Ecoregions within the Brazilian Pampa biome reflected in Odonata species assemblies

Based on vegetation composition, previous studies of the Pampa biome in southern Brazil have defined seven ecoregions within the area. Here, we test this ecoregion approach studying the semi‐aquatic insect group Odonata in five of these regions, aiming at comparing the ecoregions to the more traditional environmental predictors of water quality and land cover. Based on a data set of occupancy comprising 99 species distributed between 131 localities, a one‐way Permutational Multivariate Analysis of Variance was used to compare differences in the species composition between the ecoregions, followed by a Principal Component Analysis to visualize the variation. The composition varied significantly between all groups tested, and the ordination explained 61.8% of the variance. A partial redundancy analysis of ecoregions, land cover and water quality variables explained 71% of the variance in Odonata community structure. Ecoregion was the most important predictor, followed by water quality and land cover. Within these species assemblies, we could select certain species that were representative of a given ecoregion, to which their distribution within the Pampa biome was entirely or mainly confined. Of 24 representative species 41.7% were rare, while the rest were more abundant and, hence, easier to detect. We suspect that the differences found between the Pampa ecoregions might be due to geology, as such factors may be strong determinants of biodiversity. Specific ecological requirements at the family and genus levels also seemed to act selectively on the species compositions within the ecoregions. Today, the Pampa is highly fragmented due to agricultural activities such as rice cultivation, extensive cattle farming and forest plantations. We suggest that an ecoregion‐based approach to the implementation of conservation measures may be the best way to help these distinct species assemblies survive.     

Do riparian forest strips in modified forest landscapes aid in conserving bat diversity?

Agricultural practices lead to losses of natural resources and biodiversity. Maintaining forests alongside streams (riparian forest strips) has been used as a mechanism to minimize the impact of clearing for agriculture on biodiversity. To test the contribution of riparian forest strips to conserve biodiversity in production landscapes, we selected bats as a biodiversity model system and examined two dimensions of diversity: taxonomic and functional. We compared bat diversity and composition in forest, with and without stream habitat, and in narrow forest riparian strips surrounded by areas cleared for agriculture. We tested the hypothesis that riparian forest strips provide potential conservation value by providing habitat and serving as movement corridors for forest bat species. Riparian forest strips maintained 75% of the bat species registered in forested habitats. We found assemblage in sites with riparian forest strips were dominated by a few species with high abundance and included several species with low abundance. Bat species assemblage was more similar between sites with streams than between those sites to forests without stream habitat. These results highlight the importance of stream habitat in predicting presence of bat species. We registered similar number of guilds between forest sites and riparian forest strips sites. Relative to matrix habitats, stream and edge habitats in riparian forest strips sites were functionally more diverse, supporting our hypothesis about the potential conservation value of riparian forest strips. Results from this study suggest that maintaining riparian forest strips within cleared areas for agricultural areas helps conserve the taxonomic and functional diversity of bats. Also, it provides basic data to evaluate the efficacy of maintaining these landscape features for mitigating impacts of agricultural development on biodiversity. However, we caution that riparian forest strips alone are not sufficient for biodiversity maintenance; their value depends on maintenance of larger forest areas in their vicinity.     

Diversity and distribution of aquatic insects in Southern Brazil wetlands: implications for biodiversity conservation in a Neotropical region

The selection of priority areas is an enormous challenge for biodiversity conservation. Some biogeographic methods have been used to identify the priority areas to conservation, and panbiogeography is one of them. This study aimed at the utilization of panbiogeographic tools, to identify the distribution patterns of aquatic insect genera, in wetland systems of an extensive area in the Neotropical region (approximately 280 000km2), and to compare the distribution of the biogeographic units identified by the aquatic insects, with the conservation units of Southern Brazil. We analyzed the distribution pattern of 82 genera distributed in four orders of aquatic insects (Diptera, Odonata, Ephemeroptera and Trichoptera) in Southern Brazil wetlands. Therefore, 32 biogeographic nodes corresponded to the priority areas for conservation of the aquatic insect diversity. Among this total, 13 were located in the Atlantic Rainforest, 16 in the Pampa and three amongst both biomes. The distribution of nodes showed that only 15% of the dispersion centers of insects were inserted in conservation units. The four priority areas pointed by node cluster criterion must be considered in further inclusions of areas for biodiversity conservation in Southern Brazil wetlands, since such areas present species from different ancestral biota. The inclusion of such areas into the conservation units would be a strong way to conserve the aquatic biodiversity in this region.     

