# Forest-to-grassland conversion reduces ground beetles (Coleoptera: Carabidae) diversity in tropical Andean montane habitats

Anderson Arenas-Clavijo, James Montoya-Lerma, Pierre Moret

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Anderson Arenas-Clavijo, James Montoya-Lerma, Pierre Moret. Forest-to-grassland conversion reduces ground beetles (Coleoptera: Carabidae) diversity in tropical Andean montane habitats. Journal of Insect Conservation, 2026, 30 (1), pp.18. 10.1007/s10841-025-00739-8. hal-05510327

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# Forest-to-grassland conversion reduces ground beetles (Coleoptera: Carabidae) diversity in tropical Andean montane habitats

Anderson Arenas-Clavijo<fnref n="1" /> ID · James Montoya-Lerma<fnref n="2" /> ID · Pierre Moret<fnref n="3" /> ID

Received: 23 March 2025 / Accepted: 28 November 2025

© The Author(s) 2025

**Abstract** The deforestation of natural habitats due to agricultural expansion and human settlements is a major driver of global biodiversity loss. This study assessed the impact of natural habitat conversion into grasslands on ground beetle (Coleoptera: Carabidae) assemblages along two elevational gradients (~1900–3000 m asl) in the southwestern tropical Andes of Colombia. Sampling was performed using pitfall traps, litter sifting, and nocturnal hand collection at six localities, each comprising a natural forest area and a deforested grassland area. Environmental variables were recorded to characterize land use. We found that these conditions differed markedly between forests and grasslands, with forests exhibiting more stable temperature and humidity, and greater leaf litter depth. Ground beetle richness and abundance were consistently higher in forests, while grasslands were dominated by a few generalist species. Assemblages in forests showed high species turnover, including a large proportion of flightless, microendemic species, whereas grassland communities were homogenized across cordilleras and elevations.

**Implications for insect conservation** Our study highlights the detrimental effects of deforestation on carabid diversity and assemblage structure. The observed loss of diversity implies a high risk of extinction for flightless, endemic forest specialists restricted to small areas. These findings emphasize the importance of preserving natural forest areas at different elevations and on different slopes, highlighting the need for taxonomic studies to inform targeted conservation strategies in tropical montane ecosystems.

**Keywords** Diversity assessment · Tropical forests · Grassland · Land use · Mountain · Endemism

## Introduction

It is well established that deforestation caused by the expansion of agricultural frontiers and/or the establishment of human settlements leads to the loss of biodiversity in terrestrial habitats (Ponce-Reyes et al. 2013; Franco et al. 2019; Sánchez-Bayo and Wyckhuys 2019; Cardoso et al. 2020). The Tropical Andes are a biodiversity hotspot (Myers et al. 2000; Larsen et al. 2011) and home to a high number of microendemic species (Murienne et al. 2022).

In this region, habitat transformation has led to a loss of around 75% of natural forest (Bare 2014). The high rate of deforestation, driven primarily by agricultural expansion and cattle ranching (Hoekstra et al. 2005; Armenteras et al. 2013; Sánchez-Cuervo and Aide 2013), is further exacerbated by mining, tourism, and urban development (Portillo-Quintero and Sánchez-Azofeifa 2010). Collectively, these factors contribute to rapid landscape transformation, degradation, and fragmentation. For instance, a recent satellite-based study in natural parks of Colombia, reported a loss of 32,511 hectares of natural cover between 2021 and 2022 (Gualdrón-Díaz & Corredor Gil, 2023), alongside an increase in fragmented forests with secondary vegetation (40,710 ha) and in transformed cover (32,432 ha), which included burned zones, cleared grasslands, and mosaics interspersed with natural spaces.

These changes reflect direct interactions between human activities and the natural environment, driven by socio-economic and environmental factors (Lambin et al. 2003). Human intervention has shaped productive landscapes of

<figure><img src="image_2.png" /></figure>

<footnote>Anderson Arenas-Clavijo
anderson.arenas@correounivalle.edu.co</footnote>

<footnote n="1">¹ Grupo de Investigación en Biología, Ecología y Evolución de Artrópodos, Universidad del Valle, Cali, Colombia</footnote>

<footnote n="2">² Grupo de Investigación en Ecología de Agroecosistemas y Hábitats Naturales, Universidad del Valle, Cali, Colombia</footnote>

<footnote n="3">³ Laboratoire TRACES UMR 5608 5, Université Toulouse Jean Jaurès, Toulouse, France</footnote>
high economic, cultural, and ecological value; however, growing anthropogenic pressure jeopardizes their long-term sustainability (Palang et al. 2017). Land-use change directly influences patterns of insect biodiversity loss, as evidenced by large-scale analyses of deforestation impacts (Van Klink et al. 2020). In Colombia, most human settlements are concentrated in the tropical Andes, exerting significant pressure on this unique ecosystem (Armenteras et al. 2011), where over 69% of the original natural vegetation cover has been transformed for agricultural use (Etter et al. 2006). Consequently, there is an urgent need to investigate how understudied taxonomic groups respond to these dramatic environmental transformations.

Insects are among the organisms most affected by the transformation of natural habitats. Owing to their ecosystem services, they have been widely used to assess the impact of anthropogenic activities across a great variety of environments (Gerlach et al. 2013). Among them, ground beetles (Coleoptera: Carabidae) include both predatory and granivore species, whose assemblages rapidly respond to land use changes (Kromp 1999; Rainio and Niemelä 2003).

In Neotropical lowlands, it has been reported that ground beetle assemblages are less diverse in completely transformed grassland habitats than in natural habitats (Cajaiba et al. 2018). In Colombia, previous studies performed at very local scales show discrepancies in the relationship between the level of perturbation and ground beetle diversity (Arenas-Clavijo and Armbrecht 2019; Ariza et al. 2021). Conversely, surveys conducted along broad elevational gradients in the Andes of Ecuador and Peru have demonstrated that richness and structure of ground beetle assemblages vary with elevation (Moret 2009; Maveety et al. 2011, 2014). However, these studies have been confined to natural habitats.

