Macroinvertebrate diversity in alpine lakes: Effects of altitude and catchment properties

Lakes are common features of alpine landscapes, and the attention given to alpine lakes has increased recently in response to increased recognition of the important role that these freshwaters play as sensible indicators of climate change. Despite this general research interest, there is nevertheless a general lack of information about zoobenthos especially of lakes in the Alps, and only few published data are available, which has made it nearly impossible to draw general conclusions in respect to benthic community structure, profundal and/or littoral food webs. This paper aims to explore the relationships between main environmental/catchment properties of 55 lakes and their littoral benthic fauna across three regions of the Alps. We provide updated information on relative abundance, species richness, distribution and ecology of macroinvertebrates which occur and are typical in the littoral of high-mountain lakes of the Alps. These lakes were located in the Lago-Maggiore Watershed (Italy and Switzerland), in South Tyrol (Italy) and in North/East Tyrol (Austria), between 1840 and 2796 m a.s.l., in catchments undisturbed by human activities. As the studied lakes are situated above the tree line, they were characterised by low nutrient levels indicating an oligotrophic status. Lake water chemistry corresponded closely to the geo-lithology of the catchment and some parameters (especially nutrient concentrations) differed between the regions. The macroinvertebrates were dominated by insects which to a high degree were chironomid larvae and pupae. Other insect orders were typical cold stenotherm species of Ephemeroptera, Plecoptera and Trichoptera. Non-insect macroinvertebrates contributed to the 144 taxa found. Other than lake size and catchment area, the faunal parameters exhibited a clearer pattern along altitude. Macroinvertebrates per sample increased with higher elevation, reached their maximum in lakes between 2400 and 2600 m a.s.l., but decreased strongly above 2600 m. The altitudinal pattern of species richness and Shannon diversity resembled each other being highest between 2001 and 2200 m a.s.l., but decreased when going lower and higher, respectively. Various patterns and trends along altitude were also evident when individual groups were analysed within the individual sampling regions. The somewhat conflicting trends of various biocoenotic indices let assume that factors other than altitude are also responsible for the structure of faunal assemblages in the littoral of alpine lakes. Six variables ("bare rocks" and "nitrate", "alkalinity", "ammonia" and "peat bog") were selected by the CCA analysis where these three groups of lakes were identified: (1) lakes with a higher alkalinity (higher pH, conductivity, ion concentration), a higher relative vegetation cover (compared to the "bare rocks" on the opposite side) and lower nitrate levels; (2) lakes with a higher portion of "bare rocks" in their catchments and higher nitrate levels; and (3) a smaller group of lakes with higher ammonia levels and a boggy environment. Geographical patterns seemed to have weak effects on the presence of taxa while catchment properties had evident impacts on macroinvertebrate communities in these lakes. In this way, the present study contributes to the overall understanding of environmental settings and effects on high mountain lake ecosystems and assists in refining research and conservation strategies for an important landscape aspect in the Alps.