Relationship of butterfly diversity with nectar plant species richness in and around the Aokigahara primary woodland of Mount Fuji,central Japan

We examined the relationships between the diversities of vegetation, adult nectar plants, and butterflies in and around the Aokigahara primary woodland on the northwestern footslopes of Mount Fuji, central Japan. The results showed that the nectar resource utilization by adult butterflies was significantly biased to herbaceous plants, especially to perennials, compared to woody species, although most of the study area was in and near a primary woodland. There were greater nectar plant species in sites with greater plant species richness. Among the butterfly community indices analyzed, the strongest correlation was detected between butterfly species richness and nectar plant species richness at each site. Another close correlation was detected between the species richness of nectar plants and herbaceous plants at each site. These results suggest that herbaceous plant species richness in a habitat plays a central role in its nectar plant species richness, and the nectar plant richness is a highly important factor supporting its adult butterfly species richness. Consequently, we propose that the maintenance and management of herbaceous plant species richness in a butterfly habitat, which lead to those of its nectar plant species richness, are very important for conservation of butterfly diversity even in and around woodland landscapes of temperate regions.     

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.     

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.     

Connectivity compensates for low habitat quality and small patch size in the butterfly <Emphasis Type="Italic">Cupido minimus</Emphasis>

Habitat size, habitat isolation and habitat quality are regarded as the main determinants of butterfly occurrence in fragmented landscapes. To analyze the relationship between the occurrence of the butterfly Cupido minimus and these factors, patch occupancy of the immature stages in patches of its host plant Anthyllis vulneraria was studied in the nature reserve Hohe Wann in Bavaria (Germany). In 2001 and 2002, 82 A. vulneraria patches were surveyed for the presence of C. minimus larvae. The occurrence was largely affected by the size of the food plant patches. In a habitat model that uses multiple logistic regression, the type of management and habitat connectivity are further determinants of species distribution. Internal and temporal validation demonstrate the stability and robustness of the developed habitat models. Additionally, it was proved that the colonization rate of C. minimus was significantly influenced by the distance to the next occupied Anthyllis patch. Concerning long-term survival of (meta-) populations in fragmented landscapes, the results show that lower habitat quality may be compensated by higher connectivity between host plant patches. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Depauperation and divergence of plant‐specialist herbivore assemblages in a fragmented tropical landscape

1. Our understanding of the structure and spatial organisation of biological assemblages in human‐modified tropical landscapes has critical importance to improve conservation actions. Investigations on this topic have focused on local (α) diversity patterns, overlooking the changes in species turnover (β diversity) between sites, and its consequences on total (γ) diversity. 2. This study assessed the differences in α, β and γ diversities of galling insects and their host plants (saplings) in a fragmented Atlantic forest landscape in northeast Brazil. Both assemblages were recorded in 30 plots (total of 0.1 ha for each forest type) located in the interior and on the edges of a large fragment and small forest fragments (10 plots per forest type). 3. α diversity of host plants and galling insect assemblages was significantly higher in interior (reference) plots than in edge and fragment plots. Yet, both assemblages showed higher β diversity in fragment and edge plots than in reference plots – a finding potentially associated with the hyperdynamism of fragmented forests and consistent with the landscape divergence hypothesis. 4. However, biotic differentiation of host plant and galling insects was not great enough to compensate the loss of α diversity, and thus γ diversity, because most host plant and galling insect species in forest fragments were also registered in reference plots. Our findings indicate that, despite each small forest fragment being very dissimilar from each other, they have low importance for the conservation of plant assemblages and their specialized herbivores at landscape scale.     

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.     

