Cheetahs of the deep sea: deep foraging sprints in short-finned pilot whales off Tenerife (Canary Islands)

1. Empirical testing of optimal foraging models for breath-hold divers has been difficult. Here we report data from sound and movement recording DTags placed on 23 short-finned pilot whales off Tenerife to study the foraging strategies used to catch deep-water prey. 2. Day and night foraging dives had a maximum depth and duration of 1018 m and 21 min. Vocal behaviour during dives was consistent with biosonar-based foraging, with long series of echolocation clicks interspersed with buzzes. Similar buzzes have been associated with prey capture attempts in other echolocating species. 3. Foraging dives seemed to adapt to circadian rhythms. Deep dives during the day were deeper, but contained fewer buzzes (median 1), than night-time deep dives (median 5 buzzes). 4. In most deep (540-1019 m) daytime dives with buzzes, a downward directed sprint reaching up to 9 m s(-1) occurred just prior to a buzz and coincided with the deepest point in the dive, suggestive of a chase after escaping prey. 5. A large percentage (10-36%) of the drag-related locomotion cost of these dives (15 min long) is spent in sprinting (19-79 s). This energetic foraging tactic focused on a single or few prey items has not been observed previously in deep-diving mammals but resembles the high-risk/high-gain strategy of some terrestrial hunters such as cheetahs. 6. Deep sprints contrast with the expectation that deep-diving mammals will swim at moderate speeds optimized to reduce oxygen consumption and maximize foraging time at depth. Pilot whales may have developed this tactic to target a deep-water niche formed by large/calorific/fast moving prey such as giant squid.     

Different modes of acoustic communication in deep‐diving short‐finned pilot whales (Globicephala macrorhynchus)

Toothed whales use a pneumatic sound generator to produce echolocation and communication sounds. Increasing hydrostatic pressure at depth influences the amplitude and duration of calls but not of echolocation clicks. Here we test the hypothesis that information transfer at depth might be facilitated by click‐based communication signals. Wild short‐finned pilot whales (27) instrumented with multisensor DTAGs produced four main types of communication signals: low‐ and medium‐frequency calls (median fundamental frequency: 1.7 and 2.9 kHz), two‐component calls (median frequency of the low and high frequency components: 2 and 9 kHz), and rasps (burst‐pulses with median interclick interval of 21 ms). Rasps can be confused with foraging buzzes, but rasps are shorter and slower, and are not associated with fast changes in body acceleration nor reduced acoustic output of buzzes, characteristic of prey capture attempts. Contrary to calls, the energy flux density of rasps was not significantly affected by depth. This, and a different information content, may explain the observed increase in the relative occurrence of rasps with respect to calls at depth, and supports the hypothesis that click‐based communication signals may facilitate communication under high hydrostatic pressure. However, calls are produced at depth also, indicating that they may carry additional information relevant for deep‐diving animals, including potential communication among whales diving at the same time in this highly social deep‐diving species.     

Shallow food for deep divers: Dynamic foraging behavior of male sperm whales in a high latitude habitat

Groups of female and immature sperm whales live at low latitudes and show a stereotypical diving and foraging behavior with dives lasting about 45 min to depths of between 400 and 1200 m. In comparison, physically mature male sperm whales migrate to high latitudes where little is known about their foraging behavior and ecology. Here we use acoustic recording tags to study the diving and acoustic behavior of male sperm whales foraging off northern Norway. Sixty-five hours of tag data provide detailed information about the movements and sound repertoire of four male sperm whales performing 83 dives lasting between 6 and 60 min. Dives ranged in depth between 14 and 1860 m, with a median depth of 175 m, and 92% of the surfacings lasted less than 15 min. The four whales clicked for an average 91% (SD = 10) of the dive duration, where the first usual click was produced at depths ranging between 4 and 218 m and the last usual click at depths ranging between 1 and 1114 m. Echolocation buzzes, which are used as an indication of prey capture attempts, were emitted at depths between 17 and 1860 m, during both the descent and ascent phase of deep dives. The foraging behavior varied markedly with depth, with the timing and duration of prey capture attempts during shallow dives suggesting that the whales target more sparsely distributed prey. In contrast, deep dives involve frequent prey capture attempts and seem to target more dense food layers. The evidence of exploitation of different food layers, including epipelagic prey, is consistent with the hypothesis that male sperm whales may migrate to high latitudes to access a productive, multi-layered foraging habitat.     

