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Nesting in dense aggregations is common in central place foragers, such as group-living birds and insects. Both environmental heterogeneity and behavioral interactions are known to induce clustering of nests, but their relative importance remains unclear. We developed an individual-based model that simulated the spatial organization of nest building in a gregarious digger wasp, Bembix rostrata. This process-based model integrates environmental suitability, as derived from a microhabitat model, and relevant behavioral mechanisms related to local site fidelity and conspecific attraction. The drivers behind the nesting were determined by means of inverse modeling in which the emerging spatial and network patterns from simulations were compared with those observed in the field. Models with individual differences in behavior that include the simultaneous effect of a weak environmental cue and strong behavioral mechanisms yielded the best fit to the field data. The nest pattern formation of a central place foraging insect cannot be considered as the sum of environmental and behavioral mechanisms. We demonstrate the use of inverse modeling to understand complex processes that underlie nest aggregation in nature.

Tracking devices have become small enough to be widely applied to arthropods to study their movement. However, possible side effects of these devices on arthropod performance and behaviour are rarely considered. We performed a systematic review of 173 papers about research in which tracking devices-radio frequency identification (RFID), harmonic radar and radio telemetry tags-were attached to terrestrial arthropods. The impact of such tags was quantified in only 12% of the papers, while in 40% the potential impact was completely disregarded. Often-cited rules of thumb for determining appropriate tag weight had either no empirical basis or were misconstrued. Several properties of a tracking device (e.g. weight, balance, size, drag) can affect different aspects of an arthropod's life history (e.g. energy, movement, foraging, mating). The impact can differ among species and environments. Taken together, these tag effects can influence the reliability of obtained movement data and conclusions drawn from them. We argue that the impact of tracking devices on arthropods should be quantified for each (a) study species, (b) tag type, and (c) environmental context. As an example, we include a low-effort impact study of the effect of an RFID tag on a digger wasp. Technological advancements enable studying the movement of arthropods in unprecedented detail. However, we should adopt a more critical attitude towards the use of tracking devices on terrestrial arthropods. The benefits of tracking devices should be balanced against their potential side effects on arthropods and on the reliability of the resulting data.

Understanding connectivity among populations in fragmented landscapes is of paramount importance in species conservation because it determines their long-term viability and helps to identify and prioritize populations to conserve. Rare and sedentary species are particularly vulnerable to habitat fragmentation as they occupy narrow niches or restricted habitat ranges. Here, we assess contemporary interpopulation connectedness of the threatened, myrmecophilous butterfly, Maculinea alcon, in a highly fragmented landscape. We inferred dispersal, effective population sizes, genetic diversity and structure based on 14 locations of M. alcon in Belgium and the Netherlands using data from 12 microsatellite loci. Despite the reported sedentary behaviour of M. alcon, we observed moderate levels of contemporary dispersal between patches, but only in landscapes where populations were located within a distance of 3 km from neighbouring populations. Estimates of effective population sizes (Ne) were very low(ranging from1.6 to 17.6) and bottleneck events occurred in most of the studied populations. We discuss the functional conservation units delineated based on a former mark-release-recapture study, and formulate appropriate conservation strategies to maintain viable (meta)populations in highly fragmented, anthropogenic landscapes.

Ecotones are often species-rich and harbour specific resources and environmental conditions for invertebrates. Despite their functional significance for conservation, they are often not explicitly included in biotope typologies relevant to conservation policy and management (e.g., the European Habitats Directive). The Ilex Hairstreak (Satyrium ilicis) is a species of European conservation concern and a typical ecotone species inhabiting gradients from open (e.g., heathland, grassland) to closed vegetation (e.g., woodland). Here, we investigated its occurrence and habitat use at different spatial scales in Flanders (north Belgium). At a regional scale, species distribution modelling predicted 1,152 grid cells of 1 × 1 km2 to be suitable of which 190 were presumed to occur within colonization capacity (±2.5 km). At a local scale, adult butterflies were more abundant on sites sheltered by bushes and small trees and with nectar sources in the vicinity of tall oak trees (mate locating sites). For egg-laying, females preferred oaks of intermediate height (50–150 cm) with many low branches at some distance from the nearest woodland edge (12 m). Additionally, Alder Buckthorn (Frangula alnus) was abundant as well as a herb layer of 10–15 cm. 73 % of the eggs were parasitized and parasitism occurred more often within sites where small oaks were very abundant. Making use of our results, we suggest conservation measures at different scales for this endangered ecotone species: policy measures at a regional level to delineate functional conservation units and local management measures using a resource-based approach.

