Air, Soil and Water Research
ORiginAL ReSeARch
Open Access
Full open access to this and
thousands of other papers at
http://www.la-press.com.
estimates of Biomass and Fixed carbon at a Rainforest
in panama
Reinhardt Pinzón1, José Fábrega1, David Vega1, erick n. Vallester1, Rafael Aizprúa2,
Francisco R. López-Serrano3 Fred L. Ogden4 and Kleveer espino1
1
centro de investigaciones hidráulicas e hidrotecnicas (cihh) de la Universidad Tecnológica de Panamá (UTP), Panama
city, Panama. 2Flora Tropical, Villa Zaita, Las cumbres, calle circunvalación, Panama city, Panama. 3Universidad de
castilla La Mancha, campus Universitario, Albacete, Spain. 4University of Wyoming, Laramie, WY USA.
corresponding author email: reinhardt.pinzon@utp.ac.pa
Abstract: This paper presents an estimation of the quantity of carbon ixed in trees in a one hectare (ha) plot at the Cerro Pelado-Gamboa
Hydrology Tropical Observatory, which is located in the province of Colon, Panama. The estimation of carbon ixed in trees may provide signiicant information on carbon lux due to water circulation, which may ultimately enable evaluation of the carbon cycle. All
trees larger than 10 cm diameter at breast height (DBH) in the plot were investigated. Carbon ixed within these trees was estimated
using a parameterized formula. Tree biomass estimations for the plot were 97.21 Mg ha−1. We identiied a rare arboreal pear species
(Euphorbiaceous) with higher carbon density than other trees in the plot. The presence of this apparently unique species may be due to
speciic soil characteristics. The method was evaluated by comparing the results with a second study performed in 2011, which resulted
in an estimate of net new carbon (biomass) increment (NNCI), which gives 3.88 Mg ha−1 year−1. In general, the estimation of the biomass and associated carbon content found in this investigation are useful comparative data for economic evaluation of tropical forests
in terms of capacity to capture carbon.
Keywords: biomass, carbon, climate change, net new carbon increment, Panama, rainforest, re-growth
Air, Soil and Water Research 2012:5 79–89
doi: 10.4137/ASWR.S9528
This article is available from http://www.la-press.com.
© the author(s), publisher and licensee Libertas Academica Ltd.
This is an open access article. Unrestricted non-commercial use is permitted provided the original work is properly cited.
Air, Soil and Water Research 2012:5
79
Pinzón et al
Introduction
The Kyoto Protocol, established during the United
Nations Framework Convention on Climate Change
(UNFCCC) in 1997,1 spotlighted the need for a
quantitative understanding of the global capacity
of forests to capture and ix carbon. During the
convention, shortcomings in data were identiied. The
cause of these shortcomings was principally the lack
of standardized algorithms for estimating the volume
of woody biomass in forests.
Schlegel et al2 deined the major source of carbonixing biota and ixed-carbon storage in forests as
above-ground biomass (AGB). Several methods have
been proposed for determining AGB, which vary signiicantly in precision. Chave et al3 compared simple
allometric models and proposed speciic estimating methods for four tropical ecosystems (dry forest, humid forest, swamp, and very humid forest)
and developed two equations for each type of forest,
one using the total height of individual trees and the
other for when tree height is not known (in addition to
other variables). The estimating methods proposed by
Chave et al3 are well accepted as having reasonable
precision. This latter study gave a re-evaluation of
the quality and the robustness of these models across
2000 tropical trees harvested in 27 sites. The analytical power of these models depends on how well they
are validated using tree biomass data obtained directly
from destructive crop experiments. The validation
was done based on a large dataset collected at sites
from dry woodlands to hyperhumid closed-canopy
forest and secondary to old-growth forest.4 The goodness of it of the Chave et al model was measured by
the residual standard error of the it and by a penalized
likelihood criterion. This model estimated accurately
the above-ground biomass at most sites; however, the
authors included different kinds of forest, such secondary and old-growth forest. Wood speciic gravity
was a key predictive variable in all these models. Its
signiicance may not be clear if one is interested in
guessing the biomass in an old-growth forest dominated by hardwood species, spanning a narrow range
of wood densities. Baker et al5 have shown that disregarding differences in wood density should result
in poor overall calculation biomass of the above
ground stand. Finally, regression models should not
be used beyond their range of validity.3 The methods
employ basal diameter or diameter at breast height
80
(DBH), the wood speciic density, and the total height
of each individual. Because variables such as wood
density vary between species and region, it is important to choose an appropriate model.
