Oliver-D. Finch

Macroecological patterns of spider species richness across Europe

We analysed the pattern of covariation of European spider species richness with various environmental variables at different scales. Four layers of perception ranging from single investigation sites to the whole European continent were selected. Species richness was determined using published data from all four scales. Correlation analyses and stepwise multiple linear regression were used to relate richness to topographic, climatic and biotic variables. Up to nine environmental variables were included in the analyses (area, latitude, elevation range, mean annual temperature, local variation in mean annual temperature, mean annual precipitation, mean July temperature, local variation in mean July temperature, plant species richness). At the local and at the continental scale, no significant correlations with surface area were found, whereas at the landscape and regional scale, surface area had a significant positive effect on species richness. Factors that were positively correlated with species richness at both broader scales were plant species richness, elevation range, and specific temperature variables (regional scale: local variation in mean annual, and mean July temperature; continental scale: mean July temperature). Latitude was significantly negatively correlated with the species richness at the continental scale. Multiple models for spider species richness data accounted for up to 77% of the total variance in spider species richness data. Furthermore, multiple models explained variation in plant species richness up to 79% through the variables mean July temperature and elevation range. We conclude that these first continental wide analyses grasp the overall pattern in spider species richness of Europe quite well, although some of the observed patterns are not directly causal. Climatic variables are expected to be among the most important direct factors, although other variables (e.g. elevation range, plant species richness) are important (surrogate) correlates of spider species richness.

Ground beetle diversity in ancient woodland remnants in north-western Germany (Coleoptera, Carabidae)

We investigated ground beetle communities (Coleoptera, Carabidae) in ancient woodland remnants in north-western Lower Saxony, Germany. A total of 90 pitfall traps was exposed in a stratified design in 10 stands of mature oak–beech and oak–hornbeam forests in the year 2003. Overall, 47 species (10,676 individuals) were recorded. Among these were the two relict species Carabus glabratus and Abax parallelus, and 14 further eurytopic forest species. Eleven species exhibited a high frequency and were found in all of the ten stands. Multiple linear regressions showed several significant relationships at two scales for species richness of different groups of carabids and for several of the measured environmental factors. Forest area, litter depth, amount of dead wood, distance to forest edge, and soil moisture were found to be key factors determining species richness. Furthermore, recent disturbance by logging reduced the number of forest species. According to direct gradient analyses soil moisture and litter depth have greatest influence on species communities of both, forest species and widespread species. Habitat suitability models for the two recorded relict species, A. paralellus and C. glabratus, were developed using logistic regression. The presence of A. parallelus in the mature ancient woodland remnants depends mainly on higher values of soil moisture, whereas for C. glabratus none of the measured environmental variables appeared to be key factors. Implications for the conservation of carabid assemblages in mature ancient woodlands include the advice to spread out logging over long periods of time and over various woodlands in order to keep the stand disturbance at a long-term low level. Variation in logging practices may help to conserve diverse structures. Afforestation with non-native tree species should be avoided in the managed ancient woodlands. Finally, especially the preservation of a high soil moisture seems to be important to conserve typical carabid communities.

The parasitoid complex and parasitoid‐induced mortality of spiders (Araneae) in a Central European woodland

The parasitoid complex of spiders was studied in three different sites of a forest and, for comparison, in open habitats in north‐western Lower Saxony (Germany). Qualitative data and, for four spider species, detailed quantitative rearing data were obtained. At least 25 parasitoids of spiders were recorded (23 species of Hymenoptera and two species of Diptera). External parasitoids of spiders were rare in the woodlands; only up to 1% of the spiders collected by hand searching and sweep netting had ectoparasitoids. During the investigation 23 parasitoid species were reared from spider egg masses, 10 of them occurring in woodlands. Two species were secondary parasitoids. Several parasitoid–host relationships were recorded for the first time. Egg masses of Floronia bucculenta were parasitized up to 5% by two species of the genus Gelis. Up to 10% of the egg masses of Linyphia hortensis were parasitized by Aclastus species. Egg masses of the spider genus Ero were parasitized by three species, causing an overall mortality of 40%. For egg masses of Agroeca spp. parasitization at some locations was up to 60%. An overview of the Central European parasitoid complex of spiders is provided.

Indicators of species richness at the local scale in an alpine region: a comparative approach between plant and invertebrate taxa

Studies investigating congruent variations in species richness patterns in alpine habitats are scarce. We investigated the potential of using the indicator taxa approach for species richness in alpine habitats of the Scandes (Norway). In four areas, we investigated seven functional and taxonomic terrestrial groups of organisms and evaluated their contribution to the species diversity. The function of each group as a surrogate for the overall species diversity or for the diversity of another taxon was analysed. Three groups of invertebrates (spiders without Lycosids, Lycosids only, and ground beetles), three groups of plants (shrubs, graminoids, and herbs), and lichens were used for a cross-taxon analysis of species diversity. Congruence in species richness was restricted to several significant results, with vascular plants and spiders (Araneae) being best suited as surrogate taxa in alpine habitats of the Scandes. In the cross-taxon analyses they showed strongest significant positive correlations, covering the total species richness of the alpine habitats best. Species counts in one group account for up to 70% of the variation in total species richness. We found only limited evidence for an ideal, efficient biodiversity indicator taxon that could be applied without restrictions at different alpine habitats in low and middle alpine areas. Thus, our results suggest that it is very important to use more than one taxon as indicator for species richness in terrestrial alpine habitats. This should facilitate future conservation planning in alpine areas.

