Environmental gradients and succession patterns of carabid beetles (Coleoptera: Carabidae) in an Alpine glacier retreat zone

Accelerated by global warming, retreating glaciers leave behind spatially ordered moraines with underlying primary succession and disturbance. Current knowledge of primary succession comes mainly from studies of vegetation dynamics. Information about above-ground macroinvertebrates is still scarce. We used carabid beetles (Coleoptera; Carabidae) as indicator taxon to assess the effects of (1) terrain age (species turnover along the proglacial chronosequence) and (2) small-scale habitat architecture (vegetation cover, surface texture) on the carabid assembly. For this purpose, 33 sampling sites with pitfall traps were installed throughout the glacier foreland Morteratsch (Engadine, Switzerland), adjacent sparse forests serving as reference sites. With a total of 33 carabid species on the foreland and another 2 on the reference sites, the study area yielded a very high carabid species diversity compared to other glacier forelands. In general, the age of deglaciation proved to be a highly significant predictor for the carabid distribution, especially for particularly discriminant species. Observed species richness and activity densities showed bimodal patterns with a steep increase within the first ca. 40?years, a decline between around 40–90?years, and a further increase towards the terminal moraine. There was no evidence of dispersal-stochasticity: distinct clusters of sites with similar species composition were found. Microhabitat suitability proved to be a secondary effect, embedded in a temporal framework of primary succession. Surface cover with litter, herbs and dwarf-shrubs turned out to be the crucial habitat factors. Habitat loss as a result of climate warming will primarily affect cold-stenotopic carabids, but may potentially be absorbed by active selection for cooler microhabitats.     

Influence of climate changes on animal communities in space and time: the case of spider assemblages along an alpine glacier foreland

The impact of global warming in space and time is described for species assemblages of wandering spiders along the alpine glacier foreland of the Forni Valley (Northern Italy). We tested the effect of environmental variables (e.g. elevation, age of glacier retreat, vegetation cover, debris cover) on species richness and on species composition of spiders. Age of glacier retreat was the only significant variable influencing spider species assemblages in the valley. A spatially structured distribution of species and species assemblages along the chronosequence of glacier retreat was evidenced. The threshold abruptly differentiating two groups of species richness and species composition fell between sites deglaciated 100 and 155 years before the analysis. Latitudinal shifts towards the poles in species ranges at the global scale in response to climatic changes are known, and an altitudinal shift in species range should be expected for spiders at the local scale of the Forni Valley. Such a shift is present in spider species assemblages, although not as an expected gradual change in species richness and composition, but with a threshold effect after one century of glacier retreat. We discuss our results in the light of plausible future scenarios due to global warming, the consequence of further glacier retreats onto spiders, and caveats for monitoring studies.     

Life in harsh environments: carabid and spider trait types and functional diversity on a debris‐covered glacier and along its foreland

1. Patterns of species richness and species assemblage composition of ground‐dwelling arthropods in primary successions along glacier forelands are traditionally described using a taxonomic approach. On the other hand, the functional trait approach could ensure a better characterisation of their colonisation strategies in these types of habitat. 2. The functional trait approach was applied to investigate patterns of functional diversity and life‐history traits of ground beetles and spiders on an alpine debris‐covered glacier and along its forefield in order to describe their colonisation strategies. 3. Ground beetles and spiders were sampled at different successional stages, representing five stages of deglaciation. 4. The results show that the studied glacier hosts ground beetle and spider assemblages that are mainly characterised by the following traits: walking colonisers, ground hunters and small‐sized species. These traits are typical of species living in cold, wet, and gravelly habitats. The diversity of functional traits in spiders increased along the succession, and in both carabids and spiders, life‐history traits follow the ‘addition and persistence model’. Accordingly, there is no turnover but there is an addition of new traits and a variation in their proportion within each species assemblage along the succession. The distribution of ground beetles and spiders along the glacier foreland and on the glacier seems to be driven by dispersal ability and foraging strategy. 5. The proposed functional approach improves knowledge of the adaptive strategies of ground‐dwelling arthropods colonising glacier surfaces and recently deglaciated terrains, which represent landforms quickly changing due to global warming.     

