Dung beetle (Coleoptera: Scarabaeinae) persistence in Amazonian forest fragments and adjacent pastures: biogeographic implications for alpha and beta diversity

The aim of this study is to analyze the effects of habitat loss and forest replacement by cattle pasture on the alpha and beta diversity, abundance, biomass and species composition of dung beetles with different dispersal ability. Dung beetles were captured in 19 forest fragments and neighbouring pastures. Forest fragment area ranged from 3.7 to 4825 ha and in this study were grouped into four categories: small, medium, large and control forest. A total of 35,048 dung beetles representing 101 species were collected. Forest fragments had the highest richness with 81 species, followed by pasture with 58 species. Replacement of forest by pasture reduced species richness; however, due to the proximity and connectivity of these areas with Cerrado patches, pastures also had high species richness, but species composition was independent of adjacent fragments. Small fragments had lower abundance and species richness than our other habitat categories, even pastures. Our results highlight that proximity and connectivity with Cerrado areas influenced the patterns of alpha and beta diversity of dung beetles in fragments and pastures. We highlight that the ability to cross the pasture matrix is a strong adaptive trait for species living in human-modified landscapes. Consequently, species with these abilities are less susceptible to the effects of forest fragmentation and local extinction. Our results reinforce the importance of considering the biogeographic location and distribution pattern of species in forest fragmentation studies.     

Roadside connectivity does not increase reptile abundance or richness in a fragmented mallee landscape

The effect of isolation and the importance of dispersal in establishing and maintaining populations in fragments of remnant habitat remain poorly understood. Nevertheless, environmental connectivity is likely to be important for ensuring the long‐term preservation of biodiversity in extensively cleared landscapes. In this study, we compared reptile communities in large conservation parks with those in small woodland remnants 6.5–12 km from the parks, on the Eyre Peninsula, South Australia, Australia. We assessed the impact of fragmentation on the abundance, richness and habitat preferences of reptiles, and examined whether connection to linear roadside vegetation altered reptile communities in small woodland remnants. Of the 31 reptile species, 12 were restricted to conservation parks and six to habitat fragments in farmland. There was a substantial reduction in reptile species richness and abundance in farmland fragments. Direct connection of remnant vegetation to roadside corridors did not affect abundance of common species in the farmland fragments, although species richness was lower in isolated remnants in one of our two study regions. The habitat preference of the scincid lizard Menetia greyii differed between farmland fragments, where they were regularly found on dunes and roadsides, and conservation parks, where they were rare and not detected on dunes. We suggest that habitat fragmentation may have altered interspecific interactions, enabling an expansion of habitat use in the farming landscape. Significantly lower abundance of four common species in farmland settings compared with reserves indicated that existing corridors and small fragments provide inadequate connectivity over larger distances. To counter this effect, large reserves may need to be less than 10 km apart.     

Landscape structure and land use history influence changes in island plant composition after 100 years

Aim We investigated how current and historical land use and landscape structure affect species richness and the processes of extinction, immigration and species turnover. Location The northern part of the Stockholm archipelago, Baltic Sea, Sweden. We resurveyed 27 islands ranging from 0.3 to 33 ha in area. Methods We compared current plant survey data, cadastral maps and aerial photographs with records obtained from a survey in 1908, using databases and a digital elevation model to examine changes in plant community dynamics in space and time. We examined the effects of local and landscape structure and land use changes on plant species dynamics by using stepwise regression in relation to eight local and three landscape variables. The eight local variables were area, relative age, shape, soil heterogeneity, bedrock ratio, number of houses, forest cover change, and grazing 100 years ago. The three landscape variables were distance to mainland, distance to closest island with a farm 100 years ago, and structural connectivity. Hanski’s connectivity measure was modified to incorporate both connectivity and fragmentation. Results The investigated islands have undergone drastic changes, with increasing forest cover, habitation, and abandonment of grassland management. Although the total species richness increased by 31% and mean island area by 23%, we found no significant increase in species richness per unit area. Local variables explain past species richness (100 years ago), whereas both local and landscape variables explain current species richness, extinctions, immigrations and species turnover. Grazing that occurred 100 years ago still influences species richness, even though grazing management was abandoned several decades ago. The evidence clearly shows an increase in nitrophilous plant species, particularly among immigrant species. Main conclusions This study highlights the importance of including land use history when interpreting current patterns of species richness. Furthermore, local environment and landscape patterns affect important ecological processes such as immigration, extinction and species turnover, and hence should be included when assessing the impact of habitat fragmentation and land use change. We suggest that our modified structural connectivity measure can be applied to other types of landscapes to investigate the effects of fragmentation and habitat loss.     

