Open access journal: http://periodicos.uefs.br/index.php/sociobiology ISSN: 0361-6525 DOI: 10.13102/sociobiology.v69i3.8099Sociobiology 69(3): e8099 (September, 2022) Introduction Metacommunities are a set of local communities that are linked by dispersal of multiple interacting species (Wilson, 1992). Local metacommunities are shaped by speciation, dispersal, abiotic conditions, biotic interactions, and ecological drift processes (Vellend, 2010; 2016). Such processes, which shape the structure and dynamics of local communities, may occurs as a result of habitat fragmentation (Chase et al., 2020). Habitat fragmentation isolate natural habitats, change their abiotic conditions, resource availability and quality (Arroyo-Rodríguez et al., 2017; 2020; Fahrig, 2003; Schmidt et al., 2013; Sobrinho et al., 2003; Sobrinho & Schoereder, 2007). Communities from isolated habitats Abstract Habitat fragmentation changes biological communities and its spatiotemporal dynamics – which may lead to either biotic homogenization or heterogenization along time and space. Both processes can occur by addition, replacement or loss of species within communities, altering compositional similarity across the landscape. We investigated which of these two processes (biotic homogenization or heterogenization) occurs, and its possible underlying mechanism, over 15 years in an Atlantic Forest landscape using ants as model organisms. We sampled ants in 17 forest fragments across three different years, compared their composition similarity, species richness, and species richness of groups classified according to their habitat preferences. We sampled a total of 132 ant species. Ant communities in fragments diverged over time, suggesting they experienced an idiosyncratic structuring process. This biotic heterogenization occurred through an additive process, as ant species richness increased over time, mainly due to an increase of generalist ant species, and a decrease of forest specialist ant species. These changes occurred despite the higher forest cover in the landscape along years. Since different species can perform different functions in ecosystems, this biotic heterogenization may have implications for ecosystem functioning. Investigating how disturbances structure biological communities over time, especially those performing important ecosystem functions, can shed light to our understanding of possible changes in ecosystem functions and consequently for forest regeneration. Sociobiology An international journal on social insects Vanessa S. Ribeiro¹, Lucas N. Paolucci¹, José H. Schoereder¹, Ricardo R. de C. Solar² Article History Edited by Wesley Dáttilo, Mexico Received 25 April 2022 Initial acceptance 26 April 2022 Final acceptance 12 August 2022 Publication date 07 September 2022 Keywords Biotic Heterogenization, Biotic Homogenization, Habitat Fragmentation, Community Divergence, Formicidae, Tropical Rain Forest. Corresponding author Vanessa Soares Ribeiro Department of General Biology Federal University of Viçosa Av. Peter Henry Rolfs s/nº, Campus Universitário, CEP: 36570 000 Viçosa, Minas Gerais, Brasil. E-Mail: soaresribeirov@gmail.com have smaller populations, different migration patterns and distribution, and richness and composition (Arroyo-Rodríguez et al., 2020; Bender et al., 1998; Fahrig, 2019; Lasmar et al., 2021; Madureira et al., 2012; Sobrinho & Schoereder, 2007; Vellend, 2010). These changes in species dynamics within fragments, may lead communities to become either more similar among each other or more dissimilar – i.e., they can cause a biotic homogenization (McKinney & Lockwood 1999) or biotic heterogenization, respectively (Laurance et al., 2007; Socolar et al., 2016). Biotic homogenization may occur due to two mechanisms: additive and subtractive. Additive homogenization occurs when new species immigrate from other regions and become present in most fragments in the landscape. 1 - Department of General Biology, Federal University of Viçosa, Viçosa-MG, Brazil 2 - Department of Genetics, Ecology and Evolution, Federal University of Minas Gerais, Belo Horizonte-MG, Brazil RESEARCH ARTICLE - ANTS Divergence of Ant Communities Over Time in a Fragmented Atlantic Rain Forest Landscape mailto:soaresribeirov@gmail.com Vanessa S. Ribeiro et al. – Divergence of ant communities over time across fragments2 Subtractive homogenization occurs when particular species with a distribution restricted to a few fragments disappear from all of them and consequently from the landscape (Socolar et al., 2016). Therefore, biotic homogenization occurs through immigration and extinction, which in turn impact species distribution and abundance across local sites (Olden, 2006). Lower connectivity in landscapes after fragmentation benefits species with a high dispersal ability, that can establish in isolated fragments and contribute to biotic homogenization (Chase et al., 2020). Changes in resource availability due to habitat fragmentation can benefit species that have more similar needs and are more adapted to disturbances (generalist species), altering local species composition due to invasion and extinction (McKinney & Lockwood, 1999; Olden et al., 2004; Tabarelli et al., 2012). Biotic