Conservation strategies for Araucaria Forests in Southern Brazil: assessing current and alternative approaches

Efforts to protect the remaining Araucaria Forest fragments in Southern Brazil have focused on large forests dominated by the threatened species Araucaria angustifolia. However, such an approach can miss significant biodiversity by ignoring smaller forest fragments scattered throughout the landscape, many of which exist on small farms. Here, I evaluate the method used to define a set of natural reserves in Southern Brazil to assess if it captures and preserves the significant biodiversity that exists in this unique biome. I used a relatively simple field sampling methodology and statistical analyses to evaluate floristic and structural parameters, comparing tree species diversity within a large forest fragment (with and without A. angustifolia dominance) to smaller fragments in the surrounding landscape. While stands dominated by A. angustifolia showed primary forest‐like physiognomy, they did not have the levels of diversity expected for old‐growth forests. Traditional forestry parameters, especially basal area and number of individuals, are useful for differentiating forest types in terms of structure. While no single diversity index provides a complete picture of the composition and variation of diversity in forest fragments, when taken together and assessed using extrapolation and comparative statistics, they provide a meaningful method for comparing and identifying high priority areas for conservation. Future conservation strategies should look to alternative approaches for assessing biodiversity across the landscape, while including small‐scale landowners in conservation efforts.     

Structural implications of traditional agricultural landscapes on the functional diversity of birds near the Korean Demilitarized Zone

Bird assemblages are sensitive to changes in landscape composition and the environment, such as those that result from drought. In this study, the relationship between landscape composition and avian functional diversity in traditional agricultural ecosystems in the Civilian Control Zone (CCZ) of Korea was examined. In addition, the resilience of biodiversity to changes in landscape elements resulting from drought conditions was investigated. The traditional agricultural landscape (TAL) of the sites studied was divided into three types: TAL 1 had a high proportion of rice paddies, TAL 2 included large forest areas, and TAL 3 represented areas with drylands. Of these, TAL 1 showed the highest species richness and functional richness, but these measures were most vulnerable to drought. Meanwhile, TAL 2 showed that the bird communities were more tolerant under drought event. This study shows that to conserve and enhance the diversity of birds in traditional agricultural landscapes of Northeast Asia, active management of forest areas is needed to protect bird populations. In addition, commercial pressures to develop this area will require urgent biodiversity conservation plans to protect the unique biodiversity of the Korean CCZ. This study thus provides landscape management guidance for conservation planning.     

Patterns in the organization of Cerrado pond biodiversity in Brazilian pasture landscapes

There is a worldwide concern on the loss of pond biodiversity in human dominated landscapes. Nevertheless, agricultural activities appear to increase pond number in the Brazilian Cerrado through damming streams for cattle raising. These man-made ponds may represent important landscape features, but their importance to regional biodiversity has not yet been studied. Here, we evaluated differences in alpha and beta diversity under a multi-taxonomic approach, as well as tested pond size as the main driver of local species richness. We also assessed the importance of environmental heterogeneity through the analysis of the regional species accumulation curves (SAC). The overall result suggests that species turnover was the major component of regional biodiversity for all groups. Major physical and chemical water conditions had no effects on algae, macrophytes, water bugs, and birds species richness. Pond size had a significant effect on Odonata and fish species richness, while water beetles and amphibians were influenced by trophic conditions. Results from regional SAC show variations among different taxonomic groups regarding landscape heterogeneity: only algae, fish, and birds do not reached to an asymptote and had higher z-values. Our results highlight the importance of ponds for biodiversity conservation in increasingly agricultural landscapes in central Brazil.     