Deep-diving beaked whales dive together but forage apart

Echolocating animals that forage in social groups can potentially benefit from eavesdropping on other group members, cooperative foraging or social defence, but may also face problems of acoustic interference and intra-group competition for prey. Here, we investigate these potential trade-offs of sociality for extreme deep-diving Blainville′s and Cuvier''s beaked whales. These species perform highly synchronous group dives as a presumed predator-avoidance behaviour, but the benefits and costs of this on foraging have not been investigated. We show that group members could hear their companions for a median of at least 91% of the vocal foraging phase of their dives. This enables whales to coordinate their mean travel direction despite differing individual headings as they pursue prey on a minute-by-minute basis. While beaked whales coordinate their echolocation-based foraging periods tightly, individual click and buzz rates are both independent of the number of whales in the group. Thus, their foraging performance is not affected by intra-group competition or interference from group members, and they do not seem to capitalize directly on eavesdropping on the echoes produced by the echolocation clicks of their companions. We conclude that the close diving and vocal synchronization of beaked whale groups that quantitatively reduces predation risk has little impact on foraging performance.     

Echolocation in Blainville’s beaked whales (Mesoplodon densirostris)

Here we use sound and movement recording tags to study how deep-diving Blainville’s beaked whales (Mesoplodon densirostris) use echolocation to forage in their natural mesopelagic habitat. These whales ensonify thousands of organisms per dive but select only about 25 prey for capture. They negotiate their cluttered environment by radiating sound in a narrow 20° field of view which they sample with 1.5–3 clicks per metre travelled requiring only some 60 clicks to locate, select and approach each prey. Sampling rates do not appear to be defined by the range to individual targets, but rather by the movement of the predator. Whales sample faster when they encounter patches of prey allowing them to search new water volumes while turning rapidly to stay within a patch. This implies that the Griffin search–approach–capture model of biosonar foraging must be expanded to account for sampling behaviours adapted to the overall prey distribution. Beaked whales can classify prey at more than 15 m range adopting stereotyped motor patterns when approaching some prey. This long detection range relative to swimming speed facilitates a deliberate mode of sensory-motor operation in which prey and capture tactics can be selected to optimize energy returns during long breath-hold dives.     

Investigation of foraging habits and prey selection by humpback whales (Megaptera novaeangliae) using acoustic tags and concurrent fish surveys

Tags containing acoustic time-depth transmitters (ATDT) were attached to four humpback whales near Kodiak, Alaska. Tags allowed for whale dive depths to be recorded in real time. Acoustic and mid-water trawl surveys were conducted concurrent with tagging efforts within the study area to quantify available fish resources and describe potential prey selection by humpback whales. Recorded dives were grouped through visual assessment and t -tests. Dives that indicated likely foraging occurred at a mean maximum depth of 106.2 m with 62% of dives occurring between 92 m and 120 m. Acoustic backscatter from fish surveys was attributed to potential humpback prey based on known target strength values and 10 net tows. Capelin comprised 84% of the total potential prey abundance in the region followed by age 0 (12%) and juvenile pollock (2%), and eulachon (<1%). Although horizontally segregated in the region, both capelin and age 0 pollock were distributed at depths exceeding 92 m with maximum abundance between 107 m and 120 m. The four-tagged humpbacks were found to forage in areas with greatest capelin densities but bypassed areas of high age 0 pollock abundance. The location and diving behavior of tagged whales suggested that whales were favoring capelin over pollock as a prey source.     

Blue whales respond to simulated mid-frequency military sonar

Mid-frequency military (1–10 kHz) sonars have been associated with lethal mass strandings of deep-diving toothed whales, but the effects on endangered baleen whale species are virtually unknown. Here, we used controlled exposure experiments with simulated military sonar and other mid-frequency sounds to measure behavioural responses of tagged blue whales (Balaenoptera musculus) in feeding areas within the Southern California Bight. Despite using source levels orders of magnitude below some operational military systems, our results demonstrate that mid-frequency sound can significantly affect blue whale behaviour, especially during deep feeding modes. When a response occurred, behavioural changes varied widely from cessation of deep feeding to increased swimming speed and directed travel away from the sound source. The variability of these behavioural responses was largely influenced by a complex interaction of behavioural state, the type of mid-frequency sound and received sound level. Sonar-induced disruption of feeding and displacement from high-quality prey patches could have significant and previously undocumented impacts on baleen whale foraging ecology, individual fitness and population health.     