heterogenization may also occur due to species introduction (additive mechanisms) or species loss (subtractive mechanisms) (Socolar et al., 2016). Contrary to homogenization, heterogenization is likely driven mainly by idiosyncratic events, and if so local communities diverge along time after disturbance (Audino et al., 2014; Laurance et al., 2007; Socolar et al., 2016). For instance, additive heterogenization occurs when each fragment experiences a unique successional pattern, being colonized by different species from neighbouring fragments (Arroyo-Rodríguez et al., 2013) which depends on landscape connectivity and dispersal ability of organisms (Mouquet & Loreau, 2003). Subtractive heterogenization occurs if certain species are more prone to extinction due to new abiotic conditions, different biotic interactions and stochastic events, favouring different species across fragments (Socolar et al., 2016). In this case, changes in species groups - which represent the role that species play in the ecosystem - can also occur due to specialist species extinction and/or generalist species invasion (Tabarelli et al., 2012). Habitat fragmentation can thus shape community structure across space and time by changing migration and extinction patterns, and local conditions and resources (Chase et al., 2020; Dornelas et al., 2014; Solar et al., 2015). To better conserve and restore impacted communities it is necessary to understand how and why community structuring vary over time (Arroyo-Rodríguez et al., 2013; Chase et al., 2020), but it is still unclear how biotic homogenization and heterogenization contribute to shape biological communities in human-driven fragmented landscapes (Arroyo-Rodríguez et al., 2020; Chase et al., 2020). Available studies suggest that metacommunities can suffer both processes in human-altered landscapes (Chase et al., 2020), and as a demand for conservation, it is necessary to understand how these processes contribute to community structuring in tropical forests, which are under high anthropogenic pressure. Additionally, anthropic pressure can favor non-native species invasion and exclude native species in forests, which first leads to biotic heterogenization, as initially species invades few fragments, but these species spread and can lead to biotic homogenization (McKinney & Lockwood, 1999). Here we investigated processes driving ant community composition in a fragmented Atlantic Forest landscape, a hotspot Brazilian biome that suffered a severe fragmentation process: there is about 15% of the original cover remaining, almost half of which is located < 100 m from the edge, and most remaining fragments are < 50 ha (Ribeiro et al., 2009). We studied ants (Hymenoptera: Formicidae) because they are highly sensitive to habitat fragmentation (Lasmar et al., 2021; Vasconcelos et al., 2006), reflect impacts on other animal and plant species (Majer et al., 1997), and constitute an important ecological group of insects in tropical forests due to their large biomass. Additionally, ants play important roles in several ecological processes such as seed dispersal, invertebrate predation, soil bioturbation and herbivory (Ribas et al., 2012). We studied 17 secondary fragments of Atlantic Forest in southeastern Brazil throughout a 15-year interval. We investigated whether ant communities across these forest fragments experienced homogenization or heterogenization (or if neither process occurred), and which underlying mechanism (subtractive or additive) was responsible for such changes. We also evaluated whether generalist ant species are responsible for possible changes in species composition, by investigating whether species proportion of this group is higher in fragments. Materials and methods Study site We carried out this study in the Atlantic Forest of southeastern Brazil, in the municipality of Viçosa, Minas Gerais state (20°45’S, 42°50’W). According to landowners of the region and previous studies, the region was covered by forests up to the 19th century, but during the 1930’s and 1940’s an accelerated process of fragmentation has begun and lasted until the 1960’s (Schmidt et al., 2013; Sobrinho & Schoereder 2007). Most current fragments in the region resulted from natural recovery, and are between 20 and 60 years old, although some are up to 100 years old (Ribon et al., 2003). We documented changes in land-use types in our study region to better understand how different land-use cover could influence our results. During the years in which the samples were collected, there was an overall increase in forest cover in the study region (Table 1: Supplementary Material; MapBiomas, 2020). Ant sampling We sampled ant species from 17 fragments (Figure 1) in three years: 1995, 1996 and 2010 at the same points in all years, and always in the rainy season (from November to March). Fragments varied in size from three to 300 hectares (detailed in Table 2: Supplementary Material). We used pitfall traps of 19 cm in diameter and 11 cm high buried at Sociobiology 69(3): e8099 (September, 2022) 3 the ground level, containing water and detergent. Traps were baited with human feces and cattle carrion, and remained in the field for 48 h. These baits were