TL;DR: This article describes the echolocation behavior of insect-eating bats and shows how differing circumstances such as habitat type, foraging mode, and diet favor different signal types, and outlines the perceptual tasks that must be performed by foraging bats.
Abstract: B (order Chiroptera) are ecologically more diverse than any other group of mammals. Numerous morphological, physiological, and behavioral adaptations of sensory and motor systems permit bats access to a wide range of habitats and resources at night. The more than 750 species of the suborder Microchiroptera occupy most terrestrial habitats and climatic zones and exploit a great variety of foods, ranging from insects and other arthropods, small vertebrates, and blood to fruit, leaves, nectar, flowers, and pollen. Echolocation is one of the adaptations that make bats so successful. Echolocating animals emit signals of high frequency (mostly ultrasonic) and analyze the returning echoes to detect, characterize, and localize the reflected objects. Sophisticated echolocation systems have evolved only in the bat suborder Microchiroptera and in dolphins. Less efficient systems have been reported for a few species of the bat suborder Megachiroptera and for some birds (Henson and Schnitzler 1980). Bats use echolocation for orientation in space, that is, for determining their position relative to the echo-producing environment. In addition, many bats, especially those that hunt for flying insects, use echolocation to detect, identify, and localize prey. Bats use a wide variety of species-specific signal types differing in frequency structure, duration, and sound pressure level (SPL). In addition, signal structure varies depending on the echolocation task confronting the bat. Search signals that are emitted when bats search for prey differ from approach signals that are emitted when they approach prey. The echolocation signals and hearing systems of bats are well adapted for gathering behaviorally relevant information (e.g., Schnitzler and Henson 1980, Neuweiler 1989, Fenton 1990, Denzinger et al. forthcoming). In this article we describe the echolocation behavior of insect-eating bats and show how differing circumstances such as habitat type, foraging mode, and diet favor different signal types. To demonstrate relationships between echolocation and ecological conditions, we outline the perceptual tasks that must be performed by foraging bats and discuss the suitability of typical elements of echolocation signals for solving such problems. We then define habitat types according to the problems they impose on bats and relate the observed variability in signal structure to ecological constraints set by habitat type and foraging mode. Perceptual problems for foraging bats Foraging bats confront a multitude of problems when flying to their hunting grounds and searching for prey. These problems differ depending on where bats hunt, what they eat, and how they acquire their food. For example, bats hunting for insects in the open encounter conditions different from those that search for prey near the edges of vegetation, in vegetation gaps, in dense forest, or near the ground. The problems also differ depending upon whether they capture moving prey in flight (aerial mode) or mostly stationary prey from surfaces such as leaves or ground (gleaning mode) or water (trawling mode). Foraging bats must detect, classify, and localize an insect and discriminate between echoes of prey and echoes of unwanted targets such as twigs, foliage, or the ground, referred to as clutter echoes, or simply “clutter.”For many bats echolocation delivers all of the information they need to catch an insect.
TL;DR: This conceptual framework calls for a new view on field data from bats orienting and foraging in different types of habitats according to the ecological constraints in which foraging bats operate, four distinct functional groups or guilds within each group, signal design and echolocation behavior are rather similar.
Abstract: Field research on echolocation behavior in bats has emphasized studies of food acquisition, and the adaptive value of sonar signal design as been considered largely in the context of foraging. However, echolocation tasks related to spatial orientation also differ among bats and are relevant to understanding signal structure. Here, we argue that the evolution of echolocation in bats is characterized by two key innovations: first, the evolution of echolocation for spatial orientation and, second, a later transition for prey acquisition. This conceptual framework calls for a new view on field data from bats orienting and foraging in different types of habitats. According to the ecological constraints in which foraging bats operate, four distinct functional groups or guilds can be defined. Within each group, signal design and echolocation behavior are rather similar.
TL;DR: Echolocating bats use different information-gathering strategies for hunting prey in open, uncluttered environments, in relatively open environments with some obstacles, and in densely cluttered environments.
Abstract: Echolocating bats use different information-gathering strategies for hunting prey in open, uncluttered environments, in relatively open environments with some obstacles, and in densely cluttered environments. These situations differ in the extent to which individual targets such as flying insects can be detected as isolated objects or must be separated perceptually from backgrounds. Echolocating bats also differ in whether they use high-resolution, multidimensional images of targets or concentrate specifically on one particular target dimension, such as movement, to detect prey.
TL;DR: An overlap-free “window” within which pipistrelles may detect potential prey and which allows predictions of minimum distances to prey and clutter-producing objects is proposed.
Abstract: We studied the echolocation and hunting behavior of three aerial insectivorous species of bats (Vespertilionidae: Pipistrellus) in the field in order to characterize the signals used by the bats and to determine how call structure varies in relation to habitat structure (“uncluttered” versus “cluttered” space). We documented free-flying, naturally foraging wild pipistrelles in various habitats using multiflash stereophotography combined with simultaneous sound recordings. Then we reconstructed the bat's flight position in three-dimensional space and correlated it with the corresponding echolocation sequences. In all three species of pipistrelles, signal structure varied substantially. In echolocation sequences of the search phase we found a consistent association of signal types with habitat types. In uncluttered habitats (obstacles more than 5 m from the bat) pipistrelles emitted almost exclusively narrowband signals with bandwidths less than 15 kHz. In cluttered habitats (obstacles less than 5 m from the bat) they switched to signals with bandwidths of more than 15 kHz. Wideband signals were also used when the bats were turning in cluttered and uncluttered spaces and for an instant after turning away from obstacles. Prey detection occured only when the outgoing signal did not overlap with the returning echo from potential prey. The bats also avoided overlap of echoes from potential prey and obstacles. Based on the results of this study, we propose an overlap-free “window” within which pipistrelles may detect potential prey and which allows predictions of minimum distances to prey and clutter-producing objects.
TL;DR: Measurements confirm theoretical expectations and earlier observations that atmospheric attenuation is progressively more severe at higher frequencies and that the atmosphere acts as a low-pass filter for conducting sounds in the frequency range used for echolocation by bats.
Abstract: The absorption of sound propagating through the atmosphere under laboratory conditions of 25 degrees C and 50% relative humidity was measured at frequencies from 30 to 200 kHz. The attenuating effect on the passage of ultrasonic sounds through air ranged from 0.7 dB/m at 30 kHz. These measurements confirm theoretical expectations and earlier observations that atmospheric attenuation is progressively more severe at higher frequencies and that the atmosphere acts as a low-pass filter for conducting sounds in the frequency range used for echolocation by bats. Different species of bats use different portions of this range of frequencies, and bats emitting sonar signals predominantly above 100 kHz encounter especially severe attenuation of over 3 dB/m. With the greatly restricted operating distances for echolocation at such high frequencies, bats using these higher frequencies must be under compelling ecological pressures of a higher priority than long-range detection of targets.