Functional and phylogenetic structure of forest and savanna bird assemblages across spatial scales

Ecological and evolutionary mechanisms that drive community assembly vary in space and time. However, little is known about how such mechanisms act in contrasting habitats. Here, we estimated the functional and phylogenetic structure of forest and savanna bird assemblages across different spatial scales to understand: 1) the mechanisms that govern the structure of assemblages in these habitats; 2) the relationship between phylogenetic and functional structure; and 3) the influence of species richness on the functional and phylogenetic structure of assemblages. We used a null model where forest and savanna bird species were allowed to occur in the same null assemblages and other where species were separated based on their habitats. According to the first null model, forest bird assemblages were functionally and phylogenetically clustered at all spatial scales, whereas savanna bird assemblages generally showed random functional and phylogenetic structure. These results can be explained by the low dispersal rate of forest species across of the patchy habitats and the widespread distribution of savanna species. However, in the second null model, both forest and savanna bird assemblages showed random functional and phylogenetic structure at regional and local scales. This suggests that trait‐based assembly might not play an important role in both habitats and across different spatial scales. In addition, the phylogenetic and functional structure of assemblages were not correlated, evidencing that caution is necessary when using phylogenetic relationships as a surrogate to functional distances among species. Finally, the relationships between species richness and functional and phylogenetic structure indicated that an increase in the number of species can promote both clustering and overdispersion, depending on the studied habitat and scale. Our study shows that integrating different types of habitat, spatial scales and biodiversity components in a single framework can shed light on the mechanisms that determine the community assembly.     

A phylogenetic and trait‐based analysis of community assembly in a subtropical forest in central China

Despite several decades of study in community ecology, the relative importance of the ecological processes that determine species co‐occurrence across spatial scales remains uncertain. Some of this uncertainty may be reduced by studying the scale dependency of community assembly in the light of environmental variation. Phylogenetic information and functional trait information are often used to provide potentially valuable insights into the drivers of community assembly. Here, we combined phylogenetic and trait‐based tests to gain insights into community processes at four spatial scales in a large stem‐mapped subtropical forest dynamics plot in central China. We found that all of the six leaf economic traits measured in this study had weak, but significant, phylogenetic signal. Nonrandom phylogenetic and trait‐based patterns associated with topographic variables indicate that deterministic processes tend to dominate community assembly in this plot. Specifically, we found that, on average, co‐occurring species were more phylogenetically and functionally similar than expected throughout the plot at most spatial scales and assemblages of less similar than expected species could only be found on finer spatial scales. In sum, our results suggest that the trait‐based effects on community assembly change with spatial scale in a predictable manner and the association of these patterns with topographic variables, indicates the importance of deterministic processes in community assembly relatively to random processes.     

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co‐occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within‐community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.     

Phylogenetic and morphological relationships between nonvolant small mammals reveal assembly processes at different spatial scales

The relative roles of historical processes, environmental filtering, and ecological interactions in the organization of species assemblages vary depending on the spatial scale. We evaluated the phylogenetic and morphological relationships between species and individuals (i.e., inter‐ and intraspecific variability) of Neotropical nonvolant small mammals coexisting in grassland‐forest ecotones, in landscapes and in regions, that is, three different scales. We used a phylogenetic tree to infer evolutionary relationships, and morphological traits as indicators of performance and niche similarities between species and individuals. Subsequently, we applied phylogenetic and morphologic indexes of diversity and distance between species to evaluate small mammal assemblage structures on the three scales. The results indicated a repulsion pattern near forest edges, showing that phylogenetically similar species coexisted less often than expected by chance. The strategies for niche differentiation might explain the phylogenetic repulsion observed at the edge. Phylogenetic and morphological clustering in the grassland and at the forest interior indicated the coexistence of closely related and ecologically similar species and individuals. Coexistence patterns were similar whether species‐trait values or individual values were used. At the landscape and regional scales, assemblages showed a predominant pattern of phylogenetic and morphological clustering. Environmental filters influenced the coexistence patterns at three scales, showing the importance of phylogenetically conserved ecological tolerances in enabling taxa co‐occurrence. Evidence of phylogenetic repulsion in one region indicated that other processes beyond environmental filtering are important for community assembly at broad scales. Finally, ecological interactions and environmental filtering seemed important at the local scale, while environmental filtering and historical colonization seemed important for community assembly at broader scales.     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

