Claudia Drees

Value of semi-open corridors for simultaneously connecting open and wooded habitats: a case study with ground beetles.

To counteract habitat fragmentation, the connectivity of a landscape should be enhanced. Corridors are thought to facilitate movement between disconnected patches of habitat, and linear strips of habitat connecting isolated patches are a popular type of corridor. On the other hand, the creation of new corridors can lead to fragmentation of the surrounding habitat. For example, heathland corridors connect patches of heathland and alternatively hedgerows connect patches of woodland. Nevertheless, these corridors themselves also break up previously connected patches of their surrounding habitat and in so doing fragment another type of habitat (heathland corridors fragment woodlands and woodland strips or hedgerows fragment heathlands). To overcome this challenge we propose the use of semi-open habitats (a mixture of heathland and woodland vegetation) as conservation corridors to enable dispersal of both stenotopic heathland and woodland species. We used two semi-open corridors with a mosaic of heathland and woody vegetation to investigate the efficiency of semi-open corridors for species dispersal and to assess whether these corridors might be a suitable approach for nature conservation. We conducted a mark-recapture study on three stenotopic flightless carabid beetles of heathlands and woodlands and took an inventory of all the carabid species in two semi-open corridors. Both methodological approaches showed simultaneous immigration of woodland and heathland species in the semi-open corridor. Detrended correspondence analysis showed a clear separation of the given habitats and affirmed that semi-open corridors are a good strategy for connecting woodlands and heathlands. The best means of creating and preserving semi-open corridors is probably through extensive grazing.

Multiple Glacial Refugia of the Low-Dispersal Ground Beetle Carabus irregularis: Molecular Data Support Predictions of Species Distribution Models

Classical glacial refugia such as the southern European peninsulas were important for species survival during glacial periods and acted as sources of post-glacial colonisation processes. Only recently, some studies have provided evidence for glacial refugia north of the southern European peninsulas. In the present study, we combined species distribution models (SDMs) with phylogeographic analyses (using mitochondrial DNA = mtDNA) to investigate if the cold-adapted, stenotopic and flightless ground beetle species, Carabus irregularis, survived the Last Glacial Maximum (LGM) in classical and/or other refugia. SDMs (for both a western European and for a Carpathian subgroup) were calculated with MAXENT on the basis of 645 species records to predict current and past distribution patterns. Two mtDNA loci (CO1 and ND5, concatenated sequence length: 1785 bp) were analyzed from 91 C. irregularis specimens to reconstruct the phylogeography of Central and eastern European populations and to estimate divergence times of the given lineages. Strong intra-specific genetic differentiation (inter-clade ΦST values ranged from 0.92 to 0.99) implied long-term isolation of major clades and subsclades. The high divergence between the nominate subspecies and the Carpathian subspecies C. i. montandoni points to two independent species rather than subspecies (K-2P distance 0.042 ± 0.004; supposed divergence of the maternal lineages dated back 1.6 to 2.5 million years BP) differing not only morphologically but also genetically and ecologically from each other. The SDMs also inferred classical as well as other refugia for C. irregularis, especially north of the Alps, in southeastern Europe and in the Carpathians. The coincidences between the results of both methods confirm the assumption of multiple glacial refugia for the studied species and the usefulness of combining methodological approaches for the understanding of the history of low-dispersal insect species.

Multiple Glacial Refuges of Unwinged Ground Beetles in Europe: Molecular Data Support Classical Phylogeographic Models

Since the 1930s, several European zoologists have developed scenarios for glacial refuges and postglacial expansions, mainly based on studies of the morphological differentiation of populations and distribution patterns of species. For example, Holdhaus described the distribution of blind euedaphic and troglobitic beetles restricted to an area South of a well-defined line crossing the Southern Europe from West to East. In these areas, where many endemic animal and plant species occur, other species that are currently more widely distributed in Europe were probably able to survive the glacial period(s). Molecular analyses of 77 populations of the silvicolous ground beetle Carabus auronitens support the existence of these postulated refuge areas. Genetic differentiation of C. auronitens provides good evidence for multiple refuges, which are, however, situated further North than previously assumed. Furthermore, genetic differentiation is more pronounced in the areas South of the “Holdhaus line” than in the areas North of it.

