Effects of macro and micro-environmental factors on the species richness of terrestrial tardigrade assemblages in an Iberian mountain environment

Tardigrade communities are affected by micro and macro-environmental conditions but only micro-environmental variables, and altitudinal gradients have been studied. We review previous reports of altitudinal effects and evaluate the influence by interacting macro- (climate, soils, biome, and others) and micro-environmental (vegetation, moss and leaf litter) factors on tardigrade assemblages at the Sierra de Guadarrama mountain range (Iberian Central System Mountains, Spain). Terrestrial tardigrade assemblages were sampled using standard cores to collect leaf litter and mosses growing on rocks. General Linear Models were used to examine relationships between Tardigrada species richness and abundance, and macro- and micro-environmental variables (altitude, habitat characteristics, local habitat structure and dominant leaf litter type, and two bioclimatic classifications). Variation partitioning techniques were used to separate the effects of altitude and habitat variation, and to quantify the independent influences of climate and soil, vegetation structure and dominant type of leaf litter. Altitude shows a unimodal relationship with tardigrade species richness, although its effect independent of habitat variation is negligible. The best predictors for species richness were bioclimatic classifications. Separate and combined effects of macro-environmental gradients (soil and climate), vegetation structure and leaf litter type are important determinants of richness. A model including both macro- and micro-environmental variables explained nearly 60% of tardigrade species richness in micro-scale plots. Abundance was significantly related only to soil composition and leaf litter type. Tardigrade abundance was not explained by macro-environmental gradients analysed here, despite a significant correlation between abundance and richness.