Adaptation to urban environments.

Despite widespread evidence of urban evolution, the adaptive nature of these changes is often unclear. We review different phenotypic and molecular lines of evidence used for assessing urban adaptation, discussing the benefits and limitations of each approach, and rare examples of their integration. We then provide a synthesis of local adaptation to urban and rural environments. These data were drawn from phenotypic reciprocal transplant studies, the majority of which focus on insects and other arthropods. Broadly, we found support for local adaptation to urban and rural environments. However, there was asymmetry in the evidence for local adaptation depending on population of origin, with urban adaptation being less prevalent than rural adaptation, suggesting many urban populations are still adapting to urban environments. Further, the general patterns were underlain by considerable variation among study systems; we discuss how environmental heterogeneity and costs of adaptation might explain system-specific variation in urban-rural local adaptation. We then look to the future of urban adaptation research, considering the magnitude and direction of adaptation in context of different agents of selection including urban heat islands, chemical pollutants, and biotic interactions.