Advanced Ecology

(Biol 551)

Tamrat Bekele (PhD)

Professor of Vegetation Ecology
Department of Plant Biology and
Biodiversity Management
 Course Management

1. Lectures
2. Individual Reference Readings and Classroom and
Independent Discussions***

Course Evaluation
3. Assignments
4. Final test
 Selected References

1. Principles of terrestrial Ecosystem Ecology

2. Philosophy of Ecology

3. Biodivesrity , Ecosystem Functioning &

Human Wellbeing
Contd.

Introduction

Ecology: the study of the environmental

relationships that determine the geographical
distribution and abundance of organisms.
 Advanced Ecology (Biol 551)
Introduction

Pertinent ecological questions:

- What is the nature of the relationship?
- How does the relationship influence the
geographical distribution of the organism?
- How does the relationship influence the
abundance of the organism?
Important Ecological concepts

General Overview
• The Ecosystem Resources Nature Reserve
• Food Webs • Species Diversity • The Community
• Island • Stochastic • Population Cycles
Biogeography Processes • Pattern
Theory • Competition • Life-history
• Succession • Density-dependent Strategies
• Ecological Regulation • Predator-prey
Adaptation • Natural Interactions
• Bioaccumulation in Disturbance • The
Food Chains • Niche Diversity/Stability
• Energy Flow • Limiting Factors Hypothesis
• Environmental • Habitat Restoration • Ecosystem Fragility
Heterogeneity • Materials Cycling • Plant-herbivore
• Coevolution • Carrying Capacity Interactions
• Conservation of • • Socioecology
 Some Ecological Concepts underlying Important

Environment/Conservation Management Plans

E.g., Restoration Ecology

1. Population and ecological genetics:

2. Community ecology:

3. Biodiversity and ecosystem functioning:

4. Invasive species and community invasibility:

5. Ecophysiological and functional ecology:
6. Demography, population dynamics, metapopulation
ecology:
7. Evolutionary ecology:
8. Fine-scale heterogeneity:
9. Food webs:
10. Ecological dynamics and trajectories:
11. Modeling and simulations:
12. Research design and statistical analysis:
13. Macroecology:
14. Paleoecology, climate change:

Therefore,
“What would be Advanced Ecology”?
 Outline
1. Introduction
2. Ecological gradients
3. Species diversity
4. Ecosystem function
5. Ecosystem services
6. Vegetation processes
7. Alien invasive species
8. Restoration ecology
9. Field ecological methods
10. Biodiversity and global climate change
11. Ecological Modeling
 Expectations of the Course:

Personal reflections:
• Altitude
• Rainfall
• Temp.(max/min)
etc
Equatorial Westerlies

Indian Ocean Stream

 II) Basic concepts of Vegetation Ecology

1) Terminology of Gradients

• Variation in vegetation and ecological factors along gradients

is a major field of interest in modern vegetation ecology.
Contd.

The following are commonly used terminologies of gradient as

proposed by Whittaker (1967, 1978):

• An ecological gradient
: is the gradual change in any ecological factor.
Ecological factors are often correlated with each other, and
act on the plants in combination.
Contd.

• Complex gradient

A complex of ecological factors varying more or less in

parallel make up a complex gradient.
e.g. nutrient (Ca, Mg, Mn) gradient.
Contd.

• Coenocline

Vegetation change (in composition) along this complex

gradient is known as Coenocline.
Contd.

• Ecocline
The coenocline (variation in vegetation) is mostly correlated
with, and partly caused by, the variation in an underlying
complex gradient.

If we consider the coenocline as a function of a known

complex-gradient and want to emphasize both the
vegetational and ecological variation, we use the concept
ecocline.
Contd.

• Ecotone
An ecotone is a relatively sharp change in one, two or three of
complex gradient, coenocline and ecocline.
Contd.

• Ecological factors differ greatly in the way they influence

plants.

• Among the various schemes followed in the classification of

ecological gradients, Austin (1980) put forward the following;
- emphasizing their way of influencing the plants rather than
their source:
contd

•Indirect environmental gradients

e.g. Altitude, OM, Soil depth, and Local topography
Contd.

