Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

Examining functional diversity in two plant communities

under ecological restoration in Farallones de Cali, Colombia

Ricardo Cardona1; https://orcid.org/0000-0002-0833-7586

Alba Marina Torres González1; https://orcid.org/0000-0002-3010-2505

1. Departamento de Biología, Universidad del Valle, Calle 13 # 100 – 00, Cali, Valle del Cauca, Colombia; crcardonav@

gmail.com (*Correspondence), alba.torres@correounivalle.edu.co

Received 08-III-2024. Corrected 02-I-2025. Accepted 07-V-2025.

ABSTRACT

Introduction: Functional diversity is crucial in understanding species performance and monitoring ecological

restoration processes. In Valle del Cauca, Colombia, where only a small percentage of tropical dry forest remains,

ongoing ecological restoration efforts are vital. However, monitoring of restoration efforts is typically not

conducted.

Objective: To compare the functional diversity of two plant communities, restored two and eight years ago, in

the Loma Larga reserve, Colombia.

Methods: We assessed nine functional traits in the five most significant species of each community. The analysis

included contrast tests for functional trait differences, as well as functional diversity indices.

Results: The 8-year community displayed greater values for maximum height, diameter at breast height, and

specific leaf area. Conversely, the 2-year community exhibited higher leaf thickness. Moreover, the 8-year com-

munity presented the highest values in the functional indices: richness, evenness, divergence, dispersion, and

specialization.

Conclusions: Ecological restoration had a positive impact on plant communities, as evidenced by increased

functional diversity and structural complexity in the 8-year community compared to the 2-year community. This

suggests that ecological succession processes advance significantly over time, leading to more resilient and func-

tionally diverse communities. The analysis of functional traits stands out as an effective tool for monitoring the

success of restoration and guiding future efforts in critically threatened ecosystems such as tropical dry forests.

Keywords: ecological succession; functional ecology; functional traits; plant communities; tropical dry forest.

RESUMEN

Examinando la diversidad funcional en dos comunidades vegetales

bajo restauración ecológica en Farallones de Cali, Colombia

Introducción: La diversidad funcional se usa para entender el desempeño de las especies y monitorear procesos

de restauración ecológica. En el Valle del Cauca, Colombia, solo queda un pequeño porcentaje del bosque seco

tropical original, por lo que los planes de restauración ecológica son necesarios, pero no se realiza monitoreo

de estos.

Objetivo: Comparar la diversidad funcional de dos comunidades vegetales, con dos y ocho años desde la restau-

ración ecológica en la reserva Loma Larga, Colombia.

Métodos: Se midieron nueve rasgos funcionales para las cinco especies más importantes de cada comunidad.

Los análisis incluyeron pruebas de contraste para diferencias entre los rasgos funcionales, e índices de diversidad

funcional.

https://doi.org/10.15517/rev.biol.trop..v73i1.59149

ECOLOGÍA TERRESTRE

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

INTRODUCTION

Functional diversity (FD) examines the

connection between ecosystem diversity, struc-

ture, and function. It assesses biological success

by comparing functional traits (FTs) among

species and calculating FD indices across

communities (Mason & de Bello, 2013). FTs

encompass individual measurements of mor-

phological, physiological, and phenological

characteristics that can be extrapolated to the

community level (Córdova-Tapia & Zambrano,

2015). These traits enable the evaluation of spe-

cies’ growth, reproduction, and survival, as well

as their responses to environmental factors and

their impact on ecosystem processes (Tedesco

et al., 2023). Salgado-Negret (2016) stated that

FTs provide insights into the responsiveness of

plant communities to climate change and the

effects of environmental gradients on inter-

specific variation. This author also pointed

out that plant FTs are categorized into veg-

etative, foliar, and regenerative traits. Vegeta-

tive traits offer information about persistence,

competition, and longevity; foliar traits are

linked to establishment; and regenerative traits

are associated with dispersal and colonization

(Rosell et al., 2022).

