. 12.6) suggest low levels of biodiversity at the baseline. The
studies and species documented on the farm are summarized in Table 12.4.

[Link] Trees

Pinzacuá’s transformation has relied heavily on planting trees and managing woody
natural regeneration on degraded pastures for different purposes. Riparian forests
were established to improve and protect the water supply; forestry plantations were
established in the steepest parts of the farm that were not apt for grazing; and
different silvopastoral arrangements were implemented to reduce cattle heat stress
and improve productivity. As a result, overall tree diversity has increased dramati-
cally over time. The farm also propagates several tree species and shares seedlings
with others interested in improving on-farm tree cover.
To date, studies of tree diversity on the farm have identified 45 different tree
species as adults or seedlings across the different land uses (Table 12.4). Thirty-six
woody species have been identified across the different forest fragments, 17 as adult
trees and 28 as regenerating seedlings. Eight tree species were also recorded in the
mixed plantation (Calle and Méndez 2017, unpublished data, Giraldo et al. 2019),
and 16 additional tree species have been recorded in the silvopastures.
A survey of woody vegetation structure and composition in 20 recovering forests
protected during the RISEM project included two young riparian forests in Pinzacuá.
The study found five species in the canopy and 25 species regenerating in the farm’s
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 321

Table 12.4 Biodiversity studies and species reported in Pinzacuá

Woody plants Author and methods
11 species Cedrela odorata, Cestrum sp., Cordia 1000 m2 transects (Calle and
219 alliodora, Gliricidia sepium, Inga edulis, Méndez 2017, unpublished data)
individuals Leucaena leucocephala, Maclura tinctoria,
Psidium guajava, Senna spectabilis,
Vachellia macracantha, Vernonanthura
patens
Anacardium excelsum, Erythrina
poeppigiana, Syagrus sancona (recorded
outside the plots)
16 species Anacardium excelsum, Bauhinia picta, 1000 m2 transects (Giraldo et al.
Cecropia angustifolia, Cedrela odorata, 2019, unpublished data)
64 individuals Cordia alliodora, Cupania americana,
Erythrina fusca, Erythrina poeppigiana,
Gliricidia sepium, Juglans neotropica,
Montanoa quadrangularis, Nectandra
turbacensis, Psidium guajava, Samanea
saman, Solanum aphyodendron
Woody regeneration
28 species Anacardium excelsum, Bauhinia picta, 1000 m2 transects (Calle and
834 Cecropia angustifolia, Celtis schippii, Méndez 2017, unpublished data)
individuals Cinnamomum triplinerve, Citrus sinensis,
Cordia alliodora, Cupania americana,
Erythrina poeppigiana, Eugenia sp, Ficus
insipida, Ficus tonduzii, Hymenaea
courbaril, Inga edulis, Maclura tinctoria,
Myrcia sp, Nectandra lineata, Nectandra
turbacencis, Ocotea macropoda,
Oroepanax cecropifolium, Persea ameri-
cana, Sapium laurifolium, Senna
spectablis, Sorocea trophoides,
Tetrochidium rubrinervium, Trichilia
pallida, Zanthoxylum rhoifolium
Dung beetles
11 species Canthidium sp., Delthochilum sp., Pitfall traps
97 individuals Dichotumius alyattes, Dichotomius sp., (Giraldo et al. 2019, unpublished
Eurysternuspp; —mexicanus, Eurysternus data)
plebejus, Ontherus azteca, Onthophagus
acuminatus, Onthophagus nasutus,
Onthophagus sp., Oxysternon
conspicillatum
Birds
34 species Amazilia tzacatl, Amazona ochrocephala, Point counts
104 Anthracothorax nigricollis, Ara severus, (Giraldo et al. 2019, unpublished
individuals Bubulcus ibis, Coccyzus pumilus, Coereba data)
flaveola, Columbina talpacoti, Dendroplex
picus, Icterus nigrogularis, Lepidocolaptes
souleyetii, Melanerpes formicivorus,
Melanerpes rubricapillus, Myiarchus
apicalisa, Myiodynastes maculatus,
(continued)
322 I. Montes-Londoño et al.

Table 12.4 (continued)

Woody plants Author and methods
Myiopagis viridicata, Picumnus
granadensisa, Pionus chalcopterus, Pionus
menstruus, Piranga rubrab, Pitangus
sulphuratus, Polioptila plúmbea,
Pyrocephalus rubinusb, Setophaga
petechiab, Sicalis flaveola, Synallaxis
azarae, Thamnophilus multistriatus,
Thraupis episcopus, Thraupis palmarum,
Tiaris olivaceus, Todirostrum cinereum,
Troglodytes aedon, Tyrannus
melancholicus, Zimmerius chrysops
89 species Bubulcus ibis, Camptostoma obsolete, Point counts (Sánchez and
28 families Columbina talpacoti, Columbina minuta, Camargo 2015)
1069 Dendroica petechia, Egretta thula,
individuals Eucometis penicillate, Leptopogon
superciliaris, Myiodynastes maculatus,
Phaeothlypis fulvicauda, Pachyramphus
rufus, Pygochelidon cyanoleuca,
Pyrocephalus rubinus, Sporophila
schistacea, Thraupis episcopus,
Tyrannulus elatus, Tyrannus
melancholicus, Vireo olivaceus,
Xiphorhynchus susurrans, among others
Butterflies
16 species Actinote ozomene, Anartia amathea, Visual encounter surveying
Anthanassa Drusilla, Caligo eurilochus, (VES) (Montes-Londoño 2019,
Ceratinia tutia, Cissia pompilia, Dryas unpublished data)
iulia, Eurema daira, Heliconius
charitonia, Heliconius erato, Papilio
thoas, Pteronymia artena, Pteronymia
picta, Siproeta stelenes, Timolus echion,
Urbanus proteus
Mammals
8 species Sciurus granatensis, Didelphis Observations by managers and
marsupialis, Dasyprocta spp., Dasypus workers
novemcinctus, Cerdocyon thous, Puma
yagouaroundi, Cuniculus paca, Leopardus
pardalis
Reptiles
6 species Chironius monticola, Bothriechis Observations by managers and
schlegelii, Micrurus mipartitus, Oxybelis workers
fulgidus, Oxyrhopus petola, Spilotes
pullatus
a
Endemic species
b
Migratory species
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 323

bamboo forest, and another 11 species in the canopy and 11 species regenerating in
the mixed planted forest (Calle and Méndez 2017, unpublished data) (Table 12.4).
Pinzacuá’s bamboo forest had the highest abundance of woody regeneration with
785 seedlings, including many species that are not currently represented in its forest
canopy. The study suggests that these riparian sites that were retired from cattle
production over a decade ago are slowly recovering, and the total number of woody
species will likely increase over time (Calle and Holl 2019).

[Link] Birds

Birds play a number of important roles in both natural and agricultural ecosystems:
they contribute to natural pest control by consuming pest insects and organisms; they
pollinate flowers helping with fruit production; they disperse seeds facilitating
natural regeneration; and they add aesthetic and cultural value to landscapes (Giraldo
et al. 2019).
Two different bird studies have been conducted in Pinzacuá. Sánchez and
Camargo (2015) examined the relationships between bird diversity and landscape
structure in four farms with different land uses (i.e., conventional treeless pastures,
silvopastoral systems and agroforestry systems) connected to bamboo forests. The
highest bird diversity and abundance were recorded in Pinzacuá, where 1069
individuals belonging to 89 species and 28 families, including 3 migratory and
3 endemic species (Table 12.4), were sighted in the silvopastoral systems connected
to bamboo forests. The structural and functional attributes of these systems create
connectivity to the forest edges, effectively providing food and habitat resources
needed for the permanence of bird species, both local and migratory.
The abundance of birds in the Tyrannidae family (180 species) suggests that
silvopastoral systems offer optimal conditions for insectivorous species. These birds
are known to reduce insect pest outbreaks and herbivory in multi-strata cocoa and
coffee agroforestry systems (De Beenhouwer et al. 2013; Maas et al. 2013; Peters
and Greenberg 2013; Poch and Simonetti 2013). Although the same could be true in
silvopastoral systems, where the main pests are ticks and flies, the role of insectiv-
orous birds in controlling pests in these systems has not been studied.
Another study comparing bird diversity across three different land uses in
Pinzacuá (i.e., forest, silvopastoral systems, and fodder hedges) recorded 104 indi-
viduals belonging to 34 species (Giraldo et al. 2019) (Table 12.4). Most species were
open area generalists but at least four were forest dwellers; three species were
migratory and three others were endemic. Silvopastoral systems with scattered
trees had the highest bird diversity, followed closely by the forest; they also hosted
a variety of common generalists that have been displaced by the elimination of
coffee shade and overall tree cover across the region.
The presence of birds with a wide range of habitat preferences in Pinzacuá
suggests that the complex and perennial habitat diversity that was intentionally
created in the farm facilitates bird movement in fragmented agroecosystems and
can provide important habitat for farmland species and migratory birds (Philpott and
Bichier 2012; Vílchez et al. 2013).
324 I. Montes-Londoño et al.

[Link] Dung Beetles

Dung beetle communities are shaped by both dung availability and vegetation
structure. Because beetle assemblages can be severely modified by farming intensi-
fication practices and the elimination of tree cover, dung beetles are useful indicators
of land use change and pasture health (Davis et al. 2004; Giraldo et al. 2011). In
grazing landscapes, dung beetles mediate important ecosystem functions: they
rapidly remove dung piles from the pasture surface by incorporating them into the
soil and improving its fertility; they reduce soil compaction and improve soil
structure; and they interrupt the life cycles of flies and parasites that lay their eggs
on dung piles and affect cattle health (Giraldo et al. 2019).
A study comparing dung beetle diversity across three land uses in Pinzacuá found
97 individuals of 11 species, of which 24 individuals of five species were in the
pastures with scattered trees (silvopastures); 20 individuals of seven species were in
the fodder hedges; and 53 individuals of eight species were in the planted forest
(Table 12.4). Species diversity was twice as high in the forest than in the
silvopastures, and similar in silvopastures and fodder hedges (Giraldo et al. 2019).
Structural complexity, high canopy cover, and the presence of a leaf litter layer that
retains moisture explain the higher abundance and richness in the forests, as well as
the diverse assemblage in the pastures with high tree density.

[Link] Other Species

No formal studies of mammals or reptiles have been conducted to date on Pinzacuá.

However, squirrels, opossums, agoutis, and armadillos are common on the farm,
crab eating fox (Cerdocyon thous) and jaguarundi (Puma yagouaroundi) have been
occasionally seen, and lowland pacas (Cuniculus paca) and ocelots (Leopardus
pardalis) are likely present as they have been spotted in neighboring farms. In
addition, three species of snakes including the false coral (Oxyrhopus petolarius),
the green vine snake (Oxybelis fulgidus), and the yellow rat snake (Spilotes pullatus)
are also on site.
Overall, Pinzacuá’s efforts to increase vegetation cover and complexity by
planting and protecting a variety of tree species have not only increased tree
diversity, but they have provided adequate resources for other wildlife species,
thereby transforming this previously degraded farm into an island of biodiversity
amidst an otherwise impoverished landscape. Furthermore, Pinzacuá has become a
site for biodiversity research and has contributed to our understanding of how the
incorporation of trees can aid in the recovery of degraded farmlands.
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 325

Table 12.5 Productivity indicators for two intensive cattle ranching systems in Colombia
High input treeless pastures, Cimitarra, Silvopastoral system,
Productivity indicators Santandera Pinzacuá
Stocking rate (animal unitsb 7 5.5
ha1)
Meat production 600 600
(gr animal1 day1)
Conception rate 80% 95%
(excluding first time
pregnancies)
Veterinary care costs 4% of total input cost 0.5% of total input cost
(vet, pest control, vitamins,
etc.)
Calf mortality rate 5% 3%
a
Personal communication with Antonio José Uribe, CIPAV
b
1 animal unit ¼ 450 Kg

12.4.3 Impact on Productivity

In terms of productivity, the silvopastoral systems implemented over the past

20 years have allowed the farm to support cattle stocking rates similar to those
seen in high-input cattle grazing systems, while also eliminating the use of chemical
fertilizers completely. During this period fodder biomass production in the pastures
has increased from two tons ha1 in 1997 when fertilization ceased, to 11 tons ha1
in the silvopastoral systems. Compared to high-input grazing systems, Pinzacuá’s
calf mortality rate is 2% lower and the birth rate is 10% higher, with a calving
interval of 13 months (Table 12.5) (Reyes et al. 2017). In addition, the shade
provided by trees has reduced the risk of heat stress, creating a more comfortable
environment in which animals stay hydrated and maintain their normal feeding
behavior (Broom et al. 2013). Shade also provides habitat for predators that control
ticks, flies, and other common cattle pests, thereby reducing the incidence of disease.
These improvements in livestock’s’ bodily condition reduce costs in veterinary care
and medications. Overall, the direct and indirect benefits provided by trees are
reflected in better animal welfare and higher productivity.

12.4.4 Impact on Ecosystem Services

[Link] Water Quality and Quantity in Restored Streams

The re-establishment of riparian corridors in the farm’s previously unprotected

steams has also impacted water quality. The upstream town of Alcalá discharges
raw sewage from approximately 1400 people into the main stream that flows through
Pinzacuá at a rate of 10.6 L s1. An evaluation of the stream’s self-purification
326 I. Montes-Londoño et al.

capacity was conducted in 2010, approximately 10 years after 5 to 10-meter-wide

strips were fenced off and reforested with guadua (Guadua angustifolia) on both
streambanks. Analyses of three composite water samples taken at 400-m intervals
showed the following removal efficiencies for five pollutants along the 800-m
stretch: 52% for nitrogen, 49% for phosphorus, 88% for chemical oxygen demand
(COD), 92% for total coliforms and 97% for fecal coliforms (Montes Londoño and
Rodríguez Susa 2011). Compared to similar studies, the stream appears to retain its
self-purification capacity thanks in part to resemblance of the restored forests to the
basin’s historical guadua forest cover. Despite the lack of baseline data, workers and
landowners report substantial changes in water quality parameters such as color,
odor, turbidity and the presence of foam. Thus, riparian restoration appears to have
had a positive effect on the water, underscoring the links between riparian area
protection, land management practices, and water quality.

[Link] Climate Change Mitigation

Whereas greenhouse gas (GHG) emissions, carbon stocks and carbon sequestration
have not been directly measured at Pinzacuá, numerous studies from tropical and
sub-tropical Latin America highlight the potential of tree-based grazing systems for
climate mitigation. Silvopastoral systems can contribute in two important ways: via
carbon sequestration in both the woody biomass and the soils, and via an improved
animal diet that reduces emissions from enteric fermentation (Ibrahim et al. 2007;
Arias Giraldo et al. 2009; Mesa Arboleda 2009; Amézquita et al. 2010; Naranjo et al.
2012; Montagnini et al. 2013; Aynekulu et al. 2019).
Silvopastoral systems store more carbon than grass-only systems because of their
higher amount of above and belowground biomass. A study conducted in the
RISEM project farms estimated a total carbon storage of 153 Mg C in 1 ha of
silvopasture consisting of improved grass planted with 83 native trees (Arias Giraldo
et al. 2009). Silvopastures also perform better in their total GHG balance, as shown
in a study comparing four different types of pasture systems in Colombia. The two
conventional systems—degraded treeless pastures and improved treeless pastures—
were net GHG sources, emitting 3.2 and 3.3 tons CO2e ha1 year1 respectively.
Meanwhile, the two silvopastoral systems—one combining improved grasses with
Leucaena leucocephala (60,000 shrubs ha1) and a similar one that also had mixed
species of timber trees—were net sinks, respectively removing 8.8 and 26.6 ton
CO2e ha1 year1 (Naranjo et al. 2012).
Sequestration potential varies with factors such as soil type, tree species, stand
age, and management (Nair et al. 2010). However, these studies suggest that given
the farm’s baseline condition of mostly degraded soils and treeless pastures, and the
significant increase in tree cover over the past two decades, Pinzacuá has become a
provider of climate mitigation services.
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 327

12.5 Discussion and Conclusions

12.5.1 Pinzacuá as an Island of Biodiversity

Twenty years ago, Pinzacuá began a gradual transformation of its land use and
management practices with the goal of improving productivity and ensuring eco-
nomic viability. The results are evident.
Increasing farm productivity was the key first step that allowed the farm to reduce
the production area to accommodate more areas for conservation. Pasture mono-
cultures were replaced with live fences, silvopastures, riparian buffers, and connec-
tivity corridors, transforming the farm into a mosaic of vegetation structures
associated to different land uses. Today, the added diversity and complexity pro-
vides refuge for birds, bats, and other mammals; native flowering trees offer
resources for wildlife and a variety of beneficial insects; canopy cover creates
favorable conditions for ground-dwelling ants and dung beetles that require moisture
and tree litter to move through the pasture; trees support the web of underground
biodiversity that underpins soil fertility; and riparian corridors and high tree cover
provide connectivity throughout the farm. In short, Pinzacuá has become a high-
quality matrix that serves as permanent habitat or temporary refuge for a variety of
taxa that strive to persist in this highly fragmented landscape. As seen in the satellite
images, Pinzacuá stands out, literally, as an island of vegetation in an otherwise
treeless landscape (Calle 2020).
Beyond the physical transformation, Pinzacuá’s process is the expression of a
deep cultural change that began at a personal level and eventually became a family
project. However, Pinzacuá alone is not enough, and its impact will be limited unless
other farms make similar management decisions and intentional efforts to integrate
more trees and change harmful practices.

12.5.2 Challenges and Setbacks

While transitioning from input-based conventional systems to agroecological-based

systems can entail a productivity trade-off, silvopastoral systems may be a notable
exception. The extensive cattle production model currently used across Latin Amer-
ica is known for its low productivity (0.6 animal units ha1 (FAO 2006) and high
environmental impact. By contrast, the higher productivity of silvopastoral systems
is rooted precisely in their environmental benefits, and both cannot be decoupled
(Murgueitio et al. 2011; Chará et al. 2019). The question then is, if the systems is so
good why are more cattle ranchers not adopting it? The answer is simple:
silvopastoral systems are knowledge and management intensive, and they require
a significant up-front investment that takes time to recover. Whereas Pinzacuá’s
owner was able to keep the farm afloat during the transition period by drawing from
another business, many farmers are not in a position to do the same. Unfortunately,
328 I. Montes-Londoño et al.

the structures needed to overcome these entry barriers are currently not in place in
most countries. In recent years, however, the RISEM and other projects have shown
that economic incentives and technical assistance can effectively address these
barriers and deliver lasting impacts (Calle 2020; Pagiola et al. 2020).
Pinzacuá’s transformation clearly illustrates the potential to increase productivity
while restoring and conserving the natural capital, a process known as ecological
intensification (Gaba et al. 2014). However, this transformation requires a radical
shift in mindset, from one of controlling nature to one of learning from nature, as
well as perseverance in facing challenges. Below we describe some of the failed
trials and unexpected setbacks faced at Pinzacuá, as well as some of the emerging
challenges.
One of the earliest failures was the implementation of an intensive system with
Leucaena leucocephala, a nutritious nitrogen-fixing fodder tree/shrub (Murgueitio
et al. 2011). Although expensive to implement, the system’s high productivity can
quickly offset the investment. However, Pinzacuá’s soils proved to be too acidic for
Leucaena, and the time and resources invested were lost. A number of failures were
related to tree-planting. For example, different versions of corrals built with barb
wire or bamboo and efforts to spray cattle dung and other substances to deter
herbivores failed to keep leaf-cutter ants and cattle at bay. Eventually better protec-
tion methods were developed through trial and error. The high mortality of healthy-
looking tree seedlings, also frustrating early on, was later attributed to roots coiling
in the small bags used by commercial nurseries. Careful sourcing of plant material
and direct seeding techniques eventually helped overcome the problem. Pinzacuá
even faced challenges with the propagation of Gliricidia sepium live fences, which is
commonly done using plant stakes. After strong winds repeatedly unrooted the new
fences, the farm modified its propagation approach planting Gliricidia directly from
seed to obtain stronger roots.
Besides tree-planting, other setbacks are related to human factors. After
transitioning to agroecological methods, the use of chemical inputs –even fertilizers
to facilitate tree growth– has become a difficult choice to make. Although this has led
to creative workarounds like the use of charcoal residue as biochar for the soils,
developing and perfecting new methods has been time consuming. Perhaps one of
the most recurring and persisting frustrations is related to training and retaining of
farm workers. Silvopastoral systems represent a radical departure from conventional
ranching and requires a completely different mindset. Training a new worker to do
things differently requires an investment of time and resources, and losing a trained
worker represents a huge setback. To alleviate labor, Pinzacuá has also experimented
with agri-voluntourism but so far volunteers have been ill-prepared for the difficult
field tasks, and often just interested in how the experience looks on their resume and
on social media.
Over the past 20 years, Pinzacuá has faced a number of setbacks and learned
many important lessons. Moving forward, the main challenges appear to be less
related to increasing or maintaining productivity, and more to the numerous local
socio-economic factors that threaten the farm’s long-term sustainability. Today, the
most significant production cost in the farm is labor. Compared to conventional
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 329

extensive production, silvopastoral systems require more labor for activities such as
herd rotation and tree maintenance. Production costs can be diluted either by
increasing productivity per unit area or by increasing the size of the herd (Holmann
et al. 2003). Although higher productivity in silvopastoral systems allows for higher
stocking rates and lower costs, labor costs are barely covered at the current herd size
and herd expansion is limited by the size of the farm.
Pinzacuá’s proximity to an urban center raises other cost-related concerns. As
land prices increase, so does the opportunity cost of using the property for
conservation-friendly farming, and the pressure to plant cash crops or sell to
developers increases. Proximity to town also exacerbates the labor shortage as
younger generations increasingly reject farming in favor of urban employment
(Calle 2020). Here, as elsewhere in the world, rural life has been downgraded and
youth are concerned with the lack of future and opportunities in the agricultural
sector.
The potential to diversify production and increase farm revenue through the
integration of high-value timber and non-timber products is often highlighted as
one of the benefits of silvopastoral systems (Somarriba 1997; Pezo and Ibrahim
1998). However, Pinzacuá has faced a different reality. While sales of bamboo posts
alone can be up to ten times more profitable than cattle breeding, the excessive
paperwork and bureaucracy required for harvesting and transporting planted timber
make it difficult to realize these diversified revenue streams. The lack of developed
value chains and local markets for both planted native hardwoods and non-timber
forest products currently limits access to these other sources of revenue that could
potentially complement income from livestock. Thus, although ongoing trials with
vanilla, pepper, and dinde show promise, their future will depend on the develop-
ment of local markets.
Being an island of biodiversity in a degraded landscape is certainly reason for
pride, but at times it also feels like an uphill battle. The farm has suffered the
spillover effects from unsustainable practices used in the surroundings. For example,
in only 2 years the apiary, which provides additional revenue, has lost 30% of the
beehives due to agrochemical drift from neighboring farms. In addition, the use of
unconventional methods elicits mockery by other farmers, increasing the sense of
isolation in an already lonely occupation.
Finally, Pinzacuá faces challenges related to monitoring of both productive and
ecological indicators. Basic farm data are essential for sound decision making,
especially in agroecological systems where management—not inputs—determines
productivity, and introducing timely changes is essential. Simple tasks such as
recordkeeping, measuring pasture capacity or tree growth, testing for diet deficien-
cies, and other forms of data collection remains challenging as the farm lacks the
technological and human capacity to systematically compile and analyze this
information.
Likewise, monitoring ecological data in a simple and cost-effective way poses a
challenge moving forward. The studies conducted to date, mostly through collabo-
rations with individual researchers, have sparked the landowners’ interest in under-
standing the underlying ecological processes. Implementing simple protocols to
330 I. Montes-Londoño et al.

consistently monitor key ecological indicators in the long-term would therefore be

desirable. For example, understanding soil dynamics and the impact of organic
management on sensitive groups such as butterflies and amphibians, is of particular
interest. Thus, the challenge will be to build long-term partnerships with universities
or research institutions in order to develop a structured long-term research plan.

12.5.3 Opportunities

Despite these challenges, the farm continues to seek new opportunities to remain
competitive while respecting its commitment to biodiversity-friendly production.
For example, Pinzacuá has already positioned itself as a demonstration farm and
hosts 400+ visitors every year, including local and international groups of school and
university students, farmers, researchers, as well as NGO and government workers.
Aside from providing additional income, these visits foster opportunities to
exchange knowledge and ideas, strengthen networks, and contribute to scale-up
sustainable land use practices across the region (Calle et al. 2013).
Pinzacuá is located within a larger region known as the Coffee Cultural Land-
scape of Colombia, an UNESCO World Heritage site and the country’s second most
popular tourist destination with approximately 100,000 visitors every year (La Patria
2019). Agritourism is already one of the main sources of income across the region,
and the farm could potentially combine nature-based tourism with biodiversity-
friendly farming. Furthermore, ecotourism and proximity to urban centers are
potential new markets for organic produce, which the farm already grows, and direct
marketing of specialty crops such as mushrooms or underutilized species such as
natural dyes or medicinal plants. But this will require efforts to identify short value
chains supplied by many small producers, and to market the services.
Finally, agri-voluntourism and scientific tourism could potentially offer an alter-
native to facilitate the collection and analysis of both productivity and ecological
data. A careful selection process to identify volunteers with the right skills set will be
required to realize this opportunity.

12.6 Conclusion

Pinzacuá’s efforts to increase vegetation cover and complexity by planting and

protecting a variety of trees have not only improved the farms’ productivity but
have also increased its biodiversity. By providing adequate resources for wildlife
species, this previously degraded farm has been transformed into an island of
biodiversity amidst an otherwise impoverished landscape. Furthermore, Pinzacuá
has become a farm that serves as an example for others interested in restorative
agriculture, and a site for biodiversity research that has contributed to our under-
standing of how agroforestry and sustainable management can aid in the recovery of
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 331

degraded farmlands. Today, Pinzacuá stands as living proof that by mimicking

natural ecosystems, the ecological interactions and synergies among the components
of the agroecosystem can be re-established, and land degradation can be reversed to
restore soil productivity, support biodiversity, and recover important ecosystem
services. The process has been long and not without setbacks, but the results have
led to an undeniable transformation of both the land and its people. As new
challenges continue to emerge, Pinzacuá will draw on the lessons learned to continue
applying agroecological principles, and perhaps more importantly, to remain open to
experimenting with new ideas.

References

Amézquita MC, Murgueitio E, Ibrahim RB (2010) Carbon sequestration in pasture and

silvopastoral systems compared with native forests in ecosystems of tropical America. In:
Abberton M, Conant R, Batello C (eds) Grassland carbon sequestration: management, policy
and economics: proceedings of the workshop on the role of grassland carbon sequestration in the
mitigation of climate change. Food and Agriculture Organization of the United Nations
(FAO), Rome
Arias Giraldo LM, Camargo García JC, Cardona Trujillo H (2009) Carbono orgánico edáfico en
rodales de guadua, Guadua angustifolia Kunth., Poaceae y en pasturas arborizadas en la zona
cafetera de Colombia. In: Murgueitio E, Cuartas C, Naranjo J (eds) Ganadería del futuro:
Investigación para el desarrollo. Cali, Fundación CIPAV, pp 246–261
Aynekulu E, Suber M, Zomer R, Mboi D, Arango J, Rosenstock TS (2019) Mitigation benefits from
expansion of trees on rangeland: an analytical proof of concept for Colombia. CCAFS working
paper no. 295. CGIAR Research Program on Climate Change, Agriculture and Food Security
(CCAFS). Wageningen. Available online at: [Link]
Becerra I, Becerra M, Montes N (2017a) Dinámica de mercado: departamento de Quindío. Unidad
de Planificación Rural Agropecuaria, UPRA, Bogotá
Becerra I, Becerra M, Montes N (2017b) Dinámica de mercado: departamento de Caldas. Unidad de
Planificación Rural Agropecuaria, UPRA, Bogotá
Becerra I, Becerra M, Montes N (2017c) Dinámica de mercado: departamento de Risaralda. Unidad
de Planificación Rural Agropecuaria, UPRA, Bogotá
Broom DM, Galindo FM, Murgueitio E (2013) Sustainable, efficient livestock production with high
biodiversity and good welfare for animals. Proc R Soc Biol Sci 280:2013–2025
Calle A (2008) What makes an early adopter? Transforming landscapes one farmer at a time. Trop
Resour 27:7–14
Calle A (2020) Can short-term payments for ecosystem services deliver long-term tree cover
change? Ecosyst Serv 42(C). [Link]
Calle A, Holl KD (2019) Riparian forest recovery following a decade of cattle exclusion in the
Colombian Andes. For Ecol Manage 52(August):10.1016/[Link].2019.117563
Calle Z, Méndez LE (2017) Assessment of the regenerating woody vegetation in Pinzacua.
Unpublished data
Calle Z, Piedrahita L (2007) ¿Cómo diseñar estrategias para el manejo de plantas de interés para la
conservación en paisajes ganaderos? Agroforestería en las Américas 45:117–122
Calle Z, Murgueitio E, Chará J, Molina CH, Zuluaga AF, Calle A (2013) A strategy for scaling-up
intensive silvopastoral systems in Colombia. J Sustain For 32(7):677–693
Calle Z, Molina C CH, Molina D CH, Molina D EJ, Molina E JJ, Murgueitio C B, Murgueitio C A,
Murgueitio R E (2022) A highly productive biodiversity island within a monoculture landscape:
El Hatico nature reserve (Valle del Cauca, Colombia). In: Montagnini F (ed) Biodiversity
332 I. Montes-Londoño et al.

islands: strategies for conservation in human-dominated environments. Topics in Biodiversity

and Conservation, Springer, Cham, pp 279–304
Camargo JC, Cardona GA (2005) Análisis de fragmentos de bosque y guaduales. In: Consultancy
report – regional integrated silvopastoral approaches to ecosystem management project. The
World Bank, GEF, CIPAV, p 38
Chará J, Reyes E, Peri P, Otte J, Arce E, Schneider F (2019) Silvopastoral systems and their
contribution to improved resource use and sustainable development goals: Evidence from Latin
America. FAO, CIPAV and Agri Benchmark, Cali. Retrieved from [Link]
[Link]/fileadmin/templates/res_livestock/docs/2018_Ulaanbataar/Silvopastoral_
Systems_and_their_contribution_to_improved_resource_use_and_SDG.pdf
Chazdon RL, Harvey C, Komar O, Griffith DM, Ferguson BG, Martínez-Ramos M, Morales H,
Nigh R, Soto-Pinto L, van Bruegel M, Philpott S (2009) Beyond reserves: a research agenda for
conserving biodiversity in human-modified tropical landscapes. Biotropica 4:142–153
Chízmar Fernández C (2009) Plantas comestibles de Centroamérica. Instituto Nacional de
Biodiversidad – INBio. Santo Domingo de Heredia, Costa Rica
Cioffi G, Morales-Escobar L, Braca A, De Tommasi N (2003) Antioxidant chalcone glycosides and
flavanones from Maclura (Chlorophora) tinctoria. J Nat Prod 66:1061–1064
Comisión Conjunta (2008) Plan de Ordenación y Manejo Cuenca Hidrográfica (POMCH) del Rio
La Vieja. Corporación Autonóma Regional del Quindío, Corporación Autonóma Regional de
Risaralda, Corporación Autonóma Regional del Valle del Cauca, Cali
Cordero J, Boshier DH (2003) Descripción de Especies: Maclura tinctoria. In: Cordero J, Boshier
DH (eds) Arboles de Centroamérica: un manual para extensionistas. Oxford Forestry Institute
(OFI, Oxford University, Oxford, UK) and Centro Agronómico Tropical de Investigación y
Enseñanza (CATIE), Turrialba, pp 690–697
DANE (2014) Tercer Censo Nacional Agropecuario 2014 Tercer CNA. Departamento
Administrativo Nacional de Estadística, Gobierno Nacional, Bogotá DC
Davis ALV, Scholtz CH, Dooley PW, Bham N, Kryger U (2004) Scarabaeine dung beetles as
indicators of biodiversity, habitat transformation and pest control chemicals in agro-ecosystems.
S Afr J Sci 100:415–424
De Beenhouwer M, Aerts R, Honnay O (2013) A global meta-analysis of the biodiversity and
ecosystem service benefits of coffee and cacao agroforestry. Agric Ecosyst Environ 175:1–7
Delgado M, Pérez C (2018) Proyecciones de actividad económica regional 2017-2021.
Fedesarrollo, Bogotá
Espinal S (1977) Zonas de Vida y Formaciones Vegetales de Colombia, vol. XIII, no. 11. Instituto
Geográfico Agustín Codazzi–IGAC, Bogotá
Etter A (1998) General ecosystem Map of Colombia (1:1,500,000). Instituto Alexander von
Humboldt, Bogotá
FAO (2006) Informe Pecuario. Subdirección de Políticas y Apoyo en Materia de Publicación
Electrónica. FAO, Rome
Gaba S, Bretagnolle F, Rigaud T, Philippot L (2014) Managing biotic interactions for ecological
intensification of agroecosystems. Front Ecol Evol 2. [Link]
Giraldo C, Escobar F, Chará JD, Calle Z (2011) The adoption of silvopastoral systems promotes the
recovery of ecological processes regulated by dung beetles in the Colombian Andes. Insect
Conserv Divers 4:115–122. [Link]
Giraldo C, Hernández M, Giraldo AM, Calle A, Mendivil J, Quevedo C, Velásquez A, Perdomo A,
Castaño K, Giraldo NV, Chará J (2019) Resultados de monitoreo de biodiversidad – Finca de
Referencia Pinzacuá. Proyecto Ganadería Colombiana Sostenible. CIPAV, Cali
Griscom BW, Adams J, Ellis PW, Houghton RA, Lomax G, Miteva DA, Schlesinger WH, Shoch D,
Siikamäki JV et al (2017) Natural climate solutions. Proc Natl Acad Sci 114(44):11645–11650.
[Link]
Harvey CA, Villanueva C, Esquivel H, Gómez R, Ibrahim M, López M, Martinez J, Muñoz D,
Restrepo C, Saénz JC, Villacís J, Sinclair FL (2011) Conservation value of dispersed tree cover
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 333

threatened by pasture management. For Ecol Manag 261(10):1664–1674. [Link]

1016/[Link].2010.11.004
Holmann F, Rivas L, Carulla J, Rivera B, Giraldo LA, Guzman S, Martinez M, Medina A, Farrow A
(2003) Evolution of milk production systems in tropical latin america and its interrelationship
with markets: an analysis of the Colombian case (online). Livest Res Rural Dev 15-
(9) Available at: [Link]
Ibrahim M, Chacón M, Cuartas C, Naranjo J, Ponce G, Vega P, Casasola F, Rojas J (2007) Carbon
storage in soil and biomass in land use systems of ranchlands of Colombia, Costa Rica and
Nicaragua. Agroforestería en las Américas 45:27–36
IUCN Commission on Ecosystem Management (2020) Nature based solutions. Retrieved from
[Link]
based-solutions on May 24, 2020
Kattan GH, Alvarez-López H (1996) Preservation and management of biodiversity in fragmented
landscapes in the Colombian Andes. In: Schelhas J, Greenberg R (eds) Forest patches in tropical
landscapes. Island Press, Washington, D. C
Kattan GH, Franco P, Rojas V (2004) Biological diversification in a complex region: a spatial
analysis of faunistic diversity and biogeography of the Andes of Colombia. J Biogeogr 31:
1829–1839
Kremen C, Merenlender AM (2018) Landscapes that work for biodiversity and people. Science
362(6412):eaau6020
La Patria (2019) Las cuentas del turismo en el Eje Cafetero. Monday, April 15, 2019. Retrieved
from [Link] on
February 25, 2020
López Murillo E (2015) El valor del suelo. Columna de opinión La Cronica del Quindío. Retrieved
on June 1, 2020 from [Link]
valor-del-suelo-cronica-del-quindio-op-11685
Maas B, Clough Y, Tscharntke T (2013) Bats and birds increase crop yield in tropical agroforestry
landscapes. Ecol Lett 16:1480–1487
Martins MM, Setz EZF (2000) Diet of buffy tufted-eared marmosets (Callithrix aurita) in a forest
fragment in southeastern Brazil. Int J Primatol 21:467–476
Mejía Cubillos J (2013) Perfil económico del Eje Cafetero. Análisis con miras a la competitividad
territorial. Adaptación del informe final de la consultoría “Competitividad territorial de la
Ecorregión Eje Cafetero. Análisis desde el perfil económico del Eje Cafetero como aporte a la
línea de base de la Agenda de Desarrollo Sostenible” realizada para Universidad Tecnológica de
Pereira. MPRA Paper No. 43873. Retrieved on June 1, 2020 from [Link]
[Link]/43873/
Mendenhall C, Karp D, Meyer C, Hadly E, Daily G (2014) Predicting biodiversity change and
averting collapse in agricultural landscapes. Nature 509:213–217. [Link]
nature13139
Mesa Arboleda HF (2009) Balance de gases de efecto invernadero en un modelo de producción de
ganadería doble propósito con alternativas silvopastoriles en Yaracuy, Venezuela. MS thesis.
CATIE, Turrialba, 225 p
Mittermeier RA, Turner WR, Larsen FW, Brooks TM, Gascon C (2011) Global biodiversity
conservation: the critical role of hotspots. In: Zachos FE, Habel JC (eds) Biodiversity hotspots.
Springer, London, pp 3–22
Moebius-Clune BN, Moebius-Clune DJ, Gugino BK, Idowu OJ, Schindelbeck RR, Ristow AJ, van
Es HM, Thies JE, Shayler HA, McBride MB, Kurtz KSM, Wolfe DW, Abawi GS (2016)
Comprehensive assessment of soil health – the Cornell framework, Edition 3.2. Cornell
University, Ithaca
Montagnini F, Ibrahim M, Murgueitio E (2013) Silvopastoral systems and climate change mitiga-
tion in Latin America. Bois et forêts des Tropiques 316:3–16
Montes-Londoño (2019) Monitoring of species richness of butterfly in Pinzacua through Visual
Encounterying surveying. Unpublished data
334 I. Montes-Londoño et al.

Montes Londoño I, Rodríguez Susa MS (2011) Evaluación de la capacidad de autodepuración de

una quebrada con bosque de guadua ribereño en la cuenca del Río La Vieja. Undergraduate
thesis. Universidad de Los Andes, Bogotá
Montes-Londoño I, Montagnini F, Ashton MS (2017) Allometric relationships and reforestation
guidelines for Maclura tinctoria, an important multi-purpose timber tree of Latin America. New
For. [Link]
Murgueitio E, Calle Z, Uribe F, Calle A, Solorio B (2011) Native trees and shrubs for the productive
rehabilitation of tropical cattle ranching lands. For Ecol Manag 261(10):1654–1663
Nair PKR, Nair VD, Kumar BM, Showalter JM (2010) Carbon sequestration in agroforestry
systems. Adv Agron 108:237–307
Naranjo JF, Cuartas CA, Murgueitio E, Chará J, Barahona R (2012) Balance de gases de efecto
invernadero en sistemas silvopastoriles intensivos con Leucaena leucocephala en Colombia.
Livestock research for rural development, 24, Article #150. Retrieved May 31, 2020 from http://
[Link]/lrrd24/8/[Link]
Pagiola S, Ríos AR (2013) Evaluation of the impact of payments for environmental services on land
use change in Quindío, Colombia. PES learning papers. World Bank, Washington DC
Pagiola S, Honey-Rosés J, Freire-González J (2020) Assessing the permanence of land-use change
induced by payments for environmental services: evidence from Nicaragua. Trop Conserv Sci
13:1940082920922676
Perfecto I, Vandermeer J (2010) The agricultural matrix as alternative to the land sparing/agriculture
intensification model. PNAS 107:5786–5791
Peters VE, Greenberg R (2013) Fruit supplementation affects birds but not arthropod predation by
birds in Costa Rican agroforestry systems. Biotropica 45:102–110
Pezo D, Ibrahim M (1998) Sistemas Silvopastoriles. Materiales de Enseñanza no. 44/CATIE
CATIE-GTZ Turrialba, 276 pp
Philpott SM, Bichier P (2012) Effects of shade tree removal on birds in coffee agroecosystems in
Chiapas, Mexico. Agric Ecosyst Environ 149:171–180
Piotto D, Montagnini F, Kanninen M, Ugalde L, Viquez E (2004) Forest plantations in Costa Rica
and Nicaragua: performance of species and preferences of farmers. J Sustain For 18(4):59–77
Poch TJ, Simonetti JA (2013) Ecosystem services in human-dominated landscapes: insectivory in
agroforestry systems. Agrofor Syst 87:871–879
Prevedello JA, Almeida-Gomes M, Lindenmayer DB (2018) The importance of scattered trees for
biodiversity conservation: a global meta-analysis. J Appl Ecol 55:205–214
Rangel A (1949) Maderas industriales de Colombia Caribb For 10(3):161–162
Reyes E, Bellagamba A, Molina JJ, Izquierdo L, Deblitz C, Chará J, Mitchell L, Romanowicz B,
Gómez M, Murgueitio E (2017) Measuring sustainability on cattle ranches. Working paper on
Silvopastoral Systems. Agribenchmark, CIPAV, Fedegan, World Animal Protection. Retrieved
from [Link]
Rivers MC, Barstow M, Mark J (2017) Maclura tinctoria. The IUCN red list of threatened species
2017: e.T61886731A61886745. Retrieved from [Link]
[Link]. Feb 29, 2020
Rodríguez N, Armenteras D, Morales M, Romero M (2004) Ecosistemas de los Andes
Colombianos. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt,
Bogotá
Roig JT (1974) Plantas medicinales, aromáticas o venenosas de Cuba. Ed Ciencia y Técnica,
Instituto del Libro. La Habana, Cuba
Sadeghian S, Murgueitio E, Mejía C, Rivera JM (2001) Ordenamiento ambiental y reglamentación
del uso y manejo del suelo en la zona cafetera. En: Suelos del eje cafetero. Universidad
Tecnológica de Pereira, GTZ, Fondo Editorial del Departamento de Risaralda, Pereira, pp
96–108
Sánchez Gómez EL, Camargo García JC (2015) Diversidad de avifauna en paisajes rurales de la
cuenca del río La Vieja, Eje Cafetero de Colombia. Recursos Naturales y Ambiente:83–89
Somarriba E (1997) Pastoreo bajo plantaciones forestales. Agroforestería en las Américas 4:26–28
12 Hacienda Pinzacuá: An Example of Regenerative Agriculture Amidst. . . 335

Suárez A, Williams-Linera G, Trejo C, Valdez-Hernández JI, Cetina-Alcala VM, Vibrans H (2012)

Local knowledge helps select species for forest restoration in a tropical dry forest of central
Veracruz, Mexico. Agrofor Syst 85:35–55
Toro Zuluaga G (2005) Eje Cafetero colombiano: compleja historia de caficultura, violencia y
desplazamiento. Documento preparado para el Congreso 2004 de la Asociación de Estudios
Latinoamericanos (LASA), Las Vegas, Nevada, octubre 7-9, 2004. Revista de Ciencias
Humanas UTP 35: 127–149
Tscharntke T, Clough Y, Wanger TC, Jackson L, Motzke I, Perfecto I, Vandermeer J, Whitbread A
(2012) Global food security, biodiversity conservation and the future of agricultural intensifi-
cation. Biol Conserv 151:53–59
Unidad de Planificación Rural Agropecuaria, UPRA (2019a) Gestión de información agropecuaria
y planificación del desarrollo agropecuario: Qundío. Retrieved on June 1, 2020 from https://
[Link]/file/d/1Wnmc5v8CGRi0q0-Hz65bAqIV_4yjZL0V/view
Unidad de Planificación Rural Agropecuaria, UPRA (2019b) Gestión de información agropecuaria
y planificación del desarrollo agropecuario: Caldas. Retrieved on June 1, 2020 from https://
[Link]/file/d/11013KahlDDAN628zQrY6lR0OcQBvQaD9/view
Unidad de Planificación Rural Agropecuaria, UPRA (2019c) Gestión de información agropecuaria
y planificación del desarrollo agropecuario: Risaralda. Retrieved on June 1, 2020 from https://
[Link]/file/d/1cgTexaHDDyXOrl8QhhLeH4bjx3aackm4/view
United Nations Development Programme (UNDP) (2004) Informe Regional de Desarrollo
Humano. PNUD, Manizáles
van Breugel M, Hall JS, Craven DJ, Gregoire TG, Park A, Dent DH, Wishnie MH, Mariscal E,
Deago J, Ibarra D, Cedeño N, Ashton MS (2011) Early growth and survival of 49 tropical tree
species across four sites differing in soil fertility and rainfall. For Ecol Manag 261:1580–1589
Vílchez S, Harvey C, Sáenz JC, Casanoves F, Carvajal JP, Villalobos JG, Hernandez B, Medina A,
Montero J, Merlo D, Sinclair FL (2013) Consistency in bird use of tree cover across tropical
agricultural landscapes. Ecol Appl 24:158–168
Wishnie MH, Dent DH, Mariscal E, Deago J, Cedeño N, Ibarra D, Condit R, Ashton MS (2007)
Performance of 24 tropical tree species in relation to reforestation strategies in Panama. For Ecol
Manag 243:39–49
Chapter 13
Islands of Trees in Long-Fragmented
Landscapes in Great Britain

Keith J. Kirby

Abstract The British countryside contains scattered ‘islands’ of ancient woodland

and groups of ancient trees in ‘seas’ of managed farmland and plantations of
non-native trees. However, do they function like islands for the plants and insects
particularly associated with these patches? Is the absence of species from newer
islands because of dispersal limitations, or because their habitat requirements are not
met in newly formed sites?
Past fragmentation may mean remnant patches hold more species than can be
sustained in the longer-term. However, if the fragmentation of the original wildwood
happened hundreds, even thousands of years ago, the woodland species that have
survived and are valued today, of necessity must be able to live in relatively small
patches with a predominance of edge conditions. Some can also be found associated
with individual trees and hedges scattered through the intervening farmland.
The island analogy is useful but has limitations in guiding future conservation
practice. The extent and pattern of patches as perceived by researchers, i.e. what is
mapped as woodland or wood-pasture, may be very different to the actual patch size
used by the species. In managed landscapes there is the possibility of creating or
restoring habitats in the places that appear to offer the best opportunities for allowing
species movement through a landscape. Colonisation credits should be as important
a focus as extinction debts. This might involve translocating ground flora plants to
new woods and deliberately damaging mature trees to expedite the decay processes
normally associated with ancient trees.

Keywords Ancient woodland · Parkland · Saproxylic beetles · Vascular plants ·

Veteran trees · Wood-pasture · Woodland management

K. J. Kirby (*)
Department of Plant Sciences, University of Oxford, Oxford, UK
e-mail: [Link]@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 337
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
338 K. J. Kirby

13.1 Introduction

During the 1980s, a new law, The Wildlife and Countryside Act, was introduced in
Great Britain. This law gave much stronger protection to the series of sites identified
by the state nature conservation agencies as important for biodiversity (Sheail 1998).
These areas (examples of woodland, heathland, meadow, wetland, etc.) covered
about 7% of the land surface. However, despite this and subsequent stronger
legislation to protect such sites, many species continued to decline at a national
scale (Defra 2017). It was recognised that conservation needed to work across the
countryside as a whole.
The principles and approaches of landscape ecology, e.g. Forman and Godron
(1986) gained ground amongst the conservation movement in Great Britain during
the 1980s. Ideas based around island biogeography theory seemed particularly
appropriate to the British conditions. Frequently there are small patches of habitat
with a high biodiversity value (the “islands”) distributed amongst large areas of low
biodiversity habitat (the “seas”) consisting of arable crops, highly-improved grass-
land or monocultural plantations of introduced trees managed for production of food
and fibre (Peterken 2002) (Fig. 13.1). The enduring influence of landscape ecolog-
ical concepts can be seen in Fig. 13.2 based on an influential report on the state of

Fig. 13.1 A view of Devon, south-west Britain, showing ‘islands’ of woodland in an agricultural
‘sea’. (Photo: Keith Kirby)
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 339

Fig. 13.2 Enhancing ecological networks based on Making Space for Nature. (Lawton et al. 2010)
Approaches needed include: (a) improving the quality of habitat patches; (b) making existing sites
bigger; enhancing connectivity through (c) a new woodland corridor or (d) stepping stone patches;
(e) creating new woods; reducing pressures on sites through (f) establishing grass-scrub buffers and
ecotones, or (g) more general biodiversity enhancement in the surrounding farmland.

England’s protected sites produced for the Department of Environment, Food and
Rural Affairs (Defra) (Lawton et al. 2010).
This chapter explores the degree to which patches of ancient woodland and wood-
pasture behave as ‘island’ habitats in the cultural landscapes of Great Britain.
Peterken (1977, 1993) and Rackham (1976, 2003) summarised the conservation
importance of ancient woodland for ecological and cultural reasons. The significance
of veteran and ancient trees, especially oak in Britain – often occurring in more open
conditions such as parkland – for a range of different species groups such as fungi,
lichens and saproxylic invertebrates has been highlighted by Rose (1976, 1993),
Harding and Rose (1986) and contributions in Farjon (2017).
My focus is on two species groups: vascular plants that have a high affinity with
ancient woodland; and saproxylic invertebrates such as the stag beetle, Sinodendron
cylindricum, that are strongly associated with a continuity of ancient trees such as are
often found in wood-pastures (Table 13.1). However, the real world is more variable
in space and time than the conditions and assumptions that underpinned some
landscape models.
340 K. J. Kirby

Table 13.1 Some terms used in this paper to describe the tree and woodland resource in Britain
‘Wood’, ‘woodland’ and ‘forest’ are used generally to describe tree-covered lands.
‘Forest’ (capital F) is also used in a more specialised sense where it refers to land subject to Forest
law in the medieval period; the land to which it is applied might or might not be covered by trees.
‘Wood-pasture’ refers to landscapes grazed by domestic stock or deer with an open tree cover.
‘Wildwood’ is used for the pre-Neolithic landscape (prior to c.7000 years BP) when tree-cover
was probably at its most extensive in Britain.
‘Ancient woods’ (or ancient forests) are those where there has been continuous woodland cover
since 1600 AD in England and Wales and 1750 AD in Scotland.
‘Coppicing’ refers to the practice of repeatedly cutting trees close to ground level, at intervals of
between five and thirty years, then allowing regrowth of multiple stems from the stump to provide
the next harvest. ‘Pollarding’ is a similar process, but the cut is made at 2-3 m above the ground so
that the regrowth is out of the reach of browsing animals.
‘Ancient trees’ are those old for their species with features such as cavities or a hollow trunk, bark
loss over sections of the trunk and a large quantity of dead wood in the canopy. The broader term
‘veteran trees’ includes younger individuals that have developed similar characteristics, perhaps
due to adverse growing conditions or injury.
‘Hedges/hedgerows’ are lines of (usually thorny) shrubs managed to provide boundaries between
fields. Large mature trees are often scattered along the hedge.
‘Saproxylic species’ are those that live on or in dead and decaying wood, with deadwood beetles
being one of the most studied groups.

13.2 An Ancient Island Pattern

During the Holocene, much of Britain came to be tree-covered below about 300 m
altitude, reaching its maximum extent about 7000 years BP (Kirby and Watkins
2015). Thereafter, tree cover dropped as farming developed and land was cleared for
crops and for livestock grazing. By 1000 years BP there may have been less than
20% cover; clearance continued and c.100 years BP woodland cover was about
4–5%. There had been some large-scale planting of new forests in the nineteenth
century and this accelerated during the twentieth century such that woodland cover is
currently about 13% (Forestry Commission 2019). However, patches of woodland
composed of native trees, mainly broadleaved species, tend to be small, less than
50 ha.
Early studies confirmed the expectation that larger patches of native trees do tend
to contain more species than smaller patches for butterflies, birds, and vascular plants
(Moore and Hooper 1975, Shreeve and Mason 1980, Usher et al. 1992). The history
of the patches is also important. More of the specialised woodland vascular plant
species occur in patches that have existed for several hundred years at least (ancient
woods, see Table 13.1 and Peterken (1977)). The distribution of ancient woods
across Britain has been mapped by the Nature Conservancy Council and its succes-
sors, as illustrated in Fig. 13.3 (Goldberg et al. 2011, Goldberg 2015).
Ancient woods do contain generalist plant species such as brambles (Rubus
fruticosus) that are widespread through the rest of the landscape. However ancient
woods are distinctive in being richer than more recently created woodland of a
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 341

Fig. 13.3 The distribution of ancient and recent woodland in part of eastern England. The
intervening land is arable and improved grassland. In the last 150 years the development of the
recent woodland means that the ancient woods are now less isolated than they were. (Based on data
from the Ancient Woodland Inventory (Spencer and Kirby 1992))

similar extent in a suite of vascular plants that are generally more shade-tolerant,
stress-tolerant and poor-colonisers (often apparently reliant on ants for dispersal,
e.g. Anemone nemorosa, Mercurialis perennis, Primula vulgaris) compared to the
non-woodland flora (Peterken 1974, 2000, Kirby et al. 2012, Kimberley et al. 2013).
Peterken and Game (1984) showed that for these vascular plants new woods that had
developed adjacent to ancient woodland were richer than more remote new woods,
suggesting isolation as an important factor. Similar findings have been reported from
elsewhere in north-western Europe, e.g. Brunet and Von Oheimb (1998).
Britain also has many areas of open ancient wood-pasture, with large-crowned,
open-grown trees, generally old parks and Royal Forests, grazed by deer or livestock
(Kirby and Perry 2014). These may or may not be mapped as woodlands depending
on the extent of canopy cover (Fig. 13.4), but they include many of the sites of
highest value for epiphytic lichens (e.g Lobaria pulmonaria, L. amplissima,
Thelotrema lepadinum) and for saproxylic invertebrates (e.g. Ampedus
cinnabarinus, Gnorimus variabilis, Limoniscus violaceus) (Harding and Rose
1986). Both specialist lichens and invertebrates appear to be very specific in their
requirements for dead wood conditions and have poor rates of dispersal (Rose 1993,
Scheidegger and Werth 2009, Irmler et al. 2010). For these species, the extent of the
site (hectares) is less important than the number of suitable habitat trees, which may
be scattered or clustered across a larger or smaller grazed area. There is still a broad
relationship between site size and richness because large sites tend to have more
342 K. J. Kirby

Fig. 13.4 (a) Glenamara Park, Cumbria: an upland wood-pasture showing areas of scattered and
clustered veteran trees; (b) Moccas Park, Herefordshire: a lowland wood-pasture with scattered
trees. (Photos: Keith Kirby)
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 343

trees and a greater variety within the tree population in terms of species and
structural forms, but the area effect is an indirect one.

13.3 Real Islands Versus Tree Patches

Real islands are surrounded by a matrix habitat, the sea or another water body, that
cannot support the terrestrial island species. By contrast, the matrix habitats (farm-
land, plantations) around ancient woods and parklands often can support some
woodland or wood-pasture species albeit generally at low population levels. Spe-
cialist woodland flora do occur in and spread along hedges between fields; ancient
trees, with associated decaying wood fauna, may similarly occur in fields and along
boundaries (Fig. 13.5).
During the last two centuries, particularly since the end of World War II, there
have been losses of many of these microhabitats in the matrix through agricultural
intensification and increased fragmentation of patches by major roads and railway
lines, e.g. Peterken and Allison (1989). This has increased the effective isolation of
the woodland and wood-pasture patches. However, there have also been substantial
increases in woodland and tree cover as a consequence of changing forestry policies

Fig. 13.5 A farmed landscape near Oxford, with abundant hedges and hedgerow trees that might
act as habitat for woodland species. (Photo: Keith Kirby)
344 K. J. Kirby

during the twentieth century that should eventually increase the available habitat
extent and reduce effective isolation between woods. An increase in new woodland
is a poor substitute for the loss of long-established or ancient woods, but new farm
woods can within a century start to harbour woodland specialist vascular plants
(Usher et al. 1992, Harmer et al. 2001).
Saproxylic species may benefit from the aging of oak trees established in the
nineteenth Century along hedges and in small woods, or from those left to grow on
as timber trees in woods that were once cut-over regularly as coppice, but in most
cases were largely neglected in the late twentieth century. These trees are or will
soon be moving into the age and size classes where decaying wood starts to form
hollows in the main trunk.

13.4 Recent Versus Long-Fragmented Systems

Fragmentation models often start with a large more-or-less continuous block of

habitat – say a forest – which is then split up into small isolated blocks leading to
the loss of species with large area requirements; a loss of ‘interior’ species and an
increase in ‘edge’ species (the latter generally seen as less valuable); and an ongoing
loss of other species for a while – the extinction debt – because the smaller patches
created by fragmentation initially contain more species than can be sustained in the
reduced area. How well does this model fit with current-day ancient woods and
wood-pastures?
In Britain there is debate as to what was the nature of the ‘continuous habitat’ –
the wildwood of the pre-Neolithic era – from which our ancient woodland and wood-
pasture have been derived. If much of the country were closed forest (Peterken
1996), perhaps similar to that in Bialowieza National Park, Poland (Falinski 1986),
then the initial clearance and fragmentation effects would have led to the sorts of
major changes in species richness suggested by models and studies of recent
fragmentation of tropical forests. However, if the wildwood was a more open
mixture of blocks of tree cover, scrub and open grassland, such as proposed by
Vera (2000), then the contrast between the wildwood and modern landscapes
(particularly those prior to the mid-twentieth century agricultural intensification)
would be much less and we might suppose that the changes in species abundances
would be correspondingly lower. This debate has not been fully resolved.
The initial clearance and fragmentation happened centuries or millenia ago. Any
species that required large blocks of continuous woodland would be unlikely to have
survived into the historic period. We have lost most of the larger wild mammals
(Alces alces, Ursus arctos, Bos primigenius, Lynx lynx, Canis lupus, Sus scrofa,
Castor fiber) at least in part from reductions in their potential habitat area (Yalden
1999), but our ancient woods have then had 500–1000 years to adapt to their
absence. Even some very small woods may hold diverse and valued assemblages
of ancient woodland plants. Furthermore, these may now be threatened by the recent
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 345

Table 13.2 Ellenberg’s Light Scores as modified for British conditions by Hill et al. (2004) for
British woodland specialist vascular plants (Kirby et al. 2012)
Ellenberg’s Number of woodland specialist
score Meaning (Ellenberg 1988) species with that score
1 Plants in deep shade, may be less than 1%, 0
seldom more than 30% relative light.
2 Between 1 and 3 2
3 Shade plants, mostly less than 5% relative light, 111
but also in lighter places.
4 Between 3 and 5 39
5 Plants of half shade, rarely in full shade but 48
generally more than 10% relative light.
6 Between 5 and 7. 36
7 Plants generally in well-lit places but also occur 23
in partial shade.
8 Light loving plants, rarely found where there is 7
less than 40% relative light.
9 Plants in full light, found only in full sun; rarely 0
in less than 50% relative light.

spread of large herbivores (mainly deer, both native and introduced) and wild boar
(escaped from farms), e.g. Kirby (2001), Sims et al. (2014).
Our native broadleaved woods are small. Riutta et al. (2014) calculated that even
if woods less than 2 ha were excluded, about 45% of the total woodland area of
England was within 60 m of an open edge. Many of the plants that we seek to
conserve are ‘edge’ species, found in partially open conditions, rather than deep
shade interior species (Table 13.2).
Similarly, many of the best places for saproxylic beetle assemblages are where the
oaks are growing in relatively open, sunny conditions – edge type environments,
with a different suite of species to that found where ancient trees occur in dense
stands. This is one argument in favour of the Frans Vera’s model of the wildwood. In
his model transitions from open to closed cover, i.e. ‘edge’ could have been
widespread as part of the natural state.
High levels of edge conditions may not be therefore a negative feature of British
ancient woods and wood-pastures, as they have been a characteristic that has existed
for such a long time. However, there are new threats associated with edges that have
developed in the last century. The most significant of these are the potential for
increased eutrophication at edges (from emissions from roads, farmland etc); pesti-
cide drift in from adjacent farmland; and increased drying-out of the edge zone under
climate change (Pitcairn et al. 2002, Gove et al. 2007, Riutta et al. 2012).
When large areas of forest are broken-up, leaving smaller patches, these patches
may initially contain more species than can be sustained in the long-term, so that
they then gradually lose species. This phenomenon is known as ‘extinction debt.’
Over much of Britain, however, ancient woods and patches of ancient trees have
been relatively stable in size and isolated in the landscape for centuries. Their species
346 K. J. Kirby

richness is likely to be in equilibrium with their current area. Peterken and Game
(1984) did not find any evidence for an extinction debt in woods that had been
recently reduced in area compared to ancient woods whose extent had long been
stable, although Vellend et al. (2006) did find such an effect in ancient woods studied
in Belgium. Even if, at the species level there is spread to new sites, this may include
only part of the genetic variation that is present in the source population
(Scheidegger and Werth 2009), such that there could still be a genetic debt.
Of greater significance for British conditions may be how to make the most of the
‘colonisation credit’ that is building up. Over much of Britain the extent of
broadleaved woodland has been increasing over the last century, but not all the
species that this larger area can support have yet colonised it. Spread of specialist
woodland plants to new sites does occur but can be slow even when they are directly
adjacent to an ancient woodland source (Peterken and Game 1984, Rackham 2003).
Should we therefore deliberately sow such species into new woods e.g. Francis and
Morton (2001), Worrell and Francis (2003)? Conservation managers have also been
experimenting with whether the processes of wood decay in mature trees can be
speeded up through management (Lonsdale 2013). The trees are deliberately dam-
aged by cutting into the bark in order to bring forward the time when they might be
suitable for colonisation by the saproxylic species characteristic of ancient trees.
There are periodic discussions as to whether it is better to have a single large
patch as a reserve or several smaller patches (SLOSS debate (Tjørve 2010)). In long-
fragmented landscapes where the species assemblages are in equilibrium with patch
area some of the arguments for preferring the single large reserve are less relevant.
Collections of small woods often contain more species than a single site of the same
total area because they are, for example, spread across different soil types or are in
different ownerships and hence have different past management and woodland
structures. Recent research results suggest that small patches can have greater
importance for biodiversity conservation than previously anticipated (Montagnini
et al. 2022). Results from the Lacandona rainforest of Mexico, where the effect of
forest patch size on species density of different taxonomic groups was examined, add
to the increasing evidence that, on a per-sample area basis, small patches are
valuable for conservation of forest-specialist species, and are not the near-exclusive
habitat of generalist species (Arroyo-Rodríguez et al. 2022).

13.5 Natural Disturbances Versus Regular Human

Intervention

Pickett and Thompson (1978) discuss the idea of the minimum dynamic area in
nature reserve design. A reserve needs to be large enough that it can always include
patches of recent disturbance, whatever form that disturbance takes. Many woodland
plants including some ancient woodland species are mainly found in the temporary
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 347

Fig. 13.6 Recently cut areas often show increased flowering of many woodland specialist plants,
the yellow of Lamiastrum galeobdolon contrasting with the blue of Hyacinthoides non-scripta.
(Photo: Keith Kirby)

open gaps (Fig. 13.6), so ideally woods should be big enough that there are always
some trees falling and creating gaps.
This concept can be difficult to apply with natural woodland disturbance patterns
because gap creating events are generally unpredictable in size and frequency.
Where woods are managed and the disturbance is from felling the trees, then the
minimum dynamic area can be specified more precisely. For example, many ancient
woods in England were managed by the coppice system in the past where stands
were cut on rotations of between about 5 and 30 years. Often key species are found
mainly in the first few years after cutting (Buckley and Mills 2015a, b) (Fig. 13.6). If
we assume that a minimum gap extent of 1 ha is required for the key species and that
gaps remain suitable as habitat for 2 years, then an area of 1 ha must be felled at least
every 2 years. If the trees are managed on a coppice rotation of 20 years that implies
a minimum woodland area of 10 ha; but if the trees are to be grown for 100 years as
high forest, 50 ha of woodland would be needed. If the woodland extent is fixed
(which is usually the case) the procedure can be used to calculate the frequency of
gap creation cutting needed to sustain the gap species.
In the example given above, the rotation length in a 20-ha wood should not be
more than 40 years. Most ancient woods are less than 20 ha (Spencer and Kirby
1992) so the shift from coppice management (5–30 year rotations) to high forest
348 K. J. Kirby

(60–120 year rotations) over the last 80 years has generally been accompanied by
reductions in the more light-demanding elements of the vascular plant flora because
the frequency of gap creation has been reduced, e.g. Kirby et al. (2005).
An additional consideration is how close a patch that is becoming suitable needs
to be to a source patch for it to be useful. The butterfly Melitaea athalia in south-east
England was commonly found in recently cut coppiced areas where one of its larval
food plants, Melampyrum pratense, grew and flowered in abundance for a few years
before declining as the canopy reformed. However, while adults readily colonised
new patches within 300 m of a source patch, they rarely did so when the patches
were 600 m or more apart (Warren 1987).
The same approach can be taken for any other groups of species that are found
largely in one stage in the tree/stand life cycle. For saproxylic species, the minimum
dynamic area is that that can contain sufficient trees across all generations to ensure
there are some that have developed sufficient maturity to replace the current ancient
trees when they finally die (Kirby 2015). For example, if the aim is to sustain two
ancient trees (>400 years old) per hectare and on average half the trees in any age
cohort die, then this might mean 32 trees per hectare (<100 years old), 16 trees
(101–200 years old), 8 trees (201–300 years old); and 4 trees (301–400 years old).
Most wood-pasture sites in Britain do not contain such a balanced population
structure. The potential younger age classes of trees as do exist in the countryside
may be too far from the current ancient trees for reliable colonisation to be assumed,
although rare long-distance colonisation events cannot be ruled out, partly because
they can be very difficult to detect (Jonsell et al. 2003).

13.6 Cost-Dispersal Distances and Developing Networks

Arable fields, i.e. those lands that are under use for crops, or hold vegetation of low
stature that allows for ploughing, are likely to be less favourable for the spread to
new sites of both woodland specialist plants and saproxylic beetles than, for exam-
ple, fields of tall grass with scattered scrub. Measures of how ‘joined-up’ the
landscape is, that is, the likelihood of species being able to move from one site to
another, should take into account the nature of the intervening landscape as well as
the absolute distance between patches (Adriaensen et al. 2003). This is then used to
measure the effective l distance between two woodland patches to give a better idea
of the degree of connectivity within a landscape. Such analyses might suggest
whether species populations in a group of woods were likely to be linked through
exchange of individuals and genes; or they might be used in locating new woodland
to provide the most useful stepping stones or linkages for species dispersal through
the landscape (Catchpole 2006, Latham et al. 2013).
The results from such modelling can inform the various initiatives seeking to
establish networks of sites across the country (Crick et al. 2020). For example, which
of the current habitat patches may be most critical in allowing species to spread,
where are there significant gaps or bottlenecks that hinder dispersal? Practical
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 349

conservation action can be targeted to key locations, for example to improve the
condition of a group of old trees that are an important stepping stone between two
important wood-pastures, to create new woodlands and hedges to provide potential
links between ancient woods, or to encourage the growth of new trees along existing
hedges where currently the trees are widely spaced.

13.7 Conclusions

Landscape ecology has proved a useful framework for looking at how individual
patches of woodland or wood-pasture relate to the ecological processes taking place
across the surrounding landscape. Among the lessons that have emerged from their
application to our long-fragmented cultural landscapes are the following.
The extent and pattern of patches as perceived by researchers, i.e. what is mapped
as woodland or wood-pasture, may be very different to the actual patch size used by
the species. The effective patch size may be smaller because a species uses only part
of the woodland cycle, e.g. the vascular plants found mainly in the short-lived gap
phase, or the beetles found only in the trees over 300 years-old in a parkland; or
larger because a species can also use part of the adjacent matrix. Patch boundaries,
their extent and pattern in the landscape are species specific.
The processes of species colonisation of new woodland are not simply the reverse
of those that apply when a continuous piece of habitat is broken up: if a piece of
ancient woodland is converted to an arable field, the specialist woodland plant
species may be lost overnight. If an arable field is converted to woodland, it may
take centuries for some of those specialist species to colonise that area. A 400-year-
old tree can be felled in a day, but it will take many years before a replacement young
tree has grown sufficiently to build up the equivalent levels of decaying wood habitat
(Watts et al. 2020).
In old cultural landscapes, most of the land has been managed in the past and
species occurrence may be closely linked to how habitats such as ancient woods and
wood-pastures have been treated. Some species may survive in smaller patches than
might be expected under more natural conditions because controlled disturbances
may allow a smaller minimum dynamic area.
In managed landscapes there is the possibility of creating or restoring habitats in
the places that appear to offer the best opportunities for allowing species movement
through a landscape. In planning restoration strategies, colonisation credits should
be as important a focus as extinction debts. This might involve translocating ground
flora plants to new woods and deliberately damaging mature trees to expedite the
decay processes normally associated with ancient trees.
It is difficult to detect and allow for rare long-distance dispersal, but even so we
can increase the likelihood of its success simply by building up source populations in
existing woods and wood-pastures through improved habitat management; and
increasing the extent of new woodland and wood-pasture patches in the target area.
350 K. J. Kirby

References

Adriaensen F, Chardon JP, De Blust G, Swinnen E, Villalba S, Gulinck H, Matthysen E (2003) The
application of ‘least-cost’ modelling as a functional landscape model. Landsc Urban Plan 64:
233–247
Arroyo-Rodríguez V, Arasa-Gisbert R, Arce-Peña N, Cervantes-López MJ, Cudney-Valenzuela SJ,
Galán-Acedo C, Hernández-Ruedas MA, San-José M, Fahrig L (2022) The importance of small
rainforest patches for biodiversity conservation: a multi-taxonomic assessment in the Lacandona
region, Mexico. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Topics in Biodiversity and Conservation, Springer, Cham,
pp 41–60
Brunet J, Von Oheimb G (1998) Migration of vascular plants to secondary woodlands in southern
Sweden. J Ecol 86:429–438
Buckley G, Mills J (2015a) Coppice silviculture: from the mesolithic to 21st century. In: Kirby KJ,
Watkins C (eds) Europe’s changing woods and forests: from wildwood to managed landscapes.
CABI, Wallingford, pp 77–92
Buckley G, Mills J (2015b) The flora and fauna of coppice woods: winners and losers of active
management or neglect. In: Kirby KJ, Watkins C (eds) Europe’s changing woods and forests:
from wildwood to managed landscapes. CABI, Wallingford, pp 129–139
Catchpole RDJ (2006) Planning for biodiversity – opportunity mapping and habitat networks in
practice: a technical guide. English Nature Research Reports Number 687, Northminster House,
Peterborough PE1 1UA
Crick H, Crosher I, Mainstone C, Taylor S, Wharton A, Langford P, Larwood J, Lusardi J,
Appleton D, Brotherton P, Macgregor N (2020) Nature networks evidence handbook, Natural
England Research Report 81. Natural England, Peterborough
Defra, Department for Environment, Food and Rural Affairs (2017) Wild bird populations in the
UK 1970 to 2016. Defra, London
Ellenberg H (1988) The vegetation ecology of Central Europe. Cambridge University Press,
Cambridge
Falinski JB (1986) Vegetation dynamics in temperate lowland primeval forests. W. Junk, Dordrecht
Farjon A (2017) Ancient oaks in the English landscape. Royal Botanic Gardens, Kew, Surrey. For-
estry Commission (2019) Forestry Statistics 2019. Forestry Commission, Edinburgh
Forman RTT, Godron M (1986) Landscape ecology. Wiley, New York
Francis JL, Morton A (2001) Enhancement of amenity woodland field layers in Milton Keynes.
British Wildlife 12:244–251
Goldberg EA (2015) The UK’s ancient woodland inventory and its use. In: Kirby KJ, Watkins C
(eds) Europe’s changing woods and forests: from wildwood to managed landscapes. CABI,
Wallingford, pp 326–336
Goldberg EA, Peterken GF, Kirby KJ (2011) Origin and evolution of the ancient woodland
inventory. British Wildlife 23:90–96
Gove B, Power SA, Buckley GP, Ghazoul J (2007) Effects of herbicide spray drift and fertilizer
overspread on selected species of woodland ground flora: comparison between -term and long-
term impact assessments and field surveys. J Appl Ecol 44:374–384
Harding PT, Rose F (1986) Pasture-woodlands in Lowland Britain. Institute of Terrestrial Ecology
Abbots Ripton, Huntingdon
Harmer R, Peterken GF, Kerr G, Poulton P (2001) Vegetation changes during 100 years of
development of two secondary woodlands on abandoned arable land. Biol Conserv 101:291–
304
Hill MO, Preston CD, Roy DB (2004) PLANTATT: attributes of British and Irish plants. Biological
Records Centre (NERC), Cambridge
Irmler U, Arp H, Nötzold R (2010) Species richness of saproxylic beetles in woodlands is affected
by dispersion ability of species, age and stand size. J Insect Conserv 14:227–235
13 Islands of Trees in Long-Fragmented Landscapes in Great Britain 351

Jonsell M, Schroeder M, Larsson T (2003) The saproxylic beetle Bolitophagus reticulatus: its
frequency in managed forests, attraction to volatiles and flight period. Ecography 26:421–428
Kimberley A, Blackburn GA, Whyatt JD, Kirby KJ, Smart SM (2013) Identifying the trait
syndromes of conservation indicator species: how distinct are British ancient woodland indica-
tor plants from other woodland species? Appl Veg Sci 16:667–675
Kirby KJ (2001) The impact of deer on the ground flora of British broadleaved woodland. Forestry
74:219–229
Kirby KJ (2015) What might a sustainable population of trees in wood-pasture sites look like?
Hacquetia 14:43–52
Kirby KJ, Perry SC (2014) Institutional arrangements of wood-pasture management: past and
present (UK). In: Hartel T, Pleininger T (eds) European wood-pastures in transition: a social-
ecological approach. Earthscan/Routledge, Abingdon, pp 254–270
Kirby KJ, Watkins C (eds) (2015) Europe’s changing woods and forests: from wildwood to
managed landscapes. CABI, Wallingford
Kirby KJ, Smart SM, Black HJ, Bunce RGH, Corney PM, Smithers RJ (2005) Long-term
ecological changes in British woodland (1971-2001), English Nature Research Report 653.
English Nature, Peterborough
Kirby KJ, Pyatt DG, Rodwell JS (2012) Characterization of the woodland flora and woodland
communities in Britain using Ellenberg values and functional analysis. In: Rotherham ID,
Jones M, Handley C (eds) Working and walking in the footsteps of ghosts: volume 1 the
wooded landscape. Wildtrack Publishing, Sheffield
Latham J, Sherry J, Rothwell J (2013) Ecological connectivity and biodiversity prioritisation in the
terrestrial environment of Wales. CCW Staff Science Report No. 13/3/3, Terrestrial Ecosystem
Group, Countryside Council for Wales, Bangor
Lawton JH, Brotherton PNM, Brown VK, Elphick C, Fitter AH, Forshaw J, Haddow RW,
Hilborne S, Leafe RN, Mace GM, Southgate MP, Sutherland WJ, Tew TE, Varley J, Wynne
GR (2010) Making space for nature: a review of England’s wildlife sites and ecological
network. Defra, London
Lonsdale D (2013) Ancient and other veteran trees:further guidance on management. The Tree
Council, London
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity Islands: Strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human-dominated environments. Topics in Biodiversity and Conservation,
Springer, Cham, pp 1–35
Moore NW, Hooper MD (1975) On the number of bird species in British woods. Biol Conserv 8:
239–250
Peterken GF (1974) A method for assessing woodland flora for conservation using indicator
species. Biol Conserv 6:239–245
Peterken GF (1977) Habitat conservation priorities in British and European woodlands. Biol
Conserv 11:223–236
Peterken GF (1993) Woodland conservation and management, 2nd edn. Chapman and Hall,
London
Peterken GF (1996) Natural woodland: ecology and conservation in northern temperate regions.
Cambridge University Press, Cambridge
Peterken GF (2000) Identifying ancient woodland using vascular plant indicators. British Wildlife
11:153–158
Peterken GF (2002) Reversing the habitat fragmentation of British woodlands. WWF-UK,
Godlaming
Peterken GF, Allison H (1989) Woods, trees and hedges: a review of changes in the British
countryside. Nature Conservancy Council, Peterborough
Peterken GF, Game M (1984) Historical factors affecting the number and distribution of vascular
plant species in the woodlands of Central Lincolnshire. J Ecol 72:155–182
352 K. J. Kirby

Pickett STA, Thompson JN (1978) Patch dynamics and the design of nature reserves. Biol Conserv
13:27–37
Pitcairn C, Skiba U, Sutton M, Fowler D, Munro R, Kennedy V (2002) Defining the spatial impacts
of poultry farm ammonia emissions on species composition of adjacent woodland groundflora
using Ellenberg Nitrogen Index, nitrous oxide and nitric oxide emissions and foliar nitrogen as
marker variables. Environ Pollution 119:9–21
Rackham O (1976) Trees and woodland in the British landscape. Dent, London
Rackham O (2003) Ancient woodland: its history, vegetation and uses in England (revised ed.).
Castlepoint Press, Dalbeattie
Riutta T, Slade EM, Bebber DP, Taylor ME, Malhi Y, Riordan P, Macdonald DW, Morecroft MD
(2012) Experimental evidence for the interacting effects of forest edge, moisture and soil
macrofauna on leaf litter decomposition. Soil Biol Bioche 49:124–131
Riutta T, Slade EM, Morecroft MD, Bebber DP, Malhi Y (2014) Living on the edge: quantifying
the structure of a fragmented forest landscape in England. Landsc Ecol 29:949–961
Rose F (1976) Lichenological indicators of age and environmental continuity in woodlands. In:
Brown DH, Hawksworth DL, Bailey RH (eds) Lichenology: progress and problems. Academic,
London
Rose F (1993) Ancient British woodlands and their epiphytes. British Wildlife 5:83–93
Scheidegger C, Werth S (2009) Conservation strategies for lichens: insights from population
biology. Fungal Biol Rev 23:55–66
Sheail J (1998) Nature conservation in Britain - the formative years. The Stationery Office, London
Shreeve T, Mason C (1980) The number of butterfly species in woodlands. Oecologia 45:414–418
Sims NK, John EA, Stewart AJ (2014) Short-term response and recovery of bluebells
(Hyacinthoides non-scripta) after rooting by wild boar (Sus scrofa). Plant Ecol 215:1409–1416
Spencer JW, Kirby KJ (1992) An inventory of ancient woodland for England and Wales. Biol
Conserv 62:77–93
Tjørve E (2010) How to resolve the SLOSS debate: lessons from species-diversity models. J
Theoretical Biol 264:604–612
Usher MB, Brown AC, Bedford SE (1992) Plant species richness in farm woodlands. Forestry 65:
1–13
Vellend M, Verheyen K, Jacquemyn H, Kolb A, Van Calster H, Peterken GF, Hermy M (2006)
Extinction debt of forest plants persists for more than a century following habitat fragmentation.
Ecology 87:542–548
Vera FWM (2000) Grazing ecology and forest history. CABI, Wallingford
Warren M (1987) The ecology and conservation of the heath fritillary butterfly, Mellicta athalia. III.
Population dynamics and the effect of habitat management. J Appl Ecol:499–513
Watts K, Whytock RC, Park KJ, Fuentes-Montemayor E, Macgregor NA, Duffield S, Mcgowan PJ
(2020) Ecological time lags and the journey towards conservation success. Nature Ecol Evol 4:
304–311
Worrell R, Francis J (2003) Establishing woodland flora in broadleaf and conifer woodlands. Scott
For 57:203–210
Yalden D (1999) The history of British mammals. Poyser, London
Chapter 14
Natural Landscape of the Pampa Region
in Santa Fe Province, Argentina:
Environmental Resilience and Opportunity
for Changing the Agri-Food Paradigm

Libertario Hugo González, Germán Neffen, Victoria Benedetto,

Marta Sánchez Miñarro, Andrea García, Ricardo Biasatti, Pablo Rimoldi,
Cristian Alesio, and Daniel Paiz

Abstract The Argentinean humid pampas were once a large grassland biome.
Starting when the European colonizers arrived five centuries ago, and lasting until
present, the original grassland ecosystems suffered an extreme transformation and
have now almost completely disappeared. Today this region is under pressure from
an “agriculture without farmers” dependent on external inputs which are increas-
ingly scarce. This has serious social, environmental and economic implications. The

The human being disconnected from nature, conceived it as an object of knowledge, then as an
object of domination and finally as a mere commodity. [...] With the advance of modernity, a
paradigm of the individual was consolidated whose relationship with the “others” is one of
domination and exploitation, a concept of the individual as an uprooted being, with weakened
and fragmented collective ties (Gauna Zotter and Rey 2017)

L. H. González (*) · V. Benedetto · A. García

Experiment Station (EEA) INTA Oliveros, Santa Fe, Argentina
e-mail: [Link]@[Link]
G. Neffen
Agricultural Producer, Pergamino, Buenos Aires, Argentina
M. Sánchez Miñarro
Economist and Director of the School of Social Entrepreneurships, Municipality of Rosario,
Santa Fe, Argentina
R. Biasatti
Faculty of Veterinary Sciences, National University of Rosario, Santa Fe, Argentina
P. Rimoldi
Chair of General Zoology, Faculty of Agricultural Sciences, National University of Rosario,
Santa Fe, Argentina
C. Alesio · D. Paiz
Chair of Biology and Ecology, Faculty of Veterinary Sciences, National University of Rosario,
Santa Fe, Argentina

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 353
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
354 L. H. González et al.

landscape matrix has been increasingly alternating its seasonal composition between
chemical fallows and monocultures of wheat, corn, and soybean. If allowed to
continue, this artificialization of ecosystems, dependent on fossil fuels, chemical
inputs, and biotechnology, represents a serious threat to present and future social and
environmental wellbeing. We have identified surviving land areas that persisted in
the increasingly anthropic landscape of this region, constituting patches of biodiver-
sity referred to in this chapter as “islands of resilience.” In this chapter, we propose a
multidisciplinary approach which utilizes the existing resilience islands in rural areas
and in bordering urban zones. We analyze a case study located in the city of Las
Rosas (Santa Fe province, Argentina) and the land’s current and potential ecological
wellbeing. We examined several socioeconomic and agroecological strategies asso-
ciated with the production of local healthy food supplied to simplified marketing
frameworks. These strategies are developed with production-consumption, commu-
nity, and cooperative practices in mind. In this case study, we are guided by
principles of cooperation, mutual support and solidarity, which build the foundation
for an eco-social and resilient transition to more widespread biodiversity friendly
agricultural landscapes.

Keywords Agri-food system · Agroforestry systems · Biodiversity Islands ·

Common goods · Landscape ecology · Production models · Sustainability

14.1 Introduction

The term “islands of resilience” arises from empirical observation of the recurrent
vegetation that stands out in industrial agriculture’s landscape matrix. The tree,
shrub and herbaceous composition of these islands is unexpectedly dynamic, making
the islands a symbol of resistance to a dominant agricultural production model that
transformed natural ecosystems. These islands or patches have varied forms, includ-
ing circular, ovoid and irregular. They contrast with the extensive degraded check-
erboard plains of the humid pampas of southern Santa Fe, Argentina (Fig. 14.1).
In addition to disrupting human-dominated land, resilience islands also reflect the
complex structure and functionality of ecosystems that maintain stability, despite
external disturbances. Satellite images of these resilience islands can allow observers
to develop their own reasoning for the persistence of the patches. But it is in situ,
when travelling along provincial and national routes, when one can appreciate their
presence and observe the different stages of plant succession in which these relics are
found. This chapter provides a detailed description of the history of the landscape
and the current state of ecological wellbeing of the islands in order to offer a logical
explanation for their persistence.
In this chapter we describe the change over time of the natural landscape matrix of
the Argentinean humid pampas. This land transformation is depicted through the key
historical events of the southern region of the Santa Fe province, from the original
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 355

Fig. 14.1 Island of resilience inserted in the matrix of the dominant landscape. (Photo: Pablo
Olive)

Fig. 14.2 Ecosystems recovery capacity in the Pampean region of Santa Fe, Argentina. (Photos:
Ricardo Biasatti) (see text for explanation)

predominance of the pastures to the implementation of an extensive system of

monocultures. This chapter utilizes research and information on the flora and
fauna that are crucial to aiding in the regeneration of resilience islands. In addition,
the chapter integrates agroecological proposals based on the experience of the
agroecology module of the National Institute of Agricultural Technology (INTA,
Oliveros Experiment Station) (Benedetto et al. 2017). These publications and pro-
posals demonstrate the capacity of ecosystems to recover their structural and func-
tional parameters after land degradation due to external pressures (Biasatti et al.
2013, 2016) (Fig. 14.2).
The images in Fig. 14.2 correspond to the physiognomy of the landscape that is
the subject of this work. The first image represents the matrix of the landscape that
predominates in the region, in which the agro-productive activity has entirely
modified the environment. The second image shows a representative example of
356 L. H. González et al.

an island of biodiversity. The third image shows the role of biological corridors
(a stream in the area) contributing to landscape connectivity. For our case study, this
capacity for recovery or resilience is a starting point for developing a new model of
the pampean rurality, including agricultural production areas with greater biodiver-
sity and landscape connectivity.

14.2 Passages from the History of the Santa Fe Pampa

Region

14.2.1 The New Configuration of the Landscape Matrix

Our analysis of landscape transformation in the Pampa region included climate

change as an influencer of structural change in the landscape matrix, and humans
in their role as decision makers on whether to work with nature or oppose it. In the
study region, indigenous people used to work with nature. Evidence of mega-fauna
grazing activities, dated at circa (ca.) 12,200–10,000 years, proves that the inhabi-
tants of the Pampean zone were hunters (Politis et al. 2014). Their assimilation to the
territory and movement in search of food was part of the pre-Hispanic culture before
the Spanish conquest of the region.
The year 1492 was a turning point in the history of this hemisphere of the world.
America did not appear on any map prior, so the Spanish and Portuguese, as the first
European occupants, named the continent and initiated a territorial organization
similar to that already existing in Spain and Portugal (Mignolo 2007). The
European arrival in the Americas was a crucial event that led to a history of
European imperial expansion and developed the existence of a European lifestyle
that became prominent in Argentine society. The Americas emerged in the European
consciousness as a great expanse of land to be conquered with a people to be
evangelized and exploited (Mignolo 2007).
The start of imperialistic expansion into this region led to the first signs of the
fragmentation of its pristine landscape. This event initiated the fragmentation seen
throughout the history of the Pampa, where the colonizers began to transform the
unaltered South American landscape as they had similarly done in their own land in
Europe. This fragmentation was due in part to the introduction of new animal species
such as cattle and horses by the Spanish upon their arrival, which significantly
altered the ecosystems of the pampean grassland biome (Brailovsky 2006).
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 357

14.2.2 Territory Configuration or Landscape

Fragmentation?

In two and a half centuries of colonial hegemony, the independence movement,

which started in 1810 in Argentina, grew significantly due to an economic power
imbalance between the locals and the colonizers (Donghi 2014). At the beginning of
the nineteenth century, a new configuration of the economy and politics was
structured around Buenos Aires, Argentina, in the context of an economic junction
favorable to the trade of cattle hides. The litoral region (riverine coastal area) which
had been exhausted as a supplier of leathers for overseas markets, was replaced by
the Buenos Aires markets, which profited as the cattle industry rapidly expanded,
providing an opportunity for territorial expansion (Donghi 2000).
As the wealth and power of Buenos Aires increased, its political decision makers
planned and executed military campaigns of extermination and subjugation of the
native inhabitants, with the aim of expanding the border that was under their control.
The result of these campaigns was an immense increase of available land, which the
Argentine government publicly offered in “emphyteusis”. This system consists of a
contract for the use of the land in exchange for taxing the production. This implies
the temporary cession of the land, but use of this land ultimately became perpetual,
requiring very low tax payments (Donghi 2000).
However, these newly acquired lands were only granted to landowners of the
lands of old colonization, political and military leaders of the provinces, and holders
of urban mercantile wealth. None of the lands’ original inhabitants were given
parcels. These elite groups formed the social nucleus of the landowning class
(Donghi 2000). In this way, the system of latifundia (mega-landowners) was con-
solidated, resulting in the fragmentation of the landscape matrix, as proprietors
implemented barriers such as fences to contain the cattle and to delimit private
property.

14.2.3 Benefit or Penalty? Geopolitics of Sacrifice

At the end of the nineteenth century, the Argentine government, led by President
Nicolás Avellaneda, continued its military campaigns, taking the lands of the
aborigines to be exploited. However, as the Argentine government expanded its
control over land in the region, the available territory to be conquered diminished.
Finally, on October 5, 1878, Law 947 was passed in order to obtain funds for the
“Conquest of the Desert” to further expand the territory under Argentine control (Del
Corro 2003). Avellaneda affirmed that the elimination of Native American
populations in the frontiers was meant to allow Argentine people to “populate the
desert.” He defined settling as the replacement of the original inhabitants who were
not linked to the national and international markets by others who were. Avellaneda
358 L. H. González et al.

further explained: “to govern is to populate, and to populate is to change nomads for
sedentary salaried workers to develop a new livestock production model”
(Brailovsky and Foguelman 2009).
Under the impetus of a sustained British demand for foods and raw materials, the
leading sectors oriented the use of natural resources to meet global market demands
(Brailovsky and Foguelman 2009). Thus, between 1860 and 1930 Argentina entered
the global market as a producer of meat, wool and cereals. Argentina’s growing
involvement in international markets led the country to be considered “the granary of
the world” (Brailovsky and Foguelman 2009). The same president, two years before
Law 947 was passed, passed law N
817, which regulated immigration and coloni-
zation, helping to provide the region with a labor force and further grow Argentina’s
economy (Sili and Soumoulou 2011).
As a consequence, at the beginning of the twentieth century, the Argentine
agricultural sector had been separated into two distinct facets. One was the ranchers
who owned the land, most of them dedicated to cattle raising. The other was small
and medium-sized farmers who produced cereals and lived in the colonies, either
using their own land or renting out plots in the large ranches (Sili and Soumoulou
2011).

14.2.4 History of Agriculture in the Santa Fe Pampa

The history of agriculture in the Santa Fe province of Argentina is strongly tied to the
history of the work of immigrant families and their descendants (Cloquell et al.
2007). These actors played a leading role in the transformation of the landscape,
which is evident in their productive activities and in the increase in population over
time. Over almost 40 years, the proportion of immigrants grew from 10% of the total
population in 1858 to 42% in 1895 (Cloquell et al. 2007).
While older production methods did contribute to the fragmentation of the
landscape matrix, they were not as harmful as subsequent production methods
utilized at the end of the 20th and beginning of the twenty-first century. In contrast,
the first settlers’ belief toward nature was that they not only “lived in a territory,” but
they “lived from the territory” they inhabited. This ideology explains why these
original landowners did not decimate their habitat through their productive actions
because of their care for the environment.

[Link] Argentina as “The Granary of the World”

The expansion of Argentina’s borders, the introduction of machinery, and massive

immigration caused exportation of cereals and flax to increase, allowing Argentina to
prosper greatly and become known as “the granary of the world” (Volkind 2009).
Unlike the United States or Canada, Argentina’s agricultural policy did not encour-
age small and medium independent producers, but rather aimed at incorporating the
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 359

land into the national agricultural economy by creating incentives for owners to put it
into production (Bidaseca and Lapegna 2006). This resulted in an asymmetric
relationship between the producer, who leases land, and the owner, who signs
short-term leases and has the flexibility to frequently increase rental fees (Cloquell
et al. 2007). However, producers held the belief that they were lessees only tempo-
rarily, as one must first lease the property before eventually owning it. Their goal
was to transition from being a part of the labor force to being owner-producers,
thereby transitioning their immigrant status (Cloquell et al. 2007).
It is important to note that some contemporaries of the time, such as the geogra-
pher and naturalist Kropotkin, in his 1892 book “The Conquest of Bread,” warned of
what was happening in that region of the world. He explained that as the productivity
of land increases, the workers’ profits do not increase to the same degree as the
profits of the landowners and the government. This inequity results from an increase
in taxes towards the farmers as their production rises (Kropotkin 1977).
This era could have been a time in Argentina’s history for equitable redistribution
of land among farmers, in consonance with the social struggles that were taking
place, in which the access, use, and occupation of the land for production and
habitation were disputed. However, in the framework of the international economic
crisis of 1930, a mass migration of farmers occurred from the countryside to the city
due to their financial fragility and dependence on international prices, resulting in the
concentration of land into the hands of few (Cloquell et al. 2007).
Towards 1940, a new government policy enabled productive units under lease to
be bought by the tenant producer, either by benefits of credit policies, or by tacit
agreements due to the situation (Cloquell et al. 2007). Despite the struggle for
control of this land, the Pampean landscape was still relatively untamed up to this
point in time. However, the land was subsequently quickly transformed and
fragmentated. The islands of resilience, which are the subject of this chapter, are
surviving relics of this process, and they are exemplified by our case study, the farm
located in Las Rosas, home to the Fontana family, in the south of the Santa Fe
province.
These trends of transformation and fragmentation continued to the end of the
1950s, when the national political and economic situation changed abruptly. This
societal change was brought about by an international context that, concluding the
post-war reconstruction stage, sought to reduce the role of the government in the
national economy and to promote the liberation of economic relations (Cloquell et al.
2007). Along with this change, new strategic scientific and technological advances
were promoted. Agricultural production was given strong incentives to fulfill its
traditional role as a provider of foreign exchange to the national economy. These
incentives launched a new phase of capital accumulation that furthered Argentina’s
role in the international market (Cloquell et al. 2007).
360 L. H. González et al.

[Link] The “Modernization” of the Agricultural Sector

Around 1960, the region experienced increases in production and productivity based
on the application of the industrial-based technological model. This began the
process of “modernization,” which led the agriculture sector to maximize the process
of capital accumulation, with different impacts on the social actors of the agriculture
industry (Cloquell et al. 2007).
In our case study island of resilience in Las Rosas, the recollections of a family
member, Angel Fontana, serve to illustrate this transition. As Angel explains, in his
farm they used to share equipment with neighbors, using herbicides and planting
non-hybrid seeds. Later on, due to the challenges of harvesting crops in the 1970s, in
addition to the impacts of weeds, insects, and other pests, they incorporated dairy
farming to have a more regular income. Over time, they began to rely more on
additional contract workers and rental equipment to face increasing challenges.
As Angel relates, “In those days, in some farms they worked the land with horses,
but not in ours where there was already a small tractor, which we shared with other
family members so that they could work on their own fields. In the early days, some
seed was selected at the time of harvest for replanting, but little by little the
commercial hybrid seed gained ground. Herbicides were there as long as I can
remember, but the fertilizers arrived much later. Being almost a teenager, we started
with my father a small dairy farm to get a monthly income that made it possible for
us not to wait until the agricultural harvest. Milk delivery was not a problem, since
the creameries sent the truck to the house. This experience was from 1970 to 1980,
when I saw different crops grow, potatoes, watermelons, melons and fruit trees. Over
time it was impossible to fight off weeds, insects and other pests. In the sixties,
someone used to pass by with a vehicle to get eggs, milk, and chickens and offered in
exchange sugar, and oil in smaller quantities because we used pork fat (the product
of periodic meatings). Then I saw the incorporation of wheat, corn and soybean
planting. The soil was broken and planting was implemented with the use of zero-
tillage or direct sowing. Through the years everything changed. From having our
own tools we moved on to the work done by a contractor and rental that continues to
this day” (Angel Fontana, personal communication).
This first-person experience provides evidence of the gradual incorporation of
new agricultural technologies, collectively known as “the green revolution”. The
integration of these technologies was promoted by government agencies, facilitating
the advancement of private companies into the region. This integration of new inputs
and technologies, which was more prevalent in crops than in cattle raising, began in
the second half of the 1950s, grew in the 1970s and 1980s, and was accentuated in
the 1990s (Pizarro 2003).
With the start of the green revolution, new agricultural practices began to be
utilized across the landscape matrix in order to increase profits. Among these new
practices, the increasingly common use of monocultures and limited productive
diversification led to a destructive occupation of the land. In fact, other qualities of
the land, such as its social or cultural benefits, were disregarded as productive value
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 361

became the dominant productive focus. In this mindset for the use of the territory,
land was considered socially disposable, simply designating it as “sacrifice areas” for
the sake of progress (Svampa and Viale 2014).
Agribusiness, and especially transgenic soybeans (Glycine max), is currently the
core of Argentina’ s extractive matrix. Currently, the country is among the four main
world producers of transgenic soybeans with almost 24 million hectares cultivated.
Since the end of the 1990s, when the use of transgenic soy was approved, the
expansion of agribusiness was tied to the use of genetically-modified organisms
(GMOs), which led to a global restructuring of the traditional agrarian ways
(Svampa and Viale 2020).
This new agricultural model spread not only in the Pampas region, but also in
marginal areas, such as in the northern and coastal regions of Argentina. Today it
occupies 23 to 33 million hectares of land, of which 90% is dedicated to soybeans.
The great concentration of transgenics and agrochemicals across this soybean-
dominated land presents dangers to the country due to prolonged exposure to these
chemicals (Svampa and Viale 2020). The massive scale of this exposure within the
territory and on the bodies of people constitutes a true model of “bad-development”
similar to the Chernobyl case, which illustrates the dangers and their disregard by
governments and economic actors involved (Svampa and Viale 2020).

14.2.5 Resilience of the Santa Fe Pampean Ecosystem

Throughout the history of the Pampa plains of the Santa Fe province, there have been
significant changes to the landscape, the results of which will still play out over years
to come. The era in which human actions on their environment started to dominate
the landscape led to the designation of the time of the Anthropocene (Herrero 2017).
During this era, the Pampean grassland biome was transformed into an
agroecosystem in which the economic-financial goals were prioritized. This priori-
tization disregarded the multiple dimensions that must be addressed for a productive
system that is sustainable over time and that allows humans to monitor their actions
on the environment and its natural commons such as water, soil, air, and
biodiversity.
Our case study shows that the establishment of resilient systems in the Pampean
grasslands, with the aim of restoring biotic communities congruent with the original
landscape physiognomy, has already been observed in the region. While we were
performing our work for this chapter, some restrictions on land use were established.
This allowed for the development of plant secondary succession that in relatively
short periods of time (2 to 4 years) has shown the system’s capacity to recover
spontaneous biodiversity, increase soil coverage, and allow a successive occupation
of the space with progressive substitution of species (Biasatti et al. 2013, 2016).
362 L. H. González et al.

Other observations of these resilience islands have revealed their potential for
developing new specific ecological niches and a variety of new habitats or micro-
habitats according to local soil, hydrological and even climatic variations (Biasatti
et al. 2013, 2016). This evidence of the land’s persistence encourages us to assess the
viability of this land for the development of agroecologically-based production
practices. These new practices would help strengthen food sovereignty and security,
improving human welfare and benefitting other forms of life in the context of the
dominant agricultural matrix. In addition, this proposal could be extrapolated to
other regions of the country and to other parts of the world which are withstanding
similar ecological conditions.
The next sections of the chapter offer a characterization of the islands of resil-
ience, focusing on their flora and fauna, and arguing for their integration into the
landscape matrix. We propose to apply agroecological principles to crop manage-
ment around the islands, and we relate experiences of local markets using agroeco-
logical production. The path we choose, among the different paths that intersect in
human development, can undoubtedly determine the future of our species for
centuries to come (Bookchin 2019).

14.3 The Islands of Resilience and their Context in South

Santa Fe
14.3.1 Recovering Areas with Real Potential for Hosting
Biodiversity

In this section, islands of resilience are defined from the perspective of landscape
ecology as “patches”, as they are areas with greater biodiversity than the surrounding
anthropic matrix, which is dominated by an extensive homogenized and simplified
territory associated with the agro-industrial production model or system of soy
cultivation (Forman and Grodon 1986, Biasatti et al. 2007). These patches can
take many forms, such as the one described in this chapter, as well as marginal
lands, floodable lands, and lands with limitations for agriculture.
Our case study provides evidence of the depopulation of rural areas due to the
development of soybean production methods over the last decades. As previously
explained, in the Argentinean Humid Pampas, farming originally consisted of mixed
production, with the producer’s family living in the countryside and producing a
variety of goods and foods. The arrival of soybean cultivation introduced a new
model of production with less reliance on people, which led to the migration of the
rural settler and the abandonment of their homes and surroundings. This new model
of agricultural production, which heavily relied on the soybean, replaced diverse
cropping systems and became dominant.
These abandoned settlements were largely left untouched, which allowed for the
development of plant secondary succession, sheltered by the infrastructure that had
been left behind and its generally wooded surroundings. The abandoned areas
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 363

contained exotic species such as fruit trees and others that would provide shade,
shelter from the winds and some wood products, such as firewood for cooking and
heating.
Not all of these abandoned relics have been preserved. However, it is still
common to observe these “patches” in the pampas landscape of monoculture,
where nature reveals its biotic potential. This shows the capacity of resilience of
the Pampean grasslands to recolonize these relics through successive occupation of
space with spontaneous species, starting with pioneers and then moving into stages
of greater forest maturity.
The province of Santa Fe has several examples that show the resilience of the
Pampean grasslands. Government strategies have created several biological corri-
dors, rescuing relics of the original ecosystems. These have quickly shown their
effectiveness for restoration, not only of plant species, but also of animals, including
birds and mammals of medium and large size. These species also can take advantage
of corridors to move around the landscape and establish themselves in recovered
spaces.
The conservation model that utilizes these strategies is based on the configuration
of a “reticulated system for the conservation of biological diversity” through the
implementation of biological corridors around lotic water bodies (rivers and streams)
or communication routes, roads, and railways (Cracco and Guerrero 2004, Biasatti
et al. 2013). The implementation of this proposal was carried out by preventing
human intervention in the areas of study and through the monitoring and manage-
ment of spontaneous secondary succession. In none of the case studies was second-
ary succession induced by the planting of species or the transfer of animals to
increase biological richness. Instead, succession was self-generated by the favorable
conditions created by protective barriers. Corridors also aided in ecological succes-
sion by connecting previously isolated relics which developed into existing patches,
or “islands of resilience.”
Due to the large-scale agricultural use of the humid Pampa, human intervention
has led to a significant transformation, including the creation of “resiliency islands”
in an agricultural landscape. Human domination in this region has reached levels that
exceed 95% of the available land used for productive purposes or for urban areas.
The large scale of human occupancy has transformed the landscape into a new
anthropic and homogeneous matrix. Contrary to other regions, the strategy necessary
to conserve biodiversity and restore land in the humid pampas requires the fragmen-
tation of the anthropized matrix. This phenomenon was studied and developed under
the concept of “inverse fragmentation,” within the framework of a strategy aimed at
establishing mechanisms compatible with the joint practices of production and
conservation (Biasatti et al. 2019).
“Resiliency islands” are key elements of this strategy because they help recover
spaces with real potential to host biodiversity. Many animal species typical of the
Pampean grasslands, such as the Pampean fox (Lycalopex gymnocercus), the wild
cat (Leopardus geoffroyi), or the coipo (Myocastor coypus) are threatened because
they do not cross cultivated environments. However, the presence of natural vege-
tation corridors allows these species to reach these biodiversity patches, aiding in
364 L. H. González et al.

Fig. 14.3 The island of resilience studied and its immediate environment, highlighting the contrast
in the configuration of the landscape of the region. (Google image)

species longevity (Rimoldi and Chimento 2018, Biasatti et al. 2019). Plant species
can also thrive in these areas, moving from one habitat patch to another as they are
carried by wind or animals that move seeds or propagules (eggs, larvae, vegetatively
reproducing plant parts) through corridors. This dispersal mode has the additional
advantage of enriching the corridors. Biological corridors facilitate the movement of
native species even in landscapes that are severely hostile due to their high degree of
homogenization and habitat simplification generated by the expansion of the agri-
cultural frontier.
Since the aesthetic or ethical appeal for protection alone may not suffice, the
conservation of “resiliency islands” requires the application of sustainable produc-
tive practices. In addition to providing potential productive benefits, these islands
contribute to economic development by maintaining species and their genetics,
aiding in species diversification and promoting variability in an otherwise uniform
landscape (Biasatti et al. 2016, Biasatti et al. 2017).
The previously mentioned model of a “reticulated system for the conservation of
biological diversity” constitutes a strategy that relies on islands of resilience inte-
grated into a network of connectivity devices designed as biological corridors. This
relationship between patches and corridors creates a system capable of strengthening
both the dispersion and the colonization of spontaneous species in a highly
anthropized productive region that invests material and energy in maintaining
broad spaces with minimal biodiversity (i.e. crops). “Resiliency islands,” by their
definition, are isolated from other biodiversity, as can be seen in Fig. 14.3. However,
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 365

Fig. 14.4 The island of resilience and its potential links to strengthen a system that increases
connectivity, allows the transfer of spontaneous species, and promotes biodiversity and coexistence
with alternative production systems. (Satellite image, Google Earth font)

these patches can be part of a large, interconnected web of islands that utilize
corridors (Fig. 14.4). This network constitutes a viable and effective alternative for
establishing a new system of production that allows for the conservation of regional
biodiversity. From the spatial unit of the island of resilience that is the subject of this
chapter (see Fig. 14.1) it is possible to extend the scope by zooming out to a regional
scale in which the sum of other islands linked by biological corridors constitutes a
viable and effective alternative for establishing new models of production and
conservation, as can be seen in Fig. 14.3.

14.3.2 Case Study in the Center-South of the Province

of Santa Fe

Our case study is an island of resilience that covers an area of approximately one
hectare, located in the south-central province of Santa Fe (32
310 0600 S 61
330 2600 W),
near the town of Las Rosas and a few meters from National Route
No. 178 (Fig. 14.5). With regards to purely anthropic structures, there is a mill and
366 L. H. González et al.

Fig. 14.5 Province of Santa Fe in Argentina. (Google Image)

a small “tapera” (abandoned rural housing), which can potentially be used as a

refuge by the local fauna.
This area of study corresponds to the “humid pampas” or “pampas grasslands”
biome which is in a transition to an “espinal” (thorny scrub) biome. The region has a
temperate climate of marked seasonality, average temperatures between 17–18
C,
an annual rainfall close to 1000 mm, and soils with a low slope and a well-developed
A horizon. The land in this area has good quality and productive potential, which has
led to its transformation for agricultural purposes.
In the studied island of resilience, the dominant tree species is the paper mulberry
(Broussonetia papyrifera). Other less abundant species include eucalyptus (Euca-
lyptus sp.), paradise tree (Melia azedarach), evergreen (Ligustrun sp.), three-thorned
acacia (Gleditsia triacanthos), olive tree (Olea europaea), peach tree (Prunus
persica), fig tree (Ficus carica), Cereus sp., ceibo (Erythrina crista-galli), ombú
(Phytolacca dioica), and palo borracho (Chorisia sp.). At one edge of the island
there is a small group (about 20m2) of canes (Arundo donax). Young individuals of
cina cina (Parkinsonia aculeata) and carob tree (Prosopis sp.) were planted to enrich
the patch with native species and fruit trees like apple trees (Malus domestica) for
their food production (Table 14.1). Below the tree canopy and in areas without tree
cover, communities of herbaceous species cover the soil.
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 367

Table 14.1 Plant species recorded in island of resiliency in Las Rosas, Santa Fe, Argentina
Family Scientific name Origin/ Observations
Amaranthaceae Amaranthus hybridus L. ssp. Hybridus Native/ Direct
observation
Apiaceae Bowlesia incana Ruiz &Pav. Native/ Direct
observation
Asteraceae Ambrosia cumanensis Kunth Exotic/ Direct
observation
Artemisia absinthium L. Exotic/ Direct
observation
Carduus acanthoides L. Exotic/ Direct
observation
Carduus nutans L. Exotic/ Direct
observation
Conyza bonariensis (L.) Cronquist var. Native/ Direct
bonariensis observation
Cynara scolymus L. Exotic/ Direct
observation
Helianthus tuberosus L. Exotic/ Direct
observation
Sonchus oleraceus L. Exotic/ Direct
observation
Boraginaceae Borago officinalis L. Exotic/ Direct
observation
Brassicaceae Raphanus sativus L. Exotic/ Direct
observation
Cactaceae Cereus argentinensis Britton & Rose Native/ Direct
observation
Commelinaceae Commelina erecta L. Native/ Direct
observation
Cannaceae Canna indica L. Native/ Direct
observation
Convolvulaceae Ipomoea grandifolia (Dammer) O´Donell Native/ Direct
observation
Euphorbiaceae Euphorbia peplus L. Exotic/ Direct
observation
Fabaceae Parkinsonia aculeata L. Native/ Direct
observation
Erythrina crista-galli L. var. crista-galli Native/ Direct
observation
Prosopis alba Griseb. var. alba Native/ Direct
observation
Gleditsia triacanthos L. Exotic/ Direct
observation
Malvaceae Sida rhombifolia L. Native/ Direct
observation
Ceiba speciosa (A. St.-Hil.) Ravenna Native/ Direct
observation
(continued)
368 L. H. González et al.

Table 14.1 (continued)

Family Scientific name Origin/ Observations
Meliaceae Melia azedarach L. Exotic/ Direct
observation
Moraceae Broussonetia papyrifera (L.) Vent. Exotic/ Direct
observation
Morus alba L. Exotic/ Direct
observation
Ficus carica L. Exotic/ Direct
observation
Oleaceae Olea europaea L Exotic/ Direct
observation
Phytolaccaceae Phytolacca dioica L. Native/ Direct
observation
Poaceae Bromus catharticus Vahl var. Catharticus Native/ Direct
observation
Sorghum halepense (L.) Pers. var. Halepensé Exotic/ Direct
observation
Cynodon dactylon (L.) Pers. var. Dactylon Exotic/ Direct
observation
Nassella neesiana (Trin. &Rupr.) Barkworth Native/ Direct
observation
Arundo donax L. Exotic/ Direct
observation
Rosaceae Malus domestica Borkh. Exotic/ Direct
observation
Prunus persica (L.) Batsch var. Pérsica Exotic/ Direct
observation
Rhamnaceae Hovenia dulcis Thunb. Exotic/ Direct
observation
Solanaceae Salpichroa origanifolia Baill. Native/ Direct
observation
Cestrum parqui L’Hér. Native/ Direct
observation
Urticaceae Parietaria debilis G. Forst. Exotic/ Direct
observation
Urtica urens L. Exotic/ Direct
observation
Verbenaceae Aloysia citrodora Palau Native/ Direct
observation
Aloysia polystachya (Griseb.) Moldenke Native/ Direct
observation
Vitaceae Vitis vinifera L. Exotic/ Direct
observation
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 369

Biological corridors in this area that help increase connectivity are relatively
simple yet successful interventions to promote the growth of spontaneous commu-
nities while also creating small refuge areas. These corridors often use plot edges or
rural roads to create the naturally interconnected network described above without
affecting the surrounding agricultural fields (Fig. 14.2; Biasatti et al. 2013).
The location of the island of study is within a larger system of similar islands
connected through potential biological corridors, which allows for each patch to
remain connected. This network provides an opportunity to study the relationship
between landscape ecology, agroforestry systems, and agroecology models with
scientific and epistemological support. The study of the intersection of these disci-
plinary fields provides valuable information that can be integrated into conservation
strategies. This research adds to the already rich and diverse knowledge that those
disciplinary fields supply, which serves to provide recommendations for combined
systems that integrate trees, shrubs, and wild plants with crops and domestic animals.
This intersection lays the foundation for local food security and food sovereignty
(Montagnini et al. 2015).

14.4 The Role of Islands of Resilience in Wildlife

Conservation

Loss of animal species diversity is a critical problem in both community ecology and
conservation biology (Moreno et al. 2011). Its study has gained greater relevance
globally in recent years as the issue has worsened due to human activities (Moreno
et al. 2011).
As previously explained, the agricultural production model in this region of
Argentina has had severe effects on biodiversity. The model of maximizing agricul-
tural profits has led to a significant decrease in wellbeing of the natural environment
and a change in the structure and functioning of the local ecosystem. Anthropic
actions have had a clear effect on species’ populations, wildlife in particular,
affecting the abundance and range of species (Dirzo et al. 2014). Wildlife species
have different levels of sensitivity to human alterations, depending on their space
requirements, feeding needs, and reaction to anthropization (e.g., Fox and Fox 2000,
Smith et al. 2000, Poiani et al. 2001, Abba et al. 2007).
The effect of anthropization on fauna is a multidimensional phenomenon, the
study of which requires a multidisciplinary approach to best understand the dynam-
ics and structure of species and predict their maintenance in modified environments.
One important contributor to the perseverance of species in a rapidly anthropizing
region is the presence of resiliency islands. These relics have the potential to serve as
ecological sanctuaries and thus provide the minimum resources necessary to meet
the biological needs of wildlife.
370 L. H. González et al.

