Behavioral Responses

In addition to the direct effects on diversity and abundances, species' responses to anthropogenic habitat modification and disturbance can manifest as behavioral changes, which may include disruptions to species' dispersal, movement, activity patterns, and interspecific interactions (Fischer and Lindenmayer 2007). Few studies so far have addressed these issues for tropical bats (Fig. 3.3b).

Although a number of studies have reported movement distances and space use for a variety of tropical bat species (not reviewed here), few have explicitly addressed these phenomena in anthropogenically modified landscapes. Mark–recapture and radiotracking studies in the Neotropics suggest that in areas where landscape connectivity is relatively high, bats may regularly traverse open areas between forest fragments or between fragments and continuous forest. Evidence for interhabitat movements comes from landscapes with agricultural matrices (Estrada et al. 1993a; Estrada and Coates-Estrada 2002; Bianconi et al. 2006; Medina et al. 2007; Mendes et al. 2009; Trevelin et al. 2013) or from those with a more inhospitable aquatic matrix (Albrecht et al. 2007; Meyer and Kalko 2008a). Recapture data from a study in a fragmented landscape in Malaysia also indicate long-distance between-habitat movements for some cave-roosting species (Struebig et al. 2008). Whether a species is able to move over fragmented landscapes may be linked to the species' foraging ecology (Albrecht et al. 2007; Henry et al. 2007b). Overall, these studies were fundamental in determining the general capacity of tropical bats to move across human-modified habitats. However, they provide mostly circumstantial evidence and cannot establish whether anthropogenic disturbance elicits direct behavioral responses in bats that manifest as changes in movement distances or patterns of space use. Better insights into how habitat modification influences movement behavior can be gained through detailed radiotracking or long-term banding studies that compare movement patterns for species with different autecologies. Such studies, although difficult and costly to implement, would ideally compare continuous forest with fragments or other disturbed habitats.

Studies that have assessed behavioral changes to habitat modification in terms of effects on temporal activity patterns have followed such a rigorous approach. Disturbance-related changes in resource abundance, diversity, or predictability can be assumed to potentially alter temporal activity of species that exploit those resources (Presley et al. 2009b). Presley et al. (2009a) found no interspecific differences in activity patterns of eight abundant frugivorous bats in primary lowland Amazonian rain forest. However, for five species, activity patterns differed between primary or secondary forest and agricultural habitats, whereby bats in larger agricultural areas exhibited reduced crepuscular activity compared to those in undisturbed forest. Elsewhere in Amazonia, Castro-Arellano et al. (2009) detected no differences in activity levels for nectarivores and gleaning animalivores in response to RIL. Conversely, understory frugivores (Carollia spp.) decreased activity at dusk. Another study found reduced activity by some frugivores in small forest clearings created by tree removal, although the overall effects of RIL on activity patterns of frugivores were negligible (Presley et al. 2009b). In all cases, the curtailment of activity in open areas at twilight or during periods of high lunar illumination was best explained by increased predation risk (SaldañaVázquez and Munguía-Rosas 2013). Habitat modification and disturbance may consequently influence energy budgets of bats as they have less time available for foraging, with possible negative repercussions for their ability to meet daily energy requirements.

Human disturbance may also affect roosting behavior and roost site selection. In fragmented rain forest in Mexico, Evelyn and Stiles (2003) found that both sexes of cavity-roosting Sturnira lilium selected large-diameter trees in mature forest stands, as did females of the foliage-roosting Artibeus intermedius, whereas males of the latter species roosted in secondary forest. These findings underscore that preferences in terms of roosting and foraging habitat are not necessarily correlated and point to the importance of preserving mature forest patches in human-dominated landscapes for meeting the roosting requirements of tree-cavityroosting species.

Key research needs:

• More studies, particularly in the Paleotropics, that assess the extent to which human-driven habitat change affects bat behavior in terms of roosting and foraging ecology.

• Research that addresses how such behavioral changes translate into fitness con-

sequences (e.g., in terms of survival, reproductive success, physiology) that may affect long-term population persistence.

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