Impacts of forest fragmentation on orchid bee (Hymenoptera: Apidae: Euglossini) communities in the Chocó biodiversity hotspot of northwest Ecuador

Habitat loss is a major driver of bee declines worldwide, and is of key relevance in the tropics given high deforestation rates, but we continue to have a poor understanding of the impact of land-cover change on tropical bee communities. Orchid bees (Apidae: Euglossini) are critical long-distance pollinators and may be highly susceptible to forest fragmentation given their reliance on forest habitat. Previous studies on the impact of forest fragmentation on euglossines have been geographically limited, have largely ignored β-diversity, and have not compared fragments with continuous forest. To contribute to addressing these gaps, we sampled male euglossine bees in 18 forest fragments (area range: 2.5–33 ha) and at eight locations within a large (3500 ha) continuous forest in the Chocó biodiversity hotspot of Ecuador during the dry season in 2014. We assessed how euglossine abundance, richness, and evenness related to fragment area, isolation, and edge:area ratio. We also compared fragments to continuous forest, in terms of α- and β-diversity. In fragments, a single species (Euglossa tridentata) comprised 78% of captures, and we found no significant effect of fragment area, isolation, or edge on abundance, richness, or evenness among fragments. Forest fragments and continuous forest differed in both community composition and evenness, but not in abundance or species richness. Spatial turnover (β-diversity) showed a non-significant trend toward changing more rapidly in continuous forest relative to fragments. These results underscore the conservation value of continuous forest for orchid bee diversity.     

Relative importance of habitat area, connectivity, management and local factors for vascular plants: spring ephemerals in boreal semi-natural grasslands

Using species and environmental data from an extensive grassland area in south-western Finland, we investigated the effect of patch area and connectivity, management and local habitat variables on the occurrence of spring-flowering vascular plants and their richness in boreal agricultural landscapes. Generalized linear models (GLM) and variation partitioning were used to study the explanatory power of the three groups of variables and their combined contributions on the richness and occurrence of six spring-flowering plant species. Generalized additive models (GAMs) and associated cross-validation tests were used to evaluate the predictability of the species occurrence and richness patterns. Present-day grassland patch area and connectivity were important predictors for occurrence and richness of the studied plant species. In addition, local habitat factors, especially radiation, accounted for major fractions of occurrence patterns of the studied species. Hybrid models including variables from all three variable groups had higher explanatory power and predictive capability than partial models. However, performance of the separate single-species models varied considerably between the six study species. Exclusion of radiation or connectivity from the hybrid models decreased their predictive performance, suggesting that these factors are of particular importance for grassland plant species at their northern range margins. When developing conservation and management planning for grassland plant species in Northern Europe, attention should be paid to well-connected networks of grassland patches including large, steeply-sloped patches with a favorable microclimate.     

Landscape-scale habitat fragmentation is positively related to biodiversity,despite patch-scale ecosystem decay

Positive effects of habitat patch size on biodiversity are often extrapolated to infer negative effects of habitat fragmentation on biodiversity at landscape scales. However, such cross-scale extrapolations typically fail. A recent, landmark, patch-scale analysis (Chase et al., 2020, Nature 584, 238–243) demonstrates positive patch size effects on biodiversity, that is, ‘ecosystem decay’ in small patches. Other authors have already extrapolated this result to infer negative fragmentation effects, that is, higher biodiversity in a few large than many small patches of the same cumulative habitat area. We test whether this extrapolation is valid. We find that landscape-scale patterns are opposite to their analogous patch-scale patterns: for sets of patches with equal total habitat area, species richness and evenness decrease with increasing mean size of the patches comprising that area, even when considering only species of conservation concern. Preserving small habitat patches will, therefore, be key to sustain biodiversity amidst ongoing environmental crises.     

