Acta Oecologica 114 (2022) 103814

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Acta Oecologica
journal homepage: www.elsevier.com/locate/actoec

Functional diversity and species diversity in flooded and unflooded

tropical forests
Lilia Roa-Fuentes a, *, Lisney A. Villamizar-Peña b, Jhon A. Mantilla-Carreño b,
M. Alejandra Jaramillo c
a
Pontificia Universidad Javeriana, Departamento de Ecología, Transv.4◦ No.42-00. Edificio J. Rafael Arboleda, Código Postal, 110231, Bogotá, D.C., Colombia
b
Universidad Industrial de Santander, Ciudad Universitaria, Cra. 27 #9, Bucaramanga, Santander, Colombia
c
Grupo de Investigación DIVERSITAS, Facultad de Ciencias Básicas y Aplicadas, Universidad Militar Nueva Granada, Km 2 Cajicá-Zipaquirá, Cajicá, Cundinamarca,
Colombia

A R T I C L E I N F O A B S T R A C T

Keywords: Functional diversity and species diversity provide information to understand differences between plant com­
Functional richness munities from a contrasting environment and the complementary on regional scale. Environmental variation
Functional divergence determines which plants can persist in the community via species sorting processes; forest exposed to extended
Functional evenness
inundation each year has shown divergence in plant community composition, structure, and functional diversity,
Magdalena valley
compared with unflooded dense forest. Here, we researched flooded and unflooded forests in the upper Mag­
dalena River Valley (Colombia) to quantify and compare the plant species diversity and plant functional di­
versity. We calculated three components of functional diversity (i.e., functional richness, functional evenness,
and functional divergence), we also used a measure of functional richness weighted by species abundance. Our
results show that the plant community in the unflooded forest has higher stem density and lower basal area than
flooded forests. Also, we show that tree communities in the flooded forests had higher specific leaf area, but
lower wood density, suggesting an acquisitive resource strategy. Species diversity and functional diversity were
higher in the plant community from the unflooded forest. Our findings of the functional traits and the functional
diversity components reinforce the idea that the variation between flooded and unflooded forests is going beyond
the species list in the plant communities. However, it is still unclear whether any general patterns exist regarding
differences between flooded and unflooded tropical forests.

1. Introduction Functional diversity is defined as the value, range, distribution, and

Plant species composition, plant diversity and the factors affecting
the species distribution have been well studied in tropical forests.
However, this knowledge neither shows us the variation of plant attri­
butes, resulted from the historical pressure of the environment, nor their
effect on ecosystem functioning (Hedberg et al., 2013). Alternatively,
functional ecology (i.e., trait-based ecology) can reveal a predictive
relationship between plant community and environmental gradients
using functional traits, i.e., traits affecting the species’ fitness (Shipley
et al., 2016). Plant functional traits are relevant to define the strategies
of plants to use resources. In such a way, that value of plant functional
traits can be used to define the ecological role of species (Díaz et al.,
1999; Lavorel and Garnier, 2002) and its response to environmental
changes (Lavorel et al., 1997).
relative abundance of the functional traits in a community (Díaz et al.,
2007). Despite the theoretical attention on functional diversity, much
remains to be understood about its pattern across environmental gra­
dients and different ecosystem types at the local scale (Lohbeck et al.,
2012; Thakur and Wright, 2017). Variation of abiotic conditions at a
local scale might determine which plant functional traits will facilitate
the persistence of species in a community (Hobbs and Norton 2004).
Therefore, abiotic conditions exert control within and between com­
munities affecting the dispersion of trait values (Grime, 2006).
Forests with extended and variable inundations each year (flooded
forest) show different plant community composition, structure, and
functional diversity than the unflooded forest (Baraloto et al., 2011;
Lawson et al., 2015; Homeier et al., 2017; Stevenson et al., 2018).
However, studies that compare flooded and unflooded forests show