used because, in addition to ants, samples were also intended to collect other invertebrates. Although the use of bait is not common in ant community studies, it is known that at least 60% of total ant species are captured using this method (Przybyszewski et al., 2020). Furthermore, the same method was repeated in all sampled and years. Each sampling point was constituted by a set of two traps, each with one bait type, three meters apart. According to Schoereder et al. (2004a), we used a sampling effort proportional to each fragment size: in fragments larger than 93 hectares, 48 traps were placed, totaling 24 sampling points; in fragments between 30 and 61 hectares, 24 traps in 12 sampling points were used; and in fragments between 3 and 9 hectares, 8 traps at 4 sampling points were used. Traps were placed at least 20 m from the edge of the fragments, and the distance between each sampling point was at least 30 m. Ant species were identified to the smallest possible level and deposited in the ant collection of the Laboratório de Sistemática e Biologia de Coleoptera of the Universidade Federal de Viçosa, Brazil. To investigate whether different groups of ant species are responsible for possible changes in species composition, we categorized sampled ant species as generalists, forest specialists, open-habitat specialists or unclassified, according to environments in which species are typically found. Accordingly, forest specialist species rarely or never occur in savanna, and open-habitat specialists rarely or never occur in forest. Generalist species frequently occur in both types of habitats. This classification was only attributed for species where enough information is available. An ant taxonomist (Julio M.C. Chaul) provided the classifications. Statistical analysis To test whether ant communities in the studied fragments experienced a process of biotic homogenization or heterogenization, we first compared species composition similarity across all pairs of fragments from each year separately, using the Jaccard similarity index. Afterwards we compared the average similarity across years using a GLMM (Generalized Mixed Model), with fragment pair identity as a random effect, and the year of sampling as a fixed effect. As we expected larger changes in the mean similarity indices between the years 1995 - 2010 than between 1995 - 1996, the years were included in the model as a continuous variable. We used a binomial error distribution because these data are strictly bounded which values are between 0 and 1 (Crawley, 2013), and a contrast analysis to check if ant composition similarity differed among years. We also investigated whether the found process (i.e., biotic homogenization or heterogenization process) was driven by species addition or subtraction mechanisms by analyzing changes in ant species richness over time across fragments. Changes in ant species composition driven by an increase in ant species richness in fragments suggest an additive mechanism. Alternatively, a decrease in ant species richness in each fragment suggests a subtractive mechanism. For that, we built a GLMM with Poisson error distribution, using fragments as a random effect and the sampling year as a fixed effect. We used a contrast analysis to check for differences among years. We used the proportion of ant species richness from each group (generalists, forest specialists and open-habitat specialists species) as a response variable to test whether changes in ant species richness of these groups could have Fig 1. Map of sampled Atlantic rain forest fragments in Viçosa region, Minas Gerais state, Brazil. The red crosses show the location of the 17 fragments sampled in 1995, 1996 and 2010. Vanessa S. Ribeiro et al. – Divergence of ant communities over time across fragments4 caused biotic homogenization or heterogenization, through additive or subtractive mechanisms. The proportion of ant species from each group was obtained by dividing the number of species from each group by the total number of species sampled in each fragment and year. We adjusted a GLMM with a Binomial error distribution, using proportion of ant species richness from each group as the response variable, forest fragments as a random effect, and year as a fixed effect. We also used a contrast analysis to check for species richness differences among years. We used the R software (R-core team 2021) to perform all analyzes. We used the packages vegan 2.5-6 (Oksanen et al., 2019) for species composition, lme4 1.1-26 (Bates et al., 2015) to perform GLMMs, and DHARMa 0.3.3.0 (Hartig, 2020) to perform residual analysis, check model suitability and distribution. Results We sampled a total of 132 ant species, distributed in eight subfamilies: Dolichoderinae; Dorylinae; Ectatomminae; Formicinae; Heteroponerinae; Myrmicinae; Ponerinae and Pseudomyrmicinae (Table 3: Supplementary Material). Subfamilies Dorylinae and Heteroponerinae was the only subfamily that decreased the species number in the year 2010, when compared to 1995 and 1996. Dorylinae had only one species, Neivamyrmex alfaroi, which was sampled in 1995 and 1996. Only two species were from the Heteroponerinae: Heteroponera dolo, which was sampled in all years, and