FD has emerged as an approach for moni-

toring and evaluating anthropogenic impacts

on ecosystems, as it is more sensitive to envi-

ronmental changes than species loss alone

(Tedesco et al., 2023). FD has been used to

assess the recovery of degraded areas during

restoration processes by analyzing FTs and

comparing FD indices (Qin et al., 2016; Rosell

et al., 2022). Using FTs in restoration studies

and projects has gained attention due to the

ease of measuring soft traits and the important

insights that they provide (Carlucci et al., 2020).

For example, tree height (Ht) and diameter at

breast height (DBH) are used to understand

carbon storage dynamics within plant com-

munities (Mensah et al., 2016). Similarly, leaf

thickness (LT) correlates with hydric balance

and herbivory protection (Sandoval-Granillo

& Meave, 2023), while specific leaf area (SLA)

is linked to light capture efficiency and water

regulation (Mensah et al., 2016).

Building on these trait-based insights,

functional diversity is quantified using indices

that capture different aspects of trait distribu-

tion within communities. The main FD indices

proposed by Mason et al. (2005) include rich-

ness (FRic), evenness (FEve), and divergence

(FDiv). Other indices such as specialization

(FSpe) (Villéger et al., 2010) and dispersion

(FDis) (Laliberté & Legendre, 2010) have also

been proposed. These indices provide insights

into community functioning patterns and

their variations along environmental gradients

(Mason & de Bello, 2013). For example, high

species richness and FEve are related to a low

likelihood of losing functional groups in a

plant community (Fonseca & Ganade, 2001).

Although the FD field of research has increased

in recent years, there are still information gaps

Resultados: La comunidad de 8 años presentó mayores valores de altura máxima, diámetro a la altura del pecho,

y área foliar específica. La comunidad de 2 años exhibió mayores valores de grosor foliar. La comunidad de 8

años presentó los mayores valores en los índices funcionales: riqueza, equitatividad, divergencia, dispersión, y

especialización.

Conclusiones: La restauración ecológica tuvo un impacto positivo en las comunidades vegetales, evidenciado por

una mayor diversidad funcional y complejidad estructural en la comunidad de 8 años en comparación con la de

2 años. Esto sugiere que los procesos de sucesión ecológica avanzan significativamente con el tiempo, resultando

en comunidades más resilientes y funcionalmente diversas. El análisis de rasgos funcionales se destaca como

una herramienta efectiva para monitorear el éxito de las restauraciones y guiar futuros esfuerzos en ecosistemas

críticamente amenazados como el bosque seco tropical.

Palabras clave: sucesión ecológica; ecología funcional; rasgos funcionales; comunidades vegetales; bosque seco

tropical.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

in several ecosystems globally. Most conserva-

tion studies have focused on tropical moist

forests, and tropical dry forests (TDFs) remain

poorly understood in comparison (Siyum,

2020). Furthermore, TDFs are critically endan-

gered, and the use of FD may give insights into

the resilience of these ecosystems that are fac-

ing climate change and anthropic disturbances

(Dexter et al., 2018).

Colombia’s TDF has experienced a signifi-

cant decline, with over 90 % of its original cover

lost due to agricultural and livestock expansion,

deforestation, and mining (Ruíz et al., 2023).

The TDF ecosystem is highly fragmented and

critically endangered, with Valle del Cauca hav-

ing 6 303 ha of remaining natural state TDF

and 53 184 ha of small, unconnected fragments

(García & González-M., 2019). In the Loma

Larga reserve, situated in the foothills of the

Farallones de Cali, patches of TDF face various

stressors, such as the invasion of species like

bracken fern (Pteridium aquilinum (L.) Kuhn),

forest fires, grazing, and deforestation. Ecologi-

cal restoration efforts have aimed to mitigate

TDF loss in the Loma Larga area. These efforts

have included revegetation using native plant

species and suppressing bracken fern in the res-

toration zones. However, systematic monitor-

ing of these restoration areas to evaluate their

progress and outcomes has yet to be conduct-

ed (Corredor, G., personal communication).

