Distribution and abundance of dung beetles in fragmented landscapes

Related species utilising similar resources are often assumed to show similar spatial population structures and dynamics. This paper reports substantial ecological variation within a set of Aphodius dung beetles occurring in the same patchily distributed resource, livestock dung in pastures. We show how variation in habitat and resource selectivity, in the rate of movements between pastures, and in the distribution of local population sizes all contribute to interspecific differences in spatial population structures. Local dung beetle assemblages are compared between two landscapes with different densities of pastures. In one of the landscapes, we contrast the abundances and regional distributions of Aphodius before and after 15 years of rapid habitat loss. Different species show very dissimilar responses to changes in the structure of the landscape. Our results suggest that generalist Aphodius species, and specialist species with high dispersal powers, occur as large "patchy" populations in the landscape. In contrast, a strict pasture specialist species with limited dispersal powers (A. pusillus) forms classical metapopulations. At the community level, interspecific differences in spatial population structures make the local community composition a function of the structure of the surrounding landscape.     

Disentangling the ‘brown world’ faecal–detritus interaction web: dung beetle effects on soil microbial properties

Many ecosystem services are sustained by the combined action of microscopic and macroscopic organisms, and shaped by interactions between the two. However, studies tend to focus on only one of these two components. We combined the two by investigating the impact of macrofauna on microbial community composition and functioning in the context of a major ecosystem process: the decomposition of dung. We compared bacterial communities of pasture soil and experimental dung pats inhabited by one (Aphodius), two (Aphodius and Geotrupes), or no dung beetle genera. Overall, we found distinct microbial communities in soil and dung samples, and that the communities converged over the course of the experiment. Characterising the soil microbial communities underlying the dung pats revealed a significant interactive effect between the microflora and macrofauna, where the diversity and composition of microbial communities was significantly affected by the presence or absence of dung beetles. The specific identity of the beetles had no detectable impact, but the microbial evenness was lower in the presence of both Aphodius and Geotrupes than in the presence of Aphodius alone. Differences in microbial community composition were associated with differences in substrate usage as measured by Ecoplates. Moreover, microbial communities with similar compositions showed more similar substrate usage. Our study suggests that the presence of macrofauna (dung beetles) will modify the microflora (bacteria) of both dung pats and pasture soil, including community diversity and functioning. In particular, the presence of dung beetles promotes the transfer of bacteria across the soil–dung interface, resulting in increased similarity in community structure and functioning. The results demonstrate that to understand how microbes contribute to the ecosystem process of dung decomposition, there is a need to understand their interactions with larger co‐occurring fauna.     

Consistent effects of consumer species loss across different habitats

Our knowledge of the effects of consumer species loss on ecosystem functioning is limited by a paucity of manipulative field studies, particularly those that incorporate inter‐trophic effects. Further, given the ongoing transformation of natural habitats by anthropogenic activities, studies should assess the relative importance of biodiversity for ecosystem processes across different environmental contexts by including multiple habitat types. We tested the context‐dependency of the effects of consumer species loss by conducting a 15‐month field experiment in two habitats (mussel beds and rock pools) on a temperate rocky shore, focussing on the responses of algal assemblages following the single and combined removals of key gastropod grazers (Patella vulgata, P. ulyssiponensis, Littorina littorea and Gibbula umbilicalis). In both habitats, the removal of limpets resulted in a larger increase in macroalgal richness than that of either L. littorea or G. umbilicalis. Further, by the end of the study, macroalgal cover and richness were greater following the removal of multiple grazer species compared to single species removals. Despite substantial differences in physical properties and the structure of benthic assemblages between mussel beds and rock pools, the effects of grazer loss on macroalgal cover, richness, evenness and assemblage structure were remarkably consistent across both habitats. There was, however, a transient habitat‐dependent effect of grazer removal on macroalgal assemblage structure that emerged after three months, which was replaced by non‐interactive effects of grazer removal and habitat after 15 months. This study shows that the effects of the loss of key consumers may transcend large abiotic and biotic differences between habitats in rocky intertidal systems. While it is clear that consumer diversity is a primary driver of ecosystem functioning, determining its relative importance across multiple contexts is necessary to understand the consequences of consumer species loss against a background of environmental change. Synthesis The roles of species may vary with environmental context, making it difficult to predict how biodiversity loss affects ecosystem functioning across multiple habitats. We tested how natural algal assemblages in two distinct intertidal habitats responded to the removal of different combinations of key consumer species. Despite an initial habitat‐dependent effect of consumer loss, habitat type did not modify the longer‐term responses of algal assemblages to either the identity or number of consumer species removed. Our findings show that, in certain systems, consumer diversity remains a primary driver of ecosystem functioning across widely different environmental contexts.     