Changes in Diversity, Abundance, and Structure of Soil Bacterial Communities in Brazilian Savanna Under Different Land Use Systems

The Brazilian Savanna, also known as “Cerrado”, is the richest and most diverse savanna in the world and has been ranked as one of the main hotspots of biodiversity. The Cerrado is a representative biome in Central Brazil and the second largest biome in species diversity of South America. Nevertheless, large areas of native vegetation have been converted to agricultural land including grain production, livestock, and forestry. In this view, understanding how land use affects microbial communities is fundamental for the sustainable management of agricultural ecosystems. The aim of this work was to analyze and compare the soil bacterial communities from the Brazilian Cerrado associated with different land use systems using high throughput pyrosequencing of 16S rRNA genes. Relevant differences were observed in the abundance and structure of bacterial communities in soils under different land use systems. On the other hand, the diversity of bacterial communities was not relevantly changed among the sites studied. Land use systems had also an important impact on specific bacterial groups in soil, which might change the soil function and the ecological processes. Acidobacteria, Proteobacteria, and Actinobacteria were the most abundant groups in the Brazilian Cerrado. These findings suggest that more important than analyzing the general diversity is to analyze the composition of the communities. Since soil type was the same among the sites, we might assume that land use was the main factor defining the abundance and structure of bacterial communities.     

Odonate species occupancy frequency distribution and abundance–occupancy relationship patterns in temporal and permanent water bodies in a subtropical area

This paper investigates species richness and species occupancy frequency distributions (SOFD) as well as patterns of abundance–occupancy relationship (SAOR) in Odonata (dragonflies and damselflies) in a subtropical area. A total of 82 species and 1983 individuals were noted from 73 permanent and temporal water bodies (lakes and ponds) in the Pampa biome in southern Brazil. Odonate species occupancy ranged from 1 to 54. There were few widely distributed generalist species and several specialist species with a restricted distribution. About 70% of the species occurred in <10% of the water bodies, yielding a surprisingly high number of rare species, often making up the majority of the communities. No difference in species richness was found between temporal and permanent water bodies. Both temporal and permanent water bodies had odonate assemblages that fitted best with the unimodal satellite SOFD pattern. It seems that unimodal satellite SOFD pattern frequently occurred in the aquatic habitats. The SAOR pattern was positive and did not differ between permanent and temporal water bodies. Our results are consistent with a niche‐based model rather than a metapopulation dynamic model.     

The impacts of habitat disturbance on adult and larval dragonflies (Odonata) in rainforest streams in Sabah,Malaysian Borneo

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Conserving the Brazilian semiarid (Caatinga) biome under climate change

The Caatinga is a semiarid biome of the northeast of Brazil with only 1?% of its territory currently conserved. The biome’s biodiversity is highly threatened due to exposure to land conversion for agricultural and cattle ranch. Climate forecasts predict increases in aridity, which could pose additional threats to the biome’s biodiversity. Here, we ask if the remnants of natural vegetation in Caatinga biome, where endemic terrestrial vertebrate species occur, are likely to retain more climatic suitability under climate change scenarios than other less pristine areas of the biome. In order to assess changes in climate suitability across individual species ranges, ensemble forecasting was used based on seven bioclimatic envelope models, three atmosphere–ocean general circulation models, and two greenhouse emission gas scenarios for 2020, 2050, and 2080. We found that most species will gain climatic suitability in the natural vegetation remnants of the Caatinga. Such gains are even greater than the expected to occur within random sets of areas with size similar to the natural vegetation remnants. Our results suggest that natural vegetation remnants will likely play a role of climate refuges for endemic vertebrate species, so efforts should be concentrated in these regions.     

Comparative effects of urban and agricultural land transformation on Odonata assemblages in a biodiversity hotspot

Rivers of the Cape Floristic Region (CFR) biodiversity hotspot are threatened by land transformation. This region is a centre of endemism for many taxa, including Odonata. These insects are highly sensitive to changes in physical habitat structure, which makes them good bioindicators, and this led to the development of the Dragonfly Biotic Index (DBI). We investigated the effects of local agricultural and urban land transformations on Odonata species richness, assemblage composition and DBI scores in three CFR rivers. A total of 48 sites were selected and categorized as natural, agricultural or urban land use. Adult male Odonata and four environmental variables were recorded over two seasons. Land transformation significantly influenced Odonata assemblage composition but did not always significantly reduce species richness. Average vegetation height also affected Odonata assemblage composition and decreased species richness. Agricultural and urban sites had Odonata assemblages differing from those in the natural areas. Agricultural and urban local land use types reduced opportunities for some endemic species but provided for the persistence and establishment of widespread, generalist species, as indicated by great changes in DBI scores. Mitigating the adverse influences of land transformation through establishment of protected areas is essential for the conservation of rare taxa, particularly in an area with a high number of endemic species.     