Extensively managed or abandoned urban green spaces and their habitat potential for butterflies

Urban green spaces are often subject to intensive management and therefore only provide habitat for very few plant and animal species. Many studies have already proven that less intensive management of near-natural grassland habitats can increase species diversity, for example of butterflies. However, there is still a lack of knowledge as to what extent less intensive management of urban green spaces can promote urban butterfly diversity. Therefore, we examined parameters that determine the number of butterfly species in less maintained or even abandoned urban green spaces. We expected butterfly species richness and observation numbers to be affected by different management regimes. Butterfly surveys were carried out on 56 sites in the city of Hannover, Germany, in three consecutive years. The sites were subject to different management regimes. We explored parameters such as area size, the diversity of nectar plant species and habitat diversity for each site. A total of 34 species were recorded, with a minimum of 0 and a maximum of 17 species per site and year. The extreme drought and heat in 2018 clearly influenced the results of our study and this was reflected in massive decreases of observation numbers for some species. Less intensive management showed preferable effects in comparison to intensive management, but area size and nectar plants were more important factors influencing butterfly species richness and number of observations. Our results suggest that site parameters such as area size, nectar plant species richness or number of habitat types had stronger influences on butterflies than variations of extensive management. We conclude that less intensive urban green space management for good butterfly habitats is not a question of grazing, mowing or temporary abandonment, but instead must aim to increase nectar plant and habitat diversity.     

Local and landscape effects on butterfly density in northern Idaho grasslands and forests

Understanding butterfly response to landscape context can inform conservation management and planning. We tested whether local-scale resources (host and nectar plants, canopy cover) or landscape context, measured at two scales, better explained the densities of four butterfly species. The density of Coenonympha tullia, which has host plants strongly associated with grassland habitats, was positively correlated with the amount of grassland in 0.5- and 1-km radius landscapes and only occurred in forests when they bordered grasslands. For the other species, Celastrina ladon, Cupido amyntula, and Vanessa cardui, local-scale resources better explained butterfly densities, emphasizing the importance of local habitat quality for butterflies. These three species also used host plants that were distributed more heterogeneously within and among habitat types. Our findings demonstrate the importance of host plant spatial distributions when determining the scale at which butterfly density relates to resources, and we recommend that both these distributions and landscape context be evaluated when developing butterfly monitoring programs, managing for species of concern, or modeling potential habitat.     

Effects of environment,habitat configuration and forest continuity on the distribution of forest plant species

In present day European landscapes many forest plant species are restricted to isolated remnants of a formerly more or less continuous forest cover. The two major objectives of this study were (1) to determine the relative importance of habitat quality (mainly in terms of soil parameters), habitat configuration (patch size and isolation) and habitat continuity for the distribution of herbaceous forest plant species in a highly fragmented landscape and (2) to examine if groups of species with different habitat requirements are affected differently. Deciduous forest patches in northwestern Germany were surveyed for the presence of a large set of forest species. For each patch, habitat quality, configuration and continuity were determined. Data were analysed by Redundancy Analysis with variation partitioning for effects on total species composition and multivariate logistic regression for effects on individual species, for two different data sets (base‐rich and base‐poor forest patches). Overall, we found strong effects of habitat quality (particularly of soil pH, water content and topographic heterogeneity in the base‐rich forest patches; and of calcium content and disturbance in the base‐poor patches), but only relatively weak effects of habitat configuration and habitat continuity. However, a number of species were positively affected by patch area and negatively affected by patch isolation. Furthermore, the relative importance of habitat configuration tended to be higher for species predominantly growing in closed forests compared to species occurring both in the forest and in the open landscape.     

Contrasting effects of habitat quantity and quality on moth communities in fragmented landscapes

Habitat loss is commonly identified as a major threat to the loss of global biodiversity. In this study, we expand on our previous work by addressing the question of how lepidopteran species richness and composition vary among remnants of North American eastern deciduous forest located within agricultural or pastoral landscapes. Specifically, we tested the relative roles of habitat quantity (measured as stand area and percent forest in the greater landscape) and habitat quality (measured as tree species diversity) as determinants of moth species richness. We sampled >19 000 individuals comprising 493 moth species from 21 forest sites in two forested ecoregions. In the unglaciated Western Allegheny Plateau, the species richness of moths with woody host plants diminished as forest stand size and percent forest in the landscape decreased, but the total species richness and abundance of moths were unaffected by stand size, percent forest in the landscape, or tree species diversity. In contrast, the overall species richness and abundance of moths in the glaciated North Central Tillplain were affected primarily by tree species diversity and secondarily by forest size. Higher tree species diversity may reduce species loss from smaller forest stands, suggesting that small, diverse forests can support comparable numbers of species to those in less diverse, large stands. Smaller forests, however, contained a disproportionate number of moth species that possess larvae known to feed on herbaceous vegetation. Thus, although woody plant feeding moths are lost from forests with changes in stand area, new species appear capable of recolonizing smaller fragments from the surrounding habitat matrix. Our study further suggests that when species replacement occurs, local patch size and habitat quality may be more important than landscape context in determining the community structure of forest Lepidoptera.     