Diversity, distribution and larval habitats of North Swedish blackflies (Diptera: Simuliidae)

1. Blackfly species richness and community structure were analysed at fifty-six sites in northern Sweden in two seasons. The sites were situated in a wide range of streams and rivers from small springbrooks, bog streams and lake-outlet streams to medium-sized forest rivers and large rivers draining montane regions.
2. Thirty-nine blackfly species were found, with between two and thirteen species per site. Neither species richness nor abundance could be related to the environmental variables measured.
3. An analysis of labral fan size of blackflies indicated a clear trend for the prevalence of larvae with small fans in large rivers and larvae with larger fan size in small streams. Similarly, fan size related to current velocities so that large fans were associated with slow current velocities and small fans with high velocities.
4. A strong relationship existed between species composition and habitat, as seen in ordination by non-metric multidimensional scaling. The relationship found between fan size and habitat size-related variables, such as channel width, depth, velocity and substratum particle size, along with longitude and altitude, in partial least squares regression analysis offered an explanation of the species composition–habitat relationship.
5. In addition to testing that distributions of blackfly larvae reflect morphological traits, we tested two general hypotheses pertaining to distribution patterns: (a) that blackfly communities show bimodal distributions; and (b) that their distributions are nested. Neither of these two hypotheses was supported by our observations. However, widespread blackfly species were locally more abundant than those found at relatively few sites, thus showing a positive abundance–occupancy relationship.     

Longitudinal changes in macroinvertebrate assemblages along a glacial river system in central Iceland

1. Macroinvertebrate assemblages were studied in the glacial river West-Jökulsá, originating from the Hofsjökull Ice Cap in central Iceland at an altitude of 860 m. Sampling sites were distributed from the source to 45 km downstream at 160 m a.s.l. Comparative studies were carried out on non-glacial rivers and tributaries in the area, at similar altitudes and distances from the glacial source.
2. Detrended correspondence analysis (DCA) demonstrated that species composition of benthic macroinvertebrates was related to the distance from the glacier. Assemblages at sampling sites furthest from the glacier were similar in species composition to sites in non-glacial rivers. Temporal variation was small compared with longitudinal zonation.
3. Based on canonical correspondence analysis (CCA) of data from the main glacial river, distance from the glacier, altitude, bryophyte biomass and the Pfankuch Index of channel stability were the measured explanatory variables having a significant effect on the structure of macroinvertebrate assemblages, accounting for 31% of the total variation in the data set. When data from all the rivers were analysed, altitude, bryophyte biomass, channel slope, suspended sediment concentration and maximum water temperature explained 21% of the variance.
4. Macroinvertebrate communities were in general agreement with the predictions of the conceptual model of Milner & Petts (1994) for the upstream reaches. The assemblages consisted mainly of Orthocladiinae and Diamesinae (Chironomidae), although other taxa such as Simuliidae, Plecoptera and Trichoptera were also found in low numbers. Shredders were lacking from the benthic communities, apparently because of continued glacial influence in the river even 45 km downstream from the glacier and lack of allochthononus inputs from riparian vegetation.     

Effects of catchment perturbations by logging and wildfires on zooplankton species richness and composition in Boreal Shield lakes

SUMMARY 1. Forest logging and wildfires are important perturbations of the boreal forest, but their effects on lake biota remain largely unknown. Here, we test whether zooplankton species richness and species assemblages differed among three groups of lakes in Eastern Canada characterised by different catchment conditions: logged in 1995 ( n =9); burnt in 1995 ( n =9); unperturbed ( n =20). Lakes were sampled in June, July and September 1 year after catchment perturbations.
2. Cumulative species richness in reference lakes averaged 46 (33–60) of which 63% were rotifers. Mean cumulative species richness and mean diversity in logged and burnt lakes did not differ from those in reference lakes.
3. Lake species assemblages were described by the density of 62 species (41 rotifers and 21 crustaceans). Among-group differences in species assemblages were not significant. Eighteen per cent of the total variability in species assemblages could be explained by 13 environmental factors, among which dissolved oxygen concentration and cyanobacteria biovolume were the most important. About 5% of species assemblage variability was attributed to covariation between environmental factors and time of sampling, while 4.1% was attributed to temporal variation.
4. Variations in zooplankton species richness and assemblages in Boreal Shield lakes are important, both among lakes and among sampling dates. They seem to depend on environmental factors unrelated to catchment-based perturbations, at least on the short-term of 1 year.     