In this context, it is essential to examine how habitat transformation influences these assemblages across different elevations and slopes of the Andes to gain a more comprehensive understanding of biodiversity dynamics and trajectories. Therefore, the objective of this study was therefore to compare the response of carabid assemblages to the transformation of natural montane forest habitats into grasslands in the tropical Andes of Colombia, at a regional scale, encompassing two cordilleras (i.e. mountain ranges) separated by an inter-Andean valley. We hypothesized that (1) as a result of harsher environmental conditions associated to the lack of tree cover, ground beetle richness and abundance are expected to be lower in anthropogenic grasslands than in natural forests, regardless of the location of the sampled sites within the cordilleras; (2) beta diversity is also expected to be lower in grasslands than in forests, as grassland assemblages are partly structured by species loss and community homogenization, and (3) differences in

species richness and community composition are expected between cordilleras, especially among forests, reflecting their distinct biogeographic trajectories.

# Materials and methods

## Study area

This study was conducted at six sites of Valle del Cauca, southwestern Colombia. Three sampling sites were located between 1896 and 2980 m asl in the Western Cordillera (Table 1, WC1, WC2 and WC3), within the “Páramo de los Farallones de Cali district” bounded to the east by the Cauca River valley and to the west by the Chocó-Darién biogeographic region (Jiménez-Rivillas et al. 2018). Three other sites were located between 2149 and 2788 m asl in the Central Cordillera (Table 1, CC1, CC2 and CC3), within the “Páramos Quindío district” which is limited by the valleys of the Cauca and Magdalena rivers (Jiménez-Rivillas et al. 2018; Fig. 1). The Spanish word “cordillera” refers here to the roughly parallel mountain ranges, oriented N-S, that structure the orography of the Northern Andes.

At each sampling site, six plots measuring 10×10 m were established, three in a forest area (natural habitat) and three in grasslands (transformed habitat) (Table 1). In some cases, grasslands were interspersed with forest patches due to logistical constraints, namely the presence of illegal armed groups, which prevented us from collecting data outside secured areas. Plots were spaced at least 50 m apart and located at least 50 m away from the forest-grassland edge. This distance was considered sufficient to minimize the influence of edge-specialist species (Fernandes Martins et al. 2012). The elevational difference among plots within a given locality did not exceed 200 m (Table 1). Each location was sampled twice (see below).

## Sampling methods

Ground beetles were collected between 2021 and 2023 (Table 1) using three methods: pitfall trapping, litter sifting, and nocturnal hand collection. These complementary techniques were chosen to avoid incomplete sampling (Knapp et al. 2020).

For pitfall trapping, five 500 ml containers were installed at each plot, forming a square 10 m on each side, with four containers at the corners and the fifth one at the center. Pitfall traps were buried flush with the soil surface and filled to two-thirds full with a solution of 5% acetic acid, water, liquid soap, and salt. To prevent trap flooding, artificial roofs were set with wooden sticks and plastic plates. The traps operated for 25 to 37 days (Table 1).
<table><caption>**Table 1** Altitudinal range, geopolitical location, and visit dates for the sampled localities</caption><thead><tr><th>Geographical Formation</th><th>Code</th><th>Municipality</th><th>Locality</th><th>Land Use</th><th>Plot</th><th>Coordinates</th><th>Elevation (m. asl)</th><th>Visit dates</th><th>Total number of days (Pitfall traps)</th></tr></thead><tbody><tr><td rowspan="10">Western Cordillera</td><td rowspan="10">WC1</td><td rowspan="10">Trujillo</td><td rowspan="10">Finca Bellavista</td><td>Fo</td><td>1</td><td>4.17052, -76.40692</td><td>2078</td><td rowspan="10">05-20.xi.2021/27.i-14.ii.2023</td><td rowspan="10">33</td></tr><tr><td></td><td>2</td><td>4.170472, -76.406547</td><td>2066</td></tr><tr><td></td><td>3</td><td>4.170676, -76.406291</td><td>2060</td></tr><tr><td rowspan="3">Gr</td><td>1</td><td>4.170898, -76.405768</td><td>2036</td></tr><tr><td>2</td><td>4.171115, -76.404836</td><td>2009</td></tr><tr><td>3</td><td>4.171854, -76.402507</td><td>1896</td></tr><tr><td rowspan="6">WC2</td><td rowspan="6">Finca Gratinianos</td><td>Fo</td><td>1</td><td>4.17167, -76.42377</td><td>2389</td><td rowspan="6">04-17.xii.2021/13-28.i.2023</td><td rowspan="6">28</td></tr><tr><td></td><td>2</td><td>4.170681, -76.422261</td><td>2357</td></tr><tr><td></td><td>3</td><td>4.170473, -76.421192</td><td>2304</td></tr><tr><td rowspan="3">Gr</td><td>1*</td><td>4.174473, -76.422104</td><td>2232</td></tr><tr><td>2</td><td>4.174247, -76.42194</td><td>2220</td></tr><tr><td>3*</td><td>4.174601, -76.420602</td><td>2159</td></tr><tr><td rowspan="6">WC3</td><td rowspan="6">Monteloro</td><td>Fo</td><td>1</td><td>4.22977, -76.436461</td><td>2906</td><td rowspan="6">09-23.iv.2022/18.ii-04.iii.2023</td><td rowspan="6">28</td></tr><tr><td></td><td>2</td><td>4.230349, -76.437073</td><td>2940</td></tr><tr><td></td><td>3</td><td>4.23099, -76.43749</td><td>2967</td></tr><tr><td rowspan="3">Gr</td><td>1*</td><td>4.231009, -76.438768</td><td>2976</td></tr><tr><td>2*</td><td>4.2303767, -76.438708</td><td>2980</td></tr><tr><td>3</td><td>4.232994, -76.433144</td><td>2806</td></tr></tbody></table>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Geographical Formation</th><th>Code</th><th>Municipality</th><th>Locality</th><th>Land Use</th><th>Plot</th><th>Coordinates</th><th>Elevation (m. asl)</th><th>Visit dates</th><th>Total number of days (Pitfall traps)</th></tr></thead><tbody><tr><td rowspan="10">Central Cordillera</td><td rowspan="10">CC1</td><td rowspan="10">Pradera</td><td rowspan="10">Reserva El Triunfo</td><td>Fo</td><td>1</td><td>3.46325, -76.14793</td><td>2206</td><td rowspan="10">26.iv-10.v.2022/28.ii-13.iii.2023</td><td rowspan="10">27</td></tr><tr><td></td><td>2</td><td>3.463455, -76.148147</td><td>2224</td></tr><tr><td></td><td>3</td><td>3.463945, -76.14823</td><td>2241</td></tr><tr><td rowspan="3">Gr</td><td>1</td><td>3.459228, -76.149392</td><td>2149</td></tr><tr><td>2</td><td>3.459943, -76.149864</td><td>2207</td></tr><tr><td>3</td><td>3.4620755, -76.147504</td><td>2160</td></tr><tr><td rowspan="6">CC2</td><td rowspan="6">El Cerrito</td><td rowspan="6">Reserva El Pailón</td><td>Fo</td><td>1</td><td>3.705204, -76.064732</td><td>2473</td><td rowspan="6">04-19.iii.2022/22.iv-05.v.2023</td><td rowspan="6">28</td></tr><tr><td></td><td>2</td><td>3.70637, -76.06286</td><td>2541</td></tr><tr><td></td><td>3</td><td>3.706564, -76.061814</td><td>2584</td></tr><tr><td rowspan="3">Gr</td><td>1</td><td>3.70155, -76.066713</td><td>2385</td></tr><tr><td>2</td><td>3.700382, -76.066565</td><td>2416</td></tr><tr><td>3</td><td>3.706525, -76.063034</td><td>2558</td></tr><tr><td rowspan="6">CC3</td><td rowspan="6"></td><td rowspan="6">Reserva Madhú</td><td>Fo</td><td>1</td><td>3.642971, -76.143677</td><td>2788</td><td rowspan="6">22.vii-07.viii.2022/17.vi-08.vii.2023</td><td rowspan="6">37</td></tr><tr><td></td><td>2</td><td>3.642817, -76.143816</td><td>2781</td></tr><tr><td></td><td>3</td><td>3.637424, -76.149907</td><td>2614</td></tr><tr><td rowspan="3">Gr</td><td>1</td><td>3.637148, -76.151704</td><td>2590</td></tr><tr><td>2</td><td>3.637505, -76.150501</td><td>2610</td></tr><tr><td>3</td><td>3.637686, -76.149616</td><td>2623</td></tr></tbody></table>

