Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

Altitudinal gradients drive strong species turnover

in epigeous ant communities of the Colombian Andes

Vianny Plaza-Ortega1*; https://orcid.org/0000-0003-0886-1393

Yamid Mera-Velasco2; https://orcid.org/0000-0002-5221-8824

Inge Armbrecht3; https://orcid.org/0000-0003-0574-2559

James Montoya-Lerma3; https://orcid.org/0000-0003-2122-1323

1. Programa de doctorado en Ciencias-Biología, Universidad del Valle, Campus Meléndez Calle 13 # 100-00, Cali, Valle

del Cauca, Colombia; vianny.plaza@correounivalle.edu.co (*Correspondence)

2. Departamento de Biología, Facultad de Ciencias Naturales, Exactas y de la Educación, Universidad del Cauca, Sede

Tulcán Popayán, Cauca, Colombia; ymera@unicauca.edu.co

3. Departamento de Biología, Sección Entomología, Universidad del Valle, Campus Meléndez Calle 13 # 100-00, Cali,

Valle del Cauca, Colombia; inge.armbrecht@correounivalle.edu.co, james.montoya@correounivalle.edu.co

Received 07-VII-2025. Corrected 25-VIII-2025. Accepted 15-IV-2026.

ABSTRACT

Introduction: Epigeous ants are widely used as bioindicators to test ecological and biogeographic hypotheses. In

tropical regions, elevation is a key driver of biodiversity patterns due to the rapid ecosystem shifts with increas-

ing altitude.

Objective: To examine the distribution and turnover of epigeous ants along an altitudinal gradient (1 000-2 800

m asl) on the opposite slopes of the Western and Central Andean cordilleras in the tropical belt of Colombia,

South America.

Methods: Intensive field sampling of ants was conducted, and Zeta diversity was used as an analytical metric to

assess spatial diversity components within a multi-site partitioning framework, given its sensitivity to richness

differences among habitat types.

Results: A total of 204 ant species were recorded: 128 on the Western Cordillera and 131 on the Central

Cordillera. Myrmicinae was the most diverse subfamily, represented across all elevational bands. The mid eleva-

tion band (1 500 m asl) exhibited the highest diversity in both cordilleras, while the 2 800 m asl band showed the

lowest. The 1 500 m band on the Central Cordillera harbored 71 species, in contrast, the 2 800 m band on the

Western Cordillera had only two species. Ant community composition on opposite flanks of the Western and

Central Cordilleras showed a high turnover, with only three species occurring across all five elevational bands in

both ranges. The ant communities were dominated by rare species or those with low capture frequencies, likely

reflecting the interaction of biotic and abiotic factors specific to each elevational life zone.

Conclusions: These findings confirm that each altitudinal gradient is home to specific ant communities that

respond to the environmental, historical, and biogeographical conditions of each mountain range. Likewise, it

is confirmed that environmental and spatial factors determine the composition of ants and the replacement rate

of the community.

Key words: ant diversity; environmental filtration; distribution patterns; strategic ecosystems; tropical rainforest.

https://doi.org/10.15517/wfnx9b94

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

INTRODUCTION

Colombia is located within the tropical

belt and features a unique geographic position

that harbors a remarkable diversity of ecosys-

tems, ranging in elevation, from sea level to

high Andean peaks (Pérez-Escobar et al., 2022;

Rangel-Ch., 2015). This altitudinal complexity

is shaped by the presence of the three Andean

cordilleras (Western, Central, and Eastern) that

traverse the country (Iriarte, 2006; Myers et al.,

2000; Narváez-Vidal et al., 2024). The climatic

shifts driven by this mountainous geomor-

phology pose adaptive challenges for ecto-

thermic organisms, including ants, a highly

diverse and ecologically dominant insect group

in tropical ecosystems (Folgarait, 1998). Ants

are highly sensitive to changes in temperature,

humidity, precipitation, habitat transforma-

tion, and anthropogenic pressures (Guerrero

& Sarmiento, 2010; Kaspari & Majer, 2000;

Stuble et al., 2013).

Globally, ants inhabiting elevational gradi-

ents have been the subject of numerous studies,

and Colombia is not the exception. Regional

studies (v.g. Guerrero & Sarmiento, 2010) on

the Northwestern slope of the Sierra Nevada

de Santa Marta, and more localized efforts,

such as those conducted in Tatamá National

Natural Park (Idarrága-Giraldo, 2021) and Far-

allones de Cali National Natural Park (Narváez-

Vidal et al., 2024), consistently report a decline

in species richness with increasing elevation.

These contributions are valuable not only for

their findings but also for their spatial rep-

licability and their capacity to reveal sharp

environmental changes over relatively short

distances (Kaspari & Majer, 2000; Longino &

Branstetter, 2019; Longino & Colwell, 2011;

Sanders, 2002). Beyond richness patterns, other

RESUMEN

Gradientes altitudinales impulsores del fuerte recambio de especies en las comunidades

de hormigas epigeas de los Andes colombianos

Introducción: Las hormigas epigeas se utilizan ampliamente como bioindicadores para comprobar hipótesis eco-

lógicas y biogeográficas. En las regiones tropicales, la altitud es un factor clave en los patrones de biodiversidad

debido a los rápidos cambios en los ecosistemas a medida que aumenta la altitud.

Objetivo: Examinar la distribución y el recambio de las hormigas epigeas a lo largo de un gradiente altitudinal

(1 000-2 800 m.s.n.m.) en las laderas opuestas de las cordilleras occidental y central de los Andes, en el cinturón

tropical de Colombia, Sudamérica.

Métodos: Se llevó a cabo un muestreo intensivo de hormigas en el campo y se utilizó la diversidad Zeta como

métrica analítica para evaluar los componentes de la diversidad espacial dentro de un marco de partición multi-

sitio, dada su sensibilidad a las diferencias de riqueza entre los tipos de hábitat.

