Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
Limited evidence of coupling between above and belowground
functional traits in tropical dry forest seedlings
Rosa Arrieta-González
; https://orcid.org/0000-0002-3169-5015
Juan Paez
; https://orcid.org/0000-0002-0774-8391
Yamileth Dominguez-Haydar
; https://orcid.org/0000-0002-3103-9878
Beatriz Salgado-Negret
*; https://orcid.org/0000-0002-3103-9878
1. Departamento de Biología, Universidad del Atlántico, Puerto Colombia-Atlántico, Barranquilla, Colombia;
rarrieta@unal.edu.co, yamilethdominguez@mail.uniatlantico.edu.co
2. Departamento de Biología, Universidad Nacional de Colombia, Bogotá, Colombia;
bsalgadon@unal.edu.co (*Correspondence)
3. Grupo de Investigación en Química y Biología, Universidad del Norte, Barranquilla, Colombia;
jpaezg93@gmail.com, bsalgadon@unal.edu.co
Received 06-IV-2021. Corrected 15-VI-2021. Accepted 28-VI-2021.
Introduction: Water availability is one of the main factors determining the distribution of woody species in
the tropics. Although the functional mechanisms that determine the species tolerance to water deficit have been
extensively studied in adult individuals, the responses of early ontogenetic stages have been less explored.
Objective: To identify functional strategies and trait correlations between different seedlings’ dimensions (leaf,
stem, and root). We expect limited coordination between above and below-ground functional traits due to a
single conservation-acquisition trade-off cannot capture the variability of functions and environmental pressures
to which the root system is subjected.
Methods: We measured 12 functional traits belonging to 38 seedling species in a tropical dry forest in Colombia.
We explored the relationships between pairs of traits using Pearson correlations, and to obtain an integrated view
of the functional traits, a principal component analysis (PCA) was performed.
Results: The results showed limited evidence of linkage between above- and below-ground traits, but we did
find significant correlations between traits for the continuum of conservative and acquisitive strategies. Root
traits related to water and nutrient take capacity formed an orthogonal axis to the acquisitive-conservative
Conclusions: Our results showed that dry forest seedlings have different functional strategies to cope with water
deficit. The incorporation of root traits helps to explain new functional strategies not reported for leaf and stem
traits. This study contributes to understanding the mechanisms that explain species coexistence and is particu-
larly relevant for predicting future forest trajectories.
Key words: acquisitive strategy; Colombia; conservative strategy; functional traits; water deficit; water storage.
Arrieta-González, R., Paez, J., Dominguez-Haydar, Y., &
Salgado-Negret, B. (2021). Limited evidence of coupling
between above and belowground functional traits in
tropical dry forest seedlings. Revista de Biología Tropical,
69(2), 763-771. https://doi.org/10.15517/rbt.v69i2.46549
Water availability is one of the main envi-
ronmental factors determining the establish-
ment and distribution of tree species at local
and regional scales (Engelbrecht et al., 2007;
Markesteijn et al., 2011; Méndez-Alonzo et
al., 2012). Different studies have shown that
the performance of species varies in response
to water availability, and this is explained
Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
through their functional traits (Engelbrecht
et al., 2007; Poorter & Markesteijn, 2008;
Markesteijn et al., 2011; González-M et al.,
2020). For instance, species adapted to drought
have developed functional traits that allow
them to maintain growth despite low soil water
potentials. Among the most studied traits are
stomatal closure and leaf deciduousness, which
allows plants to avoid dehydration during the
driest periods of the day or year (Markes-
teijn et al., 2011; Méndez-Alonzo et al., 2012).
Water storage in tissues to maintain xylem
water transport and low levels of dehydration
(Borchert, 1994; Scholz et al., 2007; Pineda-
García et al., 2013), investment in deep roots to
access more humid soils (Paz et al., 2015), or
the presence of dense tissues that do not dehy-
drate and promote high resistance to cavitation
(Bonal & Guehl, 2001; Comita & Engelbrecht,
2014). Despite this knowledge, studies are
still needed that incorporate traits associated
with other plant structures and functions, such
as root traits.