Food acquisition by competing surfperch on a patchy environmental gradient

Synopsis Black surfperch, Embiotoca jacksoni, and striped surfperch, Embiotoca lateralis, coexisted along steep sloping rocky habitats at Santa Cruz Island, California. The range of depths occupied (to 15 m) was characterized by a strong gradient in abundance of prey and a changing mosaic of substrate types from which surfperch harvested food. Availability of prey and diversity of benthic substrates were greatest in shallowest areas and both declined with increasing depth. Individuals of both surfperch species were residential within a narrow range of depths, with the result that different segments of their populations were consistently exposed to different foraging environments. These two phenomena (residential behavior combined with a gradient in availability of resources) resulted in variation in foraging behaviors and diets among individuals that resided at different depths. The pattern of within-population variation differed between the surfperch species. Black surfperch individuals achieved similar taxonomic diets and expended similar foraging effort at all depths, but deep-water foragers captured much less prey biomass per unit effort. The taxonomic composition of striped surfperch diets differed among depths, and although similar amounts of prey biomass were captured everywhere, individuals in deep areas expended much greater effort to obtain that level of food return. For both species, habitat profitability (food return to foraging effort) declined with depth. The difference in habitat profitability appeared to influence fitness components of both surfperches. Individuals occupying deep habitats were about 5% shorter in standard length than conspecifics of the same chronological age living in shallow areas; the disparity in body size resulted in an estimated difference in clutch size of 10–18%.     

Food-related bray calls in wild bottlenose dolphins (Tursiops truncatus)

Because cetaceans are difficult to study in the wild, little is known about how they use their sounds in their natural environment. Only the recent development of passive acoustic localization systems has enabled observations of the communication behaviour of individuals for correlation with their surface behaviour. Using such a system, I show that bottlenose dolphins in the Moray Firth, Scotland, produce low-frequency bray calls which are clearly correlated with feeding on salmonids. The production of these calls is followed by fast approaches by conspecifics in the area. In animals which use sound as a foraging tool, it is difficult to distinguish between food calls which have evolved because of their role in attracting conspecifics, and food manipulation or searching calls which may attract conspecifics as a by-product. However, the low-frequency structure of the bottlenose dolphin bray suggests that it evolved because of a role in manipulating prey rather than in attracting conspecifics. This conclusion suggests that dolphins exploit the perceptual systems of their prey to facilitate capture.     

Following a foraging fish-finder: diel habitat use of Blainville's beaked whales revealed by echolocation

Simultaneous high resolution sampling of predator behavior and habitat characteristics is often difficult to achieve despite its importance in understanding the foraging decisions and habitat use of predators. Here we tap into the biosonar system of Blainville''s beaked whales, Mesoplodon densirostris, using sound and orientation recording tags to uncover prey-finding cues available to echolocating predators in the deep-sea. Echolocation sounds indicate where whales search and encounter prey, as well as the altitude of whales above the sea-floor and the density of organisms around them, providing a link between foraging activity and the bio-physical environment. Tagged whales (n = 9) hunted exclusively at depth, investing most of their search time either in the lower part of the deep scattering layer (DSL) or near the sea-floor with little diel change. At least 43% (420/974) of recorded prey-capture attempts were performed within the benthic boundary layer despite a wide range of dive depths, and many dives included both meso- and bentho-pelagic foraging. Blainville''s beaked whales only initiate searching when already deep in the descent and encounter prey suitable for capture within 2 min of the start of echolocation, suggesting that these whales are accessing prey in reliable vertical strata. Moreover, these prey resources are sufficiently dense to feed the animals in what is effectively four hours of hunting per day enabling a strategy in which long dives to exploit numerous deep-prey with low nutritional value require protracted recovery periods (average 1.5 h) between dives. This apparent searching efficiency maybe aided by inhabiting steep undersea slopes with access to both the DSL and the sea-floor over small spatial scales. Aggregations of prey in these biotopes are located using biosonar-derived landmarks and represent stable and abundant resources for Blainville''s beaked whales in the otherwise food-limited deep-ocean.     