Butterflies as an indicator group of riparian ecosystem assessment

Riparian ecosystems play an important role in modulating a range of ecosystem processes that affect aquatic and terrestrial organisms. Butterflies are a major herbivore in terrestrial ecosystems and are also common in riparian ecosystems. Since butterflies use plants for larval food and adult nectar sources in riparian ecosystems, butterfly diversity can be utilized to evaluate riparian ecosystems. We compiled butterfly data from 33 sites in three riparian ecosystem types across the country and compared butterfly diversity in terms of number of species and quality index in relation to riparian environmental variables. Number of butterfly and plant species was not different among three riparian habitat types. Additionally, there was no significant ecological variable to distinguish the butterfly communities on three riparian habitats. Non-metric multi-dimensional scaling ordination showed that butterfly communities in three riparian ecosystem types differed from each other, and butterfly riparian quality index was the main variable for butterfly assemblages. Five indicator species for moor and another five species for riverine riparian ecosystems were identified. Three and one indicator species for moor and riparian ecosystems, respectively, were plant specialists, while 44 butterflies were general feeders, feeding on a wide range of hostplants in several habitats. These results suggest that butterfly species use actively riparian habitats for nectar and larval food, and the butterfly riparian quality index can be employed to track faunal change in riparian habitats, which are frequently threatened by disturbances such as water level and climate changes, and invasive species.     

Barriers to dispersal of rain forest butterflies in tropical agricultural landscapes

Fragmentation of natural habitats can be detrimental for species if individuals fail to cross habitat boundaries to reach new locations, thereby reducing functional connectivity. Connectivity is crucial for species shifting their ranges under climate change, making it important to understand factors that might prevent movement through human‐modified landscapes. In tropical regions, rain forests are being fragmented by agricultural expansion, potentially isolating populations of highly diverse forest‐dependent species. The likelihood of crossing habitat boundaries is an important determinant of species dispersal through fragmented landscapes, and so we examined movement across rain forest‐oil palm plantation boundaries on Borneo by using relatively mobile nymphalid butterflies as our model study taxon. We marked 1666 individuals from 65 species, and 19 percent (100/527) of recaptured individuals crossed the boundary. Boundary crossing was relatively frequent in some species, and net movement of individuals was from forest into plantation. However, boundary crossing from forest into plantation was detected in less than 50 percent (12/28) of recaptured species and was dominated by small‐sized butterfly species whose larval host plants occurred within plantations. Thus, while oil palm plantations may be relatively permeable to some species, they may act as barriers to the movement of forest‐dependent species (i.e., species that require rain forest habitat to breed), highlighting the importance of maintaining forest connectivity for conserving rain forest species.     

Ecological Restoration Treatments Increase Butterfly Richness and Abundance: Mechanisms of Response

