Conserving butterflies in fragmented plantation forests: are edge and interior habitats equally important?

Edge effects are increasing in forest-dominated landscapes worldwide, due to increased fragmentation by other land uses. Understanding how species respond to edges is therefore critical to define adequate conservation measures. We compared the relative importance of interior and edge habitats for butterflies in a landscape composed of even-aged pine plantations interspersed with semi-natural habitats. Butterfly assemblages were surveyed simultaneously at the edge and the interior of 68 patches belonging to four main habitat types: herbaceous firebreaks, clearcuts and young pine stands, older pine stands, and deciduous woodlands. Butterfly species richness was higher at edges than in interior habitats, especially for pine stands. Assemblage composition differed significantly between edge and interior habitats, except for firebreaks. Of the 23 most abundant butterfly species, seven were significantly more abundant in one or all edge habitat types, five in interior habitats, and 11 species showed no edge-interior preference. Modelling the presence of individual species in edge habitats revealed the importance of habitat variables such as the abundance of nectar and host-plants, but also of the abundance of the same species in the adjacent interior habitat. Moreover, our results suggest that most species use several, different habitat types to find supplementary or complementary resources, including micro-climatic refuges to escape hot temperatures during summer. The use of adjacent edge and interior habitats by butterflies is probably a key process in such mosaic landscapes and underlines the importance of landscape heterogeneity for butterfly conservation.     

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.     

Dung beetle persistence in human-modified landscapes: Combining indicator species with anthropogenic land use and fragmentation-related effects

Identifying and making use of ecological indicators becomes an essential task in the conservation of tropical systems, mainly in fragmented landscapes where land use intensification and habitat loss are confounding factors in the detection of species’ responses to human-caused disturbance. We aimed to analyze the importance of anthropogenic land use and fragmentation-related effects on dung beetle (Coleoptera: Scarabaeinae) persistence according to the interior–exterior non-linear gradient (forest + matrix) in a fragmented Atlantic Forest landscape used to sugar cane production and cattle ranching/farming. We offer scores for a comprehensive set of community-level attributes, from beetle abundance to taxonomic and ecological composition (i.e. species body size), including a list of indicator species of different forest habitats and adjacent matrix. Dung beetles were surveyed by traps across forest interiors (i.e. core forest areas) and edges of a primary forest, small fragments, sugar cane fields and pastures in a total of 60 sites. Indicator analyses were conducted across the landscape, using two well-established methods (IndVal and SIMPER). Our results suggest that (1) cross-habitat taxonomic distinctness is associated with the presence of indicator species, (2) some species benefit or are dependent of open habitats created by human-disturbances, such as forest edges (e.g. Canthon nigripennis) and matrices (e.g. Canthon aff. piluliformis, Dichotomius nisus and Trichilum externepunctatum), (3) although landscape habitats exhibit reduced beta diversity, dung beetle assemblages are spatially organized in response to the presence of both forest habitats and matrix and fragment area, (4) forest interior supports beetle assemblages biased toward large-bodied species, (5) accordingly forest interior, forest edges and matrix support taxonomically distinct assemblages, both contributing to the bulk of species richness at landscape level, (6) the response of dung beetles to the interior–exterior non-linear gradient (i.e. forest edge + matrix) reveals a similar pattern regardless of the nature of the matrix, and (7) there is no within-habitat variation in beetle abundance and species richness associated with distance from forest edge. Given that there is a high number of forest-dependent or forest-interior specialist species (e.g. Aphengium aff. sordidum, Ateuchus aff. alipioi, Dichotomius mormon, Ontherus aff. erosus and Onthophagus aff. clypeatus) dung beetle persistence in human-modified landscape is highly dependent on the presence of core areas, although edge-affected and matrix habitats may be complementary. This information is essential to permit a better prospect for dung beetle persistence in human-modified landscapes as they continue to move toward edge-dominated landscapes with intensively managed matrices.     