Relative influence of size,connectivity and disturbance history on plant species richness and assemblages in fragmented grasslands

Questions: What is the relative influence of size, connectivity and disturbance history on plant species richness and assemblages of fragmented grasslands? What is the contribution of small fragments to the conservation of native species pool of the region? Location: Tandilia's Range, Southern Pampa, Argentina. Methods: Cover of plants was registered within 24 fragments of tall‐tussock grassland remnants within an agricultural landscape using modified Whittaker nested sampling. We analysed the influence of site variables related to disturbance history (canopy height, litter thickness) and fragment variables (size, connectivity) on species richness (asymptotic species richness, slope of the species–area curve) as well as on species assemblages by multiple regressions analysis and canonical correspondence analyses, respectively. Cumulative area was used for analysing whether small fragments or large fragments are more important to species diversity in the landscape. Results: Asymptotic species richness was significantly influenced by site variables, in particular by Paspalum quadrifarium's canopy height, but not by fragment variables. Species assemblages were also affected by site variables (12.2% of total variation), but no additional portion of the species assemblage variability was significantly explained by fragment size and connectivity. Sampling of several small fragments rendered more exotic and native species than sampling of few large fragments of the same total area. Conclusions: Our results agree with previous studies reporting low sensitivity of species diversity to size and isolation of grassland fragments in fragmented landscapes and high sensitivity of species diversity to local variables. The higher capture of regional native species pool by small grassland fragments than by few larger ones of equivalent accumulated area highlights the value of small fragments for conservation.     

Effects of habitat fragmentation and black-tailed prairie dogs on urban avian diversity

Urbanization and habitat fragmentation have the potential to influence bird communities. In addition, these phenomena, as well as ongoing lethal control measures, have also greatly reduced the range of the black-tailed prairie dog (Cynomys ludovicianus) since the beginning of the 20th century. Although prairie dogs are highly interactive species that can influence avian communities, few studies have investigated whether these interactions persist in urban settings. Our goal was to investigate the relative impacts of habitat fragmentation and prairie dogs on bird communities within an urban matrix. We performed bird surveys on 20 habitat fragments (10 colonized by prairie dogs, 10 uncolonized by prairie dogs) distributed throughout the Denver metropolitan area, and calculated Shannon–Weiner diversity and richness of all birds and native species, as well as total counts of grassland birds and raptors. Diversity, richness, and counts of many species increased with increasing fragment connectivity, and decreased on fragments isolated for longer periods of time. Avian diversity and richness did not differ between fragments with and without prairie dogs, suggesting that this element of the ecological role of prairie dogs is not fully retained in urban habitat. Future studies of the role of prairie dogs as keystone species in urban systems should include other taxa as well as consider the influence of the urban matrix surrounding prairie dog habitat. Our results emphasize that conservation of urban avian diversity should focus on landscape connectivity as well as local habitat features.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

宏生态尺度上景观破碎化对物种丰富度的影响

生物多样性的地理格局及其形成机制是宏生态学与生物地理学的研究热点。大量研究表明,景观尺度上的生境破碎化对物种多样性的分布格局具有重要作用,但目前尚不清楚这种作用是否足以在宏生态尺度上对生物多样性地理格局产生显著影响。利用中国大陆鸟类和哺乳动物的物种分布数据,在100 km×100 km网格的基础上生成了这两个类群生物的物种丰富度地理格局,进一步利用普通最小二乘法模型和空间自回归模型研究了物种丰富度与气候、生境异质性、景观破碎化的相关关系。结果表明,景观破碎化因子与鸟类和哺乳动物的物种丰富度都具有显著的关联关系,其方差贡献率可达约30%—50%(非空间模型)和60%—80%(空间模型),略低于或接近于气候和生境异质性因子。方差分解结果显示,景观破碎化因子与气候和生境异质性因子的方差贡献率的重叠部分达20%—40%。相对鸟类而言,景观破碎化对哺乳动物物种丰富度的地理格局具有更高的解释率。     

