The relationship between multi-scale habitat heterogeneity and farmland bird abundance

It has recently been proposed that losses in farmland habitat heterogeneity may have been a primary driver of the profound declines exhibited by many farmland bird species in recent decades. However, it has yet to be demonstrated which facets of heterogeneity and what spatial scales are most important for birds. Here we analyse the relationship between abundance and features of landscape heterogeneity at three spatial scales (1, 9 and 25 km2) for 32 bird species commonly associated with farmland. Heterogeneity was quantified using three contrasting indices reflecting 1) the spatial mixing of land uses, 2) variation in field sizes and 3) the density of field boundaries. The spatial mixing of land-uses explained, on average, the most variation in, and was most likely to be positively associated with, abundance at all spatial scales. The majority of species (6675%, depending on the spatial scale) were more common in heterogeneous landscapes overall; however, migrants, those under a high level of conservation concern and farmland specialist species tended to be less abundant in more heterogeneous landscapes at all scales. Ground-nesting species were also more likely to be found in more homogeneous habitats than non-ground-nesters, but only at the finest spatial scale. Relationships between abundance and heterogeneity were generally consistent across spatial scales; however, species of high conservation concern had more variable associations compared with other species. These results highlight a potential role for farmland habitat heterogeneity in determining the abundance of many farmland species but suggest that population responses to an increase in heterogeneity would not be unanimously positive and would probably have negative impacts on some species, notably those that are already threatened.