Vocal foragers and silent crowds: context-dependent vocal variation in Northeast Atlantic long-finned pilot whales.
ABSTRACT: Vocalisations form a key component of the social interactions and foraging behaviour of toothed whales. We investigated changes in calling and echolocation behaviour of long-finned pilot whales between foraging and non-foraging periods, by combining acoustic recordings and diving depth data from tagged individuals with concurrent surface observations on social behaviour of their group. The pilot whales showed marked vocal variation, specific to foraging and social context. During periods of foraging, pilot whales showed more vocal activity than during non-foraging periods (rest, travel). In addition to the expected increase in echolocation activity, call rates also increased, suggesting that pilot whales communicate more during foraging. Furthermore, calls with multiple inflections occurred more often immediately before and after foraging dives and during the early descent and late ascent phases of foraging dives. However, these calls were almost never detected at diving depths of the tagged whale beyond 350 m. Calls with no or few inflections were produced at all times, irrespective of diving depth of the tagged whale. We discuss possible explanations for the distinct vocal variation associated with foraging periods. In addition, during non-foraging periods, the pilot whales were found to be more silent (no calling or echolocation) in larger, more closely spaced groups. This indicates that increased levels of social cohesion may release the need to stay in touch acoustically.Social toothed whales rely on vocalisations to find prey and interact with conspecifics. Species are often highly vocal and can have elaborate call repertoires. However, it often remains unclear how their repertoire use correlates to specific social and behavioural contexts, which is vital to understand toothed whale foraging strategies and sociality. Combining on-animal tag recordings of diving and acoustic behaviour with observations of social behaviour, we found that pilot whales produce more calls during foraging than during non-foraging periods. Moreover, highly inflected calls were closely associated to the periods around and during foraging dives. This indicates enhanced communication during foraging, which may, for example, enable relocation of conspecifics or sharing of information. Whales reduced their vocal activity (calling and echolocation) at increased levels of social cohesion, indicating that in certain behavioural contexts, closer association (i.e. more closely spaced) may release the need to stay in touch acoustically.
Project description:Diving behaviour of short-finned pilot whales is often described by two states; deep foraging and shallow, non-foraging dives. However, this simple classification system ignores much of the variation that occurs during subsurface periods. We used multi-state hidden Markov models (HMM) to characterize states of diving behaviour and the transitions between states in short-finned pilot whales. We used three parameters (number of buzzes, maximum dive depth and duration) measured in 259 dives by digital acoustic recording tags (DTAGs) deployed on 20 individual whales off Cape Hatteras, North Carolina, USA. The HMM identified a four-state model as the best descriptor of diving behaviour. The state-dependent distributions for the diving parameters showed variation between states, indicative of different diving behaviours. Transition probabilities were considerably higher for state persistence than state switching, indicating that dive types occurred in bouts. Our results indicate that subsurface behaviour in short-finned pilot whales is more complex than a simple dichotomy of deep and shallow diving states, and labelling all subsurface behaviour as deep dives or shallow dives discounts a significant amount of important variation. We discuss potential drivers of these patterns, including variation in foraging success, prey availability and selection, bathymetry, physiological constraints and socially mediated behaviour.
Project description:Echolocating toothed whales produce powerful clicks pneumatically to detect prey in the deep sea where this long-range sensory channel makes them formidable top predators. However, air supplies for sound production compress with depth following Boyle's law suggesting that deep-diving whales must use very small air volumes per echolocation click to facilitate continuous sensory flow in foraging dives. Here we test this hypothesis by analysing click-induced acoustic resonances in the nasal air sacs, recorded by biologging tags. Using 27000 clicks from 102 dives of 23 tagged pilot whales (Globicephala macrorhynchus), we show that click production requires only 50?µL of air/click at 500?m depth increasing gradually to 100?µL at 1000?m. With such small air volumes, the metabolic cost of sound production is on the order of 40?J per dive which is a negligible fraction of the field metabolic rate. Nonetheless, whales must make frequent pauses in echolocation to recycle air between nasal sacs. Thus, frugal use of air and periodic recycling of very limited air volumes enable pilot whales, and likely other toothed whales, to echolocate cheaply and almost continuously throughout foraging dives, providing them with a strong sensory advantage in diverse aquatic habitats.
