The NATURA 2000 database as a tool in the analysis of habitat selection at large scales: factors affecting the occurrence of pine and stone martens in Southern Europe

Most studies on habitat selection among animals are conducted at local scales, whereas reliable determination of species requirements at larger spatial scales can be problematic. We used data available for NATURA 2000 sites to determine the habitat requirements of two relatively widespread and common species—pine marten Martes martes and stone marten Martes foina—in Southern Europe. Using presence-absence data, we applied statistical models at two spatial scales. At the local scale (within the dispersal distances of the species), sites occupied by martens were compared with unoccupied sites using buffers of different sizes, whereas at the regional scale, unoccupied sites were selected randomly. To adjust for spatial autocorrelation of data, penalised quasi-likelihood approximations were used. Both species of martens demonstrated preferences for areas with higher proportions of forest cover and steeper terrain. At the local, but not at the regional, scale, pine martens occurred at lower elevations, whilst stone martens occurred at higher elevations. We found that climatic variables (mean temperature, precipitation) had no significant effect on the presence of the martens. The results of our analyses generally confirmed findings of previous studies on habitat selection of both marten species in Southern Europe. This demonstrates the utility of data collected for NATURA 2000 sites for use in various analyses such as conservation planning and evaluating the impact of climate change on the distribution of animal species.     

A Method for Evaluating Outcomes of Restoration When No Reference Sites Exist

Ecological restoration typically seeks to shift species composition toward that of existing reference sites. Yet, comparing the assemblages in restored and reference habitats assumes that similarity to the reference habitat is the optimal outcome of restoration and does not provide a perspective on regionally rare off-site species. When no such reference assemblages of species exist, an accurate assessment of the habitat affinities of species is crucial. We present a method for using a species by habitat data matrix generated by biodiversity surveys to evaluate community responses to habitat restoration treatments. Habitats within the region are rated on their community similarity to a hypothetical restored habitat, other habitats of conservation concern, and disturbed habitats. Similarity scores are reinserted into the species by habitat matrix to produce indicator ( I ) scores for each species in relation to these habitats. We apply this procedure to an open woodland restoration project in north Mississippi (U.S.A.) by evaluating initial plant community responses to restoration. Results showed a substantial increase in open woodland indicators, a modest decrease in generalists historically restricted to floodplain forests, and no significant change in disturbance indicators as a group. These responses can be interpreted as a desirable outcome, regardless of whether species composition approaches that of reference sites. The broader value of this approach is that it provides a flexible and objective means of predicting and evaluating the outcome of restoration projects involving any group of species in any region, provided there is a biodiversity database that includes habitat and location information.     

Importance of soil nutrients in the distribution of forest communities on a large geographical scale

Aim Soil nutrient content plays a key role in plant growth through mineral nutrition and toxicity. Its impact on plant species and community distribution is studied on a large geographical scale through surrogates like topography or geology. We investigated the importance of soil pH and C:N ratio, as direct nutritional gradients, to determine, with climatic factors, the spatial distribution of plant communities over large territories. Location We studied the distribution of six beech habitats of the NATURA 2000 network throughout France (550,000 km²). Methods Models were calibrated with 2108 floristic plots classified in the NATURA 2000 system and including climatic and topographic variables and soil nutritional measurements carried out in a laboratory. Logistic regression was used to model habitat distribution according to environmental variables. Climatic layers, a digital elevation model and maps of soil pH and nitrogen content, created using plant indicator values and large floristic databases, were used to map the sites suitable for beech communities. Distribution models were evaluated with an independent set of 2091 phytosociological plots. Results pH and nitrogen supply were the key distribution drivers for four of the six beech communities on a national scale. Their use in the distribution models distinguished within homogeneous climatic territories a gradient of nutritional conditions from acidic areas, suitable for nutrient‐poor beech communities, to calcareous areas suitable for nutrient‐rich ones. Predicted maps of beech habitats fit the spatial distribution of validation plots. Main conclusions Soil pH and nitrogen supply strongly improve predictions of forest community distribution carried out with climatic variables on a broad geographical scale. They allow delineation of areas with nutritional conditions suitable for each community, as well as the realization of predictive high‐resolution maps over large areas useful for sustainable and conservation management. Nomenclature Tutin & Heywood (2001 ) Flora Europaea. Cambridge University Press, Cambridge.     