Landscape context modulates alien plant invasion in Mediterranean forest edges

Natural habitats in human-altered landscapes are especially vulnerable to biological invasions, especially in their edges. We aim to understand the influence of landscape and local characteristics on biological invasions by exploring the level of plant invasion and alien species traits in forest edges in highly urbanized landscapes. We identified all plant species in 73 paired plots in the edge and 50 m towards the interior of the forest. We explored the association between alien species richness and similarity in species composition between edge and interior plots with landscape and local variables, using generalized linear models and variance partitioning techniques. Then, we performed Fourth-corner analyses to explore the association between alien plant traits and local and landscape variables. In contrast to native species richness, alien species richness was more affected by the surrounding landscape than by the local characteristics of the edge. Road proximity was positively associated with alien species richness and proportion and was its most important correlate, whereas disturbance was negatively associated with native species richness and was its most influential factor. Alien plant traits were also primarily associated with landscape characteristics. For instance, species of Mediterranean origin and introduced for agriculture were associated with higher agriculture use in the landscape. Our findings suggest that risk analyses of habitat vulnerability to invasion must consider the landscape context in order to successfully predict highly invaded areas and identify potentially successful invaders.     

The role of various meadow margin types in shaping carabid and staphylinid beetle assemblages (Coleoptera: Carabidae,Staphylinidae) in meadow dominated landscapes

Less intensively managed semi-natural habitats, e.g., field and meadow margins like hedgerows, are thought to be crucial landscape components for maintaining biodiversity in highly disturbed and intensively managed agricultural landscapes. In this study, we focused on the effects of three meadow margin types on activity-density, species richness and species composition of carabid and staphylinid beetles recorded by pitfall traps in Central European landscapes dominated by intensively managed meadows. Carabid activity-density was significantly higher in meadows than in meadow margins and within meadow margins their activity-density increased from grassy meadow margins via shrubby ones to woody meadow margins. We found that recorded species richness of both carabid and staphylinid beetles was not significantly affected by habitat identity (meadow margin or neighbouring meadow) and meadow margin type. Recorded species composition of both investigated taxa was significantly affected by habitat identity and interaction between habitat identity and meadow margin type (i.e. it differed between particular meadow margin types). Assemblages inhabiting various meadow margin types were more dissimilar between each other than assemblages from neighbouring meadows. Meadow margins within grassland dominated landscapes maintain local species richness by hosting different species from those living in surrounding meadows. Dissimilarity of carabid and staphylinid assemblages from meadows neighbouring both sides of particular meadow margin did not differ between meadow margin types. Our results indicate that semi-natural habitats play an important role in maintaining biodiversity not only in agricultural landscapes dominated by arable fields, but also in those dominated by meadows.     

Forest–farm edge effects on communities of ground beetles (Coleoptera: Carabidae) under different landscape structures

In fragmented landscapes, ecological processes may be significantly influenced by edge effects, but few data are available for edge effects across forest–farmland edges. We investigated patterns of species richness, abundance, and species composition in ground beetles across forest–farm edges in two different agro-forest landscapes in Korea. Nine and five sites were selected from Hwaseong, a fragmented landscape, in 2011 and 2012, respectively, while eight sites were selected from Hoengseong, a relatively well-protected landscape, in 2012. Ground beetles were collected by pitfall trapping. Species richness was higher in the surrounding habitat than in the forest interior or edge in both Hwaseong and Hoengseong. However, in Hwaseong, species richness of the forest edge was similar to that of the forest interior, while in Hoengseong forest edge species richness was intermediate between that of the forest interior and surrounding areas. In addition, non-metric multidimensional scaling based on the combined data of both locations showed that the species composition of ground beetles in the forest edge was more similar to that of the forest interior than the surrounding areas, although some open-habitat species occurred at the forest edges. Three characteristic groups (forest specialists, edge-associated species, and open-habitat species) of ground beetle species were detected by indicator value analysis. In our study, ground beetle assemblages differed in the forest edges of two agro-forest landscapes, suggesting that the edge effect on biota can be influenced by landscape structure.     

Responses of wasp communities to urbanization: effects on community resilience and species diversity

Urbanization is one of the most extreme and rapidly growing anthropogenic pressures on the natural world. It is linked to significant impacts on biodiversity and disruptions to ecological processes in remnant vegetation. We investigated the richness and abundance of wasps in a highly fragmented urban landscape in Sydney, Australia, comparing assemblages in small urban remnants to edges and interiors of continuous areas of vegetation. We detected no difference in wasp abundance or species richness between remnant types indicating that communities are highly resilient to the effects of urbanization at this scale. However, Chao 2 estimates of predicted species richness indicate that edge sites would support a greater richness and abundance of species compared to small and interior sites. Although families were represented evenly across the sites, interior and edge sites supported more species within families. Wasp composition was significantly affected by the temporal variation and trap location (arboreal or ground), particularly at the family level demonstrating high species turnover and discrimination in vertical space. These sampling effects and temporal inconsistencies highlight the hazards of relying on one-off snapshot surveys and uncorrected datasets for assessments of diversity and responses to urban landscapes. The strong resilience of wasp communities to urbanization when assessed at coarse scales indicates that responses at finer spatial and taxonomic scales are critical to understanding the maintenance of ecosystem function in highly modified landscapes.     

