How do urban green space designs shape avian communities? Testing the area–heterogeneity trade-off

In cities, green areas are essential for biodiversity conservation, with land cover heterogeneity being a decisive factor. Yet, as heterogeneity increases for a given green area, the patch size of land covers automatically decreases, as the area available for individual species, especially habitat specialist species. This relationship, known as the area-heterogeneity trade-off, is expected to lead to a unimodal relationship between species richness and land cover heterogeneity, and contrasted effects between species according to their level of urban avoidance.
We investigated the potential consequences of this trade-off on birds in green areas selected along an urban intensity gradient in six European cities. Using a European database on bird occurrences in nesting habitats, we defined a continuous gradient of urban avian avoidance.
We confirmed the marked area-heterogeneity trade-off in urban green areas but found no effect of land cover heterogeneity on total avian richness at green area level. However, both land cover heterogeneity and patch size were positively associated with richness of urban avoider species, indicating that urban avoiders fared better in green areas with large and heterogeneous patches. Total richness was also higher in green areas surrounded by an urban matrix composed of a variety of land covers.
To protect urban bird avoiders, which are most at risk in cities, green area managers and urban planners should thus be aware that land cover heterogeneity is not a panacea if patch sizes are too small. To conserve avian richness, we stress the importance of maintaining large vegetated areas as well as heterogeneity in land covers within the urban matrix.