Monitoring is a crucial component of restora-

tion processes, as it enables the evaluation and

potential adjustment of activities to enhance

the long-term effectiveness and success of the

restoration program (Ruíz et al., 2023).

Since FD is an effective tool for evaluating

restored areas, this research aimed to evaluate

and compare functional diversity traits and

indices between two restored plant communi-

ties in the Loma Larga reserve, situated in the

piedmont of Los Farallones de Cali. Addition-

ally, this study sought to assess the progress of

the restoration process in the examined plant

communities by utilizing functional diversity as

an analytical tool.

MATERIAL AND METHODS

Study area: We conducted the research

in the ecological restoration areas of the Loma

Larga Reserve (3°20’ N & 76°33’ W), located

in El Peón village, foothills of the Farallones

de Cali, township of Pance, municipality of

Santiago de Cali, Valle del Cauca, Colombia.

Fieldwork took place between March and May

2022. The reserve encompasses an altitudinal

range of 1 100 to 1 350 m.a.s.l., with average

temperatures between 17 and 24 °C, and annual

rainfall from 1 220 to 1 640 mm (Sardi et al.,

2018). The area represents a transitional zone

between the TDF and premontane rainforest

(Sardi et al., 2018). The plant species composi-

tion in the area includes Jacaranda caucana

Pittier, Calliandra pittieri Standl., and Cecropia

angustifolia Trécul, among others (Botina &

García, 2005; Sardi et al., 2018).

Experimental design: We chose two zones

within the ecological restoration areas to con-

duct our study, including an older community

that had undergone restoration eight years ago

and a more recent community that had under-

gone restoration two years ago. Native plant

species were planted in both zones, while intro-

duced species, specifically Clitoria fairchildiana

R.A. Howard and Acacia mangium Willd., were

used in the 8-year community. Ten rectangu-

lar plots measuring 25 x 4 m were randomly

established in each community, resulting in a

total area of 1 000 m2 per community (Rangel

& Velázquez, 1997). We calculated the Impor-

tance Value Index (IVI) in each community

following the methodology proposed by Rangel

and Velázquez (1997). We sampled individuals

with a DBH greater than 1 cm and followed the

measurement approach described by Leverett

and Bertolette (2013). Likewise, we identified

the species using the species guide by Botina

and García (2005), with validation performed

at the CUVC Herbarium of the Universidad

del Valle.

To identify the most significant species,

we ranked them using their IVI. Based on this

ranking, we selected the top five species with

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

the highest IVI in each community, ensuring

that these five species collectively represented at

least 50 % of the total IVI within each commu-

nity. We then randomly selected five individu-

als from each of the five previously identified

species per community, following the method-

ology outlined by Salgado-Negret et al. (2016),

resulting in a total of 25 individuals sampled

per community. Finally, we used reflective tape

to mark these individuals and took measure-

ments for three vegetative traits, five foliar

traits, and two regenerative traits.

Functional traits: The vegetative FTs

assessed for each species included bark type,

classified according to Ramírez-Padilla and

Goyes-Acosta (2004). We measured Ht and

DBH following the approach proposed by Lev-

erett and Bertolette (2013). Regarding foliar

FTs, we employed the methodology outlined by

Pérez-Harguindeguy et al. (2013). We collected

ten leaves from each individual, for a total of 250

leaves per community. We characterized leaf

type and the presence of indumentum for each

species. Additionally, we measured LT using

a REXBETI digital micrometer, dry weight

using a Mettler Toledo® analytical balance, and

leaf area using ImageJ software (Schneider et

al., 2012). We calculated SLA following the

methodology proposed by Salgado-Negret et

al. (2016); for species with compound leaves, we

randomly selected a leaflet for measurements

(Pérez-Harguindeguy et al., 2013). For regen-

erative FTs, we characterized fruit type and

dispersal syndrome based on a literature review,

following guidelines by Pérez-Harguindeguy et

al. (2013).