The Macedonian Grayling is listed as critically endangered in the recent IUCN Red List of European butterflies because of its extreme rarity and habitat loss due to quarrying. This categorisation was, however, based on rather limited knowledge on its actual distribution, popu- lation size and habitat requirements. In 2012, we conducted field surveys to acquire more information. We found the species at six new sites extending its known range of suitable habitat to just under 10 km 2 . The daily population size was estimated using capture-mark-recapture method in the most densely populated part of the Pletvar pass site at more than 650 individuals. Adults proved to be extremely sedentary, not moving far even within the continuous habitat on the same slope. Oviposition was observed on dry plant material and in a rock crevice close to the potential larval host plant Festuca sp. Quarrying is confirmed to be the main threat to the habitat of the Macedonian Grayling with five out of seven populated sites containing active marble quarries. Due to the enlargement of the known area of occupancy, its threat status would now be estimated at endangered. Despite the restricted knowledge about its distribution and trends in the population size, the IUCN criteria proved to be applicable to determine the threat status of a rare and localized butterfly such as Pseudoc- hazara cingovskii. Its original assessment of being called the most threatened butterfly in Europe resulted in imme- diate research project and subsequent actions that will undoubtedly help to conserve it in the future.

Aim Invasive alien species (IAS) are recognized as major drivers of biodiversity loss, but few causal relationships between IAS and species declines have been documented. In this study, we compare the distribution (Belgium and Britain) and abundance (Belgium, Britain and Switzerland) of formerly common and widespread native ladybirds before and after the arrival of Harmonia axyridis, a globally rapidly expanding IAS

Red Lists are used to assess the extinction risk of species based on quantitative IUCN criteria. For the compilation of a new Red List of butterflies in Flanders (north Belgium), we collated ca 800,000 distribution records and applied the IUCN Red List criteria to this small region (ca 135,00 km2). We also explored the effect of spatial resolution on the outcome of the Red List assessment by alternatively using 1 × 1 km2 and 5 × 5 km2 grid cells for geographic range size and trend calculations. We determined conservation hot spots in Flanders based on the Red List status of the species composition in each grid cell. The new Red List classified 20 butterflies (out of 68 resident species) as Regionally Extinct, six as Critically Endangered, five as Endangered, seven as Vulnerable and seven as Near Threatened. The remaining 23 species were classified as Least Concern. Using coarse instead of fine grain grid cells would have classified ten species in a lower Red List category. Compared with the previous Red List, nine species were classified in a lower and 12 in a higher threat category. In total, 218 1 × 1 km2 grid cells were considered as (very) high butterfly conservation priority sites. The application of the new IUCN criteria in a small region such as Flanders resulted in a Red List that offered challenging opportunities for the conservation of butterflies in particular and biodiversity in general.