Do riparian reserves support dung beetle biodiversity and ecosystem services in oil palm-dominated tropical landscapes?

Agricultural expansion and intensification are major threats to global biodiversity, ecological functions, and ecosystem services. The rapid expansion of oil palm in forested tropical landscapes is of particular concern given their high biodiversity. Identifying management approaches that maintain native species and associated ecological processes within oil palm plantations is therefore a priority. Riparian reserves are strips of forest retained alongside rivers in cultivated areas, primarily for their positive hydrological impact. However, they can also support a range of forest‐dependent species or ecosystem services. We surveyed communities of dung beetles and measured dung removal activity in an oil palm‐dominated landscape in Sabah, Malaysian Borneo. The species richness, diversity, and functional group richness of dung beetles in riparian reserves were significantly higher than in oil palm, but lower than in adjacent logged forests. The community composition of the riparian reserves was more similar to logged forest than oil palm. Despite the pronounced differences in biodiversity, we did not find significant differences in dung removal rates among land uses. We also found no evidence that riparian reserves enhance dung removal rates within surrounding oil palm. These results contrast previous studies showing positive relationships between dung beetle species richness and dung removal in tropical forests. We found weak but significant positive relationships between riparian reserve width and dung beetle diversity, and between reserve vegetation complexity and dung beetle abundance, suggesting that these features may increase the conservation value of riparian reserves. Synthesis and applications: The similarity between riparian reserves and logged forest demonstrates that retaining riparian reserves increases biodiversity within oil palm landscapes. However, the lack of correlation between dung beetle community characteristics and dung removal highlights the need for further research into spatial variation in biodiversity–ecosystem function relationships and how the results of such studies are affected by methodological choices.     

Fruit‐feeding Butterfly Communities are Influenced by Restoration Age in Tropical Forests

Currently, a large‐scale restoration project aims to restore around 15 million hectares of Atlantic Forest in Brazil. This will increase forest cover and connectivity among remnant sites as well as restore environmental services. Currently, studies on recovery of fauna in restored areas of the Atlantic Forest are practically nonexistent. To address this knowledge vacuum, our study compares diversity patterns of fruit‐feeding butterflies in three forest areas with different restoration ages (11, 22, and 54 years), and uses a native forest area as reference. Results showed butterfly communities in maturing restored areas becoming more similar to the ones found in the native forest, with an increase in the proportional abundance of forest species, and a decrease of edge and grassland species. Moreover, we found a higher diversity among sites at the intermediate restoration age, with a community composed of both grassland and forest species. Butterfly species composition differed significantly among sites, showing interesting patterns of potential species replacement over time. Our results indicate that, although restored sites were located in a fragmented landscape, they provide suitable habitats for recolonization by fruit‐feeding butterfly assemblages. Hence, restored areas can be considered important habitat for forest animal species, increasing local biodiversity and, possibly, restoring some of the ecosystem services provided by them.     

Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats

Agricultural landscapes can be defined as mosaics of landscape elements which are affected by farming practices. Woodland habitats, even though they are managed, are amongst the most stable elements of agricultural landscapes and can play a key role in the maintenance of biodiversity. This study of the ground beetle (carabid) communities of woodlands and woody linear features in a Scottish agricultural landscape shows that these habitats contribute significantly to the overall landscape diversity of these beetles. Communities in woods and hedgerows display the same species diversity and are both characterized by the presence of forest species. The main factors constraining carabid communities in both environments are the grazing intensity and, to a lesser extent, the type of soil. Heavily grazed locations are characterized by the occurrence of grassland species while forest species are restricted to ungrazed locations. At the landscape scale, the distribution of the forest species is limited by spatial isolation, indicating that there are insufficient functional links between woodland habitats in the study area. Isolation could be compensated for either by a better control of grazing so that linear features can be used as dispersal corridors for forest carabids or by planting more linear features and woods in the area.     