Table 14.2 Animal species (mammals) recorded in island of resiliency in Las Rosas, Santa Fe,
Argentina
Common name Scientific name Origin/ Observations
Comadreja overa Didelphisal biventris Native/ Indirect records
Peludo Chaetophractus villosus Native/ Indirect records
Zorro Lycalopex gimnocercus Native/ Indirect records
Gato montés Leopardus geoffroyi Native/ Indirect records
Zorrino Conepatus chinga Native/ Indirect records
Hurón menor Galictis cuja Native/ Indirect records
Ratón del pastizal Akodon azarae Native/ Indirect records
Colilargo chico Oligoryzomys flavescens Native/ Indirect records
Laucha manchada Calomys laucha Native/ Indirect records
Laucha bimaculada Calomys musculinus Native/ Indirect records
Cuis Cavia aperea Native/ Indirect records

14.4.1 Measuring Biodiversity in Resiliency Islands

In order to measure the diversity of terrestrial vertebrates, indirect records such as

footprints or caves/burrows left by the transit of some mammals were utilized. The
methodology used was the delineation and examination of linear transects in search
of signs of wildlife activity. These observations were made at an average speed of
500 m/hour, which allowed the observer to carefully examine the area of study. The
field work was carried out between late winter and early spring.
Due to the presence of caves/burrows, we can say that several of the observed
mammals can be considered “engineers” of the ecosystems because they influence
the availability of resources for other species through changes in the state of the
biotic or abiotic materials used to build their shelters (Jones et al. 1994). Although
the concept of “ecological engineers” has been refuted because of its possible
application to all organisms, it is useful as a reference for studying biotic interactions
caused by changes to a habitat (Jones and Gutiérrez 2007). Excavating mammals, for
example, modify their environment by creating entrances and conduits in the ground
that cause changes related to the soil formation processes and increase the availabil-
ity of shelter for other vertebrate species (Jones et al. 2006). Particularly in highly
anthropized ecosystems, caves or burrows represent a valuable resource as a refuge
to avoid predators and mitigate worsening environmental conditions. It is common
to observe the reuse of a cave over time by individuals of the same or different
species (Roldan and Saurthier 2016). The species that occupy these caves can save
time and energy intended for building their own shelter or protecting themselves. For
example, foxes take refuge mainly in caves and holes of different types and sizes that
may belong to hairy armadillos (Chaetophractus villosus) or skunks (Conepatus
chinga).
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 371

Utilizing these shelters as well as other footprints and signs, observations were
made of a total of 11 species of mammals belonging to 8 families (Didelphidae,
Dasypodidae, Canidae, Felidae, Mephitidae, Mustelidae, Cricetidae and Caviidae)
and 4 orders (Didelphimorphia, Cingulata, Carnivora and Rodentia). These observed
mammals include the following: 1 marsupial species, 1 xenarthran (an exclusively
American placental mammal in a clade that include anteaters, armadillos, and
sloths), 4 carnivores, and 5 rodents (Table 14.2). For amphibians and reptiles, no
samples were taken during the period of observation, but tracks have shown the
presence of the overo lizard (Tupinambis merianae).
Regarding birds, the records obtained allowed us to determine the presence of
39 species, which included 9 orders and 18 families (Table 14.3). This field work
was carried out between late winter and early spring and focused on pre-established
transects. Each transect had a fixed-radius of 10 meters and was 20 meters high,
separated from each other by a distance of 25 meters. Intervals for counting
specimens were 15 minutes long. Counts were made in the morning, starting
20 minutes after sunrise and up to no more than 4 hours after sunrise. The results
allowed us to determine that the largest number of individuals observed belong to the
order Passeriformes, which contributed 22 species, followed by the order
Falconiformes and Columbiformes, with 4 species.
Bird nests also played an important role in the evaluation of species richness. The
construction of nests for reproduction, hibernation and/or rest is a very common
activity in birds. Bird nests are a large determining factor in the reproductive success
of the nesters because of the role nests play in sexual selection, protection against
potential predators, and maintenance of suitable temperature and pH level, which
prevents the proliferation of pathogens. Due to the importance of nests in bird
survival, the presence of nests in the study area led us to conclude that the existing
resources satisfy the previously mentioned needs, which are nearly nonexistent in
the surrounding matrix.
The existence of these small green patches in a dominant matrix that is highly
disturbed by anthropic activity provides an opportunity for the recovery of favorable
environmental conditions for the conservation of biodiversity. However, in order to
enhance conservation, it is fundamental to establish the basis for future monitoring
of the species involved in this study. Tracking species interactions within the
ecosystem is crucial in evaluating the importance of the resiliency island in animals’
feeding and breeding habits and therefore assessing the ecological importance of the
islands in the region.
372 L. H. González et al.

Table 14.3 Animal species (Birds) recorded in island of resiliency in Las Rosas, Santa Fe,
Argentina
Common name Scientfic name Origin/ Observations
Paloma Doméstica Columba livia Exotic/ Direct Observation
Paloma Manchada Patagioenas maculosa Native/Directly Observed
Paloma Picazuro Patagioenas picazuro Native/Directly Observed
Torcaza Común Zenaida auriculata Native/Directly Observed
Pirincho Guira guira Native/Directly Observed
Chiflón Syrigmasi bilatrix Native/Directly Observed
Picaflor Verde Chlorostilbon lucidus Native/Directly Observed
Tero común Vanellus chilensis Native/Directly Observed
Milano Blanco Elanus leucurus Native/Directly Observed
Taguató Común Rupornis magnirostris Native/Directly Observed
Carpintero real Común Colaptes melanolaimus Native/Directly Observed
Carpintero Campestre Colaptes campestroides Native/Directly Observed
Carancho Caracara plancus Native/Directly Observed
Chimango Phalcoboenus chimango Native/Directly Observed
Halconcito Colorado Falco sparverius Native/Directly Observed
Halcón Plomizo Falco femoralis Native/Directly Observed
Cotorra Myiopsitta monachus Native/Directly Observed
Hornero Furnarius rufus Native/Directly Observed
Piojito Gris Serpophaga nigricans Native/Directly Observed
Benteveo Común Pitangus sulphuratus Native/Directly Observed
Suiriri Real Tyrannus melancholicus Native/Directly Observed
Churrinche Pyrocephalus rubinus Native/Directly Observed
Monjita Coronada Xolmis coronatus Native/Directly Observed
Monjita Cabeza Negra Microspingus melanoleucus Native/Directly Observed
Tijereta Tyrannus savana Native/Directly Observed
Golondrina Barranquera Pygochelidon cyanoleuca Native/Directly Observed
Tacuarita Azul Polioptila dumicola Native/Directly Observed
Ratona Común Troglodytes aedon Native/Directly Observed
Calandria Grande Mimus saturninus Native/Directly Observed
Calandria Real Mimus triurus Native/Directly Observed
Gorrión Común Passer domesticus Exotic/ Direct observation
Cachilo Ceja Amarilla Ammodramus humeralis Native/Directly Observed
Chingolo Zonotrichia capensis Native/Directly Observed
Tordo Pico Corto Molothrus rufoaxillaris Native/Directly Observed
Tordo Renegrido Molothrus bonariensis Native/Directly Observed
Tordo Músico Agelaioides badius Native/Directly Observed
Corbatita Común Sporophila caerulescens Native/Directly Observed
Jilguero Dorado Sicalis flaveola Native/Directly Observed
Misto Sicalis luteola Native/Directly Observed
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 373

14.5 Proposals for Promoting Alternative Agro-Food

Systems

14.5.1 An Agroecological Approach for Rethinking Food

Production from Islands of Resilience

Landscape ecology conceptualizes landscapes as complex, adaptive systems com-

posed of ecological and social processes interacting at different levels (Zacagnini
2014). There are many other fields of study that consider the potential interaction
between ecological systems and agricultural practices. Agroecology is one such field
that focuses on agricultural systems and the role ecological processes can and should
have in productive models (Levin 2022). Due to agroecology’s focus on production
and conservation, it is possible to address the socioeconomic, cultural, ecological,
and ethical complexity of food, fiber, and energy production, as well as processing
systems, by providing a set of principles for planning local food systems (Wezel
et al. 2009). Therefore, agroecology serves as an alternative to current production
models that would transform the way we produce and consume food into a more
sustainable practice (Nyéléni Boletin 2013, Levin 2022).
Permaculture, regenerative agriculture, agroforestry, biodynamics, and natural
agriculture are models with a strong ecological component in their conceptualiza-
tions. These practices integrate agricultural needs of local communities with sus-
tainable farming practices to better protect local ecosystems. Therefore, these
various models of sustainable agriculture can be classified as agroecological strate-
gies. With regards to the pampean grasslands, it is appropriate, and potentially
beneficial, to use these agricultural practices that provide diversity and require
engagement from producers, as opposed to current mechanized agricultural prac-
tices. In order to protect biodiversity, it is necessary to promote productive processes
that incorporate landscape ecology and agroecology practices to maintain production
while benefitting the local ecosystems.
Developing a new productive system that utilizes the islands of resilience and
local ecology requires an understanding of the biological interactions in the ecosys-
tems. For example, the design must consider effects on the soil, nutrient recycling,
biodiversity, and the optimal use of energy. Permaculture practices provide efficient
energy use by locating and designing landscape elements according to their capacity
of use. Areas that are visited daily or with greater frequency are located near homes
or other central components of a landscape, while less-visited sites are located farther
away, thereby minimizing energy spent in transit and protecting more productive
areas from unintentional impacts. This design is related to the multifunctionality of
the ecosystem and the complementary relationships between its components.
Another key design point is the promotion of secondary ecological succession to
reestablish areas of greater biodiversity, whether spontaneous or induced, in accor-
dance with the local soil characteristics (Fig. 14.6).
374 L. H. González et al.

Fig. 14.6 This diagram represents a reticulated system to contribute to the conservation of
biological diversity, in order to integrate it with the productive systems. (Satellite image intervened
by architect Mercedes Machado)

With regards to the reticulated system for biodiversity conservation, spaces

between corridors can be illustrated in diagrams linking multifunctional components
of systems. These designs can integrate permaculture design components, laid out in
concentric rings or strips that relate to the various production systems (Biassatti et al.
2013, 2017). These productive systems can include vegetables, legumes, cereals,
fruit trees, pastures with animal integration or agroforestry systems.
Moving towards sustainable food production through agroecological principles
requires several simultaneous transitions at different scales and in different contexts,
such as social, biological, economic, cultural, institutional and/or political (Tittonell
2019). Appropriate agroecological practices differ according to initial site conditions
due to differing needs and associated conservation strategies (Marasas 2012).
Through careful design and planning, we can use these “islands,” that could be
considered “abandoned,” as central elements of ecosystem conservation practices,
achieving desired goals for the ecosystem. Many of these transformations are already
occurring as young people and adults are organizing in pursuit of a healthier
environment, a better quality of life for their children, and a harmonious relationship
with nature.
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 375

14.5.2 Promotion of Productive Practices in the Islands

of Resilience

Management of islands of resilience should promote biodiversity with emphasis on

soil improvement, using the famous “living soil” approach developed by agronomist
Ana Primavesi in “Ecological Soil Management” (Primavesi and Molina 1984).
Agroecological approaches are described elsewhere in this book, such as the chapter
“Regenerative Agricultural Systems as Biodiversity Islands” (Levin 2022).
Here, we propose land management tactics that can sustainably benefit produc-
tion outside the “islands,” improving soil health and biodiversity management. This
will serve as a guide that can be useful for planning sustainable productive systems.

[Link] Soil Management

Soil is a living system, and its characteristics are properties that emerge from the
networks of interacting parts, not explainable by any single component (Restrepo
et al. 2000). The physical, chemical and biological properties of a soil are
interdependent and generate favorable bio-structural and nutrient conditions, as
well as biological interactions that support the health of plants and the food chain.
Many agro-industrial production systems are unable to store stable carbon in the
soil due to bare soil, the lack of sufficient photosynthetic capacity of planted crops,
and/or the use of synthetic fertilizers or other chemicals which inhibit plant-soil
microorganism associations. The soil must be treated as a living organism and
covered with actively growing plants which provide root exudates and organic
substances that are critical to the cycling of nutrients and surface organic matter,
processes which are necessary for maintaining healthy soil and therefore increasing
the wellbeing of the ecosystem as a whole (Jones 2008).
Successful soil management balances the inputs and outputs of organic matter by
keeping the soil covered as well as maintaining living roots within the soil to
promote the normal functions of living soil without the use of chemical inputs
(Benedetto et al. 2019). Rotating crops that add nutrients and improve soil function-
ing also add structure.
The incorporation of cover crops (CC), plants covering the soil for the purpose of
its protection and improvement rather than to be harvested, is one of the most
commonly adopted practices of soil conservation. CC contribute significant amounts
of organic matter to the soil, increasing the soil’s microbial life. In addition, CC can
reduce soil and nutrient losses due to surface runoff (Capurro 2018). However, the
soil may still be threatened by mechanical disruption from the harvest of products or
by water use by plants.
An optimal CC mixture depends on the sequence in which crops are to be planted,
and on the landowner’s objectives. CC compositions can be modified to address
specific crop needs or other soil constraints. For example, if the previous crop is a
legume, the CC could be a mixture of grasses. If the soil is compacted, we can add
376 L. H. González et al.

species with deep tap roots that make channels and break up the dense soil layers.
Several species that can help loosen soil include wild turnip (Brassica rapa L.),
fodder cabbage (Brassica napus x B. oleracea cv Interval), cow’s tongue (Rumex
crispus), and chicory (Cichorium intybus). In some cases, a light grazing of grass
species can be integrated, contributing manure, biological pest control, and also
aiding in weed control.

[Link] Cropping Systems

Several techniques can improve cropping systems, including intercropping, strip

cropping, and multistrata cropping. Each has its own benefits to production and
conservation. In addition to improving productive capabilities of crops,
implementing a sequence of diverse crops generates favorable environments for
insect populations, increasing biological interactions among different components of
biodiversity, which further promotes key ecological functions and processes.

[Link] Promotion of Biodiversity

The promotion of biodiversity contributes to the health of the agro-ecosystem. The

different strategies for protecting the ecosystem contribute to preserving and increas-
ing predatory and parasitic insects, conserving healthy levels of pest species, includ-
ing birds, as well as micro and mesofauna. The presence of multistrata borders,
hedges and patches of plants in landscape design establishes refuge zones for insects,
as well as barriers to threatening anthropized areas.
Biodiversity corridors, integrating species from the Umbelliferae, Brassicaceae
(Cruciferaceae), Asteraceae (Compositaceae) and Fabaceae (Leguminosae) families
can be established to attract pollinators and other beneficial insects, further increas-
ing biodiversity.

14.5.3 Key Factors for Achieving Ecosystem Recovery

Land tenure is a key factor to consider in the design of productive agro-ecosystems.

With guaranteed tenure, the producer is able to sustainably plan for long-term
ecological recovery, utilizing long-lived, regionally appropriate species. Public
policies linked to the promotion, support, strengthening and financing of agroeco-
logical food production are also necessary. Producers decide to make changes to
their production strategy depending on many factors such as their presence on the
farm, personal observations, and farmers’ connection to changes in the local envi-
ronment. They also often work towards their goals of becoming independent of
external inputs, demonstrating opinions on appropriate uses of agriculture technol-
ogies, which often contrast with predominant trends.
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 377

Understanding nature and ecological principles should guide us in the design and
management of agroecosystems, discarding a priori designed formulas or
one-dimensional responses. The complexity of the relationships and interactions
within the environment and between species, including humans, should be the
foundation for planning agroecological models of food production. This growing
emphasis on relationships in nature is reflected in the many and varied experiences
that are described in the following section.

14.6 Integrating Ecology and Economy in the Region

In most societies around the world, the dominant economic system has relied on a
similar method of anthropic transformation of natural landscapes, which has
increased significantly in scale over the last three centuries. This reliance on
anthropization has had serious negative impacts on the environment, which has
perverted the original objectives of the economy as an agent of transforming nature
to satisfy human needs. This system becomes further distorted by the fact that human
society is a subordinate system to the environment, which provides all of its shelter
and resources. In fact, the biased and erroneous hierarchization of the economy over
the environment compromises the sustainability of current and future societies. As
the Romanian economist Georgescu-Rogen states, “the only characteristic that
differentiates humanity from all other species. . . is that we are the only species
that in its evolution has violated biological limits” (Georgescu-Roegen 1977).
Therefore, it is crucial to incorporate bioeconomic practices into the current
economic systems, integrating economic activities into natural ecological contexts
(Passet 1996). The bioeconomic practices have been developed by “ecological
economists” such as José Manuel Naredo (2015) or Joan Martínez Alier (2005).
The original definition of the Greek term “economy” (“oikos” ¼ home or household,
“nemein” ¼ management and dispensation) refers to the practice of managing
resources, materials, and natural goods for the support and welfare of the family
home (Aristotle, Politics, Book I, Chap. III) (Aristóteles 2005). This definition
illustrates the fundamentals of the economy in which humans and society depend
on nature and its resources for survival and social reproduction. Humans are
therefore responsible for designing an “instituted process of interaction between
them and their natural environment, in order to provide themselves with material
means to satisfy their needs,” (Polanyi 1976). Therefore, the “primary objective of
economic activity is the conservation of the human species” (Georgescu-Roegen
1971). However, the human species will fail to be conserved if the natural environ-
ment, in which human life originates, develops and endures, is destroyed.
Historically, several communities used a balanced logic of exchange with the
natural environment that included principles such as reciprocity, redistribution and
exchange. Today, many economies consider these practices, which were harmoni-
ously and wisely linked to the surrounding environment, primitive (Polanyi 2011).
378 L. H. González et al.

Over time, the center of the economy in most societies around the world has shifted to
the market and capital, which are both increasingly financialized and speculative,
further neglecting the importance of the environment.
However, in many countries around the world, and in Argentina in particular,
despite threats to the environment posed by profit-focused economies, family
agroecosystems that produce healthy food continue to exist. They allow us to
reconnect with the original empirical and substantive sense of the economy. Present
mainly in increasingly environmentally threatened rural and peri-urban areas, they
become small resilient strongholds in areas where an economy linked to the extrac-
tive model dominates, which seriously compromises the health of the environment,
people and local economies.
These local economic systems linked to family agroecosystems are centered on
people and sustainable productive work. They link production, transformation,
distribution and consumption in order to meet the needs of community members
while still considering the wellbeing of the ecosystem. In this sense, they involve
“forms of production and reproduction of their material conditions of existence, based
on a balanced metabolism with nature” (Toledo and González de Molina 2007). In
these family agroecosystems, the household unit is integrated to the production
process, forming a nucleus of social management of the agroecosystem (Petersen
et al. 2017). The creation of this nucleus is the foundation for implementing diver-
sified economies, integrating diverse and complementary activities linked to the
possibilities and limitations of the territory and maintaining a reduced economic
and productive scale, since the primary purpose of the system is the continuation of
the family economic unit.
Our proposal is to recover and promote these economic and agroecological
experiences with local communities, mainly in the humid pampa region of Santa
Fe, Argentina. This would require the use of the interstitial spaces provided by the
“islands of resilience” and the peri-urban ranges or strips of the cities. The proposal
also urges for the economy to be restructured to incorporate ecologically-mindful
practices, and particularly, agroecology.
Locally based agro-food systems are also a key important aspect of this new
proposed economic system, and further demonstrate the need to redesign local
economic systems for the purposes of increasing healthy food production for the
community. By shifting control of the economy and of the various components of
the agroecosystem to actors of the local community, agro-food systems would be
given more consideration because of the numerous benefits they provide to commu-
nity members. This will further aid in the proposal to integrate “resiliency islands”
and the larger ecosystem into the economy in order to increase wellbeing of both the
community and local biodiversity.
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 379

14.6.1 The Potential of Local Food Systems

The current global food system is unsustainable from an economic, social and
environmental perspective. It is based on the concentration of the food supply into
the hands of a small number of actors who decide what we eat, where we acquire that
food, and what prices we pay. They also have the ability to introduce increasingly
unhealthy, non-nutritional, and processed food, which may travel long distances,
unleashing serious social and environmental impacts. These are the “mileage foods”
described by the organization Friends of the Earth ([Link]) in their 2012
article called “Mileage foods: CO2 emissions from food imports into Spain”. In
addition to challenges with food supply and demand, the global trend of urbaniza-
tion, a large-scale societal transition from periods of progressive expansion and
growth to the unsustainable mega-cities, threatens food supply, generates waste,
and promotes high levels of energy and materials use. Therefore, it is vital for
economies to be centered more regionally, relocating food production and consump-
tion to the local level, within the framework of a true eco-social transition.
In many areas in the Santa Fe Pampa, the local demand for food could be supplied
by the diversified production of healthy and fresh food coming from the region itself.
This goal could only be achieved if the conditions were enabled for the local
economy to be adequately strengthened and enhanced. However, in most rural
areas of the region, there is currently a dependence on food from distant places,
which negatively affects regional food security. Local communities are losing
autonomy and capacity to influence their own food production and consumption
models as they rely more on global food production and a food culture that is
increasingly homogeneous, and less healthy and diverse.
Therefore, we need to recover and energize fair, healthy, sustainable and sover-
eign food production systems. This transformation must occur at the local level,
revitalizing towns, villages and rural areas, identifying, characterizing, and enhanc-
ing local “islands of resilience,” while supporting local, agroecology-based food
systems.

14.6.2 Examples of Local Food Strategies in the Province

of Santa Fe, Argentina

In economic systems focused on food production, potential exists for varied and
creative initiatives that link primary production, product transformation or value
addition, distribution, marketing and consumption. For instance, in the Province of
Santa Fe, a digital platform is currently being used which was implemented in
Rosario, called EcoAlimentate, the purpose of which is “connecting producers and
consumers.” The platform maps existing farms in Argentina that sell directly to
consumers, providing their location, reviews and contact information (https://
[Link]/). This increases transparency and connection between local
people and producers.
380 L. H. González et al.

One such farm is “El Hornerito,” an agroecological farm that has both educational
and productive goals and is managed cooperatively by a group of farmers in the town
of Totoras, in the south-central part of the province of Santa Fe ([Link].
[Link]). This project carries out primary production (milk, eggs,
garden vegetables, chickens, corn, etc.) and value addition of products (cheese,
jams, etc.) with an agroecological approach. In addition, the farm sells its products
directly to the local community through unique marketing strategies, such as the sale
of pre-ordered baskets of seasonal vegetables.
Another example located in the southcentral region of the province is “El Manso”
farm, a business operated by a family of five (Granja El Manso). This farm is a small-
scale operation, raising chickens and laying hens outdoors, fed with a balanced diet
of chickpeas, corn and ground peas. The operation also utilizes organic compost,
created from pruning waste and other organic residues from the farm and surround-
ing areas. In addition, the family’s income is diversified and supplemented by
vegetables grown in its garden, consumed on site. Some of these products are
commercialized in the farm’s store “Las 3 Ecologías” in the center of the city of
Rosario (1551 Julio Cortázar street).
The “Common Land” project ([Link] which originated
in 2012 in the town of Lucio V. López, 44 km northwest of Rosario and 132 km from
Santa Fe, the provincial capital, produces a variety of foods without agrochemicals.
It is a self-management and cooperative project developed to produce, add value to,
and market healthy foods (vegetables, seedlings, honey, etc.) grown with an agro-
ecological approach. In addition to the original products, some agroecological pro-
ducers involved with this project in the “green belt” of Rosario (or peri-urban belt in
“agroecological transition”) also have diversified their commercial offerings. This
project enables customers in Rosario to place orders in advance, which are then
delivered on a weekly basis. This allows producers to gradually engage local
consumers, as well as better plan and organize their production and offerings of
healthy food. Producers are also able to better engage with their consumers and tell
the stories of their sustainable methods of production and their benefits to local
ecologies and economies. The project recently has grown to have a fixed sales
location, its own concessionary station in the food area of the promenade known
as “Mercado del Patio” ([Link] a large store strate-
gically located in front of the bus terminal in Rosario.
Popular, traditional food fairs (markets), which originated in the northeast region
of Argentina, can also be utilized to promote a local foods movement. In these rural
fairs, local producers can directly connect with their consumers, generating bonds of
trust and transparency in the construction of prices and traceability of their food. In
urban areas of the region (such as the cities of Rosario or Santa Fe), there are also
“consumer groups,” in which neighbors or consumers are organized, driving the
supply of local agroecological producers who are typically linked through periodic
weekly or biweekly commitments of healthy consumer baskets.
Other more complex forms of engaging production and consumption locally are
consumer cooperatives or producer and consumer cooperatives (called
“prosumers”). In prosumer cooperatives some of the consumers are also local
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 381

producers themselves and are committed members of the social organization that
constructs the system supply, distribution logistics and demand. There are also some
worker cooperatives that operate as distributors and marketers of agroecological
products, such as Pronoar ([Link] or In lak’ ech Almacen
Natural (In lak’ ech Natural Store, 1967 Brown Street, in Rosario).
It is increasingly imperative that governments play an active role in promoting
sustainable food policies. Municipalities must engage with groups of producers and
consumers, aid in the direct transportation and purchase of products from local
producers for school lunchrooms or vulnerable social groups. In addition, local
authorities should generate institutional and regulatory mechanisms that guarantee
the safety of foods and improve their marketing. They should support agroecological
production which guarantees socio-environmental health. An example of this is
found in the Municipality of Bellavista, province of Corrientes, which has passed
a law recognizing the “agroecological label,” derived from the participatory guar-
antee system – or SPG – carried out by the organized network of public and private
actors and the local community itself.
Other examples of creative and innovative mechanisms for supporting local foods
systems can be found in Europe. In Spain, for example, the government allows for
the establishment of “hubs” that concentrate the logistics and distribution of healthy
food, eliminating unnecessary and/or speculative intermediaries and building fair
prices. In Catalonia (Spain), the design and implementation of agroecological food
banks offers a healthy food supply as an alternative to conventional “food banks,”
which involve large distributors of industrialized food with low nutritional value.
Finally, “cooperative supermarkets,” in several places in the United States and
Europe, also offer a valuable alternative to traditional food markets. They are self-
managing and involve participation of consumers, with their focus being the provi-
sion of ecological food to urban consumers. The institutional precedent is the “Park
Slope Food Coop,” in Brooklyn ([Link] which presents a
model that has been exported and adapted to countries like France or Spain (coop-
erative supermarket “La Louve” in Paris and “La Osa,” in Madrid, https://
[Link]/ and [Link] respectively). These coops are alternatives to
traditional capitalist supermarkets, helping organic producers and promoting healthy
food consumption.

14.6.3 Towards Regionalized Food Systems

All these projects show socio-productive and economic strategies that facilitate the
production and consumption of healthy and agroecological foods (primary and/or
with added value) at the local level. Furthermore, these are locally based food
systems in which there are few intermediaries or speculative actors, directly
(or nearly directly) linking producers and consumers, allowing for the building of
trust and transparency between both.
382 L. H. González et al.

Current global food supply chains sustain their dynamics with a few concentrated
agro-industrial actors (often large globalized economic groups) and multiple inter-
mediaries, inserting high quantities of processed, expensive foods with low nutri-
tional value into markets. In contrast, the models proposed in this section are based
on regional food systems with an agroecological approach, rooted in community.
Food systems benefit the local and regional economies by boosting socio-productive
conditions, improving the social structure, and recovering local food culture. They
also provide multiple environmental and social benefits to the region and local
communities.

14.7 Conclusions

This chapter condenses the history of the Pampean grassland biome, where a series
of significant political, economic and social events led to the fragmentation of the
landscape matrix, homogenizing and simplifying the natural ecosystems. As in other
regions of the planet, in this part of the world, the domination and subjugation of
nature prevailed historically, considered as a repository of resources at the service of
an economy based on accumulation, growth and unlimited consumption. In contrast,
in this chapter we propose a strategy of balanced integration between human
communities and ecosystems, in order to embark on another path that will gradually
reverse the degradation of natural resources and make it possible to sustain life in a
broad sense for present and future generations.
The identification of “islands of resilience” in the landscape matrix surrounding
industrial agriculture represents an interesting discovery of enclaves where flora and
fauna are able to retreat, protect themselves and reproduce. For example, in the case
study that we discussed, 23 families of tree, shrub and herbaceous plants, 11 species
of mammals and 39 species of birds were observed, as well as the presence of
burrows and nests. All of these are indicators of ecological success.
Moreover, islands of resilience can contribute to strengthening the economy and
endogenous development of local communities. Thus, the sustainable management
of local agroecosystems leads to the development of territorialized food systems that
link the production of healthy food with local consumption. The “islands of resil-
ience” and the peri-urban fringes of our cities are potential interconnected “nodes” of
production, transformation, organization and distribution of healthy food to the
surrounding communities. Our research showed a wide range of creative strategies
based on local economic circuits. These strategies include practices and actors such
as prosumers, consumer cooperatives, local food fairs or small local family-owned
farms. In all of them, producers, consumers, local government, social organizations
and other relevant local actors link, agree and collaborate to build regional, endog-
enous and healthy food systems, while forging new socioeconomic and community
links.
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 383

We cannot ignore the fact that human beings today are faced with extraordinary
problems that are radically different from anything they have faced before in human
history. They must ask themselves whether organized human society can survive in a
recognizable form. And the answer cannot be delayed (Chomsky 2020). No matter
how much this society is disguised as green or how many speeches there are about
the need for an ecological perspective: the way society actually functions cannot be
transformed unless it undergoes a deep structural transformation, namely by
replacing competition with cooperation, and the pursuit of economic profit with
relationships based on mutual solidarity and concern (Bookchin 2019).

References

Abba AM, Cassini MH, Vizcaíno SF (2007) Effects of land use on the distribution of three species
of armadillos (Mammalia, Dasypodidae) in the pampas, Argentina. J Mammal 88:502–507
Aristóteles (2005) Política, Libro I, Capítulo III. Ediciones Istmo (Colección Fundamentos; Serie
Clásicos del pensamiento político), Madrid
Benedetto V, Gamundi JC, Timoni R, Cardozo F, Longo AD, Guglielmone P, Aradas Díaz ME
(2017) Experimental module of agroecological transition. Magazine to Improve Production N

56. INTA Oliveros, Santa Fe, Argentina
Benedetto V, Aradas Díaz M E, Cardozo FV, Longo AD, Timoni RE (2019) Experience of
extensive Agroecological production with integral and multidimensional approach in INTA
EEA Oliveros, Santa Fe. Book of abstracts, 1st Argentine Congress of Agroecology. National
University of Cuyo. Mendoza, Argentina. ISBN 978-987-575-210-8
Biasatti NR, Bedetti F, Colomar G, Di Nucci D, Marc LB, Romano MC, Spiaggi EP (2007) El
impacto de la actividad agropecuaria sobre la conservación de la biodiversidad en la pampa
húmeda. Rev. UNR/AMBIENTAL N
7, Comité Universitario de Política Ambiental, SECyT
(UNR), Rosario, Argentina, 203p. ISBN 950-817-116-2
Biasatti NR, Avogradini F, Rapalino M (2013) Gridded system for biodiversity conservation in the
province of Santa Fe, Argentina. Mag Argentine Assoc Landscape Ecol 4(2):181–189
Biasatti NR, Rozzatti JC, Fandiño B, Pautaso A, Mosso, E, Marteleur G, Algañaraz N, Giraudo A,
Chiarulli C, Romano M, Ramirez Llorens P, Vallejos L (2016) Las eco-regiones: su
conservación y las áreas naturales protegidas de la provincia de Santa Fe. Secretaría de Medio
Ambiente. Imprenta UNL, Santa Fe. Primera Edición, 244 p. ISBN 978-987-23776-2-5
Biasatti NR, Marc L, Rimoldi P, Spiaggi E (2017) Los valores ambientales de ecosistemas
relictuales en la pampa húmeda: caso de “El Espinillo” y el tramo medio del río Carcarañá en
Santa Fe”. Centro de Estudios Ambientales (CEAV). Facultad de Ciencias Veterinarias.
Universidad Nacional de Rosario (UNR). UNR Editora, Rosario, 62p. ISBN 978-987-702-
262-9
Biasatti NR, Rimoldi P, Cabrera L (2019) Desafíos de la conservación biológica en el espinal y la
pampa húmeda santafesina, los mamíferos del sur de Santa Fe. Ministerio de Medio Ambiente.
Provincia de Santa Fe, Santa Fe, Argentina.106p. ISBN978-987-45488-3-2
Bidaseca K, Lapegna P (2006) El grito de Alcorta revisitado: cultura y sentimientos en la acción
colectiva. Anuario 2006_09 [Link] 09/11/2006 06:04 am, p 309
Bookchin M (2019) La próxima revolución. Las asambleas populares y la promesa de la democracia
directa. Virus Editorial, Barcelona ISBN 978-84-92559-96-1
Brailovsky A (2006) Ecological history of Ibero-America: from the Mayas to Don Quixote. 1st ed,
Editorial Capital Intelectual, Buenos Aires, 240 p ISBN 987-1181-45-0
384 L. H. González et al.

Brailovsky A, Foguelman D (eds) (2009) Green memory. Ecological History of Argentina. 7th
Edition. ISBN 987-1138-30-X. Ed Sudamericana, Buenos Aires, Argentina
Capurro JE (2018) Efectos de un cultivo de cobertura sobre propiedades edáficas y uso del agua en
ambientes con erosión hídrica y monocultivo de soja, en el sur de la provincia de Santa Fe. Tesis
de Magister en Manejo y Conservación de Recursos Naturales. Facultad de Ciencias Agrarias.
Universidad Nacional de Rosario, Rosario, Argentina
Chomsky N (2020) Climate crisis and the global green new Deal: the political economy of saving
the planet. Ed Capital Intelectual, Buenos Aires, 188 p, ISBN 978-987-614-607-4
Cloquell S, Albanesi R, Propersi P, Preda G, De Nicola M (2007) Rural families. The end of a story
in the beginning of a new agriculture. Editorial Homo Sapiens, Rosario, 200 p. ISBN 978-950-
808-524-5
Cracco M E, Guerrero S (2004) Aplicación del enfoque ecosistémico a la gestión de corredores en
América del Sur. Memorias taller regional. 3 al 5 de junio, Quito, Ecuador
Del Corro F (2003) Apropiación de la tierra a los aborígenes y genocidios en el Rio de la Plata.
Rebanadas de Realidad, Buenos Aires 11/08/03
Dirzo R, Young S, Galetti M, Ceballos G, Isaac N, Collen B (2014) Defaunation in the
Anthropocene. Science 345:401. [Link] [Link].
org/content/345/6195/401/suppl/DC1
Donghi TH (2000) From the Independence revolution to the Rosista confederation (Argentine
History, [Link]). Ed Paidos Iberoamerica, Buenos Aires, Argentina, ISBN 978-950-127-723-4
Donghi TH (2014) Revolution and war: building a leading elite in Argentina creole. Siglo
Veintiuno Publishing House, Mexico, 480 p, ISBN 978-987-629-379-2
Forman RT, Grodon M (1986) Landscape ecology. Wiley, New York
Fox B, Fox M (2000) Factors determining mammal species richness on habitat islands and isolates:
habitat diversity, disturbance, species interactions and guild assembly rules. Glob Ecol Biogeogr
9:19–37
Friends of the Earth (2012) Food miles: CO2 emissions from food imports into Spain ([Link].
com)
Gauna Zotter J, Rey M (2017) Tecnología y Sociedad. Ed EDUCA, Buenos Aires. Available in:
[Link]
[Link]
Georgescu-Roegen N (1971) The entropy law and the economic process. Princeton University
Press, Princeton
Georgescu-Roegen N (1977) Bioeconomics. Princeton University Press, Princeton
Herrero A (2017) Navigating through the turbulent times of the Anthropocene. Political ecology.
International debate notebooks, June 2017. ISSN: 1130-6378. Eds: Martínez Alier J, Puig
Ventosa I, Monjo A. Ed Fundació ENT / Icaria editorial, Barcelona
Jones C (2008) The liquid carbon pathway. Australia. Extract from Australian Farm Journal
Ed. 338, [Link]
Jones CG, Gutiérrez JL (2007) On the purpose, meaning, and usage of the ecosystem engineering
concept. In: Ciddington K, Byers JE, Wilson WG, Hastings A (eds) Ecosystem engineers: plants
to protists. Academic, Cambridge, MA, pp 3–24
Jones CG, Lawton JH, Shachak M (1994) Organisms as ecosystem engineers. Oikos 69:373–386
Jones CG, Gutierrez JL, Groffman PM, Shachak M (2006) Linking ecosystem engineers to soil
processes: a framework using the Jenny state factor equation. Eur J Soil Biol 42:39–53
Kropotkin P (1977) The conquest of bread. Jucar Editions, Gijón. ISBN 84-334-1555-7
Levin B (2022) Regenerative agricultural systems as biodiversity islands. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments, Series:
Topics in Biodiversity and Conservation. Springer, Cham, pp 61–88
Marasas M (2012) El camino de la transición agroecológica. Ediciones INTA, Ciudad Autónoma de
Buenos Aires, Argentina. ISBN 978-987-679-104-5 1
14 Natural Landscape of the Pampa Region in Santa Fe Province, Argentina:. . . 385

Martínez Alier J (2005) The environmentalism of the poor. Environmental conflicts and languages
of valuation, 3rd ed. Antrazyt, Icaria Editorial 207, Barcelona ISBN: 978-84-7426-743-3
Mignolo W D (ed) (2007) The idea of Latin America. Editorial Gedisa, S.A., Buenos Aires. ISBN:
978-84-9784-094-1
Mollison B (1994) Introduction to permaculture. Tagari Publications, Stanley, Tasmania, Australia.
ISBN 0 908228 09
Montagnini F, Somarriba E, Murgueitio E, Fassola H, Eibl B (Eds) (2015) Sistemas Agroforestales.
Funciones productivas, socioeconómicas y ambientales. Serie Técnica Informe Técnico
402, CATIE, Turrialba, Costa Rica. Fundación CIPAV. Cali, Colombia, 454p
Moreno C, Barragán F, Pineda E, Pavón N (2011) Reanálisis de la diversidad alfa: alternativas para
interpretar y comparar información sobre comunidades ecológicas. Revista Mexicana de
Biodiversidad 82:1249–1261
Naredo JM (2015) Economic roots of ecological and social deterioration. Ed Siglo XXI, Madrid
Nyéléni Boletin (2013) Número 13, Marzo 2013, [Link] - info@[Link]
Passet R (1996) Principios de bioeconomía. Fundación Argentaria. Visor distribuciones S.A.,
Colección Economia y Naturaleza, Madrid-Bilbao
Petersen P, Marçal da Silveira L, Bianconi Fernandes G, Gomes de Almeida S (2017) Método de
Análise Econômico-Ecológica de Agroecossistemas. AS-PTA, Rio de Janeiro, 246 p
Pizarro JB (2003) La evolución de la producción agropecuaria pampeana en la segunda mitad del
siglo XX. Revista Interdisciplinaria de Estudios Agrarios, numero 18, 1er semestre 2003,
Universidad de Buenos Aires, Buenos Aires
Poiani K, Merrill M, Chapman K (2001) Identifying conservation-priority areas in a fragmented
Minnesota landscape based on the umbrella species concept and selection of large patches of
natural vegetation. Conserv Biol 15:513–522
Polanyi K (1976) El sistema económico como proceso institucionalizado. In: Godelier M,
Antropología y economía. Anagrama, Barcelona, pp 155–178
Polanyi K (2011) The great transformation. The political and economic origins of our time.
Ed. Economic Culture Fund, Mexico
Politis G, Gutierrez MA, Scabuzzo C (2014) Chapter 7: The vertebrates of the site Arroyo Seco
2. Palaeo-environmental implications in current status of the investigations in the archaeological
site Arroyo Seco 2, Tres Arroyos County, 1st ed. National University of the Center of the
Province of Buenos Aires. Faculty of Social Sciences, Tandil, 514 p, ISBN 978-950-658-338-5
Primavesi A, Molina JS (1984) Ecological soil management: agriculture in tropical regions. Ed El
Ateneo, Buenos Aires, 5th edition, 499p
Restrepo JM, Ángel SD, Prager M (2000) Agroecology. Centro para el Desarrollo Agropecuario y
Forestal, Inc (CEDAF), Santo Domingo, Dominican Republic. ISBN 99934-8-002-9
Rimoldi PG, Chimento NR (2018) Diversidad de mamíferos nativos medianos y grandes en la
cuenca del rio Carcarañá, provincia de Santa Fe (Argentina). Rev Mus Argentino Cienc Nat
20(2):333–341
Roldan AV, Sauthier UD (2016) Madrigueras de Dolichotis patagonum como recurso para otros
vertebrados en Península Valdés. Mastozoología Neotropical 23(2):515–520
Sili M, Soumoulou L (2011) The land issue in Argentina. Conflicts and dynamics of use, tenure and
concentration. Ministry of Agriculture, livestock and fisheries. Presidency of the Nation, Buenos
Aires
Smith W, Solow A, Chu C (2000) An index of the contribution of spatial community structure to the
species accumulation curve. Ecology 81:3233–3236
Svampa M, Viale E (2014) Bad development. The Argentina of extractivism and dispossession.
Katz Publishers, Buenos Aires. ISBN Argentina: 978-987-1566-92-1
Svampa M, Viale E (2020) The ecological collapse is here. XXI Century Publishers, Buenos Aires,
296p. ISBN 978-987-801-027-4
Tittonell P (2019) Agroecological transitions: multiples scales, levels and challenges. Mendoza,
Argentina. Rev. FCA UNCUYO 51 (1): 231-246. ISSN (online) 1853 8665
386 L. H. González et al.

Toledo VM, González de Molina M (2007). El metabolismo social: las relaciones entre la sociedad
y la naturaleza. In: Garrido F, González de Molina M (Ed.). El paradigma ecológico en las
ciencias sociales. Icaria Editorial. Barcelona, España. ISBN 978-84-7426-756-3, p. 85–112
Volkind P (2009) El acuerdo de 1920 entre la Federación Agraria Argentina y la Federación Obrera
Regional Argentina (IX Congreso): alcances y limites en el marco de la conflictividad agraria de
la época. Revista Interdisciplinaria de Estudios Agrarios n
31-2do semestre de 2009. Buenos
Aires, Argentina
Wezel A, Bellon S, Doré T, Francis C, Vallod D, David C (2009) Agroecology as a science, a
movement and a practice. A review. Agronomy Sustain Dev 29(4):503–515. [Link]
1051/agro/2009004
Zaccagnini ME (2014) Manual of good practices for the conservation of soil and biodiversity and its
ecosystem services. UNDP, Secretary of the Environment and Sustainable Development of the
Nation, INTA, Buenos Aires. ISBN 978-987-1560-55-4
Chapter 15
Residential Garden Design for Urban
Biodiversity Conservation: Experience from
Panama City, Panama

Helen R. C. Negret, Rafael Negret, and Irene Montes-Londoño

Abstract Urban expansion encroaches on natural areas causing habitat and species
loss. However, cities can offer ecological spaces that harbor high proportions of
regional and local species. In addition to public urban green spaces, private residen-
tial gardens are important for biodiversity conservation particularly if spatially
arranged to maximize habitat-patch sizes and minimize isolation from remnants of
native habitat in the city. Urban growth is projected to increase considerably,
including in biodiversity hotspots, many of which are in developing tropical coun-
tries. In urban areas of these countries, residential “ornamental” gardening is not as
widespread as in temperate developed countries where a multimillion-dollar industry
supports garden design and maintenance. This case study discusses residential
garden design frameworks for tropical biodiversity conservation that, if adopted at
scale, could channel private finance to conservation in urban areas. It documents the
establishment and management of a residential ornamental garden designed to
protect native fauna and flora in an urban landscape in Panama City, Panama. It
describes the design elements and records the positive impact on biodiversity over
15 years in a 1700 m2 property. Grass areas were reduced by 80%, and 64% of the
property was planted, increasing vascular plant species from 10 to at least 180 and
birds from 9 to 157 species. Management approaches, and challenges of increasing
habitat alongside human wellbeing benefits from the garden, are presented. Recom-
mendations and required attitude changes are outlined for garden practitioners, urban
planners and policymakers to replicate the design elements of this biodiversity
garden island in Panama City, and beyond.

H. R. C. Negret (*)
Retired/former environmental advisor, United Nations Development Program, Panama City,
Panama, Republic of Panama
R. Negret
Retired/former environmental advisor, Interamerican Development Bank, Panama City,
Panama, Republic of Panama
I. Montes-Londoño
Hacienda Pinzacuá, Alcalá, Valle del Cauca, Colombia

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 387
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
388 H. R. C. Negret et al.

Keywords Attitude changes · Biodiversity gardens · Garden practitioners · Tropical

gardens · Urban green spaces · Urban policy and planning

15.1 Introduction

15.1.1 Urbanization and Biodiversity

Urban expansion is a major driver of habitat conversion. It transforms habitats

critical for biodiversity, decreasing and fragmenting natural patches by altering
their size, shape, and their connectivity in the landscape (Ricketts 2001; Alberti
2005; McDonald et al. 2008). Consequently, urbanization poses one of the largest
threats to global biodiversity (Luck et al. 2009; Seto et al. 2012; McDonald et al.
2013). Growth models project the global total urban area to triple by 2030, with
respect to 2000 levels. Across the world this expansion will be 4
0.8 times in
biodiversity hotspots,1 with local and global ecological consequences (Seto et al.
2012; Güneralp and Seto 2013; McDonald et al. 2013). In absolute area, South
America is expected to have the largest increase of urban land in biodiversity
hotspots with an increase of more than 100,000
25,000 km2, corresponding to
nearly a 3.5
0.5-fold increase. The largest proportional increase, of about
14
three-fold, is forecasted to be in Mid-Latitudinal Africa (Güneralp and Seto
2013). Biodiversity loss is inevitable with this urban expansion; however, its extent
will depend on urban and landscape planning approaches in public and private land.

15.1.2 Biodiversity in Cities

Urbanization fragments natural ecosystems and alters ecosystem processes, but also
creates social and economic opportunities to build unique ecological spaces that can
conserve native species (Muller et al. 2013). Studies have shown that globally
declining taxa can attain high densities in urban habitats (McFrederick and LeBuhn
2006; Osborne et al. 2008). In this context, many taxonomic groups have 50% or
more of the regional or even national species assemblage found in cities (Sec CBD
2012; McDonald et al. 2013). Two-thirds of plant species occurring in urban areas
tend to be native to the region of each city; the proportion of native bird species being
considerably higher than that of other taxa (94%) (McDonald et al. 2013).

1
Biodiversity hotspots have high endemism, are threatened by human inhabitation and have been
designated as priority areas for conservation (Myers et al. 2000). Of the 36 biodiversity hotspots
identified globally, many are in the tropics and all contain urban areas. Panama City is within the
Mesoamerica Corridor Biodiversity Hotspot.
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 389

Thus, cities can play important roles in the conservation of biodiversity. Within
cities, urban green spaces (UGS) can be critical for native biodiversity conservation,
providing habitat that supports threatened and specialist species, and which warrant
conservation (Aronson et al. 2014; Ives et al. 2016; Lepczyk et al. 2017). UGS
comprise all-natural, semi-natural, and artificial ecological systems within and
around a city (Cilliers et al. 2013). They include a range of habitat types from public
parks with well conserved remnant patches of native vegetation, urban wastelands,
private gardens/yards, to engineered green infrastructure such as green roofs
(Aronson et al. 2017). These UGS vary in their conservation value, depending on
the biodiversity present, but all could represent opportunities to contribute to con-
servation if designed and managed appropriately.

15.1.3 Urban Residential Gardens and Biodiversity

Conservation

In this case study, a distinction is made between an urban residential garden and a
homegarden. The latter is the agroforestry practice of planting a mixed patch of
livelihood-oriented perennial and annual species within a clearly bounded area near
the homes in rural or urban landscapes. The role of homegardens in biodiversity
conservation is important in agricultural landscapes (Kumar and Nair 2006;
Montagnini 2006; Galluzzi et al. 2010). Agricultural production in these
homegardens in tropical cities constitutes a substantial element of food production,
while also playing a role in urban biodiversity conservation (González-García and
Gómez Sal 2008).
In contrast, residential gardens, especially in urban settings, focus primarily on
the provision of other benefits such as aesthetic, therapeutic, and sensory pleasures.
These gardens, referred to by a variety of names such as residential, domestic,
ornamental and private, are the focus of this article. The term residential garden is
used hereafter in this article.
In some cities, residential gardens make up a large part of the urban area and its
UGS. In the UK, residential gardens typically represent 27% of urban land and up to
47% of the UGS area, e.g. in Leicester (Goddard et al. 2010). In developing
countries, residential gardens are also an important resource for ornamental and
conservation purposes, as well as for food. In Latin America, for example, household
“patios” make up 85% of overall UGS area in Nicaragua (González García and
Gómez-Sal 2008) and 45% of UGS in Asunción, Paraguay, which constitutes 23%
of the city’s land area (Yanosky et al. 2014).
Though private green spaces have traditionally been less studied than public ones
due to access difficulties (Goddard et al. 2010), some literature has discussed their
contribution to species diversity in urban areas. In the UK for example, 22.7 million
households (87% of homes) have gardens (Thompson and Head 2014), harboring an
estimated 28.7 million trees, at least 4.7 million nest boxes and up to 3.5 million
390 H. R. C. Negret et al.

ponds (Davies et al. 2009). A study of 61 residential gardens in the city of Sheffield,
UK, found 4000 species of invertebrates, 80 species of lichens, and more than 1000
species of plants (Gaston et al. 2007).
Habitat patch size, quality, pattern and connectedness in the landscape are
important for the survival of several taxa, so, to achieve long term conservation
from residential gardens, land use planning, policy and coordination of actors is
needed across an entire urban space (Evans et al. 2009; Goddard et al. 2010).
However, urbanization rates are highest in those regions of the world where there
are generally under-resourced urban governance arrangements (Muller et al. 2013)
and where financial constraints and livelihood options may continue to place priority
on food-production focused gardening. A multimillion-dollar industry has arisen to
support the design and maintenance of residential gardens particularly in the north-
ern hemisphere.2 Although figures on financial investment by private citizens are not
easily available for all countries, much of urban expansion will occur in developing
Middle-Income Countries, such as all but one of the countries in Latin America.
Mobilizing more gardeners to adopt biodiversity conservation garden design and
management in these countries could harness a largely untapped resource for
achieving local and national conservation targets and provide opportunities for
private citizens to become stewards of biodiversity and ecosystem services within
city boundaries. For this to occur, several challenges need to be overcome including
knowledge gaps on biodiversity garden design and individuals’ attitudes to garden-
ing and biodiversity conservation.

15.1.4 Garden Design for Biodiversity Conservation

There is increasing data in the literature on the role of residential gardens, and in
general UGS, in biodiversity conservation (Goddard et al. 2010; Aronson et al.
2017); however, this focuses mainly on temperate countries in the northern hemi-
sphere and with a high proportion in the United States (Muller et al. 2013). This is
not the case in the developing world, particularly in the tropics where there is a
scarcity of empirical, systematically obtained data on biodiversity conservation in
urban residential gardens (Gaston et al. 2005; Jaganmohan et al. 2012). Nonetheless,
it can be expected that urban residential gardens in the tropics play a role in
biodiversity conservation as well. Further studies can provide guidance for scaling
up this practice. Our case study reports empirical observations from our residential
garden, providing information that could be useful to scientists and practitioners in
understanding and promoting these ecosystems in tropical urban settings.

2
In the UK alone, households spent around £7.5 billion (or US$9.21 billion) on garden goods in
2017, equivalent to 1% of household spending (Oxford Economics 2017). In the USA, in 2018 77%
of American gardeners reported spending a record US$47.8 billion on lawn and garden retail sales,
with an average household expenditure of US$503 (NGS 2018).
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 391

Studies in temperate countries show that form and management of gardens are
essential variables influencing the benefits they provide, including biodiversity
(Cameron et al. 2012). Historically, the legacy of green spaces intended to deliver
aesthetic and recreational benefits has been the simplification of habitats (Davies
et al. 2019). For example, pruning and removing trees, shrubs, and leaf litter,
dramatically simplifies the garden structure (Aronson et al. 2017). Gardeners often
clear away dead wood for potential safety and aesthetic reasons, yet this negatively
affects species that rely on coarse woody debris (e.g., woodpeckers; Kane et al.
2015). Furthermore, very few green spaces are designed and managed to deliver
synergistic biodiversity conservation and wellbeing and health benefits (Davis et al.
2019). Guidelines for amateur gardeners could support a broad range of biodiversity
conservation, in balance with the aesthetic and wellbeing values of the garden.
Garden design is a complete package that defines a vision and provides guidance
and a road map for developing a garden over time.3 Classical garden design books
for residential gardens follow a variety of schools, or styles, largely developed in
temperate climates4 and provide practical, off-the-peg options that may not neces-
sarily include the conservation of natural assemblages across different taxa. Garden
books for the tropics (e.g. Warren 1991; Wijaya 1999; 2007; Neal 2012) often focus
on showcasing renowned gardens or listing plants that can be used in tropical
settings, and may not always propose specific garden design elements and guidance
for the establishment of new gardens specifically for biodiversity conservation.
Hence there is a need for a residential garden design framework for the tropics that
addresses conservation of the broad range of biodiversity, from wildlife to native
plants within urban settings. This chapter is a step towards filling these gaps.
The case study presented in this chapter describes the establishment and man-
agement of an urban residential garden for biodiversity conservation. It reports the
observations of biodiversity increases over 15 years and provides design guidelines
with biodiversity conservation elements alongside those from traditional garden
design. In this sense, it is one of the first documented long-term experiences of
converting a barren grassland residential property into a garden rich in native species
in a tropical developing country where there are still only incipient urban biodiver-
sity conservation actions. By describing the main elements of this garden design
framework and how it could be replicated, the case study also provides an example
of how private initiatives can contribute to biodiversity conservation strategies in
tropical developing countries.

3
Garden design includes three elements (Newbury 2000): (i) Definition of function, style and feel of
the garden (ii) Costs of initial construction (borders), garden features (paths, ponds), plantings
(trees, bushes, annuals), and maintenance; (iii) Physical nature of the garden plot (temperature,
rainfall, soil, slopes, drainage, aspect)
4
Garden design schools include formal with axial symmetry and small numbers plant species;
Japanese with simple styles and intellectual and philosophical elements; sensory with sounds and
colors to stimulate people with disabilities; entertainment with features for outdoor living; and
wildlife with food and shelter for animals (Newbury 2000).
392 H. R. C. Negret et al.

15.2 Garden Location, Design and Establishment

15.2.1 Location of Garden and Surrounding Environment

The residential garden is situated in Clayton, in the Ancon District of Panama City. It
is located near two national parks in an area that was an American army base called
Fort Clayton until its return to the Panamanian Government at the start of 2000. The
area was designated, through Law 21,1997, as a “garden city” which advocated the
pursuit of knowledge and housed the “City of Knowledge” (Gaceta Official 1997;
Conniff and Bigler 2019). The army base headquarters, and most of the nearby
military houses, were built on steep slopes refilled by heavy clay and stones from the
Panama Canal excavation. The houses had no formally established residential
gardens and were joined by communal open spaces of grass dotted with high
Royal Palm trees.5
Two years after the transfer of the base to the government of Panama, in 2002, the
army houses were sold, and an increasing number of families moved from the city
center to Clayton. Since 2000, the population of Ancon has jumped from 11,700 to
an estimated 79,301 in 2020. The annual growth rate from 2000 to 2010 was 10.3%,
compared to just 0.34% in Panama City Center (National Institute of Statistics and
Census, [Link] Since 2002, a series of new housing develop-
ment projects with small gardens were started, often with buildings encroaching into
forest fragments. Active hunting of wildlife is practiced by construction workers,
many of whom immigrated to the city from the interior of the country where hunting
still occurs in some rural areas. Nonetheless, there is still significant tropical forest
habitat in and around Clayton and other large blocks in the nearby Metropolitan and
Camino de Cruces National Parks.
Panama City is known for its warm climatic conditions with annual average
temperatures of around 27  C and a 24 (minimum) to 32  C (maximum) range.
Relative humidity is high (80 to 100%) with a total precipitation of 1930 mm, most
of which falls over 8 months (May to December) with a monthly average of over
200 mm. The remaining 4 months (January to April) are very dry (less than 20 mm
monthly precipitation) with strong winds and high evapotranspiration (WMO 2019).
The garden faces east-southeast.

15.2.2 Garden Design Approach

When we bought the property in 2005, the house, and two neighboring homes, were
set in an expanse of grass covering 9000m2, located about 60 m away from a patch of

5
The Royal Palm scientific name is Roystonea regia, native to Mexico and Central America. From
here on all plants and animals will referred to with the common name. Scientific names are shown in
Table 15.1
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 393

Table 15.1 Scientific names of species cited in text

Cited Animals Cited Plants
English
common
Taxa Type Scientific name names
Birds Ortalis Gray-headed Bushes Sanchezia Sanchezia
cinereiceps Chachalaca especiosa
Buteo nitidus Gray Hawk Brunfelsia Yesterday,
paiciflora Today &
Tomorrow
Gampsonyx Pearl Kite Megaskepasma Brazilian red
swainsonii erythrochlamys cloak
Piaya cayana Cuckoo Duranta erecta Pigeon berry
Squirrel
Amazilia Rufous-tailed Stacytarpheta Snakeweed
tzacatl Hummingbird jamiacensis
Trogon Violaceous Ixora coccinea Jungle
violaceus Trogon and varieties geranium
Trogon Slaty tailed Bougainvillea Bougainvillea
massena Trogon spp.
Momotus Blue-crowned Rusellia Firecracker
momota Motmot equisetiformis
Thamnophilus Barred Ant Flowers Allamanda Golden
doiatus shrike cathartica Trumpet
Myozetetes Social Lantana Lantana
granadensis Flycatcher camara
Zimmerius Paltry Hibiscus spp Rose Mallow
vilissimus Tyrannulet
Vireo Yellow-green Ruella simplex Mexican
flavoviridis Vireo petunia
Contopus Western Cuphea Mexican
sordidulus Wood Pewee hyssopifolia heather
Euphonia Yellow- Alpinia spp Red, Pink and
luteicapilla crowned Shell Ginger
Euphonia
Troglodytes House Wren Heliconia spp Parrot´s Beak
aedon
Dendroica Yellow Foliage Spathiphyllum Peace Lily
petechia Warbler Tall, willis
Thraupis Blue-gray medium Dieffenbachia Dumbcane
episcopus Tanager and ground amoena
Thraupis Palm Tanager cover Tradescantia Wandering
palmarum spp Jew
Piranga rubra Summer Calathea spp. Prayer plants
Tanager
Piranga Scarlet Philodendron Climbing and
olivacea Tanager spp. non-climbing
(continued)
394 H. R. C. Negret et al.

Table 15.1 (continued)

Cited Animals Cited Plants
English
common
Taxa Type Scientific name names
Ramphocelus Crimson- Arachis pintoi Pinto peanut
dimiditus backed
Tanager
Turdus grayi Clay-colored Maranta Prayer plant
Thrush leuconeura
Seiurus Northern Palms Roystonea Royal Palm
noveboracensis Water Thrush regia
Quiscalis Grackle Bactris Peach fruit
mexicanus gasipaes Palm
Tityra Masked tityra Elaeis oleifera American Oil
semifasciata (Cotinga) Palm
Icterus galbula Baltimore Trees Calliandra Pink powder
Oriole surinamensis puff Tree
Icterus spurius Orchard Caesalpinia Peacock
Oriole pulcherrima flower Tree
Basiliscus Basilisk Tabebuia rosea Pink trumpet
basiliscus Lizard Tree
Reptiles Iguana iguana Green Iguana Fruit trees Theobroma Cacao
cacao
Bothrops asper Fer-de-lance Annona Soursop
muricata
Micrurus Central Myrciaria Jabuticaba
nigrocinctus American cauliflora
Coral
Mammals Sciurus Variegated Carica papaya Papaya
variegatoides Squirrel
Dasyprocta Agoutis Psidium Guava
punctata guajava
Cuniculus paca Paca Mangifera Mango
indica
Saguinus Geoffroy’s Vines Thunbergia sp Kings mantle
geoffroyi tamarin
Tamandua Northern Petrea sp Sandpaper
mexicana anteater vine

forest connected to the Camino de Cruces National Park (approx. 4000 ha), a
kilometer away. This park, in turn, is adjacent to the Soberanía National Park
(22,104 ha). Our property was an island in a sea of grass with no physical connection
to the forest and its biodiversity. As far as we observed, there were only a few
mammals (such as agoutis and coatis), surprisingly few birds and insects, and no
flowers. As professional ecologists, life-long ornithologists, and keen gardeners, we
decided to take on the challenge of designing and establishing a diverse garden
reflecting and protecting the rich biodiversity of the surrounding area.
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 395

A key long-term goal was to increase connectivity with the native forest patch,
providing a wildlife haven within the garden and a safe transit across the garden to a
second smaller patch of forest behind two other houses nearby. We developed a
framework to guide a garden design that would contribute to biodiversity conserva-
tion whilst fulfilling other important aesthetic, recreational, therapeutic, and spiritual
attributes of gardens. This framework draws from our professional work in biodi-
versity conservation and includes elements from different gardening schools. It
adopts a gardening approach that replicates processes that are known to increase
connectivity in natural habitats such as hedgerows and large canopy trees. It uses
aspects of the English garden such as paths with curves to entice the walker to
discover new species and spaces at each bend. Japanese elements include a Zen-style
pool with wide underwater steps to sit on and contemplate the forest. Elements from
modernistic tropical designs include those of the Brazilian landscaper Roberto Burle
Marx, which uses tropical native plants, especially bromeliads and heliconias, and
landscaping to create structured elements that capture tropical exuberance.
The process aimed to create a structured design in the short and medium term, so
that the garden did not look like a “work-in-progress” but rather, had a specific and
satisfying feel and character in each phase. Thus, from the start we defined the long-
term vision and identified five main elements of design to repeat and expand over the
next 15 years and beyond as we gradually achieved our goal. These elements are
hedgerows, large flowerbeds/islands with mixes of species native to Panama or other
neotropical countries, trees, paths, and bodies of water. Figure 15.1 demonstrates the
integration of these five elements into the landscape design of our garden. Together,
these elements provide various benefits for wildlife and people (Table 15.2).

15.2.3 Garden Design Framework

Windbreaks and hedgerows have a role in tropical forest regeneration particularly if

they are connected to forest (Harvey 2000; Chacon and Harvey 2006; Leon and
Harvey 2006). Applying this same principle to our residential urban garden, we first
started work at one edge of three properties that bordered the forest patch. These
properties, collectively covering 9000 m2, were largely grassland. Starting at the
edge of the forest, and using native forest trees species, we planted a hedgerow that
separated these properties from others nearby and from a main road. It has been left
to develop naturally, and now has 20 m high trees. We then planted narrower
hedgerows along the borders of our property and the adjacent properties, again
starting at the edge of the forest. We opted for creating hedgerows, not hedges, to
create greater complexity (species mix and dimensional structure) and optimize
biodiversity conservation (Daniels and Kirkpatrick 2006; Smith et al. 2006a, b;
Gonzalez-Garcia et al. 2009). Normally used in rural areas, a hedgerow is a row of
usually untended, closely planted bushes or trees of different species. A hedge, by
contrast, is a thicket of bushes, usually of one species and trimmed regularly to form
a fence between two portions of land or parts of the garden. If designed carefully
396 H. R. C. Negret et al.

both provide food, shelter, and nesting sites for wildlife, and act as a corridor for their
safe passage between different habitats (Peoples Trust for Endangered Species 2019;
Woodland Trust 2019). As in other ecosystems, ecological corridors in the urban
context can play an effective role for the distribution of organisms with low dispersal
capabilities and should be included in planning strategies (Vergnes et al. 2013).
The next step was to plant large flowerbeds or islands of different types of flowers
and shrubs to attract native fauna, provide diverse habitats and nesting sites, and
facilitate movement across the garden and between the hedgerows. To create these
islands, native or naturalized species were used that attract a wide variety of
pollinators including insects, hummingbirds, and bats. Mixes of hardy flowering

Fig. 15.1 Garden Habitat changes over fifteen years

(1) Growth of Hedgerows and Flowerbed-Islands in the Garden. Top image First row, left to right:
Garden in 2006, 2012; Second row, left to right: Garden in 2015, 2020
(2) The Five Elements of Garden Design Framework. Bottom image. Left column, top to bottom.
Mixed Flower Islands, Trees (Pink powder puff and Peacock flower tree), Water: fountain in
bromeliad area. Right column, top to bottom: Hedgerow, Path with bench and resting area at the end
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 397

Fig. 15.1 (continued)

bushes were favored as these withstand tropical downpours and require low main-
tenance. As the aim was to optimize biodiversity conservation, we used many
different species. However, in keeping with ornamental gardening advice, in some
flowerbeds, we clumped several plants of one species together for creating
more visual impact. For the same reason, we also created smaller flowerbeds planted
entirely with only one species.
A third element of the framework was to plant trees, especially those with larger
canopies and edible fruits. These were planted strategically to improve connectivity
between and within the larger flowerbed islands and hedgerows and to increase
structural complexity. Also, they were planted between existing trees and the forest
so as provide canopy level connectivity with the forest. The selection of the locations
and structure of the tree species was made to optimize different conservation
strategies and to provide fruit for wildlife and people.
398 H. R. C. Negret et al.

Table 15.2 The Five Elements of the Garden Design Framework and their Benefits
Species Examples
Framework Main Ecological (Scientific names
Element Area established Function Human wellbeing in Table 15.1)
Hedgerow 480m2 outside Structurally com- Sound and wind- Trees
garden plex corridors breaks e.g. American oil
480 m2 in 4 gar- connecting dif- Privacy palm; pink trumpet
den border ferent habitats Visual diversity tree
hedgerows; each Shelter from Scientific knowl- Large bushes
with ~ area of predators edge e.g. Sanchezia
120m2. Individ- Food & nesting Multidimensional sp. (flowers attract
ual hedgerows sites complexity and hummingbirds);
have 58 plant Carbon sink sensorial Yesterday, today
spp.; development and tomorrow;
30 spp.,18 spp. Brazilian red cloak
and 22 spp.(aver- Vines
age is 32 spp.). e.g. Sandpaper
vine; Kings mantle
Medium height
Heliconias,
Dumbcane, Gin-
gers
Lower foliage
e.g. Philodendron;
Calatheas
Flowerbed/ 4 large islands Safe transit Visual beauty Flowering bushes:
islands (>12m2) with across garden from plant colors e.g. Pigeon berry,
mixed species. between hedge- and mixes and large Ixora, Hibis-
Total area rows variety of pollina- cus, Snakeweed,
219m2; ~ indi- Nesting habitats tor species includ- Bougainvillea
vidual area 55m2; and food ing hummingbirds Flowers: Peace lil-
with ~12spp. Shelter from and bees ies, small
each strong wind and Creativity (shapes Heliconias, Mexi-
heavy rain and mixes) can heather, Mexi-
Olfactory diver- can petunias
6 smaller islands sity Small shrubs:
(<12m2) totaling Alignment with Golden trumpet,
55 m2; ~ area different garden- Lantana, Fire-
8.5m2; ~ 4spp. ing schools cracker
Flowers: Mexican
heather
Ground cover:
Tradescantia spp.,
small prayer plants
18 m long Lantana spp.
flowerbeds with Heliconia spp.
fewer species, Mexican petunias,
total 291 m2; dwarf jungle
~16m2; ~2spp. / geraniums
flowerbed
Golden trumpet
Hibiscus
(continued)
Table 15.2 (continued)
Species Examples
Framework Main Ecological (Scientific names
Element Area established Function Human wellbeing in Table 15.1)
4 small
flowerbeds 56m2;
~2.65m2; ~3spp.
Trees in 14 existing at Canopy level Sensory stimulus Flowering trees:
garden start. connectivity with movement, light Pink powder puff,
including in 64 new trees forest and shade, visual peacock flower
hedgerows planted from Increased habi- beauty Fruit trees:
31spp. (including tats, nesting sites Fruit trees Jabuticaba,
21 fruit trees from and food papaya, soursop,
9 spp. and 3 self- Carbon sink breadfruit, guava,
planting Royal Shade mango, Banana,
Palms) coconut, cacao
5 stand-alone Palms: Peach fruit
bushes palm, American oil
palm
Paths and 144m2 paths Paths provide Access to, and Paths of granite
resting 6 seating areas / transit across discovery of, gar- stones used in
places decks garden for small den diversity and building and
100m2 Burle mammals spaces cement
Marx bromeliad e.g. agoutis Spaces for relaxa- Resting places of
garden with Seating areas/ tion, meditation, cobblestones with
27 bromeliad decks provide observation rockeries of large
species feeding spaces rocks unearthed
from flowerbeds
and pool excava-
tions
Decks from
ceramic and coral-
line tiles
Painted benches,
aluminum chairs
and tables
Water Garden fountain Drinking and Sensory benefits, Garden fountain:
Swimming pool cooling water for therapeutic 3-tiered trickle-
birds and small sounds, visual down fountain
mammals interest, and facil- attracting tanagers
Hunting ground itates wildlife (blue-grey, crim-
for insectivo- sighting son-backed), and
rous animals; Pool provides small mammals
attracts migra- exercise reducing e.g. agoutis
tory birds; stress and relaxa- Pool: 12x 4 meters
increases tion, reflection headed with raised
humidity in dry and meditation waterfall, sub-
season areas merged steps fac-
ing pool for
relaxation and
meditation.
Attracts green
iguana; basilisk
lizard (Jesus Christ
lizard); social fly-
catcher birds
400 H. R. C. Negret et al.

The fourth element of the framework was garden features, including pathways and
resting places. An important element of a garden is to have easy and safe access to its
different corners, so we gradually installed paths made from porous material such as
stones, pebbles and bricks for drainage. Since more effort will likely be put into the
management and conservation of a garden where enjoyment and wellbeing can be
derived, we created resting places with benches or tables along the paths. Over time, as
new trees grew, shade became too dense for grass, so we laid pebbles dotted with
shade-loving species. In the largest of these areas, we created a bromeliad garden using
unearthed rocks and driftwood collected from beaches. This provided visual diversity
and new habitats for many animals, especially invertebrates.
The fifth element of the framework is water. Annual rainfall in Panama is high,
but four months are very dry, hot, and windy, creating harsh conditions for many
animals and plants. We first installed a small garden fountain in the pebbled area. It
provides drinking and bathing water for different species and a soft background
murmuring that contributes to human wellbeing. We later added a swimming pool to
a steep, exposed, grassy slope that was hard to mow. The pool is headed by a
waterfall that provides additional soothing sounds. It is surrounded by multiple plant
species that attract many animals including native bees and hummingbirds and has
provided a new hunting ground for others such as lizards and fly-catchers. We have
even seen iguanas drink from the pool during the dry season (Fig. 15.4).

15.2.4 Observations on Biodiversity Changes in the Garden

Over 15 Years

Over 15 years, the property has been transformed from a sea of grass to a garden with
at least 180 vascular plant species from 120 genera and 48 families (varieties and
cultivars are not included). There has been an 80% reduction of grass-areas and,
using mainly those native to the Neotropics, the planting of 64% of the total area as
hedgerows, mixed and single plant species flowerbeds, and 64 trees (Figs. 15.2). The
different garden-design elements have provided biodiversity and human wellbeing
benefits (Tables 15.2 and 15.3).
In 2005, the only wildlife we observed in the garden were a few agoutis,
occasionally groups of coatis and grass seed-eating birds. Over the years we have
seen an increase in wildlife species in the garden, except for coatis which have
diminished considerably. In mammals, new species include the paca, the northern
anteater, Geoffroy’s tamarin, the nine-banded armadillo, and variegated squirrels.
We have observed that these mammals move to and from the larger forest patch
along the corridors formed by hedgerows. The number of agoutis in the hedgerows
and large flowerbeds also have increased considerably. Between 10 and 20 agoutis
can be seen feeding in the garden in the early mornings. We have seen an increase in
the litter size of agoutis that reproduce in the garden. The litter of agouti is usually
1 or 2 offspring (Smythe 1978). For the first 5 years, this was the case in the female
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 401

Fig. 15.2 Diagrammatic Illustration of the Increase in Area of the Different Garden Design
Elements from 2005 to 2020
402 H. R. C. Negret et al.

Table 15.3 Examples of Successful Plant-types Used in the Garden Design Framework. (All
native to the Americas except for those marked * which are Asian in origin, now naturalized
throughout the tropics)
Plant Type Examples and Ecological Functions and Human wellbeing
Flowering bushes and trees Pigeon berry: (Neotropical) plants of different sizes with
yellow berries, with white, pink or blue flowers. Dwarf
plants good for low borders and standard size for shaping
into hedges. Attracts bees as pollinators in gardens and
vital for 70% of crops that feed 90% of global population.
Sanchezia: Plants native to South America; flowers year-
round with colored leaves, bracts and large orange or
yellow tube-shaped flower that attract many pollinators,
especially hummingbirds. Grows well in dappled shade in
the garden as medium size bushes. Good for hedgerows
Yesterday, today and tomorrow: With blooms lasting
3 days starting violet-purple as a signal to pollinators that
the flower is full of nectar and pollen; then fading to pale
lavender-blue and then white that are no longer attractive,
increased pollination efficiency.
Colorful hardy bracts withstand- Heliconia: e.g. Parrot’s beak flowers, false birds paradise,
ing the heavy rains with 195 known species most native to tropical America,
provide color, aesthetic shapes and lush foliage year-
round; propagate very quickly, attract hummingbirds and
are low maintenance. Some rhizomes are medicinal.
Bougainvillea: (native to South America) visual beauty
from large colorful bracts (pink, magenta, purple, red,
orange, white) surrounding 3 simple waxy white or pale
yellow flowers; thorny stems provide predator protection
for some nesting birds. Leaves are medicinal.
Red and pink ginger*: Provides long-lasting color and
yearlong lush foliage affording protection for wildlife
transit. Once mature, ginger produces offshoots from the
flower bracts that enable easy propagation. Although
Asian in origin now naturalized throughout the tropics.
Hardy flowers withstanding Jungle geranium (Ixora*): Colorful, tropical shrub with
heavy rains numerous named cultivars differing in flower color (white,
yellow, pink, orange) and plant size (dwarf through king),
easy to maintain, good for borders, foundation, large
flowerbeds or standalone shrubs of small tree. Attracts
numerous native bee species. Native to tropical and sub-
tropical areas throughout the world (although world center
of diversity is Tropical Asia).
Mexican heather: Small hardy flowers, attracts numerous
native bee species.
Delicate flowers but with copious, Lantana: Small, tubular blooms with abundant nectar
year-round flowering attract many pollinators particularly butterflies and colorful
Euglossa bees.
Snakeweed: Year-round flowering, attracts pollinators for
native bees, especially colorful Euglossine bees, hum-
mingbirds and butterflies; leaves are toxic to insects so
maintains shape well, can be used for medicinal tea.
Requires frequent pruning and can be well shaped to
different heights.
(continued)
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 403

Table 15.3 (continued)

Plant Type Examples and Ecological Functions and Human wellbeing
Firecracker: Colorful year-round flowers (red or yellow)
that attract pollinators including hummingbirds. Tolerates
full sun to partial shade, grows quickly and has dense
foliage good for overhanging walls. Provides visual
diversity due to weeping form and thin leaves.
Bromeliads Provide visual diversity, are easy to maintain and useful in
tree trunks or shady steep spaces, excellent for stony rest-
ing spaces for meditation. Guzmania spp. and Nidularium
have colored long lasting flowers; Tislandia and Vriesa
have smaller but brightly colored flowers. Other genera
such as Neoregelia, and Crypanthus have small flowers
but have colorful leaves year-round. Some such as pine-
apple have edible fruits. Complex architecture and role in
water reservoirs make bromeliads biodiversity amplifiers,
providing habitats for frogs, insects and spiders. 3499 of
the 3500 species are from the Neotropics (Baench and
Baench 1994).
Foliage Ground cover e.g. pinto peanut and small Maranta prayer
plants, provide year emerald color. Tradescantia spp.
provides purple and pink colors.
Medium height foliage Larger Prayer plants, including
the Calathea genus, provide a variety of shapes and colors
of leaves and flourish in shady conditions
Taller foliage Dumbcane is hardy and shade-loving with
large, oblong, and cream or light-yellow leaves with deep
green spots and stripes. Can reach up to 6 feet; Peace lilies:
Large green leaves year-round and white cup shaped
flowers.
Climbing and ground: Philodendrum, glossy leaves, fast
growing.
Fruit trees Peach palm (Pisba), soursop, mangos, guava, and
jabuticaba provide feeding sources for a variety of animals
and food for humans if people manage to pick them before
they are eaten by wildlife.
High volume flowering tree Pink and red powder puff trees provide food for many
species of insects, birds, butterflies and moths. Flowers
come out in the early morning and attract daytime polli-
nators, and again early evening for nocturnal species. The
flower-laden, umbrella-shape provides shade for garden
tables.
Peacock flower tree: Seed pods provide food for birds such
as parrots and squirrels.

agoutis we saw in the garden with their offspring. At least 3 pairs now breed in the
garden in burrows under the pool and along one hedgerow. In two of these pairs, the
litter size has increased to three, which is normally only found in food rich and safe
environments or captivity (Smythe 1978; Aliaga–Rossel et al. 2008). Also, we have
404 H. R. C. Negret et al.

observed an increase in litter size in the Geoffroy’s tamarin that started coming near
the house in 2014 once the new trees provided some canopy connectivity with the
forest 50 m away. In 2015 the pair had one offspring, in 2017 two offspring and this
year (2020) a further two (Fig. 15.3 (3)).

Fig. 15.3 Examples of Wildlife Conservation Effectiveness

(1) Examples of Bird Species Observed in the Garden as Habitat Diversity Increased. Top left
image. Left column, top to bottom: Resident birds: Blue-gray tanager and Social Flycatcher. Right
column, top to bottom: Migrant birds: Baltimore Oriole, Summer Tanager, Yellow Warbler
(2) Forest-dwelling Bird Species Observed in Garden as Habitat Diversity Increased. Top right
image. Top left to right, Blue Crowned Motmot, Barred Ant Strike. Bottom left to right: Slaty tailed
Trogon, Violaceous Trogon
(3) Observed Increased Reproductive Success (increased litter size). Bottom left image. Three
Young Agoutis Feeding (top) and Geoffroy’s Tamarin with Young (bottom)
(4) Examples of Increased Species of Bee Pollinators. Bottom right image. Right column top to
bottom, Euglossine Bee spp. on Hibiscus and Snakeweed flowers. Left column, Native Bees on
Peacock Flower Tree, Jungle Geranium and Periwinkle flowers.
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 405

The insect biota has also increased. In 2005, the main insects in the garden were
mosquitos, ants, fireflies and a few butterflies. Now countless species of insects can
be seen flying between the canopy of trees and understories in the hedgerows and
large bushes in flowerbeds. Many of these insects are pollinators, such as 10 species
of native bees (Fig. 15.3 (4)), at least 7 wasp species, and a range of butterflies
including the blue morpho. Butterflies in the genus Morpho are noticeable compo-
nents of neotropical forests and fly at either canopy or understory height with
characteristic gliding and flapping flight patterns (DeVries et al. 2010). We now
frequently see this forest species at midday flapping along the hedgerows and across
the garden along flowerbed islands.
The most dramatic and well registered increase in wildlife has been in bird
species. Bird watching is a constant activity for us and has included, on average,
one-hour observation periods from the same spots and times, twice a day, 5 days a
week over the 15 years. In 2005, some nine species frequented the garden. These
were mainly grass seed eaters. We have now recorded 157 bird species in the garden
from 43 families and 101 genera. Based on our observation and knowledge of birds,
along with bird literature (Angehr 2010; Audubon 2019), we have categorized these
birds into three main groups. The first group is native bird species that permanently
feed in the garden, often also nesting in it. Examples of this group are Tanagers
(crimson-backed, blue-and-grey, and palm), Euphonias, House Wren, Paltry
Tyrannulet, Social Flycatcher and Rufous-tailed Hummingbird.
The second group is native bird species that frequent the garden temporarily for
feeding or at different times in the year, but do not nest in it. These include Cotingas,
Cuckoo Squirrel, Great Kiskadee, Streaked Flycatcher, Tropical Kingbird, Gray-
headed Chachalaca and birds of prey such as the Gray Hawk and Pearl Kite. It also
includes bird species that normally dwell and hunt in the forest and are now seen in
the garden. An example of these are the Violaceous and Slaty tailed Trogons. They
usually live in the forest, feeding on insects and fruits, but after seven years, started
coming into our garden where they are now seen often. Another forest species that
now comes into our garden is the Blue-crowned Motmot. A third is the Barred Ant
shrike that feeds on insects in tangled and dense mid-story vegetation but that has
recently started visiting the garden (Figs. 15.3 (1) and (2)).
The third group includes migratory birds that are known to stop over in Panama
(Angehr 2010). We have seen migratory birds in the garden in September through
November coming from the northern hemisphere autumn to the southern summer,
and in April and May on the return journey. Of the 31 migratory species we have
identified during our bird observation, some use the garden as temporary stop overs,
staying for a few days or weeks as they fly south (e.g. Western-wood Pee-wee, some
Warblers, Baltimore and Orchard Orioles). Other migratory species stay more time
in the garden (e.g. the Summer Tanager) until flying back north, and in a few cases
nest here (e.g. the Yellow-green Vireo). We do not know if other individuals of these
species stop over before flying on to more southern sites. Our bird watching
observations have recorded migrant bird diversity increasing for the first 8 years,
but in the last 7 years there has been a reduction in some migratory birds such as the
Baltimore and Orchard Oriole, several warbler species, the Scarlet Tanager and
406 H. R. C. Negret et al.

Northern Water Thrush. This may reflect the loss of avifauna in North America and
the steep decline in migrating birds from the use of pesticides and habitat loss there
and along their routes (Eng et al. 2019; Rosenberg et al. 2019).
Observations of increased reproductive success and number of insects and polli-
nator species, such as native bees and hummingbirds, suggest the effectiveness of the
garden as a biodiversity island for wildlife. Insect biota in tropical rain forests is
distributed between the forest canopy and understory that form a vertical, structur-
ally complex continuum with differing temperature, humidity, light levels and plant
life (DeVries et al. 2010). Structural complexity and vascular plant composition in
gardens also have been shown to be correlated to native biodiversity, especially
invertebrate richness (Smith et al. 2006a, b). This appears to be the case in our
structurally complex hedgerows and large flowerbeds that now house a plethora of
insects. Their growth also roughly correlates to the observed increase in forest
mammals in the garden. Increased vegetation and connectivity to the forest most
likely has facilitated the movement of these mammals through the sheltered areas
rich in food. We also suggest that the increase in litter size of agoutis and tamarins
may be attributed to the increase of suitable habitat, fruits and seeds in the garden.
This is also the case with birds. The increase in bird species and sightings of forest
birds in the garden became more frequent as hedgerows and flowerbeds grew and
structural complexity increased, suggesting their effectiveness in habitat diversifica-
tion and connectivity with the forest.

15.3 Challenges

15.3.1 Challenges in Establishing our Garden

There have been several challenges in creating diverse habitat and increasing
biodiversity. One has been overcoming the physical limitations of the garden lot
and the climate in Panama. Soil enrichment has been a main concern as soil in the
garden was poor and had high clay content. Much of the soil for the first flowerbeds
was brought in from nurseries. Since then, we have used humus-rich soil formed
from decomposed garden plant material left for several seasons under the forest
cover.
Whilst temperatures permit all year growing, water shortage in the dry season is
acute. Animals such as agoutis dig up roots or gnaw trunks for water thus requiring
the netting of some plants. Although the amount of foliage is greatly reduced in the
dry season, many plants can be kept alive by watering daily. Many species flower in
the dry season, but are not all able to withstand the extremely heavy rains of the wet
season. Placing the most delicate plants in sheltered environments or in beds under
the house eaves, enables flowering across seasons. In open spaces, we have planted
small bushes with hardy flowers, or plants with colorful bracts that can withstand the
rain and provide year-round color. Also, in the dry season, winds are high and blow
palm leaves down, causing damage to plants and potentially to humans.
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 407

New palms have been located only on hedgerows. The native Royal Palms are
self-seeding and fast growing. Although the agoutis help control their spread by
eating large numbers of their seeds, several have grown in the outer edges of the
forest with the danger of displacing other species. We now remove younger seed-
lings and reforest the forest edge with other native species. For this, we collect seeds
from forest species.
Overcoming these challenges required time and knowledge, often obtained from
trial and error. These challenges were exacerbated by a lack of well-stocked nurser-
ies. In general, non-native species are preferred over native species in urban designs
because of their ornamental qualities rather than their ecological function (Quigley
2011), which leads to a homogenization of plants available in nurseries worldwide
(Ignatieva 2011). In 2005 there was little variety of native plants available in
nurseries in Panama, and many were from other regions. Nursery staff were unable
to provide guidance on suitable native plants for different light, soil and temperature
conditions. Books on tropical plants from nearby countries that described their
characteristics, growth, and management were available (e.g., Gentry 1996;
Zuchowski and Forsyth 2007). While these are important as field guides for natu-
ralists, they are not user-friendly gardening books with information on mixes of
species to be used in urban residential gardens.
City infrastructure and human activity can change the local light, temperatures
and hydrologic systems (Güneralp and Seto 2013) and native plantings often fail
because species are not adapted to urban environments (Quigley 2011). Even within
the garden, these conditions have changed over time, particularly with the growth of
large canopy trees changing the light and to some extent temperature conditions of
original planting sites, thus requiring relocation of plants. Our experiences have
provided several recommendations of plant-types that can be used in different
garden-design elements (Table 15.3).
Associated with lack of knowledge is the challenge of using native versus
naturalized plant species to maintain the balance between optimizing biodiversity
and enhancing more traditional aesthetic values. Native plant species grow espe-
cially well as they are suited to both the weather conditions of the area and have been
in co-evolution with native insects and animals. However, many of the most visually
attractive and more well-known garden plants come from other regions and have
been the focus of horticulturalist cultivars and gardening guidance.
Some of these species are so widely used in gardening that they have become
naturalized in many parts of the world, adjusting to the local environment and
sustaining populations without direct intervention from humans. While some can
help in the process of ecological restoration, some exotic plants can however become
invasive, with adverse effects on the local ecosystem (Parrota et al. 1997; Senbeta
et al. 2002; Carpenter et al. 2004; McNamara et al. 2006; Berens et al. 2008; Lamb
and Madsen 2012). As much as possible, we use only species native to Panama or
other Neotropical countries. The exception is some species that are naturalized
throughout the tropics and have been established in Panama for many years such
as the Alpinia (ginger) and Ixora (jungle geranium) varieties and cultivars. Both
provide high visual impact and appear to offer shelter and food for native fauna. We
408 H. R. C. Negret et al.

have observed wildlife moving freely through ginger plants near the forest and
ixoras attract a wide range of native bees.
Another challenge that will affect the long-term sustainability of our garden
biodiversity and the replication of our design framework at scale is the attitude
and practices of neighboring homeowners. The first hedgerows were established in
cooperation with keen gardening neighbors that shared costs and helped with
planting. The gardens of the current surrounding homeowners are less well-tended
and their owners may not have biodiversity conservation in mind. Many are regu-
larly sprayed with pesticides to keep insects and vermin at bay and to decrease the
spread of tropical diseases such as dengue. Pesticide use is detrimental to biodiver-
sity. Recent work indicates that the population dynamics of pollinator communities
are less resilient to chemical control of plants and insects in fragmented urban
habitats (Muratet and Fontaine 2015). Few detailed studies have examined impacts
of the amount and type of chemical applications in urban areas, and more should be
done to promote the use of alternatives to chemical control. The use of pesticides and
herbicides in public areas should be managed with the conservation of adjacent
residential gardens in mind.
In pursuit of our biodiversity conservation goal, we have kept the use of pesti-
cides and fungicides to a minimum. The proximity to the forest with large nests of
leaf cutter ants has made this a challenge. When large numbers of ants have come
into the garden, stripping entire trees and bushes, we have used limited amounts of
site-specific insecticide to reduce populations. The control of grasshoppers, cater-
pillars and green iguanas has also been a challenge. As they contribute to the overall
biodiversity of the garden, we have not used manufactured pesticides to control their
populations. The use of natural repellents (garlic, tobacco and pepper), manual
removal, and replenishment of plants has let us maintain populations of these
herbivores and enjoy flowering plants and butterflies. However, we have had to
change our attitude, relinquishing the vision of pristine, shiny whole leaves depicted
in traditional gardening books, and accept the intrinsic beauty of insect-eaten leaves
as part of the attraction of our biodiverse garden (Fig. 15.4).

15.3.2 Challenges in Scaling-up Biodiversity Gardens

Individual residential gardens can contribute to biodiversity conservation in cities,

particularly when garden design and maintenance frameworks are developed accord-
ingly. Their contribution will depend in part on the number of gardens within a city
that adopt a biodiversity garden design approach. There are few examples of this
available in the tropics, but this case study has provided a number of lessons that
could be used for scaling up. Nonetheless, many knowledge gaps and questions
remain and could be addressed with input from the scientific community. For
example, we recorded a significant increase in bird species in the garden over
15 years and hypothesize that this is related to increased habitat that provides food
and shelter. However, an important element of sampling is to determine what
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 409

Fig. 15.4 Sharing the Garden Space with Wildlife

Top left image: Green Iguana drinking from pool and eating garden plants.
Top Right Image: Caterpillar eating Golden Trumpet leaves
Bottom image. Insect eaten leaves (part of the intrinsic beauty of a biodiversity garden)

constitutes a count: a single individual of a species or a viable population (Muller

et al. 2013). For this, more systematic and uniform monitoring of populations is
required.
It is also important to define potential wildlife carrying capacities within the
garden space. In this case study, our own observations seem to indicate that bird
population increases have plateaued and several native species such as the Clay-
colored Thrush, Palm tanagers, Blue and Gray tanager, Gray-headed Chachalaca and
Crimson-backed Tanagers are increasing. The effect these have on displacing other
native species remains to be seen. The recent absence of the Tody Flycatcher from
the garden could be a sign of such displacement.
410 H. R. C. Negret et al.

Similarly, research and community collaboration are needed to identify methods

to control the spread of more aggressive birds in cities. In Clayton, for example, we
have observed a growing population of the Great-tailed grackle (Talingos) preying
on young iguanas and other lizards and on eggs in the nests of native birds. They
were absent from the area until 4 years ago, but can now be seen in new housing
development areas during construction, and in nearby gardens where we have
observed them eating pet food left outside. Panama City has several scientific
institutions that could provide this type of research, including its national universi-
ties, branches of international ones such as Florida State and McGill, and the
Smithsonian Tropical Research Institute.
Garden-size is another factor that will affect the impact on biodiversity. The
relatively small spaces of some gardens may not guarantee enough foraging space or
breeding habitats for long-term survival of some wildlife. Small, stand-alone gardens
may be able to support small populations of some species, but increased isolation
tends to reduce population and gene-flow among patches with the risk of deteriora-
tion of gene pools (McDonald et al. 2013). In this case study we had direct
intervention over 1700m2 and an indirect impact over 9000m2 via our hedgerows
connecting these areas to forest patches adjacent to a large national park. Impacts on
biodiversity could therefore be expected to be considerable. However, residential
gardens in Panama City are rarely this size, although several new housing develop-
ments have gardens and big apartment complexes are creating large ornamental
gardens in residential common spaces.
The location of gardens relative to each other and across the city also will affect
their contribution to biodiversity. Improving connectivity between gardens and other
UGS within city limits will increase the positive contribution from small garden
sizes. For this, the role of residential gardens should be recognized in policy
frameworks and biodiversity strategies to facilitate incentives and focus priority
action in specific urban settings (Rudd et al. 2002). Their role should also be
integrated into urban land use planning and monitored. In this case, our garden is
in an area designated by law as a “garden city” with land-use planning and norms
that favor the role of gardens in conservation. Although the law, for example,
requires garden fences to leave gaps for wildlife transit, this is largely ignored by
homeowners, and enforcement is missing.
Networks of like-minded gardeners that adopt similar approaches to managing
their properties can also enhance benefits over larger scales. The importance of
networks of gardens for conserving avian diversity is well-documented (White et al.
2005; Palomino and Carrascal 2006). Similarly, networks for planting native vege-
tation in gardens adjacent to creeks have been advocated for enhancing habitat
connectivity in riparian corridors in New South Wales, Australia (Parker et al.
2008). In this case study, we were able to plant hedgerows along bordering proper-
ties with likeminded gardeners, but this needs to be extended across larger areas.
Homeowner associations can provide a vehicle for setting up networks (Lerman
et al. 2012) and could be promoted in new housing developments here. Similarly,
NGOs have a role, for instance, the USA National Audubon Society offers the
Healthy Yard Pledge for gardeners to commit to specific management principles.
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 411

The Panama Audubon Society could support networks for monitoring of bird species
and populations in gardens, including migratory species using these as stop-over
sites.
In addition to garden networks, research has shown that all UGS could interact
synergistically to support biodiversity when clustered together (Colding 2007), and
that the presence of adjacent gardens can increase the species richness of urban parks
(Chamberlain et al. 2004). Our garden is situated near the visitor center of one of the
most visited parks in the city, the Metropolitan National Park. Our garden’s location
next to the park could serve to showcase how biodiversity friendly gardens adjacent
to parks could contribute to the conservation of their biodiversity. The visitor center
is also close to two large plant nurseries. Nursery staff could be trained on native
plant species and on the conditions for growing them in residential gardens here.
Ideally, they should avoid stocking exotic plants that have the potential to become
invasive and focus on species native to the country or region. They could also be
stocked with user-friendly guidelines, in local languages, on garden design and
species that contribute to biodiversity conservation. Such guidelines still need to
be developed.
For these upscaling approaches to gain traction, undoubtedly the most important
challenge is that of the choices and attitudes of individuals. Neighbors and city
planners’ attitudes and behavior will determine whether or not our garden will
remain as a biodiversity island, and whether other property owners adopt similar
approaches. The choices people make in terms of what they plant, and how they
structure and manage their garden (behavior), will influence biodiversity outcomes.
Landscape aesthetics and lifestyle factors often take precedence over ecological
concerns (Larson et al. 2009; Kiesling and Manning 2010). For behavioral change
to occur, a first step is to overcome knowledge gaps. Attitudes determine behaviors,
but actual behavior change is the final step in a transition from increasing knowl-
edge, to attitude change, to the intention to change behavior, and finally to actual
behavior change itself (Orams 1999).
This case study provides an example of what can be achieved when choices are
made that favor biodiversity conservation. In Panama, and in many other tropical
developing countries, closing knowledge gaps needs to be complemented by
strengthening awareness around biodiversity and residential gardening. Multimedia
awareness programs are needed on the role gardens play in biodiversity conservation
and in the provision of ecosystem services on which the city depends (Gómez-
Baggethun et al. 2013). This includes biodiversity conservation as a natural solution
in the fight against climate change and in controlling its effects, such as flooding, in
urban areas. Recent research has presented ways of engaging citizens in exploring
biodiversity and ecosystem services in residential gardens (Beumer and Martens
2015), and has shown that learning-focused initiatives can enhance biodiversity on
private property (Diduck et al. 2020). For this and for self-guided learning, a wider
range of supportive materials are needed. This includes developing user-friendly
gardening design books in Spanish.
Awareness building and behavioral change are a long-term endeavor, but there
are some favorable conditions for this in Panama. Plants grow very quickly here, and
change occurs in a relatively short timescale, thus avoiding one of the challenges
412 H. R. C. Negret et al.

faced in biodiversity-focused attitude-change initiatives in the UK (van Heezik et al.

2012). Ultimately, however, behavioral change will only come about once individ-
uals experience first-hand the many benefits that gardens and gardening provide to
human wellbeing, including the aesthetic, therapeutic, and spiritual values of a
garden (Lin et al. 2017; Raymond et al. 2019). In addition to biodiversity benefits,
the design elements we developed in our garden have provided benefits important in
our wellbeing. These include food provision (fruit trees) and exercise (gardening and
swimming), as well as benefits from observing and listening to the more subtle
voices of the garden. There is increasing recognition of positive associations
between species diversity and psychological and physical well-being (Barton and
Pretty 2010; Aerts et al. 2018); of beneficial aspects of the biodiversity and spiritual
well-being nexus (Irvine et al. 2019); and of the correlation between biodiversity and
spiritual and aesthetic experiences (De Lacy and Shackleton (2017). In our experi-
ence, spiritual and aesthetic benefits from the garden have risen alongside the
increase in habitats and biodiversity. These include mindfulness benefits such as
meditation and relaxation; sensory benefits such as light and shadow, tactile, olfac-
tory, and auditive senses; and scientific benefits such as knowledge, observation and
garden science. All of these augment our fulfillment from the garden and sustain our
commitment to its continued management.

15.4 Conclusions

The case study presents the changes in biodiversity over 15 years in a residential
garden established using a design framework to enhance biodiversity conservation.
The design framework included creating hedgerows and large flowerbeds of native
and naturalized species managed with a minimal amount of pesticide. New habitat
patches were established across and along the garden, creating connectivity between
these and the forest, reducing fragmentation and providing corridors for wildlife.
Thus, as well as increasing biodiversity in the garden, this design framework has
likely reduced the edge effect on forest species, potentially contributing to the long-
term survival of the native forest remnant.
Our observations and supporting literature have led us to hypothesize that the
increase in wildlife that we have observed is a consequence of the creation of a
structurally complex and plant species-rich garden that has been successful in
supporting high levels of biodiversity. But this hypothesis needs to be tested.
Replication and complementary studies are needed. The garden is in a privileged
location near a native forest patch. It is not universally replicable, but there are large
intact forest patches in Panama City that could be favorable to replicating the
experience, especially in Clayton and in new urbanizations such as Panama Pacific,
Sendero de Cruces, Portanorte, and Green City.
Whilst direct replication may be restricted to areas closer to native forest, smaller
garden spaces further away are important for creating new habitats and conserving
many native species. However, to optimize the contribution of all gardens to
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 413

conservation, independent of their size, specific biodiversity considerations should

be included in the garden design framework. These have been outlined in the garden
design presented in this case study. Recommendations for plants that can be used in
the different elements of the garden design have been provided.
To achieve uptake of this approach, and take it to scale, recommendations have
been made in the previous section that could be applied in Panama City. Thus far,
scaling up has been documented in developed countries where gardening cultures
and urban planning are strong. Ironically, this does not often correlate with the
richest biodiversity under threat and where urban expansion is likely to be highest.
Promoting residential gardens as urban islands of biodiversity in biodiversity
hotspots in the tropics could bring multiple biodiversity benefits in threatened
areas, along with improved physical and mental health to large parts of urban
populations.

Acknowledgements We thank our parents for instilling in us the love of nature and gardens and
their importance in human wellbeing. We would also like to thank Alexander Coles from the
Florida State University in Panama, who first inspired us to share the design elements of our garden
with his students. Thanks also to all others that have showed interest in the garden, not least of
which is our gardener, Poto, who has helped us for 15 years with the most strenuous of our tasks,
and those that have visited our Instagram account @voicesofthegarden.

References

Aerts R, Honnay O, Van Nieuwenhuyse A (2018) Biodiversity and human health: mechanisms and
evidene of the positive health effects of diversity in nature and green spaces. Br Med Bull 127:
5–22
Alberti M (2005) The effects of urban patterns on ecosystem functions. Int Reg Sci Rev 28:168–192
Aliaga-Rossel E, Kays RW, Fragoso JMV (2008) Home-range use by the Central American agouti
(Dasyprocta punctata) on Barro Colorado Island, Panama. J Trop Ecol 24:367–374
Angehr GR (2010) The birds of Panama. A field guide. Comstock Publishing Associates, Cornell
University Press, Ithaca
Aronson MFJ, La Sorte FA, Nilon CH (2014) A global analysis of the impacts of urbanization on
bird and plant diversity reveals anthropogenic drivers. P Roy Soc B 281:20133330
Aronson MFJ, Lepczyk CA, Evans K (2017) Biodiversity in the city: key challenges for urban
space management. Front Ecol Environ 15(4):189–196
Audubon (2019). [Link] Accessed July 2019
Baench U, Baench U (1994) Blooming Bromeliads. Tropical Beauty Publishers, Nassau
Barton J, Pretty J (2010) What is the best dose of nature and green exercise for improving mental
health? A multi-study analysis. Environ Sci Technol 44:3947–3955
Berens DG, Farwig N, Schaab G, Böhning-Gaese K (2008) Exotic guavas are foci of Forest
regeneration in Kenyan farmland. Biotropica 40:104–112
Beumer C, Martens P (2015) Biodiversity in my (back)yard: towards a framework for citizen
engagement in exploring biodiversity and ecosystem services in residential gardens. Sustain Sci
10(1):87–100
Cameron RWF, Blanusa T, Taylor JE, Salisbury A, Halstead AJ, Henricot B, Thompson K (2012)
The domestic garden – its contribution to urban green infrastructure. Urban For Urban Greening
11(2):129–137
414 H. R. C. Negret et al.

Carpenter FL, Nichols JD, Sandi E (2004) Early growth of native and exotic trees planted on
degraded tropical pasture. For Ecol Manag 196:367–378
Chacon M, Harvey CA (2006) Live fences and landscape connectivity in a neotropical agricultural
landscape. Agrofor Syst 68:15–26
Chamberlain DE, Cannon AR, Toms MP (2004) Associations of garden birds with gradients in
garden habitat and local habitat. Ecography 27:589–600
Cilliers S, Cilliers J, Lubbe R, Siebert S (2013) Ecosystem services of urban green spaces in African
countries – perspectives and challenges. Urban Ecosyst 16:681–702
Colding J (2007) Ecological land-use complementation for building resilience in urban ecosystems.
Landsc Urban Plan 81:46–55
Conniff ML, Bigler GE (2019) Modern Panama: from occupation to crossroads of the Americas.
Cambridge University Press, Cambridge
Daniels GD, Kirkpatrick JB (2006) Does variation in garden characteristics influence the conser-
vation of birds in suburbia? Biol Conserv 133:326–335
Davies Z, Fuller R, Loram A, Irvine K, Sims V, Gaston K (2009) A national scale inventory of
resource provision for biodiversity within domestic gardens. Biol Conserv 142:761–771
Davies ZG, Dallimer M, Fisher JC, Fuller RA (2019) Biodiversity and health: implications for
conservation. In: Marselle M, Stadler J, Korn H, Irvine K, Bonn A (eds) Biodiversity and health
in the face of climate change. Springer, Cham, pp 283–294
De Lacy P, Shackleton C (2017) Aesthetic and spiritual ecosystem services provided by urban
sacred sites. Sustainability 9(9):1–14
DeVries PJ, Penz CM, Hill RI (2010) Vertical distribution, flight behavior and evolution of wing
morphology in Morpho butterflies. J Animal Ecol 79:1077–1085
Diduck AP, Raymond CM, Rodela R, Moquin R, Boerchers M (2020) Pathways of learning about
biodiversity and sustainability in private urban gardens. J Environ Plan Manage 63(6):
1056–1076
Eng ML, Stutchbury BJM, Christy A, Morrissey CA (2019) A neonicotinoid insecticide reduces
fueling and delays migration in songbirds. Science 365(6458):1177–1180
Evans KL, Newson SE, Gaston KJ (2009) Habitat influences on urban avian assemblages. Ibis 151:
19–39
Galluzzi G, Eyzaguirre P, Negri V (2010) Home gardens: neglected hotspots of agro-biodiversity
and cultural diversity. Biodivers Conserv 19:3635–3654
Gaston KJ, Smith RM, Thompson K, Warren PH (2005) Urban domestic gardens (II): experimental
tests of methods for increasing biodiversity. Biodivers Conserv 14(2):395–413
Gaston KJ, Cush P, Ferguson S, Frost P, Gaston S, Knight S, Loram A, Smith RM, Thompson K,
Warren PH (2007) Improving the contribution of urban gardens for wildlife: some guiding
propositions. British Wildlife 18(3):171–177
Gazeta Official NO 23323 Republic of Panama (1997). [Link] /gacetas/
[Link] Accessed January 2020
Gentry AH (1996) A field guide to the families and genera of Woody plants of North west South
America. Conservation International, Washington DC
Goddard M, Dougill A, Benton T (2010) Scaling up from gardens: biodiversity conservation in
urban environments. Trends Ecol Evol 25:90–98
Gómez-Baggethun E, Gren A, Barton DN, Langemeyer J, McPhearson T, O’Farrell P,
Andersson E, Hamstead Z, Kremer P (2013) Urban ecosystem services. In Elmqvist et al.
(eds) Urbanization, biodiversity and ecosystem services: challenges and opportunities: a global
assessment. Springer, Dordrecht, pp. 201–277
González-García A, Gómez Sal A (2008) Private urban greenspaces or “patios” as a key element in
the urban ecology of tropical Central America. Hum Eco 36:291–300
Gonzalez-Garcia A, Bellure J, Gomez Sal A (2009) The role of urban greenspaces in fauna
conservation: the case of the iguana Ctenosaura similis in the ¨patios¨ of Leon city, Nicaragua.
Biodivers Conserv 18:1909–1020
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 415

Güneralp B, Seto KC (2013) Futures of global urban expansion: uncertainties and implications for
biodiversity conservation. Environm Res Letters 8:014025
Harvey CA (2000) Colonization of agricultural windbreaks by forest trees: effects of connectivity
and remnant trees. Ecol Appl 10(6):1762–1773
Ignatieva M (2011) Plant material for urban landscapes in the era of globalization: roots, challenges,
and innovative solutions. In: Richter M, Weiland U (eds) Applied urban ecology: a global
framework. Blackwell Publishing, Hoboken, pp 139–161
Irvine KN, Hoesly D, Bell-Williams R, Warber SL (2019) Biodiversity and spiritual well-being. In:
Marselle M, Stadler J, Korn H, Irvine K, Bonn A (eds) Biodiversity and health in the face of
climate change. Springer, Cham, pp 213–250
Ives CD, Lentini PE, Threlfall CG, Ikin K, Shanahan DF, Garrard GE, Bekessy SA, Fuller RA,
Mumaw L, Rayner L, Rowe R, Valentine LE, Kendal D (2016) Cities are hotspots for threatened
species. Global Ecol Geog 25:117–126
Jaganmohan M, Vailshery LS, Gopal D, Nagendra H (2012) Plant diversity and distribution in
urban domestic gardens and apartments in Bangalore. India Urban Ecosyst 15:911–925
Kane, Warren PS, Lerman SB (2015) A broad scale analysis of tree risk, mitigation and potential
habitat for cavity-nesting birds. Urban For Urban Green 14(4):1137–1146
Kiesling FM, Manning CM (2010) How green is your thumb? Environmental gardening identity
and ecological gardening practices. J Environ Psychol 30:315–327
Kumar BM, Nair PKR (eds) (2006) Tropical homegardens: a time-tested example of sustainable
agroforestry. Springer, Dordrecht
Lamb D, Madsen SJ (2012) What is Forest landscape restoration? In: Stanturf J, Lamb D, Madsen P
(eds) Forest landscape restoration: integrating natural and social sciences. World Forests XV,
Springer, Dordrecht, pp 3–24
Larson KL, Casagrande D, Harlan SL, Yabiku ST (2009) Residents’ yard choices and rationales in a
desert city: social priorities, ecological impacts, and decision tradeoffs. Environ Manag 44:921–
937
Leon MC, Harvey CA (2006) Live fences and landscape connectivity in a neotropical agricultural
landscape. Agrofor Syst 68(1):15
Lepczyk A, Aronson MFJ, Evans KL (2017) Biodiversity in the city: fundamental questions for
understanding the ecology of urban green spaces for biodiversity conservation. BioSci 67:799–
807
Lerman SB, Turner VK, Bang C (2012) Homeowner associations as a vehicle for promoting native
urban biodiversity. Ecol Soc 17(4):45
Lin BB, Gaston KJ, Fuller RA, Wu D, Bush R, Shanahan DF (2017) How green is your garden?
Urban form and socio-demographic factors influence yard vegetation, visitation, and ecosystem
service benefits. Landsc Urban Plan 157:239–246
Luck GW, Smallbone LT, O’Brien R (2009) Socioeconomics and vegetation change in urban
ecosystems: patterns in space and time. Ecosystems 12:604–620
McDonald RI, Kareiva P, Forman R (2008) The implications of urban growth for global protected
areas and biodiversity conservation. Biol Conserv 141:1695–1703
McDonald RI, Marcotullio PJ, Güneralp B (2013) Urbanization and global trends in biodiversity
and ecosystem services. In: Elmqvist T et al (eds) Urbanization, biodiversity and ecosystem
services: challenges and opportunities. Springer, Dordrecht, pp 60–81
McFrederick QS, LeBuhn G (2006) Are urban parks refuges for bumble bees Bombus spp.
(Hymenoptera: Apidae)? Biol Conserv 129:372–382
McNamara S, Viet Tinh D, Erskine PD, Lamb D, Yates D, Brown S (2006) Rehabilitating degraded
forest land in Central Vietnam with mixed native species plantings. For Ecol Manag 233:358–
365
Montagnini F (2006) Homegardens of Mesoamerica: biodiversity, food security and nutrient
management. In: Kumar BM, Nair PKR (eds) Tropical Homegardens: a time-tested example
of sustainable agroforestry. Springer, Dordrecht, pp 61–84
416 H. R. C. Negret et al.