Relative contributions of local and regional factors to species richness and total density of butterflies and moths in semi-natural grasslands

Metapopulation theory predicts that species richness and total population density of habitat specialists increase with increasing area and regional connectivity of the habitat. To test these predictions, we examined the relative contributions of habitat patch area, connectivity of the regional habitat network and local habitat quality to species richness and total density of butterflies and day-active moths inhabiting semi-natural grasslands. We studied butterflies and moths in 48 replicate landscapes situated in southwest Finland, including a focal patch and the surrounding network of other semi-natural grasslands within a radius of 1.5 km from the focal patch. By applying the method of hierarchical partitioning, which can distinguish between independent and joint contributions of individual explanatory variables, we observed that variables of the local habitat quality (e.g. mean vegetation height and nectar plant abundance) generally showed the highest independent effect on species richness and total density of butterflies and moths. Habitat area did not show a significant independent contribution to species richness and total density of butterflies and moths. The effect of habitat connectivity was observed only for total density of the declining butterflies and moths. These observations indicate that the local habitat quality is of foremost importance in explaining variation in species richness and total density of butterflies and moths. In addition, declining butterflies and moths have larger populations in well-connected networks of semi-natural grasslands. Our results suggest that, while it is crucial to maintain high-quality habitats by management, with limited resources it would be appropriate to concentrate grassland management and restoration to areas with well-connected grassland networks in which the declining species currently have their strongest populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Landscape‐ and small‐scale determinants of grassland species diversity: direct and indirect influences

Species richness is influenced both by mechanisms occurring at landscape scales, such as habitat availability, and local‐scale processes, that are related to abiotic conditions and plant–plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined effect of factors measured at multiple spatial scales. In this study, we quantified the simultaneous influence of historical landscape‐scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small‐scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had significant direct and indirect (i.e. through the modification of another factor), effects on species richness. At the landscape scale, we found a direct positive influence of historical habitat availability on species richness, and indirect positive influence of past human population (via its effects on historical habitat availability). At small scales, we found a positive direct influence of light heterogeneity and shrub cover on species richness. Conversely, we found that small‐scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative effect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively influenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also influenced species richness. In contrast to the positive heterogeneity–diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. The combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.     

Functional composition and phenology of fruit-feeding butterflies in a fragmented landscape: variation of seasonality between habitat specialists

For butterflies, tolerance to the matrix may be an important criterion of habitat occurrence in fragmented landscapes. Here we examine the relative effects of habitat fragmentation and the surrounding agricultural matrix on the functional composition of fruit-feeding butterflies of the Atlantic rain forest in southeastern Brazil. Generalized linear models were used to detect the effects of landscape metrics on butterfly richness and abundance of the total assemblage and functional groups. Circular statistics were used to analyze the patterns of monthly abundance of the total assemblage and functional groups in the forest remnants and the surrounding matrices. In total, 650 butterflies representing 57 species were captured; species composition differed significantly between the forest fragments and the surrounding matrices. We recorded 22 forest specialists, 18 matrix specialists, 11 common species with matrix preference and six common species with forest preference. Forest connectivity favored the richness of forest specialists, while habitat fragmentation enhances the richness and abundance of matrix-tolerant species. Circular analysis revealed that forest specialists were more abundant in the rainy season while matrix-tolerant species proliferated in the dry season. Although maintaining connectivity of forest fragments may increase the mobility and dispersion of forest species, our results showed that landscape fragmentation modify butterfly assemblage by promoting an increase of matrix tolerant species with detriment of forest specialists.     