Gradients in the number of species at reef-seagrass ecotones explained by gradients in abundance

Gradients in the composition and diversity (e.g. number of species) of faunal assemblages are common at ecotones between juxtaposed habitats. Patterns in the number of species, however, can be confounded by patterns in abundance of individuals, because more species tend to be found wherever there are more individuals. We tested whether proximity to reefs influenced patterns in the composition and diversity ('species density' = number of species per area and 'species richness' = number of species per number of individuals) of prosobranch gastropods in meadows of two seagrasses with different physiognomy: Posidonia and Amphibolis. A change in the species composition was observed from reef-seagrass edges towards the interiors of Amphibolis, but not in Posidonia meadows. Similarly, the abundance of gastropods and species density was higher at edges relative to interiors of Amphibolis meadows, but not in Posidonia meadows. However, species richness was not affected by proximity to reefs in either type of seagrass meadow. The higher number of species at the reef-Amphibolis edge was therefore a consequence of higher abundance, rather than species richness per se. These results suggest that patterns in the composition and diversity of fauna with proximity to adjacent habitats, and the underlying processes that they reflect, likely depend on the physiognomy of the habitat.     

Environmental filtering is the main assembly rule of ground beetles in the forest and its edge but not in the adjacent grassland

In a fragmented landscape,transitional zones between neighboring habitats are common,and our understanding of community organizational forces across such habitats is important.Edge studies are numerous,but the majority of them utilize information on species richness and abundance.Abundance and taxonomic diversity,however,provide little information on the functioning and phylogeny of the co-existing species.Combining the evaluation of their functional and phylogenetic relationships,we aimed to assess whether ground beetle assemblages are deterministically or stochastically structured along grassland-forest gradients.Our results showed different community assembly rules on opposite sides of the forest edge.In the grassland,co-occurring species were functionally and phylogenetically not different from the random null model,indicating a random assembly process.Contrary to this,at the forest edge and the interior,co-occurring species showed functional and phylogenetic clustering,thus environmental filtering was the likely process structuring carabid assemblages.Community assembly in the grassland was considerably affected by asymmetrical species flows (spillover)across the forest edge:more forest species penetrated into the grassland than open-habitat and generalist species entered into the forest.This asymmetrical species flow underlines the importance of the filter function of forest edges.As unfavorable,human-induced changes to the structure,composition and characteristics of forest edges may alter their filter function,edges have to be specifically considered during conservation management.     

Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis

Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This 'landscape-divergence hypothesis', if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.     

Effects of structural diversity and identity of patches of habitat on diversity of benthic assemblages

A fundamental problem in ecology, regardless of habitat or system, is understanding the relationship between habitats and assemblage of organisms. It is commonly accepted that differences in composition and surrounding landscape of habitats affect the diversity of assemblages, although there is not much empirical evidence because of difficulties of manipulating structure in many habitats. These relationships were examined experimentally, using habitats of artificial turfs that are colonized by diverse assemblages of gastropods. Each habitat was made of nine sub‐habitats, which were sampled individually to allow tests of hypotheses about the effect of type of habitat and the influence of other adjacent sub‐habitats on the colonizing assemblage. Turf habitats were deployed for 8 weeks on a rocky shore after which they were collected and the colonizing assemblages of gastropods sampled. Independently of the types of turfs combined to form different habitats, there were more species where there was more than one type of component in a habitat (i.e. structural diversity). The type of habitat (i.e. structural identity) itself had little or no influence on the colonizing assemblage. The number of species colonizing short‐sparse and short‐dense turfs was influenced by which type of habitat was adjacent. Thus, when units of one type (e.g. short‐sparse turf) were added to a patch of habitat of long‐sparse turfs, the number of species in short‐sparse turfs was greater than in patches of the same type. This also increased total number of taxa in the whole patch of habitat. These results show how diversity of gastropods colonizing heterogeneous patches of habitat is influenced not only by the number of types of sub‐habitats, but also by interactions with surrounding sub‐habitats. These findings reiterate the importance of investigating the role of structure of habitats and of their surrounding landscapes across different systems, irrespective of their size or associated assemblages of organisms.     