Integrating habitat diversity into species conservation in the Mediterranean mosaic landscape

Biodiversity conservation requires strategies that encompass a variety of land uses and habitat diversity. In this study, we used sites of high ecological interest identified on the basis of the distribution of priority plant and vertebrate species to assess the implications of habitat diversity for conservation management in the Mediterranean mosaic landscape. 40% of the priority species occur in open habitats that depend on continued human presence and low-intensity land-use activities. Furthermore, 70% of the sites have more than one species habitat and 15% of sites contain more than four different species habitats. By explicitly integrating localized habitat variation, conservation planning can address the multiple conservation issues at stake in regions of high landscape diversity to provide clear and effective management objectives.     

Predicting habitat distribution and frequency from plant species co-occurrence data

Aim Species frequency data have been widely used in nature conservation to aid management decisions. To determine species frequencies, information on habitat occurrence is important: a species with a low frequency is not necessarily rare if it occupies all suitable habitats. Often, information on habitat distribution is available for small geographic areas only. We aim to predict grid‐based habitat occurrence from grid‐based plant species distribution data in a meso‐scale analysis. Location The study was carried out over two spatial extents: Germany and Bavaria. Methods Two simple models were set up to examine the number of characteristic plant species needed per grid cell to predict the occurrence of four selected habitats (species data from FlorKart, http://www.floraweb.de ). Both models were calibrated in Bavaria using available information on habitat distribution, validated for other federal states, and applied to Germany. First, a spatially explicit regression model (generalized linear model (GLM) with assumed binomial error distribution of response variable) was obtained. Second, a spatially independent optimization model was derived that estimated species numbers without using spatial information on habitat distribution. Finally, an additional uncalibrated model was derived that calculated the frequencies of 24 habitats. It was validated using NATURA2000 habitat maps. Results Using the Bavarian models it was possible to predict habitat distribution and frequency from the co‐occurrence of habitat‐specific species per grid cell. As the model validations for other German federal states were successful, the models were applied to all of Germany, and habitat distribution and frequencies could be retrieved for the national scale on the basis of habitat‐specific species co‐occurrences per grid cell. Using the third, uncalibrated model, which includes species distribution data only, it was possible to predict the frequencies of 24 habitats based on the co‐occurrence of 24% of formation‐specific species per grid cell. Predicted habitat frequencies deduced from this third model were strongly related to frequencies of NATURA2000 habitat maps. Main conclusions It was concluded that it is possible to deduce habitat distributions and frequencies from the co‐occurrence of habitat‐specific species. For areas partly covered by habitat mappings, calibrated models can be developed and extrapolated to larger areas. If information on habitat distribution is completely lacking, uncalibrated models can still be applied, providing coarse information on habitat frequencies. Predicted habitat distributions and frequencies can be used as a tool in nature conservation, for example as correction factors for species frequencies, as long as the species of interest is not included in the model set‐up.     

The contribution of common and rare species to plant species richness patterns: the effect of habitat type and size of sampling unit

Understanding how overall patterns of spatial variation in species richness are affected by distributional patterns of species has been an area of growing concern. In the present study, we investigated the relative importance of common and rare species as contributors in overall plant species richness. We further examined if the effects of common or rare species in richness patterns are affected by the size of the sampling units and if the observed patterns hold at different habitats. We used a dataset of 5,148 higher plant species distributed across 16,114 sampling plots located in 240 sites of the NATURA 2000 network of Greece. We ranked all species based on the number of sites they occupied and we developed a common to rare and a rare to common sequence. We correlated those sequences with cumulative species distributions. We performed this analysis in nine different sizes of sampling units and in three different datasets referring to (a) all habitat types together, (b) coniferous habitats only and (c) alpine habitats only. Our analysis showed that despite the proportionally higher numbers of restricted species, widespread species make a greater contribution to overall richness patterns and that this observed pattern does not depend on the size of the sampling units. Moreover, the observed pattern stands for different habitat types. Our findings support the generality of this pattern and highlight the importance of widespread species as adequate indicators of biodiversity patterns at various habitat types. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecological and biodiversity gradients across alpine dry grassland habitats: implications for an endangered species

Dry grasslands are of great interest for nature conservation in Europe, because they have a central role in the conservation of numerous rare and endangered species. In this study carried out in the Brenta mountain group (Italian alps), we investigated the effect of environmental factors mainly controlled by topography, on the biodiversity trends across different dry grassland habitats where the threatened alpine stenoendemic Erysimum aurantiacum grows. Plant community data and ecological factors were analysed by means of a multi‐habitat CCA approach and by analysis of biodiversity gradients in 7 natural and semi‐natural habitats. We found that species turnover and biodiversity patterns vary as a function of multi‐factorial ecological gradients. For the single habitats, elevation gradient was the main factor explaining compositional variation, followed by inclination and proportion of exposed rock surface. Despite its endangered status, E. aurantiacum showed a relatively high degree of ecological plasticity across these semiarid grassland habitats that probably allows it to survive in different environments, including in some cases those impacted by human activities. This prompts for habitat‐ more than species‐level conservation actions. According to their characteristics and threats, habitat‐specific management practices are recommended for long term conservation of plant species communities in the different ecological niches.     