* Corresponding author.
E-mail address: lilia.roa@javeriana.edu.co (L. Roa-Fuentes).

https://doi.org/10.1016/j.actao.2022.103814
Received 15 December 2020; Received in revised form 30 November 2021; Accepted 17 January 2022
Available online 3 February 2022
1146-609X/© 2022 Published by Elsevier Masson SAS.
L. Roa-Fuentes et al.
unambiguous patterns. When forest structure is compared between
flooded and unflooded forests, the trend has shown that stem density
and basal area are higher in the unflooded forest (Baraloto et al., 2011).
In contrast, the results reported by Kurzatkowski et al. (2015) show that
stem density and basal area are not dependent on the flood level;
therefore, the results are not conclusive for all flooded ecosystems.
Similarly, when plant functional traits are compared between flooded
and unflooded forests, the picture is not clearer. Mommer et al. (2006)
and Fortunel et al. (2014) reported that unflooded forest shows higher
wood density and lower specific leaf area than the flooded forest, indi­
cating a conservative resource plant strategy (Swenson and Enquist
2009). Such a strategy is linked with slow growth and high leaf dry
matter content (LDMC), which have been related to reducing cavitation
risk (Chave et al., 2009).
Conversely, other reports show that flooded forests have higher
wood density, pointing out a trade-off between plant flood resistance
and rapid resource acquisition. (Lawson et al., 2015; Kurzatkowski et al.,
2015). The plant conservative vs. acquisitive resource in the flooded and
unflooded forest could be an approach to the stability of the plant
community. Dynamics of water and resource inputs in flooding forests
may favor the traits related to resource acquisition (Fischer et al., 2016),
similarly as an early successional state. In contrast, the unflooded forest
could be exhibiting a conservative resource acquisition as a
late-successional state.
This apparent ambiguity highlights some integrative ideas; first,
functional traits incorporate many physiological processes interrelated
and can be highly plastic (Paine et al. 2015). Second, the flooding dy­
namic is an indirect gradient that includes multiple varied factors, which
push plants to respond. Parolin (2009) suggested that the plant’s strat­
egy in flooded forests exhibits a great diversity of morphological dif­
ferences. Evidence for the early life-history stage (seeds) show
differences between the plant communities in flooding and unflooding
forest (Lopez, 2001). Also, trait values in organs involved in the oxygen
exchange (e.g., leaf and wood) can affect the persistence of plant species
in a flooded forest (Fischer et al., 2016; Mommer et al., 2006; Fortunel
et al., 2014). The evidence suggests high morphological diversity in the
flooded forest in favor of high functional diversity, whit a low species
diversity. However, the available evidence on functional diversity and
its attributes in flooded and unflooded forests is inconclusive.
Despite these significant gaps and challenges in defining a pattern for
plant responses to flood dynamics, we need to increase our knowledge in
the flooded and unflooded forest, which will allow us to provide valu­
able suggestions for conservation in the face of climate change.
In Magdalena Valley (Colombia), hydrologic fluctuations shaped
vegetation and have formed a mosaic of unflooded and flooded forests.
Plant diversity is scarcely studied, and no national parks protect lowland
forests in the Magdalena Valley (Stevenson et al., 2018). In this region,
forests are strategic because their services support human well-being
(Angel-Escobar et al., 2014).
This study evaluates the pattern of plant species and functional di­
versity values between flooded and unflooded forests in the Magdalena
Valley. We explored the plant species diversity by calculating the
abundance weighed diversity index, and the stand structure by
measuring the stem density and basal area. The functional diversity was
measured by determining the plant functional groups and the plant
functional diversity (i.e., functional richness, functional evenness, and
functional divergence) for both forest types. In addition, we calculated
the community-based weighted functional diversity index (wFDc) in the
unflooded forest than in the flooded forest.
We expect two main trends: 1. Plant communities with conservative
resource strategy in the unflooded forest; and 2. Higher values of func­
tional diversity and community-based weighted functional diversity
index (wFD) in the unflooded forest than in the flooded forest.
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Acta Oecologica 114 (2022) 103814
2. Materials and methods
2.1. Study area
The study area is located on the floodplain of Barbacoas River in
Yondó municipality, Colombia (06◦ 48′ 2N, 74◦ 12′ W), part of the upper
Magdalena River basin. Barbacoas is a strategic ecosystem for water
flow regulation, due to its strategic location in the lower Magdalena
River in the inter-Andean valley. Also, floodplain of Barbacoas River is a