Project description:During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.
Project description:In contrast to their terrestrial call, the offshore call of penguins during their foraging trips has been poorly studied due to the inaccessibility of the foraging site-the open ocean-to researchers. Here, we present the first description of the vocal behaviour of penguins in the open ocean and discuss the function of their vocal communication. We deployed an animal-borne camera on gentoo penguins (Pygoscelis papua) and recorded their foraging behaviour during chick guarding. From the video recordings, we collected 598 offshore calls from 10 individuals in two breeding seasons (2014-2015 and 2015-2016), and we analysed the acoustic characteristics and behavioural contexts of these calls, including diving patterns, group association events, and foraging behaviour. The offshore calls varied in their dominant frequency and length, and penguins produced calls of different lengths in succession. Group associations were observed within one minute following an offshore call in almost half of the instances (43.18%). Penguins undertook dives of shallower depths and shorter durations after producing an offshore call than those before producing an offshore call. Our findings show that penguins may use vocal communication in the ocean related with group association during foraging trips.
Project description:Cuvier's beaked whales exhibit exceptionally long and deep foraging dives. The species is little studied due to their deep-water, offshore distribution and limited time spent at the surface. We used LIMPET satellite tags to study the diving behaviour of Cuvier's beaked whales off Cape Hatteras, North Carolina from 2014 to 2016. We deployed 11 tags, recording 3242 h of behaviour data, encompassing 5926 dives. Dive types were highly bimodal; deep dives (greater than 800 m, n = 1408) had a median depth of 1456 m and median duration of 58.9 min; shallow dives (50-800 m, n = 4518) were to median depths of 280 m with a median duration of 18.7 min. Most surface intervals were very short (median 2.2 min), but all animals occasionally performed extended surface intervals. We found no diel differences in dive depth or the percentage of time spent deep diving, but whales spent significantly more time near the surface at night. Other populations of this species exhibit similar dive patterns, but with regional differences in depth, duration and inter-dive intervals. Satellite-linked tags allow for the collection of long periods of dive records, including the occurrence of anomalous behaviours, bringing new insights into the lives of these deep divers.
Project description:Air-breathing marine predators that target sub-surface prey have to balance the energetic benefit of foraging against the time, energetic and physiological costs of diving. Here we use on-animal data loggers to assess whether such trade-offs can be revealed by the breathing rates (BR) and timing of breaths in long-finned pilot whales (Globicephela melas). We used the period immediately following foraging dives in particular, for which respiratory behavior can be expected to be optimized for gas exchange. Breath times and fluke strokes were detected using onboard sensors (pressure, 3-axis acceleration) attached to animals using suction cups. The number and timing of breaths were quantified in non-linear mixed models that incorporated serial correlation and individual as a random effect. We found that pilot whales increased their BR in the 5-10 min period prior to, and immediately following, dives that exceeded 31 m depth. While pre-dive BRs did not vary with dive duration, the initial post-dive BR was linearly correlated with duration of >2 min dives, with BR then declining exponentially. Apparent net diving costs were 1.7 (SE 0.2) breaths per min of diving (post-dive number of breaths, above pre-dive breathing rate unrelated to dive recovery). Every fluke stroke was estimated to cost 0.086 breaths, which amounted to 80-90% average contribution of locomotion to the net diving costs. After accounting for fluke stroke rate, individuals in the small body size class took a greater number of breaths per diving minute. Individuals reduced their breathing rate (from the rate expected by diving behavior) by 13-16% during playbacks of killer whale sounds and their first exposure to 1-2 kHz naval sonar, indicating similar responses to interspecific competitor/predator and anthropogenic sounds. Although we cannot rule out individuals increasing their per-breath O2 uptake to match metabolic demand, our results suggest that behavioral responses to experimental sound exposures were not associated with increased metabolic rates in a stress response, but metabolic rates instead appear to decrease. Our results support the hypothesis that maximal performance leads to predictable (optimized) breathing patterns, which combined with further physiological measurements could improve proxies of field metabolic rates and per-stroke energy costs from animal-borne behavior data.
Project description: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 (<i>Physeter macrocephalus; N</i> = 46), blue whales (<i>Balaenoptera musculus; N</i> = 8), and fin whales (<i>B. physalus; N</i> = 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.