Twenty years ago, Desender and Turin (1989) analysed the changes in the composition of carabid beetles in four NW European countries between the periods

Being ectotherms, insects are predicted to suffer more severely from climate change than warm-blooded animals. We forecast possible changes in diversity and composition of butterflies, grasshoppers and dragonflies in Belgium under increasingly severe climate change scenarios for the year 2100. Two species distribution modelling techniques (Generalised Linear Models and Generalised Additive Models), were combined via a conservative version of the ensemble forecasting strategy to predict present-day and future species distributions, considering the species as potentially present only if both modelling techniques made such a prediction. All models applied were fair to good, according to the AUC (area under the curve of the receiver operating characteristic plot), sensitivity and specificity model performance measures based on model evaluation data. Butterfly and grasshopper diversity were predicted to decrease significantly in all scenarios and species-rich locations were predicted to move towards higher altitudes. Dragonfly diversity was predicted to decrease significantly in all scenarios, but dragonfly-rich locations were predicted to move upwards only in the less severe scenarios. The largest turnover rates were predicted to occur at higher altitudes for butterflies and grasshoppers, but at intermediate altitudes for dragonflies. Our results highlight the challenge of building conservation strategies under climate change, because the changes in the sites important for different groups will not overlap, increasing the area needed for protection. We advocate that possible conservation and policy measures to mitigate the potentially strong impacts of climate change on insect diversity in Belgium should be much more pro-active and flexible than is the case presently.

Aim To analyse the effect of the inclusion of soil and land-cover data on the performance of bioclimatic envelope models for the regional-scale prediction of butterfly (Rhopalocera) and grasshopper (Orthoptera) distributions. Location Temperate Europe (Belgium). Methods Distributional data were extracted from butterfly and grasshopper atlases at a resolution of 5 km for the period 1991-2006 in Belgium. For each group separately, the well-surveyed squares (n = 366 for butterflies and n = 322 for grasshoppers) were identified using an environmental stratification design and were randomly divided into calibration (70%) and evaluation (30%) datasets. Generalized additive models were applied to the calibration dataset to estimate occurrence probabilities for 63 butterfly and 33 grasshopper species, as a function of: (1) climate, (2) climate and land-cover, (3) climate and soil, and (4) climate, land-cover and soil variables. Models were evaluated as: (1) the amount of explained deviance in the calibration dataset, (2) Akaike's information criterion, and (3) the number of omission and commission errors in the evaluation dataset. Results Information on broad land-cover classes or predominant soil types led to similar improvements in the performance relative to the climate-only models for both taxonomic groups. In addition, the joint inclusion of land-cover and soil variables in the models provided predictions that fitted more closely to the species distributions than the predictions obtained from bioclimatic models incorporating only land-cover or only soil variables. The combined models exhibited higher discrimination ability between the presence and absence of species in the evaluation dataset. Main conclusions These results draw attention to the importance of soil data for species distribution models at regional scales of analysis. The combined inclusion of land-cover and soil data in the models makes it possible to identify areas with suitable climatic conditions but unsuitable combinations of vegetation and soil types. While contingent on the species, the results indicate the need to consider soil information in regional-scale species-climate impact models, particularly when predicting future range shifts of species under climate change.

Abstract National or regional conservation strategies are usually based on available species distribution maps. However, very few taxonomic groups achieve a full coverage of the focal region. Distribution data of well-mapped taxonomic groups could help predict the distribution of less well-mapped groups and thus fill gaps in distribution maps. Here, we predict the distribution of five heathland butterflies in Flanders (north Belgium) using typical heathland bird distribution data as predictor variables. We compare predictions with those using only biotope or a combination of both biotope and bird data as variables. In addition, we test the transferability of 'bird', biotope and combined models to the Netherlands, an ecologically similar region. Transferability was tested in three separate sandy regions in the Netherlands at different distances from the region in which the models were built. For each of the five heathland butterflies, we applied logistic regressions on ten random model sets and tested the models on ten random evaluation sets within Flanders. We used the area under the curve (AUC) of the receiver-operating characteristics (ROC) plots to estimate model accuracy. Overall, bird models performed significantly better than biotope models but were not significantly different from the combined models in Flanders. In the Netherlands, the transferred biotope and the combined models performed better than the transferred 'bird models'. We conclude that on a local scale, birds can, to some extent, serve as proxies for biotope quality, but that biotope models are more robust when transferred to another region.