Leaf litter decomposition rates in degraded and fragmented tropical rain forests of Borneo

Previously extensive tracts of primary rain forest have been degraded by human activities, and we examined how the effects of forest disturbance arising from habitat fragmentation and commercial selective logging affected ecosystem functioning in these habitats by studying leaf litter decomposition rates in litter bags placed on the forest floor. The rain forests of Borneo are dominated by trees from the family Dipterocarpaceae, and we compared leaf litter decomposition rates of three dipterocarp species at eight forest fragment sites (area 3–3529 ha) that had different histories of disturbance pre‐fragmentation: four fragments had been selectively logged prior to fragmentation and four had been formed from previously undisturbed forest. We compared these logged and unlogged forest fragments with sites in continuous forest that had been selectively logged (two sites) and fully protected and undisturbed (two sites). After 120 d, undisturbed continuous forest sites had the fastest rates of decomposition (52% mass loss). Forest fragments formed from unlogged forest (32% mass loss) had faster decomposition rates than logged forest fragments (28% mass loss), but slower rates than continuous logged forest (39% mass loss). Leaves of a light‐demanding species (Parashorea malaanonan) decomposed faster than those of a shade‐tolerant species (Hopea nervosa), but decomposition of all three dipterocarp species that we studied responded similarly to logging and fragmentation effects. Reduced decomposition rates in logged and fragmented forest sites may affect nutrient cycling and thus have detrimental consequences for forest regeneration. Conservation management to improve forest quality should be a priority, particularly in logged forest fragments.     

Dung Beetle Community and Functions along a Habitat-Disturbance Gradient in the Amazon: A Rapid Assessment of Ecological Functions Associated to Biodiversity

Although there is increasing interest in the effects of habitat disturbance on community attributes and the potential consequences for ecosystem functioning, objective approaches linking biodiversity loss to functional loss are uncommon. The objectives of this study were to implement simultaneous assessment of community attributes (richness, abundance and biomass, each calculated for total-beetle assemblages as well as small- and large-beetle assemblages) and three ecological functions of dung beetles (dung removal, soil perturbation and secondary seed dispersal), to compare the effects of habitat disturbance on both sets of response variables, and their relations. We studied dung beetle community attributes and functions in five land-use systems representing a disturbance gradient in the Brazilian Amazon: primary forest, secondary forest, agroforestry, agriculture and pasture. All response variables were affected negatively by the intensification of habitat disturbance regimes, but community attributes and ecological functions did not follow the same pattern of decline. A hierarchical partitioning analysis showed that, although all community attributes had a significant effect on the three ecological functions (except the abundance of small beetles on all three ecological functions and the biomass of small beetles on secondary dispersal of large seed mimics), species richness and abundance of large beetles were the community attributes with the highest explanatory value. Our results show the importance of measuring ecological function empirically instead of deducing it from community metrics.     

Landscape size affects the relative importance of habitat amount,habitat fragmentation,and matrix quality on forest birds

It is important to understand the relative effects of landscape habitat loss, habitat fragmentation, and matrix quality on biodiversity, so that potential management options can be appropriately ranked. However, their effects and relative importance may change with the size of the landscape considered because the multiple (and potentially conflicting) ecological processes that are influenced by landscape structure occur at different spatial scales (e.g. dispersal, predation, foraging). We estimated the relative effects of habitat loss, habitat fragmentation, and matrix quality (measured as the amount of forest, the proportion of forest area contained in large core forests, and the density of roads respectively) on fragmentation‐sensitive forest birds in southern Ontario, Canada using a range of landscape sizes (0.8–310 km²). We used three complementary statistical approaches to estimate relative effects of these correlated landscape factors – 1) multiple regression, 2) information theoretic (AIC) estimates of the most parsimonious model, and 3) multi‐model inference to average effects across all supported models. We controlled for spatial autocorrelation, local habitat, roadside sampling bias, time of day, season, habitat heterogeneity, and the interaction between the effects of habitat amount and fragmentation. We found that relative effects of habitat amount and fragmentation were scale dependent; habitat amount had a consistently positive effect that was consistent over more than two orders of magnitude in landscape area (~1–300 km²). In contrast, the effects of habitat fragmentation depended on the size of the landscape considered. Indeed, for veery Catharus fuscescens, habitat fragmentation had positive effects at one scale and negative effects at another. The effects of matrix quality were generally weak and changed little with scale. For the number of fragmentation sensitive species and the presence of veery, habitat amount was most important in large landscapes and habitat fragmentation in small landscapes but for the presence of ovenbird Seiurus aurocapilla, habitat amount was most important at all scales.     