• Direct environmental gradients

The environmental factor has a physiological influence on
plants without being a resource for plant growth.
e.g.
- pH (influences the availability of various nutrients);
- temperature,
- aspect, and
- inclination/slope
Conyd.

• Resource gradient
The environmental gradient is an essential resource for
plat growth, for which competition may occur.
e.g. Nutrients
Gradient

Ecological gradient The gradual change in any ecological factor

Complex gradient Ecological factors varying more or less in parallel

Coenocline Vegetation change (in composition) along a complex

gradient

Ecocline Refers to both the change in vegetation and

underlying ecological variation

Ecotone Sharp change in one, two or three of complex

gradients
 Basic concepts of Vegetation Ecology (Contd)

2) The Concept of the Species Diversity

Objective: To introduce one of the important concepts that is

employed in the description of and analysis of vegetation
data……

• the meaning of diversity,

• methods for measuring and assessing diversity, and
• ecological interpretation of different levels of diversity.
Contd.

Species richness:
- a count of the number of plant species in a quadrat, area or
community;
- often equated with diversity;

• Most methods for measuring diversity actually consist of two

components.
- species richness,
- relative abundance (evenness or unevenness) of species within the
sample or community.
Contd.

• Diversity is thus measured by recording the

- number of species and

- their relative abundance.

• The two components of species richness and evenness may be

examined separately or combined into some form of index.
Contd.

Types of Diversity

Two types of diversity: alpha and beta diversity.

i. Alpha diversity
• the mean number of species within a chosen area or
community such as one type of woodland or
grassland,
i.e. at the within-community level.
Contd.

• The measure is dependent on sample plot size.

• Why?

- because increase in sample plot area invariably leads

to an increase in the environmental heterogeneity and
thus to increased number of species per sample plot.
ii. Beta diversity
: the difference in species diversity between areas or communities.

Beta diversity is thus sometimes called habitat diversity because it

represents differences in species composition between very
different areas or environments and the rapidity of change of
those habitats.

a.k.a.. habitat diversity

Contd.

Measuring Biodiversity
• Measuring BD requires determining which elements of BD
are present in the area of interest.

• i.e., ideally this requires a complete inventory, including

genes, species and ecosystems.

• But logistical constraints become limiting!

Contd.

i) Species richness: a simple, commonly used measure of

diversity.

• The number of species present in a given area.

• An ecosystem with 10 species is more diverse than

another where there are only 8 species.
Contd.

ii) Evenness: relative abundance of different species.

H= -Σ pi log pi

• Where pi is a measure of the importance of the ith

species.

• Evenness = H / Hmax, where Hmax is the maximum

possible value of H.
Contd.

For instance,
• Consider the following scenario;

Vegetation “A” -

- Species richness =10 species;

- Species abundance = 10 individuals each;

- Total = 100 individuals;

Vegetation “B” –

- Species richness =10 species;

- Species abundance = 9 spp. (2 indiv. each), and 1spp (82 ind.);

- Total = 100 individuals;

 Synopsis

Diversity scales (R. Whittaker, 1960):

• Devised a simple system for classifying the scales at which
diversity occurs;

i. Alpha Diversity
ii. Beta Diversity, and
iii. Gamma Diversity
Contd.
Alpha Diversity:
• The diversity that exists within an ecosystem;
e.g., species richness in a forest,
Beta Diversity:
• Among habitat diversity;
e.g., species diversity between a forest and the adjacent
meadow,
Gamma Diversity:
• Geographic-scale diversity;
e.g., species diversity among differing habitats on one island and
another one a thousand km away,
 Diversity indices

• A large number of indices of diversity have been devised, each of

which seeks to express the diversity of a sample or quadrat by a
single number.

• the simple totaling of species numbers to give species richness.

• However, of the indices that combine species richness with

relative abundance, probably the most widely used is the
Shannon index.
 Diversity indices (cf. Kent and Cocker)

• Shanon-Wiener Index

• Equitability or Evenness Index

• McIntosh Index
Various formulae to measure diversity:
cf
Issued books.
Equitability or evenness index: (Check!)

Equitability J = H’ / H’max = ∑ pi ln pi / ln s

(evenness)
where s = the umber of species

pi = the proportion of individuals or the

abundance of the ith species expressed as a
proportion of total cover

ln = log basen
Contd.