Heteroponera dentinodis, sampled only in 1996. Subfamily Myrmicinae had the highest number of species in all years, representing 44% of total species. The average similarity of ant species composition among fragments differed over time (χ² [2, N = 17] = 72.036, p < 0.001, Figure 2). Ant species richness was higher in 2010 than in 1995 and 1996 (χ² [1, N = 17] = 9.13, p = 0.002, Figure 3), which did not differ each other. Among the species sampled, 28% were classified as generalist, 34% forest specialist, 3% open-habitat specialist, and 35% of the species could not be classified into the three categories. Fig 3. Average ant species richness across studied fragments, located in southeastern Brazilian Atlantic rain forest. In 2010, ant species richness was higher than in 1995 and 1996 (p = 0.002). Each point represents a fragment sampled in the correspondent year. The black lozenges represent the average and the bars represent the standard error. Different letters represent significant differences at alfa = 0.05. Fig 2. Average similarity of ant species composition across studied fragments, located in southeastern Brazilian Atlantic rain forest. Each point represents a pairwise comparison between fragments using the Jaccard similarity index. To compare averages, we used a mixed model considering each forest fragment identity as a random effect (see main text). The black lozenges represent the average, and the bars represent the standard error. Different letters represent significant differences at alfa = 0.05. Fig 4. Average proportion of generalist ant species richness across studied fragments, located in southeastern Brazilian Atlantic rain forest. In 2010 it was higher than when compared to 1995 and 1996 (p = 0.01). Points represent the proportion of generalist ant species from each fragment in each year. The black lozenges represent the average and the bars represent the standard error. Different letters represent significant differences at alfa = 0.05. Sociobiology 69(3): e8099 (September, 2022) 5 The proportion of generalist species increased over time (χ² [2, N = 17] = 9.0914, p = 0.01, Figure 4). However, the proportion of forest specialist species was higher in 1995 compared to 1996 and 2010 (χ² [2, N = 17] = 32.264, p < 0.001, Figure 5), while open-habitat specialist species was similar among years (χ² [2, N = 17] = 1.76, p = 0.41, Figure 6). Discussion Here we documented a 15-year long biotic hetero- genization of ant communities across a fragmented rain forest landscape, suggesting ant communities experienced an idiosyncratic structuring process. The average ant species richness in fragments increased over time, so this biotic heterogenization probably occurred through an additive mechanism. Important implications for conservation biology emerge from this idiosyncrasy, since each fragment may contain a different set of species, including those with key roles in the ecosystem, interactions, and also invasive species (Laurance et al., 2007). Biotic heterogenization occurred due to an increase of ant species richness in 2010, which was ~20% higher in comparison to 1995. In the same period, the forested area in our study landscape increased by 3% and agricultural area decreased by 4.2% (Table S.2: Supplementary Material). Habitat structure can be an environmental filter strongly related to changes in ant richness and composition in forest fragments (Castro et al., 2020). The increase of forest cover probably causes a higher habitat availability and quality for ant species in our study region (Schmidt et al., 2013; Sobrinho et al., 2003), which in turn reduces the extinction risk to forest species, and allows species to establish across fragments. Furthermore, forest recovery likely contributed to the biotic heterogenization we observed because it increased the proportion of different habitat (i.e., forest and agriculture) available in the landscape. As ant communities markedly differ across habitat types in our study region (Schmidt et al., 2013), some ant species from agriculture can enter forest remnants (Thomas, 2020) and alter their species composition (Sobrinho et al., 2003). Despite higher forest area in 2010, forest specialist ant species richness decreased over time. Specialist ant species are more vulnerable to disturbed environments (Andersen, 2019), and our results showed that these species were not reestablished in our fragments across the years. Contrarily, the number of generalist species increased about 254% in 2010 compared to 1995 – for instance, species usually found in disturbed savannas as Ectatomma planidens and Linepithema humile (Ramos et al., 2003) were sampled only in 2010, suggesting that generalist species and from open areas invaded fragment forests along years, despite the higher forest cover. This result supports recent findings that edge and matrix effects can increase ant species richness in local scales due to an invasion of generalist ant species (Lasmar et al., 2021), which are less vulnerable to impacts due to their wider use of resources (Andersen, 2019). Invasions of ant species from open areas into forests can occur after other anthropogenic disturbances, as due to recurrent forest fires and consequent