The role of habitat connectivity and landscape geometry in experimental zooplankton metacommunities

Theory predicts that inter-patch dispersal rates and patterns of patch heterogeneity both have the potential to alter patterns of local and regional species diversity. To test this, we manipulated both rates of habitat connectivity and the geometric arrangement of habitat heterogeneity within regions of experimental zooplankton communities. We found no effects of habitat geometry on any metric of species diversity or composition. Additionally, we found no effect of habitat connectivity rate on local species diversity. We did, however, find that increasing connectivity led to a decrease in regional diversity, as well as an increase in the percent similarity of local communities within regions. Of all of the species in these communities, the relatively large cladoceran Ceriodaphnia reticulata significantly responded to the treatments, and had a higher probability of achieving high densities when connectance was high. As such, we suggest that this species played a large role in driving the increased local community similarity and decreased regional species richness as connectivity increased. These findings are in opposition to previous experimental studies of metacommunities, but support the notion that increased connectance among local patches may decrease regional diversity when patches are heterogeneous.     

Forest Fragment and Breeding Habitat Characteristics Explain Frog Diversity and Abundance in Singapore

Habitat loss and fragmentation can have severe negative and irreversible effects on biodiversity. We investigated the effects of forest fragmentation on frog diversity in Singapore because of its high rates of deforestation and the demonstration that frogs are some of the most sensitive species to habitat degradation. We surveyed frog species in 12 forest fragments varying from 11 to 935 ha. We compared differences in species richness, abundance, and Shannon's index in relation to forest fragment size, connectivity (distance between fragments), and breeding habitat heterogeneity. A total of 20 species from 12 genera and five families were encountered in 12 fragments. Larger fragments and those closer to larger fragments had higher species richness. Abundance, however, was not correlated with forest area or connectivity, but we found fewer individual frogs in the larger fragments. We also found that breeding habitat heterogeneity best explained frog species diversity and abundance in forest fragments. Fragments with a high diversity of breeding habitats had more species. We found no evidence to suggest that abundance and diversity are strongly correlated, particularly in disturbed areas, but that breeding habitat heterogeneity is an under-appreciated factor that should be considered when prioritizing areas for anuran conservation. Enriching breeding habitat heterogeneity, creating corridors between fragments, and reforesting degraded areas are some of the most beneficial strategies for preserving urban frog biodiversity.     

Mixed effects of habitat fragmentation on species richness and community structure in a microarthropod microecosystem

Abstract.  1. Theory is unclear about the optimal degree of isolation of habitat fragments where the aim is to maximise species richness. In a field-based microecosystem of Collembola and predatory and non-predatory mites, moss patches of the same total area were fragmented to varying degrees. The habitat was left for several months to allow the communities to approach a new state of equilibrium.
2. The species richness (in particular of predatory mites) of a given area of habitat was greater when it was part of a large mainland area than part of an island, in agreement with theory.
3. Conversely, species richness and abundance were largely unaffected by fragmentation of a fixed area of island habitat. In this case, it is suggested here that the advantages of several small patches (e.g. reduced impact of environmental stochasticity, wider range of habitats overall) were equally balanced by the advantages of a single large patch (e.g. reduced effect of demographic stochasticity, wider range of habitats within a single patch, reduced edge effect), or that both effects were small.
4. The shapes of rank–abundance curves were similar among the levels of fragmentation of a fixed area of island habitat, implying that fragmentation had little impact on community structure. Conversely, the species composition of non-predatory mites varied weakly, but significantly, by fragmentation.     

Habitat fragmentation and species diversity in competitive communities

Habitat loss is one of the key drivers of the ongoing decline of biodiversity. However, ecologists still argue about how fragmentation of habitat (independent of habitat loss) affects species richness. The recently proposed habitat amount hypothesis posits that species richness only depends on the total amount of habitat in a local landscape. In contrast, empirical studies report contrasting patterns: some find positive and others negative effects of fragmentation per se on species richness. To explain this apparent disparity, we devise a stochastic, spatially explicit model of competitive species communities in heterogeneous habitats. The model shows that habitat loss and fragmentation have complex effects on species diversity in competitive communities. When the total amount of habitat is large, fragmentation per se tends to increase species diversity, but if the total amount of habitat is small, the situation is reversed: fragmentation per se decreases species diversity.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.     