In Panama, there are few studies that measure carbon luxes in humid tropical rainforests or that measure the amount of carbon present in the biomass.
Very little is known about tropical forest biomass, and
the few studies that are available have covered small
areas because they required destructive sampling of
the forest.6 Forests in Panama contain approximately
4800 species of trees and shrubs.7
The main objectives of the study reported in this
paper are (1) to evaluate the quantity and diversity of
tree species, (2) to determine the actual biomass and
carbon storage, and (3) to estimate the net new carbon (and biomass) increment in this secondary forest
stand.
Method
Study area
The study was conducted at Cerro Pelado-Gamboa
(S.1),8 Colon Province, Panama (9°7′28″N,
79°42′9″W), at an elevation of approximately 160 to
200 m above sea level. For more than 70 years, the
US Army used this place as a pilot site for materials,
equipment, medicines, and for evaluating conditions
in a humid tropical forest. On September 23, 2002,
Resolution No. 115-02, ARI (Autoridad de la
Región Interoceánica) attached to the Universidad
de Panamá (UP) and the Universidad Tecnológica
de Panamá (UTP), extended for 20 years the use
and administration of former Gamboa Tropical Test
Center, covering an area of 751.45 hectares, for the
development of scientiic and technological research
on a tropical environment. During early Panama
Canal operations, Cerro Pelado was clear-cut and can
now be considered well-established second growth.
There is little old-growth forest to be found in the
Canal corridor due to human activity that greatly
accelerated after 1870.
cerro Pelado-gamboa climate
Movement of the inter-tropical convergence zone
(ITCZ) and topography are more or less responsible
for the rainfall patterns in Gamboa and, indeed, all of
Panama. Broadly, the ITCZ is responsible for the rainy
seasons closely associated with tropical rain forests.
Air, Soil and Water Research 2012:5
Estimates of biomass and ixed carbon
In Panama, the rainy season generally persists from
May until December and the dry season from late
December until late April. Panama additionally has
a rainfall gradient from the Caribbean to the Paciic,
with the Atlantic side receiving signiicantly more
rainfall. The Caribbean receives much more rainfall
because the moisture-laden trade winds are primarily oriented from the northeast during the dry season.
Gamboa is located in almost exactly the middle of the
isthmus and exhibits a marked seasonality in rainfall
with an average annual value of 2148 mm.
A detailed climatologically record exists at
Gamboa because of Panama Canal operations with
rainfall records dating back as far as 1897. Gamboa
receives less than 20 mm of rainfall during the months
of February and March. October is the wettest month
receiving 306 mm of rain on average. In Panama,
there is frequently a period of slightly decreased rainfall in July and August known as the veranillo de San
Juan. Veranillo de San Juan can last anywhere from
one to four weeks and is generally accompanied by an
increase of wind speed,9 but may not be as prevalent
in Gamboa. The temperature variability is remarkably small throughout the year with a mean temperature of 26.4 °C. The relative humidity increases and
solar radiation decreases as would be expected with
the onset of the wet season. Interestingly, the average daily wind speed decreases dramatically with the
onset of the wet season.10
cerro Pelado-gamboa geological
features
Panama Isthmus is located on a complex tectonic
plate, which rests on a microplate named Block of
Panama. On this microplate, four lithospheric plates
concur: the Caribbean located to the north, Cocos to
the southwest, Nazca to the south, and South America
to the east and southeast. The Block of Panama’s geographic location between both continents and two
oceans has been important for both scientiic and
economic reasons. Historically, the Panama Isthmus
was considered a commercial path of goods that used
to come from South America and Europe due to the
Spanish conquest during the sixteenth and seventeenth centuries. Following the successful interoceanic canal construction through the isthmus, Panama
became a major naval route for the world. Therefore,
it was necessary to dam the Chagres River. The Cerro
Air, Soil and Water Research 2012:5
Pelado-Gamboa area is located on a sharp hillslope
on this river. The Cerro Pelado areas are affected by
the Gatuncillo Formation, which is covered by clay
shales, lutites, quartz sandstone, algaelike, and foraminifer limestone. The Gatuncillo Formation is a
common geological formation of the Superior Middle
Eocene (S.2).11 Pre-Terciary is another geologically
important formation that predominates in this area
where lavas, basaltic tuff, and andesites are present
and intrusive dioritic and dacitics rocks are found.8
Half of the Panama Canal Basin has been deforested, and the oficial policy in this area is to reforest in anticipation of regaining ecosystem services.