habitat for threatened plant species as Cariniana pyriformis Miers,
Couratari guianensis Aubl. and Caryocar amygdaliferum Mutis, which
are difficult to find in other protected reserves in the country (Calderón
et al., 2002). The flooding period in the region and our study plots is
from 4 to 8 months for a year in the rainy season from April to
November, during flooding soil may be covered by a sheet of water, the
forest is considered as a black water forest (Angel-Escobar et al., 2014).
In 2014 floodplain of Barbacoas River was designated as a protected
area.
Vegetation in Magdalena River Valley corresponds to wet tropical
forest (WTF) in the Holdridge life zone system (Holdridge et al., 1971).
The climate is warm and humid, with an average annual temperature of
28 ◦ C. Annual rainfall ranges between 5000 mm and 7000 mm, with
maximum precipitation from April to May and October through
November (Angel-Escobar et al., 2014). Furthermore, the floodplain of
Magdalena River is farmed intensively (Lewis et al., 2006), and near our
study site, the principal driver of forest loss is the use of pasture for cattle
ranching. Nevertheless, our study area still contains a relatively undis­
turbed mosaic of flooded (FF) and unflooded forest (UF). Site use history
was assigned based on local expert knowledge (Fig. 1). The selected sites
have the same use history without evidence of logging.
2.2. Field sampling
We carried out floristic inventories in two environments FF and UF.
We selected sites using geographic information system tools and Rap­
idEye satellite imagery. Sites were located following the local guide and
plots in FF and UF were installed. Sites in each forest type were selected
considering they exhibit as much environmental homogeneity as
possible regarding soil, topography, and land-use history; in the same
way, we considered forest structure homogeneity.
Data were collected from June until August 2014. We used ten plots
(50 × 2 m) per forest type (i.e., FF and UF, respectively). The location of
each transect is summarized in Fig. 1. Flooded forest plots were located
along the major river in backwater swamps. Diameter at breast height
(DBH) of all tree individuals with DBH >2.5 cm (Gentry 1988) was
measured. We sampled ten healthy, mature, and fully expanded leaves
from tree crown of each individual in the plots, and we collected only
sun-exposed leaves. Leaves for each tree were stored in water-saturated
conditions, following protocol used by Pérez-Harguindeguy et al.
(2013). To sample wood density (WD) in great individuals, we used an
increment borer (5 mm diameter) to extract trunk core sections. To
avoid the negative effect on small trees, we sampled branch sections
from smaller individuals inside plots (Chave et al., 2005). Wood cores
and branch sections were oven-dried at 70 ◦ C until they achieved a
constant weight (about 48 h). Botanical specimens were dried using
standard herbarium techniques (Mori et al., 1989) and deposited in
Universidad Industrial de Santander Herbarium (UISH).
We select plant traits that reflect a gradient of slow (conservative) to
fast (acquisitive) strategies in terms of investment and use of nutrients
and other resources (Wright et al., 2004; Díaz et al., 2016). For leaf area
(LA) measurements, the leaf petiole was cut; then, the leaf blade was
measured and photographed. Images were stored and processed using
software ImageJ 1.49 (Schneider et al., 2012). Leaves were weighed for
fresh mass and later oven-dried at 70 ◦ C for 48 h and weighed for dry
mass. Specific leaf area (SLA cm2 g− 1) influences the canopy expansion
and growth through its effect on total leaf area per plant affecting light
L. Roa-Fuentes et al.
Fig. 1. Map of the study region (left) and site localities (right) in the upper Magdalena River basin; study plot localization (in the boxes) of flooded forests rep­
resented with black points and unflooded forests represented with white points.
interception and light use efficiency (Kumer et al., 2012) and was
calculated as the ratio of leaf area to leaf dry mass, excluding petioles
and rachises for compound-leaved species. Leaf thickness was measured
in four places between major veins with a digital micrometer (Mitutoyo
293-240-30). Leaf dry matter content (LDMC, mg g–1) another func­
tional marker of plant strategy for resource use (Garnier et al., 2001) was
calculated as oven-dry mass (mg) of a leaf, divided by its water-saturated
fresh mass (g). Wood density (WD, g cm− 3) reflects carbon allocation to
structural support was estimated as the ratio of the oven-dry mass (g) of
a wood sample divided by its green volume (cm3). Volume was deter­
mined by using the geometrical dimensions of wood core (Chave et al.,
2005). For functional diversity analysis, we used mean trait value for
each species.
2.3. Analysis
To explore tree species composition and forest structure, we first
calculated species richness (S) as the number of species sampled in each
plot for each forest type. To avoid under sampling-bias, we calculated
species richness adjusting the number of individuals and the number of