Edge effects across boundaries between natural and restored jarrah (Eucalyptus marginata) forests in south‐western Australia

Edge effects are a widespread and ubiquitous ecological phenomenon, yet they remain poorly studied across edges between restored and natural forests. To address this lack of knowledge, we studied vertebrate communities across edges between 3‐year old restored mine‐pits and adjacent unmined forest in the jarrah (Eucalyptus marginata) forest of south‐western Australia. We found that mammal communities showed no edge response but reptile communities did. Overall reptile abundance and Morethia obscura abundance were higher in unmined forest along edges, Egernia napoleonis abundance was lower in unmined forest along edges, while Pogona minor abundance was lower in restored mine‐pits along edges. Predictive models were unable to predict species edge responses, due to the lack of knowledge of the ecology of jarrah forest reptiles, but proved useful in identifying potential ecological mechanisms behind observed edge responses and suggested that potential mechanisms were likely different for each species. Our study is the first to show edge responses in both habitats forming the edge between restored and natural forests, emphasizing the importance of studying both habitats forming the edge. Our results also suggest that, despite being poorly studied, edge responses are common across edges between restored and natural forest and result from a variety of ecological mechanisms. An increased understanding of the ecological mechanisms driving edge responses across edges between restored and natural forests will improve our ability to integrate restored areas into cross‐landscape management and, ultimately, improve our ability to manage landscapes for biodiversity conservation.     

Influence and value of different water regimes on avian species richness in arid inland Australia

Riparian habitats in arid landscapes are recognised for their structurally diverse vegetation and diverse bird species assemblages. In the extensive semi-arid and arid centre of Australia, riparian woodland habitats are impacted by pastoral land-use which may negatively influence vegetation structure and avian species composition. However, pastoralism has promoted the establishment of artificial water bodies, so that additional riparian vegetation may occur in the landscape. In this study, we surveyed the importance of different water regimes (i.e. artificial lakes, natural waterholes, desert sites) together with their associated vegetation on avian species richness in north-western New South Wales, Australia. Our results show that bird species richness was highest at water locations, in particular at artificial lakes. Avian species richness was negatively associated with distance to water bodies, both in desert vegetation types and in the riparian vegetation type along dry creeks. Moreover, riparian habitats supported larger avian assemblages and especially those of sedentary bird species compared to the surrounding shrub-steppe landscape. This indicates that artificial water bodies may be of significance for arid zone bird species and might gain in importance with changing water availabilities due to climatic changes.     

Edge contrast does not modulate edge effect on plants and pollinators

Edge contrast, is one of the main determinants of edge effects. This study examines the response of plant and pollinator diversity (bees and butterflies) to forest edge contrast, i.e. the difference between forests and adjacent open habitats with different disturbance regimes. We also investigated a potential cascading effect from plants to pollinators and whether edge structure and landscape composition mediate the relationship between edge contrast and beta diversity of pollinators. We sampled 51 low-contrast edges where forests were adjacent to habitats showing low levels of disturbance (i.e. grey dunes, mowed fire-breaks, orchards, grasslands) and 29 high-contrast edges where forests were adjacent to more intensively disturbed habitats (i.e. tilled firebreaks, oilseed rape) in three regions of France. We showed that plant diversities were higher in edges than in adjacent open habitat, whatever the edge contrast. However, plant beta diversity did not differ significantly between low and high-contrast edges. While we observed higher pollinator diversities in adjacent habitats than in low-contrast edges, there were no significant differences in pollinator beta diversity depending on edge contrast. We did not observe a cascading effect from plants to pollinators. Plant and bee beta diversities were mainly explained by local factors (edge structure and flower cover) while butterfly beta diversity was explained by surrounding landscape characteristics (proportion of land cover in grassland).     