Muller N, Ignatieva M, Nilon CH, Werner P, Zipperer WC (2013) Patterns and trends in urban
biodiversity and landscape design. In: Elmqvist T et al (eds) Urbanization, biodiversity and
ecosystem services: challenges and opportunities. Springer, Dordrecht, pp 149–200
Muratet A, Fontaine B (2015) Contrasting impacts of pesticides on butterflies and bumblebees in
private gardens in France. Biol Conserv 182:148–154
Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hotspots
for conservation priorities. Nature 403(6772):853–858
National Gardener Survey (2018) A comprehensive study of consumer gardening practices, trends
and products sales. GardenReseachcom [Link]
2018-edition/. Accessed January 2020
Neal N (2012) Gardener’s Guide to Tropical Plants (Gardener’s Guides). Cool Springs Press,
Minneapolis
Newbury T (2000) The ultimate garden designer. Seven Dials Cassell & Co., London
Orams MB (1999) The effectiveness of environmental education: can we turn tourists into
‘greenies’? Prog Tour Hosp Res 3:295–306
Osborne JL, Martin AP, Shortall CR, Todd AD, Goulson D, Knight ME, Hale RJ, Sanderson RA
(2008) Quantifying and comparing bumblebee nest densities in gardens and countryside
habitats. J Appl Ecol 45:784–792
Oxford Economics (2017) The economic impact of ornamental horticulture in the UK pp 50 www.
[Link]/recent-releases/3b5ce883-cc72-4cf9-910e-be267fe93f46 Accessed
January 2020
Palomino D, Carrascal LM (2006) Urban influence on birds at regional scale. A case study with the
avifauna of northern Madrid Province. Landsc Urban Plan 77:276–290
Parker K, Head L, Chisholm L, Feneley N (2008) A conceptual model of ecological connectivity in
Shellharbour local government area, New South Wales, Australia. Landsc Urban Plan 86:47–59
Parrotta JA, Turnbull JW, Jones N (1997) Catalyzing native forest regeneration on degraded
tropical lands. For Ecol Manag 99:1–7
Peoples Trust for Endangered Species (2019). [Link]
leaflets/hedgerow-guide-web-version/. Accessed February 2019
Quigley MF (2011) Potemkin gardens: biodiversity in small designed landscapes. In: Niemelä J
(ed) Urban ecology: patterns, processes, and applications. Oxford University Press, New York
Raymond CM, Diduck AP, Buijs A, Boerchers M, Moquin R (2019) Exploring the co-benefits (and
costs) of home gardening for biodiversity conservation. Int J Justice Sustain 24(3):258–273
Ricketts TH (2001) The matrix matters: effective isolation in fragmented landscapes. Am Nat
158(1):87–99
Rosenberg KV, Adriaan M, Dokter AM, Blancher PJ, Sauer JR, Smith AC, Smith PA, Stanton JC,
Panjabi A, Helft L, Parr M, Marra PP (2019) Decline of the North American Avifauna. Science
366(6461):120–124
Rudd H, Vala J, Schaefer V (2002) Importance of backyard habitat in a comprehensive biodiversity
strategy. A connectivity analysis of urban green spaces. Restor Ecol 10:368–375
Secretariat of the Convention on Biological Diversity (2012) Cities and biodiversity outlook.
Montreal. [Link]
Senbeta F, Beck E, Lüttge U (2002) Exotic trees as nurse-trees for the regeneration of natural
tropical forests. Trees 16:245–249
Seto KC, Güneralp B, Hutyra LR (2012) Global forecasts of urban expansion to 2030 and direct
impacts on biodiversity and carbon pools. Proc Nat Acad Sci 109(40):16083–16088
Smith R, Gaston K, Warren P, Thompson K (2006a) Urban domestic gardens (VI): environmental
correlates of invertebrate richness. Biodivers Conserv 15:2415–2438
Smith R, Thompson K, Hodgson JG, Warren PH (2006b) Urban domestic gardens (IX): Compo-
sition and richness of the vascular plant flora, and implications for native biodiversity. Biodiv
Conserv 129:312–332
Smythe N (1978) The natural history of the Central American agouti (Dasyprocta punctata). In:
Smithsonian contributions to zoology 257. Smithsonian Institution Press, Washington DC
15 Residential Garden Design for Urban Biodiversity Conservation:. . . 417

Thompson K, Head S (2014) Gardens as a resource for wildlife. Wildlife Gardening Forum Nov
2019. [Link] Accessed November 2019
van Heezik YM, Dickinson KJM, Freeman C (2012) Closing the gap: communicating to change
gardening practices in support of native biodiversity in urban private gardens. Ecol Soc 17(1):34
Vergnes A, Kerbiriou C, Clergeau P (2013) Ecological corridors also operate in an urban matrix. A
test case with garden shrews. Urban Ecosyst 16:511–525
Warren W (1991) Tropical garden plants. Thames and Hudson, London
White J, Antos M, Fitzsimons J, Palmer G (2005) Non uniform bird assemblages in urban
environments: the influence of streetscape vegetation. Landsc Urban Plan 71:123–135
Wijaya M (1999) Tropical garden design. Archipelago Press and Wijaya Words, Singapore
Wijaya M (2007) Modern tropical garden design. Thames and Hudson, London
Woodland Trust (2019). [Link]
[Link] Accessed February 2019
World Meteorological Organization (2019) World weather information service – Panama City.
[Link] Accessed December 2019
Yanosky A, Ferreira A, Jent A, Brítez Sánchez FJ, Cabral H, Del Castillo H, Benítez Stanley MA,
Velilla M, Ruiz Diaz M, Mereles Haidar NM, Rodas Insfran OI, Cacciali Sosa P, Clay RP,
Granada López YE (2014) Libro Verde de Asunción: Justificación técnica y biológica para la
declaración de Asunción como Capital Verde de Iberoamérica de 2015. Asociación Guyra
Paraguay
Zuchowski W, Forsyth T (2007) Tropical plants of Costa Rica: a guide to native and exotic Flora.
Zona Tropical Publications, Cornell University Press, Ithaca
Chapter 16
Biodiversity Islands at the World’s
Southernmost City: Plant, Bird and Insect
Conservation in Urban Forests
and Peatlands of Ushuaia, Argentina

Rosina Soler, Julieta Benítez, Francisco Sola, and María Vanessa Lencinas

Abstract Ushuaia, a coastal city located in Patagonia, Argentina, amid unique natural
habitats such as sub-antarctic Nothofagus forests and Sphagnum peatlands, preserves
green areas of natural ecosystems within the urban matrix. Forests and peatlands have
persisted as natural islands of different sizes, conservation status and anthropic distur-
bances. These urban green areas are underappreciated and administratively abandoned,
and several threats could lead to their irreversible transformation (e.g., replacement by
new neighborhoods, urban waste deposits). However, natural green urban areas can
serve as biodiversity islands that make substantial contributions to conserve local biota
and also to house new species assemblages (different from those in pristine areas),
bringing native forest and peatland species closer to the local residents and visitors.
These green urban islands can also help to preserve traditional cultural values, enhance
education and generate local touristic attractions. In this chapter, we assessed the
assemblages of plants and birds in urban native forests, and plants and insects in
urban peatlands of Ushuaia city considering two levels of urbanization surrounding
these green areas, and analyzed their variations compared with assemblages in nearby
similar non-urban (low level of disturbance) ecosystems. Despite some differences in
richness, abundance, biodiversity indices, and species composition of vascular plants,
birds and insects, natural urban and peri-urban forests and peatlands still conserve
several species and characteristics similar to those of unmodified ecosystems. How-
ever, the presence of new species, mainly plants, introduced into these urban patches
has modified the original communities, establishing new assemblages that may con-
tribute positively to urban biodiversity islands.

R. Soler (*) · J. Benítez · M. V. Lencinas

Centro Austral de Investigaciones Científicas (CONICET), Ushuaia, Tierra del Fuego,
Argentina
e-mail: [Link]@[Link]
F. Sola
Instituto de Ciencias Polares, Ambiente y Recursos Naturales (ICPA), Universidad Nacional de
Tierra del Fuego, Ushuaia, Tierra del Fuego, Argentina

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 419
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
420 R. Soler et al.

Keywords Endemic species · Green areas · Nothofagus · Patagonia · Refuge

habitats · Sphagnum · Tierra del Fuego

16.1 Introduction

Cities located in mountainous regions and coastal areas are one of the most attractive
destinations for tourism. These areas are still perceived as natural treasures, offering
striking landscapes and a wide variety of highly attractive plant and animal species.
In Patagonia, one of the main attractions are the pristine landscapes surrounding the
cities and the possibility to be in close contact with wildlife through the broad
spectrum of existing opportunities for outdoor activities. In addition, natural areas
within the urban matrix (i.e., parks or wild areas containing original vegetation) are
of great value to residents and visitors, as urban green areas provide a number of
ecosystem services to the community. These ecosystem services include regulating
(e.g., noise reduction, modulation of temperature, removal of air pollution, protec-
tion of water quality), supporting (e.g., increased biodiversity, habitat, soil formation
and storage and cycling of nutrients), cultural (recreation, enhancement of property
value, community cohesion, source of knowledge), and provisioning (e.g., food,
water, fuel) services (Millennium Ecosystem Assessment 2005; Gómez-Baggethun
and Barton 2013). To benefit from ecological services provided by urban green areas
it is necessary to incorporate these overlooked areas in the urban planning of our
cities. The places where people live and work need to be designed so that they can
serve to promote contact with the natural world (Miller 2005). Even those areas
whose original biological composition has been profoundly modified are still impor-
tant for people. The benefits provided by green urban areas, even those that are
somewhat altered, are invaluable (Thomas and Geller 2013).

16.1.1 Biodiversity in Urban Green Areas

From the perspective of island biogeography (MacArthur and Wilson 1967), urban
green areas can be considered as a type of ecological island more or less isolated
from the other green habitats by the surrounding urban landscape (McGregor-Fors
et al. 2011). Urban green areas, even with some levels of disturbance, are considered
to function as biodiversity islands providing potential habitat refuge for different
organisms within the urban matrix (Matthies et al. 2017; Montagnini et al. 2022).
They are also important in maintaining key ecosystem services (e.g., water purifi-
cation and regulation) which are necessary for human well-being and public health
in cities (Gómez-Baggethun and Barton 2013). Biodiversity is essential for
maintaining ecosystem function (Maestre et al. 2012; Reich et al. 2012) therefore
it is important to maintain urban green areas as undisturbed as possible.
Plants play a significant role in animal diversity because they constitute the first
trophic level, i.e. they are the primary producers of energy that is used by other
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 421

members of the food chain. Poorly managed urban green areas have lower biodi-
versity compared to undisturbed ones (Burghardt et al. 2010; Tallamy et al. 2010).
Similarly, poorly managed landscapes can promote the introduction of non-native
plants (e.g., Rozzi et al. 2003; Marzluff et al. 2008; Ceplová et al. 2017; Lencinas
et al. 2017) which can have negative effects on native plant diversity and microhab-
itat conditions (Ceplová et al. 2017). Non-native plants sustain significantly fewer
insects (e.g., caterpillars) than native-plants (Tallamy et al. 2010). Insects play a
significant role in supporting further biodiversity because they are eaten by other
animals (e.g., birds) and represent the main energy source in critical stages of the life
cycle of many species (e.g., many birds feed their hatchlings with insects). Further-
more, insects are involved in soil formation and organic matter decomposition,
which affect both the diversity and structure of the vegetation (Hartley and Jones
2008).
The integration of green urban areas for biodiversity conservation in city planning
is among the most difficult problems to resolve by public and local policies (Murphy
1999; Nilon et al. 2017). Even the effort to keep the native biota intact in urban green
areas may represent an unlikely goal to achieve for local administrations. This is the
case of young and developing cities which suffer from intense and regular distur-
bances (e.g., constant building, new streets) which significantly alter local biota in
green areas (Lososová et al. 2011). In contrast, green areas in less urbanized habitats
(e.g., those that suffer irregular and less strong disturbances) support rich species
diversity and native biota (Zerbe et al. 2003; Ceplová et al. 2017).
In order to identify conservation priorities and/or prevent future environmental
problems in cities it is necessary to understand how urbanization influences different
biological groups. It is important to assess the potential role of biodiversity islands to
conserve local biota and ultimately avoid its contamination, transformation and total
loss inside an urban matrix.

16.1.2 Case Study: Ushuaia City and its Urban Green Areas

Ushuaia, a coastal city founded in 1884, is one of the southermost urban settlements
in the world, and the most populated city (56,900 inhabitants, INDEC 2010) at such
latitude. It is the political and administrative capital of the Argentinian province of
Tierra del Fuego, Antarctica and Islands of the South Atlantic, located in the
biogeographic region of the Andean-Patagonian forest (Fig. 16.1.)
Ushuaia’s urban development has had a short and hurried history since its
foundation, marked by a penal colony established in 1896 (closed down in 1947)
that started to bring people to the city, and by the Industrial Promotion Law (1972)
that stimulated regional economic development. During 1972–1990, the city
underwent a population boom, which tripled its number of residents and expanded
the urban area into the surrounding natural forests to settle new neighborhoods. After
the 1990s, the unexpected increase of the population strongly promoted building-up
areas (houses, schools, etc.) and installing public services, which reduced the
422 R. Soler et al.

Fig. 16.1 Map of Tierra del Fuego, Argentina, showing Ushuaia city, which is located on the
coastal Nothofagus forests of the Beagle Channel

Fig. 16.2 Inside view from urban Nothofagus forests (a) and Sphagnum peatlands (b) in Ushuaia
city, Tierra del Fuego, Argentina. (Photos: Julieta Benítez and Francisco Sola)

forested areas around the unplanned urban layout, a situation that continues in
present days. Therefore, existing forest patches inside the city are currently remnants
of old or secondary native forests (~50 years old) originated from old cuts or fires
(Fig. 16.2). The urbanization growth also gradually surrounded and isolated the
existing peatlands within the city (Fig. 16.2). Small peatlands (<0.3 ha) were filled
with soil, rubble and rocks, and therefore irreversibly transformed to building areas.
But the larger ones, deemed non-suitable for building were left as “brown areas”
immersed in the urbanization, exposed to degrading uses (e.g., substrate extraction,
artificial drainage, urban waste accumulation, refuge of abandoned domestic ani-
mals, human trespassing). A similar situation is currently found in some patches of
urban forests, which are also used to install gardens or squares, or exceptionally
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 423

preserved areas (e.g., the Parque Yatana Urban Reserve, located in the center of
Ushuaia).
Due to the unplanned nature of the city’s sprawl, recreational and protected areas
are also unplanned and scarce, resulting in the misuse and/or overuse of pre-existent
natural patches inside and outside the city. This is the consequence of a cultural and
social necessity, where the lack of accessible leisure outlets is compensated with
available natural resources, resulting in its unregulated use and eventual deteriora-
tion. There is still the possibility to restore these sites as natural biodiversity refuges
while still making them available to the public under sustainable conditions.
Although Ushuaia city is surrounded by natural ecosystems, the natural green
areas in the most densely urbanized area barely reach 10% of the surface. While
these forests and peatlands inside the urban area were not created consciously to
increase the proportion of green areas available to residents, they can provide the
areas of green spaces that are necessary for a healthier and more sustainable city.
Unfortunately, these urban natural green areas are underappreciated and admin-
istratively abandoned. Moreover, we have a limited understanding of how green
spaces are used for refuge, foraging and reproduction by different animal species.
Urban green areas, especially if they are patches of pre-existing native forests and
peatlands, retain part of the local diversity. In this chapter we aim to determine the
role of natural urban green areas as biodiversity islands within Ushuaia city as a case
study from Patagonia, Argentina. We present results of studies on species assem-
blages (species richness, relative abundance, diversity, evenness and similarity
indices) of understory vascular plants and birds in forests, and vascular plants and
insects in peatlands of Ushuaia city. We also compare species composition of
patches inside the urban matrix (two levels of urbanization surrounding these
green areas) with similar and near undisturbed patches outside the urban matrix
(non-urban patches). We propose these natural urban islands could potentially serve
as places to promote conservation of local biodiversity, bringing native forest and
peatland species closer to people. This can contribute to biodiversity conservation in
a broad sense, which could also help to preserve traditional cultural values, improve
education and enhance touristic attractions.

16.2 Methods
16.2.1 Study Area

Ushuaia city is located in the north bank of the Beagle Channel, Argentina
(Figs. 16.1 and 16.3) in the southern end of the South American Cone (54
480 S,
68
190 W). Mountains and glaciers surround the city, and the natural vegetation is
primarily comprised of forests that reach the coast. These native forests were the
main source of firewood and timber since the city foundation. Most of them are
deciduous forests dominated by Nothofagus pumilio (lenga, Nothofagaceae), which
develop mainly in mountain slopes and valleys. Mixed deciduous-evergreen forests
424 R. Soler et al.

Fig. 16.3 Location of the study area in Ushuaia city, Tiera del Fuego, Argentina. Circles ¼ forest
patches, squares ¼ peatlands. The color indicates the urban categories studied: urban (white), peri-
urban (gray) and non-urban (black)

of lenga and N. betuloides (guindo) also occur on the coast and mid slopes and
include other woody species such as Drymis winterii (canelo), Embothrium
coccineum (notro) and Maytenus magellanica (leña dura) as small trees in the
understory (Moore 1983). On the other hand, N. antarctica (ñire) is found surround-
ing peatbogs and inhabiting mixed coastal forests. The vascular plants in the
understory usually include few shrubs (e.g., Berberis and Ribes) that produce
berries, and abundant forbs, graminoids, fungi, mosses, liverworts and ferns. In the
whole landscape, forest masses are intermingled with peatlands and grasslands of
different size and composition according to elevation above sea level. Peatlands
formed by accumulation of Sphagnum spp. mosses during geological times support
other species of vascular plants like Cyperaceae (e.g., Carex spp.), Astelia sp., and
Drosera sp. (Moore 1983). The climate in Ushuaia city is sub-antarctic (Tuhkanen
1992) with short, cold summers (7
C average temperature) and long winters (1
C
average temperature) with regular snow and frost. Mean annual rainfall is 611 mm
near the coast and more than 1000 mm in the mountains.
For our study, we classified forests and peatlands in Ushuaia city according to the
current degree of surrounding urbanization (i.e., their respective locations in the
urban matrix): (1) Urban: patches inside the urban matrix, 100% surrounded by
buildings, streets or waste disposal areas; (2) Peri-urban: peripheral patches but still
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 425

Table 16.1 Summarized description (patch level) of native forests and peatlands studied in the
urban landscape of Ushuaia, Tierra del Fuego (Argentina)
Urban Peri-urban Non-urban
General location Surrounded by Partially surrounded Not surrounded by
urbanization by urbanization urbanization
Area Forests 9–3 ha 10–9 ha > 10 ha
Peatlands 0.3–12 ha
Main anthropic Recreation, educative, Recreation, sports, Trekking, very
use and frequency passage, frequent sporadic sporadic
Main Household and recrea- Recreational waste Without
contamination tional waste contamination
Domestic species Dogs, cats, horses, very Dogs and horses, Almost none, very
and frequency of frequent frequent sporadic
visit

inside the urban matrix, and partially surrounded (approximately 50%) by the urban
matrix; and (3) Non-urban: patches outside the urban matrix (i.e., without direct
contact with urbanization but still close to the city), with low-impact recreational use
(Fig. 16.3). The degree of urbanization is also determined by the main anthropic uses
of such natural habitats, the use frequency or intensity, and consequently, the
potential pollution and habitat degradation (Table 16.1).

16.2.2 Sampling Design: Surveys of Plant, Bird and Insect

Species

For each forest type, we selected four forest patches (replicates) located at the same
elevation range (38 to 170 m above sea level (m.a.s.l.). These were at least 150 m
apart from each other and from other types of structures, as well as at least 50 m from
the edge of the same forest type, in order to avoid the influence of other environ-
ments. In the case of peatlands, we selected 12 urban, 4 peri-urban and 4 non-urban
locations (elevation range 20 to 200 m.a.s.l.) separated by at least 150 m from each
other.
Vegetation surveys were carried out in each patch (forests and peatlands) during
2016–2017, using a circular plot of approximately 30 m radius centered in each
sampling site. In each plot, we identified vascular plants (Dicotyledonae,
Monocotyledonae and Pteridophytae) at the species level (Moore 1983; Correa
1969–1998). We estimated ground cover of vascular plants using a modification
of the relevé method of Braun-Blanquet proposed by Pauchard et al. (2000). The
relevé method involves describing recognizable units in the vegetation of a region by
the characterization of the vegetation in a single representative standard plot—a
relevé—within each unit. The relevés from many units are then analyzed to develop
descriptions and classifications of the vegetation in the study region and have been
increasingly used in other kinds of vegetation studies as a practical, relatively fast
426 R. Soler et al.

means of collecting information on vegetation (Minnesota Department of Natural

Resources 2013). Ground cover was estimated for each species separately and then
values were added to obtain family and total vascular plant cover. The complemen-
tary covers to reach 100% ground cover (data not shown) were: bryophyte (mosses
and liverworts), woody debris (> 3 cm diameter), and bare soil including litter.
Species richness was calculated as the total number of vascular plant species
identified in each replica of treatments. Voucher specimens were deposited in the
Herbarium of Tierra del Fuego at the Centro Austral de Investigaciones Científicas
(Austral Center of Scientific Research, CADIC-CONICET) in Ushuaia, Argentina.
Bird surveys in forest patches were conducted during the 2016–2017 breeding
season in a 4-h period at sunrise, always under similar climatic conditions (i.e.,
avoiding rainy or foggy days). We identified bird assemblages by using the point-
count method with unlimited distances, which had been used widely in previous
studies in Tierra del Fuego (see Deferrari et al. 2001; Lencinas et al. 2005, 2014,
2019). We selected one observation point in each forest patch (n ¼ 12), with two
observers during each survey conducting direct sightings (with binoculars) during
13 min of effective observation period (leaving 2 min prior to recording to allow for
birds’ habituation to the observers). We registered the birds to the species level
according to the classification proposed by Remsen et al. (2019). From these
surveys, we calculated bird species richness and relative frequency (number of
individuals). Forests patches were revisited three times per month for bird observa-
tions during the complete breeding season (from October to February). Species
richness represent the total number of species recorded during that period, and the
relative abundance was calculated as the sum of the total number of individuals
observed for the same species during visits.
Epigean insects were collected in peatlands during January–February 2016.
Sampling was done using pitfall traps (plastic containers 12 cm in diameter and
14 cm height) buried and filled to 1/3 of their volume with soapy water (300 ml) to
trap and kill arthropods which fell in. We used pitfall traps in sets of five in each plot,
and contents of the five traps in a plot were pooled and used as a single sample in
order to calculate total capture per sample (Lencinas et al. 2014). Traps were
arranged placing one at the center and the remaining four at 5 m from the first,
and at 90 degrees from each other, and were left open at ground level for one week
before being collected. We obtained 20 samples from 100 pitfall traps. All samples
were taken to the laboratory for specimen identification and quantification. Identifi-
cations were performed under a binocular dissecting microscope to genus or species
level when possible (Roig-Juñent et al. 2002; Marvaldi and Lanteri 2005; Posadas
2012). Due to a lack of complete taxonomic data on Patagonian beetles, some
specimens could not be determined to the species level. We employed the recogniz-
able taxonomic unit or morphospecies concept (Oliver and Beattie 1993; Gerlach
et al. 2013) when the former could not be determined (hereafter, “species”). This has
been demonstrated to be a good tool for insect diversity studies in Patagonian
ecosystems, such as Nothofagus forests (Lencinas et al. 2014; Sola et al. 2016;
Cárcamo et al. 2019).
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 427

16.2.3 Statistical Analyses

Data collected were used to determine species richness, relative abundance (ground
cover for plants, and number of individuals for birds and insects), Shannon-Wiener
diversity (H0 ) and Pielou evenness (J’) indices of plants and birds in each forest, and
plants and insects in each peatland types (Pielou 1975). We used one-way ANOVAs
after validation of the statistical assumptions of homoscedasticity (homogeneity) and
normality, to evaluate species richness, relative abundance, Shannon-Wiener diver-
sity and Pielou evenness indices of vascular plants, birds and insects, considering the
category of forests and peatlands (urban, peri-urban and non-urban) as the main
factor.
Also, differences in assemblage composition of plants, birds and insects along the
urban categories were examined by the non-Metrical Multidimensional Scaling
(NMDS) ordination method using a manual methodology, with a Bray-Curtis
distance and 250 iterations. A Monte Carlo test was used to evaluate stress in
randomized data; probability was presented for each axis.
Finally, we calculated the abundance-based Chao-Sørensen similarity index
(Chao et al. 2005) to determine differences among categories of urbanization, both
for forests and peatlands, analyzing each taxonomic group separately. This index,
varying between zero and one, considers explicitly the relative abundance of both
common and rare species, and estimates the extent of shared species taking into
account unseen shared species, based on the number of observed rare (singletons and
doubletons, as well as unique and duplicate), shared species between two sites. The
use of this index is recommended when study sites could be under-sampled and
contain only a substantial fraction of the rare species (Chao et al. 2005). We used the
software EstimateS (Colwell 2009) for each pairwise evaluation and compared
averages and standard deviation for each urbanization category.

16.3 Results and Discussion

Plants, birds and insects have been widely used as bio-indicators of human impact on
natural ecosystems (Lencinas et al. 2005, 2014, 2017; Gerlach et al. 2013; Rozzi and
Jiménez 2014; Contador et al. 2015) and urban biodiversity islands (Noreika et al.
2015; Matthies et al. 2017). These three groups, due to high latitude and extreme
climatic conditions of Southern Patagonia, sustain the greatest biological diversity in
the natural ecosystems, with many endemic species and guilds (Roig-Juñent et al.
2002; Díaz 2012; Rozzi and Jiménez 2014; Cárcamo et al. 2019). Plants are among
the more characteristic and recognized organisms for people, both in forests and
peatlands, while birds are the most conspicuous groups in forests, and arthropods are
more visible and easily distinguishable in peatlands (open areas).
428 R. Soler et al.

16.3.1 Vascular Plants

In Ushuaia city, vascular plants were affected differently in forest patches and
peatlands. Species richness did not strongly differ along the urbanization categories,
but the relative abundance of vegetation and species diversity indexes were strongly
modified in urban and peri-urban peatlands (Table 16.2). Urbanization significantly
increased the relative abundance of vascular plants, due to the increase of some plant
species favored by microclimatic modifications (e.g., light, soil moisture).
According to our data, forbs and graminoids such as Gunnera magellanica, Acaena
magellanica, and Carex sp. doubled their cover, while other species (e.g., the native
fern Blechnum penna-marina and the exotic herbs Achilea millefolium, Rumex
crispus, Taraxacum officinale, Poa annua), which were absent in peri-urban and
non-urban peatlands, entered in the urban ones. Soil drainage, building and protec-
tion against extreme temperatures and winds could favor herbaceous development
on peatlands (Pinceloup et al. 2020).

Table 16.2 Species richness, relative abundance, Shannon-Wiener diversity (H0 ) and Pielou
evenness (J’) indices of vascular plants, birds and insects occurring in urban, peri-urban and
non-urban forests and peatlands of Ushuaia city
Type Richness Abundance H0 J’
Forest plants Urban 16.0 67.2 4.2 0.82
Peri-urban 19.5 48.2 2.4 0.83
Non-urban 15.5 37.5 2.2 0.86
F (p) 0.40 (0.679) 0.80 0.15 (0.859) 0.15 (0.865)
(0.480)
Forest birds Urban 9.8 93.7 b 1.9 0.82
Peri-urban 10.0 62.5 a 2.0 0.88
Non-urban 7.8 39.8 a 1.8 0.87
F (p) 1.54 (0.266) 13.98 (0.002) 0.98 (0.411) 0.71 (0.518)
Peatland plants Urban 10.7 89.6 b 1.4 a 0.60 a
Peri-urban 8.0 86.7 b 1.1 a 0.53 a
Non-urban 13.5 66.2 a 2.1 b 0.81 b
F (p) 1.59 (0.232) 14.5 (<0.001) 3.81 (0.0502) 4.53 (0.048)
Peatland insects Urban 5.6 22.4 a 1.17 0.68
Peri-urban 8.3 44.3 ab 1.38 0.65
Non-urban 8.8 92.2 b 1.29 0.60
F (p) 1.75 (0.206) 9.34 0.18 (0.838) 0.13 (0.883)
(0.002)
F (p) Fisher statistic with probability in parentheses. Different letters show differences by Tukey test
at p < 0.05. Values followed by different letters in each column and for each factor are signifi-
cantly different according to the Tukey test at a p  0.05
Relative abundance shows: vascular plants¼ total ground cover (%), birds ¼ average of total
observations along five months during the breeding season (n
individuals), and insects ¼ average
of adult individuals collected in pitfall traps for 15 days in the summer season (n
individuals)
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 429

Fig. 16.4 Results of analysis of differences in assemblage composition of plants examined by the
Non-Metrical Multidimensional Scaling (NMDS) ordination method, based on understory plants
composition in different urban categories of forest patches (a) and peatlands (b)

On the other hand, urban and peri-urban forests showed changes in their com-
munity composition in some patches (Fig. 16.4a), by the introduction of exotic
cultivated shrubs (Ribes idaeus and Ribes rubrum) and potentially invasive forbs
(Ranunculus acris and Hypochaeris radicata), which could be intentionally planted
by people or escaped from home gardens, orchards and parks. Urbanization also
differently influenced the Shannon diversity and Pielou evenness indices for vascu-
lar plants in forests patches and peatlands. While both indices were quite similar in
forest patches along the urban categories, Shannon diversity and Pielou evenness
indices of urban peatlands decreased by half compared to those in non-urban
peatlands. Plant composition in urban peatlands was similar to vegetation in peri-
urban areas but differed from non-urban peatlands located far from urbanization
(Fig. 16.4b).

16.3.2 Birds

With respect to birds, urban assemblages in Ushuaia’s forests are conformed by bird
species commonly observed in Nothofagus forests (Deferrari et al. 2001; Lencinas
et al. 2005), which correspond to thirteen families of Passeriformes, and one family
of five orders (Charadriiforme, Falconiforme, Pelecaniforme, Piciforme,
Psittaciforme (Benítez et al. 2020). Some species detected in these urban forests
are endemic to Patagonia (i.e. Aphrastura spinicauda, Phrygilus patagonicus,
Pygarrichas albogularis, Enicognathus ferrugineus and Campephillus
430 R. Soler et al.

magellanicus) (Deferrari et al. 2001). Insectivore and omnivore birds are the most
important groups, being more than a half of all resident species (Benítez et al. 2020).
In our study, species richness remained similar in peri-urban and non-urban
patches (Table 16.2). However, bird abundance significantly increased in urban
forests, with slight changes in peri-urban and non-urban forest patches. This was
explained by the high abundance of generalist bird species and/or the occurrence of
non-specialist forest birds in altered urban patches (pers. obs.). We observed that
urban forests offered different food sources (e.g., waste, fruits/seeds) and nesting
alternatives (e.g., open places, canopy openings) compared to native forests, which
coincides with other studies (Marzluff and Rodewald 2008). This favored the
occurrence of bird species from open habitats such as Vanellus chilensis (Marín
2014).
The availability of extra food and greater hunting possibilities favored the
occurrence of scavenging raptors such the Milvago chimango, as was also reported
by Biondi et al. (2005). Moreover, urbanization in Ushuaia increased the frequency
of exotic bird species (e.g., Passer domesticus). Contrary to this, native birds such as
woodpeckers and hole-nesters birds (C. magellanicus, P. albogularis, Troglodytes
aedon and A. spinicauda) showed an increasing trend in richness and abundance
from the urban to the periphery environments. These charismatic species prefer
forests with dense vegetation (Vergara and Schlatter 2006), but they can be seen
while feeding in urban habitats (pers. obs.). Analyzing the whole species assem-
blages of urban forests (Fig. 16.5a), great differences in biodiversity along the
urbanization categories were possible to distinguish. Plant and bird communities
in urban forests differed from the communities of non-urban forests (Figs. 16.4a and
16.5a).

Fig. 16.5 Results of analysis of differences in bird abundance in different urban forests (a), and
insect abundance in different urban peatlands (b), examined by the Non-Metrical Multidimensional
Scaling (NMDS) ordination method
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 431

16.3.3 Invertebrates

Peatland invertebrate communities are almost unknown, and even more the effects
that anthropic disturbances have on their diversity. Recent surveys in urban and
non-urban peatlands of Ushuaia city (Sola et al. 2018) indicate the presence of a wide
variety of invertebrates: snails (Gastropoda), spiders (Araneae), and harvestmen
(Opiliones), mites (Acari), woodlice (Isopoda), and many orders of insects. Due to
their diversity, abundance, easy sampling, and rapid response to environmental
changes, insects are useful bio-indicators of habitat degradation (Bolger et al.
2000; Gerlach et al. 2013; Moreno et al. 2013), and have been widely used to
study the effects of habitat modification even in south Patagonia (Sola et al. 2016;
Lencinas et al. 2014, 2017, 2019; Cárcamo et al. 2019). In particular, the study of
ground dwelling insects has important functional implications because of their
ecological roles, as they include predators, scavengers, and herbivores.
According to our results, the main insect orders present in all categories included
beetles (Coleoptera), moths (Lepidoptera), true bugs (Heteroptera), springtails
(Collembola), leafhoppers (Auchenorrhyncha), and wasps (Hymenoptera). Insect
richness in peatlands (including Coleoptera, Lepidoptera, Heteroptera,
Auchenorrhyncha, Collembola and Hymenoptera) remained similar throughout the
urbanization categories (Table 16.2). However, insect abundance significantly
decreased in urban peatlands compared with those in non-urban; while peri-urban
peatlands showed intermediate values. The most affected groups were Coleoptera,
Auchenorrhyncha and Collembola, which reduced by half the number of individuals
within these orders. All three of these groups are weaker fliers (in the case of beetle
species sampled), making them more susceptible to local impacts. Furthermore,
Auchenorrhyncha are herbivorous and although the ecology of local species is
unknown, they could be susceptible to changes observed in local flora at each site.
Collembolans, as other detritivores, can be associated with available organic matter;
a fall in their abundance could be indicative of reduced productivity, and may also
result in slowed decomposition rates and nutrient cycling, essential in these cold,
high latitude landscapes. Finally, beetles sampled fell under the categories of
scavengers or predators (e.g., Curculionidae, Carabidae) indicating a higher trophic
position. These can ultimately serve as indicators of the health of plants and prey in
the ecosystem, both in diversity and abundance. Although richness values recorded
in our study did not significantly differ among the three urbanization levels, they
hide changes in assemblages (Fig. 16.5b). Nevertheless, the proximity between
points entails the occurrence of some species both in urban and non-urban sites.
The Chao-Sørensen similarity index varied among taxonomic groups and urban
categories (Fig. 16.6), showing different tendencies among urbanization levels.
Plants showed less similarity for all urban categories in forests (0.57 in average)
than in peatlands (0.73 on average), with the lower values in urban patches. Greatest
similarities in plants occurred in peri-urban forests and in non-urban peatlands,
where we also found the lowest variation. In contrast, birds showed great similarity
values (0.85 in average) in all urban categories, with slightly higher values in urban
432 R. Soler et al.

Fig. 16.6 Abundance-

based Chao-Sorensen
similarity index
(mean  SE) for forest
plants and birds, and
peatland plants and insects
among the urban categories
(urban, peri-urban, and
non-urban)

than in peri-urban than in non-urban forests. On the other hand, insects showed high
similarity only in non-urban peatlands (0.90), with very low values in urban and
peri-urban patches (0.26 in average), highlighting the greater sensitivity of this group
compared with plants and birds.

16.3.4 Main Factors Affecting Biodiversity in Urban Forests

and Peatlands in Ushuaia City

Despite some differences in richness, abundance, biodiversity indices, and compo-

sition for vascular plants, birds and insects, urban and peri-urban forests and
peatlands still conserve several species and characteristics similar to non-urban
ecosystems. However, the occurrence of new species, mainly plants, introduced
into these urban ecosystems, modified the original communities into new assem-
blages, which offer another food supply to the fauna (i.e., birds and insects)
(Marzluff and Rodewald 2008). The probability of greater dispersion of these
newly introduced exotic plants in the non-urban ecosystem could depend on the
mobility of dispersing agents, and on the connectivity and distance between urban
forests and peatland patches to non-urban ecosystems. Previous studies affirm that
urban green areas are ecological islands isolated from others in and around the city
(Ceplová et al. 2017), but this depends on the size of cities and green areas. In
addition, the relative importance of local- versus larger-scale variables in influencing
urban biodiversity remains under discussion and will vary with the scale dependen-
cies of different taxa (Goddard et al. 2010). The influence of local factors, especially
patch size and quality, for urban forests and peatlands in Ushuaia city has not been
analyzed, either for urban planning or conservation strategies.
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 433

The preservation of forests and peatlands as biodiversity islands within the urban
matrix generates benefits for the environment and also for citizens living there,
promoting ecological and socio-cultural functions (Rozzi et al. 2012) that would
be completely lost if such natural islands were not preserved. Unfortunately, the link
between the identity of inhabitants and care of regional habitats in Ushuaia is weak
(Orzanco 1999; Kizman 2012) and much of the ancestral knowledge about natural
ecosystems and their species has been lost. Moreover, the inadequate planning of
public services (e.g., streets, sewage) to attend to the growth rate of the city, and
bulky waste management are considered the main conflicts between urbanization
and sustainability in Ushuaia city (Orzanco 1999). All these factors negatively
influence the structure and functioning of biodiversity islands in the city (Fig. 16.7).
Because urban development is guided by human values, there is a need to inform
people about the relevance of natural ecosystems supporting multiple services that
increase human well-being and encourage caring for the environment. Ushuaia is a
unique city in Argentina where peatlands are part of the urban space. However, their
value as part of the natural heritage has not yet been recognized by society, and
peatlands are considered obstacles to urban development rather than areas of natural,
recreational and/or educational interest. However, peatlands were recently declared
Environmental Reserves for their conservation and protection as Historical and
Cultural Heritage by the Municipality of Ushuaia (Environment Office of Ushuaia
2018).
Although in Ushuaia urban forests and peatlands are connected to the natural
undisturbed forests and peatlands occurring in the surrounding landscape, especially
by birds and also by some insects (e.g., wasps) and some plants, this is generally not
common knowledge for local people (Environment Office of Ushuaia 2018). This
generates a twofold problem: on one hand, urban green areas do not receive the
attention they rightfully deserve and are frequently modified irreversibly; and on the
other, natural landscapes on the periphery are in danger from freely introduced

Fig. 16.7 Conceptual framework to understand the main threats (top panels) affecting biodiversity
(species richness and abundance) in urban forests and peatlands in Ushuaia city. Biodiversity
islands serve as refuge for some native species that manage to survive despite structural changes
and isolation from other natural environments, although they are also prone to invasion by
introduced species (e.g., exotic plants) dispersed from homegardens, parks, etc
434 R. Soler et al.

exotics, either by negligence (weeds, insects) or intent (ornamental plants, pets). We

recommend controlling and avoiding the installation of exotic plants, especially trees
and bushes with fruits eaten by birds, in these urban green areas, to limit their spread
and/or invasion to undisturbed environments.

16.4 Conclusions and Recommendations

Our results confirm that the urban forests and peatlands of Ushuaia are valuable
reservoirs of biodiversity, even under anthropogenic influence. We recommend
preserving existing natural green areas for their role as biodiversity islands in the
urban landscape, due to their function as a refuge for native plants, birds and insects.
In order to increase biodiversity and improve the connectivity of biodiversity islands
within Ushuaia city we consider necessary the preservation and/or delimitation of
new natural green areas (e.g., within new neighborhoods). Moreover, we highlight
their value compared with parks and squares since these are non-natural, and mostly
contain exotic vegetation (e.g., Salix sp., Populus sp., Pseudotsuga sp.).
According to our results, the urban peatlands would be more susceptible to
deterioration and loss of species typical of these habitats than the urban forests.
Therefore, it is important to implement urgent conservation plans to avoid the loss of
these remaining natural habitats (indeed, some of the studied green areas have
nowadays already been transformed and urbanized). Moving forward, the level
and magnitude of the urbanization impact (i.e., distance to the urban center, trans-
formation degree in the surroundings) on biodiversity deserve more analyses.
Urban islands must contribute to strengthening the link between society and
nature, while still offering refuge, food and habitat for plant and animal species.
These ecosystems also play an important role in maintaining the minimum environ-
mental (e.g., carbon sinks, water purification) and socio-cultural functions (e.g.,
recreation, sense of belonging).
We also recommend including environmental education programs to facilitate the
understanding of the importance of these biodiversity islands in the city, spread
knowledge about native species (especially those inconspicuous or less charismatic),
and promote the sustainable use of these environments. For local governments and
planners, it is essential to plan the retention of natural green spaces when planning
new neighborhoods, buildings, schools and parks. Moreover, conserving, designing,
and managing urban green spaces requires balancing human perceptions, needs, and
use with ecological requirements for preserving and enhancing biodiversity.

References

Benítez J, Pizarro JC, Blazina AP, Lencinas MV (2020) Response of bird communities to native
forest urbanization in one of the southernmost city of the world. Urban Urban Gree 126887.
[Link]
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 435

Biondi LM, Bó MS, Favero M (2005) Dieta del chimango (Milvago chimango) durante el periodo
reproductivo en el sudeste de la provincia de Buenos Aires, Argentina. Ornitol Neotrop 16:31–
42
Bolger DT, Suarez AV, Crooks KR, Morrison SA, Case TJ (2000) Arthropods in urban habitat
fragments in southern California: area, age and edge effects. Ecol Appl 10:1230–1248
Burghardt KT, Philips CR, Tallamy DW, Shropshire KJ (2010) Non-native plants reduce abun-
dance, richness, and host specialization in Lepidoptera communities. Ecosphere 1(5):11. https://
[Link]/10.1890/ES10-00032.1
Cárcamo JR, Contador T, Gañán M, Convey P, Kennedy J, Rozzi R (2019) Altitudinal gradients in
Magellanic sub-Antarctic lagoons: the effect of elevation on freshwater macroinvertebrate
diversity and distribution. Peerj 7:e7128–e7128. [Link]
Ceplová N, Kalusová V, Lsosová Z (2017) Effects of settlement size, urban heat island and habitat
type on urban plant biodiversity. Landsc Urban Plan 159:15–22
Chao A, Chazdon RL, Colwell RK, Shen TJ (2005) A new statistical approach for assessing
similarity of species composition with incidence and abundance data. Ecol Lett 8:148–159
Colwell RK (2009) EstimateS: statistical estimation of species richness and shared species from
samples. [Link]
Contador T, Kennedy JH, Rozzi R, Villarroel JO (2015) Sharp altitudinal gradients in Magellanic
sub-Antarctic streams: patterns along a fluvial system in the Cape Horn biosphere reserve. Polar
Biol 38:1853–1866
Correa MN (1969–1998) Flora Patagónica. Colección Científica INTA Tomo 8. Partes II, III,
IVb, V, VI y VII. Buenos Aires
Deferrari G, Camilión C, Martínez Pastur GM, Peri PL (2001) Changes in Nothofagus pumilio
forest biodiversity during the forest management cycle. 2: birds. Biodivers Conserv 10:2093–
2108
Díaz S (2012) Biología y conservación de la cachaña (Enicognathus ferrugineus) en Argentina. El
Hornero 27:17–25
Environment Office of Ushuaia (2018) Plan de manejo de la Reserva Natural Urbana Bahía
Encerrada, Municipalidad de Ushuaia, Tierra del Fuego, 98 p
Gerlach J, Samways M, Pryke J (2013) Terrestrial invertebrates as bioindicators: and overview of
available taxonomic groups. J Insect Conserv 17:831–850
Goddard MA, Dougill AJ, Benton TG (2010) Scaling up from gardens: biodiversity conservation in
urban environments. Trends Ecol Evol 25:90–98
Gómez-Baggethun E, Barton DN (2013) Classifying and valuing ecosystem services for urban
planning. Ecol Econ 86:235–245
Hartley SE, Jones TH (2008) Insect herbivores, nutrient cycling and plant productivity. In: Weisser
WW, Siemann E (eds) Insects and ecosystem function. Springer, Berlin, pp 27–52
INDEC (2010) Censo Nacional De Población, Hogares y Viviendas 2010. Documento
Metodológico Preliminar. Available at: [Link]
Accessed 1 Apr 2020
Kizman S (2012) Diagnóstico participativo: desarrollo de actividades sustentables. Parque Nacional
Tierra del Fuego. APN, Tierra del Fuego. [Link]
Lencinas MV, Martínez Pastur G, Medina M, Busso C (2005) Richness and density of birds in
timber Nothofagus pumilio forests and their unproductive associated environments. Biodivers
Conserv 14:2299–2320
Lencinas MV, Martínez Pastur G, Gallo E, Cellini JM (2014) Decreasing negative impacts of
harvesting over insect communities using variable retention in southern Patagonian forests. J
Insect Conserv 18:479–495
Lencinas MV, Sola FJ, Martínez Pastur G (2017) Variable retention effects on vascular plants and
beetles along a regional gradient in Nothofagus pumilio forests. For Ecol Manag 406:251–265
Lencinas MV, Sola FJ, Cellini JM, Peri PL, Martínez Pastur GJ (2019) Land sharing in South
Patagonia: conservation of above-ground beetle diversity in forests and non-forest ecosystems.
Sci Total Environ 690:132–139
436 R. Soler et al.

Lososová Z, Horsák M, Chytry M, Cejka M, Danihelka T, Fajmon J et al (2011) Diversity of

Central European urban biota: effects of human-made habitat types on plants and land snails. J
Biogeogr 38:1152–1163
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press,
Princeton
Maestre FT, Castillo-Monroy AP, Bowker MA, Ochoa-Hueso R (2012) Species richness effects on
ecosystem multifunctionality depend on evenness, composition and spatial pattern. J Ecol 100:
317–330
Marín M (2014) Distribución, fenología reproductiva, e historia natural del queltehue (Vanellus
chilensis) en la zona central de Chile. Bol Mus Nac Hist Nat 63:119–126
Marvaldi AE, Lanteri AA (2005) Key to higher taxa of South American weevils based on adult
characters (Coleoptera, Curculionoidea). Rev Chil Hist Nat 78:65–87
Marzluff J, Rodewald A (2008) Conserving biodiversity in urbanizing areas: nontraditional views
from a bird’s perspective. Cities Environ 1(2):6–27
Marzluff JM, Bradley G, Shulenberger E, Endlicher W, Alberti M, Bradley G, Ryan C,
ZumBrunnen C, Simon U (eds) (2008) Urban ecology. An international perspective on the
interaction between humans and nature. Springer, Berlin
Matthies SA, Rüter S, Schaarschmidt F, Prasse R (2017) Determinants of species richness within
and across taxonomic groups in urban green spaces. Urban Ecosyst 20:897–909
McGregor-Fors I, Morales-Pérez L, Schondube JE (2011) Does size really matter? Species–area
relationships in human settlements. Divers Distrib 17:112–121
Millennium Ecosystem Assessment (2005) Ecosystems and human well-being. Island Press,
Washington
Miller JR (2005) Biodiversity conservation and the extinction of experience. Trends Ecol Evol 20:
430–434
Minnesota Department of Natural Resources (2013) A handbook for collecting vegetation plot data
in Minnesota: The relevé method. 2nd ed. Minnesota Bio-logical Survey, Minnesota Natural
Heritage and Nongame Research Program, and Ecological Land Classification Program. Bio-
logical Report 92. St. Paul, Minnesota Department of Natural Resources
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity Islands. Strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human-dominated environments. Topics in Biodiversity and Conservation,
Springer, Cham, pp 1–35
Moore DM (1983) Flora of Tierra del Fuego. Anthony Nelson
Moreno ML, Fernández MG, Molina SI, Valladares G (2013) The role of small woodland remnants
on ground dwelling insect conservation in Chaco Serrano, Central Argentina. J Insect Sci 13:40.
[Link]
Murphy D (1999) Challenges to biological diversity in urban areas. In: Wilson EO
(ed) Biodiversity. National Academy Press, Washington, pp 71–76
Nilon CH, Aronson MFJ, Cilliers SS, Dobbs C, Frazee LJ, Goddard MA, O’Neill KM, Roberts D,
Stander EK, Werner P, Winter M, Yocom KP (2017) Planning for the future of urban biodi-
versity: a global review of city-scale initiatives. Bioscience 67:332–342
Noreika N, Pajunen T, Kotze DJ (2015) Urban mires as hotspots of epigaeic arthropod diversity.
Biodivers Conserv 24(12):2991–3007
Oliver I, Beattie AJ (1993) A possible method for the rapid assessment of biodiversity. Conserv
Biol 7(3):562–568
Orzanco MG (1999) Problemas ambientales detectados por la población de Ushuaia (Tierra del
Fuego, Argentina). Invest Geogr 40:85–98
Pauchard A, Ugarte E, Millán J (2000) A multiscale method for assessing vegetation baseline of
environmental impact assessment (EIA) in protected areas of Chile. In: McCool SF, Cole DN,
Borrie WT, O’Loughlin J (eds) Wilderness science in a time of change conference. USDA
Forest Service, Rocky Mountain Research Station (United States), pp 111–116
Pielou EC (1975) Ecological diversity. Wiley, New York
16 Biodiversity Islands at the World’s Southernmost City: Plant, Bird. . . 437

Pinceloup N, Poulin M, Brice M-H, Pellerin S (2020) Vegetation changes in temperate

ombrotrophic peatlands over a 35 years period. PLoS One 15(2):e0229146. [Link]
1371/[Link].0229146
Posadas P (2012) Species composition and geographic distribution of Fuegian Curculionidae
(Coleoptera: Curculionoidea). Zootaxa 3303:1–36
Reich PB, Tilman D, Isbell F, Mueller K, Hobbie S, Flynn D (2012) Impacts of biodiversity loss
escalate through time as redundancy fades. Science 336(6081):589–592
Remsen JV, Areta JI, Cadena CD, Claramunt S, Jaramillo A, Pacheco JF, Robbins MB, Stiles FG,
Stotz DF, Zimmer KJ (2019) A classification of the bird species of South America. American
Ornithologists’ Union. [Link]
Roig-Juñent S, Crisci JV, Posadas P, Lagos S (2002) Áreas de distribución y endemismo en zonas
continentales. In: Costa C, Vanin SA, Lobo JA, Melic A (eds) Red Iberoamericana de
Biogeografía y Entomología Sistemática (PrIBES). Sociedad Entomológica Aragonesa, Zara-
goza, pp 247–266
Rozzi R, Jiménez JE (2014) Magellanic sub-antarctic ornithology: first decade of long-term bird
studies at the Omora Ethnobotanical Park, Cape Horn biosphere reserve, Chile. University of
North Texas Press, Dallas
Rozzi R, Massardo F, Silander J, Dollenz O, Connolly B, Anderson C, Turner N (2003) Native and
exotic tree species in the squares of the Magellan Region. An Inst Patagon 31:27–42
Rozzi R, Armesto JJ, Gutiérrez JR, Massardo F, Likens GE, Anderson CB, Poole A, Moses KP,
Hargrove E, Mansilla AO, Kennedy JH, Willson M, Jax K, Jones CG, Callicott JB, Arroyo
MTK (2012) Integrating ecology and environmental ethics: earth stewardship in the southern
end of the Americas. Bioscience 62(3):226–236
Sola FJ, Peri PL, Huertas L, Martínez Pastur GJ, Lencinas MV (2016) Above-ground arthropod
community structure and influence of structural-retention management in southern Patagonian
scrublands, Argentina. J Insect Conserv 20:929–944
Sola FJ, Soler R, Huertas A, Lencinas MV (2018) Urbanización y diversidad de invertebrados en
turbales de Tierra del Fuego. In: Proc XXVIII Reunión Argentina de Ecología, 29 Oct - 2 Nov,
Mar del Plata, Argentina
Tallamy DW, Ballard M, D’Amico VD (2010) Can alien plants support generalist insect herbi-
vores? Biol Invasions 12:2285–2292
Thomas K, Geller L (2013) Urban forestry: toward an ecosystem services research agenda. The
National Academies Press, Washington
Tuhkanen S (1992) The climate of Tierra del Fuego from vegetation geographical point of view and
its ecoclimatic counterpartes elsewhere. Acta Bot Fenn 145:1–64
Vergara PM, Schlatter RP (2006) Aggregate retention in two Tierra del Fuego Nothofagus forests:
short-term effects on bird abundance. For Ecol Manag 225:213–224
Zerbe S, Maurer U, Schmitz S, Sukopp H (2003) Biodiversity in Berlin and its potential for nature
conservation. Landsc Urban Plan 62:139–148
Chapter 17
Paradise Lot: A Temperate-Climate Urban
Agroforestry Biodiversity Island

Eric Toensmeier

Abstract Paradise Lot is a garden and home site occupying 400 square meters in the
cold temperate subhumid northeastern United States. Emerging from the permacul-
ture movement, its agroforestry approach emphasizes perennial crops. The goal is to
provide ecosystem services while producing food. Data are presented on plant
biodiversity, including the numbers of species that are native and/or provide impor-
tant functions including nectar for pollinators, wildlife cover, nitrogen fixation and
groundcover. Anecdotal evidence is provided suggesting that the site is a haven for
invertebrates and offers some benefits to vertebrates despite its small size. This case
study demonstrates that food production and ecosystem function can go hand in
hand, with the wide variety of plants incorporated here providing various ecological
roles and human uses. Located in an urban area with a depauperate flora and fauna,
the site serves as a biodiversity island.

Keywords Agroforestry · Food forest · Homegarden · Perennial crops ·

Permaculture · Urban agriculture

17.1 Introduction

Paradise Lot is an urban agroforestry garden, inspired by the permaculture move-

ment, which serves as a biodiversity island. Designed and managed by the author
and Jonathan Bates, the project has been documented in several books (Jacke and
Toensmeier 2005; Toensmeier and Bates 2013).
Agroforestry systems incorporate woody plants into agriculture in a variety of
ways. Paradise Lot practices both tree intercropping (growing annual crops and
woody plants together) and multistrata agroforestry (perennial systems with multiple
layers of crops) (Toensmeier 2016).

E. Toensmeier (*)
Global Evergreening Alliance, Perennial Agriculture Institute, Yale University,
Holyoke, MA, USA

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 439
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
440 E. Toensmeier

Toensmeier and Bates were very involved in the world of permaculture as they
began the project (Toensmeier and Bates 2013). Permaculture is an international and
regional movement that disseminates and practices a worldview, using a best
practices framework and design systems (Ferguson and Lovell 2013). Ecosystem
mimicry is one of permaculture’s core practices, and it is at the heart of Jacke and
Toensmeier (2005), which Toensmeier was writing in 2004 when the garden was
established. This work lays a foundation for “edible forest gardens”, a home-scale
version of multistrata agroforestry drawing heavily on permaculture design and
eastern forest ecosystem mimicry. This form of garden has become known as a
“food forest” and has spread rather rapidly in the last two decades, due in part to
appreciation of its multifunctional approach that provides ecosystem services while
producing food (Clark and Nicholas 2013).
While this agroforestry practice is relatively new in temperate climate, it has been
practiced for thousands of years in the tropics. The “tropical homegarden” produces
highly biodiverse perennial and annual crops, and often small livestock, right around
the home. Homegardens have been described by scientists as “the epitome of
sustainability” for their impressive impacts on biodiversity and other ecosystem
services (Nair 2006; Nair and Kumar 2006). Indeed, they have been ranked as
having the highest biodiversity of any anthropogenic ecosystem (Nair 2006).
These systems were a major, though little credited, inspiration and model for the
early innovators of permaculture (Ferguson and Lovell 2014).
At the garden level, edible forest gardens strive to mimic the architecture (layers,
density, patterns, and diversity), social structure (niches, relationships, and commu-
nities), soil ecology and fertility dynamics, and successional dynamics of natural
ecosystems. In particular, it is aggrading mid-successional ecosystems that are the
most important model, as in humid temperate climates these feature high rates of
biomass growth and high diversity of perennial food plants (Jacke and Toensmeier
2005). This is reflected in the central importance of polycultures (growing more than
one species in the same place, while attempting to minimize competition and
maximize cooperation). Toensmeier and Bates (2013) observed wild species asso-
ciations and replicated them, sometimes with slight modifications, in the garden. For
example, elderberry (Sambucus canadensis) often serves as a trellis for groundnut
(Apios americana) in the wild, and this association has been used as the basis for a
polyculture in the garden, albeit with improved, higher-yielding forms of each.

17.2 Site History

The site has been documented in two books. Jacke and Toensmeier (2005) uses the
site as a case study of ecological site design. Toensmeier and Bates (2013) is a
garden memoir telling the story of the garden’s inspiration and development.
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 441

17.2.1 Goals

The garden designers began the process of clarifying goals before identifying a site,
and continued the process once moving in in 2004. These goals are published in
Jacke and Toensmeier (2005), for which the site was a design process case study, and
in Toensmeier and Bates (2013), which includes some revised goals.
The initial goal was to create a demonstration site to test the “edible forest
garden” model in the northeastern United States. Specific goals included: “creating
an intensively managed backyard foraging paradise;” “a mega-diverse living ark of
useful and multifunctional plants from our own bioregion and around the world;” “to
bring our dead and blighted backyard to life. . . creating a lush, semiprivate oasis that
inspires our neighbors to plant their own;” and “to serve as a refuge in our
biologically impoverished neighborhood” (Jacke and Toensmeier 2005). To expand
a bit on the biodiversity oasis aspect of the goals, Toensmeier and Bates (2013)
write: “It was too much to hope that moose and great blue herons would come to
visit, but we did want to create a haven for smaller forms of wildlife, particularly
those birds, insects and amphibians that help control pests.” It was hoped that the
garden would serve as an inspiration to gardeners in the neighborhood.
In terms of food production, the goal was to have as long a harvest season as
possible for production of fruits and vegetables. Emphasis was on perennial crops in
diverse polyculture systems. The design incorporated nitrogen-fixing plants to aid in
restoration of degraded soils and maintain fertility, plants to attract pollinators and
beneficial insects, groundcovers to serve as a living mulch and suppress weeds, and
thicket-forming and evergreen plants for wildlife cover. Other elements included
beds for annual crops, a greenhouse, a pond for aquatic vegetable production, and
abiotic elements including patio, toolshed, compost piles, and vehicle access (Jacke
and Toensmeier 2005).
In species selection, an emphasis was placed on multifunctionality. If a species
performs more than one ecosystem function and/or provides more than one human
use, it saves space – a key issue in this small, intensive garden. This kind of
multifunctionality is a key principle of permaculture (Ferguson and Lovell 2014).

17.2.2 Site Analysis and Assessment

The Paradise Lot site is located in Holyoke, Massachusetts, in the northeastern

United States. Climate is cold temperate subhumid with winter lows usually no
lower than 12  C and summer highs usually below 38  C. Precipitation is around
1000 mm, with no dry season. The frost-free growing season is roughly 165 days
(Toensmeier and Bates 2013).
Holyoke is a “rust belt” city, a former industrial center in the western part of the
state. Most homes have lawns with some ornamental woody plants. Right across the
street from the site is a brownfield, the abandoned and contaminated location of a
442 E. Toensmeier

Fig. 17.1 The neighborhood context. The site (flagged as 145 Brown Avenue) seen in context of a
neighborhood comprised mostly of lawns with some trees. Brownfield complex across street to
right, and steep wooded sloped above railroad tracks. (Image: Google Maps)

former dry-cleaning operation (contamination risk to Paradise Lot is likely minimal

as the issue is buried tanks, and the brownfield is significantly lower elevation).
Behind the brownfield is a steep wooded slope that provides a bit of a wildlife
corridor between two densely settled neighborhoods. Little food is grown in the
neighborhood and there are no visible plantings for wildlife and minimal landscap-
ing with native plants (author personal observation) (Fig. 17.1).
The site for the garden was barren in 2004 when the project began (Fig. 17.2). The
previous house had burned down. Fill (mineral soil with extremely low organic
matter content) was brought in after construction to level the yard and parking lot.
The property occupies 400 square meters including the house and parking, with the
back yard where most of the garden is located occupying 195 square meters
(Toensmeier and Bates 2013).
Testing determined that the site features three distinct soil types. The original soil,
to the west end of the garden, is sandy and was pH 5.3 in 2004. It contained low
levels of lead. The land between this area and the house consists of highly compacted
clay with construction debris including bricks, concrete, and asphalt. The remainder
of soils were gravel and sand fill (Toensmeier and Bates 2013).
Shade was another key limitation in the design. Buildings to the south shade
much of the garden in winter, and trees overhanging from the property to the north
provide shade in the summer. Only one small area offered sun year-round, which
became the site of the greenhouse (Toensmeier and Bates 2013).
Del Tredici (2010) is a field guide to plants growing spontaneously in urban areas.
It includes notes on the ecological conditions tolerated by these species and thus
permits their use as indicator species to provide information about urban sites.
Shading the northern end of the garden from across the fenceline were two mature
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 443

Fig. 17.2 The site in 2004 before development of the garden. The Paradise Lot site was a typical
degraded urban yard with major disturbances from recent construction. (Photo: E Toensmeier)

Norway maples (Acer platanoides), which produced many seedlings in the yard
below. Del Tredici describes this species as tolerating compacted soils and road salt.
The areas of recently arrived “fill” soil were virtually free of vegetation, with a few
plants of the annual lamb’s quarters (Chenopodium album), which del Tredeci notes
is common in disturbed landscapes and tolerant of compacted soils. The original
sandy acid soil was dominated by crabgrass (Digitaria ischaemum):
“[a] disturbance-adapted colonizer of bare ground; tolerant of contaminated and
compacted soil” (Del Tredici 2010). Fencelines were tangled with vines of bitter-
sweet (Celastrus orbicularis), a noted aggressive species. Though these (mostly
reviled) species were not desired by Bates and Toensmeier, they observed that the
maples were the only place in the garden where birds congregated.

17.2.3 Design

The design process used is laid out in Jacke and Toensmeier (2005). It involves goals
clarification, site analysis and assessment, and the development of a design concept.
This concept is fleshed out into greater levels of detail (schematic and detailed
design), followed by more detailed designs for specific polycultures and infrastruc-
ture elements.
444 E. Toensmeier

Fig. 17.3 Map of the site. The initial design for the garden as shown in Jacke and Toensmeier
(2005). (Image: Dave Jacke based on work by the author)

The design matches patches with particular conditions with suitable plants spe-
cies or other design elements. For example, an area with full shade in summer and
compacted clay soil is utilized for a toolshed. These patches must also fit into the
overall design concept.
The Paradise Lot design is shown in Fig. 17.3. Along the north and west edges,
taller trees are planted to mimic a forest edge. In sunnier areas, beds for annual crops
alternate with lower-growing woody crop polycultures. Near the house, a kiwi trellis
and pond are located, along with toolshed and compost and mulch piles.

17.2.4 Evolution and Management

Overall, the project has been successful in achieving its goals, with the exception of
adoption of ecological practices by neighbors. Figure 17.4 illustrates the increase in
biomass and habitat diversity since 2004. Fruit is available outdoors daily for about
six months of the year (plus greenhouse citrus in winter), and vegetables are
available for 9–10 months outdoors and throughout the winter in the greenhouse.
Chickens (Gallus gallus), silkworms (Bombyx mori), worms (Eisenia fetida), and
native black soldier fly larvae (Hermetia illucens) have all been raised at various
times, with chickens providing eggs and the invertebrates providing compost and
food for the chickens. Lead in the sandy soil type was remediated by buffering the
pH, increasing organic matter, and mulching (in line with University recommenda-
tions), and has since fallen to safe levels according to ongoing testing.
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 445

Fig. 17.4 The site in 2019 demonstrating increased biomass and biodiversity. Paradise Lot in fall
2019. Note 10-meter tall American persimmon at right. (Photo: E Toensmeier)

While the initial goals were largely met by 2009, the herbaceous perennial layer
was poorly organized, with excessive competition from some aggressive plants. The
initial enthusiasm for crop biodiversity for its own sake was dampened by the reality
that some crops did not taste good, while others did not thrive in the site conditions
(particularly the initially degraded soils). The site goals were amended with the
following: “to grow the things we like to eat, that grow well for us, and assemble
them in functioning polycultures” (Toensmeier and Bates 2013).
Over time, the goal to develop the collection of native plants became increasingly
important. This extended to cultivating species that had been part of the Eastern
Agricultural Complex, a suite of crops first domesticated several thousand years ago
in what is today Missouri (Toensmeier and Bates 2013). As we found new resources
on native plants as providers of food and habitat (Roth 2006, Tallamy 2007, Mäder
et al. 2011, 2014), it was noted that the garden already included many of these
species as a result of research carried out by Jacke and Toensmeier (2005). At the
same time, additional species recommended in these references were incorporated
into the garden.
In 2017, co-founding designer and gardener Jonathan Bates left the site to begin a
farming operation. At this time the goals were again revised to simplify manage-
ment. These included: further culling of undesirable or unproductive species;
446 E. Toensmeier

planting more large perennial grasses for mulch production; expanding plantings for
beneficial insects and pollinators; increasing rainwater capture and storage (city
water is the main source of irrigation at moment); developing raised beds for annual
crops; and testing new approaches to pathway management (author’s personal
experience). A quantitative assessment on hours of labor required (compared to
lawn management) has not been performed but would be a useful contribution.
The initial expectation was that similar gardens would be planted throughout the
neighborhood, as neighbors were inspired by the example. This has not occurred,
perhaps because so many neighbors rent and do not have tenure agreements that
permit gardening. Toensmeier and Bates speculate that this form of extreme
“ecofunctional” gardening is perhaps too far from the norms of landscape manage-
ment, and have advocated for “suburban landscape mimics” that incorporate the
same elements but have an aesthetic more in line with American norms, even though
this involves some reduction in ecosystem function. Perhaps other tenure models
like leasing of yards might encourage adoption as well. At the same time, thousands
of gardeners from all over the world have visited the site, and many have gone on to
implement similar designs (Toensmeier and Bates 2013).
The project continues to present trade-offs. The high diversity and intensity create
a high labor demand, as does the ongoing experimentation and “editing.” Inclusion
of non-native food plants, offering high yields of desirable crops, is in dynamic
tension with the desire to maximize use of native multipurpose species. Native food
plants are often at a lower level of domestication than exotic species, exhibiting
lower yields and/or stronger flavors in some cases. For example, the native nut shrub
chinquapin (Castanea pumila) was replaced with peaches (Prunus persica), which
require more work but offer much higher and more palatable yields. Maintaining a
high level of plant biodiversity is a priority, but presents an ongoing management
challenge. Some species, both native and non-native, have become ongoing weed
issues – yet new species are continually brought to the garden.

17.3 Plant Species

Data for this site are available on plant species but only anecdotal information is
available on wildlife, so the focus here is on plants, both as biodiversity themselves
and in their roles in providing food and habitat for wildlife. Data on uses of plants are
also presented, to demonstrate that in this case habitat and human use need not be
incompatible goals.
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 447

17.3.1 Plant Biodiversity

The garden contains 59 botanical families, 130 genera, and 191 perennial or self-
sowing species. In 2004, when planting began, there were perhaps five species
present. This represents a substantial increase in botanical diversity, though not all
of it is native.
Novel ecosystems, comprised of non-historic assemblages of native and
non-native organisms, have become an area of study in recent decades (Hobbs
et al. 2013). In some cases, novel ecosystems have been shown to provide similar
and even improved ecosystem services when compared to historic ecosystems of the
same region (Davis et al. 2011, Hallet 2013). Multistrata agroforestry systems like
those practiced at Paradise Lot can be seen as anthropogenic novel ecosystems.
Table 17.1 shows the plant species present in 2019. Taxonomy, food uses, and
ecological functions are noted. Additional uses occur in the garden (e.g., medicinal
plants, bamboo for staking and trellising), but are not documented here. Almost all
species now present were brought to the site intentionally, with the exceptions of
Lactuca canadensis (spontaneous) and Aster cordifolia (present from the outset).
Two species are present but undesired, currently in the process of eradication:
Thladiantha dubia and Houttyunia cordata. More than a hundred other species
have been attempted and either failed to succeed or were eradicated based on their
poor flavor or other undesirable traits.

17.3.2 Ecological Functions

The garden was designed with five ecological functions in mind (native plant
species, nectary plants, wildlife habitat, nitrogen fixation, and groundcover), and
species have been selected accordingly. Table 17.2 shows the number and percent-
age of plants in the garden providing these ecological functions, broken down by
form (woody plants, vines, and herbaceous plants). Woody plants include trees,
shrubs, and bamboos. Vines includes woody lianas as well as herbaceous perennial
vines, and a few self-seeding annuals. Herbaceous plants are mostly perennial and
include forbs, grasses, and more, along with a few self-sowing and broadcast
annuals. The ecological functions of all species in the garden are shown in
Table 17.1. In addition to these, some annual crops are produced in outdoor beds
each year. Winter annuals and subtropical perennial crops including Citrus species
are grown in the greenhouse.
The first category examined here is native plants, which are presumed more likely
to have pre-existing relationships with native animals (Jacke and Toensmeier 2005).
Overall, 46% of the species in the garden are native to the eastern United States, and
many were collected from plants growing within a 15-min driving radius of the site.
448 E. Toensmeier

Table 17.1 Plant species present in 2019

Latin name Common name Botanical family Food uses Ecological functions
Woody Plants
Amelanchier spp. “Autumn bril- Rosaceae Fruit Native
liance”
serviceberry
Amelanchier spp. “Regent” Rosaceae Fruit Native, nectary, wild-
juneberry life cover
A. stolonifera Running Rosaceae Fruit Native, nectary, wild-
serviceberry life cover
Amorpha False indigo Fabaceae Native, nitrogen fixa-
fruticosa tion, nectary,
Asimina triloba Pawpaw Annonaceae Fruit Native
Calycanthus Carolina Calycanthaceae Spice Native, wildlife cover
floridus allspice
Castanea Chinese Fagaceae Nuts
mollisima chestnut
Cercis Redbud Fabaceae Vegetable Native
canadensis
Corylus spp. Hazelnut Corylaceae Nuts Nectary, wildlife cover
Cudrania Che fruit Moraceae Fruit
tricuspidata
Diospyros American Ebenaceae Fruit Native
virginiana persimmon
Elaeagnus Goumi Elaeagnaceae Fruit Nitrogen fixation,
multiflora nectary
Hamamelis Witch hazel Hamamelidaceae Native
virginiana
Indigofera Kirilow’s Fabaceae Nitrogen fixation, nec-
kirilowii indigo tary, wildlife cover
Lespedeza Bush clover Fabaceae Vegetable Nitrogen fixation
bicolor
Lindera benzoin Spicebush Lauraceae Spice Native
Lycium chinense Edible-leaf goji Solanaceae Fruit, wildlife cover
vegetable
Malus domestica Dwarf apple Rosaceae Fruit Nectary
Morus alba White mulberry Moraceae Fruit,
vegetable
M. alba tartarica Tartarian Moraceae Vegetable
mulberry
M. macrura Himalayan Moraceae Fruit
mulberry
Phyllostachys Running Poaceae Vegetable Wildlife cover
aureosulcata bamboo
P. Nigridolaria Running Poaceae Vegetable Wildlife cover
bamboo
P. nuda Running Poaceae Vegetable Wildlife cover
bamboo
(continued)
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 449

Table 17.1 (continued)

Latin name Common name Botanical family Food uses Ecological functions
Prunus besseyi Sand cherry Rosaceae Fruit Native, nectary, wild-
life cover
P. persica Peach Rosaceae Fruit Nectary
P. tomentosa Nanking cherry Rosaceae Fruit Nectary
Pyrus communis European pear Rosaceae Fruit Nectary
Pyrus spp. Asian pear Rosaceae Fruit Nectary
Ribes spp. Jostaberry Grossulariaceae Fruit
R. nigrum Black currant Grossulariaceae Fruit
R. odoratum Clove currant Grossulariaceae Fruit Native, wildlife cover
R. rubrum Red currant Grossulariaceae Fruit
R. uva-crispa Gooseberry Grossulariaceae Fruit Wildlife cover
Rubus idaeus Raspberry Rosaceae Fruit Native, nectary, wild-
life cover
R. occidentalis Black raspberry Rosaceae Fruit Native, nectary, wild-
life cover
Salix spp. Native willow Salicaceae Native, nectary
Sambucus Elderberry Adoxaceae Fruit, Native, nectary
canadensis vegetable
Senna hebecarpa Wild senna Fabaceae Native, nitrogen fixa-
tion, nectary, wildlife
cover
Tilia cordata Littleleaf linden Malvaceae Vegetable
Toona sinensis Chinese toon Meliaceae Vegetable
Vaccinium Halfhigh Ericaceae Fruit Native
corymbosum blueberries
Viburnum opulus Highbush Adoxaceae Fruit Native, nectary
cranberry
x Sorbopyrus Shipova Rosaceae Fruit
Zizyphus jujuba Jujube Rhamnaceae Fruit
Vines
Actinidia arguta Hardy kiwifruit Actinidiaceae Fruit
Amphicarpa Ground bean Fabaceae Beans, Groundcover, native
bracteata tubers
Apios americana Groundnut Fabaceae Tubers Native, nectary, nitro-
gen fixation
A. priceana Price’s Fabaceae Beans, Native, nectary, nitro-
groundnut tubers gen fixation
Dioscorea Chinese yam Dioscoreaceae Bulbils,
polystachya tubers
Hablitzia Caucasian Amaranthaceae Vegetable
tamnoides spinach
Humulus lupulus Hops Cannabidaceae Vegetable,
tea
Melothria Cucumber Cucurbitaceae Vegetable Groundcover, native
pendula berry
(continued)
450 E. Toensmeier

Table 17.1 (continued)

Latin name Common name Botanical family Food uses Ecological functions
Passiflora Maypop Passifloraceae Fruit, Native, nectary
incarnata vegetable
Phaseolus Thicket bean Fabaceae Beans Native, nectary, nitro-
polystachyos gen fixation
Strophostyles Woolly bean Fabaceae Beans Native, nectary, nitro-
helvola gen fixation
S. umbellata Perennial Fabaceae Beans Native, nectary, nitro-
woolly bean gen fixation
Thladiantha Manchu Cucurbitaceae Fruit, Nectary
dubia tubergourd vegetable
Vitis riparia Riverbank Vitaceae Fruit, Native
grape vegetable
V. vinifera x Grape Vitaceae Fruit, Native
labrusca vegetable
Herbs
Agastache Anise hyssop Lamiaceae Tea Native, nectary
foeniculum
Allium Perennial leek Alliaceae Culinary, Nectary
ampeloprasum vegetable
A. canadense Wild garlic Alliaceae Culinary, Native
vegetable
A. cepa Walking onion Alliaceae Culinary,
proliferum vegetable
A. cernuum Nodding wild Alliaceae Culinary, Native, nectary
onion vegetable
A. fistulosum Welsh onion Alliaceae Culinary, Nectary
vegetable
A. tricoccum Ramps Alliaceae Culinary, Native
vegetable
A. tuberosum Garlic chives Alliaceae Culinary, Groundcover, nectary
vegetable
A. ursinum Ramsons Alliaceae Culinary,
vegetable
A. victorialis Victory onion Alliaceae Culinary,
vegetable
A. vineale Field garlic Alliaceae Culinary,
vegetable
Amorpha nana Dwarf false Fabaceae Native, nectary, nitro-
indigo gen fixation
Anthriscus Woodland Apiaceae Culinary, Nectary
sylvestris chervil vegetable
Aquilegia Columbine Ranunculaceae Vegetable Native
canadensis
Artemisia annuua Sweet Annie Asteraceae
Asarum Wild ginger Aristolochiaceae Groundcover, native
canadense
(continued)
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 451

Table 17.1 (continued)

Latin name Common name Botanical family Food uses Ecological functions
Asclepias Poke milkweed Apocynaceae Vegetable Native, nectary
exaltata
A. incarnata Swamp Apocynaceae Vegetable Native, nectary
milkweed
A. speciosa Showy Apocynaceae Vegetable Native, nectary
milkweed
A. syriaca Common Apocynaceae Vegetable Native, nectary
milkweed
A. tuberosa Butterfly weed Apocynaceae Vegetable Native, nectary
Asparagus Asparagus Asparagaceae Vegetable
officinalis
Aster cordifolia Heartleaf aster Asteraceae Native, nectary
A. novae-angliae New England Asteraceae Native, nectary
aster
A. scaber Chwinamul Asteraceae Vegetable
Astragalus Licorice Fabaceae Tea Groundcover, nitrogen
glycyphyllos milkvetch fixation
Atriplex hortensis Orach Amaranthaceae Vegetable
Brassica napus Kale Brassicaceae Vegetable
B. rapa Mustard Brassicaceae Vegetable
Bunias orientalis Turkish rocket Brassicaceae Vegetable Nectary
Bunium Earth chestnut Apiaceae Vegetable, Nectary
bulbocastanum root
Camassia Camass Asparagaceae Root
quamash
Camassia Wild hyacinth Asparagaceae Root Native
scillioides
Campanula Bluebells Campanulaceae Vegetable Groundcover
porschariskayana
Campanula Rampion Campanulaceae Root
rapunculus
Cardamine Toothwort Brassicaceae Culinary Native
diphylla
Ceanothus New Jersey tea Rhamnaceae Tea Native, nectary, nitro-
americanus gen fixation
Chamaenerion Fireweed Onagraceae Vegetable Native, nectary
angustifolium
Chasmanthium River oats Poaceae Native, nectary, wild-
latifolium life cover
Chenopodium Good king Amaranthaceae Vegetable
bonus-henricus Henry
Chrysogonum Green and gold Asteraceae Groundcover, native,
virginianum nectary
Crambe maritima Sea kale Brassicaceae Vegetable Nectary
Crocus sativus Saffron crocus Iridaceae Culinary
(continued)
452 E. Toensmeier

Table 17.1 (continued)

Latin name Common name Botanical family Food uses Ecological functions
Cryptotaenia Honewort Apiaceae Vegetable Native, nectary
canadensis
Dennestedia Hay-scented Dennestediaceae Groundcover, native
punctilobula fern
Desmodium Tick trefoil Fabaceae Native, nitrogen
canadense fixation
D. glutinosum Pointed-leaved Fabaceae Groundcover, native,
tick trefoil nitrogen fixation
Diplotaxis Sylvetta arugula Brassicaceae Vegetable Nectary
muralis
Disporum Chinese fairy Liliaceae Vegetable
cantonense bells
Dryopteris Spiny wood Dryopteridaceae Roots
expansa fern
Dystaenia Korean celery Apiaceae Vegetable Nectary
takesimiana
Eurybia Largeleaf aster Asteraceae Vegetable Groundcover, native,
macrophylla nectary
Festuca ovina sheep’s fescue Poaceae Groundcover
Fragaria Strawberry Rosaceae Fruit Groundcover
anasana
F. moschata Musk Rosaceae Fruit Groundcover
strawberry
Fragaria vesca Wood Rosaceae Fruit Groundcover, native
strawberry
Gaultheria Wintergreen Ericaceae Fruit, tea Groundcover, native
procumbens
Helianthus Sunchoke Asteraceae Root Native, nectary, wild-
tuberosus life cover
Hibiscus Swamp Malvaceae Vegetable Native, nectary
moscheotus hibiscus
Hosta spp. Edible hosta Asparagaceae Vegetable Groundcover
Houyttunia Houttyunia Saururaceae Vegetable Groundcover
cordata
Hydrophyllum Waterleaf Boraginaceae Vegetable Groundcover, native,
spp. nectary
Impatiens Touch me not Balsaminaceae Native, nectary
capensis
Kalimeris indica Yomena aster Asteraceae Vegetable Nectary
Lactuca Wild lettuce Asteraceae Vegetable Native, nectary
canadensis
Ligularia fischeri Gomchee Asteraceae Vegetable Nectary
Lilium superbum Turk’s cap lily Liliaceae Root Native
Lobelia Cardinal flower Lobeliaceae Native, nectary
cardinalis
(continued)
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 453

Table 17.1 (continued)

Latin name Common name Botanical family Food uses Ecological functions
L. siphilitica Blue lobelia Lobeliaceae Native, nectary
Matteuccia Ostrich fern Onocleaceae Vegetable Native
struthiopteris
Melissa Lemon balm Lamiaceae Culinary, Nectary
officinalis tea
Mentha piperita Peppermint Lamiaceae Culinary, Nectary
tea
Mentha spp. Apple mint Lamiaceae Culinary, Nectary
tea
Mertensia Oysterleaf Boraginaceae Vegetable Native, nectary
virginiana
Miscanthus Giant Poaceae Vegetable Wildlife cover
giganteus miscanthus
Monarda Bergamot Lamiaceae Culinary Native, nectary
fistulosa
Musa basjoo Japanese fiber Musaceae Vegetable
banana
Muscari Italian grape Asparagaceae Vegetable
botryoides hyacinth
M. neglectum Musk hyacinth Asparagaceae Vegetable,
root
Myrrhis odorata Sweet cicely Apiaceae Vegetable Nectary
Nelumbo nucifera Chinese lotus Root, nut
Oenanthe Water celery Apiaceae Vegetable Groundcover, nectary
javanica
Osmunda Cinnamon fern Osmundaceae Vegetable Native
cinnamomea
O. japonica Japanese royal Osmundaceae Vegetable
fern
Pachysandra Allegheny Buxaceae Groundcover, native
procumbens spurge
Phacelia Purple phacelia Boraginaceae Groundcover, native,
bipinnatifida nectary
Physalis Clammy Solanaceae Fruit Groundcover, native
heterophylla groundcherry
P. longifolia Longleaf Solanaceae Fruit Groundcover, native
groundcherry
Phyalis spp. Perennial Solanaceae Fruit Groundcover
groundcherry
Pleioblastus Dwarf fernleaf Poaceae Groundcover
distichus bamboo
Podophyllum Mayapple Berberidaceae Fruit Native
peltatum
Polygonatum Giant Solo- Asparagaceae Vegetable, Native
commutatum mon’s seal roots
Mountain mint Lamiaceae Tea Native, nectary
(continued)
454 E. Toensmeier

Table 17.1 (continued)

Latin name Common name Botanical family Food uses Ecological functions
Pycnanthemum
hyssopifolium
P. muticum Mountain mint Lamiaceae Tea Native, nectary
Pycnanthemum Mountain mint Lamiaceae Tea Native, nectary
spp
Rheum palmatum Turkish rhubarb Polygonaceae Vegetable
R. x cultorum Rhubarb Polygonaceae Vegetable
Rudebeckia Sochan Asteraceae Vegetable Native, nectary
laciniata
Rumex acetosa Sorrel Polygonaceae Vegetable
Sagittaria Arrowhead Alismataceae Vegetable, Native
latifolia root
Scilla siberica Siberian squill Asparagaceae Vegetable
Sedum ternatum Woodland Crassulaceae Vegetable Groundcover, native,
stonecrop nectary
Sium suave Water parsnip Apiaceae Vegetable, Native, nectary
root
Solanum Black Solanaceae Fruit, Native
ptycanthum nightshade vegetable
Solidago spp. Goldenrod Asteraceae Native, nectary
Stachys affinis Chinese Lamiaceae Root Groundcover
artichoke
Symphytum Large-flowered Boraginaceae Groundcover, nectary
grandiflorum comfrey
Symphytum spp. “Hidcote blue” Boraginaceae Groundcover, nectary
comfrey
S. x uplandicum Russian Boraginaceae Nectary
comfrey
Trifolium repens White clover Fabaceae Vegetable Groundcover, nectary,
nitrogen fixation
Tulipa clusiana Clusiana tulip Liliaceae Root
Tulipa spp. “Purissima” Liliaceae Vegetable
hybrid tulip
Typha spp Cattail Typhaceae Vegetable, Native
root
Vaccinium Lowbush Ericaceae Fruit Native
angustifolium blueberry
Verbena hastata Blue vervain Verbenaceae Seeds, tea Native, nectary
Viola sororia Common blue Violaceae Vegetable Groundcover, native
violet
Waldsteinia Barren Rosaceae Groundcover, native
fragaroides strawberry
Zingiber mioga Mioga ginger Zingiberaceae Vegetable
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 455

Table 17.2 Plant species performing ecological functions

% of # of % of
# of Woody Woody # of % of Herbaceous Herbaceous
Function plants plants Vines Vines plants plants
Native plant 20 45% 10 67% 58 44%
species
Nectary 19 43% 7 47% 54 41%
plants
Wildlife 17 39% 0 0% 2 2%
cover
Nitrogen 5 11% 6 40% 6 5%
fixation
Groundcover 0 0% 2 13% 31 23%

Plants with flowers attractive to pollinators and beneficial insects were also
prioritized, and a flowering calendar was created to ensure that pollen and nectar
are available throughout the growing season. Species were selected using listings
from sources including Jacke and Toensmeier (2005) and Mäder et al. (2011 and
2014). Nectary species of this kind constitute 42% of those grown in the garden.
Wildlife cover is defined here as being provided by vegetation that is at least
100 cm tall and either evergreen, forming multistemmed thickets, or both (Jacke and
Toensmeier 2005). Wildlife cover is provided by 10% of the species in the garden,
predominantly woody plants.
Nitrogen fixation is an important tool for restoring degraded sites and maintaining
fertility (Jacke and Toensmeier 2005). Though only 9% of species in the garden fix
nitrogen, they are densely planted to address nitrogen requirements.
Groundcovers provide a living mulch to suppress weed germination and maintain
optimal conditions for healthy soil (Jacke and Toensmeier 2005). This function is
provided by 17% of the species in the garden.

17.3.3 Human Uses

Food production has been the primary human use for which plants have been
selected at Paradise Lot. Table 17.3 shows the number and percentage of plants
providing edible parts. These include: fruits (excluding those used as vegetables);
vegetables including leaves, shoots, flowers, and fruits used as vegetables
(Toensmeier et al. 2020); roots and tubers including bulbs, corms and rhizomes;
nuts, seeds and dry beans; and culinary and spice plants and teas. The edible uses of
all species in the garden are shown in Table 17.3. There are many non-edible uses for
plants in the garden, such as medicinal plants and bamboo for garden stakes and
trellises, but these are not analyzed here.
456 E. Toensmeier

Table 17.3 Number of plant species providing human uses

% of # of % of
# of Woody Woody # of % of Herbaceous Herbaceous
Use plants plants Vines Vines plants plants
Fruit 29 66% 4 27% 10 8%
Vegetable 7 16% 6 40% 65 49%
Roots & 0 0% 4 27% 16 12%
tubers
Nuts, seeds 2 5% 5 33% 1 1%
& beans
Culinary, 2 5% 0 0% 31 23%
spice & tea

Eighty-seven percent of the 191 plant species in the garden are edible. This
includes 23% with edible fruit, 41% used as vegetables, 10% with edible roots and
tubers, 4% with edible nuts, seeds, or beans, and 17% with culinary, spice, or tea
uses. Some of these species produce more than one edible category, for example
Lycium chinese has both edible leaves and fruit.

17.4 Anecdotal Observations on Animal Biodiversity

Several elements were designed into the garden to improve habitat for animals,
primarily insects and birds. Access to water is provided via an artificial pond. Water
is an important habitat element for arthropods and vertebrates (Jacke and Toensmeier
2005). Virtually no pesticides are used at all. The exception is kaolin clay powder,
which is non-toxic and an important control for plum curculio (Conotrachelus
nenuphar), which otherwise devastates pome and stone fruits (author personal
experience). The garden is also designed with an irregular texture, with open
areas, thickets, and forest-like areas to maximize habitat diversity. This “lumpy
texture” has been shown to minimize pest damage by providing niches for many
birds and other predators (Jacke and Toensmeier 2005).
Formal data have not been collected on animal diversity. However, some anec-
dotal observations will be shared here.

17.4.1 Arthropods

Invertebrate density and diversity were low in 2004 and began to increase dramat-
ically in 2005. Dead logs, leaf and wood chip mulch, and flowers were observed to
be strong attractants of invertebrates. As of 2019, large and diverse populations of
arthropods have been observed in the garden. Unlike birds and most other verte-
brates, a small garden of this size appears to be able to provide sufficient habitat for
many insects to live their entire life there. For example, Chinese mantis (Tenodera
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 457

aridifolia) has been present in the garden for many years. Each year over a dozen
oothecae (egg cases) are laid in the garden. The nymphs emerge in spring and appear
to live their entire lives in the garden, which would seem to be large enough to
provide all the space needed for multiple adult mantises.

17.4.2 Birds

Initially the great majority of birds in the garden were urban species like starlings
(Sturnus vulgaris) and house sparrows (Passer domesticus). As habitat (larger
woody plants and the installation of the pond) and food availability (insects, seeds
and fruit) increased over several years, bird diversity and abundance increased.
Migrating birds now often stop to visit the garden including Baltimore oriole (Icterus
galbula), multiple species of warbler (Parulidae), and wood thrush (Hylocicha
mustelina). The presence of the wood thrush is notable as it is a forest understory
species (Jacke and Toensmeier 2005). The garden alone would appear to be too
small to provide sufficient territory for birds, though a flicker (Colaptes auratus)
nested in a neighboring tree and fed in the garden daily one summer.

17.4.3 Mammals

Mammals in the garden represent typical small to mid-sized urban residents. Grey
squirrels (Sciurus carolinensis) are year-round residents. Frequent visitors, often
only visible at night, are skunks (Mephitis mephitis) and opossums (Didelphis
virginiana). Occasional visitors include red fox (Vulpes vulpes), star-nosed mole
(Condylura cristata), and raccoon (Procyon lotor). The very high populations of
neighborhood cats (Felis catus), which hunt in the garden severely limit populations
of smaller rodents. Bats, likely Eptesicus fuscus, can be seen hunting at night over
the garden in summer.

17.4.4 Herptiles

Though introduction of tadpoles of the native American toad (Anaxyrus americanus)

was a failure due to predation by cats, a lone wood frog (Rana sylvatica) has come to
the pond in spring several years to unsuccessfully call for a mate. Several sightings
of the red-backed salamander (Plethodon cinereus) may be the same individual or
represent a larger population. It may have arrived in nursery pots. A DeKay’s
brownsnake (Storeria dekayi), a small slug-eating snake, was greeted enthusiasti-
cally by the gardeners but it was not seen again and may well have been consumed
by cats.
458 E. Toensmeier

17.5 Conclusions

This case study demonstrates that food production and ecosystem function can go
hand in hand, with the wide variety of plants incorporated here providing various
ecological roles and human uses. While the scale of the garden is too small to
provide significant habitat for vertebrate wildlife, it is quite successful at serving the
needs of arthropods and offers some benefits to neighborhood and migrating birds
and mammals.
Researchers are invited to use the site to quantify the impacts on invertebrates and
other wildlife, as only anecdotes are possible here. Research to compare this
temperate site to tropical homegardens is also suggested. Its embracing of the
novel ecosystem concept is controversial and also worth of study.
Development of similar, perhaps simpler models on public land in cities could
increase biodiversity impacts. Demonstration and instruction in development of
streamlined systems, for example modular systems for annual crops, small fruits
and tree fruits, pollinator plantings, could increase adoption to the levels necessary to
reintroduce some ecosystem services and habitat into biologically barren neighbor-
hoods. Should this model be more widely adopted, even in simplified form, it could
potentially offer benefits to people while providing important islands, and potentially
corridors, of biodiversity in urban areas.

References

Clark KH, Nicholas KA (2013) Introducing urban food forestry: a multifunctional approach to
increase food security and provide ecosystem services. Landsc Ecol 28:1649–1669
Davis MA, Chew MK, Hobbs RJ, Lugo AE, Ewel JJ, Vermeij GJ, Brown KH (2011) Don’t judge
species on their origins. Nature 474:153–154
Del Tredici P (2010) Wild urban plants of the Northeast: a field guide. Comstock Publishing, Ithaca
Ferguson RS, Lovell ST (2013) Permaculture for agroecology: design, movement, practice, and
worldview. A review. Agron Sustain Dev 34:251–274
Ferguson RS, Lovell ST (2014) Permaculture for agroecology: design, movement, practice, and
worldview. Agron Sustain Dev 34:251–274.
Hallet LM (2013) Towards a conceptual framework for novel ecosystems. In: Hobbs RJ, Higgs ES,
Hall CM (eds) Novel ecosystems: intervening in the new world order. John Wiley, Oxford, pp
16–28
Hobbs RJ, Higgs ES, Hall CM (2013) Introduction: why novel ecosystems? In: Hobbs RJ, Higgs
ES, Hall CM (eds) Novel ecosystems: intervening in the new world order. John Wiley, Oxford,
pp 1–8
Jacke D, Toensmeier E (2005) Edible forest gardens. Chelsea Green Publishing, White River
Junction
Mäder E, Shepherd M, Vaughn M, Black SH, LeBuhn G (2011) The xerces society guide to
attracting native pollinators: protecting North America’s bees and butterflies. Storey Publishing,
North Adams
Mäder E, Hopwood J, Morandin L, Vaughn M, Black SH (2014) The xerces society guide to
farming with native beneficial insects: ecological pest control solutions. Storey Publishing,
North Adams
17 Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity Island 459

Nair PK (2006) Whither homegardens? In: Nair PK, Kumar BM (eds) Tropical homegardens.
Springer, Dordrecht
Nair PK, Kumar BM (2006) Introduction. In: Nair PK, Kumar BM (eds) Tropical homegardens.
Springer, Dordrecht
Roth S (2006) Bird-by-bird gardening: the ultimate guide to bringing in your favorite birds, year
after year. Rodale, Emmaus
Tallamy DW (2007) Bringing nature home: how you can sustain wildlife with native plants. Timber
Press, Portland
Toensmeier E (2016) The carbon farming solution: a global toolkit of perennial crops and
regenerative agriculture practices for climate change mitigation and food security. Chelsea
Green Publishing, White River Junction
Toensmeier E, Bates J (2013) Paradise lot: two plant geeks, one-tenth of an acre, and the making of
an edible garden oasis in the city. Chelsea Green Publishing, White River Junction
Toensmeier E, Ferguson R, Mehra M (2020) Perennial vegetables: a neglected resource for
biodiversity, carbon sequestration, and nutrition. PLOS One, in press
Chapter 18
Contribution to the Domestication
and Conservation of the Genetic Diversity
of Two Native Multipurpose Species
in the Yabotí Biosphere Reserve, Misiones,
Argentina

Fernando Niella, Patricia Rocha, Ariel M. Tuzinkievicz, Ricardo Buchweis,

Christian Bulman Hartkopf, Peggy Thalmayr, José González,
Florencia Montagnini, and Sandra Sharry

Abstract In this chapter we describe a strategy for domestication and conservation

of the genetic variability of two native species of the Interior Atlantic Forest, or Selva
Paranaense, Peltophorum dubium and Enterolobium contortosiliquum. Both species
are leguminous trees, commonly used for forest restoration projects. In tropical and
subtropical forests, tree species are vulnerable to habitat fragmentation and popula-
tion reductions. The resulting negative genetic effects, such as loss of genetic
variability and inbreeding depression, can affect the long-term survival of forest
species, leading to their further vulnerability or extinction. While in situ strategies
such as protected areas and biodiversity islands may be an option for biodiversity
conservation, in practice there are many challenges to these strategies in humid

F. Niella (*) · P. Rocha · A. M. Tuzinkievicz · R. Buchweis · C. Bulman Hartkopf

School of Forest Science (FCF), Misiones National University (UNaM), Eldorado,
Misiones, Argentina
e-mail: lpv@[Link]
P. Thalmayr
IBS-CONICET, Iguazú, Misiones, Argentina
J. González
Puerto Laharrague S. A, Montecarlo, Misiones, Argentina
e-mail: jgonzalez@[Link]
F. Montagnini
School of the Environment, The Forest School, Yale University, New Haven, CT, USA
e-mail: fl[Link]@[Link]
S. Sharry
School of Agriculture and Forest Science, La Plata National University (UNLP),
La Plata, Buenos Aires, Argentina
National University of Rio Negro, Viedma, Argentina

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 461
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
462 F. Niella et al.

subtropical ecosystems, where there is high tree species diversity and low abundance
per species (0.1–1 trees/ha). Trees in remnants of forests, which can serve as
biodiversity islands, must have high genetic diversity so that they can persist through
time by adapting to disturbances. There is a great need to expand the genetic basis of
collection of species’ propagation material, so that the germplasm available for
restoration programs includes the largest possible genetic diversity. To address this
need, we established short- and long-term provenance and progeny trials and a
vegetative reproduction methodology to produce propagation material to ensure
the genetic diversity of these two species for domestication, including for restoration
and enrichment. Our results lay the foundation for the conservation of genetic
variability of P. dubium and E. contortosiliquum and contribute to the design of a
possible biodiversity island strategy for these species.

Keywords Conservation strategy · Forest restoration · Genetic variability · Interior

Atlantic forests · Native tree species · Selva paranaense · Vegetative propagation
techniques

18.1 Introduction

The Yabotí Biosphere Reserve, with a total of 235,900 hectares, is located in the
province of Misiones, Argentina and is part of the Atlantic Forest. The Atlantic
Forest extends over the Atlantic coast of Brazil and is one of the most diverse
ecosystems in the world, with about 15,000 species of vascular plants, of which at
least 48% are endemic (Myers et al. 2000; Martini et al. 2007; Murray-Smith et al.
2009). The Atlantic Forest extends into the interior of Brazil and into the province of
Misiones in the northeast of Argentina and eastern Paraguay, where it is known as
Interior Atlantic Forest or Selva Paranaense (Paranaense forest) or Selva Misionera.
The Interior Atlantic Forests, together with the northwest montane forest of the
country known as the Yungas (mostly in the provinces of Salta, Jujuy and Tucumán),
represent the most biodiverse regions of Argentina. The Atlantic Forest was included
as one of the original biodiversity hotspots or critical zones in the widely cited
publication of Ernst Mayr in 1988, along with 10 other regions of the globe, which
have now been extended to 36 regions worldwide. Designation as a hotspot region
should meet two strict criteria: a) it must contain at least 1500 endemic species of
vascular plants (0.5% of the total vascular plants identified on Earth); and b) it must
have experienced habitat loss of at least 70% of its original area. More than 50% of
the world’s plant species are endemic to the 36 biodiversity critical areas mentioned
above and once covered 15.7% of the Earth’s surface. Now, they are reduced to
2.3% of the Earth’s surface and include many endemic species, which face a growing
threat of extinction (Reed et al. 2011).
The growing population, the advance of the livestock agricultural frontier, and the
extraction of timber trees have been the most frequent causes of the drastic decrease
in surface area and consequent vulnerability of the 36 critical areas or hotspots.
Given this situation, and as part of the commitments made on a global scale at the
18 Contribution to the Domestication and Conservation of the. . . 463

biodiversity convention in 1992 in Rio de Janeiro, the Global Strategy for Plant
Conservation (GSPC 2010) agreement was signed in 2002 by more than 180 coun-
tries, with the aim of developing actionable policies to support plant conservation.
GSPC Objective 8 requires that “at least 75% of threatened plant species are in ex
situ collections and at least 20% available for recovery and restoration programs”, as
outlined by the United Nations Development Program (Reed et al. 2011).
The recognition of islands of biodiversity provides an important step for restora-
tion and can contribute to reversing fragmentation and increasing biodiversity
around the world (Montagnini et al. 2022). Several authors, such as Ratnam et al.
(2014) and Thomas et al. (2014), have suggested strategies to increase resilience in
forest restoration initiatives. Such measures include increasing tree population size
and genetic diversity, maintaining forest cover in the landscape for genetic and
geographical connectivity between tree populations, promoting genetic improve-
ment of tree species, and enhancing the identification and protection of evolutionary
shelters (Bhagwat et al. 2012; Pauls et al. 2013).
In tropical and subtropical forests, tree species are vulnerable to habitat fragmen-
tation and population reductions. The resulting negative genetic effects, such as loss
of genetic variability and inbreeding depression, affect the long-term survival of
forest species, leading to their further vulnerability or extinction (Maina and Howe
2000). While in situ strategies such as protected areas and biodiversity islands may
be an option for biodiversity conservation, in practice there are many challenges with
these strategies in humid subtropical ecosystems, where there is high tree species
diversity and low abundance (0.1–1 trees/ha of the same species).
In studies comparing fruits of the species Enterolobium cyclocarpum obtained
from trees located in continuous forests and from isolated trees from pasture grazing
areas, Rocha and Aguilar (2001) infer that habitat fragmentation alters the mecha-
nisms through which plants regulate the quality of their offspring by increasing self-
pollination rates in isolated trees and producing less fruit, which are then usually
abortive. For this reason, isolated trees produce progeny that is less vigorous than the
progeny of mother trees in continuous forests, which is another adverse effect of
forest fragmentation. Therefore, intraspecific genetic variability is directly related to
the resilience of the ecosystem and its ability to cope with abrupt changes in the
climate and its habitat to ensure its long-term survival.

18.1.1 Importance of Genetic Diversity in Restoration

and Conservation Programs

Genetic diversity is the basis of an organism’s ability to adapt to changes in its

environment through natural selection. Populations with little genetic variation are
more vulnerable to the emergence of new pests or diseases, pollution, climate
change, and habitat destruction due to human activities or other catastrophic events.
The inability to adapt to changing conditions greatly increases the risk of extinction.
464 F. Niella et al.

Reed and Frankham (2003) found that the correlation between genetic diversity at
the population level and the adaptability of the population to changing environmen-
tal conditions was highly significant, concluding that the loss of heterozygosity1 has
a strong effect on the adaptability of the population. This is consistent with the
provisions of the International Union of Conservation of Nature (IUCN) on the
status of importance of the conservation of genetic diversity (McNeely et al. 1990).
Genetic diversity is one of the three major components of biodiversity, but is still
overlooked in most plans for conserving biodiversity. Therefore, the implementation
of genetic criteria into the Red List assessment process will help to define more
precisely the conservation status of the species (Bruford and Segelbacher 2018).
Meanwhile, there is also a commitment from the international community to
restore hundreds of millions of hectares of degraded forest landscapes, following the
Strategic Plan for Biodiversity 2011–2020, including the Aichi Biodiversity Targets
(CBD 2018). However, the successes and failures of past restoration efforts remain
in many cases undocumented and uncommunicated. Case studies show that failures
may have been much more common than successes (Wuethrich 2007; Godefroid
et al. 2011). The causes of restoration failures can be multiple. One reason, often
ignored, is the inadequate consideration of the source and genetic quality of forest
reproductive material (Godefroid et al. 2011; Le et al. 2012).
The impact of logging on the structure of forest species populations depends to a
large extent on the degree of disruption and intensity of logging. The threat to
genetic diversity posed by commercial logging correlates with the abundance of a
species in each forest management unit. Natural regeneration, while allowing the
transmission of genetic information to the next generation, does not guarantee
adaptive and non-adaptive change of structures during the regeneration phase
(Rajora and Pluhar 2003). When the number of seed trees left in the logged forest
is low and dysgenic selection is heavily practiced according to the market demands,
the tree phenotype left in the remaining population is skewed, leading to a gradual
process of genetic erosion of the entire population. Without genetic diversity,
evolution is impossible, and adaptation decreases, which can result in local extinc-
tions. Processes such as natural selection, genetic drift,2 and genetic flow3 collec-
tively affect the genetic diversity of populations and either promote or hinder local
and wide-ranging adaptation.
Particularly in tropical and subtropical regions, the genetic diversity of tree
species is a key component of the functioning of forest ecosystems (Ratnam et al.
2014). In tropical species, such as for example Dipteryx odorata (hermaphroditic
and pollinated by insects), and Bagassa guianensis (dioecious and mainly wind-
pollinated), studies of the impact of forest management have shown that selective

1
The presence of different alleles at one or more loci are not the same (Acquaah 2012).
2
The random fluctuations of gene frequencies in a population such that the genes amongst offspring
are not a perfectly representative sampling of the parental genes (Schlegel 2010).
3
Introduction of genetic material from one population of species to another, thereby, changing the
composition of the gene pool of the receiving population (White et al. 2007).
18 Contribution to the Domestication and Conservation of the. . . 465

logging induced an asynchrony in flowering, limited the flow of genes, and induced
inbreeding, even in species that were cross-pollinated. In this manner, the regener-
ation of their offspring is affected with serious consequences, especially for tree
species managed by selective logging. This is especially true in situations where the
remaining forests have only few trees of reproductive age and lack pollinators, which
contribute to the flow of genes (Ratnam et al. 2014).

18.1.2 Genetic Considerations for Restoration Programs

for the Interior Atlantic Forest

The high intensity of some methods of logging, as mentioned above, can modify
breeding patterns in residual trees and result in increasingly inbred seeds through
self-fertilization or crossbreeding between closely related individuals (biparental
inbreeding), compromising the role of a tree population as a source of seeds. The
risk of inbreeding must be seriously considered in activities dealing with genetic
resources, use of germplasm in practical forestry and tree improvement. In such
cases, for restoration and/or enrichment programs, the use of germplasm obtained
from similar ecological conditions, even if they are not from local sources, may be a
better option than resorting to fragmented nearby forests or isolated trees (Navarro
Pereyra 2002; Thomas et al. 2014).
Various general guidelines for seed collection that ensure a minimum of genetic
diversity for restoration programs have been published, e.g., works by The
Australian Network for Plant Conservation Inc. (Vallee et al. 2004), the University
of California (Rogers and Montalvo 2004), the World Agroforestry Centre (ICRAF)
(Kindt et al. 2006), and the Royal Botanic Gardens, Kew (2003), among others. It is
essential that the collection of germplasm captures a representative sample of the
genetic diversity of species to be used in restoration projects in a way that takes into
consideration the natural distribution of the species as much as possible. A set of
general guidelines for tree seed harvesting are intended to ensure a minimum level of
genetic diversity. For example, as a general rule, a minimum of 30 randomly selected
trees must be sampled for a completely cross-pollinated species (Rogers and
Montalvo 2004).
In the province of Misiones, Argentina, the extraction of wood from forests and
the harvesting and planting of yerba mate (Ilex paraguariensis A. St -Hil) began at
the end of the nineteenth century. Forestry activities (mostly plantations of fast-
growing exotic species such as Pinus spp) were consolidated by the end of the
twentieth century (Ferrero 2005), along with plantations of tea, tobacco, and yerba
mate monocultures, and more recently livestock production. In the area of study
(Yabotí Biosphere Reserve) historically the selective cutting of timber species was
done using low impact management practices, such as the Minimum Cutting Diam-
eters (MCDs) that were established by provincial legislation.
466 F. Niella et al.

The selective cutting method following the MCD requirement for forest
harvesting is widely used in tropical and subtropical ecosystems. However, in
many cases, this type of management is suited to industrial processing technologies
and market demands, while not considering the reproductive biology of the individ-
ual trees, and the appropriate number of remaining seedlings, among other factors, to
ensure the persistence in perpetuity of the harvested species. Therefore, a manage-
ment program focused exclusively on the MCD does not result in ecologically
sustainable forest management in the long term (Montagnini et al. 1998; Putz et al.
2000; Jennings et al. 2001; Sist et al. 2003).
Among the species that are vulnerable and/or threatened by excessive habitat
fragmentation and extraction in the Interior Atlantic Forest, Peltophorum dubium
(Spreng.) Taub and Enterolobium contortosiliquum (Vell.) Morong, are two impor-
tant species that have been long used in restoration projects. These species are both
hermaphrodites and pollinated by insects. Mori et al. (2013), who studied the
reproductive system in natural populations of P. dubium, observed that the crossings
were not random, and that the species was not self-incompatible, but presented a
typical system of mixed mating, combining crosses with self-fertilization. Inbreed-
ing was also detected in both the parental generation and in its offspring, although
they observed evidence for natural selection between the juvenile and adult phase
against inbred individuals.
There is a great need to expand the genetic basis of species propagation material,
so that the germplasm available for restoration programs includes the largest possible
genetic diversity. In this chapter, we propose a strategy for domestication and
conservation of the genetic variability of two native species, Peltophorum dubium
and Entorolobium contortosiliquum, to provide germplasm for enrichment and/or
restoration projects of native forests. Our methods could also be used for other
species, to produce propagation material with genetic diversity for domestication,
enrichment, and/or restoration programs. This work in progress is expected to
contribute to a strategy for establishing or enriching biodiversity islands.

18.2 Case Study: Provenance and Progeny Trials

for the Domestication and Conservation of Genetic
Diversity of Peltophorum dubium (Caña Fístola)
and Enterolobium contortosiliquum (Timbó)
in the Yabotí Biosphere Reserve, Misiones, Argentina

The species, provenance, and progeny trials aim to ascertain genetic variation at
different levels: between species, between geographical regions, between stands and
between individuals, and they are the first logical step of genetic improvement for
any species (White et al. 2007). Therefore, as a case study, we proposed to assess and
evaluate the genetic resources of two native species of the Interior Atlantic Forests,
P. dubium and E. contortosiliquum, as a source of germplasm for enrichment and/or
18 Contribution to the Domestication and Conservation of the. . . 467

restoration projects of native forests. The species, P. dubium and

E. contortosiliquum, both belonging to the legume family, are of great interest to
the local forest industry due to the quality of their wood, which has led to high levels
of extraction and in turn, genetic erosion. Currently, there is also an increasing
interest in planting them in small- and medium-sized farms, in compliance with
provincial regulations. These species can facilitate the production of honey, have
good properties for restoration of degraded areas, can be used in agroforestry or
silvicultural consociated systems, and have dendro-energy potential. Local towns
have high demand for these species for use as ornamentals in landscaping projects in
urban areas, since they are representative of the Misiones Forest (Eibl et al. 1998;
Barth et al. 2008; Alexandre et al. 2009; Eibl et al. 2017). As seen, local communities
consider both as multipurpose species.
Initial field trials to assess their behavior under different silvicultural conditions,
have identified P. dubium as a heliophyte species, with rapid initial growth and good
behavior in open monoculture plantations. In the west Chaco, field trials have shown
that P. dubium can reach 45 cm in diameter at 20 years of age (Gómez and Cardozo
2003). In Misiones, plantation trials of E. contortisiliquum have shown that this
species can reach 25 cm in diameter at 14 years of age (Montagnini et al. 2006),
demonstrating fast growth even when genetic material with little or no domestication
was used. Similar results have been obtained in forest enrichment experiments
carried out under canopy of native forest San Pedro, Misiones (Eibl et al. 1993)
and in San Ignacio in plantations under the canopy of Pinus spp. stands (Crechi et al.
2016).
The main goal of the present case study was to develop a working strategy
(Fig. 18.1) for domestication and conservation of genetic variability of two native
species of the Interior Atlantic Forests, P. dubium and E. contortosiliquum, which
contemplated the following activities (short, medium and long term): 1) selection
and marking of seed trees using the mother seed tree method (White et al. 2007)
covering the ranges of both species in Argentina, and subsequent seed harvest from
these mother trees; 2) study of morphometric characteristics of fruits and seeds of
provenance and progeny of P. dubium and E. contortosiliquum; 3) establishment of
provenance and progeny trials of P. dubium and E. contortosiliquum in the green-
house (short-term) and in the field (long-term) to analyze the genetic diversity of
their adaptive characteristics; and 4) development of mini-clonal gardens of prove-
nance and progeny of P. dubium and E. contortosiliquum for the conservation and
propagation of propagules with genetic diversity, for domestication, enrichment
and/or restoration projects.
Below, we describe each of the activities4 that were carried out.

4
The studies developed in the activities described in this chapter were carried out using appropriate
statistical designs and analyses, which will be described in future publications.
468 F. Niella et al.

Selection and marking of seed

trees in natural range
Harvesting seeds from
provenances and progenies P.
dubium and E.
contortosiliquum

Mini-clonal garden of Genetic diversity for Study of morphological

provenance and progeny domestication, enrichment markers from
propagation provenances and progeny
and restoration programs
(To provide genetic of native species of the (To study the genetic
diversity for regional Paranaense Forest structure of populations)
nurseries)

Establishment of short-term
greenhouse and long-term field
trials of provenances and
progeny.
(To study the genetic structure
and parameters of adaptive
characteristics)

Fig. 18.1 Proposed strategy for the domestication and conservation of genetic diversity of
P. dubium (Caña fístola) and E. contortosiliquum (Timbó) in the Yabotí Biosphere Reserve,
Misiones, Argentina (Own source)

18.2.1 Selection and Marking of Seed Trees and Harvest

of Seeds from Provenances and Progenies
of P. dubium and E. contortosiliquum

A survey of seed trees, and seed harvests were carried out in much of the natural
distribution range of these species in Argentina. Seeds were harvested from individ-
uals in native populations of the northwestern and northeastern region of Argentina
(latitudes 23
280 to 32
08´ South, and longitude 54
300 to 65
18´ West), over an
area of 1,025,000 km2. We were able to harvest seeds from 54 trees of
E. contortosiliquum and 35 trees of P. dubium from the provinces of Misiones,
Corrientes, Entre Rios, Formosa, Tucuman, Salta, and Jujuy.
The information survey in the field and subsequent processing of the data were
carried out as follows: 1- Identification and contact with owners who possessed
isolated individuals or remnant forest with the species under study. 2- Visit to the
property to be able to identify, mark and take the following data from the seed trees:
diameter at breast height (DBH) (cm), shaft height (meters), total height (meters),
treetop diameter (meters), phenological state, sanitary state, environmental variables
and other observations; photograph of the selected tree; information on roads to
18 Contribution to the Domestication and Conservation of the. . . 469

Fig. 18.2 Seed and fruits of one progeny of P. dubium. (Photo: Y. López)

reach the site and location coordinate data with GPS. The information was processed
using a Geographic Information System (GIS).

18.2.2 Study of Morphometry of Fruits and Seeds, from

the Provenances and Progenies of P. dubium
and E. contortosiliquum

A minimum of 2000 seeds per tree were harvested from each geo-referenced tree.
The fruit and seeds morphometric variables of weight, width, length, number of
seeds per fruit, number of seeds per kg of fruits, and number of fruits per kg were
measured for P. dubium, from the material collected, as described in point 2.1.1
(Fig. 18.2).
Seeds of P. dubium and E. contortosiliquum (Fig. 18.3) were also used to study
the germination power, seedling morphometry and survival in the nursery under
controlled conditions of temperature and humidity, and they were also used for the
field progeny trials. The collection sites, i.e. the provinces, are the provenances and
the progeny from each tree registered in this study, are progeny groups. The
morphometric characteristics of fruits and seeds, as well as germination and seedling
growth, were evaluated and measured in the Vegetative Propagation Laboratory of
the Faculty of Forestry Sciences (UNaM), in Eldorado (Misiones).
470 F. Niella et al.

Fig. 18.3 Seed and fruits of one progeny of E. contortosiliquum. (Photo: Y. Lopez)

18.2.3 Establishment of Short-Term Greenhouse, Long-Term

Under-Canopy Enrichment Field Trials, and Mini
Clonal Gardens of P. dubium and E. contortosiliquum

[Link] Short-Term Greenhouse Trials

To perform germination power tests, the seeds of each provenance and progeny of
each species were scarified with sandpaper (150 granulometry) and disinfected with
sodium hypochlorite (NaClO) at 50% (v/v) for 15 minutes, with 4 subsequent rinses
with distilled water. The seeds were germinated in 90 cm3 HIKO™ trays, in winter
with semi-controlled conditions of humidity (micro spray irrigation) and temperature
(between 20 and 30
C). The substrate used was composted pine bark, with
application of Plantacote Plus slow release fertilizer™ (3 kg/m3).
From each offspring, seed batches of between 25 to 40 seeds were taken, and
germinations counts were made after 4, 6, 10, 24, 25 and 31 days. Within 10 days of
planting the germination and emergency stabilized, therefore this was considered the
period to assess the germination capacity. Seedling morphometry and survival were
assessed at 60 and 120 days. Seedling height (cm), neck diameter (ND, mm), and
number of internodes were measured.
18 Contribution to the Domestication and Conservation of the. . . 471

[Link] Long-Term Under Canopy Enrichment Field Trial

of P. dubium and E. contortosiliquum

For the establishment of the under-canopy long-term field trial of provenances and
progenies of P. dubium and E. contortosiliquum, the site selected is in the Munic-
ipality of El Soberbio, Province of Misiones, in the Yabotí Biosphere Reserve
(latitude 27o08’320 ‘South and longitude 53o58’570 ‘West), belonging to the Com-
pany El Moconá SA and managed by Puerto Laharrague SA.
The site has a subtropical climate that corresponds to a Cfa classification
according to Köppen (1936), without a dry season, with average temperatures
ranging from 12 to 31
C and with more than 2000 mm of annual precipitation.
Soils are deep, red Ultisols classified as Kandiudults. For the preparation of the site,
the strips were opened mechanically and manually, ranging from 3 to 3.5 m wide and
90 to 190 m long, with 20 m of undisturbed area between strips.
Seedling establishment was done in a completely randomized block design with
20 replicate blocks. For each 3 m-wide enrichment blocks on the opened strips of
forest, the distance between blocks was 20 m and the seedlings were set in rows with
3 m between seedlings (Fig. 18.4). The following characteristics were evaluated:

Fig. 18.4 Long term field

trial establishment in
enrichment blocks of
P. dubium. (Photo:
P. Thalmayr)
472 F. Niella et al.

survival at the first, second and third year, diameter at breast height (DBH), total
height, and trunk shape. The morpho-physiological variables (leaf size, stomatal
density, photosynthesis, transpiration, phenology), will be evaluated at the fourth,
seventh year and at half of the rotation age. The field experiments are being
maintained up to 3 years post-establishment, with periodic control of weeds and
ants, and replacement of failed seedlings.

