A conceptual framework for the colonisation of urban areas: the blackbird Turdus merula as a case study

Despite increasing interest in urban ecology the factors limiting the colonisation of towns and cities by species from rural areas are poorly understood. This is largely due to the lack of a detailed conceptual framework for this urbanisation process, and of sufficient case studies. Here, we develop such a framework. This draws upon a wide range of ecological and evolutionary theory and the increasing number of studies of how the markedly divergent conditions in urban and rural areas influence the traits of urban populations and the structure of urban assemblages. We illustrate the importance of this framework by compiling a detailed case study of spatial and temporal variation in the urbanisation of the blackbird Turdus merula. Our framework identifies three separate stages in the urbanisation process: (i) arrival, (ii) adjustment, and (iii) spread. The rate of progress through each stage is influenced by environmental factors, especially human attitudes and socio-economic factors that determine the history of urban development and the quality of urban habitats, and by species' ecological and life-history traits. Some traits can positively influence progression through one stage, but delay progression through another. Rigorous assessment of the factors influencing urbanisation should thus ideally pay attention to the different stages. Urbanisation has some similarities to invasion of exotic species, but the two clearly differ. Invasion concerns geographic range expansion that is external to the species' original geographic range, whilst urbanisation typically relates to filling gaps within a species' original range. This process is exemplified by the blackbird which is now one of the commonest urban bird species throughout its Western Palearctic range. This is in stark contrast to the situation 150 years ago when the species was principally confined to forest. Blackbird urbanisation was first recorded in Germany in 1820, yet some European cities still lack urban blackbirds. This is especially so in the east, where urbanisation has spread more slowly than in the west. The timing of blackbird urbanisation exhibits a marked spatial pattern, with latitude and longitude explaining 76% of the variation. This strong spatial pattern contrasts with the weaker spatial pattern in timing of urbanisation exhibited by the woodpigeon Columba palumbus (with location explaining 39% of the variation), and with the very weak spatial pattern in timing of black-billed magpie Pica pica urbanisation (in which location explains 12% of the variation). Strong spatial patterns in timing of urbanisation are more compatible with the leap-frog urbanisation model, in which urban adapted or imprinted birds colonise other towns and cities, than with the independent urbanisation model, in which urban colonisation events occur independently of each other. Spatial patterns in isolation do not, however, confirm one particular model. Factors relating to the arrival and adjustment stages appear particularly likely to have influenced the timing of blackbird urbanisation. Spatial variation in the occurrence of urban populations and the timing of their establishment creates opportunities to assess the factors regulating urbanisation rates, and how the composition of urban assemblages develops as a result. These are major issues for urban ecology.