Land cover and its manipulation can have collateral
impacts, both positive and negative. The positive
impacts are referred to as ecosystem services, which
include carbon storage, water quantity and quality,
and biodiversity. Some negative impacts are water
wasting, soil erosion, and wildires.12 Basically, the
Cerro Pelado area in Gamboa is formed by mature
secondary forest. Moreover, inside the Panama Canal
watershed there are different kinds of forests and uses,
such as disturbed forest, mature forest, mangroves,
agricultural uses, and so on (S.3).13
A topographic map with contours of the Cerro
Pelado-Gamboa area and the study plot are shown in
S.4.13 Basically, the west side of the plot area shows
a smooth slope, while the east side is more steeply
sloped. Topography of the zone is very varied.
Practically lat lands (0% to 3% slope), slightly wavy
hills (3% to 15% slope), and lands that are markedly
inclined, with slopes between 30% and 60%. The
topography of the area where this investigation was
carried out show contour lines ranging from 40 meters
to 154 meters.13 True triple canopy conditions typical
of pristine tropical catchments are not found on Cerro
Pelado, with the situation more accurately described
as a two and a half layer canopy. Trees reaching 25 m,
with several larger trees reaching 35 m, dominate
the average canopy height. By far the most prevalent trees are various species of palm combined with
emerging deciduous trees form the secondary canopy
layer. Cerro Pelado-Gamboa is reachable from a path.
From the top of the path, the hill immediately rises
up to a point that forms two lat saddle areas before
again rising to the highest peak formally known as
Cerro Pelado. Portions of the saddles are remarkably
lat given the topography.10
81
Pinzón et al
Forest inventory
One hectare (ha) of forest was established as a permanent
plot for long-term studies. This permanent plot allows
for the estimation of the parameters for carbon content
analyses, the collection of data on the behavior and
distribution of the species that are found in the plot, and
modeling of future changes.14 The long axis of the 1-ha
plot is oriented toward magnetic north to ease placement
of markers. Within the plot, a grid of 25 quadrants of
20 m × 20 m was marked. In turn, every 20 m × 20 m
area was subdivided into 4 sub-quadrants of 10 m × 10 m
and into 16 sub-quadrants of 5 m × 5 m. Compasses
were used to delimit the plot. Conventional 50 m metric
tapes were used to measure distances. Each 20 m corner
was marked by a 1.9 cm polyvinyl chloride (PVC) pipe
with permanent markings and pink lagging. The 10 m
corners were marked with 1.3 cm PVC pipe. The 5 m
corners were marked with orange lagging. Within each
plot, trees selected for measurement (DBH $ 10 cm)
were marked with aluminum tags ixed to the tree with
aluminum nails.
DBH was taken of the principal trunk at a height
of 1.30 m from the level of the adjacent soil or base
of the tree. In the case that the trunk was irregular at a
height of 1.30 m (due to knots or protruding growths),
a measurement was taken at the point where the trunk
became more cylindrical.
The four different measurements taken for each
tree with DBH $ 10 cm were total height, commercial height, crown deep, and crown diameter. These
are described below.
1. Total height is the vertical distance from the level
of the adjacent soil up to the highest point where
the canopy of the tree is projected.
2. Commercial height is the measurement from the
level of the soil (base of the tree) up to the point
where the lowest branches occur.
3. Canopy height is the difference between the commercial height and the total height.
4. Canopy radius (r) is the distance from the center or imaginary axis of the silhouette generated
by the canopy of a tree towards any point of the
perimeter of this circumference. For practical reasons r = D/2, where D represents the diameter of
a tree of perfectly circular canopy. The radius of
canopy is measured using a conventional longitudinal tape.
82
For trees with more than one trunk smaller 1.30 m,
the principal trunk diameter was measured if the
tree’s DBH was greater than 10 cm. In this case, the
tree was deined as a multiple stem, and additional
parameters including total height, commercial height,
deep and diameter of crown were taken, as established
by Schlegel et al.2 Two inventories were performed.
The irst inventory was done in February 2008 and
the second one in February 2011. The commercial
height and the total height were measured using an
electronic clinometers model Haglöf (HEC-MD). All
the individuals were identiied in situ. When doubts
existed about a species’ identiication, voucher specimens were collected for evaluation by the University of Panama botanist. More than 95% of the trees
were successfully identiied in situ. To determine the
species, each sample was examined meticulously
when the trees were tall to assure the identiication.