Australian Tropical and Subtropical Rain Forest Community Assembly: Phylogeny,Functional Biogeography,and Environmental Gradients

To compare community assemblage patterns in tropical northeastern and subtropical central eastern Australia across selected gradients and scales, we tested the relationship of species traits with phylogenetic structure, and niche breadth. We considered phylogenetic relationships across current‐day species in assemblages in relation to rain forest species pool sizes, and trait values along gradients including elevation and latitude. Trait values were quantified across scales for seed size, leaf area, wood density and maximum height at maturity for 1137 species and 596 assemblages using trait gradient analysis (TGA). Local assemblages of subtropical species had narrower trait ranges, and higher niche breadth values than corresponding assemblages of tropical species. Leaf size and seed size increased at low latitudes, and community phylogenetic structure was most strongly correlated with seed traits in the subtropics, reflecting dispersal and re‐colonization processes. Elevation accounted for little of the variance in community phylogenetic structure or trait variation across local and regional scales. Stable moist forest areas retained many species from ancestral rain forest lineages across a range of temporally conserved habitats; species within assemblages were less related; and rain forest assemblages had higher functional diversity, but lower niche breadth. This suggests that on average, assemblages of species in stable areas had greater trait variation and narrower distributions. Historic and recent rain forest contraction and re‐expansion can result in recolonized areas that are dominated by species that are more related (phylogenetically) than by chance, have smaller, widely dispersed seeds, and greater niche breadth (broader distributions).     

The assembly of tropical tree communities – the advances and shortcomings of phylogenetic and functional trait analyses

Tropical tree communities present one of the most challenging systems for studying the processes underlying community assembly. Most community assembly hypotheses consider the relative importance of the ecological similarity of co‐occurring species. Quantifying this similarity is a daunting and potentially impossible task in species‐rich assemblages. During the past decade tropical tree ecologists have increasingly utilized phylogenetic trees and functional traits to estimate the ecological similarity of species in order to test mechanistic community assembly hypotheses. A large amount of work has resulted with many important advances having been made along the way. That said, there are still many outstanding challenges facing those utilizing phylogenetic and functional trait approaches to study community assembly. Here I review the conceptual background, major advances and major remaining challenges in phylogenetic‐ and trait‐based approaches to community ecology with a specific focus on tropical trees. I argue that both approaches tremendously improve our understanding of tropical tree community ecology, but neither approach has fully reached its potential thus far.     

The Nature of Seedling Assemblages in a Fragmented Tropical Landscape: Implications for Forest Regeneration

Whether aging forest fragments are able to recover original assemblages or progressively move toward impoverished successional stages remains as an open question. This study tested the hypothesis that seedling assemblages in forest fragments differ from those across mature forest stands and examined to what extent the uncovered patterns supported the notion that edge‐affected habitats tend to support impoverished tree assemblages dominated by pioneer species. We contrasted a series of small forest remnants (3–91 ha) to old‐growth stands located in the largest (ca 3500 ha) and best preserved forest remnant in northeastern Brazil and found that tree seedling assemblages inhabiting forest fragments exhibited reduced species richness (up to 50%) at different spatial scales in comparison to seedling assemblages in mature forest and adult assemblages in both fragments and mature forest stands. Moreover, ordination analyses clearly segregated fragment seedling assemblages in taxonomic/functional terms and segregation correlated to the richness of pioneer species. Seedlings of pioneer species and those bearing medium‐sized seeds (0.6–1.5 cm) increased in fragments, whereas large‐seeded species (1.5–3.0 cm) were reduced by more than a half. Such a multiple‐scale replacement of the old‐growth flora by pioneers was also confirmed by an indicator species analysis and the resulting pioneer indicator species. Our results suggest that small forest fragments support impoverished and distorted seedling assemblages. This floristic/functional drift implies that forest remnants or edge‐affected habitats tend to be dominated by a small set of pioneer tree species rather than supporting a substantial portion of the old‐growth flora as do mature forest stands.     

