Effects of local climate and habitat heterogeneity on breeding-bird assemblages of semi-natural grasslands

The intensification of agricultural land-use, abandonment and afforestation have caused severe loss and degradation of nutrient-poor, semi-natural grasslands across Europe. Calcareous grasslands have an outstanding value for nature conservation due to their highly diverse flora and fauna, including birds. However, knowledge of environmental factors driving the composition of bird communities in calcareous grasslands is still scarce. The aim of our study was to compare the breeding-bird assemblages of calcareous grasslands and the second most common semi-natural grassland type in the study area (Diemel Valley, Central Germany), mesic grasslands, along an elevation (climate) gradient. For each grassland type, we randomly selected 27 square-shaped plots with a size of 5 ha each and a cover of the focal grassland type of at least 50% within the plot. Our study revealed that both the number and density of threatened species of breeding birds were higher in plots of calcareous grasslands than in those of mesic grasslands. Based on multivariable generalised linear mixed-effects models, the most important predictors of overall species richness and density of breeding birds in plots of calcareous grasslands were the area of juniper-rich calcareous grassland and Shannon habitat diversity. By contrast, the number and density of threatened species were in both grassland types mainly driven by breeding-season temperature. Heterogeneous, juniper-rich calcareous grasslands seem to be a key habitat for bird assemblages with high species richness and density. These grasslands usually consisted of small-scale habitat mosaics providing suitable nesting sites and foraging grounds in close proximity. By contrast, mesic grasslands had rather homogeneous swards and shrubs or trees were normally restricted to the edges of the grassland patches. Based on our study, increasing habitat heterogeneity at the landscape scale and within grasslands is the key to enhancing species richness and density of breeding birds.