Edge effects on ant community structure and species richness in an agricultural landscape

The effect of sharp edges between three different types of land use on the species richness and structure of ant communities was examined in an agricultural landscape within Central Hesse, Germany. Species richness and nest densities of ants at the centres and the edges of meadows, crop fields, and fallow land were recorded by hand sampling during 1997 and 1998. Edges between different land-use types did not increase ant species richness at the landscape scale, nor were they unique habitats for a specialised ant fauna. Nonetheless, most species shared ecotonal effects in the way that their relative abundance either decreased (e.g. Myrmica scabrinodis) or increased (e.g. Lasius niger, Lasius flavus) at the edges, resulting in different community structure between edges and centres of the land-use types. This was influenced by two major factors: (i) the boundary contrast between the neighbouring habitats (i.e. in terms of disturbance caused by agricultural practices), and (ii) the response of different species to changing abiotic conditions. High nest densities of aggressive species with large colonies occurred along edges. We hypothesise that this can significantly reduce edge permeability for surface-dwelling arthropods.     

Markedly Divergent Tree Assemblage Responses to Tropical Forest Loss and Fragmentation across a Strong Seasonality Gradient

We examine the effects of forest fragmentation on the structure and composition of tree assemblages within three seasonal and aseasonal forest types of southern Brazil, including evergreen, Araucaria, and deciduous forests. We sampled three southernmost Atlantic Forest landscapes, including the largest continuous forest protected areas within each forest type. Tree assemblages in each forest type were sampled within 10 plots of 0.1 ha in both continuous forests and 10 adjacent forest fragments. All trees within each plot were assigned to trait categories describing their regeneration strategy, vertical stratification, seed-dispersal mode, seed size, and wood density. We detected differences among both forest types and landscape contexts in terms of overall tree species richness, and the density and species richness of different functional groups in terms of regeneration strategy, seed dispersal mode and woody density. Overall, evergreen forest fragments exhibited the largest deviations from continuous forest plots in assemblage structure. Evergreen, Araucaria and deciduous forests diverge in the functional composition of tree floras, particularly in relation to regeneration strategy and stress tolerance. By supporting a more diversified light-demanding and stress-tolerant flora with reduced richness and abundance of shade-tolerant, old-growth species, both deciduous and Araucaria forest tree assemblages are more intrinsically resilient to contemporary human-disturbances, including fragmentation-induced edge effects, in terms of species erosion and functional shifts. We suggest that these intrinsic differences in the direction and magnitude of responses to changes in landscape structure between forest types should guide a wide range of conservation strategies in restoring fragmented tropical forest landscapes worldwide.     

Dynamic effects of ground-layer plant communities on beetles in a fragmented farming landscape

Vegetation effects on arthropods are well recognized, but it is unclear how different vegetation attributes might influence arthropod assemblages across mixed-agricultural landscapes. Understanding how plant communities influence arthropods under different habitat and seasonal contexts can identify vegetation management options for arthropod biodiversity. We examined relationships between vegetation structure, plant species richness and plant species composition, and the diversity and composition of beetles in different habitats and time periods. We asked: (1) What is the relative importance of plant species richness, vegetation structure and plant composition in explaining beetle species richness, activity-density and composition? (2) How do plant-beetle relationships vary between different habitats over time? We sampled beetles using pitfall traps and surveyed vegetation in three habitats (woodland, farmland, their edges) during peak crop growth in spring and post-harvest in summer. Plant composition better predicted beetle composition than vegetation structure. Both plant richness and vegetation structure significantly and positively affected beetle activity-density. The influence of all vegetation attributes often varied in strength and direction between habitats and seasons for all trophic groups. The variable nature of plant-beetle relationships suggests that vegetation management could be targeted at specific habitats and time periods to maximize positive outcomes for beetle diversity. In particular, management that promotes plant richness at edges, and promotes herbaceous cover during summer, can support beetle diversity. Conserving ground cover in all habitats may improve activity-density of all beetle trophic groups. The impacts of existing weed control strategies in Australian crop margins on arthropod biodiversity require further study.     