Resultados: Se registraron un total de 204 especies de hormigas: 128 en la cordillera occidental y 131 en la cordi-

llera central. Myrmicinae fue la subfamilia más diversa, representada en todas las bandas altitudinales. La franja

de altitud media (1 500 m.s.n.m.) presentó la mayor diversidad en ambas cordilleras, mientras que la franja de 2

800 m.s.n.m. mostró la menor. La franja de 1 500 m en la Cordillera Central albergaba 71 especies, en contraste,

la franja de 2 800 m en la Cordillera Occidental sólo tenía dos especies. La composición de la comunidad de

hormigas en los flancos opuestos de las cordilleras Occidental y Central mostró una alta rotación, con solo tres

especies presentes en las cinco bandas altitudinales de ambas cordilleras. Las comunidades de hormigas estaban

dominadas por especies raras o con bajas frecuencias de captura, lo que probablemente refleja la interacción de

factores bióticos y abióticos específicos de cada zona de vida altitudinal.

Conclusiones: Estos hallazgos confirman que cada gradiente altitudinal alberga comunidades específicas de

hormigas que responden a condiciones ambientales, históricas y biogeográficas de cada cordillera, así mismo,

se confirma que los factores ambientales y los espaciales determinan la composición de hormigas y la tasa de

reemplazo de la comunidad.

Palabras clave: diversidad de hormigas; filtración ambiental; patrones de distribución; ecosistemas estratégicos;

bosque tropical.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

ecological dimensions, such as the composi-

tional turnover and the ecological mechanisms

driving these changes, remain essential areas of

inquiry, especially in the face of critical chal-

lenges like climate change (Ripple et al., 2024).

In ants, community richness is shaped

by factors such as area, temperature, humid-

ity, and soil nutrients, with temperature being

a key predictor of assemblage composition,

along with biotic interactions and historical

factors (Longino & Colwell, 2011). In complex

landscapes like the Andes, it becomes feasible

to evaluate species turnover across elevations

in different mountain ranges simultaneously,

providing insights into their response to cli-

matic variation. This is particularly important

given the ongoing threat of habitat loss due to

anthropogenic development, especially within

certain altitudinal belts. Combined with cli-

mate change, such pressures could result in

local, regional, or even global extinctions, espe-

cially among endemic species (Colwell et al.,

2008; Hethcoat et al., 2019; Longino & Colwell,

2011; Parra-Sanchez et al., 2025). Understand-

ing the patterns and drivers of species turnover

across elevational belts is, therefore, essential

to update conservation and restoration actions

(e.g., connectivity) in a megadiverse coun-

try like Colombia, which is also dramatically

affected by climatic global change.

Ants may exhibit diverse patterns of rich-

ness and composition along elevational gradi-

ents, shaped by specific biological adaptations

and ecological processes. Two main distribu-

tion patterns have been described: a mid-

elevation peak in species richness (e.g., Sanders,

2002), and a monotonic decline with increasing

altitude (Fisher, 1996; Stevens, 1992). The first

pattern may result from factors such as higher

productivity, larger available area, or the overlap

of species’ elevational ranges around the center

of the gradient, leading to great richness, even in

the absence of the productivity effects (Colwell

& Lees, 2000). The second pattern aligns with

Rapoport’s rule, which suggests that species

at higher elevations have broader elevational

ranges due to greater environmental variability,

although fewer species are able to tolerate such

conditions. In contrast, lower elevations, with

more stable climates, allow more species to

coexist within narrower ranges (Stevens, 1992).

Longino and Colwell (2011), suggest these pat-

terns result from a combination of historical,

evolutionary, and ecological factors, many of

which are influenced by paleoclimatic events

such as the current interglacial period, which

has shifted species ranges in upslope. Further-

more, lowland species often experience a lack

of competition at their lower range boundaries,

enabling unique assemblages that occupy broad

environmental niches. In contrast, highland

species inhabit more restricted climatic zones

and exhibit faster species replacement with

elevation (Gaston, 2000; Gaston & Spicer, 2001;

Janzen, 1967; Stevens, 1992).

Despite these general patterns, distribution

of ants along elevational gradients remains con-

troversial. While some studies show a mono-

tonic decline in species richness with elevation

(Fisher, 1996; Guerrero & Sarmiento, 2010;

Longino & Colwell, 2011; Narváez-Vidal et

al., 2024), others report richness peaks at mid-

elevations, a pattern considered common in

nature (Longino & Branstetter, 2019; Sanders,

2002). Even the interpretation of Rapoport’s

rule is debated: Rahbek (1995) argues that it

describes a monotonic decline in diversity with

elevation, whereas Sanders (2002) and Pérez-

Escobar et al. (2022) contend it may also help

explain mid-elevation richness peaks.

Given the ongoing controversy and the

contrasting patterns of species diversity, exam-

ining compositional combinations along altitu-

dinal gradients can be useful for understanding

biodiversity responses to ecological changes

and the availability of biodiversity resources.

Parra-Sánchez et al. (2025), suggest the impor-

tance and urgency of understanding how com-

munity assembly occurs in natural habitats and

how it is restructured under the influence of

both natural and anthropogenic drivers across

different spatial scales (local, landscape, and

regional). At the landscape scale, anthropic

transformation and subsequent land-use

changes modify the structural and functional

connectivity of natural habitats through habitat

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loss and fragmentation (Fahrig, 2003; Parra-

Sanchez et al., 2025). The reduction in con-

nectivity interrupts recolonization processes,

hence favoring the replacement of species with

restricted distribution ranges by those with

wider distribution ranges (Fahrig, 2002; Fahrig,

2003; Fahrig, 2017).

Building on the previous discussion, our

study employs zeta diversity as an analytical

framework to quantify ant species distribution

and turnover across multiple sites and spatial

scales (McGeoch et al., 2019), incorporating

the influence of natural drivers such as altitude

and temperature. Zeta diversity allows a more

detailed understanding of biodiversity change

(Hui & McGeoch, 2014) by generating a series

of values (ζ1, ζ2, ζ3, ..., ζN), where each value

represents the number of species shared among

different combination of sites, encompassing

rare, intermediate, and common species that

drive compositional turnover (Hui et al., 2018;

Latombe et al., 2018; Parra-Sanchez et al., 2025).

