Abstract
For this project, the sequestration and storage
capacity of carbon were quantified in canopy trees along with their
epiphytes in a fragment of a cloud forest where oak trees - Quercus
humboldtii - represent the largest part of the whole forest. It
was found that each of the components of the epiphyte-host tree
system shows different percentages of C concentration. In the case of
Oak trees, it was found that the branches with diameters lower than 5
cm. have the largest capacity in capturing C (40.13%), followed by
the wood contained in the trunk and in branches with diameters higher
than 5 cm (38.75%) and fresh leaves and dead leaves show (35.95%) and
(34.05%) of C retention respectively. In the case of epiphytes
growing on these trees, it was discovered that the lichens and
bryophytes yielded a 43% of C, being the
component with more capacity of retention of C in the epiphyte -
host tree system; the bromeliads had a capacity of 38.82% also
presenting a bigger capacity of capture of C than that of the
components of their host tree, excepting the upper branches.
As for the C stored in the biomass it was
found that in the four evaluated trees and their epiphytes, there was
32 066 kg in 0.0938 ha.
Key words: Neotropic,
Cloud Forest, Carbon sequestration, Canopy, Oaks, Epiphytes
Resumen
Se cuantificó la capacidad de captura y almacenamiento de
carbono en árboles de dosel y sus epífitas en un
fragmento de bosque de niebla dominado por árboles de Quercus
humboldtii. Se encontró que cada componente del sistema
epífitas - hospederos presenta diferentes porcentajes de
concentración de C, en el caso de los árboles de roble
se obtuvo que las ramas con diámetros inferiores a 5 cm. son
las que mayor capacidad de captura de C presentan (40.13%), seguido
por la madera contenida en el tronco y en las ramas con diámetros
mayores a 5 cm (38.75%), las hojas frescas (35.95%) y la hojarasca
(34.05%). En el caso de las epífitas que crecen sobre estos
árboles se encontró que los líquenes y briófitos
con el 43 % son el componente que más capacidad de retención
de C tuvieron en el sistema epífitas - hospederos; las
bromelias tuvieron una capacidad de 38.82% presentando también
una mayor capacidad de captura de C que la de los componentes de su
hospedero, exceptuando las ramas superiores. En cuanto al C
almacenado en la biomasa se encontró que en los cuatro árboles
evaluados y sus epífitas, hubo 32 066 kg en 0.0938 ha
Palabras claves:
Neotrópico, Bosque de niebla, Captura de carbono, Dosel,
Robles, Epífitas
Introduction
Tropical forests and temperate zones sequester
and store more carbon than any other terrestrial ecosystem does. In
addition, these ecosystems contribute to the 90% of the annual flow
of C between the atmosphere and the ground (Brown et al. 1993,
Dixon et al. 1994). This fact has generated a special concern
about the great importance of these forests as C stock places (Husch
2001), since they store big amounts of greenhouse gases, especially
CO2 (Macera et al. 2000).
In the case of tropical forests, these have
received special attention due to their wide covering on a great part
of the terrestrial surface (Terborgh 1985), the rapid transformation
rate into commercial plantations and pastures (Vitousek et al.
1987), and their contributions to the C cycles as well as their
potential impacts in the global weather (Brown & Lugo 1982). In
the last decades, the amount of C found in land vegetation has
generated a special concern, so different conservation and
reforestation strategies have being posed in order to face this
situation (Macera et al. 2000).
Eleven percent of these tropical forests are
represented by montane and submontane forests. They are spread all
over America, Africa, Southwest of Asia, and Pacific islands
(Doumenge et al. 1995). In America, Montane and submontane
forests are located in Central America, the Caribbean (Labastille &
Pool 1978) as well as in the tropical Andes in the northern part of
South America (UNESCO 1981). In this continent, the biggest extension
of montane forests is located in Peru, followed by Colombia, Bolivia,
Ecuador, and Venezuela.