Butterfly edge effects are predicted by a simple model in a complex landscape

Edge responses have been studied for decades and form a critical component of our understanding of how organisms respond to landscape structure and habitat fragmentation. Until recently, however, the lack of a general, conceptual framework has made it difficult to make sense of the patterns and variability reported in the edge literature. We present a test of an edge effects model which predicts that organisms should avoid edges with less-preferred habitat, show increased abundance near edges with preferred habitat or habitat containing complementary resources, and show no response to edges with similar-quality habitat that offers only supplementary resources. We tested the predictions of this model against observations of the edge responses of 15 butterfly species at 12 different edge types within a complex, desert riparian landscape. Observations matched model predictions more than would be expected by chance for the 211 species/edge combinations tested over 3 years of study. In cases where positive or negative edge responses were predicted, observed responses matched those predictions 70% of the time. While the model tends to underpredict neutral results, it was rare that an observed edge response contradicted that predicted by the model. This study also supported the two primary ecological mechanisms underlying the model, although not equally. We detected a positive relationship between habitat preferences and the slope of the observed edge response, suggesting that this basic life history trait underlies edge effects and influences their magnitude. Empirical evidence also suggested the presence of complementary resources underlies positive edge responses, but only when completely confined to the adjacent habitat. This multi-species test of a general edge effects model at multiple edge types shows that resource-based mechanisms can explain many edge responses and that a modest knowledge of life history attributes and resource availability is sufficient for predicting and understanding many edge responses in complex landscapes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Positive edge effects on forest-interior cryptogams in clear-cuts

Biological edge effects are often assessed in high quality focal habitats that are negatively influenced by human-modified low quality matrix habitats. A deeper understanding of the possibilities for positive edge effects in matrix habitats bordering focal habitats (e.g. spillover effects) is, however, essential for enhancing landscape-level resilience to human alterations. We surveyed epixylic (dead wood inhabiting) forest-interior cryptogams (lichens, bryophytes, and fungi) associated with mature old-growth forests in 30 young managed Swedish boreal forest stands bordering a mature forest of high conservation value. In each young stand we registered species occurrences on coarse dead wood in transects 0-50 m from the border between stand types. We quantified the effect of distance from the mature forest on the occurrence of forest-interior species in the young stands, while accounting for local environment and propagule sources. For comparison we also surveyed epixylic open-habitat (associated with open forests) and generalist cryptogams. Species composition of epixylic cryptogams in young stands differed with distance from the mature forest: the frequency of occurrence of forest-interior species decreased with increasing distance whereas it increased for open-habitat species. Generalists were unaffected by distance. Epixylic, boreal forest-interior cryptogams do occur in matrix habitats such as clear-cuts. In addition, they are associated with the matrix edge because of a favourable microclimate closer to the mature forest on southern matrix edges. Retention and creation of dead wood in clear-cuts along the edges to focal habitats is a feasible way to enhance the long-term persistence of epixylic habitat specialists in fragmented landscapes. The proposed management measures should be performed in the whole stand as it matures, since microclimatic edge effects diminish as the matrix habitat matures. We argue that management that aims to increase habitat quality in matrix habitats bordering focal habitats should increase the probability of long-term persistence of habitat specialists.     

Fruit-feeding butterflies in edge-dominated habitats: community structure,species persistence and cascade effect

As old-growth forests are converted into edge-affected habitats, a substantial proportion of tropical biodiversity is potentially threatened. Here, we examine a comprehensive set of community-level attributes of fruit-feeding butterfly assemblages inhabiting edge-affected habitats in a fragmented Atlantic forest landscape devoted to sugar cane production. We also explored whether the consequences of habitat loss and fragmentation can interact and cause cascading ecosystem changes, with the pervasive simplification of tree assemblages inhabiting edge-dominated habitats, altering fruit-feeding butterfly persistence. Butterflies were sampled in three forest habitats: small fragments, forest edges and patches of forest interior of a primary forest fragment. Assemblage attributes, including taxonomic composition, correlated to some patch (patch size) and landscape (such as forest cover) metrics as well as habitat structure (tree density and richness). Fruit-feeding butterfly assemblages in the forest interior differed from those in small fragments due to an increased abundance of edge-specialist species. On the other hand, several forest-dependent species were missing in both small fragments and forest edges. Our results suggest that edge-affected habitats dominated by pioneer tree species support taxonomically distinct assemblages, including the presence of disturbance-adapted species, and butterfly community structure is highly sensitive to fragmentation- and plant-related variables, such as forest cover and pioneer tree species. In this way, while the establishment of human-modified landscapes probably results in the local extirpation of forest-dependent species, it allows the persistence of disturbance-adapted species. Thus, forest-dependent species conservation and the plant–animal interaction webs they support could be improved by retaining a significant amount of core forest habitat.