Unlike pairwise beta diversity, equivalent

to zeta diversity of order two (Magurran &

Henderson, 2010; Shimadzu et al., 2015), which

tends to be biased toward the contribution of

rare species and may overestimate their influ-

ence (Jost, 2007), zeta diversity allows multi-

site assessments of assemblage structure, less

biased by rare species and providing insights

into the rate and nature of compositional turn-

over. As zeta order increases, rare species are

progressively excluded, enabling the detection

of the differential contribution of the entire

spectrum of species to compositional turnover

across a range of orders (McGeoch et al., 2019;

Parra-Sanchez et al., 2025).

In this context, the present study aims

to understand how epigeous ant communi-

ties vary in species richness, composition and

species turnover along an elevational gradient

(1 000-2 800 masl) by examining two parallel

mountain ranges adjacent to the Cauca River

Valley in Colombia. Specifically, we assessed

how environmental variation, particularly ele-

vation and temperature, influences these pat-

terns, and whether species with specialized

habits and narrow environmental tolerances

could be more vulnerable to climate change.

MATERIALS AND METHODS

Study area: The study area is in South-

western Colombia, in the central region of the

Valle del Cauca department. This inter-Andean

valley separates two neighboring mountain

ranges (hereafter referred to as “Cordilleras”)

involving an altitudinal gradient ranging from 1

000 masl (the elevation of the valley floor where

the Cauca River flows South to North), up to 2

800 masl (the highest elevation involved in this

study). The valley is flanked by the opposing

slopes of the Central and Western cordilleras

(Cordilleras “Central” and “Occidental” are the

Spanish proper names of these two mountain

ranges) (Fig. 1). The study area spans four

ecosystem types (Table 1), from tropical dry

forest (Bs-T) at the lowest elevations to lower

montane very humid forest (Bmh-MB) at the

highest based on the life zone classification sys-

tem of Holdridge as adapted for Colombia by

Espinal (1968). Using the Cauca River valley as

a central reference, five altitudinal bands were

selected along “mirror” slopes on each Cordil-

lera, ranging from 1 000 to 2 800 masl. Con-

sequently, five sampling sites were established

on each slope at approximately 500 m intervals

(Fig. 1). During sampling, temperature and

relative humidity were recorded hourly for 15

consecutive days using data loggers (Elitech

model RC-51H, Table 1). All reported data cor-

responds to ants collected in forested habitats

or natural, unmanaged ecosystems specific to

each elevation.

Epigeous ant sampling: Ants are highly

conspicuous organisms in soil ecosystems, par-

ticipating in a wide variety of ecological inter-

actions (Del Toro et al., 2012; Donoso, 2017).

Their role is crucial in key processes such as

herbivory, predation, scavenging, the establish-

ment of mutualistic relationships with other

organisms and recirculation of nutrients in the

soil (Urrego-Sánchez & Camero, 2020; Schultz

& McGlynn, 2000). Because of their significant

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

Fig. 1. Location of the study area: Colombia and Valle del Cauca Department (upper left corner), sampling sites along five

elevational bands on the opposite slopes of the Western and Central Andes and the geographical valley of the Cauca River.

The cordilleras profiles are shown at the lower left corner of the figure. Note. The map was created by the author based on

a digital elevation model (DEM) obtained from Colombia en mapas (Instituto Geográfico Agustín Codazzi [IGAC], 2024).

Table 1

Environmental characterization of sampling localities.

Localities Code Cordillera Coordinates Altitude

(m asl) Landscape Life zone T °C

Mean

% HR

Mean

Latitude Longitude

Fca. Monteloro 5W Western 4.2297 -76.4364 2 800 Mountain Bmh-MB 12.97 89.5

RNSC Madhú 5C Central 3.6429 -76.1436 2 800 Mountain Bmh-MB 14.92 97.3

Fca. Gratinianos 4W Western 4.1716 -76.4237 2 400 Mountain Bmh-T 16.27 89.69

RNSC El Pailón 4C Central 3.7052 -76.0647 2 400 Mountain Bmh-T 15.23 85.64

RNSC El Silencio 3W Western 4.1704 -76.4064 2 000 Mountain Bmh-T 18.29 88.43

RNSC El Triunfo 3C Central 3.4632 -76.1479 2 000 Mountain Bmh-T 17.45 88.03

RNF Bosque de Yotoco 2W Western 3.8776 -76.4377 1 500 Mountain Bh-T 21.91 89.29

Fca. El Canto ícaro 2C Central 3.5037 -76.1912 1 500 Mountain Bh-T 21.05 84.9

Rest. El Mirador 1W Western 3.8782 -76.4143 1 000 Geographic valley Bs-T 22.15 82.65

PNR. El Vínculo 1C Central 3.8343 -76.2954 1 000 Geographic valley Bs-T 25.11 73.82

All acronyms come from Spanish according to Espinal (1968): Bmh-MB = Very humid low montane forest, Bmh-T = Very

humid tropical forest, Bh-T = Humid tropical forest, Bs-T = Dry tropical forest.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

biomass, susceptibility to environmental stress,

and crucial role in terrestrial ecosystems, epi-

geous ants are recognized by many researchers

as a vital indicator group for assessing environ-

mental changes (Andersen et al., 2002; Donoso,

2017; Fowler & Delabie, 1995; Urrego-Sánchez

& Camero, 2020). Hence, we focused on this

stratum thus pursuing the objectives of this

study. At each elevation, three plots (10 m²)

were randomly established, with a minimum

distance of 50 m between them. Within each

plot, five pitfall traps were installed, each filled

with a solution of 5 % acetic acid, one pound

of salt per 20 l of solution, and 5 gr liquid soap.