A review of dystrophic lake and pool habitat in Europe: An Irish perspective

Freshwater lakes and pools contained within peatlands are unique habitats that support rare and specialised species. Despite this, these ecosystems have been overlooked in conservation and management practices. One of these habitats, ‘3160 Natural dystrophic lakes and ponds’, is protected under the European Union (EU) Habitats Directive with a concerning proportion of these habitats having an “unfavourable-bad” or an “unfavourable-inadequate” conservation status across Europe. Our current understanding of the key physico-chemical and ecological features of this habitat is inadequate which is hindering the implementation of effective conservation measures. This review summarises the current knowledge of this protected lake habitat as defined under the EU Habitats Directive. With a focus on Ireland, we demonstrate how the current monitoring and assessment methods used to characterise and assess the structure and function and conservation status of this habitat, which relies largely on the use of macrophyte community composition and surrogate physico-chemical data collected under the EU Water Framework Directive, is ineffective. We propose the incorporation of further or alternative ecological metrics including, but not limited to, algae and macroinvertebrates which are needed to improve our understanding of the structure and function of this priority lake habitat. In addition, application of such data via ecological metrics would allow for the quantification of biodiversity and species rarity metrics which would aid in identifying sites of conservation importance.     

Butterfly diversity at the ecotone between agricultural and semi‐natural habitats across a climatic gradient

Aim Understanding the response of species to ecotones and habitat edges is essential to designing conservation management, especially in mosaic agricultural landscapes. This study examines how species diversity and composition change with distance from semi‐natural habitats, over ecotones into agricultural fields, and how within‐site patterns of community transition change across a climatic gradient and differ between crop types. Location A total of 19 sites in Israel where semi‐natural habitats border agricultural fields (wheat fields or olive groves) distributed along a sharp climatic gradient ranging between 100 and 800 mm mean annual rainfall. Methods  We performed butterfly surveys in 2006. We analysed species richness (α‐diversity), diversity, community nestedness and species turnover (β‐diversity) within sites and between sites (γ‐diversity). We also assessed where species of conservation concern occurred. Results In wheat sites, richness and diversity declined abruptly from ecotones to fields and remained homogenously poor throughout the fields, regardless of climate. In olive sites, despite the sharp structural boundary, richness and diversity remained high from the semi‐natural habitat to the grove margins and then declined gradually into groves. Species of conservation concern occurred across all habitats at olive sites, but none were found inside wheat fields or at their ecotones. The contrast in community structure between semi‐natural habitats and fields was affected by both climate and field type. Irrigation in arid regions did not augment species diversity. Main conclusions Our results indicate that consideration of crop type, within a climatic context, should receive high priority in biodiversity conservation in agricultural areas. In ‘hostile’ crops, such as wheat, we suggest favouring a combination of high‐intensity management and wide margins over less intensive management without margins, which may merely aid generalist butterfly species. The scarcity of butterflies in arid irrigated fields suggests a need to carefully assess the effects of irrigation and agrochemicals on species’ communities.     

Quantitatively defining the conservation status of Natura 2000 forest habitats and improving management options for enhancing biodiversity

The main goal of Natura 2000 network is to guarantee the favourable conservation status of habitats and species ensuring European biodiversity. As a result, certain forest areas have been included in this network listed as 9230-Quercus pyrenaica habitat and 9340-Quercus ilex subsp. rotundifolia forest habitat. These areas were previously used for firewood extraction or livestock grazing and browsing. Nowadays these habitats are coppice forests with asexual regeneration, which is far from the desired conservation status. Traditional timber harvesting plans do not take account of the new objectives required for these Natura sites, which attempt to ensure biodiversity and recreational uses instead of simply focusing on timber production. This paper proposes a flexible methodology (applied to the study area “Dehesa Boyal” in ávila, Spain) for managing Natura 2000 forest sites by stands for sustainable forest management and the new requirements. The methodology has two phases. The first, “Division of the forest area into stands”, defines homogeneous patches of vegetation distinct in species composition, physiognomic structure and future management. The second, “Conservation status assessment of stands”, quantifies the conservation status of each previously classified stand considering a series of factors such as: functional health, restoration, floral richness and structure. A total value integrating the conservation status of stands is then calculated for the habitat. Both phases use Geographic Information System tools for managing information and visualizing results. The proposed methodology provides forest managers with a good knowledge of the territory and subsequently enables them to take appropriate conservation measures to maintain biodiversity.     