Above- and below- ground biomass
estimation
Estimations of above-ground dry biomass (AGB)
followed the methodology proposed by Brown and
Lugo6 based on allometric regression equations that
were itted for tropical ecosystems by Chave et al.3
These allometric equations were applied for species
and for each inventory and then added to obtain the
total biomass per plot. In the case of humid tropical forests with individual height data, the following
equation 1 was used.
(AGB)est = exp (−2.977 + ln (ρD2H))
≡0.0509 x ρD2H
(1)
where AGB is above-ground dry biomass (given
Mg ha−1), D is the stem diameter, ρ is the speciic
density of the wood,15 and H is the total height of the
individual tree. Values for each specie’s wood speciic
gravity (density) were obtained from Chave et al.15
and FAO.1 When a value for the speciic density of
any species was not listed in the literature, a standard
value of 0.60 g/cm3 proposed by FAO16 for the ecosystems of America was used.
The direct determination of the below-ground
biomass (BGB) traditionally has been carried out
through destructive methods, which are typically
labor-intensive and inappropriate for forests within
natural reserves. In this study, BGB was estimated
Air, Soil and Water Research 2012:5
Estimates of biomass and ixed carbon
using AGB estimates. To determine BGB, AGB was
multiplied by a coeficient established by Cairns et al.17
The coeficient used was a root/shoot ratio of 0.24 for
tropical forests, indicating 24% of radicular (root) biomass compared to above ground biomass. The fraction of carbon contained in the total dry biomass is an
additional parameter. Values typically used range from
45% to 53%, but a considerable number of countries
use 50% value by default. Although the average heavy
value of the forest is 49%,18 in our estimations we used
50% as this is the value more commonly used.
net new carbon (biomass)
increment (nnci)
The net new carbon increment (NNCI) is that part of the
gross primary productivity (GPP) (integrated over the
annual period) that is retained by the vegetation as
new growth. The NNCI is used to estimate the carbon
stock in the canopy.19 Therefore, this concept differs
from the net primary productivity (NPP) that is usually
mentioned because it does not take into account
losses other than autotrophic respiration.20 However,
if the time scale considered was short, then we could
accept both concepts to be comparable. Moreover,
the NNCI is estimated by the difference (increment)
between total biomass (carbon) for each inventory
and divided for the period of time among them. In
the present study, the NNCI was obtained from the
comparison between the 2008 and 2011 inventories,
that is, NNCI = (C2011 − C2008)/3.
Result and Discussion
Forest inventory in 2008
Within the 1 ha plot studies, a total of 384 individual
trees with DHB $ 10 cm were identiied (S.5). A total
of 40 species of trees belonging to 31 families were
identiied of which the family Fabaceae/Mimosoidea
(4 species) contributed the largest number of species
to the inventory. The species Pera arborea Pear
(Euphorbiaceous) was the most common individual
with 80 individuals, followed by Oenocarpus mapora
(Arecaceae) with 65 individuals and Amaioua
corymbosa (Rubiaceae) with 60 individuals. Mean
diameter was 20.07 cm, average total height was
19.10 m, average commercial height was 12.42 m,
and average crown radius was 2.77 m. Forty multiple
stems were measured corresponding mainly to clonal
palm Oenocarpus mapora (Arecaceae) with 38 stems.
The two remaining multiple stems included an Pera
arborea Pear (Euphorbiaceae) species and a Roupala
montana (Proteaceae). Regarding the distribution of
sizes of 384 trees mapped, 249 had a DBH between 10
and 20 cm, 70 trees had a DBH between 20 and 30 cm,
and 65 trees had a DBH greater than 30.0 cm (S.6).
Biomass estimation in 2008
The total estimated dry biomass (Table 1) calculated
the amount of carbon present in the 1-ha study plot by
using equation (1). The species that contributed the
greatest biomass in the study plot were arboreal Pera
arborea with 45.72 Mg ha−1 (AGB) and 10.97 Mg ha−1
(BGB), Enterolobium schomburgkii with 25.29 Mg ha−1
(AGB) and 6.07 Mg ha−1 (BGB) and Vantanea depleta
with 19.38 Mg ha−1 (AGB) and 4.65 Mg ha−1 (BGB).
The AGB of trees found in the Cerro Pelado plot compared favorably with other forests in Panama and
Costa Rica, as indicated in (S.7).16,21,22
Forest inventory in 2011
The inventory was repeated after the initial study.
The methodology was identical to that used in the
irst study. An added beneit was that the results from
the second estimation were compared with the irst,
allowing a irst time quantiication of the net new carbon increment (NNCI).