Project description:Here, we describe the diving behavior of sperm whales (Physeter macrocephalus) using the Advanced Dive Behavior (ADB) tag, which records depth data at 1-Hz resolution and GPS-quality locations for over 1 month, before releasing from the whale for recovery. A total of 27 ADB tags were deployed on sperm whales in the central Gulf of California, Mexico, during spring 2007 and 2008, of which 10 were recovered for data download. Tracking durations of all tags ranged from 0 to 34.5 days (median = 2.3 days), and 0.6 to 26.6 days (median = 5.0 days) for recovered tags. Recovered tags recorded a median of 50.8 GPS-quality locations and 42.6 dives per day. Dive summary metrics were generated for archived dives and were subsequently classified into six categories using hierarchical cluster analysis. A mean of 77% of archived dives per individual were one of four dive categories with median Maximum Dive Depth >290 m (V-shaped, Mid-water, Benthic, or Variable), likely associated with foraging. Median Maximum Dive Depth was <30 m for the other two categories (Short- and Long-duration shallow dives), likely representing socializing or resting behavior. Most tagged whales remained near the tagging area during the tracking period, but one moved north of Isla Tiburón, where it appeared to regularly dive to, and travel along the seafloor. Three whales were tagged on the same day in 2007 and subsequently traveled in close proximity (<1 km) for 2 days. During this period, the depth and timing of their dives were not coordinated, suggesting they were foraging on a vertically heterogeneous prey field. The multiweek dive records produced by ADB tags enabled us to generate a robust characterization of the diving behavior, activity budget, and individual variation for an important predator of the mesopelagos over temporal and spatial scales not previously possible.
Project description:<h4>Background</h4>Why a variety of social animals emit foraging-associated calls during group foraging remains an open question. These vocalizations may be used to recruit conspecifics to food patches (i.e. food advertisement hypothesis) or defend food resources against competitors (food defence hypothesis), presumably depending on food availability. Insectivorous bats rely heavily on vocalizations for navigation, foraging, and social interactions. In this study, we used free-ranging big-footed myotis (Myotis macrodactylus Temminck, 1840) to test whether social calls produced in a foraging context serve to advertise food patches or to ward off food competitors. Using a combination of acoustic recordings, playback experiments with adult females and dietary monitoring (light trapping and DNA metabarcoding techniques), we investigated the relationship between insect availability and social vocalizations in foraging bats.<h4>Results</h4>The big-footed myotis uttered low-frequency social calls composed of 7 syllable types during foraging interactions. Although the dietary composition of bats varied across different sampling periods, Diptera, Lepidoptera, and Trichoptera were the most common prey consumed. The number of social vocalizations was primarily predicted by insect abundance, insect species composition, and echolocation vocalizations from conspecifics. The number of conspecific echolocation pulses tended to decrease following the emission of most social calls. Feeding bats consistently decreased foraging attempts and food consumption during playbacks of social calls with distinctive structures compared to control trials. The duration of flight decreased 1.29-1.96 fold in the presence of social calls versus controls.<h4>Conclusions</h4>These results support the food defence hypothesis, suggesting that foraging bats employ social calls to engage in intraspecific food competition. This study provides correlative evidence for the role of insect abundance and diversity in influencing the emission of social calls in insectivorous bats. Our findings add to the current knowledge of the function of social calls in echolocating bats.
Project description:Interactions between individuals of different cetacean species are often observed in the wild. Killer whales (Orcinus orca) can be potential predators of many other cetaceans, and the interception of their vocalizations by unintended cetacean receivers may trigger anti-predator behavior that could mediate predator-prey interactions. We explored the anti-predator behaviour of five typically-solitary male sperm whales (Physeter macrocephalus) in the Norwegian Sea by playing sounds of mammal-feeding killer whales and monitoring behavioural responses using multi-sensor tags. Our results suggest that, rather than taking advantage of their large aerobic capacities to dive away from the perceived predator, sperm whales responded to killer whale playbacks by interrupting their foraging or resting dives and returning to the surface, changing their vocal production, and initiating a surprising degree of social behaviour in these mostly solitary animals. Thus, the interception of predator vocalizations by male sperm whales disrupted functional behaviours and mediated previously unrecognized anti-predator responses.