FORAGING DEPTHS OF SEA OTTERS AND IMPLICATIONS TO COASTAL MARINE COMMUNITIES

We visually observed 1,251 dives, of 14 sea otters instrumented with TDRs in southeast Alaska, and used attribute values from observed dives to classify 180,848 recorded dives as foraging (0.64), or traveling (0.36). Foraging dives were significantly deeper, with longer durations, bottom times, and postdive surface intervals, and greater descent and ascent rates, compared to traveling dives. Most foraging occurred in depths between 2 and 30 m (0.84), although 0.16 of all foraging was between 30 and 100 m. Nine animals, including all five males, demonstrated bimodal patterns in foraging depths, with peaks between 5 and 15 m and 30 and 60 m, whereas five of nine females foraged at an average depth of 10 m. Mean shallow foraging depth was 8 m, and mean deep foraging depth was 44 m. Maximum foraging depths averaged 61 m (54 and 82 for females and males, respectively) and ranged from 35 to 100 m. Female sea otters dove to depths ≤20 m on 0.85 of their foraging dives while male sea otters dove to depths ≥45 m on 0.50 of their foraging dives. Less than 0.02 of all foraging dives were >55 m, suggesting that effects of sea otter foraging on nearshore marine communities should diminish at greater depths. However, recolonization of vacant habitat by high densities of adult male sea otters may result in initial reductions of some prey species at depths >55 m.     

The diving behavior of blue and fin whales: is dive duration shorter than expected based on oxygen stores?

Many diving seabirds and marine mammals have been found to regularly exceed their theoretical aerobic dive limit (TADL). No animals have been found to dive for durations that are consistently shorter than their TADL. We attached time-depth recorders to 7 blue whales and 15 fin whales (family Balaenopteridae). The diving behavior of both species was similar, and we distinguished between foraging and traveling dives. Foraging dives in both species were deeper, longer in duration and distinguished by a series of vertical excursions where lunge feeding presumably occurred. Foraging blue whales lunged 2.4 (+/-1.13) times per dive, with a maximum of six times and average vertical excursion of 30.2 (+/-10.04) m. Foraging fin whales lunged 1.7 (+/-0.88) times per dive, with a maximum of eight times and average vertical excursion of 21.2 (+/-4.35) m. The maximum rate of ascent of lunges was higher than the maximum rate of descent in both species, indicating that feeding lunges occurred on ascent. Foraging dives were deeper and longer than non-feeding dives in both species. On average, blue whales dived to 140.0 (+/-46.01) m and 7.8 (+/-1.89) min when foraging, and 67.6 (+/-51.46) m and 4.9 (+/-2.53) min when not foraging. Fin whales dived to 97.9 (+/-32.59) m and 6.3 (+/-1.53) min when foraging and to 59.3 (+/-29.67) m and 4.2 (+/-1.67) min when not foraging. The longest dives recorded for both species, 14.7 min for blue whales and 16.9 min for fin whales, were considerably shorter than the TADL of 31.2 and 28.6 min, respectively. An allometric comparison of seven families diving to an average depth of 80-150 m showed a significant relationship between body mass and dive duration once Balaenopteridae whales, with a mean dive duration of 6.8 min, were excluded from the analysis. Thus, the short dive durations of blue whales and fin whales cannot be explained by the shallow distribution of their prey. We propose instead that short duration diving in large whales results from either: (1) dispersal behavior of prey; or (2) a high energetic cost of foraging.     

Behavioral context of echolocation and prey-handling sounds produced by killer whales (Orcinus orca) during pursuit and capture of Pacific salmon (Oncorhynchus spp.)

Availability of preferred salmonid prey and a sufficiently quiet acoustic environment in which to forage are critical to the survival of resident killer whales (Orcinus orca) in the northeastern Pacific. Although piscivorous killer whales rely on echolocation to locate and track prey, the relationship between echolocation, movement, and prey capture during foraging by wild individuals is poorly understood. We used acoustic biologging tags to relate echolocation behavior to prey pursuit and capture during successful feeding dives by fish-eating killer whales in coastal British Columbia, Canada. The significantly higher incidence and rate of echolocation prior to fish captures compared to afterward confirms its importance in prey detection and tracking. Extremely rapid click sequences (buzzes) were produced before or concurrent with captures of salmon at depths typically exceeding 50 m, and were likely used by killer whales for close-range prey targeting, as in other odontocetes. Distinctive crunching and tearing sounds indicative of prey-handling behavior occurred at relatively shallow depths following fish captures, matching concurrent observations that whales surfaced with fish prior to consumption and often shared prey. Buzzes and prey-handling sounds are potentially useful acoustic signals for estimating foraging efficiency and determining if resident killer whales are meeting their energetic requirements.     