Review Refugial Areas and Postglacial Colonizations in the Western Palearctic

Past climatic oscillations, such as the glacial-interglacial cycles of the Pleistocene (Williams et al. 1998) led to severe worldwide altitudinal and latitudinal range shifts of taxa and ecosystems (Hewitt 2004). For a reconstruction of the glacial and postglacial history of species, data obtained from methods ranging from evolutionary genetics to the analysis of distribution patterns and studies of pollen and subfossil remains of invertebrates and vertebrates are essential (de Lattin 1967; Coope 1970, 1978, 1994; Varga 1977; Huntley and Birks 1983; Taberlet et al. 1998; Willis et al. 1995; Hewitt 1999, 2000; Willis and van Andel 2004; Schmitt 2007; Varga and Schmitt 2008). The synthesis of all these analyses reveals the existence of different paradigm patterns in the Western Palearctic. In the following, we give a brief overview of these patterns.

Review: The Dark Side of Relict Species Biology: Cave Animals as Ancient Lineages

Due to their fascinating biology and phenomena belonging to the realm of scientific curiosity, cave animals have been objects of study for zoologists for numerous decades. This chapter not only focuses on the extremes (e.g., absence of eyes, specialization to extreme environments), but also serves as an introduction to understand the geographic distribution patterns and history of these highly diverse ecological groups with their relict characteristics. After an introduction to the subterranean environment in Sect. 1, we briefly review the biology and ecology of cave animals with their regressive and progressive evolutionary tendencies in order to understand the innate reasons for restricted distribution patterns (Sect. 2). In Sect. 3, we summarize the main aspects of our knowledge regarding the distribution of these species, especially in the Holarctic; and finally in Sect. 4, we highlight the relict characteristics of cave animal distribution and the ancient phylogenetic splits between cave and surface lineages.

Poleward range expansion without a southern contraction in the ground beetle Agonum viridicupreum (Coleoptera, Carabidae).

Abstract We investigated the extent of poleward shifts in the distribution range of Agonum viridicupreum due to climate change in the western Palaearctic. Species’ records were obtained from extensive literature sources as well as from collections, and consistent amateur entomologists’ recordings. Within the general geographic range of the species, we analyzed in detail two parts of both, the northern and southern distribution range boundaries: (1 and 2) north-western Germany (leading or high-latitude edge), (3) Israel and (4) southern Italy (rear or low-latitude edge). Temporal changes in the occurrence data of the species indicated a northward shift of the leading edge of a minimum of 100 km within the last 50 to 100 years. In contrast, according to the data gathered, the rear edge has not changed during the last decades. Further studies are needed in order to fully understand the underlying mechanisms of the different behaviour of leading and rear range edges of Agonum viridicupreum in the current context of global change. Despite our incomplete understanding, chronosequences of the occurrence of the given species have the potential to optimize climate niche modelling to predict trends in the distribution range in the future.

Does isolation affect phenotypic variability and fluctuating asymmetry in the endangered Red Apollo

Isolated populations represent one of the main focuses in conservation biology. Long-term isolation often causes losses of genetic diversity and as a consequence might reduce individual fitness. Morphometric characters can be used as suitable markers to analyse ecological stress and individual fitness of local populations. Asymmetry in bilateral symmetry is used as a measure for developmental instability of populations and is often negatively correlated with population size and low genetic diversity. As a study system, we selected the endangered butterfly Parnassius apollo, which occurs in small and isolated remnant populations in Central Europe, but also in fairly large metapopulations in the Alps. We analysed wing morphometrics (shape and size characters) of 812 individuals representing (1) already extinct, (2) highly isolated and (3) still interconnected populations. Seventeen landmarks on veins were used to analyse morphological variances in the wing shape. Our data show significant deviations between landmarks on the left and right wing side within individuals and strong morphological variance among individuals. The highest morphological variability could be found for individuals in the Alps, however, the level of asymmetry was very similar for all populations analysed. The higher morphological variability found in the Alps can be interpreted as a consequence of the higher level of the genetic diversity detectable for this region. Analysis on morphological variance of P. apollo individuals of the Mosel valley using time series ranging from 1895 until today showed no significant rise in asymmetry and no decline of morphological variability over time, although, anthropogenic habitat destruction has caused severe bottlenecks in this population.