In Colombian submontane and montane forests, where
generally a co-dominance is presented among several species, it can
be found homogeneous woods dominated by Oaks - Quercus
humboldtii- (Lozano & Torres 1974). These forests are located
in the three Colombian mountain ranges, covering areas from 1 100m up
to 3 450m high (Cavelier et al. 2001). In the past, these
forests covered big areas of land. However, nowadays,
the presence of the oak in Colombia is limited to discontinuous
fragments that put it in certain threat degree (UNESCO 2001).
So far, studies about the capacity of capturing carbon in canopy
trees (Q. humboldtii) and their epiphytes had not been
carried out. Therefore, the following article is an approach to know
the role of the Oak trees as well as their epiphytes that they hold
in the storing process of C; this with the purpose of showing one of
the many environmental services that this ecosystem provides.
Materials and Methods
Study Area
The project was carried out in the Macanal Reserve
located in the eastern Colombian Andean Mountain, in a town called
Bojaca, located 27km far from Bogota. The study area is a montane
cloud forest which is in a precipitous area with steep slopes, and it
is located at 2700m above the sea level. The vegetation presents
different levels of human intervention, and this is shown in the
fragments of mature forests where mature Oak trees (Q.
humboldtii) are the predominant species.
The climatology data of the area is registered by the Acapulco
weather station of the "Instituto de Hidrología,
Meteorología y Estudios Ambientales (IDEAM)". According to
the 13 year-old data analysis (from 1990 to 2002), it has a
precipitation with a bimodal behavior with an annual average of
61.5mm of rainfall; in which the most rainfall months that are March,
April, and May in the first term of the year, and October and
November in the second term. The average annual temperature is about
13º C, with the highest temperature on May with about 13.4º
C and the minimum on July with about 12.6º C. The relative
annual humidity average is about 91.9% for April, June, and July
present the highest percentage with about 93%.
Methods
This study was carried out in an oak forest where
neither wood mining, nor any other kind of forest profit have taken
place. The methods used for measuring the tree volume were not
destructive at all. In fact, in order to measure the trees and the
epiphytes, five platforms were built over the upper surface of the
four Canopy trees - Q. humboldtii- to an average height of
20m and 23m. By means of a simple rope technique and tree-climbing
equipment, it was easy to have access to different parts of the four
chosen trees such the trunk and the crown as well as to the
platforms.
The approximate biomass of each evaluated tree was calculated from
the tree volume and the wood density. For each tree, the trunk and
the branches volume were calculated. For this, the length and the
diameter of each trunk and branch were measured, and after that, we
calculated the volume with the cylinder equation. In addition, the
wood density was obtained from fragments of trunks and branches. For
these samples, it was taken advantage of the natural falling of
canopy oaks that were near the study area.
The biomass found in green leaves and upper
branches of the crown was calculated from the gathering of six
branches of different trees with approached dimensions of: 2m x 2m x
2m. All the leaves were taken off these branches in order to
be dried and weighed. Subsequently, the number of branches
that showed a covering of 2m x 2m x 2m was counted in each
tree. With this, it was made an approximation of the number of
branches and the quantity of biomass that these held.
The total biomass of bromeliads and organic matter
placed on the bromeliads was evaluated in the selected trees taking
into account the following parameters: 1) species, 2) number
of individuals from each species in each host
tree, 3) age class of the epiphytes. This was established from the
gathering of 115 individuals that came in different sizes and
the seven species that lived in the place. The diameter and the
height were measured from these individuals, and then, leaves were
taken off. Once the plant was stripped off, the bromeliad leaves were
separated from the organic matter placed on the plant. Both samples
were dried to a temperature of 70º C up to the point of
obtaining a homogeneous drying to be weighted to obtain the biomass
quantity after all. The analysis of bromeliads was carried out by
visual estimation after having established the size.
Likewise, the biomass of lichens and bryophytes held in the crown
tree was calculated from the selection of 32 branches of well-known
volume with distinct diameters and located in different points of the
crown of different trees. The lichens and bryophytes biomass held in
the trunk was evaluated from six trunk parts of mature trees. All
lichens and bryophytes were taken off to the 38 branches and trunk
fragments, so that they could be dried to a temperature of 70º C
with the purpose of obtaining the biomass value. The total biomass of
lichens and bryophytes was calculated starting from generalizating
about the samples of the whole tree volume.