[Link] Mini-Clonal Gardens of Provenances and Progenies

of P. dubium and E. contortosiliquum

To ensure the long-term conservation and production of provenances and progenies

of P. dubium, a mini-clonal garden was established, using seedlings produced for the
short-term greenhouse experiment, as described above. The methodology described
by Niella et al. (2013, 2014, 2016a, b) was used for the mini-stumps management
and rooting of mini-cuttings (Figs. 18.5 and 18.6).

Fig. 18.5 Greenhouse

mini-stumps garden
(P. dubium). (Photo:
F. Niella)
18 Contribution to the Domestication and Conservation of the. . . 473

Fig. 18.6 Rooted mini-cutting (P. dubium). (Photo: F. Niella)

18.3 Results and Discussion

18.3.1 Morphometry of Fruits and Seeds from Provenances

and Progenies of P. dubium and E. contortosiliquum

For P. dubium, the morphometric variables of weight, width, length, number of

seeds per fruit, number of seeds per kg of fruits, and number of fruits per kg showed
significant differences among provenances and progenies (Table 18.1, Figs. 18.2 and
18.3). These data agree with the results of a study of morphological variation of
fruits and seeds of Prunus nepaulensis Steud. in Meghalaya, India (Shankar and
Synrem 2012). The weight of the fruit showed the greatest variation (37.94%)
among provenances. Positive correlations were found between fruit weight and
length, and lower correlations among weight, fruit width, and number of seeds per
fruit (data not shown).
Significant differences among provenances and progenies of P. dubium were
observed for all seed morphometric variables (Table 18.2). Of these, seed length
varied the most. Seed weight was positively correlated with germination percentage.
In this study, the association between the size of the P. dubium seeds, expressed
in terms of length and width, with the size of the seedlings with 4 months of growth
was verified. These data agree with other studies that demonstrate a correlation
between seed size with seedling size, verifying that larger seeds produce more
 474

Table 18.1 Means of seed morphometric variables for P. dubium provenance and progenies
Provenances Progenies REP PF (g) AF (mm) LF (mm) N
SEM Sem kg1 Fruits kg1
MISIONES CF31 100 0.23a 15.70b 71.72bc 1.45bcde 6.21 h 4.76i
MISIONES CF23 100 0.20abcd 14.37 cd 80.51a 1.56bcd 7.40fgh 5.27ghi
MISIONES CF32 100 0.20bcde 14.87de 68.84cde 1.42cdef 7.12fgh 5.65fghi
MISIONES CF25 100 0.17efg 17.28a 66.53def 1.21efg 7.81efgh 6.40 cdfg
FORMOSA CF22 100 0.22ab 13.09 fg 67.75cdef 1.70ab 7.84efgh 5.13hi
FORMOSA CF21 100 0.19bcde 14.16 cd 65.11efg 1.60abcd 9.62 cd 6.18cdefg
FORMOSA CF24 100 0.16fgh 12.72 g 64.18 fg 9.91bc 9.91bc 7.46ab
FORMOSA CF20 100 0.15gh 12.84 g 64.62efg 11.39ab 11.39ab 7.08abc
FORMOSA CF30 100 0.14 h 14.66c 70.58bcd 1,16 g 8.05defg 7.73a
JUJUY CF34 100 0.21abc 15.88b 67.84cdef 1.67cbc 7.71efgh 5.27ghi
JUJUY CF17 100 0.18cdef 15.55b 74.38b 1.55abcd 8.52cdef 5.85efgh
TUCUMÁN CF28 100 0.16fgh 13.56ef 71.01bc 8.77cdef 8.76 cdef 6.72bcde
TUCUMÁN CF33 100 0.17efg 14.08cde 70.38bcd 1.8a 11.68a 6.18cdefg
SALTA CF27 100 0.18cdefg 15.88b 67.71cdef 1.17 fg 6.77gh 5.97defgh
CORRIENTES CF29 100 0.17defg 13.15 fg 61.76 g 1.37 cdefg 9.24 cde 6.85abcd
Adapted from Tuzinkievicz (2019)
REP (replications). Means with a common letter are not significantly different (p > 0.05). Test: Tukey Alpha-0.05. PF (g) (fruit weight in grams), AF (fruit width
in mm), LF (fruit length in mm), No. SEM (number of seeds per fruit), Sem kg1 (seeds per kg) and fruits kg1 (fruits per kg)
F. Niella et al.
18 Contribution to the Domestication and Conservation of the. . . 475

Table 18.2 Means of seed morphometric variables for P. dubium provenance and progenies
Provenance Progenies REP PS 1000 (g) Sem kg1 AS (mm) LS (mm)
MISIONES CF31 100 60.20a 16.79i 4.49bc 10.16a
MISIONES CF23 100 51.40 cd 20.06fgh 3.98ef 9.48bc
MISIONES CF32 100 55.71b 1824hi 4.06def 9.20 cd
MISIONES CF25 100 54.21bc 19.11ghi 4.04def 8.77ef
FORMOSA CF22 100 51.22bcd 19.42fghi 4.52ab 9.30bc
FORMOSA CF21 100 50.46cde 20.49fgh 4.76a 8.37 fg
FORMOSA CF24 100 49.93de 20.27fgh 4.24 cd 8.77ef
FORMOSA CF20 100 33.14j 32.30a 3.56 h 8.50efg
FORMOSA CF30 100 35.44ij 29.35b 3.85 fg 8.87de
JUJUY CF34 100 46.58ef 21.89ef 4.02def 9.29bcd
JUJUY CF17 100 40.99gh 24.76 cd 4.02def 8.13 g
TUCUMÁN CF28 100 48.33def 21.68efg 3.88f 9.68b
TUCUMÁN CF33 100 37.85hi 27.27bc 4.18de 9.61bc
SALTA CF27 100 41.27gh 24.95 cd 4.15de 8.76ef
CORRIENTES CF29 100 44.43 fg 23.16de 3.60gh 9.24 cd
Adapted from Tuzinkievicz (2019)
PS 1000 (g) (weight of 1000 seeds in grams). Sem kg1 (seeds per kg). AS (mm) (seed width in
mm). LS (mm) (seed length in mm). REP (number of replications). Means with a common letter are
not significantly different (p > 0.05). Test: Tukey Alpha-0.05

vigorous seedlings, and with better survival. (Frazao et al. 1985; Haig and Westoby
1991; Pastorino and Gallo 2000; Ursulino Alves et al. 2005).
For the species E. contortosiliquum, the studies of fruit and seed morphometry are
still in process.

18.3.2 Short-Term Greenhouse, Long-Term Under Canopy

Trials, and Mini Clonal Gardens of Provenances
and Progenies of P. dubium and E. contortosiliquum

[Link] Short-Term Greenhouse Trials

For P. dubium the results of germination capacity tests stabilized within 10 days of
planting. Seeds from the Jujuy and Entre Ríos provinces had the highest germination
rates. The greatest variation in germination was recorded for seeds from Misiones
and the smallest for seeds from Jujuy. Germination power and survival rate
presented high coefficients of variation both within and between progenies from
different provenances, except progenies from Jujuy (Tuzinkievicz 2019). Seed
weight was positively correlated with germination rate. The emergence speed in
this study was an average of 6.67 AD (average days), compared to the results
observed by Espínola Areco and Rodriguez Espínola (2010), where the P. dubium
seeds registered an emergency speed of 6.85 AD.
476 F. Niella et al.

At 60 to 120 days of growth, P. dubium seedlings showed significant differences

among provenances and progenies for seedling height (HT cm) (P value: 0.001) and
root collar diameter (RCD mm) (P-value: 0.001). Progenies CF10 and CF6 from the
Corrientes provenance had the highest mean heights at 60 and 120 days of nursery
growth. Progenies CF13 and CF15 from the Misiones provenance had the lowest
mean heights at 60 and 120 days of nursery growth. The Misiones provenance had
both the minimum and maximum mean RCD, with progenies CF26 having the
maximum RCD at 60 days and CF31 at 120 days, while CF13 showed the minimum
RDC at 60 and 120 days of growth. Overall, positive correlations were found
between seedling height at 120 days and seed width and length, and between fruit
width and seedling height of seedlings at 60 days, suggesting that the larger the fruits
and seeds, the larger the seedlings.
While there were no significant differences in seedling survival among prove-
nances, there were significant differences in survival among progenies (Table 18.3).
Nevertheless, Misiones presented the highest number of progenies mortality. Sur-
vival was negatively correlated with seedling size at 120 days of growth, suggesting
that as the plants grew, the survival in the nursery decreased if the seedling density in
the tray was high. According to Villagra (2012), P. dubium seedlings are character-
ized by a high light requirement and fast growth. Therefore, since the tray was not
thinned, competition could have been one of the problems. This also agrees with
work on the evaluation of containers for the production of Eucalyptus globulus
Labill, in which they concluded that in some cases, such as those obtained for
P. dubium, the greenhouse low survival rate could be explained by the accelerated
growth of the seedlings, which suppress the growth of other individual seedlings
(Molina et al. 1992).
Significant differences were found among progeny for germination and seedling
growth of E. contortosiliquum, with seedling survival per progeny ranging from 0 to
77% at 60 days of growth (Table 18.4). Preliminary results showed that on the third
day after planting, more than 50% of progeny had at least one seedling germinated,
and that the average germination rate was 23.3% after 120 days of growth (Buchweis
2019). The germination rate in general was low for the species (23%), which could
be attributed to the fact that many of the seed trees from which the seeds were
collected were isolated. This is in accordance with findings by Rocha and Aguilar
(2001), who studied the reproductive biology of E. cyclocarpum, a species of the
same genus, and concluded that trees that grow in continuous forests were almost six
times more likely to produce more seeds than those from isolated trees. Moreover,
the seeds from the continuous forests are more vigorous than those of trees that grow
isolated, probably due to increasing self-pollination rates in isolated trees, thereby
producing less fruit, which are usually abortive.
Significant differences among progenies were found, both for seedling height and
RCD. The Misiones province had the progenies with the highest mean seedling
heights and RCDs followed by progenies from the Corrientes and Tucumán prov-
inces. The overall mean seedling height was 37.3 cm, mean RCD was 5.48 mm, and
mean survival was 44.85% after 120 days of growth (Buchweis 2019).
18 Contribution to the Domestication and Conservation of the. . . 477

Table 18.3 Mean green- Provenances Progenies %SUVGH %SUVFIELD

house (%SUVGH) and field
MISIONES CF 12 30ab 80ab
(%SUVFIELD) survival
capacity at 12 months for MISIONES CF13 0b NA
P. dubium provenances and MISIONES CF14 0b NA
progenies MISIONES CF15 0b 100a
MISIONES CF16 0b NA
MISIONES CF 18 57.5ab 90ab
MISIONES CF 19 7.5b 100a
MISIONES CF 23 42.5ab 71ab
MISIONES CF 25 60ab 75ab
MISIONES CF 26 45.7ab 81ab
MISIONES CF 31 65ab 86ab
MISIONES CF 32 50ab 86ab
MISIONES CF 35 40ab 75ab
CORRIENTES CF 6 50ab 80ab
CORRIENTES CF 7 63ab 85ab
CORRIENTES CF8 0b 100a
CORRIENTES CF 9 11.4b 82ab
CORRIENTES CF 10 66.7ab 90ab
CORRIENTES CF 11 5ab 100a
CORRIENTES CF 29 30ab 80ab
ENTRE RIOS CF 1 14.3b 92ab
ENTRE RIOS CF 2 16b 50b
ENTRE RIOS CF 3 65ab 85ab
ENTRE RIOS CF 4 80a 63ab
ENTRE RIOS CF 5 62.5ab 81ab
FORMOSA CF 20 15b 69ab
FORMOSA CF 21 14b 80ab
FORMOSA CF 22 60ab 80ab
FORMOSA CF24 0b 100a
FORMOSA CF 30 5b 100a
JUJUY CF 17 42.5ab 40b
JUJUY CF 34 55ab 60ab
TUCUMÁN CF 28 12.5b 65ab
TUCUMÁN CF 33 62.5ab 45b
SALTA CF 27 65ab 62ab
Means with a common letter are not significantly different
(p > 0.05). Test: Tukey Alpha-0.05

[Link] Long-Term Under Canopy Enrichment Field Trials

One year after establishment of the field trial, the survival rates of P. dubium
progenies differed significantly, with a mean of 79% (Table 18.3, Fig. 18.4). Prog-
enies of E. contortosiliquum showed no significant differences in field survival
478 F. Niella et al.

Table 18.4 Mean green- Provenance Progenies %SUVGH %SUVFIELD

house (%SUVGH) and field
Misiones TB1 45 89ab
(%SUVFIELD) survival
capacity at 12 months for Misiones TB2 22.5b NA
E. contortisiliquum prove- Misiones T28 75a 80ab
nances and progenies Misiones T29 77.5a 70bc
Misiones T30 55ab 70bc
Misiones T31 10b NA
Misiones T32 15b NA
Misiones TB8 7.5b NA
Corrientes TB3 0 NA
Corrientes TB4 0 NA
Corrientes TB5 35ab 78ab
Corrientes TB6 5 NA
Corrientes TB7 0 NA
Tucumán TB9 52.5ab 67bc
Tucumán TB10 25ab 78ab
Tucumán TB11 32.5ab NA
Tucumán TB12 17.5b NA
Tucumán TB13 0 NA
Tucumán TB14 0 NA
Tucumán T15 62.5ab 100ab
Tucumán T16 22.5b NA
Tucumán T17 5b NA
Tucumán T18 5b NA
Tucumán T19 22,5b NA
Jujuy T21 5b NA
Jujuy T22 5b NA
Jujuy T23 5b NA
Jujuy T24 47.5ab 60bc
Jujuy T25 12.5b NA
Jujuy T26 15b NA
Salta T20 45ab 70bc
Formosa T27 15b NA
Adapted from Buchweis (2019)
Means with a common letter are not significantly different
(p > 0.05). Test: Tukey Alpha-0.05

capacity, with a 70% mean survival (Table 18.4, Fig. 18.7). According to Widiyatno
et al. 2014, the variable of survival rate could be used to select the best family for
breeding programs in the future because this variable showed the adaptation of the
family toward the extreme condition in their early establishment.
18 Contribution to the Domestication and Conservation of the. . . 479

Fig. 18.7 Seedling at the

first-year evaluation field
trial (E. contortosiliquum).
(Photo: R. Buchweis)

[Link] Mini-Clonal Gardens of Provenances and Progenies

of P. dubium and E. contortosiliquum

A mean of 927 mini-cuttings/m2/year was produced for both species, based on three
collections per year; with the mean survival and rooting capacity of mini-cuttings
both above 85% after 120 days of growth in greenhouse conditions (Niella et al.
2014). This technique offers great opportunities for improving the quality of planting
stock, through its capability to produce any quantities of plantation materials in
homogenous quality at any time (Yasman and Natadiwirya 2001).

18.4 Conclusions

At this stage of the study, it can be concluded that:

1. In spite of the low density of individuals/ha, the number of seed trees selected for
germplasm collection of the species P. dubium and E. contortosiliquum met the
480 F. Niella et al.

minimum of 30 randomly selected trees to be sampled according to Rogers and

Montalvo (2004).
2. Fruits and seeds of P. dubium showed differences among provenances and
progenies, and positive correlations between morphometric variables of fruits
and seeds and the height and survival of seedlings at 60 and 120 days.
3. Short-term greenhouse trials showed significant differences among provenances
and progenies of P. dubium for height at 60 and 120 days, and positive correla-
tions between fruit and seed size and height. Field survival, 12 months after
establishment differ significantly among progenies of P. dubium; nevertheless, no
difference was observed on E. contortosiliquum.
4. The strategies presented in this case study agree with the ex-situ conservation and
population improvement methods proposed by Navarro Pereyra (2002), which
include: vegetative propagation of mother trees; import of seeds from other
populations outside the one to be improved; establishment of conservation
gardens that include at least twenty non-related progenies, and development of
low intensity breeding programs for the tropical species.
5. The presented results are preliminary and long-term studies are required for both
species. However, this study lays the foundation for understanding genetic
variability in P. dubium and E. contortosiliquum and provides methods for the
propagation of germplasm to ensure genetic diversity of these species for domes-
tication, enrichment, and/or restoration programs. It will also contribute to delin-
eation of a proper design for a biodiversity island strategy for P. dubium and
E. contortosiliquum.

Acknowledgements The authors thank the Sustainable Management of Natural Resources Pro-
ject, Ministry of Agriculture, Livestock, Fisheries and Food (MAGPyA), the International Bank for
Reconstruction and Development (BIRF 7520 AR)/Project PIA10031, the General Secretariat of
Science and Technology and the School of Forest Science (FCF) of Misiones National University
(UNaM), Misiones, Argentina, as well as the Puerto Laharrague Company, and the forest technician
Federico Robledo for his contribution to field trials establishment.

References

Acquaah J (2012) Principles of plant genetics and breeding. John Willey & Sons LTD, West Sussex
Alexandre RS, Gonçalves FG, Rocha AP, Pereira M, Lemes E (2009) Tratamentos físicos y
químicos na superaçâo de dormência em sementes de Enterolobium contortisiliquum (Vell.)
Morong. Agrária 4:156–159
Barth SR, Eibl B, Montagnini F (2008) Adaptability and species growth. Forestry and environ-
mental technical conferences of the FCF-National University of Misiones and EEA Monte
Carlo. INTA, Eldorado, Misiones, pp 1–16
Bhagwat SA, Nogué S, Willis KJ (2012) Resilience of an ancient tropical forest landscape to 7500
years of environmental change. Biol Conserv 153:108–117
Bruford M, Segelbacher G (2018) IUCN SSC Conservation Genetics Specialist Group. 2018
Report. [Link]
[Link]
18 Contribution to the Domestication and Conservation of the. . . 481

Buchweis R (2019) Study of growth and survival of seedlings in nursery and in the field of progeny
of Enterolobium contortisiliquum from northeast Argentina. Undergraduate’s thesis for the
Forestry Engineering Degree, Faculty of Forest Sciences of the Misiones National University.
[Link]
Crechi E, Keller A, Vera C, Domecq C, Henning A (2016) Growth of 3 native hardwood species in
open field and under pine canopy at 16 years, in Misiones, Argentina (Cordia trichotoma,
Balfourodendron riedelianum, Enterolobium contortisiliquum). XVII Jornadas Técnicas
Forestales y Ambientales. Posadas, Misiones. Producción Forestal Primaria. Pp. 78–80
Eibl B, Morandi F, Muñoz D, Martinez L (1993) Enrichment in sashes with native forest species in
San Pedro, Misiones, Argentina. VII Jornadas Técnicas: Ecosistemas Forestales Nativos, Uso,
Manejo y Conservación. ISIF-FCF. Eldorado. Misiones Actas I:268–277
Eibl B, Morandi F, Muñoz D, Martínez L (1998) El potencial de las especies nativas en programas
de plantación. En: VI Jornadas Técnicas, Serie Técnica No 6, Ecología de Especies Nativas de la
Selva Subtropical Misionera. Universidad Nacional de Misiones, Facultad de Ciencias
Forestales. Eldorado, Misiones, Argentina. Mayo 1998. Pp.19–26
Eibl B, Montagnini F, Lopez M, Lopez LN, Montechiesi R, Esterche E (2017) Organic yerba mate,
Ilex paraguariensis, in association with native tree species: a sustainable production
alternative. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity conser-
vation on food sovereignty, advances in agroforestry 12. Springer, Cham, pp 261–281
Espínola Areco NL, Rodríguez Espínola HN (2010) Efecto de los incleendios forestales sobre la
emergencia de semillas de tres especies forestales nativas en condiciones in vitro. Investigación
Agraria 12(1):29–34
Ferrero B (2005) Study of territorial management and natural resources, of the rural population of
the Area of Influence of the Yabotí-Argentina Biosphere Reserve. Looking for alternatives for
sustainable local development around a Biosphere Reserve. Programa Man and Biosphere,
UNESCO. [Link]/MAB/doc/mys/2004/[Link]. (12 de diciembre 2012) 184 pp
Frazao DAC, Costa SD, Coral FS, Azevedo JA, Figueiredo FJC (1985) Influência do peso da
semente no desenvolvimento e vigor de mudas de cacau (Theobroma cacao L.). Revista de
Agricultura 60(1):2–16
Godefroid S, Piazza C, Rossi G, Buord S, Stevens AD, Aguraiuja R, Cowell C, Weekley CW,
Vogg G, Iriondo J, Johnson I, Dixon B, Gordon D, Magnanon S, Valentin B, Bjureke K,
Koopman R, Vicens M, Virevair M, Vanderborght T (2011) How successful are plant species
reintroductions? Biol Conserv 144(2):672–682
Gómez C, Cardozo F (2003) Enrichment of the native forest with ibira puita guazu (Peltophorum
dubium spreng). Informe interno INTA-Centro Regional Chaco-Formosa E.E.A Sáenz Peña.
Campos Anexo Presidencia de la Plaza
GSPC (2010) Global Strategy for Plant Conservation ([Link]
Haig D, Westoby M (1991) Seed size, pollination costs and angiosperm success. Evol Ecol 5(3):
231–247. [Link]
Jennings SB, Brown ND, Boshier DH, Whitmore TC, Lopes J (2001) Ecology provides a pragmatic
solution to the maintenance of genetic diversity in sustainably managed tropical rain forests. For
Ecol Manage 154:1–10
Kindt R, Lillesø JPB, Mbora A, Muriuki J, Wambugu C, Frost W, Beniest J, Aithal A, Awimbo J,
Rao S, Holding-Anyonge C (2006) Tree seeds for farmers: a toolkit and reference source. World
Agroforestry Centre, Nairobi, Kenya. [Link]
20seeds/Documents%20of%20Toolkit/PDF/[Link]. (Accessed April 2014)
Köppen W (1936) Das geographische System der Klimate. In: Köppen W, Geiger R (eds)
Handbuch der Klimato - logie. Gebrüder Borntraeger, Berlin, pp 1–44
Le HD, Smith C, Herbohn J, Harrison S (2012) More than just trees: assessing reforestation success
in tropical developing countries. J Rur Stud 28(1):5–19
Maina G, Howe HF (2000) Inherent rarity in community restoration. Conserv Biol 14:1335–1340
Martini AMZ, Fiaschi P, Amorin AMA, Paixa˜o JL (2007) A hot-point within a hot-spot: a high
diversity site in Brazil’s Atlantic Forest. Biodivers Conserv 16:3111–3128
482 F. Niella et al.

McNeely JA, Miller KR, Reid WV, Mittermeier RA, Werner TB (1990) Conserving the world’s
biological diversity. World Conservation Union, World Resources Institute, Conservation
International, World Wildlife Fund–US and the World Bank, Washington, D.C.
Molina MPB, Barros DR, Ipinza RC (1992) Análisis de distintos contenedores para la producción
de plantas de Eucalyptus globulus Labill. Ciencia e Investigación Forestal 8(2):169–193
Montagnini F, Eibl B, Woodward C, Szczipanski L, Ríos R (1998) Tree regeneration and species
diversity following conventional and uniform spacing methods of selective cutting in a sub-
tropical humid forest reserve. Biotropica 30(3):349–361
Montagnini F, Levin B, Berg KE (2022) Biodiversity Islands. Strategies for conservation in human
dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies for conservation
in human dominated environments. Springer, Cham, pp 1–35
Montagnini F, Eibl B, Fernández R (2006) Rehabilitation of degraded lands in Misiones. Argentina
Bois et Forets des Tropiques 288(2):51–65
Mori E, Sebbenn A, Tambarussi E, Gurie R (2013) Sistema de reprodução em populações naturais
de Peltophorum dubium. Sci For, Piracicaba 41(99):307–317
Murray-Smith C, Brummitt NA, Oliveira-Filho AT, Bachman S, Moat J, Lughadha EMN, Lucas EJ
(2009) Plant diversity hotspots in the Atlantic Forest in Brazil. Conserv Biol 23:151–163
Myers N, Mittermeier RA, Mittermeier CG, Fonseca GAB, Kent J (2000) Biodiversity hotspots for
conservation priorities. Nature 403:853–858
Navarro Pereyra CM (2002) Genetic resources of Cedrela odorata L. and their efficient use in
Mesoamerica. Academic Dissertation in Forest Tree Breeding. Faculty of Agriculture and
Forestry of the University of Helsinki. ISBN: 952–10–0791-5 (PDF)
Niella F, Eibl B, Rocha P, Dummel C, Keller HA (2016a) Strategy for the conservation, domes-
tication and rescue of plant genetic resources native to the Paranaense forest. XVII Jornadas
Técnicas Forestales y Ambientales. FCFUNaM-INTA-COIFORM. Posadas, Misiones-
Argentina. Pp. 302–304. [Link]
Niella F, Rocha P, Eibl B, Bohren A, Ayala L, Conti P, Franco M (2016b) Development of clonal
propagation techniques for the sustainable use of multipurpose native forest species and/or
nitrogen fixators and non-wood forest products. In: Forest Research 2011–2015: Applied
Research Projects. Propagation section. Buenos Aires: Ministerio de Agroindustria. Unidad
para el cambio rural-UCAR. Pp. 386–389
Niella F, Rocha P, Eibl B, Bohren A, Ayala L, Conti P, Franco M, Radins M (2013) Development
of clonal propagation techniques for the sustainable use of multipurpose native forest species
and/or nitrogen fixers and non-wood forest products. 4to Congreso Forestal Argentino y
Latinoamericano, Iguazú, Misiones, Argentina
Niella F, Rocha P, Eibl B, Schoffen C, Martinez M, Conti P, Franco M, Ayala L (2014) Clonal
propagation of Peltophorum dubium (cane fistola), Myrocarpus frondosus (incense), and
Cordia trichotoma (Peteribi) for conservation and domestication. Revista Forestal Yvyrareta
21(2014):43–50. [Link]
2014-n21
Pastorino MJ, Gallo LA (2000) Variación geográfica en peso de semilla en poblaciones naturales
argentinas de “Ciprés de la Cordillera”. Bosque (Valdivia) 21(2):95–109
Pauls SU, Nowak C, Bálint M, Pfenninger M (2013) The impact of global climate change on
genetic diversity within populations and species. Mol Ecol 22(4):925–946
Putz FE, Redford KH, Robinson JG, Fimbel R, Blate GM (2000) Biodiversity conservation in the
context of tropical forest management. In: The World Bank environment department, biodiver-
sity series impact studies paper 75. The World Bank, Washington DC, USA
Rajora OP, Pluhar SA (2003) Genetic diversity impacts of forest fires, forest harvesting and
alternative reforestation practices in black spruce (Picea mariana). Theor Appl Genet 106:
1203–1212
Ratnam W, Rajora OP, Finkeldey R, Aravanopoulos F, Bouvet JM, Vaillancourt RE, Kanashiro M,
Fady B, Tomita M, Vinson C (2014) Genetic effects of forest management practices: global
synthesis and perspectives. For Ecol Manage 333:52–65
18 Contribution to the Domestication and Conservation of the. . . 483

Reed BM, Sarasan V, Kane M (2011) Biodiversity conservation and conservation biotechnology
tools. In Vitro Cell Dev Biol Plant 47:1–4
Reed DH, Frankham R (2003) Correlation between fitness and genetic diversity. Conserv Biol
17(1):230–237
Rocha O, Aguilar G (2001) Reproductive biology of the dry forest tree Enterolobium cyclocarpum
(Guanacaste) in Costa Rica: a comparison between trees left in pastures and trees in continuous
forest. Amer J Bot 88(9):1607–1614
Rogers DL, Montalvo AM (2004) Genetically appropriate choices for plant materials to maintain
biological diversity, report to the USDA Forest Service. University of California, Rocky
Mountain Region, Lakewood
Royal Botanic Gardens, Kew (2003) A field manual for seed collectors. Seed Collecting for the
Millennium Seed Bank Project, Royal Botanic Gardens, Kew
Schlegel RHJ (2010) Dictionary of plant breeding, 2nd edn. CRC Press Taylor & Francis Group,
Boca Raton
Shankar U, Synrem YIL (2012) Variation in morphometric traits of fruits and seeds of prunus
nepaulensis steud in Meghalaya, India. Trop Ecol 53(3):273–286
Sist P, Fimbel R, Douglas S, Nasi R, Chevallier A (2003) Towards sustainable management of
mixed dipterocarp forests of Southeast Asia: moving beyond minimum diameter cutting limits.
Environ Conserv 30(4):364–374
Thomas E, Jalonen R, Loo J, Boshier D, Gallo L, Cavers S, Bordacs S, Smith P, Bozzano M (2014)
Genetic considerations in ecosystem restoration using native tree species. For Ecol Manage 333:
66–75
Tuzinkievicz A (2019) Geographical variation of morphometric characteristics of fruits, seeds and
seedlings, germination power and survival in the origins and progeny of Peltophorum dubium
(Spreng.) Taub. of the northwest and coastal region of Argentina. Master’s thesis in Forestry
Sciences, Faculty of Forestry Sciences of the Misiones National University. [Link].
[Link]
Ursulino Alves E, de Lucena R, Alcântara B, Pereira de Oliveira A, Ursulino Alves A, de Paula DC
(2005) Influência do tamanho e da procedência de sementes Mimosa caesalpiniifolia Benth.
Sobre a germinação e vigor. Revista Árvore 29:877–885. [Link]
S0100-67622005000600006
Vallee L, Hogbin T, Monks L, Makinson B, Matthes M, Rossetto M (2004) Guidelines for the
translocation of threatened plants in Australia, 2nd edn. Australian Network for Plant Conser-
vation, Canberra, Australia
Villagra M. (2012) Plasticidad morfológica y fisiológica de especies arbóreas del Bosque Atlántico
en respuesta a cambios en la disponibilidad de luz y nutrientes. Tesis doctoral. Facultad de
Ciencias Exactas y Naturales. Universidad de Buenos Aires. 139pp
White T, Adams W, Neale D (2007) Forest genetics. CABI Publishing, CAB International,
Wallingford, UK, p 682
Wuethrich B (2007) Biodiversity. Reconstructing Brazil’s Atlantic rainforest. Science 315:1070–
1072
Yasman I, Natadiwirya M (2001) Dipterocarp plantation: the strategy and the approaches of PT
Inhutsni I. In: Tielges BA, Sastrapradja SD, Rimbawanto A (eds) In-situ and ex-situ conserva-
tion of commercial tropical trees. ITTO-Gadjah Mada University, Yogiakarta, Indonesia, pp
407–412
 Part IV
Safeguarding the Environmental,
Economic, and Social Benefits
of Biodiversity Islands
Chapter 19
How Community-Led Action Can Advance
the Development of Biodiversity Islands

Brett Levin

Abstract Community-led action can contribute to the development of biodiversity

islands. Biodiversity islands constitute areas of high biodiversity nested within
human-dominated landscapes. Community action towards the development of bio-
diversity islands may be longstanding within cultures or learned and applied from
ethical, philosophical, scientific, cultural, or economic motivations. Land access and
long-term decision-making power provide the basis for communities to maintain
areas of high biodiversity within degraded landscapes on private, public, and
indigenous lands. The legal system and governing process of the presiding people
and culture determine the tools that can aid in the establishment and protection of
these areas. Some useful legal and financial tools may include land trusts, conser-
vation easements, supportive zoning, novel financial resource pooling, and strong
indigenous land rights. Community-developed biodiversity islands may be governed
through a vast array of methods. Management of biodiversity islands requires
methods that enhance or maintain biodiversity outcomes through time. This can be
achieved through a broad array of community land uses and techniques. Examples of
grassroots community action for the advancement of biodiversity conservation
practices are numerous, diverse, and worldwide. Indigenous, religious, governmen-
tal, nonprofit, and for-profit organizations are capable of further expanding
community-led action for the development of biodiversity islands. Examples of
community-led biodiversity islands are described, including spiritual and religious
sites, public areas, agricultural systems, and beyond. No matter the governing and
organizational approach, considerations of social and environmental justice remain
an important factor in how biodiversity islands are recognized, developed, and
managed through time.

Keywords Advocacy · Conservation easements · Governance · Indigenous

management · Land trusts · Sacred sites

B. Levin (*)
Dietrick Institute for Applied Insect Ecology, Ventura, CA, USA
e-mail: [Link]@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 487
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
488 B. Levin

19.1 Introduction

Biodiversity islands are areas of high biodiversity nested within ecologically

degraded, human-dominated landscapes (Montagnini et al. 2022). Community-led
action can advance the development and perpetuity of biodiversity islands. Through
the empowerment of traditional biodiversity conservation practices, grassroots com-
munity action, and the utilization of legal and financial tools, community-led
management can protect intact sections of land where plants and animals can thrive
without ecologically degenerative interference from human activity (Montagnini
et al. 2022). Such areas of high biodiversity within otherwise degraded landscapes
can be found in rural areas, for example, in agroforestry systems (Montagnini and del
Fierro 2022) as well as in suburban and urban contexts such as in homegardens,
residential gardens, and urban green infrastructure (Toensmeier 2022; Negret et al.
2022; Soler et al. 2022).
In many instances, traditional community-based approaches to landscape man-
agement include biodiversity enhancing outcomes which can be considered biodi-
versity islands. Examples include sacred sites, sacred groves, community forests,
church forests, and other sites that are protected and managed by local communities
(Bhagwat and Rutte 2006; Ceperley et al. 2010; Moeliono et al. 2015; Baez Schon
et al. 2022). The term “biodiversity island” in many cases is used to refer to these
indigenous and traditional biodiversity conservation practices, highlighting results
and understandings around procedures that advance appreciation and empowerment
of such systems.
This chapter highlights the commons, which are the cultural and environmental
resources accessible to all members of a society, including ecological assets such as
air, water, and ecosystem health. These resources are held in common and not
privately owned. Emphasis is placed on advances in community-based conservation,
as governance approaches that tend to appreciate social complexities, utilize demo-
cratic and inclusive decision-making processes, and advance pluralistic frameworks.
Grassroots community action and legal tools such as conservation easements
through partnerships with non-profit Land Trusts can also aid in the development
of biodiversity islands. Examples of grassroots efforts for the advancement of
conservation practices are numerous, diverse, and worldwide. Legal frameworks,
methods of enforcement, modes of implementation, and levels of community
engagement are site and context dependent, but through these efforts of
community-led action, biodiversity islands may be protected and managed
through time.

19.1.1 Community Defined

A community can be defined as a group of individuals living in the same area or who
have a characteristic in common (Berkes 2004, 2017). The scale and scope of this
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 489

definition can encompass a significant range of variables dependent on culture and

perspective, from humancentric to biocentric and animistic (Harvey 2005). Such an
inclusive definition of community may naturally lead to sustainable uses of land
which protect biodiversity and provide for human habitation through time. One such
example includes the aboriginal people of Australia, where worship of nature and the
abiotic features of the landscape leads to a great respect for conservation and the
value nature holds. In Oaxaca, Mexico, low intensity forest use and traditional
rotational agriculture are spurred by deep connections to the land. This creates a
more diverse forest-agriculture mosaic where traditional forest management harmo-
nizes livelihoods and sustained biodiversity conservation (Berkes 2017). The
Kenyah Dayak people of Indonesia manage subsistence orchards as gifted foresters.
Their gardens look wild but are carefully cultivated (Peters 2018). Such morally and
spiritually motivated biocentric approaches towards landscape management are
present within many indigenous groups around the world. When considering
community-led action towards biodiversity, such varied perspectives produce inter-
esting considerations; the relinquishment of humans as the main component of
community to humans as listeners or collaborators; the shift from anthropocentrism
to biocentrism; and pathways towards rights of nature in the dialogue for community
conservation (Prieto Méndez 2013).
From a less metaphysical and more human centric perspective, one cannot deny
the abiotic factors of climate, topography, geology, soil, and precipitation in deter-
mining the species present and historical development of humans in an area (Dunson
et al. 1991). Patterns of settlement are interrelated with the presence or absence of
natural resources. Transportation corridors tend to follow ease through topographies.
Agriculture and cultivation are inextricably linked to soils, water resources, and
climate. From this vantage point, biological and physical components of a place are
inextricably linked with community, whether centrally focusing on the human
dimensions or expanding into the biophysical.

19.2 Motivations for Managing for Biodiversity

Through Time

For numerous indigenous and traditional communities, the motivations for conserv-
ing and managing biodiversity through time are an integral part of the culture.
Interactions with the landscape flora and fauna are learned and passed down over
generations, which maintain or enhance biodiversity. These interactions form the
basis of Traditional Ecological Knowledge (TEK) and become nested in culture
(Gadgil et al. 1993; Inglis 1993). In ecologically degraded human-dominated land-
scapes, learning from, protecting, properly valuing, and/or restoring such
community-led cultural practices may allow for areas of higher biodiversity, or
biodiversity islands, to persist or emerge (Berkes 2017). This is not to advocate
notions of the “noble savage” and attributes of romantic primitivism across all
490 B. Levin

traditional or indigenous cultures (Raymond 2007). Instead, here a focus on scien-

tific findings, supported by historical ecology, demonstrate longstanding ethno-
ecological knowledge leads to improved biodiversity outcomes in numerous com-
munities worldwide (Berke et al. 1994; Balée 1998).
Several cases exist where improved biodiversity outcomes result from expanded
perspectives of community, inclusive of genetic diversity, habitat diversity, land-
scape diversity, and temporal diversity (Salick 2012). One such example is
evidenced in Maranhão, Brazil, with landscape management of increased biodiver-
sity by indigenous people focusing on sustainable extraction of specific forest
products, and the nurturance of specific forest plant species (Balée 1993). The loss
of such long-standing land-based culture may result in biological degradation, even
when newly implemented management objectives are advanced to achieve ‘conser-
vation’ or ‘biodiversity goals’ (Crosby 2004). Another example relates to the
traditional agricultural management of Pima farmers in the Sonoran Desert. There,
many emerged volunteer plant species were left in the field for a broad range of
reasons including more diverse food yields, to host edible insect larvae, for protec-
tive hedges, and to provide shade and cycle nutrients. Such intensive and intentional
practices have become a rarity in the now conventionally cultivated fields of the
region (Nabhan et al. 1989). These examples highlight cultural practices which
maintain or enhance biodiversity over time, and suggest that protecting and
empowering such practices can play a key role in the development and expansion
of biodiversity islands.
For other cultures and societies, learned ethical, scientific, philosophical or
economic motivations may drive actions for community-led biodiversity conserva-
tion within a landscape. From a western perspective, there is a broad range of
literature and media that has helped advance such notions. Books such as Aldo
Leopold’s A Sand County Almanac (1949), Rachel Carson’s Silent Spring (1962),
James Lovelock’s Gaia Hypothesis (1979), E.O. Wilson’s Biophilia (1984), and
Murray Bookchin’s Ecology of Freedom (1982) are just a few highlighted examples
that will be described below.

19.2.1 Philosophical and Ethical Motivations

Aldo Leopold’s “land ethic” brought forth an advancement in ethics as a process of

ecological evolution through a moral responsibility for western humanity to care for
the natural world (Leopold 1949). From this vantage point, the moral sense of right
and wrong and how community is defined expanded beyond humans to include the
soils, trees, grasses, and all species and biophysical components of the environment.
Leopold’s “community concept” argues true community-led action must then be
built upon consideration of these human and non-human pieces. From this stand-
point, where humans alter a landscape in ways that diminish non-human life through
time, an ethical violation has occurred. Bringing this vantage point into the realm of
western consciousness provides a foundation for people to come together towards
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 491

the protection of species where landscapes would otherwise become degraded. This
learned ethical approach towards conservation, biodiversity, and their inherent
values has helped shape the recommendations of community organizations, planning
bodies, and governments. Throughout the United States, many recent conservation
projects which may be recognized as biodiversity islands within a landscape can be
attributed to the realization of the philosophical underpinnings of Aldo Leopolds’
work. Today, his legacy continues through The Aldo Leopold Foundation (www.
[Link]). Located on his home site in Baraboo, Wisconsin, the organization
was founded in 1982 by his children as a non-profit organization to continue the
practices incorporated by his land ethic. The foundation owns and manages his
property to conduct education and outreach for land stewardship programs. There
are also other academic and research institutions throughout the United States, run
through the Forest Service, that function as centers of wilderness education, agri-
culture, and programs, celebrating Leopold’s legacy and promoting his ideals and
teachings towards a land ethic.
James Lovelock’s Gaia Hypothesis, which proposed that the biosphere is a
complex and self-regulating organism, advanced philosophical motivations towards
conservation of biodiversity (Lovelock 1979). This hypothesis promulgated the
notion that living matter on earth collectively determines and controls the material
conditions necessary for the regulation of life. From this premise, philosophy
regarding the role of humanity as part of Gaia permeated the realm of western
environmental consciousness. Are the negative environmental impacts of humans
a destabilizing force which will eventually be self-regulated? Can humans persist as
part of a global community of living organisms in a fashion compatible and
harmonious with self-regulation? Many of us are still seeking answers to these
questions, though the notion of humans as a force within the planetary community
towards improved biodiversity was advanced by Lovelock’s work.
E.O. Wilson’s Biophilia emphasized the philosophical underpinnings of the
human compulsion and innate drive towards interactions with other forms of life.
Wilson argued that “the connections that human beings subconsciously seek with the
rest of life are inherent in our biology.” It is argued that modern western society has
disengaged the masses in recognizing and engaging with their love of life-giving
systems. This mindset is responsible for ecological degradation and biodiversity loss
(Wilson 1984). To reverse these outcomes, Wilson argues a philosophical motiva-
tion towards conservation of species through reconnecting with our inherent
biophilic nature. Through this teaching and mindset, community groups and orga-
nizations have worked together to advance conservation projects, many of which
may be considered biodiversity islands.
Murray Bookchin’s book “Ecology of Freedom” also advanced philosophical
motivations towards community-led action towards greater biodiversity through the
message of social ecology, linking human relationships within society to human
relationships towards the land and natural systems (Bookchin 1982). In societies
where domination and exploitation are the norm, one may recognize similar patterns
imposed on the landscape. These outcomes may also manifest in a downtrodden
psyche of the individual. It is suggested that societies built upon foundations of
492 B. Levin

mutual respect among people may materialize a more uplifted psyche of the indi-
vidual and permeate into a more harmonious societal relationship with the landscape
and natural resources. Though criticized as utopian by some, the greater message
purveyed is that through such reciprocal positive mechanisms of right relationship
between people to people and people to place, the individual spirit may be lifted, and
greater freedom emerge. Through this philosophical concept, further motivations
towards greater biodiversity outcomes have entered the western canon of environ-
mental thought, in conjunction with advances in environmental ethics, as discussed
in the previous section.
More recently, evolution in environmental philosophy and ethics advanced by
Stephen Kellert, Mary Evelyn Tucker, and John Grimm, among others, has contin-
ued to shape a mindset towards a more interconnected worldview of community. In
Mary Evelyn Tucker and Brian Swimme’s “Journey of the Universe,” the intercon-
nection and interrelation of all beings brings a biophilic understanding towards the
basis of community (Swimme and Tucker 2011). Stephen Kellert advanced an
ethical imperative within design to incorporate both human and ecological consid-
erations within any project (Kellert 2018). “The Value of Life” is an exploration of
the actual and perceived importance of biological diversity for human beings and
society (Kellert 1997). Stephen R. Kellert identifies ten basic values, which he
describes as biologically based, inherent human tendencies that are greatly
influenced and moderated by culture, learning, and experience. Drawing on
20 years of original research, he considers the universal basis for how humans
value nature. Differences in those values vary by gender, age, ethnicity, occupation,
and geographic location. He discusses how environment-related activities affect
values, variation in values relating to different species, how values vary across
cultures, and their policy and management implications. Throughout his book
“Nature by design: The practice of biophilic design,” Kellert argues that the preser-
vation of biodiversity is fundamentally linked to human well-being in the largest
sense, as he illustrates the importance of biological diversity to the human sociocul-
tural and psychological condition (Kellert 2018).

19.2.2 Scientific Motivations

Rachel Carson advanced biological conservation and subsequent conservation pro-

jects through the scientific underpinnings of “Silent Spring” her famous book that
pinpointed the environmental harms brought about by widespread and unregulated
pesticide use (Carson 1962). Carson’s effective and open communications of the loss
of biological diversity brought regulation, ignited public discourse, bolstered the
formation of the United States Environmental Protection Agency (EPA), and
inspired a generation of environmentalists. This heightened awareness and pro-
foundly shaped the western mindset regarding the environment.
Advances in conservation biology, regarding the measurement, distribution,
abundance, and loss of species through time create another motivating factor for
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 493

conserving biodiversity (Hawksworth 1995). As it becomes clear that more habitat is

lost due to deforestation and habitat destruction, science-based motivations for
community-led efforts for biodiversity conservation grow (Hawksworth 2010a, b).
As such, knowledge regarding inventories for understanding baseline terrestrial
biodiversity for estimating both local and global species diversity can allow for
both rapid assessment and comparison of species diversity across geographies
(Colwell and Coddington 1994). Numerous textbooks, journals, and publications
have been created to further advance knowledge in these realms (Hawksworth
2012). As it becomes clear that species loss is greater, motivations to implement
practices for conservation are bolstered. In many instances, communities may
advance and aid in the scientific understanding of conservation or utilize scientific
methodology to implement best practices in biodiversity conservation.
Overall, there can be a strong motivation towards the advancement of biodi-
versity islands from the academic and scientific realm. Biodiversity islands are the
subject of study from many different angles, from edge effect to island biogeo-
graphy (Montagnini et al. 2022). Deepening the scientific understanding of forest
patch dynamics also provides a motivation for conservation of such landscape
features.

19.2.3 Grassroots Motivations

Other motivations for biodiversity conservation may arise from grassroots actions
and through counterculture. In communities where, for a variety of reasons, institu-
tional or governmental engagement has been ineffective, grassroots education and
action may emerge to bring forth greater biological diversity through community
efforts in degraded, human-dominated landscapes. This can be seen in various forms
and is distinctly recognized through development of grassroots community garden-
ing, urban agriculture, and in the permaculture movement worldwide (Veteto and
Lockyer 2008; Ferguson and Lovell 2014).
Gardens are known to have the potential to foster significant diversity in cultural,
biological, and agro-biological ways (Galluzzi et al. 2010; Goddard et al. 2010;
Negret et al. 2022). In urban centers throughout the United States and beyond,
community gardens have emerged, bringing forth both resilient food systems and
positive biodiversity outcomes from grassroots formation (Clarke and Jenerette
2015; Di Pietro et al. 2018). Additionally, the permaculture movement is responsible
for the development of thousands of projects throughout the globe which enhance
biodiversity outcomes (Toensmeier 2022). Built upon ethical principles, permacul-
ture and the training course known as a Permaculture Design Certificate (PDC) offer
students a method for designing biologically diverse “permanent” agriculture sys-
tems and explore methods for sustained human habitation on a landscape (Mollison
1988). Similar local agroecological trainings, workshops, and community-led
494 B. Levin

courses provide a grassroots approach to such training, especially in those cases

where institutional and formal education fails to provide this information.

19.2.4 Economic Motivations

Economic motivations may also drive community-led action towards the develop-
ment of biodiversity islands. Although historically economic motivations have often
led to exploitation and landscape degradation, such motivations can also be a force
for conservation of biodiversity.
Examples of profitable biodiversity enhancing community enterprise models span
a diverse range of geographies, cultures, products, and services offered (Hay-Edie
and Halverson 2006). This includes sustainable ecotourism, where visitors pay to
stay and engage with local cultures and traditions. Though complex and sometimes
difficult to obtain, the biodiversity of ecotourism sites can be significantly higher
than that of the surrounding human-dominated and degraded landscape (Gossling
1999; Chung et al. 2018). Examples of other community enterprises which support
similar biodiversity outcomes have been shown to produce a broad range of products
such as basketry, incense, herbal medicines, and teas (Jarrett et al. 2017; Rocha et al.
2017).
Agricultural production may also play an important role in community enterprise
development which supports biodiversity conservation within a broader degraded
landscape (Badgley 2018). Sustainable agricultural models in which biodiversity
islands are developed through collectives and community action include crops such
as cacao, coffee, tea, yerba mate, dried fruits, and various other species (Erisman
et al. 2016; Dudley and Alexander 2017; Hunter et al. 2017; Montagnini and del
Fierro 2022). Sustainable timber production has also been a method of enterprise
creation that can respect and support biodiversity conservation within degraded
landscapes in addition to positive economic outcomes (Carey et al. 1999).

19.3 Land Access, Tenure, and Long-Term Control

for Pockets of Biodiversity

For any community action towards the development of biodiversity islands to take
place, long term control, tenure, and decision-making authority over the land base
are paramount. Community land management without legal authority may prove
difficult into posterity. This is documented in numerous case studies worldwide,
from local government closure of community gardens on vacant urban land to the
removal of indigenous people from ancestrally managed lands (Springer 2009;
Holmes and Cavanagh 2016). Acquiring land title or the ability to manage lands
in response to governmental territorial expansions may prove particularly
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 495

challenging for societies without robust property laws or in cultures without con-
cepts of land ownership. In societies where property rights are recognized and
biodiversity outcomes are a community objective, people can find innovative ways
to engage with legal frameworks to gain and maintain land access and tenure through
time. Examples of such methods include use of land trusts and conservation ease-
ments, supportive zoning, inventive financial resource pooling, robust laws
supporting indigenous land rights, and the use of public lands which encourage
community-led biodiversity conservation. These methods are discussed in the fol-
lowing section.

19.3.1 Conservation Easements and Community Land Trusts

Conservation easements are one tool that may be advocated for by a community for
the development of biodiversity islands. Conservation easements are voluntary legal
agreements between landowners and a land trust or government agency that protects
conservation values on a property by permanently limiting uses of the land and
offering tax incentives to the landowner. A land trust or community land trust is a
non-profit organization that acquires land or conservation easements through support
from donations or government funding. The opportunity for land trusts to maintain
and protect biodiversity through such conservation easements is significant
(Rissman et al. 2007; Wilson 2011). In many instances, a land trust acts as a
conservation organization to help draft, implement, and ensure compliance with an
easement while prioritizing community and conservation goals. Community land
trusts are a subset of land trusts which specifically focus on the development and
conservation of community assets for community benefit. Such a model may be
particularly valuable towards the development of biodiversity islands. There are
several examples around the world where conservation easements are used for the
protection of biodiversity in the USA, Canada, Australia, Costa Rica, and other
countries (Alexander and Hess 2012).
Conservation easements may include restrictions on development and land uses
such as recreation, forestry, agroforestry, or agriculture. These easements, which
provide conservation value in addition to opportunities for land-based revenue
generation, are known as working land conservation easements. Throughout the
world, communities can support and develop community land trusts to assist in the
development of community-led biodiversity islands to transition private lands into
long-term community conservation projects. In Chap. 1 (Montagnini et al. 2022) an
example is shown of a Land Trust that received a donation from a family whose
members preferred that their land was preserved and used for recreation instead of
selling it. This is just one example where through land trusts and conservation
easements community action provides protection of a biodiversity island and judi-
cious management for posterity.
496 B. Levin

19.3.2 Innovative Financing for Biodiversity Islands

The use of innovative financing to acquire property is another method for long-term
community-led land control which can support the development of biodiversity
islands. A group of community members or a community organization may pool
financial resources to acquire land outright through a cash purchase or a loan. This is
seen in numerous countries throughout the world including lands and resources
surrounding Juan Castro Blanco National Park in Costa Rica (Castro-Arce and
Vanclay 2020). This community pooling of resources is also known as
crowdsourcing. While there has been significant advancement and literature on the
use of crowdsourced data collection in relation to biodiversity conservation, such as
monitoring bird species and populations, there is much opportunity for the use
of internet platforms as tools for further crowdsourced funding for the acquisition
of property which may enhance biodiversity outcomes through time. The advent of
multiple online platforms which allow for numerous smaller donations or invest-
ments to develop into projects, provides a groundwork for the advancement of
crowdfunding towards biodiversity conservation (Gallo-Cajiao et al. 2018).

19.3.3 Indigenous Land Rights Protecting Biodiversity

Strong indigenous land rights can allow for biodiversity islands to emerge or persist
throughout otherwise degraded landscapes. Dominion within current or historically
inhabited and managed indigenous lands can support sovereignty, knowledge shar-
ing, cultural empowerment, and positive biodiversity outcomes through time (Lang-
ton and Rhea 2005; Erikson 2008; Sobrevila 2008; Garnett 2018; Baldwin and
Beazley 2019; Beller et al. 2020). Native title (Australia), Indian title (United States),
Customary title (New Zealand) or Indigenous or Aboriginal title are common law
doctrines which strengthen sovereignty of historical land rights through recognition
by governments. While such rights may be powerful, many indigenous peoples
emphasize indigenous rights that do not necessitate state sanctions to exist (Gilbert
2016).
Secure and long-term indigenous management of landscapes are worth highlight-
ing in relation to community-led action toward the development of biodiversity
islands. Research on this topic is steeped in complexity. Not all indigenously
managed lands promote biodiversity, particularly in instances where resources
were overexploited through time (Raymond 2007). Additionally, there are many
examples where indigenous sovereignty has been withdrawn in the name of biodi-
versity conservation (Baldwin and Beazley 2019; Beller et al. 2020). With these
considerations in mind, this section seeks to highlight the potential power and
opportunity of indigenous land rights as a tool for community-led biodiversity
islands to emerge and persist in otherwise degraded landscapes.
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 497

19.4 Governance and Management of Biodiversity Islands

Once land tenure is achieved, proper governance and subsequent management of

biodiversity islands is essential for long term success. Such governance and man-
agement can increase biodiversity through time, allow community members to have
a voice in decision-making, and avoid resource degradation, species loss, and a
tragedy of the commons. Governance can take many varied forms across a range of
scales depending on culture and context. Commonly, hierarchical decision making
dominates organizational governance. While hierarchical governance is effective to
achieve certain ends, alternative egalitarian governance structures may also be
effective for community managed lands (Leventon et al. 2019). This chapter high-
lights three methods of land governance aligned with community-led action that may
support high levels of biodiversity within a larger degraded landscape through
commons, cooperatives, and community based conservation approaches.
Governance as commons, where land is held equally by all community members,
allows for egalitarian ownership of a space. Common resources governed and
adapted to local conditions with clear boundaries, measured outcomes, defined
rules and mechanisms for conflict resolution, and self-determination prove resilient.
Such governance allows for nested enterprises to emerge and common resource
pools to persist through time. While some argue that community-owned resources
degrade through time due to motivations of self-interest leading to overuse, deemed
a tragedy of the commons, countless examples of long-term community managed
resources refute such claims (Hardin 1968; Ostrom 1990). As applied to biodiversity
islands, these governance strategies and other types of community governance are
foundational support for management through time.
Cooperatives as an organizational approach can also provide a more egalitarian
governance and management method to community-fostered biodiversity islands. In
such approaches, ownership and decision making tend to be more equitably distrib-
uted throughout the organization, and outcomes, yields, and profits are then distrib-
uted accordingly. Globally, examples of agricultural cooperatives that value and
incorporate agrobiodiversity into their cultivation practices are numerous (Méndez
et al. 2007). Cooperative organizations which incorporate ethos of biodiversity
conservation span many industries and both non-profit and for-profit ventures.
Such systems can enhance social capital, which has shown to maintain positive
biodiversity outcomes within degraded landscapes (Pretty and Smith 2004).
Advances in community-based conservation (CBC) may also provide the mech-
anisms toward effective development, governance, and management of biodiversity
islands through time. CBC seeks to align development and conservation outcomes
emergent from the community. Criticism of such methods of conservation have
emerged due to the preponderance of linear thinking about development, often
considering a single variable such as Gross Domestic Product (GDP) and situations
where conservation and human habitation are seen as incompatible. Approaches that
consider humans as part of the ecosystem and promote wide scale participation in
ecosystem management provide a more feasible context from which CBC may
498 B. Levin

emerge (Otto et al. 2013). The integration of concepts of common property, tradi-
tional ecological knowledge, environmental ethics, political ecology, and environ-
mental history can further support the longevity of projects, providing knowledge of
the past, reflection of the present resources, opportunities among stakeholders, and a
context for visioning future goals.
The importance of fluid and adaptive collaborative approaches which recognize
multiple stakeholders and development goals may also contribute significant value
to CBC projects (Berkes 2004). Participatory consent-driven decision making as
exemplified by the frameworks of Holacracy,1 provides an example of participatory
governance which may be applied to further mature CBC (Robertson 2015). The
examples of CBC projects throughout the world that have increased biodiversity
within degraded landscapes are extensive (Otto et al. 2013). For this reason, it is
important to note that CBC can provide a strong avenue for community-led action
towards the development of biodiversity islands.

19.5 Community Engagement and Advocacy Towards

Biodiversity Islands

Community engagement and advocacy are other important forces towards the
development of biodiversity islands. Intercultural and interpersonal communication,
education and political engagement which support the appreciation and values of
biodiversity within a landscape are foundational. Interpersonal communication
through talking and engaging with neighbors and community members and fostering
good relationships is an essential first step in building community coalitions for
biodiversity conservations. Once achieved, intercultural communication, with dif-
ferent belief systems, histories, stories, and relationships to landscape and place, can
then build the foundations from which a more biodiverse landscape may arise within
a community (Pretty et al. 2009).
Environmental education focused around topics of biodiversity are shown to be
effective to engage, empower, and bridge science and social issues (Van Weelie and
Wals 2002). When this knowledge is imparted on youth, a next generation may
prioritize issues of biodiversity conservation. When this knowledge is shared with
politicians through actions of lobbying and political engagement, innovative legal
tools and frameworks may be adopted or developed by governments. Scheduling
meetings with local, regional, and national representatives, attending governmental
meetings, writing letters, and coalition building are strategic tactics for empowering
government action.

1
Holacracy is a method of decentralized management and organizational governance, in which
authority and decision-making are distributed throughout self-organizing teams rather than being
vested in a management hierarchy.
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 499

19.6 Examples of Community-Led Conservation Strategies

Examples of grassroots community action for the advancement of conservation

practices are numerous, diverse, and worldwide (de Boef et al. 2013; Otto et al.
2013). Ancient traditional cases of biodiversity islands within the landscape are
exemplified by religious and sacred sites worldwide. Other examples span urban and
rural settings, agrarian and nomadic cultures, and incorporate various organizational
approaches.
Sacred groves, or natural sites that are dedicated to ancestral and spiritual deities,
are found throughout the world. These sites often harbor and protect greater
biodiversity than surrounding degraded landscapes (Bhagwat and Rutte 2006;
Khan et al. 2008). Church forests, ancient temples, and dedicated sections of rivers
and waterways have been protected and maintained by cultures throughout the globe
for religious purposes for millennia (Baez Schon et al. 2022). Deeply held
intergenerational importance of such sites provides strong community incentive
for conservation through time. Other examples of sacred sites that serve as biodi-
versity islands include the Mizorom sacred groves in Northeastern India, sacred
pools called Íbú ódó protected by Tchabè communities along the Ouèmé and Okpara
Rivers of Central Benin (West Africa), sacred cacao groves of the Maya, and other
sacred groves in Zimbabwe, Ghana, Thailand, China, and Nepal (Gómez-Pompa
et al. 1990; Gadgil et al. 1993; Bhagwat and Rutte 2006; Ceperley et al. 2010). These
biodiversity islands share common outcomes where community action enhances and
protects biodiversity through time in human-dominated landscapes. Although the
original purpose in most cases is not to create biodiversity islands, biodiversity
islands are a consequence of their actions. Other community-protected sites include
the village forests in Indonesia, known as Hutan Desa, which are legally recognized
for the ecosystem services and benefits to society they provide. Their management
and protection are guided by traditional communal governance as well (Moeliono
et al. 2015).
Traditional methods of community-based biodiversity conservation which can
maintain or build biodiversity islands may also include agrarian and nomadic
cultures. The Yanesha at the headwaters of the Amazon basin in Central Peru have
hunted, gathered, and farmed sustainably for thousands of years with little evidence
of biodiversity degradation (Salick 1989). As deforestation and unsustainable devel-
opment expand, community-led traditional landscape management provides the
opportunity for biodiversity islands to emerge. In some instances, levels of biodi-
versity within traditional communities can increase through agricultural production
as documented throughout Amazonia (Erikson 2008). Community-fostered swidden
systems from Southwest China to the Ecuadorian Amazon to Madagascar can be
managed in accordance with biodiversity enhancing outcomes (Rerkasem et al.
2009; Xu et al. 2009). In other examples, such as in the Southwestern-United States,
species composition of agricultural plots is actively managed to embrace diversity
and complexity in both production species and management of surrounding habitat
(Nabhan 2000).
500 B. Levin

Moreover, community-led biodiversity islands exist in highly varied geographies

catalyzed by diverse organizational structures. Rural community action from
Zapatistas in southern Mexico to community restoration projects in Northern Ethi-
opia to the newly developing global concept of Ecosystem Restoration Camps, all
seek to enhance biodiversity outcomes in rural settings through community efforts
([Link] Sigman 2022). Non-profit organiza-
tions such as Eco-agriculture Partners and The Forest Dialogue also aim to help
engage community action for biodiversity in rural settings through methods of
community engagement within integrated landscape management (https://
[Link]/, [Link] In suburban settings
homegardens, community garden projects, and ecovillages act as biodiversity
islands in the fragmented landscape (Negret et al. 2022; Toensmeier 2022). Enter-
prises such as Permaculture Artisans, a California-based landscaping company,
develop properties into productive gardens, creating biodiversity islands on the
suburban landscape ([Link]). In the urban context, the
rise of urban forestry, urban community gardens, and educational centers which
support these ends continue to grow in popularity (Soler et al. 2022). These
examples in varied scales of population density, diverse cultures, and various
locations, provide excellent lessons for community-led action advancing the crea-
tion, establishment, and maintenance of biodiversity islands.

19.7 Conclusion

Community-led action towards the development of biodiversity islands demands

looking beyond just the human factors of place to include all living beings and
biophysical components that make up the community. Motivations for biodiversity
conservation are diverse across cultures. In many instances, historical ecology and
indigenous and traditional philosophies of the region may empower decision making
and cultural practices towards the development and preservation of biodiversity
islands. When empowered, communities across a range of contexts may effectively
mobilize towards protecting, creating, and expanding islands of biodiversity. Land
access and long-term decision-making authority become essential in the longevity of
such pockets of biodiversity. Various legal and financial strategies such as conser-
vation easements, community land trusts, and novel methods of crowdsourced
funding can further aid in successful outcomes. Once biodiversity islands emerge
within a landscape, egalitarian management and governance approaches have
proven effective through time. Community advocacy may further bolster related
positive outcomes. By employing these tactics, community-led biodiversity islands
can rapidly multiply and continue to span the globe, as protected pockets of
biodiversity.
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 501

References

Alexander L, Hess GR (2012) Land trust evaluation of progress toward conservation goals. Conserv
Biol 26(1):7–12
Badgley C (2018) Biodiversity and sustainable agriculture. Promoting biodiversity in food
systems. In: Hawkins IW (ed) Promoting biodiversity in food systems. CRC Press, Boca
Raton, pp 165–182
Baez Schon M, Woods CL, Cardelús CL (2022) Sacred church forests in northern Ethiopia:
biodiversity and cultural Islands. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Springer, Cham, pp 531–549
Baldwin RF, Beazley KF (2019) Emerging paradigms for biodiversity and protected areas. Land
8(3). [Link]
Balée W (1993) Indigenous transformation of Amazonian forests: an example from Maranhão,
Brazil. L’Homme 33:231–254
Balée W (1998) Advances in historical ecology. Columbia University Press, New York. 429pp
Beller EE, McClenachan L, Zavaleta ES, Larsen LG (2020) Past forward: recommendations from
historical ecology for ecosystem management. Global Ecol Conserv 21. [Link]
[Link].2019.e00836
Berkes F (2004) Rethinking community-based conservation. Conserv Biol 18(3):621–630
Berkes F (2017) Environmental governance for the Anthropocene? Social-ecological systems,
resilience, and collaborative learning. Sustainability 9(7):1232. [Link]
su9071232
Berkes F, Folke C, Gadgil M (1994) Traditional ecological knowledge, biodiversity, resilience and
sustainability. In: Perrings CA, Mäler KG, Folke C, Holling CS, Jansson BO (eds) Biodiversity
conservation. Springer, Switzerland AG, pp 269–287
Bhagwat SA, Rutte C (2006) Sacred groves: potential for biodiversity management. Front Ecol
Environ 4(10):519–524
Bookchin M (1982) The ecology of freedom. Cheshire Books, Palo Alto, p 480
Carey AB, Lippke BR, Sessions J (1999) Intentional systems management: managing forests for
biodiversity. J Sustain For 9(3–4):83–125
Carson R (1962) Silent Spring. Houghton Mifflin, Boston
Castro-Arce K, Vanclay F (2020) Transformative social innovation for sustainable rural develop-
ment: an analytical framework to assist community-based initiatives. J Rural Studies 74:45–54
Ceperley N, Montagnini F, Natta A (2010) Significance of sacred sites for riparian forest conser-
vation in Central Benin. Bois Forêts Tropiques 303:5–23
Chung MG, Dietz T, Liu J (2018) Global relationships between biodiversity and nature-based
tourism in protected areas. Ecosyst Serv 34:11–23
Clarke LW, Jenerette G (2015) Biodiversity and direct ecosystem service regulation in the com-
munity gardens of Los Angeles, CA. Landsc Ecol 30(4):637–653
Colwell RK, Coddington JA (1994) Estimating terrestrial biodiversity through extrapolation. Philos
Trans R Soc Lond B Biol Sci 345(1311):101–118
Crosby AW (2004) Ecological imperialism: the biological expansion of Europe, 900–1900. Cam-
bridge University Press, Cambridge
de Boef WS, Subedi A, Peroni N, Thijssen M, O’Keeffe E (eds) (2013) Community biodiversity
management: promoting resilience and the conservation of plant genetic resources. Routledge,
Abington
Di Pietro F, Mehdi L, Brun M, Tanguay C (2018) Community gardens and their potential for urban
biodiversity. In: Gladtron S, Granchamp L (eds) The urban Garden City. Springer, New York,
pp 131–151
Dudley N, Alexander S (2017) Agriculture and biodiversity: a review. Biodiversity 18(2–3):45–49
Dunson WA, Travis J (1991) The role of abiotic factors in community organization. Amer Nat
138(5):1067–1091
502 B. Levin

Erickson CL (2008) Amazonia: the historical ecology of a domesticated landscape. In: Isbell W
(ed) Silverman H. The Handbook of South American Archaeology Springer, New York, pp
157–183
Erisman JW, van Eekeren N, de Wit J, Koopmans C, Cuijpers W, Oerlemans N, Koks BJ (2016)
Agriculture and biodiversity: a better balance benefits both. Agric & Food 1(2):157–174
Ferguson RS, Lovell ST (2014) Permaculture for agroecology: design, movement, practice, and
worldview. A review. Agron Sustain Dev 34(2):251–274
Gadgil M, Berkes F, Folke C (1993) Indigenous knowledge for biodiversity conservation.
Ambio:151–156
Gallo-Cajiao E, Archibald C, Friedman R, Steven R, Fuller RA, Game ET, Morrison TH, Ritchie
EG (2018) Crowdfunding biodiversity conservation. Conserv Biol 32(6):1426–1435
Galluzzi G, Eyzaguirre P, Negri V (2010) Home gardens: neglected hotspots of agro-biodiversity
and cultural diversity. Biodivers Conserv 19(13):3635–3654
Garnett ST, Burgess ND, Fa JE, Fernández-Llamazares Á, Molnár Z, Robinson CJ, Watson JE,
Zander KK, Austin B, Brondizio ES, Collier NF (2018) A spatial overview of the global
importance of indigenous lands for conservation. Nat Sustain 1(7):369
Gilbert J (2016) Indigenous peoples’ land rights. International Law Brill Publishers, Leiden
Goddard MA, Dougill AJ, Benton TG (2010) Scaling up from gardens: biodiversity conservation in
urban environments. Trends Ecol Evol 25(2):90–98
Gómez-Pompa A, Flores JS, Fernández MA (1990) The sacred cacao groves of the Maya. Lat Amer
Antiq 1(3):247–257
Gössling S (1999) Ecotourism: a means to safeguard biodiversity and ecosystem functions? Ecol
Econ 29(2):303–320
Hardin G (1968) The tragedy of the commons. Science 162(3859):1243–1248
Harvey G (2005) Animism: respecting the living world. Wakefield Press, Adelaide
Hawksworth DL (ed) (1995) Biodiversity: measurement and estimation. Chapman and Hall,
London
Hawksworth DL (ed) (2010a) Management and the conservation of biodiversity. Springer,
New York
Hawksworth (ed) (2010b) Methods and practice in biodiversity conservation. Springer, New York
Hawksworth DL (2012) Books on biodiversity and conservation. Biodivers Conserv 21(14):
3745–3769
Hay-Edie T, Halverson E (2006) Community action to conserve biodiversity: linking biodiversity
conservation with poverty reduction. GEF Small Grants Programme, UNDP, New York. sgp.
[Link]
Holmes G, Cavanagh CJ (2016) A review of the social impacts of neoliberal conservation:
formations, inequalities, contestations. Geoforum 75:199–209
Hunter D, Guarino L, Spillane C, McKeown PC (eds) (2017) Routledge handbook of agricultural
biodiversity. Routledge, Abington
Inglis J (ed) (1993) Traditional ecological knowledge: concepts and cases. IDRC, Ottawa
Jarrett C, Cummins I, Logan-Hines E (2017) Adapting indigenous agroforestry systems for
integrative landscape management and sustainable supply chain development in Napo,
Ecuador. In: Integrating landscapes: agroforestry for biodiversity conservation and food sover-
eignty. Springer, Cham, pp 283–309
Kellert SR (1997) The value of life: biological diversity and human society. Island Press,
Washington DC
Kellert SR (2018) Nature by design: the practice of biophilic design. Yale University Press, New
Haven
Khan ML, Khumbongmayum AD, Tripathi RS (2008) The sacred groves and their significance in
conserving biodiversity: an overview. Int J Ecol Environ Sci 34(3):277–291
Langton M, Rhea ZM (2005) Traditional indigenous biodiversity-related knowledge. Aust Acad &
Res Libr 36(2):45–69
19 How Community-Led Action Can Advance the Development of Biodiversity Islands 503

Leopold A (1949) A Sand County almanac, and sketches here and there. Oxford University Press,
New York
Leventon J, Schaal T, Velten S, Loos J, Fischer J, Newig J (2019) Landscape-scale biodiversity
governance: scenarios for reshaping spaces of governance. Environ Policy Gov 29(3):170–184
Lovelock J (1979) Gaia: a new look at life on earth. Oxford University Press, Oxford
Méndez V, Gliessman SR, Gilbert GS (2007) Tree biodiversity in farmer cooperatives of a shade
coffee landscape in western El Salvador. Agric Ecosyst Environ 119(1–2):145–159
Moeliono M, Mulyana A, Adnan H, Manalu P, Yuliani L (2015) Village forests (hutan desa):
empowerment, business or burden? World Agroforestry Centre-ICRAF, Southeast Asia
Regional Office, Bogor
Mollison B (1988) Permaculture: a Designer’s manual. Tagari Publications, Sisters Creek
Montagnini F, del Fierro S (2022) Functions of agroforestry systems as biodiversity islands in
productive landscapes. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Springer, Cham, pp 89–116
Montagnini F, Levin B, Berg KE (2022) Biodiversity Islands. Strategies for conservation in human-
dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies for conservation
in human dominated environments. Springer, Cham, pp 1–35
Nabhan GP (2000) Native American management and conservation of biodiversity in the Sonoran
Desert bioregion. In: Minnis PE, Elisens WJ (eds) Biodiversity and native America. University
of Oklahoma Press, Norman, pp 29–44
Nabhan G, Hodgson W, Fellows F (1989) A meager living on lava and sand? Hia ced O’odham
food resources and habitat diversity in oral and documentary histories. J Southwest:508–533
Negret HRC, Negret R, Montes-Londoño I (2022) Residential garden Design for Urban Bio-
diversity Conservation: Experience from Panama City, Panama. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments.
Springer, Cham, pp 387–417
Ostrom E (1990) Governing the commons: the evolution of institutions for collective action.
Cambridge University Press, Cambridge
Otto J, Zerner C, Robinson J, Donovan R, Lavelle M, Villarreal R, Salafsky N, Alcorn J,
Seymour F, Kleyneyer C, Pearl M (2013) Natural connections: perspectives in community-
based conservation. Island Press, Washington D.C
Peters CM (2018) Managing the wild: stories of people and plants and tropical forests. Yale
University Press, New Haven
Pretty J, Smith D (2004) Social capital in biodiversity conservation and management. Conserv Biol
18(3):631–638
Pretty J, Adams B, Berkes F, de Athayde S, Dudley N, Hunn E, Maffi L, Milton K, Rapport D,
Robbins P, Sterling E, Stolton S, Tsing A, Vintinnerk E, Pilgrim S (2009) The intersection of
biological diversity and cultural diversity: towards intersection. Conserv Soc 7:100–112
Prieto Méndez JM (2013) Derechos de la naturaleza. Fundamento, contenido y exigibilidad
jurisdiccional. Corte Constitucional del Ecuador, CEDEC, Quito. 280pp
Raymond H (2007) The ecologically noble savage debate. Ann Rev Anthropol 36:177–190
Rerkasem K, Lawrence D, Padoch C, Schmidt-Vogt D, Ziegler AD, Bruun TB (2009) Conse-
quences of swidden transitions for crop and fallow biodiversity in Southeast Asia. Hum Ecol
37(3):347–360
Rissman AR, Lozier L, Comendant T, Kareiva P, Kiesecker JM, Shaw MR, Merenlender AM
(2007) Conservation easements: biodiversity protection and private use. Conserv Biol 21(3):
709–718
Robertson BJ (2015) Holacracy: the new management system for a rapidly changing world. Henry
Holt and Company, New York
Rocha P, Niella F, Keller H, Montagnini F, Metzel R, Eibl B, Kornel J, Romero F, López L,
Araujo J, Barquinero J (2017) Ecological indigenous (EIK) and scientific (ESK) knowledge
integration as tool for sustainable development in indigenous communities. Experience in
Misiones, Argentina. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity
conservation and food sovereignty, advances in agroforestry 12. Springer, Cham, pp 235–260
504 B. Levin

Salick J (1989) Ecological basis of Amuesha agriculture, Peruvian upper Amazon. Adv
Econ Bot:189–212
Salick J (2012) Indigenous peoples conserving, managing, and creating biodiversity. In: Harlan JR,
Gepts P, Famula TR, Bettinger RL, Brush SB, Damania AB, McGuire PE, Qualset CO (eds)
Biodiversity in agriculture: domestication, evolution, and sustainability. Cambridge University
Press, Cambridge, pp 426–445
Sigman E (2022) Safeguarding the benefits of biodiversity islands in northern Ethiopia in the midst
of political change. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Springer, Cham, pp 647–674
Sobrevila C (2008) The role of indigenous peoples in biodiversity conservation World Bank,
Washington D.C.
Soler R, Benítez J, Solá F, Lencinas MV (2022) Biodiversity islands at the world’s
southernmost city: plant, bird and insect conservation in urban forests and peatlands of Ushuaia,
Argentina. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Springer, Cham, pp 419–437
Springer J (2009) Addressing the social impacts of conservation. Conserv Soc 7(1):26–29
Swimme B, Tucker ME (2011) Journey of the universe. Yale University Press, New Haven
Toensmeier E (2022) Paradise lot: a temperate urban multistrata agroforestry island of
biodiversity. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Springer, Cham, pp 439–459
Van Weelie D, Wals A (2002) Making biodiversity meaningful through environmental education.
Int JSci science Edu 24(11):1143–1156
Veteto JR, Lockyer J (2008) Environmental anthropology engaging permaculture: moving theory
and practice toward sustainability. Cult Agric 30(1–2):47–58
Wilson EO (1984) Biophilia. Harvard University Press, Cambridge MA
Wilson R (2011) Documenting and protecting biodiversity on land trust projects. Land Trust
Alliance, Washington DC
Xu J, Lebel L, Sturgeon J (2009) Functional links between biodiversity, livelihoods, and culture in a
Hani swidden landscape in Southwest China. Ecol Soc 14(2). [Link]
02916-140220
Chapter 20
Priorities, Perspectives, and Use of a
Community Forest by Surrounding
Residents in Mayagüez, Puerto Rico:
Protecting the Forest for Its Services

Gabriela M. Morales-Nieves

Abstract Community forests are important for protection of lands, reduction of

deforestation, conservation of biodiversity, and carbon sequestration while provid-
ing socioeconomic benefits to those living around them. In the insular Caribbean,
community forests do not seem to be used widely. Given that it is a region with high
biodiversity and endemism, while being also the sub region in Latin America and the
Caribbean with highest population density, protection of lands in fragmented land-
scapes is especially important. To expand the adoption of community forests by
communities, it is important to understand the interest of the residents in managing
them and the opportunity they see in community forests. This research used inter-
views with people living around a community forest in Mayagüez, Puerto Rico to
understand local perceptions of and interactions with the forest. Interviewees were
asked which activities they would be interested in participating in the forest,
perceived ecosystem services and what should be the management priorities and
services the forest should provide to the community. The most important ecosystem
service for residents was air quality, followed by recreation. Motives that discour-
aged the use of the forest by residents were the presence of non-desirable species and
lack of knowledge regarding the functioning of the forest. The forest is currently a
meeting space for community activities such as celebrations, workshops and the
selling of food and other products. Although conservation of biodiversity is just one
motivation for managing the forest as a community, the values that people assign to
the forest contribute to its preservation as a biodiversity island within a rural-urban
landscape.

Keywords Caribbean · Community forestry · Conservation · Ecosystem services ·

Human perceptions · Participation · Urban expansion

G. M. Morales-Nieves (*)
Yale University, School of the Environment, The Forest School, New Haven, CT, USA
e-mail: [Link]-nieves@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 505
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
506 G. M. Morales-Nieves

20.1 Introduction

Community forestry can be broadly defined as government-approved or recognized

initiatives that have the objective of increasing the role of surrounding communities
in governing and managing forest resources to provide social and economic benefits
while promoting the sustainable management of said forest lands (Bowler et al.
2012; Gilmour 2016). Community forests have been considered as an essential
component for the conservation of lands and reducing deforestation rates (Porter-
Bolland et al. 2012; Lambrick et al. 2014), maintaining biodiversity (Souza et al.
2016), and sequestering carbon (Wood et al. 2019), while providing different
socioeconomic benefits to surrounding residents (Bridhikitti and Khadka 2019).
However, mixed results have been found in cases when biodiversity conservation
goals may conflict with interest in using the forest resources. There are often some
trade-offs between activities that enhance biodiversity conservation, like controlling
wildlife hunting, and those that could have negative impacts on biodiversity and
ecosystem functions, like for example, removing or selecting certain plant species
and replacing for others of commercial value (Shrestha and Shrestha 2010;
Rasolofoson et al. 2015). Even though the term “community forestry” is broad and
used in varying contexts, most of them coincide in that they “aim to protect forests
and benefit local livelihoods” (Rasolofoson et al. 2015).
In the Caribbean islands region, literature on participatory management of
resources seems to focus more on those in coastal ecosystems (Geoghegan and
Renard 2002; McConney and Pena 2012; Smith 2012; Dalton et al. 2012). No
scientific literature has been found on community forests in the Caribbean islands,
but there is some gray literature and reports on community-based forest manage-
ment, such as a report by the Caribbean Natural Resources Institute (CANARI) and
the Food and Agriculture Organization (FAO) (2012). As in other parts of the world,
with climate change, the Caribbean islands are expected to experience changes in
temperatures, precipitation patterns (Pulwarty et al. 2010) and increased intensity of
hurricanes (Holland and Bruyère 2014; Knutson et al. 2015). This is especially
important given their small physical dimensions, their high ranking in vulnerability
to global disasters, and their high dependency on natural resources for the tourism
industry, among other aspects (Pulwarty et al. 2010).
The Caribbean region is also a world biodiversity hotspot and it is important for
biodiversity conservation due to its high levels of endemism and impending threats
related to increases in human populations and encroachment on forests and other
natural areas (Myers et al. 2000; Anadon-Irizarry et al. 2012). The Caribbean Islands
comprise the most densely populated sub region in Latin America and the Carib-
bean, given 2017 data obtained from the World Bank (DataBank | The World Bank)
found in Appendix A. Nine of ten of the most densely populated countries within
Latin America and the Caribbean are from the insular Caribbean. Therefore,
maintaining biodiversity islands in a human-dominated landscape is especially
important for the conservation of high biodiversity in a region with limited territorial
extent.
20 Priorities, Perspectives, and Use of a Community Forest by. . . 507

To expand the adoption of community forests as a tool for ecosystem and

biodiversity protection and also for economic development in the Caribbean Islands,
it is important to understand the interests of communities surrounding forested lands
in managing them and also the opportunities seen in them. This chapter examines the
use of a community-managed forest in Mayagüez, Puerto Rico as a case study. This
research aimed to understand how the surrounding community used or would use the
forest for, what ecosystem services or benefits they felt the forest provided to the
community, and what the management priorities for the forest should be. It uses
these findings to discuss the potential of community forests for conservation and
offers recommendations for management in other similar contexts in the region and
elsewhere.

20.1.1 Recent Trends in Land Use History of Puerto Rico

and the Caribbean

[Link] Puerto Rico

To understand the results of this case study, it is necessary to contextualize the land
use history of Puerto Rico and the Caribbean. By the nineteenth century, with
consistent increases in the population in Puerto Rico post colonization, agriculture,
comprised mostly of sugarcane, coffee, plantain, cotton and rice, had spread
throughout. In addition, by that time pasturelands covered about 55% of the land
(Grau et al. 2003). By the 1930s, agriculture and cattle production represented about
45% of the gross national product (GNP). In the late 1940s, “post war efforts to
promote industrialization shifted the economy from agriculture to light industry,”
with agriculture comprising less than 5% of the GNP by 1980 (Grau et al. 2003).
Since agricultural land abandonment due to this transition in economy from mainly
agricultural to industry and service-based in current times, Puerto Rico gained from
5–6% forest cover in the 1940s to 50–57% by 2009 (Álvarez-Berríos et al. 2013;
Brandeis and Turner 2013), remaining in a steady state up to 2014 (Marcano-Vega
2017).
During the first decade of the 2000s, urban cover also increased. Urban expansion
was the major cause for deforestation in Puerto Rico during this time and the
previous decade, with the loss of agricultural lands as the major land use change,
however with an overall increase in woody vegetation recovery (Álvarez-Berríos
et al. 2013). Although urbanization contributed to woody regrowth in the past, recent
patterns of urban expansion are the major driver of deforestation and forest frag-
mentation in the island. Urbanization seems to compete and replace the same lands
as agriculture in Puerto Rico, since the appropriate characteristics for urbanization
are very similar to those that are needed for intensive agriculture (e.g. lower eleva-
tions, flat topography, proximity to roads), with 60% of total development by 2003
occurring where the most productive lands for agriculture are located (López et al.
2001; Grau et al. 2003; Martinuzzi et al. 2007).
508 G. M. Morales-Nieves

By 2016, 16.1% of Puerto Rico’s land cover was under some form of protection,
whether under state, federal (U.S.A.), private or non-profit organization supervision.
Of these protected lands, 27% are considered forest lands and 56% are forested
wetlands (Castro-Prieto et al. 2019).

[Link] Caribbean Region

By 2012 in the Greater Antilles forest cover ranged from less than 4% for Haiti, to
more than 64% for Puerto Rico (González and Scalley 2016). In the Lesser Antilles,
most islands range between 36–61% forest cover, with Barbados having 19–31%
and St. Lucia and St. Vincent and the Grenadines having more than 64%. Similarly,
north of the Lesser Antilles, Bahamas and Turks and Caicos range mostly around
36–61% forest cover.
Another study has found that in St. Kitts, Nevis, Grenada and Barbados, forest
cover has increased from 50% to 95% after the 1950s (Helmer et al. 2008). At the
same time, cultivated lands have decreased by 59–99%, and developed land area has
increased, especially at lower elevations, reflecting a very similar land use change
pattern as in Puerto Rico, due to similar reasons. This study also found that these
islands, though their humid forest, dry forests and mangroves cover has increased,
are under very little protection and therefore are high conservation priorities.
Jamaica and Trinidad and Tobago were found to have net forest loss between
2001 and 2010 (Aide et al. 2013). The same authors also found that Haiti had an
increase in woody vegetation during the same time and that overall, the Caribbean
had experienced a net gain in woody vegetation between 2001 and 2010.

20.1.2 Community Forests

Motivations behind forest conservation by a community can vary from considering

that forests are sacred sites (Ceperley et al. 2010), an opportunity for the economic
development of a community, or a vehicle for increased recognition of autonomy,
and a sense of conservation commitment (Moeliono et al. 2015; Ruiz-Mallén et al.
2015). After centuries of colonization, and in response to failure of centralized
government management of forested lands, during the 1970s and 1980s many
countries in the global South and the New World re-adopted or supported small-
holder and community-based forestry as it became increasingly recognized in
international arenas (Gilmour 2016).
In the same report, Gilmour explains that community-based forestry regimes can
range from participatory conservation, where stakeholders do not own the land but
participate in some form of decision-making regarding a forest but with little
authority, to private ownership by a community. In the private ownership regime
20 Priorities, Perspectives, and Use of a Community Forest by. . . 509

form, most rights to access, management and even compensation belong to the
households, groups or communities that own the land. Finally, Gilmour (2016)
provides estimates of forest lands under some sort of community-based forestry
regime globally, which range between 200 and 505 million hectares.

20.1.3 Community Forests in Latin America

and the Caribbean

Especially during the 1980s and 1990s, many efforts from indigenous communities
to get their territorial autonomy recognized resulted in the establishment and legal
recognition of community-forests throughout Latin America (Ruiz-Mallén et al.
2015). México is probably the country with the most internationally recognized
community forestry and history, with around 80% of the country’s forests under
some type of legal jurisdiction by local communities (Gilmour 2016). More recent
efforts include the Programa de Manejo Florestal Comunitário e Familiar,
established in 2009 in Brazil.
Gilmour (2016) also reported that around 272 million hectares are under a
community-based forestry regime in Latin America, although none of the countries
listed are in the Caribbean islands. In collaboration with the FAO, the Caribbean
Natural Resources Institute developed a synthesis of community-based forestry in
the Caribbean (CANARI 2012). In this report, they use 14 case studies in countries
or territories like Haiti, Cuba, St. Kitts and Nevis and Grenada. Many of these case
studies seem to be in the form of a collaborative regime, where the community has
some sort of participation in the management of forests, some type of extraction
rights on state-owned forests, or permission of use within a state-owned forest, with
some groups requesting autonomy or management control, or on privately-
owned land.
All projects include livelihood improvement and capacity-building as primary or
secondary objectives. Primary objectives range from watershed rehabilitation to
plantation timber production. In Puerto Rico, the Department of Natural Resources
has collaborative agreements with different organizations to perform activities
related to “restoration, education, ecotourism, agroecology and scientific research”
in different protected areas that the state owns (Acuerdos Colaborativos 2015). A
very emblematic case of a co-managed forest is Bosque del Pueblo in Adjuntas,
Puerto Rico, where the community organization Casa Pueblo successfully halted a
mining project in the mountainous interior of the main island and proposed and
established a protected forest instead (Massol González et al. 2006).
510 G. M. Morales-Nieves

20.1.4 Río Hondo Community Forest: Establishment

and History

The Río Hondo Community Forest (RHCF) is located in Barrio Río Hondo of the
municipality of Mayagüez, in western Puerto Rico (18
.174455 N,
67
.132376 W). The Río Hondo Community Forest is 27.5 hectares in size. In
its recently published land management plan, it is described as a community forest in
the rural-urban interface. The land where the forest is now located had been used for
growing sugar cane decades prior and later as pasture land for cattle and horses. By
1975, the Barrio Río Hondo land area only had 4% forest cover. By 2010, 95% of the
land was covered by secondary forest that is no more than 40 years old after
agricultural activity was abandoned. In the Barrio Río Hondo area, urban expansion
has increased even though population has declined between 2000 and 2010 (Castro-
Prieto et al. 2017).
This land was privately owned and was intended to be turned into a housing
project in 2007. Upon learning about this, different community members organized
to put a halt to said project. The community started renting the land from the
landowners and organizing activities on the land. They originally had intentions of
building a theater-museum and other infrastructure as well as dedicating land to
serve as demonstration sites of different historical agricultural practices in Puerto
Rico. However, that idea later turned into keeping the land as forest. Its main
governing body is the Río Hondo Community Forest Board, which is composed
of volunteers, mainly residents of the community, who have an interest in
maintaining the forest, organizing activities and carrying out all proposed projects
within their land management plan.
The RHCF land management plan was created under a proposed co-management
agreement between the RHCF Community Board, the Mayagüez Municipality and
the United States Forest Service. In 2018, the Mayagüez Municipality and the United
States Forest Service, under the Community Forest and Open Space Conservation
Program (How the Community Forest Program Works | US Forest Service), pro-
vided the funding to formally purchase the entire property (Fig. 20.1). The official
title of the land belongs to the Mayagüez Municipality, protecting the land into
perpetuity and ensuring that all activities are in line with the statutes established by
the Community Forest and Open Space Conservation Program (Figueroa Vázquez,
Program Manager for State and Private Forestry for Puerto Rico and the US Virgin
Islands, US Forest Service, pers. comm.).

[Link] Biodiversity in the Río Hondo Community Forest

As described in the land management plan, a total of 23 tree species have been
identified in the RHCF (Rodríguez Candelaria et al. Working Document), eight of
which are considered introduced and the rest native (Appendix B). Different stands
within the forest have different dominant tree species, like Albizia procera
20 Priorities, Perspectives, and Use of a Community Forest by. . . 511

Fig. 20.1 Walking along a trail that runs through an early-successional part of the property
bordering households with members of the Community Forest Board and my research assistant.
(Photo: Gabriela Morales-Nieves)

(non-native), Spathodea campanulata (non-native), Senna siamea (non-native),

Cupania americana (native) and Guarea guidonia (native), with a distinct amount
of native tree species currently in the understory of all these stands.
Thirty-four bird species have been identified in the forest, nine of which are
endemic to Puerto Rico. In addition, 4 reptile species and 3 amphibians (one endemic)
have also been identified in the Río Hondo Community Forest (Appendix C).

20.2 Methods

I first identified all houses that directly surrounded the forest (134 total) and selected
a set of 70 houses randomly using ArcGIS software, aiming for 50 random houses,
while making space for those that were possibly unoccupied or non-residential
structures (Fig. 20.2). Either through previous agreement or by visiting, I conducted
semi-structured interviews to record residents’ interests, desires and values for the
forest. I gathered basic household data on the size, ages and other social demo-
graphics of the household. I asked both closed and open-ended questions. Household
residents were asked about different benefits they felt they received from the
512 G. M. Morales-Nieves

Fig. 20.2 Aerial image of the Río Hondo Community Forest. The green dots represent all potential
households that directly border the forest. (Image: ArcMap)

adjacent forest, what opportunities they see in this forest, what activities they would
like to be a part of, and what crops and tree species they would choose to plant in the
forest. Finally, I used the six goals set in the recently published land management
plan for the forest and asked residents to rank these goals from most important to
least important for them. The complete goals can be found in Appendix D.
Using MAXQDA software, I categorized the responses to the questions
(e.g. positive vs. negative perceptions) to be analyzed quantitatively, following
Garen et al. (2009). After coding the responses, I analyzed the proportion of
residents that touched on a specific topic in an individual question. Other relevant
comments made outside the formal survey were recorded and used to supplement or
support formal responses to questions in the survey.
I calculated the means for each goal that I asked residents to rank, using six as the
most important and one as the least important goal. I used MINITAB software to
perform a one-way ANOVA to analyze if there was a significant difference in how
interviewees organized the goals in order of importance; in other words, if there was
a common pattern among interviewees in the order of importance of the goals. The
ANOVA was followed by a Tukey Pairwise Comparisons test to determine which
means were significantly different from which.
20 Priorities, Perspectives, and Use of a Community Forest by. . . 513

20.3 Results

Out of the 70 houses I reached out to, I performed interviews in 37 households. Some
of the 70 houses were not occupied, were commercial buildings rather than homes,
did not have residents there at the moment, or had residents that did not want to be
interviewed. Twenty-four of the interviewees identified as female, with the rest
(13) identifying as male. Out of the 37 interviewees, 34 of them owned the house
they lived in. The average age of interviewees was 59 years (16, s.d.). The average
age time of residence in the household was 32 years ( 21, s.d.). The amount of
people living in the household ranged between 1 and 6 people. Including all
members in each household surveyed, most people living around the community
forest are above 40 years old (Fig. 20.3).

20.3.1 Participation and Visits to the Río Hondo Community

Forest

Of the 37 residents that were interviewed, 19 people had visited the forest and 18 had
not. However, two of the 18 that had not visited the forest mentioned that another
member of the household had visited or participated in activities hosted at the forest.
Besides mentioning whether they had visited or not, at least 2 other interviewees
expressed that they had spent time in the land before it was declared a protected area,

Fig. 20.3 Age distribution by gender of residents in households surveyed around the Río Hondo
Community Forest
514 G. M. Morales-Nieves

since they had lived in the community for many years. There was no difference in
responses between visiting or not having visited the forest when analyzed by gender.
In terms of what was the purpose of their visits to the forest, 17 responded it was
to participate in specific activities. These responses were grouped into common
categories in Table 20.1.
The most notable common activities were social gatherings and parties. These
happen mainly in a common, more open area at the entrance to the property. Many
residents specifically mentioned a community Christmas party that is thrown every
year and added that they enjoyed it. For example, one resident mentioned: “When I
most enjoy [going to the forest] is when they bring music and people have fun.” It
should be noted that some of these activities happen at the same time. Parties on
occasion also include interpretative hikes in the trails, for example.

20.3.2 Interest in Activities and Other Recommendations

Seventeen interviewees recommended activities beyond those mentioned among the

ones that currently are organized in the forest, most of them related to recreation
(Table 20.2). Within this topic, 3 people consistently mentioned the development of
trails specifically for mountain biking: “Well I am a cyclist and I think it would be
good if they would incentivize cycling in trails”.
Within recreation, residents also recommended developing more infrastructure to
promote it, like gazebos and resting areas. Residents also recommended social
activities like bingos and barbeques. Four of the 5 educational activities
recommended were meant for children: “[I would recommend] a summer camp
where they show children the value of crops and where it is educational, subsidized
by the [Mayagüez] municipality to promote agriculture”. Finally, residents
recommended to perform different commercial activities, mainly selling forest
products like trees and added value products from harvested crops.

20.3.3 Benefits and Ecosystem Services

The most common theme that arose in conversations with interviewees was air
quality. Even more so, 5 people specifically referred to the forest as a “lung” to
the community: “That forest is a lung for this area. If they didn’t protect it, they
(original landowners) were going to cut it down to build houses and put concrete.”
When asked what benefits they felt the forest provided to them or the community,
residents, including some that had not actually visited the forest, mentioned a
variety, including benefits they may not feel they receive currently, but more so as
potential in the future: “It could be a source of employment for people that are
unemployed.”
 20

Table 20.1 Categories of responses to the open question regarding for what purpose residents have visited the Río Hondo Community Forest. Each interviewee
might have mentioned more than one category
Social gatherings and Purchasing of Informational and Planning Arts and
parties products Meetings Educational Volunteering Recreation workshops
8 7 4 3 3 3 2
Priorities, Perspectives, and Use of a Community Forest by. . .
515
516 G. M. Morales-Nieves

Table 20.2 Interest in activities that the Río Hondo Community currently hosts. Each interviewee
might have mentioned more than one category.
Sales and Recreational Be part of
exhibition Educational activities the
Related to of crops walks or (e.g. hiking, Related community
agriculture and crafts tours camping) to crafts forest board Volunteering
30 28 26 25 17 14 10

Others, though less frequently, mentioned the forest as serving aesthetic purposes
(“The forest is beautiful”) or as a source of tranquility (“Air purification, for the
birds. . . peacefulness. That land serves for a lot of things”). Finally, 6 residents did
not perceive any benefit or service that the forest provided to them or the community,
the majority of them because they had not visited or were unfamiliar with the forest.
The different categories in which people mentioned perceived benefits or eco-
system services provided by the forest are summarized in Table 20.3. There is
overlap between perceived benefits and observed positive impressions of the forest.
However, there were certain comments that expressed general enthusiasm about the
forest, like the following:
“[I would be a] promoter so that the forest always serves a purpose.”
“[. . .] but the forest is wonderful.”
“[. . .] and I love that in there.”

20.3.4 Negative Impressions and Concerns

Within the different questions and additional comments, concerns or conflicting

desires regarding the forest’s purpose were also captured. There was also some
misinformation in the knowledge people had of the forest. For example, there was
not much clarity around where funding for operations came from, who owned the
forest and therefore, what activities were allowed or not.
People expressed concerns about certain activities or events being charged to the
public and lack of participation of the community in events or meetings. Two main
topics surfaced, that will be described in more detail below, which were first, about
non-native species or those that are perceived as a nuisance and second, about the
concept of cleanliness of the forest.

[Link] Perspectives on Non-native and Non-beneficial Plants Species

In conversation with residents, many of them associated the presence of the forest as
housing different species of both plants and wildlife that they consider a nuisance for
different reasons. For example, there is a vine that grows in the area, commonly
called “picapica” (Mucuna pruriens, considered “probably native” which has hairs
 20

Table 20.3 General categories of perceived benefits or ecosystem services provided by the Río Hondo Community Forest. Interviewees might have mentioned
more than one category
Air Recreation and Don’t know or Food and medicine Wildlife and Aesthetics and Physical
quality tourism Education perceive any source habitat relaxation Employment protection
11 9 6 6 6 5 2 2 1
Priorities, Perspectives, and Use of a Community Forest by. . .
517
518 G. M. Morales-Nieves

that line the seed pod and that upon contact or release with the wind can attach and
cause irritation and itchiness to the skin (Acevedo-Rodríguez 2005). Other species of
concern were iguanas (Iguana iguana, non-native) and red-tailed boas (Boa con-
strictor, non-native), especially the former as a threat for getting into backyards and
eating residents’ crops.
Finally, residents mentioned the tree species Albizia procera and Senna siamea
(both referred to as ‘Acacia’ and both non-native) as a species of little worth or as a
nuisance. “Around there what there is is ‘picapica’, iguanas and ‘acacia’, which is
not even useful for coal.” Three residents mentioned that trees bordering the forest
could potentially fall after a hurricane and could damage their houses or property due
to how close they are to the border with their land. “I have a small ranch and I would
like for [the people that manage the forest] to cut down some Acacia trees that may
fall on the ranch.”

[Link] “Cleanliness” of the Forest

Another common theme in discussion with residents was that of the forest needing
cleaning or fixing in order to fulfill its purpose:
“If they clean it and they set it well then maybe it can serve as a tourism attraction or if they
make more trails.”
“Well if they fix it, it will be a forest that serves for internal tourism. It is abandoned; it’s been
like that for more than 10 years.”

Residents also expressed concern about some areas that have overgrown or that
reduce mobility for walking. There was also a desire to reduce the presence of
‘picapica’ along the borders of the forest and concern about people dumping trash
along the road portion that runs through the forest: “Residents have requested them
to clean at least the borders [of the forest] to reduce the ‘picapica’, which is horrible.”

20.3.5 Ranking of Goals in the Management Plan

Three people refused to answer the section of ranking goals, two of them explaining
that they thought all goals were equally important. On the other hand, there is a
significant difference between what the remaining interviewees chose as the most
and the least important goals (Table 20.4). When analyzing the Tukey Pairwise
Comparisons, the first goal, which referred to maintaining the forest and planting
trees for the purpose of improving air quality and reducing pollution, was the highest
ranked and significantly different than 3 other goals: one that promoted participation
of different sectors in the forest (18c), one that aims to strengthen cultural and social
development of the community through the forest (18d), and one that aims to
increase resiliency of the community in the face of climate change and economic
crises (18e) (Table 20.5). The rest of the goals were not significantly different from
each other.
20 Priorities, Perspectives, and Use of a Community Forest by. . . 519

Table 20.4 Mean ranking for each of the six goals identified in the RHCF’s Land
Management Plan
Goal N Mean Standard deviation 95% CI
18a 34 4.50 1.52 (3.94, 5.06)
18b 34 3.68 1.99 (3.12, 4.24)
18c 34 3.35 1.74 (2.76, 3.89)
18d 34 3.32 1.60 (2.53, 3.65)
18e 34 3.09 1.32 (2.50, 3.62)
18f 34 3.06 1.70 (2.79, 3.91)
Pooled StDev ¼ 1.66
The means are significantly different (see Appendix D for description of goals)

Table 20.5 Tukey Pairwise Question N Mean Grouping

Comparisons of all six goals
18a 34 4.50 A
using a 95% confidence
interval 18b 34 3.68 A B
18f 34 3.35 A B
18c 34 3.32 B
18d 34 3.09 B
18e 34 3.06 B
The grouping column represents which goal is different from
which. Means that do not share a letter are significantly different.
In this case, goal 18a is different from goals 18c, 18d, 18e. Goals
18c, 18d and 18e, however, are not significantly different from
each other (see Appendix D for description of goals)

20.4 Discussion

The Río Hondo Community Forest is a small protected forest that has undergone a
similar pathway of land use history to many other parts of Puerto Rico and the rest of
the Caribbean Islands. However, it is the first forest to receive funding from the
United States Forest Service’s Community Forest and Open Space Conservation
Program in the Caribbean to purchase the property and maintain it as protected land,
co-managed by its surrounding community. The surrounding community is one
where many of their residents have been living for a large part of their lifetime.
Therefore, the community forest has the opportunity to build long-lasting relation-
ships with residents in a community that does not have fast turnover.
520 G. M. Morales-Nieves

20.4.1 Demographic Trends

The results showed that there is a large number of residents that are above 40 years of
age. This is perhaps a reflection of a more country-wide trend, where many Puerto
Ricans, especially due to an ongoing economic crisis and consequent migratory
wave since 2006 that has been accentuated after the passage of hurricane María, have
migrated from Puerto Rico. Even though data by age for these dates are not yet
available, between 2013 and 2017, 30.2% of the people that migrated were between
1–19 years old, 58.4% were 20–59 years old and 11.4% were above 60 years old
(Junta de Planificación de Puerto Rico 2019).
The Center for Puerto Rican Studies in a research brief explains that “since
Hurricane María, migration intensified, especially among families with children”
and that “this pattern has shifted the Island’s demographic structure”. Finally, it
reports that “population decline [. . .] was 129,848 in Puerto Rico immediately
following the storm” (Center for Puerto Rican Studies 2019). Age distribution is
important to know if the RHCF board wishes to develop activities and curriculum
catered to the age of most likely visitors.

20.4.2 Addressing Negative Perceptions

and Recommendations

Even though I did not ask interviewees why they had not visited the forest, some
mentioned the following reasons: fear of wildlife (specifically snakes), not being
interested, being busy with work and childcare, not knowing of the existence of the
project, not knowing where to look for information, and finally, not perceiving the
forest as functional or open to the public. Similarly, other research has found that
around half of interviewed local residents have not visited protected areas that are
adjacent to their homes (Pérez-Verdin et al. 2004; Moorman 2006).
Regardless of whether they had visited the forest or not, 33 of the 37 interviewees
expressed interest in participating in at least one of the activities that the RHCF board
already organizes. This implies that rather than lack of interest from community
members, perhaps increased communication about the activities, setting weekly
hours for visits and making other offerings available could increase participation
of residents that surround the community forest. Increasing opportunities of sur-
rounding residents to learn about the ecology of the forest and its establishment may
increase their knowledge and valuation of the forest as a protected area (Moorman
2006). This effort of increased communication is a good opportunity since, beyond
just participating in activities, 14 interviewees mentioned interest in forming part or
getting involved in the development or management activities and events with the
RHCF board.
20 Priorities, Perspectives, and Use of a Community Forest by. . . 521

The perception of the forest not being open or functional, or interviewees prefer-
ring to visit the forest if it was “clean” has to do with different values that people
attribute to natural areas, and probably with aesthetics and what is visually pleasing,
as well as with the presence of trash, especially on the sides of the road that crosses the
forest. Other studies have demonstrated that attributes such as removal of dead trees,
removal of understory, improved visibility, greater crown density, less leaf litter and
improved visual distance contribute to perceived aesthetic improvements and land-
scape quality (Tyrväinen et al. 2003; Chen et al. 2015) and are usually related to
factors such as accessibility and security of the area (Tyrväinen et al. 2003).
Ecological and aesthetic management goals can compete with each other, espe-
cially when some management for ecological purposes, like improved wildlife
habitat, can be perceived as not aesthetically pleasing. The forest therefore might
be perceived as not functional and as potentially -but not currently- optimal for
tourism and recreation because the current forest structure is heterogeneous and has a
dense understory, even more so due to the creation of canopy openings after
hurricane María. For the same reason the forest also has many snags, and tree
species that have associated negative perceptions, like Albizia procera.
Although not documented in the survey, informal conversations with residents
shed light in some confusion in terms of the ownership of the forest and what
management practices are allowed or prohibited. Some people referred to the forest
as belonging to the U.S. federal government and some as belonging to the Mayagüez
municipality, and overall they were not quite sure of the role of the community in
making decisions on the management of the forest. This should be an important
clarification to make in further outreach and communications coming from the
RHCF board, as protected areas that are “perceived to have a level of mutual benefit
and co-management generate more support toward conservation and recreational use
goals” (Buta et al. 2014).

20.4.3 Exploring the Potential of Positive Perceptions

and Values for Sustaining Conservation

The results on purpose of interviewee’s visits to the forest highlight the importance
of the forest to be used as a common place for social gatherings for the community.
Meanwhile, the results on the activities that residents would go for, shows that many
of the activities that the RHCF board already hosts would be attended by residents
that surround the forest, especially those that are dedicated to agriculture, sales,
education and recreation. Many of the recommendations by interviewees, like
increased recreational infrastructure, are present in the goals of the forest’s manage-
ment plan. This highlights that the participatory process of developing this plan
captured many interests of the community, which have been emphasized in this
research project. However, the implementation of these goals is contingent on
sourcing funding and the voluntary labor (for now) of members of the RHCF
board and others.
522 G. M. Morales-Nieves

In terms of air quality, it has been found that urban trees can remove air pollution,
like particulate matter, ozone, nitrogen dioxide, sulfur dioxide and carbon monoxide,
in amounts of around 80 pounds per acre per year (~90 kg/ha) at least in the United
States (Nowak and Heisler 2010). Urban trees also have been found to lower local
temperatures, which can also result in lower emission of pollutants and improved air
quality. The reduction in local temperature also results in lower energy consumption
from air conditioning from surrounding households. However, trees can also be a
source of pollutants, like pollen and volatile organic compounds (Nowak and Heisler
2010; Leung et al. 2011; Baumgardner et al. 2012). Considering the hard edge of the
forest with neighboring households, and in consideration with comments about the
“cleanliness” of the forest, the RHCF could contemplate managing the understory at
least in the borders of the forest to reduce the presence of species like the “picapica”
(Mucuna pruriens).
This research also highlighted how much people value the forest for the perceived
benefit of air quality improvement. As an interviewee mentioned: “If they didn’t
protect it, they (original landowners) were going to cut it down to build houses and
put concrete”, while another mentioned: “Because of the trees, for the air. If you
notice, there are no trees in all of this area”. This demonstrates that residents value
the forest for its mere presence in the area. Different perhaps from many other
community forests in Latin America, surrounding residents are not highly dependent
on the forest resources for their livelihoods, like for fuelwood, food and medicine.
They mostly value it or perceive benefits from more passive uses, like air quality and
recreation.
Research in other community forests has shown that they provide opportunities
for nature-based recreation and for income either through the direct charging of
services associated with the forest or indirectly from visitor expenditure in surround-
ing businesses (Birch et al. 2014). The RHCF is in a great position to expand
recreation as a service of the forest since many of its current activities are based
on recreation and education and many residents in this research highlighted recre-
ation, both as a perceived benefit of the forest and as activities that they would
participate in.
Beyond these two main services, the forest can also provide other services to the
community such as serving as a source of food. Among both perceived services and
even more so among activities that residents would visit the forest for, were those
that were related to food and medicine. Other community forests, even those that are
of small scale such as the RHCF, have been proven to serve as a source of food and
addressing food insecurity (Paudel 2018; Bridhikitti and Khadka 2019). In case
studies from other community-based forestry projects in the Caribbean, results show
that skills, knowledge and capacity building; strengthened organizational capacity
and empowerment; and increased financial income, have been the main impacts and
benefits (CANARI 2012).
The goal to maintain and improve the quality of life of the community through the
conservation of the RHCF as the highest ranked goal highlighted once again the
intrinsic value of the presence of the forest for surrounding residents. On the other
20 Priorities, Perspectives, and Use of a Community Forest by. . . 523

hand, the competing rankings for the rest of the goals shows that all goals delineated
in the management plan are equally important, and no pattern demonstrated other
goals to be more important for people, made more evident with two interviewees
declining to rank them at all.
Other research has found common attributes that contribute to successful man-
agement of community forests and influence surrounding residents to engage in
conservation projects. These attributes I would recommend be considered by those
managing the RHCF based on my findings and are summarized below:
• Interest and positive attitudes of surrounding residents toward forest management
along with strong, open, inclusive, involvement and participation (Pagdee et al.
2006; CANARI 2012; Bridhikitti and Khadka 2019), with an emphasis on youth
(CANARI 2012)
• Incentives for forest use for fulfilling basic community needs (Pagdee et al. 2006)
• Market and state economic incentives, human and financial resources to provide a
new source of income (Pagdee et al. 2006; Ruiz-Mallén et al. 2015)
• Clear ownership to be able to use and manage the forest (Pagdee et al. 2006)
• Clearly defined forest boundary (Pagdee et al. 2006)
• Strong leadership and effective local organizations (Pagdee et al. 2006; CANARI
2012)
• Continuous transfer of knowledge and use of traditional techniques to harvest
forest products (Pagdee et al. 2006)
• A developed sense of place, historical connection, belonging and cultural identity
of community members as well as social cohesion (CANARI 2012; Ruiz-Mallén
et al. 2015)
• Building trust and maintaining open and frequent communication with govern-
ment partners for transparency and accountability (CANARI 2012)
The protection of this forest, besides being important for the ecosystem services it
provides for people, can also provide benefits for biodiversity in the area. As
mentioned by an interviewee, it is the most important ecosystem service of all:
“But the most important thing for the community is to leave space for habitat for
animals, and for bees to pollinate plants.” This forest can be a refuge for wildlife and
plant life, even when used for different purposes. Agroforests and managed second-
ary forests, like the RHCF, have been found to have high plant diversity, including
threatened species and often have similar basal area than that of old-growth forests,
all while improving local livelihoods (Souza et al. 2016). Perhaps a combined effort
on education and demonstration of the uses of non-native species like Albizia
procera for fuelwood (Lugo et al. 1990) for example, to be then replaced with
assisted planting of native species and other desirable species by the community
could increase forest use, while improving ecosystem function and services. As
mentioned by Castro-Prieto et al. (2017) there seems to be increasing development
pressures around the RHCF, therefore its protection as a biodiversity island could be
considered increasingly relevant.
524 G. M. Morales-Nieves

20.5 Conclusions

People living around the Río Hondo Community Forest in Mayagüez, Puerto Rico
valued the forest for its mere presence: as a source of clean air, recreation or
socializing, beauty and education. Even though nobody opposed the presence of
the forest, some people had concerns about the status of the forest, especially in
relation to the presence of wildlife and plant species that are perceived as a nuisance;
dense understory, and lack of park-like infrastructure.
Given similar demographic trends of the community to other parts of Puerto Rico
and ecologic traits of the forest itself, if this model of community forest were to be
replicated in other areas, one could anticipate similar expectations, perceived bene-
fits and potential uses by other communities. However, as the land is guaranteed to
remain as forest into perpetuity, the RHCF within their efforts, according with what
is already outlined in their management plan, and with additional resources and
funding, could expand and teach about other potential or increased uses, like very
small-scale timber harvesting, agroforestry, or other. They could also promote more
active sources of income for surrounding community residents, like tourism and
development of value-added products.
This project is a case study that can be comparable to other forests with similar
land use history in many other parts of Puerto Rico and the Caribbean and could be
used as an example to be replicated taking into account the findings in this research,
along with attributes that make community-based forestry successful. Based on
observed wildlife and tree species found, including endemic ones, this forest serves
as a biodiversity island within a fragmented landscape in the rural-urban interface in
western Puerto Rico.