The role of various meadow margin types in shaping carabid and staphylinid beetle assemblages (Coleoptera: Carabidae,Staphylinidae) in meadow dominated landscapes

Less intensively managed semi-natural habitats, e.g., field and meadow margins like hedgerows, are thought to be crucial landscape components for maintaining biodiversity in highly disturbed and intensively managed agricultural landscapes. In this study, we focused on the effects of three meadow margin types on activity-density, species richness and species composition of carabid and staphylinid beetles recorded by pitfall traps in Central European landscapes dominated by intensively managed meadows. Carabid activity-density was significantly higher in meadows than in meadow margins and within meadow margins their activity-density increased from grassy meadow margins via shrubby ones to woody meadow margins. We found that recorded species richness of both carabid and staphylinid beetles was not significantly affected by habitat identity (meadow margin or neighbouring meadow) and meadow margin type. Recorded species composition of both investigated taxa was significantly affected by habitat identity and interaction between habitat identity and meadow margin type (i.e. it differed between particular meadow margin types). Assemblages inhabiting various meadow margin types were more dissimilar between each other than assemblages from neighbouring meadows. Meadow margins within grassland dominated landscapes maintain local species richness by hosting different species from those living in surrounding meadows. Dissimilarity of carabid and staphylinid assemblages from meadows neighbouring both sides of particular meadow margin did not differ between meadow margin types. Our results indicate that semi-natural habitats play an important role in maintaining biodiversity not only in agricultural landscapes dominated by arable fields, but also in those dominated by meadows.     

Carabids of differently aged reforested pinewoods and a natural pine forest in a historically modified landscape

We studied the responses of carabid beetles to Pinus sylvestris ageing in four plantations aged 2–3, 10–12, 40–45 and 80 years, and one mature natural pine forest – in NW Spain. In 2003, a total of 16,866 carabid individuals (50 species) were collected using pitfall traps. Overall carabid abundance and species richness differed significantly among the five age classes with the highest values occurring in the youngest age class. The carabid assemblages of the initial stages of the ageing sequence differed considerably from the later stages that were relatively similar to each other – a consequence of differences in species dominance at specific age classes. Yet, the carabid assemblage of the 80-year-old stage did not approach that of the natural pine forest, but, unexpectedly, the 40–45-year-old age class did. Canopy cover and soil pH were the main environmental variables that affected the carabid distribution. We also detected species-level responses to the ageing process, with open habitat species more abundantly collected at the youngest stages and forest specialists at the older stages. We conclude that carabid beetles responded predictably to the pine plantation ageing process, and that although pine forests in NW Spain do not seem to provide habitat for unique species, they act as secondary habitats for forest specialist species.     

Determinants of local spider (Araneidae and Thomisidae) species richness on a regional scale: climate and altitude vs. habitat structure

Abstract.  1. The habitat heterogeneity hypothesis states that the more complex the habitat, the higher the species richness. The present study analyzes the effect of local factors on regional spider ( Araneidae and Thomisidae ) richness. The main objective is to disentangle the relative importance of habitat structure and other environmental variables.
2. Fifteen territorial units of 1 km² were sampled to obtain reliable and comparable inventories of the two spider families. Richness values were modelled using general regression models and a set of climate, topographic and vegetation structure variables. Pure and joint effects were computed via variation partitioning.
3. The results highlight the great importance of vegetation complexity, especially of grass and sub-shrub cover, in determining spider species richness.
4. The maximum temperature is the only climate variable significantly related to species richness, although its effect is combined with that of spatial and vegetation structure variables.
5. These results support the habitat heterogeneity hypothesis, and highlight the importance of taking vegetation complexity into account when managing habitats and where spider conservation is desired.     