The capacity of carbon sequestration in each component was determined
in the laboratory by means of oxidation by using a dicromato
potassium solution mixed with sulphuric acid measured
colorimetrically. To sum up, 36 tests were made that were distributed
in the following way: six samples of wood taken from the trunk, six
samples of branches, six samples of green leaves, six samples of dead
leaves, six samples of non-vascular epiphytes which included lichens
and bryophytes, and, six samples of bromeliads. The sample C content
was carried out at the "Corpoica - Tibaitata" soils laboratory.
Results
The aerial biomass
of oak trees and their epiphytes
The evaluated aerial biomass was made up by the
trunks, branches and leaves of each tree studied besides all the
group of epiphytes that they held. The selected trees presented an
average height of 25m and 26m, coverings between 209m2
and 255m2,
and diameters between 0.86m and 1.70m (Table 1). The biggest
percentage of biomass was found in the wood contained in the trunk,
and the branches with a diameter higher than 5cm, where values
fluctuated between 15 000kg and 25 000kg (Table 2). The branches with
a diameter lower than 5cm were the ones that less biomass presented
with an average between 58% kg and 78kg. Finally, it was found
that the biomass of the green leaves had an average between 20kg and
28kg (Table 3).
In the case of the epiphytes associated to the oak tree and to the
organic matter placed on the bromeliads, it was found that the
biggest biomass was provided by the lichens and bryophytes with
values between 36kg and 63kg, continued by the bromeliads whose
biomass fluctuated between 6kg and 39kg, and finally, the organic
matter placed on the bromeliads with values between 4kg and 23kg,
which was made up mostly by dead oak tree leaves (Table 2).

Table 1.Sampled oak trees - Quercus humboldtii- characteristics

Table 2.Oak tree biomass distribution in each one of its components and the epiphytes distribution that grow in the crown
including the organic matter accumulated on it.
Sequestration and
Storage Capacity of C
The capacity of sequestration of carbon fluctuated
in the different components of the oak tree. The values were between
35.95% and 40.13 % (Table 3). The component with the biggest capacity
of C capture was that of the branches with diameters lower than 5cm
with 40.13%, continued by the wood
contained in the trunk and in the branches with superior diameters to
5cm with 38.75%; then, the fresh leaves
with 35.95%, and finally, the dead leaves produced by the tree with
34.05% (Table 3)
On the other hand, in the epiphytes grown in the outer canopy it was
found that the lichens and bryophytes presented 43% of C. These
showed to be the component with the biggest C capacity of capturing
inside the epiphytes epiphyte-host tree system. Bromeliads captured
38.82% showing a higher capacity of capture than of the components of
their host tree, except by the superior branches. (Table 3)

Table 3. C (%) capture capacity in oak trees (Q. humboldtii) as well as the epiphytes held on them. n = number of
individuals; X = average value; SD = standard deviation.
As for the C stored in the biomass it was found
that in the four evaluated trees and their epiphytes, there was 32
066kg in 0.0938ha - corresponding to the sum of the coverings of the
four evaluated trees -. The biggest
quantity of C was found in the trunk and the branches higher than 5cm
with values between 6 161 and 9 543kg. On the other hand, the
branches lower than 5cm captured between 23.36kg and 30.99kg.
Finally, the green leaves captured between 7.38kg and 9.78kg.
Concerning the epiphytes, the lichens, and the bryophytes, the values
were between 15.65kg and 26.95kg. The bromeliads had between 2.6kg
and 15.2kg while dead leaves had between 1.62kg and 7.66kg (Table 4).

Table 4. Carbon content of tree stock in each component of the oak tree, their associated epiphytes and the intercepted organic matter.
Discussion
One of the main factors that is affecting the
montane cloud forests is Global warming which is directly associated
to the growing presence in the atmosphere of greenhouse gases and the
destructive emissions of the ozone layer (Hamilton 2001). In
addition, the deforestation processes have reduced the original
montane woods coverage in South America, and in fact, they are
considered along with the tropical dry woods, to be the most
threatened ecosystems in the world (Cavalier et al.