• What is the ecological importance of species richness?

(Case 1):

- An ecosystem with more species receives priority in

conservation. i.e., considering all other things being equal.
e.g.,
Evenness: relative abundance of different species.

Ecosystem A B C

Tid 40 120 80

Zigba 30
60 60

Woyra 20 20 60

Kerero 10 - -

Richness 4 3 3

Evenness 0.92 0.88 0.99

Contd.

Evenness:
• In the above Table – ecosystem C is more diverse than B.

• (i.e., because the three species have similar levels of

abundance, or high evenness).

• Cf. A team of 5 men and 5 women is more diverse than a

team with 8 men and 2 women! Because it is more even!
e.g.,
Evenness: relative abundance of different species.

Ecosystem A B C

Tid 40 120 80

Zigba 30
60 60

Woyra 20 20 60

Kerero 10 - -

Richness 4 3 3

Evenness 0.92 0.88 0.99

H 0.56 0.39 0.47

Contd.

• What is the ecological importance of evenness?

(Case 2):

- Reduction in evenness is of concern to conservation

biologists, because this would make uncommon species less
common, while common species become more common.

(Cf. losing a woman from the team that had 8 men and only 2
women)
Contd.

BD and Spatial Scales

• “Extinction” vs “Local extinction”

• Risks of extinction at different spatial scales ….

Key consideration when deciding which species are a high

priority.

• The larger the scale at which an extinction is likely to

occur, the more important it is to try to prevent it.
Contd.

Interpretation of diversity indices

– Most interpretation of diversity indices is still based

primarily on species richness rather than both species

richness and evenness.

– Arguments over the ecological significance of high diversity

still abound.
Contd.

Most of these arguments assume that diversity is equated solely

with species richness and take no account of the relative species
abundances and evenness.

i. Conservation evaluation

ii. Diversity/stability

iii. Diversity/productivity
Contd.

i. Conservation evaluation
• Most ecologists consider high species richness to be

- a desirable property of any community or ecosystem and

this criterion has dominated most methods for ecological
and conservation evaluation techniques.

• Species richness has been shown to be the most frequently

cited criterion in a review of several methods of
conservation assessment (see Usher, 1986).
Contd.

A point of distinction:
• Two types of diversity are recognized by Systematists:
- Taxonomic diversity, and
- Phylogenetic diversity
Contd.

Phylogenetic diversity: conceived as variety in phylogenetic

distinctiveness, measured in terms of discriminating
morphological or genetical features;

Taxonomic diversity: measured on the basis of of the

taxonomical key to the taxa within the group, and thus
Taxonomic distinctiveness is taken as a measure of
phylogenetic distinctiveness;
Taxonomic distinctiveness:
• TD vs Species Diversity

e.g., Ginkgo biloba

(Just one species, in one genus, in one family. In one order,
The Ginkgoales

• How do we quantify this taxonomic distinctiveness?

Measurement for Taxonomic distinctiveness:
– Informative taxonomic statements (col. I)
– Weight for the distinctiveness (col. Q)
– Transforming values to give “1” to the least
distinctive species (col. W)
– Relative distinctiveness as a percentage (col. P)
Taxonomic distinctiveness: This
approach is not in fact developed for measuring
species diversity, but for creating a basis upon which
areas can be compared as to their conservation
value.

• HOW?
Contd.

ii. Diversity/stability

• The relationship between diversity and stability has been long-

running.

• There was a long held view among some ecologists that high
species diversity leads to community stability.

• This has been laid to rest. There is no direct/automatic

relationship between high diversity or species richness and
community/ecosystem stability.
Contd.

• Many species-poor communities are extremely stable, while

many species-rich communities show considerable instability.

• Pielou (1975) discussed this problem at length and showed that

ecosystem and community stability is ultimately dependent on
environmental stability.
Contd.

iii. Diversity/productivity

• species diversity increases towards productive

environments.
 Factors affecting species diversity

i. Niche relations
Alpha diversity

• Most discussions of diversity

are at the local community
scale. (within-community
diversit
ii. Habitat diversity

– habitat diversity (within-

community diversity)
– Overall diversity is
increased at this next scale.
 iii. Mass effects

At still higher level of scale,

- diversity is accentuated by the flow of
propagules of individuals from areas
of high diversity (core areas) to
unfavorable areas, where they

achieve viability.
• These marginal species from between differing plant
adjacent areas around one communities or habitat types.
particular habitat or
community type can greatly
increase species diversity.