From a conservation point of view, species’ tolerances towards disturbance are often generalised and lack reference to spatial scales and underlying processes. In order to investigate how average typical species react to habitat fragmentation and disturbance, we adopted a multi-species approach to address occupancy patterns of five specialised dune arthropods (butterflies Hipparchia semele, Issoria lathonia; grasshopper Oedipoda caerulescens; spiders Alopecosa fabrilis, Xysticus sabulosus) in recently fragmented coastal dune habitats which are subjected to varying levels and modes of local disturbance, i.e. trampling by cattle or people. Occupancy patterns were assessed during two successive years in 133 grey dune fragments of the Flemish coastal dunes (Belgium, France). By treating species as a random factor in our models, emphasis was placed on generalisations rather than documenting species-specific patterns. Our study demonstrates that deteriorating effects of local disturbance on arthropod incidence cannot be interpreted independent of its landscape context, and appear to be more severe when patch area and connectivity decrease. When controlled for patch area and trampling intensity, the probability of species occupancy in poorly connected patches is higher under cattle trampling than under recreation. Incidences additionally decrease with increasing intensity of cattle trampling, but increases with trampling by tourists. This study provides evidence of mode- and landscape-dependent effects of local disturbance on species occupancy patterns. Most importantly, it demonstrates that trampling of sensitive dune fragments will lead to local and metapopulation extinction in landscapes where trampling occurs in a spatially autocorrelated way, but that the outcome (spatial patterns) varies in relation to disturbance mode, indicating that effects of disturbance cannot be generalised.

ABSTR AC T Woodpastures (open, grazed woodlands with a mosaic of grassland, shrub and tree patches) are of high biological and cultural value and have become a threatened ecosystem in Europe. Spontaneous tree regeneration in the presence of large herbivores, is an essential process for management and restoration of this structurally diverse habitat. We examined the suitability of five vegetation types (grasslands, ruderal vegetations, tall sedges, rush tussocks and bramble thickets), grazed by large herbivores, for tree regeneration. We hypothesized that bramble thickets and tall herb communities operate as safe sites for palatable tree species through the mechanism of associational resistance. We set up a field experiment with tree seedlings in grazed and ungrazed conditions and recorded mortality and growth of seedlings of two palatable tree species (Quercus robur and Fraxinus excelsior) during three growing seasons. In the same experiment, we studied the effect of a two year’s initial time gap before grazing. Bramble thickets were suitable safe sites for survival and growth of seedlings of both species. Tall sedges, soft rush tussocks and ruderal vegetations with unpalatable or spiny species provided temporal protection, allowing seedlings to survive. Tree regeneration in livestock grazed grassland was highly constrained. Rabbits may undo the nursing effects of bramble thickets. The first year’s survival is of major importance for the establishment of trees. Subsequent grazing affects growth rather than survival. A two year’s initial time gap before grazing, had positive effects on survival, but did not enhance outgrowth of unprotected trees.

Since 1998, SCK•CEN, in partnership with IBA s.a. and many European research laboratories, is designing a multipurpose Accelerator Driven System (ADS) – MYRRHA – and is conducting an associated R&D support programme. MYRRHA aims to serve as a basis for the European experimental ADS to provide protons and neutrons for various R&D applications. Besides an overall configuration of the MYRRHA reactor internals, the description in this paper is limited to the mechanical design of the main components of the Primary System and Associated Equipment (vessel and cover, diaphragm, spallation loop, sub-critical core, primary cooling system, emergency cooling system, in-vessel fuel storage and fuel transfer machine), the conceptual design of the robotics for In-Service Inspection and Repair (ISIR), together with the remote handling for operation and maintenance (O&M).