Edge-Related Loss of Tree Phylogenetic Diversity in the Severely Fragmented Brazilian Atlantic Forest

Deforestation and forest fragmentation are known major causes of nonrandom extinction, but there is no information about their impact on the phylogenetic diversity of the remaining species assemblages. Using a large vegetation dataset from an old hyper-fragmented landscape in the Brazilian Atlantic rainforest we assess whether the local extirpation of tree species and functional impoverishment of tree assemblages reduce the phylogenetic diversity of the remaining tree assemblages. We detected a significant loss of tree phylogenetic diversity in forest edges, but not in core areas of small (<80 ha) forest fragments. This was attributed to a reduction of 11% in the average phylogenetic distance between any two randomly chosen individuals from forest edges; an increase of 17% in the average phylogenetic distance to closest non-conspecific relative for each individual in forest edges; and to the potential manifestation of late edge effects in the core areas of small forest remnants. We found no evidence supporting fragmentation-induced phylogenetic clustering or evenness. This could be explained by the low phylogenetic conservatism of key life-history traits corresponding to vulnerable species. Edge effects must be reduced to effectively protect tree phylogenetic diversity in the severely fragmented Brazilian Atlantic forest.     

Mosquitoes of field and forest: the scale of habitat segregation in a diverse mosquito assemblage

Knowledge of the distribution of arthropod vectors across a landscape is important in determining the risk for vector‐borne disease. This has been well explored for ticks, but not for mosquitoes, despite their importance in the transmission of a variety of pathogens. This study examined the importance of habitat, habitat edges, and the scale at which mosquito abundance and diversity vary in a rural landscape by trapping along transects from grassland areas into forest patches. Significant patterns of vector diversity and distinct mosquito assemblages across habitats were found. The scale of individual species' responses to habitat edges was often dramatic, with several species rarely straying even 10 m from the edge. The present results suggest blood‐seeking mosquito species are faithful to certain habitats, which has consequences for patterns of vector diversity and risk for pathogen transmission. This implies that analysts of risk for pathogen transmission and foci of control, and developers of land management strategies should assess habitat at a finer scale than previously considered.     

Using functional traits to investigate the determinants of crustacean zooplankton community structure

Understanding the various processes contributing to community assembly is among the central aims of ecology. As a means of exploring this topic we quantified the relative influences of habitat filtering and competition in establishing patterns of community functional trait diversity across a landscape of lakes. Habitat filtering has been invoked in shaping community structure when co‐occurring taxa are more similar in their traits than expected by chance (under‐dispersion), and competition has been inferred as a structuring agent when co‐occurring taxa are less similar (over‐dispersion). We tested these hypotheses in crustacean zooplankton communities using a functional trait‐based approach based on five traits defining zooplankton feeding and habitat preferences across 51 lakes spanning several large limnological gradients. In general, zooplankton communities were functionally less diverse than random assemblages created from the same regional species pool. Furthermore, functional diversity was strongly correlated with variables related to lake productivity, suggesting that access to resources was the chief habitat filtering process constraining zooplankton functional diversity. This pattern was driven by the predominantly herbivorous cladocerans as opposed to the more commonly omnivorous, and sometimes carnivorous, copepods.     

The Seedling Community of Tropical Rain Forest Edges and Its Interaction with Herbivores and Pathogens1

Edge effects alter biotic interactions and forest regeneration. We investigated whether edge creation affected the seedling community and its interactions with herbivores and leaf‐fungal pathogens. In forest edges and interior sites in Chiapas, Mexico, we counted all woody seedlings and species (10–100 cm tall) present in 1‐m² plots, measured their size (height and leaf number), and examined them for the occurrence of herbivory and pathogen damage. We investigated relationships between levels of damage and size, species richness and density. Species composition and biotic damage varied greatly among sites and habitats (edge vs interior). Late‐successional species dominated the community, but richness was lower in interior sites and species similarity was greater among edges than among interiors. Nearly all species (95%) present at edges and interiors showed herbivory damage, whereas 76 percent of the species in edge plots and 68 percent in interior plots showed pathogen damage. Although leaf area damaged by herbivores was similar between habitats (average 9.2%), pathogen damage was three times greater in edge plots (1.85%) than in interior plots (0.57%). Size was positively related to biotic damage at both habitats. Relationships between herbivory and pathogen damage and between pathogen damage and leaf number were significant only for edges. Biotic damage was not related to density or species richness. Overall, plant community structure was similar between habitats, but biotic damage was enhanced at edges. Thus, disease spread at edges may arise as a threat to tropical rain forest vegetation.     