canopy opening (Paolucci et al., 2017). In the studied fragments, one of the explanations for the invasion of generalist species in small fragments is the similarity between Fig 6. Average proportion of open-habitat specialist ant species richness across studied fragments, located in southeastern Brazilian Atlantic rain forest. Open-habitat specialist species was similar among years in fragments located in southeastern Brazilian Atlantic rainforest (p = 0.41). Points represent the proportion of open-habitat specialist ant from each fragment sampled in each year. The black lozenges represent the average and the bars represent the standard error. Different letters represent significant differences at alfa = 0.05. Fig 5. Average proportion of forest specialist ant species richness across studied fragments, located in southeastern Brazilian Atlantic rain forest. In 1995 it was higher when compared to 1996 and 2010 in fragments located in southeastern Brazilian Atlantic rain forest (p < 0.001). Points represent the proportion of forest specialist ant species from each fragment sampled in each year. The black lozenges represent the average and the bars represent the standard error. Different letters represent significant differences at alfa = 0.05. Vanessa S. Ribeiro et al. – Divergence of ant communities over time across fragments6 these fragments and the surrounding matrix (Schoereder et al., 2004b), and here we show that such invasions can contribute to a biotic heterogenization of ant communities. We did not test what were the consequences of additive biotic heterogenization for the ecosystem functions performed by ants in this landscape, although the loss of functions due to changes in ant species composition has been found before (Del Toro et al., 2012). What we do know is that the environmental filters imposed by habitat fragmentation can change ant taxonomic diversity, but not necessarily the functional diversity (Castro et al., 2020). Thus, even if biotic heterogenization occurs in the fragments, ecosystem functions promoted by ants can still be maintained, but this is an area for further investigations. Our results have important implications for decisions regarding rain forests conservation, since ants are an ecologically dominant group, that play several ecosystem functions, and also bioindicators that can represent changes in other taxonomic groups (Andersen & Majer, 2004). Forest fragments under a natural recovery process can experience idiosyncratic trajectories regarding environmental conditions, human pressure and thus colonization / extinction of plant species (Laurance et al., 2007), and our results point to a parallel process for the associated fauna. The knowledge about ant species composition changes in remaining fragments and also about how these can impact ecosystem functions performed by ants are important to understand natural forest recovery, given several functions performed by ants relate to this process, as seed dispersal, herbivory and predation of insect herbivores. In addition, we hope that identifying empirically that fragmentation drives communities divergence leads to further investigations on other types of disturbances and their consequences for biological communities over time. Acknowledgments We thank A. C. Maradini for her data survey assistance, J. Chaul for assisting with ant species identification and habitat classification. C. Ribas, L. Rattis, T. Sobrinho and I. Gomes discussed and improved the first draft of the manuscript. Two anonymous referees and the Editor provided valuable suggestions that improved the manuscript. We thank the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) and Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG) for supporting this study. JHS was supported by a Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) grant. Authors contribution All authors contributed in the stages of: conceptualization, methods, validation, formal analysis, writing, review and editing, visualization, supervision. J.H. Schoereder was responsible for research, resources and procurement of funding. References Andersen, A.N. (2019). Responses of ant communities to disturbance: five principles for understanding the disturbance dynamics of a globally dominant faunal group. Journal of Animal Ecology, 88: 350-362. doi: 10.1111/1365-2656.12907 Andersen, A.N. & Majer, J.D. (2004). Ants show the way down under: invertebrates as bioindicators in land management. Frontiers in Ecology and the Environment, 2: 291-298. doi: 10.1890/1540-9295(2004)002[0292:ASTWDU]2.0.CO;2 Arroyo-Rodríguez, V., Fahrig, L., Tabarelli, M., Watling, J.I., Tischendorf, L., Benchimol, M., Cazetta, E., Faria, D., Leal, I.R., Melo, F.P.L., Morante-Filho, J.C., Santos, B.A., Arasa-Gisbert, R., Arce-Peña, N., López, M.J.C., Cudney- Valenzuela, S., Galán-Acedo, C., San-José, M., Vieira, I.C.G., Slik, J.W.F., Nowakowski, A.J. & Tscharntke T. (2020). Designing optimal human-modified landscapes for forest biodiversity conservation. Ecology Letters, 23: 1404- 1420. doi: 10.1111/ele.13535 Arroyo-Rodríguez, V., Rös, M., Escobar, F., Melo, F.P.L., Santos, B.A., Tabarelli, M. & Chazdon, R. (2013). Plant β-diversity in fragmented rain forests: testing floristic homogenization and differentiation hypotheses. Journal of Ecology, 101: 1449-1458. doi: 10.1111/1365-2745.12153 Arroyo-Rodríguez, V., Saldaña-Vázquez, R.A., Fahrig, L. & Santos, B.A. (2017). Does forest fragmentation cause an increase in forest temperature? Ecological Research, 32: 81- 88. doi: 10.1007/s11284-016-1411-6 Audino, L.D., Louzada, J., Comita, L. (2014). Dung beetles as indicators of tropical forest restoration success: is it possible to recover species and functional diversity? Biological Conservation, 169: 248-257. doi: 10.1016/j.biocon.2013.11.023 Bates, D., Martin, M., Bolker, B. & Walker, S. (2015). Fitting linear mixed-effects models using lme4. Journal of Statistical Software, 67: 1-48. doi: 10.18637/jss.v067.i01 Bender, D.J., Contreras, T.A. & Fahrig, L. (1998). Habitat loss and population decline: a meta-analysis of the patch size effect. Ecology, 79: 517-533. doi: 10.1890/0012-9658(1998) 079[0517:HLAPDA]2.0.CO;2 Castro, F.S. de, Da Silva, P.G., Solar, R.R.C., Fernandes, G.W. & Neves F.S. (2020). Environmental drivers of taxonomic and functional diversity of ant communities in a tropical mountain. Insect Conservation and Diversity, 13: 393-403. doi: 10.1111/ icad.12415 Chase, J.M., Jeliazkov, A., Ladouceur, E. & Viana, D.S. (2020). Biodiversity conservation through the lens of metacommunity ecology. Annals of the New York Academy of Sciences, 1469: 86-104. doi: 10.1111/nyas.14378 Crawley, M.J. (2013). The R book. Second edition. Chichester, West Sussex, UK: Wiley. Sociobiology 69(3): e8099 (September, 2022) 7 Del-Toro, I., Ribbons, R.R., Pelini, S. (2012). The little things that run the world revisited: a review of ant- mediated ecosystem services and diservices (Hymenoptera: Formicidae). Myrmecological News, 17: 133-146. Dornelas, M., Gotelli, N.J., McGill, B., Shimadzu, H., Moyes, F., Sievers, C. & Magurran, A.E. (2014). Assemblage time series reveal biodiversity change but not systematic loss. Science, 344: 296-299. doi: 10.1126/science.1248484 Fahrig, L. (2003). Effects of habitat fragmentation on biodiversity. Annual Review of Ecology, Evolution, and Systematics, 34: 487-515. doi: 10.1146/annurev.ecolsys.34.011802.132419 Fahrig, L. (2019). Habitat fragmentation: a long and tangled tale. Global Ecology and Biogeography, 28: 33-41. doi: 10.11 11/geb.12839 Hartig, F. (2020). DHARMa: residual diagnostics for hierarchical (multi-level/mixed) regression models. R package version 0.4.1. https://CRAN.R- project.org/package=DHARMa Lasmar, C.J., Queiroz, A.C.M., Rosa, C., Carvalho, N.S., Schmidt, F.A., Solar, R.R.C., Paolucci, L.N., Cuissi, R.G. & Ribas, C.R. (2021). Contrasting edge and pasture matrix effects on ant diversity from fragmented landscapes across multiple spatial scales. Landscape Ecology, 36: 2583-2597. doi: 10.1007/s10980-021-01258-y Laurance, W.F., Nascimento, H.E.M., Laurance, S.G., Andrade, A., Ewers, R.M., Harms, K.E., Luizão, R.C.C. & Ribeiro, J.E. (2007). Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis. PLoS ONE, 2: e1017. doi: 10.1371/journal.pone.0001017 Leal, I.R., Filgueiras, B.K.C., Gomes, J.P., Iannuzzi, L. & Andersen, A.N. (2012). Effects of habitat fragmentation on ant richness and functional composition in Brazilian Atlantic forest. Biodiversity and Conservation, 21: 1687-1701. doi: 10.1007/s10531-012-0271-9 Madureira, M.S., Sobrinho, T.G. & Schoereder, J.H. (2012). Why do ant species occur in the matrix and not in the forests? invasion from other habitats or expansion from forest gaps (Hymenoptera: Formicidae). Sociobiology, 59: 1137-1149. doi: 10.13102/ sociobiology.v59i4.513 Majer, J.D., Delabie, J.H.C. & McKenzie, N.L. (1997). Ant litter fauna of forest, forest edges and adjacent grassland in the Atlantic Rain Forest Region of Bahia, Brazil. Insectes Sociaux, 44: 255-266. doi: 10.1007/s000400050046 McKinney, M.L. & Lockwood, J.L. (1999). Biotic homogenization: a few winners replacing many losers in the next mass extinction. Trends in Ecology and Evolution, 14: 450- 453. doi: 10.1016/S0169-5347(99)01679-1 MapBiomas – Coleção v.6. Série Anual de Mapas de Cobertura e Uso de Solo do Brasil. Acessado em 13/09/2021: https:// plataforma.brasil.mapbiomas.org/ Mouquet, N. & Loreau, M. (2003). Community patterns in source-sink metacommunities. The American Naturalist, 162: 544-557. doi: 10.1086/378857 Oksanen, J., Blanchet, F.G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, P., Stevens, M.H.H., Szoecs, E. & Wagner, H. (2020). vegan: community ecology. Package. R package version 2.5-7. https://CRAN.R-project.org/package=vegan Olden, J..D. (2006) Biotic homogenization: a new research agenda for conservation biogeography. Journal of Biogeography, 33: 2027-2039. doi: 10.1111/j.1365-2699.2006.01572.x Olden, J.D., Poff, N.L., Douglas, M.R., Douglas, M.E. & Fausch, K.D. (2004). Ecological and evolutionary consequences of biotic homogenization. Trends in Ecology and Evolution, 19: 18-24. doi: 10.1016/j.tree.2003.09.010 Paolucci, L.N., Schoereder, J.H., Brando, P.M. & Andersen, A.N. (2017). Fire-induced forest transition to derived savannas: cascading effects on ant communities. Biological Conservation, 214: 295-302. doi: 10.1016/j.biocon.2017.08.020 Przybyszewski, K.R., Silva, R.J., Vicente, R.E., Freitas, J.V.G., Pereira, M.J.B., Izzo, T.J., Tonon, D.S. (2020). Can baited pitfall traps for sampling dung beetles replace conventional traps for sampling ants?” Sociobiology, 67: 376- 387. doi: 10.13 102/sociobiology.v67i3.5201. R Core Team (2021). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria https://www.R-project.org/. Ramos, L.S., Filh, R.Z.B., Delabie, J.H.C., Lacau, S. & Marinho C.G.S. (2003). Comunidades de formigas (Hymenoptera: Formicidae) de serapilheira em areas de cerrado ‘stricto sensu’ em Minas Gerais. Lundiana: International Journal of Biodiversity, 4: 95-102. doi: 10.35699/2675-5327.2003.21859 Ribas, C.R., Campos, R.B.F., Schmidt, F.A. & Solar, R.R.C. (2012). Ants as indicators in Brazil: a review with suggestions to improve the use of ants in environmental monitoring programs. A Journal of Entomology, 2012: 1-23. doi: 10.1155/ 2012/636749 Ribeiro, M.C., Metzger, J.P., Martensen, A.C., Ponzoni, F.J. & Hirota, M.M. (2009). The Brazilian Atlantic Forest: how much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation, 142: 1141-1153. doi: 10.1016/j.biocon.2009.02.021 Ribon, R., Simon, J.E. & De Mattos, G.T. (2003). Bird extinctions in atlantic forest fragments of the Viçosa Region, southeastern Brazil. Conservation Biology, 17: 1827-1839. doi: 10.1111/j.1523-1739.2003.00377.x Schmidt, F.A., Ribas, C.R. & Schoereder, J.H. (2013). How predictable is the response of ant assemblages to natural forest recovery? Implications for their use as bioindicators. Vanessa S. Ribeiro et al. – Divergence of ant communities over time across fragments8 Ecological Indicators, 24: 158-166. doi: 10.1016/j.ecolind. 2012.05.031 Schoereder, J.H., Galbiati, C., Ribas, C.R., Sobrinho, T.G., Sperber, C.F., DeSouza, O., & Lopes-Andrade, C. (2004a). Should we use proportional sampling for species-area studies? Journal of Biogeography, 31: 1219-1926. doi: 10.1111/j.1365- 2699.2004. 01113.x Schoereder, J.H., Sobrinho, T.G., Ribas, C.R., Campos, R.B.F. (2004b). Colonization and extinction of ant communities in a fragmented landscape. Austral Ecology, 29: 391-398. doi: 10.1111/j.1442-9993.2004.01378.x Sobrinho, T.G., Schoereder, J.H., Sperber, C.F. & Madureira M.S. (2003). Does fragmentation alter species composition in ant communities (Hymenoptera: Formicidae)? Sociobiology 42: 1-14. Sobrinho, T.G. & Schoereder, J.H. (2007). Edge and shape effects on ant (Hymenoptera: Formicidae) species richness and composition in forest fragments. Biodiversity and Conservation, 16: 1459-1470. doi: 10.1007/s10531-006- 9011-3 Socolar, J.B., Gilroy, J.J., Kunin, W.E. & Edwards, D.P. (2016). How should beta-diversity inform biodiversity conservation? Trends in Ecology & Evolution, 31: 67-80. doi: 10.1016/j.tree.2015.11.005 Solar, R.R.C., Barlow, J., Ferreira, J., Berenguer, E., Lees, A.C., Thomson, J.R., Louzada, J., Maués, M., Moura, N.G., Oliveira, V.H.F., Chaul, J.C.M., Schoereder, J.H., Vieira, I.C.G., Nally, R.M. & Gardner, T.A. (2015). How pervasive is biotic homogenization in human-modified tropical forest landscapes? Ecology Letters, 18: 1108-1118. doi: 10.1111/ ele.12494 Tabarelli, M., Peres, C.A. & Melo, F.P.L. (2012). The ‘few winners and many losers’ paradigm revisited: emerging prospects for tropical forest biodiversity. Biological Conservation, 155: 136-140. doi: 10.1016/j.biocon.2012.06.020 Thomas, C.D. (2020). The development of anthropocene biotas. Philosophical Transactions of the Royal Society B: Biological Sciences, 375: 20190113.doi: 10.1098/rstb.2019.0113 Vasconcelos, H.L., Vilhena, J.M.S., Magnusson, W.E. & Albernaz, A.L.K.M. (2006). Long-term effects of forest fragmentation on Amazonian ant communities. Journal of Biogeography, 33: 1348-1356. doi: 10.1111/j.1365- 2699.2006.01516.x Vellend, M. (2010). Conceptual synthesis in community ecology. The Quarterly Review of Biology, 85: 183-206. doi: 10.1086/652373 Vellend, M. (2016). The theory of ecological communities. Princeton University Press. doi: 10.1515/9781400883790 Wilson, D.S. (1992). Complex interactions in metacommunities, with implications for biodiversity and higher levels of selection. Ecology, 73: 1984-2000. doi: 10.2307/1941449. https://doi.org/10.2307/1941449 Sociobiology 69(3): e8099 (September, 2022) 9 Supplementary Material Table S1. Area size in hectare and percentage of total area (in parentheses) showing changes in land uses in Viçosa municipality -MG, Brazil, classified in 1995, 1996 and 2010. Class / Year 1995 1996 2010 Forest 6874.023 (22.96%) 6844.791 (22.86%) 7782.602 (25.99%) Agriculture 22046.48 (73.64%) 22040.04 (73.61%) 20794.64 (69.45%) Unvegetated area 997.3359 (3.33%) 1033.261 (3.45%) 1340.006 (4.48%) Water 22.10562 (0.07%) 21.85455 (0.07%) 22.69341 (0.08%) Size total 29939.95 (100%) Source: MapBiomas Table S2. Proportional size in hectare of fragments area, number of traps installed and points sampled in Viçosa, state of Minas Gerais, Brazil. Geographic coordinates Fragment size Fragment size 1995 Fragment size 2010 Traps Points 20º48’22.2’’S/42º51’10.9’’W > 93 hectares 298.92 350.14 48 24 20º49’31.0’’S/42º55’44.5’’W > 93 hectares 93.37 88.52 48 24 20º45’24.2’’S/42º51’38.8’’W 30 to 61 hectares 60.52 79.57 24 12 20º47’29.5’’S/42º53’26.9’’W 30 to 61 hectares 