Functional traits can be coupled to define
functional strategies, and tree species exposed
to water deficit are distributed along a con-
tinuum of traits related to tissue investment
and a hydraulic safety-efficiency trade-off
(Méndez-Alonzo et al., 2012; Pineda-García
et al., 2015; González-M et al., 2020). At one
extreme, there are species with high invest-
ment in dense tissues, dense stems with narrow
xylem vessels, high leaf dry matter content,
and leaf longevity. These species with dense
and hydraulically safe tissues, known as “con-
servative” species, have low growth rates but
high survival (Poorter et al., 2008). The other
extreme are the “acquisitive” species charac-
terized by thin, short-lived leaves with high
investment in nutrients and less dense stems
with wide xylem vessels. These soft tissues
and their high hydraulic efficiency result in
high growth rates but low survival (Poorter et
al., 2008). Root trait dimension has been less
explored. One hypothesis postulate that roots
follow the “acquisition-conservation” trade-off
analogous to leaf economic spectrum (Weems-
tra et al., 2016). However, recent studies have
found mixed evidence (Valverde-Barrantes et
al., 2015; Withington et al., 2006; Weemstra et
al., 2016), showing decoupling between above
and below ground traits (Withington et al.,
2006; Weemstra et al., 2016; Freschet et al.,
2020). Recent work showed a trade-off in C
investment between deep roots and water stor-
age tissues associated with successional and
old-growth forest species, respectively (Paz
et al., 2015). These studies invite to discuss
the correlations between functional traits and
whether coordination between traits reported
in adults develops from the first stages of
plant development.
Understanding the response mechanisms
to drought is particularly relevant in seedlings
because regeneration allows for the inference
of forest successional trajectories and ulti-
mately is key to determining forest resilience
(Osuri et al., 2017). Additionally, seedlings are
the most sensitive to environmental changes as
they have poorly developed roots that do not
reach deep soil layers (Engelbrecht & Kursar,
2003) and must compete for water and nutri-
ents with established adult individuals (Lewis
& Tanner, 2000). Furthermore, seedlings face
light limitations in the understory that reduces
their photosynthetic and growth rates (Romo,
2005); and in turn, they have limited carbo-
hydrate reserves to invest in organs such as
leaves or roots (Poorter & Markesteijn, 2008).
These characteristics, added to the fact that
many tropical species can spend decades under
understory (Hubbell et al., 1999), make this
ontogenetic state the bottleneck for establishing
individuals and forests of the future.
Tropical dry forest is an ideal model for
understanding how tree species cope with
drought. First, it presents a strong dry season
where soil water deficit limits tree species’
growth and establishment (Murphy & Lugo,
1986; González-M et al., 2019). Second, strate-
gies to cope with water deficit have been well-
studied in adult individuals, but it is unclear
whether these strategies are fully developed
from the seedling ontogenetic stage. And third,
high fragmentation rates in this ecosystem can
amplify the climatic severity due to the edge
Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
effects with effects on its structure and diver-
sity (González-M et al., 2019). Understanding
the functional mechanisms that determine seed-
ling establishment will strengthen our ability
to predict and understand these forests’ future
under climate change scenarios in which great-
er seasonality and reductions in rainfall are
expected (Dai, 2013; Feng et al., 2013).
This study evaluated the variation of 12
functional traits above and belowground in 38
species of dry forest seedlings. The selected
traits are related to the different strategies of
plants to cope with drought, such as investment
in dense tissues, efficiency in root foraging
of water and nutrients, and the water stor-
age capacity (Markesteijn & Poorter, 2009;
Méndez-Alonzo et al., 2012; Pineda-García et
al., 2013; Paz et al., 2015; Salgado-Negret et
al., 2016). The main objective was to identify
the functional strategies and the correlation
among pairs of traits belonging to different
plant dimensions (leaf, stem, and root). We
expected species are distributed along with two
extreme strategies, acquisitive and conserva-
tive. However, we expect low coordination
between above and below-ground functional
traits due to a single conservation-acquisition
trade-off cannot capture the variability of func-
tions and environmental pressures to which the
root system is subjected (Valverde-Barrantes et
al., 2015; Weemstra et al., 2016).