samples collected (rarefaction) (Gotelli and Colwell, 2011). The Shan­
non–Wiener index (H) as an indicator of species diversity weighted by
species abundance was used. We computed a one-way analysis of vari­
ance ANOVA test to examine the difference between forest type for plant
structure and taxonomic diversity. To describe the forest’s structure, we
calculated stem density as the number of individuals per area. To
calculate species’ importance to each forest, we used the importance
value index (IVI) as the sum of the species’ relative density, relative
frequency, and relative basal area (Curtis and McIntosh, 1950). To
compare the forests’ composition, we used a similarity analysis (ANO­
SIM) (Clarke 1993). ANOSIM was completed using a matrix of plots per
species abundance.
To select traits for functional diversity analysis, we evaluate the
degree to which plant traits determine plant species’ relative abun­
dance, as was proposed by Shipley et al. (2016). We carried out a
Pearson correlation between all species’ relative abundance and SLA,
LDMC, WD, leaf thickness and LA values to each forest type. A signifi­
cant relationship (p < 0.05) was assumed to quantify the functional
importance of used traits in our specific context. There was a clear
systematic relationship between plant traits and species abundance in
both forest types. In flooding forest, SLA (p = 0.012), LDMC (p =
0.04594), WD (p = 0.004), leaf thickness (p < 0.001), and LA (p = 0.05),
showed a relationship with species’ relative abundance. In concordance,
in unflooded forest SLA (p = 0.0289), LDMC (p = 0.05), WD (p = 0.002),
leaf thickness (p = 0.02), and LA (p = 0.03), showed a positive linear
relationship with species relative abundance, and then such traits were
used as functional traits for functional diversity analysis.
To characterize functional diversity and compare between FF and UF
forest, we identified plant functional groups (PFGs) using a hierarchical
3
Acta Oecologica 114 (2022) 103814
conglomerate analysis, Ward clustering algorithm and Euclidean dis­
tances. PFGs were determined using a matrix containing standardized
plant traits was used (SLA, LDMC, leaf thickness, LA, and WD) for each
species. The standardization was carried out using the stand. x argument
in the dbFD function (Package ‘FD’, R version 2.13.1.) function to
standardized to mean 0 and unit variance. Finally, we used a principal
component analysis (PCA) to visualize trait distributions among species.
Then, we compared the number of species and the traits means of each
PFGs between forests type.
Three components of functional diversity were explored by calcu­
lating functional richness (FRic), evenness (FEve), and divergence
(FDiv) (Villéger et al., 2008) using the dbFD function (Package ‘FD’, R
version 2.13.1). Also, to describe functional diversity, we calculated
community-based weighted functional diversity index (wFDc), which
was computed as the total branch length of the functional dendrogram,
including species community pool (Pla et al., 2012). The dendrogram
was computed using Ward clustering algorithm and Euclidean distances.
Finally, we calculated community-weighted means (CWMs) for each
plot and all traits (Casanoves et al., 2011) to represent the state of a trait
in the community (Pla et al., 2012). To examine differences between
forest types for CWMs of each trait, we computed a one-way ANOVA test
(ANOVA). Statistical analyses were carried out using R version 2.13.1.
(R Core Team, 2018).
Since some functional diversity metrics are correlated with species
richness (Laliberte and Legendre, 2010), it was not possible to compare
functional diversity components between plots and forests that differ in
their number of species. To solve such constrain, we used a null-model
type similarity profile routine (SIMPROF; Clarke et al., 2008) with
9999 permutation and Euclidean distance as a similarity measure. We
permuted the values of each functional diversity metric between plots
and forest. We used a functional diversity component per plot matrix,
and we interpreted the absence of structure (p > 0.05) as a similarity
expected under the null hypothesis that all functional diversity com­
ponents were calculated from the same forest. All SIMPROF analysis was
conducted in PRIMER software (Clarke and Gorley 2006).
3. Results
3.1. Species diversity and forest structure
We registered foliar and stem traits for 183 individuals comprising
53 species (116 individuals) in UF and 45 species (67 individuals) in FF.
This data included 31 families in UF and 27 in FF. Results show a sig­
nificant difference in stem density and the basal area between forests,
with a higher density recorded in UF (Table 1) but a larger basal area in
FF (Table 1). Results also show that species richness is higher in UF
compared to FF, a pattern that is also seen in species diversity (H) values
(Table 1). Similarly, when the rarefy curve was calculated, the species
richness was greater in UF compared to FF. In contrast, the basal area
L. Roa-Fuentes et al. Acta Oecologica 114 (2022) 103814