Dung beetle species interactions and multifunctionality are affected by an experimentally warmed climate

While substantial effort has been invested in modelling changes in species distribution with climate change, less attention has been given to how climate warming will affect interactions among co‐occurring species, and the cascading functional consequences. In this study, realistic dung beetle communities were subjected to an experimental warming treatment and the net effect on the functions of dung decomposition (in terms of dung mass) and plant productivity (in terms of biomass production of ryegrass grown on soil from underneath the dung pats) were examined. A priori, we hypothesized that the largest tunneling species would be functionally dominant, and be differently affected by experimental warming compared to pat‐dwelling, smaller species. In terms of dung decomposition, the largest beetles did prove to be the functionally most important, with the qualitative pattern unaffected by experimental warming. In contrast, for plant productivity all species appeared equally important under ambient conditions. However, the effects of single species on plant productivity were reduced as temperature increased: In a warmed climate, a combination of both tunneling and pat‐dwelling species came the closest to returning ecosystem functioning to levels found in the ambient treatment. These results suggest different roles for different species, and highlight the importance of maintaining multiple species within an ecosystem – particularly when systems are perturbed.     

Understanding the effect of landscape fragmentation and vegetation productivity on elephant habitat utilization in Amboseli ecosystem,Kenya

Understanding factors affecting the distribution of the African elephant is important for its conservation in increasingly human‐dominated savannah landscapes. However, understanding how landscape fragmentation and vegetation productivity affect elephant habitat utilization remains poorly understood. In this study, we tested whether landscape fragmentation and vegetation productivity explain elephant habitat utilization in the Amboseli ecosystem in Kenya. We used GPS (Global Positioning System) telemetry data from five elephants to quantify elephant habitat utilization. Habitat utilization was determined by calculating the time elephants spent within a unit area. We then used generalized additive models (GAMs) to model the relationship between time density and landscape fragmentation, as well as vegetation productivity. Results show that landscape fragmentation and vegetation productivity significantly (P < 0.05) explain elephant habitat utilization. A significant (P < 0.05) unimodal relationship between vegetation productivity and habitat utilization was observed. Results suggest that elephants spend much of their time in less fragmented landscapes of intermediate productivity.     

Influence of habitat complexity and landscape configuration on pollination and seed-dispersal interactions of wild cherry trees

Land-use intensification is a major cause for the decline in species diversity in human-modified landscapes. The loss of functionally important species can reduce a variety of ecosystem functions, such as pollination and seed dispersal, but the intricate relationships between land-use intensity, biodiversity and ecosystem functioning are still contentious. Along a gradient from forest to intensively used farmland, we quantified bee species richness, visitation rates of bees and pollination success of wild cherry trees (Prunus avium). We analysed the effects of structural habitat diversity at a local scale and of the proportion of suitable habitat around each tree at a landscape scale. We compared these findings with those from previous studies of seed-dispersing birds and mammals in the same model system and along the same land-use gradient. Bee species richness and visitation rates were found to be highest in structurally simple habitats, whereas bird species richness—but not their visitation rates—were highest in structurally complex habitats. Mammal visitation rates were only influenced at the landscape scale. These results show that different functional groups of animals respond idiosyncratically to gradients in habitat and landscape structure. Despite strong effects on bees and birds, pollination success and bird seed removal did not differ along the land-use gradient at both spatial scales. These results suggest that mobile organisms, such as bees and birds, move over long distances in intensively used landscapes and thereby buffer pollination and seed-dispersal interactions. We conclude that measures of species richness and interaction frequencies are not sufficient on their own to understand the ultimate consequences of land-use intensification on ecosystem functioning.     