46.16 51.12 24 12 20º46’17.6’’S/42º52’11.7’’W 30 to 61 hectares 39.21 100.47 24 12 20º47’56.9’’S/42º50’41.3’’W 30 to 61 hectares 30.13 36.90 24 12 20º44’34.0’’S/42º49’53.3’’W 3 to 9 hectares 8.57 9.24 8 4 20º49’17.0’’S/42º53’51.4’’W 3 to 9 hectares 4.05 4.72 8 4 20º48’55.9’’S/42º52’49.9’’W 3 to 9 hectares 3.15 9.98 8 4 20º49’07.3’’S/42º55’58.9’’W 3 to 9 hectares 4.99 4.15 8 4 20º46’36.2’’S/42º50’28.2’’W 3 to 9 hectares 8.00 6.58 8 4 20º43’20.2’’S/42º51’40.7’’W 3 to 9 hectares 7.72 7.19 8 4 20º43’30.9’’S/42º51’42.5’’W 3 to 9 hectares 7.97 5.66 8 4 20º43’20.3’’S/42º49’33.3’’W 3 to 9 hectares 6.91 5.09 8 4 20º42’50.7’’S/42º49’11.6’’W 3 to 9 hectares 5.56 6.44 8 4 20º47’08.8’’S/42º53’33.4’’W 3 to 9 hectares 4.21 6.12 8 4 20º48’52.7’’S/42º51’56.0’’W 3 to 9 hectares 5.77 2.32 8 4 Vanessa S. Ribeiro et al. – Divergence of ant communities over time across fragments10 Subfamily / species 1995 1996 2010 Dolichoderinae Dolichoderus attelaboides FS X X X Linepithema humile G X Linepithema iniquum FS X X X Linepithema leucomelas FS X X Linepithema sp.1 X Linepithema sp.2 X Linepithema sp.5 X X Linepithema sp.6 X Linepithema sp.7 X Linepithema sp.8 X Linepithema sp.9 X Linepithema sp.10 X Dorylinae Neivamyrmex alfaroi FS X X Ectatomminae Ectatomma edentatum G X X X Ectatomma permagnum FS X X X Ectatomma planidens OH X Ectatomma sp.3 X X X Ectatomma sp.4 X Ectatomma sp.5 X Gnamptogenys acuminata FS X Gnamptogenys horni FS X X X Gnamptogenys mina FS X X X Gnamptogenys regularis FS X Gnamptogenys striatula G X X X Gnamptogenys sp.3 X X Gnamptogenys sp.4 X X Gnamptogenys sp.8 X Gnamptogenys sp.9 X Formicinae Brachymyrmex depilis G X X Brachymyrmex sp.1 G X Brachymyrmex sp.2 G X Brachymyrmex sp.4 G X Brachymyrmex sp.5 G X Brachymyrmex sp.6 G X Brachymyrmex sp.8 G X Formicinae Camponotus atriceps G X X X Camponotus crassus OH X X Camponotus genatus G X X X Camponotus melanoticus G X X X Camponotus novogranadensis G X X Camponotus rufipes G X X X Camponotus sp.2 X X Camponotus sp.7 X X X Camponotus sp.10 X X X Nylanderia fulva G X X X Nylanderia sp.1 X X X Nylanderia sp.2 X Heteroponerinae Heteroponera dentinodis G X Heteroponera dolo FS X X X Myrmicinae Acanthognathus rudis G X X Acromyrmex niger G X X X Acromyrmex rugosus G X Acromyrmex subterraneus G X X X Acromyrmex sp.2 G X Acromyrmex sp.3 G X Acromyrmex sp.4 G X Acromyrmex sp.5 G X Acromyrmex sp.7 G X Acromyrmex sp.10 G X X Acromyrmex sp.15 G X Acromyrmex sp.21 G X Apterostigma jubatum FS X X X Apterostigma sp.3 FS X X X Apterostigma sp.4 FS X X X Apterostigma sp.7 FS X Atta sexdens OH X X X Basiceros disciger FS X Basiceros sp.1 FS X X X Cardiocondyla emeryi OH X X Cephalotes atratus FS X X X Cephalotes ustus X Table S3. Ant species sampled in the years of 1995, 1996 and 2010. The letter G represents the species classified as generalists, letters FS as forest specialist and letters OH those species classified as open-habitat specialist. Species that do not have any letter were unclassified. Subfamily / species 1995 1996 2010 Sociobiology 69(3): e8099 (September, 2022) 11 Myrmicinae Crematogaster distans G X Crematogaster nigropilosa G X Crematogaster ocuta G X Crematogaster sp.1 G X Crematogaster sp.5 G X Eurhopalothrix sp.1 FS X Hylomyrma balzani FS X X X Hylomyrma reitteri FS X X X Hylomyrma sp.1 X X X Megalomyrmex drifti FS X Megalomyrmex gnomus FS X Megalomyrmex goeldii FS X X X Megalomyrmex iheringi FS X X X Megalomyrmex incisus FS X X Megalomyrmex modestus FS X X Megalomyrmex pusillus FS X X Megalomyrmex sp.2 X Octostruma rugifera FS X X X Oxyepoecus plaumanni FS X Oxyepoecus reticulatus FS X X Procryptocerus convergens FS X Procryptocerus marginatus FS X X X Procryptocerus sp.2 X Rogeria ciliosa FS X Rogeria sp.1 X X X Rogeria sp.2 X Strumigenys apreciatta FS X X Trachymyrmex aspersus G X X Trachymyrmex oetkeri FS X Trachymyrmex sp.1 X X X Trachymyrmex sp.2 X X X Trachymyrmex sp.4 X Trachymyrmex sp.5 X Trachymyrmex sp.6 X Trachymyrmex sp.7 X Trachymyrmex sp.8 X Ponerinae Hypoponera foreli G X Hypoponera sp.1 X X X Hypoponera sp.2 X X X Hypoponera sp.3 X X Hypoponera sp.4 X Hypoponera sp.6 X X Leptogenys sp.1 FS X X X Neoponera lenis FS X X X Neoponera magnifica FS X X X Neoponera metanotalis FS X X X Neoponera venusta FS X Neoponera sp.4 X X Neoponera sp.6 X Neoponera sp.7 X X X Neoponera sp.8 X Odontomachus chelifer G X X X Odontomachus minutus FS X X X Odontomachus sp.2 X X Pachycondyla harpax FS X X X Pachycondyla impressa FS X X Pachycondyla striata FS X X X Pseudomyrmicinae Pseudomyrmex tenuis G X X Pseudomyrmex sp.1 X X X Pseudomyrmex sp.2 X X X Pseudomyrmex sp.3 X Table S3. Ant species sampled in the years of 1995, 1996 and 2010. The letter G represents the species classified as generalists, letters FS as forest specialist and letters OH those species classified as open-habitat specialist. (Continuation) Subfamily / species 1995 1996 2010Subfamily / species 1995 1996 2010 _Hlk106809476 _Hlk106809635 _Hlk106809976 _Hlk106726296 _Hlk106869915 _Hlk106883699 _Hlk106884382 _Hlk106884047 _Hlk63952304 _Hlk106810693 _Hlk106811609 move80695823 _Hlk106816192 _Hlk109056316 _Hlk110601015 _Hlk110601278 _Hlk108794325 _Hlk106816258 _Hlk106886543 _Hlk106816279 _Hlk109057517 _Hlk110602194 _Hlk47515972 _Hlk110602427