For quantitative FTs, we conducted two-

sample tests to compare the mean values

between the communities. We used Mann-

Whitney tests to compare LT, Ht, and SLA;

and a Student’s t-test to compare DBH. These

statistical analyses were performed using

PaST (Hammer et al., 2001). For functional

diversity analysis, we computed a PCA analy-

sis, as well as the FRic, FEve, FDiv, FDis,

and FSpe indices using the “mFD” package

(Magneville et al., 2022) in R 4.4.1 with RStudio

(R Core Team, 2024).

RESULTS

Community composition: The composi-

tion of plant communities differed between the

8-year and 2-year restoration areas at the land-

scape level. The 8-year community featured a

closed canopy with abundant leaf litter in the

understory and scattered patches of bracken

fern. In contrast, the 2-year community had

an open canopy with minimal leaf litter and

was dominated by grass cover. A total of 494

individuals were identified across both areas,

with 277 individuals belonging to 30 species

in the 2-year community and 217 individuals

belonging to 31 species in the 8-year commu-

nity (SMT 1).

Figure 1 shows the distribution of diamet-

ric classes in both communities. The 2-year

community had a higher proportion of indi-

viduals in diameter classes I (79 %) and II (16

%), with a significant decline in class III (5 %),

and no individuals in classes IV and above. In

contrast, the 8-year community had a lower

percentage of individuals in classes I (58 %) and

II (13 %), with a gradual decrease from class III

to VIII: 8 % in class III, 8 % in class IV, 5 % in

class V, 4 % in class VI, 3 % in class VII, and 1

% in class VIII. There were no individuals in

class IX, and only one Ladenbergia oblongifo-

lia (Mutis) L. Andersson individual in class X

(0.5 %). Both communities exhibited a reverse

J-shaped distribution pattern (Fig. 1).

Regarding the IVI, the five species with

highest IVI (55.2 % in total) in the 2-year com-

munity were Miconia rubiginosa (Bonpl.) DC.-

20.1 %, Miconia minutiflora (Bonpl.) DC.-12.5

%, Erythroxylum citrifolium A.St.-Hil.-9.4 %,

Miconia prasina (Sw.) DC.-7.4 %, and Persea

caerulea (Ruiz & Pav.) Mez.-5.7 %. In the 8-year

community, the five species with the highest

IVI (63.2 %) were Didymopanax morototoni

(Aubl.) Decne. & Planch.-25.2 %, Henriettea

seemannii (Naudin) L.O. Williams-11.4 %, M.

minutiflora-9.9 %, C. fairchildiana-8.6 %, and

Eugenia egensis DC-8.1 %. The only species

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

shared by both communities was M. minuti-

flora. The selected species were also the most

abundant (SMT 1), except for Genipa ameri-

cana L., whose individuals were smaller and

had a limited impact on the IVI.

Functional traits and diversity indices:

Table 1 summarizes the evaluated FTs, includ-

ing qualitative traits with their corresponding

character states for each species, and quantita-

tive traits with average values and standard

deviations for each species. The most frequent

character states for the five qualitative FTs

evaluated in the two communities were sim-

ple leaf, glabrous indumentum, fissured bark,

berry-like fruit, and zoochorous dispersal.

Regarding quantitative FTs, the contrast tests

revealed significantly higher mean values of Ht

Fig. 1. Comparison of the diameter class distribution in each community.

Table 1

Summary of functional traits in the Loma Larga Reserve.