Table 1
Forest structure and woody plant species diversity and functional diversity in flooded and non-flooded forests in Magdalena River valley-Colombia. Tree with DBH
≥2.5 cm were sampled in 10 (50 × 2 m) plots in each forest type. BA: Basal area; LA: leaf area; SLA: specific leaf area, LDMC: Leaf dry matter content; WD: wood
density; FRic: functional richness; FEve: functional evenness; FDiv: functional divergence; wFDc: community-based weighted functional diversity index. Different
capital letters indicate means are significantly different among forest types. F and P are the statistics and significance values from ANOVA test. Pi and P are the statistics
and significance values from SIMPROF analysis.
Unflooded Forest Flooded Forest F P

Mean S.E. Mean S.E.

Forest structure
Stem density (ind/m2) 0.16A ± 0.014 0.13B ± 0.01 7.938 0.036
BA (m2/ha) 5.4B ± 0.11 10.02A ± 0.3 9.79 0.03

Species Diversity

S (Species richness) 13.8A ± 0.08 9.5B ± 0.89 12.81 0.002

S (rarefaction) 8.2A ± 0.23 7.4B ± 0.22 5.53 0.03
H′ (Shannon Wiener) 2.47A ± 0.07 2.12B ± 0.08 6.42 0.020

CWM-Trait values

LA (cm2) 250.67A ± 43.48 219.11A ± 23.78 0.351 0.56

SLA (m2/Kg) 14.21 B ± 0.39 16.88 A ± 1.02 4.86 0.040
LDMC (mg/g) 387.15 ± 9.21 390.14 ± 17.89 0.027 0.87
Leaf thickness (μm) 984.55 ± 189.78 772.04 ± 193.05 0.53 0.46
WD (g/cm3) 0.64 A ± 0.012 0.59 B ± 0.017 7.24 0.007

