Adaptability of Species to Urban Areas: General Trends, Species-Specific Differences and Future Research

Urban areas can provide suitable habitat for a variety of species, albeit an anthropogenically altered habitat (McKinney 2006). However, our general understanding of what influences a species' success in urban environments is limited. Arguably the conservation of species such as bats in urban areas is dependent upon this knowledge (Fenton 1997). Within this book chapter, we reviewed the existing literature on bats in urban areas. In addition, we combined published data in a metaanalysis to evaluate and derive general patterns in the response of bats to urban development.

Our meta-analysis revealed that, in general, habitat use of bats decreases in urban areas. A high degree of urbanisation had a stronger negative effect on overall habitat use of bats compared to an intermediate degree of urban development. However, habitat use in intermediate urban development was much lower compared with natural areas. This is alarming, as it is generally thought that small towns and suburban landscapes could potentially provide suitable habitat for a wide range of species (McKinney 2006), including bats. The combination of habitats with different complexity in smaller urban developments should lead to greater complementarity at a local scale and should favour species diversity and abundance. Some of the publications in our meta-analysis dataset indeed report a higher bat diversity, activity (Hourigan et al. 2010; Threlfall et al. 2011, 2012b) and feeding activity (Jung and Kalko 2011; Threlfall et al. 2012a) at intermediate levels of disturbance compared to natural or urban habitats. Other studies reported that any urban land cover, even if low-density residential, can decrease bat activity and species richness (Hourigan et al. 2006; Gonsalves et al. 2013; Luck et al. 2013), and even deter individual species (Jung and Kalko 2010; Gonsalves et al. 2013; Luck et al. 2013). Altogether, this strongly suggests regional differences in the intensity of urban development and points towards an interacting effect of the surrounding landscape (see Coleman and Barclay 2011).

Results from recent urban bat studies suggest that bats of some families (e.g. molossids Jung and Kalko 2011) are better pre-adapted for life in an urban environment compared to others (e.g. rhinolophids Stone et al. 2009; Threlfall et al. 2011). Our analysis also indicated a family-specific effect of urbanisation and confirmed the negative response of Rhinolophidae to urban development across the Old World. However, the responses of Molossidae and Vespertilionidae, which are known to frequently roost in man-made structures in North and South America, did not reveal consistent associations with either urban or natural areas across continents. This might be due to the high morphological and behavioural heterogeneity within these families. We believe that the likely explanation for our results is that the response to urbanisation is dictated by the behavioural and morphological traits of species, regardless of geographic region or phylogeny. In particular, species foraging in open space seem to persist in urban areas, as due to their wing morphology (high aspect ratio and wing loading) they might be able to commute large distances between roosting sites and feeding areas (Jung and Kalko 2011). Thus traits predicting species mobility have been associated with urban tolerance (Jung and Kalko 2011; Threlfall et al. 2012a), and the ability to forage around street lights (see Rowse et al., Chap. 7 this volume). In addition, traits that allow for flexible roost and foraging strategies confer an advantage for urban-tolerant species. Our current results support these findings and thus suggest that adaptability of bats to urban environments (or disturbance in general) might be correlated with, and reflected by, species behavioural flexibility. Advancement of knowledge in this area will assist with conservation efforts of bat species globally, and potentially allow development of a predictive framework for assessing the impacts of urban development on bats.

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