8-year community

Species IVI (%) Lf Im Bk Ft Sd LT (mm) SLA (cm2/g) Ht (m) DBH (cm)

D. morototoni 25.2 Co Pu Le Be Zo 0.27 ± 0.05 68.6 ± 33.4 10.2 ± 1.6 14.3 ± 5.5

H. seemannii 11.4 Si Pu Fi Be Zo 0.13 ± 0.02 209.9 ± 35.4 5.7 ± 2.0 8.8 ± 5.3

M. minutiflora 9.9 Si Gl Fi Be Zo 0.09 ± 0.02 196.9 ± 49.5 6.0 ± 0.5 5.8 ± 0.8

C. fairchildiana 8.6 Co Gl Le Lg Au 0.10 ± 0.01 226.2 ± 38.2 7.9 ± 2.3 15.6 ± 9.0

E. egensis 8.1 Si Gl Fi Be Zo 0.14 ± 0.02 117.1 ± 23.6 4.3 ± 1.2 3.1 ± 3.1

Trait’s mean value 0.15 ± 0.07 163.7 ± 71 6.3 ± 2.9 9.5 ± 7.0

2-year community

Species IVI (%) Lf Im Bk Ft Sd LT (mm) SLA (cm2/g) Ht (m) DBH (cm)

M. rubiginosa 20.1 Si Pu Fi Be Zo 0.49 ± 0.08 70.7 ± 13.1 3.4 ± 0.4 6.1 ± 2.1

M. minutiflora 12.5 Si Gl Fi Be Zo 0.17 ± 0.03 130.6 ± 42.7 3.1 ± 2.0 3.1 ± 2.9

E. citrifolium 9.4 Si Gl Fi Be Zo 0.25 ± 0.02 11.2 ± 2.4 3.5 ± 1.1 2.5 ± 1.0

M. prasina 7.4 Si Pu Fi Be Zo 0.20 ± 0.02 84.5 ± 22.0 4.9 ± 1.3 6.0 ± 3.0

P. caerulea 5.7 Si Gl Le Dr Zo 0.21 ± 0.02 9.17 ± 12.8 2.5 ± 0.3 2.1 ± 0.6

Trait’s mean value 0.27 ± 0.12 99.0 ± 31.7 4.1 ± 1.7 3.9 ± 2.4

Lf: Leaf type, Co: Compound leaves, Si: Simple leaves; Im: Indumentum presence, Gl: Glabrous, Pu: Pubescent; Bk: Bark

type, Le: Lenticellate, Fi: Fissured; Ft: Fruit type, Lg: Legume, Be: Berry, Dr: Drupe; Sd: Seed dispersal, Au: Autochory, Zo:

Zoochory; LT: Leaf thickness; SLA: Specific leaf area; Ht: Height; DBH: Diameter at breast height.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

(Fig. 2A), DBH (Fig. 2B), and SLA (Fig. 2C) in

the 8-year community. Finally, the 2-year com-

munity exhibited a significantly higher mean

LT value (Fig. 2D).

The position occupied by each species in

the functional space is shown in Fig. 3A. The

cumulative percentage of variance explained by

PC1 and PC2 was 73.8 %. The values obtained

for the FD indices (2-year community vs. 8-year

community) were: 0.012 vs. 0.032 for FRic (Fig.

3B), 0.539 vs. 0.916 for FEve (Fig. 3C), 0.867

vs. 0.907 for FDiv (Fig. 3D), 0.308 vs. 0.784

for FDis (Fig. 3E), and 0.424 vs. 0.782 for FSpe

(Fig. 3F). All indices exhibited higher values in

the 8-year community compared to the 2-year

community. Among the indices, FRic had the

lowest values. The smallest difference between

the two communities was obtained for the FDiv

index, whereas the greatest differences between

the two communities were found for the FDis,

FEve, and FSpe indices.

DISCUSSION

Plant communities: The two areas studied

exhibited a reverse J-shaped curve regarding

the diametric classes, which is typical of early

successional forests. As time progresses, the

abundance of individuals shifts towards higher

diameter classes due to biomass accumulation.