Litter sifting and screening involved collecting 2 L of leaf litter from five randomly selected locations within each plot. The litter was passed through a 1 × 1 cm mesh sieve and carefully inspected to extract carabid specimens.

Active nocturnal hand collection complemented these methods by targeting carabids during their activity period. This technique involved two experienced researchers searching for 30 min per plot, and was performed once per visit, between 19:00 and 23:00, providing 120 min of cumulative search effort per plot across the two visits. Specimens were collected on the ground, on the surface of the leaf litter and on vegetation up to a height of 170 cm.

## Taxonomic treatment

All collected specimens were processed and stored in 70% ethanol in the laboratory of the Research Group on Biology, Ecology, and Evolution of Arthropods at Universidad del Valle, Cali, Colombia. Ground beetles were identified to the finest level possible, using taxonomic keys and revisions of supraspecific taxa (Reichardt 1968, 1972; Noonan 1973; Mateu 1979; Shpeley and Ball 2000; Moret 2003; Ball and Shpeley 2009; Erwin and Zamorano 2014; Boyd and Erwin 2016). When necessary, comparisons were made with type specimens housed at the Muséum National d'Histoire Naturelle (Paris, France). Representative samples were deposited in the Entomological Museum at Universidad del

<figure><img src="image_3.png" /></figure>

<footnote>*Grassland plots interspersed with forests</footnote>
<figure><img src="image_4.png" /><figcaption>**Fig. 1** Location map. Top left: location of Colombia, bottom left: location of Valle del Cauca within Colombia, top right: location of sampling sites, bottom right: example of plot layout within a site</figcaption></figure>

Valle (MUSENUV). Specimens were collected under permit number 1070, issued by the National Environmental Licensing Authority of Colombia.

## Land use characterization

To assess the structural and environmental characteristics of the study sites, several variables were measured within each plot. The number of standing plants with a diameter at breast height (DBH) ≥ 10 cm was recorded, the standard threshold used to define trees (Searle and Chen 2017; González-Melo et al. 2024). The depth of the leaf litter layer was measured using a metal ruler inserted at random points within a 50 cm radius around each pitfall trap.

Environmental conditions were monitored using temperature and humidity loggers (Elitech model RC-51 H; Elitech Technology Inc., San Jose, CA, USA), installed in one forest plot and one grassland plot per locality. These devices were attached to a tree (in forests) or a post (in grasslands) ca. 1.3 m above ground. Air temperature and humidity were recorded hourly throughout the pitfall trap deployment period.

## Data analysis

To evaluate differences in environmental conditions between land uses, we fitted linear mixed models that included land

use and cordillera as fixed factors, and locality as a random effect. This approach aimed to reduce type-1 error inflation. We developed two sets of models: one that included the interaction Cordillera × Land Use, and another that did not. In all cases, model comparisons revealed no significant differences between interaction and non-interaction models; therefore, we selected models without interactions (Table 2, Supplementary Material 1).