Acknowledgements I would like to thank my advisors, Drs. Florencia Montagnini and Mark
Ashton; Dr. Oscar Abelleira for connecting me with this wonderful forest; and Drs. Amity Doolittle,
Eva Garen, Kathleen McGinley and Germán Ramos for their additional advice in the design of this
project. I also thank Magaly Figueroa, from the US Forest Service’s International Institute of
Tropical Forestry; Betsy Acevedo and all the members of the RHCF Community Board for their
advice, for their time to meet with me and for the great voluntary service they provide to the broader
community in managing this forest. I also thank Yale University’s Tropical Resources Institute and
Sustainable Food Program for providing funding for this research project. Finally, I thank Camila
Martínez for her assistance during data collection.
20 Priorities, Perspectives, and Use of a Community Forest by. . . 525

Appendices

Appendix A: Population Density in 2017 of Countries in Latin

America (LA) and the Caribbean ©

Population density (people per

Country name LA or C sq. km of land area)
Saint Maarten (Dutch part) C 1193
St. Martin (French part) C 672
Barbados C 666
Aruba C 585
Haiti C 398
Puerto Rico C 375
Curacao C 361
Grenada C 326
El Salvador LA 308
Virgin Islands (U.S.) C 306
St. Lucia C 297
St. Vincent and the Grenadines C 282
Trinidad and Tobago C 270
Jamaica C 270
Cayman Islands C 264
Dominican Republic C 218
Antigua and Barbuda C 217
St. Kitts and Nevis C 200
British Virgin Islands C 197
Guatemala LA 158
Cuba C 109
Costa Rica LA 97
Dominica C 95
Honduras LA 84
Ecuador LA 68
Mexico LA 64
Panama LA 55
Nicaragua LA 53
Colombia LA 44
Turks and Caicos Islands C 39
The Bahamas C 38
Venezuela LA 33
Brazil LA 25
Chile LA 25
Peru LA 25
Uruguay LA 20
Paraguay LA 17
(continued)
526 G. M. Morales-Nieves

Population density (people per

Country name LA or C sq. km of land area)
Belize LA 16
Argentina LA 16
Bolivia LA 10
Guyana LA 4
Suriname LA 4

Appendix B: Tree Species Found in Río Hondo Community

Forest (Obtained from Rodríguez Candelaria et al. Working
Document)

_________________________________________________________________.

Albizia proceraa Ceiba pentandra Randia acuelata

Andira inermis Cinamomun montanum Schefflera morotononi
Artocarpus altilisa Cordia laevigata Senna siameaa
Casearia aculeata Cupania americana Spathodea campanulataa
Casearia decandra Delonix regiaa Swietenia macrophyllaa
Casearia guianensis Guarea guidonia Terminalia catappaa
Casearia sylvestris Inga laurina Zanthoxylum martinicense
Cecropia peltata Melicocus bijogatusa
a
Non-native species

Appendix C: Wildlife Species Observed in the RHCF Between

2009 and 2017 (Obtained from Rodríguez Candelaria et al.
Working Document)

Bird Species
Icterus dominicensisa Turdus plumbeus Mimus polyglottos
Melanerpes portorricencisa Lonchura punctulata Patagioenas squamosa
Spindalis portorricensisa Butorides virescens Coereba flaveola
Myiarchus antillaruma Coccyzus vieillotia Anthracothorax dominicus
Todus mexicanusa Vireo latimeri Falco sparverius
Dendroica discolor Loxigilla portorricencisa Buteo jamaicencis
Mniotilta varia Vireo altiloquus Anthracothorax viridisa
Parula americana Euplectes franciscanus Tyrannus dominicensis
Dendroica tigrina Columbina passerina Brotogeris verscicolurus
(continued)
20 Priorities, Perspectives, and Use of a Community Forest by. . . 527

Bird Species
Icterus icterus Margarops fuscatus Crotophaga ani
Zenaida asiatica Tiaris bicolor Tyrannus caudifasciatus
Zenaida aurita
Reptiles
Ameiva exsul Anolis spp. Iguana iguana
Alsophis spp.
Amphibians
Bufo marinus Eleutherodactylus spp.a Leptodactylus albilabris
a
Endemic species

Appendix D: The Six Goals Identified During the Creation

of the RHCF Land Management Plan (Obtained from
Rodríguez Candelaria et al. Working Document)

18a. Maintain and improve the quality of life of the community through the
conservation of the RHCF (for example, through the planting and maintenance
of trees that improve air quality and reduce pollution).
18b. Provide economic benefits to communities through the generation of employ-
ment, maintenance of clean conditions of the forest and being self-sufficient by
selling products, services and activities.
18c. Promote the participation of members of the community, community organi-
zations, educational institutions, government entities and environmental conser-
vation organizations.
18d. Strengthen the social development of the community by highlighting the
customs and the cultural, historic, folkloric, generational and traditional elements
that represent and integrate the community with the objective of creating and
contributing to the cultural development.
18e. Increase the resiliency of the community to face the challenges of climate
change and economic crises (for example, install a rainwater collection system
to share in times of lack of water, etc.)
18f. Promote the forest as a recreational space for the individual, family and
community well-being (for example, the maintenance and creation of new walk-
ing trails, maintenance of areas for camping, etc.)
528 G. M. Morales-Nieves

References

Acevedo-Rodríguez P (2005) Vines and climbing plants of Puerto Rico and the Virgin Islands.
Smithsonian Inst. Contr US Nat Herb 51:1–483
Acuerdos colaborativos (2015) In: DRNA. [Link]
acuerdos-colaborativos/. Accessed 12 Feb 2020
Aide TM, Clark ML, Grau HR, López-Carr D, Levy MA, Redo D, Bonilla-Moheno M, Riner G,
Andrade-Núñez MJ, Muñiz M (2013) Deforestation and reforestation of Latin America and the
Caribbean (2001–2010). Biotropica 45:262–271. [Link]
00908.x
Álvarez-Berríos NL, Redo DJ, Aide TM, Clark ML, Grau RH (2013) Land change in the greater
Antilles between 2001 and 2010. Land 2:81–107. [Link]
Anadon-Irizarry V, Wege DC, Upgren A, Young R, Boom B, Leon YM, Arias Y, Koenig K,
Morales AL, Burke W (2012) Sites for priority biodiversity conservation in the Caribbean
Islands biodiversity hotspot. J Threat Taxa:2806–2844. [Link]
2806-44
Baumgardner D, Varela S, Escobedo FJ, Chacalo A, Ochoa C (2012) The role of a peri-urban forest
on air quality improvement in the Mexico City megalopolis. Environ Pollut 163:174–183.
[Link]
Birch JC, Thapa I, Balmford A, Bradbury RB, Brown C, Butchart SHM, Gurung H, Hughes FMR,
Mulligan M, Pandeya B, Peh KS-H, Stattersfield AJ, Walpole M, Thomas DHL (2014) What
benefits do community forests provide, and to whom? A rapid assessment of ecosystem services
from a Himalayan forest. Nepal Ecosyst Serv 8:118–127. [Link]
03.005
Bowler DE, Buyung-Ali LM, Healey JR, Jones JP, Knight TM, Pullin AS (2012) Does community
forest management provide global environmental benefits and improve local welfare? Front
Ecol Environ 10:29–36. [Link]
Brandeis TJ, Turner JA (2013) Puerto Rico’s forests, 2009. USDA Forest Service, Asheville
Bridhikitti A, Khadka B (2019) Assessing factors to successful management for small-scale
community forest under threat of urban growth: in a case of Ban Na Kham Noi community
forest, Mukdahan, Thailand. J Sustain For 0:1–17. [Link]
1631184
Buta N, Holland SM, Kaplanidou K (2014) Local communities and protected areas: the mediating
role of place attachment for pro-environmental civic engagement. J Outdoor Recreat Tour 5–6:
1–10. [Link]
CANARI (2012) Community forestry in the Caribbean: a regional synthesis. Caribbean Natural
Resources Institute (CANARI). Laventille, Trinidad and Tobago
Castro-Prieto J, Gould W, Ortiz-Maldonado C, Soto-Bayó S, Llerandi-Román I, Gaztambide-
Arandes S, Quinones M, Cañón M, Jacobs KR (2019) A comprehensive inventory of protected
areas and other land conservation mechanisms in Puerto Rico. IITF-Gen Tech Rep 50. IITF, Rio
Piedras, Puerto Rico
Castro-Prieto J, Martinuzzi S, Radeloff VC, Helmers DP, Quiñones M, Gould WA (2017) Declin-
ing human population but increasing residential development around protected areas in Puerto
Rico. Biol Conserv 209:473–481. [Link]
Center for Puerto Rican Studies (2019) Puerto Rico two years after hurricane María. Hunter College
CUNY, New York
Ceperley N, Montagnini F, Natta A (2010) Significance of sacred sites for riparian forest conser-
vation in Central Benin. Bois Forets Trop 303(1):5–23
Chen Y, Sun B, Liao S, Chen L, Luo S (2015) Landscape perception based on personal attributes in
determining the scenic beauty of in-stand natural secondary forests. Ann For Res 59:91–103.
[Link]
20 Priorities, Perspectives, and Use of a Community Forest by. . . 529

Dalton T, Forrester G, Pollnac R (2012) Participation, process quality, and performance of marine
protected areas in the wider Caribbean. Environ Manag 49:1224–1237. [Link]
s00267-012-9855-0
DataBank | The World Bank. [Link] Accessed 22 Jan 2020
Garen EJ, Saltonstall K, Slusser J, Mathias S, Ashton MS, Hall JS (2009) An evaluation of farmers’
experiences planting native trees in rural Panama: implications for reforestation with native
species in agricultural landscapes. Agrofor Syst 76(1):219–236
Geoghegan T, Renard Y (2002) Beyond community involvement: lessons from the insular Carib-
bean. Parks 12(2)
Gilmour DA (2016) Forty years of community-based forestry: a review of its extent and effective-
ness. FAO, Rome
Gonzalez G, Scalley TH (2016) Building a collaborative network to understand regional forest
dynamics and advance forestry initiatives in the Caribbean. Caribb Nat 2016 Spec Issue 1:245–
256
Grau HR, Aide TM, Zimmerman JK, Thomlinson JR, Helmer E, Zou X (2003) The ecological
consequences of socioeconomic and land-use changes in postagriculture Puerto Rico. Bio
Science 53:1159–1168. [Link]
CO;2
Helmer EH, Kennaway TA, Pedreros DH, Clark ML, Marcano-Vega H, Tieszen LL, Schill SR,
Carrington CMS (2008) Land cover and forest formation distributions for St. Kitts, Nevis,
St. Eustatius, Grenada and Barbados from decision tree classification of cloud-cleared satellite
imagery. Caribb J Sci 44:175–198
Holland G, Bruyère CL (2014) Recent intense hurricane response to global climate change. Clim
Dyn 42:617–627. [Link]
How the Community Forest Program Works | US Forest Service. [Link]
managing-land/private-land/community-forest/program. Accessed 10 Feb 2020
Junta de Planificación de Puerto Rico (2019) Resumen Económico De Puerto Rico. Junta de
Planificación de Puerto Rico, San Juan. [Link]
Knutson TR, Sirutis JJ, Zhao M, Tuleya RE, Bender M, Vecchi GA, Villarini G, Chavas D (2015)
Global projections of intense tropical cyclone activity for the late twenty-first century from
dynamical downscaling of CMIP5/RCP4.5 scenarios. J Clim 28:7203–7224. [Link]
1175/JCLI-D-15-0129.1
Lambrick FH, Brown ND, Lawrence A, Bebber DP (2014) Effectiveness of community forestry in
prey long forest, Cambodia. Conserv Biol 28:372–381. [Link]
Leung DYC, Tsui JKY, Chen F, Yip W-K, Vrijmoed LLP, Liu C-H (2011) Effects of urban
vegetation on urban air quality. Landsc Res 36:173–188. [Link]
2010.547570
López T d M, Aide TM, Thomlinson JR (2001) Urban expansion and the loss of prime agricultural
lands in Puerto Rico. Ambio 30:49–54
Lugo AE, Wang D, Herbert Bormann F (1990) A comparative analysis of biomass production in
five tropical tree species. For Ecol Manag 31:153–166. [Link]
90158-8
Marcano-Vega H (2017) Forests of Puerto Rico, 2014. USDA Forest Service, Asheville
Martinuzzi S, Gould WA, Ramos González OM (2007) Land development, land use, and urban
sprawl in Puerto Rico integrating remote sensing and population census data. Landsc Urban
Plan 79:288–297. [Link]
Massol González A, González E, Massol Deyá A, Deyá Díaz T, Geoghegan T (2006) Bosque del
Pueblo, Puerto Rico: Cómo la lucha antiminera cambió la política forestal desde la base
comunitaria. International Institute for Environment and Development, London
McConney P, Pena M (2012) Capacity for (co)management of marine protected areas in the
Caribbean. Coast Manag 40:268–278. [Link]
Moeliono A, Mulyana M, Adnan A, Manalu H, Yuliani P, Balang L (2015) Village forests (hutan
desa): empowerment, business or burden? World Agroforestry Centre (ICRAF), Bogor
530 G. M. Morales-Nieves

Moorman RS (2006) Benefits of local residents visiting La Selva Biological Station, Costa Rica.
Environ Conserv 33:89–99. [Link]
Myers N, Mittermeier RA, Mittermeier CG, Fonseca GAB d, Kent J (2000) Biodiversity hotspots
for conservation priorities. Nature 403:853–858. [Link]
Nowak D, Heisler G (2010) Air quality effects of urban trees and parks. Res Ser Monogr Ashburn
VA Natl Recreat Parks Assoc Res Ser Monogr 44:1–44
Pagdee A, Kim Y, Daugherty PJ (2006) What makes community forest management successful: a
meta-study from community forests throughout the world. Soc Nat Resour 19:33–52. https://
[Link]/10.1080/08941920500323260
Paudel J (2018) Community-managed forests, household fuelwood use and food consumption. Ecol
Econ 147:62–73. [Link]
Pérez-Verdin G, Lee ME, Chávez DJ (2004) Outdoor recreation in a protected area in southern
Durango, Mexico: analysis of local residents’ perceptions. Soc Nat Resour 17:897–910. https://
[Link]/10.1080/08941920490505310
Porter-Bolland L, Ellis EA, Guariguata MR, Ruiz-Mallén I, Negrete-Yankelevich S, Reyes-García
V (2012) Community managed forests and forest protected areas: an assessment of their
conservation effectiveness across the tropics. For Ecol Manag 268:6–17. [Link]
1016/[Link].2011.05.034
Pulwarty RS, Nurse LA, Trotz UO (2010) Caribbean Islands in a changing climate. Environ Sci
Policy Sustain Dev 52:16–27. [Link]
Rasolofoson RA, Ferraro PJ, Jenkins CN, Jones JPG (2015) Effectiveness of community forest
management at reducing deforestation in Madagascar. Biol Conserv 184:271–277. [Link]
org/10.1016/[Link].2015.01.027
Rodríguez Candelaria I, López Méndez C, Rivera Sanantonio J, García Santiago D, Crespo Vélez S,
Pérez Méndez A, Abelleira Martínez O, Ramos Cartagena G (n.d.) (Working Document) Plan
de Co-Manejo del Bosque Comunitario de Río Hondo en Mayagüez, Puerto Rico: Empresa
Comunitaria con Propósito Social, Ambiental, Cultural y Recreativo. Mayagüez, Puerto Rico.
Bosque Comunitario de Río Hondo-Mayagüez-Facebook
Ruiz-Mallén I, Schunko C, Corbera E, Rös M, Reyes-García V (2015) Meanings, drivers, and
motivations for community-based conservation in Latin America. Ecol Soc 20:3. [Link]
10.5751/ES-07733-200333
Shrestha B, Shrestha S (2010) Biodiversity conservation in community forests of Nepal: rhetoric
and reality. Int J Biodivers Conserv 2:98–104
Smith SL (2012) Toward inclusive co-management: factors influencing stakeholder participation.
Coast Manag 40:327–337. [Link]
de Souza SEXF, Vidal E, Chagas G d F, Elgar AT, Brancalion PHS (2016) Ecological outcomes
and livelihood benefits of community-managed agroforests and second growth forests in
Southeast Brazil. Biotropica 48:868–881. [Link]
Tyrväinen L, Silvennoinen H, Kolehmainen O (2003) Ecological and aesthetic values in urban
forest management. Urban For Urban Green 1:135–149. [Link]
8667-00014
Wood A, Tolera M, Snell M, O’Hara P, Hailu A (2019) Community forest management (CFM) in
south-west Ethiopia: maintaining forests, biodiversity and carbon stocks to support wild coffee
conservation. Glob Environ Change 59:1–11. [Link]
Chapter 21
Sacred Church Forests in Northern
Ethiopia: Biodiversity and Cultural Islands

Mabel Baez Schon, Carrie L. Woods, and Catherine L. Cardelús

Abstract Sacred forests, protected due to their religious importance, form a vast
network of informal and often inadvertent (shadow) conservation sites worldwide.
Despite socioeconomic and political pressures increasing deforestation worldwide,
these shadow conservation sites are remarkable in their ability to resist these
pressures through their cultural, religious, and social significance. In this chapter
we discuss our research on the sacred forests of the Ethiopian Orthodox Tewahido
Church (EOTC), the dominant religion in Ethiopia. Ethiopia has 11% forest cover
distributed in a patchwork of forest fragments which are, in many regions, the only
repository of biodiversity, including many endemic and endangered species. In the
South Gondar Region (14,607 km2) of northern Ethiopia, around 1022 of these
forest fragments are sacred forests that surround churches of the EOTC. Despite their
small size, on average 5.2 hectares (range 1.7–148.9 ha), they constitute 100% of the
forests in the region. High levels of biodiversity, including endangered and endemic
taxa, endow these forests with great value. They provide essential ecosystem
services to the surrounding community and are integral to the rituals and culture of
the EOTC. Far from being static cultural and religious relics scattered throughout the
Ethiopian landscape, these sacred church forests are complex and dynamic socio-
ecological systems. We discuss EOTC forests as islands of biocultural diversity,
their central role in local communities, the ecosystem services they provide, and the
threats they face. We base our observations on a 10-year (2010–2020) interdisci-
plinary research project where we explore the mechanisms of the religious manage-
ment of EOTC church forests.

M. Baez Schon (*)

School of Forest, Fisheries, and Geomatics Sciences, University of Florida,
Gainesville, FL, USA
e-mail: mabelbaez@ufl.edu
C. L. Woods
Department of Biology, University of Puget Sound, Tacoma, WA, USA
C. L. Cardelús
Department of Biology, Colgate University, Hamilton, NY, USA

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 531
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
532 M. Baez Schon et al.

Keywords Biocultural conservation · Church forests · Ethiopian Orthodox

Tewahido Church · Sacred forests · Sacred natural sites · Shadow conservation

21.1 Introduction

21.1.1 Sacred Natural Sites

Sacred natural sites (SNS) are defined as “areas of land or water having special
spiritual significance to peoples and communities” (Oviedo et al. 2005, p. 3).
Sacredness can be interpreted in different ways, and is attributed to natural sites by
people across different religious backgrounds and faiths—from indigenous and
traditional to institutionalized religions (Verschuuren et al. 2010; Pungetti 2012).
As such, the definition of SNS is an open working definition, acknowledging the fact
that interpretation of what areas are sacred, and why, is not fixed (Verschuuren et al.
2010). In some indigenous traditions, for example, all nature is sacred and animals
and people are related (Harmon and Putney 2003; Pungetti 2012). Sacred natural
sites can be rock formations (e.g. Uluru, Australia), mountain ranges (e.g. Mt.
Hakusan, Japan), or whole ecosystems (e.g. Tanchara Wetlands, Ghana). Water is
an essential resource that is often depicted as sacred (Altman 2002). Sacred water
sites include sacred pools (e.g. Benin) (Ceperley et al. 2010), rivers (e.g. the Ganges
River, India), lakes, wells, and springs (e.g. sacred fertility springs in Kyrgyzstan)
(Altman 2002; Verschuuren and Wild 2012). Natural areas may also have sacred
energies, sacred species, sacred ancestral burial grounds, contain individual sacred
trees, encompass entire forest groves (e.g. Indian sacred groves), and/or be sacred at
specific times of the year (Bhagwat and Rutte 2006; Delgado et al. 2012;
Pungetti 2012).
Sacred natural sites are ubiquitous and found on every continent except Antarc-
tica (Bhagwat and Rutte 2006; Rutte 2011). Sacred forests, a type of SNS, are often
small in size but together make up an important network of forest conservation areas.
These are often referred to as “shadow” conservation areas. Shadow conservation
areas are protected indirectly through community use rather than directly through
legal protection designation. Notably, they are not protected for the explicit conser-
vation of biodiversity (Bhagwat and Rutte 2006; Cardelús et al. 2017; Kent and
Orlowska 2018). Sacred forests are some of the oldest forms of conserved areas and
are important for biodiversity at the local, regional, and national scales comparable
to, and complementary to, protected areas (PA) (Bhagwat and Rutte 2006; Tengö
et al. 2007; Verschuuren et al. 2010). The religious, cultural, and biological impor-
tance of sacred forests are well known and documented (Verschuuren et al. 2010;
Pungetti et al. 2012).
The origins of sacred forests are as diverse as their various examples. In areas
with extensive agricultural development, sacred forests are likely as old as agricul-
ture (Bhagwat and Rutte 2006). As land was cleared for farming, humans set aside
parcels of forested land that were considered aesthetically pleasing or were reli-
giously valuable. As populations grew, and agricultural practices intensified,
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 533

economic and sociocultural pressures increased to use these fragments in a more

utilitarian manner. As communities began to use these land areas as resources,
benefitting from the ecosystem services that they provided, the cultural services
the land could provide were increasingly threatened (Bhagwat and Rutte 2006).
Ecosystem services can be divided into 4 categories—(1) provisioning services
(food, water, timber), (2) regulating services (regulation of climate, floods, water
quality), (3) cultural services (recreation, aesthetic enjoyment, and spiritual fulfill-
ment), and (4) supporting services (soil formation, photosynthesis, and nutrient
cycling) (Millenium Ecosystem Assessment 2010). To prevent the overexploitation
of ecosystem services and natural resources that the forest fragments provided,
“social fences” grew around them (Bhagwat and Rutte 2006). Social fences are
described as informal community conservation mechanisms that prevent
overexploitation of forest resources and ensure their perpetuity (Colding and Folke
2001; Bhagwat and Rutte 2006). Thus, while every ecosystem service may not have
been the primary purpose for protecting these sacred forests, their very presence had
utilitarian value when the landscape changed. This argument is supported by
research on the landscape characteristics of protected areas (PAs) and SNS in
Italy, in which the network of PAs and SNS showed little overlap. Sacred natural
sites were found more often in extensively agricultural and peri-urban settings, while
PAs where located farther from human settlements (Frascaroli et al. 2019).
While sacred forests are not officially considered conservation mechanisms in the
International Union for Conservation of Nature (IUCN) Protected Area network,
sacred forests are a form of community forest management, which currently play,
and have traditionally played, an integral role in the conservation of biodiversity and
should be considered in conservation initiatives (Bhagwat and Rutte 2006; Dudley
and Higgins-Zogib 2012; Montagnini et al. 2022). Their role in conservation of
biocultural (biological and cultural) diversity was acknowledged in the 2005 United
Nations Educational, Scientific and Cultural Organization (UNESCO) international
symposium in Tokyo which resulted in the joint IUCN-UNESCO guidelines for
management of SNS (Wild and McLeod 2008). Sacred natural sites are currently
recognized under UNESCO’S World Heritage Sites (Wild and McLeod 2008).
Ultimately sacred forests are protected due to their spiritual, historical, and cultural
value (Bhagwat and Rutte 2006), also providing benefits to the surrounding area by
serving as biodiversity islands.
In this chapter we discuss the sacred forests of the Ethiopian Orthodox Tewahido
Church (EOTC) (henceforth: EOTC church forests), the dominant religion in Ethi-
opia. EOTC church forests are islands of biocultural diversity due to their central role
in local communities, the ecosystem services they provide, and the threats the
otherwise denuded landscape faces. Despite their designation as SNS, they have
received comparatively little attention relative to other well-known SNS, such as
sacred groves in India (Ormsby and Bhagwat 2010; Notermans et al. 2016) or
western Africa (e.g., SNS in Ghana, Ormsby 2012). Our observations are based on
extensive ecological and ethnographical field research conducted by an interdisci-
plinary research team in the South Gondar Administrative Zone (SGAZ), Amhara
National Regional State, Ethiopia. We researched 46 church forests over 10 years
534 M. Baez Schon et al.

(2010–2020) to document and evaluate the relationship of the church forests and the
communities that inhabit and depend on them (Cardelús et al. 2013; Klepeis et al.
2016; Scull et al. 2017; Woods et al. 2017; Kent and Orlowska 2018; Orlowska and
Klepeis 2018; Cardelús et al. 2019; Woods et al. 2020; Cardelús et al. 2020).

21.2 History of Sacred Natural Sites in Ethiopia and Their

Shadow Conservation

Sacred church forests of northern Ethiopia have a fascinating history believed to

have originated in the fourth century AD (Wassie 2002; Karbo 2013). Community-
driven “shadow conservation” through traditional institutions has allowed these
sacred forests to persist to today (Dudley et al. 2009; Cardelús et al. 2017). This
contrasts with the negative perception that exaggerates historical anthropogenic
degradation and deforestation in Ethiopia. The most pervasive story is that forest
cover decreased from 40% to 4% in the late nineteenth and early twentieth centuries
as detailed by McCann (1997). The first published reference to this dramatic forest
loss was published, uncited, in the Food and Agriculture Organization’s “Agricul-
ture in Ethiopia” (1961) (McCann 1997). This narrative took hold abroad, as well as
within Ethiopia. In 1985, the Ethiopian Relief and Rehabilitation Commission
officially connected the Ethiopian famine to the 36% reduction in forest cover
from 1885 to 1985. The story of Ethiopia’s degradation was directly attributed to
Ethiopians’ mismanagement of their natural resources. While there is strong evi-
dence of human reduction of forest cover, many of the current forests exist because
of human intervention (McCann 1997; Sigman 2022).
The earliest known landscape descriptions of the Ethiopian highlands from the
1830s highlight a lack of forest cover. The presence of trees was not an indication of
remnant trees spared from deforestation, but rather of careful management. Today,
across the Ethiopian highlands, forests are almost exclusively found with an Ethio-
pian Orthodox Tewahido Church (EOTC) at their center (Wassie 2002). The persis-
tence of these sacred church forests in the region is because of their shadow
conservation by the community (Cardelús et al. 2017; Kent and Orlowska 2018).
A precise number of EOTC church forests is unknown; however estimates place
them at around 35,000 throughout Ethiopia (McCann 1997; Berhane-Selassie 2008).
While Ethiopia is ethnically, culturally, and linguistically diverse, there persists a
remarkable sacred connection between the Ethiopian peoples and their forests, a
cultural phenomenon in which there is EOTC Christian, non-Christian, and indige-
nous veneration of sacred trees and forests (Berhane-Selassie 2008; Doffana 2019).
EOTC church forests have proven to be resilient within the landscape. A recent
study compared changes in the northern Ethiopian highlands sacred forests and
surrounding forest cover using aerial photographs taken during the Italian occupa-
tion of Ethiopia (1935–1941) against modern (2014–2016) aerial photographs.
While they found a decrease in the forest cover surrounding the sacred forests and
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 535

adjacent agricultural lands, they did not find a decrease in EOTC church forest size
(Scull et al. 2017). During this time period, only four EOTC church forests
disappeared. These findings corroborate those of McCann (1997) and show remark-
able persistence of forest patches due to local stewardship. Specifically, the gradual
deforestation of buffer zones surrounding sacred forests, but not of the sacred forests
themselves, highlights the religious value of SNS and their importance to the EOTC
community (Scull et al. 2017). The elimination of buffer zones also presents a new
threat to these islands of biodiversity as they are more susceptible to “edge effects,”
which are characterized by increased air temperature, vapor pressure deficits, UV
radiation, wind penetration, evapotranspiration, and tree mortality, along with
decreased soil moisture and changes in nutrient availability that diminish the pro-
ductivity and integrity of forests (Kapos 1989; Malcolm 1994; Scariot 2000).
To examine the resilience of EOTC church forests in Ethiopia, it is important to
understand the significant social and political changes within Ethiopia since the
1930s. Following the return of emperor Haile Salassie in 1941, marking the end of
the Italian occupation (1935–1941), the EOTC experienced a process of centraliza-
tion. Under Salassie’s rule, the church and state were united. In 1974, the Marxist
Derg regime overthrew emperor Haile Salassie. While the Derg regime maintained
ties to the Ethiopian Church, largely due to its importance to Ethiopians, in 1974 the
church and state were officially separated. The Public Ownership of Rural Land
Proclamation on 1975 further weakened the EOTC, by ending the landlord system
through the nationalization of all rural land (Keeley and Scoones 2000; Ancel and
Ficquet 2015; Zerga 2016). This had significant implications for sacred lands
throughout Ethiopia, and specifically for the EOTC lands. The proclamation led to
the redistribution of non-forested church land to landless individuals, though ulti-
mately land ownership remained in the hands of the State (Keeley and Scoones
2000; Ancel and Ficquet 2015; Zerga 2016).
The restructuring of land tenure systems was a significant blow to EOTC income
and land security as 5–50% of productive land used for agriculture and economic
activities was lost (Yigremew 2002). Historically, the narrative of degradation
reflected and supported a broader movement of land centralization in the name of
preventing the overuse of the common resources (Zerga 2016).
The Derg Regime was overthrown by the current government in 1991. Despite
promises to change land-tenure systems throughout Ethiopia, under the current
government, land tenure has remained essentially the same. The post-revolutionary
government introduced an ethnic territoriality system known as “Kilil”, which often
ignores local land claims, such as those held by the EOTC (Berhane-Selassie 2008).
Currently, the EOTC only has usufruct rights to their lands, and these sacred church
sites are not officially recognized by the state (Berhane-Selassie 2008). While the
church forests de facto belong to the church, by law, they belong to the state.
Despite the 1974 separation of church and state, Orthodox Christianity is the most
popular religion in Ethiopia (41%), followed by Islam, (35%), Protestantism (20%)
and traditional/indigenous religions followed by a minority of the population (3%)
(FDRE Population Census Commission 2008). The distribution of religion through-
out Ethiopia is clumped, with many regions overwhelmingly Muslim or Orthodox
536 M. Baez Schon et al.

(FDRE Population Census Commission 2008). Across northern Ethiopia, where our
research is based, the majority (83%) of the population is Orthodox Christian and
EOTC church forests are at the center of social and religious life; essential for both
religious and secular outreach (e.g., FDRE Population Census Commission 2008;
Klepeis et al. 2016; Cardelús et al. 2017; Orlowska and Klepeis 2018). We chose
Northern Ethiopia as our focus of study for three main reasons: (1) Orthodox
Christianity is dominant in the region, (2) all of the region’s forests are EOTC
church forests (Cardelús et al. 2017) and (3) northern Ethiopia is home to the
country’s second largest city, Bahir Dar, which finds itself at the center of much of
the country’s rapid economic and infrastructure development plans (Fig. 21.1).

Fig. 21.1 Map of the study area in South Gondar Administrative Zone (SGAZ), Amhara National
Regional State, Ethiopia. (Reproduced from Cardelús et al. (2019))
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 537

21.3 Community Use of EOTC Church Forests in Northern

Ethiopia

21.3.1 Religious Functions

Sacred forests, of the EOTC and other religions, are an integral part of Ethiopian
culture and are considered the interface between the divine and the mortal (Berhane-
Selassie 2008; Kent and Orlowska 2018). Each sacred forest is believed to be located
in a divinely chosen place (Kent and Orlowska 2018). Some stories proclaim that
when Jesus was crucified in Jerusalem and his blood dispersed throughout the world,
areas anointed with his blood were deemed suitable for the establishment of EOTC
churches. Other stories state that the founding priest of each church, while wander-
ing felt a stone or wooden replica of the tabot, the Ark of the Covenant given to
Moses at Mount Sinai, grow incredibly heavy as they reached the sacred spot upon
which the church should be constructed (Klepeis et al. 2016). The main function of
the church at the center of the forest is to house the sacred tabot.
For the EOTC community, the forests themselves are not the objects of worship,
nor are they protected to maintain biodiversity. Rather, their main function is to act
as a shield to the church and the tabot, which is generally found in the center of the
church in the middle of the forest. The forest becomes more sacred as one nears the
tabot and acts as a shroud to protect it and the church within which it is contained
(Klepeis et al. 2013). Thus, in EOTC tradition, the forests themselves may not have
the intrinsic religious value often associated with sacred groves in other contexts
where individual trees are sacred (Verschuuren et al. 2010).
EOTC church forests are also the resting place for the dead, both men and women
(Kent and Orlowska 2018). Within the EOTC tradition, the souls of the dead are not
properly at rest until they are buried within a sacred forest (Klepeis et al. 2016).
Church forests also have springs with holy water, either inside the forest itself, or on
adjacent land. This sacred water is used by community members to cure ailments,
from headaches to complex neurological and psychological problems. The strength,
and thus capacity for curing ailments, varies among holy springs.
EOTC church forests are also training sites for young priests and provide shelters
for monks and nuns (Kent and Orlowska 2018). In the traditional EOTC church
schools, young boys are housed in wooden huts within the church forests. They are
trained to reach a high level of asceticism, or avoidance of all forms of indulgence by
spending hours daily learning Ge’ez, an ancient Semitic language used in EOTC
liturgy, practicing EOTC rituals, and begging for food (Kent and Orlowska 2018).
Learning the teachings of the EOTC, regardless of whether a boy becomes a priest, is
a way to increase their social standing in the community (Kent and Orlowska 2018).
EOTC church forests also provide shelter for monks and nuns who renounce all
worldly things, fast extensively, and practice sexual abstinence in order to achieve a
level of asceticism and purity that makes them closer to the divine (Kent and
Orlowska 2018; Orlowska and Klepeis 2018). Women can become nuns only after
menopause as menstruation is considered impure (Kent and Orlowska 2018). Monks
538 M. Baez Schon et al.

and nuns are often elders whom turn to the church for shelter and food. In the church,
however, as monks and nuns, they maintain their respectability (Orlowska and
Klepeis 2018, pg 6).

21.3.2 Financial and Social Functions of EOTC Church

Forests

The Public Ownership of Rural Lands Proclamation of 1975 decreased the abilities
of churches to perform income-generating activities such as selling grass on
non-forested lands. This significantly reduced church income, requiring a diversifi-
cation of church revenue production. In particular, priests became more reliant on
donations from the community and sacred forests became supplemental sources of
income through the cultivation of fast growing and economically profitable plants
such as Eucalyptus, Cupressus, and coffee (Klepeis et al. 2016; Cardelús et al.
2019). The church forests also provide community members with other products
such as honey, medicinal plants, and wood for fuel (Cardelús et al. 2017).
Church forests are critical areas for community interactions. The forest has a
series of paths and clearings where community members congregate throughout the
week to meet in “Mehabirs”, or social subgroups of the congregation. Areas where
Mehabirs meet are called “Mehabir clearings”. These gathering spaces are often the
only shaded meeting spots in the community. Mehabir meetings vary in purpose
from planning burials to hosting feasts (Klepeis et al. 2016). Throughout the week,
community members use the space to seek peace, preach, and volunteer for com-
munity service. Due to the centrality of church forests in communities, government
stakeholders also use these areas for community outreach, including for health
campaigns (Fig. 21.2) (Orlowska and Klepeis 2018) and even the local HIV positive
support group (Authors’ Pers. Obs.). Overall, Klepeis et al. (2016) describe these
forests as the centers of the community, a place of interaction between humans and
the divine, a meeting place for the community, and a place that facilitates interactions
among community members and secular authorities. EOTC church forests are
therefore important for the conservation of forests, as well as for the role they play
as anchors that sustain a community’s cultural identity, making them biocultural
islands (Bhagwat and Rutte 2006; Cardelús et al. 2012). As such, despite the
top-down nature of the EOTC in Ethiopia, each EOTC church forest is unique due
to its specific uses by the surrounding community.

21.3.3 Governance

Forest use is regulated in three main ways—through “wugz” or “wugzet” (a curse),

by hired guards, or by community enforcement (Klepeis et al. 2016; Kent and
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 539

Fig. 21.2 An EOTC church forest near Bahir Dar illustrates how sacred forests can function as the
centers of the community. (Photo: C. Cardelús)

Orlowska 2018). The “wugzet” is a curse executed by the priests which stipulates
that any transgressor of forest use would face the wrath of God (Klepeis et al. 2016).
Protection mechanisms also include fines, time in jail and excommunication, with
punishments escalating. Other deterrents come in the form of official fines for cutting
trees, allowing cattle into the forest, etc. These rules are not evenly enforced among
forests, illustrating the strong effect of community on forest management (Klepeis
et al. 2016; Reynolds et al. 2017). In fact, variation in plant species composition was
also found to vary across “Woreda” (or district) regardless of size, which indicates a
strong and direct connection between the local community management of their
church forests, their species composition, and the health and long-term persistence of
these forests (Woods et al. 2020).
Given the historical structuring of governance of the EOTC, in which the
Patriarch of the church holds the most power, there is a hybrid top-down and
bottom-up forest governance system that relies on both the Patriarch and the locals
(Klepeis et al. 2016). EOTC church forests have continued to serve as biodiversity
refuges integral to the Ethiopian cultural identity (Berhane-Selassie 2008). It is
evident that religion in this case acts as an effective “social fence” (Bhagwat and
540 M. Baez Schon et al.

Rutte 2006), and the combination of religious and utilitarian worth makes them
valuable to the community, ensuring their preservation.

21.4 Landscape Analysis Status

21.4.1 EOTC Church Forest Status and Edge Effects

Across the South Gondar Administrative Zone (SGAZ) of northern Ethiopia

(14,607 km2), there are around 1022 EOTC forests; surprisingly, these forests are
almost uniformly distributed at around 2 km intervals (Cardelús et al. 2013). While
these forests are protected, their small average size (approximately 5 ha) and high
level of human activity has resulted in high disturbance; on average forests have 56%
disturbance, characterized by trails, structures, graves, plantations (e.g., coffee,
Eucalyptus), etc. (Cardelús et al. 2019). Across 46 sacred forests, disturbance was
not found to be correlated with proximity to population centers but was correlated
with forest size: the smaller the forest, the greater the disturbance. This disturbance
negatively impacts plant species richness, forest biomass, and forest regeneration
(Cardelús et al. 2019). Despite the high levels of disturbance, EOTC church forests
are valuable repositories of biodiversity in an otherwise denuded landscape (Aerts
et al. 2016; Woods et al. 2020). The resilience of these forests is compromised by the
slow decay of the surrounding matrix of shrubs and remnant trees, not by defores-
tation within the forest itself (Fig. 21.3; Scull et al. 2017).
The resilience of forests in South Gondar is also seen in other EOTC church
forests in South Western highlands where maintenance of the church forests has
continued to be demonstrated despite forest area decreasing by 37% concurrently
with agriculture increasing by 109% (1995–2010) (Daye and Healey 2015). The loss
of the landscape matrix, which acted as limited buffers, exposes these forests to
greater edge effects, threatening their integrity and regeneration capacity. Further-
more, it eliminates refugia for pollinators and dispersers, such as perch sites for
birds, and reduces movement of dispersers between forests, limiting the dispersal of
fruits and seeds, compromising forest regeneration (Cordeiro et al. 2009; Woods
et al. 2020). While until now sacredness was enough to protect these forests, the loss
of the matrix is compromising the forests and the communities that rely on them.

21.4.2 Forest Regeneration

These church forests are the remaining “Noah’s arks” for many indigenous species
in the region. Up to 168 woody plant species have been recorded in these forests,
95% of which are endemic (Wassie et al. 2010). However, the presence of large
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 541

Fig. 21.3 An example of an

EOTC sacred forest in the
South Gondar
Administrative Zone
(SGAZ) (38.028 N,
11.903 E) shown in 1938
and 2015 (DigitalGlobe, by
way of ESRI). Note there is
minimal decline of woody
biomass within the church
forest compared to a
significant decline in woody
biomass in the area
surrounding the church
forest. Scale is 1:10,000.
(Source: Scull et al. 2017)

canopy trees could be a mirage of a healthy forest if there are no seedlings of

old-growth trees in the understory.
Tree regeneration in these forests is compromised by human activity and cattle
entering the forests off the main trails (Cardelús et al. 2019; Wassie et al. 2010), as
well as by the limited number of tree species represented in the seed bank (Teketay
and Granström 1995; Wassie and Teketay 2006). In response to these disturbances,
and to better demarcate church land, some EOTC church forests have built a small
wall around their church, which has resulted in a marginally healthier seedling
community (Woods et al. 2017).
542 M. Baez Schon et al.

21.5 Threats to EOTC Church Forests: Ecological, Social,

and Cultural

Ethiopian Orthodox Tewahido church forests are important examples of how SNS
can be islands of biocultural diversity in human modified landscapes. As with many
SNS that have been historically protected, shifts in the social, cultural, and economic
landscapes can challenge the very traditional institutions which have ensured their
longstanding protection. Ethiopia has one of the fastest growing economies in the
world (Shiferaw 2017) with significant investment from foreign governments, par-
ticularly China (Chakrabarty 2016; US Department of State 2019). This has led to an
increase in infrastructure projects throughout the country, including roads as well as
a planned highway that would connect Djibouti with Bahir Dar in the South Gondar
Administrative Zone, where our research takes place (Chakrabarty 2016; Cheru
2016).
These shifts in economic development have the potential to impact sacred natural
sites in Ethiopia through their effect on the local population (Chandrakanth et al.
2004; Daye and Healey 2015; Reynolds et al. 2017). For example, in a chronological
study (2002 and 2014) of four EOTC church forests in the South Gondar Adminis-
trative Zone, Reynolds et al. (2017) found that increasing wealth is often associated
with more elaborate cement burial tombs and larger churches. These changes
increase disturbance within the already small church forests (Klepeis et al. 2016;
Reynolds et al. 2017). The replacement of native trees with economically beneficial
Eucalyptus around and within church forests has reduced soil quality and
compromised future forest expansion (Cardelús et al. 2019; Cardelús et al. 2020).
We also found native species planted within the forests for economic benefit,
including Juniperus, used as building material, coffee (Coffea), and Rhammus
which is used for making the traditional beer called “Tella” (Cardelús et al. 2019).
However, due to the small size of EOTC church forests, even low levels of
disturbance have significant effect on their ecological integrity and regenerative
capacity.
Alongside rapid economic growth are social and cultural changes (Reynolds et al.
2017). In the EOTC church forest context, the most notable example of weakening
community protection of church forests is the shift away from the expectations that
local communities and church leaders are responsible for forest protection to the
expectancy that church leaders and secular authorities are in charge (Reynolds et al.
2017). Due to their isolation, high levels of disturbance, and low seedling recruit-
ment, EOTC church forests may be ecologically “living dead” such that when the
last large trees die, so will the forests (Cardelús et al. 2019). Despite this, EOTC
church forests are still at the center of social and religious life. Church forests still
have several factors aiding in their protection. They have “grassroots participation,
have sociocultural legitimacy, and have long demonstrated ecological efficacy”
(Sheridan 2008, pg 10), which are all essential for sustainable conservation.
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 543

21.6 Global SNS: Social, Economic, and Cultural Threats

and Opportunities

Sacred natural sites (SNS) historically have served as successful forms of local
biocultural conservation, their protection rooted in local beliefs and the strength of
their religious leaders and guardians. However, SNS are not static and have adapted
and changed responding to economic, social and ecological pressures (Sheridan and
Nyamweru 2008; Ormsby 2012; Bossou et al. 2020). For some SNS these changes
have brought increased protection (See: Ormsby 2012), while many SNS are
threatened by social and religious change through the displacement of traditional
beliefs and power as well as by ambiguous property rights (Chandrakanth et al.
2004; Tengö et al. 2007; Ormsby 2012; Bossou et al. 2020).
The dynamic nature of SNS can provide sources of adaptation that allow for
continued or even increased protection, despite ongoing social and cultural changes
that may threaten traditional beliefs and power. In some cases, external support may
strengthen local customs (Ormsby and Edelman 2010; Ormsby 2012). An example
of this is the Tafi Atome Monkey Sanctuary, in Ghana (Ormsby and Edelman 2010;
Ormsby 2012). According to the locals, upon migrating to Tafi Atome they brought
with them their fetish, a local god, which they placed within the forest. This forest
immediately became sacred and protected by the community. Soon thereafter vil-
lagers noticed monkeys had followed them to their new home, a subspecies of the
mona monkey (Cercopithecus mona mona). The villagers in Tafi Atome believed
that the monkeys were representatives of the gods and therefore considered them
sacred and taboo to kill them. The sacred forest was protected for more than
200 years; however, the arrival of Christianity directly contradicted these local
customs. In an attempt to directly challenge traditional religious beliefs, local priests
encouraged the killing of the sacred monkeys and exploitation of the forest for
personal economic gains. In 1996, in order to provide a counter economic incentive
to protect the forest, the Taffi Atome community in collaboration with the Ghana
Tourist board and the Nature Conservation Research Centre—a leader in developing
rural ecotourism and community protected areas— established the Tafi Atome
Monkey Sanctuary. The ecotourism project reaffirms local traditions, protects the
sacred forest and the monkeys, and is also an economic boom for the town (Ormsby
and Edelman 2010; Ormsby 2012). This is an example of an SNS which, with
external help, stopped the erosion of local traditions and increased protection of their
sacred forest.
Sacred natural sites have the capacity to adapt and persist on the landscape; it is
important to note, however, that physical persistence does not guarantee ecological
integrity. A well-known example of this is the Ganges River of India. The Ganges
River is one of the most sacred rivers in Hinduism and considered the physical
manifestation of the divine goddess, Ganga, also providing water to 40% of India’s
population (Das and Tamminga 2012; Sachdeva 2017). The Ganges Basin is one of
the most populated areas of the world, and lack of widespread and adequate
sanitation infrastructure makes the Ganges one of the most heavily polluted rivers
544 M. Baez Schon et al.

in the world and a potential health hazard to users (Das and Tamminga 2012).
Despite the high levels of pollution, the Ganges remains sacred—bathing in it, for
example, is thought to cleanse the soul (Sachdeva 2017). The juxtaposition of
pollution and sacredness can prove challenging to conservation when the ecological
integrity of the SNS is disconnected from their sanctity in a way that can challenge
their very existence.

21.7 The Role of SNS in Preserving Biodiversity:

Conservation and Management

Ethiopian Orthodox Tewahido church forests are central to the religious, ecological,
and social fabric of the Ethopian Orthodox Tewahido church. Far from being fixed
cultural and religious relics spread throughout the Ethiopian landscape, these sacred
forests are complex and dynamic socio-ecological systems (Sheridan 2008). The role
of these forests within Ethiopia reflects larger global patterns whereby SNS have
successfully preserved biocultural diversity through protection by and for their
communities (Verschuuren et al. 2010).
The role of SNS in preserving biodiversity through culturally legitimate and
successful community forest management practices make them attractive to those
who seek to increase the network of protected habitats (Sheridan 2008; Wild and
McLeod 2008; Verschuuren et al. 2010). However, SNS are often inadvertent
(shadow) conservation sites which may limit their ability to protect biodiversity
and ecosystem services. Conservation initiatives involving SNS need to be collab-
orative, anchored within the community, involve all relevant stakeholders, and
follow guidelines such as IUCN’S Sacred Natural Sites: Guidelines for Protected
Areas Managers (Wild and McLeod 2008, (Bossou et al. 2020). This approach
would ensure that conservationists take into account the traditional beliefs that have
historically protected these forests and are important for the local cultural identity.
Sacred groves across Africa maintain unique and diverse communities of flora
and fauna. In Guinea-Bissau in western Africa, sacred groves hold a unique avifauna
of forest specialists and insectivorous birds relative to other forest types in the region
(Kühnert et al. 2019) and in Benin, riparian forests around sacred pools contain a
diversity of plant species endemic to western Africa (Ceperley et al. 2010). How-
ever, sacred groves are currently under threat from increased demand for arable land
due to increasing human populations, changes to cultural traditions and beliefs, and
climate change (Ormsby and Bhagwat 2010). Agricultural expansion remains the
largest threat to the conservation of sacred groves. In Benin, a replacement of
shifting cultivation in forest patches to cashew as a cash crop has resulted in a
reduction in biodiversity and increased food insecurity (Temudo and Abrantes
2014).
The maintenance of local-level resource management through continued tradi-
tional beliefs and social cohesion rather than the reliance on external development or
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 545

conservation initiatives are essential to the conservation of sacred groves. Examples

of successful local-level resource management include, for example, the sacred
forest of Nvangkpe on the border of Nigeria and Cameroon which has the largest
and rarest trees in the region and has survived because it is valued for its spiritual,
ancestral and magical worth amidst the pressures of agricultural expansion
(Kamanda et al. 2003). In Benin, the appreciation of sacred pools by local commu-
nities and the continued education of newly arrived migrants about social norms has
conserved the riparian forests around these pools from overfishing and cattle grazing
(Ceperley et al. 2010). These conservation successes reinforce the importance of
local custom and local reinforcement of traditions.
However, conservation and forest management in the absence of community
input and inclusion of indigenous narratives of protection will likely not succeed
(Bongers et al. 2006; Orlowska and Klepeis 2018; Temudo 2012). There are many
cautionary tales in which good intentions were not successful. For example in
Guinea-Bissau in western Africa, the intention of Non-Governmental Organization
(NGO) conservationists to create a National Park (known as “fortress conservation”)
led to a weakening of local institutions and social cohesion because local conserva-
tion management practices were denied and local stakeholders fought each other to
secure their livelihoods (Temudo 2012). Thus, the continued practices of local
communities for sacred forest protection must be supported.

21.8 Conclusion

Sacred natural sites may not widely be considered as important contributors to

biodiversity conservation, but in some regions where religion occupies a central
role, such as the South Gondar Region of northern Ethiopia, they are biodiversity
islands. Due to their spiritual status, sacred forests in South Gondar are readily
protected by the Ethiopian Orthodox Tewahido Church and the surrounding com-
munities, which has allowed forest cover to remain at relatively constant levels as
nearby land has been cleared for a growing agriculture economy. While SNS are not
primarily protected because of ecological concerns, they do offer a potentially
successful alternative to more traditionally recognized conservation strategies, par-
ticularly in regions where there may be few options. However, the successful
conservation requires cooperation between the church, nearby communities, and
any other relevant groups such as government actors who have ties to the SNS.
Without involvement from all parties that rely on or use the land and ensure its
protection, the rising economic and sociocultural pressures of the region will lead to
the degradation of the few remaining islands of biodiversity in Ethiopia, potentially
having serious implications to the land’s environmental and spiritual significance.

Acknowledgements We are grateful to the respected Ethiopian priests and monks who allowed us
access to their sacred church forests. We would like to acknowledge the students of the Cardelús
and Scull labs at Colgate University, the Woods lab at the University of Puget Sound, and the
546 M. Baez Schon et al.

Tsegay lab at the University of Bahir Dar, Ethiopia, for their contributions to the lab and field work.
We want to thank Ben Everett-Lane for his editorial work and feedback on our revisions. We also
want to thank our reviewers Dr. Alison Ormsby and Dr. Keith Kirby. This work was funded through
the Dynamics of Coupled Natural Human Ecosystems Program of the National Science Foundation
[Award No. 1518501] and by the National Science Foundation Graduate Research Fellowship
[Award No. DGE 1842473]. Initial work was funded by The Picker Interdisciplinary Science
Institute at Colgate University: [Link]
ary-science-institute.

References

Aerts R, Van Overtveld K, November E, Wassie A, Abiyu A, Demissew S, Daye DD, Giday K,
Haile M, TewoldeBerhan S, Teketay D, Teklehaimanot Z, Binggeli P, Deckers J, Friis I,
Gratzer G, Hermy M, Heyn M, Honnay O, Paris M, Sterck FJ, Muys B, Bongers F, Healey
JR (2016) Conservation of the Ethiopian church forests: threats, opportunities and implications
for their management. Sci Total Environ 551–552:404–414. [Link]
2016.02.034
Altman N (2002) Sacred water: the spiritual source of life. Paulist Press
Ancel S, Ficquet É (2015) The Ethiopian orthodox Tewahido church (EOTC) and the challenges of
modernity. In: Prunier G, Ficquet E (eds) Understanding contemporary Ethiopia: monarchy,
revolution, and the legacy of Meles Zenawi. Hurst and Company, London, pp 65–91
Berhane-Selassie T (2008) The socio-politics of Ethiopian sacred groves. In: Sheridan MJ,
Nyamweru C (eds) African sacred groves: ecological dynamics and social change. Ohio
University Press, Athens
Bhagwat SA, Rutte C (2006) Sacred groves: potential for biodiversity management. Front Ecol
Environ 4:519–524. [Link]
Bongers F, Wassie A, Sterck F, Bekele T, Teketay D (2006) Ecological restoration and church
forests in northern Ethiopia. J Dry Lands 1:35–44
Bossou B, Lokossou O, Assongba M, Agbangla M (2020) Saving the sacred. Yokohama
Cardelús CL, Lowman MD, Eshete AW (2012) Uniting church and science for conservation.
Science 80(335):2006–2008
Cardelús CL, Scull P, Hair J, Baimas-George M, Lowman MD, Eshete AW (2013) A preliminary
assessment of ethiopian sacred grove status at the landscape and ecosystem scales. Diversity 5:
320–334. [Link]
Cardelús CL, Scull P, Wassie Eshete A, Woods CL, Klepeis P, Kent E, Orlowska I (2017) Shadow
conservation and the persistence of sacred church forests in northern Ethiopia. Biotropica 49:
726–733. [Link]
Cardelús CL, Woods CL, Mekonnen AB, Dexter S, Scull P, Tsegay BA (2019) Human disturbance
impacts the integrity of sacred church forests, Ethiopia. PLoS One 14:1–14. [Link]
1371/[Link].0212430
Cardelús CL, Mekonnen AB, Jensen KH, Woods CL, Baez MC, Montufar M, Bazany K, Tsegay
BA, Scull P, Peck WH (2020) Edge effects and human disturbance influence soil physical and
chemical properties in sacred church forests in Ethiopia. Plant Soil 329–342. [Link]
1007/s11104-020-04595-0
Ceperley N, Montagnini F, Natta A (2010) Significance of sacred sites for riparian forest conser-
vation in Central Benin. Bois Forets des Trop 303(1):5–23. [Link]
303.a20450
Chakrabarty M (2016) Ethiopia – China economic relations. A Classic Win – Win Situation? 7:
226–248
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 547

Chandrakanth MG, Bhat MG, Accavva MS (2004) Socio-economic changes and sacred groves in
South India: protecting a community-based resource management institution. Nat Resour Forum
28:102–111. [Link]
Cheru F (2016) Emerging southern powers and new forms of South–South cooperation: Ethiopia’s
strategic engagement with China and India. Third World Q 37:592–610. [Link]
1080/01436597.2015.1116368
Colding J, Folke C (2001) Social taboos: “invisible” systems of local resource management and
biological conservation. Ecol Appl 11:584–600. [Link]
0584:STISOL][Link];2
Cordeiro NJ, Ndangalasi HJ, McEntee JP, Howe HF (2009) Disperser limitation and recruitment of
an endemic African tree in a fragmented landscape. Ecology 90:1030–1041. [Link]
1890/07-1208.1
Das P, Tamminga KR (2012) The ganges and the GAP: an assessment of efforts to clean a sacred
river. Sustainability 4:1647–1668. [Link]
Daye DD, Healey JR (2015) Impacts of land-use change on sacred forests at the landscape scale.
Glob Ecol Conserv 3:349–358. [Link]
Delgado F, Escobar C, Verschuuren B, Hiemstra W (2012) Sacred natural sites, biodiversity and
Well-being: the role of sacred sites in endogenous development in the COMPAS network.
Sacred Nat Sites Conserv Nat Cult 188–197. [Link]
Doffana ZD (2019) Sacred sites and ancestor veneration in Sidama, Southwest Ethiopia: a socio-
ecological perspective. Cogent Soc Sci 5. [Link]
Dudley N, Higgins-Zogib L (2012) Protected areas and sacred nature: a convergence of beliefs. In:
Pungetti G, Oviedo G, Hooke D (eds) Sacred species and sites: advances in biocultural
conservation. Cambridge University Press, New York, pp 36–45
Dudley N, Higgins-Zogib L, Mansourian S (2009) The links between protected areas, faiths, and
sacred natural sites. Conserv Biol 23:568–577. [Link]
01201.x
FDRE Population Census Commission (2008) Summary and statistical report of the 2007 popula-
tion and housing census: population size by age and sex. Addis Ababa
Frascaroli F, Zannini P, Acosta ATR, Chiarucci A, d’Agostino M, Nascimbene J (2019) Sacred
natural sites in Italy have landscape characteristics complementary to protected areas: implica-
tions for policy and planning. Appl Geogr 113:102100. [Link]
102100
Harmon D, Putney A (2003) The full value of parks: from the economics to the intangible. Rowman
& Littlefield Publishing Group, Lanham
Kamanda B, Angu AK, Nguinguiri J-C (2003) The social value of the Nyangkpe scared forests of
south West Province of Cameroon. In: Harmon D, Putney AD (eds) The full value of parks:
from economics to intangible. Rowman & Littlefield Publishing Group, Lanham, pp 77–89
Kapos V (1989) Effects of isolation on the water status of Forest patches in the Brazilian Amazon. J
Trop Ecol 5:173–185
Karbo T (2013) Religion and social cohesion in Ethiopia. Int J Peace Dev Stud 4:43–52. [Link]
org/10.5897/ijpds2013.0164
Keeley J, Scoones I (2000) Knowledge, power and politics: the environmental policy-making
process in Ethiopia. J Mod Afr Stud 38:89–120. [Link]
Kent EF, Orlowska I (2018) Accidental environmentalists. Worldviews Environ Cult Relig 22:113–
142. [Link]
Klepeis P, Scull P, Lalonde T, Svajlenka N, Gill N (2013) Changing forest recovery dynamics in the
northeastern United States. Area 45:239–248. [Link]
Klepeis P, Orlowska IA, Kent EF, Cardelús CL, Scull P, Wassie Eshete A, Woods C (2016)
Ethiopian church forests: a hybrid model of protection. Hum Ecol 44:715–730. [Link]
10.1007/s10745-016-9868-z
548 M. Baez Schon et al.

Kühnert K, Grass I, Waltert M (2019) Sacred groves hold distinct bird assemblages within an
Afrotropical savanna. Glob Ecol Conserv 18:e00656. [Link]
e00656
Malcolm JR (1994) Edge effects in central Amazonian Forest fragments author. Ecol Soc Am 75:
2438–2445
McCann JC (1997) The plow and the Forest: narratives of deforestation in Ethiopia, 1840-1992.
Environ Hist 2:138–159
Millenium Ecosystem Assessment (2010) Ecosystems and human Well-being: biodiversity synthe-
sis. Ecosystems 285. [Link]
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity Islands: strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human-dominated environments, Series: Topics in Biodiversity and Con-
servation. Springer, Cham, pp 1–35
Notermans C, Nugteren A, Sunny S (2016) The changing landscape of sacred groves in Kerala
(India): a critical view on the role of religion in nature conservation. Religions 7. [Link]
10.3390/REL7040038
Orlowska I, Klepeis P (2018) Ethiopian church forests: a socio-religious conservation model under
change. J East African Stud 12:674–695. [Link]
Ormsby A (2012) Cultural and conservation values of sacred forests in Ghana. In: Sacred species
and sites: advances in biocultural conservation. Cambridge Univ Press, Cambridge, pp 335–350
Ormsby AA, Bhagwat SA (2010) Sacred forests of India: a strong tradition of community-based
natural resource management. Environ Conserv 37:320–326. [Link]
S0376892910000561
Ormsby A, Edelman C (2010) Community-based ecotourism at Tafi atome monkey sanctuary, a
sacred natural site in Ghana. In: Pungetti G, Oviedo G, Hooke D (eds) Sacred natural sites:
conserving nature and culture. EarthScan, London, pp 233–243
Oviedo G, Jeanrenaud S, Otegui M (2005) Protecting sacred natural sites of indigenous and
traditional peoples: an IUCN perspective
Pungetti G (2012) Sacred species and sites: dichotomies, concepts and new directions in biocultural
diversity conservation. In: Pungetti G, Oviedo G, Hooke D (eds) Sacred species and sites:
advances in biocultural conservation. Cambridge University Press, New York, pp 13–27
Pungetti G, Oviedo G, Hooke D (eds) (2012) Sacred species and sites advances in biocultural
conservation. Cambridge University Press, New York
Reynolds TW, Stave KA, Sisay TS, Eshete AW (2017) Changes in community perspectives on the
roles and rules of church forests in northern Ethiopia: evidence from a panel survey of four
Ethiopian orthodox communities. Int J Commons 11(355–387). [Link]
ijc.707
Rutte C (2011) The sacred commons: conflicts and solutions of resource management in sacred
natural sites. Biol Conserv 144:2387–2394. [Link]
Sachdeva S (2017) The influence of sacred beliefs in environmental risk perception and attitudes.
Environ Behav 49:583–600. [Link]
Scariot A (2000) Seedling mortality by litterfall in Amazonian forest fragments. Biotropica 32:662–
669. [Link]
Scull P, Cardelús CL, Klepeis P, Woods CL, Frankl A, Nyssen J (2017) The resilience of Ethiopian
church forests: interpreting aerial photographs, 1938–2015. L Degrad Dev 28:450–458. https://
[Link]/10.1002/ldr.2633
Sheridan MJ (2008) The dynamics of African sacred groves: ecological, social & symbolic
processes. In: Sheridan MJ, Nyamweru C (eds) African sacred groves: ecological dynamics
and social change. Ohio University Press, Athens
Sheridan M, Nyamweru C (2008) African sacred groves: ecological dynamics & social change.
James Currey Publisher, Oxford
Shiferaw A (2017) Productive capacity and economic growth in Ethiopia. CDP Backgr Pap No
34 ST/ESA/2017/CDP/34 6:1–17
21 Sacred Church Forests in Northern Ethiopia: Biodiversity and Cultural Islands 549

Sigman E (2022) Safeguarding the benefits of biodiversity islands in northern Ethiopia in the midst
of political change. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments, Series: Topics in Biodiversity and Conservation. Springer,
Cham, pp 647–674
Teketay D, Granström A (1995) Soil seed banks in dry Afromontane forests of Ethiopia. J Veg Sci
6:777–786. [Link]
Temudo MP (2012) “The white men bought the forests”: conservation and contestation in Guinea-
Bissau, Western Africa. Conserv Soc 10:354–366. [Link]
Temudo MP, Abrantes M (2014) The cashew frontier in Guinea-Bissau, West Africa: changing
landscapes and livelihoods. Hum Ecol 42:217–230. [Link]
Tengö M, Johansson K, Rakotondrasoa F, Lundberg J, Andriamaherilala JA, Rakotoarisoa JA,
Elmqvist T (2007) Taboos and forest governance: informal protection of hot spot dry forest in
southern Madagascar. Ambio 36:683–691. [Link]
TAFGIP][Link];2
US Department of State (2019) 2019 investment climate statements: Ethiopia. [Link]
Verschuuren B, Wild R (2012) Sacred natural sites- sources of biocultural diversity
Verschuuren B, Wild R, McNeely J, Oviedo G (2010) Sacred natural sites: conserving nature and
culture. Earth Scan, Routledge
Wassie A (2002) Opportunities, constraints and prospects of Ethiopian orthodox tewahido churches
in conserving forest resources: the case of churches in South Gonder, Northern Ethiopia.
Swedish Univeristy of Agricultural Sciences, Uppsala
Wassie A, Teketay D (2006) Soil seed banks in church forests of northern Ethiopia: implications for
the conservation of woody plants. Flora Morphol Distrib Funct Ecol Plants 201:32–43. https://
[Link]/10.1016/j.flora.2005.04.002
Wassie A, Sterck FJ, Bongers F (2010) Species and structural diversity of church forests in a
fragmented Ethiopian Highland landscape. J Veg Sci 21:938–948. [Link]
1654-1103.2010.01202.x
Wild R, McLeod C (2008) Sacred natural sites: guidelines for protected area managers. IUCN,
Gland. [Link]
Woods CL, Cardelús CL, Scull P, Wassie A, Baez M, Klepeis P (2017) Stone walls and sacred
forest conservation in Ethiopia. Biodivers Conserv 26:209–221. [Link]
s10531-016-1239-y
Woods CL, Bitew Mekonnen A, Baez-Schon M, Thomas R, Scull P, Abraha Tsegay B, Cardelús
CL (2020) Tree community composition and dispersal syndrome vary with human disturbance
in sacred church forests in Ethiopia. Forests 11:1082. [Link]
Yigremew A (2002) Review of landholding systems and policies in Ethiopia under the different
regimes. Ethiopian economic association,/Ethiopian economic policy research institute, Addis
Ababa, Ethiopia. [Link]
Zerga B (2016) Land resource, uses, and ownership in Ethiopia: past, present and future. Int J Sci
Res Eng Technol 2:2395–2566
Chapter 22
Beyond the Island: Integrated Approaches
to Conserving Biodiversity Islands
with Local Communities

Michael S. Esbach, Mahi Puri, Robinson Botero-Arias, and Bette A. Loiselle

Abstract This chapter highlights how graduate students can work with local com-
munities and stakeholders to improve biodiversity conservation and rural develop-
ment outcomes both in and outside of biodiversity islands with examples from
Ecuador, India, and Brazil. In the first case study, Michael Esbach draws on his
long-term work with the Indigenous Cofán people in the Ecuadorian Amazon to
describe the creation of a biodiversity island within the Cofán territory of Zábalo. In
the second case study, Mahi Puri focuses on a human-dominated landscape in India
with a high occurrence of carnivores. Finally, Robinson Botero-Arias tells the story
of the development of sustainable use plans and conservation of wildlife in an
aquatic ecosystem in the Amazon. These case studies demonstrate the “learning
and action” platform of the interdisciplinary Tropical Conservation and Develop-
ment (TCD) program at the University of Florida (UF), which stresses the impor-
tance of working with partners and local communities to design problem-oriented
research to address challenges at the intersection of conservation and development.
Through formal and informal methods, TCD embodies the lessons learned from the
examples presented in this chapter, which include the linkage between biodiversity
conservation and human well-being; the necessity of understanding local needs,
cultural values, and motivations; and engaging stakeholders in partnerships that
center participatory research targeted at finding solutions that make the goals of
local actors and biodiversity conservation more compatible.

M. S. Esbach
Tropical Conservation and Development Program, Center for Latin American Studies,
University of Florida, Gainesville, FL, USA
High Meadows Environmental Institute, Princeton University, Princeton, NJ, USA
M. Puri · R. Botero-Arias · B. A. Loiselle (*)
Tropical Conservation and Development Program, Center for Latin American Studies,
University of Florida, Gainesville, FL, USA
Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL, USA
e-mail: bloiselle@latam.ufl.edu

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 551
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
552 M. S. Esbach et al.

Keywords Interdisciplinary approaches · Partnership · Participatory research ·

Stewardship

22.1 Introduction

The past several decades have seen significant changes in the spatial distribution of
intact forests and the patterns of land use surrounding these landscapes (Hansen et al.
2013; Kim et al. 2015; Grantham et al. 2020). These processes have resulted in
numerous biodiversity islands: biologically diverse, ecologically intact refugia that
are often surrounded by a matrix of human dominated and transformed land.
Conserving biodiversity islands, including the ecological functions and services
found within, becomes increasingly challenging as they become smaller and more
isolated from each other (as per MacArthur and Wilson 1967; Damschen et al. 2019;
Montagnini et al. 2022). Scientific studies have often focused on island character-
istics and the ways in which the matrix can both positively and negatively impact
these islands (e.g., Renjifo 2001). These studies, however, often fail to integrate
important social, economic, and cultural contexts, which may drive conservation
outcomes within and surrounding biodiversity islands. Ignoring these human dimen-
sions also makes bridging the gap between theory and practice extremely challeng-
ing. In this chapter, we argue that scientific research needs to move beyond the island
in order to support conservation within the island. By embracing multi-stakeholder
participatory processes and partnerships, researchers can develop studies that sup-
port conservation science and local action (Duchelle et al. 2009; Kainer et al. 2019).
We focus on the role graduate (doctoral) students can play in strengthening
such work.
Designing and conducting problem-oriented dissertation studies that respond to
local concerns, meet requirements of academic advisors and committees, and bridge
theory and practice is exceedingly difficult to achieve. Success will depend on a
number of underlying mechanisms that empower and prepare graduate students to
collaborate with local actors in research processes, integrate social and biological
sciences, and communicate research results to a number of audiences, including
peer-reviewed journals and local stakeholders. The interdisciplinary learning and
action framework of the Tropical Conservation and Development (TCD) program at
the University of Florida (UF) was developed to catalyze and enable a generation of
young scientists to embrace new approaches to biodiversity conservation research
(Kainer et al. 2019). This framework focuses on the development of theoretical
foundations and skills necessary to work across disciplines and effectively collabo-
rate and communicate with local stakeholders. Several TCD courses focus on the
development and application of key skills (e.g., communication, facilitation, conflict
management), which empower students to effectively listen, learn, and act with key
local actors (Duchelle et al. 2009).
22 Beyond the Island: Integrated Approaches to Conserving. . . 553

Dissertation studies offer an opportunity to put these skills to use. Student

researchers are encouraged to establish partnerships with local communities and
governmental or non-governmental organizations to better understand the
social-ecological context and to more effectively generate and exchange knowledge.
Below we identify important principles for effective partnerships. The case studies
that follow highlight these principles in practice.
• Build Social Capital: In order to build strong relationships, it is important to
spend time with your partners, often outside the context of a specific research
agenda. By focusing on issues with a clear motivation for partners (e.g., access to
certain kinds of information or training), students can establish trust and a record
of engagement prior to initiating a research project.
• Acknowledge Diversity: It is important to recognize that different parties will have
different motives, objectives, and values. Student research should consider mech-
anisms for incorporating this diversity, assessing trade-offs, and creating equita-
ble conflict resolution mechanisms (Gavin et al. 2018).
• Share Power: Effective partnerships require the sharing of power between
parties. In addition to utilizing diverse participatory methods (e.g., Reason and
Bradbury 2001; Reed 2008), students are encouraged to co-design and implement
research with local partners. Sharing power in this way encourages joint respon-
sibility and active relationship management (Mead 2013).
• Encourage Long-Term Commitments: It is important to recognize that effective
partnerships are built upon long-term commitments that can be resource-intensive
and transcend the timeline of graduate research. We encourage students to
carefully consider this prior to initiating new relationships. At the same time,
students may be able to build upon relationships they already have or work in
collaboration with ongoing opportunities. This allows them to capitalize on
previous work and provide continuity to that progress.
The remainder of this chapter uses case studies to highlight how graduate students
can leverage diverse skills and partnerships to conduct research in support of
biodiversity islands. The three case studies are diverse and include (1) efforts by
an Amazonian Indigenous community to simultaneously maintain subsistence liveli-
hoods and wildlife populations in Ecuador, (2) efforts to establish a sustainable
working landscape surrounding a tiger reserve which benefits small landowners
while increasing suitable habitat for large predators in India, and (3) management
and sustainable harvest of a threatened top predator by local communities in a
sustainable development reserve in the Brazilian Amazon. While different, each
case study highlights common themes: leveraging existing local partnerships, work-
ing across diverse stakeholders with different objectives, and student researchers
utilizing their unique professional skills, experience, and history in their particular
study area.
554 M. S. Esbach et al.

22.2 Case Studies

We present three case studies in the following section. In the first case study,
Michael Esbach draws on his long-term work with the Indigenous Cofán people in
the Ecuadorian Amazon to describe the creation of a biodiversity island within the
Cofán territory of Zábalo. The emergence and persistence of this reserve area
represents a social process whereby the Cofán developed coordinated, collective
arrangements to maintain resources within their territory from generation to gener-
ation. Using ethnographic research and common property theory, he examines Cofán
common property institutions, critical components of Indigenous tenure regimes that
shape conservation outcomes like biodiversity islands. In the second case study,
Mahi Puri focuses on a human-dominated landscape in India with a high occurrence
of carnivores. Limited research on carnivore ecology outside protected areas (PAs)
has hindered conservation efforts. Concurrently, for marginalized farmers, losses
due to fluctuating weather patterns, conflict with wild animals, and minimal govern-
ment support reinforces issues of food insecurity. Combining ecological models on
species distribution with concepts from environmental economics, Mahi explores
potential solutions that could contribute to biodiversity conservation in partnership
with private landowners in shared landscapes. The final case study tells the story of
the development of sustainable use plans and conservation of wildlife in an aquatic
ecosystem in the Amazon. Here, Robinson (Robin) Botero-Arias has taken on a
leadership role as part of the staff of the Mamiraua Institute for Sustainable Devel-
opment in Brazil. As part of his dissertation research and building on his and other
long-term population ecology studies of black caiman, a large crocodilian species, he
describes the social, ecological, and political context of the Mamiraua Sustainable
Development Reserve and the efforts to develop a caiman harvesting program with
local communities.

22.2.1 Indigenous Stewardship of a Biodiversity Island

in Ecuador

The Amazon Basin is the largest tropical rainforest in the world, widely recognized
for its biological and cultural diversity (Jenkins et al. 2013). This cultural diversity
includes over 370 Indigenous ethnicities that have inhabited and managed the region
for millennia (Denevan 1992). Despite significant challenges, including coloniza-
tion, disease, and dispossession, Indigenous peoples have successfully established
property rights to over 25 percent of the Amazon (RAISG 2016). Within these
territories, Indigenous peoples have created biodiversity islands that host high
biodiversity (Pretty et al. 2009; Stevens 2014), prevent deforestation and wildfires
(Gray and Coates 2010; Blackman et al. 2017, Schleicher et al. 2017), and maintain
the culture and livelihoods of Indigenous peoples (Garnett et al. 2018).
22 Beyond the Island: Integrated Approaches to Conserving. . . 555

The social processes that create biodiversity islands within Indigenous territories
are complex and understudied (Bremner and Lu 2006). In response, the case study
presented here focuses on the emergence and maintenance of a biodiversity island
within the Cofán territory of Zábalo in the Ecuadorian Amazon. This research was
made possible through a long-term partnership between Michael Esbach and the
Cofán people. His approach to partnership was informed by previous experiences
working with Indigenous peoples across the globe, as well as through diverse
learning experiences provided through UF’s TCD program. This approach, as
noted in the introduction, focuses on building social capital, acknowledging diverse
motives and objectives, power sharing, and a long-term engagement.
Prior to initiating his research, Michael regularly visited with Cofán people in
order to build social capital and learn more about the history of Zábalo. During this
time he focused on supporting Cofán initiatives not directly related to his dissertation
research. By focusing on Cofán needs and interests, such as securing funds for their
ongoing initiatives, he was able to establish a record of engagement and build trust
with the community. Over time, Michael developed a plan of research in partnership
with the Cofán that responded to their interests and future goals. Throughout this
process, he managed expectations by recognizing that each party should benefit, but
in potentially different ways. For instance, while Michael’s research was aligned
with Cofán interests, he also recognized the importance of paying Cofán residents
for their time and support and seeking out training opportunities to further their local
capacity. At the same time, Michael benefitted from the Cofán’s expertise and
participation in his dissertation. Ultimately, Michael and his Cofán partners were
able to explore local resource management institutions, ecological knowledge, and
trends in animal populations over time. This work directly supports the Cofán’s
desire to improve self-determination over resource management within their terri-
tory. This is important because it shows the effectiveness of local institutions in
maintaining animal populations over time. Beyond these accomplishments, Michael
continues to work with the Cofán on a number of initiatives, deepening his partner-
ship with the Cofán.
The remainder of this case study is dedicated to results from this research
partnership, which explores how the Cofán created institutions to conserve and
protect a biodiversity island within their territory of Zábalo. One of five Cofán
territories with legal title, Zábalo is located in the northeastern Ecuadorian Amazon
within the Cuyabeno Wildlife Reserve (Fig. 22.1). Like the majority of Indigenous
territories in the Ecuadorian Amazon, Zábalo is owned collectively by the Cofán.
This communal tenure regime signals that resources within the territory are managed
through formal and informal rules known as common property institutions (Bremner
and Lu 2006). These institutions govern how resources are managed in Zábalo,
allowing Cofán residents to enforce rules among themselves and exclude external
individuals from using resources found within the territory (Ostrom 1990).
Zábalo was founded by residents from Dureno, another Cofán community, in the
early 1980s in response to agrarian reform initiated by the Ecuadorian state which
opened the Cofán’s ancestral lands to colonization. This process supported the
emergence of the Cofán common property institutions that regulate resource use
556 M. S. Esbach et al.

Fig. 22.1 Map of Zábalo in the Ecuadorian Amazon depicting the biodiversity island stewarded by
the Cofán people. Resource use activities are prohibited within this area, and additional, specific
rules have been institutionalized within the hunted area. Map and illustration created by Michael
Esbach using freely available data and images

and promote conservation. Research suggests that for these institutions to emerge,
people need to recognize that resources important to their livelihoods are becoming
scarce and that they are capable of reducing the pressures that create this scarcity
(Bromley and Cernea 1989; Lu 2001). As colonists increasingly encroached on their
traditional territory and degraded the surrounding environment, the Cofán realized
that their forest-based livelihood was dependent upon an intact environment and
fought to preserve it through three primary methods: (1) a secure land title, (2) the
ability to exclude outsiders, and (3) the importance of managing resource use.
22 Beyond the Island: Integrated Approaches to Conserving. . . 557

Across the Amazon, one of the primary drivers for Indigenous peoples to secure
legal property rights are external pressures on important resources (Lu 2001). Zábalo
residents first filed for legal property rights in 1981. It was not until 1992, however,
with the expansion of the Cuyabeno Wildlife Reserve, that the Cofán were able to
come to an agreement with the Ministry of Environment (MAE). This
co-management agreement established the Cofán’s legal title to 85,000 hectares in
exchange for their management of the area. The Cofán would go on to sign
additional agreements with the MAE, increasing their territorial rights to approxi-
mately 145,000 hectares. Today there are about 180 people living in Zábalo,
representing nearly 40 families. While some of these families live in more isolated
areas, most are clustered into four locations along the Aguarico River. Zábalo
residents hunt across their territory, fish within both white-water and black-water
rivers, and practice agriculture near their homes and within individual plots scattered
along the Aguarico River.
With their land title secure, Cofán leaders crafted approaches to defend their
territory from outside threats. This included the creation of the Cofán park guard
program, which focuses on patrolling Zábalo’s boundaries, clearing trails, and
manning guard stations in order to keep outsiders from extracting resources. Cofán
leaders have demonstrated the effectiveness of this program, and as a result, the
national government signed two official documents (Acuerdo No. 138 in 2002 and
Registro Official No. 5 in 2003) in support of the Cofán park guard program.
Currently, all Zábalo residents participate in the guard program, conducting patrols
and clearing boundaries on a yearly basis. Group of men and women are deployed to
different regions of the territory to conduct these responsibilities, and different
families are assigned to live in guard stations for months at a time.
In addition to a secure land title and the ability to exclude outsiders, the Cofán
also developed a sophisticated resource management system within Zábalo called
se’pi’cho (see Cepek 2012). Over the past 30 years, the community of Zábalo has
experimented with se’pi’cho in order to manage their subsistence activities in a way
that sustains communal resources. These efforts have resulted in an effective system
of adaptive management whereby daily subsistence acts as a type of monitoring.
Residents, individually and collectively, develop detailed knowledge related to
changes in resources, including the distribution of animal populations and their
abundance across the territory. This knowledge is shared annually during a commu-
nity meeting when Zábalo residents discuss their observations. During this meeting,
they have the opportunity to create and/or revise rules based on their knowledge.
Proposed changes will be voted on by the community and take written form if the
majority approve. The rule is then applied to all community members and enforced
strictly through different fine structures and public acknowledgement when a resi-
dent breaks a rule. The interaction of these components—where observation repre-
sents a form of monitoring and knowledge production that informs rules with
revisions over time based on additional observations—is the crux of adaptive
management.
558 M. S. Esbach et al.

Se’pi’cho rules currently take diverse forms, including users, activities, and
species-specific restrictions (see Cepek 2012). The Cofán have also developed
spatial boundaries which recognize areas for human use, primarily along the
Aguarico River (e.g., houses, gardens) and subsistence activities where hunting,
fishing, and gathering are permitted, as well as a reserve area which acts as a source
for wildlife populations (Fig. 22.1). This reserve area functions as a biodiversity
island where animals and plants are protected from human use, but tourism and
scientific research activities are allowed. Most of the land base in the territory of
Zábalo comprises a biodiversity island whereby community members utilize diverse
strategies, from adaptive management to the exclusion of outsiders, to sustain
biodiversity that is important to Cofán livelihoods and identity. The creation and
sustainability of biodiversity islands within Indigenous territories, therefore, is a
social process whereby Indigenous peoples secure and enforce communal property
rights and develop institutions to manage resources in culturally appropriate ways.

22.2.2 Buffering Impacts to a Biodiversity Island in India

India is a megadiverse country with approximately 20% of its land forested

(Mittermeier et al. 2004). Less than five percent of its land is protected, however.
Most of India’s biodiversity islands, or PAs, were created from princely hunting
grounds or reserve forests established during the colonial era (Rangarajan 1996). As
a result, the approximately 800 PAs scattered across the country are small,
fragmented, and isolated, making them less than optimal in terms of their ecological
capacity and potential (Nayak et al. 2020). These PAs are also vulnerable to
expanding infrastructure, agricultural development, and economic activities such
as mining, dams, and commercial logging (Jayadevan et al. 2020).
The foundation of PAs in India is the Wildlife Protection Act (1972), which
spurred the creation of many biodiversity islands through the relocation of human
settlements (Rangarajan 2005; Rangarajan and Shahabuddin 2006). Limited science-
based conservation planning, along with the exclusion of local communities from
decision-making (Kothari et al. 1995), has contributed to conflicts over the manage-
ment of resources. Suspension of the traditional rights of local communities over
forests and restrictions on access has resulted in resentment towards the government
and conservation policies (Maikhuri et al. 2001). In addition to these challenges, the
high density of people and livestock around PAs has resulted in numerous human-
wildlife conflicts (Bagchi and Mishra 2006; Karanth et al. 2012). Unsustainable
resource use has also contributed to forest degradation and exacerbated negative
impacts on wildlife populations (Shankar et al. 1998; Kumar and Shahabuddin 2005;
Davidar et al. 2010).
Given their mixed success in achieving ecological objectives, exclusion of local
peoples, and mounting development threats, government and non-government orga-
nizations across India have started experimenting with alternative conservation
strategies (Saberwal and Rangarajan 2003; Sinha et al. 2012). In this case study,
22 Beyond the Island: Integrated Approaches to Conserving. . . 559

Mahi Puri first explores the conservation potential of legally sanctioned community
reserves and other voluntary community-led initiatives, and presents results from her
own research working directly with private landowners living in a buffer around a
biodiversity island.
In 2003, an amendment to the Wildlife Protection Act allowed for the establish-
ment of community reserves. Today, there are over 100 community reserves where
local users have greater access to resources which are administered by local people
and elected village institutions (Kothari 2006). While limited research has been
conducted on these reserves, results show that they significantly outperform open-
access areas in terms of biodiversity conservation and can play a complementary role
to strict PAs (Shahabuddin and Rao 2010). In terms of governance, community
reserves have had varying degrees of success due to bureaucratic inefficiency
(Pathak 2006). Despite being included in the country’s PA system, several reserves
continue to face economic and political constraints.
Other forms of communally managed forests exist in India that do not have the
same legal status as community reserves. Across the country, community forest
resource (CFR) rights have been recognized under the Forest Rights Act (2006),
allowing communities to conserve and manage forest resources. Management plans
developed by communities are diverse, responding to local environments and needs.
Communities have initiated measures for soil and water conservation, carried out fire
management, imposed restrictions on grazing activities in areas under assisted
natural regeneration, removed invasive species, and even set aside areas to allow
wildlife presence. Communities have experimented with short-rotation species
(including bamboo and various native fruiting trees) that provide annual economic
returns and have been able to generate substantial incomes through the sustainable
harvest of non-timber forest products. These initiatives have contributed to employ-
ment creation (especially for the landless) and empowerment of local village-level
institutions and women’s collectives (for detailed case studies see Agarwal and
Saxena 2018). While some CFR areas have seen tremendous success, many others
continue to be challenged by a lack of tenure rights, power imbalances with the forest
authorities, conflicts with neighboring villages, and crop damage by wild animals.
In addition to community-led initiatives, there are also private landowners and
organizations assisting in forest regeneration and restoration. In the Kodagu district
of Karnataka, Save Animals Initiative (SAI) Sanctuary is a private forest managed by
a non-profit trust. Elsewhere, ecotourism-based initiatives such as Kumaon Maati
and Gorukana channel their revenues back into conservation or for the benefit of the
rural poor (Puri et al. 2019). However, most of these projects have been independent
initiatives, conducted at small spatial scales and with limited involvement of local
people. The need to expand wildlife habitat, and increased effectiveness of the
existing PA network must be reconciled with the fact that the land outside PAs is
largely privately owned. This mandates incorporating the social context in ecolog-
ical studies in order to reduce the gap between conservation goals and their current
feasibility.
560 M. S. Esbach et al.

Natural regeneration on degraded or modified lands is a slow and long-term

process. The process can be sped up through active restoration and intensive
management. Adoption of agroforestry systems that support food production and
provide other resources (such as fodder, firewood) is a potential strategy that can
benefit local livelihoods and concurrently enhance the biodiversity value of the
hostile landscape surrounding PAs by maintaining connectivity across landscapes
and reducing pressures on PAs (Montagnini et al. 2022). However, the success of
such an intervention is contingent upon the decisions and choices of individual
landowners (Knight et al. 2010). Within this context, Mahi Puri undertook a study
rooted in a social-ecological systems approach (Liu et al. 2007; Carter et al. 2014) to
examine the (1) habitat use patterns of a carnivore assemblage (including tigers,
leopards, and wild dogs) in a human-dominated landscape, and (2) willingness of
farmers to modify existing land-use by adopting agroforestry, if provided with a
monetary incentive.
The study was conducted in the administrative buffer of Pench Tiger Reserve, a
biodiversity island in central India (Fig. 22.2). The site was selected based on Mahi’s
long-term work in the region, specifically her research on landscape connectivity and
human-carnivore conflicts. Prior discussions with the local Forest Department and
the partnering NGO also helped in identifying specific objectives for the region. The
field studies were comprised of camera trap surveys in approximately 500 square
kilometers of multi-use forests, and interviews with over 600 farmers. At the data
collection stage, Mahi employed members from the local communities. Engagement
of local people in the research project helped in building trust with other local
farmers who participated in the surveys. The larger goal of this research project
was to prioritize restoration in the landscape and establish an incentive-based
conservation initiative, which integrates ecological values with the preferences of
key decision-makers (i.e., private landowners).
The results from the study demonstrate that at fine spatial scales, habitat use by
carnivores was positively influenced by the composition and configuration of the
landscape. With regard to modifying current land-use, farmer decision-making was
positively influenced by their education and current income from agriculture, and
negatively influenced by household size. In addition, the percentage of forest cover
around the landholding and recurring incidents of crop damage by wild herbivores
contributed to people’s willingness to adopt agroforestry. Variables such as people’s
perceptions of benefits from agroforestry, perceived risks, as well as social norms
were also important driving factors. Interdisciplinary studies such as this could be
useful in exploring the potential for carnivore conservation and collaboration with
local stakeholders within a forest-agriculture matrix. This could provide valuable
information for identifying areas where conservation interventions might be pursued
and succeed. Ultimately, it could aid in landscape level planning that integrates
agricultural lands with natural areas, concurrently contributing to farmer incomes
through an incentive-based program.
Moving forward, bridging the gap between scientific research and conservation
action will require a combination of both bottom-up and top-down approaches.
Participatory methods that consider local needs and preferences are an essential
component for supporting local stewardship. To overcome economic barriers of
22 Beyond the Island: Integrated Approaches to Conserving. . . 561

Fig. 22.2 Pench Tiger Reserve in central India is surrounded by a buffer area that supports the
conservation of key carnivore species and the livelihoods of local farmers. Map and illustration
created by Michael Esbach using freely available data and images

restoration, government incentives, private investments, and support from NGOs

will be crucial. An important tool in this regard is payment for ecosystem services
(PES). Unlike many countries around the world where government support exists, a
formal PES system is currently absent in India. Another important consideration for
the program’s effectiveness is the need for structured mechanisms such as insurance
and compensation schemes. These mechanisms may reduce and/or mitigate conflicts
as people practice agroforestry and their interactions with wildlife increase. In the
long-term, improved habitat quality that encourages wildlife presence can create
opportunities to promote alternative forms of tourism in such landscapes. This could
help offset tourism pressure on PAs, diversify existing tourism activities, and
contribute to income generation. Ultimately, integrating restored lands through
community-led initiatives and wildlife-friendly agriculture on private lands with
the country’s extant PA network has the potential to minimize some developmental
challenges, thereby accomplishing the dual objectives of improving landscape
conservation and securing local livelihoods.
562 M. S. Esbach et al.

22.2.3 Sustainable Management of an Aquatic Biodiversity

Island in Brazil

The Brazilian Amazon supports high biodiversity and immense reservoirs of carbon
and water (Mittermeier et al. 2005; Ritter et al. 2017). The aquatic wildlife and river
systems found across this region, which can be viewed as “islands” in forested
landscapes, are extremely important from both social and ecological perspectives
(Antunes et al. 2016). To safeguard these resources, Brazil has developed an
extensive PA network which covers approximately 28% of the Amazon region.
This network includes different categories of PAs which utilize different approaches
to conserve biodiversity, including strict nature conservation, sustainable develop-
ment, tourism, scientific research, and education (Fearnside 2003; Silva 2005; de
Queiroz 2010). Tailored to unique social-ecological contexts, this system is able to
include local people in PA governance and support the sustainable use of natural
resources through management planning and zoning systems (e.g., strictly protected
zones and harvest areas) (Gillingham 2001; Rylands and Brandon 2005; de Queiroz
2010). This case study focuses on Sustainable Development Reserves (SDR), a
Brazilian PA category that integrates local stakeholders into conservation priorities.
In this context, Robinson Botero-Arias discusses his work with black caiman
(Melanosuchus niger) and local communities within the Mamiraua SDR.
Brazil’s SDRs utilize both scientific and local knowledge to create conservation
priorities and management strategies. By facilitating the involvement of local people
and utilizing their knowledge base, SDRs integrate diverse knowledge systems to
establish guidelines for natural resource use and zoning of activities within a specific
area. The zoning plan, for example, must take into consideration the subsistence
needs of local communities, together with ecological requirements for sustainable
use and biodiversity conservation (Castello 2008; Castello et al. 2009; Marioni et al.
2013a, b). The Mamiraua SDR has implemented a successful pirarucu (Arapaima
gigas) sustainable management program with local stakeholders. In response to
overfishing by external parties, this program focused on building a participatory
monitoring system of the fishery (Castello et al. 2009). This system has effectively
incorporated local communities in the conservation of this large fish species. Rec-
ognizing the success of the pirarucu participatory management program, caimans
were identified as an ideal target species for further participatory action given local
needs and conservation threats. Given Robin’s leadership role in international
crocodilian conservation and knowledge of local systems, his dissertation research
in collaboration with Mamiraua SDR was designed to advance this sustainable
management goal.
Populations of black caimans have suffered from past overexploitation. Intense
hunting began between 1940 and 1950 and then continued, though less intensively,
through the 1970s. Until the 1990s, the black caiman was considered one of the most
threatened Neotropical crocodilians due to unsustainable use, habitat loss and
degradation, and over-hunting by local fishermen (Da Silveira and Thorbjarnarson
1999; Thorbjarnarson and Da Silveira 2000; Marioni et al. 2013a, b). Given the
22 Beyond the Island: Integrated Approaches to Conserving. . . 563

importance of black caiman to local livelihoods, as both a source of food and

income, the Brazilian government sought ways to sustainably harvest this species.
A key first step was the down-listing of the black caiman from CITES (Convention
on International Trade in Endangered Species) Appendix I to Appendix II in 1999;
this change signaled international support for establishing a caiman management
system. Shortly thereafter, the Brazilian State of Amazonas initiated an event in
partnership with scientific and local communities that aimed to develop a caiman
conservation, monitoring, and harvest management program. The Mamiraua SDR
was chosen as the site for a pilot program, and was chosen for two main reasons:
(1) the existence of ongoing monitoring actions with evidence of a high abundance
of black caimans, and (2) a demonstrated capacity of local communities to engage in
effective, sustainable use of natural resources (such as with the pirarucu management
program) (Botero-Arias et al. 2010).
Five years after the creation of Mamiraua SDR in 1996, local populations of black
caiman began to recover, suggesting that supervised harvests might successfully
generate income for local residents without negative, long-term impacts on the wild
population. Robinson Botero-Arias has been collaborating with the Caiman Con-
servation and Management Research Program at the Mamiraua Institute for Sustain-
able Development since 2007. There he has supported processes to establish a
framework for legal caiman harvesting. This legal framework was strengthened
via two resolutions in 2011 that established (1) technical procedures for caiman
management in PAs, and (2) criteria for the harvest and processing of crocodilians,
both in the State of Amazonas. The foundation of this framework is a participatory
process that integrates conservation principles with local knowledge and needs.
Specifically, adaptive management principles help maintain stable and healthy
caiman populations over time by incorporating feedbacks from caiman monitoring,
various sustainability indicators, and local participation. Through this system, black
caiman populations have demonstrated significant resilience to harvest (Botero-
Arias and Regatieri 2013; Marioni et al. 2013a, b). The sustainable use of Amazo-
nian caimans has become an important management strategy to conserve the species
and preserve its habitats.
By working with local communities, government institutions, and other
researchers, Robin has supported the sustainability of caiman harvests within the
Mamiraua SDR. Participatory processes have proven critical to these efforts. By
working directly with local people that have historically benefited from the caiman
harvest to develop different zones (e.g., harvesting areas, reserve areas; Fig. 22.3),
his work has supported local decision-making that directly supports local needs. His
work also points to the importance of adaptive management performed through
partnerships between local communities and researchers that draw on different
knowledge systems and values. This partnership has supported the integration of
biological research on harvested species with the social and economic needs of local
peoples through a constant participatory evaluation process that combines sustain-
ability indicators and caiman population monitoring. Robin’s work has also sought
to support local communities by connecting them with regional supply chains so that
they can develop a fair market for caiman with competitive pricing. This way, local
564 M. S. Esbach et al.

Fig. 22.3 The Mamiraua Sustainable Development Reserve functions as a biodiversity island
focused on the conservation and sustainable management of black caiman and pirarucu. Map and
illustration created by Michael Esbach using freely available data and images

people can get the best prices for their harvested meat, which simultaneously
reinforces local benefits and motivation to sustainably manage caimans and their
habitat.
Management systems for the use of the natural resources within SDRs have
positioned local socio-cultural values at the center of the decision-making process,
with the complementary aim of conserving biodiversity and decreasing illegal
activities threatening wildlife populations. Despite its success over the past decade,
caiman management in the State of Amazonas is still considered experimental. As
such, Robin will continue to work with his partners in Mamiraua SDR through his
affiliation with the Mamiraua Institute for Sustainable Development to conduct
further collaborative research focused on producing information about the
populations to be exploited, the long-term sustainability of their harvest, and various
social and organizational challenges that arise when working in a complex social-
ecological system.
22 Beyond the Island: Integrated Approaches to Conserving. . . 565

22.3 Conclusions

Conserving biodiversity islands, like those described in this chapter, requires the
development of approaches that facilitate multi-stakeholder inclusion and participa-
tion in order to address underlying research needs and integrate research findings
into participatory decision-making processes. Graduate students can play an impor-
tant role in developing and advancing such interdisciplinary research like that
described here which responds to local concerns while contributing to finding viable
solutions for biodiversity conservation in complex social-ecological systems that
characterize “islands” of native habitat.
The case studies presented here represent the dissertation work of three graduate
students with considerable knowledge of local systems in which they conducted
their research. Although the contexts are different, each student took on research
questions that addressed local needs in partnership with local communities, conser-
vation NGOs, and/or government actors. The case studies highlight (1) the impor-
tance of existing partnerships with local communities and NGOs to identify issues
threatening biodiversity, (2) application of interdisciplinary problem-oriented
research to better understand threats and potential solutions, and (3) current or
proposed mechanisms to share results with local stakeholders to advance solutions
appropriate to particular social-ecological contexts. Outcomes from their research
included enhanced understanding of mechanisms by which Indigenous peoples
successfully manage wildlife populations in community reserves in Ecuador, iden-
tification of opportunities for increasing habitat for large carnivores in rural India that
generate alternative income sources via more wildlife-friendly agriculture, and the
development of participatory management systems for the sustainable use of aquatic
wildlife populations in Amazonian PAs. This work was supported by the learning
and action platform emphasized through the TCD program at UF. TCD graduate
students like Michael, Mahi, and Robin were provided opportunities to enrich their
professional skills by working across disciplines and engaging diverse stakeholders
to develop problem-oriented research questions in partnership with local peoples,
NGOs, and other partners and with whom they could jointly develop more holistic
solutions to the conservation of biodiversity islands.

References

Agarwal S, Saxena AK (2018) People’s forests: is community forest resource governance the future
of India’s jungles? Centre for Science and Environment, New Delhi
Antunes AP, Fewster RM, Venticinque EM, Peres CA, Levi T, Rohe F, Shepard GH (2016) Empty
forest or empty rivers? A century of commercial hunting in Amazonia. Sci Adv 2(10):e1600936
Bagchi S, Mishra C (2006) Living with large carnivores: predation on livestock by the snow leopard
(Uncia uncia). J Zool 268(3):217–224
Blackman A, Corral L, Lima ES, Asner GP (2017) Titling indigenous communities protects forests
in the Peruvian Amazon. Proc Natl Acad Sci 114(16):4123–4128
566 M. S. Esbach et al.

Botero-Arias R, Regatieri SA (2013) Construindo as bases para um Sistema de Manejo

Participativo dos Jacarés Amazônicos. Série Prot, Tefé, Amazonas, Brazil
Botero-Arias R, Marmontel M, Queiroz HL (2010) Projeto de manejo experimental de Jacarés no
Estado Do Amazonas: Abate de Jacarés no setor Jarauá - Reserva de Desenvolvimento
Sustentável Mamirauá, Dezembro de 2008. Scientific Magazine UAKARI 5(2):49–57
Bremner J, Lu F (2006) Common property among Indigenous peoples of the Ecuadorian Amazon.
Conserv Soc 4(4):499–521.
Bromley DW, Cernea MM (1989) The management of common property natural resources: some
conceptual and operational fallacies. World Bank discussion paper 57, World Bank,
Washington DC
Carter NH, Viña A, Hull V, McConnell WJ, Axinn W, Ghimire D, Liu J (2014) Coupled human and
natural systems approach to wildlife research and conservation. Ecol Soc 19(3):43
Castello L (2008) A socio-ecological synthesis on the conservation of the pirarucu (Arapaima) in
floodplains of the Amazon. PhD dissertation, State University of New York, Syracuse
Castello L, Viana JP, Watkins G, Pinedo-Vasquez M, Luzadis VA (2009) Lessons from integrating
fishers of arapaima in small-scale fisheries management at the Mamirauá Reserve. Amazon
Environ Manage 43(2):197–209
Cepek ML (2012) A future for Amazonia: Randy Borman and Cofán environmental politics.
University of Texas Press, Texas.
Da Silveira R, Thorbjarnarson JB (1999) Conservation implications of commercial hunting of black
and spectacled caiman in the Mamiraua sustainable development reserve, Brazil. Biol Conserv
88:103–109
Damschen EI, Brudvig LA, Burt MA, Fletcher RJ Jr, Haddad NM, Levey DJ, Orrock JL, Resasco J,
Tewksbury JJ (2019) Ongoing accumulation of plant diversity through habitat connectivity in
an 18-year experiment. Science 365(6460):1478–1480
Davidar P, Sahoo S, Mammen PC, Acharya P, Puyravaud JP, Arjunan M, Garrigues JP, Roessingh
K (2010) Assessing the extent and causes of forest degradation in India: where do we stand?
Biol Conserv 143(12):2937–2944
de Queiroz HL (2010) Protected areas in the Amazonian Várzea and their role in its conservation:
the case of Mamirauá Sustainable Development Reserve (MSDR). In: Junk W, Piedade M,
Wittmann F, Schöngart J, Parolin P (eds) Amazonian floodplain forests. Ecological studies
(analysis and synthesis). Springer, Dordrecht, pp 465–483
Denevan WM (1992) The pristine myth: the landscape of the Americas in 1492. Ann Assoc Am
Geogr 82(3):369–385
Duchelle AE, Biedenweg K, Lucas C, Virapongse A, Radachowsky J, Wojcik DJ, Kainer KA
(2009) Graduate students and knowledge exchange with local stakeholders: possibilities and
preparation. Biotropica 41(5):578–585
Fearnside PM (2003) Conservation policy in Brazilian Amazonia: understanding the dilemmas.
World Dev 31(5):757–779
Garnett ST, Burgess ND, Fa JE, Fernández-Llamazares Á, Molnár Z, Robinson CJ, Leiper I (2018)
A spatial overview of the global importance of indigenous lands for conservation. Nat Sustain
1(7):369–374
Gavin MC, McCarter J, Berkes F, Mead ATP, Sterling EJ, Tang R, Turner NJ (2018) Effective
biodiversity conservation requires dynamic, pluralistic, partnership-based approaches. Sustain-
ability 10:1846
Gillingham S (2001) Social organization and participatory resource management in Brazilian
Ribeirinho communities: a case study of the Mamirauá sustainable development reserve.
Amazonas Soc Nat Res 14(9):803–814
Grantham HS, Duncan A, Evans TD, Jones KR, Beyer HL, Schuster J, Walston JC, Robinson JG,
Callow M, Clements T, Costa HM, DeGemmis A, Elsen PR, Ervin J, Franco P, Goldman E,
Goetz S, Hansen A, Hofsvang E, Jantz P, Jupiter S, Kang S, Langhammer P, Laurance WF,
Lieberman S, Linkie M, Malhi Y, Maxwell S, Mendez M, Mittermeier R, Murray NJ,
Possingham H, Radachowsky J, Saatchi S, Samper C, Silverman J, Shapiro A, Strassburg B,
22 Beyond the Island: Integrated Approaches to Conserving. . . 567

Stevens T, Stokes E, Taylor R, Tear T, Tizard R, Venter O, Visconti P, Wang S, Watson JEM
(2020) Anthropogenic modification of forests means only 40% of remaining forests have high
ecosystem integrity. Nat Commun 11:5978
Gray M, Coates J (2010) ‘Indigenization’and knowledge development: extending the debate. Int
Soc Work 53(5):613–627
Hansen MC, Potapov PV, Moore R, Hancher M, Turubanova SA, Tyukavina A, Thau D, Stehman
SV, Goetz SJ, Loveland TR, Kommareddy A, Egorov A, Chini L, Justice CO, Townshend JRG
(2013) High-resolution global maps of 21st-century forest cover change. Science 342:850–853
Jayadevan A, Nayak R, Karanth KK, Krishnaswamy J, DeFries R, Karanth KU, Vaidyanathan S
(2020) Navigating paved paradise: evaluating landscape permeability to movement for large
mammals in two conservation priority landscapes in India. Biol Conserv 247:108613
Jenkins CN, Pimm SL, Joppa LN (2013) Global patterns of terrestrial vertebrate diversity and
conservation. Proc Nat Acad Scie USA 110(28):2602–2610
Kainer KA, Binnqüist CL, Dain JL, Jaimes BC, Castillo PN, Basulto RG, Contreras FM (2019)
Leading by listening, learning by doing: modeling democratic approaches to conservation
leadership in graduate education. J Environ Stud Sci 9(2):206–217
Karanth KK, Gopalaswamy AM, DeFries R, Ballal N (2012) Assessing patterns of human-wildlife
conflicts and compensation around a Central Indian protected area. PLoS One 7:e50433
Kim D, Sexton JO, Townshend JR (2015) Accelerated deforestation in the humid tropics from the
1990s to the 2000s. Geophys Res Lett 42:1e7
Knight AT, Cowling RM, Difford M, Campbell BM (2010) Mapping human and social dimensions
of conservation opportunity for the scheduling of conservation action on private land. Conserv
Biol 24:1348–1358
Kothari A (2006) Community conserved areas: towards ecological and livelihood security. Parks
16(1):3–13
Kothari A, Suri S, Singh N (1995) Conservation in India: a new direction. Econ Polit Wkly 30:
2755–2766
Kumar R, Shahabuddin G (2005) Effects of biomass extraction on vegetation structure, diversity
and composition of forests in Sariska Tiger Reserve. India Environ Conserv 32(3):248–259
Liu J, Dietz T, Carpenter SR, Folke C, Alberti M, Redman CL, Schneider SH, Ostrom E, Pell AN,
Lubchenco J, Taylor WW (2007) Coupled human and natural systems. Ambio 36(8):639–649
Lu F (2001) The common property regime of the Huaorani Indians of Ecuador: implications and
challenges to conservation. Human Ecol 29(4):425–447
MacArthur RH, Wilson EO (1967) Island biogeography. Princeton University Press, Princeton
Maikhuri RK, Nautiyal S, Rao KS, Saxena KG (2001) Conservation policy–people conflicts: a case
study from Nanda Devi Biosphere Reserve (a world heritage site). India For Pol Econ 2(3):
355–365
Marioni B, Botero-Arias R, Fonseca-Junior SF (2013a) Local community involvement as a basis for
sustainable crocodilian management in protected areas of Central Amazonia: problem or
solution? Tropical Conserv Sci 6(4):484–492
Marioni B, Farias I, Verdade LM, Bassetti L, Coutinho ME, de Mendonça SH, Vieira TQ,
Magnusson WE, Campos Z (2013b) Avaliação do risco de extinção do jacaré-açu
Melanosuchus niger (Spix, 1825) no Brasil. Biodiv Brasil 3(1):31–39
Mead ATP (2013) Sharing power: Māori and protected areas. In: Katene S, Mulholland M (eds)
Governance and leadership in future challenges for Māori: He Kōrero Anamata. Huia Pub-
lishers, Wellington, pp 195–204
Mittermeier RA, Gil PR, Hoffmann M, Pilgrim J, Brooks T, Lamoreux CM, Da Fonseca GAB
(2004) Hotspots revisted: Earth’s biologically wealthiest and most threatened ecosystems.
CEMEX, México, pp 99–103
Mittermeier RA, Da Fonseca GA, Rylands AB, Brandon K (2005) A brief history of biodiversity
conservation in Brazil. Conserv Biol 19:601–607
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity Islands: strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
568 M. S. Esbach et al.

for conservation in human-dominated environments, Topics in Biodiversity and Conservation.

Springer, Cham, pp 1–35
Nayak R, Karanth KK, Dutta T, Defries R, Karanth KU, Vaidyanathan S (2020) Bits and pieces:
Forest fragmentation by linear intrusions in India. Land use policy:104619
Ostrom E (1990) Governing the commons: the evolution of institutions for collective action.
Cambridge University Press, Cambridge
Pathak N (2006) Community conserved areas in South Asia. Protected Areas Programme 56
Pretty J, Adams B, Berkes F, De Athayde SF, Dudley N, Hunn E, Pilgrim S (2009) The
intersections of biological diversity and cultural diversity: towards integration. Conserv Soc
7(2):100–112
Puri M, Karanth KK, Thapa B (2019) Trends and pathways for ecotourism research in India. J
Ecotour 18(2):122–141
RAISG (Rede Amazônica de Informação Socioambiental Georreferenciada) (2016) Amazonia
2016: protected areas and indigenous territories. Amazon Network of Geo-Referenced Socio-
Environmental Information. [Link]
Rangarajan M (1996) The politics of ecology: the debate on wildlife and people in India, 1970-95.
Econ Polit Wkly:2391–2409
Rangarajan M (2005) India’s wildlife history: an introduction. Orient Blackswan, Hyderabad
Rangarajan M, Shahabuddin G (2006) Displacement and relocation from protected areas: towards a
biological and historical synthesis. Conserv Soc 4(3):359–378
Reason P, Bradbury H (2001) Handbook of action research: participative inquiry and practice. Sage
Publications, London
Reed MS (2008) Stakeholder participation for environmental management: a literature review. Biol
Conserv 141:2417–2431
Renjifo LM (2001) Effect of natural and anthropogenic landscape matrices on the abundance of
SubAndean bird species. Ecol Appl 11:14–31
Ritter CD, McCrate G, Nilsson RH, Fearnside PM, Palme U, Antonelli A (2017) Environmental
impact assessment in Brazilian Amazonia: challenges and prospects to assess biodiversity. Biol
Conserv 206:161–168
Rylands AB, Brandon K (2005) Brazilian protected areas. Conserv Biol 19(3):612–618
Saberwal VK, Rangarajan M (2003) Battles over nature: science and the politics of conservation.
Orient Blackswan, Hyderabad
Schleicher J, Peres CA, Amano T, Llactayo W, Leader-Williams N (2017) Conservation perfor-
mance of different conservation governance regimes in the Peruvian Amazon. Sci Rep 7(1):
1–10
Shahabuddin G, Rao M (2010) Do community-conserved areas effectively conserve biological
diversity? Global insights and the Indian context. Biol Conserv 143(12):2926–2936
Shankar U, Murali KS, Shaanker RU, Ganeshaiah KN, Bawa KS (1998) Extraction of non-timber
forest products in the forests of Biligiri Rangan Hills, India. 4. Impact on floristic diversity and
population structure in a thorn scrub forest. Econ Bot 52(3):302–315
Silva M (2005) The Brazilian protected areas program. Conserv Biol 19(3):608–611
Sinha BC, Qureshi Q, Uniyal VK, Sen S (2012) Economics of wildlife tourism – contribution to
livelihoods of communities around Kanha tiger reserve, India. J Ecotour 11:207–218
Stevens S (ed) (2014) Indigenous peoples, national parks, and protected areas: a new paradigm
linking conservation, culture, and rights. University of Arizona Press, Tucson
Thorbjarnarson J, Da Silveira R (2000) Secrets of the flooded Forest. Nat Hist 109(2):70–79
Chapter 23
Agroecology and Forest Conservation
in Three Types of Land Reform
Communities in the Cacao Region of Bahia,
Brazil

Kathleen R. Painter, Robert Buschbacher, Luiz Carlos Souto Silva,

and Emerentina Costa e Silva

Abstract To address conflicts between human occupation and forest conservation

in the cacao region of southern Bahia, Brazil, the Jupará Agroecological Movement,
in partnership with WWF, promotes an agroecological model of land reform. This
chapter presents results of this initiative, and assesses its contribution to two kinds of
biodiversity islands: diversified agroforestry systems and remnants of natural forest.
The chapter utilized interviews and visits to 60 families in three communities with
different forms of land occupation: “contested” and “negotiated” models of land
reform, and a “traditional community” created by escaped slaves (quilombo). Par-
ticipants and non-participants in the Jupará program are compared, with forest cover
changes documented using remote sensing imagery. Jupará promotes agroecological
practices, community organization and commercialization. Jupará participants
adopted three key agroecological practices: substituting organic fertilizer and pest
control for agrochemicals, investing in contour erosion strips, and implementing
agroforestry systems that incorporate biodiversity and ecological services. Mainte-
nance of native forest varies by community type. Land reform communities on
former cacao estates have fertile soil and can be highly productive while maintaining
a diverse native forest overstory (cabruca), also protecting remnants of native forest.

K. R. Painter
Institute for Social and Economic Development (ISED) Solutions, Garden City, ID, USA
[Link]
R. Buschbacher (*)
School of Forest Fisheries and Geomatics Sciences and Tropical Conservation and
Development Program, University of Florida, Gainesville, FL, USA
e-mail: rbusch@ufl.edu
L. C. Souto Silva
Jupará Agroecological Movement and COOPASB Cooperative, Bahia, Brazil
E. Costa e Silva
Associação Agricola do Projeto de Assentamento Frei Ventuy, Bahia, Brazil

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 569
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
570 K. R. Painter et al.