• This effect also explains the

high species diversities of many
transitional zones or ecotones
 iv. Ecological equivalency

• At the largest scale, diversity is

increased by ecological equivalency.

• In one part of the world, a certain

collection of plants will have evolved
and comprise a particular
community.
 iv. Ecological equivalency
• Elsewhere in the world, in a very
similar environment, an equivalent
but totally different set of species can
have evolved to make up the
communities in that region.

• Thus, diversity is maintained simply

by the spatial separation of the two
locations and their very different
species compositions.
Contd.

Environmental limiting factors:

Why do certain species grow together in a particular

environment?

• Usually because they have similar requirements for

existence in terms of environmental factors such as
light, temperature, water, drainage and soil nutrients.
Contd.

• i.e., they have similar requirements for existence in

terms of environmental factors such as
 light,

 temperature,

 water,

 drainage ,and

 soil nutrients;

• They may also share the ability to tolerate the activities

of animals and humans such as grazing, burning, cutting
or trampling.
2) The 3)Concept of Plantthe
The Concept of PlantTypesCommunity
Community

Vegetation: The aggregate of all the plants found in an area.

– The simplest concepts of vegetation types have been based on

physiognomy, i.e.,

- The general physical structure and appearance of the

vegetation
Contd.
Contd.

Plant community types:

• Imagine an ecologist standing on a hilltop and

surveying a landscape dominated by natural or semi-
natural vegetation somewhere:-

 The main difference in the landscape will be those of

plant communities.
Contd.
Contd.

• Major distinctions will be made on the basis of

physiognomy or the growth form of the vegetation.

For instance;
- woodland as opposed to scrublad or grassland.

 These units will also represent the major subdivisions of

the landscape in functional terms as ecosystems.
…%...
Contd.

Awash Valley at the edge of ANP

Contd.

Sanati Plateau - BNP

Contd.

Bamboo Forest- BSG

Contd.

• More subtle changes in the landscape will also be

evident in variations in color between areas of
vegetation with perhaps the same physiognomy.

• These color variations will be a reflection of

differences in:
- plant species composition, and
- stage of development.
Contd.

Blue Nile
Contd.

• A considerable part of plant ecology and vegetation

science is concerned with methods for actually
characterizing and defining these areas as different
plant communities.

• The concept of the plant community is, therefore,

absolutely fundamental to the whole discipline of
vegetation ecology.
Contd.

The plant community can be defined as:

“ the collection of plant species growing together in a

particular location that show a definite association or
affinity with each other”
Contd.

Formal description and classification of communities

generally focuses on features such as:

• Floristic composition,
• Floristic structure, and
• Relative abundance
Contd.

i.e.,
• The community is not described by simply listing all the
species which compose it, …… because each are not of
equal importance.

• Instead, a community is characterized by detailing those

species which most contribute to its unique structure and
composition.
Contd.

NB. A community (or vegetation in general) is not equivalent

to the flora of an area.

cf.
• Flora refers to a list of species or to the plant taxa
occurring in that area.

• A flora in a document form may range from:

– a floristic checklist to

– a complete taxonomic treatment with keys –

morphological and nomenclatural information.
 Contd.

Floras, as a rule, do not give information on:

i. the combinations (communities) in which the species
occur in nature,

ii. their abundance, importance, or uniqueness –

all species have equal weight.
 Contd.

• Vegetation ecology is concerned with:

i) - identifying the plant communities (the vegetation) on an

area, but also with

ii) - determining how they are related to one another and to

the environmental factors.
Contd.

Approaches to Description of Vegetation

The Traditions
Introduction
The multiplicity of approaches and theories in
contemporary vegetation science can only be
understood by knowledge of its past.
Contd.

• The history of vegetation ecology before 1960 is

intimately connected with description of vegetation,
for which classification has been the major tool.