Seasonal dynamics of waterbird assembly mechanisms revealed by patterns in phylogenetic and functional diversity in a subtropical wetland

Despite growing interest in using phylogenetic and functional methods to understand community assembly, few studies have examined how these methods can be used to assess seasonal variation in assembly mechanisms among migrant species. Migration can rapidly alter the relative influence of stochastic processes, species interactions, or environmental factors in shaping communities across seasons. Here, we describe seasonal dynamics in the phylogenetic and functional diversity of waterbirds in Mai Po Wetland, a subtropical region with significant and predictable temporal variation in climate and migratory bird density. Phylogenetic α diversity varied seasonally, exhibiting a clustered structure (indicative of environmental filtering) in summer, and over‐dispersed structure (indicative of biotic filtering) in winter. Phylogenetic diversity in spring and autumn exhibited a more intermediate, random structure, consistent with stochastic arrivals and departures of migrants. Functional diversity was clustered in spring but showed over‐dispersion in the other three seasons. Phylogenetic β diversity in summer and winter assemblages was characterized by two distinct groups, while spring and autumn assemblages were mixed. Our results suggest that waterbird assemblages were primarily shaped by interspecific competition in winter, while random processes tended to shape assemblages in spring and fall. Environmental factors played a more important role in summer, during periods of high heat stress. In addition, species co‐occurrence patterns were significantly more strongly related to phylogenetic similarity in winter than in summer. Our results suggest that the relative importance of assemblage mechanisms can vary seasonally in response to changing environmental conditions, suggesting that studies attempting to infer a single dominant assembly mechanism may ignore important assembly processes. Temporal shifts in assembly mechanisms may play an important role in maintaining diversity in subtropical and temperate wetlands and perhaps also in other dynamic systems.     

Co‐declining mammal–dung beetle faunas throughout the Atlantic Forest biome of South America

The millennial–scale evolutionary relationships between mammals and dung beetles have been eroded due to several drivers of contemporary biodiversity loss. Although some evidence of co‐decline has been shown for mammals and dung beetles at some Neotropical sites, a biome‐scale analysis for the entire Atlantic Forest of South America would strengthen our understanding of how relictual sets of mammal species can affect dung beetle co‐occurrences and co‐declines. We therefore collated hundreds of assemblages of both dung beetles and medium‐ to large‐bodied mammals throughout the world's longest tropical forest latitudinal gradient to examine to what extent mammal assemblages may exert a positive influence on dung beetle species composition and functional assembly, and whether this relationship is scale dependent. We also collated several climatic and other environmental variables to examine the degree to which they shape mammal–dung beetle relationships. The relationships between local mammal and dung beetle faunas were examined using regression models, variation partitioning, dissimilarity indices and ecological networks. We found a clear positive relationship between mammal and dung beetle species richness across this forest biome, indicating an ongoing process of mammal–dung beetle niche‐mediated co‐decline. We found a strong relationship between the species composition of both taxa, in which dung beetle species dissimilarity apparently track changes in mammalian dissimilarity, typically in 80% of all cases. Co‐variables such as phytomass and climatic variables also influenced mammal–dung beetle patterns of co‐decline along the Atlantic Forest. We conclude that dung beetle diversity and community assembly are shaped by the remaining co‐occurring mammal assemblages and their functional traits, and both groups were governed by environmental features. We emphasize that ecosystem‐wide effects of mammal population declines remain poorly understood both quantitatively and qualitatively, and curbing large vertebrate defaunation will ensure the persistence of co‐dependent species.     

