Perch time allocation and feeding efficiency of flycatching Rhinolophus formosae: an optimal foraging behavior?

Background: Flycatching bats are species-rare and comprise predominantly horseshoe bats (Rhinolophidae). Their hang-and-wait foraging mode and long constant-frequency echolocation calls offer advantages in energetics and prey detection, and may enable them apt to foraging optimally, yet not much is known about the foraging behavior of flycatching bats. Thus we assessed the perch use and foraging performance in the field by one of the largest horseshoe bats, Rhinolophus formosae, and offered insights on their perch time allocation. Results: The perching-foraging behaviors of the bats did not differ significantly between forest settings, but the residence and giving-up time, mean attack, and attack rate were higher in the late spring-early summer, whereas the mean capture, capture rate, and attack efficiency were lower in the late summer when volant juveniles joined the nocturnal activity. The bats maintained flycatching and exhibited largely similar attack rates through the night with peak residence time around the midnight, but the capture rate and attack efficiency both reduced toward midnight and then increased toward the hours right before dawn. The attack rate was negatively correlated to the number of perches used and perch switch; by contrast, the capture rate was positively correlated with both factors. The total residence time at a site increased but mean residence time per perch decreased as the number of perches used and perch-switch increased. The giving-up time was inversely correlated to the attack rate and attack efficiency, and decreased with an increasing capture rate. Conclusions: The bats increased perch switch at lower attack rates in early spring, but switched less frequently in late spring and prime summer months when insect abundance is higher. By scanning through a broad angular range for prey detection, and switching more frequently among perches, R. formosae foraged with an increased capture rate, and were able to remain at the site longer by slightly reducing their mean residence time per perch. Our results concur with the predictions of optimal foraging theory for patch selection and offer implications for further exploration of the foraging behavior of flycatching horseshoe bats.