2002; Ewers and Didham 2006). Accordingly, in several cases positive SA-relationships have been observed

for habitat-specific species, but not for total species numbers (Lövei et al. 2006; Magura et al. 2001; Vries de et al. 1996). Several hypotheses have been proposed to explain the SAR, two of the most prominent being the ‘area per se hypothesis’ (Preston 1960; MacArthur and Wilson 1967) and the ‘habitat heterogeneity hypothesis’ (Williams 1964). The area per se hypothesis is based on assumptions that probabilities of extinction and colonization will generally be lower and Adavosertib in vitro higher, respectively, in larger areas, while the habitat heterogeneity hypothesis assumes that habitats will be more diverse in larger areas and therefore more species will be able to live in them. Both hypotheses probably partially explain the SAR, although it is difficult to distinguish their relative effects (Connor and McCoy 1979). Efforts to evaluate their relative importance have had varying results (e.g., Báldi 2008; Kallimanis et al. 2008). Attempts have also been made to unify the two hypotheses (Triantis et al. 2003). In this study, the beetle assemblages of 13 sand pits in east-central Sweden were examined

to evaluate the effects of the area of sand pits on the number and composition of species they host. A positive SAR was expected for the selleck inhibitor target species, i.e., specialist species of open sandy habitats (here termed sand species). The effects of four additional habitat characteristics were also tested: the proportion of sand material at the surface, vegetation cover, tree cover and edge habitat. As beetles are a very diverse group we specifically analyzed carabids (Carabidae) in order to see if they could be used as an indicator of diversity for the whole order. Carabids could be useful indicators as they are well known (taxonomically and ecologically), they can be easily and cost-efficiently sampled by pitfall traps, and they include many species confined to habitats in an early

successional stage (Ljungberg 2002; Rainio and Niemelä 2003). Finally, we used our data to draw conclusions with respect to conservation measures for sand pits. More specifically we addressed the following questions: Does the area of sand pits influence beetle ID-8 species number and composition? Does the surrounding matrix influence SAR? Do other examined variables (proportion of sand material, vegetation cover, tree cover and edge habitat) influence beetle diversity? Can carabids be used as a diversity indicator for all beetle species in sand pits? Based on our results, what recommendations can be made for species conservation in sand pits? Materials and methods Study region and study sites The study focused on 13 sites located along three eskers (Enköpings-, Vattholma- and Uppsalaåsen) in Uppsala County, east-central Sweden (Fig. 1).