Functional Diversity Pi P

FRic 1.57 ± 0.42 0.25 ± 0.18 0 1

FEve 0.71 ± 0.03 0.73 ± 0.03 0 1
FDiv 0.69 ± 0.01 0.75 ± 0.02 0 1
wFDc 22.50A ± 3.34 17.58B ± 1.5 0.24 0.01

value was lower in UF compared with FF (Table 1). We found significant
differences between forest types when species compositions were
compared (R = 0.18; p = 0.024). We found 12 species accounting for
50% of total IVI in the UF forest and 12 species accounting for 56% of the
total IVI in FF. Both forest types shared four species (Annona sp., Cav­
anillesia platanifolia (Bonpl.), Kunth, and Pterocarpus cf. officinalis). One
specie was exclusive for UF (Brownea ariza Benth.), and four species
were exclusive for FF (Luehea seemannii Triana and Planch, Swartzia sp.,
Theobroma glaucum H. Karst, and Trichanthera gigantea (Bonpl.) Nees)
(Table 2; Supporting information, Appendix 1-2). We reported IVIs for
species that together account for more than 50% of total IVI in each
forest type (Table 2). A complete list of species with IVIs values is in
Supporting Information (Appendix 1-2).
3.2. Functional diversity
The presence of four plant functional groups (PFGs) was shown
(Supporting information, Appendix 3A). PCA allowed the explanation of
separation between PFGs (Supporting Information, Appendix 3B). The
first principal component (PC1) summarizes 34.12% of the variation

Table 2
Species frequency, density, dominance, and IVI (importance value index) of woody plant species in the unflooded and flooded forest. The last column indicates the
plant functional group (PFG).
Unflooded Forest (UF)

Species Family Frequency (%) Density (%) Dominance (%) IVI* PFG

Annona sp. Annonaceae 2.90 2.55 14.36 19.80 Thin-Con.

Lecythis mesophylla SA Mori Lecythidaceae 2.90 2.55 11.33 16.78 Small-Acq.
Clathrotropis brunnea Amshoff Fabaceae 3.62 3.06 9.29 15.97 Small-Acq.
Pseudolmedia laevigata Trécul Moraceae 5.07 5.61 3.51 14.19 Small-Acq.
Cavanillesia platanifolia (Bonpl) Kunth Malvaceae 0.72 0.51 11.69 12.92 Large-Acq.
Pouteria cf caimito (Ruiz & Pav) Radlk Sapotaceae 4.35 6.63 1.30 12.28 Thin-Con.
Gloeospermum sp. Violaceae 4.35 7.14 0.35 11.84 Small-Acq.
Brownea ariza Benth Fabaceae 5.07 5.61 0.98 11.66 Thin-Con.
Pouteria cf multiflora (A. DC.) Eyma Sapotaceae 3.62 4.08 2.66 10.37 Small-Acq.
Pterocarpus cf. officinalis Jacq Fabaceae 1.45 1.02 5.84 8.31 Small-Acq.
Virola flexuosa A.C. Sm. Myristicaceae 2.17 3.57 2.46 8.20 Small-Acq.
Hura crepitans L. Euphorbiaceae 2.17 1.53 4.46 8.17 Thin-Con.
Flooded Forest (FF)
Cavanillesia platanifolia (Bonpl.) Kunth Malvaceae 2.63 2.52 29.26 34.41 Large-Acq.
Annona sp. Annonaceae 5.26 5.88 17.97 29.11 Thin-Con.
Spondias purpurea L. Anacardiaceae 3.95 3.36 9.93 17.24 Large-Acq.
Swartzia sp. Fabaceae 5.26 3.36 3.77 12.39 Thin-Con.
Pterocarpus cf officinalis Jacq. Fabaceae 2.63 1.68 7.31 11.62 Small-Acq.
Gustavia superba (Kunth) O. Berg. Lecythidaceae 3.95 5.88 1.32 11.15 Large-Acq.
Pseudoxandra sp. Annonaceae 3.95 6.72 0.44 11.11 Thick-Con.
Jacaratia sp. Caricaceae 2.63 3.36 3.95 9.94 Small-Acq.
Theobroma glaucum Karsten Malvaceae 3.95 4.20 0.23 8.38 Small-Acq.
Luehea seemannii Triana & Planch. Malvaceae 2.63 3.36 1.65 7.65 Large-Acq.
Trichanthera gigantea (Bonpl) Nees Acanthaceae 1.32 5.88 0.21 7.41 Small-Acq.
Calyptranthes killipii Standl. Myrtaceae 2.63 4.20 0.26 7.09 Small-Acq.