Evolutionarily significant units in a flightless ground beetle show different climate niches and high extinction risk due to climate change

Species distribution models (SDMs), especially those basing on climatic parameters, have frequently been used to project future species ranges and to develop conservation strategies. As suggested by several authors, we considered both different dispersal abilities and different evolutionarily significant units (ESUs, as determined in an earlier genetic survey). For our study species, the flightless ground beetle Carabus irregularis, SDMs for two ESUs from the western and the Carpathian area of the distribution range showed immense, and deviating future range contractions reflecting divergent ecological requirements. As minimal dispersal SDMs resulted in a stronger decline of future ranges than the maximal dispersal models, low dispersal ability tended to strengthen the already high vulnerability of the cold-adapted mountain species to global warming. Areas shown in our maximal dispersal models as offering climatically suitable habitats for C. irregularis in the future should be considered as potential areas of action in future conservation planning (e.g. assisted migration or assisted colonisation). Thus, both dispersal scenarios and different (if applicable) ESUs should be considered when developing SDMs as useful tools for species conservation strategies adapted to species’ performance and differentiation patterns.

Land-use type and intensity differentially filter traits in above- and below-ground arthropod communities

Along with the global decline of species richness goes a loss of ecological traits. Associated biotic homogenization of animal communities and narrowing of trait diversity threaten ecosystem functioning and human well-being. High management intensity is regarded as an important ecological filter, eliminating species that lack suitable adaptations. Below-ground arthropods are assumed to be less sensitive to such effects than above-ground arthropods. Here, we compared the impact of management intensity between (grassland vs. forest) and within land-use types (local management intensity) on the trait diversity and composition in below- and above-ground arthropod communities. We used data on 722 arthropod species living above-ground (Auchenorrhyncha and Heteroptera), primarily in soil (Chilopoda and Oribatida) or at the interface (Araneae and Carabidae). Our results show that trait diversity of arthropod communities is not primarily reduced by intense local land use, but is rather affected by differences between land-use types. Communities of Auchenorrhyncha and Chilopoda had significantly lower trait diversity in grassland habitats as compared to forests. Carabidae showed the opposite pattern with higher trait diversity in grasslands. Grasslands had a lower proportion of large Auchenorrhyncha and Carabidae individuals, whereas Chilopoda and Heteroptera individuals were larger in grasslands. Body size decreased with land-use intensity across taxa, but only in grasslands. The proportion of individuals with low mobility declined with land-use intensity in Araneae and Auchenorrhyncha, but increased in Chilopoda and grassland Heteroptera. The proportion of carnivorous individuals increased with land-use intensity in Heteroptera in forests and in Oribatida and Carabidae in grasslands. Our results suggest that gradients in management intensity across land-use types will not generally reduce trait diversity in multiple taxa, but will exert strong trait filtering within individual taxa. The observed patterns for trait filtering in individual taxa are not related to major classifications into above- and below-ground species. Instead, ecologically different taxa resembled each other in their trait diversity and compositional responses to land-use differences. These previously undescribed patterns offer an opportunity to develop management strategies for the conservation of trait diversity across taxonomic groups in permanent grassland and forest habitats.

Living in Heterogeneous Woodlands – Are Habitat Continuity or Quality Drivers of Genetic Variability in a Flightless Ground Beetle?

Although genetic diversity is one of the key components of biodiversity, its drivers are still not fully understood. While it is known that genetic diversity is affected both by environmental parameters as well as habitat history, these factors are not often tested together. Therefore, we analyzed 14 microsatellite loci in Abax parallelepipedus, a flightless, forest dwelling ground beetle, from 88 plots in two study regions in Germany. We modeled the effects of historical and environmental variables on allelic richness, and found for one of the regions, the Schorfheide-Chorin, a significant effect of the depth of the litter layer, which is a main component of habitat quality, and of the sampling effort, which serves as an inverse proxy for local population size. For the other region, the Schwäbische Alb, none of the potential drivers showed a significant effect on allelic richness. We conclude that the genetic diversity in our study species is being driven by current local population sizes via environmental variables and not by historical processes in the studied regions. This is also supported by lack of genetic differentiation between local populations sampled from ancient and from recent woodlands. We suggest that the potential effects of former fragmentation and recolonization processes have been mitigated by the large and stable local populations of Abax parallelepipedus in combination with the proximity of the ancient and recent woodlands in the studied landscapes.