The total estimated dry biomass calculated the
amount of carbon present in the 1 ha study plot (S.8).
Calculated biomass changed from 194.43 Mg ha−1 to
217.73 Mg ha−1, and increased by 10% from 2008 to
2011. The species that contributed the greatest aboveground biomass in the study plot were arboreal Pear
(27.16%), Enterolobium schomburgkii (15.77%), Vantanea
Table 1. Above-ground biomass (AgB), below-ground biomass (BgB), total biomass and total carbon in the 1 ha plot, 2008.
site
AGB
(Mg ha−1)
BGB
(Mg ha−1)
Total biomass
(Mg ha−1)
carbon total
(Mg ha−1)
cerro Pelado
Panama
156.80
37.63
194.43
97.21
Air, Soil and Water Research 2012:5
83
Pinzón et al
Table 2. nnci in different localities.
site of study
nncI
(Mg c ha1 y1)
Type of forest
Life zone
cerro peladoa
Magdalena terrace and slope, colombiab
Limon, costa Ricac
3.88
2.6–2.2
3.1
Secondary
Secondary
Secondary
Tropical rainforest
Tropical evergreen
Tropical rainforest
a
nnci present study; bFölster et al;23 cchacon et al.24
depleta (12.30%), Matayba apetala (10.76%),
and Oenocarpus mapora (3.99%) (S.9). The rest
of the species combined contribute 30.01% of the
175.59 Mg ha−1 above ground biomass estimated (S.10).
net new carbon (biomass)
increment (nnci)
The results from the second estimation were compared
with the irst, allowing a quantiication of net new
carbon (biomass) increment (NNCI) (Table 2).23,24
An NNCI value of 3.88 Mg ha−1 year−1 was obtained
using NNCI = (C2011 − C2008)/3 = (108.86–97.21)/3.19
NNCI, 3.88 Mg ha−1 year−1, is larger compared
with those obtained by Fölster et al (2.2–2.6 Mg ha−1
year−1, Table 2)23 whose study that was conducted in
a secondary rainforest in Colombia. This discrepancy may be related to the presence of Pera arborea
Pear species (Euphorbiaceous) as predominant species that have higher carbon density than the other
trees in the plot due to speciic soil characteristics.
Because the maximum NNCI value reported by
Fölster et al23 was 3.8 Mg ha−1 year−1 in a primary
rainforest, our larger NNCI value may suggest that
our study area has similar characteristics as a primary rainforest. In addition to the larger NNCI value
compared with a primary rainforest, López-Serrano25
demonstrated that NNCI value tends to be smaller
soon after the forest ire, which means that the initial stage of a secondary rainforest may have similarly small NNCI. Therefore, our larger NNCI value,
3.8 Mg ha−1 year−1, may be expected not to be in the
initial stage, which also suggests that the characteristics of our study area has reached a steady state, like
a primary rainforest.
conclusions
The estimation of the biomass and associated carbon content found in this investigation are useful
84
comparative data for economic evaluation of tropical
forests in terms of capacity to capture carbon.
A rare Pera arborea Pear species (Euphorbiaceous)
with higher carbon density than other trees in the plot
was found. The presence of this apparently unique
species may be due to speciic soil characteristics.
A irst estimation of the NNCI gives 3.88 Mg ha−1
year−1. Moreover, our larger NNCI value may suggest
that our study area has similar characteristics as a primary rainforest. Therefore this comparative study can
be used as baseline for new future research developments at Cerro Pelado-Gamboa that will allow Pera
arborea there ine NNCI values for the 750 ha for
over longer time periods.
Acknowledgments
We would like to acknowledge the editorial services
provided by Sustainable Sciences Institute (SSI), in
particular the revisions and comments made by Eng.
Dana Brock an SSI volunteer. Also, the authors would
like to acknowledge Dr. Keisuke Nakayama from
Kitami Institute of Technology in Japan for his wise
comments and suggestions. Furthermore, we express
our gratitude to the editor and the two anonymous
referees of this paper for their beneicial comments.
Moreover, thanks to all CIHH members who contributed and helped carry out this project to its successful
completion.
Author contributions
Conceived and designed the experiments: RA, RP,
FRLS. Analysed the data: RA, RP, DV, FRLS, KE.
Wrote the irst draft of the manuscript: RP. Contributed to the writing of the manuscript: JF, RA, and
FRLS. Agree with manuscript results and conclusions:
RP, JF, DV, ENV, KE, RA, FRLS, FLO. Jointly
developed the structure and arguments for the paper:
RP, JF, FRLS. Made critical revisions and approved
Air, Soil and Water Research 2012:5
Estimates of biomass and ixed carbon
inal version: FRLS, FLO. All authors reviewed and
approved of the inal manuscript.