• The history of classification shows an early divergence

into several schools, and many of the schools later
became fragmented into minor branches.
 Contd.

Phytosociology

• In the past, different ecologists may have suggested a formal

methodology for the description and classification of

vegetation,

• However, the best known and most universally accepted

methodology is the phytosociology of Braun-Blanquet (1965).

 Contd.

All descriptive traditions except the British and American share

some fundamental principles:

• subjective classification into units, plant

communities, and

• filing of these communities into a formal hierarchy.

These are the fundamental tenets of the branch of

vegetation ecology known as phytosociology.
 Contd.

This methodology requires complete inventory of all species

composing the vegetation (full floristic inventory) and
provides a simple, rapidly applicable method for describing
vegetation.
Contd.

Phytosociology can be described in terms of three

basic steps:

i. The analytic phase: the first is the field inventory,

in which stand descriptions called relevés are
made from representative samples (plots) of the
vegetation of interest.
Contd.

• The field method a.k.a. the analytic phase: subjective

selection of homogenous stands of vegetation
(homogeneity judged subjectively or, more rarely, by
some statistical method), subjective selection of one or
more sample plots within the stand, and

• analysis of the vegetation within these plots (or stand

descriptions called reléves), are made from
representative samples (plots) of the vegetation of
interest.
Contd.

ii. The synthetic phase, arrangement of sample plots into

tables showing similarities and differences in species
composition;

• This step involves recognition of regularly occurring

species combinations, initially called “communities” in
the collection of all reléves done in a region (or
sometimes from similar but other regions as well)
Contd.

iii. The syntaxonomical phase, the abstraction (in the

investigator’s mind) of plant communities (hierarchically
arranged) from the ordered tables of sample plots in
accordance with the rules of the tradition in question.

• This step involves interpretation of the resulting

‘communities’ as vegetation units at different levels in a

system.
Contd.

• The units are given Latin names, with endings reflecting

their level within the resulting hierarchy or syntaxonomy.
Contd.

The Braun-Blanquet Approach

• European phytosociology has its roots in early 19th century

vegetation description.

• Among the many regional schools emerging at the start of

the 20th century, the Braun-Blanquet Approach stands out as
the leading approach, getting support from most of
Europe.
Contd.

This is the approach named after its founder, J. Braun-

Blanquet (1884-1980), developing in Zurich (Switzerland)

and Montpellier (France) during the 1900s and 1910s.

Contd.

The Braun-Blanquet approach is also called:

• “the floristic-sociological approach”,

• “the Zurich-Montpellier school”, and
• “the Middle European-Mediterranean school”
 Contd.

Basic Principles:

The essence of the school may be summarized in three ideas:

1. Plant communities are conceived as types of

vegetation, recognized by their floristic
composition.

• The full species compositions of communities better

express their relations to one another than and
environment than any other characteristic.
Contd.

2. Amongst the species that make up the species composition

of a community, some are more sensitive expression of a
given relationship than others.
Contd.

For practical classification (and indication of environment)

the approach uses:
- those species whose ecological relationships make them most
effective indicators;

- Such species are known as diagnostic species (character

species, differential species, and constant companions).
Contd.

3. Diagnostic species are used to organize communities into a

hierarchical classification of which the association is the
basic unit.
(cf. species in taxonomic classification)
Contd.

Structuring Processes
i.e. pattern and process in vegetation

Four main structuring processes in vegetation may be

considered:
i. Interspecific interactions
ii. Destabilizing factors
iii. Stress
iv. Chance
 Contd.

1) Interspecific interactions
Several kinds of interspecific interactions occur between plants.
Plants with strong competitive ability have been termed Competitors.

- Characteristics of competitors:

• small proportion of annual production devoted to reproduction,

• perennation by dormant buds and diaspors

• photosynthetic activity correlated with vegetative growth, etc.

Contd.

Interspecific competition occurs with various intensities.

The following are types of Interspecific competitions, in order
of increasing intensity:

i. competition for empty space

ii. competition for space, and
iii. competition for resources in short supply.
 Contd.

• A typical example of competition for empty space occurs

during the invasion of bryophytes and lichens on fallen,
decorticated logs;

• Competition between plants for space occurs as neighbor

interactions or localized competition.