This solution was prepared once and used

throughout the sampling period. Traps were

operated for 15 days, buried at ground level,

and camouflaged with surrounding vegetation.

Sampling was complemented by manual col-

lection (1 hour/person per plot), targeting all

accessible substrates, including soil, leaf litter,

understory vegetation, decomposing trunks,

and hollow or dry twigs. Additionally, leaf lit-

ter was collected from 1 m2 at four randomly

selected points within each plot and processed

by visual screening.

Each altitudinal band comprised 42 sam-

ples, totaling 420 samples across both gradients.

All individuals were counted and identified to

the highest possible taxonomic resolution using

identification keys from Fernández-Castiblan-

co et al. (2019) and Feitosa and Dias (2024),

as well as direct comparison with reference

collection specimens from the Entomological

Museum of the Universidad del Valle (MEUV),

for its acronym in Spanish). Taxonomic ver-

ification was conducted with support from

expert myrmecologists when needed. Species

were functionally classified as forest specialists,

generalists, or open-habitat specialists, follow-

ing the Laboratório de Ecologia de Formigas

(UFAC, 2023) application (SMT1). Forest spe-

cialists are defined in this study as species that

depend exclusively on well-preserved forest

habitats, while generalists are capable of inhab-

iting both forested and altered environments,

and finally, open-habitat specialists are restrict-

ed to more open or disturbed vegetation types.

All specimens were preserved following

standard entomological protocols and depos-

ited in the MEUV collection under collecting

permit No. 120, issued by Colombian Ministry

of Environment and Sustainable Development

(August 24, 2015). An incidence matrix (pres-

ence-absence) of samples was elaborated for

each elevational band, incorporating environ-

mental variables associated with each study plot.

Statistical analyses: All statistical analyses

were conducted exclusively on worker ants.

Rescaled zeta diversity was used to calculate

diversity components at spatial scales under the

same multi-site diversity partitioning frame-

work (Hui et al., 2018; McGeoch et al., 2019),

as this metric is more sensitive to richness dif-

ferences inherent to each habitat types. There-

fore, comparisons focused on zeta diversity

differences between mountain ranges and for-

est zones along the altitudinal gradient (Hui

& McGeoch, 2014; McGeoch et al., 2019). To

complement the analyses, the effective number

of species was calculated based on Hill numbers

(Cultid-Medina & Escobar, 2019; Jost et al.,

2010; Moreno, 2001), and Pearson correlation

was used to verify whether diversity decreased

or increased with altitude. Species composition

and distribution patterns across the altitudinal

gradient were explored through non-metric

multidimensional scaling (NMDS) to detect

structures or clusters in the data, followed by a

PERMANOVA test with 999 permutations.

To account for spatial structure, we gen-

erated a matrix of spatial variables based on

geographic distances using the Principal Coor-

dinates of Neighbor Matrices (PCNM) method

(Borcard & Legendre, 2002), incorporating

spatial distances among altitudinal bands and

between the flanks of the mountain ranges.

This method extracts positive eigenvectors

from a Euclidean distance matrix of sampling

locations, based on multiple coefficients of

determination (R²). These eigenvectors were

then used as covariates in a partial Redundan-

cy Analysis (pRDA) following Blanchet et al.

(2008). The pRDA was applied to a Hellinger-

transformed community matrix (species per

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

site), wich reduces the influence of double

absences (Legendre, 2008). In this analysis,

the matrix of environmental variables (Alti-

tude, Leaf litter depth, PAP30, T mean, Tmax,

Tmin, HR mean, HRmax, HRmin), previously

centered and standardized using Z-scores, was

used as the explanatory matrix. The signifi-

cance of the pRDA was tested by ANOVA, and

variable selection was performed using the

forward selection method with a cut-off value

of alpha = 0.05 and based on the R2 coefficient.

To assess whether the ant community

follows a pattern of monotonic decline with

altitude, we applied the approach proposed

by Sanders (2002) and Longino and Colwell

(2011). This involved estimating the altitudinal

range for each species as the difference between

its maximum and minimum recorded eleva-

tions, assuming continuous presence across

that interval. All Statistical analysis were con-

ducted in R version 4.3.3. (R Core Team, 2023),

using the packages zetadiv (Latombe et al.,

2018), vegan (Oksanen et al., 2022), and iNEXT

(Hsieh et al., 2022).

RESULTS

A total of 32 321 individuals represent-

ing 204 species/morphospecies were recorded,

from which 45 were identified as morpho-

species given the extreme difficulty to reach

the specific identity for being highly complex

taxonomic genera groups. The genera with the

highest number of morphospecies were Phei-

dole and Camponotus, which are megadiverse

genera. Identification, mainly in Pheidole, is

based on the soldier caste, and it is common

to collect workers in the field (Fernández-

Castiblanco et al., 2019). From the described

total species, more than half were present in

each of the mountain ranges, this is 128 were

recorded in the Western Cordillera, while 131

in the Central Cordillera. Among all these, 51

species/morphospecies were shared between

both ranges, and 80 and 77 were exclusive to

the Central and Western cordilleras, respec-

tively. Of all the species reported, Linepithema

neotropicum (Wild, 2007) and Linepithema

piliferum (Mayr, 1870) were found across all

five altitudinal bands in both ranges. However,

Pheidole sp. 9 is expected to occur along the full

altitudinal gradient of the Central cordillera, as

it was present in all bands except those at 2 000

and 2 400 masl.

Both elevation gradients exhibited trun-

cated bell-shaped richness curves (SMF1), with

peaks above 1 500 masl (with 59 and 71 spe-

cies in the Western and Central Cordilleras,

respectively), followed by the marked decrease

in the valley zone at 1 000 masl (with 50 and

61 species in the Western and Central Cordil-

leras, respectively). At this mid-elevation belt,

the most diverse subfamilies were, in order of

richness: Myrmicinae, Ponerinae and Formici-

nae (consistent with patterns observed in other

studies). In contrast, the lowest species richness

occurred at 2 800 masl in both mountain flanks,

with only two and six species recorded in the

Western and Central Cordilleras, respectively.