Land reform communities on less fertile soils require initial forest clearing. As these
communities develop, agroforestry systems increase; native forest cover initially
declines, but there is potential to recover and reconnect some remnants. These results
draw attention to contextual factors beyond the local scale of project implementa-
tion. While diversified agroforestry biodiversity islands were viable in all commu-
nities, protection of native forest remnants depends on public policies and
consideration of broader historical, geographic and biophysical drivers of land use
dynamics.

Keywords Agroforestry · Biodiversity islands · Cabruca · Forest conservation ·

Jupará agroecological movement · Land reform · Quilombo · WWF

23.1 Introduction

23.1.1 Competing Conservation Paradigms

According to a prevalent conservation paradigm, large blocks of undisturbed habitat

are the most effective approach to achieve conservation goals (Kramer et al. 1997;
Noss et al. 1999; Wuerthner et al. 2015; Dinerstein et al. 2017). Yet the social and
biological assumptions and implications of this approach are complex, and its
normative character is often unacknowledged. Biologically, one consequence of
the “large, intact, contiguous blocks” paradigm is that protected areas tend to be in
remote, steep, infertile and inaccessible regions that are less suitable for human
occupation (Hunter and Yonzon 1993; Scott et al. 2001; Joppa and Pfaff 2009, 2011;
Barnes et al. 2018; Putz 2020). The species composition of such regions is not likely
to be as rich as, nor representative of, the more anthropocized flat, lowland, fertile
habitats. Furthermore, the ecology of so-called “intact” natural areas may in fact be
highly influenced by past and present large and small-scale human activities
(Denevan 1992; Cronon 1996; Heckenberger et al. 2008; Heckenberger and Neves
2009; Pivello 2011; Ross 2011; Clement et al. 2015; Boivin et al. 2016; Levis et al.
2017, 2020; de Souza et al. 2018; Franco-Moraes et al. 2019; Shepard et al. 2020).
To illustrate the normative aspect of the large-block protection paradigm, we can
consider that while contemporary conservationists generally abhor fragmentation,
not very long ago “edge” habitats and ecotones were considered particularly valu-
able because they incorporate a diversity of habitats and consequently high local
species richness (Leopold 1933 cited in Silbernagel 2003; Odum 1971). In part, this
shift in emphasis is a normative change from valuing game species to seeking to
protect rare species that require a large range and are particularly vulnerable to
extinction (Laurance 2008; Laurance et al. 2018). Yet a recent review of frag-
mentation research (Fahrig 2017) found that biodiversity effects of habitat fragmen-
tation are usually confounded with effects of habitat loss and that 76% of significant
effects of fragmentation were positive. Arroyo-Rodríguez et al. (2022) carried
out a multi-taxonomic assessment of small patches of Lacandona rainforest in
Mexico and found that species density was as great in patches as in continuous
23 Agroecology and Forest Conservation in Three Types of Land. . . 571

forest, including for forest-specialist species. They conclude that preservation of all
forests is important to biodiversity, especially preservation of small patches in
human-dominated areas.
Besides these biological considerations, many human populations live in high
biodiversity areas, and an exclusionary conservation paradigm creates an inherent
conflict between conservation advocates and local people who depend on use of land
and management of natural resources for their livelihoods. The potential power and
wealth imbalances between external conservation advocates and local communities
raises ethical concerns and calls into question the long-term political viability of
conservation interventions (Chapin 2004; West et al. 2006; Brockington et al. 2008;
Brockington and Wilkie 2015; Holmes and Cavanagh 2016; Kopnina 2016; Kohler
and Brondizio 2017; Ybarra 2018).
An alternative conservation paradigm of community-based conservation has been
developed since the 1980s. There is strong empirical evidence that community
control of resources and habitat leads to equal or better conservation outcomes
than state-tenure fully protected areas (Charnley and Poe 2007; Bray et al. 2008;
Lele et al. 2010; Soares-Filho et al. 2010; Nelson and Chomitz 2011; Porter-Bolland
et al. 2012; Radachowsky et al. 2012; Wood et al. 2019; Elleason et al. 2020; Vélez
et al. 2020). Nevertheless, integrating biodiversity conservation with socioeconomic
development is challenging and context-specific. Furthermore, the alternative para-
digms or approaches may very well be complementary rather than competitive. The
concept of biodiversity islands provides ample room for both fully protected areas
and biodiversity protection within areas of human occupation (Montagnini
et al. 2022).

23.1.2 Key Questions and Overview of the Chapter

The purpose of this chapter is to present the experience, results and lessons from the
Jupará Agroecological Movement, which worked with land reform communities to
address the apparent conflict between forest protection and human occupation in the
cacao region of southern Bahia, Brazil. Using the perspective of biodiversity islands,
we analyze whether the initiative was effective in terms of land sharing and land
sparing (i.e. creation and maintenance of biodiversity islands through agroforestry
production systems, and maintenance of remnants of intact native forest within the
occupied landscape).
The implementation and effectiveness of any community-based conservation
initiative is highly context specific. Therefore, following this introduction Sect.
23.2 presents a historical overview of human occupation of the cacao region in
southern Bahia, and then describes the land reform process occurring in this region.
This overview provides the geographical, biological and social context of the
apparent conflict between forest conservation and human occupation in this region.
Section 23.3 presents the Jupará Agroecological Movement (hereafter shortened
to Jupará), which emerged, with the support of World Wildlife Fund (WWF), to
572 K. R. Painter et al.

promote an alternative, agroecological model of land reform in 35 land reform and

traditional agricultural communities throughout the cacao region of southern Bahia.
Jupará’s approach and results varied according to the characteristics of each com-
munity, and this study focuses on three communities representative of different types
of land occupation, which are also presented in Sect. 23.3.
Section 23.4 provides an overview of the research methodology, Sect. 23.5
presents the results of Jupara’s work in terms of adoption of agroecological prac-
tices, and Sect. 23.6 presents the dynamics of native forest and agroforestry cover in
the different communities. These results are presented both as a comparison between
participants and non-participants in Jupará activities from within the same commu-
nities, and by comparison among the three types of community.
We conclude with reflections on the viability of Jupará’s approach for creating the
two types of biodiversity islands within land reform communities in southern Bahia
(Sect. 23.7). Does the Jupará Agroecological Movement present a viable “proof-of-
concept” for agroecological production, community organization and commerciali-
zation to achieve forest conservation? How do the historical, ecological and geo-
graphic differences between different types of communities enable or limit the
adoption of agroecological practices and the protection of forest remnants? And
what are the implications of these findings for mobilizing public policy to produce
positive outcomes for ecosystems and communities?

23.2 Regional Context: Human Occupation and Land

Reform in the Cacao Region

23.2.1 Cacao and Human Occupation of Brazil’s Atlantic

Forest

Brazil’s Atlantic Forest is considered a global “hot spot” for biodiversity conserva-
tion (Ribeiro et al. 2011). While overall only 12% of this biome maintains its forest
cover (including regenerating areas and degraded forest, Ribeiro et al. 2011), it
covers a vast region with varied topography, vegetation, human occupation and
conservation status. WWF divides the Atlantic Forest into nine ecoregions (Olson
and Dinerstein 1998). Two of these, the Serra do Mar Coastal Forest and the Coastal
Forests of Bahia, maintain relatively higher levels of forest coverage, the former
because of relatively low human occupation, and the latter because of how humans
occupied and used the forest. In the Serra do Mar Coastal Forest ecoregion, steep
topography drove early colonization of southern Brazil to “leapfrog” from the
heavily occupied coast (e.g. Santos and Rio de Janeiro) to the inland plateau that
includes major cities such as São Paulo and Curitiba (Morse 1974). In contrast, the
Coastal Forests of Bahia ecoregion, in particular in the cacao region of southern
Bahia, has maintained large blocks of forest because of an occupation model based
on cacao agroforestry with an overstory of native forest (cabruca system).
23 Agroecology and Forest Conservation in Three Types of Land. . . 573

While the port city of Salvador was a major colonial city founded in 1564,
indigenous resistance, steep topography and inhospitable climate kept southern
Bahia’s forests relatively intact as late as 1820 (Adonias Filho 1978). At that
point, cacao cultivation took off, and by 1895 cacao was the economic basis for
the entire state of Bahia. Between 1895 and 1930, cacao was consolidated as the
second greatest source of wealth in all of Brazil. Adonias Filho (1978) refers to the
first period as the pioneer phase (desbravadores), while the second period was
dominated by the “cacao barons” (coronéis) who controlled large estates. The latter
period was immortalized by Jorge Amado in novels such as Terras do Sem Fim
(1942, translated as The Violent Land, 1945).
While cacao cultivation produced great wealth for some, it required a large mass
of salaried workers who lived in precarious social and economic conditions.
According to Mascarenhas et al. (1999), the Human Development Index in the
cacao region of Southern Bahia (5.1) was worse than in three other cacao producing
regions of Brazil (5.8–6.5 in Rondonia, Pará and Espiritu Santo), much worse than
Indonesia (7.0), Ecuador (8.0) and Malaysia (9.0), and only slightly better than the
Ivory Coast (4.0) and Ghana (5.0). Brazil has one of the most unequal distributions
of both wealth and land in the world. As of 1979, the region’s Gini coefficient was
0.62, with 9% of rural properties being larger than 200 hectares and occupying 49%
of the region, while 57% of the properties were less than 50 hectares and occupied
16% of the region (Trevizan 1982). However, productivity was inversely propor-
tional to property size – smaller properties produce relatively more cacao (Trevizan
1982).
Traditionally, cacao was cultivated exclusively in the cabruca system. Cacao
(Theobroma cacao) is an understory tree native to the Amazon which requires shade.
Cabruca is the system of planting cacao underneath the canopy of native old-growth
forest supplemented by some cultivated forest species such as jackfruit (Artocarpus
heterophyllus) and cajá (Spondias mondim), which leaves most of the native forest
biomass and ecological services intact. From the 1930s to the 1950s, cacao produc-
tion declined for multiple economic and biological reasons, including the Great
Depression, lack of investment, soil exhaustion and cacao diseases (Adonias Filho
1978). In 1957, a federal agency was created to provide research, credit, rural
electrification and other services to strengthen the cacao economy (CEPLAC –
Comissão Executiva do Plano da Lavoura Cacaueira, Executive Commission for
Cacao Farming Plan). CEPLAC developed a technical package that included
homogenous plantations of cacao with a monoculture overstory of either rubber or
the leguminous tree Erythrina, and included use of chemical fertilizers, herbicides
and insecticides.
According to Almeida et al. (2001), this high-input technical package offered a
160% increase in private profit to the landowner, but a lower “social return”
measured as the broader distribution of benefits throughout society. Notably, the
traditional, low-input system provided greater employment and thus Almeida et al.
(2001) calculated that in this case the social return was three times greater than the
private profit. Mascarenhas et al. (1999) indicate that labor is 67% of the production
cost of cacao (varying from 54 to 84%, depending on which level of technology
is used).
574 K. R. Painter et al.

In the 1990s, cacao production in southern Bahia went into crisis. High global
prices in the 1970s stimulated increasing production in other tropical countries, so
that by 1992 prices were at a low point. At the same time, the Brazilian government
overvalued its currency as a strategy to control inflation (Plano Real), further
lowering the price in Reais. Low prices meant less investment in maintaining
cacao plantations, and when witches broom, a fungal disease (Crinipellis
perniciosa), invaded the region in 1989, it quickly spread out of control (Alger
and Caldas 1994). Large landowners saw their income decline, government revenue
decreased, and masses of salaried rural workers became unemployed. Eventually,
land prices of cacao estates declined, and landowners were eager to sell out. Today
much of the cacao produced in Bahia is produced by small and medium-sized
landholders, because of the labor intensity required to control the disease (Rice
and Greenberg 2000).

23.2.2 Land Reform and the Pressure on Forests

Land reform (also known as agrarian reform) has been on Brazil’s political agenda as
a mechanism of addressing poverty and inequality since the era of industrialization
following World War II. The Catholic church has been active with this issue since the
1960s (Ecclesiastic Base Communities, CEB) and 1970s (Pastoral Land Commis-
sion, CPT, created in 1975). Land reform social movements, such as the Movement of
Landless Rural Workers (MST), the Worker’s Party (PT), and the Rural Worker’s
department within the national union umbrella organization CUT (Central Única dos
Trabalhadores or Unified Workers’ Central), all emerged between 1979 and 1985
(Deere and León 1999). In the 1990s, hundreds of thousands of landless poor engaged
in hundreds of land occupations throughout the country. The Fernando Henrique
Cardoso administration (1995–2002) settled 525,000 families on 18 million hectares.
This was a rather chaotic process with a focus on numerical targets, with limited
support for credit or technical assistance, the key factors for success identified by a
global analysis of agrarian reform experiences throughout the world (Smith 2001).
The land reform process in the cacao region of Southern Bahia had a very violent
trajectory until 1998, when conditions changed dramatically. Key to understanding
this is the fact that, in Brazilian land reform generally, only 5% of projects were
initiated by the government (i.e. by INCRA, the National Agency for Colonization
and Land Reform), with the other 95% initiated by land reform movements such as
MST, who organized a group of settlers and petitioned INCRA’s cooperation (Cullen
et al. 2005). The process of staking a claim and establishing a settlement involved
occupation of a suitable property (acampamento), either directly or on the side of the
road adjacent to it. Such occupations could take months to years to be regularized by
INCRA, who would expropriate the land and compensate the landowner if it could be
shown to be “unproductive” and/or if enough political pressure was applied. During
occupation, settlers could be forcibly removed (despejo), either by landowners’
“hired guns” and/or by the police; in many cases settlements suffered multiple
removals but persisted for months and years until land was expropriated.
23 Agroecology and Forest Conservation in Three Types of Land. . . 575

During this “contested” land reform, settlers had a strong motivation to occupy
areas of forest, protected areas, remote and infertile areas, because in all of these,
landowner resistance would be less. In the case of protected areas, settlers might be
peaceably removed by offering an alternative piece of land. In the case of large cacao
estates, landowners might be open to negotiation if occupation occurred on an
uncultivated forest reserve versus established plantations. Inevitably, converting
newly occupied forest into productive farms required extensive forest clearing.
Around 1998, the cacao crisis (as explained in Sect. 23.2.1) permitted a transition
from this “contested” situation to what we call a “negotiated” form of land reform.
At that point, the price of cacao had crashed and productivity had declined as well
(due to what some settlers sardonically referred to as “Saint Witches Broom”).
Landowners were willing to cede land to expropriation, in which case they might
even negotiate compensation (paid by INCRA) far in excess of the market price.
This was a tipping point, and agrarian reform rapidly switched from painfully violent
occupation of marginal lands to peaceful distribution of the region’s best lands, since
the major cacao estates were on the best soils and had received infrastructure
investments. In some cases, such as that of Fazenda Liberdade in the municipality
of Maraú, former paid laborers making less than minimum wage abruptly became
collective owners of lands that they had lived and worked on for decades.
Converting laborers to owners produced a virtuous cycle of positive incentives for
them to invest in the labor- and managerially-intensive control of Witches Broom,
thus increasing productivity and quality of cacao.

23.3 Study Context: Jupará Approach and Types

of Communities

23.3.1 The Jupará Agroecological Movement

The Jupará Agroecological Movement originated in a “collective” of church and

rural worker union organizations that supported land reform in the cacao region in
the 1980s. In the same region and time period, WWF had been successfully working
to create and consolidate the 11,400-hectare Una Biological Reserve, the only
legally protected habitat of the endangered golden-headed lion tamarin
(Leontopithecus chrysomelas) (Saracura 1997). Subsequent WWF-sponsored
research showed that the Reserve was not large enough to support a minimum viable
population of this species (Dietz et al. 1996), and in 1995 WWF provided an initial
grant to Jupará to explore alternatives to forest clearing in land reform communities
near the Una Reserve.1 Jupara’s goal was to incorporate conservation in a productive
agricultural landscape, while simultaneously seeking to improve social and eco-
nomic well-being.

1
WWF’s initial funding went from 1995 to 1998, with an 18-month funding gap and a second phase
of support from 1999 to 2003 (Buschbacher 2008).
576 K. R. Painter et al.

WWF and Jupará set the following program goals: implementation of agroeco-
logical practices, organic certification, increased family incomes, and maintenance
of 30% of community areas in forests and 40% in agroforestry systems
(Buschbacher 2008). The overall goal was to substitute slash and burn agriculture,
prevalent among smallholders in areas of tropical, moist forest, with a more resilient
and sustainable form of family agriculture. Jupará’s agroecological production
system substituted natural products and ecological techniques such as compost,
green manure, contour erosion barriers, and multicropping for the use of fire and
agrochemicals. A total of 10 agroecological practices were promoted (Table 23.1).
Jupará’s conceptual model, developed jointly with WWF, integrated production,
community organization, commercialization and forest conservation (Fig. 23.1).
Jupará took a holistic, agroecological approach. On-the-ground activities integrated
technical assistance on agroecological production with a methodology based on
empowerment, autonomy and solidarity, including a strong focus on gender equity.
Jupará recognized that economic viability would depend on improving the terms
of trade for inputs and products (commercialization in Fig. 23.1). A cooperative
(COOPASB – Cooperativa dos Pequenos Produtores e Produtoras Agroecologistas
do Sul da Bahia Ltda.) was formed to negotiate better prices at scale. In addition,
Jupará proposed small agroindustries to enhance market value and access for diverse
crops, such as cacao, fruits, guaraná (Paullinia cupana) and manioc. This included
processing (i.e. fermentation and drying of cacao and guaranaa seeds and dehydra-
tion of fruits), packaging, and value-added products (tapioca cakes from manioc,
liqueurs, and jellies and frozen pulp from many kinds of fruit, and eventually
chocolate).
Community organization and leadership was supported by Jupará to strengthen
cooperation within communities on common production practices, management of
agroindustrial enterprises, and articulation with government and other partners.
Finally, the conceptual model considered forest conservation as an inherent
characteristic of the agroecological system being promoted. Organizations that
wished to participate in Jupará agreed to eliminate forest clearing, prioritize organic
agriculture, and commit to quantitative targets for diversified agroforestry produc-
tion systems and intact forest (40% and 30% of total area, respectively). While the
project did not target specific species nor measure biodiversity, reducing forest
clearing was expected to conserve biodiversity and ecological services. In addition,
relative to either of the prevalent alternatives of slash and burn subsistence agricul-
ture or high-chemical-input cacao cultivation, organic and agroforestry-based pro-
duction systems are inherently biodiverse and have conservation value (Schroth
et al. 2004; McNeely and Schroth 2006; Bhagwat et al. 2008; Jose 2009, 2012;
Palacios Bucheli and Bokelmann 2017; Haggar et al. 2019; Santos et al. 2019;
Udawatta et al. 2019; Marsden et al. 2020; Montagnini et al. 2022; Montagnini and
del Fierro 2022). Protected natural forests and agroforestry-based production sys-
tems are the two types of biodiversity island addressed by this chapter.
23 Agroecology and Forest Conservation in Three Types of Land. . . 577

Table 23.1 Description of agroecological production practices promoted by Jupará

1. Compost. Composting of organic materials such as manure, cacao shells and food scraps to
produce organic fertilizer
2. Homemade liquid organic fertilizer. A type of compost tea, using manure and other organic
ingredients, that Jupará extensionists taught farmers to make for themselves
3. Green manures. Nitrogen fixing plants that are planted either between crops or during a fallow
period and allowed to decay on site. Can be used to improve soil quality or control erosion;
farmers also report using green manures to control ants. Species commonly used were Canavalia
(Canavalia enisformis) and pigeon pea (Cajanus cajan)
4. Commercial organic fertilizer. Jupará produced this fertilizer at a central location and sold it
to farmers at cost
5. Mulching. Use of mulches such as leaves or cacao shells, usually around the base of trees, to
add nutrients and control erosion
6. Multicropping. More than one species is planted in each field. Tree crops might be planted
with annual crops or several tree crops planted together
7. Crop rotation. Rotation of the location of crops from year to year. An annual crop should be
followed by a crop from a different family. In agroforestry systems, annual crops are gradually
replaced by perennial and tree crops
8. Contour erosion barriers. Planting along the contour, with the creation of some kind of barrier
to control erosion and gradually form terraces perpendicular to the slope. The barrier is usually
created by piling up the organic residue (weeds and fallow vegetation hand cleared in preparation
for planting) in contour rows. Typically, annual crops are planted in the cleared area between the
rows of residue, and tree crops are planted in the rows. This way of treating the organic residue is
an alternative to burning; it is labor-intensive but promotes long-term soil productivity.
9. Elimination of agrochemicals. Agrochemicals commonly used in Bahia include chemical
fertilizers and pesticides to eliminate leafcutter ants. Jupará hoped to replace these with organic
products
10. Elimination of fire. Farmers in Bahia commonly burn to clear forest or clear fallowed fields
for planting. Jupará hoped to replace burning with intensive, permanent agroforestry systems and
the use of contour barriers
Source: Painter (2006)

23.3.2 Three Types of Settlements in the Cacao Region

of Bahia, Brazil

The Jupará-WWF project worked in 35 communities in 17 municipalities throughout

the cacao region of southern Bahia (Fig. 23.2). The historical origins, and conse-
quently the social, biological and geographical characteristics of these communities
fall into three clearly differentiated categories. This study looks at one representative
community of each type (Table 23.2): a traditional quilombo community (Lagoa
Santa); a “contested” land reform settlement (Fortaleza); and a “negotiated” land
reform settlement (Cascata).
Jupará received WWF support from 1995 to 2003; the dynamics of land reform in
this region changed around 1998, and Jupará adapted accordingly. Initially they
worked with settlements from “contested” land reform. These were mainly forest
578 K. R. Painter et al.

Fig. 23.1 Conceptual model jointly developed by WWF and the Jupará Agroecology Movement.
(Figure elaborated by Carolina Jordão)

remnants that had been passed over by cacao-driven occupation in the nineteenth and
early twentieth centuries because of disadvantageous soils and geographies. Forta-
leza is typical of this pattern and was selected as a priority area by Jupará and WWF
because it is in the same municipality as the Una Biological Reserve.
23 Agroecology and Forest Conservation in Three Types of Land. . . 579

Fig. 23.2 Location of three agricultural communities within the Cacao Region of Southern Bahia,
Brazil. (Map elaborated by Felipe Veluk Gutierrez using MapBiomas)

After 1998, Jupará continued to work with many of these communities but added
several others that were located on cacao estates, such as Cascata, that became
accessible to “negotiated” land reform after the cacao crisis around 1998.
580 K. R. Painter et al.

Table 23.2 Main social, historical and biophysical characteristics of three family agriculture
communities in the cacao region of southern Bahia, Brazil
Community Lagoa Santa Fortaleza Cascata
Origin history Traditional; Agrarian reform (contested) Agrarian reform
quilombo (negotiated)
Origin date 1800s Mid-1970sa 1998
Average 9 ha 22 hab 11 hac
landholding
Accessibility Isolated Isolated On major highway
Soils Moderate; Oxisols; Moderate; Oxisols; Colônia Fertile; Alfisols;
Colônia formation formation Cepec formation
Production Highly diversified Diversified Cacao
system
% of income 45 67 75
from farming
Percent natu- 8% 36% 40%
ral forest
Percent 75% 39% 46%
agroforestry
Forest Maintain and expand Increase forest on private lands; Maintain agrofor-
tendency agroforestry loss of community forest estry and forest
reserve
Source: Based on data from Painter (2006)
a
The area was occupied starting in the 1970s and expropriated by INCRA in 1986 which gave the
50 occupying families collective land rights. However, it was not until 1998 that INCRA began the
process of sub-dividing the area into defined lots for each family
b
This includes 17 hectare per family lot + 5 hectare per family in the community forest reserve
c
All the land is held collectively, but each family is assigned 4–5 hectare of cacao plantation to work

The third type of community was not explicitly involved in official land reform,
but is a historical remnant of a kind of spontaneous land reform that was self-
generated by slaves who escaped from plantations, Jesuit missions, and the slave
trade in the nineteenth century. Lagoa Santa was one community of this type where
Jupará worked intensively because of the protagonism of a dynamic local leader,
André Jesus de Conceição (known as Catixa). These “traditional communities”
make an intriguing comparison because they can be conceived as an extrapolation
into the future of the contested land reform type of community (e.g. Fortaleza)
because they were also settled in remote remnants of forest with minimal capital
or support, and have now been occupied for generations.

[Link] Lagoa Santa

Lagoa Santa is a traditional agricultural community located in the municipality of

Ituberá. Lagoa Santa is a quilombo community, meaning a community that was
originally founded by a group of escaped slaves in the nineteenth century. Resident
families have farmed this land for several generations, resulting in numerous land
23 Agroecology and Forest Conservation in Three Types of Land. . . 581

divisions among farmers’ descendants and, consequently, small landholdings. Land-

holding size ranges from 2 to 25 hectares, with an average size of 9 hectares. Many
of the larger landholdings are shared by extended families. Little native forest
remains in this area. Soil quality is moderate. Terrain is very hilly. While fairly
close to the municipal headquarter city of Ituberá, access to Lagoa Santa is difficult.2
Farmers in Lagoa Santa maintain highly diverse agroforestry systems, producing
mainly rubber, cloves, piassava (Attalea funifera) palm fiber for roofing material,
and small amounts of cupuaçu (Theobroma grandiflorum), açaí palm (Euterpe
oleracea), guaraná, cacao, black pepper, and manioc flour. On average, 67% of
family income comes from agriculture and the remaining 33% comes from off-farm
sources, including employment in Ituberá and on nearby plantations.

[Link] Fortaleza

Fortaleza is a land reform settlement in the municipality of Una and a neighbor of the
Una Biological Reserve. Fifty families were settled on 1102 hectares, with an
average lot size of 17 hectares. Approximately 250 hectares were designated com-
munity areas including a small forest reserve and a community cacao plantation.
Families began settling the area in the mid-1970s when the area was completely
forested. Most settlers grew up on their parents’ farms, usually also in Southern
Bahia, or worked on nearby rubber plantations before settling in Fortaleza. INCRA
(Instituto Nacional de Colonização e Reforma Agraria, the government agency
responsible for land reform) officially divided the land and gave titles to these
landholders only in 1997. At that time, and as part of the titling process, some
families were moved from their original plot of land to a different plot: in some
cases, a new, completely forested plot. In other places, agroforestry plantations that
had been cultivated by one family were allocated to a different family; any compen-
sation had to be arranged informally between these families. At the time of this
research, many of the original farmers in Fortaleza were getting older and had
children who were married and beginning their own families. In some cases, the
land has been unofficially divided among several sons. In one case, 23 people were
surviving on the production of one 17 hectare parcel. In other cases, the children of
these farmers worked off-farm in Una or on nearby plantations. Farmers in Fortaleza
produced mainly rubber and cacao as cash crops, as well as some manioc, corn, fruit
and vegetable crops for consumption. Few families owned livestock beyond one
mule for hauling cacao and a few chickens. On average, 47% of family income came
from agricultural production and 53% came from off-farm sources, usually employ-
ment in the nearby town of Una, or retirement pensions. Most of the agricultural
income came from rubber.

2
Because of their origin as refuges of escaped slaves, quilombos occur in areas of difficult access.
While Lagoa Santa is only about 5 miles from Ituberá, due to very hilly terrain it currently takes
about 30 min by car and a good part of a day by foot.
582 K. R. Painter et al.

[Link] Cascata

Cascata, in the municipality of Aurelino Leal, was an abandoned cacao plantation

that was sold for land reform, settling 40 families in 1998. The land titles were
granted not to individuals but to the community association, although each family
was assigned an area of 4–5 hectares of the plantation to maintain and harvest. In
addition, about 180 hectares of Atlantic Forest were designated as a community
forest reserve. While at the time of this research Lagoa Santa and Fortaleza lacked
electricity, potable water, basic sanitation, and regular public transportation services,
Cascata is located along a major highway, allowing for easy access to public
transportation, local markets, secondary education and health services. Because
Cascata was established on a former plantation, much infrastructure was already in
place, including a dozen workers’ houses occupied by resident families, a large
building for meetings and events, a church and school, production areas for
processing cacao, and a large cacao dryer. Families enjoy well-constructed brick
homes, electricity, running water, modern sanitary facilities, and a public telephone.
Cascata has fertile soils and produces cacao almost exclusively; the cacao plantation
was well established when families arrived, but heavily disease-infested, and cultural
practices focused on recuperation of these plantations.

23.4 Methods Used to Assess Agroecological Adoption

and Forest Conservation

In order to evaluate the long-term efficacy of the Jupará Agroecological Movement’s

model of land reform, and to assess the extent to which it is able to create and
maintain two kinds of biodiversity islands (diversified agroforestry systems and
remnants of natural forest), in 2005 a University of Florida (UF) Master’s student
and participant of UF’s Tropical Conservation and Development program conducted
research in collaboration with Jupará to address the following questions:3 Are
families that participated in the Jupará project meeting the quantitative goals of
40% of their area in diversified agroforestry production systems and 30% in intact
forest? How do family dynamics and size of landholding influence project imple-
mentation? Are project results consistent in different types of communities? (Painter
2006).

3
Painter collected and analyzed all data presented in this chapter. Buschbacher was her thesis
advisor. He worked at WWF until 2000, was the main WWF collaborator with Jupará from 1994 to
2000, and wrote an unpublished case study on the Jupará project which is the basis for much of the
regional description in this chapter (Buschbacher 2008). Luiz Carlos Souto Silva and Emerentina
Costa e Silva are the leaders of Jupará, and facilitated the design and implementation of Painter’s
field work. Souto provided information on Jupará agroindustry and commercialization activities
post-2008.
23 Agroecology and Forest Conservation in Three Types of Land. . . 583

Painter (2006) interviewed and made site visits to 60 families in the 3 represen-
tative communities described above (10 families in Cascata, 20 in Lagoa Santa, and
30 in Fortaleza). In Cascata, where management practices are collective and uniform
among families, data were analyzed for the settlement as a whole. For Lagoa Santa
and Fortaleza, an index of participation was used in order to differentiate and
compare results between families who were active participants in the Jupará agro-
ecological movement and those who were not (Painter 2006). In Fortaleza, land use
change over time was assessed using Landsat imagery (1986 and 2001) analyzed in
ArcGIS; participatory mapping with GPS reference points and a digital map pro-
vided by INCRA were used to align remote sensing imagery to properties (Painter
2006).

23.5 Adoption of Agroecological Production Practices

in Different Settlement Types

The Jupará project promoted a set of 10 agroecological production practices

(Table 23.1). The results of the interviews conducted by Painter in 2005 show that
these 10 practices could be partitioned into 3 groups: three that were widely
implemented by both participants and non-participants in the Jupará program;
three that were rarely implemented by any of the farmers; and three practices that
were differentially implemented by Jupará project participants and not by
non-participants (Fig. 23.3).
In 2005, 10 years after initiation of the Jupará project, multi-cropping, crop
rotation and mulching were implemented by 70–100% of all farmers, without
clear differentiation between Jupará participants and non-participants. These can
be considered well established practices in this region (Painter 2006).
On the other hand, relatively few farmers used green manure, homemade liquid
organic fertilizer, or organic compost, whether they were Jupará participants or not.
Green manures were used by less than 20% of both participants and non-participants
in both communities. This practice was not as vigorously promoted as others (many
farmers had never even heard of the practice), and seeds were not readily available.
The few farmers who did report using green manures reported using them as a pest
control method against leaf-cutter ants, rather than as a source of fertilizer. Both the
homemade liquid organic fertilizer and organic compost require animal manure as a
key ingredient; very few of the farmers in these communities have livestock, and
“importing” the manure proved infeasible.
In this context, Jupará changed their strategy to produce their own “organo-
mineral fertilizer” in the industrial zone of Ilhéus, a central city in the cacao region
(See Fig. 23.2) where appropriate inputs (ground rock rich in phosphorus and
micronutrients, ash, and composted organic matter) could be obtained (Buschbacher
2008). This fertilizer can build up soil organic matter content and contribute to long-
584 K. R. Painter et al.

Fig. 23.3 Adoption of 10 targeted agroecological practices in 2005 by participants and

non-participants of Jupará Agroecology movement in two communities of southern Bahia, Brazil

term soil fertility, but it was slightly more expensive than commercial chemical
fertilizers and also less concentrated (greater weight and volume), which made it
more difficult to transport and apply. These disadvantages were cited by several
farmers who chose not to use it, but nevertheless the use of Jupará’s organic fertilizer
was one of the agroecological practices that were differentially implemented by
Jupará participants versus non-participants. There was considerable overlap between
the use of Jupará’s organic fertilizer and the practice of total elimination of agro-
chemicals including pesticides, herbicides and chemical fertilizer.
The third practice that was much more widely adopted by Jupará participants than
non-participants was the construction of contour erosion barriers. Contour erosion
barriers are labor intensive and require some technical expertise to create. The fact
that Jupará participants widely adopted this practice indicates that Jupará was
successful in providing technical expertise and motivating farmers to invest in a
labor-intensive but ecologically beneficial process that enhances long-term
23 Agroecology and Forest Conservation in Three Types of Land. . . 585

productivity. In Fortaleza, around 55% of Jupará participants used contour erosion

barriers, 65% used Jupará’s organic fertilizer, and nearly 90% reported no use of
agrochemicals; in each case less than 20% of non-participants adopted these prac-
tices (Fig. 23.3). A similar difference of 50% of participants versus less than 20% of
non-participants created contour erosion barriers in Lagoa Santa, whereas only about
25% of Lagoa Santa participants used Jupará’s organic fertilizer and totally elimi-
nated agrochemicals (versus less than 10% of non-participants).
The use of fire to prepare fields, generally considered ecologically undesirable,
was persistent among Jupará participants and non-participants alike. Because inter-
views considered the use of fire as a binary yes/no variable, the data do not
distinguish how fire is used or how extensively. Certain agroecological practices
are incompatible with the use of fire, such as the contour erosion strips which form a
vegetative barrier and is a labor-intensive investment that would be destroyed by fire.
In addition, agroforestry systems are long-term investments and once established not
likely to be intentionally burned. Thus, it is most likely that burning was used only
(or mainly) for the initial establishment of agricultural fields, whether from native
forest or successional regrowth forest (i.e. previously abandoned farmland). The key
to eliminating this practice is continuous cropping once fields are established
(i.e. agroforestry systems). Thus, the data on maintenance of forest cover and extent
of agroforestry systems (Sect. 23.6) are probably much better indicators of ecolog-
ical and biodiversity values than whether or not fire is completely eliminated from
the repertoire of agricultural practices.
Both biophysical factors and production systems are quite different in Cascata,
the former cacao plantation (Table 23.2). The cacao agroforestry system (cabruca,
with native forest overstory) was already established but heavily attacked by
Witches Broom disease; thus, efforts were focused on pruning diseased branches,
managing shade and humidity with overstory thinning and repair of drainage ditches,
weeding, organic fertilization, replanting of overstory gaps, and grafting disease-
resistant clones of cacao (Buschbacher 2008). All Cascata farmers were Jupará
participants and the community received a collective organic certification in 2000.
All used Jupará’s organic fertilizer and none used agrochemicals. Crop rotation and
contour erosion barriers were less utilized in the cabruca system, but 50% of farmers
did use the homemade liquid organic fertilizer for foliar application (Painter 2006).
Cascata does have some livestock used to transport cacao from the fields to the
processing center for fermentation and roasting, and in 2004 the community built a
biogas facility that converted manure to gas for the dryer and other machinery
processing cacao and fruits, with the residue used as fertilizer. Only one farmer
reported using fire (probably for a small area of subsistence crops, not for the cacao
plantation).
586 K. R. Painter et al.

23.6 Forest Conservation in Different Settlement Types

Although Jupará participants implemented a different subset of agroecological

practices when compared with non-participants, differences in forest cover mainte-
nance between these groups were small and not statistically significant (Painter
2006). However, there were major differences between communities, and these are
illustrative of how history, land tenure, biophysical characteristics and production
systems influence forest dynamics and conservation.
In both Lagoa Santa and Fortaleza, much of the forest cover was converted to
agricultural use long before the Jupará project began. The traditional quilombo
community of Lagoa Santa has had long occupation, with landholdings subdivided
across generations, such that average landholdings are a relatively small 9 hectare
per family. Only 8% of this area remains in native forest, far lower than in either of
the other communities; yet far more of this community is in agroforestry systems,
which cover 75% of the entire community. These agroforestry systems are highly
diverse, with rubber, cloves, and piassava palm fiber (Attalea funifera) as major
crops, often intercropped with cacao, banana, plantain, açai, cupuaçu, black pepper,
guaraná, passionfruit, pineapple, manioc, timber species and others. Anecdotally,
Jupará participants may have more diverse agroforestry systems than
non-participants; certainly, the more prevalent use of contour erosion strips is
indicative of more intensive management.
Fortaleza was a typical land reform settlement of the “contested” period when
cacao estates were heavily defended and land occupations occurred opportunistically
in remote, unsettled areas of forest. When Fortaleza was occupied in the mid-1970s,
it was fully forested. By 2005, it had 36% of forest cover, plus 39% in agroforestry
(Painter 2006). Agroforestry systems in Fortaleza are diverse but less diverse than in
Lagoa Santa. Rubber and cacao are the main cash products; manioc and other short-
lived crops are produced during the early stages of agroforestry system establish-
ment, and piassava, açai, cupuaçu and other species are also present.
From the time of occupation until 1998 when the federal land reform agency
INCRA actually demarcated individual lots, the community had set aside 20% of its
area as a community forest reserve. Rather than formalize the land division that had
been established during the first 20 years of occupation, INCRA decided unilaterally
to allocate the territory, such that eight of the 50 families were given lots in fully
forested areas within the community forest reserve. The families that were relocated
into the forest reserve had to clear part of that forest to produce. In spite of this, forest
cover from remote sensing images showed that total community forest cover
remained almost constant between 1986 and 2001 (the latest cloud-free image
available in 2005): natural forest cover declined by 2% while agroforestry and
successional forest (which could not be distinguished in the remote sensing imagery)
increased by 3% (Painter 2006, Fig. 23.4). In fact, while forest cover decreased on
the “new lots” of community members who were relocated to the forest reserve,
forest cover increased on the older lots by 20% among Jupará participants and nearly
10% among non-participants. Much of this increase in forest cover occurred along
23 Agroecology and Forest Conservation in Three Types of Land. . . 587

Fig. 23.4 Percent change in forest cover, agroforestry, and cleared areas among participants and
non-participants of the Jupará Agroecology movement, in new lots established within the forest
reserve, and in the remaining forest reserve of Fortaleza Land Reform Settlement, Bahia, Brazil,
1986–2001

two riparian corridors in the center of the community, consistent with Jupará
recommendations. Furthermore, the community forest reserve was never fully intact;
about 5% had been cleared by 1986, which increased to 6% in 2001 (as opposed to
28% cleared in the areas that were transformed into family lots).
The case of Cascata, which was an established cacao plantation that was
converted into a land reform settlement, is quite different. Upon establishment as a
land reform community, the estate already had 46% of its area in agroforestry
systems, specifically cabruca cacao. In addition, 40% of the settlement area was
intact forest. This is rather typical for cacao estates, in which landowners claimed
and controlled as much land as they could and gradually implemented cacao
plantations in the most accessible and least steep portions. Although no remote
sensing analysis was performed, there was no major observable change to the
coverage of either the native forest or agroforest (Painter 2006).
588 K. R. Painter et al.

23.7 The Effectiveness of Jupara’s Agroecological Land

Reform Model and Its Applicability for Regional
Conservation and Development

This final reflection will look at Jupará’s impact in terms of agroecological produc-
tion practices, the social and economic viability of this approach to land reform, and
the larger implications for regional forest conservation. Does the Jupará Agroeco-
logical Movement present a viable “proof-of-concept” for agroecological produc-
tion, community organization and commercialization to achieve forest conservation?
How do the historical, ecological and geographic differences between different types
of communities enable or limit the adoption of agroecological practices and the
protection of forest remnants? And what are the implications of these findings for
mobilizing public policy to produce positive outcomes for ecosystems and
communities?

23.7.1 Viability of Jupará’s Agroecological Model

In terms of agroecological production practices, Jupará was able to introduce and

implement diversified cropping systems with organically enriched soils that will be
productive for decades in all three communities. The adoption of organic practices
(Jupará organic fertilizer and elimination of agrochemicals) and long-term invest-
ment in diversified agroforestry (contour erosion barriers) by Jupará participants
demonstrates a significant shift towards more sustainable agriculture.
Furthermore, Jupará was able to capitalize on the agroecological production and
community organization to implement a series of commercialization activities
between 2002 and 2012. Jupará’s Cooperative COOPASB exported one container
of organic cacao directly to Europe with WWF’s support in 2002 and continued
selling certified organic cacao to regional cacao trading companies thereafter. In
addition, food processing centers (“Casas de Agroaproveitamento”) in six different
communities produced dehydrated fruit and fruit pulp for juices, including jackfruit
(Artocarpus heterophyllus), cajá (Spondias mombin), banana, plantain, pineapple
and jenipapo (Genipa americana). Sales agreements for these products were nego-
tiated with government programs for food acquisition from family agriculture.4
Jupará, COOPASB and participating communities managed resources from federal
agencies INCRA, CONAB5 and the Secretariat for Territorial Development, and

4
PAA – Programa de Aquisição de Alimentos (Food Acquisition Program), destined for poor
populations and national food stocks and PNAE – Programa Nacional de Alimentação Escolar
(National Program of School Nutrition).
5
CONAB – Companhia Nacional de Abastecimento, implements the PAA and executed a purchase
agreement with COOPASB which included advanced payment that was used for capital investment
in the food processing centers.
23 Agroecology and Forest Conservation in Three Types of Land. . . 589

attained production levels of 80–100 tons per year of dehydrated fruits and 4–5 tons
per month of fruit pulp, providing revenue of 3.5 million Brazilian Reais per year
and partial employment to 400 community members, with priority given to women
and youth.6
Finally, environmental conservation through the land sharing type of biodiversity
island is evident in the agroecological practices. These increase productivity and
long-term sustainability of the production system (Sandhu et al. 2010; Powlson et al.
2011; Palm et al. 2014; Reganold and Wachter 2016), while enhancing biodiversity
and ecological services both on and off site. The practices decrease manufacturing,
transport and sale of toxic chemicals, as well as toxic exposure of both producers and
consumers. They also increase the quality of water runoff, by reducing erosion,
leaching of nutrients and toxic chemical load (Sebesvari et al. 2012; Cambardella
et al. 2015). Organic matter and soil organisms are noticeably more abundant in
contour erosion strips where weeds and other organic material are piled to decay,
when compared to adjacent land between the strips, and especially when compared
to sites prepared by burning (authors’ personal observation). The use of organic pest
control maintains the biodiversity of natural pest predators and such systems are
inherently bird-friendly (Bengtsson et al. 2005; Crowder et al. 2010).
Taken together, these positive results for agroecological production, community
organization, commercialization and land sharing clearly demonstrate the viability
of Jupará’s model of land use in agricultural communities of the cacao region of
southern Bahia. Jupará’s accomplishments, especially the integrated model includ-
ing the biodigestor implemented in the community of Cascata, received national and
global recognition: the Social Technology award from the Bank of Brazil Founda-
tion in 2005, “Best Practice for Local Management” in the category of “Environ-
mental management, combating poverty, changing patterns of production and
consumption” from the Caixa Economica Federal bank, and “Best Practice in
Conservation of the Natural Environment” from UNESCO in 2006.7

23.7.2 Application of Jupará’s Model for Forest

Conservation at Landscape Scale

To understand the broader implications of Jupará’s model of agroecological land

reform for regional forest conservation (i.e. the land sparing type of biodiversity
island), the comparison among the three different communities is decisive. When
given the opportunity to occupy a productive part of the landscape (i.e. the cacao
estate at Cascata), there was no necessity to clear forest and the large forest remnant
covering 40% of this settlement was maintained. The 46% of the settlement that is in

6
Each center employed about 20 workers, which was rotated among 80 community members,
i.e. 1 week per month each.
7
[Link]
590 K. R. Painter et al.

cabruca cacao absorbed the community’s labor and provided the economic basis for
a prosperous quality of life. This result could be undermined in the future by
population growth or increasing economic aspirations (e.g. Jevons paradox – see
Alcott 2005; Polimeni et al. 2015; York and McGee 2016). Nevertheless, Jupará’s
actions to increase the productivity of the forest overstory by processing the wide-
spread jackfruit and cajá, diversifying this overstory with other economically valu-
able species, and converting animal manure into biogas, as well as proposals to
produce artisanal chocolate and develop agritourism with access to native forest and
a waterfall, would all be beneficial to the sustainability of this system and its
biodiversity and ecological services.
The case of Fortaleza is in some ways opposite to Cascata. Landless poor
occupied an area of forest in the 1970s and had to clear the forest to produce. Poor
soils made it more difficult to produce the more nutrient-demanding cacao as
intensively as in Cascata, and poor logistics and limited markets made it difficult
to commercialize the full array of potential agroforestry crops.8 The slash and burn
model of tropical forest agriculture would lead to continuation of this initial forest
loss, but by basing production on diversified agroforestry (primary income sources
are rubber and cacao) these farmers achieved an equilibrium with a mix of agrofor-
estry, remnant forest, and some cleared areas (Fig. 23.4). In fact, our data show that
forest cover on some parts of the settlement is increasing, more so among Jupará
participants than non-participants (Fig. 23.4). We observed that community mem-
bers gave priority to reforesting connected riparian areas (Painter 2006). Unfortu-
nately, the public policy implemented by INCRA relocated some families onto the
community forest reserve, and the cycle of initial forest clearing began again.
Lagoa Santa may represent the future tendency of Fortaleza extrapolated through
several generations. The biophysical and geographical situation of these two com-
munities is similar (Table 23.2), but Lagoa Santa has had much longer occupation.
Subdivision of landholdings across generations has led to more intensively managed
agroforestry systems that are more diverse in Lagoa Santa than in Fortaleza, but very
little remaining forest (8% in Lagoa Santa compared to 36% in Fortaleza).
Notably, these differences in socioeconomic and conservation outcomes across
communities are not a function of Jupará efforts, but rather due to the broader
historical, geographic and biophysical characteristics of each community. Cascata
had the most successful economic and conservation outcomes because of favorable
soils, location, and pre-existing land use. In addition, Cascata maintained a collective
organization, with all community members participating in Jupará, versus about half
of community members from other communities participating in Jupará. Unlike
Lagoa Santa and Fortaleza, where farmers work individually on their own widely-
separated landholdings, Cascata Land Reform Settlement was created on an existing
plantation, with an established, continuous plantation of cacao and a centralized

8
Cascata is on a main road with access to public transport, whereas the other two sites are much
more difficult to access. This matters both for access to markets and access to other jobs for off-farm
income, and potentially less pressure for the land to support everyone in the family.
23 Agroecology and Forest Conservation in Three Types of Land. . . 591

Fig. 23.5 Conceptual model modified to consider the type of land occupation and macro-systemic
factors. (Figure elaborated by Carolina Jordão)

hamlet of housing and processing facilities. This enabled implementation of the

biogas digester, centralized processing of cacao to ensure high quality, and collective
commercialization.
These findings lead us to propose a revised version of Jupará’s conceptual model,
expanding it to look beyond the specific actions within the community that Jupará
can influence directly (Fig. 23.1), to incorporate the larger-scale system within which
Jupará’s efforts are embedded (Fig. 23.5). Agroecological practices, from production
systems to community organization and commercialization, vary dramatically
between settlements in productive areas (i.e. former cacao estates such as Cascata)
and settlements within forested, inaccessible and less fertile areas. Among the latter,
there are significant differences in landholding size, demographics, and production
systems between long-established “traditional communities” such as quilombos and
newly established “contested land reform communities” (i.e. Lagoa Santa versus
Fortaleza). All of these differences reflect the cacao region’s history, biophysical
variability, and geographic implications for markets and logistics. Finally, the
establishment, implementation and viability of all of these settlement types is
strongly affected by public policies.
The broader systemic perspective on regional development and land reform
(Fig. 23.5) calls attention to the complexity of the challenges of conservation and
development. One critical yet under-studied aspect of sustainability and livelihoods
of these family farmers is the complex set of linkages between rural and urban areas.
For example, 25–55% of family income in these communities comes from sources
other than agriculture, whether off-site labor or retirement or other payments
592 K. R. Painter et al.

(Table 23.2). The successful production and commercialization system described

below (André Catixa) is based on access to the weekly municipal market. These are
some of the complexities that come into focus when we broaden our perspective to a
larger systemic scale.

23.7.3 Public Policy Influence on Implementation

of Agroecological Practices and Forest Conservation

The potential for maintenance of native forest within agricultural settlements in

southern Bahia varies from the forest rich cacao estates to the traditional communi-
ties which are rich in agroforestry but have little native forest. Within this panorama,
public policy can be critical. The pre-1998 approach to land reform – reactive
legalization following occupation of economically marginal forest rather than pro-
active distribution of existing productive lands – catalyzed forest clearing. Various
INCRA policies exacerbated this outcome, such as considering private lands with
more than 50% forest as “unproductive” and thus subject to occupation and expro-
priation, and demarcating lands that exceed the legally required minimum of 20%
forest as areas to be farmed.9 The outrageous case of Fortaleza where 8 of the
50 families were moved internally within the settlement by INCRA onto lots that
were wholly natural forest illustrates the negative impact of these policies. While in
one part of the community families were increasing the area of forest along riparian
corridors, in another part INCRA resettled people with no option but to clear forest.
But with the transition of the cacao economy that occurred in 1998, land reform
began to occur in the most productive part of southern Bahia’s landscape. As a result,
it is now possible to reconcile land reform with conservation in both kinds of
biodiversity islands, agroforestry production systems and forest remnants. Cascata
exemplifies this, and at one point Jupará incorporated several such settlements
throughout the region. These communities contain important forest remnants, with
the potential to amplify their impact by taking a regional perspective. Just as farmers
within Fortaleza protected a corridor of forest in a riparian area, the geography and
topography of the entire cacao region is amenable to creating regional corridors.
Cacao estates open onto roads and back up into steep slopes; cabruca agroforestry is
concentrated near the entrance and considerable forest remnants are left in the
interior. These forest remnants could be linked to those of nearby landholdings
with similar layouts and designated as regional corridors/biodiversity islands.
The current situation of André Catixa, the community leader from Lagoa Santa, is
an exemplary case of how government support can be integrated with the Jupará
agroecological approach. Through the agency of community members, the state

9
Even though this is contrary to Atlantic Forest Decree 750/93. “Ficam proibidos o corte, a
exploração e a supressão de vegetação primária ou nos estágios avançado e médio de regeneração
da Mata Atlântica”
23 Agroecology and Forest Conservation in Three Types of Land. . . 593

Fig. 23.6 Catixa (André Jesus de Conceição) in his family’s irrigated production system for edible
greens, with a small portion of his agroforestry plot immediately behind, Lagoa Santa community.
Far background is neighboring land that does not participate in Jupará. (Photo: Robert
Buschbacher)

government has formally recognized Lagoa Santa as a quilombo community, with

accompanying land rights; some non-quilombo landowners are still present but over
time these should be expropriated and removed. Public policies have provided
support for housing, and Catixa’s extended family lives in a community of six
houses; initial construction of these was sub-standard but community members
have repaired and expanded these and currently live in dignified conditions. Credit
to quilombo members enabled the family to install an irrigation system, and they
currently have a small, terraced plot that produces a variety of edible greens
(Fig. 23.6) that the family harvests weekly, cleans, packages, and sells at the Ituberá
Saturday market. The family has access to education, internet, a car, and motorcy-
cles. The remainder of their land is in diversified agroforestry systems, with multiple
plots divided among different family members. Planting is responsive to market
opportunities, with recent increase in açai planting and diversification into high-
value exotic fruits such as mangosteen and rambutan.
André Catixa was able to leverage government support to achieve exemplary
results, but unfortunately the Jupará extension model has not yet been widely
adopted by the government. Rather than being supportive or even proactive, gov-
ernment extension agents who prepare family agriculture loan projects, and the
594 K. R. Painter et al.

banks who process them, have to be coaxed and guided to enable agroforestry-based
projects. The case of land demarcation in Fortaleza shows INCRA working directly
contrary to forest conservation. The Jupará Agroecological Movement has demon-
strated social, economic and production feasibility, but until this approach is
mainstreamed in existing governmental programs, its implementation will depend
on necessarily-limited external funding and favorable macro conditions.

23.8 Conclusions

The goal of the Jupará Agroecological Movement was to demonstrate and imple-
ment a socially, environmentally and economically sustainable model of land reform
adapted to the historical, biophysical and socioeconomic conditions in the cacao
region of southern Bahia. WWF supported this initiative in the belief that it could
reduce the rate of forest clearing and conserve biologically important habitat. Their
partnership was guided by a conceptual model (Fig. 23.1) which integrated agro-
ecological production practices, community organization, and commercialization of
multiple, value-added products, to achieve both land sharing in the organic agrofor-
estry plots and land sparing in protected remnants of native forest.
Jupará was able to introduce and implement diversified cropping systems with
organically enriched soils that will be productive for decades in all three communi-
ties. The adoption of organic practices (Jupará organic fertilizer and elimination of
agrochemicals) and long-term investment in diversified agroforestry (contour ero-
sion barriers) by Jupará participants demonstrates a significant transition. In addi-
tion, Jupará was able to demonstrate economic viability in the context of government
investments in land reform communities and policies for food acquisition from
family agriculture. Finally, the organic, agroforestry-based production systems
were clearly compatible with the land-sharing type of biodiversity island. These
positive results for agroecological production, community organization, commer-
cialization and land sharing clearly demonstrate the viability of Jupará’s model of
land use in agricultural communities of the cacao region of southern Bahia.
In terms of the regional applicability of this model and its implications for the
land sparing type of biodiversity island, our analysis shows that larger-scale contex-
tual factors – ranging from the type of land occupation to broader systemic
variables – determine the type of productive system, how land can be utilized, and
thus the feasibility of maintaining forest cover. We thus propose a revised conceptual
model (Fig. 23.5) that recognizes the different types of community land-use occu-
pation and considers broader systemic social, political, economic and biophysical
factors. These driving variables were not taken into account when WWF and Jupará
participants agreed on quantitative targets for 30% forest10 and 40% agroforestry

10
The 30% target was conceived as an incremental improvement on the Brazilian Forest Code’s
requirement, albeit with limited to no enforcement, that all landholdings in the Atlantic Forest
maintain 20% forest cover.
23 Agroecology and Forest Conservation in Three Types of Land. . . 595

coverage. There was never an enforcement mechanism related to these targets, nor
even a measured baseline or monitoring of forest coverage prior to Painter (2006).
This was because Jupará conceived standing forest as a valued asset, not as an
opportunity cost (Buschbacher 2008). Results show that the agroforestry targets
were achievable on their economic merits, but that protection of forest remnants
depends on enabling conditions, as illustrated by the three types of community
analyzed in this chapter. One implication is that Jupará could develop more viable
targets for forest protection by taking existing conditions into account at the level of
communities or individual landholdings (i.e. the fact that Lagoa Santa has only 8%
forest cover or that some farmers in Fortaleza were moved to landholdings with
100% forest). More broadly, the key finding of this study with regard to the land
sparing type of biodiversity island within land reform communities of southern
Bahia is that it does occur in some areas, but that supportive public policies are
essential to achieve large-scale maintenance of natural forest within the broader
landscape.
Public policies can support diversified production systems, maintenance of forest
remnants, and socioeconomic well-being of family farmers in two ways: by provid-
ing more support (i.e. financing, technical assistance, and public purchasing) and by
ensuring that existing forms of support align with agroecological land use. The
example of André Catixa shows the positive outcomes from well-aligned policies,
capacity and motivation for agroecological land use. The case of land demarcation in
Fortaleza shows INCRA working directly contrary to forest conservation. The
Jupará Agroecological Movement has demonstrated social, economic and produc-
tion feasibility, but until this approach is mainstreamed in existing governmental
programs, its implementation will depend on necessarily-limited external funding
and favorable macro conditions.

References

Alcott B (2005) Jevons’ paradox. Ecol Econ 54(1):9–21

Alger K, Caldas M (1994) The declining cocoa economy and the Atlantic Forest of southern Bahia,
Brazil: conservation attitudes of cocoa planters. Environmentalist 14(2):107–119
Almeida FA, Mascarenhas GCC, Midlej RR (2001) Estudo da cadeia agroindustrial do cacau. In:
RCMT V, Teixeira Filho A (eds) Cadeias Produtivas no Brasil: Análise de Competitividade, vol
1, 1st edn. Embrapa, Brasília, pp 110–135
Amado J (1942) Terras do Sem Fim. Livraria Martins, São Paulo
Arroyo-Rodríguez V, Arasa-Gisbert R, Arce-Peña N, Cervantes-López MJ, Cudney-Valenzuela SJ,
Galán-Acedo C, Hernández-Ruedas MA, San-José M, Fahrig L (2022) The importance of small
rainforest patches for biodiversity conservation: a multi-taxonomic assessment in the Lacandona
region, Mexico. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments, Topics in Biodiversity and Conservation. Springer, Cham,
pp 41–60
Barnes MD, Glew L, Wyborn C, Craigie ID (2018) Prevent perverse outcomes from global
protected area policy. Nat Ecol Evol 2(5):759–762
Bengtsson J, Ahnström J, Weibull AC (2005) The effects of organic agriculture on biodiversity and
abundance: a meta-analysis. J Appl Ecol 42(2):261–269
596 K. R. Painter et al.

Bhagwat SA, Willis KJ, Birks HJB, Whittaker RJ (2008) Agroforestry: a refuge for tropical
biodiversity? Trends Ecol Evol 23(5):261–267
Boivin N, Zeder MA, Fuller DQ, Crowther A, Larson G, Erlandson JM, Denham T, Petraglia MD
(2016) Ecological consequences of human niche construction: examining long-term anthropo-
genic shaping of global species distributions. Proc Natl Acad Sci 113(23):6388–6396
Bray DB, Duran E, Ramos VH, Mas JF, Velazquez A, McNab RB, Barry D, Radachowsky J (2008)
Tropical deforestation, community forests, and protected areas in the Maya Forest. Ecol Soc
13(2):56
Brockington D, Wilkie D (2015) Protected areas and poverty. Philos Trans R Soc B Biol Sci
370(1681):20140271
Brockington D, Duffy R, Igoe J (2008) Nature unbound: conservation, capitalism and the future of
protected areas. Earthscan, London/Sterling
Buschbacher R (2008) Reforma Agrária Ecológica na Região Cacaueira do Sul da Bahia: O Projeto
Jupará de Conservação e Desenvolvimento. Unpublished manuscript, 92pp
Cambardella CA, Delate K, Jaynes DB (2015) Water quality in organic systems. Sust Agric Res
4(3):60–69
Chapin M (2004) A challenge to conservationists. World Watch 17(6):17–31
Charnley S, Poe MR (2007) Community forestry in theory and practice: where are we now? Annu
Rev Anthropol 36:301–336
Clement CR, Denevan WM, Heckenberger MJ, Junqueira AB, Neve EG, Teixeira WG, Woods WI
(2015) The domestication of Amazonia before European conquest. Proc R Soc B Biol Sci
282(1812):20150813
Cronon W (1996) The trouble with wilderness: or, getting back to the wrong nature. Environ Hist
1(1):7–28
Crowder DW, Northfield TD, Strand MR, Snyder WE (2010) Organic agriculture promotes
evenness and natural pest control. Nature 466(7302):109–112
Cullen L Jr, Alger K, Rambaldi DM (2005) Land reform and biodiversity conservation in Brazil in
the 1990s: conflict and the articulation of mutual interests. Conserv Biol 19(3):747–755
de Souza JG, Schaan DP, Robinson M, Barbosa AD, Aragão LE, Marimon BH Jr, Marimon BS, da
Silva IB, Khan SS, Nakahara FR, Iriarte J (2018) Pre-Columbian earth-builders settled along the
entire southern rim of the Amazon. Nat Commun 9(1):1–10
Deere CD, León M (1999) Towards a gendered analysis of the Brazilian agrarian reform. Occa-
sional Paper #16, Latin American Studies Consortium of New England. University of Connect-
icut, Storrs
Denevan WM (1992) The pristine myth: the landscape of the Americas in 1492. Ann Assoc Am
Geogr 82(3):369–385
Dietz JM, de Sousa SN, Billerbeck R (1996) Population dynamics of golden-headed lion tamarins
Leontopithecus chrysomelas in Una Reserve, Brazil. Dodo J Wildl Preserv Trusts 32:115–122
Dinerstein E, Olson D, Joshi A, Vynne C, Burgess ND, Wikramanayake E et al (2017) An
ecoregion-based approach to protecting half the terrestrial realm. Bioscience 67(6):534–545
Elleason M, Gua Z, Deng Y, Jiang A, Goodale E, Mammides C (2020) Strictly protected areas are
not necessarily more effective than areas in which multiple human uses are permitted. Ambio
2020:1–16
Fahrig L (2017) Forty years of bias in habitat fragmentation research. In: Kareiva P, Marvier M,
Silliman B (eds) Effective conservation science. Oxford University Press, Oxford
Filho AA (1978) Sul da Bahia: Chão de Cacau (Uma Civilização Regional). Editora Civilização
Brasileira, Rio de Janeiro
Franco-Moraes J, Baniwa AF, Costa FR, Lima HP, Clement CR, Shepard GH Jr (2019) Historical
landscape domestication in ancestral forests with nutrient-poor soils in northwestern Amazonia.
For Ecol Manag 446:317–330
Haggar J, Pons D, Saenz L, Vides M (2019) Contribution of agroforestry systems to sustaining
biodiversity in fragmented forest landscapes. Agric Ecosyst Environ 283:106567
Heckenberger M, Neves EG (2009) Amazonian archaeology. Annu Rev Anthropol 38:251–266
23 Agroecology and Forest Conservation in Three Types of Land. . . 597

Heckenberger MJ, Russell JC, Fausto C, Toney JR, Schmidt MJ, Pereira E, Franchetto B, Kuikuro
A (2008) Pre-Columbian urbanism, anthropogenic landscapes, and the future of the Amazon.
Science 321(5893):1214–1217
Holmes G, Cavanagh CJ (2016) A review of the social impacts of neoliberal conservation:
formations, inequalities, contestations. Geoforum 75:199–209
Hunter ML, Yonzon P (1993) Altitudinal distributions of birds, mammals, people, forests, and
parks in Nepal. Conserv Biol 7(2):420–423
Joppa LN, Pfaff A (2009) High and far: biases in the location of protected areas. PLoS One 4(12):
e8273. [Link]
Joppa LN, Pfaff A (2011) Global protected area impacts. Proc R Soc B Biol Sci 278(1712):
1633–1638
Jose S (2009) Agroforestry for ecosystem services and environmental benefits: an overview.
Agrofor Syst 76(1):1–10
Jose S (2012) Agroforestry for conserving and enhancing biodiversity. Agrofor Syst 85(1):1–8
Kohler F, Brondizio ES (2017) Considering the needs of indigenous and local populations in
conservation programs. Conserv Biol 31(2):245–251
Kopnina H (2016) Half the earth for people (or more)? Addressing ethical questions in conserva-
tion. Biol Conserv 203:176–185
Kramer R, van Schaik C, Johnson J (eds) (1997) Last stand: protected areas and the defense of
tropical biodiversity. Oxford University Press, Oxford
Laurance WF (2008) Theory meets reality: how habitat fragmentation research has transcended
island biogeographic theory. Biol Conserv 141(7):1731–1744
Laurance WF, Camargo JL, Fearnside PM, Lovejoy TE, Williamson GB, Mesquita RC, Meyer CF,
Bobrowiec PE, Laurance SG (2018) An Amazonian rainforest and its fragments as a laboratory
of global change. Biol Rev 93(1):223–247
Lele S, Wilshusen P, Brockington D, Seidler R, Bawa K (2010) Beyond exclusion: alternative
approaches to biodiversity conservation in the developing tropics. Curr Opin Environ Sustain
2(1–2):94–100
Leopold A (1933) Game management, 1986 edn. University of Wisconsin Press, Madison, 481pp
Levis C, Costa FR, Bongers F, Peña-Claros M, Clement CR, Junqueira AB, Neves EG, Tamanaha
EK, Figueiredo FO, Salomão RP, Castilho CV (2017) Persistent effects of pre-Columbian plant
domestication on Amazonian forest composition. Science 355(6328):925–931
Levis C, Peña-Claros M, Clement CR, Costa FR, Alves RP, Ferreira MJ, Figueiredo CG, Bongers F
(2020) Pre-Columbian soil fertilization and current management maintain food resource avail-
ability in old-growth Amazonian forests. Plant Soil 450:29–48
Marsden C, Martin-Chave A, Cortet J, Hedde M, Capowiez Y (2020) How agroforestry systems
influence soil fauna and their functions – a review. Plant Soil 453(9):29–44
Mascarenhas GCC, Midlej RR, Trevizan SP, Afonso JM, Andrade Filho EN, de Medeiros JX
(1999) O cluster do cacau no sul da Bahia. In: Haddad PR (ed) A Competitividade do
Agronegócio e o Desenvolvimento Regional no Brasil: Estudo de Clusters. CNPq/EMBRAPA,
Brasilia, pp 59–123
McNeely JA, Schroth G (2006) Agroforestry and biodiversity conservation–traditional practices,
present dynamics, and lessons for the future. Biodivers Conserv 15(2):549–554
Montagnini F, del Fierro S (2022) Functions of agroforestry systems as biodiversity islands in
productive landscapes. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments, Topics in Biodiversity and Conservation. Springer, Cham,
pp 89–116
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity islands: strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity Islands: strategies
for conservation in human-dominated environments, Topics in Biodiversity and Conservation.
Springer, Cham, pp 1–35
Morse RM (1974) From community to metropolis: a biography of São Paulo, Brazil. Octagon Press,
London
598 K. R. Painter et al.

Nelson A, Chomitz KM (2011) Effectiveness of strict vs. multiple use protected areas in reducing
tropical forest fires: a global analysis using matching methods. PLoS One 6(8):e22722
Noss RF, Dinerstein E, Gilbert B, Gilpin M, Miller BJ, Terborgh J, Trombulak S (1999) Core areas:
where nature reigns. In: Soulé ME, Terborgh J (eds) Continental conservation: scientific
foundations of regional reserve networks. Island Press, Washington, DC, pp 99–128
Odum EP (1971) Fundamentals of ecology, 3rd edn. Saunders, Philadelphia
Olson DM, Dinerstein E (1998) The Global 200: a representation approach to conserving the
Earth’s most biologically valuable ecoregions. Conserv Biol 12(3):502–551
Painter KR (2006) Effectiveness of a non-governmental organization (NGO) extension program for
forest conservation and sustainable agroforestry in southern Bahia, Brazil. Master’s thesis,
University of Florida, Gainesville, FL
Palacios Bucheli VJ, Bokelmann W (2017) Agroforestry systems for biodiversity and ecosystem
services: the case of the Sibundoy Valley in the Colombian province of Putumayo. Int J
Biodivers Sci Ecosys Serv Manag 13(1):380–397
Palm C, Blanco-Canqui H, DeClerck F, Gatere L, Grace P (2014) Conservation agriculture and
ecosystem services: an overview. Agric Ecosyst Environ 187:87–105
Pivello VR (2011) The use of fire in the Cerrado and Amazonian rainforests of Brazil: past and
present. Fire Ecol 7:24–39
Polimeni JM, Mayumi K, Giampietro M, Alcott B (2015) The Jevons paradox and the myth of
resource efficiency improvements. Routledge, Oxfordshire/New York
Porter-Bolland L, Ellis EA, Guariguata MR, Ruiz-Mallé I, Negrete-Yankelevich S, Reyes-García V
(2012) Community managed forests and forest protected areas: an assessment of their conser-
vation effectiveness across the tropics. For Ecol Manag 268:6–17
Powlson DS, Gregory PJ, Whalley WR, Quinton JN, Hopkins DW, Whitmore AP, Hirsch PR,
Goulding KW (2011) Soil management in relation to sustainable agriculture and ecosystem
services. Food Policy 36(Supplement 1):S72–S87
Putz FE (2020) Science-to-conservation disconnections in Borneo and British Columbia. For Chron
96(1):20–26
Radachowsky J, Ramos VH, McNab R, Baur EH, Kazakov N (2012) Forest concessions in the
Maya Biosphere Reserve, Guatemala: a decade later. For Ecol Manag 268:18–28
Reganold JP, Wachter JM (2016) Organic agriculture in the twenty-first century. Nat Plants 2(2):
15221
Ribeiro MC, Martensen AC, Metzger JP, Tabarelli M, Scarano F, Fortin MJ (2011) The Brazilian
Atlantic forest: a shrinking biodiversity hotspot. In: Zachos FE, Habel JC (eds) Biodiversity
hotspots: distribution and protection of conservation priority areas. Springer, Berlin/Heidelberg,
pp 405–434
Rice RA, Greenberg R (2000) Cacao cultivation and the conservation of biological diversity.
AMBIO 29(3):167–173
Ross NJ (2011) Modern tree species composition reflects ancient Maya “forest gardens” in
northwest Belize. Ecol Appl 21(1):75–84
Sandhu HS, Wratten SD, Cullen R (2010) Organic agriculture and ecosystem services. Environ Sci
Pol 13(1):1–7
Santos PZF, Crouzeilles R, Sansevero JBB (2019) Can agroforestry systems enhance biodiversity
and ecosystem service provision in agricultural landscapes? A meta-analysis for the Brazilian
Atlantic forest. For Ecol Manag 433:140–145
Saracura VF (1997) Reserva Biológica de Una: Plano de Manejo. ICMBio, Brasilia. [Link]
[Link]/portal/images/stories/imgs-unidades-coservacao/REBIO%[Link]
Schroth G, da Fonseca GAB, Harvey CA, Gascon C, Vasconcelos HL, Izac AMN (eds) (2004)
Agroforestry and biodiversity conservation in tropical landscapes. Island Press, Washington,
DC/Covelo/London
Scott JM, Davis FW, McGhie RG, Wright RG, Groves C, Estes J (2001) Nature reserves: do they
capture the full range of America’s biological diversity? Ecol Appl 11(4):999–1007
23 Agroecology and Forest Conservation in Three Types of Land. . . 599

Sebesvari Z, Le HTT, Van Toan P, Arnold U, Renaud FG (2012) Agriculture and water quality in
the Vietnamese Mekong Delta. In: Renaud FG, Künzer C (eds) The Mekong Delta system.
Springer, Dordrecht, pp 331–361
Shepard GH Jr, Neves E, Clement CR, Lima H, Moraes C, dos Santos GM (2020) Ancient and
traditional agriculture in South America: tropical lowlands. In: Oxford research encyclopedia of
environmental science. Oxford University Press, New York
Silbernagel J (2003) Spatial theory in early conservation design: examples from Aldo Leopold’s
work. Landsc Ecol 18:635–646
Smith T (2001) Beyond land rights. Newsweek International. January 21, 2002, p. 26
Soares-Filho B, Moutinho P, Nepstad D, Anderson A, Rodrigues H, Garcia R, Dietzsch L, Merry F,
Bowman M, Hissa L, Silvestrini R (2010) Role of Brazilian Amazon protected areas in climate
change mitigation. Proc Natl Acad Sci 107(24):10821–10826
Trevizan SP (1982) Estrutura fundiária e produtividade na região cacaueira da Bahia. Boletim
Técnico 103. Centro de Pesquisas do Cacau (CEPLAC), Bahia, Brasil. pp 3–20
Udawatta PR, Rankoth L, Jose S (2019) Agroforestry and biodiversity. Sustainability 11(10):2879
Vélez MA, Robalino J, Cardenas JC, Paz A, Pacay E (2020) Is collective titling enough to protect
forests? Evidence from Afro-descendant communities in the Colombian Pacific region. World
Dev 128:104837
West P, Igoe J, Brockington D (2006) Parks and peoples: the social impact of protected areas. Annu
Rev Anthropol 35:251–277
Wood A, Tolera M, Snell M, O’Hara P, Hailu A (2019) Community forest management (CFM) in
south-west Ethiopia: maintaining forests, biodiversity and carbon stocks to support wild coffee
conservation. Glob Environ Chang 59:101980
Wuerthner G, Crist E, Butler T (eds) (2015) Protecting the wild: parks and wilderness, the
foundation for conservation. Island Press, Washington, DC/Covelo/London
Ybarra M (2018) Green wars: conservation and decolonization in the Maya forest. University of
California Press, Oakland
York R, McGee JA (2016) Understanding the Jevons paradox. Environ Soc 2(1):77–87
Chapter 24
Preserving Biodiversity in Appalachian
Mixed Mesophytic Forests Through
the Permit-Based Harvest of American
Ginseng and Other Forest Botanicals

Karam Sheban

Abstract Appalachian mixed mesophytic forests are home to some of the world’s
most diverse temperate forests; 90% of their plant biodiversity is in the understory.
Many of these understory plants are of great cultural and/or commercial significance,
making their protection vital from ecosystem, sociocultural, and economic perspec-
tives. Due to development and forest fragmentation across the Eastern United States,
large, relatively undisturbed tracts of Appalachian mixed mesophytic forests exist
primarily in federally protected national parks and forests. Protected national forests
are managed under a multiple-use mandate; they exist to protect native forest and
also to provide services to the public such as recreation and harvestable forest
products. A limited number of national forests have permit-based harvest programs
for medicinal and culinary understory plants, some of which, like American ginseng,
are commercially valuable. Demand for these plants and concern over their conser-
vation status, however, is resulting in national forests shrinking their permit-based
harvest programs or eliminating them altogether. Contrary to this trend, I argue that a
renewal of the relationship between people and culturally significant understory
plants—through the expansion of permit-based harvest programs on national forest-
land as well as through the intentional cultivation of forest herbs (referred to as forest
farming)—presents the best way forward to reestablish viable populations of under-
story herbs in the mature second growth forests of the Appalachian region. In
conjunction with forest farming of understory herbs, permit-based harvesting is an
effective mechanism for safeguarding the benefits of biodiversity islands in mixed
mesophytic forests.

Keywords Deciduous forests · Eastern USA · Forest botanicals · Forest farming ·

Herbs · Understory

K. Sheban (*)
School of Forestry and Environmental Studies, Yale University, New Haven, CT, USA
e-mail: [Link]@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 601
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
602 K. Sheban

24.1 Introduction

The Appalachian mixed mesophytic forest is an ecoregion within the more extensive
deciduous forests ranging across the Eastern United States. This forest type repre-
sents the epicenter of temperate forest biodiversity in the U.S. and is characterized by
the wealth of species occupying each stratum of the forest, from the canopy down
into the understory. The canopy of Appalachian Mixed Mesophytic Forests is
incredibly diverse, with a single site often supporting more than 30 tree species
(Loucks et al. n.d.). The understory is the main source of floral biodiversity in these
forest systems, however, accounting for 90% of the plant diversity in the region
(Gillium 2007).
While mixed mesophytic forest used to cover a much greater extent across North
America, its range has been reduced considerably, and the remaining tracts have
experienced widespread degradation; it is estimated that 95% or more of this habitat
has been converted or degraded at some point over the past 200 years (Loucks et al.
n.d.). Forest cover generally has increased across the Eastern U.S. in this timeframe,
with the abandonment of cropland, pasture, and other cleared lands, although the
trend has begun to reverse in the twenty-first century (Drummond and Loveland
2010); however, these second-growth forests do not contain the same species
associations or diversity as are seen in primary forests (Braun 1951; Gilliam
2014). In particular, populations of slow-growing herbaceous understory species
take many years to reach reproductive maturity, struggle to disperse widely, and in
most cases do not recover fully in secondary forests (Duffy and Meier 1992).
Herbaceous understory plants provide not only tremendous ecological value, but
sociocultural value as well. The understory of Appalachian mixed mesophytic
forests is home to over 50 species that have been harvested for hundreds or
thousands of years for use in indigenous, traditional, and folk medicine, cooking,
and spiritual practices (Burkhart and Jacobson 2009; Chamberlain 2005). A number
of these plants—such as American ginseng (Panax quinquefolius), goldenseal
(Hydrastis canadensis) and ramps (Allium triccocum)—have significant commercial
value; prices paid for ginseng plants harvested in the wild can reach over $1,000 per
pound (Hamilton 2017).
The remaining intact tracts of Appalachian mixed mesophytic forest can be
considered biodiversity islands, “area[s] of high biodiversity within ecologically
degraded, human dominated landscapes” (Montagnini et al. 2022). These tracts are
largely contained within state and federally controlled public lands, such as national
parks and national forests across the Appalachian region (Loucks et al. n.d.).
Of these remaining tracts, only the national forests allow varying degrees of
permit-based understory plant harvesting. This is in keeping with the principles
laid out in the Multiple-Use Sustained-Yield Act of 1960 passed by the United States
Congress, a guiding piece of legislation that dictates that national forests are
managed for a variety of uses, including timber harvesting, livestock grazing,
watershed protection, fish and wildlife, and recreation. Along with the harvest of
firewood and the hunting of animals such as deer, squirrels, and turkeys, the
permitted harvesting of understory plants on select national forests takes place
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 603

during a designated season. These permits are open to the public for a nominal fee
and are limited to one permit per person per season. Permit-based plant harvesting
confers many advantages: it creates a legal pathway for plant resource access and
use; generates funds for the monitoring of plant harvest levels and the allocation of
personnel; engages the public in the conservation and management of forest biodi-
versity; and elevates the profile of understory plants within the national forest
system.
In this chapter I will argue that the permit-based harvest of understory plants in
national forests represents a strategic and just way to conserve and restore
populations of dispersal-limited understory plants and to protect these threatened
islands of mixed mesophytic biodiversity, while simultaneously preserving Appala-
chian traditions and traditional livelihoods. Critically, I will suggest that this strategy
is contingent on the parallel development of markets for understory herbs through
the expansion of intentional cultivation operations using the agroforestry practice of
forest farming. This chapter will start with an overview of mixed-mesophytic forests
and the unique biodiversity they house. It will cover some of the threats facing the
biodiversity of these forests. The chapter will briefly touch on the history of
understory plant harvesting in the Appalachians, focusing on American ginseng as
a case study in how biodiversity, the management of public lands, market forces, and
cultural traditions can collide. And lastly, the chapter will consider how shifting the
supply chain balance of commercially valuable herbs from wild-harvested toward
sustainably cultivated products could support a permit-based allowable harvest on
public lands, achieving multiple social and ecological benefits.

24.2 The Appalachian Mixed Mesophytic Forest

The Appalachian mixed mesophytic forest of the Eastern United States was first
described by the forest ecologist and University of Cincinnati professor E. Lucy
Braun in 1916 (Braun 1916). Using Braun’s nomenclature, deciduous forests in the
Eastern U.S. are organized into forest associations, defined by the dominant tree
species found within them. Mixed mesophytic forests represent the oldest and most
complex of these forest associations; “the epicenter of highest development of the
Eastern deciduous forest” (Rosson 2008). They are situated in the geographic center
of Eastern U.S. deciduous forests. From this mixed forest or the preceding forests of
the Tertiary period, all other forest associations within eastern deciduous forests
have arisen (Braun 1951).
Appalachian mixed mesophytic forest ranges from Northwest Alabama and East-
Central Tennessee through Eastern Kentucky, Western North Carolina, the majority
of West Virginia, up to Southeast Ohio and Southwest Pennsylvania (Fig. 24.1).
Within this range, the fullest expression of this forest type is seen in ravines, gorges,
coves, and valleys where the topography is varied enough to support a multitude of
species (Braun 1951; Rosson 2008) (Fig. 24.2). While the majority of forest asso-
ciations in the U.S. are dominated by two or three primary canopy species,
604 K. Sheban

Fig. 24.1 Appalachian mixed mesophytic forests range map. (Source: USGS Ecoregions Index
Map, [Link]

ecological complexity is the defining feature of mixed mesophytic forests, with up to

30 tree species on a single site.1 In addition to floral diversity, these forests house
incredibly diverse fungi, songbird, salamander, land snail, and beetle communities.
Their freshwater communities “are the richest temperate freshwater ecosystems in
the world, with high species richness and endemism in mussels, fish, crayfish, and
other invertebrates” (Loucks et al. n.d.).

1
For an incomplete list of tree, shrub, and herbaceous species found in Appalachian Mixed
Mesophytic Forests see appendix at the end of this chapter.
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 605

Fig. 24.2 The Red River Gorge in the Daniel Boone National Forest, a national forest in the heart
of the Appalachian mixed mesophytic range. (Photo: [Link])

Mixed mesophytic forests once covered much of the temperate regions of the
Northern Hemisphere, across what is now North America and the deciduous regions
of China, though their range has shrunk considerably due to the pressures of
colonization and development. In the Eastern United States, it is estimated that
half or more of the present forest cover has been cut some time in the past three
centuries as a result of agricultural expansion, development, and harvesting of
fuelwood and other resources (Drummond and Loveland 2010). Beginning in the
early 1900s as settlement and agriculture moved westward, the Eastern United States
experienced a gain in forest cover, including natural forest regeneration across the
range of mixed mesophytic forest. But that trend has reversed, as logging, urbani-
zation, and the urban sprawl of low-density settlement have begun affecting forest
regrowth. The Eastern U.S. experienced a 2.3% decline in forest cover between 1973
and 2000—a total loss of over 165,000,000 hectares—with regions of mixed
mesophytic forest, such as the Piedmont and the Central and Southwestern Appala-
chians experiencing declines in forest cover at an even faster rate (Drummond and
Loveland 2010). Since 2000, the trend has largely continued and projections suggest
forest cover across the Eastern U.S. will continue to decline in the coming decades
(Adams et al. 2019).
The contemporary trend of forest decline indicates the urgency of preserving the
remaining forests, despite many of these being the result of secondary growth.
Mature second growth forest can contain high levels of biodiversity, as well as
eventually achieve an old-growth structure. E. Lucy Braun observed that, on the best
606 K. Sheban

sites within the mixed mesophytic forest association, the regrowth may return to a
similar species composition of an old growth forest if the forest was only cut once
and not utilized for other purposes such as agriculture (Braun 1951). More chronic
disturbance, such as a century of use as pasture or for agriculture, can leave behind a
significantly transformed landscape, more susceptible to colonization by inva-
sive species (Webb et al. 2000), in turn suppressing growth and reproduction of
native species (Miller and Gorchov 2004).

24.3 The Herbaceous Layer: Challenges and Opportunities

Only recently have scientists began to appreciate that the herbaceous layer in
deciduous forests—their greatest source of biodiversity—is much more sensitive
to disturbance than the canopy layer (McCarthy 2014). In a landmark paper, Duffy
and Meier (1992) concluded that herb populations in mixed mesophytic forests
would take several centuries to recover after a major disturbance. Additional
research has documented the difficulty understory species face in recolonizing
former forest habitat after a disturbance such as logging or agriculture (Holmes
and Matlack 2018).
The challenges to recolonization are myriad. Understory herbs are limited in their
sexual reproduction due to small crops of seed, which are often poorly dispersed by
gravity or insects, and generally have low germination rates (Duffy and Meier 1992).
For species that reproduce asexually, the ability to move via vegetative sprouting is
limited to less than 1 m per decade (Sobey 1977), and it takes years for an individual
plant to reach sexual maturity for many species (Bierzychudek 1982). On top of
these challenges, predation by deer and other animals can present a serious challenge
to population establishment (Alverson et al. 1988; McGraw and Furedi 2005).
The restoration of understory populations and the preservation of existing wild
populations provide two critical approaches to protecting understory biodiversity in
a landscape facing the dual pressures of deforestation and fragmentation (Drum-
mond and Loveland 2010; Loucks et al. n.d.). While important, the first approach—
restoring populations of threatened understory herbs—presents some challenges.
Restoration plantings require funding, personnel, and botanical expertise, all of
which are particularly limited when it comes to restoration of understory herb
species. Additionally, the reintroduction of native understory plants into the wild
requires a seed or transplant source, which for many understory species simply is not
available. For species such as American ginseng, for which planting stock is
available, the introduction of ginseng seed with an industrial provenance (primarily
produced on large industrial ginseng farms in Wisconsin and Quebec) into geo-
graphic regions across the plant’s native range raises concerns about maladaptation
and the corruption of local genetics (Schluter and Punja 2002). There is a critical
need for native plant nurseries to provide planting stock for restoration plantings,
private forest farming operations, and local botanical gardens and sanctuaries.
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 607

The second approach to preserving existing wild populations of understory

plants—what McCoy et al. (2019) refers to as the in-situ conservation of medicinal
and social-use understory taxa—represents the most cost-effective path forward.
While expertise and funding are required for this approach as well, starting with an
existing botanical resource eliminates some of the initial challenges facing protection
efforts. It is critical that in-situ conservation and restoration take place on private
forestland, and this is occurring through a variety of mechanisms which we will
cover later in detail. United Plant Savers, a nonprofit organization headquartered in
Rutland, Ohio, has established a network of “botanical sanctuaries” on private lands,
providing a framework for preserving habitat and populations of threatened native
species with an emphasis on medicinal understory plants ([Link].
org). The Appalachian Beginning Forest Farmers Coalition, a consortium of univer-
sities, government agencies, and nonprofit organizations, promotes the cultivation of
at-risk understory species in forest farming systems. These programs are based on
the premise of “Conservation Through Cultivation,” which posits that by cultivating
and selling commercially valuable understory plants, pressure can be taken off wild
populations as the supply chain transitions towards sustainably grown products
([Link]) (Burkhart and Jacobson 2009; Burkhart
2011). This approach has additional benefits such as maintaining private lands as
forest, generating land-based income, and strengthening the local forest and farming
sectors by diversifying the array of crops producers can cultivate. Given contempo-
rary land use trends, these methods of plant conservation on private lands are
becoming more important than ever and adoption rates of both practices are growing.

24.4 Human Use and Harvest of Understory Plants

in the Appalachians

Protecting public lands is critical. It would be misguided, however, to suggest that

the only way to preserve biodiversity islands within public lands is to create reserves
where access to plant material is sealed off from the public. This is a well-established
pattern on public lands in the U.S.: adopting a protectionist approach toward natural
resources in the name of saving them. While the U.S. is frequently lauded interna-
tionally for its public land conservation, the protectionist model it employs has been
criticized. Embodied by the National Park Service (NPS), this approach to public
lands conservation is sometimes referred to as the “Yellowstone model,” and has
been exported around the world. A wealth of academic literature, however, suggests
that this model of closing off of public land has been harmful to vulnerable rural
populations and provides national economic benefits at the expense of local com-
munities while incurring high costs for park protection and failing to effectively
conserve biodiversity (Schelhas 2001). Even if this approach to land management
was resulting in irrefutably positive changes in biodiversity—which, in the case of
understory plants, is dubious—there is an increasing consensus among academics,
608 K. Sheban

policy makers, and communities around the world that considering simply the
ecological effects of a particular policy divorced from its broader social, economic,
and political implications is not enough (Ban et al. 2013).
It is important to note that the distribution of medicinal and culinary understory
plants across the Eastern United States is inextricably linked to human use over
millennia (Abrams and Johnson 2017). This is important for several reasons, but
particularly in the context of how and to whom we allow or disallow access to plant
material in the name of saving it. Historically, the harvest of understory plants for
personal use and for income generation has taken place across a great forest-
commons, with place-based, plant-specific knowledge passed down generationally.
This was true prior to the colonization of North America and has remained true
across the Appalachian region, throughout the mixing of traditional plant-based
medicinal knowledge from old-world cultures—such as Spanish, African, British,
and French—with indigenous practices (Light 2008). These traditions persist
throughout the Appalachian region, where the biological diversity of mixed meso-
phytic forests and relative isolation kept plant-based medicinal and cultural tradi-
tions alive (Alwhaibi et al. 2017). Across the range of mixed mesophytic forest, the
knowledge of how to harvest and use plants, along with practices of good steward-
ship, was transferred intergenerationally. For understory herbs, this included prac-
tices such as only harvesting mature plants with ripe berries, planting seeds before
harvesting the plant, and leaving behind the most productive plants (“mother”
plants) to replenish local populations (Chittum et al. 2019).
Considering the historical and contemporary interactions between people and
understory plants in the mixed mesophytic forests of the Appalachian region, it is
important to acknowledge that humans have and continue to function as dispersal
agents for the understory biodiversity of the region. Humans have been harvesting
and replanting medicinal and culinary herbs for millennia; this stewardship relation-
ship is one of the main ecological dynamics responsible for the “natural” distribution
of many understory plants. In fact, given the long history of human-plant interac-
tions, some have made the argument that it is impossible to disentangle anthropo-
genic influence from the observed ecology of understory plants in the Appalachian
region and beyond, and that it would be misguided to predict the future of plants
without considering the role humans may play (Turner and McGraw 2015;
Albuquerque et al. 2018).
When considering historical human-plant interactions in forests, the question
arises as to whether these interactions have positive or negative impacts on biodi-
versity. There is evidence that direct seeding of understory plants improves germi-
nation rates (Filyaw and Sheban 2015) and that plant reintroduction into mature
second growth forests provides an avenue for circumventing the aforementioned
dispersal limitations that plague understory herbs. It may be that a renewal of this
relationship with understory plants presents the best way forward to reestablish
viable populations of understory herbs in the mature second growth forests of the
Appalachian region. The time to act is now—these traditional and indigenous
economies and this way of life are disappearing (Cavender and Beck 1995) along
with the primary forests on which they have historically depended. Shifting patterns
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 609

Fig. 24.3 A mature

American Ginseng plant.
(Photo by Karam Sheban)

of land ownership, the closing off of public space, the unsustainable harvest of plant
resources, and the reduction of undisturbed forest habitat are moving the baseline on
both forest health and forest livelihoods.

24.5 American Ginseng as a Bellwether: The Future

of Understory Plants in Appalachian Mixed
Mesophytic Forests

24.5.1 American Ginseng Biology and Brief History

When considering the cultural, economic, and ecological forces acting upon under-
story biodiversity across mixed mesophytic forests, American ginseng provides a
particularly useful case study in understanding both threats and opportunities
confronting forest herbs. American ginseng (Panax quinquefolius, Araliaceae) is a
long-lived perennial herb, reaching a maximum height of 50 cm (Anderson et al.
2002) (Fig. 24.3).
A mature ginseng plant has three to four palmately lobed leaves, each with three
or five leaflets, radiating from a central aerial stem, which itself grows out of the
underground root. Ginseng diggers commonly refer to the leaves of a ginseng plant
as its “prongs,” and the number of prongs provide an indication of the plant’s
610 K. Sheban

Fig. 24.4 The lifecycle of American ginseng. (Source: Rural Action ([Link]))

maturity (Anderson et al. 2002). The plant develops leaves in early spring, and its
flower emerges midsummer. The fruit ripens around late August, producing a cluster
of red berries, each carrying one or two seeds, and the leaves die back in late summer
to early fall (Fig. 24.4). Ginseng roots form arbuscular mycorrhizal relationships,
which help the plant access soil nutrients (Anderson et al. 2002).
Ginseng naturally occurs from Southern Quebec and Ontario, to South Dakota,
throughout the Midwest, and south to Georgia and Oklahoma (Anderson et al. 2002)
(Fig. 24.5). It tends to occur in moderately to highly shaded hardwood forests on
slopes under a canopy of mesic trees species, such as sugar maple (Acer saccharum),
red oak (Quercus rubra), American beech (Fagus grandifolium), shagbark and
mockernut hickory (Carya ovata and C. tomentosa), and white ash (Fraxinus
americana). These trees are characteristic of the ravines, coves, gorges, and valleys
that Braun identified as home to the fullest diversity of mixed mesophytic forest
(Braun 1951). Ginseng prefers moist but well-drained sites and can grow under a
range of edaphic conditions, though research has indicated that high calcium levels
are particularly important in promoting plant growth and development (Davis and
Persons 2014).
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 611

Fig. 24.5 Native range of American ginseng (Panax quinquefolius), from the USDA Plants
Database

American ginseng has been used culturally and medicinally for thousands of
years across the globe. In indigenous and folk medicine in North America and in
traditional medicine across Eastern Asia, American ginseng (along with Korean
ginseng, Panax ginseng, and other ginseng species) has been used to treat a wide
range of health conditions and ailments, from the common cold to erectile disfunc-
tion, as well as being used generally to combat stress and boost energy. Ginseng’s
purported ability to mitigate stress led to its categorization—in the parlance of
modern herbal medicine—as an adaptogen, a substance “with a relatively high
degree of safety that help[s]. . .organism[s] adapt to various types of stress”
(Dharmananda 2002). These traditional uses have inspired contemporary examina-
tions of the plant’s medicinal potential. Modern pharmacology has attributed a range
of medicinal properties to American ginseng, the majority of which are thought to
derive from a family of bioactive phytochemical compounds, known as
ginsenosides, which are abundant in the plant’s leaves, stems, and roots (Szczuka
et al. 2019). These compounds are actively being studied for their potential
anti-diabetic, anti-obesity, anti-cancer, anti-aging, and antimicrobial properties, as
well as their ability to alleviate the symptoms of a wide range of other conditions
(Szczuka et al. 2019).
Ginseng is unique among understory plants as “the most economically important
wild-harvested native medicinal plant in the United States” (Division of Scientific
612 K. Sheban

Authority 2018). It has long been harvested and used in North America, but it is also
the centerpiece of major international trade between North America and China, the
primary importer of American ginseng. This trade drives the significant harvest of
ginseng plants from the wild; nearly 8.5 million ginseng plants were harvested from
the wild in 2017, according to data reported by states and Tribes with export
programs (Division of Scientific Authority 2018).
The extensive harvesting of wild ginseng to supply the international market has
been ongoing for over 300 years, since the first shipment of ginseng roots from North
America to China in 1717 (Parsons 2016). Due to concerns around the plant’s
conservation, ginseng was listed as a controlled species under Appendix II of the
Convention on International Trade in Endangered Species (CITES) in 1975, signi-
fying an acknowledgement that ginseng was “not necessarily now threatened with
extinction but that [it] may become so unless trade is closely controlled” (https://
[Link]/eng/app/[Link]).
The challenge presented by forest botanical species, but especially American
ginseng—as its commercial value is orders of magnitude greater than other mer-
chantable temperate herb species—is a strong incentive to harvest the plant when it is
encountered. For a potential harvester, leaving the plant in the ground passes the
opportunity on to another harvester. And unlike some other non-timber forest prod-
ucts (NTFPs)—such as collecting maple sap to produce syrup or gathering berries or
tree nuts—harvesting a ginseng root, the merchantable part of the plant, is lethal.
Considering that American ginseng is dispersal-limited and typically requires a
minimum of 5 years to reach reproductive maturity, its high commercial value and
the lethality of harvest has created conservation challenges across ginseng’s entire
range. And yet, as put by Curt Freese in his book “Harvesting Wild Species,” “[r]
egardless of what conservationists think, much of the world depends on wild species
for an array of products, whether for food, fiber, or medicine. Thus, in many cases, the
question is not whether to use wild species, but rather how to move from a system of
use that is clearly not sustainable toward one that is better” (Freese 1997).

24.5.2 Permit-Based Harvesting of Ginseng

Currently, the harvest of wild ginseng plants is allowed in 19 states—Alabama,

Arkansas, Georgia, Illinois, Indiana, Iowa, Kentucky, Maryland, Minnesota, Mis-
souri, New York, North Carolina, Ohio, Pennsylvania, Tennessee, Vermont, Vir-
ginia, West Virginia, and Wisconsin—and also in the territory of the Menominee
Indian Tribe of Wisconsin. While the monitoring and regulation of American
ginseng falls under the federal purview of the U.S. Fish and Wildlife Service, the
states bear the bulk of actual oversight, regulating the harvest, sale, and certification
of wild ginseng roots within their jurisdiction. The purpose of these regulations is to
promote the regeneration of plant populations. To achieve this end, the states and
tribes enforce a harvest season, a minimum harvest size (3 or 4 leaves) and/or age
(5 to 10-year-old plants), and certain sustainable practices such as planting the seeds
of harvested plants on site (Division of Scientific Authority 2018).
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 613

The proliferation of private property across the United States and the increasing
ownership of forest and farm lands by absentee landowners (Petrzelka et al. 2013)
has shifted control over and access to plant material across large swaths of the
Eastern United States. Now more than ever, public lands provide one of the only
avenues for legal plant harvest for landless Americans. The trend of restricting
access to plant resources that have traditionally existed in a commons continues
today. Most states (17 out of 19) do not allow harvest on state lands, and only five
states allow the harvesting of plants on national forest land. Within these states, the
number of national forests that allow the harvesting of plants is shrinking. In 2016,
the Daniel Boone National Forest in Kentucky shut down its permit-based harvest
program due to conservation concerns (Division of Scientific Authority 2018).
Since 2013, in national forests where ginseng harvesting is still permitted, the
number of permits issued, the allowable harvest quantity, and the length of the harvest
season have all been reduced (Division of Scientific Authority 2018). This is not
without some scientific justification. Understory plant populations are being affected
by the harvest of wild populations for commercial gain, but without baseline data on
population numbers for various merchantable understory plants, it is difficult to say
exactly how significant that effect is. Regarding ginseng, there is research showing
that on lands that have an allowable ginseng harvest there is a higher proportion of
juvenile compared to adult (i.e., sexually mature) plants than on lands where harvest
is not allowed (Cruse-Sanders and Hamrick 2004; Young et al. 2009; McGraw et al.
2013). And beyond ginseng, regular harvests of large quantities of other understory
medicinal and edible plants—such as goldenseal (Hydrastis canadensis), ramps
(Allium triccocum), and black cohosh (Actaea racemosa)—raise questions of sus-
tainability. A recently developed initiative out of Virginia Tech University,
RootReport, has begun to track the harvest quantities for 14 of the more commonly
traded native forest medicinal species ([Link] Data
collection efforts such as this will be necessary for understanding the long-term
ecological trends of forest understory species going forward.
While additional research on forest herbs other than ginseng is certainly needed,
in the meantime, McGraw et al. (2013) suggest that because we know more about
ginseng than other understory herbs, it can be used as a “phytometer”—a species that
can be used as a measure of physiological responses to various environmental
factors—to “better understand how environmental changes are affecting many
lesser-known species that constitute the diverse temperate flora of Eastern North
America” (McGraw et al. 2013). Ginseng’s economic arc may provide an indication
of the commercial future of other understory species as well. As the herbal products
market continues to grow—the international market for herbal medicine was valued
at $60 billion dollars in 2019 (Ahmad Khan and Ahmad 2019)—and as wild plant
populations become increasingly scarce, demand and prices can be expected to rise
broadly for all merchantable plants. Indeed, there has been a recent spike in prices
paid for goldenseal, resulting in shifting demand for the herb and likely having an
effect on wild populations (Tanner Filyaw, personal communication). Thus,
although ginseng’s history, cross-cultural significance, and high value make it
unique among understory plants, we can likely expect other species to follow suit,
both ecologically and commercially.
614 K. Sheban

In the face of these trends—increasing demand for plant material and decreasing
access of formerly communal plant resources—and in considering how best to
protect biodiversity islands within the forests of the Eastern United States, the
U.S. is reaching an inflection point. If decreasing forest biodiversity in Appalachian
mixed mesophytic forests can be understood broadly as the consequence of coloni-
zation, development, and climate change, who should bear the costs associated with
its protection? Are there ways to preserve and even enhance this biodiversity while
also deviating from the historic trend of restricting the access to resources by local
communities and landless Americans? Facilitating a targeted set of social, economic,
and ecological changes in order to maintain and even expand the permit-based
harvest of plant material on public lands represents a way to achieve this.
It has been shown that allowing ginseng harvest on national forestland can impact
the proportion of adult to juvenile wild plants (McGraw et al. 2013), but could it be
possible that an allowable harvest also results in a greater number of juvenile plants
overall? If best practices are required and then followed by legal harvesters on public
land—only harvesting mature plants with ripe berries, replanting those berries on
site—this would be the expected result. Such a consideration has the potential to
subvert the narrative of harvest necessarily undermining long-term population
survival.

24.5.3 Factors to Consider in Allowing Plant Harvest

on Public Lands

In light of this scientific and management ambiguity in how harvest affects wild
American ginseng populations, several factors should be considered when weighing
the value of an allowable harvest on public land. First, the establishment of a permit-
based harvest on national forestland generates revenue which could be used to fund
critical long-term population monitoring efforts. This research is essential for under-
standing how understory plant populations are responding to myriad changes in their
environment, including climate change, herbivory, and harvest pressure.
Population monitoring is taking place in the Nantahala and Pisgah National
Forests in North Carolina, Cherokee National Forest in Tennessee, and Wayne
National Forest in Ohio (Division of Scientific Authority 2018; Tanner Filyaw,
personal communication). In the Wayne National Forest, Forest Product Permits
are separated into one permit for ginseng and one permit for six other species: Blue
Cohosh (Caulophyllum thalictroides), Black Cohosh (Actaea racemosa), White
Snakeroot (Ageratina altissima), Goldenseal (Hydrastis canadensis), Bloodroot
(Sanguinaria canadensis), and Wild Ginger (Asarum canadense). Permits cost $20
each and, with between 60 and 100 permits issued each year, the program generates
between $1,000 and $2,000 annually (Tanner Filyaw, personal communication).
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 615

Fig. 24.6 Forest farming ramps; a demonstration led by Rural Action, a nonprofit in Southeast
Ohio. (Photo by Karam Sheban)

While this is not much, it is enough to fund monitoring efforts on a portion of the
Wayne National Forest. Funds generated from understory plant harvest permits in
Wayne National Forest are also directed towards education efforts in partnership
with Rural Action, a local nonprofit with a history of working with commercially
valuable forest herbs ([Link]), among other groups. This partnership
funds workshops that educate local landowners and community members about the
sustainable harvest and management of at-risk forest herbs. Promoting “Conserva-
tion through Cultivation,” landowners are taught how to forest farm ginseng and
other commercial species such as ramps (Fig. 24.6). Programs such as these raise the
profile of understory diversity, which may otherwise go largely unnoticed and
unfunded in a national forest.
Illegal harvest or “poaching” of ginseng and other wild plants is a major issue
across the Appalachian region, and it takes place on both public and private lands.
There are many factors responsible for rates of ginseng poaching across the region,
from poverty and drug use (Pokladnik 2008; Young et al. 2011) to false perceptions
that stealing ginseng will lead to a quick windfall, promoted by television shows
such as the History Channel’s Appalachian Outlaws. This television show and
others like it (National Geographic’s Smokey Mountain Money) rely on derogatory
stereotypes and depict a lawless, get-rich-quick ginseng economy across the Appa-
lachian region. There are indications that these shows may have been partly respon-
sible for an uptick in rates of ginseng poaching on private lands after their debut in
2014 (Arnold 2016; Hamilton 2017).
616 K. Sheban

The establishment of an allowable harvest on public lands creates a legal avenue

for harvesting ginseng and other medicinals for people who may not own or have
access to private lands for plant material. Given the reality of poaching, the estab-
lishment of plant monitoring efforts represents an effective approach to evaluating
the extent to which poaching is occurring on national forestland. Such efforts could
be partially or fully funded through harvest permits. Additionally, education about
the ecology and sustainable harvest of understory plants—also promoted through
permit-based harvesting—represents a systemic way to combat the theft of valuable
understory herbs.
Currently permit-based harvesting of understory plants is generating only a small
amount of revenue due to the limited number of permitting programs and the prices
paid for understory plants. As demand for herbal products continues to grow, it is
important to consider how permit-based harvesting might evolve to support har-
vesters and plant populations in tandem.

24.6 Forest Farming as a Means to Support and Grow

Permit-Based Harvesting Programs

The majority of traded American botanical products are supplied to markets through
wild-harvesting (Dentali 2007; Dentali and Zimmerman 2012), with some of the
most prominent species being harvested from forests. Scholars, farmers, herbalists,
conservation groups, government agencies, and nonprofit organizations have called
for the intentional cultivation of native North American forest plants, particularly
through systems such as forest farming. Agroforestry practices such as forest
farming confer advantages over field-based cultivation (Bannerman 1997; Robbins
1998, 1999; Gladstar and Hirsch 2000; Burkhart and Jacobson 2009), including
fewer costs and the potentials for higher product quality, reduced disease pressure,
and income generation while preserving or even increasing forest diversity (Teel and
Buck 1998; Hill and Buck 2000; Rao et al. 2004).
I propose that an additional advantage of the intentional cultivation of medicinal
and culinary forest plants in forest farming systems is its potential to support the
permit-based harvest of these same plants on public lands. This is achieved by
stabilizing supply chains, taking pressure off of wild plant populations, and increas-
ing prices for forest-based medicinal and culinary plants. Improving the price points
for a range of understory species would facilitate greater revenue generation on
public lands, allowing for enhanced investment in long-term monitoring, plant
protection, and education and outreach efforts around native biodiversity. Thus,
increasing the support for, as well as the overall viability of, forest farms on private
lands is a necessary step in the protection of remaining islands of native plant
biodiversity while allowing for continued access to plant resource commons. This
could be accomplished by ensuring that forest farmers are receiving prices that
support the cultivation of multiple understory species, rather than a select few.
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 617

However, in order for the transition from wild-harvesting to sustainable cultivation

of understory herbs to equitably impact people who rely on native forest biodiversity
for livelihoods or cultural expression, it must provide opportunities for the most
economically disadvantaged. Increasing prices for understory plants must be trans-
lated into greater investment in NTFP permit-harvesting programs on public lands,
rather than higher prices and greater harvest pressure leading to these programs
being shut down.
Currently, prices paid for American ginseng are extremely high when compared
to other wild-harvested understory species. Research suggests that ginseng and
goldenseal are the only medicinal herb species in the Eastern U.S. that can profitably
be forest farmed, while the price points for other medicinals remain too low to justify
the expense (Burkhart and Jacobson 2009). This dramatic imbalance in price creates
uneven pressures on wild plant populations. Freese (1997) suggests this phenome-
non can be observed in wild-harvested natural commodities around the world. He
cites the example of Latin American mahogany, which commands a very high price
compared to the lumber of species which are considered “lesser-known”, placing
inordinate demand on one species while undervaluing others. In this scenario, one
strategic conservation response would be to grow markets and demand for these
lesser-known species to spread harvest pressure evenly across a range of species
(Freese 1997). While industrial buyers understand that relying on wild-collected
herbal material creates sustainability challenges (as well as issues with adulteration
due to a lack of supply chain transparency and traceability (Jiang et al. 2011), many
are not willing or able to pay prices high enough to encourage the cultivation of
additional understory species.
Recently, however, this has begun to change. United Plant Savers (www.
[Link]) administers a third-party certification system, called “Forest
Grown Verification” (FGV), designating that a given forest farmer is intentionally
and sustainably cultivating plants rather than harvesting them from wild populations.
This certification structure is the result of years of meetings between forest farmers,
Technical Service Providers, and industry groups seeking solutions to supply chain
challenges. Designed in the spirit of Organic certification, the program seeks similar
beneficial outcomes: increased profitability, reduced inputs, greater economic and
ecological resilience, and increased employment potential (Jouzi et al. 2017). Forest
Grown Verification is supported by herbal products companies, such as Oregon-
based Mountain Rose Herbs ([Link]), through the introduc-
tion of FGV product lines for multiple herb species, including ginseng, goldenseal,
and black cohosh. The program is financed partly through consultation and annual
renewal fees, while verified growers are paid a premium for their
sustainably-cultivated product. Industry buyers like Mountain Rose Herbs that
establish product lines for FGV material are critical partners in establishing an herbal
products supply chain based increasingly on cultivated products.
Programs such as FGV have the potential to make forest farming increasingly
profitable, as industry buyers begin to pay forest farmers prices that more accurately
reflect what their cultivated plants are worth. In turn, this increased value is
supported by consumers, willing to pay a premium for a product they believe is
618 K. Sheban

higher quality and produced in a way that is worth supporting. This is the model of
ethical consumption that has helped sustain the growth of organically certified foods
across the globe (Johnston et al. 2011; Sebastiani et al. 2013). Public-lands
harvesting is supported as a consequence, through boosting regional plant
populations and raising overall awareness of at-risk understory biodiversity. At the
same time, higher price-points for lesser-known species could allow for greater
permit fees on public lands, generating more revenue, which could feed into long-
term monitoring and educational programs.