Are distribution patterns linked to dispersal mechanism? An investigation using pond invertebrate assemblages

1. Dispersal ability influences the distribution and abundance of organisms, but empirical investigations of the relationship between dispersal ability and the composition of ecological assemblages are scarce. Here, we compare between-site variation in the species richness and community composition of actively and passively dispersing pond invertebrates.
2. Coleoptera (active dispersers) and microcrustacea (passive dispersers) were sampled over a season from 16 ponds within a 4-km radius in south-west England. Species richness and community composition were related to environmental variables using regression and Canonical Correspondence Analysis (CCA), respectively.
3. Coleopteran species richness was significantly and positively correlated with pond permanence and maximum area, whereas microcrustacean species richness was relatively equal across sites and did not correlate with environmental variables. The frequency of species' occurrence between sites was the same for both groups, which suggests that active and passive dispersers exhibited the same degree of dispersal.
4. Between-site variation in community composition was non-random for both groups, with pond permanence and area, together, explaining similar proportions of between-site variation for Coleoptera. Permanence was correlated most strongly with microcrustacean community composition and a high proportion (25%) of microcrustacean species were more numerous in smaller, more ephemeral ponds.
5. These data suggest that, at small spatial scales, Coleoptera which can undertake multiple dispersal events, are more likely to colonise large, more permanent ponds than passively dispersing microcrustacea. For microcrustacea, other traits (in this case those permitting existence in ephemeral habitats) may over-ride the influences of dispersal in driving between-site variation in species composition.     

Even the Smallest Non-Crop Habitat Islands Could Be Beneficial: Distribution of Carabid Beetles and Spiders in Agricultural Landscape

Carabid beetles and ground-dwelling spiders inhabiting agroecosystems are beneficial organisms with a potential to control pest species. Intensification of agricultural management and reduction of areas covered by non-crop vegetation during recent decades in some areas has led to many potentially serious environmental problems including a decline in the diversity and abundance of beneficial arthropods in agricultural landscapes. This study investigated carabid beetle and spider assemblages in non-crop habitat islands of various sizes (50 to 18,000 square metres) within one large field, as well as the arable land within the field, using pitfall traps in two consecutive sampling periods (spring to early summer and peak summer). The non-crop habitat islands situated inside arable land hosted many unique ground-dwelling arthropod species that were not present within the surrounding arable land. Even the smallest non-crop habitat islands with areas of tens of square metres were inhabited by assemblages substantially different from these inhabiting arable land and thus enhanced the biodiversity of agricultural landscapes. The non-crop habitat area substantially affected the activity density, recorded species richness and recorded species composition of carabid and ground-dwelling spider assemblages; however, the effects were weakened when species specialised to non-crop habitats species were analysed separately. Interestingly, recorded species richness of spiders increased with non-crop habitat area, whereas recorded species richness of carabid beetles exhibited an opposite trend. There was substantial temporal variation in the spatial distribution of ground-dwelling arthropods, and contrasting patterns were observed for particular taxa (carabid beetles and spiders). In general, local environmental conditions (i.e., non-crop habitat island tree cover, shrub cover, grass cover and litter depth) were better determinants of arthropod assemblages than non-crop habitat island size, indicating that the creation of quite small but diversified (e.g., differing in vegetation cover) non-crop habitat islands could be the most efficient tool for the maintenance and enhancement of diversity of ground-dwelling carabids and spiders in agricultural landscapes.     