At this elevation, only five of the nine subfami-

lies reported in this study were present, with

Dolichoderinae and Dorylinae being the most

species-rich (five species each), and Formicinae

the least diverse.

The subfamily Myrmicinae was the most

diverse among both altitudinal gradients. It

was present at all elevations and showed a great

representation at 1 000 and 1 500 masl (lower

elevations). In contrast, Agroecomyrmecinae

and Amblyoponinae were the least represented,

each with only one species, restricted to low

elevations. Changes in the composition along

the elevation gradient were mainly driven by

species turnover within genera. The sampled

ant species belonged to 59 genera, of which four

(Camponotus, Linepithema, Pheidole, Neopone-

ra) included species occurring both in lowlands

and highlands sites. The most specious genera

were Pheidole (40 spp.), Camponotus (18 spp.),

Pseudomyrmex (11 spp.), Linepithema (11 spp.),

Strumigenys (11 spp.), Crematogaster (9 spp.),

Solenopsis (9 spp.), Cyphomyrmex (8 spp.) and

Hypoponera (8 spp.). Conversely, only three

genera, Neivamyrmex, Parathrachymyrmex and

Rophalotrix, were found exclusively above 2 400

masl each with low diversity and abundance. At

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

elevations between 1 000-1 500 masl, only 31

genera were recorded.

A total of 29 % of the species were forest

specialists, 20 % generalists, and 2 % open-

habitat specialists (SMT1). The remaining 49 %

could be assigned to any category due to a lack

available information, either because they are

not catalogued or because of their taxonom-

ic resolution at the morphospecies or family

level. Similarly, SMF2 shows that forest special-

ists and generalists exhibited greater diversity

at lower elevations, with richness decreasing

above 2 400 masl.

Altitudinal variation in species rich-

ness: In both mountain ranges, ant diver-

sity decreased with increasing elevation. The

decline was steeper in the Western Cordillera

than in the Central Cordillera, especially, for

hill numbers at q = 0 (Fig. 2). Concerning order

numbers q = 1, q = 2, which give more weight

to the common and most abundant species, the

slopes were less pronounced compared to q =

0 (Fig. 2).

Across both mountain ranges, the great-

est number of shared ant species was found

between the 1 000-1 500 masl bands, 17 for the

Central and 11 for the Western range (Fig. 3). In

contrast, the highest elevations (2 400 and 2 800

masl) showed no shared species in the Western

Cordillera, and only one species, Labidus spin-

inodis (Emery, 1890) was shared in the Central

Cordillera. These results highlight strong com-

positional differentiation at high elevations, a

pattern corroborated by the NMDS analyses

(stress = 0.1159, Fig. 4) and pRDA (R2 = 26.42,

p-value = 0.001, Fig. 5).

At mid-elevation (2 000 masl), the West-

ern Cordillera showed higher similarity in the

composition between 2 000 and 2 400 masl

(13 shared species), while similarity declined

sharply below 2 000 m, with only six species

shared between 1 500 and 2 000 masl: Crema-

togaster nigropilosa (Mayr, 1870), Hypoponera

aff. punctatissima (Roger, 1859), Myrmelachista

zeledoni (Emery, 1896), Pheidole biconstricta

(Mayr, 1870), Labidus coecus (Latreille, 1802)

and Linepithema sp. In the Central Cordillera no

Fig. 2. Pearson’s correlation tests between diversity and altitude for the different Hill numbers, performed separately for both

mountain ranges: Western (top row) and Central (bottom row). In each case, response plots corresponding to values of q =

0, q = 1 and q = 2 are presented, arranged from left to right in that order.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

Fig. 3. Venn’s diagram shows the similarity of the species community between the altitudinal gradient. A. The Eastern flank

of the Central Cordillera. B. The Western flank of the Western Cordillera.

Fig. 4. Non-metric multidimensional scaling for the ant community of the Eastern flanks of the Western Cordillera and

Western flanks of the Central Cordillera.

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species are shared between 2 000-2 400 m, and

only four species, C. nigropilosa, H. aff. puncta-

tissima, M. zeledoni, P. biconstricta, were found

between elevations 1 500-2 000 masl. Notably,

species composition was more similar between

cordilleras at these mid-elevations (Fig. 3).

Species at lower and mid elevations

(1 000-2 000 masl) tended to have broader

altitudinal ranges, while those at high ele-

vations (2 000-2 800 masl), exhibited

narrower distributions.

Differentiation and structure of the com-

munity: The NMDS ordination model for both

mountain ranges yielded a stress of 0.1159, indi-

cating congruence in the associations of the ant

assemblages for the sampled flanks. The poly-

gons delimit the composition related to each

mountain range, thus the interaction between

the mountain ranges largely explains the varia-

tion in the composition of the species (PER-

MANOVA: F = 1.92, R2 = 0.0665, p < 0.011),

and the differences between ant communities

between the flanks of the mountain ranges (Fig.

4). However, the proportion of shared species is

perhaps equal to or like that of dissimilar spe-

cies between both flanks evaluated.

Environmental and spatial factors affect-

ing community composition: The permutation

test (F = 2.70, p = 0.001) confirmed a significant

association between ant composition on the

Western flanks of the Central Cordillera and

the Eastern flank of the Western Cordillera (R2

= 0.2642, p < 0.05, Fig. 5). The pRDA model

explained 60.05 % of the variation in ant inci-

dence between mountain ranges, with 24.08 %

of the variance explained by covariates (spatial

Fig. 5. pRDA analysis for the ant communities of two altitudinal gradients in the central and Western mountain ranges

and the Cauca River geographic valley. Altitude is presented in meters above sea level. HRmax: Maximum values of relative

humidity. HRmin: Minimum values of relative humidity. Tmax: Maximum temperature values in degrees Celsius. Tmin:

Minimum temperature values in degrees Celsius. PAP30: Perimeter at chest height greater than 30.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

distances between sampling sites) and an addi-

tional 35.97 % corresponding to environmen-

tal variables (Altitude, PAP30, Tmax, Tmin,

HRmax, HRmin). The Redundancy Analysis

(RDA1 and RDA2 axes), accounted for 72.46 %

of the variance explained (41.45 and 31.01 %,

respectively). This indicates that the environ-

mental gradients represented by these two axes

are the main drivers of community structuring.