24.7 Discussion

New scientific research has reinvigorated the call to better understand, monitor, and
support understory herb populations (Spicer et al. 2020). Forest farming and public-
harvest programs both represent approaches to raise the profile of understory herbs
and to advance conservation, cultural use, and positive socioeconomic development.
Transitioning the supply of forest botanicals from wild-harvested toward
sustainably-cultivated plants represents a promising opportunity to preserve public
access to plants on public lands while supporting biodiversity islands in mixed
mesophytic forests. This entails formalizing the supply chains (or value chains) for
commercial understory species. Belcher and Schreckenberg (2007) describe this
process in their analysis of the commercialization of non-timber forest products.
They provide a useful framework for thinking about the characteristics of NTFPs,
drawing a distinction between NTFPs and smallholder agricultural products
(Table 24.1). By bringing medicinal and culinary herbs into cultivation, landowners
are moving these plants away from the authors’ categorization of NTFPs and
towards that of smallholder agricultural products (see the authors’ note in Figure 7
on the nature of a spectrum between these two categories).
According to Belcher and Schreckenberg, bringing NTFPs under cultivation can
introduce novel challenges, both ecological and socioeconomic. They suggest that
“[i]ntensively managed NTFP production systems may completely displace the
natural vegetation within the management unit,” though acknowledge that “the
impacts at the landscape or plot level are less clear for some of the less intensively
managed cases” and that “[i]n most cases ecological and biodiversity impacts have
not been measured or even estimated” (Belcher and Schreckenberg 2007). These
concerns, however, may be less germane to understory medicinal and culinary herb
forest farming systems in Eastern North America. Few forest farming operations are
intense enough to require the removal of competing vegetation. The majority of
small landowners introduce native understory plants into their forestland and simply
allow them to grow—an approach to forest farming referred to as “wild-simulated
production” (Carroll and Apsley 2013). These systems are often mixed and diverse,
both spatially and temporally, containing introduced native plants within an assem-
blage of other native species, producing a range of products and value throughout the
year and over the years. Multistrata agroforestry is often seen as a practice restricted
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 619

Table 24.1 Key differences in the value chains of NTFPs and smallholder agricultural productsa
Factor NTFPs Smallholder agricultural products
Resource Collection areas for wild harvested Fields usually close to or in walking
biology NTFPs often distant from the home distance of home
Resource Low-density production means bulking- Cultivation leads to higher density;
biology up becomes very important usually many producers in one area
Resource Usually “wild” or relatively unimproved Known varieties and availability of
biology leading to problems of inconsistent qual- inputs allow for more uniform
ity, sometimes highly dependent on the production
vagaries of weather
Resource Insecure tenure over collection areas leads Individual tenure, therefore ability
tenure to risk of over-exploitation; inability to to exclude others, provides incen-
manage the resource (to improve quality tive to invest in the resource
and/or quantity)
Resource – Traditional knowledge only, little formal Many staple and minor agricultural
Knowledge research products subject of agricultural
Base research and extension programs
Policy issues Little relevant policy in support of com- Supportive policies in place,
mercialization; usually restricts harvest including credit provision, exten-
and/or transport and sale of NTFPs sion, research
Market “Thin markets“– Often few buyers for the Many different buyers at different
structure total product from a production area scales; producers have more options
for trading
Market Very little available; channeled through Often widely available
information intermediaries
Production Often a supplementary activity, therefore Usually a more consistent part of
volumes production varies as producers choose livelihoods, leading to more pre-
between different livelihood opportunities dictable production volumes
Destination Very diverse, faddish, frequently “lux- Better known markets and more
markets ury” goods and niche markets predictable
Intellectual May be critical for medicinal products Can be an issue with respect to
property and, if active ingredients are synthesized propagation of improved varieties
rights issues away from original source, requiring
negotiation of benefit-sharing agreements
Note: For the purpose of the discussion, the table polarizes the two extremes, although some NTFPs
are not cultivated as smallholder crops, and some smallholder crops are produced for niche markets,
giving them similar characteristics to NTFP value chains
a
Adapted from Belcher and Schreckenberg in Development Policy Review, Issue 25, Volume
3, 2007, page 363

to the tropics, but it is important to recognize that a practice such as the wild-
simulated approach to forest farming described in this chapter can serve as an
example of multistrata agroforestry within Eastern temperate forests. Aside from
herbs, other products that can be integrated into these multistrata systems include
tree syrups, tree bark, firewood, timber, and mushrooms.
Belcher and Schreckenberg identify land tenure, grower organization, and a clear
understanding of the biology and ecology of specific understory species as con-
straints to the successful cultivation of historically wild-harvested species (Belcher
620 K. Sheban

and Schreckenberg 2007). The authors caution that “domestication of NTFPs is

constrained by the need for secure tenure (usually at the individual level) and,
depending on the requirements of the plant, some technical skills and investment
capital” and that “[t]hese conditions can prevent landless and other poor people from
participating” (Belcher and Schreckenberg 2007). Increasing quality and consis-
tency of cultivated products can also disadvantage those who continue to wild
harvest. They also point out that “many NTFPs represent important sources of
‘safety-net’ (a resource that households can turn to in times of need), subsistence
(for households’ own consumption) or “cash” and that increased commercialization
of the resource can lead to overexploitation or shifting control of property rights such
that local people who have historically used the resource are ultimately worse off
(Belcher and Schreckenberg 2007).
These considerations need to be front-of-mind as we transition botanical supply
chains away from depending primarily on the wild-harvest of herbal material from
mixed mesophytic forests and Eastern deciduous forests in general. There is a need
for more research into understory species aside from American ginseng, which is
relatively well-studied. This research should be connected to organizational efforts
such as the Appalachian Beginning Forest Farmers Coalition, as well as University
Extension, local nonprofits, conservation districts, and other Technical Service Pro-
viders, as these organizations provide an effective structure to disseminate informa-
tion and organize forest farmers around the production, verification, and marketing
of their products. For landless Americans, innovative opportunities for forest farm-
ing are being explored through contract growing and leasing arrangements on
private family and corporate-owned lands (for example, on a coal company’s land
in West Virginia) (Tanner Filyaw, personal communication).
The continued access to plant material through permit-based harvesting on public
lands also remains a critical way to ensure equality of access. A more dynamic,
robust, and resilient harvest program on public lands should be integrated into the
overall strategic plan of national forests in the region to support conservation of the
remarkable biodiversity of remaining Appalachian mixed mesophytic forests. Fur-
ther research into and education around novel and sustainable harvest practices will
help address overexploitation. The development of product lines of dried ginseng
leaf represents an opportunity for non-lethal harvest based on research indicating
that ginseng stems and leaves may have higher concentrations of ginsenosides than
the root itself (Kang and Kim 2016; Zhang et al. 2020). Research indicating that the
harvest of ramp leaves is not lethal to the plant and represents a sustainable form of
harvesting that still allows for enjoyment of the resource is similarly promising
(Dion et al. 2016).
Expanding markets and access to plant material on public lands should be
accompanied by stricter regulations to protect plant biodiversity. This requires better
education for law enforcement, adaptive policy that supports forest farmers, and
certification structures that support cultivated plant material in the herbal products
market. Shifting the minimum harvest age from 5 to 10 years for wild ginseng has
been proposed, as well as the implementation of new harvest regulations supporting
overall population growth, prohibiting the harvest of not just the youngest ginseng
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 621

plants, but the most reproductive plants as well, supporting overall population
growth (McGraw et al. 2013). Training law enforcement to recognize the value of
forest plants and understand that their theft is not a trivial matter is also a critical
component of protecting native biodiversity and supporting forest farmers. Ginseng
growers routinely have thousands, even hundreds of thousands of dollars of herbal
material stolen from them, and often no legal recourse to recover their financial loses
(Arnold 2016; Johannsen 2002). Law enforcement can provide important protection
for both wild and cultivated plants but must understand their financial value. At the
policy level, forest farmers could be supported through the creation of a designation
under CITES for the export of “wild-simulated” ginseng. This would allow ginseng
growers in states without an export program to certify their product as cultivated,
while still maintaining the high value of ginseng roots grown under natural
conditions.
It is also crucial that, along with direct investment in private forest farming
systems and botanical markets, we have policies in place that support the overall
viability of diverse farming livelihoods and of forestland ownership. As the number
of farms in the U.S. shrinks, the average age of farmers grows, and the average farm
size increases (MacDonald 2020), we need policies that support small-scale agricul-
tural operations growing a diverse array of products. Programs that allow forestland
owners to gain economic livelihoods from the sustainable management and retention
of forests are critical as well—programs such as The Nature Conservancy’s new
Family Forest Carbon Program, seeking to allow small forestland owners access to
carbon markets. Currently, a very small number of private forestland owners receive
Payments for Ecosystem Services (PES). Only 6% of private forestland owners in
the U.S. have participated in a cost-share program, 2% have conservation easements
on their land, and a similar number receive payments for hunting leases or entrance
fees. Payments for wetland mitigation, representing 38% of all ecosystem services
payments applied to forest ecosystems, were accessed by .00002% of private forest
landowners in the United States (Mercer et al. 2011). Increasing access to PES
programs would allow for more diverse and resilient forest ownership, of which
forest farming could be a part, helping to avoid the overharvest of commercially
valuable understory plants.
Additionally, from a cultural perspective there is a need for a shift toward a
conservation ethic concerning understory plants. For many Americans, the most
significant reference point they may have for American ginseng is the reality
television show Appalachian Outlaws, which attracted nearly three million viewers
in 2014 (Hufford 2014). This show has been criticized for projecting cultural
stereotypes suggesting the harvest of ginseng from the mountainous Appalachians
is a lawless endeavor, where clandestine treachery and the evasion of law enforce-
ment is endemic to local culture (Hufford 2014). As a counterbalance to this negative
imagery, there is a need for a comparable monetary investment in education around
the conservation and sustainable harvest of ginseng and other understory plants, as
well as a reinvigoration of the stewardship practices that have accompanied plant
harvesting for generations. This cultural shift will require continued access to and
engagement with these plants with permit-based harvesting providing an equitable
point of contact through our national forest system.
622 K. Sheban

In the context of the COVID-19 pandemic, this work has taken on increased
urgency. Early research has suggested that markets for nutraceutical products,
including medicinal herbs, have been growing rapidly in response to the crisis as
consumers seek out traditional and alternative forms of medicine and as medical
researchers search for new plant-based compounds to treat COVID infections
(Ayseli et al. 2020; Tahir et al. 2020). Natural products companies found themselves
overwhelmed by orders for alternative medicines and herbal products (see Mountain
Rose Herbs’ temporary suspension of new orders, [Link]
about/new-order-closure).

24.8 Conclusion

With projections for the future of forest cover in the U.S. suggesting it may continue
to decline, and with climate change resulting in large-scale plant species migrations,
the protection of remaining native biodiversity islands has never been more impor-
tant. As the primary source of biodiversity in Eastern deciduous forests, understory
plants should be a focal point for preservation efforts (Spicer et al. 2020). Develop-
ment and implementation of an effective management strategy for American ginseng
could establish precedents for similar programs directed toward other increasingly
valuable and threatened forest understory species. There is a great opportunity to
enlist the public in the protection of these plants through the creation of policies and
incentive structures that support the livelihoods of those who live in and around the
biodiverse Appalachian mixed mesophytic Forests. Permit-based harvesting of
understory plants represents one important way to engage the public in forest
conservation programs. This can only work, however, if harvest on public lands is
strengthened simultaneously with the intentional cultivation, or forest farming, of
valuable medicinal and culinary plants using an approach of “Conservation through
Cultivation” to ensure that plant populations expand rather than dwindle.
Recent years have demonstrated how public health and geopolitical events—
exemplified by COVID-19—can directly impact patterns of natural resource use. If
the uncertainty of the future leads more people to traditional foods and medicines,
and thereby to the plants that support these traditions, we must ensure we have laid
the groundwork to support an equitable and sustainable future for both plants and
people. Protecting and expanding our existing islands of plant biodiversity, as an
investment in the plants themselves, the people that use them, and the generations to
come, is a project worthy of a national effort.
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 623

Appendix: Some Common Species of Appalachian Mixed

Mesophytic Forests

Canopy

Scientific name Common name

Fagus grandifolia American beech
Liriodendron tulipifera Tulip poplar/yellow poplar/tulip tree
Tilia americana American basswood
Acer saccharum Sugar maple
Aesculus flava Yellow buckeye/common buckeye, sweet buckeye
Quercus rubra Red oak
Quercus alba White oak
Tsuga canadensis Eastern hemlock
Betula lenta Black birch
Prunus serotina Black cherry
Magnolia acuminata Cucumber magnolia
Fraxinus americana White ash
Acer rubrum Red maple
Nyssa sylvatica Black gum/sour gum/tupelo
Juglans nigra Black walnut
Carya spp. Hickory species

Midstory

Scientific name Common name

Cornus florida Flowering dogwood
Magnolia spp. Magnolia species
Oxydendrum arboreum Sourwood/sorrel tree
Acer pensylvanicum Striped maple
Cercis canadensis Eastern redbud
Carpinus caroliniana Ironwood/American hornbeam
Ostrya virginiana American Hophornbeam
Ilex opaca American holly
Amelanchier arborea Service berry/shadbush
Lindera benzoin Spicebush
Hamamelis virginiana Witch hazel
Asimina trilobal American pawpaw
Hydrangea arborescens Smooth hydrangea/wild hydrangea
Cornus alternifolia Alternative-leaved dogwood/green osier
Rhododendron maximum American rhododendron/great laurel
(continued)
624 K. Sheban

Scientific name Common name

Viburnum acerifolium Maple-leaved dogwood
Ribes cynosbati Eastern prickly gooseberry
Pyrularia pubera Buffalo nut
Stewartia ovata Mountain camellia
Sambucus canadensis American black elderberry/common elderberry
Euonymus americanus Strawberry bush/bursting-heart
Clethra acuminata Mountain pepper bush
Aralia spinosa Prickly ash/devil’s walking stick

Understory

Scientific name Common name

Trillium grandiflorum White trillium
Trillium erectum Red trillium/wake robin/bethroot
Erythronium americanum Trout lily
Cypripedium calceolus Lady’s slipper orchid
Viola spp. Violet species
Sanguinaria canadensis Bloodroot
Stylophorum diphyllum Celandine poppy/woods poppy
Panax quinquefolius American ginseng
Delphinium tricorne Dwarf larkspur
Hydrophyllum spp. Waterleaf species
Phacelia bipinnatifida Fernleaf phacelia/spotted phacelia
Phlox divaricata Wild blue phlox/wild sweet William
Synandra hispidula Guyandotte beauty
Hydrastis canadensis Goldenseal/yellowroot
Allium triccocum Ramps/wild leeks
Anemone lancifolia Lanceleaf anemone/mountain thimbleweed
Anemone quinquefolia Wood anemone
Anemonella thalictroides Rue anemone
Actaea pachypoda White baneberry/doll’s eyes
Actaea racemosa Black cohosh/black snakeroot
Caulophyllum thalictroides Blue cohosh/squaw root
Claytonia virginica Virginia springbeauty
Claytonia caroliniana Carolina springbeauty
Dicentra canadensis Squirrel corn
Dicentra cucullaria Dutchman’s breeches
Cardamine douglassii Limestone bittercress/purple cress
Tiarella cordifolia Heartleaf foamflower/false miterwort
Dryopteris goldieana Giant wood fern
(continued)
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 625

Scientific name Common name

Phegopteris hexagonoptera Broad beech fern
Adiantum pedatum Northern maidenhair fern
Athyrium pycnocarpos Narrow-leaved-spleenwort/glade fern
Athyrium thelypteroides Silvery glade fern/silvery spleenwort
Osmunda claytoniana Interrupted fern
Symphyotrichum cordifolius Common blue wood aster/heartleaf aster
Eurybia divaricatus White wood aster
Solidago caesia Blue-stemmed/wooden goldenrod
Solidago latifolia Broadleaved/zigzag goldenrod
Eupatorium rugosum White snakeroot/white sanicle/richweed

References

Abrams MD, Johnson SE (2017) The impact of native American activity on vegetation and soil
charcoal in the eastern US. Glob J Archaeol Anthropol 1. [Link]
01.555554
Adams AB, Pontius J, Galford G, Gudex-Cross D (2019) Simulating forest cover change in the
northeastern U.S.: decreasing forest area and increasing fragmentation. Landsc Ecol 34:2401–
2419. [Link]
Ahmad Khan MS, Ahmad I (2019) Herbal medicine: current trends and future prospects. In: Ahmad
Khan MS, Ahmad I, Chattopadhyay D (eds) New look to phytomedicine. Academic, Cam-
bridge, MA, pp 3–13. [Link]
Albuquerque UP, Gonçalves PHS, Ferreira Júnior WS, Chaves LS, da Silva Oliveira RC, da Silva
TLL, dos Santos GC, de Limra Araújo E (2018) Humans as niche constructors: revisiting the
concept of chronic anthropogenic disturbances in ecology. Perspect Ecol Conserv 16:1–11.
[Link]
Alverson WS, Waller DM, Solheim SL (1988) Forests too deer: edge effects in northern Wisconsin.
Conserv Biol 2:348–358
Alwhaibi M, Goyat R, Kelly KM (2017) The use of herbal remedies among mothers of young
children living in the Central Appalachian region. Evid Based Complement Alternat Med.
[Link]
Anderson RC, Anderson MR, Houseman G (2002) Wild American Ginseng. Native Plants J 3:93–
105
Arnold C (2016) As demand for ginseng soars, poachers threaten its survival. Natl Geogr. URL
[Link]
seng-high%2D%2Dpoachers-threaten-its-survival/. Accessed 2 Feb 2020
Ayseli YI, Aytekin N, Buyukkayhan D, Aslan I, Ayseli MT (2020) Food policy, nutrition and
nutraceuticals in the prevention and management of COVID-19: advice for healthcare pro-
fessionals. Trends Food Sci Technol 105:186–199. [Link]
Ban NC, Mills M, Tam J, Hicks CC, Klain S, Stoeckl N, Bottrill MC, Levine J, Pressey RL,
Satterfield T, Chan KM (2013) A social–ecological approach to conservation planning: embed-
ding social considerations. Front Ecol Environ 11:194–202. [Link]
Bannerman JE (1997) Goldenseal in world trade: pressures and potentials. HerbalGram J Am Bot
Counc 41:51
Belcher B, Schreckenberg K (2007) Commercialisation of non-timber forest products: a reality
check. Dev Policy Rev 25:355–377. [Link]
626 K. Sheban

Bierzychudek P (1982) Life histories and demography of shade-tolerant temperate forest herbs: a
review. New Phytol 90:757–776
Braun EL (1916) The physiographic ecology of the Cincinnati region. Ohio State University,
Columbus
Braun EL (1951) Deciduous forests of eastern North America. Blakiston Press, Philadelphia
Burkhart EP (2011) “Conservation through cultivation:” economic, socio-political and ecological
considerations regarding the adoption of ginseng forest farming in Pennsylvania [doctoral
dissertation]. Penn State University
Burkhart EP, Jacobson MG (2009) Transitioning from wild collection to forest cultivation of
indigenous medicinal forest plants in eastern North America is constrained by lack of profit-
ability. Agrofor Syst 76:437–453. [Link]
Carroll C, Apsley D (2013) Growing American ginseng in Ohio: an introduction (Fact Sheet). The
Ohio State University, Columbus
Cavender AP, Beck SH (1995) Generational change, folk medicine, and medical self-care in a rural
Appalachian community. Hum Organ 54:129–142
Chamberlain JL (2005) Conserving the Appalachian medicinal plant industry. In: Proceedings of
the Appalachian opportunities medicinal and aromatic plants conference. Presented at the
Appalachian opportunities medicinal and aromatic plants: producing, using, and marketing
herbs and non-timber forest products, Beckley, West Virginia, p 13
Chittum HK, Burkhart EP, Munsell JF, Kruger SD (2019) A pathway to a sustainable supply of
forest herbs in the eastern United States. HerbalGram J Am Bot Counc 124:60–77
Cruse-Sanders JM, Hamrick JL (2004) Genetic diversity in harvested and protected populations of
wild American ginseng, Panax quinquefolius L. (Araliaceae). Am J Bot 91:540–548. [Link]
org/10.3732/ajb.91.4.540
Davis J, Persons S (2014) Growing and marketing ginseng, goldenseal and other woodland
medicinals, 2nd edn. New Society Publishers. [Link]
Dentali S (2007) Tonnage survey of select North American wild-harvested plants, 2004–2005.
American Herbal Products Association, Silver Spring
Dentali S, Zimmerman M (2012) Tonnage survey of select North American wild-harvested plants,
2006–2010. American Herbal Products Association, Silver Spring
Dharmananda S (2002) The nature of ginseng: traditional use, modern research, and the question of
dosage. HerbalGram J Am Bot Counc 54:34–51
Dion P-P, Bussières J, Lapointe L (2016) Sustainable leaf harvesting and effects of plant density on
wild leek cultivation plots and natural stands in southern Quebec, Canada. Agrofor Syst 90:979–
995. [Link]
Division of Scientific Authority (2018) General advice for the export of wild and wild-simulated
American ginseng (Panax quinquefolius) legally harvested during the 2018 harvest season in
19 states and one tribe with a CITES Export Program for American ginseng. U.S. Department of
the Interior, Fish and Wildlife Service, Marion
Drummond MA, Loveland TR (2010) Land-use pressure and a transition to forest-cover loss in the
eastern United States. Bioscience 60:286–298. [Link]
Duffy DC, Meier AJ (1992) Do Appalachian herbaceous understories ever recover from
clearcutting? Conserv Biol 6:196–201
Filyaw T, Sheban K (2015) Restoring populations of American ginseng, goldenseal, and ramps on
the Wayne National Forest – round two (final grant report to the National Forest Foundation
No. AH-903) Rural Action, Plains, OH, [Link]
Freese C (ed) (1997) Harvesting wild species: implications for biodiversity conservation. Johns
Hopkins University Press, Baltimore. [Link]
Gilliam FS (ed) (2014) The herbaceous layer in forests of eastern North America. Oxford University
Press, Oxford
Gillium F (2007) The ecological significance of the herbaceous layer in forest ecosystems.
Bioscience 57:845–858
Gladstar R, Hirsch P (eds) (2000) Planting the future: saving our medicinal herbs. Healing Arts
Press, Rochester
24 Preserving Biodiversity in Appalachian Mixed Mesophytic Forests. . . 627

Hamilton J (2017) Spreading the ginseng gospel: case study in ginseng production and promotion
from Watauga County Cooperative Extension. In: Leopold S, Ormsby A (eds) Proceedings: the
future of ginseng and forest botanicals. Presented at the future of ginseng and forest botanicals
symposium, Morgantown, West Virginia, pp 109–118
Hill D, Buck L (2000) Forest farming practices. In: Garrett H, Rietveld W, Fisher R (eds) North
American agro-forestry: an integrated science and practice. American Society of Agronomy,
Madison
Holmes MA, Matlack GR (2018) Assembling the forest herb community after abandonment from
agriculture: long-term successional dynamics differ with land-use history. J Ecol 106:2121–
2131. [Link]
Hufford M (2014) Review of Appalachian outlaws. J Appalach Stud 20:234–239. [Link]
10.5406/jappastud.20.2.0234
Jiang B, Ma C, Motley T, Kronenberg F, Kennelly EJ (2011) Phytochemical fingerprinting to
thwart black cohosh adulteration: a 15 Actaea species analysis. Phytochem Anal PCA 22:339–
351. [Link]
Johannsen KL (2002) Root Rustlers. Mother Jones. URL [Link]
politics/2002/07/root-rustlers/. Accessed 23 Apr 2020
Johnston J, Szabo M, Rodney A (2011) Good food, good people: understanding the cultural
repertoire of ethical eating. J Consum Cult 11:293–318. [Link]
1469540511417996
Jouzi Z, Azadi H, Taheri F, Zarafshani K, Gebrehiwot K, Van Passel S, Lebailly P (2017) Organic
farming and small-scale farmers: main opportunities and challenges. Ecol Econ 132:144–154.
[Link]
Kang O-J, Kim J-S (2016) Comparison of ginsenoside contents in different parts of Korean ginseng
(Panax ginseng C.A. Meyer). Prev Nutr Food Sci 21:389–392. [Link]
2016.21.4.389
Light PD (2008) A history of southern and Appalachian folk medicine. J Am Herbalists Guild 8:27–
38
Loucks C, Olson D, Dinerstein E, Weakley A, Noss R, Stritholt J, Wolfe K (n.d.) Appalachian
mixed mesophytic forests | ecoregions | WWF. World Wildl Fund. URL [Link]
[Link]/ecoregions/na0402. Accessed 19 Jan 2020
MacDonald JM (2020) Tracking the consolidation of U.S. agriculture. Appl Econ Perspect Policy
42:361–379. [Link]
McCarthy BC (2014) The herbaceous layer of eastern old-growth deciduous forests. In: Gilliam FS
(ed) The herbaceous layer in forests of eastern North America. Oxford University Press, Oxford
McCoy J-A, Young JH, Nifong JM, Hummer K, DeNoma J, Avendaño-Arrazate CH, Greene SL,
Kantar MB (2019) Species for medicinal and social use with an emphasis on Theobroma cacao
L. (Cacao), Nicotiana tabacum L. (Tobacco), Actaea racemosa L. (Black Cohosh), and Humulus
lupulus L. (Hops). In: Greene SL, Williams KA, Khoury CK, Kantar MB, Marek LF (eds) North
American crop wild relatives, volume 2: important species. Springer, Cham, pp 645–692.
[Link]
McGraw JB, Furedi MA (2005) Deer browsing and population viability of a forest understory plant.
Science 307:920–922
McGraw JB, Lubbers AE, der Voort MV, Mooney EH, Furedi MA, Souther S, Turner JB, Chandler
J (2013) Ecology and conservation of ginseng (Panax quinquefolius) in a changing world. Ann
N Y Acad Sci 1286:62–91. [Link]
Mercer DE, Cooley D, Hamilton K (2011) Taking stock: payments for forest ecosystem services in
the United States. Ecosystem Marketplace. Forest Trends, [Link]
Miller KE, Gorchov DL (2004) The invasive shrub, Lonicera maackii, reduces growth and
fecundity of perennial forest herbs. Oecologia 139:359–375. [Link]
004-1518-2
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity islands. Strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands. Strate-
gies for conservation in human-dominated environments. Series: Topics in Biodiversity and
Conservation, Springer, Cham, pp 1–35
628 K. Sheban

Parsons CM (2016) The natural history of colonial science: Joseph-François Lafitau’s discovery of
ginseng and its afterlives. William Mary Q 73:37–72
Petrzelka P, Ma Z, Malin S (2013) The elephant in the room: absentee landowner issues in
conservation and land management. Land Use Policy 30:157–166. [Link]
landusepol.2012.03.015
Pokladnik RJ (2008) Roots and remedies of ginseng poaching in Central Appalachia. Antioch
University, Keene
Rao MR, Palada MC, Becker BN (2004) Medicinal and aromatic plants in agroforestry systems.
Agrofor Syst 61:107–122. [Link]
Robbins C (1998) American ginseng: the root of North America’s medicinal herb trade. TRAFFIC
52–54. Washington
Robbins C (1999) Medicine from US wildlands: an assessment of native plant species harvested in
the United States for medicinal use and trade and evaluation of the conservation and manage-
ment implications. TRAFFIC North America (traffi[Link])
Rosson JF (2008) Current forest conditions of older stands of the mixed mesophytic forest region on
the Appalachian plateaus province of eastern Kentucky. In: Proceedings of the 16th central
hardwoods forest conference, pp 213–222
Schelhas J (2001) The USA national parks in international perspective: have we learned the wrong
lesson? Environ Conserv 28:300–304. [Link]
Schluter C, Punja ZK (2002) Genetic diversity among natural and cultivated field populations and
seed lots of American ginseng (Panax quinquefolius L.) in Canada. Int J Plant Sci 163:427–439.
[Link]
Sebastiani R, Montagnini F, Dalli D (2013) Ethical consumption and new business models in the
food industry: evidence from the Eataly case. J Bus Ethics 114:473–488. [Link]
1007/s10551-012-1343-1
Sobey DG (1977) The structure and rate of growth of the rhizomes of some forest herbs and dwarf
shrubs of the New Brunswick-Nova Scotia border region. Can Field-Nat 91:377–383
Szczuka D, Nowak A, Zakłos-Szyda M, Kochan E, Szymańska G, Motyl I, Blasiak J (2019)
American ginseng (Panax quinquefolium L.) as a source of bioactive phytochemicals with
pro-health properties. Nutrients 11. [Link]
Tahir AH, Javed MM, Hussain Z (2020) Nutraceuticals and herbal extracts: a ray of hope for
COVID-19 and related infections (review). Int J Funct Nutr 1:1. [Link]
2020.6
Teel WS, Buck LE (1998) From wildcrafting to intentional cultivation: The potential for producing
specialty forest products in agroforestry systems in temperate North America. North American
Conference on Enterprise Development Through Agroforestry: Farming the Agroforest for
Specialty Products. Minneapolis, Minnesota
Turner JB, McGraw JB (2015) Can putative indicator species predict habitat quality for American
ginseng? Ecol Indic 57:110–117
Webb SL, Dwyer M, Kaunzinger CK, Wyckoff PH (2000) The myth of the resilient forest: case
study of the invasive Norway Maple (Acer platanoides). Rhodora 102:332–354
Young JA, King TL, van Manen FT, Eackles MS (2009) Habitat characterization, genetic diversity,
and population abundance of American ginseng. U.S. Geological Survey final report to the
U.S. Fish and Wildlife Service, Division of Scientific Authority, Arlington, Virginia
Young JA, van Manen FT, Thatcher CA (2011) Geographic profiling to assess the risk of rare plant
poaching in natural areas. Environ Manag 48:577–587. [Link]
9687-3
Zhang F, Tang S, Zhao L, Yang X, Yao Y, Hou Z, Xue P (2020) Stem-leaves of Panax as a rich and
sustainable source of less-polar ginsenosides: comparison of ginsenosides from Panax ginseng,
American ginseng and Panax notoginseng prepared by heating and acid treatment. J Ginseng
Res. [Link]
Chapter 25
Farmer Perceptions of Tropical Dry Forest
Restoration Practices on the Azuero
Peninsula of Panama – Implications
for Increasing Biodiversity
in a Human-Dominated Landscape

Vicente Vásquez, Cristina Barber, Yassine Dguidegue, T. Trevor Caughlin,

Roxana García, and Ruth Metzel

Abstract Patches of second-growth forest are biodiversity islands with critical

importance in agricultural landscapes, providing ecosystem services that support
human livelihoods and promote biodiversity, no matter the size. Natural regeneration
presents an opportunity for forest landscape restoration (FLR) at low cost. However,
this is not always the chosen strategy for the smallholder farmers that control about
80% of farms worldwide and are the primary decision-makers for large land areas
with restoration potential. Understanding the perceptions of farmers on restoration
strategies can increase our capacity to implement restoration projects. We surveyed
64 Panamanian land managers in Los Santos province, where 53% of farmers are
smallholders, to better understand how they perceive restoration strategies (refores-
tation, regeneration, and assisted natural regeneration) and their preferences. Partic-
ipants were most confident defining reforestation, associating it with tree planting.
Farmers associated natural regeneration with independently growing vegetation, and
assisted natural regeneration with human intervention and tree planting. Farmers
were polarized in their preference for land clearing, with 36% preferring it as a first
option and 42% preferring it as a last option. Over half of participants ranked letting
vegetation grow and tree planting among their first and second options. High
percentages of tree cover were associated with low preference for land clearing.
Large farm size was associated with higher preference for natural regeneration.
Expanding natural regeneration requires clearly defining and standardizing this

V. Vásquez (*) · Y. Dguidegue

University of the Ozarks, Clarksville, AR, USA
e-mail: vvsqu937@[Link]
C. Barber · T. T. Caughlin
Boise State University, Boise, ID, USA
R. García · R. Metzel
Fundación Pro Eco Azuero, Pedasí, Los Santos, Panamá

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 629
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
630 V. Vásquez et al.

practice. Understanding farmers’ preferences and knowledge of restoration practices

will allow the best FLR strategies to be implemented in each specific case to achieve
desired restoration goals.
Abstract (Spanish) Los parches de bosque secundario son islas de biodiversidad
con importancia crítica para paisajes agrícolas, porque proveen servicios
ecosistémicos que mantienen los medios de vida humanos y promueven la
biodiversidad. La regeneración natural ofrece una oportunidad de restauración del
paisaje forestal (FLR, por sus siglas en inglés) a bajo costo. No obstante, esta no es
siempre la estrategia de restauración preferida por los productores que manejan
fincas de pequeña escala que representan el 80% de las fincas en el mundo. Dichos
productores son los principales actores en la toma de decisiones sobre una
proporción significativa de tierra con potencial para restauración. Entender cómo
los pequeños productores perciben la restauración podría ayudar a mejorar la
eficiencia en la implementación de proyectos de FLR. Realizamos 64 encuestas a
productores panameños en la provincia de Los Santos, donde 53% de los
productores manejan fincas de pequeña escala. El objetivo de estas encuestas fue
comprender cómo perciben diferentes estrategias de restauración (reforestación,
regeneración natural y regeneración natural asistida), y sus preferencias. Los
participantes definieron con más confianza reforestación, que asociaron con plantar
árboles, mientras que asociaron regeneración natural con vegetación que crece por sí
misma, y regeneración natural asistida con intervención humana y plantar árboles.
Los productores tenían opiniones polarizadas respecto a limpiar toda la parcela: el
36% la evaluaron como su estrategia preferida y un 42% como menos preferida. Más
de la mitad de los encuestados clasificaron plantar árboles y dejar que la vegetación
crezca como su primera o segunda opción. Altos porcentajes de cobertura arbórea
fueron asociados a una menor preferencia por limpiar toda la parcela, y parcelas más
grandes fueron asociadas a una mayor preferencia por regeneración natural.
Expandir la regeneración natural como estrategia de restauración depende en gran
medida de alcanzar una definición común de esta práctica. Esto ayudará a integrar
los conocimientos y necesidades de los ganaderos con los objetivos de restauración,
e implementar la mejor estrategia de restauración en cada caso.

Keywords Cattle · Community-based research · Forest landscape restoration

(FLR) · Participatory · Reforestation · Regeneration · Tree planting

25.1 Introduction

Patches of second-growth forest are biodiversity islands with critical importance in

agricultural landscapes. Even small patches of second-growth forest can provide
ecosystem services that support human livelihoods and promote biodiversity (Souza
et al. 2016, Acevedo-Charry and Aide 2019). At the same time, they represent a
low-cost natural climate solution with great potential to sequester carbon (Chazdon
et al. 2016a, b, Griscom et al. 2017).
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 631

Due to the numerous benefits of forest recovery, land managers and policymakers
across the tropics are promoting forest and landscape restoration (FLR) as a way to
increase tree cover in working landscapes while supporting multiple needs of local
stakeholders (Stanturf et al. 2019). Large scale landscape restoration can contribute
to biodiversity conservation of critically endangered species. For example, in Los
Santos province, the Azuero spider monkey (Ateles geoffroyi azuerensis) is known
to need contiguous forest corridors with large width and tall canopy height, which
are in danger if forest restoration practices are not instituted (Schelegle 2011).
The focus of this chapter is on understanding farmer definitions and preferences
on how to restore the tree cover of large contiguous patches on their farm. Under-
standing their perceptions on restoration strategies can increase our capacity to
implement restoration projects, helping to integrate farmers’ knowledge and needs
with forest restoration goals, and matching each specific case to the best restoration
strategy.

25.1.1 Choice of Restoration Strategy: Natural Regeneration,

Tree Planting

A very important decision for forest restoration is whether to pursue strategies that
involve a high level of human intervention, like tree-planting, or to enable forest to
recover with minimal intervention via strategies like natural regeneration (Holl and Aide
2011). Restoration, reforestation, assisted natural regeneration, and natural regeneration
are all terms used often and with wide ranging definitions (Chazdon et al. 2016a, b).
Global, academic, or policy conceptualizations of what it means to restore, reforest, or
regenerate an area can differ from locally used definitions of these concepts, as it will be
discussed in this chapter. In this chapter we generally use the term reforestation, unless
otherwise stated, to mean “re-establishment of forest through planting and/or deliberate
seeding on land classified as forest” as defined by the FAO’s Forest Resources Assess-
ment (FAO 2020). This definition excludes natural regeneration, which is defined as the
“establishment of new forest by self-sown seed, coppice shoots or root suckers”
(Grebner et al. 2013). Assisted natural regeneration is natural regeneration with
human intervention, such as fencing, weed competition removal, enrichment planting,
and protection from fire and other threats (Shono et al. 2007).

25.1.2 Advantages of Natural Regeneration

While tree planting has long served as a default restoration strategy, recent ecolog-
ical research has revealed numerous benefits of natural regeneration, particularly
when regeneration sites are located near existing forest fragments (Griscom 2020).
When compared using meta-analyses, natural regeneration was found to, on average,
632 V. Vásquez et al.

provide greater improvements in ecosystem function than other restoration practices

(Crouzeilles et al. 2017; Jones et al. 2018; Reid et al. 2018). The financial costs of
natural regeneration can also be significantly lower than maintaining nurseries or
buying tree saplings for planting (Elliott et al. 2013). Other researchers have
suggested implementing tree-planting practices only in land areas where natural
regeneration would otherwise be slow (Strassburg et al. 2019).
Despite its potential effectiveness, natural regeneration remains underutilized by
decision-makers, farmers, and restoration practitioners (Chazdon and Uriarte 2016).
Recent research concluded that natural regeneration should be especially adopted in
areas adjacent to riparian zones (Griscom 2020). From a cost-benefit analysis
standpoint, natural regeneration should be the first restoration approach attempted
before resorting to “intensive restoration methods” (Brancalion and Holl 2020; Holl
and Brancalion 2020).
Recent work has examined the political and economic reasons why natural
regeneration is rarely a major component of restoration planning. These reasons
include a lack of economic policy to monetize naturally-regenerating forests, and
agrarian reform laws that penalize landowners for uncultivated land (Chazdon et al.
2020). Explanations for the slow adoption of natural regeneration are largely insti-
tutional. In the context of individual motivations to naturally regenerate vegetation,
the different ways in which farmers in Azuero, Panama protect valuable species for
livelihood purposes on their land has been thoroughly studied (Garen et al. 2011;
Metzel and Montagnini 2014). However, it is important to better understand land
managers’ perceptions and preference of natural regeneration compared to refores-
tation in order to implement effective restoration projects.

25.1.3 Understanding Farmers’ Preferences on Restoration

Strategies

As emphasized in major FAO reports, smallholding farmers own over 80% of farms
worldwide (Schneider 2016); therefore, they are the primary decision-makers for
large portions of land with restoration potential. Key research and policy organiza-
tions have emphasized the indispensable need to include smallholding farmers in
restoration initiatives (FAO 2017). Implementing sustainable restoration practices
requires understanding the different factors that contribute to farmers’ preferences
for restoration strategies. This includes understanding smallholding farmers’ atti-
tudes towards differing forest restoration practices, and examining existing types of
collaboration among farmers, governmental stakeholders, and environmental orga-
nizations (Thacher et al. 1996; Schneider 2016).
Understanding farmers’ preferences on restoration strategies, including percep-
tions of natural regeneration versus reforestation, can improve the capacity to
achieve FLR goals. In some cases, spontaneous tree growth in silvopastoral systems
may already be a central tenet of farm management, which could be leveraged to
support natural regeneration. Farmers maintain trees in pastoral landscapes for
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 633

diverse reasons, including cultural, economic, and agronomic motivations (Garen

et al. 2011). For example, cattle ranchers in the Ecuadorian Amazon promote
recruitment and survival of native tree species in pastures to sustain the viability of
their cattle-focused farming activities (Lerner et al. 2015). Understanding that
farmers maintain tree cover for a variety of reasons, some of which may relate to
natural regeneration, is crucial to customizing FLR in a way that responds to specific
needs of farming communities (Garen et al. 2009). However, in some cases farmers
perceive natural regeneration as “messy” and undesirable and may be more averse to
implementing it in their farmland (Zahawi et al. 2014). Participatory research with
farmers could provide ideas for implementing natural regeneration in ways that
improve aesthetic value, and ultimately longevity, of second-growth forests, provid-
ing more incentives for farmers to use natural regeneration strategies, including
those who may not have originally appreciated them.
We studied farmer preferences for restoration strategies, including reforestation,
natural regeneration, and assisted natural regeneration in a tropical landscape under-
going secondary succession. Our study area, Los Santos province in Panama, is
representative of many Latin American pastoral systems that were once extensively
deforested for cattle ranching and now present opportunities for forest recovery
(Heckadon-Moreno 2009) (Fig. 25.1). In Los Santos, 53% of farms are managed by
smallholders, as defined by the two-hectare classification used in Schneider’s global
study (2016, INEC 2010). Similar to many areas with relatively sparse tree cover,
estimates based on forest production (e.g. Hansen et al. 2013, cited by Fagan 2020)
are likely to provide potentially inaccurate assessments of tree cover change in
Azuero (Caughlin et al. 2016). Although some research showed a regional forest
loss of 220,000 ha from 2001 to 2018, other satellite remote sensing provides some
evidence that tree cover is actually increasing in Los Santos (Metzel 2010; Hansen
et al. 2013; Sloan 2015). Remote sensing detection devices with very high-resolution
imagery provide additional support for regional increases in tree cover, albeit with
high spatial variability (Tarbox et al. 2018; Caughlin et al. 2019). One explanation
for high spatial variability in tree cover change in Los Santos is heterogeneity in
farmer decision-making across a landscape that consists of smallholder farms.
In Los Santos the Panamanian government and NGOs are working to promote
large-scale restoration of land, a majority of which is privately owned. These current
restoration efforts are often centered around reforestation and require purchasing
seedlings and other materials, which limits the scope of restoration practices due to
costs. Farmer preferences are central to understanding potential techniques to increase
forest cover and to successfully implement restoration programs in the region.

25.2 Methodology

25.2.1 Study Site: Azuero Peninsula, Panama

This study was primarily conducted in Los Santos province on the Azuero peninsula
of Panama (Figs. 25.1 and 25.2). Los Santos province forms part of the eastern half
634 V. Vásquez et al.

Fig. 25.1 Pasture and secondary regrowth in the Los Santos Province, Azuero Peninsula, Panama.
(Photo: Cristina Barber)

of the peninsula, which falls within the tropical dry forest biome, defined as “a
vegetation type typically dominated by deciduous trees where at least 50% of trees
present are drought deciduous, the mean annual temperature is > 25
C, total annual
precipitation ranges between 700 and 2,000 mm, and there are three or more dry
months every year (precipitation < 100 mm)” (Sánchez-Azofeifa et al. 2005).
This study region experiences a 5-month dry season from December through
April, and a bimodal rainy season from May through November, with a short dry
period in June (Love and Spaner 2005; Heckadon-Moreno 2009).

25.2.2 Survey of Farm Managers

We developed a survey to better understand how farmers perceive restoration

strategies (reforestation, regeneration, and assisted natural regeneration), their pref-
erences for these strategies for increasing tree cover on the landscape, and the factors
that determine these preferences. The semi-structured, voluntary, anonymous survey
consisted of 19 questions about farmer characteristics, land management practices,
and knowledge and perceptions of regeneration and reforestation. This survey was
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 635

Legend
Panama City
Los Santos Province
Panama border

Locations of survey data collection

Has survey respondent ever reforested?

No
Yes
Percent forest cover (c. 2000)
0
99

Fig. 25.2 Map of survey locations. Symbol coloring represents percentage of farmers at each
location who indicated that they had reforested their land in the past

developed by one of the authors from the Azuero peninsula, and it was revised
through conversations with different landowners and after being used in a pilot
survey group from July 2019–January 2020. We wanted to ensure a high level of
comprehension of the survey by using non-technical terms when appropriate.
The survey was administered to 64 Panamanian “farm managers”, a term which
encompasses farm owners and farm administrators, from Herrera (5) and Los Santos
(59) provinces in February 2020. Survey participants were selected by going door to
door (58 surveys; 1 PM – 8:30 PM) and visiting agricultural supply stores (6 surveys;
9 AM – 1 PM) in sixteen rural communities (Fig. 25.3). Surveys varied in duration
from 10–30 minutes and were conducted orally by one of the authors and a colleague
who was born in the Azuero peninsula, using the Survey123 smartphone application
to record participant answers.
636 V. Vásquez et al.

8%
Proportion of farmers preference
11%
28%
42%
34% 27% 4thchoice
3rdchoice
31%
11%
2ndchoice
11% 37%
33%
16% 1st choice

36%
25% 25% 25%

Parcel Reforestation Natural Assisted

clearing regeneration natural
regeneration

Fig. 25.3 Percentage of farmers ranking parcel clearing, reforestation, natural regeneration (letting
vegetation grow) and assisted natural regeneration (letting vegetation grow with fencing) as their
first (most preferred), second, third or fourth (least preferred) land management strategy

25.2.3 Research Questions

[Link] Farmers’ Understanding of Reforestation and Natural
Regeneration

To gain insight into how farmers understand restoration techniques that increase tree
cover, we asked farmers to define concepts like reforestation, natural regeneration
and assisted natural regeneration. We used the tidytext (Silge and Robinson 2016)
tool to analyze their answers, including tokenizing linked words that repeat across
participant answers, and classifying these into the actions, connotations and alterna-
tive definitions associated with the concept, as well as answers indicating lack of
comprehension or response. The linked words repeated across participants to define
reforestation, natural regeneration and assisted natural regeneration were counted
and converted into percentages.

[Link] Farmers’ Preferences for Reforestation or Natural

Regeneration

To determine farmer preferences of land management options, we asked farmers to

rank their preferences for the following four strategies: (1) clear the whole plot for
cattle, (2) reforest the parcel, (3) let vegetation grow in the parcel (unassisted natural
regeneration), and (4) let the vegetation grow in specific areas with the use of fences
(assisted natural regeneration). The options in the farmers’ native language
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 637

(Spanish) were: (1) “limpiar toda su parcela para ganado”, (2) “reforestar su parcela”,
(3) “dejar que el monte se crezca” and (4) “dejar que el monte se crezca delimitando el
área con cercas”. This phrasing was adopted for the general survey to adapt these
concepts to local terminology (i.e. “monte” is commonly used to refer to vegetation).
In pre-survey consultations, “letting vegetation grow” was the phrase most evocative
of natural regeneration for the farmers consulted, and “letting vegetation grow with
fencing” was most evocative of an assisted natural regeneration practice that excludes
cattle from the system through human labor to accelerate succession.
No questions on past or future land use for the forested land were asked in the
survey, and talking about it was purposefully left to the preference of the survey
participants. Survey participants ranked the land management option strategies from
1 (most preferred) to 4 (least preferred), and we then calculated the percentage of
times that each strategy was mentioned in each preference category.
To identify which economic costs might drive farmers’ decisions, we asked
survey participants to choose from a list of resources which they would prefer to
obtain during a reforestation or natural regeneration program. The possible options
for reforestation were fencing materials, native tree seedlings, volunteer labor, and
there was an option of not answering the question. The possible options for assisted
natural regeneration were fencing materials, volunteers to build the fence, and the
option of not answering the question. Answers were counted and shown in
percentages.

[Link] Factors Driving Farmer Preferences

We also quantified the relative importance of factors driving farmer land manage-
ment preferences. We created four models including the four following management
strategies: parcel clearing, reforestation, letting vegetation grow, and letting vegeta-
tion grow with fencing. The preferences of survey participants were tested against
survey variables (Table 25.1). All variables were treated as fixed effects. We used a
Bayesian modeling framework with Hamiltonian Monte Carlo sampling to analyze
survey answers using the BRMS package (Bürkner 2017), and modeled the answers
using a cumulative regression. This kind of distribution is used for categorical data in
which the order of the categories is meaningful (McElreath 2020).

25.3 Results

25.3.1 Farmers’ Understanding and Definition

of Reforestation, Natural Regeneration and Assisted
Natural Regeneration

The number of farmers confident in defining reforestation was much greater than
those confident in defining natural regeneration and assisted natural regeneration.
The percentage of survey participants that stated not knowing or understanding
638 V. Vásquez et al.

Table 25.1 Variables included in the models to explain survey participants’ preference for
different restoration strategies: parcel clearing, reforestation, natural regeneration, and fenced
natural regeneration
Variable name Type Explanation
Farm size Continuous Farm size
Age Continuous Farmer age
Previous ref Categorical If the farmer has already reforested on their farm (yes/no)
Education Ordered Farmers educational level (primary/ basic education/ high
categorical school/ undergraduate / masters)
Self-reported tree Continuous Percentage of land reserved for trees reported by survey
cover percentage participants.
Future ref Categorical If the farmers would consider future reforestation on their
farm (yes/no)
Ref C Categorical Preferred reforestation inputs (fencing wire/ native tree
seedlings/ volunteers for fencing)
Future NR Categorical If the farmers would consider letting the vegetation to grow
on their farm in the future (yes/no)
Know ref Categorical If the farmer knows someone that has reforested (yes/no)
Notes: Continuous, categorical, and ordered categorical variables used to explain survey partici-
pant’s preference for restoration: parcel clearing, reforestation, natural regeneration, and fenced
natural regeneration. Future Ref stands for reforestation in the future, Ref C stands for the
reforestation input categories, Future NR stands for Natural regeneration in the future, and
Known ref. stands for knowing people that reforest

when asked to define the different land use alternatives was 3% for reforestation,
13% for natural regeneration and 19% for assisted natural regeneration. Of all survey
participants, 77% included the planting of trees and seedlings in their definition of
reforestation. When defining natural regeneration, 59% of survey participants
included trees, other vegetation or forest growing by itself, and 8% described it as
a process without human intervention. There were 8% of survey participants that
included tree-planting in their definition of natural regeneration. Nearly half of
survey participants defined assisted natural regeneration as a process that involves
reforesting and planting trees (48%) and nearly one fifth (17%) defined it as a
process involving human intervention. Fewer than 5% of participants mentioned
letting trees, vegetation and forest grow on their own in their definitions of assisted
natural regeneration. While 17% of survey participants defined reforestation as a
process that happens on land previously deforested, fewer than 5% of participants
defined natural regeneration or assisted natural regeneration as a process that hap-
pens on land previously deforested.
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 639

25.3.2 Farmers’ Preferences for Reforestation or Natural

Regeneration

Farmers had strong opinions about land clearing. Most farmers’ answers were
polarized between choosing parcel clearing as their most preferred (first; 36%) or
least preferred (fourth; 42%) land management option (Fig. 25.3). Letting vegetation
grow (natural regeneration) and reforestation were equally preferred as the top
choice by one fourth of participants. Over half of survey participants (58%) preferred
reforestation as their first or second option, while few (8%) chose it as their least
preferred option. Over half of survey participants (62%) chose to let the vegetation
grow (natural regeneration) as their first or second option, while few (11%) said it
was their least preferred option. Letting vegetation grow with fencing (assisted
natural regeneration) was the least preferred restoration strategy. More than half of
survey participants (59%) chose it as their third or fourth option and fewer partic-
ipants put it as their first or second choice compared to each of the three alternative
land management options (Fig. 25.3).
Some survey participants expressed the reasons they consider reforesting or
letting vegetation grow on part of their land. For reforestation reasons, they men-
tioned watershed protection, timber and fruit for their personal use, and to contribute
to the environment. Natural regeneration reasons included contributing to solving
climate change issues, protecting flora and fauna, watershed protection and allowing
the land to recover.

25.3.3 Farmers’ Preference of Inputs for Reforestation

and Natural Regeneration

Results show that for participating in a reforestation project, 72% of farmers

preferred to receive native tree seedlings, 14% preferred volunteer labor, 8% pre-
ferred fencing materials, and 6% did not answer. When asked about preference for
inputs to support an assisted natural regeneration project, 61% of farmers preferred
to receive volunteer labor, 28% preferred fencing materials, and 11% did not answer.

25.3.4 The Factors Driving Farmer Preferences for Land

Management Strategies

Our analysis of the factors driving survey participant preferences for the four
different management strategies found that a higher tree cover percentage on the
farm and greater farm size increased the probability of ranking parcel clearing in 3rd
or 4th place. The coefficients illustrate the relative effect size of the variables
(i.e. how strong is the effect of each variable compared with the others). A positive
640 V. Vásquez et al.

***
Farm Size
***

Land management strategy

Land clearing
Reforestation
Tree cover Natural regeneration
percentage Assisted natural regeneration
***

Fig. 25.4 Coefficients plot for all land management strategies. The variables that had an effect on
land management strategy preference are on the y axis. We consider that a variable has an effect on
land management preference when the 95% CI does not cross zero. The vertical line marks the zero
value. *** means that the variable has an effect on land management preference. In our preference
rankings, fourth is least preferred and first is most preferred. Therefore, a lower parameter value
(i.e. negative) indicates that the variable (farm size, tree area percentage) has a positive effect on
preferred land management strategy

coefficient indicates more probability of grading the response variable as less

desirable (3rd or 4th) and a negative coefficient indicates more probability of grading
the response variable as more desirable (1st or 2nd). The 95% credibility intervals
(CI) indicates the range of values containing the 95% of the posterior distribution
and provides an uncertainty measure. The positive sign of the effect of self-reported
tree cover percentage (relative effect size ¼ 0.17; 95% CI: 0.03 to 0.40) and farm
size (relative effect size ¼ 1.84; 95% CI: 0.00 to 4.59) indicates that more tree cover
on a farm is associated with less land clearing preference.
Larger farm size increases the probability of ranking natural regeneration in 1st or
2nd place. The negative sign of the effect of farm size (relative effect size ¼ 1.50;
95% CI: 3.65 to 0.00) on natural regeneration indicates that as farm size increases,
natural regeneration is a more attractive option. None of the variables shown in
Table 25.1 seemed to influence farmers’ choice of reforestation or assisted natural
regeneration preference as land management strategy (Fig. 25.4).

25.4 Discussion

25.4.1 Reasons Why Farmers May Prefer Reforestation

to Natural Regeneration

Including smallholder farmers in restoration initiatives requires finding a common

language and conceptualization of restoration strategies. If farmers and institutions
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 641

do not share a common understanding and vocabulary about natural regeneration,

this could impede its use as a landscape restoration strategy. Our results show that
most farmers understand natural regeneration as vegetation growing without human
intervention. If cattle farmers perceive natural regeneration as a restoration strategy
devoid of human action, they are more likely to associate it with abandoned and
unused land (Zahawi et al. 2014). Incorporating a human element to natural regen-
eration, such as fencing a parcel, fire prevention, or monitoring vegetation growth for
scientific, societal or ecosystemic purposes may allow farmers to have a greater
feeling of control and intervention in natural regeneration strategies, while at the
same time saving them the high costs of tree-planting.
The percentage of farmers that appeared unfamiliar with the term natural regen-
eration was significantly higher than expected, given that many farmers (62%) chose
letting vegetation grow as their first or second land management option, and also
considering the recent land use changes in the Azuero Peninsula (Metzel 2010; Sloan
2015; Caughlin et al. 2016). One explanation for this discrepancy is that farmers may
not perceive natural forest recovery on land that is underutilized for agriculture as
part of a restoration strategy. At the regional-scale, agricultural labor scarcity has
been proposed as a primary driver of forest cover increases, as Los Santos farmers
increasingly are employed in off-farm jobs (Sloan 2015). The combination of lack of
preference for natural forest growth relative to tree planting, combined with the labor
needed to put up fences, could explain why assisted natural regeneration ranked
lowest among management strategies in the survey of individual farmers. Despite the
cost-effectiveness of assisted natural regeneration relative to tree planting and its
potential to speed forest recovery relative to unassisted natural regeneration, there
still exists a challenge of implementing this strategy on a widespread scale in
working landscapes, including cattle pastures in Los Santos.
Reforestation may be a concept more widely understood by Azuero farmers
because most of the previous institutional restoration efforts have focused on
reforestation, and more specifically on tree-planting. Decades of forest restoration
programs, such as the Alliance for a Million Hectares (Panama’s Bonn Challenge
commitment), the Smithsonian-Yale Native Species Reforestation Project
(PRORENA, Hall et al. 2011), Yale’s Environmental Leadership and Training
Initiative (ELTI), Fundación Pro Eco Azuero’s ecological corridor restoration pro-
gram, government incentive programs to plant exotic species like teak and other
initiatives by community based organizations have all played a role in informing
farmers about reforestation and the benefits of tree planting.

25.4.2 Potential for Increased Use of Natural Regeneration

Bridging the divide between farmer and institutional definitions and understandings
of natural regeneration and assisted natural regeneration may help to increase
implementation of these strategies. Increasing use of techniques related to natural
regeneration will require facilitating the integration of different strategies in
642 V. Vásquez et al.

landscape restoration projects, as well as implementing donor/NGO success indica-

tors to monitor progress of natural regeneration projects (Chazdon and Uriarte 2016;
Chazdon et al. 2020). Fundación Pro Eco Azuero has implemented strategies like
monitoring vegetative growth together with farmers to emphasize the value of
natural regeneration and decreasing the density of planting to allow regeneration
between planted trees while still retaining some of the benefits of a reforestation
parcel. Other suggestions to promote natural regeneration may include: (1) leverag-
ing natural regeneration at the boundary of existing forest reserves to expand
secondary forest patches, (2) implementing assisted natural regeneration strategies
as part of existing agroforestry and silvopastoral systems, and (3) shifting govern-
ment and donor reporting frameworks away from use of indicators such as number
of trees planted, and toward the total area of land restored.
Expanding the implementation of natural regeneration will also largely depend on
the compatibility of individual farmers’ needs and the benefits of natural regenera-
tion strategies. Several studies have highlighted the uses and values that farmers
attribute to the presence of trees on their parcels (Griscom et al. 2009; Metzel and
Montagnini 2014; Jakovac et al. 2017). This indicates that farmers value restoration
strategies where they are able to select the species and design of vegetation on their
land. For example, farmers might prefer to have fruit trees close to their houses
(Metzel and Montagnini 2014). By contrast, plants growing in a natural regenerating
patch depend highly upon the species pool of neighboring forest patches due to
dispersal limitation, thus restraining farmer species options. Farmers often prefer to
protect rare species on their land (Lenkeek 2003), so young recruits of common
species arriving from nearby patches might be perceived as less valuable. Although
natural regeneration is a low-cost option, its implementation might be underutilized
because of the disconnect between natural regeneration outcomes and farmer
expectations.

25.4.3 Strategic Planning of Restoration Strategies:

Understanding Farmers’ Preferences

Farmers evaluate the advantages and disadvantages of a) investing in labor to clear a

plot for cattle, b) allowing vegetation to grow despite perceptions of abandonment,
or c) opting for reforestation projects that provide a perceived higher degree of
control, as well as tangible economic benefits from timber and fruit (Metzel and
Montagnini 2014; Zahawi et al. 2014; Chazdon et al. 2020).
Understanding why some farmers are willing to restore their land using refores-
tation (tree planting) and/or natural regeneration, while others prefer parcel clearing
as a predominant strategy, will assist strategic planning for where to target restora-
tion efforts. We found that two attributes related to the landscape context of farms
had strong impacts on management strategies: farm size and tree cover percentage.
Parcel clearing, in particular, was affected most strongly by farm size and tree cover
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 643

percentage. In contrast, the education level and age of survey participants had
minimal impact on land management preferences.
Our finding that higher self-reported farm tree cover percentage is associated with
a lower preference for land clearing has implications for land cover dynamics in Los
Santos. This suggests that maps of farm tree cover and size could be used to predict
willingness to forego land clearing and to contribute to strategic planning for
restoration activities. Relationships between tree cover and farmer preferences are
likely to change the rate of forest transformation, including rapid transitions between
treeless and silvopastoral management techniques (Valencia Mestre et al. 2019). An
increased willingness to support tree-intensive land management strategies when
forest is present represents an example of a positive feedback loop that could
promote regional increases in tree cover.

25.5 Conclusion

In a human dominated landscape such as the Azuero peninsula, Forest Landscape

Restoration (FLR) strategies must align with the human preferences for land uses. In
this chapter, we highlighted farmer definitions and preferences for restoration strat-
egies like reforestation, natural regeneration and assisted natural regeneration.
Smallholder farmers are crucial to the protection of the Azuero peninsula because
they own a majority of farmland, which is why their participation and knowledge on
restoration is indispensable to improve FLR capacity.
Farmers were more confident in defining reforestation, which they associated
with tree planting, and were less confident in defining natural regeneration and
assisted natural regeneration. They associated assisted natural regeneration with
definitions of reforestation (human intervention and tree planting) more than with
definitions of natural regeneration. Participants were more likely to define refores-
tation as a process occurring on previously deforested land.
Our findings indicate that the preferred management strategy among farmers was
to clear the land for cattle. However, a similar proportion of farmers would choose
clearing land for cattle as their least preferred option. This polarization may indicate
that many farmers are looking for alternatives to treeless pasture systems. Over half
of farmers chose natural regeneration and reforestation as their first and second
restoration options, but assisted forest regeneration was the least preferred option.
Assisted natural regeneration may be the least preferred option because the cost of
labor or materials to fence off parcels is too high for a practice with seemingly
limited economic benefits, especially given the exclusion of cattle from the parcel
through fencing.
Farmers may prefer natural regeneration because letting vegetation grow is
perceived as a land management strategy for unused land that involves a low
investment of time and resources. While farmers associate both reforestation and
assisted natural regeneration with human intervention, they may prefer to reforest
because of a greater familiarity with the methods, as many restoration programs
644 V. Vásquez et al.

established on the peninsula to date have focused on tree planting, or because they
can choose which species to plant, especially beneficial rare species that would not
grow in the parcel otherwise. To make natural regeneration a viable option will
require creating a broader understanding of restoration approaches among farmers in
order to fine tune their implementation within each land area based on their benefits
and obstacles to farmers.
Through the results of this research we gained insights into farmer’s understand-
ing, preferences, and drivers regarding restoration strategies to bridge their local
knowledge and practices with academic discussions and debates around restoration
alternatives. The results can be useful in the design and implementation of programs
geared to accelerate forest landscape restoration at the global scale, which is needed
to achieve the success of ambitious global agendas such as the Bonn Challenge, and
the Sustainable Development Goals: climate action (SDG 13), and the restoration
and sustainable use of terrestrial ecosystems (SDG 15).

Acknowledgments The authors would like to thank Oscar Vega of the ESRI Panama office for
GIS assistance in creating the field surveys, and Sandra Vásquez and Jesus Batista for field
implementation support.

References

Acevedo-Charry O, Aide TM (2019) Recovery of amphibian, reptile, bird and mammal diversity
during secondary forest succession in the tropics. Oikos 128:1065–1078
Brancalion PHS, Holl KD (2020) Guidance for successful tree planting initiatives. J Appl Ecol.
[Link]
Bürkner PC (2017) Brms: an R package for Bayesian multilevel models using Stan. J Stat Softw
80(1):1–28
Caughlin TT, Rifai SW, Graves SJ, Asner GP, Bohlman SA (2016) Integrating LiDAR-derived tree
height and Landsat satellite reflectance to estimate forest regrowth in a tropical agricultural
landscape. Remote Sens Ecol Conserv 2:190–203
Caughlin TT, Graves SJ, Asner GP, Tarbox BC, Bohlman SA (2019) High-resolution remote
sensing data as a boundary object to facilitate interdisciplinary collaboration. In: Perz SG
(ed) Collaboration across boundaries for social-ecological systems science: experiences around
the world. Springer International Publishing, Cham, pp 295–326
Chazdon RL, Uriarte M (2016) Natural regeneration in the context of large-scale forest and
landscape restoration in the tropics. Biotropica 48:709–715
Chazdon RL, Broadbent EN, Rozendaal DMA, Bongers F, Zambrano AMA, Aide TM,
Balvanera P, Becknell JM, Boukili V, Brancalion PHS, Craven D, Almeida-Cortez JS, Cabral
GAL, de Jong B, Denslow JS, Dent DH, DeWalt SJ, Dupuy JM, Durán SM, Espírito-Santo
MM, Fandino MC, César RG, Hall JS, Hernández-Stefanoni JL, Jakovac CC, Junqueira AB,
Kennard D, Letcher SG, Lohbeck M, Martínez-Ramos M, Massoca P, Meave JA, Mesquita R,
Mora F, Muñoz R, Muscarella R, Nunes YRF, Ochoa-Gaona S, Orihuela-Belmonte E, Peña-
Claros M, Pérez-García EA, Piotto D, Powers JS, Rodríguez-Velazquez J, Romero-Pérez IE,
Ruíz J, Saldarriaga JG, Sanchez-Azofeifa A, Schwartz NB, Steininger MK, Swenson NG,
Uriarte M, van Breugel M, van der Wal H, Veloso MDM, Vester H, Vieira ICG, Bentos TV,
Williamson GB, Poorter L (2016a) Carbon sequestration potential of second-growth forest
regeneration in the Latin American tropics. Sci Adv 2(5):e1501639
25 Farmer Perceptions of Tropical Dry Forest Restoration Practices on. . . 645

Chazdon RL, Brancalion PHS, Laestadius L, Bennett-Curry A, Buckingham K, Kumar C, Moll-

Rocek J, Guimarães Vieira IC, Wilson SJ (2016b) When is a forest a forest? Forest concepts and
definitions in the era of forest and landscape restoration. Ambio:538–550
Chazdon RL, Lindenmayer DB, Guariguata M, Crouzeilles R, Benayas JMR, Chavero EL (2020)
Fostering natural forest regeneration on former agricultural land through economic and policy
interventions. Environ Res Lett 15(4):043002
Crouzeilles R, Ferreira MS, Chazdon RL, Lindenmayer DB, Sansevero JBB, Monteiro L,
Iribarrem A, Latawiec AE, Strassburg BBN (2017) Ecological restoration success is higher
for natural regeneration than for active restoration in tropical forests. Sci Adv 3:e1701345
de Souza SEXF, Vidal E, Chagas Gde F, Elgar AT, Brancalion PHS (2016) Ecological outcomes
and livelihood benefits of community-managed agroforests and second growth forests in
Southeast Brazil. Biotropica 48:868–881
Elliott S, Blakesley D, Hardwick K, Plukham S, Saengkam D, Agbor JN (2013) Restoring tropical
forests: a practical guide. Royal Botanic Gardens, Kew
Fagan ME (2020) A lesson unlearned? Underestimating tree cover in drylands biases global
restoration maps. Glob Chang Biol 26(9):4679–4690
FAO (2017) Smallholder Forest Producer Organizations in a Changing Climate, Rome. [Link]
[Link]/3/[Link]
FAO (2020) Forest Resources Assessment Terms and Definitions. [Link]
[Link]
Garen EJ, Saltonstall K, Slusser JL, Mathias S, Ashton MS, Hall JS (2009) An evaluation of
farmers’ experiences planting native trees in rural Panama: implications for reforestation with
native species in agricultural landscapes. Agr Syst 76(1):219–236
Garen EJ, Saltonstall K, Ashton MS, Slusser JL, Mathias S, Hall JS (2011) The tree planting and
protecting culture of cattle ranchers and small-scale agriculturalists in rural Panama: opportu-
nities for reforestation and land restoration. For Ecol Manage 261(10):1684–1695
Grebner DL, Bettinger P, Sirey JP (2013) Chapter 11 – common forestry practices. In: Grebner DL,
Bettinger P, Sirey JP (eds) Introduction to forestry and natural resources. Academic, Cambridge,
pp 255–285
Griscom HP (2020) The long-term effects of active management and landscape characteristics on
carbon accumulation and diversity within a seasonal dry tropical ecosystem. For Ecol Manage
473:118296
Griscom HP, Griscom BW, Ashton MS (2009) Forest regeneration from pasture in the dry tropics of
Panama: effects of cattle, exotic grass, and forested Riparia. Restor Ecol 17:117–126
Griscom BW, Adams J, Ellis PW, Houghton RA, Lomax G, Miteva DA, Schlesinger WH, Shoch D,
Siikamäki JV, Smith P, Woodbury P, Zganjar C, Blackman A, Campari J, Conant RT,
Delgado C, Elias P, Gopalakrishna T, Hamsik MR, Herrero M, Kiesecker J, Landis E,
Laestadius L, Leavitt SM, Minnemeyer S, Polasky S, Potapov P, Putz FE, Sanderman J,
Silvius M, Wollenberg E, Fargione J (2017) Natural climate solutions. Proc Natl Acad Sci
114(44):11645–11650
Hall JS, Love BE, Garen EJ, Slusser JL, Saltonstall K, Mathias S, van Breugel M, Ibarra D, Bork
EW, Spaner D, Wishnie MH (2011) Tree plantations on farms: evaluating growth and potential
for success. For Ecol Manage 261(10):1675–1683
Hansen MC, Potapov PV, Moore R, Hancher M, Turubanova SA, Tyukavina A, Thau D, Stehman
SV, Goetz SJ, Loveland TR, Kommareddy A, Egorov A, Chini L, Justice CO, Townshend JRG
(2013) High-resolution global maps of 21st-century forest cover change. Science 342(6160):
850–853
Heckadon-Moreno S (2009) De Selvas a Potreros: La Colonización Santeña en Panamá:
1850–1980. ExedraBooks, Panamá
Holl KD, Aide TM (2011) When and where to actively restore ecosystems? For Ecol Manage 261:
1558–1563
Holl KD, Brancalion PHS (2020) Tree planting is not a simple solution. Science 368(6491):
580–581
646 V. Vásquez et al.

Instituto Nacional de Estadística y Censo (2011) VII Censo Nacional Agropecuario. Volumen II:
Características de los Productores y las Explotaciones Agrarias. Cuadro 1. Productores
agropecuarios en la República, por grupos de edad, empresas y organizaciones comunales,
según provincia, comarca indígena y tamaño de la explotación: 24 de abril de 2011, Panamá
Jakovac CC, Peña-Claros M, Kuyper TW, Bongers F (2017) Loss of secondary-forest resilience by
land-use intensification in the Amazon. J Ecol:67–77
Jones HP, Jones PC, Barbier EB, Blackburn RC, Benayas JMR, Holl KD, McCrackin M, Meli P,
Montoya D, Mateos DM (2018) Restoration and repair of Earth’s damaged ecosystems. Proc R
Soc B 285:2017–2577
Lenkeek AG (2003) Diversity makes a difference: farmers managing inter- and intra-specific tree
species diversity in Meru Kenya. PhD Diss, Wageningen University, Wageningen
Lerner AM, Rudel TK, Schneider LC, McGroddy M, Burbano DV, Mena CF (2015) The sponta-
neous emergence of silvo-pastoral landscapes in the Ecuadorian Amazon: patterns and pro-
cesses. Regional Environ Change 15(7):1421–1431
Love B, Spaner D (2005) A survey of small-scale farmers using trees in pastures in Herrera
Province, Panama. J Sustain For 20(3):37–65
McElreath R (2020) Statistical rethinking: a Bayesian course with examples in R and Stan. CRC
Press, Boca Raton
Metzel RNB (2010) From “Finca” to forest: forest cover change and land management in Los
Santos, Panama. Thesis, Princeton University, Princeton, New Jersey
Metzel R, Montagnini F (2014) From farm to forest: factors associated with protecting and planting
trees in a Panamanian agricultural landscape. Bois Et Forets Des Tropiques 322:3–15
Reid JL, Fagan ME, Zahawi RA (2018) Positive site selection bias in meta-analyses comparing
natural regeneration to active forest restoration. Sci Adv 4:eaas9143
Sánchez-Azofeifa GA, Quesada M, Rodriguez JP, Nassar JM, Stoner KE, Castillo A, Garvin T,
Zent EL, Calvo-Alvarado JC, Kalacska MER, Fajardo L, Gamon JA, Cuevas-Reyes P (2005)
Research priorities for neotropical dry forests. Biotropica 37:477–485
Schelegle S (2011) Connections in a fragmented landscape: Azuero spider monkey corridors in the
Azuero Peninsula, Panama. Thesis, Princeton University, Princeton, New Jersey
Schneider S (2016) Family farming in Latin America and the Caribbean: looking for new paths of
rural development and food security. Pap 137, FAO, [Link]
Shono K, Cadaweng E, Durst P (2007) Application of assisted natural regeneration to restore
degraded tropical forestlands. Restor Ecol 15:620–626
Silge J, Robinson D (2016) Tidytext: text mining and analysis using tidy data principles in R. JOSS
1(3). [Link]
Sloan S (2015) The development-driven forest transition and its utility for REDD+. Ecol Econ 116:
1–11
Stanturf JA, Kleine M, Mansourian S, Parrotta J, Madsen P, Kant P, Burns J, Bolte A (2019)
Implementing forest landscape restoration under the Bonn challenge: a systematic approach.
Annals For Sci 76:50
Strassburg BBN, Beyer HL, Crouzeilles R, Iribarrem A, Barros F, de Siqueira MF, Sanchez-
Tapia A, Balmford A, Sansevero JBB, Brancalion PHS, Broadbent EN, Chazdon RL, Filho
AO, Gardner TA, Gordon A, Latawiec A, Loyola R, Metzger JP, Mills M, Possingham HP,
Rodrigues RR, Scaramuzza CAM, Scarano FR, Tambosi L, Uriarte M (2019) Strategic
approaches to restoring ecosystems can triple conservation gains and halve costs. Nature Ecol
Evol 3:62
Tarbox BC, Fiestas C, Caughlin TT (2018) Divergent rates of change between tree cover types in a
tropical pastoral region. Landsc Ecol 33:2153–2167
Thacher T, Lee DR, Schelhas JW (1996) Farmer participation in reforestation incentive programs in
Costa Rica. Agr Syst 35(3):269–289
Valencia Mestre MC, Ferguson BG, Vandermeer J (2019) Syndromes of production and tree-cover
dynamics of Neotropical grazing land. Agroecol Sustain Food Syst 43(4):362–385
Zahawi RA, Reid JL, Holl KD (2014) Hidden costs of passive restoration. Restor Ecol 22:284–287
Chapter 26
Safeguarding Biodiversity Islands
in Northern Ethiopia Amidst Political
Change

Emily Sigman

Abstract This chapter concerns the safeguarding of biodiversity islands established

through community-based restoration in Northern Ethiopia, and centers particularly
on the case study of a kebele (or village, also spelled q’ebele) called Abreha we
Atsbeha. Severely degraded landscapes in the Tigray region of Northern Ethiopia
have enjoyed extensive restoration efforts in recent decades, and these efforts have
intensified further since Ethiopia joined the African Resilient Landscapes Initiative
(ARLI) and became a member of the Bonn Challenge. The resultant restored areas
serve as biodiversity islands in an otherwise highly degraded area. While praise
abounds for the restoration successes and benefits to communities like Abreha we
Atsbeha, there is scant research exploring how the restoration takes place, and
insufficient attention has been paid to the capacity of these restoration efforts—
and the biodiversity islands they enable—to survive in the midst of political change.
This chapter therefore investigates the communal labor activities that enliven suc-
cessful ecological restoration and relies on a political ecology framework to discuss
governance and restoration in this context. Overall, the chapter demonstrates that
restoration-based biodiversity islands in Abreha we Atsbeha are created and
safeguarded through a robust single-party political system. This system at once
mandates universal participation in restoration activities while limiting the partici-
pation of many community constituents in the planning of restoration activities. The
author argues that this social-ecological system needs to be adapted to ensure that
rural, restoration-based biodiversity islands in the region can continue to flourish
alongside more pluralistic and democratic political norms and institutions.

Keywords Communal labor · Dryland · Political ecology · Restoration · Social-

ecological systems · Watershed

E. Sigman (*)
Yale School of Forestry and Environmental Studies, Yale Jackson Institute for Global Affairs,
New Haven, CT, USA
e-mail: [Link]@[Link]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 647
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
648 E. Sigman

26.1 Introduction

The term ‘biodiversity islands’ tends to conjure up a certain image, one of pristine
ecosystems kept away from the destructive hand of human civilization. High levels
of degradation and destruction wrought on the planet—careless disregard for pre-
cious soils, rapacious plundering of the world’s forests, flagrant pollution of life-
giving air and water systems—do much to cement this idea. Environmental degra-
dation is a powerful driver, motivating conservationists to do all they can to
safeguard what precious few biodiversity islands have been spared from the clutches
of human shortsightedness.
Yet environmental degradation also motivates a different, concurrent response:
the push to restore. Increased awareness of humanity’s mutual dependency on intact,
connected, and diverse ecosystems has arisen in tandem with improved understand-
ings of the role that human beings have played in the stewardship of magnificent
ecosystems, from the Amazon rainforest (Levis et al. 2017) to the Great Plains of
North America (Krech 2000). Moreover, we as a global community are beginning to
appreciate and acknowledge the role that we can play in reversing the tides of
environmental degradation. We are increasingly aware of our collective capacity
to usher in a new era of flourishing. Harnessing the power of restoration, we can do
more than simply protect biodiversity islands: we can also create them.
In rural areas across Ethiopia, especially in the Northern province of Tigray,
many local communities have already long been exercising this capability. This
chapter explores biodiversity islands in the context of one community in the Tigray
Region of Northern Ethiopia, called Abreha we Atsbeha. There, rural farmers have
successfully implemented impressive soil and water conservation techniques and
pioneered new social norms that have effectively restored watersheds, catalyzed
natural plant regeneration, and over time, provisioned biodiversity islands in land-
scapes that were otherwise so severely degraded that they were deemed
uninhabitable by humans. These novel socio-ecosystems—which exist only because
of extensive and direct human mediation—pose an interesting set of questions for
those seeking strategies to safeguard biodiversity islands.
The following sections will synthesize recent developments in socio-ecological
restoration, parse out the case study, and offer commentary on the often-
understudied attributes of these kinds of restored landscapes, in particular local
labor and political dynamics. The chapter swivels around the concept of a Restora-
tion Dilemma, described in 2.3, with the goal of assuaging the tensions that arise as
restoration ambitions and initiatives progress in scale. Overall, this chapter aims to
highlight the active role that local communities can, do, and must play in creating
biodiversity islands. It imagines a future where the fruits of restoration labor can be
effectively safeguarded—and equitably savored—by all.
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 649

26.2 Restoration, Local Actors, and Biodiversity Islands

26.2.1 Global Impetus for Restoration

Recent decades have witnessed an increasing global awareness of the need to restore
landscapes on a mass scale. Deforestation and degradation have advanced so
thoroughly that it is no longer enough to conserve what vestiges of intact forest
remain; the task of safeguarding the world’s ecosystems must also be fulfilled by
efforts to actively restore what’s been destroyed.
Globally, more than 2 billion hectares of deforested and degraded lands hold
potential for restoration (Minnemeyer et al. 2014). The benefits of restoration are
multitudinous, and are often encapsulated as ‘ecosystem services’ in the field’s
contemporary literature (Daily 1997; Costanza et al. 2014). Increased biodiversity
ranks chiefly among the ecosystem services sought by restoration advocates (Mace
et al. 2012). In foundational island biogeography theory (MacArthur and Wilson
1967), biodiversity islands—which in these early studies referred to physical islands
surrounded by water—comprise areas in a landscape with biodiversity measures
significantly higher than those of the surrounding landscape. Extending this concept,
restored areas generally possess higher biodiversity metrics relative to their sur-
rounding degraded environments, that is, restored areas serve as biodiversity islands
in and of themselves in the context of degraded landscapes (Benayas et al. 2009).
Biodiversity gains in these spaces can take the form of increased plant, wildlife,
insect, and soil diversity—among many other measures—and many of these gains
are understood to be of both direct and indirect human benefit (Aerts and Honnay
2011).
‘Restored’ biodiversity islands geared primarily towards human use
(i.e. agroforests, timberlands, foraging areas, or other such zones intended to be
regularly utilized) can likewise enhance other, most restrictive biodiversity islands
(such as national parks or pristine zones). These areas can, for example, provision
connective corridors between conservation areas, serve as reservoirs for pollinators
and dispersers, contribute to increased genetic diversity needed to support healthy
species reproduction, and help stabilize large-scale soil and water dynamics, which
improve mosaic landscape function as a whole (Boesing et al. 2018). Moreover, with
careful planning and proper governance, these areas can alleviate pressure to extract
from protected zones, serving as sources of food, fuel, timber, fibers and other
materials while acting as areas of intermediate biodiversity between degraded
landscapes and zones of extreme conservation importance (Schroth et al. 2004;
Kumar 2010).
A number of restoration initiatives have emerged from this understanding, and
chief among them is the Bonn Challenge. The Bonn Challenge is a global, multi-
stakeholder effort that was launched in 2011 with the goal of restoring 150 million
hectares of degraded and deforested lands by the year 2020 and 350 million hectares
650 E. Sigman

by 2030.1 It offers institutional, policy, and technical support to developing coun-

tries, which in turn have pledged to restore a certain number of hectares. The
challenge promotes regional collaboration platforms, such as the African Resilient
Landscape Initiative (ARLI) (World Bank 2015). ARLI, in turn, supports initiatives
like the African Forest Landscape Restoration Initiative (AFR100), a country-led
effort to bring 100 million hectares of land in Africa into restoration by 2030.2
Twenty-nine countries have so far made a commitment of more than 125 million
hectares to the initiative, with Ethiopia pledging 15 million hectares, by far the
region’s largest single contribution. In exchange, development banks have contrib-
uted more than US$1 billion, with an additional $481 million pledged by the private
sector (Anderson and Piembert 2019).

26.2.2 Increasing Focus on Local-Level Community

Engagement

Early attempts in the last decades to formalize restoration studies, projects, and
initiatives worldwide tended to focus primarily on biophysical dynamics, without
incorporating—or even acknowledging—the complex political, economic, and
social forces at play in restoration projects (Brudvig 2011). More recent years
have witnessed an outpouring of literature seeking to correct the course and expand
the restoration lens to better capture and understand the intricacies of multipart
cultural-environmental landscapes (Rovere 2015).
As ecologists have progressed in their understanding of forest stand dynamics—
no longer viewing forest succession as advancing towards a ‘climax’ state, but rather
existing in a process of perpetual evolution—so too have restoration scholars relaxed
the tendency to view restoration as a reversion ‘back’ to some previous ecosystem
(Trigger et al. 2008). Such a view allows ‘restoration’ to focus on key attributes of
ecosystems—such as soil dynamics, watershed health, and biodiversity measures—
while permitting and even encouraging the emergence of novel socio-ecosystems
(Hobbs et al. 2009). At the same time, the intersecting fields of anthropology,
environmental history, and political ecology have substantially rewritten the con-
servation script in recent years, convincingly illuminating the key role that human
societies have historically played in the stewardship and even creation of ecosystems
once considered unmediated or primary forests (Fernandez-Manjarres et al. 2018).
These streams of knowledge have started to merge, each contributing new
dimensions to a growing, interdisciplinary inquiry that seeks to align the develop-
ment of human society with environmental conservation and ecosystem regenera-
tion. A Social-Ecological Systems (SoESs) framework was first proposed by Berkes
and Folke (1998) to analyze resilience in local resource management systems; since

1
Bonn Challenge. In: Bonn Challenge. [Link] Accessed 25 Jan 2020
2
AFR100 In: [Link]. [Link] Accessed 25 Jan 2020
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 651

then, the term has come to refer to a number of frameworks that expressly recognize
the complex linkages between human and ‘natural’ systems (Martin 2017), espe-
cially in restoration and resilience contexts (López et al. 2017). Socio-ecological
restoration, in turn, has emerged at the confluence of these schools of thought, as a
practical discipline working to address interconnected social and environmental
challenges (Colding and Barthel 2019).
This understanding—while stressing the need to restore ecosystems on a land-
scape-scale—has emphasized the importance of local community dynamics. This
emphasis recognizes that local communities are often the direct beneficiaries and/or
agents of restoration (Brancalion et al. 2014). It highlights the value of local
ecological knowledge (Uprety et al. 2012), and promotes community participation
(Maynard 2013), improved understanding of local power dynamics (Habtezion et al.
2015), long-term community monitoring and evaluation (Wortley et al. 2013), and
an intentional awareness and anticipation of potential unintended consequences and
pernicious outcomes (Daily and Maston 2008).
Scholars, public agencies, and non-government institutions have increasingly
adopted the perspectives and practices advanced by the social-ecological restoration
agenda, some of which now operates under the global framework promulgated by
the UN Sustainable Development Goals (SDGs). This self-professed ‘blueprint to
achieve a better and more sustainable future for all’ unambiguously understands the
interconnected nature of the world’s most pressing social and environmental prob-
lems (United Nations 2020). Many of the targets and indicators are expressly
concerned with community integration and engagement. Despite this improved
understanding, effective community-scale work remains difficult in practice, and
coordination among actors across different scales has likewise proven challenging.

26.2.3 The ‘Restoration Dilemma’

On the whole, literature from socio-ecological restoration tends to suggest that the
most effective and durable restoration projects are idiosyncratic, take place at a local
level, are culturally and ecologically site-specific, and therefore tend to resist scaling.
Yet irrefutable evidence from studies concerned with climate change, mass extinc-
tion and biodiversity loss, and reliable resource provisioning—bolstered by the
imperatives of the international development agenda—clearly articulates the need
for restoration at a global scale.
Restoration practitioners therefore face a profound dilemma. A dilemma is
defined as ‘a problem offering two possibilities, neither of which is unambiguously
acceptable or preferable’(Garner 2009). Focusing on global-scale restoration often
has pernicious outcomes at the local scale (which is not acceptable). Focusing on
local-scale restoration often stymies practitioners attempting to transform “a thou-
sand random acts of restoration” into a “coherent strategy” that can realistically meet
global needs (Covelli-Metcalf et al. 2015; Budiharta et al. 2016) (which is not
preferable).
652 E. Sigman

Escaping through the horns of this dilemma requires careful, consistent and
adaptive feedback about how national, regional, and global restoration directives
reverberate at the local level and vice versa. A firm grasp on how restoration actually
happens—the human as well as the non-human agencies involved—offers key
insights into project dynamics and vulnerabilities, and in so doing, improves pros-
pects for multi-scalar success.

26.3 Restoration in Ethiopia – The Case of Abreha We

Atsbeha

26.3.1 The Case Study in Context

Ethiopia offers fertile ground for those seeking to explore the kinds of biodiversity
islands that emerge from restoration, and in particular, for studies examining how to
safeguard such biodiversity islands in the midst of political change. Ethiopia today
comprises Africa’s second most populous country3—the most populated landlocked
country to exist anywhere in the history of the planet—and by many accounts is
home to the fastest growing economy in the world. Yet this growth occurs in the
context of a largely rural society, comprised of a roughly 80% agrarian population.4
Despite the fact that Ethiopia is essentially governed by a one-party state with a
planned economy, the country’s vast population is spread out over more than a
million square kilometers and divided into nine politically autonomous ethnic states,
each possessing unique socio-ecological features and agronomic patterns. The
country as a whole is thus best conceived as a quilted patchwork stitched of an
extraordinary diversity of landscapes, languages, and life-ways.
Restoration and conservation in this context are therefore highly suited to an
ethno-regional focus. The country often garners substantial coverage in public media
outlets and has earned a reputation among international agencies for its eye-catching
national restoration activities, such as, for example, the recent planting of 350 million
trees across the country over a period of 12 hours (UNEP 2019). Yet the attention
paid to country-level restoration blitzes such as these tends to obfuscate an under-
standing of the heterogeneity of long-term restoration activities at the federation- and
even regional-level. It is at these smaller scales where connections between restora-
tion, livelihoods, and biodiversity are made apparent, and where prospects for
safeguarding biodiversity islands are most accessible. This study is therefore

3
The population of Ethiopia is roughly 112,000,000. The most populous country in Africa is
Nigeria, with a population of roughly 201,000,000. From Ethiopia Overview. In: The
World Bank: Where we Work. (2019) [Link]
Accessed 25 Jan 2020
4
Rural population (% of total population) – Ethiopia. In: The World Bank Data Indicators 2018
revision. [Link] Accessed
25 Jan 2020
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 653

restricted to the northern Tigray region, and focuses on the dynamics of one kebele
(or community, village, also spelled ‘q’ebele’) within this context. The case study
helps concretize the ‘Restoration Dilemma’ outlined in Sect. 26.2.3, and draws out
the multi-scalar dynamics that bind participatory restoration, biodiversity provision-
ing, and socio-political change.

26.3.2 Physical, Ecological, and Social Characteristics

of Tigray

The wrinkles of humanity are etched in the broken, volcanic plateaus of Tigray. The
heart of the ancient Aksumite kingdom, one of the cradles of modern civilization,
once pumped through this region, drained by the Tekeze and Gash rivers which ran
to the Red Sea, along with the Upper Nile. This made of the empire a vast marine
trading power unlike the territorial constraints that today render the whole of
Ethiopia landlocked (McKenna 2019). The region has remained dominated by an
Orthodox Christian population (95.6% in 2007 – IHSN 2019), which has a
longstanding history as a seat of political power for the country as a whole, though
Ethiopia’s capital is in Addis Ababa, located some thousand kilometers away in the
Amhara region (Van Veen 2016). This political history—which also includes a
notable lack of prolonged European colonialism, the survival of Haile Selassie’s
Imperial Kingdom well into the twentieth century, and a punctuated, brutal commu-
nist interlude throughout the 1970s and 1980s—reverberates in contemporary Ethi-
opia and has important implications for understanding and promoting restoration and
biodiversity initiatives in Tigray and beyond.
Tigray’s topography is characterized by its relatively high elevations, ranging
from 1000 to > 3500 m in altitude, and semi-arid temperate climate, with a mean
annual precipitation of 700–1200 mm (Bard et al. 2000). Temperatures range
between 15 and 25 C, while soils range in agricultural quality, from Vertic
Cambisols in the upper reaches, to Vertic Calcisols, Vertic Cambisols, Calcaric
Phaeozems and Calcaric Regosols5 in the lower regions (Rabia et al. 2013). Despite
the difficult conditions, the highland regions served as a center of African plant
domestication and agricultural innovation, and extensive human manipulation of the
landscape has taken place for millennia (Bard et al. 2000). The Tigrean Plateau was
once vegetated with dry evergreen, montane forests and deciduous wooded grass-
lands, yet today it is mostly montane grassland (Pankhurst and Ingrams 1988).

5
A Cambisol is a soil with little or no profile differentiation. They are typically found in landscapes
with high rates of erosion, and are exploited for agriculture. A Calcisol is a soil with a layer of
migrated calcium carbonate in the soil profile. They are typically found in arid zones, and their chief
use is for animal grazing. Phaeozem is a dark soil with high base status typically exploited for
intensive agriculture. Regosols are poorly developed mineral soils in unconsolidated materials,
extensive in eroding lands in arid areas.
654 E. Sigman

This loss of forest cover is the result both of natural disasters such as drought and
landslides, and human activity, including intensive deforestation, agriculture, and
livestock grazing (Nyssen et al. 2000).
It is important to recognize that the barren landscape of present-day Tigray is not
a phenomenon of recent decades, but likely dates back to at least the seventeenth
century (Pankurst 1988). Few, if any, remnant forests exist that are not the direct
result of human intervention in the region. Many such forests are the direct result of
coordinated restoration activities undertaken in the absence of the edaphic and
hydraulic conditions that would facilitate natural regeneration. As such, the great
majority of biodiversity islands that exist in Tigray today should be understood as
the result of intentional restoration and management, which enabled the growth of
forests in an otherwise highly degraded landscape – a point which deserves great
emphasis.

26.3.3 Contemporary Restoration in Ethiopia

While the history of restoration in Ethiopia is not well studied, attempts at better
documentation of present-day restoration activities furnish a cursory understanding
of contemporary restoration dynamics. Much of the restoration is driven by a desire
to deliver ecosystem services, notably the renewal of watersheds to provision water
for agricultural systems (Gebregziabher et al. 2016). Over time, the lack of woody
vegetation on hillsides has destabilized watersheds; instead of percolating slowly
through the rocky soils, episodic rainfall rushes off the slopes, carrying with it
precious topsoil and seedbanks. This not only leads to periodic inundation, sedi-
mentation and gully formation in lowland settlements, but also to the gradual
impoverishment of valley groundwater supplies. The persistent threat of drought
was—and continues to be—compounded by this chronic incapacitation of ground-
water, which was one of many factors that led to the perilous famines witnessed
during the late twentieth century (Keller 1992). Today more than a quarter of the
population of Tigray remains chronically food insecure, requiring the continued
provision of international food aid (Alemu et al. 2014). This precariousness is further
exacerbated by the growing incidence of erratic precipitation and rising temperatures
associated with climate change (Teshome and Zhang 2019).
To reverse this trend and build resilience, international agencies and local com-
munities recognize the need to restore watersheds, which in turn requires afforesta-
tion of degraded hillsides. As a signatory of the Bonn Challenge, supported by the
AFR100, Ethiopia has committed to restore 15 million hectares, or roughly one-sixth
of its total land area, by 2025 (African Resilient Landscapes Initiative, ARLI, https://
[Link]). Tremendous efforts have been taking place across the country under the
banner of the Sustainable Land Management Programme (SLMP), a multi-
stakeholder project commissioned by the World Bank and GIZ (GIZ 2020), which
contributes to these substantial re-greening goals through programs broadly referred
to as ‘Soil and Water Conservation’ (SWC) (Hurni et al. 2016), constituting
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 655

Fig. 26.1 Free Labour Contribution Period (FLCP) activities. On left: two women use a repurposed
food sack to carry stones from the excavation site to the terracing site. On right: A mixed group of
men and women construct a stone terrace. These activities happen side by side continuously.
(Photos: E. Sigman, with permission from community members photographed)

watershed-level interventions on agricultural landscapes. Numerous additional

NGOs, research, and foreign government aid organizations have likewise made
substantial investments towards conservation and reforestation in Ethiopia. While
these are nation-wide efforts, a special emphasis has been placed on the Tigray
region.
With such severe levels of degradation, natural regeneration typically does not
take place even with the exclusion of agricultural and grazing activities on hillsides
(Aynekulu et al. 2009). Moreover, in addition to proving cost-prohibitive, refores-
tation through active tree planting has shown to have dubious benefits, as—in the
absence of underlying functional water and nutrient cycles—many trees do not
survive to catalyze forest succession (Shono et al. 2007). Extensive efforts have
therefore focused on the repair of watersheds, which—in the absence of financial and
technical capital—has been conceived through mass hillside stabilization programs.
Such programs are carried out by large groups of rural residents, often equipped with
little but their own hand tools (see Fig. 26.1), and are followed by ‘exclosure’
systems that restrict access to these areas while natural or assisted regeneration
takes places over several decades (Mekuria et al. 2017). The resultant forests,
constituting newly created biodiversity islands, in addition to provisioning water
and other ecosystem services, support populations of native wildlife and insects and
facilitate connections to other conservation areas.
The greatest contribution to restoration and the creation of biodiversity islands in
Northern Ethiopia therefore comes from rural communities. Farmers and pastoralists
are the agents that ultimately contribute the labor required to stabilize hillsides,
reverse watershed degradation, restore nutrient cycles, and catalyze functional
regeneration. They are also the agents that must significantly alter their land practices
to accommodate these programs, and consent to continued land-use restrictions
which generate, maintain, and safeguard the newly-created biodiversity islands.
656 E. Sigman

26.3.4 Abreha we Atsbeha as a Model Community

for Restoration in Ethiopia

As noted, Ethiopia has gained recognition within the international restoration com-
munity for its recent activities and successes. Since the launch of a series of
nationwide programs in 2010 (MOFED 2010), together with substantial financial
investment from interested international agencies, Ethiopia has invested more than
US$1.2 billion annually in restoration activities across the country, rehabilitating
more than 12 million hectares of land and over 3000 watersheds, and supporting
more than 1.6 million hectares under active ‘Sustainable Land Management’
(Seyoum 2016). Recent research published by the International Center for Tropical
Agriculture (CIAT) confirms that these investments have significantly reduced
runoff and soil erosion, increased crop yields, and enhanced soil organic carbon
(Tamene et al. 2018). Meanwhile, other studies have demonstrated the benefits of
coupling active restoration with community exclosures (Mekuria et al. 2018),
particularly in Tigray, which has led to increased vegetation cover and biodiversity
(Asefa et al. 2003; Mengistu et al. 2005; Mekuria et al. 2012), along with enhanced
soil fertility (Mekuria et al. 2017), water flows (Dessalew et al. 2016), and ground-
water recharge (Anwar et al. 2016).
According to some researchers (Nyssen et al. 2014), as a result of this blend of
national initiatives, foreign assistance, and local participation, “Ethiopia is now
greener than it has ever been during the last 145 years. . .human investments have
overridden the impacts of climate change.” In the midst of this movement—what
some (e.g. Dodd 2015) refer to as Ethiopia’s ‘Green Revolution’—a community
called Abreha we Atsbeha has garnered substantial recognition for its achievements
in both implementation and innovation in the intersecting fields of restoration and
conservation.
Abreha we Atsbeha—a village of about 5030 people in the Tigray Region of
Northern Ethiopia (see Fig. 26.2)—has been widely promoted by both the Ethiopian
federal government and a bevy of international research and aid organizations as a
key example of restoration best practices, both from a biophysical and socioeco-
nomic perspective (Lamond 2012).
Several journal articles and reports have been published specifically studying
Abreha we Atsbeha. Some of these detail historical, demographic, ecological, and
hydrologic conditions of the site and summarize water harvesting and restoration
techniques (e.g. Tadesse et al. 2015), while others measure labor inputs to restoration
(e.g. Hachoofwe 2012). Other reports have focused on livelihoods and economic
stability, detailing the positive impacts of restoration activities on biodiversity,
socioeconomics, production, and policy, while outlining some pathways forward
for sustainability and replication (UNDP 2013).
The majority of information on Abreha we Atsbeha, however, comes from grey
literature, much of which centers on the community’s charismatic chairman
Gibremechel Giday, better known as ‘Aba Hawi’. In 2012, Aba Hawi traveled to
Brazil to attend the Rio + 20 summit and accept the UN Equator Prize, an award
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 657

Fig. 26.2 A day at the market in Abreha we Atsbeha. In the background is the Abreha we Atsbeha
church for which the community is named. This rock-hewn edifice has been in continuous use since
at least the tenth century. (Photo: E. Sigman)

bestowed by the UN Development Program (UNDP) upon 25 initiatives in recog-

nition of their “outstanding projects working to advance sustainable development
solutions for people, nature and resilient communities”. Aba Hawi also features
prominently in a documentary called Ethiopia Rising: From Red Terror to Green
Revolution, which has won several international film awards, and has enjoyed
thousands of public screenings throughout the globe. Stories of Abreha we
Atsbeha’s success have been published in BBC (Haslam 2015), Reuters (Win
2019), The Guardian (Watson 2016), and many other national and international
news publications.6 Blog posts about Abreha we Atsbeha are common among the
websites of several major international organizations working in Ethiopia, including
the World Food Program,7 The World Agroforestry Center (Kuria et al. 2016),
World Vision, World Resources Institute (Rejj 2015) and many others. There are
also a number of short films about Aba Hawi—focusing on his role as a charismatic
leader who has championed for restoration and inspired his constituents to undertake
volunteer restoration work—and about Abreha we Atsbeha, created by visitors and
international organizations on YouTube and Vimeo.
Ethiopia’s regional and federal governments, as well as a number of international
research and development organizations, are actively working to try to ‘scale’ Abreha
we Atsbeha’s success. Yet precisely how this ‘scaling’ will take place remains
unclear. Most efforts thus far have centered on replicating specific ecological

See “Ethiopia, 30 Years on from Famine and Live Aid.” EthiopiaOnline.

6

7
Ethiopian Village Recognized At Rio + 20 For Innovative Hunger Solution. In: United Nations
World Food Program Blog. Accessed 25 Jan 2020
658 E. Sigman

interventions, spreading practices through ‘farmer-to-farmer’ trainings, or provision-

ing particular resources such as materials for check-dam construction or nurseries for
on-farm agroforestry intensification. Despite the significant attention paid to this
community, critical gaps in research remain to understand both why the community
has been so successful and how those successes could be most effectively scaled.
Filling these research gaps is key to assuaging the ‘Restoration Dilemma’ at play in
this case study, and safeguarding the biodiversity islands that have been created in
communities like Abreha we Atsbeha.

26.4 The Free Labor Contribution Period (FLCP)

26.4.1 Contributions of Coordinated Group Labor
to Restoration Programs and Biodiversity Outcomes

Concomitant with rising global awareness and promotion of participatory restoration

and conservation, coordinating bodies—particularly those emanating from interna-
tional agencies—have celebrated Ethiopia’s ‘community-based’ environmental
activities. Widespread praise abounds for what are conceived as democratic, coop-
erative, and innovative restoration initiatives like those underway in Abreha we
Atsbeha. Yet few international agencies, even those heavily vested in restoration
programs, have made systemic inquiries into how participation in restoration is
motivated and coordinated in practice, and existing studies examining cooperative
structures have so far proven simplistic.
‘Scaling’ restoration in the Ethiopian context is not simply a matter of identifying
and promoting technical landscape interventions. The agrarian nature of the county’s
overwhelmingly rural population, coupled with severe levels of environmental deg-
radation, renders scaling a highly social and political undertaking. Moreover, though
Ethiopia’s economy is growing swiftly, and despite significant monetary investments
in restoration, levels of financial and technical capital in these vast rural landscapes
remain low. In this setting, unpaid communal labor provides an effective means to
achieve needed restoration outcomes. For all of these reasons, residents of local
communities are almost invariably the agents of restoration and the stewards of
resultant restored landscapes. The way such residents are motivated, organized, and
sustained in this work should consequently be of paramount interest to those
researchers and agencies concerned with scaling restoration and safeguarding biodi-
versity islands.
It is therefore surprising to see how little attention has been paid to the coordi-
nated group labor systems that animate restoration in practice. Who organizes
people? How are they organized? What incentives motivate this participation?
Few have sought answers to these questions, despite the vital role coordinated
labor plays in achieving the highly praised restoration outcomes in Ethiopia, and
despite the central function such coordinated systems must therefore occupy in
attempts to scale existing successes.
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 659

Fig. 26.3 Newly constructed check dams in an outwash area in Abreha we Atsbeha. (Photo:
E. Sigman)

In particular, little attention has been paid to a widespread phenomenon called in

Tigrinyan “Israin Tshanta”—translating to “Twenty (20) Days”— and referred to by
others in the region as the “Free Labor Contribution Period” (FLCP). The FLCP is a
central feature of rural life in the Tigray region; it is a period lasting between
20–60 days, typically carried out during the driest season (January–March). During
this time, all able-bodied members of a community are expected to volunteer on
community projects. Communities work on a wide range of tasks during this time,
but most of them revolve around landscape restoration. Often, this includes labor-
intensive undertakings such as terracing, check-dam construction (Fig. 26.3), and
other SWC activities, organized and managed with high levels of oversight and
coordination by local and regional officials (Sigman 2019a). The origins and evolu-
tion of the FLCP remain unclear, making it difficult to parse if and how perceptions
about the FLCP have changed over time (Sigman 2019b).
It is estimated that between 2010 and 2015 more than 15 million rural dwellers in
the region contributed the unpaid labor equivalent of US $750 million annually
(Seyoum 2016). Initial assessments report that Tigray—a mountainous area approx-
imating the size of Italy—has been almost entirely terraced. Laid end-to-end the
terraces of Tigray would be longer than the Great Wall of China (Dodd 2015). All of
this has occurred in the last 20–30 years, and much of it through the apparatus of the
FLCP (see Fig. 26.4).
660 E. Sigman

Fig. 26.4 A mountainside in Abreha we Atsbeha undergoing active restoration via terracing and
‘exclosure’ programs. Vegetation and soil can cover evidence of terracing over time, obscuring
from view the human labor required to catalyze regeneration. The two images are identical; on the
right image the author has added lines to show where terracing has taken place. Human shapes in the
photo give a sense of the scale of the undertaking. (Photo and rendering: E. Sigman)

The FLCP has been central to the success of restoration in Abreha we Atsbeha
and elsewhere, and should be understood as the direct human mechanism through
which restoration activities take place. Yet despite the obvious centrality of the
FLCP to Ethiopia’s many large-scale restoration initiatives, the FLCP as a social and
political institution is rarely acknowledged. Though it is a critical feature of rural life
in Tigray, and plays a key role in translating international, federal, and regional land
management strategies to the local level, the FLCP is virtually absent from the
discussions by the international development and restoration community surround-
ing how to best design, manage, evaluate and scale landscape changes, and how to
safeguard resultant biodiversity islands.
The FLCP is a massive social and political institution, through which initiatives
like the Bonn Challenge and AFR100 reverberate, and a substantial portion of
Ethiopia’s pledged 15 million hectares of restoration will likely be facilitated
through the apparatus of this established group labor system. Studying the FLCP
furnishes a keen understanding of how large-scale projects are implemented at the
local level, making it an instructive lens through which to try to parse the ‘Restora-
tion Dilemma’. Such an understanding can help develop projects that are compatible
with local systems, assess how the responsibilities and benefits of such projects flow
through a community, and highlight possible synergies or incongruences that
emerge at the intersection of landscape restoration and political power. These
insights in turn improve prospects for scaling up restoration projects and minimizing
threats to the region’s newly established biodiversity islands.
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 661

26.4.2 Mechanisms of the Free Labor Contribution Period

Theories and methods from political ecology supported the author’s immersive field
visits to Ethiopia in 2017 and 2018, oriented with the express purpose of
documenting the FLCP and illuminating its centrality in restoration and conservation
(Sigman 2019b). These visits focused on deepening an understanding of the labor
dynamics at play during the FLCP in Abreha we Atsbeha, hypothesizing that this
knowledge would provide a more accurate picture of what restoration actually
entails, and therefore improve prospects for linking local level successes to global
goals through scaling efforts (i.e., to assuage the ‘Restoration Dilemma’).
Participant observation, purposive sampling, interviews with key informants, and
other qualitative methods undertaken during visits coinciding with the local FLCP,
led to the creation of a local governance map (Fig. 26.5). This map provided a

Fig. 26.5 ^The Kilil is a region, or state, of Ethiopia. There are 9 Kilils in Ethiopia, divided ethno-
linguistically
^^Woredas are districts within Kilils. There is an assigned woreda liaison responsible for
A/Atsbeha, who occasionally comes to kebele meetings and also assigns and manages A/Atsbeha’s
3 on-site development agents
*The Kebele is “the village” over which the chairman, Aba Hawi, presides. Aba Hawi is elected by
a 208-person parliament (made up of the wahayo or “cells), of which all are members of the TPLF.
Aba Hawi then appoints a 14-person cabinet: 3 wudaba members (people responsible for coordi-
nating with the Woreda), 3 propaganda ministers, 2 finance ministers, 3 ministers of men’s
associations, and 3 ministers of women’s associations. He also selects from this same pool various
committees (including watershed committee), and kushet leaders. Everyone in the cabinet and
committees are TPLF members
**The Kushet is the sub-village unit, determined geographically. Each Kushet is assigned a leader
from the 14-person cabinet. All kushet leaders are party members. Kushet leaders meet with group
leaders every Friday, and with committee members every Saturday
***The ‘Got’ is a neighborhood unit, determined geographically. Gots do not have assigned
representatives, but rather serve as organization units, mostly for the watershed committee when
determining where to do restoration activities
****Groups consist of about 25–30 people and can be made up of both party and non-party
members. Each contain a ‘cell’ which is made up of 4 party members: cell leader, representative
of the cell leader, secretary, and finance. The cell leader represents the group at Friday meetings
662 E. Sigman

conceptual documentation of the decision-making structures that govern the com-

munity as a whole, and which of those are adapted to facilitate the FLCP and other
related programs. It likewise furnished an understanding of important cleavages in
the community, chief among them political affiliation, kushet residence (i.e., geo-
graphic partitioning) and gender.
Figure 26.5 represents a strong hierarchical system exercising power from the
regional government to the household level. Figure captions explain how households
in Abreha we Atsbeha, and other Tigrinyan kebele, are partitioned into gendered
‘groups’, which send representatives both to a local parliament (which elects the
community chairman) and to regular weekly meetings, where decisions concerning
the community as a whole are regularly made. It is during these regular weekly
meetings, and through directives from the parliamentary-elected community
chairman’s chosen cabinet, that agendas concerning land use and the FLCP are
set. While the meetings themselves are participatory and democratic from the
perspective of the elected and appointed members, access to these meetings turned
out to be highly restrictive.
One of the key insights that emerged from the conceptual mapping exercise was
that, while in theory, residents of any political denomination could serve as cell
members (i.e., as parliamentary and weekly meeting representatives), in practice,
only recognized members of the ruling party (The Ethiopian People’s Revolutionary
Democratic Front – EPRDF) had ever held these positions. This homogeneity occurs
despite the fact that less than a sixth of the population in Abreha we Atsbeha are
recognized as EPRDF members.
Given Ethiopia’s recent political history, this single-party hegemony is hardly
surprising. However, recognizing that the supremacy of the party reaches all the way
down to the kebele and even household level is important, especially given the fact
that such a small portion of the population possesses access to representation and
agency. This recognition contends with the realities of ‘community-driven’ restora-
tion. What the conceptual map revealed is that restoration activities, carried out
through the FLCP, are conducted by a rural citizenry that remains systematically
removed from the decision-making processes that govern them, and possesses little
to no recourse for meaningful participation in anything other than the FLCP itself.
This is not to say that local actors do not understand the value of their work, or
appreciate the ways in which the restoration activities they undertake—and the
biodiversity islands they collectively safeguard through exclosures—benefit them
directly. Local actors benefit mostly via the increased provisioning of water, and
through ‘cut and carry’ systems, wherein citizens are allowed to enter the exclosures
on foot and hand harvest grasses and other fodder to bring back to their on-farm
livestock. Ecological literacy and favorable perceptions of the ‘water banks’ created
by restoration was high among virtually all surveyed respondents in Abreha we
Atsbeha. Rather, it is to suggest that outside perceptions of restoration in Ethiopia—
particularly those of international agencies lavishing praise on the participatory
nature of the restoration activities—typically do not comprehend the stratification
of power in local communities, and therefore fail to grasp the degree to which
participation in restoration relies on a system of mass labor mandated by an elite
political minority.
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 663

This realization raises a number of thorny questions for those interested in

promoting and scaling participatory restoration initiatives. Most germane to this
chapter is the question of longevity; such an asymmetrical system has the potential to
breed discontent, and therefore threatens to destabilize restoration programs in the
long-term. Safeguarding biodiversity islands in this context therefore requires an
explicit understanding of the governance structures that sustain them, and a whole-
sale accounting of the myriad ways international agencies and initiatives reverberate
through local configurations.

26.4.3 The Free Labor Contribution Period in a Pluralizing

Context

In early 2018, Ethiopia declared a national state of emergency following a mounting

series of protests which culminated in the resignation of Hailemariam Dessalegn, the
Prime Minister of Ethiopia and Chairman of the EPRDF.8 These events coincided
with the FLCP in Abreha we Atsbeha, whose regularly planned activities were
interrupted by a series of emergency meetings held in kebeles at the behest of
concerned district and regional party leaders. At various meetings, ranging in size
and topic, registered party members were asked to voice their grievances, and share
their ideas and opinions. A palpable aura of discontent—much of it directed at the
FLCP and related programs—permeated throughout the course of these intensive,
multi-day hearings. Decades of suppressed criticisms were unleashed as the fear of
sharing long-held negative views dissolved.
Grasping the significance of these meetings requires some understanding of
Ethiopia’s recent political history. Starting in 1974, following the famine-induced
collapse of the empire of Haile Selassie, Ethiopia was governed by a brutal military
junta known as The Derg, followed by a dictatorial regime administered by a
communist state party called People’s Democratic Republic of Ethiopia (PDRE).
Opposition to the Derg came primarily from groups based in the North, in Tigray.
These opposition movements coalesced in the late 1980s, forming a powerful
shadow government known as the Ethiopian People’s Revolutionary Democratic
Front (EPRDF) in 1988 (Rahmato 2009).
The EPRDF inflicted a total military defeat on the communist regime and
established in 1991 a new government that has ruled Ethiopia ever since. This
government has presided over nearly three decades of essentially single-party rule,
exercising considerable control over the entire federation (Feltor 2018). The EPRDF
is both the political party that committed Ethiopia to the Bonn Challenge and related
restoration targets, and the political party that oversees and administers the FLCP, one
key method through which these targets are currently met in Tigray and expected to
be met across the country. This is especially true in the Tigray region, where Abreha
we Atsbeha is located, and where the hegemony of the EPRDF is particularly strong.

8
Ethiopia declares national state of emergency. BBC News (2018)
664 E. Sigman

The EPRDF has suppressed many critical groups movements throughout its
tenure, but mass protests and unrest beginning in 2016 have proven—so far—
politically transformative. In April 2018, shortly after the conclusion of the FLCP
in Abreha we Atsbeha, notable reformist Abiy Ahmed was sworn in as the new
Prime Minister, becoming Ethiopia’s first ethnically Oromo leader, thereby weak-
ening the stronghold of Tigrinyan power. Ahmed swiftly launched a wide program
of internationally-acclaimed political and economic reforms, and was awarded the
2019 Nobel Peace Prize for his work in brokering reconciliation with Eritrea,
Tigray’s tumultuous northern neighbor. Most recently, Sahle-Work Zewde was
installed as Ethiopia’s first female president, and Africa’s only serving female
head of state. The international community has generally lauded Ethiopia’s recent
progress towards a more open, pluralistic society and improved international
relations.
Yet the single-party, restrictive government of Ahmed and Sahle-Work’s pre-
decessors remains central in places like Abreha we Atsbeha, and still resides at the
core of rural institutions like the FLCP. During the emergency meetings held in lieu
of 3 days of FLCP activities in Abreha we Atsbeha, many community members
levied significant complaints and accusations against the party: its structure, its
leaders, and specifically, the restoration and conservation programs it engenders.
These events, along with the governance hierarchies described in Fig. 26.5, made
clear that this autocratic system of government, especially at the local level—is the
force that is overwhelmingly responsible for current restoration programs. Likewise,
it is the same authority that sets norms and bylaws around continued safeguarding of
the community’s hard-earned biodiversity islands.
Scaling Abreha we Atsbeha’s ecological success currently involves replicating
these obscured socio-political dynamics. At the same time, there is little guarantee
that such programs—invariably bound up in the politics of the last several decades—
will prove compatible with the evolution of the country’s governance systems as a
whole. Safeguarding biodiversity islands therefore requires an explicit recognition of
the FLCP, the socio-political structures that animate it, the misalignment between
these existing structures and the advancement of Ethiopia’s social, political and
economic systems as a whole.

26.5 Sustaining Restoration and Safeguarding Biodiversity

Islands Amidst Social and Political Change
26.5.1 Acknowledging Labor Realities in Restoration
and Conservation Programs

This chapter has repeatedly stressed the central role that coordinated group labor
plays in the rural regions of Tigray. This centrality is a reflection of the region’s
manifest socio-ecological conditions: the landscape is semi-arid, mountainous, and
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 665

highly degraded and the communities living there chronically lack financial capital
and equipment. Establishing biodiversity islands where prolonged environmental
degradation limits the ability for ecosystems to regenerate naturally requires sub-
stantial acts of human intervention, and this intervention must fit within the existing
limits of community capacities.
These human interventions, moreover, must be thoughtfully planned and exe-
cuted. Restoration of watersheds and the creation of biodiversity islands is a multi-
farious undertaking, requiring not only ecological but also sociological expertise and
coordination. If human labor is the ultimate and direct mechanism through which
restoration is achieved, then those concerned with promulgating successful restora-
tion must be expressly concerned with the dynamics of that labor. The paucity of
attention paid to this critical aspect of restoration in the region suggests that
international and federal agencies alike largely take labor—including mass coordi-
nated labor like the FLCP—for granted.
The previous section detailed the operations of a substantial local hierarchy,
supported by a hegemonic regional authority that has been historically empowered
by a single-party federal government. This hierarchy coordinates rigorous restora-
tion activities among large segments of rural society, organizing, incentivizing, and
monitoring individual and group performance in a calculated, iterative, and—from
many perspectives—effective manner. Owing to this hierarchy, inhabitants of local
communities abide by social norms dictating that their contributions be made
reliably, compulsorily, and without pay. Acceptance of these bylaws thus substan-
tially reduces the costs associated with restoration and thereby makes it possible in
this otherwise limited context. Furthermore, the maintenance of this hierarchy and
system of labor control is presently critical to the continued creation and protection
of biodiversity islands in Tigray.
It is difficult to overstate the point that biodiversity islands in this region are
generally not forest remnants to be protected against human encroachment, but
rather are sites reflecting extraordinary feats of intentional restoration. The continued
provisioning of such exclosure systems requires enormous community buy-in,
strong bylaws against harvesting infractions, and high levels of ecological literacy
that entail an understanding of the connections between hillside vegetation and
lowland agriculture.
At the same time, these exclosure systems are not akin to national parks or
protected areas; they typically remain the communal property of the kebele. Such
ecosystems therefore depend not only on the regeneration of woody species and the
encouragement of native animals and insects, but also on the continued renewal of
community consent and respect for use-norms. The landscape of Tigray is a
cautionary tale: systems don’t always regenerate automatically. This is as true of
the ecosystems as of the socio-political systems that are bound in the biodiversity
islands of Abreha we Atsbeha. Paying attention to community labor programs is key
for pinpointing critical vulnerabilities and building capacity around them before the
system reaches a tipping point.
666 E. Sigman

26.5.2 Recognizing Complexity in Local-Level Motivations

Precisely because people are not compensated financially for their contributions in
the FLCP, scholars and practitioners must carefully consider the other motivations at
play in these spaces. The author conducted interviews in Abreha we Atsbeha, where
people generally displayed an impressive understanding of the complex watershed
dynamics in the area, and were motivated by the understanding that restoring
vegetation to hillsides would secure water supplies in their agricultural fields
(Sigman 2019a). They also appreciated other benefits of restoration, such as
increased biodiversity—especially wild animals—and access to cooler microcli-
mates. At the same time, individuals also reported that they participated in the
FLCP because they feared punishment if they withheld their labor or complained
(Sigman 2019b).
This punishment could come in the form of social castigation or perceived
favoritism of others by those in power, suggesting strong local norms around
group participation. Additionally, participation in the FLCP is a prerequisite for
membership in Tigray’s welfare system, called the Productive Safety Net Program
(PSNP). Through the PSNP, vulnerable members of society can receive subsistence
amounts of food or money in exchange for an extension of their labor contribution to
restoration activities. However, those members who are physically able to work must
still contribute their labor without compensation during the FLCP in order to be
eligible for the program. There is thus a complexity of compulsion-based motiva-
tions at play, which affect different segments of society asymmetrically, and may
constitute a major obstacle to effective scaling.
There is a wide range of perceptions and motivations surrounding restoration and
biodiversity islands in the area. These constitute not only different perceptions
between different people, but individual people can have blended motivations—
both positive and negative—for participating. Continued adherence to exclosure
zones (i.e., the safeguarding of biodiversity islands) is driven by both positive and
negative motivations; people can appreciate biodiversity islands (or exclosures) and
the work it takes to create and maintain them while still holding negative views about
the process. As demonstrated in the emergency meetings, these negative views are
linked with feelings of disempowerment. Empowerment of stakeholders in the
community therefore plays a key role in safeguarding biodiversity islands against
these negative perceptions.