Few ecosystem restoration studies evaluate whether arthropods are important components of ecosystem recovery. We tested the hypothesis that ponderosa pine restoration treatments would increase adult butterfly species richness and abundance as a direct result of increased understory diversity and abundance. To examine mechanisms that potentially affect adult butterfly distribution, we quantified host plant frequency, nectar plant abundance, and insolation (light intensity) in restoration treatment and control forests. This study is unique, because this is the first invertebrate monitoring in ponderosa pine forest restoration treatments in the U.S. Southwest and also because these treatments are the first replicated ponderosa pine restoration treatments at a landscape scale. Three patterns emerged: (1) butterfly species richness and abundance were 2 and 3 times greater, respectively, in restoration treatment units than in paired control forests 1 year after treatment, and 1.5 and 3.5 times greater, respectively, 2 years after treatment, ordination of control and treatment sampling units using butterfly assemblages showed significant separation of control and restoration treatment units after restoration treatment; (2) host plant and nectar plant species richness showed little difference between treated and control forests even 2 years after treatment; and (3) insolation (light intensity) was significantly greater in treated forests after restoration. We suggest that changes in the butterfly assemblage may occur due to light intensity effects before plant community changes occur or can be detected. Butterfly assemblage differences will have additional cascading effects on the ecosystem as prey for higher trophic levels and through plant interactions including herbivory and pollination.     

Offspring performance and recruitment of the pioneer tree Acacia caven (Fabaceae) in a fragmented subtropical dry forest

The process of habitat fragmentation results in the breaking apart of originally continuous habitats, causing multiple changes in biotic and abiotic interactions. Alterations in resource availability and in mutualistic and antagonistic plant–animal interactions may impact plant offspring quantity and quality. Currently, several old fragmented systems evidence a process of flora homogenization, where shade‐tolerant species are replaced by pioneer light‐demanding species. Notably, the relationship between quantity and quality parameters of plant offspring production and the successful recruitment of pioneer species in fragmented forests has been poorly explored. Here, we assess population size, sapling recruitment and offspring performance of one of the most widespread tree species of subtropical South America, the native pioneer Acacia caven (Fabaceae). Population size of adults and saplings increased from small to continuous forests, whereas the sapling recruitment per adult tree (sapling/adult ratio) showed no significant differences among forests of different size. Seedling performance was negatively related to forest area and population size, implying potential superior competitive ability of seedlings produced in smaller populations compared to larger ones. Our results show that A. caven is resilient to habitat fragmentation effects, which may be ascribed to a set of advantageous ecological traits such as outcrossing, massive flowering, generalist pollination, drought resistance, rapid growth and re‐sprouting. Thus, this pioneer tree benefits from the availability of vacant sites and resources released by declining plant populations of other species, eventually becoming the dominant species in fragmented habitats. Pioneer native plant species with ecological traits such as A. caven may represent the silent successful survivors and new colonizers of fragmented habitats, the ubiquitous landscapes of the future.     

Differing responses to landscape change: implications for small mammal assemblages in forest fragments

Human modification of landscapes typically results in many species being confined to small, isolated and degraded habitat fragments. While fragment size and isolation underpin many studies of modified landscapes, vegetation characteristics are less frequently incorporated. The relative influence of biogeographic (e.g. size, isolation) and vegetation parameters on assemblages is poorly understood, but critical for conservation management. In this study, a multiple hypothesis testing framework was used to determine the relative importance of biogeographic and vegetation parameters in explaining the occurrence of an assemblage of small mammals in 48 forest fragments in an agricultural landscape in south-eastern Australia. Fragment size and vegetation characteristics were consistently important predictors of occurrence across species. In contrast, fragment isolation was important for just one native species. Differing abilities of species to move through the landscape provide a reasonable explanation for these results. We conclude that for effective conservation of assemblages, it is important to: (1) consider differing responses of species to landscape change, and (2) move beyond a focus primarily on spatial attributes (size, isolation) to recognise that landscape change also has profound effects on habitat composition and quality.     

The asymmetric relationships of the distribution of conspecific saplings and adults in forest fragments