We studied the habitat use and mobility of the Grayling butterfly (Hipparchia semele) and the Blue-Winged Grasshopper (Oedipoda caerulescens), two threatened insects within spatially structured blond and grey dunes in a nature reserve along the Belgian coast. Although both species occur in the same biotope, H. semele were more abundant in open, dynamic sites with a relatively high amount of bare sand, while O. caerulescens preferred sheltered, more stable environments with a lower amount of bare sand. Unlike H. semele, substrate use varied in accordance to body colouration in O. caerulescens, especially on cold days, with light-coloured animals being more abundant on sand and dark-coloured animals more abundant on moss. During a mark-recapture-study, we marked 493 Grayling butterflies and 1289 Blue-Winged Grasshoppers. On average, both sexes of H. semele were equally mobile (about 150 m/day; maximum recorded distance of about 1700 m) while male O. caerulescens were significantly more mobile than females (daily average 47 vs. 5 m; maximum distances observed for O. caerulescens were about 800 m). The importance of habitat heterogeneity (within and among patches) and the consequences of habitat use and mobility of both species for the conservation of typical coastal dune habitats are discussed. The complementary use of species-specific information to site-based management measures is advocated.

Aim We evaluate differences between and the applicability of three linear predictive models to determine butterfly hotspots in Belgium for nature conservation purposes. Location The study is carried out in Belgium for records located to Universal Transverse Mercator (UTM) grid cells of 5 x 5 km. Methods We first determine the relationship between factors correlated to butterfly diversity by means of modified t-tests and principal components analysis; subsequently, we predict hotspots using linear models based on land use, climate and topographical variables of well-surveyed UTM grid cells (n = 197). The well-surveyed squares are divided into a training set and an evaluation set to test the model predictions. We apply three different models: (1) a 'statistically focused' model where variables are entered in descending order of statistical significance, (2) a 'land use-focused' model where land use variables known to be related to butterfly diversity are forced into the model and (3) a 'hybrid' model where the variables of the 'land use-focused model' are entered first and subsequently complemented by the remaining variables entered in descending order of statistical significance. Results A principal components analyses reveals that climate, and to a large extent, land use are locked into topography, and that topography and climate are the variables most strongly correlated with butterfly diversity in Belgium. In the statistically focused model, biogeographical region alone explains 65% of the variability; other variables entering the statistically focused model are the area of coniferous and deciduous woodland, elevation and the number of frost days; the statistically focused model explains 77% of the variability in the training set and 66% in the evaluation set. In the land use-focused model, biogeographical region, deciduous and mixed woodland, natural grassland, heathland and bog, woodland edge, urban and agricultural area and biotope diversity are forced into the model; the land use-focused model explains 68% of the variability in the training set and 57% in the evaluation set. In the hybrid model, all variables from the land use-focused model are entered first and the covariates elevation, number of frost days and natural grassland area are added on statistical grounds; the hybrid model explains 78% of the variability in the training set and 67% in the evaluation set. Applying the different models to determine butterfly diversity hotspots resulted in the delimitation of spatially different areas. Main conclusions The best predictions of butterfly diversity in Belgium are obtained by the hybrid model in which land use variables relevant to butterfly richness are entered first after which climatic and topographic variables were added on strictly statistical grounds. The land use-focused model does not predict butterfly diversity in a satisfactory manner. When using predictive models to determine butterfly diversity, conservation biologists need to be aware of the consequences of applying such models. Although, in conservation biology, land use-focused models are preferable to statistically focused models, one should always check whether the applied model makes sense on the ground. Predictive models can target mapping efforts towards potentially species-rich sites and permits the incorporation of un-surveyed sites into nature conservancy policies. Species richness distribution maps produced by predictive modelling should therefore be used as pro-active conservation tools.

We illustrate the strong decrease in the number of butterfly species in Flanders (north Belgium) in the 20th century using data from a national butterfly mapping scheme. Nineteen of the 64 indigenous species went extinct and half of the remaining species are threatened at present. Flanders is shown to be the region with the highest number of extinct butterflies in Europe. More intensive agriculture practices and expansion of house and road building increased the extinction rate more than eightfold in the second half of the 20th century. The number of hot spots decreased considerably and the present-day hot spots are almost exclusively in the northeast of Flanders. Species with low dispersal capacities and species from oligotrophic habitats decreased significantly more than mobile species or species from eutrophic habitats. We discuss these results in a northwest European context and focus on concrete measures to preserve threatened butterfly populations in Flanders.