• Competition for resources in short supply has been

demonstrated for several plant groups in laboratory
experiments. However, the importance of this kind of
competition in natural vegetation is hard to assess.
Contd.

2) Destabilizing factors

Destabilizing factors may be divided into two categories:

• Disturbance, and
• Fluctuations
The word ‘disturbance’ is used in 2 different ways in the literature
of plant ecology.
The first use covers:
• various accidental injuries suffered by mature or seedling
plants (e.g. landslide; burial by sand; fire, or storm damage)
• defoliation by herbivores. or
• intervention by humans (e.g. ploughing arable land; road
construction, etc.)

This usage is defined as “the mechanisms which limit the plant

biomass by causing its partial or total destruction”
 Contd.

• The second usage covers processes leading to the creation of

bare ground, loose soil, or light gaps, which constitute micro-
sites in which recruitment can occur.

• The most important aspects of the disturbance regime are:

- its spatial scale (the characteristic patch size of

disturbance)
- the frequency of disturbance
- the intensity of the disturbance
 Contd.

Specific adaptations related to disturbance and micro-site

creation can be summarized in two categories:

The first is,

• morphological, phenological or physiological attributes that
increase the probability of getting into (and staying in) a
newly created micro-site; and

• traits that enable a plant to resist disturbance, and to keep

other individuals out,
 Contd.

The second set of disturbance-related traits concern the ability

of an individual plant to keep others out.

e.g. tolerance of factors such as fire, grazing,

trampling and drought enable a plant to maintain
its tenure of a site in the face of severe disturbance
.
Contd.

Other traits may enable an established individual to exclude

potential invaders either by:

– casting a dense shade,

– allelochemical effects,

– the production of deep persistent leaf litter,

– harboring high densities of invertebrate

herbivores that kill seedlings beneath the canopy,
– dessicating the soil surface by virtue of a shallow,
but highly competitive root system.
 Contd.

• Disturbance :

- encompasses sudden events impacting the vegetation by

increasing the density-independent mortality.

• Fluctuations :
- are used to denote more or less predictable variation in
environmental factors affecting density-independent mortality.
 Contd.

Plants with high tolerance to disturbance are termed ruderal.

Ruderals share a lot of characteristics:
– Short life span,
– High frequency of sexual reproduction,
– Large proportion of annual production devoted to vegetative
growth,
– Perennation by dormant diasporas, and
– Potentially high photosynthetic activity, coincident with
periods of high potential productivity,
 Contd.

The most conspicuous features of plants from low-nutrient

environments are:
– Their small size,
– Their tendency to have small, leathery, long-lived leaves, and
– Their high shoot:root ratios;
 Contd.

3) Stress
• Stress is a commonly occurring environmental impact on
vegetation that adversely affects production, mortality or
establishment.

Grime (1979) defines stress as:

“the external constraints which limit the rate of dry

matter production of all or part of the vegetation”
 Contd.

As opposed to disturbance and fluctuation, stress constantly

impacts the plants.
Stress tolerators are characterized by:
– Long or very long life spans
– Long-lived, often perennial leaves (or other photosynthetic
parts),
– Ever-greenness
– Small proportion of annual production devoted to
reproduction,
– Slow growth,
 Contd.

Stress is connected with end-points of environmental gradients: e.g.

– temperature stress increases towards high altitudes in the
mountains, leading to gradual reduction in species number
with increasing altitude;

– toxicity stress may occur near the high pH end of a nutrient

gradient;
 Contd.

 Stress reduces competition by

- inhibiting dominants from monopolizing critical

resources,
- reducing shading and probably also
- reducing diffuse competition.
 Contd.

4) Chances
– Chance factors, that is random processes in vegetation,
have important contributions in determining vegetation
structure.

– When species responses to environmental gradients are

recorded, only 50-90% of the variation in species
abundances is normally accounted for as response to
gradients.
 Contd.

• The rest represents random or apparently random variation,

often termed noise.

Factors contributing to randomness are:

– the intraspecific genetic variability, manifest in

differences in life history parameters (flowering,
biomass, reproduction), and, for instance,
» success of pollination and
» dispersal