Subsequently, a Principal Component Analysis (PCA) was performed to evaluate the contribution of the environmental variables to the variance observed among plots. Following the PCA, multicollinearity was tested using a variance inflation factor (VIF) analysis, which revealed high levels of multicollinearity. To address this issue, variables were grouped by land use, allowing the effects of highly correlated variables to be synthesized into broader categories and thereby reducing VIF values. These analyses were conducted using the ‘FactoMineR’ (Lê et al. 2008) and ‘factoextra’ (Kassambara and Mundt 2020) packages.

We calculated the abundance-based sample coverage for the ground beetles collected in the whole study area. Sample coverage represents the proportion of individuals in a given assemblage that are part of the observed species within a sample (Chiu 2023). For this purpose, we utilized the ‘iNEXT’ package (Chao et al. 2014). Following this, we assessed the impact of habitat transformation on ground beetle assemblages by comparison of species richness and
<table><caption>**Table 2** Summary of LMM tests for environmental variables measured in forests and grasslands from six localities at central (3) and Western (3) cordilleras from Colombia</caption><thead><tr><th>Variable</th><th>Effect tested</th><th>Estimate</th><th>SE</th><th>t-value</th><th>p-value</th></tr></thead><tbody><tr><td rowspan="2">T<sub>max</sub></td><td>Land use (Grassland)</td><td>6.8521</td><td>0.3645</td><td>18.797</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>-0.6898</td><td>1.6706</td><td>-0.413</td><td>0.701</td></tr><tr><td rowspan="2">T<sub>min</sub></td><td>Land use (Grassland)</td><td>0.07661</td><td>0.122</td><td>0.628</td><td>0.5305</td></tr><tr><td>Cordillera (Western)</td><td>0.5713</td><td>1.8220</td><td>0.314</td><td>0.7695</td></tr><tr><td rowspan="2">T<sub>avg</sub></td><td>Land use (Grassland)</td><td>3.4642</td><td>0.1983</td><td>17.470</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>-0.0599</td><td>1.7263</td><td>-0.035</td><td>0.9740</td></tr><tr><td rowspan="2">T<sub>diff</sub></td><td>Land use (Grassland)</td><td>6.7310</td><td>0.3711</td><td>18.139</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>-1.2109</td><td>0.5817</td><td>-2.082</td><td>0.101</td></tr><tr><td rowspan="2">RH<sub>max</sub></td><td>Land use (Grassland)</td><td>0.3755</td><td>0.4010</td><td>0.936</td><td>0.350</td></tr><tr><td>Cordillera (Western)</td><td>0.3091</td><td>0.9305</td><td>0.332</td><td>0.758</td></tr><tr><td rowspan="2">RH<sub>min</sub></td><td>Land use (Grassland)</td><td>-22.175</td><td>1.289</td><td>-17.198</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>5.200</td><td>1.292</td><td>4.024</td><td>0.024</td></tr><tr><td rowspan="2">RH<sub>avg</sub></td><td>Land use (Grassland)</td><td>-10.8889</td><td>0.7739</td><td>-14.070</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>2.9105</td><td>0.7737</td><td>3.762</td><td>&lt;0.001</td></tr><tr><td rowspan="2">RH<sub>diff</sub></td><td>Land use (Grassland)</td><td>22.653</td><td>1.119</td><td>20.235</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>-4.727</td><td>1.579</td><td>-2.994</td><td>0.038</td></tr><tr><td rowspan="2">PBH</td><td>Land use (Grassland)</td><td>-7.8889</td><td>0.7268</td><td>-10.854</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>0.5556</td><td>1.4572</td><td>0.381</td><td>0.722</td></tr><tr><td rowspan="2">LLD</td><td>Land use (Grassland)</td><td>-8.4572</td><td>0.7764</td><td>-10.892</td><td>&lt;0.001</td></tr><tr><td>Cordillera (Western)</td><td>2.6706</td><td>0.9638</td><td>2.771</td><td>0.0503</td></tr></tbody></table>

Variable descriptions: T<sub>max</sub> Maximum daily temperature, T<sub>min</sub> Minimum daily temperature, T<sub>avg</sub> Average daily temperature, T<sub>diff</sub> Difference between maximum and minimum daily temperatures, RH<sub>max</sub> Maximum daily relative humidity, RH<sub>min</sub> Minimum daily relative humidity, RH<sub>avg</sub> Average daily relative humidity, RH<sub>diff</sub> Difference between maximum and minimum daily relative humidity, PBH Number of plants with a perimeter breast height ≥ 30 cm, LLD Average depth of the leaf litter layer

abundance (Jost and González-Oreja 2012). Alpha diversity has been one of the most evaluated parameters in studies on neotropical ground beetles, due to its straightforward interpretation (Maveety et al. 2011; Ariza et al. 2021).

To assess the effect of land uses on richness and abundance, we constructed hierarchized general linear mixed

models (GLMM) with negative binomial distribution, considering plots within each land use, as well as localities within each cordillera as random effects. GLMM were constructed independently for each cordillera, since each one presents specific historical management conditions, and they belong to different biogeographic regions (see “Study Area”). Additionally, we compared localities within each cordillera regardless of land use, by treating plots within each land use at a given locality as random effects. Because localities were situated at different elevations, which can influence species richness and abundance, this approach accounted for potential elevational effects, as previously observed in natural sites (Moret 2009; Maveety et al. 2011, 2014).

These analyses were conducted using the package ‘lme4’ (Bates et al. 2015), and multiple comparisons between levels of each factor were performed with ‘lmerTest’ (Kuznetsova et al. 2017) and ‘emmeans’ (Lenth et al. 2025) packages. Since pitfall traps did not operate for the same duration at every plot, abundance was standardized by dividing the number of collected individuals per sampling days at each locality, and multiplying by the maximum number of days that pitfall traps operated in the locality with more days of pitfall sampling (CC3; 37 days). These values were rounded to the nearest integer and added to the sum of specimens collected through litter sifting and nocturnal hand collection.