Study area and species selection: This
study was carried out in a tropical dry forest
located in Caribbean region in Colombia (Bras-
ilar, municipality of San Jacinto, department of
Bolívar - 9°53’40” N, 75°10’57” W), located at
an altitude between 352 and 602 m a.s.l.
We selected 38 seedling species belonging
to 32 genera and 21 families (14 deciduous
and 24 evergreen species; Digital appendix
1). The species selection was based on their
high abundance in the study area, and together
they represented more than 80 % of the total
abundance of seedlings registered in 96 plots
of 1 m
previously established for the area
(Salgado-Negret, data unpublished). For abun-
dant species, between 4 and 15 individuals
(57.8 %) and rare species, between 1 and 3
individuals (42.1 %), were sampled. All indi-
viduals were located in similar light conditions
(canopy cover varied between 72.1 % and 98.4
%) and similar relative soil humidity (6.36 and
21.1 %). The individuals had a height between
40 and 70 cm. Harvest of each seedling was
carried out by digging with a shovel 50 cm
around the stem to avoid losing the fine roots.
The sampled seedlings were stored in plastic
bags, labeled, and transported to the field
station for processing.
Functional traits: In total, 12 functional
traits belonging to the leaf, stem, and root
dimensions were measured, following the
protocols of Poorter and Markesteijn (2008),
Pérez-Harguindeguy et al. (2013), Paz et
al. (2015) and Salgado-Negret et al. (2016).
Leaf traits were estimated for five leaves per
individual. Each leaf was weighed fresh and
scanned (Canon Canoscan LiDE 220) to esti-
mate the leaf area (LA, mm
). Leaf thickness
(LT, µm) was measured for each fresh leaf
with a digital micrometer (Mitutoyo reference
293-240-30) in three different sections of the
leaf, and avoiding the main veins. The leaves
were dried at 70 ºC for 48 hours to estimate the
dry weight, and from which the specific leaf
area (SLA, cm
/g) and leaf dry matter content
(LDMC, mg/g) were calculated.
Stem density (SD, g/cm
) was estimated
by dividing the dry mass (dried at 70 ºC for 48
hours) and the green volume calculated by the
water displacement method (Salgado-Negret
et al., 2016). The stem water content (SWC,
mg/g) was estimated by dividing the weight of
the fresh water-saturated stem sample by the
oven-dry weight. With the hydrated roots, the
fresh weight of the whole root was estimated,
then they were dried at 70 ºC for 48 hours to
estimate the dry weight. Using an architectural
criterion, the roots were separated into primary
and secondary roots. The root dry matter con-
tent (RDMC, mg/g) was calculated as the
relationship between the dry weight and the
Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
fresh weight of the secondary roots. The root
water content (RWC, g/g) was determined as
the relationship between the fresh weight and
the dry weight of the primary root. The sec-
ondary-to-primary-root-mass ratio (SPRMR,
g/g), was calculated as the secondary root mass
per unit primary root mass (Markesteijn &
Poorter, 2009). The aboveground-to- below-
ground -length ratio (AUL, cm) was calculated
as the ratio between seedling height (cm) and
maximum root depth (MRD, cm), both traits
were estimated with a tape measure. Finally,
leaf, stem and root mass fractions (LMF, SMF,
RMF, g/g) were calculated as dry mass of each
organ per unit dry plant mass). At the end, the
LMF and SMF (LSMF, g/g) were summed as
an indicator of the aboveground biomass of
the plant.
To explore the relationships between pairs
of traits, Pearson correlations using the Bon-
ferroni correction were performed. In order
to obtain an integrated view of the functional
traits, a principal component analysis (PCA)
was performed using the 38 species and the
12 functional traits to identify the functional
strategies of the species. All analysis were
performed using R statistical language (R Core
Team, 2015).
Significant correlations were reported
between traits of different seedlings’ dimen-
sions (Fig. 1). Functional strategies were
observed in the PCA (Fig. 1, Table 1), and the
first two ordering axes explained 61.4 % of the
species’ variation as a function of the function-
al traits. PCA reported two functional continua;
the first was dominated by the “trade-off”
between resource conservation and acquisition.