Funding
The authors are grateful for the inancing granted
by Secretaría Nacional de Ciencia y Tecnología e
Innovación (SENACYT) (endorsed Proposal—code
COL07-011). This research was partially supported
by the US Army Research Ofice through grant
W911NF-07-1-0389 to the University of Wyoming.
This contribution has been co-funding by Spanish
government (MICINN, AGL2011-27747 research
project) and FEDER funds.
competing Interests
Author(s) disclose no potential conlicts of interest.
Disclosures and ethics
As a requirement of publication author(s) have
provided to the publisher signed conirmation
of compliance with legal and ethical obligations
including but not limited to the following: authorship
and contributorship, conlicts of interest, privacy and
conidentiality and (where applicable) protection of
human and animal research subjects. The authors
have read and conirmed their agreement with the
ICMJE authorship and conlict of interest criteria.
The authors have also conirmed that this article is
unique and not under consideration or published in
any other publication, and that they have permission
from rights holders to reproduce any copyrighted
material. Any disclosures are made in this section.
The external blind peer reviewers report no conlicts
of interest.
References
1. Organización de las Naciones Unidas para la Agricultura y la Alimentación
(FAO). FRA 2000, Directrices para la evaluación en los países tropicales
y subtropicales. Roma, Italia: Organización de las Naciones Unidas para la
Agricultura y la Alimentación, Departamento de Montes; 1998.
2. Schlegel B, Gayoso J, Guerra J. Manual de Procedimientos para Inventarios
de Carbono en Ecosistemas Forestales. Valdivia, Chile: Universidad Austral
de Chile; 2001. Proyecto FONDEF D98I1076.
3. Chave J, Andalo C, Brown S, et al. Tree allometry and improved estimation of
carbon stocks and balance in tropical forests. Oecologia. 2005;145:87–99.
4. Sherman RE, Fahey TJ, Martinez P. Spatial pattern of biomass and aboveground net primary productivity in a mangrove ecosystem in the Dominican
Republic. Ecosystems. 2003;6:384–98.
5. Baker TR, Phillips OL, Malhi Y, et al. Variation in Wood density determines spatial patterns in Amazonian forest biomass. Glob Change Biol.
2004;10:545–62.
Air, Soil and Water Research 2012:5
6. Brown S, Lugo A. Biomass of tropical: a new estimate based on forest
volumes. Science, New Series. 1984;223(4652):1290–3.
7. Correa A, Mireya D, Galdames C, Stapf M. Catálogo de las Plantas
Vasculares de Panamá. Universidad de Panamá, Instituto Smithsonian de
Investigaciones Tropicales. República de Panamá: Editora Novo Art; 2004.
8. Mojica A. Private Communication, 2009.
9. Espinosa J. Veranillo de San Juan within the Panamá Canal Watershed.
Balboa Heights, Panamá: Panamá Canal Commission; 1998.
10. Niedzialek JM. Unusual Hydrograph Characteristics, Upper Río Chagres,
Panama, 2007 [dissertation]. Storrs: University of Connecticut; 2007.
11. Stewart RH, Stewart JL, Woodring WP. Geologic Map of Panama Canal
and Vicinity, Republic of Panama. Washington, DC: Department of the
Interior, United States Geological Survey; 1980.
12. Stallard RF, Ogden FL, Elsenbeer H, Hall J. Panama Canal Watershed
Experiment: Agua Salud Project. Water Resources IMPACT. 2010;12(4):
18–20.
13. Serrano E, Nuñez M. Cuantiicación de Flujo de CO2 en suelo [Undergraduate
thesis]. Panama City: Universidad Tecnológica de Panamá; 2009.
14. Burslem D, Garwood NC, Thomas SC. Tropical forest diversity: the plot
thickens. Science. 2001;291(5504):606–7.
15. Chave J, Condit R, Lao S, Caspersen J, Foster R, Hubbell S. Spatial and
temporal variation of biomass in a tropical forest: results from a large census
plot in Panama. J Ecol. 2003;91:240–52.
16. Food and Agricultural Organization of the United Nations. Estimating
Biomass and Biomass Change of Tropical Forest—a Primer. Rome, Italy:
Food and Agricultural Organization of the United Nations; 1997. FAO Forestry Paper No. 134.