Landscape-scale connectivity and fragment size determine species composition of grassland fragments

As a consequence of agricultural intensification and habitat fragmentation since the mid-20th century, biological diversity has declined considerably throughout the world, particularly in Europe. We assessed how habitat and landscape-scale heterogeneity, such as variation in fragment size (small vs. large) and landscape configuration (measured as connectivity index), affect plant and arthropod diversity. We focused on arthropods with different feeding behaviour and mobility, spiders (predators, moderate dispersal), true bugs (mainly herbivores and omnivores with moderate dispersal), wild bees (pollinators with good dispersal abilities), and wasps (pollinators, omnivores with good dispersal abilities). We studied 60 dry grassland fragments in the same region (Hungarian Great Plain); 30 fragments were represented by the grassland component of forest-steppe stands, and 30 were situated on burial mounds (kurgans). Forest-steppes are mosaics of dry grasslands with small forests in a matrix of plantation forests. Kurgans are ancient burial mounds with moderately disturbed grasslands surrounded by agricultural fields. The size of fragments ranged between 0.16–6.88 ha (small: 0.16–0.48 ha, large: 0.93–6.88 ha) for forest-steppes and 0.01–0.44 ha (small: 0.01–0.10 ha and large: 0.20–0.44 ha) for kurgans. Fragments also represented an isolation gradient from almost cleared and homogenous landscapes, to landscapes with relatively high compositional heterogeneity. Fragment size, connectivity, and their interaction affected specialist and generalist species abundances of forest-steppes and kurgans. Large fragments had higher species richness of ground-dwelling spiders, and the effect of connectivity was more strongly positive for specialist arthropods and more strongly negative for generalists in large than in small fragments. However, we also found a strong positive impact of connectivity for generalist plants in small kurgans in contrast to larger ones. We conclude that besides the well-known effect of enhancing habitat quality, increasing connectivity between fragments by restoring natural and semi-natural habitat patches would help to maintain grassland biodiversity.     

Environmental and vegetation variables have a greater influence than habitat fragmentation in structuring lizard communities in remnant urban bushland

Abstract The expansion of urban areas and adjacent farming land into natural landscapes modifies habitats and produces small isolated pockets of native vegetation. This fragmentation of the natural habitat subdivides animal communities, reduces population sizes and increases vulnerability to extinction. In this paper we investigate whether fragmentation decreases lizard species richness, composition, overall abundance and abundance at the species level. Urban remnants consisting of five small (< 10 ha) and four large (> 10 ha) fragments of natural bushland were paired with continuous bushland areas located near Hobart, Tasmania, Australia. These remnants were surveyed six times, using pitfall traps, from November 2001 to March 2002. Lizard species richness and abundance were not significantly influenced by habitat fragmentation or fragment size. Egernia whitii was the only lizard species significantly influenced by fragment size, and was only present in large fragments and continuous bush. Vegetation type and structure as well as environmental variables (geology and aspect) influenced the structure of reptile communities. Lizard species that were able to use a number of different habitat types were found to persist at most sites, irrespective of fragment size. Edge environment did not significantly influence lizard species richness or abundance in remnant areas. Lizard species richness was significantly lower in sites that had a high ratio of exotic to native plant species. Therefore, if remnants continue to be invaded by exotic plants, lizard species that require native plant communities will become increasingly vulnerable to local extinction. Our results suggest that lizard species requiring specialized habitats, such as E. whitii, may persist in large urban remnants rather than small urban remnants because large reserves are more likely to encompass rare habitats, such as rocky outcrops. Habitat heterogeneity, rather than size, may be the key to their persistence.