Adaptations for Substrate Gleaning in Bats: The Pallid Bat as a Case Study

Substrate gleaning is a foraging strategy in which bats use a mixture of echolocation, prey-generated sounds, and vision to localize and hunt surface-dwelling prey. Many substrate-gleaning species depend primarily on prey-generated noise to hunt. Use of echolocation is limited to general orientation and obstacle avoidance. This foraging strategy involves a different set of selective pressures on morphology, behavior, and auditory system organization of bats compared to the use of echolocation for both hunting and navigation. Gleaning likely evolved to hunt in cluttered environments and/or as a counterstrategy to reduce detection by eared prey. Gleaning bats simultaneously receive streams of echoes from obstacles and prey-generated noise, and have to segregate these acoustic streams to attend to one or both. Not only do these bats have to be exquisitely sensitive to the soft, low frequency sounds produced by walking/rustling prey, they also have to precisely localize these sounds. Gleaners typically use low intensity echolocation calls. Such stealth echolocation requires a nervous system that is attuned to low intensity sound processing. In addition, landing on the ground to hunt may bring gleaners in close proximity to venomous prey. In fact, at least 2 gleaning bat species are known to hunt highly venomous scorpions. While a number of studies have addressed adaptations for echolocation in bats that hunt in the air, very little is known about the morphological, behavioral, and neural specializations for gleaning in bats. This review highlights the novel insights gleaning bats provide into bat evolution, particularly auditory pathway organization and ion channel structure/function relationships. Gleaning bats are found in multiple families, suggesting convergent evolution of specializations for gleaning as a foraging strategy. However, most of this review is based on recent work on a single species - the pallid bat (Antrozous palli dus) - symptomatic of the fact that more comparative work is needed to identify the mechanisms that facilitate gleaning behavior.