26.5.3 Understanding ‘Political Monoculture’ as a Threat

to Biodiversity

Best practices in socio-ecological restoration demand that all stakeholders be

able to actively influence and make decisions about the restoration activities that
impact their daily lives. Yet in Abreha we Atsbeha—perhaps Ethiopia’s most
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 667

famous socio-ecological restoration case study—the majority of people who dedi-

cate substantial time, labor, and resources to the work of restoration have very little
access to the decision-making structures that would afford them a meaningful voice
in the deliberation process. The nature, duration, location, and timing of restoration
activities in Abreha we Atsbeha’s FLCP, for example, are largely dictated by
decisions made by Aba Hawi (the community chairperson) and his cabinet. The
cabinet—relying on information gleaned from the kushet9 meetings and taking into
account directives from the woreda—will come up with a plan for the FLCP, which
is then approved by the parliament. This entire process remains confined to regis-
tered party members, though the work of the FLCP necessarily involves many
people who do not have a voice in the party system. Participatory restoration projects
call for exactly the opposite structure, with full participation of all involved com-
munity members at each stage of the planning process.
Knowledge of this local political process is critical for understanding how the
FLCP operates, and for thinking about how agency, attitudes, and voluntary labor
are bound up in the restoration activities the FLCP advances. Within Abreha we
Atsbeha’s participatory restoration program, representation and agency among local
stakeholders is essentially limited to EPRDF party members, despite the fact that
registered members comprise only a sixth of the population. This asymmetry in
representation, and its implications for long-term management of restoration pro-
grams, is therefore concerning.
The administrative system which has historically enabled restoration and
safeguarded biodiversity islands in the region can be thought of as a political
‘monoculture’. There exists one elite party, which despite comprising only a fraction
of the population, dictates all of the activities of that population, in particular the
labor-intensive activities associated with restoration, and the restrictive norms
governing community exclosure from newly established biodiversity islands. This
concentrated local political power may be seen as a consequence of the region’s
unique political history, and its extant behaviors perhaps justified to many by the
exigencies of extreme land degradation coupled with the looming threat of famine.
For these reasons and others, this system has been able to operate on the margins of
the rights concerning representation and labor conditions which are otherwise
typically espoused by the international development agenda.
However, those seeking to replicate this case study and safeguard the biodiversity
islands in the region should recognize that intensive manual labor, ironclad bylaws,
and political hegemony have played a central role in its evolution. The task of
organizing thousands of people to give up communal grazing lands, volunteer up
to two months of hard labor, and wait decades to reap the benefits of their work has
been won in part by charisma and solidarity, and also in part by intense social
pressures, by fear of castigation, and by a local political system that restricts much
meaningful participation to all but a hand-selected elite.

9
See Figure 26.1 caption for description of ‘kushet’ and ‘woreda’ units
668 E. Sigman

This elite is bound up in what—for three decades—has been a recalcitrant,

‘political monoculture’ across the country. Just as diversity is a critical component
of ecological resilience, so too is diversity a critical component of socio-political
resilience. Restoration and conservation systems that rely in large part on political
hegemony may be vulnerable to political changes, whether as a result of acute
emergencies or long-term stressors.
While political hegemony may in fact be central to the success Abreha we
Atsbeha has enjoyed in the past, Ethiopia’s political system is becoming increasingly
pluralistic at the national level, and—as emergency meetings held during the
author’s fieldwork activities in 2018 demonstrated—this shift is having growing
impacts at the local level. Those interested in continuing Ethiopia’s restoration
success must therefore begin to think about how such large-scale and labor-intensive
projects can continue or be supplemented if local political institutions start to divest
power to a wider range of voices and agendas.

26.5.4 Need for Reassessment and Realignment Among

International Coordinating Organizations

Abreha we Atsbeha is undoubtedly a case study in successful dryland restoration

across a number of metrics. Through decades of thoughtful, ecologically-sound and
socio-economically sensitive interventions and innovations, the community of
Abreha we Atsbeha has effectively stewarded a resilient biophysical ecosystem in
their kebele. This effort has provided substantial social and economic benefit to the
community as a whole.
At the same time, Abreha we Atsbeha’s ecological and economic success is
linked to the perpetual reifying of its social and political systems. The international
community may be inadvertently bolstering these systems in Abreha we Atsbeha
through its unexamined praise of the community’s restored biodiversity islands.
Moreover, it may be unwittingly promoting the replication of such systems in other
areas in the attempts to scale Abreha we Atsbeha’s successes in order to meet the
demands of the Bonn Challenge and other such initiatives. A healthy debate on
whether or not international sustainability and development organizations want to be
associated with these kinds of political systems cannot even begin until there is a
greater degree of reflection on the existence of these systems—and indeed their
centrality—in the work of restoration in the first place.
Thorny questions on alignment between potentially conflicting development
priorities aside, there is a pressing issue of practicality. With the country’s recent
national reforms, civil society in Ethiopia seems to be moving in a number of
positive directions, including provisioning a more democratic, open and pluralistic
public sphere. Could this increased plurality and openness pose an unrecognized
threat to restoration and conservation programs built on hegemony? Can FLCP-
mediated restoration be modified to be more inclusive in this new political
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 669

landscape, perhaps by embracing more voices in the decision-making process? If

not, what replaces the FLCP? What safeguards the exclosure zones? In short, how
will restoration weather democratization? (NB: this chapter was authored prior to the
surge in political violence that began sweeping the Tigray region in 2021 and,
unfortunately, has reversed the country's progression towards peaceful pluralism.
Recent events pose an even more urgent question: how will restoration weather
another civil war?)
Those concerned with safeguarding biodiversity islands like those in Abreha we
Atsbeha would do better to stay ahead of these questions than behind them. This can
start with acknowledgement of local labor realities, recognition of complexity in
local-level motivations, an understanding of relationships between ‘political mono-
culture’ and landscape-level biodiversity, and better alignment between international
coordinating organizations, national initiatives, and local actors.

26.6 Conclusion

High levels of environmental degradation worldwide have spurred an increased

appreciation for the contributions restoration can and must make towards ameliorat-
ing a number of interconnected global challenges. Restored areas serve as biodiver-
sity islands in otherwise degraded landscapes, improving ecosystem connectivity
and function, and offering a number of irreplaceable services to human societies
around the globe. Recent years have witnessed mounting enthusiasm for restoration
and an expanding consciousness surrounding the intricacies of socio-ecosystems.
Substantial initiatives like the Bonn Challenge and the UN Sustainable Development
Goals have elevated this awareness to the international arena, and have catalyzed
mass-scale restoration movements while provoking research on local-level behav-
iors and impacts.
While this upsurge in awareness and activity has promised great benefits—such
as improved food and water security, increased rural economic development, climate
change adaptation and mitigation, and biodiversity protection—the realities of
restoration in practice are still unfolding. Chief among the challenges faced by
restoration practitioners is a phenomenon introduced here as the ‘Restoration
Dilemma’. The dilemma posits that restoration is highly site-specific and therefore
resists scaling, and yet, degradation tends to be so pronounced that restoration
usually must be scaled in order to realize its promised benefits (Sigman 2021).
To explore possibilities for assuaging the ‘Restoration Dilemma’, the present
chapter focused on a case study from Northern Ethiopia, in the community of Abreha
we Atsbeha. Here, a local community successfully re-vegetated substantial areas of
highly degraded landscapes through integrative watershed management and hillside
exclosures. This local-level restoration occurs in the context of a significant nation-
wide restoration platform called the Sustainable Land Management Program
(SLMP)—the largest of its kind in Africa—and has been supported by a number
of multinational restoration initiatives, including The Bonn Challenge.
670 E. Sigman

The restoration activities of the village, and the biodiversity islands they enabled,
have been widely lauded by the international community, and many have offered
Abreha we Atsbeha as an example of how effective, coordinated, multi-scale
restoration and conservation might proceed throughout the region and across the
globe. Yet, as this analysis demonstrates, an insufficient amount of attention had
been paid to the local-level dynamics that enable these restored landscapes. In
particular, there has been little to no appreciation of the substantial ‘Free Labor
Contribution Periods’ that were key to the creation of current biodiversity islands
and remain central to the restoration programs in the region.
The expectation of substantial amounts of uncompensated rural labor exists in the
context of a hegemonic, single-party ‘political monoculture’. This stands at odds
with the present-day realities of a country that is rapidly becoming politically
pluralized. Sustaining restoration and safeguarding biodiversity islands amidst social
and political change in Ethiopia therefore requires: (1) acknowledgement of local
labor realities; (2) recognition of complexity in local-level motivations; (3) under-
standing relationships between ‘political monoculture’ and landscape-level biodi-
versity, and; (4) better alignment between international coordinating organizations,
national initiatives, and local actors.
As the international community looks to the future of restoration, and together
shapes the aims and aspirations of the global restoration movement, we must pay
attention to governance and to power. Just as we work to promote diversity in our
ecosystems, so too should we explore questions of political plurality and gover-
nance. This exploration should begin with an appreciation for the substantial and
complex human labor dynamics that enable biodiversity islands, and should moti-
vate us to probe deeper into the political structures that motivate, organize, and
conscript them. In this way, we may begin to untangle the ‘Restoration Dilemma’
and, with care, enable diverse political, social, and ecological systems to thrive.

Acknowledgements This work would not have been possible without the generosity, assistance,
knowledge, and hospitality offered by Aba Hawi, his family, and the many gracious citizens of
Abreha we Atsbeha. Additionally, Cathy Watson, Niguse Hagazi, and Dr. Kiros Hagdu of the
World Agroforestry Centre (ICRAF) supported the initial stages of this project, and were instru-
mental in facilitating later independent research. Abrha Berhane (Mekelle University) and Zufan
Tsadik provided key translation and additional field analysis. Dr. Gordon Geballe (Yale FES)
provided advice and resources at the planning stages, and Dr. Florencia Montagnini (Yale FES)
served as advisor throughout the research process, and offered constructive editing. This work was
supported by the National Science Foundation Graduate Research Fellowship Program under Grant
No. DGE1122492, as well as by the Yale Tropical Resources Institute, the Yale Jackson Institute
for Global Affairs, and Yale FES. The author also gratefully acknowledges the financial support of
the CGIAR Research Program on Forests, Trees and Agroforestry and the CGIAR Trust Fund
Donors.
Any opinions, findings, and conclusions or recommendations expressed in this material are
those of the author’s and do not necessarily reflect the views of supporters.
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 671

References

Aerts R, Honnay O (2011) Forest restoration, biodiversity and ecosystem functioning. BMC Ecol
11:29. [Link]
Alemu A, Getahun M, Bezabih T (2014) Comprehensive Food Security and Vulnerability Analysis
(CFSVA). Ethiopia Central Statistical Agency/World Food Program
Anderson W, Pembert N (2019) Across Africa, restoring land is climate action. AFR100 Blog
Anwar AA, Dessalew WA, Seifu AT, Mekuria W, Mulugeta A, Tammo SS (2016) Effectiveness of
land and water management interventions in reducing runoff and soil erosion in the north-
western highlands of Ethiopia. Paper presented at First African ESP Conference; Nairobi, Kenya
Asefa D, Oba G, Weladji R, Colman J (2003) An assessment of restoration of biodiversity in
degraded high mountain grazing lands in northern Ethiopia. Land Deg & Dev 14:25–38. https://
[Link]/10.1002/ldr.505
Aynekulu E, Denich M, Tsegaye D (2009) Regeneration response of Juniperus procera and Olea
europaea subsp cuspidata to exclosure in a dry afromontane forest in northern Ethiopia. Mount
Res and Dev 29:143–152. [Link]
Bard K, Coltorti M, DiBlasi CM, Dramis F, Fattovich F (2000) The environmental history of Tigray
(northern Ethiopia) in the middle and late Holocene: a preliminary outline. Afr Archaeol Rev 17:
65–86. [Link]
Benayas J, Newton A, Diaz A, Bullock J (2009) Enhancement of biodiversity and ecosystem
services by ecological restoration: a meta-analysis. Science 325:1121–1124. [Link]
1126/science.1172460
Berkes F, Folke C (1998) Linking social and ecological systems: management practices and social
mechanisms for building resilience. Cambridge University Press, Cambridge
Boesing AL, Nichols E, Metzger JP (2018) Biodiversity extinction thresholds are modulated by
matrix type. Ecography 41(9):1520–1533
Brancalion P, Cardozo I, Camatta A (2014) Cultural ecosystem services and popular perceptions of
the benefits of an ecological restoration project in the Brazilian Atlantic forest. Restor Ecol 22:
65–71. [Link]
Brudvig L (2011) The restoration of biodiversity: where has research been and where does it need
to go? Am J Bot 98:549–558. [Link]
Budiharta S, Meijaard E, Wells J (2016) Enhancing feasibility: incorporating a socio-ecological
systems framework into restoration planning. Environ Sci Policy 64:83–92. [Link]
1016/[Link].2016.06.014
Colding J, Barthel S (2019) Exploring the social-ecological systems discourse 20 years later. Ecol
Soc. [Link]
Costanza R, de Groot R, Sutton P, van der Ploeg S, Anderson SJ, Kubiszewski I, Farber S, Turner
RK (2014) Changes in the global value of ecosystem services. Glob Environ Chang 26:152–158
Covelli-Metcalf E, Mohr J, Yung L (2015) The role of trust in restoration success: public
engagement and temporal and spatial scale in a complex social-ecological system. Restor
Ecol 23:315–324. [Link]
Daily G (1997) Nature’s services. Island Press, Washington, DC
Daily G, Matson P (2008) From theory to implementation. PNAS 105:9455–9456. [Link]
10.1073/pnas.0804960105
Dessalew WA, Anwar AA, Seifu AT, Mekuria W, Charles FN, Tammo SS (2016) Enhancing
ecosystem services in the upper Blue Nile Basin, Ethiopia through the implementation of soil
and water conservation measures. Presented at First African ESP Conference, Nairobi
Dodd M (2015) Ethiopia rising: Red terror to Green Revolution. 1080 Films, Abreha we Atsbeha,
Ethiopia
Felter C (2018) Ethiopia: East Africa’s emerging giant. Council on Foreign Relations Backgrounder
Fernández-Manjarrés JF, Roturier S, Bilhaut AG (2018) The emergence of the social-ecological
restoration concept. Restor Ecol 26(3):404–410
Garner B (2009) Garner’s modern American usage. Oxford University Press, USA, Oxford
672 E. Sigman

Gebregziabher G, Abera DA, Gebresamuel G, Giordano M, Langan S (2016) An assessment of

integrated watershed management in Ethiopia. Colombo, Sri Lanka: International Water Man-
agement Institute (IWMI). 28p. (IWMI Working Paper 170). [Link]
GIZ (2020) Sustainable use of rehabilitated land for economic development (SURED). In: [Link].
[Link] Accessed 25 Jan 2020
Habtezion S, Adelekan I, Aiyede E (2015) Earth system governance in Africa: knowledge and
capacity needs. Curr Op in Envi Sust 14:198–205. [Link]
Hachoofwe, EM (2012) Local ecological knowledge of trees on farms, constraints and opportunities
for further integration in Tigray Region, northern Ethiopia: A case study of smallholder farmers
in Abreha Wa Atsbeha and Adi gudom. MSc thesis in Agroforestry. Gwynedd, UK: Bangor
University
Haslan C (2015) Turning Ethiopia’s desert green. BBC News
Hobbs R, Higgs E, Harris J (2009) Novel ecosystems: implications for conservation and restoration.
Trends Ecol Evol 24:599–605. [Link]
Hurni H, Berhe WA, Chadhokar P, Daniel D, Gete Z, Grunder M, Kassaye G (2016) Soil and water
conservation in Ethiopia: guidelines for development agents. Second revised edition. Bern,
Switzerland: Centre for Development and Environment (CDE), University of Bern, with Bern
Open Publishing (BOP). 134 pp
Keller E (1992) Drought, war, and the politics of famine in Ethiopia and Eritrea. J Mod Afr Stu 30:
609–624. [Link]
Krech S (2000) The ecological Indian. W.W. Norton, New York
Kumar P (ed) (2010) The economics of ecosystems and biodiversity ecological and economic
foundations. Earthscan, London and Washington, 422pp
Kuria A, Winowiecki, L, Ouya D (2016) Fresh water, the reward of land restoration, flows in
Ethiopia’s dry zone. In: Agroforestry World. [Link]
6/11/10/water-reward-land-restoration-flows-ethiopias-dry-zone/. Accessed 29 Jan 2020
Lamond, G (2012) Local knowledge training using the AKT5 software and methodology at Mekelle
University, funded by the AfricaRISING project: a report on a two week research study in the
village of Abreha We Atsbeha in Tigray Region, northern Ethiopia. Nairobi, Kenya: World
Agroforestry Centre
Levis C, Costa F, Bongers F (2017) Persistent effects of pre-Columbian plant domestication on
Amazonian forest composition. Science 355:925–931. [Link]
López DR, Cavallero L, Easdale MH, Carranza CA, Ledesma M, Peri PL (2017) Resilience
management at landscape level: an approach to tackle social-ecological vulnerability of agro-
forestry systems. In: Montagnini F (ed) Integrating landscapes: agroforestry for biodiversity
conservation and food sovereignty, advances in agroforestry 12. Springer, Cham, pp 127–148
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press,
Princeton, 203 pp
Mace G, Norris K, Fitter A (2012) Biodiversity and ecosystem services: a multilayered relationship.
Trends Ecol Evol 27:19–26. [Link]
Martin D (2017) Ecological restoration should be redefined for the twenty-first century. Restor Ecol
25:668–673. [Link]
Maynard C (2013) How public participation in river management improvements is affected by
scale. Area 45:230–238. [Link]
McKenna A (2019) Tigray, historical region, Ethiopia. Encyclopedia Brittanica Online, https://
[Link]/place/Tigray-historical-region-Ethiopia
Mekuria W, Veldkamp E (2012) Restoration of native vegetation following exclosure establishment
on communal grazing lands in Tigray, Ethiopia. App Veg Sci 15:71–83. [Link]
j.1654-109x.2011.01145.x
Mekuria W, Langan S, Noble A, Johnston R (2017) Soil restoration after seven years of exclosure
management in northwestern Ethiopia. Land Deg & Dev 28:1287–1297. [Link]
1002/ldr.2527
26 Safeguarding Biodiversity Islands in Northern Ethiopia Amidst Political Change 673

Mekuria W, Wondie M, Amare T (2018) Restoration of degraded landscapes for ecosystem services
in North-Western Ethiopia. Heliyon 4:e00764. [Link]
Mengistu T, Teketay D, Hulten H, Yemshaw Y (2005) The role of enclosures in the recovery of
woody vegetation in degraded dryland hillsides of central and northern Ethiopia. J Arid Env 60:
259–281. [Link]
Ministry of Finance and Economic Development (MoFED) (2015) The Federal Democratic Repub-
lic of Ethiopia Growth and Transformation Plan (GTP). Ministry of Finance and Economic
Development (MoFED), Addis Ababa
Minnemeyer S, Laestadius L, Sizer N (2014) Atlas of forest and landscape restoration
opportunities. In: World Resources Institute. [Link]
and-landscape-restoration-opportunities. Accessed 29 Jan 2020
Nyssen J, Haile M, Moeyersons J (2000) Soil and water conservation in Tigray (Northern Ethiopia):
the traditional dagat technique and its integration with introduced techniques. Land Deg & Dev
11:199–208. [Link]
Nyssen J, Frankl A, Haile M (2014) Environmental conditions and human drivers for changes to
north Ethiopian mountain landscapes over 145 years. Sci Total Environ 485-486:164–179.
[Link]
Pankhurst R, Ingrams L (1988) Ethiopia engraved. Kegan Paul International, London
Rabia AH, Afifi RR, Gelaw AM, Bianchi S (2013) Soil mapping and classification: a case study in
the Tigray Region, Ethiopia. J Agri Env Int Dev 107(1):73–99. [Link]
20131.81
Rahmato D (2009) 2 peasants and agrarian reforms: the unfinished quest for secure land rights in
Ethiopia. Land Rights 33
Reij C (2015) How Ethiopia went from famine crisis to Green Revolution. In: World Resources
Institute Blog. Accessed 25 Jan 2020
Rovere A (2015) Review of the science and practice of restoration in Argentina: increasing
awareness of the discipline. Restor Ecol 23:508–512. [Link]
Schroth G, Fonseca G, Harvey C (2004) Agroforestry and biodiversity conservation in tropical
landscapes. Island Press
Seyoum Y (2016) Forest landscape restoration experiences in Ethiopia. AFR100 Conference, Addis
Ababa
Shono K, Cadaweng E, Durst P (2007) Application of assisted natural regeneration to restore
degraded tropical forestlands. Restor Ecol 15:620–626. [Link]
2007.00274.x
Sigman, E (2019a) The role of group labor in socioecological restoration: perspectives from Abrha
we Atsbeha, Fourth Annual World Agroforestry Congress, Montpellier, France
Sigman E (2019b) Who decides? Restoration policies, local governance, and group labor: a case
study from Ethiopia. Yale Environmental Leadership and Training Initiative. Yale University,
New Haven
Sigman E (2021) The Dilemma of Scale: competing imperatives for global restoration. Restor Ecol,
29: e13408. [Link]
Tadesse A, Gebrelibranos T, Geberehiwot M, Mezgebu A (2015) Characterization and impact
assessment of water harvesting techniques: a case study of Abreha Atsbeha watershed, Tigray,
Ethiopia. Open Agricultural Resources (OER) Africa
Tamene L, Abera W, Woldearegay K, Tibebe D, Tadesse M, Admassu Z, Yaekob T, Tefere H,
Alene T, Dubale W, Amede Y, Hailu H, Demeke G, Mekonnen K, Thome P (2018) Land
restoration initiatives and their performances in Ethiopia: a systematic assessment based on
meta-data analysis. AfricaRISING
Teshome A, Zhang J (2019) Increase of extreme drought over Ethiopia under climate warming. Adv
Met 2019:1–18. [Link]
Trigger D, Mulcock J, Gaynor A, Toussaint Y (2008) Ecological restoration, cultural preferences
and the negotiation of ‘nativeness’ in Australia. Geoforum 39:1273–1283. [Link]
1016/[Link].2007.05.010
674 E. Sigman

United Nations (2020) About the sustainable development goals. In: United Nations Sustainable
Development. [Link]
Accessed 25 Jan 2020
United Nations Development Program (2013) Abrha Weatsbha Community, Ethiopia. NY,
New York
United Nations Environment Program: News and Stories (2019) Ethiopia plants over 350 million
trees in a day, setting new world record
Uprety Y, Asselin H, Bergeron Y (2012) Contribution of traditional knowledge to ecological
restoration: practices and applications. Écoscience 19:225–237. [Link]
3-3530
Van Veen E (2016) Perpetuating power: Ethiopia's political settlement and the organization of
security. Chapter 1: dynamics of political power in Ethiopia: Past and Present
Watson C (2016) No one leaves anymore’: Ethiopia’s restored drylands offer new hope. The
Guardian
Win T L (2019) With fruits and green shoots, Tigray swaps image of famine for resilience. Reuters
World Bank (2015) NEPAD launches initiative for the resilience and restoration of African
landscapes
Wortley L, Hero J, Howes M (2013) Evaluating ecological restoration success: a review of the
literature. Restor Ecol 21:537–543. [Link]
 Part V
Conclusions
Chapter 27
Conclusions: Challenges and Opportunities
in Implementing Biodiversity Islands

Florencia Montagnini, Ryan T. Smith, Benjamin Everett-Lane,

Sara del Fierro, and Dara Albrecht

Abstract This concluding chapter presents the lessons learned from the chapters in
the four previous parts of this book: (I) Introduction; (II) Biodiversity Islands
Establishment and Management: Challenges and Alternatives; (III) Biodiversity
Islands Across the Globe: Case Studies; and (IV) Safeguarding the Environmental,
Social, and Economic Benefits of Biodiversity Islands. Constraints limiting the
adoption of Biodiversity Islands (BI), include conceptual, biophysical, economic,
political, social, and cultural factors. Opportunities for increasing the implementa-
tion of BI are presented, particularly chances for working with groups from private
conservation initiatives, such as those representing local communities, indigenous
peoples, and conservation organizations. Examples of policies promoting agroecol-
ogy are discussed, as well as current trends in conservation which support the BI
concept. Despite the challenges posed to BI, local motivation, political will, and the
right educational campaigns, can allow economically prosperous human communi-
ties and biodiversity to thrive harmoniously within shared landscapes. Many inter-
national efforts are currently underway, creating sustainable and dynamic BI within
human-dominated environments. BI are a critical strategy for conservation in the
twenty-first century while having the added benefit of contributing to climate
adaptation and resiliency solutions. This book serves as a tool for policy makers,
practitioners, and researchers interested in increasing the implementation of Biodi-
versity Islands.

F. Montagnini (*) · R. T. Smith

School of the Environment, The Forest School, Yale University, New Haven, CT, USA
e-mail: fl[Link]@[Link]
B. Everett-Lane · D. Albrecht
Yale University, Yale College, New Haven, CT, USA
S. del Fierro
United States Department of Agriculture (USDA), Washington, DC, USA

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 677
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
678 F. Montagnini et al.

Keywords Agroforestry · Biodiversity conservation · Community action ·

Ecosystem services · Indigenous communities · Private reserves · Social-ecological
systems

27.1 Introduction

There are common claims of a “sixth mass extinction” happening today with the loss
of species currently at a rate at least 1000 times the background rate (Pimm et al.
2014; Pimm 2021). The direct causes include the loss of habitats, the introduction of
invasive exotic species, over-harvesting of biodiversity resources, and homogeniza-
tion of species in agriculture. The common factor of all these elements is that they are
mostly human-driven. The economic and social root causes behind biodiversity loss,
include demographic changes, overconsumption and production patterns, economic
growth, macro-economic policies and structures, social change, and development
([Link]
[Link]., [Link]
Worldwide, industrialized agriculture grows at the expense of natural areas such
as forests and savannas, for example through “invading” forests to grow soybeans in
Latin America and palm oil in Indonesia. Commercial agriculture generated nearly
70% of deforestation in Latin America between 2000 and 2010 (FAO 2015a; Martin
2020). Southeast Asia, the region in the world that has suffered the greatest rate of
deforestation, lost 30% of its forest cover over the last 40 years (Afelt et al. 2018).
Tropical cloud forests, one of the world’s most species- and endemism-rich terres-
trial ecosystems, are threatened by direct human pressures and climate change, with
substantial species losses worldwide, especially in readily accessible places (Karger
et al. 2021, Newcomer et al. 2022).
Human population as well as per-capita resource consumption are expected to
continue to rise, driving expanded urbanization, land use change, increased demand
for agricultural land, and deforestation related activities such as unsustainable
logging and mining. Industrial agriculture, focused on maximizing production
through monocultural cropping engineered to provide maximum yields, continues
to expand. However, it is dependent on advanced plant breeding, specialized (and
costly) machinery, and agrochemicals for fertilization and control of weeds and
pests. This approach contaminates and depletes agricultural soils, resulting often in
soil degradation (Bern 2018). The planet is losing ~0.3% per year of its capacity to
produce food due to soil degradation, currently jeopardizing the food and nutrition
security of one third of the world’s population (FAO 2015a).
Deforestation is linked to increased agricultural areas and poorly managed urban
growth (Afelt et al. 2018). As forest ecosystems and their habitats are lost, displaced
organisms along with their pathogens move from forests to anthropic environments
and from animals to humans, thus creating breeding grounds for the emergence of
new diseases. The COVID-19 pandemic is a call for attention to the fact that
production models prevalent today are contributing, in multiple ways, to this health
emergency (Martin 2020).
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 679

The “One Health” concept recognizes that human health is connected to animal
health and to the environment (Afelt et al. 2018). The modern incursion of humans
into the forest, involving changes to natural ecosystems, causes imbalances and
frequently leads to the appearance and spread of zoonotic diseases and even pan-
demics like COVID-19. The main routes of transmission of zoonosis differ widely
according to the specific underlying factors leading to emerging infectious diseases
(Loh et al. 2015). This knowledge can be used to develop more effective strategies
for controlling newly emerged diseases, taking into account the different underlying
pressures leading to land use change.
Anthropized rural environments are characterized by a wide diversity of land-
scapes comprising houses, barns, fields, orchards, and woodlands of differing
density, and can provide an acceptable habitat for a large range of small animal
species which can carry a variety of pathogens next to human dwellings (Afelt et al.
2018). Anthropized rural environments are also likely to increase human exposure
and risk of contracting insect-borne diseases, particularly as changing climate
conditions facilitate expanded ranges for disease carrying insects in the tropics.
Thus, it is crucial to educate and raise awareness about the risks associated with
anthropized environments (Afelt et al. 2018).
The advance of industrialized agriculture also threatens the traditional production
systems of indigenous peoples, local communities, and small and medium-sized
producers that underpin agriculture. These traditional systems and smaller producers
provide food for 70–80% of the world’s population, as well as the collection of
ancestrally associated knowledge, the preservation of genetic diversity and its
territorial management systems (FAO 2015b, 2019; Gadgil et al. 2021; Montagnini
and Berg 2019; Pimm 2021). Growing concern about the food and nutritional
insecurity of much of the world’s population, together with the impacts of the
COVID-19 pandemic, have led to the inclusion of this problem in the international
political agenda. To this end, it is important to promote alternative ways to produce
food, as have been presented in several chapters in this book, e.g. Calle et al.,
González et al., Levin (a), Montagnini et al., Montagnini and del Fierro, Montes-
Londoño et al., Painter et al., and Toensmeier. We must focus on more sustainable
agricultural techniques, including regenerative agriculture, to produce food without
depleting soil and damaging the climate (Bern 2018). We need a revaluation of
traditional cultivation practices which for hundreds of years have given sustainment
to local peoples (and continue to do so) with diverse, nutritious, and culturally
appropriate foods (Gadgil et al. 2021; Pimm 2021). Likewise, it is important to
promote the use of biological controls that can gradually replace the toxic chemicals
that are now used in industrial agricultural systems.
It is possible to reconcile agricultural production with biodiversity conserva-
tion, when nature is part of human-dominated landscapes, truly sharing space
by virtue of coexistence (Calle et al. 2022; Crespin and Simonetti 2019; Levin
2022a; Montagnini and del Fierro 2022; and other chapters mentioned
above). Sustainable agricultural management techniques geared toward harmo-
nizing ecosystem productivity and biodiversity conservation can contribute to mit-
igating or reversing detrimental effects of human impacts on landscapes while
680 F. Montagnini et al.

ensuring that agricultural productivity can meet the needs of human inhabitants for
generations (Montagnini and Berg 2019; Montagnini et al. 2022). To face these
issues, today’s environmental, social, economic, and political circumstances require
innovative responses that are appropriate to the emerging conditions (Berlyn 2021;
Mc Neeley 2021).
Biodiversity islands (BI)—ecological refuges where plants and animals thrive
without major interference from human activity—can contribute to the provision of
ecological, economic, and social benefits at the ecosystem, landscape, and global
levels. They can exist in an assortment of human-dominated landscapes (e.g.,
agricultural, wetland, urban) ranging in size from square meters to many square
kilometers. In the following sections we summarize examples presented in this book
of BI from throughout the world, discuss their challenges, and suggest viable
alternatives in their implementation and management at each scale.

27.2 Key Messages from This Book

27.2.1 Part I, Introduction

The first part of the book establishes the framework for understanding the complex-
ities of biodiversity islands and the variety of strategies that can be used to establish
them. The Introduction defines the term “biodiversity island” (BI) as a unique type of
ecological refuge whose design depends on its purpose, as well as on the spatial
distribution of reserves throughout the landscape, degree of landscape degradation,
species present, and location within the urban-rural spectrum. BI can contribute to
the ecological strength of a land area and make local agricultural areas more resilient,
for example by converting them into agroforestry systems (AFS) using various
agroecological strategies. Land use systems in a forestry matrix, as in many indig-
enous sacred sites and AFS, can also be part of BI.

27.2.2 Part II, Biodiversity Islands Establishment

and Management: Challenges and Alternatives

Design strategies for BI depend on landscape use within the matrix of habitat
fragmentation. Integrated landscape management (ILM), including sustainable agri-
culture, agroforestry and community led action, may provide a framework for
implementation of BI in complex landscape matrices. An experiment evaluating
edge effects by Arroyo-Rodríguez et al. (2022) shows that natural forest patches of
all sizes can benefit the ecosystem, and even small patches are valuable for conser-
vation of forest-specialist species. The next chapters discuss AFS strategies, includ-
ing regenerative agriculture, the integration of agricultural productivity and
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 681

biodiversity conservation, silvopastoral systems (SPS), incorporating reforestation

into livestock farms, and riparian buffers, which safeguard aquatic and riparian
environments from harmful agricultural practices. These techniques can lead to
agricultural systems containing significantly higher biodiversity than the surround-
ing area, showing that BI can exist in protected areas as well as in human-managed
landscapes. However, the right species should be reintroduced to a landscape during
restoration. The section concludes with a chapter by Eibl et al. (2022) stressing the
necessity of seed source certification to ensure the quality of genetic material and
species reintroduced to a landscape during restoration.

27.2.3 Part III, Biodiversity Islands Across the Globe: Case

Studies

The third part presents a total of 11 case studies where varied agroecological
strategies were applied in the formation or conservation of BI in human-dominated
landscapes. These case studies include (1) forest islands surrounded by flood-prone
savannah-dominated landscapes of Paraguayan Chaco utilized for livestock produc-
tion; (2) secondary forests that have persisted in the highly deforested landscape of
the Ucayali region of the Peruvian Amazon; (3) the integrated network of conserved
areas in Monteverde, Costa Rica which facilitate species movement across BI; (4) El
Hatico Nature Reserve, a model of restoration and utilization of agricultural practices
for sustainable production surrounded by a largely monoculture Colombian land-
scape; (5) Hacienda Pinzacuá, a restored, regenerative agriculture farm in the
surrounding treeless central Andes of Colombia; (6) small persisting biodiverse
land patches in the British countryside; and (7) the resilient islands of Las Rosas
in the Argentinean humid pampas which represent an opportunity to propose diverse
agroecosystems and develop local productive and economic strategies. Case studies
focused on conservation in urban landscapes include: (8) residential gardens of
Panama City; (9) the urban forests and peatlands of Ushuaia, Argentina; and
(10) the perennial garden of Paradise Lot in the U.S.A. (Toensmeier 2022). The
section closes with a case study (11) describing experiments to attain high genetic
diversity in BI in Misiones, Argentina (Niella et al. 2022).
Techniques are discussed for raising livestock in landscapes vulnerable to risks
from extreme weather or human-caused encroachment. Through extensive or semi-
extensive livestock production or use of live fences, livestock’s impact on the
landscapes were minimized to allow for the conservation of BI. There may be
benefits but also limitations in the use of BI for promoting species migration, leading
to efforts to connect islands and allow species to travel greater distances. Where BI
are surrounded by monoculture farms or treeless landscapes there may be ecological
and economic benefits that they can provide to those regions. Even in urban
landscapes BI can be havens for species and nodes in an interconnected network
of land patches that allow wildlife to travel and prosper.
682 F. Montagnini et al.

The variety of case studies from different types of landscapes from several
regions of the world reveals the role BI play in conserving local flora and fauna
that has been largely diminished by anthropogenic activities. In addition, these case
studies show how these BI are able to strengthen or increase the genetic diversity of a
human-dominated landscape, as shown by Niella et al. Furthermore, there are
human-centered benefits in BI, from providing a deeper cultural connection to nature
to supplying ecosystem services that make BI profitable to farmers and nearby
communities.

27.2.4 Part IV, Safeguarding the Environmental, Economic,

and Social Benefits of Biodiversity Islands, and Part V,
Conclusions

These final two parts of the book further detail the economic, social, political, and
cultural aspects of the establishment and persistence of BI in anthropogenic land-
scapes. A variety of strategies can be used to establish BI, including local laws and
legal tools, monetary aid and other financial resources, and local culture, particularly
of indigenous communities. All these strategies rely on community-led action to
contribute to the development and subsequent management of BI. Different com-
munity members’ perspectives towards their local ecosystem provide further insights
into the deciding factors or various motivations for conservation. For example, the
priorities, perspectives and use of a community forest by the people living around
Mayagüez, Puerto Rico, led them to be willing to protect the natural forest for its
services. In general, community forests are important for protection of lands,
reduction of deforestation, conservation of biodiversity, and carbon sequestration
while providing socioeconomic well-being to those living around them (Morales-
Nieves 2022).
The values that people assign to the forest contribute to its preservation as a BI
within a rural-urban landscape, even if biodiversity is not the prime benefit. For
example, sacred forests in Ethiopia have survived despite socioeconomic and polit-
ical pressures increasing deforestation in the adjacent land area. These forests
provide vital spaces for religious practice, as well as ecosystem services that
contribute valuable resources to the community, further reinforcing the relationship
between the community and the forest. Other case studies in Ecuador, Brazil and
India underscore the importance of outside partners working directly with local
communities when implementing conservation practices, as opposed to leaving
local voices and knowledge out of the design process. The role of the community
in conservation is further demonstrated through strategies for community-managed
AFS that allow land reform to occur in a more sustainable way, maximizing social
and economic returns while minimizing forest clearance in the cacao region of
Bahia, Brazil.
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 683

Biodiversity islands can provide a variety of ecosystem services through the food
and resources grown within them, with overharvesting prevented via proper regula-
tion, as shown in the permit-based harvest of ginseng in Appalachian Mixed
Mesophytic Forests in the U.S.A. The attitudes of farmers towards various agroeco-
logical approaches can determine what strategies farmers are willing to use in order
to continue to benefit from their land, as shown in the research focusing on natural
regeneration, reforestation and assisted natural regeneration as strategies to establish
and maintain silvopastoral farms in the Azuero Peninsula of Panama. Moreover there
may be tensions between the restrictions that government control may place on the
relationship between the BI and the local community, as shown in Northern Ethio-
pia, and the positive role local community members can play in establishing and
maintaining BI. Social-ecological systems need to be adapted to ensure that rural,
restoration-based BI in the region can continue to flourish alongside more pluralistic
and democratic political norms and institutions.
Biodiversity islands can provide valuable ecosystem services to the communities
or farmers who choose to establish them, helping to maintain or improve produc-
tivity while also conserving local flora and fauna. BI should always be tailored
specifically to the landscape, needs, and resources of the ecosystem to ensure they
are effective at protecting native species and their genetic diversity. To ensure BI are
enduring, however, the local community members must be allowed and encouraged
to contribute to its design and maintenance. This leads to help people develop a more
sustainable relationship with nature. In the remainder of this Conclusions chapter,
the lessons learned are presented along with alternatives and suggestions for
addressing some of the challenges to establishing or maintaining BI.

27.3 Barriers to Implementation of Biodiversity Islands

While BI offer a promising and practical option for conserving and restoring
biodiversity across human-dominated landscapes, they are not without challenges.
Several barriers to establishing and proliferating BI at scale have been recognized
within the chapters of this book. Some of these are conceptual in nature, relating to
theoretical pitfalls of this particular framework, while others have been gleaned from
the specific challenges encountered within the case studies examined.

27.3.1 Conceptual and Biophysical

A key challenge in designing and managing BI, articulated first in this book by
Montagnini et al., and later in several other chapters (Arroyo-Rodríguez et al. 2022,
Clavo Peralta et al. 2022, Esbach et al. 2022, Kirby 2022, Laino et al. 2022, Negret
et al. 2022, Santos-Gally and Boege 2022, to name a few), is the question of
priorities and tradeoffs, many of which are inherent in any conservation approach.
684 F. Montagnini et al.

Designing a BI for the protection of one target species may come at the expense of
other species with different habitat requirements. Prescribing a specific BI design
without considering its relative priority within the broader landscape or the value of
alternative land uses that may be at stake can similarly lead to misguided planning or
even undermine wider conservation objectives. BI must be designed and undertaken
with careful attention to both broad and local contexts and objectives.
As a land-sharing approach, BI managers may feel pressure or criticism from both
sides of the conservation-production spectrum: from one side that they do not do
enough to support biodiversity requirements, while from the other side that there is
too much conservation at the expense of production and human needs. A similar
balancing challenge that applies to BI is what Sigman (2022) calls in her chapter the
“Restoration Dilemma”. That is, the need for restoration efforts, such as BI, to be
highly site-specific and therefore resistant to scaling, while at the same time needing
to be scaled up because of the magnitude of the challenge and the need for
widespread adoption to realize their full potential benefits. These types of tradeoffs
must be reckoned with in endeavoring to establish BI.
When planning BI, design and expectations may not always match with reality.
For example with ancient woodland islands in the British countryside, as described
by Kirby 2022, the extent and pattern of patches as perceived by researchers—i.e.
what is mapped as woodland—may be smaller or larger than the actual patch size
used by the species in reality. This may in turn influence the success of the BI.
Certain BI designs may involve specific ecological and physical parameters that
prevent them from being replicated elsewhere. For instance, the urban residential
gardens in Panama City detailed in Negret et al. occupy a unique proximity to a
native forest patch that allowed for the gardens, though small, to function as
BI. Efforts to establish similar residential gardens in other urban settings may not
be as successful if they do not similarly benefit from a nearby native forest patch.
Designing and managing BI becomes even more difficult in the face of climate
change. Newcomer et al. point out that there are many unknowns as conditions alter
in the context of climate change, from whether biological corridors will or will not
support species migration, to how climate impacts will affect the region’s socioeco-
nomic conditions, all of which have implications for the long-term sustainability of
a BI.

27.3.2 Economic and Political

Beyond these conceptual and biophysical issues, real economic and political con-
straints also make BI implementation and management a challenge. As with other
ecosystem services, while many of the benefits of BI are enjoyed broadly, the costs
are private, potentially making it a less appealing option for land managers without
an additional source of funding to compensate for opportunity costs or otherwise
incentivize conservation over other land uses. The BI approach seeks to promote
integrated land management, but it still faces challenges, similar to other
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 685

conservation efforts, in competing with more productive, and often less biodiversity
friendly, alternatives that may offer faster and, at least initially, more tangible
returns.
Establishing BI may have high start-up and operational costs, especially where
they involve more labor intensive approaches or ones with specific objectives like
maintaining phylogenetic diversity, such as the projects described by Santos-Gally
and Boege in their chapter on native tree islands within neotropical silvopastoral
systems. The cumulative costs related to seed collection, germinating and
transplanting seedlings, establishing a nursery, and then managing the tree islands
for competition and protection from cattle were significant in the study and these
costs may be prohibitive to other producers, especially those with limited resources.
Some of these costs may be recouped in the long run from additional benefits from
ecosystem enhancements, but the startup capital required for the initial transition
may be a barrier.
Incentive programs themselves may require a minimum level of conservation
before becoming sustained, viable approaches. The case study of American ginseng
by Sheban is a good example of a promising regulatory conservation tool of permit-
based harvesting. However, it can only be effective in supporting the understory
ginseng in the BI if supplemented by simultaneous forest farming to sufficiently
reestablish populations. Maintaining a BI may require a set of additional mecha-
nisms to be successful, which may be a challenging configuration of approaches to
orchestrate.
At a more macro level, the establishment of BI is often in competition with more
intensive production systems that have the potential to influence prices or major
infrastructure decisions, which in turn create path dependencies that may lock in
these less sustainable alternatives. In Laino et al.’s case study of livestock production
systems in Paraguay, major roads were being planned in anticipation of higher
yielding commodity production in the region, further jeopardizing the prospect of
BI establishment and maintenance. Once these types of large-scale investments are
made that enable more intensive production systems, it can become even more
challenging for a producer to change to a different type of production system not
dependent on and often in conflict with such aggressive and intrusive incursions on
the landscape.
Conservation decisions are inherently political, and in certain contexts, the
political landscape may be even more complex than the biophysical ones. Sigman
draws attention to the dynamics in Ethiopia’s Tigray region where large scale
government-led restoration projects resulting in unique BI enjoyed support from a
political “monoculture”, i.e., single party leadership, that is now losing favor as the
country becomes politically pluralized. The success of the BI is complicated by the
complex communal labor realities and motivations that enabled these projects and
which may not be available going forward to support restoration, at least in their
current form, as the political landscape changes.
Similarly, although the Jupará Agroecological Movement in Brazil demonstrated
the feasibility of a unique agroecological model of land reform, its success may be
limited to only its local context unless broader historical, geographic and biophysical
686 F. Montagnini et al.

drivers are accounted for and integrated into supportive public policies (Painter et al.
2022). If the model is not supported by a wider enabling environment, it may
continue to only rely on limited and potentially unstable external funding and its
potential to scale up to support large-scale conservation of natural forest will be
constrained.

27.3.3 Social and Cultural

In addition to these economic and political barriers, cultural and social preferences
and norms may sometimes create resistance to BI. Farmers may have different
attitudes or preferences that hinder their willingness to try new approaches like BI,
including risk aversion, unfamiliarity, or pressure from existing social trends. In
Vásquez et al.’s chapter on farmer perceptions of forest restoration practices in
Panama, the main management strategy preferred among farmers was land clearing
for cattle. Farmers tended to prefer familiar options or ones over which they have
more control, such as tree-planting, which was a well-known practice in the region
due to its predominance among various restoration programs across the Azuero
peninsula.
A related barrier is the lack of necessary information and knowledge dissemina-
tion. This applies not only to knowledge of new and innovative approaches or
models of BI, but also of past practices. The latter was the emphasis in the chapter
by Clavo Peralta et al., where subsequent waves of migration from various parts of
the country resulted in varied land uses over time in the Ucayali region of the
Peruvian Amazon. While earlier local communities passed along knowledge of
different uses of the diverse species maintained in the remaining forest fragments,
more recent settlers did not have that same knowledge and therefore undervalued
conservation and opted for practices that were more reflective of their originating
regions. Several chapters in the book emphasize the importance of indigenous
knowledge in supporting BI (Levin 2022a, b; Esbach et al. 2022), but without proper
record and dissemination of that knowledge, its relevance risks being lost.
González et al. (2022) have noted in their chapter that the challenge is not simply
to influence individual farmers’ preferences, but rather to facilitate a deeper struc-
tural transformation that would replace the existing paradigm of competition and
economic profit with one centered on cooperation and relationships based on mutual
solidarity and concern. Switching from conventional systems to more agroecological
approaches requires “a completely different mindset” that surrenders control of
nature in favor of learning from nature, as Montes-Londoño et al. (2022) describe
in their chapter on a silvopastoral case study in the Colombian Andes.
Furthermore, the success of BI is often contingent on a variety of stakeholders
coming together around shared, or at least congruous, goals. Baez Schon et al.,
referring to the sacred natural sites that serve as BI in northern Ethiopia, point out the
need for support from different involved/interested groups (e.g. the church, nearby
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 687

communities, government) if they are going to be viable alternatives in the face of

increasing economic and sociocultural pressures. Newcomer et al.’s case study of the
Monteverde Reserve Complex in Costa Rica similarly describes the storied history
of the multiple local reserves and the various sets of actors who came to be involved
in their establishment and management, including Quaker settlers, community-based
organizations, NGOs, schools, international research scientists, farmers, and eco-
tourism organizations. While the unique constellation of actors in these case studies
created just the right context for establishing this BI, the reliance on diverse
stakeholders may pose a challenge in sustaining the BI into the future, especially
if priorities and needs shift in the face of new pressures.
Many of these barriers presented throughout the book are not unique to BI, but are
in fact issues that have challenged various approaches to conservation and are
present in any undertaking that seeks to bring together sometimes divergent ecolog-
ical, social, economic, and political goals and dynamics. Nonetheless, these barriers
are not insurmountable. As many of the case studies have shown, given the right
incentives, support and enabling policies, barriers can be overcome and BI can be
scaled up to support conservation and production.

27.4 Viable Alternatives and Opportunities

for Establishment of Biodiversity Islands
27.4.1 Community-Based Opportunities

Biodiversity islands designed in partnership with local communities or indigenous

groups can show the greatest potential for long-term success (Reyes-García et al.
2019). Social factors have been identified by restoration practitioners as having a far
greater influence on the longevity of restoration projects than ecological factors, with
multi-stakeholder engagement being the greatest challenge (Nerfa et al. 2021). For
this reason, improving communication between policy makers, practitioners, and
local communities is critical for improving BI implementation. Esbach et al. show
that multi-stakeholder participation can be facilitated by intentionally integrating
local partnerships and participatory research into conservation and development
strategies. Participatory research can be fine-tuned to meet local needs while
empowering communities to play active roles in developing solutions, demonstrat-
ing that the goals of local actors and BI are compatible.
Community based natural resource management can incentivize local communi-
ties to sustainably manage resources for their long-term availability, serving as an
alternative to degradative cycles of exploitation. While community-based forestry
generally has positive environmental and income related outcomes, it can sometimes
inadvertently restrict the rights of communities to access forest resources (Hajjar
et al. 2021). Structured engagement with local communities can help design projects
688 F. Montagnini et al.

that are more biodiverse, and also meet the needs of their multiple users (Dumont
et al. 2019).
Forest and Farmer Producer Organizations such as community forestry user
groups or producer cooperatives can help generate local support for sustainably
managed BI. González et al. suggest that producer and consumer, or “prosumer”,
cooperatives, working across stakeholder levels, can help localized biodiverse agri-
food systems. Community actors (both producers and consumers, among others) are
the key to building locally managed agroecological systems. Levin (b) notes that
cooperative business structures can also improve social outcomes such as farm-
worker health and empowerment, core tenets of regenerative agriculture.
Education and capacity building are needed to further the implementation of BI
globally. The chapter by Vásquez et al. (2022) shows that many rural farmers in
Panama do not see assisted natural regeneration (ANR) as a restoration practice and
therefore training farmers on the benefits of ANR could help scale up the restoration
of degraded lands. Demonstration farms such as Hacienda Pinzacuá and El Hatico in
Colombia, and Paradise Lot in Massachusetts, U.S.A. serve as examples of height-
ened productivity and ecological functioning, which can inspire other farms to adopt
similar practices. Urban BI are also educational centers which can connect people to
land and local ecosystems, from tropical Panama to the temperate Northeastern
United States and Ushuaia, Argentina (Soler et al. 2022).
When working with local stakeholders, it is important to understand a
community’s diverse motivations for engaging in restoration and conservation, as
demonstrated in different chapters of this book. Levin (b) notes that community-led
action can be motivated by ethical, philosophical, scientific, cultural, and economic
values. Morales-Nieves reveals that air quality and recreation were the highest
priorities in an urban community forest in Puerto Rico. In Ethiopia, church forests
are managed for their spiritual use (Baez Shon et al. 2022) but, as Sigman points out,
the political “monoculture” may present a risk to restoration efforts sponsored by the
political party in power. In Monteverde, Costa Rica, the local conservation move-
ment is made more resilient by its many different motivations driving conservation
(Newcomer et al. 2022). Painter et al.’s work in the Atlantic forest of Brazil shows
that outside support can help BI meet community needs for production, conserva-
tion, and socioeconomic well-being when coupled with understandings of the
motivations of community members, nuances of land tenure, and appropriate
enabling conditions.

27.4.2 Current Restoration/Conservation Efforts Favoring

Biodiversity Islands

In spite of financial and other constraints faced by restoration and conservation

projects, many examples have recently been brought to our attention showing
evidence of a current and increasing trend of BI implementation. Several of these
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 689

efforts are nurtured along by non-government organizations (NGOs) with limited

funding and by private individuals and entities whose sole motivation relies on their
own vocation and desire to preserve nature. A recent trend for the resurgence of
agroecology strategies and experiences, as mentioned by Levin (a, b) and in the
chapters by González et al. and by Painter et al., contributes to harmonizing
production with conservation and promotes the spread of BI. In this subsection, a
few examples are presented which illustrate that the implementation of new BI is
already happening and gaining momentum. Further support and guidelines to assist
their management and persistence are needed.

[Link] Examples of Recently Launched NGOs and Private

Restoration/Conservation Projects

Several business enterprises located in critical areas in need of solid conservation

efforts, such as the Amazon, place a strong emphasis on biodiversity conservation
while promoting restoration and sustainable agriculture practices, including agro-
forestry, to improve livelihoods of local people. For example, in the Ecuadorian
Amazon, “Aliados” was formed in 2018 to build resilient community businesses
based on supporting biodiversity in the Andes and the Amazon and to connect them
to markets across the globe. Aliados restores and conserves landscapes in the
Amazon rainforest along with local communities and in partnership with NGOs,
private sectors and other key partners through promoting agroforestry and strength-
ening value chains ([Link] Their projects are located in the
upper Napo Watershed in the Andes-Amazon foothills, in two critical ecological
corridors at the crossroads of four National Parks composed of 150,000 hectares of
megadiverse cloud and rain forest. They have designed their own Theory of Change,
aiming towards reaching impact at the landscape level by combining their experi-
ence and network of community, corporate and philanthropic partners to build a
regenerative agroforestry and restoration landscape for a fundamentally new model
of doing business in the Amazon. Their actions contribute to the creation and
maintenance of BI financed by their enterprise profits as well as from charitable
donations and international funding.
Several recent efforts and initiatives worldwide to advance conservation have
surged with the support of local conservation organizations and people. For exam-
ple, the NRDC (Natural Resources Defense Council, [Link]
founded in 1970 by a group of law students and attorneys at the forefront of the
environmental movement, is actively working on the 30x30 initiative: to prevent
mass extinctions and bolster resilience to climate change, we must protect at least
30 percent of our lands, rivers, lakes, and wetlands by 2030. The 30x30 targets
“. . .will help maintain global biodiversity, preserve the integrity of ecosystems on
which we all depend, provide safe havens to help wildlife adapt to climate change,
and sustain natural systems that store carbon, such as forests, mangroves, seagrasses,
wetlands, and grasslands.” NRDC is focused on achieving these goals while
protecting diverse habitats, improving access for all people, and identifying and
690 F. Montagnini et al.

managing these areas in collaboration with indigenous communities ([Link]

[Link]/30x30-nrdcs-commitment-protect-nature-and-life-earth). Their multiface-
ted strategy involves key areas of work in the United States and abroad. The Sierra
Club and other international conservation organizations are joining this effort as well
(Brett Levin, personal communication, May 2021).
A number of other recent private conservation projects have some alignment with
the BI concept that we are promoting through this book. For example, Homegrown
National Park is a grassroots organization in the U.S.A. calling for action to restore
biodiversity and ecosystem function by planting native species and creating new
ecological networks. Their mission is to restore biodiversity and ecosystem function,
stemming from the realization that every human being on this planet needs diverse,
highly productive ecosystems to survive. They are catalyzing a collective effort of
individual homeowners, land managers, farmers, and “anyone with some soil to
plant in. . .to start a new habitat by planting native plants and removing most
invasive plants,” claiming this represents the largest cooperative conservation pro-
ject ever conceived or attempted in the country. Their goal is to reach 20 million
acres (8,093,713 hectares) of native plantings in the U.S.A., an area that represents
approximately half of the green lawns of privately-owned properties in the country
([Link]

27.4.3 Private Protected Areas (PPAs)

Private Protected Areas (PPAs) are areas of land or water that fulfill the conditions to
be considered Natural Protected Areas (NPA) by the IUCN and that are managed by
private governance (Mitchell et al. 2018, [Link] All over the world
there are families, communities and organizations that have decided to do something
to change the current loss of natural areas and biodiversity and have begun to protect
watershed headers and habitats of threatened species, restore degraded areas,
develop education strategies, and promote positive contact with nature, among
other initiatives that transform the way we relate to nature. Some of them have
used tools of voluntary conservation, such as PPAs, with a view to getting greater
formality and legal security to their ventures (Monteferri 2019). These areas hold BI
that need management guidelines to ensure their efficacy and persistence.
In recent decades, the voluntary conservation movement on private lands has
grown in different parts of the world, contributing to address the loss of biodiversity
(Roldán et al. 2010). For example, a study in South Africa showed that, if PPAs were
considered within the protected area system, results on estimations of species
diversity would almost triple (Gallo et al. 2009). The level of consolidation and
growth of voluntary conservation movements vary depending on each region. The
private conservation movement in Latin America has been growing in recent years.
At the 2018 Private Conservation Areas Congress for Latin America, this increase
was made evident with a total of 4152 protected areas covering 4,618,042 hectares
(Monteferri 2019). In Peru, there are a total of 1.5 million hectares of PPA in the
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 691

whole country, including different forms of private conservation: PPAs as well as

conservation and ecotourism concessions. Around 70% of these areas are located in
the Peruvian Amazon, combining different legal tools for private conservation
(Carolina Butrich, NGO “Conservamos por Naturaleza”, personal communication,
April 2021, [Link]
Owners and managers of PPAs often suffer from financial, logistical and other
difficulties which pose a threat to their conservation efforts, as manifested by Víctor
Zambrano, personal communication, April 2021. He owns a PPA in the Peruvian
Amazon and works in the “Comité de Gestión de la Reserva Nacional Tambopata”
(Management Committee of the Tambopata National Reserve), located in Madre de
Dios, Peru. For Víctor, the main challenge is for landowners to find ways to generate
long-term value without having to decrease land productivity and ecosystem
services.
Voluntary conservation plays a key role as it creates a culture of conservation and
makes it more accessible to all citizens. When managing land with conservation as a
major purpose, agroforestry, agrobiodiversity and silvopasture gain space in the face
of monocultures; permaculture and agricultural biodynamics become more impor-
tant than chemical fertilizers; organic farming eliminates pesticide use that is gener-
ating drastic decreases in insects in the world (Monteferri 2019). Owners of PPAs
explore ways of managing from a perspective of custodians, seeing nature as an ally
rather than an obstacle.
Given the multitude of complex social and ecological challenges, there is an
urgency to take action, and conservation at local and regional levels takes on unique
importance (Morton 2013). Leadership at the local level will play a key role in the
decades to come, as interconnectedness facilitates the replication of local initiatives,
with communities becoming increasingly more informed and eager to see change.
Voluntary conservation allows volunteers, companies, farmers, families, schools and
universities to be recognized, participate and collaborate in preservation, and facil-
itates respect of biodiversity at all scales. To ensure their persistence and successful
management, tax benefits could be provided to PPAs along with some measure of
compensation from other commercial interests that benefit from the carbon seques-
tration potential and other ecosystem services they provide.

[Link] Biodiversity Islands in Indigenous Territories

Land use systems made up of complex assemblages embedded in a forestry matrix,

as is the case in many traditional indigenous sacred sites, agroforestry systems
(Baez Schon et al. 2022; Gadgil et al. 2021), and indigenous territories can be
considered BI in themselves, as explained in the Introductory chapter of this book
(Montagnini et al. 2022). Known also as “Islands of Nature,” they can be many
square kilometers in size, generally use native forest species for sustainable food
production and biodiversity, and are integrated with the natural forest. Several
examples located in indigenous territories worldwide were well documented in a
recent report by The Intergovernmental Science-Policy Platform on Biodiversity and
692 F. Montagnini et al.

Ecosystem Services (IPBES) (United Nations IPBES 2019). The protection of these
Islands of Nature is ensured as long as the indigenous peoples’ territories and rights
to use the land are respected, which often conflicts with development goals of other
sectors. For example, protected territories inhabited by indigenous peoples in the
Peruvian Amazon are threatened by road development, oil extraction, and other
industries (Joseph Zárate, personal communication, April 2021, Zárate 2021). As oil
exploration moves from industrialized countries to other locations such as in Peru
and Ecuador, indigenous territories and their biodiversity face increasing threats.
Indigenous movements and their supporters, including the legal system in each
affected country/territory, however, are actively seeking justice and winning court
battles. For example, the Union of People Affected by Texaco (UDAPT, www.
[Link]) in Ecuador won a legal battle against the oil company, found to be guilty
of egregious pollution and irresponsibility in the Ecuadorian rainforest. Ten years
later, they are still fighting for the settlement that would benefit the tens of thousands
of indigenous people who have been impacted (Julio Prieto, personal communica-
tion, May 2021). The case is explained in detail in a recent Forces for Nature podcast
([Link]
ecuadorian-rainforest/).
In Ecuador, the Confederation of Indigenous Nationalities of the Ecuadorian
Amazon (La Confederación de las Nacionalidades Indígenas de la Amazonia
Ecuatoriana), or CONFENIAE, is the regional organization of indigenous peoples
in the Oriente region of the Ecuadorian Amazon. Nine indigenous peoples living in
the region—Quichua, Shuar, Achuar, Huaorani, Siona, Secoya, Shiwiar, Záparo,
and Cofán—are represented politically by the Confederation ([Link]
[Link]/). CONFENIAE is one of three major regional groupings that constitute the
Confederation of Indigenous Nationalities of Ecuador (CONAIE). It is also part of
the Amazon Basin indigenous organization, COICA ([Link] While
these organizations get some technical and financial assistance from several NGOs
and other groups, and the indigenous people appreciate their help, they often would
prefer greater autonomy in their decision making (Efren Nango, CONFENIAE,
personal communication, February 2021). The indigenous peoples and their allies
are challenged to not allow external forces to divide and disrespect their organization
as they advance efforts to exert their rights to the land and biodiversity, including the
embedded BI within their territories.

[Link] Examples of Ongoing Agroecological Initiatives Supporting

Biodiversity Islands

New ways of practicing agroecology to harmonize food production with conserva-

tion often lead to the spread of BI, as described in both chapters by Levin, and by
González et al. (2022). These practices and experiences carried along by different
groups of people are spreading, managing to overcome financial, technical training
and other constraints. For example, recent presentations on agroecology and biodi-
versity by local biologists and agronomists were offered to local farmers who are
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 693

transitioning to agroecological production systems, in an event taking place at the

agroecological farm “La Dorita” located in Basavilbaso, Entre Ríos province,
Argentina (Libertario González, personal communication, May 2021). These are
small BI which are delineated and protected to fulfill local needs.
Individual farmers worldwide who practice agroecology in a variety of ways,
including agroforestry, contribute to the creation of BI through their individual
efforts and often using their own financial resources, as described in the chapters
mentioned above as well as in other chapters in this volume from Calle et al., Esbach
et al., Montes-Londoño et al., Painter et al., and Toensmeier. An agricultural
producer in the state of Zulia, Venezuela who manages a 1000-hectare farm includ-
ing silvopastoral systems with buffalo, many saman (Samanea saman) trees that
produce feed for cattle, secondary forests of different ages, as well as areas with bam-
boo and oil palm, was seeking our advice on how to design and manage BI after
watching a program on CNN1 (Wilmer Morán, personal communication, April
2021). Thus, our book hopefully will provide information to help individual efforts
like this one to design, protect and manage BI.

[Link] Recent Government Policies Promoting Agroecological

Practices

In some cases agroecological transitions from conventional agriculture are already

being supported by government initiatives that go along with local agroecology
movements, even in countries like Argentina and the U.S.A., where industrial
agriculture for large scale production of grains is the norm. These initiatives are a
response to the environmental issues created by monoculture agriculture as men-
tioned earlier in this chapter.
In Argentina, where export taxes on soybean production comprise a large portion
of the Gross Domestic Product, the Argentine Society of Agroecology (SAAE) was
created in 2018 and has been pivotal in the consolidation of agroecology in the
country ([Link]
tina-de-Agroecolog%C3%ADa-126879274899065/). Brazil and Argentina are the
only two Latin American countries that have constituted a National Society of
Agroecology. The First Argentine Congress of Agroecology was held in 2019
([Link] Training events to lead the transition to agro-
ecology aimed at producers and technical personnel have been organized by the
National Institute of Agricultural Technology, INTA (INTA Procadis, [Link]
[Link]/acerca-de-procadis). In the Buenos Aires province, by April 2021 there were
350 farmers registered as agroecological, comprising a total of about 40,000 hectares

1
Interview, CNN en Español, from CNN studio in Atlanta, GA, U.S.A., aired March 27–28, 2021,
“Pandemia, Biodiversidad y Futuro” (“Pandemic, Biodiversity and the Future”) in GloboEconomía,
a CNN program with José Antonio Montenegro, a Warner Media production, [Link]
com/watch?v¼f68tzy65zq4, or search YouTube:“Cómo afectó el ser humano a la biodiversidad”
694 F. Montagnini et al.

distributed all over the province, including large and small producers (Germán
Lanzer, Director of Agroecology for the Buenos Aires province, Transition to
Agroecology course offered by INTA Procadis, April 2021).
In 2020, the Ministry of Agriculture, Fisheries and Livestock of Argentina
created the National Direction of Agroecology, within the Secretariat of Food,
Bioeconomy and Regional Development ([Link]
ar/2020/08/¼). Its primary objective is “to intervene in the design and implementa-
tion of policies, programs and projects that promote intensive and extensive
agroecologically-based primary production at all levels.” Interacting with producers,
agricultural organizations and municipal and provincial governments, they are
executing a Strategic Productive Transition Plan for agroecological implementation,
providing technical assistance, and establishing credits or tax tools for its promotion.
Also in Argentina, the National Network of Municipalities and Communities that
promote Agroecology (Renama) works with some thirty municipalities in the differ-
ent productive regions of the country ([Link]). Renama has incorporated
about 170 producers so far, with an estimated 86,000 hectares under agroecological
management. “This is not an alternative practice. On the contrary, it is the agriculture
of the coming years” (Eduardo Cerdá, Director of the Division of Agroecology, and
also president of Renama, 2020 interview with InterNos, local media from Buenos
Aires, Argentina). In the same interview he added: “There’s a strong paradigm shift.
The current production model was important at the time, when new fertilizers,
herbicides and strong technological innovations resulted in good production
increases. But the continuity of this model has brought us many problems in terms
of soil losses (according to INTA, more than 50% of the country’s soil organic matter
has been lost) and increases in herbicide-resistant weeds. This has brought along the
use of more agrochemicals to control them, which in turn has increased costs above
profits, leaving a lot of producers out of the game. In addition, producers realize what
it means to be in contact with these substances which were claimed not to generate
acute poisoning. But it turns out that the problem was their chronic toxicity, that is,
toxicity generated from long exposure to the products. This is a needed change of
paradigm from the perspective of improving nutrition and increasing resilience in the
face of the current pandemic” ([Link]
modelo-de-produccion-es-drogodependiente/).
Dissemination of agroecological practices and regenerative agriculture in all their
forms that are environmentally friendly, harmonizing productivity and environmen-
tal goods and services, is driving the promotion and persistence of BI in human
dominated environments. There is already enough traditional and scientific knowl-
edge among farmers and international and local institutions as we have described in
several chapters of this book (e.g., Baez Schon et al., Clavo Peralta et al., Levin (a),
Montagnini et al., Montagnini and del Fierro). These practices need to be promoted
using incentives, especially in the early years of implementation, until adequate
production levels are reached. These incentives can take many forms such as soft
loans, materials, and tools, and should include education and technical assistance at
all levels.
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 695

In the United States, the Department of Agriculture (USDA) has long provided
technical and financial assistance and other resources to farmers and ranchers via
agencies including the Natural Resources Conservation Service and the National
Agroforestry Center. These agencies and their programs help landowners implement
practices that conserve and restore the natural resources on their land and in their
operations. This assistance is provided through a variety of different incentive
programs including the Environmental Quality Incentives Program (EQIP), which
offers financial resources and one-on-one help to plan and implement improvements
that address natural resource concerns. The Conservation Reserve Program provides
an annual payment for removing environmentally sensitive land from agricultural
production and planting species that will improve environmental health and quality
([Link]
tion-reserve-program/; [Link]
programs/financial/eqip/).
With the United States re-entering the Paris Agreement of the United Nations
Framework Convention on Climate Change in 2021, renewed attention on its
international climate mitigation commitments may help bolster these working
lands conservation programs in recognition of the climate mitigation opportunities
they could potentially provide. Via executive order, the Biden-Harris administration
recently directed the USDA to develop a strategy to encourage the voluntary
adoption of climate-smart agricultural and forestry practices that result in additional,
measurable, and verifiable carbon reductions and sequestration. While the contours
of this strategy are still being developed, the strategy will be designed to accommo-
date all farmers, and will seek to integrate climate-smart practices into existing
programs or create new programs that strengthen markets for agriculture and forestry
products generated through climate-smart agriculture and forestry practices, includ-
ing via support of private voluntary carbon markets ([Link]
default/files/documents/climate-smart-ag-forestry-strategy-90-day-progress-report.
pdf). Agroforestry practices, including windbreaks, buffers, alley cropping,
silvopasture and multi-story cropping, as well as other agroecological practices
such as cover crops and no-till, are among the practices being considered for
inclusion in a potential new climate-smart agriculture and forestry program, which
may facilitate the establishment of BI on working lands in the U.S.A.

27.4.4 Valuing and Financing Biodiversity Conservation

A large disparity exists between the resources needed and the available funds for
financing conservation, and this has created a dire situation for the future of
biodiversity conservation, as discussed in a recent interview published by the Nature
Conservancy: it has been estimated that by 2030 the world needs as much as an
additional US$598 billion to $824 billion annually to close the financing gap
(Solberg 2021). Financing biodiversity on its own is a difficult task, considering
that the effects and payoffs of such conservation are not always tangible, and that
696 F. Montagnini et al.

conservation of biodiversity is a multidimensional global issue. Financing biodiver-

sity can become increasingly easier, however, when tangible and valued resources
such as carbon or water are incorporated into conservation strategies (Sheban 2022).
The first step towards building these tangible and effective conservation strategies
is the proper assessment and valuation of ecosystem services. Proper valuation is
essential in developing new initiatives, and programs such as Payments for Ecosys-
tem Services (PES) may be efficient in encouraging future biodiversity conservation
through compensation. PES programs have considerably increased in recent years,
although biodiversity-focused PES programs have remained limited. Compensatory
mitigation banking is also growing, while voluntary biodiversity offsets are a recent
policy development and have yet to experience large amounts of growth (Salzman
et al. 2018). Valuation systems that bundle multiple services could better encourage
sustainable land uses to farmers and protect biodiversity, even if the PES is only
provided for a single service such as water or carbon (Montagnini and Finney 2011).
Biodiversity objectives might also be incorporated into future ecological action
programs.
Another opportunity for funding conservation is through taxing certain activities
that are detrimental to worldwide ecosystems, such as imposing emissions taxes on
airline travel to discourage fossil fuel consumption or implementing a variety of
“polluter pays” taxation programs. These funds could in turn be used to finance
conservation initiatives such as the implementation and maintenance of BI. If the
financial penalties were established at sufficient levels, perhaps industries would
self-regulate to address environmental degradation and avoid paying such taxes.
Such an approach might accelerate progress for sustainability by directing pollution
tax funding toward implementing BI and carbon sequestration projects. As
Newcomer et al. described in their chapter in this volume, such an approach has
seen moderate success in Costa Rica with gas taxes being used to fund the national
PES program.
As we have described, the effort to integrate biodiversity into the market is
already growing, with investments worldwide coupled with positive environmental
outcomes gaining in popularity. Agricultural and consumer markets are increasingly
shifting products and supply chains to align with conservation goals. Because BI
establishment generally happens on fragmented landscapes, funding and acquiring
these small patches of land through government action and policy could satisfy both
land development and biodiversity conservation goals in the future, and may be
more effective than the approach of designating large amounts of land solely for
conservation initiatives. Preserving numerous small areas of land for BI conservation
rather than rendering large swaths of land economically unusable seems mutually
beneficial for governments and for landowners and can contribute to conservation
goals as well.
Voluntary conservation actions from companies, farmers, families, schools and
universities should be financially recognized as they participate and collaborate in
creating BI at all scales. Financial incentives to ensure successful management and
persistence of privately established BI may take many forms such as loans, soft
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 697

credits, and the provision of materials and tools. In particular, tax benefits could be
provided to Private Protected Areas as well as to landowners who protect sensitive
areas within their properties, for example, riparian forests that contribute to land-
scape connectivity and other ecosystem services as shown by Giraldo et al. (2022).
This type of incentive can be calculated based on the area covered by the BI, or by
valuing the ecological importance of a particular area for conservation purposes.
Other compensation could be derived from other commercial interests such as
companies or enterprises that benefit from the carbon sequestration potential and
other ecosystem services provided by BI.

27.4.5 The Framework: Current Trends in Biodiversity

Conservation Aligning with the Biodiversity Islands
Concept

There have been several trends and useful perspectives on how protected area design
issues could be addressed more effectively (McNeely 2021). Bengtsson et al. (2003)
highlighted some of the limitations of protected areas, which covered just over 11%
of the land at that time. While the landscape perspective and the integration of
resilience theory and biodiversity conservation is now much more reflected in
biodiversity policy (e.g., see IPBES 2019), improvements in conservation practice
and landscape management have recently been quite small (Bengtsson et al. 2021).
The CBD’s Strategic Plan for Biodiversity 2011–2020 was highly relevant to
protected areas as a major conservation tool (Pimm 2021; Sayer et al. 2021). Its
target 11 called for establishing at least 17% of terrestrial and inland water biomes as
protected areas, along with 10% of coastal and marine biomes. These were to be
effectively and equitably managed, ecologically representative, and well connected
as parts of systems of effective area-based conservation measures that are integrated
into wider landscapes and seascapes.
At the time, however, protected areas were considered too static when they need
to be more dynamic (Bengtsson et al. 2003, 2021; Mc Neely 2021). A dynamic
landscape approach that mimics natural disturbance regimes could include, for
example, successional lands that are recovering from over-exploitation. These
resemble the territories managed by indigenous peoples such as the shifting culti-
vation practices described by Gadgil et al. (2021). Indigenous territories or parts of
them where sustainable multistrata agroforestry and forest systems are practiced are
considered BI as defined in this book. The importance of landscape connectivity is a
current hot subject in the conservation mainstream (McNeely 2021), with detailed
guidelines prepared by an IUCN international team (Hilty et al. 2020). Landscape
connectivity is discussed in this book in relation to the effectiveness of BI.
The Aichi targets set in 2010 have led to the expansion of systems of national
parks and other categories of protected areas, and over 90 of the signatory countries
have attained the 17% target. However, some governments have established
698 F. Montagnini et al.

protected areas in degraded and marginal areas that have limited conservation value,
often because those protected areas simply were not needed for anything else. There
is still not enough evidence of the effectiveness of protected landscapes in delivering
biodiversity outcomes (Sayer 2021). The value of landscape approaches to conserve
biodiversity through management of the broader landscape within which conven-
tional protected areas are located is expected to increase (Sayer et al. 2021). This
concept aligns with the integrated landscape management approaches suggested in
this book.
The fifteenth meeting of the Convention on Biological Diversity (CBD) Confer-
ence of the Parties (COP), Kunming, China, October 2021 has established seven
thematic programs of work corresponding to some of the major biomes on the planet.
Each program establishes a vision and basic principles to guide future work. They
also set out key issues for consideration, identify potential outputs, and suggest a
timetable and means for achieving them. Implementation of these programs depends
on contributions from signatory parties, the Secretariat, and relevant intergovern-
mental and other organizations ([Link] CBD COP-15 is likely to
adopt relatively simple, aspirational, and politically attractive targets for biodiversity
conservation (Sayer et al. 2021). Civil society is likely to prefer enhanced conser-
vation measures, but people are often reluctant to accept actions that restrict their
material wellbeing, thus conservation strategies need to be scientifically sound and
aligned with the cultures and economies of local societies such as is described for BI
in this book. Research, training, and capacity building are needed to manage pro-
grams of biodiversity conservation (Sayer et al. 2021). This also pertains to BI, and
this book has set the basis for implementing and successfully managing BI using an
inclusive, landscape oriented and integrated approach.
It has been broadly recognized that solutions to the problems of biodiversity
conservation come down to working with people, their lives, aspirations, fears, and
social complexity. Although people and nature will not always peacefully coexist,
rigorous science and intelligent technology can help (Berlyn 2021; Pimm 2021).
Evidence from chapters of this book shows that BI may constitute such a solution
when properly designed and managed within the social milieu where they are
embedded.

27.4.6 The Context: Biodiversity Islands in Changing

Environments

Climate change is increasingly a threat to both natural and human systems. The
success of BI as effective conservation solutions is vulnerable to changing climate
conditions, including via impacts on habitat requirements and species ranges, as well
as the cascading effects of shifting socioeconomic demands. At the same time,
however, BI have an important role to play in contributing to climate change
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 699

solutions. Land use, including agriculture and forestry, accounts for an estimated
23% of total anthropogenic greenhouse gas emissions (Shukla et al. 2019).
Reducing deforestation and introducing, or reintroducing, trees and herbaceous
cover into working lands not only benefit biodiversity, but can have outsized impacts
on greenhouse gas emissions via conserved carbon stocks, reduced emissions, and
increased carbon sequestration. These so-called natural climate solutions, which
include conservation, restoration and improved agricultural practices, can contribute
up to one-third of the cost-effective climate mitigation we need with the potential to
remove as much as 23.8 petagrams of CO2 equivalent (Griscom et al. 2017).
Co-benefits of conservation, including the livelihood support they provide and
ecosystem services, such as watershed regulation, can also help increase the resil-
ience of communities impacted by climate change (Griscom et al. 2017). Given the
magnitude and urgency of this challenge, curbing climate change and limiting global
temperature rise to below the 2  C degree threshold set by the Paris Agreement will
require ambitious and radical changes in our production systems. As an integrated,
flexible conservation approach with great potential, BI can be an important piece of
the nature-based solutions necessary to address the climate crisis.

27.5 Conclusions: Biodiversity Islands in Action

The lessons learned from the chapters of this book form a collection of experiences
showing positive and promising results of how BI can be implemented and managed.
Through a series of informative case studies and detailed explanations of the
intricacies of BI creation, the contributions to this volume provide a comprehensive
context for the impediments and opportunities of BI implementation amidst various
societal and environmental factors. Several chapters in the book discuss the chal-
lenges that BI development face, including a lack of conceptual understanding of BI
function, incompatibility with current local societal practices or government prior-
ities, and competing economic/productive land use.
Many practical alternatives are presented that address these challenges through
the creation of BI. Several of these opportunities relate to communities’ motivations,
which is why addressing social issues and improving communication are critical for
establishing effective and lasting BI. The book presents a variety of examples of BI
created in differing circumstances, from NGO action to AFS implementation, where
involvement of local and indigenous community members plays a vital role.
BI present an opportunity for sustainable, dynamic, productive conservation,
which is why they are becoming increasingly desirable in the global conservation
movement. As societies look for alternatives to maintain economic prosperity,
provide culturally-important community spaces, and conserve local ecosystem bio-
diversity, BI are sure to become more widely used, which underscores the impor-
tance of the information described in this book as a tool for planning and
implementing BI.
700 F. Montagnini et al.

References

Afelt A, Frutos R, Devaux C (2018) Bats, coronaviruses, and deforestation: toward the emergence
of novel infectious diseases? Front Microbiol 9:Article 702. [Link]
00702
Arroyo-Rodríguez V, Arasa-Gisbert R, Arce-Peña N, Cervantes-López MJ, Cudney-Valenzuela SJ,
Galán-Acedo C, Hernández-Ruedas MA, San-José M, Fahrig, L (2022) The importance of small
rainforest patches for biodiversity conservation: a multi-taxonomic assessment in the Lacandona
region, Mexico. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Series: Topics in Biodiversity and Conservation. Springer,
Cham, pp 41–60
Baez Schon M, Woods CL, Cardelús CL (2022) Sacred Church forests in northern Ethiopia:
biodiversity and cultural islands. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in Biodiversity and Conserva-
tion, Springer, Cham, pp 531–549
Bengtsson J, Angelstam P, Elmqvist T, Emanuelsson U, Folke C, Hise M, Moberg F, Nyström M
(2003) Reserves, resilience and dynamic landscapes. Ambio 32:389–396. [Link]
1579/0044-7447-32.6.389
Bengtsson J, Angelstam P, Elmqvist T, Emanuelsson U, Folke C, Ihse M, Moberg F, Nyström M
(2021) Reserves, resilience and dynamic landscapes 20 years later. 50th anniversary collection:
biodiversity conservation. Ambio 50:962–966. [Link]
Berlyn GP (2021) Some thoughts on biodiversity and sustainability. J Environ Sci Public Health
5(3):371–373. [Link]
Bern L (2018) Den metabola pandemin: det största hotet mot en hållbar framtid. Recito Publishers,
Sweden, 481 pp [Link]
storsta-hotet-mot-en-hallbar-framtid/
Calle Z, Molina CCH, Molina DCH, Molina DEJ, Molina EJJ, Murgueitio B, Murgueitio A,
Murgueitio E (2022) A highly productive biodiversity island within a monoculture landscape:
El Hatico nature reserve (Valle del Cauca, Colombia). In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in Bio-
diversity and Conservation. Springer, Cham, pp 279–304
Clavo Peralta ZM, Vela Alvarado JW, Alvez-Valles CM (2022) Biodiversity islands and dominant
species in agricultural landscapes of the Western South Amazon, Peru. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in Biodiversity and Conservation. Springer, Cham, pp 207–235
Crespin SJ, Simonetti JA (2019) Reconciling farming and wild nature: integrating human–wildlife
coexistence into the land-sharing and land-sparing framework. Ambio 48:131–138. [Link]
org/10.1007/s13280-018-1059-2
Dumont ES, Bonhomme S, Pagella TF, Sinclair FL (2019) Structured stakeholder engagement
leads to development of more diverse and inclusive agroforestry options. Exp Agric 55(S1):
252–274. [Link]
Eibl B, Montagnini F, López LN, Romero HF, Dummel CJ, Küppers G, López MA, Lavignolle P,
Cortes CJ, De La Vega MA (2022) Conservation, connectivity and registration of seed areas in
remnants of natural reserves in the province of Misiones, Argentina. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in Biodiversity and Conservation. Springer, Cham, pp 157–181
Esbach MS, Puri M, Botero-Arias R, Loiselle BA (2022) Beyond the island: safeguarding biodi-
versity requires integrated approaches. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human dominated-environments. Series: Topics in Biodiversity and Conserva-
tion. Springer, Cham, pp 551–568
FAO (2015a) Status of the World’s Soil Resources. Main report. Prepared by Intergovernmental
Technical Panel on Soils (ITPS) [Link]
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 701

FAO (2015b) Los Pueblos Indígenas y las Políticas Públicas de Seguridad Alimentaria y
Nutricional en América Latina y el Caribe. Una visión regional. [Link]
[Link]
FAO (2019) Informe de Seguridad Alimentaria. [Link]
Gadgil M, Berkes F, Folke C (2021) Indigenous knowledge: from local to global. 50th anniversary
collection: biodiversity conservation. Ambio 50:967–969. [Link]
01478-7
Gallo J, Pasquin L, Reyers B, Cowling RM (2009) The role of private conservation areas in
biodiversity representation and target achievement within the little Karoo region,
South Africa. Biol Conserv 142(2):446–454
Giraldo LP, Chará J, Calle Z, Chará-Serna AM (2022) Riparian corridors: longitudinal biodiversity
islands in farming landscapes. In: Montagnini F (ed) Biodiversity islands: strategies for conser-
vation in human-dominated environments. Series: Topics in Biodiversity and Conservation.
Springer, Cham, pp 139–156
González LH, Neffeng G, Benedetto V, Sánchez Miñarro M, García A, Biasatti R, Rimoldi P,
Alesio C, Paiz D (2022) The anthropization of the natural landscape in the Pampa region in
Santa Fe province, Argentina. Resilience of the environment, opportunity for a change of the
agri-food paradigm. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Springer, Cham, pp 353–386
Griscom BW, Adams J, Ellis PW, Houghton RA, Lomax G, Miteva DA, Schlesinger WH, Shoch D,
Siikamäki JV, Smith P, Woodbury P, Zganjar C, Blackman A, Campari J, Conant RT,
Delgado C, Elias P, Gopalakrishna T, Hamsik MR, Herrero M, Kiesecker J, Landis E,
Laestadius L, Leavitt SM, Minnemeyer S, Polasky S, Potapov P, Putz FE, Sanderman J,
Silvius M, Wollenberg E, Fargione J (2017) Natural Climate Solutions. Proc Natl Acad Sci
114(44):11645–11650. [Link]
Hajjar R, Oldekop JA, Cronkleton P, Newton P, Russell AJ, Zhou W (2021) A global analysis of the
social and environmental outcomes of community forests. Nat Sustain 4(3):216–224
Hilty J, Worboys G, Keely A, Woodley S, Lauche B, Locke H, Carr M, Pulsford I, Pittock J, White
JW, Theobald DM, Levine J, Reuling M, Watson JEM, Ament R, Tabor GM (2020) Guidelines
for conserving connectivity through ecological networks and corridors. Best Practice Protected
Area Guidelines Series No. 30. Gland, Switzerland: IUCN
IPBES (2019) Summary for policymakers of the global assessment report on biodiversity and
ecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and
Ecosystem Services. IPBES Secretariat, Bonn. [Link]
Karger DN, Kessler M, Lehnert M, Jetz W (2021) Limited protection and ongoing loss of tropical
cloud forest biodiversity and ecosystems worldwide. Nat Ecol Evol 5:854–862. [Link]
10.1038/s41559-021-01450-y
Kirby K (2022) Islands of trees in long-fragmented landscapes in Great Britain. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in Biodiversity and Conservation. Springer, Cham, pp 337–352
Laino R, Musalem K, Laino L, Caballero A, Bueno D, Aranda L, Esquivel A, Ferreira Riveros M,
Romero Nardelli L, Cantero N, Irala R (2022) Island forests among savannahs: conservation of
biodiversity in rangelands of the Paraguayan humid Chaco. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Series: Topics in Bio-
diversity and Conservation. Springer, Cham, pp 185–205
Levin B (2022a) Regenerative agricultural systems as biodiversity islands. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in Biodiversity and Conservation. Springer, Cham, pp 61–88
Levin B (2022b) The role of community led action for the creation of biodiversity islands. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human-dominated envi-
ronments. Series: Topics in Biodiversity and Conservation. Springer, Cham, pp 487–504
Loh EH, Zambrana-Torrelio C, Olival KJ, Bogich TL, Johnson CK, Mazet JAK, Karesh W,
Daszak1 P (2015) Targeting transmission pathways for emerging zoonotic disease surveillance
702 F. Montagnini et al.

and control. Vector-Borne and Zoonotic Diseases 15 (7):432–437, [Link]

2013.1563
Martín E (2020) Antología Toxicológica del Glifosato. Naturaleza de derechos. Argentina. 5th ed.,
269 pp. Available at: [Link]
[Link]
McNeeley JA (2021) How to conserve biological diversity: perspectives from Ambio. 50th
anniversary collection: biodiversity conservation. Ambio 50:957–961. [Link]
s13280-020-01479-6
Mitchell BA, Stolton S, Bezaury-Creel J, Bingham HC, Cumming TL, Dudley N, Fitzsimons JA,
Malleret-King D, Redford KH, Solano P (2018) Guidelines for privately protected areas. Best
Practice Protected Area Guidelines Series No. 29. UICN, Gland
Montagnini F, Berg KE (2019) Feeding the world. In: Carver F, Gossington H, Manuel C (eds)
Sustainable Development Goals. Transforming our world. Witan Media Ltd, Painswick, UK,
United Nations Association – UK, London, pp 98–100. [Link]
feeding-the-world/
Montagnini F, del Fierro S (2022) Functions of agroforestry systems as biodiversity islands in
productive landscapes. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Series: Topics in Biodiversity and Conservation. Springer,
Cham, pp 89–116
Montagnini F, Finney C (2011) Payments for environmental services in Latin America as a tool for
restoration and rural development. Ambio 40:285–297
Montagnini F, Levin B, Berg KE (2022) Introduction. Biodiversity islands: strategies for conser-
vation in human-dominated environments. In: Montagnini F (ed) Biodiversity islands: strategies
for conservation in human dominated environments. Series: Topics in Biodiversity and Con-
servation. Springer, Cham, pp 1–35
Monteferri B (Ed) (2019) Áreas de conservación privada en el Perú: avances y propuestas a 20 años
de su creación. Sociedad Peruana de Derecho Ambiental, Lima, Peru
Montes-Londoño I, Calle A, Montes-Calderon A, Montes-Botero O (2022) Hacienda Pinzacuá: an
island of regenerative agriculture in the piedmont of the Colombian Central Andes. In:
Montagnini F (ed) Biodiversity islands: strategies for conservation in human-dominated envi-
ronments. Series: Topics in Biodiversity and Conservation, Springer, Cham, pp 305–335
Morales-Nieves GM (2022) Priorities, perspectives and use of a community forest by surrounding
residents in Mayagüez, Puerto Rico: protecting the forest for its services. In: Montagnini F
(ed) Biodiversity islands: strategies for conservation in human-dominated environments. Series:
Topics in Biodiversity and Conservation, Springer, Cham, pp 505–530
Morton T (2013) Hyperobjects: philosophy and ecology after the end of the world. University of
Minnesota Press, Minneapolis
Negret HRC, Negret R, Montes-Londoño I (2022) Residential garden design for urban biodiversity
conservation: experience from Panamá City, Panama. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human-dominated environments. Springer, Cham,
pp 387–417
Nerfa L, Wilson SJ, Reid JL, Rhemtulla JM (2021) Practitioner views on the determinants of
tropical forest restoration longevity. Restor Ecol 29(3):e13345. [Link]
13345
Newcomer Q, Camacho Céspedes F, Stallcup L (2022) The Monteverde Cloud Forest: evolution of
a biodiversity island in Costa Rica. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in Biodiversity and Conserva-
tion, Springer, Cham, pp 237–278
Niella F, Rocha P, Tuzinkievicz A, Buchweis R, Bulman C, Thalmayr P, González J, Montagnini F,
Sharry S (2022) Native multipurpose species in the Yaboti Biosphere Reserve, Misiones-
Argentina. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in Biodiversity and Conservation, Springer, Cham,
pp 461–483
Painter K, Buschbacher R, Souto Silva LC, Costa e Silva E (2022) Agroecology and Forest
Conservation in Three Types of Land Reform Community in the Cacao Region of Bahia,
27 Conclusions: Challenges and Opportunities in Implementing Biodiversity Islands 703

Brazil. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in Biodiversity and Conservation, Springer, Cham,
pp 569–599
Pimm S (2021) What is biodiversity conservation? 50th anniversary collection: biodiversity
conservation. Ambio 50:976–980. [Link]
Pimm SL, Jenkins CN, Abell R, Brooks TM, Gittleman JL, Joppa LN, Raven PH, Roberts CM,
Sexton JO (2014) The biodiversity of species and their rates of extinction, distribution, and
protection. Science 344:1246752
Reyes-García V, Fernández-Llamazares Á, McElwee P, Molnár Z, Öllerer K, Wilson SJ, Brondizio
ES (2019) The contributions of indigenous peoples and local communities to ecological
restoration. Restor Ecol 27(1):3–8. [Link]
Roldán M, Carminati A, Biganzoli F, Paruelo JM (2010) Las reservas privadas ¿son efectivas para
conservar las propiedades de los ecosistemas? Ecol Austral 20(2):89–221
Salzman J, Bennett G, Carroll N, Goldstein A, Jenkins M (2018) The global status and trends of
payments for ecosystem services. Nat Sustain 1:136–144
Santos-Gally R, Boege K (2022) Biodiversity islands. Native tree islands within silvopastoral
systems in a neotropical region. In: Montagnini F (ed) Biodiversity islands: strategies for
conservation in human-dominated environments. Series: Topics in Biodiversity and Conserva-
tion, Springer, Cham, pp 117–138
Sayer J, Margules C, McNeely JA (2021) People and biodiversity in the 21st century. 50th
anniversary collection: biodiversity conservation. Ambio 50:970–975. [Link]
s13280-020-01476-9
Sheban K (2022) Preserving biodiversity in Appalachian mixed mesophytic forest through permit-
based harvest of ginseng and other forest botanicals. In Montagnini F (ed) Biodiversity islands:
strategies for conservation in human-dominated environments. Series: Topics in Biodiversity
and Conservation, Springer, Cham, pp 601–628
Shukla PR, Skea J, Calvo Buendia E, Masson-Delmotte V, Pörtner HO, Roberts DC, Zhai P,
Slade R, Connors S, Van Diemen R, Ferrat M (2019) Climate change and land: an IPCC special
report on climate change, desertification, land degradation, sustainable land management, food
security, and greenhouse gas fluxes in terrestrial ecosystems. IPCC
Sigman E (2022) Safeguarding the benefits of biodiversity islands in northern Ethiopia in the midst
of political change. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in
human-dominated environments. Series: Topics in Biodiversity and Conservation, Springer,
Cham, pp 647–674
Solberg D (2021) Closing the gap. Nature Conservancy Magazine Summer 2021: 38–40, Nature.
org/Magazine
Soler R, Benítez J, Solá F, Lencinas MV (2022) Biodiversity islands at the world’s
southernmost city: plant, bird and insect conservation in urban forests and peatlands of Ushuaia,
Argentina In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in Biodiversity and Conservation, Springer, Cham,
pp 419–437
Toensmeier E (2022) Paradise Lot: A Temperate-Climate Urban Agroforestry Biodiversity
Island. In: Montagnini F (ed) Biodiversity islands: strategies for conservation in human-
dominated environments. Series: Topics in Biodiversity and Conservation, Springer, Cham,
pp 439–459
United Nations IPBES (2019) Global assessment report on biodiversity and ecosystem services.
[Link]
Vásquez V, Barber C, Dguidegue Y, Caughlin TT, García R, Metzel R (2022) Farmer perceptions
of tropical dry forest restoration practices on the Azuero Peninsula of Panama – implications for
increasing biodiversity in a human-dominated landscape. In: Montagnini F (ed) Biodiversity
islands: strategies for conservation in human dominated environments. Series: Topics in Bio-
diversity and Conservation, Springer, Cham, pp 629–646
Zárate J (2021) Wars of the interior. Granta Publications Ltd., Cambridge. [Link]
uk/Wars-Interior-Joseph-Zarate/dp/1783786159
Index

A 238, 239, 242, 254–269, 299–301, 346,

Advocacy, 498, 500 374, 388–413, 421, 423, 463, 488–490,
Agroecology, 63, 78, 119, 260, 299–301, 355, 493–500, 506, 545, 552, 554, 559, 562,
369, 373, 378, 509, 578, 584, 587, 689, 565, 571, 572, 631, 679, 681, 689,
692–694 695–698
Agro-food system, 373–378 Biodiversity-friendly landscapes, 54
Agroforestry, 11, 13, 15, 16, 21, 22, 28, 63, 66, Biodiversity gardens, 390, 408, 409
67, 70, 79, 90, 96–100, 103, 119, 140, Biodiversity islands, 4–29, 43, 56, 57, 62–82,
159, 306, 310, 311, 330, 373, 389, 90, 95, 105–107, 109, 110, 118–133,
439–458, 465, 467, 495, 524, 560, 561, 140–152, 159, 176, 188, 195, 197,
571, 572, 576, 580–582, 585–588, 590, 207–233, 238–269, 280–301, 375, 406,
592–595, 603, 616, 618, 619, 642, 657, 411, 420–434, 439–458, 463, 466, 480,
658, 670, 680, 689, 691, 693, 695, 697 488–500, 506, 523, 524, 533, 545,
Agroforestry systems (AFS), 11, 14, 15, 17, 20, 552–565, 571, 572, 576, 582, 589, 592,
21, 28, 29, 66, 67, 69, 78, 79, 89–110, 594, 595, 602, 607, 614, 618, 622, 630,
151, 164, 174, 230, 231, 312, 323, 369, 648–670, 678–699
374, 439, 447, 488, 560, 576, 577, 581, Biological corridors, 4, 13, 93–95, 98, 106, 108,
582, 585–587, 590, 593, 680, 682, 691, 122, 143, 195, 221, 230, 231, 238–240,
699 257, 259–267, 269, 281, 282, 356,
Ancient woodland, 5, 339, 341, 344, 346, 349, 363–365, 369, 684
684 Buffer zones, 4, 13–15, 28, 90, 93, 98, 106,
Andean Streams, 152 109, 535
Atlantic forest, 94, 100, 101, 158, 197, 462,
572–574, 582, 592, 594, 688
Attitude changes, 411, 412 C
Cabruca, 94, 572, 573, 585, 587, 589, 592
Caribbean, 21, 241, 248, 251, 506–509, 519,
B 522, 524–526
Biocultural conservation, 543 Cattle, 15, 20, 43, 46, 62, 70, 71, 79, 94, 95,
Biodiversity conservation, 6–8, 13–15, 17, 19, 102, 103, 108, 109, 118–127, 130–133,
24–29, 41–57, 62, 63, 77–80, 90, 98, 141, 144, 145, 148–151, 188, 192,
101, 102, 104, 106, 109, 119, 132, 133, 198–201, 209, 216–218, 220, 241, 262,
149, 158, 172, 198, 200, 201, 209, 225, 280, 281, 283–288, 296–298, 300, 301,

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 705
F. Montagnini (ed.), Biodiversity Islands: Strategies for Conservation
in Human-Dominated Environments, Topics in Biodiversity and Conservation 20,
[Link]
706 Index

Cattle (cont.) 214–219, 230, 232, 254, 260, 264, 267,

306, 308, 309, 311, 314, 320, 323–325, 493, 499, 506, 507, 534, 535, 540, 554,
327–329, 356–358, 360, 507, 510, 539, 606, 649, 654, 678, 682, 699
541, 545, 633, 636, 637, 641–643, 685, Degraded landscapes, 17, 28, 65–67, 71, 79, 81,
686, 693 82, 95, 105–107, 329, 488, 494,
Certification, 78, 80, 107, 110, 159–166, 171, 496–499, 649, 654, 669
173–176, 285, 289, 576, 585, 612, 617, Diversity, 4, 6, 7, 14, 17, 20, 42, 56, 62–64, 66,
620, 681 68–74, 77, 82, 90–96, 98, 99, 102–107,
Church forests, 9, 23, 488, 499, 532–542, 545, 118, 119, 123–125, 127–129, 131, 132,
688 143–146, 158–160, 165, 171, 176, 177,
Colombian Andes, 306–331, 686 195, 197, 208, 209, 214, 215, 217, 225,
Communal labor, 658, 685 242–244, 250, 252, 263, 266, 268, 269,
Community actions, 22–24, 488, 494, 495, 499, 280, 292–295, 306, 314, 320, 323, 324,
500 327, 363, 364, 369–371, 373, 374, 389,
Community-based research, 507, 522, 687 398–400, 402–405, 410, 412, 420, 421,
Community forestry, 506, 509, 688 423, 426–429, 431, 440, 444, 446, 447,
Connectivity, 7, 9, 12, 15–17, 19, 66, 95, 101, 456, 457, 463–468, 479, 480, 490, 492,
103, 104, 106, 109, 143, 149, 158, 159, 493, 499, 523, 533, 542, 544, 553, 554,
187, 188, 195, 197, 209, 210, 221, 570, 602, 604, 608, 610, 615, 616, 649,
229–232, 259, 261, 264, 266, 306, 323, 652, 668, 670, 679, 681–683, 690, 698
327, 339, 348, 356, 364, 365, 369, 388, Dryland, 5, 668
395, 397, 399, 404, 406, 410, 412, 432,
434, 463, 560, 669, 697
Conservation, 3–6, 8, 9, 12–17, 19–29, 42, 43, E
52–54, 56, 57, 62, 66, 72, 80, 89, 90, Eastern USA, 447
92–94, 96, 102–104, 107–110, 119, 121, Ecological restoration, 119, 121, 122, 124,
124, 127, 128, 140, 141, 143–145, 149, 126–131, 299, 301, 407
151, 152, 158–177, 186, 188, 190, 192, Ecosystem functioning, 64, 119, 124, 126, 129,
193, 195, 197–201, 208–210, 214–230, 296
232, 238–240, 242, 243, 245, 246, 252, Ecosystem services, 5, 11, 16, 20, 23, 25–29,
254–258, 261–264, 266–269, 280, 281, 63, 64, 69, 70, 81, 99, 105, 118, 119,
299, 301, 306, 314, 316, 327, 338, 339, 124, 125, 127–129, 132, 143, 160,
346, 348, 363–365, 369, 371, 373–377, 188–195, 201, 208, 219, 229, 232,
389–391, 400, 404, 408, 410–412, 252, 258, 262, 263, 267, 268, 291,
420–434, 462–468, 472, 480, 488–495, 298, 300, 306, 307, 325, 326, 331,
497, 499, 506–508, 510, 519, 521–523, 390, 411, 420, 440, 447, 458, 499,
527, 532, 533, 538, 542–545, 552, 554, 507, 514, 516, 517, 523, 533, 544,
556, 558–565, 570–572, 575, 576, 586, 561, 621, 630, 649, 654, 655,
589, 590, 592, 603, 607, 612, 613, 682–684, 691, 696, 697, 699
615–618, 620–622, 648–650, 652, Endangered species, 11, 95, 103, 128, 195, 242,
654–656, 658, 661, 664, 668, 670, 396, 563, 612, 631
680–692, 695–699 Endemic species, 252, 254, 322, 323, 427, 462,
Conservation and development, 266, 552, 582, 527
588, 589, 591, 687
Ethiopian Orthodox Tewahido Church (EOTC),
Conservation easements, 22, 23, 261, 488, 495,
533–542, 545
500, 621
Conservation strategy, 4, 6, 11, 56, 161, 200,
209, 230, 232, 299, 369, 374, 391, 397,
F
432, 499, 500, 545, 558, 696, 698
Floodable forest, 213, 362
Cover cropping, 67, 68
Flora, 163, 200, 201, 208–210, 214, 220–229,
232, 243, 244, 249, 341, 343, 348, 349,
355, 362, 382, 431, 489, 544, 613, 639,
D
682, 683
Deciduous forests, 423, 602, 603, 606, 620, 622
Deforestation, 21, 27, 42, 43, 56, 57, 62, 98, Food forest, 440
100, 103, 118, 119, 127, 199, 208–210, Forest botanicals, 602–625
Index 707

Forest conservation, 219, 264, 508, 532, 571, L

572, 576, 578, 582, 583, 586–595, Land reform, 570–595, 682, 685
622 Land trusts, 22, 488, 495, 500
Forest farming, 603, 606, 607, 615–622, 685 Landscape configuration, 7–12, 106, 356, 364,
Forest landscape restoration (FLR), 631–633, 560
643, 644, 650 Landscape ecology, 338, 349, 362, 369, 373
Forest restoration, 140, 151, 160, 319, 463, Landscape restoration, 9, 14, 94, 128, 631, 641,
630–644, 686 659, 660
Functional biodiversity, 280, 289, 296–297 La Vieja River, 141–143, 149, 150, 307, 316
Functional traits, 119, 121–125, 128, 129, 132 Livestock, 16, 42, 66, 70, 71, 79, 101, 102,
118–127, 130, 132, 133, 141, 147, 148,
151, 192, 198–201, 208, 209, 218–220,
G 229, 232, 280, 283, 289, 297–301, 308,
Garden practitioners, 390 312, 325, 329, 340, 341, 358, 440, 462,
Genetic variability, 4, 281, 463, 466, 467, 480 465, 480, 558, 581, 583, 585, 602, 654,
Germination, 98, 126, 131–133, 163, 164, 170, 662, 681, 685, 694
171, 174, 455, 469, 470, 473, 475, 476, Livestock farming, 121, 122, 130, 199
606, 608
Governance, 23, 390, 488, 497–500, 538,
539, 559, 562, 649, 661, 663, 664, M
670, 690 Markets, 26, 27, 73, 76, 80, 100, 107, 108, 119,
Grassroots conservation, 488 166, 218, 260, 285, 298, 308, 310, 314,
Green areas, 17, 19, 420–423, 432–434 316, 320, 329, 330, 357–359, 362, 378,
380–382, 464, 466, 523, 563, 575, 576,
582, 590–593, 603, 612, 613, 616, 617,
H 619–622, 657, 689, 695, 696
Habitat connectivity, 197, 209, 264, 410
Habitat fragmentation, 7, 8, 28, 101, 108, 143,
463, 466, 570, 680 N
Habitat loss, 5, 6, 101, 118, 127, 195, 197, 208, Native tree species, 9, 108, 124, 127, 130, 159,
229, 230, 250, 252, 406, 462, 562, 570 162, 164, 166, 174, 311, 511, 633
Habitat restoration, 63, 69, 70, 252 Natural reserves, 165, 171
Hedgerows, 14, 63, 68, 80, 310, 311, 340, 343, Nothofagus, 422, 423, 426, 429
395–400, 402, 403, 405–408, 410, 412
Herbs, 65, 103, 120, 121, 163, 166, 284, 428,
450, 603, 606, 608, 609, 612, 613, O
615–619, 622 Organic farming, 63, 100, 108, 691
Homegarden, 15, 72, 90–96, 106, 107, 109,
312–315, 389, 433, 440, 458, 488, 500
Human perceptions, 434 P
Parkland, 62, 339, 343, 349
Participation, 26, 257, 258, 381, 497, 509, 513,
I 514, 516, 518, 520, 523, 527, 542, 555,
Indigenous communities, 73, 509, 553, 682, 563, 565, 583, 643, 651, 656, 658, 662,
690, 699 666, 667, 687
Indigenous management, 496 Participatory, 6, 21, 161, 381, 498, 506, 508,
Intensive silvopastures, 288 521, 552, 553, 560, 562, 563, 565, 583,
Intercropping, 63, 65, 66, 376, 439 653, 658, 662, 663, 667
Interdisciplinary approaches, 552 Participatory research, 633, 687
Interior Atlantic forests, 462, 465–467 Partnership, 21, 269, 300, 330, 488, 552–555,
563, 565, 594, 615, 687, 689
Patagonia, 420, 423, 427, 429, 431
J Payments for ecosystem services (PES), 21, 25,
Jupará Agroecological Movement, 571, 572, 26, 56, 76, 80, 108–110, 257, 258, 261,
575–577, 582, 583, 588, 594, 595, 685 264, 267, 268, 309, 320, 561, 621, 696
708 Index

Perennial crops, 14, 15, 66, 90, 91, 93, 94, 98, Riparian biodiversity, 145
107, 109, 282, 290, 441, 447 Riparian buffers, 16, 91, 109, 140, 141,
Permaculture, 63, 373, 374, 439–441, 493, 500, 143–146, 150, 152, 311, 327, 681
691
Phylogenetic diversity, 119, 123, 124,
127–132, 685 S
Political ecology, 498, 650, 661 Sacred forests, 532–535, 537–541, 543–545,
Private land conservation, 267, 269 682
Private reserves, 12, 161, 162, 165, 255 Sacred natural sites, 532–534, 542–545, 686
Production, 9, 15, 19, 21, 25, 28, 62–64, 69, 70, Sacred sites, 28, 488, 499, 508, 680, 691
73, 77–82, 92–94, 96, 97, 99, 102, 103, Saproxylic beetles, 345, 348
106, 108, 118–122, 124, 132, 133, 140, Selva paranaense, 158, 462
151, 164, 165, 175, 177, 186, 197–201, Shadow conservation, 532, 534
209, 220, 221, 257, 260, 266, 280, 281, Silvopastoral systems, 9, 14–16, 66, 70, 71,
283, 285, 287, 289, 291, 297–301, 306, 102, 103, 144, 159, 197, 202, 209, 230,
308–312, 314, 318, 323, 325, 327–330, 231, 632, 642, 681, 685, 693
338, 356–360, 362, 363, 365, 366, Silvopastoral systems (SPS), 90, 91, 101,
373–382, 389, 441, 446, 455, 458, 465, 103, 119–121, 123, 127, 131, 132,
467, 472, 476, 494, 499, 507, 509, 538, 140, 151, 280, 282–288, 293–296,
557, 560, 571–574, 576, 577, 580–586, 298, 300, 306, 309, 312–315, 317,
588–595, 618–620, 633, 656, 678, 679, 320, 323, 325–329
681, 684, 685, 687–689, 691–695, 699 Silvopastures, 70, 71, 280, 282–287, 293–295,
Production models, 300, 327, 354, 358, 362, 312, 316–320, 324, 326, 327, 691, 695
369, 373, 678, 694 Social-ecological systems (SoESs), 565, 650,
Propagation material, 159–163, 165, 166, 168, 683
174–176, 466 Sphagnum, 422, 424
Stewardship, 229, 255, 269, 491, 535, 554, 555,
557, 560, 608, 621, 648, 650
Q Sugarcane, 280–282, 288–291, 293–299, 507
Quilombo, 577, 580, 586, 593 Sustainability, 15, 17, 20, 81, 90, 93–95,
121–124, 165, 209, 280, 287, 297,
299, 328, 377, 408, 433, 440, 558,
R 563, 564, 589–591, 613, 617, 656,
Rainforests, 8, 24, 27, 42–57, 100, 130, 141, 668, 684, 696
212, 239, 241, 242, 252, 253, 255, 257,
258, 261, 264, 268, 346, 554, 570, 648,
689, 692 T
Reduced agrochemical use, 63, 68, 69 Tierra del Fuego, 421, 422, 425, 426
Reforestation, 6, 160, 257, 261–263, 268, 314, Traditional ecological knowledge (TEK), 63,
318, 631–634, 636–643, 655, 681, 683 73–75, 77, 121, 489, 498
Refuge habitats, 420, 423, 434 Tree plantings, 4, 11, 12, 17, 19, 159–161, 209,
Regeneration, 6, 29, 67, 69–72, 108, 119, 127, 311, 312, 328, 631, 632, 638, 641–644,
149–151, 162, 165, 175, 209, 221, 285, 655, 686
315, 318, 320, 321, 323, 355, 395, 464, Tropical forest conservation, 127
465, 540, 541, 559, 560, 605, 612, Tropical gardens, 391, 440, 458
631–634, 636–644, 648, 650, 654, 655,
660, 665, 683, 688
Restoration, 4, 6, 9, 12, 23, 25, 28, 56, 63, 66, U
69, 72, 73, 75, 79, 90, 93, 96, 103, 108, Ucayali region, 208, 210–219, 229–231, 681,
109, 124–129, 132, 133, 140, 143, 686
148–150, 152, 158–164, 168, 170, Understory, 49, 65, 101, 317, 405, 406, 423,
174–177, 231, 281, 299, 326, 349, 363, 424, 429, 457, 511, 521, 522, 524, 541,
441, 463–467, 480, 500, 509, 559, 560, 573, 602, 603, 606–622, 624–625, 685
606, 607, 631–634, 636, 638–644, Urban agriculture, 493
648–658, 660–670, 681, 684–690, 699 Urban expansion, 388, 390, 413, 507, 510
Index 709

Urban green spaces (UGS), 389, 390, 410, 411, 509, 602, 639, 648, 650, 654–656, 661,
434 665, 666, 669, 689, 690, 699
Urban policy and planning, 390 Wildlife, 4, 13, 17–21, 63, 71, 79, 80, 92–94,
97, 98, 102, 103, 106, 109, 162, 190,
192, 195, 200, 201, 231, 244, 266, 284,
V 285, 301, 314, 324, 327, 330, 338,
Vascular plants, 243, 244, 249, 250, 339–341, 369–371, 391, 392, 395–397, 399, 400,
344, 345, 348, 349, 400, 406, 423–429, 402–406, 408–410, 412, 420, 441, 442,
432, 462 446–449, 451–453, 455, 458, 506, 516,
Vegetative propagation techniques, 467, 480 517, 520, 521, 523, 524, 526–527,
Veteran trees, 340, 342 553–555, 557–559, 561, 562, 564, 565,
602, 612, 649, 655, 681, 689
Woodland management, 346
W Wood-pasture, 339–344, 348, 349
Watershed, 5, 19, 21, 26, 63, 69, 141, 143, 144, World Wildlife Fund (WWF), 24, 199, 571,
146, 148–151, 257, 263, 264, 307, 316, 572, 575–578, 588, 594