Conservative species showed high leaf and
root dry matter contents (LDMC and RDMC)
and high stem density (SD), while acquisitive
species were related with high water content
in roots (RWC) and stems (SWC) (Fig. 1). The
traits that defined each of these strategies were
negatively correlated (i.e., decoupled; Table 2).
The second continuum separated the species
with high investment in belowground biomass
(RMF) and deep roots (MRD) from the species
with tall height above ground (AUL), shallow
roots, and high investment in secondary roots
(Fig. 1, Table 2). Leaf traits such as SLA and
LT were not correlated with other functional
traits (Table 2).
Loadings and proportion of variance of the principal
component analysis of leaf, root and stem traits of 38
species of seedlings from a dry forest in Colombia
Functional traits Comp.1 Comp.2
LDMC 0.29 0.225
SLA -0.133
LT -0.198
SD 0.319 0.359
RMF 0.307 -0.353
RWC -0.353 -0.264
SPRMR -0.323 0.255
RDMC 0.253 0.342
AUL -0.337 0.225
SWC -0.281 -0.368
MRD 0.339 -0.305
LSMF -0.307 0.353
Proportion of variance 38.6 22.8
Trait abbreviations: leaf dry matter content (LDMC),
specific leaf area (SLA), leaf thickness (LT), stem density
(SD), root mass fraction (RMF), root water content (RWC),
econdary-to-primary-root-mass ratio (SPRMR), root dry
matter content (RDMC), aboveground-to belowground
length ratio (AUL), stem water content (SWC), maximum
root depth (MRD) and leaf-stem mass fraction (LSMF).
Using 12 functional traits measured in
38 seedling species from a tropical dry for-
est, we found limited evidence of coupling
between above and belowground traits, with
the exception of tissue density (conservative
strategy) and storage capacity of stems and
roots (acquisitive strategy). Furthermore, roots’
traits relating to the ability to forage water
and nutrients formed an orthogonal axis to
the acquisitive-conservative continuum. These
results show that root traits cannot be reduced
to a single axis of variation, which is probably
Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
Fig. 1. Principal component analysis (PCA) of the average values of functional traits by species. Above-ground (green) and
below-ground functional traits (red). The percentages of variation explained by the first two axes of the ACP are included
in the graph. See trait abbreviations in Table 1.
Pearson correlations with Bonferroni correction between leaf, root and stem traits of 38 species
of seedlings from a dry forest in Colombia
LDMC 1 -0.46 -0.31 0.59** 0.05 -0.55* -0.42 0.4 -0.33 -0.4 0.28 -0.05
SLA 1 -0.43 -0.15 -0.01 0.23 0.23 -0.01 0.23 0.06 -0.23 0.01
LT 1 -0.31 0.04 0.13 0.21 -0.31 0.06 0.11 0.05 -0.04
SD 1 0.07 -0.74*** -0.27 0.58** -0.35 -0.79*** 0.19 -0.07
RMF 1 -0.30 -0.60** 0.2 -0.49 -0.11 0.68*** -1***
RWC 1 0.19 -0.64*** 0.32 0.74*** -0.36 0.30
SPRMR 1 -0.05 0.71*** 0.07 -0.61*** 0.60**
RDMC 1 -0.05 -0.67*** 0.05 -0.2
AUL 1 0.2 -0.79*** 0.49
SWC 1 -0.14 0.11
MRD 1 -0.68***
*P <0.05, **P < 0.01, ***P < 0.0001. Trait abbreviations: specific leaf area (SLA), leaf dry matter content (LDMC), leaf
thickness (LT), leaf-stem mass fraction (LSMF), stem density (SD), stem water content (SWC), root water content (RWC),
root dry matter content (RDMC), root mass fraction (RMF), secondary-to-primary-root-mass ratio (SPRMR), aboveground-
to belowground length ratio (AUL) and maximum root depth (MRD).
Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
related to the multiple functions that they fulfill
in the individual. Plants have different combi-
nations of traits that enable them to deal with
water limitations in these ecosystems.
The PCA did show that tissue density (SD,
RDMC, and LDMC) was coordinated across
plant dimensions, which has been reported
previously in seedlings (Poorter & Markesteijn,
2008; Markesteijn & Poorter, 2009; Pineda-
García et al., 2015; Weemstra et al., 2016).
