Mollusc community patterns and species response curves along a mineral richness gradient:: a case study in fens

Aim The goals of this study were to: (1) compare water conductivity and pH as proxy measures of mineral richness in relation to mollusc assemblages in fens, (2) examine the patterns of mollusc species richness along the gradient of mineral richness based on these factors, (3) model species-response curves and analyse calcicole-calcifuge behaviour of molluscs, and (4) compare the results with those from other studies concerning non-marine mollusc ecology. Location Altogether, 135 treeless spring fen sites were sampled within the area of the Western Carpathians (east Czech Republic, north-west Slovakia and south Poland; overall extent of study area was 12,000 km(2)). Methods Mollusc communities were recorded quantitatively from a homogeneous area of 16 m(2). Water conductivity and pH were measured in the field. The patterns of local species diversity along selected gradients, and species-response curves, were modelled using generalized linear models (GLM) and generalized additive models (GAM), both using the Poisson distribution. Results When the most acid sites (practically free of molluscs) were excluded, conductivity expressed the sites' mineral richness and base saturation within the entire gradient, in contrast to pH. In the base-rich sites, pH did not correlate with mineral richness. A unimodal response of local species diversity to mineral richness (expressed as conductivity) was found. In the extremely mineral-rich, tufa-forming sites (conductivity > 600 mu S cm(-1)) a decrease in species diversity was encountered. Response curves of the most common species showed clear differentiation of their niches. Significant models of either unimodal or monotonic form were fitted for 18 of the 30 species analysed. Species showed five types of calcicole-calcifuge behaviour: (1) a decreasing monotonic response curve and a preference for the really acid sites; (2) a skewed unimodal response curve with the optimum shifted towards the slightly acid sites; (3) a symmetrical unimodal model response curve with the optimum in the base-rich sites, with no or slight tufa precipitation; (4) a skewed unimodal response curve but with the optimum shifted to the more mineral-rich sites; and (5) an increasingly monotonic response curve, the optimum in the extremely base-rich sites with strong tufa precipitation. Main conclusions Conductivity is the only reliable proxy measure of mineral richness across the entire gradient, within the confines of this study. This information is of great ecological significance in studies of fen mollusc communities. Species richness does not increase with increasing mineral richness along the entire gradient: only a few species are able to dwell in the extremely base-rich sites. The five types of calcicole-calcifuge behaviour seen in species living in fens have a wider application: data published so far suggest they are also applicable to mollusc communities in other habitats.