随着土地利用方式变化的加剧,生境片段化已成为影响植物多样性的主要因子之一。通常,当成年树个体的密度越高,其周边同种幼树个体的存活率可能会下降,从而为其它物种提供了空间和资源,进而可以维持较高的局域物种多样性。因此,同种成年树和幼树个体的空间分布格局关系和作用强度可以调节植物多样性。然而,对于在片段化森林中,同种成年树和幼树个体空间分布关系的研究却很少报道,迄今尚不清楚片段化景观中同种个体的空间分布关系与物种多样性之间的联系。本研究选择千岛湖陆桥岛屿系统中的27个岛屿,基于岛屿上幼树和成年树个体的空间分布数据,利用混合效应模型分析它们之间的作用强度。同种幼树和成年树个体的空间作用强度越大,说明它们之间的负相互作用越强,即幼树和成年树个体空间分布越分散。此外,本研究分析了岛屿属性(岛屿面积、与大陆的距离和与最近岛屿的距离)与同种个体空间作用强度及物种多样性之间的关系。结果表明,同种个体的空间作用强度随着与最近岛屿距离的增加而增加。同时,物种多样性随着同种个体的空间作用强度的增加而显著增加,且岛屿面积和同种个体的空间作用强度分别解释了岛屿间物种多样性差异的26%和6%,共同解释了8%。耐阴种和非常见种比非耐阴种和常见种的同种幼树和成年树的空间分布更为分散。本研究表明,同种个体的空间分布可能会影响多度较低物种在片段化森林中的生存,反映了生物相互作用对于维持片段化森林中的植物多样性具有重要作用。 本研究也强调在检验同种密度制约时应考虑森林之间的连接度。     

Leaf-cutting ants negatively impact the regeneration of the Caatinga dry forest across abandoned pastures

The mechanisms affecting forest regeneration in human-modified landscapes are attracting increasing attention as tropical forests have been recognized as key habitats for biodiversity conservation, provision of ecosystem services, and human well-being. Here we investigate the effect of the leaf-cutting ants (LCA) Atta opaciceps on regenerating plant assemblages in Caatinga dry forest. Our study encompassed 15 Atta opaciceps colonies located in landscape patches with a gradient of forest cover from 8.7% to 87.8%, where we monitored regenerating individuals (seedlings and saplings of woody and herbaceous plants) in different habitats (nests, foraging areas, and control areas) over one year. We recorded 2,977 regenerating plant individuals, distributed among 55 species from 23 families. Herbaceous plants represented 82.1% and 58.2% of the total number of individuals and species, respectively. Species richness of both the whole and herbaceous plant assemblages increased along the forest cover gradient, but without difference between the habitats. Total plant abundance was highest in control areas followed by foraging areas and nests and this pattern held for both woody and herbaceous plants. Although forest cover did not influence the abundance of herbaceous plants and the whole plant assemblage, it positively affects woody plant abundance across control areas. Forest cover and habitat changed species composition of both the entire regenerating and the herbaceous assemblages. These results together indicate that LCA negatively impact regenerating plant assemblages, particularly in those sites with increased forest cover. As LCA proliferate in human-modified landscapes, they may prevent plant regeneration of disturbed areas.     

Species‐habitat relationships and ecological correlates of butterfly abundance in a transformed tropical landscape

Tropical butterfly conservation strategies often focus on total and/or common species richness to assess the conservation value of a patch or habitat. However, such a strategy overlooks the unique dynamics of rare species. We evaluated the species‐habitat relationships of 209 common, intermediate, and rare butterfly species (including morphospecies) across four habitat types (mature, degraded, or fragmented forest, and urban parks) and two patch sizes (<400 ha, ≥400 ha) in Singapore. Common species richness was consistent across habitat types. Intermediate species richness declined by more than 50 percent in urban parks (relative to all forest habitats), and rare species richness was reduced by 50 percent in degraded and fragmented forest and by 90 percent in urban parks (relative to mature forest). Large patches had comparable overall richness to small patches, but they supported more rare species and three times as many habitat‐restricted species over a similar area. Importantly, a number of rare species were confined to single small patches. Mixed‐effects regression models were constructed to identify habitat and ecological/life history variables associated with butterfly abundance. These models revealed that species with greater habitat specialization, rare larval host plants, few larval host plant genera, and narrow global geographic ranges were more likely to be rare species. Overall, these results demonstrate that the richness of habitat‐restricted and rare species do not follow the same spatial distribution patterns as common species. Therefore, while conserving mature forests is key, effective butterfly conservation in a transformed landscape should take into account rare and habitat‐restricted species.     

