RESUMEN
Environmental heterogeneity is an important driver of ecological communities. Here, we assessed the effects of local and landscape spatial environmental heterogeneity on ant community structure in temperate seminatural upland grasslands of Central Germany. We surveyed 33 grassland sites representing a gradient in elevation and landscape composition. Local environmental heterogeneity was measured in terms of variability of temperature and moisture within and between grasslands sites. Grassland management type (pasture vs. meadows) was additionally included as a local environmental heterogeneity measure. The complexity of habitat types in the surroundings of grassland sites was used as a measure of landscape environmental heterogeneity. As descriptors of ant community structure, we considered species composition in terms of nest density, community evenness, and functional response traits. We found that extensively grazed pastures and within-site heterogeneity in soil moisture at local scale, and a high diversity of land cover types at the landscape scale affected ant species composition by promoting higher nest densities of some species. Ant community evenness was high in wetter grasslands with low within-site variability in soil moisture and surrounded by a less diverse landscape. Fourth-corner models revealed that ant community structure response to environmental heterogeneity was mediated mainly by worker size, colony size, and life history traits related with colony reproduction and foundation. We discuss how within-site local variability in soil moisture and low-intensity grazing promote ant species densities and highlight the role of habitat temperature and humidity affecting community evenness. We hypothesize that a higher diversity of land cover types in a forest-dominated landscape buffers less favorable environmental conditions for ant species establishment and dispersal between grasslands. We conclude that spatial environmental heterogeneity at local and landscape scale plays an important role as deterministic force in filtering ant species and, along with neutral processes (e.g., stochastic colonization), in shaping ant community structure in temperate seminatural upland grasslands.
RESUMEN
Extensively managed grasslands are globally recognized for their high biodiversity value. Over the past century, a continuous loss and degradation of grassland habitats has been observed across Europe that is mainly attributable to agricultural intensification and land abandonment. Particularly insects have suffered from the loss of grassland habitats due to land-use change and the decrease in habitat quality, either due to an increase in livestock density, higher mowing frequency, and an increase in nitrogen fertilization, or by abandonment. However, only a few studies have used nationwide datasets to analyse the effects of land cover and land-use intensity on insects. It further remains largely unexplored how these effects are modulated by species traits, i.e. habitat specialisation and mobility. Using nationwide butterfly data originating from the German Butterfly Monitoring Scheme, we investigated the effect of three indicators related to land cover and agricultural land-use intensity on species richness as well as trait composition of butterfly communities. Based on agricultural census data at the municipality scale, we calculated the share of permanent grasslands (measure of habitat availability), the total livestock density (proxy for organic fertilization) and the livestock density of domestic herbivores (proxy for management intensity in grasslands) within a 2 km buffer surrounding each butterfly transect. To analyse the relationships between butterflies and indicators of land cover and land-use intensity, we applied generalised linear mixed effect models. We found a negative relationship between butterfly species richness and the livestock density of domestic herbivores. Further, the ratio of butterfly generalist to specialist species shifted towards generalists and the size of butterflies increased with higher herbivore livestock density, indicating a shift in communities towards mobile habitat generalists. Our results are in accordance with previous studies carried out across smaller geographic extents, highlighting the importance of low herbivore livestock densities to halt the loss of pollinating insects and safeguard biodiversity and associated ecosystem services in agricultural landscapes. We here demonstrate that indicators based on livestock distribution data at the municipality scale can provide insights into processes and spatial diversity patterns of butterflies at the national level. Further, we highlight potentials and limitations of using agricultural census data to quantify and assess effects of land cover and land-use intensity on butterflies, and make recommendations for further research needs.