Conserving freshwater biodiversity: The value,status and management of high quality ditch systems

Freshwater biodiversity is globally threatened and while most conservation efforts are focussed on natural and larger freshwater systems such as rivers and lakes, in many lowland agricultural landscapes artificial water bodies including ditches may be equally important as habitats for freshwater species. Ditches occur across the agricultural landscape but in particular, those associated with coastal and floodplain grazing marsh, have high conservation value. The importance of this habitat for rare and threatened species afforded priority status under the UK Biodiversity Action Plan is explored. The characteristics of ditches that have high conservation value are described and a set of targets against which such ditches can be assessed are presented. An analysis of the current condition of Sites of Special Scientific Interest (SSSI) for the wider coastal and floodplain grazing marsh habitat demonstrates the range of pressures affecting these sites and highlights that alongside generic freshwater issues such as eutrophication and non-native species, these sites have a unique set of pressures associated with their ongoing management and the vulnerable location of many sites at the coast. Wider conservation strategies for freshwater biodiversity in lowland landscapes across Europe need to factor in the different management requirements of artificial habitats such as ditches alongside more ambitious restoration projects for natural waterbodies. In low lying coastal areas the threat of coastal squeeze for many important grazing marshes will require a strategic approach to allow upstream migration of important biodiversity.     

Environmental diagnosis: Integrating biodiversity conservation in management of Natura 2000 forest spaces

The conservation of biodiversity in Europe is defined by Directive 92/43/EEC – commonly known as the Habitats Directive – relating to the conservation of natural habitats and of wild flora and fauna. This Directive established the creation of an ecological network of European protected areas – the Natura 2000 network – , and also recognised the need to manage these areas to maintain their “favourable conservation status”.This paper proposes a methodology which enables the conservation of biodiversity to be integrated into the management of Natura 2000 forest spaces. The methodology comprises an “environmental diagnosis” in three phases. The first phase evaluates the current conservation status of habitats using the following criteria: vital functions; floristic richness; forest structure; area occupied by the habitat; and recovery capacity. The second phase assesses the fragility of the space to determine the degree of vulnerability of habitats. This involves evaluating the fire hazard, erosion hazard, and the fragility of the vegetation. The last phase combines the two previous ones to generate management areas (optimum, intermediate or unfavourable) and to prioritise management actions.This methodology was applied in a protected forest area in the Natura 2000 network, located in Avila (Spain). Different management areas were generated for biodiversity conservation, and each habitat was associated to one of them. Finally, actions were prioritised and designed to raise the habitats to a “favourable conservation status”.     

Missing the target? A critical view on butterfly conservation efforts on calcareous grasslands in south-western Germany

Butterflies are strongly declining on grassland habitats of Central Europe. Therefore, the success of conservation measures on high quality grassland habitats is controversially discussed. We compared the changes in butterfly diversity and community structure on six managed calcareous grasslands with eight unmanaged vineyard fallows. We obtained strong losses of species diversity and remarkable shifts of community compositions on both habitat types. However, the changes on vineyard fallows were only slightly more severe but more stochastic than on the calcareous grasslands. The shifts in community composition with respect to functional species traits were rather similar between the two different grassland types so that complex butterfly communities evolved into generalist-dominated ones. Connectivity was higher among vineyard fallows than among calcareous grasslands. Consequently, conservation measures on calcareous grasslands only partly achieved their goal to maintain the high species diversity and functional complexity still observed in the 1970s. The negative impacts of eutrophication and monotonisation of the landscape as well as climate change are affecting all habitats, independently from management concepts. Therefore, management on conservation sites can buffer against these effects, but is not sufficient for a full compensation.     