Contemporary habitat loss reduces genetic diversity in an ecologically specialized butterfly

Aim This study investigated the influence of contemporary habitat loss on the genetic diversity and structure of animal species using a common, but ecologically specialized, butterfly, Theclinesthes albocincta (Lepidoptera: Lycaenidae), as a model. Location South Australia. Methods We used amplified fragment length polymorphism (AFLP) and allozyme datasets to investigate the genetic structure and genetic diversity among populations of T. albocincta in a fragmented landscape and compared this diversity and structure with that of populations in two nearby landscapes that have more continuous distributions of butterflies and their habitat. Butterflies were sampled from 15 sites and genotyped, first using 363 informative AFLP bands and then using 17 polymorphic allozyme loci (n = 248 and 254, respectively). We complemented these analyses with phylogeographic information based on mitochondrial DNA (mtDNA) haplotype information derived from a previous study in the same landscapes. Results Both datasets indicated a relatively high level of genetic structuring across the sampling range (AFLP, F_ST = 0.34; allozyme, F_ST = 0.13): structure was greatest among populations in the fragmented landscape (AFLP, F_ST = 0.15; allozyme, F_ST = 0.13). Populations in the fragmented landscape also had significantly lower genetic diversity than populations in the other two landscapes: there were no detectable differences in genetic diversity between the two continuous landscapes. There was also evidence (r² = 0.33) of an isolation by distance effect across the sampled range of the species. Main conclusions The multiple lines of evidence, presented within a phylogeographic context, support the hypothesis that contemporary habitat fragmentation has been a major driver of genetic erosion and differentiation in this species. Theclinesthes albocincta populations in the fragmented landscape are thus likely to be at greater risk of extinction because of reduced genetic diversity, their isolation from conspecific subpopulations in other landscapes, and other extrinsic forces acting on their small population sizes. Our study provides compelling evidence that habitat loss and fragmentation have significant rapid impacts on the genetic diversity and structure of butterfly populations, especially specialist species with particular habitat preferences and poor dispersal abilities.     

The effects of habitat fragmentation on niche requirements of the marsh fritillary, Euphydryas aurinia, (Rottemburg, 1775) on calcareous grasslands in southern UK

The marsh fritillary, Euphydryas aurinia, has declined greatly in distribution across its range within Europe, resulting in its designation as a protected species under Annex II of the 1979 Bern Convention and the EC Habitats and Species Directive. The decline has been linked to a marked reduction in the extent of suitable calcareous and wet grassland habitats, habitats which have been lost through conversion of land to agriculture or urban areas, or reduced in quality due to inappropriate management. The UK is now one of the major strongholds for this butterfly in Europe, although much of the remaining habitat is small, isolated and highly fragmented. E. aurinia populations fluctuate greatly due to the combined effects of biotic (e.g. parasitoids) and abiotic (e.g. climate change) factors. We quantified the habitat associations of larval webs of E. aurinia on fragmented versus extensive (unfragmented) calcareous grassland habitat in southern England to test the hypothesis that habitat requirements of E. aurinia are more constrained within fragmented landscapes. Within both fragmented and unfragmented landscapes the quality and quantity of its main host plant in the UK, Succisa pratensis, was positively related to numbers of E. aurinia larval webs found. The sward height was also important at predicting the distribution of larval webs in both landscapes, although the heights were greater within sites in the unfragmented (≈20 cm) compared to fragmented (≈15 cm) landscape. We also found significant effects of elevation and the cover of bare ground on numbers of larval webs. Elevation was strongly correlated with the availability of host plant, whilst bare ground was only significant on sites within the fragmented landscape, showing a negative relationship with number of larval webs. Our results further emphasise the importance of not only maintaining the habitat quality of extant calcareous grassland sites for E. aurinia in the UK, but also increasing the size and connectivity of these sites to increase the chances and rate of (re)colonisation of unoccupied but suitable habitat. In addition, we show that the habitat requirements of E. aurinia on sites in a large unfragmented landscape may be less specific and thus require less extensive management than that required to create optimal conditions necessary at smaller, more isolated sites in fragmented landscapes.