We assessed the impact of natural habitat transformation on ground beetle assemblages, by calculating the Sørensen dissimilarity index based on quantitative (standardized abundance) data (Chao et al. 2006; Legendre and De Caceres 2013; Legendre 2014) at each locality within each cordillera. Given that abundance data were highly variable among localities, species and land-use matrices were subjected to Hellinger transformation (i.e., the square root of the relative abundance of each species in a sample) prior to analyses to reduce the influence of very dominant species. From this result, total beta diversity (BDTotal) was decomposed into “richness difference” (BDRich) (sensu Legendre 2014; where richness difference does not imply nestedness), and “species replacement” (BDRepl) components, to identify which components influenced the observed differences between land uses across localities. We also calculated BDTotal, BDRich and BDRepl to compare localities within each land use to detect changes on communities within each cordillera. Additionally, we calculated the BD between cordilleras by pooling all the ground beetle communities present in both land uses at all localities. These analyses were conducted using the ‘adespatial’ package (Dray et al. 2024).

In the present study, we did not aim to assess the nestedness of the samples. With this in mind, we chose the abundance-based Sørensen index from the Podani family (for a detailed discussion, see Legendre's 2014 section “Appendix
<figure><img src="image_5.png" /><figcaption>**Fig. 2** Principal component analysis of environmental variables showing the differences between forests and grasslands. Variable descriptions: Elev: Elevation (in m. asl); Tmax: Maximum daily temperature, Tmin Minimum daily temperature, T<sub>avg</sub> Average daily temperature, T<sub>diff</sub> Difference between maximum and minimum daily temperatures, RH<sub>max</sub> Maximum daily relative humidity, RH<sub>min</sub> Minimum daily relative humidity, RH<sub>avg</sub> Average daily relative humidity, RH<sub>diff</sub> Difference between maximum and minimum daily relative humidity, PBH Number of plants with a perimeter breast height ≥ 30 cm; LLD Average depth of the leaf litter layer</figcaption></figure>

1"). Finally, we performed a nonmetric multidimensional scaling analysis to visualize the conformation of ground beetle assemblages and to identify a potential homogenization effect of land use transformation at a regional scale, by using the ‘vegan’ package (Oksanen et al. 2022). All the statistical analyses were performed in the program R V4.1.2 (R Core Team 2021) in the environment RStudio (RStudio, 2021).

Results

Land use characterization

Grasslands exhibited a wider daily temperature range than forests. Regarding relative humidity, grasslands showed lower daily minimum values, resulting in a higher daily range. The number of trees (DBH≥ 10 cm) was consistently higher in forests than in grasslands, although four grassland plots contained at least one tree. Leaf litter depth was also

consistently greater in forests than in grasslands within each
locality (Table 2, Supplementary Material 1). The princi-
pal component analysis (PCA) revealed a clear separation
between forests and grasslands, with the first two axes
explaining 86.8% of the total variance (Fig. 2). All environ-
mental variables contributed similarly to the principal com-
ponents, except for the maximum relative humidity (RHmax)
which had a lower contribution.

General trends in species richness and abundance

A total of 854 ground beetle specimens were collected, arranged in 63 morphospecies, 24 genera and 11 tribes. Fifteen morphospecies (23.8%) could be assigned to described species. The remaining morphospecies, referred to hereafter as species, could not be assigned to currently described taxa. Most of them probably represent species new to science. Platynini (21 species, 430 individuals) and Harpalini (14 species, 203 individuals) were the dominant tribes, followed by Lebiini (12 species, 36 individuals) (Supplementary Material 2). Six species accounted for over 50% of the total specimens collected: *Dyscolus* sp.7 (145 individuals), *Glyptolenoides* sp.1 (89), *Dyscolus* sp.2 (70), *Incagonum aeneum* (62), *Oxytrechus* sp.1 (56), and *Pelmatellus columbianus* (51). Conversely, 20 singletons and 7 doubletons were found (Table 3, Supplementary Material 2).

In terms of habitat, 47 species were recorded in forests (34 exclusive), while 29 species were collected in grasslands (16 exclusive, half as many as forest specialists). Localities in the Western Cordillera hosted 41 species (627 individuals), whereas those in the Central Cordillera yielded 30 species (227 individuals) (Table 3, Supplementary Material 2). Only eight species were present in both cordilleras (five in both environments, three in grasslands only). Brachypterous flightless species, hence presenting a low dispersal ability, amounted to 48.9% of the species collected in forests, vs. 17.2% in grasslands.

<table><caption>**Table 3** Variation in the species richness of ground beetles based on geographical formation, life zone, elevation, and land use in eight localities in Valle Del Cauca, Colombia</caption>
  <thead>
    <tr>
      <th rowspan="2">Geographical formation</th>
      <th rowspan="2">Locality</th>
      <th colspan="3">Forest</th>
      <th colspan="3">Grassland</th>
      <th rowspan="2">S<sub>Total</sub></th>
    </tr>
    <tr>
      <th>N</th>
      <th>S</th>
      <th>S<sub>excl</sub></th>
      <th>N</th>
      <th>S</th>
      <th>S<sub>excl</sub></th>
    </tr>
  </thead>
  <tbody>
    <tr>
      <td rowspan="3">WC</td>
      <td>WC1</td>
      <td>56</td>
      <td>12</td>
      <td>10</td>
      <td>7</td>
      <td>5</td>
      <td>3</td>
      <td>15</td>
    </tr>
    <tr>
      <td>WC2</td>
      <td>73</td>
      <td>10</td>
      <td>7</td>
      <td>81</td>
      <td>11</td>
      <td>8</td>
      <td>18</td>
    </tr>
    <tr>
      <td>WC3</td>
      <td>327</td>
      <td>13</td>
      <td>7</td>
      <td>83</td>
      <td>11</td>
      <td>5</td>
      <td>18</td>
    </tr>
    <tr>
      <td rowspan="3">CC</td>
      <td>CC1</td>
      <td>64</td>
      <td>10</td>
      <td>7</td>
      <td>25</td>
      <td>7</td>
      <td>4</td>
      <td>14</td>
    </tr>
    <tr>
      <td>CC2</td>
      <td>43</td>
      <td>7</td>
      <td>7</td>
      <td>58</td>
      <td>7</td>
      <td>7</td>
      <td>14</td>
    </tr>
    <tr>
      <td>CC3</td>
      <td>25</td>
      <td>11</td>
      <td>11</td>
      <td>3</td>
      <td>2</td>
      <td>2</td>
      <td>8</td>
    </tr>
  </tbody>
  <tfoot>
    <tr>
      <td colspan="2"><strong>Total*</strong></td>
      <td>597</td>
      <td>47</td>
      <td>34</td>
      <td>257</td>
      <td>29</td>
      <td>16</td>
      <td>63</td>
    </tr>
  </tfoot>
</table>