Ecological restoration of rich fens in Europe and North America: from trial and error to an evidence‐based approach

Fens represent a large array of ecosystem services, including the highest biodiversity found among wetlands, hydrological services, water purification and carbon sequestration. Land‐use change and drainage has severely damaged or annihilated these services in many parts of North America and Europe; restoration plans are urgently needed at the landscape level. We review the major constraints on the restoration of rich fens and fen water bodies in agricultural areas in Europe and disturbed landscapes in North America: (i) habitat quality problems: drought, eutrophication, acidification, and toxicity, and (ii) recolonization problems: species pools, ecosystem fragmentation and connectivity, genetic variability, and invasive species; and here provide possible solutions. We discuss both positive and negative consequences of restoration measures, and their causes. The restoration of wetland ecosystem functioning and services has, for a long time, been based on a trial‐and‐error approach. By presenting research and practice on the restoration of rich fen ecosystems within agricultural areas, we demonstrate the importance of biogeochemical and ecological knowledge at different spatial scales for the management and restoration of biodiversity, water quality, carbon sequestration and other ecosystem services, especially in a changing climate. We define target processes that enable scientists, nature managers, water managers and policy makers to choose between different measures and to predict restoration prospects for different types of deteriorated fens and their starting conditions.     

Large-scale spatial ecology of dung beetles

Recent modelling work shows that the composition of local communities can be influenced by the configuration of the surrounding landscape, but many of these models assume that all community members display the same type of extinction‐colonization dynamics. I use Aphodius dung beetles to test the hypothesis that interspecific differences in habitat selection and dispersal capacity may translate into differences in spatial population dynamics, even among closely related species coexisting on the same resource. If this is true, then groups of species with different characteristics would show different responses to landscape configuration. I first divided the area of Finland into a grid, and used collection records to describe regional variation in the Aphodius fauna of open cattle pastures. I then sampled dung beetles on 131 cattle farms, to examine whether the subset of species found on each farm was related to the density of pastures in the surrounding grid square. Finally, I used historical records to analyze changes in dung beetle communities during the last century, when there was great loss of pasture. Overall, I found no relationship between landscape characteristics and the total proportion of the regional species pool that was found on each farm. However, the distribution of species among guilds with different habitat specificity did relate to the configuration of the landscape, and the pattern was most pronounced in a specialists species with limited dispersal. Associations between community structure and landscape configuration were superimposed on two much larger and stronger patterns: a large‐scale latitudinal gradient in regional species richness, and a decelerating gain of species to local communities with an increasing regional species pool. I conclude that ecological variation among community members is a crucial factor in the analysis of local community composition, and that local species richness should always be conditioned on regional richness.     

At the landscape level,birds respond strongly to habitat amount but weakly to fragmentation

Aim

It is usually thought that habitat fragmentation acts negatively on species survival, and consequently, on biodiversity. Recent literature challenges whether habitat fragmentation per se affects species richness, beyond the effect of habitat area. Theoretical studies have suggested that fragmentation may matter most when the amount of available habitat is small or at intermediate levels. However, a recent review suggests that the effect of fragmentation on species richness is usually positive. Here, we dissect the richness–fragmentation relationship. What is the effect size? Does it depend upon the amount of habitat cover? How do individual species respond to fragmentation?

Methods

Applying a macroecological approach, we empirically related avian richness and the probability of occurrence (p_occ) of individual species to fragmentation (number of patches), after controlling for habitat amount in 991 landscapes, each 100‐km², in southern Ontario, Canada.

Results

Species richness was strongly related to total habitat amount, but habitat fragmentation had no detectable additional effect. Individual species’ p_occ related strongly to habitat amount. For some species, p_occ also related secondarily to habitat fragmentation within landscapes. Logistic models revealed that p_occ related significantly negatively to fragmentation after controlling for habitat amount for only ~13% of forest‐ and 18% of open‐habitat species bird species. However, p_occ related significantly positively to fragmentation for even greater proportions of species, including some red‐listed species. Fragmentation effects were not stronger at low or intermediate levels of habitat amount within landscapes.