4
L. Roa-Fuentes et al.
among traits, with LDMC and WD having the highest correlation scores
on this axis. PC2 accounted for 26.23% of the variation, with SLA having
the highest correlations scores (Table 3).
Species at the negative end of PC1 exhibited high WD and LDMC and
were classified into two PFG Thin-conservative (23 species) and Thick-
conservative (6 species) plant functional groups (Supporting Informa­
tion, Appendix 3A-4). Species at the positive end of PC1 had both high
SLA and FA values and were classified into two groups. Large-
Acquisitive (11 species) and Small-Acquisitive (24 species) (Support­
ing Information, Appendix 3A-4). When we compared the number of
species with the highest IVI (Table 2) considering the PFGs, we found for
FF and UF the functional groups as follow: Thin-conservative whit two
species (7 ind.) in FF and four species (20 Ind.) in UF, Thick-conservative
whit one specie (2 ind.) in FF, Large-Acquisitive with four species (8
ind.) in FF and Small-Acquisitive with seven species (26 ind.).
We found SLA-CWM and WD-CWM showed opposite patterns be­
tween UF and FF (Table 1). SLA-CWM was smaller, and WD-CWM was
greater, in UFs compared with FFs (Table 1). LA-CWM, LDMC-CWM and
LTh-CWM did not show differences between forest types (Table 1).
SIMPROF analysis showed that forest exhibit larger similarities than
expected under the null hypothesis (Pi = 0, P < 0.05) when three facets
of functional diversity (i.e., FRic, FEve, FDiv) were compared (Table 1).
In contrast, when wFD was compared between forest, we found struc­
ture (pi = 0.238, p < 0.05); i.e., both forests exhibited lower similarities
than expected under the null hypothesis (Table 1). UF showed the higher
wFDc value.”
4. Discussion
4.1. Species diversity and forest structure
The species richness in our study was 64, and the basal area was 7.71
m2 ha-1 (±0.205). These results fall below compared with values re­
ported in other forests in Magdalena River Valley. It is essential to
highlight that our sites have not been exposed to selective logging or
deforestation; thus, we suggest that lower plant richness and basal area
are the normal states in our study site. Pedraza and Molina (2006) re­
ported a basal area of 20 m2 ha-1, while Folster et al. (1976) reported a
basal area of 22 m2 ha-1 for trees with DBH >10 cm. Magdalena Valley
FF shows lower diversity than similar forests in Amazon (Gentry, 1988;
Stevenson et al., 2018); however, they are more diverse than other FF in
tropical regions around the world (Parolin and Wittmann 2010; Scarano
2006).
However, despite the low plant diversity shown in our study, it is in
concordance with the expected pattern, i.e., UF showed higher diversity
than FF. Such a pattern has been reported in other similar forests in the
Magdalena river basin and Amazon region (Luize et al., 2015; Haugaa­
sen and Peres 2006; Hawes et al., 2012; Stevenson et al., 2018; Aldana
et al., 2017). Lower diversity in FF is partially explained by the
ever-changing environment and waterlogging, which can be leading to a
constant replacement of plant species (Wittmann et al., 2006), making it
Table 3
Eigenvalues cumulative percent variation and eigenvectors of the first two
Principal Components (PCs) of five plant traits SLA specific leaf area, LDMC leaf
dry matter content, WD wood density. The PCA was used to define the functional
plant groups.
PC1 PC2
Eigenvalue 36.87 24.07
Cumulative percent variation 36.87 60.96
Eigenvectors of leaf traits
Leaf area 0.23 − 0.66
SLA 0.49 0.47
Leaf thickness − 0.25 − 0.35
LDMC − 0.61 − 0.12
WD − 0.52 − 0.46
5
Acta Oecologica 114 (2022) 103814
challenging to establish several species (Parolin and Wittmann 2010).
Only those species that have adaptations to this kind of environment
could survive. The high basal area in FF is associated with growth under
anoxic conditions prevalent in flooded plains (Parolin and Wittmann