The coordination of high tissue density is
an important strategy to cope with drought
because species with dense stems are gener-
ally associated with narrow xylem vessels that
allow them to reduce the risk of cavitation of
the xylem (Markesteijn et al., 2011; Méndez-
Alonzo et al., 2012). Additionally, a high den-
sity of fibers with thick cell walls may protect
the xylem from the strong negative pressures
generated by drought (Hacke et al., 2001).
Dense, smaller, and more rigid leaves, have
small transpiration surfaces that reduce the
plants’ wilting and water requirements (Poorter
et al., 2009). Examples of conservative species
are Trichilia carinata (Meliaceae), Machae-
rium lanceolatum (Fabaceae), and Chiococca
alba (Rubiaceae). Additionally, our results also
showed coordination between water storage
capacity in stems and roots. Water storage
plants have low stem density which do not
resist the xylem’s high tension during the dry
seasons, which could have acted as a selection
pressure promoting water storage in their tis-
sues (Méndez-Alonzo et al., 2012). The root
water storage is essential for the maintenance
of photosynthetic and growth rates during
dry seasons but also in the hours of greatest
daytime transpiration when there may be a
significant delay between the loss of water in
the leaves and the absorption of water by the
roots (Čermák et al., 2007). For the tropical
dry forest, seedlings can use the water stored
in the stems (SWC) to delay hydraulic failure
for months (Pineda-García et al., 2013), and
the high root water storage of seedlings (RWC)
allows them to function physiologically in
drought conditions (Poorter & Markesteijn,
2008). Piper piojoanum (Piperaceae), Piper
reticulatum (Piperaceae), and Anacardium
excelsum (Anacardiaceae) are examples of
water storage species.
The orthogonal axis to the acquisitive-
conservative continuum was related to the
roots’ ability to forage water and nutrients in
the soil. At one extreme, we found coordina-
tion in belowground traits related to explore
and forage deep soils (high RMF and MRD).
At the other end, species with shallow roots
(low MRD), high secondary root biomass (high
SPRMR), and high inversion on above ground
length (high AUL) were located. Plants in dry
forests have shown two main root strategies:
deep roots with high carbon investment to
explore the more humid soil horizons or invest-
ing in water-storing tissues (Paz et al., 2015;
Prieto et al., 2015). Because the two strategies
involve investing in the construction and main-
tenance of different types of tissues, the trade-
off is expected and has even been associated
with different successional stages (Paz et al.,
2015). Contrary to these studies, our results did
not show this dichotomy, and root water stor-
age (RWC) was decoupled from traits related
to water and nutrient foraging such as MRD,
RMF, and SPRMR. These results mean that
both species with deep roots and high invest-
ment in root biomass (high MRD and RMF)
and species with shallow roots with a high
proportion of secondary roots (high SPRMR)
can store water in their root tissues. Examples
of water storage species with shallow roots are:
Sorocea sprucei (Moraceae) and Ruellia mac-
rophylla (Acanthaceae), while Eugenia florida
(Myrtaceae) would be the species with deep and
water storage roots. Despite this orthogonality,
our results show that most of the acquisitive
species with water-storage roots had shallow
roots with high secondary roots investment.
These water-storage roots are associated with a
greater capacity for soil exploration, absorption
of nutrients and water (Eissenstat et al., 2000;
Roumet et al., 2006; Paz et al., 2015; Prieto et
al., 2015). Conversely, species with thick and
deep roots will have greater resistance to water
deficit because these provide more specialized
Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
functions for the transport of water and nutri-
ents (Paz et al., 2015; Prieto et al., 2015).
The decoupling between acquisitive-con-
servative continuum and the roots´ ability to
forage water and nutrients could have differ-
ent explanations. First, as the root system has
different functions, mechanical support, and
transport and storage of water and nutrients
(Eissenstat et al., 2000; Prieto et al., 2015), its
functional traits must be optimized to comply
with the three functions that could imply the
absence of a maximization of a trait associated
with a particular function. Second, the water
and the essential elements that plants acquire
from the soil have different properties, causing
the same trait to not be efficient in absorbing
all the elements (Weemstra et al., 2016). And
third, the physical properties of the soil exert
selective forces on the root system, which
could limit the maximum development of
traits for the acquisition of a specific limiting
resource (Weemstra et al., 2016). Under this
scenario, it seems likely that the roots have dif-
ferent strategic dimensions and that these vary
according to local conditions, which makes it
difficult to generate patterns in the correlations
between the above and belowground traits.