Individual right whales call louder in increased environmental noise

The ability to modify vocalizations to compensate for environmental noise is critical for successful communication in a dynamic acoustic environment. Many marine species rely on sound for vital life functions including communication, navigation and feeding. The impacts of significant increases in ocean noise levels from human activities are a current area of concern for the conservation of marine mammals. Here, we document changes in calling behaviour by individual endangered North Atlantic right whales (Eubalaena glacialis) in increased background noise. Right whales, like several bird and primate species, respond to periods of increased noise by increasing the amplitude of their calls. This behaviour may help maintain the communication range with conspecifics during periods of increased noise. These call modifications have implications for conservation efforts for right whales, affecting both the way whales use sound to communicate and our ability to detect them with passive acoustic monitoring systems.     

Geographic distribution of the Cross Seamount beaked whale based on acoustic detections

Beaked whales produce frequency-modulated echolocation pulses that appear to be species-specific, allowing passive acoustic monitoring to play a role in understanding spatio-temporal patterns. The Cross Seamount beaked whale is known only from its unique echolocation signal (BWC) with no confirmed species identification. This beaked whale spans the Pacific Ocean from the Mariana Archipelago to Baja California, Mexico, south to the equator, but only as far north as latitude 29°N. Within these warm waters, 92% of BWC detections occurred at night, 6% during crepuscular periods, and only 2% during daylight hours. Detections of BWC signals on drifting recorders with a vertical hydrophone array at 150 m depth demonstrated that foraging often occurred shallow in the water column (<150 m). No other species of beaked whale to date has been documented foraging in waters this shallow. Given their nocturnal, shallow foraging dives, this species appears to prefer prey that may be available in the water column only during those hours. The foraging behavior of Cross Seamount beaked whales appears to be unique among all beaked whales, and these findings contribute additional ecological and acoustic information which can help guide future efforts to identify this cryptic whale.     

Using dive behavior and active acoustics to assess prey use and partitioning by fin and humpback whales near Kodiak Island,Alaska

Near the Kodiak Archipelago, fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales frequently overlap spatially and temporally. The Gulf Apex Predator‐prey study (GAP) investigated the prey use and potential prey partitioning between these sympatric species by combining concurrent analysis of vertical whale distribution with acoustic assessment of pelagic prey. Acoustic backscatter was classified as consistent with either fish or zooplankton. Whale dive depths were determined through suction cup tags. Tagged humpback whales (n = 10) were most often associated with distribution of fish, except when zooplankton density was very high. Associations between the dive depths of tagged fin whales (n = 4) and the vertical distribution of either prey type were less conclusive. However, prey assessment methods did not adequately describe the distribution of copepods, a potentially significant resource for fin whales. Mean dive parameters showed no significant difference between species when compared across all surveys. However, fin whales spent a greater proportion of dive time in the foraging phase than humpbacks, suggesting a possible difference in foraging efficiency between the two. These results suggest that humpback and fin whales may target different prey, with the greatest potential for diet overlap occurring when the density of zooplankton is very high.     

The development of an intermediate‐duration tag to characterize the diving behavior of large whales