Bellwether of the Canaries: anthropogenic effects on the land snail fauna of the Canary Islands

Natural areas near human-modified landscapes experience factors that may affect local biodiversity at levels commensurate with natural environmental factors. The land snails of the Canary Islands provide excellent opportunities to evaluate the importance of anthropogenic agents in mediating the diversity and distribution of species. Land snails are particularly sensitive to disturbance and are an integral part of terrestrial ecosystems. This study analyzed the distributions and abundances of terrestrial macrosnail shell assemblages at 60 localities throughout the coastal scrub biome of the Canary Islands. This was accomplished using data on natural and anthropogenic variables to assess their relative importance in governing land snail diversity. A total of 34,801 dead shells represented a diverse malacofauna with highly localized endemism. Due to uncertain species identifications, samples from the 18 sites from the two easternmost islands are described, but excluded from statistical analyses. Regression tree analysis indicated that proximity to agricultural sites was the most important predictor of species diversity. Sites with no or very little agricultural area (≤ 0.167 km²) within a 1 km radius had significantly higher richness and diversity. These results have implications for Canary Islands conservation. Protected areas that are patchworks of natural and agricultural landscapes are still subject to native biodiversity loss because of anthropogenic impacts even when the footprint of agriculture is small.     

Dung beetle communities as biological indicators of riparian forest widths in southern Brazil

Riparian forests provide important habitat for many wildlife species and are sensitive to landscape change. Among terrestrial invertebrates, dung beetles have been used to investigate the effects of environmental disturbances on forest structure and diversity. Since many studies demonstrated a negative response of dung beetle communities to increasing forest fragmentation, and that most dung beetle species had a more pronounced occurrence during warmest seasons, three hypotheses were tested: (1) Scarabaeinae richness, abundance, diversity and evenness are lower in thinner riparian zone widths than in wider widths during the warmest seasons; (2) Scarabaeinae richness and abundance are positively influenced by leaf litter coverage and height and canopy cover; and (3) Scarabaeinae composition varies with the reduction in riparian vegetation and among annual seasons. We selected four fragments with different riparian zone widths in three secondary streams in southern Brazil. In each fragment, four sampling periods were carried out seasonally between spring 2010 and winter 2011. We collected dung beetles using pitfall traps with two types of bait. We collected 1289 specimens distributed among 29 species. In spring and summer, dung beetle richness was higher in fragments with the widest riparian zone than in those with a thinner riparian zone, and it did not vary between fragments in fall and winter seasons. Dung beetle abundance did not differ among fragments with different riparian zone widths, but it was higher in spring and summer than fall and winter. Richness and abundance were positively influenced by leaf litter. While dung beetle diversity was higher in fragments with wider riparian zone widths than in those with thinner widths, the evenness was similar among fragments. Dung beetle composition differed between the fragments with the widest and thinnest riparian zones, and it also varied among the seasons. Our results suggest that decreased riparian zones affect negatively to dung beetle community structure in southern Brazil. Fragments with thinner riparian zones had lower beetle richness in warmest seasons and an altered community composition. In this sense, the dung beetles are potentially good indicators of riparian forest fragmentation since some species were indicators of a particular riparian zone width. From a conservation perspective, our results demonstrate that the new Brazilian Forest Code will greatly jeopardize not only the terrestrial and aquatic biodiversity of these ecosystems, but also countless other ecological functions.     

How may agricultural matrix intensification affect understory birds in an Atlantic Forest landscape? A qualitative model on stochasticity and immigration

Understanding how different approaches of matrix management affect organisms that inhabit natural patches is crucial for biological conservation. Considering that great part of the tropical area is composed of agricultural land and that most of the land use is either intensified or on the verge to be, a relevant question is: how may agricultural intensification of the landscape matrix affect the population dynamics of understory birds? This paper describes a qualitative model based on the Qualitative Process Theory and implemented in Garp3 to provide answers to this question. We built a model using four species of endemic passerine birds in order to evaluate the case in an Atlantic Forest area, a biodiversity hotspot. The model describes a landscape composed of an extinction-resistant source patch and one target forest patch where stochastic events occur. If permeability of the matrix exceeds a given species-specific threshold, propagules coming from the source reach the target patch. Agriculture intensification affects the matrix spatial structure and reduces permeability to forest birds, thus reduces rescue effect. Additionally, we assume that some species may use the matrix as a supplementary habitat for feeding and that matrix management may affect the resource base for those species. Simulation results suggest that, if agriculture intensification continues to threat the Atlantic Forest biome, populations of sensitive species (Xiphorhynchus fuscus and Sclerurus scansor) that exist in the small forest patches (which is true for most forest remnants) will be highly susceptible to local extinctions and without further re-colonization.     