Community assembly in temperate forest birds: habitat filtering,interspecific interactions and priority effects

Recent attempts to understand the processes governing community assembly have increasingly focused on patterns of phylogenetic relatedness and functional similarity among co-existing species. Considerations of the species pool, the number and identity of functional traits and the metrics used to identify patterns have come under scrutiny as possible influences on the detection of non-random patterns. Most mechanistic explanations of community assembly based on functional and phylogenetic approaches rely on deterministic explanations, while ignoring the role of stochastic processes and historical contingency, despite the prominent historical role of both types of explanations of species coexistence. We evaluated the phylogenetic and functional structure of 20 temperate forest bird assemblages in northeastern North America. We compared three approaches for characterizing the functional structure of assemblages. Regardless of approach, assemblages were generally no different than expected by chance. In contrast, phylogenetic structures of bird assemblages were overdispersed, clumped or consistent with random assembly depending on the site. Nonetheless, we found little evidence for differences in phylogenetic structure arising as a consequence of the identity of the species pool. We identified a strong relationship between the proportion of residents and phylogenetic relatedness that was unrelated to the species richness of assemblages. Our results suggest that different assembly mechanisms may structure resident and migratory subsets of temperate breeding bird communities. Resident assemblages are likely structured by interspecific interactions and habitat filtering prior to arrival of migrants. In contrast, the composition of migrant assemblages may be a consequence of priority effects in which the presence and abundance of residents and earliest arriving species affect the ability of subsequent migrants to colonize sites. This phenomenon enhances the likelihood of multiple alternative community structures in similar environments.     

The complexity of forest borders determines the understorey vegetation

Questions

What are the most important drivers of plant species richness (gamma‐diversity) and species turnover (beta‐diversity) in the field layer of a forest edge? Does the tree and shrub species richness structure and complexity affect the richness of forest and grassland specialist species?

Location

Southeast Sweden.

Methods

We sampled 50 forest edges with different levels of structural complexity in agricultural landscapes. In each border we recorded trees, shrubs and herb layer species in a 50‐m transect parallel with the forest. We investigated species composition and species turnover in relation to the proportions of gaps in the border and the diversity of trees and shrubs.

Results

Total plant species richness in the field layer was mainly explained by the proportion of gaps to areas with full canopy cover and tree diversity. Increasing number of gaps promoted higher diversity of grassland specialist species within the field layer, resulting in open forest borders with the highest overall species richness. Gaps did however have a negative impact on forest species richness. Conversely, increasing forest species richness was positively related to tree diversity, but the number of grassland specialist species was negatively affected by tree diversity.

Conclusions

Managing forest borders, and therefore increasing the area of semi‐open habitats in fragmented agricultural landscapes, provides future opportunities to create a network of suitable habitats for both grassland and deciduous forest specialist species. Such measures therefore have the potential to increase functional connectivity and support dispersal of species in homogeneous forest/agricultural landscapes.     

Nested patterns of community assembly in the colonisation of artificial canopy habitats by oribatid mites

An observed species–area relationship (SAR) in assemblages of oribatid mites inhabiting natural canopy habitats (suspended soils) led to an experimental investigation of how patch size, height in canopy and moisture influence the species richness, abundance and community composition of arboreal oribatid mites. Colonisation by oribatid mites on 90 artificial canopy habitats (ACHs) of three sizes placed at each of three heights on the trunks of ten western redcedar trees was recorded over a 1‐year period. Fifty‐nine oribatid mite species colonised the ACHs, and richness increased with the moisture content and size of the habitat patch. Oribatid mite species richness and abundance, and ACH moisture content decreased with increasing ACH height in the canopy. Patterns in the species richness and community composition of ACHs were non‐random and demonstrated a significant nested pattern. Correlations of patch size, canopy height and moisture content with community nestedness suggest that species‐specific environmental tolerances combined with the differential dispersal abilities of species contributed to the non‐random patterns of composition in these habitats. In line with the prediction that niche‐selection filters out species from the regional pool that cannot tolerate environmental harshness, moisture‐stressed ACHs in the high canopy had lower community variability than ACHs in the lower canopy. Colonising source pools to ACHs were almost exclusively naturally‐occurring canopy sources, but low levels of colonisation from the forest floor were apparent at low heights within the ACH system. We conclude that stochastic dispersal dynamics within the canopy are crucial to understanding oribatid mite community structure in suspended soils, but that the relative importance of stochastic dispersal assembly may be dependent on a strong deterministic element to the environmental tolerances of individual species which drives non‐random patterns of community assembly.     