<footnote>*The total number of species (S) does not correspond to the arithmetic sum of the species richness of each locality but rather to the overall count, considering each land use (forest or grassland) across all localities.</footnote>

<footnote>WC Western Cordillera, CC Central Cordillera, N Abundance, S Richness, Sexcl Number of exclusive species, WC1 Bellavista, WC2 Gratinianos, WC3 Monteloro, CC1 El triunfo natural reserve, CC2 El Pailón natural reserve, CC3 Madhú natural reserve</footnote>
Forests harbored most of the individuals (69.9%), with overall abundance being highest in the Western Cordillera (73.4%). Sample coverage based on the abundance of individuals collected throughout the study was 98.4%.

## Effect of land use changes on ground beetle richness and abundance

### Western cordillera

Ground beetle richness did not differ significantly between land uses (Table 4). When localities were considered regardless of land use, richness at WC3 was 1.81 times higher than at WC1 (estimate: 0.596, *p*<0.05; Table 4). In contrast, abundance was 2.6 times higher in forests compared to grasslands (estimate: 0.957, *p*<0.001; Table 4). Among localities, abundance followed the pattern WC1<WC2<WC3, with all pairwise differences being significant (*p*<0.05; Table 4).

### Central cordillera

Richness was 1.91 times higher in forests than in grasslands (estimate: 0.648, *p*<0.05; Table 4). No significant differences in richness were observed among localities when land use was not considered. Abundance did not differ among localities, whereas forests presented 3.2 times greater abundance than grasslands (estimate: 1.18, *p*<0.001; Table 4).

## Effect of land use changes on ground beetle assemblages

Across all localities in both cordilleras, Sørensen dissimilarity between forests and grasslands was high (79.8% in WC, and 88.6% in CC). This dissimilarity was primarily driven by species replacement (BDRepl at WC=89.9% of BDTotal, BDRepl mean=71.8%; BDRepl at CC=81.5% of BDTotal, BDRepl mean=72.2%). Within land uses, BDTotal observed in forests was 77.1% in WC and 74.7% in CC, in both cordilleras, BDRepl represented more than 90% of BDTotal. A similar pattern was observed in grasslands. When comparing all ground beetle assemblages across both cordilleras, regardless of land use, BDTotal reached 77.2%, with BDRepl accounting for 94.4% of total observed dissimilarity (Table 5). NMDS analysis further supported these results, showing that forest assemblages were clearly differentiated between the two cordilleras, while grassland assemblages tended to group more closely together, regardless of the cordillera they belonged to. WC3 grasslands were slightly separated from other grasslands, without being close to the assemblage composition of forests of the same locality (Fig. 3).

## Discussion

This is the first study that evaluates the effect of land use transformation on the ground beetle assemblages at different elevations on two different mountain slopes in the

<table><caption>**Table 4** Pairwise GLMM contrasts (negative binomial) for ground beetle species richness and abundance between land uses and among localities in the Western and central cordilleras</caption><caption>Western Cordillera</caption><thead><tr><th>Response var.</th><th>Comparison type</th><th>Contrast</th><th>Estimate</th><th>SE</th><th>z-ratio</th><th>p-value</th></tr></thead><tbody><tr><td rowspan="4">Richness</td><td>Land use</td><td>Forest-Grassland</td><td>0.298</td><td>0.187</td><td>1.59</td><td>0.1119</td></tr><tr><td rowspan="3">Locality</td><td>WC1-WC2</td><td>-0.368</td><td>0.247</td><td>-1.49</td><td>0.2972</td></tr><tr><td>WC1-WC3</td><td>-0.596</td><td>0.238</td><td>-2.51</td><td>0.0325</td></tr><tr><td>WC2-WC3</td><td>-0.228</td><td>0.214</td><td>-1.07</td><td>0.5355</td></tr><tr><td rowspan="4">Abundance</td><td>Land use</td><td>Forest-Grassland</td><td>0.957</td><td>0.283</td><td>3.383</td><td>&lt;0.001</td></tr><tr><td rowspan="3">Locality</td><td>WC1-WC2</td><td>-0.964</td><td>0.399</td><td>-2.417</td><td>0.0414</td></tr><tr><td>WC1-WC3</td><td>-1.922</td><td>0.394</td><td>-4.875</td><td>&lt;0.001</td></tr><tr><td>WC2-WC3</td><td>-0.958</td><td>0.382</td><td>-2.507</td><td>0.0326</td></tr><tr><td rowspan="4">Central Cordillera</td><td>Land use</td><td>Forest-Grassland</td><td>0.647</td><td>0.263</td><td>2.462</td><td>0.0138</td></tr><tr><td rowspan="3">Locality</td><td>CC1-CC2</td><td>0.128</td><td>0.329</td><td>0.389</td><td>0.9202</td></tr><tr><td>CC1-CC3</td><td>0.386</td><td>0.349</td><td>1.105</td><td>0.5107</td></tr><tr><td>CC2-CC3</td><td>0.258</td><td>0.357</td><td>0.723</td><td>0.75</td></tr><tr><td rowspan="4">Abundance</td><td>Land use</td><td>Forest-Grassland</td><td>1.18</td><td>0.351</td><td>3.358</td><td>&lt;0.001</td></tr><tr><td rowspan="3">Locality</td><td>CC1-CC2</td><td>0</td><td>0.503</td><td>0</td><td>1</td></tr><tr><td>CC1-CC3</td><td>1.02</td><td>0.52</td><td>1.97</td><td>0.1197</td></tr><tr><td>CC2-CC3</td><td>1.02</td><td>0.52</td><td>1.97</td><td>0.1197</td></tr></tbody></table>