The unexplained proportion of the model was

39.96 %, which is attributed to noise and/or

unmeasured factors, such as biotic interactions

and site history.

Zeta diversity pattern: In Fig. 6A, the zeta

diversity decay curve illustrates high species

turnover, as well as compositional heteroge-

neity along the elevation gradient. The sharp

initial decrease indicates a high renewal and

low species overlap between elevation bands,

meaning that communities change rapidly as

altitude increases. This decline reached zeta

diversity of order 8 for both cordilleras (AIC =

5.699 for the Western and AIC = 9.558 for the

Central). The Fig. 6B shows the species reten-

tion rate. In the Central Cordillera, the species

retention increased to order 4-5, while in the

Fig. 6. Zeta diversity analysis for the ant community of the altitudinal gradient of two opposite flanks of the Western and

central mountain range and the geographic valley of the Cauca River. A. Shows the decline in zeta diversity as a function

of zeta order, showing the progressive decrease in the average number of shared species as the number of sites considered

increases. B. Species retention rate (zeta ratio) across zeta orders, indicating the probability that species will remain shared

as the number of sites increases. C. Exponential fit of zeta diversity (log scale) against zeta order, used to evaluate whether

species turnover is dominated by stochastic processes or a constant probability of local extinction. D. Power-law fits zeta

diversity (log scale) against zeta order, indicating the influence of deterministic processes, environmental structuring, or

spatial constraints on species composition.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

Western Cordillera it peaked at order two and

decreased sharply by order five. This suggests

that, above order five, the probability of species

being retained across multiple sites decreases

significantly, indicating an almost complete

species renewal along the gradient. Since this

strong compositional turnover can be partly

explained by the low prevalence of many spe-

cies in the samples, we used linear regression.

Therefore, to evaluate whether this pattern fol-

lowed a power law or an exponential decay, lin-

ear regression was used (Fig. 6C, Fig. 6D). The

turnover fits a power law distribution, in turn

showing a rapid exponential decline suggesting

a non-linear turnover following a non-scale

pattern of few shared species, which suggests

that niche differentiation, rather than stochas-

ticity, may underline the observed diversity

patterns, a finding that was also observed in the

NMDS results.

To compare different diversity metrics and

assess how the species similarity changes with

elevation, we evaluated ant richness across

opposite flanks of the Cordilleras (SMF1). Both

beta diversity (Jaccard) and normalized zeta

diversity (equivalent to Jaccard n-site similar-

ity) were considered (Fig. 7) Jaccard similarity

showed steeper increase with elevation, indi-

cating overestimation of shared species when

only pairwise comparisons are considered. In

contrast, normalized zeta diversity revealed

a more gradual decline, suggesting that spe-

cies heterogeneity across multiple elevation

bands is lower than expected. This is because

zeta diversity inherently includes all sites situ-

ated between the fixed point and a specified

elevation band in its calculation, consequently

leading to a lower count of shared species com-

pared to direct comparisons between the fixed

point and the specified band, as in Jaccard.

This implies more complex patterns of species

turnover, where different species dominate at

different elevations, a pattern that corresponds

with an increase in rare species at higher eleva-

tions and between mountain ranges.

DISCUSSION

Altitudinal variation in species richness:

In Colombia, ants inhabit nearly all ecosystems,

Fig. 7. Decline of ant diversity of the altitudinal gradient of two opposite flanks of the Western and Central cordillera and the

geographic valley of the Cauca River, using two similarity metrics: normalized zeta and Jaccard similarity.

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ranging from sea level to elevations above 3 000

masl. Although this study did not include sites

above 3 000 masl, Narváez-Vidal et al. (2024)

reported three ant species at elevations up to

3 200 masl in the Farallones de Cali National

Natural Park (Western Cordillera), but none

at 3 800 masl. This pattern was confirmed in

our study, where no ants were found in some

sampling units at 2 800 masl. Although our

study covered a wide altitudinal gradient, it

did not include sites below 1 000 masl, as such

elevations are absent from the inter-Andean

valley of the upper Cauca River basin, and it

was not possible to sample lower elevations for

topographic reasons.

There is consistent evidence supporting

a pattern of declining species richness with

increasing altitude in tropical mountains. In

this context, our findings align with those of

Narváez-Vidal et al. (2024) and other studies

(Guerrero & Sarmiento, 2010; Longino & Col-

well, 2011; Rodríguez & Lattke, 2012), which

report a decrease in richness at higher eleva-

tions. In the Western Cordillera, this decline

was more pronounced than in the Central

Cordillera (q = 0, Fig. 2). The declining pattern

likely reflects a combination of ecological traits,

evolutionary history, and biogeographic pro-

cesses that shape ant communities because life

zones sharply change in tropical regions.

For abundant species (q = 1), the most

frequent species in the Western Cordillera fol-

lowed the altitudinal gradient, whereas in the

Central Cordillera, no clear pattern of decline

with altitude was observed (Fig. 2). This behav-

ior may be explained by the hypothesis pro-

posed by Longino and Colwell (2011), who

argue that temperature is a key factor in pre-

dicting ant presence, linking local patterns to

broader global biodiversity trends.

Differentiation and structure of the com-

munity: Anthropogenic disturbance is believed

to be a key factor influencing ant composition

along altitudinal gradients, although it was not

directly measured in this study. While our sam-

pling focused on forest remnants within each

altitudinal belt, none of the sites represented

pristine conditions. The sampled forests were

structurally intact but disturbed, an expected

scenario in a region with a long history of

human settlement.