This shift facilitates the replacement of younger

individuals by older ones, reflecting the succes-

sion dynamics of a plant community (Imaña-

Encinas et al., 2021). The species with highest

IVI selected in the two communities represent

the vegetation of the TDF ecosystem in Valle

del Cauca, except for the introduced species C.

fairchildiana (Fabaceae), according to Bernal

Fig. 2. Comparison between quantitative traits in each community. A. Height. B. Diameter at breast height. C. Specific leaf

area. D. Leaf thickness. Significant differences are indicated by asterisks, where ** p ≤ 0.01, *** p ≤ 0.001.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

Fig. 3. Comparison of functional diversity indices between the two communities. A. Position of species in the functional

space. B. Functional richness. C. Functional evenness. D. Functional divergence. E. Functional dispersion. F. Functional

specialization. In Figure 3A, A denotes 2-year community species, B denotes 8-year community species, and AB denotes

species occurring in the two communities. 1A. M. rubiginosa, 2A. M. prasina, 3A. P. caerulea, 4A. E. egensis, 1B. E. citrifolium,

2B. H. seemannii, 3B. D. morototoni, 4B. C. fairchildiana, AB. M. minutiflora. Bluish green represents the 2-year community,

whilst olive yellow represents the 8-year community. The axes are the PCoA axes in the Functional Space: where PC1 is

driven by DBH (p = 0.0077); Height (p = 0.0230); Leaf type (p = 0.404) and Bark type (p = 0.0201), whilst PC2 is driven by

SLA (p = 0.0457) and Indumentum presence (p = 0.0143).

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

et al. (2016). Most of the sampled species are

pioneer plants with a shrubby habit, except for

P. caerulea (Lauraceae), D. morototoni (Aralia-

ceae), and C. fairchildiana, which are arboreal,

fast-growing species typically found in an inter-

mediate successional stage (da Silva et al., 2020;

Sardi et al., 2018). D. morototoni and C. fair-

childiana were exclusively found in the 8-year

community and exhibited the tallest height and

largest DBH among the sampled individuals.

Conversely, young P. caerulea individuals were

only present in the 2-year community, resulting

in lower Ht values for this species.

C. fairchildiana is a species native to Bra-

zil and introduced to Colombia. It has been

observed to form root nodules through symbi-

otic relationships with nitrogen-fixing bacteria,

making it suitable for restoring areas with

degraded soils (da Silva et al., 2020). These

characteristics made it an appropriate choice

for the initial stage of the restoration program

in the Loma Larga reserve. D. morototoni is

a fast-growing native tree with palmate com-

pound leaves that accumulate as leaf litter in

the understory (da Silva et al., 2020). The genus

Miconia, from the family Melastomataceae,

was represented by three species, all native

and characterized by a fast-growing shrubby

habit and high abundance in the TDF (Sardi

et al., 2018). However, only M. minutiflora

occurred in both communities. According to

the restoration plan, a combination of native

and introduced species, including C. fairchil-

diana, was initially used; the restoration plan

was subsequently refined and focused solely on

native species. According to information from

Corredor (personal communication), no one

intentionally planted Miconia or D. morototoni

species, suggesting that their presence in the

sampled areas resulted from natural establish-

ment, indicating ecosystem recovery and a link

with other patches (Ruíz et al., 2023). E. citrifo-

lium (Erythroxylaceae), E. egensis (Myrtaceae),

and H. seemannii (Melastomataceae) are native

pioneer shrubs that are characteristic of the

TDF species assemblage (Corredor-Londoño

et al., 2020). P. caerulea is a native tree, reach-

ing heights of up to 20 m, and is typical of the

TDF. This is a valuable species for its landscape

quality; it is used in various urban contexts

(Núñez-Florez et al., 2019) and is commonly

found in forests at an intermediate successional

stage (Sardi et al., 2018).

Functional traits: The two communities

differed in their structural composition, with

the 8-year community exhibiting traits typical

of a more advanced successional stage, com-

pared with the younger community. However,

the 8-year community was still at an early suc-

cessional stage when compared to intermediate

and advanced plant communities in the Colom-

bian TDF (Ruíz et al., 2023). Younger indi-

viduals tend to invest more metabolic energy

in vertical growth rather than in accumulation

and secondary growth (Matsuo et al., 2021).