Locality names are coded as follows: WC1 Finca Bellavista (1896–2078 m. asl), WC2 Finca Gratinianos (2159–2389 m. asl), WC3 Monteloro (2806–2980 m. asl), CC1 Reserva El triunfo (2149–2224 m. asl), CC2 Reserva El Pailón (2385–2584 m. asl), CC3 Reserva Madhú (2590–2788 m. asl)
<table><caption>**Table 5** Total beta diversity (based on quantitative Sørensen dissimilarity, Hellinger transformed data) between grassland and forest for each studied locality</caption><thead><tr><th colspan="4">Western Cordillera</th></tr></thead><tbody><tr><th colspan="4">Localities between land uses</th></tr><tr><th>Land Use</th><th>BDTotal</th><th>BDRich</th><th>BDRepl</th></tr><tr><td>Forest</td><td>0.7713</td><td>0.0533</td><td>0.7180</td></tr><tr><td>Grasslands</td><td>0.8210</td><td>0.1066</td><td>0.7144</td></tr><tr><th colspan="4">Land uses within localities</th></tr><tr><td>WC1</td><td>0.8040</td><td>0.1678</td><td>0.6362</td></tr><tr><td>WC2</td><td>0.8285</td><td>0.0714</td><td>0.7571</td></tr><tr><td>WC3</td><td>0.7619</td><td>0.0021</td><td>0.7598</td></tr><tr><th colspan="4">Central Cordillera</th></tr><tr><th colspan="4">Localities between land uses</th></tr><tr><th>Land Use</th><th>BDTotal</th><th>BDRich</th><th>BDRepl</th></tr><tr><td>Forest</td><td>0.7467</td><td>0.0647</td><td>0.6820</td></tr><tr><td>Grasslands</td><td>0.7488</td><td>0.1762</td><td>0.5726</td></tr><tr><th colspan="4">Land uses within localities</th></tr><tr><td>CC1</td><td>0.7817</td><td>0.1004</td><td>0.6813</td></tr><tr><td>CC2</td><td>0.8755</td><td>0.0223</td><td>0.8532</td></tr><tr><td>CC3</td><td>1</td><td>0.3683</td><td>0.6317</td></tr><tr><th colspan="4">Differences between cordilleras, considering all plots, land uses and localities</th></tr><tr><td>Dissimilarity</td><td>BDTotal</td><td>BDRich</td><td>BDRepl</td></tr><tr><td></td><td>0.7723</td><td>0.0431</td><td>0.7292</td></tr></tbody></table>

BDTotal total distance; bdrich: distance explained by The difference in species richness of ground beetles between land uses; bdrepl: distance explained by The difference in assemblage composition (i.e. The identity of The collected species in each land use within each locality). Locality names are coded as follows: *WC1* Finca Bellavista (1896–2078 m. asl), *WC2* Finca Gratinianos (2159–2389 m. asl), *WC3* Monteloro (2806–2980 m. asl), *CC1* Reserva El Triunfo (2149–2224 m. asl), *CC2* Reserva El Pailón (2385–2584 m. asl), *CC3* Reserva Madhú (2590–2788 m. asl)

tropical Andes, with a sampling coverage over 98%. Our results support the hypotheses that ground beetle richness and abundance are lower in grasslands than in forests,

regardless their geographical position and elevation within the two cordilleras, and that assemblages of ground beetles are consistently more homogeneous in grasslands than in forests. We recorded a high rate of singletons, a common situation in neotropical environments (Ariza et al. 2021; Maveety et al. 2011; Riley et al. 2016; Riley Peterson et al. 2021). The richness and abundance of Carabidae recorded in this study were lower than those reported in the Peruvian Andes (Maveety et al. 2011: 77 species), probably because the elevational range of our sampling area was smaller.

## Effect of land use changes on ground beetle richness and abundance

The PCA revealed a clear environmental differentiation between forests and grasslands, independent of plot position or elevation within the cordilleras. Forests were characterized by more stable temperature and humidity conditions, whereas grasslands exhibited greater daily fluctuations. These observations support the idea that tree cover removal directly increases local temperatures (Parsons et al. 2021), thereby reconfiguring invertebrate assemblages, through the dominance of a few habitat-generalist, often heat-tolerant species (Williams and Newbold 2020). This reconfiguration was evident in our dataset, where several fully winged species with broad distributions in the northern Andes, such as *Pelmatellus columbianus* and *Incagonum aeneum* were dominant. These two species have a very wide elevational range and are prevalent in open, anthropogenic habitats (Moret 2005), which suggests a relatively high tolerance to temperature and humidity variability.