As a result of prolonged anthropogenic

transformation, this study reports the wide-

spread presence of two generalist species

(L. neotropicum and L. piliferum), recorded

throughout the entire gradient in both moun-

tain ranges. Their ecological plasticity enables

them to persist in both natural and human-

modified habitats. Escárraga & Guerrero (2016)

also documented a broad altitudinal distribu-

tion for these species: from 0 to 2 600 masl

for L. neotropicum, and up to 2 800 masl

for L. piliferum.

Parra-Sánchez et al. (2025) found that

anthropic tensors shape the composition of

bird, beetle, and orchid communities, effective-

ly nullifying the influence of altitude on their

composition. In contrast, our study found that,

while the epigeous ant community responded

to altitudinal variation, geographical and envi-

ronmental factors, habitat transformation also

played a role in shaping community composi-

tion and structure.

Consequently, the number of habitat-based

groups decreased with altitude. This pattern

may be related to the fact that seven of the ten

sites evaluated were reserves dedicated exclu-

sively to conservation, each under different

protection categories (Table 1). These catego-

ries appeared to influence forest structure: pro-

tected reserves differed from forests located on

private farms, some of which were civil society

reserves maintained by landowners. However,

this distinction was not considered in the diver-

sity analyses, as all sampled habitats were classi-

fied as forested and structurally mature.

The high dominance of rare species (low-

frequency) and the low similarity of shared

species along altitudinal gradients suggest

that each elevation band harbours distinct ant

assemblages (Fig. 3, Fig. 4). These differences

reflect a combination of ecological, landscape,

historical, and environmental factors that vary

both with altitude and between mountain

ranges. NMDS ordination (stress 0.1159) and

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

PERMANOVA analyses (F = 1.92, R2 = 0.0665,

p < 0.011) confirm this pattern, revealing a

significant separation in species composition

between the Western and Central Cordilleras.

From an ecological theory perspective, this

pattern can be interpreted through the lens of

island biogeography (MacArthur & Wilson,

1967), wherein each mountain range functions

as an isolated unit that fosters rare species

assemblages due to limited dispersal and envi-

ronmental heterogeneity, a pattern observed in

pRDA (Fig. 5) and the Zeta diversity analysis

(Fig. 6). Similarly, the theory of environmental

filters helps explain how altitudinal gradients

and local conditions selectively shape commu-

nity composition (Kraft et al., 2015; Longino &

Colwell, 2011; Rahbek, 1995).

The observed overlap is likely attributable

to shared species with broad ecological niches,

eurytopic, or neutral responses to environ-

mental variation, while the distinct groupings

reflect habitat selection processes and the influ-

ence of geographical isolation. This suggests

that altitudinal gradients in each mountain

range are not equivalent, and that regional

diversity is maintained through their ecological

complementarity. Therefore, the generalized

prediction of decreasing ant diversity pattern

with altitude in tropical mountains is reinforced

in this study. A similar pattern was reported by

Narváez-Vidal et al. (2024) along an altitudinal

gradient in the Western Cordillera, underscor-

ing the importance of conservation strategies

that account for metacommunity structure and

the biogeographic uniqueness of the Andes

(Baselga, 2010; Leibold et al., 2004).

Environmental and spatial factors affect-

ing community composition: Using pRDA

with appropriate environmental and spatial

covariate showed that decline in species diver-

sity and the high turnover of ant communi-

ties observed along both altitudinal gradients

can be attributed to temperature and spatial

distance, as illustrated in Fig. 7 and supported

by Longino & Colwell (2011). These factors

are key drivers of ant community composi-

tion. Temperature variation is closely linked to

altitude, which in turn shapes life zones, vegeta-

tion structure, and habitat heterogeneity along

the gradient.

In tropical regions, such as Colombia,

temperature decreases at an average rate of

approximately 0.65 °C per 100 m increase in

altitude, with greater thermal variability at

higher altitudes. However, unlike temperate

zones, tropical mountains experience little sea-

sonal variation in temperature, which removes

an important ecological filter affecting organ-

isms elsewhere (Colwell et al., 2008; Janzen,

1967; Longino & Branstetter, 2019).

In addition, lowland species are usually

stenothermic, adapted to more restrictive envi-

ronmental conditions and limited by their tol-

erance to cold. For instance, the pRDA test for

Atta cephalotes¸ indicated that this species was

associated with conditions of lower altitude and

higher temperature. This physiological limita-

tion makes lowland species more vulnerable to

climate change (Longino & Branstetter, 2019;

Sanders, 2002; Stevens, 1992). Conversely,

higher altitude species are usually eurythermic

and can tolerate warmer temperatures than

they usually experience. This is the case of L.

spininodis and Neoponera carbonaria, both with

a high positive score on axis 1 and negative on

axis 2. These two species were associated with

higher elevation and lower temperature units.

Although this issue is still under debate and it

is accepted that temperature is strongly corre-

lated with ant distribution, some studies argue

that low temperatures (such as those found in

paramo ecosystems) are the ones who most

determines the pattern since this condition

reduce ant metabolic activity, larval develop-

ment, and their ability to colonize or persist in

unsuitable environments (Kunene et al., 2022).

From a macroecological perspective, these pat-

terns are explained as the result of temperature

and precipitation variation along the altitudinal

gradient (Kaspari & Majer, 2000).