This explains the lower SLA values found in

the 2-year community as well as the greater LT,

which aids in moisture retention and protec-

tion against herbivory, providing thus essential

benefits in the early stages of succession (Ruíz

et al., 2023), when plants are vulnerable to

temperature shifts, droughts, and solar irradi-

ance as occurs in the TDF (Sandoval-Granillo

& Meave, 2023).

The dissimilarities in DBH values found in

the two communities may be explained by the

rapid growth of individuals in the 8-year com-

munity, along with light heterogeneity, which

allows a higher interception of light by canopy

trees (Matsuo et al., 2021). With a direct source

of light and higher SLA values, as was seen in

the 8-year community, individuals can perform

further secondary growth that leads to dif-

ferentiation in the diametric classes (Matsuo

et al., 2021; Rosell et al., 2022). Furthermore,

plant communities in early and intermediate

successional stages are adapted to intense light

conditions, and display a wide range of qualita-

tive traits that improve dispersion and estab-

lishment (Carlucci et al., 2020). These species

are mainly light-demanding species that create

the canopy cover and the conditions for the

establishment and growth of the shade-tolerant

species that will replace them in later succes-

sion (Matsuo et al., 2021).

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It is worth emphasizing that Ht, DBH,

SLA, and LT are soft traits with straightforward,

cost-effective measurement methods, making

them valuable for incorporation into restora-

tion project planning (Carlucci et al., 2020).

These metrics provide an accurate representa-

tion of energy and carbon fluxes, as well as

trends within plant communities (Carlucci et

al., 2020; Pérez-Harguindeguy et al., 2013), and

the analysis of these patterns can reveal the

successional stage of plant communities (Mat-

suo et al., 2021; Rosell et al., 2022; Ruíz et al.,

2023). Additionally, FTs can guide adjustments

in restoration project planning. For example,

if a community exhibits lower trait values, this

might be indicative of incorrect species selec-

tion or of the influence of variables that may be

identified and managed. The key is to identify a

set of FTs that are easily measurable and highly

informative about the specific conditions and

dynamics of the community.

Functional Diversity Indices: The varia-

tion in FD indices between the two com-

munities could be attributed to the structural

complexity typically associated with advanced

successional stages, driven by the temporal gra-

dient (Ruíz et al., 2023). For instance, interme-

diate successional species such as D. morototoni

and C. fairchildiana contributed significantly to

the increased FD indices in the 8-year commu-

nity. These species exhibited higher FT values

linked to biomass accumulation, including Ht,

DBH, SLA, and presence of compound leaves

(Li et al., 2022).

The presence of D. morototoni and C.

fairchildiana in the 8-year community also

contributed to higher FSpe, which quantifies

the differentiation among species within the

community. Elevated FSpe values indicate a

community composed of specialist species that

exploit distinct ecological niches (Córdova-

Tapia & Zambrano, 2015). Consequently, the

specialized species in the older community

increased Fric, which measures the volume of

ecological space occupied by the community.

This niche differentiation enhances resilience

to environmental fluctuations and invasions

by other species, while also increasing ecosys-

tem productivity (Córdova-Tapia & Zambrano,

2015; Schleuter et al., 2010).

It is important to note that the relatively

low FRic values observed in this study result-

ed from analyzing only five species in each

community. However, the higher productiv-

ity observed in the 8-year community may be

linked to the greater divergence in resource

acquisition strategies among dominant species,

as represented by FDiv (Córdova-Tapia & Zam-

brano, 2015; Schleuter et al., 2010). Although

FDiv was slightly higher in the 8-year commu-

nity, the difference with the 2-year community

was minimal, suggesting that both communities

contained species with considerable variabil-

ity in resource acquisition strategies (Córdova-

Tapia & Zambrano, 2015). This variability was

further evidenced by FDis, which showed the

most significant differences between the two

communities. The higher FDis in the 8-year

community reflected greater FT specialization

and variability, indicating an enhanced capacity

to respond to environmental changes and sug-

gesting increased resilience (Cooke et al., 2019).

