Area and Isolation Effects
Early fragmentation studies generally emphasized the effects of area and isolation, reflecting the pervasive influence of island biogeographic theory (IBT, MacArthur and Wilson 1967) in ecology, while ignoring influences of the surrounding landscape matrix. This same pattern is also apparent within the fragmentation literature on tropical bats. Studies have found evidence for effects of both fragment area (Cosson et al. 1999; Struebig et al. 2008, 2011) and isolation (Estrada et al. 1993a; Meyer and Kalko 2008a, b) on populationand assemblage-level responses, whereas effects were weak or absent in others (Faria 2006; Pardini et al. 2009). Moreover, bat ensembles and species often respond differentially to fragment area or isolation, with responses of some taxa being particularly strong (Struebig et al. 2008; Estrada Villegas et al. 2010).
The relative importance of isolation versus area in shaping bat responses to fragmentation is governed by three main factors: the range of fragment sizes relative to isolation in the landscape, the history of landscape change (time since isolation, rate of change), and, probably most importantly, the type and quality of matrix habitats in which fragments are embedded. For instance, the high explanatory power of area relative to isolation reported by Struebig et al. (2008) likely reflects the low structural contrast between fragments and matrix (mostly rubber and oil palm plantations), limited range of isolation distances compared to area in the study system, and a possible time lag in the realization of isolation effects due to landscape change being fairly recent. In contrast, isolation rather than island area best predicted bat species richness and composition on Neotropical land-bridge islands (Meyer and Kalko 2008a) where fragments were surrounded by water.
The simplified dichotomous view of landscapes underlying IBT, albeit applicable in special cases (e.g., land-bridge islands), often fails to capture the influence that other land-cover types in the surrounding matrix can have and so may not be broadly applicable to most anthropogenically modified landscapes (Kupfer et al. 2006; Laurance 2008). After more than 40 years of research beyond the origins of IBT, it is now clear that for most animal taxa, including tropical bats, the majority of terrestrial habitat fragments are not islands in a homogeneous sea of inhospitable habitat. Indeed, island ecosystems support tropical bat biodiversity in fundamentally different ways compared to complex agricultural mosaic landscapes, the former adhering to IBT predictions of species loss, while countryside ecosystems are capable of maintaining high levels of species richness, evenness, and compositionally novel assemblages in human-made habitats (Mendenhall et al. 2014).