Our results suggest that ant communities

and/or composition are influenced by pre-

dictor variables such as geographic distances,

which in turn reflect a multi-scale process of

species turnover. For both mountain ranges,

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

environmental factors (altitude, for Western

and temperature and humidity, for Central)

were determinant in driving turnover along

each gradient, because the community struc-

ture was adjusted according to the ecological

tolerances of each species and each community

was structured according to environmental fil-

ters specific to each mountain range, as well as

dispersal limitations (Kraft et al., 2015). Our

grouping analyses coincided in supporting that

each mountain range functions as a unit semi-

isolated by a geographic barrier, such as the

inter-Andean valley; however, the composition

of myrmecofauna and the turnover of species

within (and between) each mountain range

give structure to the regional diversity (gamma

diversity). On the other hand, although not

measured here, factors such as climatic changes

during the current interglacial period, could

also be influenced by potentially causing alti-

tudinal changes in species distribution through

interspecific competition at ecological bound-

aries (Longino & Colwell, 2011).

Zeta diversity pattern: The steep drop

shows that few species are shared, suggesting

a high turnover in species (Parra-Sanchez et

al., 2025). This is consistent with niche theory,

where species respond to environmental filter-

ing by deterministic mechanisms (Kraft et al.,

2015; Leibold et al., 2004). Geographic dis-

tances also influence community composition,

especially when assessing turnover between

locations. This was evident in our zeta diver-

sity analyses: as comparisons between multiple

locations increased, species overlap decreased

and rare species became more dominant (Fig.

6) (Hui & McGeoch, 2014; McGeoch et al.,

2019; Parra-Sanchez et al., 2025).

Regarding species retention, it was observed

that the Central Cordillera harbors more gener-

alist or widely distributed species, resulting in

a more stable community with greater species

fidelity across all elevations. In contrast, the

Western Cordillera exhibits a marked decline in

species retention. This difference between the

communities of both gradients reinforces the

findings from other analyses, which indicate

that community composition is shaped by envi-

ronmental filtering specific to each mountain

range (Hui & McGeoch, 2014; Kraft et al., 2015;

Leibold et al., 2004; Longino & Colwell, 2011;

McGeoch et al., 2019).

The power-law fit reflects that commu-

nity structuring in the Central Cordillera obeys

more deterministic niche-based assemblage

processes, possibly reflecting stronger environ-

mental filtering, habitat continuity, or historical

stability. In contrast, the exponential fit in the

Western Cordillera is consistent with a more

stochastic turnover, which could result from

greater fragmentation, reduced dispersal, or

more irregular microhabitats.

Thus, Zeta diversity analyses revealed that

as the number of sites compared in this study

increases, the ant communities in both gradi-

ents are strongly influenced by rare or habitat-

restricted species. This finding supports and

reinforces the patterns observed in other analy-

ses regarding species turnover. The zeta diversi-

ty results show marked species turnover across

the two mountain ranges, albeit with contrast-

ing patterns. In the Western Cordillera, the

steep decline in zeta diversity and low species

retention suggest highly differentiated com-

munities dominated by species with restricted

distributions and low frequencies, indicating a

more stochastic assemblage shaped by limited

dispersal and high environmental heteroge-

neity. In contrast, in the Central Cordillera,

the greater persistence of species across zeta

orders and the better fit to a power-law model

reflect a more deterministic pattern, where

environmental filtering allows the presence

of generalist species with broad distributions.

These differences align with metacommunity

theory (Leibold et al., 2004), in which the

Western range approximates a “dispersal-lim-

ited” model, while the Central range exhibits

characteristics of species with broad ecological

niches that tolerate environmental variation

and are distributed across multiple sites along

the gradient.

In conclusion, our findings confirm that

species turnover along each gradient is influ-

enced by environmental, spatial, biogeographic,

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026278, enero-diciembre 2026 (Publicado May. 15, 2026)

historical, and demographic factors specific to

each mountain range. These factors shape the

composition and structure of ant communi-

ties, which may be dominated by rare, unique,

or limited-dispersal species. Such species tend

to be more sensitive to environmental varia-

tion, which, in turn, leads to the high species

replacement observed in this study. Alterna-

tively, communities could have been dominated

by common, generalist, and widely distributed

species, resulting in greater stability and pro-

moting biotic homogenization, something that

did not occur in this study.

Therefore, we emphasize the need for

conservation and restoration strategies in all

the altitudinal floors of tropical mountains, in

order to enhance landscape connectivity along

altitudinal gradients. Although not directly

measured in this study, fieldwork observations

suggest that species within elevation bands may

experience population declines due to accel-

erated habitat loss and fragmentation. These

findings highlight the urgency of applying the

precautionary principle to prevent local and

regional extinctions. Protecting Andean natural

forests and restoring connectivity across eleva-

tions would facilitate rescue effects and help

maintain metapopulation dynamics in these

highly fragmented landscapes.

In this context, adopting a preventive and

proactive approach to preserve Andean eco-

systems is imperative. Protecting natural for-

ests and restoring connectivity between them

are essential actions to mitigate the effects of

fragmentation and ensure the long-term persis-

tence of metapopulations. Preserving forests in

all altitudinal gradients and promoting ecologi-

cal connectivity will be crucial for safeguarding

ant diversity and, by extension, the integrity of

Andean ecosystems in the face of accelerating

climate change and increasing human pressure.

These measures would not only protect endem-

ic and vulnerable species but also enhance eco-

system resilience to future environmental and

anthropogenic disturbances.

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.

ACKNOWLEDGMENTS

The authors thank Minciencias and Uni-

valle for funding the program “Multiscale rela-

tionships of biodiversity in altitudinal gradients

of the tropical forest”, Code: 1106-852-70306,

Contract No. 491-2020. We also express special

thanks to the communities and administrators

of the different properties that were sampled for

their hospitality during the field days. We also

thank the biology students from Universidad

del Valle and Universidad del Quindío, as well

as biologists Nataly Forero, Diana Urcuqui,

Sebastián Forero, Javier Cuervo, Nicolás Mon-

tero and Anderson Arenas for their valuable

collaboration in the sampling. Finally, we

extend our thanks to the expert myrmecologists

Roberto José Guerrero, Emira García, Maria

Alejandra Bautista, Sabrina Amador, Adrián

Troya and Francisco Serna for their support

and corroboration in the taxonomic identifica-

tion of the collected ants.

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