The development of high‐resolution archival tag technologies has revolutionized our understanding of diving behavior in marine taxa such as sharks, turtles, and seals during their wide‐ranging movements. However, similar applications for large whales have lagged behind due to the difficulty of keeping tags on the animals for extended periods of time. Here, we present a novel configuration of a transdermally attached biologging device called the Advanced Dive Behavior (ADB) tag. The ADB tag contains sensors that record hydrostatic pressure, three‐axis accelerometers, magnetometers, water temperature, and light level, all sampled at 1 Hz. The ADB tag also collects Fastloc GPS locations and can send dive summary data through Service Argos, while staying attached to a whale for typical periods of 3–7 weeks before releasing for recovery and subsequent data download. ADB tags were deployed on sperm whales (Physeter macrocephalus; N = 46), blue whales (Balaenoptera musculus; N = 8), and fin whales (B. physalus; N = 5) from 2007 to 2015, resulting in attachment durations from 0 to 49.6 days, and recording 31 to 2,539 GPS locations and 27 to 2,918 dives per deployment. Archived dive profiles matched well with published dive shapes of each species from short‐term records. For blue and fin whales, feeding lunges were detected using peaks in accelerometer data and matched corresponding vertical excursions in the depth record. In sperm whales, rapid orientation changes in the accelerometer data, often during the bottom phase of dives, were likely related to prey pursuit, representing a relative measure of foraging effort. Sperm whales were documented repeatedly diving to, and likely foraging along, the seafloor. Data from the temperature sensor described the vertical structure of the water column in all three species, extending from the surface to depths >1,600 m. In addition to providing information needed to construct multiweek time budgets, the ADB tag is well suited to studying the effects of anthropogenic sound on whales by allowing for pre‐ and post‐exposure monitoring of the whale's dive behavior. This tag begins to bridge the gap between existing long‐duration but low‐data throughput tags, and short‐duration, high‐resolution data loggers.     

Threshold foraging by gray whales in response to fine scale variations in mysid density

The gray whale (Eschrichtius robustus) is a coastal species whose nearshore summer foraging grounds off the coast of British Columbia offer an opportunity to study the fine scale foraging response of baleen whales. We explore the relationship between prey density and gray whale foraging starting with regional scale (10 km) assessments of whale density (per square kilometer) and foraging effort as a response to regional mysid density (per cubic meter), between 2006 and 2007. In addition we measure prey density at a local scale (100 m), while following foraging whales during focal surveys. We found regional mysid density had a significant positive relationship with both gray whale density and foraging effort. We identify a threshold response to regional mysid density for both whale density and foraging effort. In 2008 the lowest average local prey density measured beside a foraging whale was 2,300 mysids/m³. This level was maintained even when regional prey density was found to be substantially lower. Similar to other baleen whales, the foraging behavior of gray whales suggests a threshold response to prey density and a complex appreciation of prey availability across fine scales.     

No shallow talk: Cryptic strategy in the vocal communication of Blainville's beaked whales

Communicating animals must balance fitness benefits against the costs of signaling, such as increased predation risk. Cetaceans communicate mainly with sound and near‐surface vocalizations can place signalers at risk from shallow‐diving top‐predators with acute hearing such as killer whales. Beaked whales are deep divers living in small cohesive groups with little social defense from predation. Little if anything is known about their acoustic communication. Here, eight Blainville's beaked whales were studied with suction cup attached DTags to provide the first report on social communication as a function of diving behavior for any of the 21 ziphiid species. Tagged whales produced two previously unrecorded signals with apparent communicative functions: (1) fast series of ultrasonic frequency modulated clicks (rasps) were recorded from six individuals, and (2) harmonically rich short whistles with a mean fundamental frequency of 12 kHz were recorded from one whale at up to 900 m depth, the deepest whistles recorded from a marine mammal. Blainville's were silent 80% of the time, whenever shallower than 170 m depth and during the prolonged (19 min) silent ascents from vocal dives. This behavior limits the ability of shallow‐diving predators to track Blainville's acoustically and may provide a striking example of the evolutionary influence of the risk of predation on animal communication.     

Social Calls Provide Tree‐dwelling Bats with Information about the Location of Conspecifics at Roosts

Animals can use signals emitted by other animals as sources of information. Auditory signals are important in communication networks, as they can potentially convey information about the location and state of conspecifics and other species over long distances. Signalling is important in fission–fusion societies, in which animals from the same social group temporarily split into subgroups and frequently change roost sites. We used playbacks of social calls of the noctule Nyctalus noctula produced in roosts, to show how bats might maintain group cohesion and to test the hypothesis that noctules can locate conspecifics when returning from foraging trips by eavesdropping on or communicating with roosting individuals. Noctules responded strongly to broadcasted social calls. Their reactions included inspections and landing on a loudspeaker broadcasting social calls and occasional social vocalisation. Responses by other bat species to the noctule social calls were negligible. The high amplitude, low‐frequency vocalisations emitted by noctules in roosts can propagate over long distances and allow group members to announce their position. Bats can extract information about the location of roosts containing conspecifics by eavesdropping or by communication. Social calls may thus be sufficient to locate conspecifics in roosts and maintain spatial associations of groups in mammals.