Biological traits of stream macroinvertebrates from a semi-arid catchment: patterns along complex environmental gradients

1. The relationships between biological traits of macroinvertebrates and environmental characteristics were investigated in streams with contrasting physical, chemical or landscape level attributes. We used an ordination technique, RLQ analysis, which links an environmental table (R) with traits table (Q) through an abundance table (L) to investigate the relationship between habitat characteristics and biological traits.
2. A major environmental axis explaining the distribution of species and their distinctive biological features was obtained. This axis included variables of anthropogenic pressure (agricultural and urban uses) and natural variability (climatic and geologic) that are strongly intercorrelated in the study area, with a clear spatial component.
3. The attributes of species from frequently disturbed systems (small size, multivoltinism, diapause, ovoviviparity, etc.) were associated with semi-arid areas whereas traits common in more stable and favourable environments (large body size, semi-voltinism, isolated eggs, etc.) were found in upland forested areas.
4. The natural climatic variation was proposed as a disturbance axis of a theoretical habitat templet (driven by the intense hydrological disturbances typical of semi-arid streams), while anthropogenic pressure (mainly intensive agriculture) and high salinity, a natural consequence of geology, was proposed as an adversity axis. Different life-histories associated with contrasting environmental features were superimposed in this habitat templet.
5. The ecological–evolutionary scenario in which stream macroinvertebrates have evolved and by which their communities are organized, is closely linked to disturbance, environmental harshness and human pressure.     

Bacterial community succession in a high‐altitude subarctic glacier foreland is a three‐stage process

Alpine glaciers are retreating rapidly, exposing foreland minerals, which develop into soils. Bacterial communities in glacier forelands exhibit high rates of turnover and undergo dramatic shifts in composition within the first 50 years after deglaciation, followed by relative stabilization and convergence. This period of microbial development occurs simultaneously with plant colonization in most systems; thus, it remains unclear whether the changes in the bacterial communities occur primarily as the result of edaphic, climatic or biotic factors. We examined bacterial community structure along two replicate chronosequences within the glacial foreland of Duke River Glacier, Yukon, Canada. This foreland is estimated to include >200 years of bare soils before an appreciable grassline, likely due to the high latitude and altitude of the glacier. This enabled us to examine bacterial community development prior to plant colonization over a longer period than previous studies. We observed three successional groups in the chronosequence: (i) an ‘early’ group in soils of less than approximately 50 years since deglaciation; (ii) an ‘intermediate’ group within bare soils, after the early period but before the grassline, containing communities with a relatively high degree of variability in composition; and (iii) a ‘grassline’ group in soils collected after plant colonization with higher diversity but lower age‐group variability in community composition. These findings suggest rapid replacement and addition of species better adapted to glacier foreland conditions followed by slower community shifts over the next 150 years and, finally, indications of a possible response to plant colonization.     

Carabid beetle (Coleoptera: Carabidae) richness and functional traits in relation to differently managed grasslands in the Alps

Species richness, composition, and functional traits of carabid beetle assemblages (Coleoptera: Carabidae) were studied in relation to different grassland management. Carabid beetles were sampled during the summers 2008 and 2009 by 165 traps located in 11 sites in the central-eastern Italian Alps. Using mixed effect models to account for potential spatial bias, we found that mown grasslands had significantly more species, a lower proportion of wingless species and a lower proportion of species with long larval development than grazed and natural grasslands. Within grazed and mown grasslands, neither cattle density nor number of cuts had any significant effect neither on species richness nor on any of the traits. The influence of grassland management can be summarised as follows: (1) grazing does not change community structure and functional traits compared to natural grasslands; (2) mowing negatively affects the carabid beetle assemblages; (3) the intensity of grazing and of cutting may not affect the structure of species assemblages of ground beetles. Our results support the hypothesis that agroecosystem practices in alpine grasslands influence carabid beetle communities. Specifically, the species with traits typical of undisturbed habitats (low dispersal abilities and long larval development) are more sensitive to perturbations (e.g. cutting). Our suggestion for agricultural and environmental planning and for conservation schemes is that the preservation of natural grasslands (e.g. forest gaps) and the implementation of grazing should be promoted during the planning of agroecosystem mosaics.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Ecomorphological diversity of freshwater fishes as a tool for conservation priority setting: a case study from a Balkan hotspot

