What is an urban bird? Trait-based predictions of urban tolerance for the world's birds are modulated by latitude and human population density

As human density increases, biodiversity must increasingly co-exist with urbanization or face local extinction. Tolerance of urban areas has been linked to numerous functional traits, yet few globally-consistent patterns have emerged to explain variation in urban tolerance, which stymies attempts at a generalizable predictive framework. Here, we calculate an Urban Association Index (UAI) for 3768 bird species in 137 cities across all permanently inhabited continents. We then assess how UAI varies as a function of ten species-specific traits and further test whether the strength of trait relationships vary as a function of three city-specific variables. Of the ten species traits, nine were significantly associated with urban tolerance. Urban-associated species tend to be smaller, less territorial, have greater dispersal ability, broader dietary and habitat niches, larger clutch sizes, greater longevity, and have lower elevational limits. Only bill shape showed no global association with urban tolerance. Additionally, the strength of several trait relationships varied across cities as a function of latitude and/or human population density. For example, the effects of body mass and diet breadth are more pronounced at higher latitudes, while the effects of territoriality and longevity were reduced in cities with higher population density. Thus, the importance of trait filters in birds varies predictably across cities, indicating biogeographic variation in selection for urban tolerance that could explain prior challenges in the search for global patterns. A globally-informed framework that predicts urban tolerance will be integral to conservation as increasing proportions of the worlds biodiversity are impacted by urbanization.