We observed higher ground beetle richness and abundance in forest areas across most sampled localities. These habitats were characterized by a greater leaf litter depth,

**Fig. 3** Nonmetric multidimensional scaling for the assemblages of ground beetles from forests (F, in green) and grasslands (G, in yellow) after Hellinger transformation. Stress = 0.071. Locality names are coded as follows: *WC1* Finca Bellavista (1896–2078 m. asl), *WC2* Finca Gratinianos (2159–2389 m. asl), *WC3* Monteloro (2806–2980 m. asl), *CC1* Reserva El Triunfo (2149–2224 m. asl), *CC2* Reserva El Pailón (2385–2584 m. asl), *CC3* Reserva Madhú (2590–2788 m. asl)

<figure><img src="image_6.png" /></figure>
higher number of trees with a DBH ≥ 10 cm, and higher mean relative humidity. Other studies have reported that ground-dwelling beetles are favored by increased leaf litter and humidity (Kirichenko-Babko et al. 2020; Fountain-Jones et al. 2015), while high temperatures and low humidity negatively affect their activity and diversity (Weiss et al. 2024; Wang et al. 2021).

The differences in richness and abundance between forests and grasslands suggest that deforestation and habitat transformation reduce species numbers, a pattern also observed for other taxonomic groups (Alroy 2017; Franco et al. 2019; Newbold et al. 2015). Moreover, the presence of forests may have a buffering effect against adverse environmental conditions, potentially mitigating species loss under climate change scenarios (Ganuza et al. 2022; Suggitt et al. 2018).

In the Western Cordillera, grasslands interspersed with forests (WC2, WC3) showed higher richness and abundance than isolated grasslands (WC1). This is likely due to an edge effect, characterized by the spillover of forest species into adjacent open habitats, and vice versa. This is supported by the presence of shared species in both land uses at WC2 (three species) and WC3 (five species) (Supplementary Material 2), a phenomenon previously reported in Carabidae (Della Rocca et al. 2021; Magura et al. 2001).

## Effects of land use changes and geography on ground beetle assemblages

The strong differentiation between forest and grassland carabid assemblages at each locality, detected by beta diversity analyses, suggests a high degree of ecological specialization among the ground beetle species present in the study area. The NMDS analysis also revealed that all grassland assemblages, regardless of their geographic location across the two cordilleras grouped closely together, even when the grassland area was interspersed with forest patches. This supports the hypothesis of a strong homogenizing effect resulting from forest-to-grassland conversion, as observed for example in the Brazilian Amazon (Solar et al. 2015).

Another notable finding was the high level of species replacement among forest sites, even within the same cordillera (Table 5). This pattern may be explained by an underlying factor that remains relatively unexplored in Colombia: the narrow geographic ranges of microendemic species. This phenomenon has been well documented in the alpine zone of the tropical Andes (Anthelme et al. 2014; Murienne et al. 2022; Moret and Faille 2024), but unpublished data indicate that a similar pattern occurs in montane forests at lower elevations, especially among brachypterous, flightless species, which represent 48.9% of all forest species in our study. This proportion is even higher than that recorded

in an intensively sampled forest of the Ecuadorian Chocó, at 1800–2000 m asl, where 27% of ground beetle species are flightless (Moret and Allegro 2025). The loss of diversity from forest to grassland therefore entails a high risk of extinction for these flightless, microendemic species that are confined to small forest patches within each cordillera.

From a conservation perspective, our study highlights the importance of considering biogeographic context alongside local effects of land-use change. The marked dissimilarity among forest areas emphasizes the need to preserve natural habitats across different slopes and elevations, and to promote environmental management practices that reduce habitat transformation without compromising the economic livelihoods of local communities (Ibarra-Isassi et al. 2021).

## Conclusions

Our study demonstrates that forest-to-grassland conversion has a pronounced negative impact on ground beetle diversity, reducing not only local species richness and abundance, but also regional diversity by homogenizing assemblages across sites. Hence, ground beetle communities become more similar across cordilleras and across localities within each cordillera.

The high beta diversity observed in forest areas, together with the large number of endemic and undescribed species, underscores the urgent need for comprehensive taxonomic and phylogenetic studies. Moreover, it highlights the importance of implementing targeted conservation strategies that prioritize the preservation of forest patches across a range of localities and slopes to maintain ecological integrity and mitigate biodiversity loss.

**Supplementary Information** The online version contains supplementary material available at https://doi.org/10.1007/s10841-025-00739-8.

**Acknowledgements** This work was funded by the “Patrimonio autónomo fondo nacional de financiamiento para la ciencia, la tecnología y la innovación Francisco José de Caldas”, contract number 80740-491-2020, program 70306 “Relaciones multiescalares de la biodiversidad en gradientes altitudinales del bosque tropical”. Special thanks to Dr. I. Armbrecht and Dr. R. González and personnel of Vicerrectoría de Investigaciones, Univalle. We specially acknowledge to our colleagues: N. Montero, V. Plazas, S. Forero, A. Betancourt, L. C. Londoño, and the students D. Hernández, and J. S. Lozano-Sterling, for their support in the field work. Thanks to small holders N. Andrades, A. Gómez, M. Mejía, L. Valencia, J. Cano, and C. Roa for their efforts in the conservation of forests at the studied localities. Finally, but not less important, we are appreciative for the participation of the anonymous reviewers and the section editor, whose insightful comments greatly improved the quality of this paper.

**Author contributions** Conceptualization, Methodology, Writing—original draft, Writing—review and editing: Anderson Arenas-Clavijo, James Montoya-Lerma, Pierre Moret; Data curation, Formal Analysis,
**Investigation and Visualization:** Anderson Arenas-Clavijo; Methodology: Anderson Arenas-Clavijo, James Montoya-Lerma, Pierre Moret; Project Administration and Funding Acquisition: James Montoya-Lerma; Supervision: James Montoya-Lerma and Pierre Moret.

**Funding** Open Access funding provided by Colombia Consortium. JML was supported by Universidad del Valle (Crossref Funder ID 501100007329; Research Organization Registry: https://ror.org/00jb9vg53).

**Data availability** No datasets were generated or analysed during the current study.

## Declarations

**Conflict of interest** The authors declare no competing interests.

**Open Access** This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

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