Biodiversity studies commonly focus on taxonomic diversity measures such as species richness and abundance. However, alternative measures based on ecomorphological traits are also critical for unveiling the processes shaping biodiversity and community assembly along environmental gradients. Our study presents the first analysis of habitat-trait-community structure in a Balkan biodiversity hotspot (Louros river, NW Greece), through the investigation of the relationships among freshwater fish assemblages’ composition, morphological traits and habitat features. In order to provide a hierarchical classification of species’ priority to protection measures, we highlight the most ecomorphologically distinct species using originality analysis. Our results suggest that the longitudinal changes of habitat variables (water temperature, depth, substrate, altitude) drive the local fish assemblages’ structure highlighting the upstream-downstream gradient. We also present evidence for environmental filtering, establishing fish assemblages according to their ecomorphological traits. The calculation of the seven available indices of ecomorphological originality indicates that Valencia letourneuxi and Cobitis hellenica, which are endemic to Louros and threatened with extinction, exhibited the highest distinctiveness; thus their protection is of great importance. The methodological approach followed and the patterns described herein can contribute further to the application of community ecology theory to conservation, highlighting the need to use ecomorphological traits as a useful ‘tool’.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Plant–pollinator network assembly along the chronosequence of a glacier foreland

Forelands of retreating glaciers offer an ideal model system to study community assembly processes during primary succession. As plants colonize the area that is freed from ice they should be accompanied by their pollinators to successfully reproduce and spread. However, little is known about the assembly of plant–pollinator networks. We therefore used quantitative network analysis to study the structure of plant–pollinator interactions at seven sites representing a chronosequence from 8 to 130 years since deglaciation on the foreland of the Morteratsch glacier (southeastern Switzerland). At these sites, individual visits of plant flowers by insects were recorded throughout the flowering season. Species richness of insect‐pollinated plants and plant‐pollinating insects, together with measures of interaction diversity and evenness, increased along the chronosequence at least for the first 80 years after deglaciation. Bees were the most frequent flower visitors at the two youngest sites, whereas flies dominated in mature communities. Pollinator generalization (the number of visited plant species weighted by interaction strength), but not plant generalization, strongly increased during the primary succession. This was reflected in a pronounced decline in network level specialization (measured as Blüthgen's H₂’) and interaction strength asymmetry during the first 60 years along the chronosequence, while nestedness increased along the chronosequence. Thus, our findings contradict niche‐theoretical predictions of increasing specialization of pollination systems during succession, but are in agreement with expectations from optimal foraging theory, predicting an increase in pollinator generalization with higher plant diversity but similar flower abundance, and an increase in diet breadth at higher pollinator densities during primary succession.     

Staphylinidae and Carabidae overwintering in wheat and sown wildflower areas of different age

Species richness and abundance of staphylinid and carabid beetles overwintering in winter wheat fields and 1- to 3-year-old wildflower areas were investigated during 2000/2001 on 16 study sites in Switzerland. Abundance and species richness of overwintering staphylinids significantly increased with successional age of the wildflower areas and were always higher in older wildflower areas than in winter wheat. A similar but less distinct pattern was observed for the abundance and species richness of carabid beetles. The influence of habitat parameters (vegetation cover, fine sand content, organic matter, pH, soil pore volume, surrounding landscape structure, habitat area) on the staphylinid and carabid assemblages based on the number of individuals per species and site was analysed using canonical correspondence analysis. Vegetation cover was the most significant parameter significantly characterizing both staphylinid and carabid assemblages. The amount of vegetation cover explained 15.7% of the variance, fine sand content accounted for 13.3% and surrounding landscape structure for 10.9% of the variance in the staphylinid assemblage. In the carabid assemblage, vegetation cover was the only significant factor, explaining 24.7% of the variance. This study showed for the first time that the significance of wildflower areas as a reservoir for hibernation for generalist predatory beetles increases with progressing successional age.