Conclusion

In earlier studies, negative effects of isolation were observed at the patch level in experimental manipulations. However, at the landscape level, avian species richness in southern Ontario apparently responds primarily to habitat amount and negligibly to fragmentation. We argue that the evidence is inconsistent with the hypothesis that reducing habitat fragmentation per se would be an effective conservation strategy for birds at the landscape level.

     

Ecosystem fragmentation drives increased diet variation in an endemic livebearing fish of the Bahamas

One consequence of human‐driven habitat degradation in general, and habitat fragmentation in particular, is loss of biodiversity. An often‐underappreciated aspect of habitat fragmentation relates to changes in the ecology of species that persist in altered habitats. In Bahamian wetlands, ecosystem fragmentation causes disruption of hydrological connectivity between inland fragmented wetlands and adjacent marine areas, with the consequent loss of marine piscivores from fragmented sections. We took advantage of this environmental gradient to investigate effects of ecosystem fragmentation on patterns of resource use in the livebearing fish Gambusia hubbsi (Family Poeciliidae), using both population‐ and individual‐level perspectives. We show that fragmentation‐induced release from predation led to increased G. hubbsi population densities, which consequently led to lower mean growth rates, likely as a result of higher intraspecific competition for food. This was accompanied by a broadening of dietary niches via increased interindividual diet variation, suggesting a negative effect of predation and a positive effect of intraspecific competition on the degree of diet variation in natural populations. Our results therefore indicate that habitat fragmentation can greatly impact the ecology of resilient populations, with potentially important ecological and evolutionary implications.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

The behavioural ecology and population dynamics of a cryptic ground‐dwelling mammal in an urban Australian landscape

Urbanization results in widespread habitat loss and fragmentation and generally has a negative impact upon native wildlife, in particular ground‐dwelling mammals. The northern brown bandicoot (Isoodon macrourus; Marsupialia: Peramelidae) is one of relatively few native Australian ground‐dwelling mammals that is able to survive within urbanized landscapes. As a consequence of extensive clearing and urban development within the city of Brisbane, bandicoots are now restricted to the mostly small (<10 ha) bushland fragments scattered across the city landscape. Our study examined the behavioural ecology of northern brown bandicoots within habitat fragments located on a major creek‐line, using mark‐recapture population monitoring and radio telemetry. Bandicoots at monitored sites were found to occur at high densities (typically one individual ha^?1), although one‐third of the populations were transient. Radio tracking revealed that bandicoots had relatively small home ranges (mean 1.5 ± 0.2 ha) comprised largely of bushland/grassland with dense, often weed‐infested ground cover. Bandicoots sheltered by day in these densely covered areas and also spent most time foraging there at night, although they occasionally ventured small distances to forage in adjacent maintained parklands and residential lawns. We suggest that introduced tall grasses and other weeds contribute to high habitat quality within riparian habitat fragments and facilitate the persistence of high density populations, comprised of individuals with small home ranges. The generalized dietary and habitat requirements of northern brown bandicoots, as well as a high reproductive output, undoubtedly facilitate the survival of the species in urban habitat fragments. Further research is required on other native mammal species in urbanized landscapes to gain a greater understanding of how best to conserve wildlife in these heavily modified environments.     

Ant community patterns in highly fragmented Chaco forests of central Argentina

The Chaco is the largest dry forest biome in South America and one of the regions most threatened by agricultural intensification. As a consequence, in several areas Chaco forests persist as forest remnants of different sizes embedded in an agricultural matrix. Ants are social insects that have key roles in ecosystem functioning, and the effects of this ongoing land use change process on ant communities are little known for this region. In the present study, we assessed the consequences of land use replacement by monocultures and forest fragmentation on ant communities. Particularly, we assessed whether patch size, patch isolation and edge effect affect species richness and composition of ground‐dwelling ants in fragmented landscapes of Chaco forests. We collected ants by combining hand collecting and pitfall traps in 17 forest fragments and the surrounding matrix from two sites in Córdoba, Argentina. Patch size and patch isolation had no effect on ant richness; however, patch isolation and, to a lesser extent, patch size altered ant species composition. The ant community was not affected by edge but it was negatively affected by the crop matrix, which reduced richness and altered species composition. These results indicate that monoculture matrices severely affect ant communities in the Chaco forests, and that the effects of other indicators of habitat fragmentation (patch size and edge effect) are subtler and less relevant. In the present context of land use change, even small fragments could have an important value for the conservation of ant diversity.     