This study found limited trait coordina-
tion between above- and belowground traits
and novel trade-offs regarding belowground
trait correlation. A remarkable finding was the
decoupling between the acquisitive-conserva-
tive continuum from the ability to forage water
and nutrients of roots. Our results showed that
dry forest seedlings have multiple strategies to
cope with drought and highlight the importance
of incorporating root traits for a comprehen-
sive understanding of the responses of plants
to environmental conditions. This variety of
functional strategies, and, therefore, the parti-
tion of niche between species, could help to
explain the high diversity of species at the local
scale reported in tropical dry forests. However,
it is essential to recognize that traits may vary
with ontogeny, which is a variation source
rarely accounted. Additionally, these results
could help predict the species´ responses to
future climatic scenarios, to allow the selection
of species capable of establishing themselves
under different environmental conditions and
guide restoration actions in the most endan-
gered terrestrial ecosystem in the Neotropics.
Ethical statement: authors declare that
they all agree with this publication and made
significant contributions; that there is no con-
flict of interest of any kind; and that we fol-
lowed all pertinent ethical and legal procedures
and requirements. All financial sources are
fully and clearly stated in the acknowledge-
ments section. A signed document has been
filed in the journal archives.
We are grateful to the expert botanist
Álvaro Idárraga for his help identifying the
species. Natalia Norden, Hermes Cuadros, and
Ana Belén Hurtado provide helpful comments
on earlier versions of this manuscript. Finally,
without the generosity of Mr. Eduardo Torres,
Mrs. Reyna, and Eduardo Torres, who opened
the doors of their home, this work would not
have been possible.
Evidencia limitada de acoplamiento entre rasgos
funcionales por encima y por debajo del suelo
en plántulas de bosque seco tropical
Introducción: La disponibilidad de agua es uno de los
principales factores que determina la distribución de las
especies leñosas en los trópicos. A pesar que los meca-
nismos funcionales que determinan la tolerancia de las
especies al déficit hídrico han sido ampliamente estudiados
en los individuos adultos, las respuestas de estados ontoge-
néticos tempranos han sido menos exploradas.
Objetivo: Identificar las estrategias funcionales y las
correlaciones de rasgos entre diferentes dimensiones de las
plántulas (hoja, tallo y raíz). Nosotros esperamos baja coor-
dinación entre los rasgos funcionales sobre y bajo el suelo
debido a que un único trade-off conservación-adquisición
de recursos, no puede capturar la variabilidad de funciones
y presiones ambientales a las que están expuestas las raíces.
Métodos: Medimos 12 rasgos funcionales pertenecientes a
38 plántulas en un bosque seco en Colombia. Exploramos
las relaciones entre pares de rasgos usando correlaciones
de Pearson, y para tener una visión integrada de los rasgos
Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(2): 763-771, April-June 2021 (Published June 30, 2021)
funcionales, usamos un análisis de componentes principa-
les (ACP).
Resultados: Reportamos limitada evidencia de acopla-
miento entre los rasgos sobre y bajo el suelo, pero
encontramos correlaciones significativas entre el continuo
de estrategias conservativas y adquisitivas. Los rasgos
de raíz relacionados con la capacidad de absorción de
agua y nutrientes formaron un eje ortogonal al continuo
Conclusiones: Nuestros resultados mostraron que las
plántulas del bosque seco tienen diferentes estrategias
funcionales para lidiar con el déficit hídrico. La incorpo-
ración de los rasgos de la raíz ayuda a explicar nuevas
estrategias funcionales no reportadas por los rasgos de hoja
y tallo. Este estudio contribuye al entendimiento de los
mecanismos que explican la coexistencia de especies y es
particularmente relevante para predecir las trayectorias de
los bosques futuros.
Palabras clave: estrategia adquisitiva; Colombia; estra-
tegia conservadora; rasgos funcionales; déficit hídrico;
almacenamiento de agua.
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