17. Cairns M, Brown S, Helmer EH, Baumgardner GA. Root biomass allocation
in the world´s upland forest. Oecología. 1997;111:1–11.
18. De Vries W, Reinds GJ, Posch M, et al. Intensive Monitoring of Forest
Ecosystems in Europe. Technical report. Brussels, Belgium: United Nations
Economic Commission for Europe; 2003.
19. Berry S, Roderick M. Changing Australian vegetation from 1788 to 1988:
effects of CO2 and land-use change. Aust J Bot. 2006;54:325–38.
20. Melillo JM, McGurie AD, Kicklighter DW, Moore IIIB, Vorosmarty CJ,
Schloss AL. Global climate change and terrestrial net primary production.
Nature. 1993;363:234–40.
21. Arcia D, Garibaldi C. Los bosques, bienes y servicios ambientales de la
Reserva Forestal El Montuoso, provincia de Herrera, Panamá. In: Garibaldi C,
editor. Diversidad Biológica y Servicios Ambientales de los Fragmentos de
Bosques en la Reserva Forestal el Montuoso, Panamá. Ciudad de Panamá:
Universal Book; 2004:173–93.
22. Drake JB, Knox RG, Dubayah RO, et al. Above-ground biomass estimation in closed canopy Neotropical forest using lidar remote sensing: factors affecting the generality of relationships. Glob Ecol Biogeogr. 2003;12:
147–59.
23. Fölster A, de las Salas G. A tropical evergreen forest site with perched
water table, Magdalena Valley, Colombia: biomass and bioelement inventory of primary and secondary vegetation. Oecologia Plantarum. 1976;11:
297–320.
24. Chacón P, Leblanc HA, Russo RO. Fijación de carbono en un bosque
secundario de la región tropical húmeda de Costa Rica. Tierra Tropical.
2007;3(1):1–11.
25. López-Serrano FR, De Las Heras J, Moya D, García-Morote FA, Rubio E.
Is the net new carbon increment of coppice forest stands of Quercus ilex
ssp. ballota affected by post-ire thinning treatments and recurrent ires? Int
J Wildland Fire. 2010;19(5):637–48.
85
Pinzón et al
supplementary Materials
200 km
9,118473°
Caribbean
Sea
1 km
Gamboa
-79,695692°
Cerro
Pelado
Charge
river
s.1. cerro Pelado-gamboa area geographic location, Panama canal watershed in Mojica et al.8
s.2. geological map of cerro Pelado-gamboa, its surroundings and study area from Stewart et al.11
86
Air, Soil and Water Research 2012:5
Estimates of biomass and ixed carbon
s.3. Land use map, cerro Pelado-gamboa in the Panama canal watershed, from Serrano et al.13
s.4. Topographic map of cerro Pelado-gamboa with contour lines and the plot area grid from Serrano et al.13
Counts
249
70
>10 < 20 cm
>20 < 30 cm
65
> 30 cm
DBH
s.6. Tree size distribution of diameter at breast height (DBh) of 384 trees in the plot cerro Pelado-gamboa, 2008.
Air, Soil and Water Research 2012:5
87
Pinzón et al
s.5. Trees by family and specie with DhB $ 10 cm, 2008.
Family
specie
Frequency
Annonaceae
Arecaceae
Bignoniaceae
Xylopia frutescens
Oenocarpus mapora
Jacaranda copaia
Tabebuia guayacan
Pachira sessilis
Cordia panamensis
Protium panamense
Hirtella americana
Calophyllum longifolium
Terminalia amazonia
Diospyros artanthifolia
Erythroxylum macrophyllum
Pera arborea
Tachigali versicolor
Abarema barbouriana
Enterolobium schomburgkii
Inga pezizifera
Inga thibaudiana
Vatairea erythrocarpa
Lindackeria laurina
Vantanea depleta
Lacistema aggregatum
Beilschmiedia pendula
Nectandra purpurea
Ocotea cernua
Byrsonima spicata
Henriettella tuberculosa
Guarea guidonia
Perebea xanthochyma
Maquira guianensis
Virola sebifera
Virola multilora
Myrsine coriacea
Myrcia gatunensis
Roupala montana
Alseis blackiana
Amaioua corymbosa
Matayba apetala
Ternstroemia tepezapote
Vochysia ferruginea
2
65
6
3
6
2
5
2
2
2
1
1
80
3
1
9
6
3
5
7
24
1
1
1
1
1
1
1
15
1
16
2
1
5
5
1
60
22
8
6
Bombacaceae
Boraginaceae
Burseraceae
chrysobalanaceae
clusiaceae
combretaceae
ebenaceae
Erythroxylaceae
euphorbiaceae
Fabaceae/cae.