Distribution and coexistence of shrews in patchy landscapes: A field test of multiple hypotheses

Despite the important role of shrews (Soricomorpha: Soricidae) in the functioning of ecosystems, as predators and prey, the effects of habitat loss and fragmentation on this guild of mammals are still unclear. We studied the distribution of 5 species (the greater white toothed shrew Crocidura leucodon; the lesser white toothed shrew Crocidura suaveolens; the pigmy shrew Sorex minutus; the Appennine shrew Sorex samniticus and the Etruscan shrew Suncus etruscus) in a fragmented landscape in central Italy.Shrews were trapped with pitfall traps made from plastic water bottles, the number of traps increased with patch size. A total of 170 individuals, of 5 species of shrews were captured. Shrews were widely distributed in our study area, however patch occupancy was determined mainly by vegetation and geometrical characteristics of the patches. Our data supports the hypotheses that patterns of habitat selection and the dynamics of seasonal abundance (habitat and temporal partitioning between similarly sized species) reduce competitive pressure, thus allowing coexistence of shrews in relatively species-rich assemblages, for such small amounts of habitat. The most important outcome of our results is the crucial role played by vegetation structure in determining distribution patterns. These results strongly suggest that measurements of the vegetation structure of habitat patches should always be included as explanatory variables when studying the distribution of shrews in fragmented landscapes.     

Experimental effects of habitat fragmentation on rove beetles and ants: patch area or edge?

The effects of habitat fragmentation may include the loss of species from isolated fragments or changes in species abundances among habitats that differ in area, structure, or edge characteristics. We measured the species richness and abundance of ground‐dwelling insects in a 1.14‐ha old field that was mowed to produce patches of unmowed vegetation which differed in size, degree of isolation, and the amount of habitat edge. Four treatments – ranging from unfragmented (169‐m²) to highly fragmented (1‐m²) patches – were replicated four times in a Latin square design, and insects were sampled twice during 1995 using 177 pitfall traps. Species richness showed a non‐monotonic response to fragmentation, with the fewest species occurring in the slightly fragmented treatment. Responses of rove beetles and ants, the most species‐rich and abundant taxa, respectively, were similar to the overall insect community but ants had a stronger and more consistent treatment effect in both sample months. Ordinations of ant and rove‐beetle assemblages using nonmetric multidimensional scaling showed that the slightly fragmented treatment differed from other treatments in species occurrence and abundance. The lower species richness in the slightly fragmented treatment was primarily due to a subset of ant and rove beetle species that showed a lower abundance than in other treatments, possibly because this treatment had the greatest amount of habitat edge. We hypothesize that the non‐monotonic species response to fragmentation was due to the differential effects of habitat edge on species movements across the habitat boundary between unmowed patches and mowed areas. A greater effect due to the amount of habitat edge rather than total patch area, at least among the range of patch sizes studied, suggests that the length of habitat edge may be quite important to the distribution and abundance of ground‐dwelling animals in fragmented habitats.     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Effects of local and landscape factors on the abundance of an endangered multivoltine butterfly at riverbanks

Plebejus argyrognomon is one of the grassland‐dwelling butterflies undergoing rapid decline in recent decades. Grassland habitats for butterflies are generally threatened by fragmentation and invasive species, hence are among the most vulnerable ecosystems. We studied the seasonal abundance of P. argyrognomon at habitat patches along the banks of the Kinugawa River in eastern Japan, to identify environmental factors suitable for population persistence of this species, including habitat patch connectivity. Results showed that the patch's host plant cover had a positive effect on abundance in all three seasons, while the shading of the host plants by surrounding non‐host plants and nearby forested area showed negative effects. Additionally, habitat patch connectivity and nectar richness could be considered as positive factors in autumn and summer, respectively. Analysis of habitat connectivity also showed that the Kinugawa River did not appear to act as a dispersal barrier for P. argyrognomon. Our findings emphasize the importance of understanding environmental factors that may vary among seasons, and such understanding could contribute to habitat management of multivoltine butterflies in fragmented landscapes.