Effects of landscape disturbance on mosquito community composition in tropical Australia

Emerging infectious diseases are considered to be a growing threat to human and wildlife health. Such diseases might be facilitated by anthropogenic land-use changes that cause novel juxtapositions of different habitats and species and result in new interchanges of vectors, diseases, and hosts. To search for such effects in tropical Australia, we sampled mosquito populations across anthropogenic disturbance gradients of grassland, artificial rainforest edge, and rainforest interior. From >15,000 captured mosquitoes, we identified 26 species and eight genera. Surprisingly, there was no significant difference in community composition or species richness between forest edges and grasslands, but both differed significantly from rainforest interiors. Mosquito species richness was elevated in grasslands relative to the rainforest habitats. Seven species were unique to grasslands and edges, with another 13 found across all habitats. Among the three most abundant species, Culex annulirostris occurred in all habitat types, whereas Verrallina lineata and Cx. pullus were more abundant in forest interiors. Our findings suggest that the creation of anthropogenic grasslands adjacent to rainforests may increase the susceptibility of species in both habitats to transmission of novel diseases via observable changes and mixing of the vector community on rainforest edges.     

Edge responses are different in edges under natural versus anthropogenic influence: a meta‐analysis using ground beetles

Most edges are anthropogenic in origin, but are distinguishable by their maintaining processes (natural vs. continued anthropogenic interventions: forestry, agriculture, urbanization). We hypothesized that the dissimilar edge histories will be reflected in the diversity and assemblage composition of inhabitants. Testing this “history‐based edge effect” hypothesis, we evaluated published information on a common insect group, ground beetles (Coleoptera: Carabidae) in forest edges. A meta‐analysis showed that the diversity‐enhancing properties of edges significantly differed according to their history. Forest edges maintained by natural processes had significantly higher species richness than their interiors, while edges with continued anthropogenic influence did not. The filter function of edges was also essentially different depending on their history. For forest specialist species, edges maintained by natural processes were penetrable, allowing these species to move right through the edges, while edges still under anthropogenic interventions were impenetrable, preventing the dispersal of forest specialists out of the forest. For species inhabiting the surrounding matrix (open‐habitat and generalist species), edges created by forestry activities were penetrable, and such species also invaded the forest interior. However, natural forest edges constituted a barrier and prevented the invasion of matrix species into the forest interior. Preserving and protecting all edges maintained by natural processes, and preventing anthropogenic changes to their structure, composition, and characteristics are key factors to sustain biodiversity in forests. Moreover, the increasing presence of anthropogenic edges in a landscape is to be avoided, as they contribute to the loss of biodiversity. Simultaneously, edges under continued anthropogenic disturbance should be restored by increasing habitat heterogeneity.     

Forest edge and diversity: carabids along forest-grassland transects

Diversity relationships of carabids in forest edges and the neighbouring forest interior and the surrounding grassland are studied. Samples were taken along three replicated forest-grassland transects using pitfall traps in the Aggtelek National Park in Hungary during 2 years. The study revealed significant edge effect on the carabids. The Shannon diversity of carabids were significantly higher in the forest edge and the grassland than in the forest interior. Carabids of the forest interior, forest edge and grassland can be separated from each other by ordinations, both on the species composition and abundance, suggesting that all three habitats have a distinct species assemblage. Moreover, indicator species analysis detected significant edge associated species; based on the specificity and fidelity of the carabids we have distinguished five groups of species: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Multiple linear regression analyses showed that relative air moisture, temperature of the ground and cover of the herbs are the most important factors determining the diversity of carabids along the transects. Our results show that the high diversity of carabids in forest edges is due to the edge-associated species and the presence of species characteristic to the adjacent habitats. The significance of forest edges in nature conservation, serving as a source habitat for dispersal processes, contributing to the recolonization of carabids after habitat destruction or other disturbance is emphasized.