Conservation priority of Italian Alpine habitats: a floristic approach based on potential distribution of vascular plant species

In the European Union, the Directive 92/43/EEC defines a number of species and habitats of community interest that are worthy to be preserved because in danger to disappear or because they are representative of the different European bio-geographical regions. In the light of the limited economic resources generally allocated to conservation efforts, there is the necessity to prioritise conservation actions in order to avoid deterioration of protected areas. To this aim, in the present study the most representative habitats of the Italian Alps are compared on the basis of vascular plant biodiversity and a conservation priority index is proposed for each habitat taking into account the potential distribution of 252 threatened vascular plant species. Rocky slopes, screes and alpine grasslands resulted to have the greatest percentage of endemic plant species so reflecting the general distributional pattern of endemic plant species at high altitudes in Eurasian mountains. The relationship between the conservation priority index and the corresponding habitat extent within the Natura 2000 network suggests that peatlands, arid grasslands, wet meadows and freshwater habitats deserve a higher priority in conservation actions. Although vascular plant biodiversity is not necessarily a surrogate of overall biodiversity of Alpine habitats, the results here reported can be used as an initial reference framework for prioritising conservation actions, so as to accomplish the provisions of Article 6 of Habitats Directive.     

Distribution of ground-dwelling beetles in fragmented tropical habitats

The conservation of biodiversity is increasingly dependent on human-altered habitats. In a fragmented forest landscape in northern Costa Rica within the Mesoamerican Biological Corridor, an area of great conservation importance, we compared the diversity and composition of ground-dwelling beetle communities in five habitat types along a gradient of increasing disturbance: primary forest, logged forest, secondary forest, plantation (Gmelina arborea) and pasture. Using pitfall trapping we captured a total of 1,877 beetles (Coleoptera), comprising 422 morphospecies in 26 families. The plantation sites had the lowest number of species followed by secondary forest and pasture. Multivariate analysis separated the beetle fauna according to land use, and suggested that only the logged forest maintains a similar species assemblage to primary forest. However, each habitat harboured a number of unique species indicating the conservation value of the mosaic of habitats found in fragmented landscapes. Our results suggest that to maintain forest beetle species diversity it is essential that areas of natural forest are conserved. However, other land uses also provide beetle habitats and in fragmented areas active management to maintain a mosaic of land uses will contribute to the conservation of beetle diversity.     

Bridging the gap between the Natura 2000 regional conservation status and local conservation objectives

Natura 2000 targets the sustainable conservation of Europe's biodiversity. An important cornerstone of Natura 2000 is the Habitats Directive, which is currently implemented across European member states. However, straightforward implementation is not obvious since the favourable conservation status of habitats and species needs to be achieved at the member state level, while conservation objectives need to be formulated at the protected site level. To bridge this gap, we propose to start from regional conservation objectives before site level objectives are formulated. These regional conservation objectives have the advantage of providing a framework according to which conservation objectives can be allocated both within and outside the protected sites of the Natura 2000 network. Especially since they all contribute to the national or regional conservation status. Recently, Flanders (northern Belgium) has adopted this approach and has quantified conservation objectives at the regional scale. As the current regional conservation status of habitats and species is mostly unfavourable, regional conservation objectives entail a drastic increase in area (42%) for habitats, and active conservation measures for 78% of the species. We are convinced that the method outlined here, may substantially contribute to a helpful discussion about implementing and streamlining Natura 2000 across European member states.     

钱江源国家公园体制试点区功能分区对黑麂保护的有效性评估

功能分区是统一协调国家公园不同保护管理目标的主要措施, 对于国家公园的有效管理有重要意义。钱江源国家公园是我国首批国家公园体制试点区之一, 被分为核心保护区、生态保育区、游憩展示区和传统利用区4个部分, 对应不同的保护管理措施。本研究分析了钱江源国家公园体制试点区现有功能分区与其首要保护对象黑麂(Muntiacus crinifrons)的适宜栖息地之间的空间关系。在红外相机调查获取的94个黑麂分布点的基础上, 结合海拔、地形、植被特征、人为活动干扰等15个环境特征变量, 采用MaxEnt模型预测国家公园内黑麂适宜栖息地的空间分布。结果表明, 黑麂倾向于出现在森林较为原始和道路密度较低的区域, 其适宜栖息地面积42.5 km ², 占国家公园总面积的16.9%。其中, 69.3%的黑麂适宜栖息地位于核心保护区, 30.4%位于生态保育区, 表明国家公园现有功能分区能很好地满足黑麂栖息地保护的需求。此结果也证明黑麂可以作为其分布范围内保存较好的亚热带森林生态系统的指示性物种。通过生境恢复、廊道建设和跨省共建促进黑麂栖息地的完整性保护, 是加强该区域黑麂栖息地保护的关键措施。     