To enable use of Empididae s.l. (Diptera) as a tool in nature conservation, a Red Data Book of this taxonomical group was generated for Flanders, Belgium. All distribution data on species in Belgium between 1887 and 1999 were gathered from collections as well as personal sampling efforts by the first two authors. This resulted in about 21,000 records of Empididae, Hybotidae, Microphoridae and Atelestidae with 16,119 records for Flanders (northern Belgium) and 4776 for Wallony (southern Belgium). All species were assigned to Red Data Book categories which are based on a combination of a rarity and a trend criterion. Rarity is expressed as the proportion of the total number of UTM 5 km squares sampled in which the species have been found since 1981. The trend criterion is interpreted as the change of the species rarity between 1887–1980 and 1981–1999. A comparable number of UTM 5 km squares was investigated during the two time periods. A total of 259 species were recorded in Flanders. Twenty-seven or 10% of them are considered as 'extinct in Flanders', 10 (4%) as 'critically endangered', 12 (5%) as 'endangered', 11 (4%) as 'vulnerable', 99 species (38%) as 'susceptible' or 'rare', 65 species (25%) as 'safe' or 'at low risk' and 34 species (13%) are assigned to the category 'data deficient' due to taxonomic problems or a lack of ecological data. Only one of the common species shows a recent decrease of more than 50% and is classified as 'nearly threatened'. Current threats in most species are related to the alteration or destruction of their favoured habitats. The results are discussed in the light of recent criticism of the use of Red Data Books in nature conservation.

The compilation of the Red Lists of butterflies in Flanders and the Netherlands was based on two criteria: a trend criterion (degree of decline) and a rarity criterion (actual distribution area). However, due to the large difference in mapping intensity in the two compared periods, a straightforward comparison of the number of grid cells in which each species was recorded, appeared inappropriate. To correct for mapping intensity we used reference species that are homogeneously distributed over the country, that have always been fairly common and that did not fluctuate in abundance too much during this century. For all resident species a relative presence in two compared periods was calculated, using the average number of grid cells in which these reference species were recorded as a correction factor. The use of a standardized method and well-defined quantitative criteria makes national Red Lists more objective and easier to re-evaluate in the future and facilitates the comparison of Red Lists among countries and among different organisms. The technique applied to correct for mapping intensity could be useful to other organisms when there is a large difference in mapping intensity between two periods.

Butterfly monitoring started in the Netherlands in 1990 and in Flanders in 1991. During the last few years butterflies have been counted at nearly 300 sites. This high number of transects makes it possible to calculate not only national, but also regional and habitat-indices for many species. Special attention is paid to Red list species in the Netherlands. In the near future the number of sites per species on this list is to be increased to at least 20. This can be achieved by a ‘Red list monitoring scheme‘, in which monitoring is restricted to the flight period of the species.

1 Research Institute for Nature and Forest (INBO), Kliniekstraat 25, B-1070 Brussels, Belgium 2 Butterfly Conservation Europe, P.O. Box 506, NL-6700 AMWageningen, The Netherlands 3 Departamento de Biologia, Universidad Autonoma de Madrid, Cantoblanco, ES-28049-Madrid, Spain 4 Department of Zoology, Croatian Natural History Museum, 10000 Zagreb, Croatia 5 UFZ–Helmholtz-Centre for Environmental Research, Department of Community Ecology, Theodor-Lieser-Str. 4, D-06120, Halle, Germany 6 Dutch Butterfly Conservation, P.O. Box 506, NL-6700 AMWageningen, The Netherlands 7 University of Ljubljana, Biotechnical Faculty, Department of Biology, Vecna pot 111, 1000 Ljubljana, Slovenia 8 Butterfly Conservation (UK), Manor Yard, East Lulworth, Dorset BH20 5QP, UK