Forest cover and heterogeneous pastures shape the diversity of predatory rove beetles in tropical riparian habitats

In tropical landscapes, forest remnants have been reduced to narrow strips of vegetation along rivers and streams surrounded by agricultural land that affects biodiversity, depending on the habitat and landscape characteristics. To assess the effect of riparian forest loss on the diversity of Staphylininae predatory rove beetles, we considered two habitat conditions (river sites with riparian vegetation and sites with heterogeneous pastures) within two micro-basin types (with >70% and <40% forest cover) in a tropical montane cloud forest landscape, Mexico. Beetles were collected using baited pitfall traps during the rainy season of 2014. No differences were found between micro-basin types and, although species richness (⁰D) was similar between habitat conditions, when the diversity of common (¹D) and dominant (²D) species was considered, sites with heterogeneous pastures were almost twice as diverse as those with riparian vegetation. All diversity measurements were greater in sites with heterogeneous pastures of either micro-basin type. Air temperature and canopy cover were the environmental variables that best explained the variation in beetle species composition. The greatest environmental differences related to species composition were detected between habitat conditions and were more evident in sites with heterogeneous pastures and low forest cover in the surroundings. The results suggest that replacing riparian vegetation with heterogeneous pastures, within micro-basins that lost between 30% and 60% of their forest cover, does not significantly reduce the diversity of predatory rove beetle but rather modifies the beetle composition. Effective formulation of management strategies to mitigate the impact of land use modification therefore requires an understanding of the interaction between vegetation remnants and landscape characteristics.     

Population Dynamics of Drosophilids in the Pampa Biome in Response to Temperature

Environmental variables such as temperature and rainfall can directly affect the community structure of dipterans. Seasonal oscillations in the abundance of species of Drosophilidae reflect differences in how tolerant populations are to climatic conditions. Over a period of 14 months, we collected samples in two habitats in the Pampa biome in the municipality of São Luiz Gonzaga, state of Rio Grande do Sul, Brazil (28°24′28″S, 54°57′39″W). The influence of environmental variables on populations of Drosophilidae was evaluated for both collecting sites by using correlation analysis. The results suggested a negative correlation between the abundances of Drosophila cardinoides Dobzhansky & Pavan, Drosophila maculifrons Duda, Drosophila melanogaster Meigen, Drosophila nigricruria Patterson & Mainland, and Zygothrica vittimaculosa Burla with temperature, which is reflected in the distribution of these species within Brazil. Our findings are important for characterizing and preserving biodiversity in this almost-unknown biome in southern Brazil given the current climate change scenario.     

Environmental drivers of species composition and functional diversity of dung beetles along the Atlantic Forest–Pampa transition zone

Human activities are causing a rapid loss of biodiversity, which impairs ecosystem functions and services. Therefore, understanding which processes shape how biodiversity is distributed along spatial and environmental gradients is a first step to guide conservation and management efforts. We aimed to determine the relative explanatory importance of biogeographic, environmental, landscape and spatial variables on assemblage dissimilarities and functional diversity of dung beetles along the Atlantic Forest–Pampa (i.e. forest–grassland) transition zone located in Southeast South America. We described each site according to their biogeographic position, environmental conditions, landscape features and spatial patterns. The compositional dissimilarity was partitioned into turnover and nestedness components of β‐diversity. Mantel tests and generalised dissimilarity models were used to relate β‐diversity and its components to biogeographic, environmental, landscape and spatial variables. Variation partitioning analysis was used to estimate the pure and shared variation in species composition and functional diversity explained by the four categories of predictors. Biome domain was the main factor causing dung beetle compositional dissimilarity, with a high species replacement between Atlantic Forest and Pampa. Biogeographic, environmental, landscape and spatial distances also affected the patterns of dung beetle dissimilarity and β‐diversity components. The shared effects of the four sets of predictors explained most of the variation in dung beetle composition. A similar response pattern was found for dung beetle functional diversity, which excluded biogeographic effects. Only the pure effects of environmental and spatial predictors were significant for species composition and functional diversity. Our results indicate that dung beetle species composition and functional diversity are jointly driven by environmental, landscape and spatial predictors with higher pure environmental and spatial effects. The forest–grassland transition zone promotes a strong species and trait replacement highly influenced both by environmental filtering and dispersal limitation.