Improving conservation and management of the imperiled Carson Valley silverpot butterfly <Emphasis Type="Italic">Speyeria nokomis carsonensis</Emphasis> (Lepidoptera: Nymphalidae) based on rapid assessments of distribution,habitat, and threats

The immediacy of conservation decisions for imperiled species often requires rapid assessments of fundamental species-specific data to appropriately guide conservation resources, regulatory actions, and mitigation activities. The goal of this paper was to inform future conservation and management decisions for the imperiled Carson Valley silverspot butterfly (Speyeria nokomis carsonensis), a species that has been subject to conservation attention, yet very little is known about its distribution and habitat. Surveys for S. nokomis carsonensis and its habitat were conducted during 2004–2005. An 88% decline in the number of historic subpopulations was observed, but 30 new subpopulations were discovered. Among the 62 sites that either had populations or were considered potentially suitable habitat, three distinct habitat types were delineated: agricultural seeps, isolated springs, and riparian meadows. Riparian sites comprised 58% of all sites and little variation was observed in the proportion of occupancy across habitat types. Disturbance was substantially greater in unoccupied sites than in sites occupied by S. nokomis carsonensis. Livestock grazing was the primary threat (50% of sites) to habitat, while water diversions were a secondary threat (7 sites). The variability in habitat and threats identifies that even for highly specialized subspecies, all habitats simply cannot be considered equal candidates for inclusion in a conservation strategy for the butterfly. Adopting proactive management policies that conserve suites of sites within each habitat type and reduce grazing intensity within S. nokomis carsonensis habitats should be a priority to reduce threats and circumvent a prospective listing pursuant to the US Endangered Species Act.     

Effects of habitat transitions on rainforest bird communities across an anthropogenic landscape mosaic

We compared bird community responses to the habitat transitions of rainforest‐to‐pasture conversion, consequent habitat fragmentation, and post‐agricultural regeneration, across a landscape mosaic of about 600 km² in the eastern Australian subtropics. Birds were surveyed in seven habitats: continuous mature rainforest; two size classes of mature rainforest fragment (4–21 ha and 1–3 ha); regrowth forest patches dominated by a non‐native tree (2–20 ha, 30–50 years old); two types of isolated mature trees in pasture; and treeless pasture, with six sites per habitat. We compared the avifauna among habitats and among sites, at the levels of species, functional guilds, and community‐wide. Community‐wide species richness and abundance of birds in pasture sites were about one‐fifth and one‐third, respectively, of their values in mature rainforest (irrespective of patch size). Many measured attributes changed progressively across a gradient of increased habitat simplification. Rainforest specialists became less common and less diverse with decreased habitat patch size and vegetation maturity. However, even rainforest fragments of 1–3 ha supported about half of these species. Forest generalist species were largely insensitive to patch size and successional stage. Few species reached their greatest abundance in either small rainforest fragments or regrowth. All pastures were dominated by bird species whose typical native habitats were grassland, wetland, and open eucalypt forest, while pasture trees modestly enhanced local bird communities. Overall, even small scattered patches of mature and regrowth forest contributed substantial bird diversity to local landscapes. Therefore, maximizing the aggregate rainforest area is a useful regional conservation strategy.     

Local species immigration,extinction, and turnover of butterflies in relation to habitat area and habitat isolation

Temporal dynamics of insect communities in terrestrial habitat fragments have been rarely studied. Here it was tested whether immigration, extinction, and turnover of butterfly species change with area and isolation of 31 calcareous grasslands. The area ranged from 0.03 to 5.14 ha, the isolation index from 2,100 to 86,000 (edge-to-edge distance 55–1,894 m). In both study years (1996, 2000), the total number of individuals (16,466, 15,101) and species (60, 54) sampled across all sites were similar and number of species increased with area in both years indicating an equilibrium. Rates of extinction (38% for habitat specialists vs. 20% for generalists) and turnover (51% vs. 35%) were higher, and rates of immigration (11% vs. 30%) were lower for habitat specialists than for generalists. Extinction and turnover rates decreased with increasing fragment size for both specialist (n =25 species) and generalist (n =36) butterflies, but specialists showed a significantly steeper decrease with increasing fragment size than generalists. Immigration rates increased with area. As a result, species number of habitat specialists declined in small habitats but not in large habitats between 1996 and 2000. No significant impact of habitat isolation on the butterfly community was found. The data suggest that large habitat fragments are of special importance for the conservation of the specialized, most endangered butterfly species. Habitat isolation appears to be less important, as butterflies can cope with the habitat mosaic in our study region.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.