Spatio-temporal Gradients between High Mountain Ecosystems of Central Norway

Abstract Spatio-temporal gradients between ecosystems of the central Norwegian high mountains are analyzed. Complex landscape ecological site analyses combined with intensive pitfall trapping are carried out in four investigation areas in two regions. Key questions are addressed for the differences among ecosystems along a broad-scaled oceanic-continental gradient. The answers are based on ecological process analysis and mapping of zoocoenoses in small catchments of two alpine altitudinal belts. A comparison of four ridge sites is presented by analyzing water and temperature balance and activity of arthropods during the driest summer month. The results do not implicate summer drought and heat as limiting factors; summer wetness and cooling are most decisive. Landscape ecological processes, like the accumulation of snow during winter, snow melting, freeze-thaw action, percolation, soil moisture variation, and temperature regimes are exemplified by long-term measurements throughout the year in a small catchment in continental eastern Norway—the driest mountain region in Scandinavia. To learn about the organization and diversity of zoocoenoses, epigeic arthropods (Araneae, Carabidae) are investigated along spatial gradients. Interrelations between distribution patterns of animals and spatio-temporal dynamics of the environmental conditions are presented. The results are framed by gradient theory in landscape ecology. Finally, the complexity of spatio-temporal gradient determination between ecosystems is discussed and summarized by a scheme of gradient principles for the Norwegian mountains.

Effects of the abandonment of alpine summer farms on spider assemblages (Araneae)

In parallel with the current effects and responses to climate change ongoing changes of land use strongly affect alpine areas. The aim of this study was to analyse spider assemblages occurring on farmlands with differing stages of land use intensity. We collected the ground-dwelling spider fauna of semi-natural grasslands in currently farmed and abandoned farmland in the alpine zone of Hardangervidda, Norway, using pitfall traps. Trapping during the whole vegetation period resulted in 1,548 individuals belonging to 39 species of spiders. Linyphiids and lycosids dominated. The total number of species differ little between sites, but there was a general trend of increasing proportions of lycosids, gnaphosids and thomisids individuals and species, and a decrease in the proportion of linyphiids with farmland abandonment. Two main groupings of spider assemblages were found using a principal component analysis: One grouping was characteristic of intensely disturbed sites, mainly dominated by typical pioneer species. The other was characteristic of abandoned and less disturbed sites, dominated by widespread species. The abandonment of summer farms led to a succession in spider assemblages, including a decline in spider species with a high ballooning activity and an increase of species with a larger body size. We suggest that the main reason lies in the absence of disturbance by grazing including differing vegetation architecture and the proportion of bare ground with abandonment. Spider assemblages react quickly to environmental changes that occur in alpine habitats after abandonment. Thus, it is necessary to include spiders and other invertebrate animal indicator groups in field studies and experiments dealing with grazing impacts on alpine habitats.

Elevational variation of reproductive traits in five Pardosa (Lycosidae) species

Abstract Differentiations in reproductive traits along climatic gradients can be substantial for a species to spread along a wide spatial range. We compared the reproductive effort allocated to first egg sacs of five species of the genus Pardosa: P. palustris (Linnaeus 1758), P. amentata (Clerck 1757), P. lugubris (Walckenaer 1802), P. hyperborea (Thorell 1872), and P. riparia (C. L. Koch 1833) along three elevation transects in central Norway. We tested whether population differences are consistent among the three transects, respectively along the elevational gradient. We assumed that the harsh environments of alpine areas would lead to adaptations in reproductive traits resulting in larger eggs but smaller clutches at higher elevations. The results show that female size and egg number were positively correlated among all species. However, no clear elevation-related trend was found. Other traits did not change consistently between species and along the elevational gradient. We assume that local microclimatic impacts on spider fitness are a crucial but poorly understood factor. Without further knowledge about adaptation and phenotypic plasticity in ectotherms, modeling of possible future reproduction biology might remain flawed.

Alpine activity patterns of Mitopus morio (Fabricius, 1779) are induced by variations in temperature and humidity at different scales in central Norway

Our research addresses questions about how micro-climate affects activity abundance of a common and widespread harvestman in an alpine ecosystem. Activity patterns of the Harvestman Mitopus morio (Fabricius, 1779) were studied along different alpine gradients in the central Norwegian Scandes. Within a nested design, we surveyed 18 alpine habitats with pitfall traps and microclimatological equipment along oceanic-continental, two elevational, and (fine-scaled) microtopographic gradients. Sites in the oceanic region of the Scandes showed generally higher abundance of M. morio than sites in the continental region. Furthermore, along the elevational gradient, middle-alpine sites showed higher abundances than low-alpine sites. These general patterns are best explained by higher humidity in the oceanic region and in the middlealpine belt. Focusing at a finer scale, i.e. one elevational level within each region, revealed partly opposing activity patterns within relatively short distances. While in the western middle-alpine belt these patterns were best explained by humidityrelated measures but now with higher activity abundance during drier conditions, in the drier eastern middle-alpine belt heat sums rather than humidity were found to be the best explanatory variables for the observed patterns. Hence, our results imply a pronounced different reaction of the two populations towards climatic variables that partly even contradict the previously described general pattern. Regardless whether these differences in activity abundance in M. morio are a form of phenotypic plasticity or adaptation, our findings stress the importance of detailed autecological knowledge combined with fine-scaled climatic measurements when aiming at predictions about possible future ecosystem structures and spatiotemporal phenomena. M. morio proves to be an ideal biogeographic model organism for understanding spatio-temporal responses of alpine ecosystems under modified climatic conditions.