2001). Concerning oak forests it has been estimated that it is
necessary at least a period of 65 years to restore the structure and
floral composition, without taking into account epiphytes, and at
least 84 years to reach the structure of a mature oak forest (Kapelle
2001).
Some years ago, before knowing the importance of
oak forest fragments as protectors of basins of great importance for
the hydric system, regulators of the regional weather conditions, and
their role in the cleaning of air among other environmental services,
the oak tree was used for carpentry purposes as well as agricultural
purposes. It also was used for doing posts, fences, house and
railroad beams, floors, bodywork, fine joinery, and coal, amongst
other uses (UNESCO 2001). These uses without appropriate handling
generated alterations in the storage of C in oak forests.
Deforestation as well as the incorrect use of soil have produced a
reduction in the flow of the C and in their percentages of storing.
This change is due to the lower biomass of pastures and certain crops
(Macera et al. 2000)
In the oaks evaluated, it was found that the
percentage of C in the biomass of the host trees and their epiphytes
was, in all the cases, inferior to 50% indicated as value for defect
for the IPCC (1996), and inferior to values reported in other studies
where it has been registered that the content of C of the wood of
conifers is between 50 and 53% while in the species of wide leaf
varies from 47 to 50% (Ramírez et al. 1997). These
differences are probably due to that it is a mature forest where the
efficiency of fixation of C is smaller, contrary to what happens in
the secondary forests that are composed of species of quick growth
(Denslow 1980), and that these can have a bigger efficiency in the
fixation of the C that the primary forests (Ortiz 1997).
In spite of the low values in the concentration of C in the canopy
oak trees and in their epiphytes, the epiphyte - host tree system
presented high values of biomass which makes that this system can
retain a significant quantity of C. However, it should be kept in
mind that this study had as objective the estimation of the biomass
of huge trees; therefore, the data that was obtained is generalized
only to mature individuals of oak, and not to big extensions of
vegetable covering, since in the forests there is not a continuity of
this type of trees neither of its epiphytes. As a result, an
overestimation of the aerial biomass can be made and, consequently,
content of C can be mistaken. Brown & Lugo (1982) affirm that the
presence of trees with big diameters can have a great influence on
the vegetable biomass as well as the quantity of C.
To calculate the percentage of the sequestration
and storage capacity of C in a forest has become a tool to keep big
extensions of vegetable covering, since the storage of C helps to
mitigate the global warming (Husch 2001). To be able to know
in a more precise way which is the quantity of C that can store a
natural forest it becomes necessary to calculate not only the wooden
quantity in living vegetation, but also the biomass of all those
herbaceous forms of life that grow in the soil as well as on the
trees, since these forms of life are abundant in the tropical
forests. In the case of the epiphytes, these are an important
component of the tropical forests since they contribute significantly
with the total of the biomass (Ingram & Nadkarni 1993), the
diversity of species (Gentry & Dobson 1987), and in the cycle of
several nutrients (Nadkarni 1984) of these ecosystems.
The forests of Quercus humboldtii have
several important elements to take into account inside projects that
imply forests like places that sequester and stock C, and
conservation. These are forests of wide distribution along the three
Colombian mountain ranges whose conserved relicts harbor big mature
trees with a high biomass that are also covered by a great epiphytes
biomass that at the same time intercepts a part of the organic matter
that falls. All this biomass accumulated in the epiphyte - host
tree system makes these forests to be potential resource to storing
significant quantities of gases of effect greenhouse effect and in
particular CO2; obtaining one more environmental service
from these ecosystems.
Acknowledgements
This project was carried out thanks to the economic support of the
English organization Rufford, the support of field equipments Idea
Wild and to the logistical support of Fernando Cortes, owner of the
the Macanal reserve. We thank Sentido Natural Corporation (SN) for
the given support through the project. Especially, we thank Néstor
García, Héctor Gasca, Yolima Perez, Luisa Alvarez and
Camilo Higuera for the manuscript revision. To Juan Carlos de Las
Casas, and all the people who participated in the program of
volunteers of SN for their company to field.
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