Additionally, the 8-year community dis-

played lower species dominance and a more

equitable abundance distribution, as indicated

by the higher FEve. This index reflects unifor-

mity in resource use among species, as well as

the distribution of species abundance within

the community (Mason & de Bello, 2013). In

contrast, the 2-year community exhibited lower

FEve values, indicating the presence of species

conglomerates where a few dominant species

overshadowed others with poor representation

(Córdova-Tapia & Zambrano, 2015; Schleuter

et al., 2010).

Applications and limitations: The pres-

ence of key species with desirable functional

traits (e.g. D. morototoni in Colombian TDF)

can be an indicator of a more advanced suc-

cessional stage. Additionally, integrating plant

community metrics, such as the IVI, FTs, and

FD indices would enhance restoration projects

and their monitoring strategies. (Carlucci et al.,

2020). Comparing plant communities across

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

temporal gradients is essential for tracking

functional trait dynamics and evaluating long-

term community development, whereas high

FD indices correlate with enhanced ecosystem

services, which provide a wide range of benefits

for local communities dependent on those eco-

systems (Matsuo et al., 2021; Ruíz et al., 2023).

However, several limitations must be

addressed. First, selecting FTs that accurately

represent community dynamics is challeng-

ing, particularly in species-rich ecosystems.

In such cases, IVI can help (Joshi et al., 2024;

Rangel & Velázquez, 1997). Second, measur-

ing FTs and obtaining FD indices often require

years of data collection to become meaningful,

and differences observed within specific com-

munities may not be generalizable to others

(Carlucci et al., 2020; Mason & de Bello, 2013;

Rosell et al., 2022). Despite these challenges,

further research is urgently needed, particularly

in vulnerable ecosystems such as tropical dry

forests, where effective and practical restoration

protocols are critical for ecological and social

resilience.

The results of this study emphasize the

positive impact of ecological restoration efforts

on the functional diversity and structural com-

plexity of plant communities in the Loma Larga

reserve. The 8-year community exhibited a

more advanced successional stage character-

ized by higher functional trait values (e.g., Ht,

DBH, and SLA) and elevated functional diver-

sity. These metrics suggest that the older com-

munity has a broader functional niche, greater

species differentiation, and more equitable dis-

tribution of species abundances compared to

the 2-year community. The results emphasize

the ecological benefits of using functional traits

as indicators to monitor the progress and effec-

tiveness of restoration initiatives. Functional

diversity analysis provides insights into the

contribution of species to ecosystem resilience,

stability, and productivity. In particular, the

presence of key species, such as D. morototoni

and C. fairchildiana, highlights their signifi-

cant role in shaping community dynamics and

enhancing ecosystem functionality.

This research contributes to the grow-

ing understanding of ecological restoration in

tropical dry forests, a critically endangered eco-

system. The temporal gradient revealed by the

comparison of 2-year and 8-year restored com-

munities highlights the importance of long-

term monitoring to evaluate the effectiveness

of restoration programs. Future efforts should

focus on refining restoration protocols by pri-

oritizing species selection based on functional

traits, improving connectivity between restored

patches, and addressing challenges such as

invasive species management. By providing

a robust framework for assessing restoration

outcomes, this study supports the development

of evidence-based strategies to restore and con-

serve tropical dry forests. It also underscores

the need for further research to explore the role

of functional diversity in enhancing ecosystem

resilience and its applications in restoration

ecology. Continuous investment in restoration

initiatives and monitoring will be critical for

safeguarding these ecosystems and the services

they provide to both biodiversity and human

communities.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict 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 acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

See supplementary material

a31v73n1-suppl1

ACKNOWLEDGMENTS

We wish to thank the Cooperativa Loma

Larga, Germán Corredor, Aida Baca, Zara Pla-

zas, Andrés Ocampo, Wilmar Torres, Olga

Lucía Torres, and Fernando Zapata.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e59149, enero-diciembre 2025 (Publicado May. 26, 2025)

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