Fabaceae/Mim.
Fabaceae/Pap.
Flacourtiaceae
humiriaceae
Lacistemataceae
Lauraceae
Malpighiaceae
Melastomataceae
Meliaceae
Moraceae
Myristicaceae
Myrsinaceae
Myrtaceae
Proteaceae
Rubiaceae
Sapindaceae
Theaceae
Vochysiaceae
s.7. Above-ground biomass (AgB) in different localities, 2008.
site of study
AGB
(Mg ha1)
Type of forest
Life zone
cerro Peladoa
Reserva Forestal Montuosob
Reserva Forestal Montuosob
156.80
163.00
235.50
Secondary
Secondary
Primary
Panama (general)c
169–945
Secondary
Zona del canal, Panamad
Barro colorado, Panamad
La Selva, costa Ricad
La Selva, costa Ricad
277.91
286.77
160.00
147.70
Secondary
Primary
Primary
Secondary
Tropical rainforest
Broadleaved
Semideciduous transition lowland rainforest
Broadleaved
Semideciduous transition lowland rainforest
Tropical rainforest
Tropical rainforest
Tropical rainforest
Very humid tropical forest
Very humid tropical forest
a
Present study; bArcia and garibaldi;21 cFAO;16 dDrake et al.22
88
Air, Soil and Water Research 2012:5
Estimates of biomass and ixed carbon
s.8. Above-ground biomass (AgB), below-ground biomass (BgB), total biomass, and total carbon in the 1 ha plot, 2011.
site
AGB
(Mg ha1)
BGB
(Mg ha1)
Total biomass
(Mg ha1)
carbon total
(Mg ha1)
cerro Pelado
Panama
175.59
42.14
217.73
108.86
s.9. Main biomass contributions by species in the 1 ha plot at cerro Pelado-gamboa, 2011.
specie
AGB
(Mg ha1)
BGB
(Mg ha1)
Pera arborea
Enterolobium schomburgkii
Vantanea depleta
Matayba apetala
Oenocarpus mapora
47.69
27.69
21.60
18.90
7.01
11.45
6.65
5.18
4.54
1.68
s.10. Biomass values from other species present in the plot, cerro Pelado-gamboa, 2011.
specie
AGB
(Mg ha1)
BGB
(Mg ha1)
Total
(Mg ha1)
Abarema barbouriana
Alseis blackiana
Amaioua corymbosa
Beilschmiedia pendula
Byrsonima spicata
Calophyllum longifolium
Cordia panamensis
Diospyros artanthifolia
Erythroxylum macrophyllum
Guarea Guidonia
Henriettella tuberculosa
Hirtella Americana
Inga pezizifera
Inga thibaudiana
Jacaranda copaia
Lacistema aggregatum
Lindackeria laurina
Maquira guianensis
Myrcia gatunensis
Myrsine coriacea
Nectandra purpurea
Ocotea cernua
Perebea xanthochyma
Pachira sessilis
Protium panamense
Roupala Montana
Tabebuia guayacan
Tachigali versicolor
Terminalia Amazonia
Ternstroemia tepezapote
Vatairea erythrocarpa
Virola sebifera
Virola multilora
Vochysia ferruginea
Xylopia frutescens
Totals
0.50
0.41
6.50
0.07
0.29
1.54
1.02
0.08
0.07
0.05
0.20
0.32
1.68
1.70
6.24
0.06
1.34
0.15
0.26
2.35
0.18
0.33
2.03
1.62
0.34
1.52
0.43
1.86
3.21
1.32
9.68
1.39
1.25
2.43
0.26
52.70
0.12
0.10
1.56
0.02
0.07
0.37
0.25
0.02
0.02
0.01
0.05
0.08
0.40
0.41
1.50
0.01
0.32
0.04
0.06
0.56
0.04
0.08
0.49
0.39
0.08
0.37
0.10
0.45
0.77
0.32
2.32
0.33
0.30
0.58
0.06
12.65
0.62
0.51
8.06
0.09
0.37
1.91
1.27
0.10
0.09
0.06
0.25
0.39
2.08
2.10
7.73
0.07
1.67
0.19
0.32
2.92
0.22
0.41
2.52
2.01
0.42
1.89
0.54
2.31
3.98
1.64
12.00
1.73
1.55
3.02
0.33
65.35
Air, Soil and Water Research 2012:5
View publication stats
89