2010).
Fabaceae, Rubiaceae, Lecythidaceae, and Moraceae were the most
species-rich families in both types of forests in our study, similar to
previous reports for this region (Idárraga and Pérez 2011; Pedraza and
Molina 2006). Fabaceae, Moraceae, and Arecaceae dominate the land­
scape in the Amazon region (both UF and FF, Wittmann et al.
2006Wittmann et al., 2006; Cano and Stevenson, 2008), while Bigno­
niaceae and Clusiaceae are dominant in flooded forests of Atlantic Forest
of Brazil (Scarano, 2006).
4.2. Functional diversity
In concordance with our expectation, tree communities in UF tend to
show lower CWM-SLA and higher CWM-WD values, which was inter­
preted as a suggestive conservative plant strategy (Chave et al., 2009;
Wright et al., 2004; Fortunel et al., 2014). These results can be mis­
interpreted in comparison with functional group analysis. In UF and FF,
most of the total species (54%, 56%, respectively) were classified as
acquisitive (Large-acquisitive and Small-acquisitive). As well as in the
group of species with the highest IVI values (67%, 75%, respectively).
The number of individuals in acquisitive groups was less in UF than in
the FF. For the plant community level, FF shows an acquisitive strategy
which contrast with the trend to low plant growth rate reported by
Homeier et al. (2017). We considered that fluctuating in FF’s water level
led it to remain in a transitional succession stage.
Higher functional diversity (wFD) in UF compared with FF had not
been reported before. The lower value of wFD in FF contrasts with
Parolin (2009) idea, which suggests that the plants’ strategy in flooded
forests exhibits a great diversity of morphological differences. However,
further research is needed to understand the variation of functional di­
versity of plant communities in the flooded and unflooded forest.
Although the functional richness (FRic) did not show differences be­
tween forest type, the most important species in UF includes species
from the four identified PFGs (Table 2), which suggests more functional
richness compared with FF (Pla et al., 2012). High values of FEve found
in both forest types (>0.7) can suggest high uniformity in species dis­
tribution across functional space, which could impact the resource uti­
lization and maximize productivity (Mason et al., 2005). Our result
highlights the importance of these forests and the need to investigate the
adaptations, especially because dry conditions are expected in global
change scenarios. Nonetheless, more research is necessary to prove such
a theoretical assumption.
5. Conclusions
In conclusion, our finding reinforces the idea that variation between
flooded and unflooded forests goes beyond the species list. The plant
community composition, structure, and plant functional diversity vary
between flooding regimes. Both species diversity and weighted func­
tional diversity are higher in the unflooded forest. We can suggest that
other factors than flooding regime affect the plants’ resource use. We
need to make an effort to know the factors controlling the plant
resource-use strategies in flooded and unflooded forests to understand
the potential effect of the droughts on climatic change.
Author contributions
LRF, LVP, JMC performed the experiments and field data collection;
LRF, analyzed the data; LRF, MAJ wrote and edited the manuscript.
L. Roa-Fuentes et al.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgments
LVP thanks Universidad Industrial de Santander for logistic support,
UISH for hosting botanical material, and staff at the Herbarium of the
Universidad de Antioquia for help with identifying plant material. We
thank Fundación Biodiversa for funding the data collection for this
research.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.actao.2022.103814.
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