Sowerby's beaked whale biosonar and movement strategy indicate deep-sea foraging niche differentiation in mesoplodont whales.

Closely related species are expected to diverge in foraging strategy, reflecting the evolutionary drive to optimize foraging performance. The most speciose cetacean genus, Mesoplodon, comprises beaked whales with little diversity in external morphology or diet, and overlapping distributions. Moreover, the few studied species of beaked whales (Ziphiidae) show very similar foraging styles with slow, energy-conserving movement during long, deep foraging dives. This raises the question of what factors drive their speciation. Using data from animal-attached tags and aerial imagery, we tested the hypothesis that two similar-sized mesoplodonts, Sowerby's (Mesoplodon bidens) and Blainville's (Mesoplodon densirostris) beaked whales, exploit a similar low-energy niche. We show that, compared with the low-energy strategist Blainville's beaked whale, Sowerby's beaked whale lives in the fast lane. While targeting a similar mesopelagic/bathypelagic foraging zone, they consistently swim and hunt faster, perform shorter deep dives, and echolocate at a faster rate with higher frequency clicks. Further, extensive near-surface travel between deep dives challenges the interpretation of beaked whale shallow inter-foraging dives as a management strategy for decompression sickness. The distinctively higher frequency echolocation clicks do not hold apparent foraging benefits. Instead, we argue that a high-speed foraging style influences dive duration and echolocation behaviour, enabling access to a distinct prey population. Our results demonstrate that beaked whales exploit a broader diversity of deep-sea foraging and energetic niches than hitherto suspected. The marked deviation of Sowerby's beaked whales from the typical ziphiid foraging strategy has potential implications for their response to anthropogenic sounds, which appears to be strongly behaviourally driven in other ziphiids.

Scaling of maneuvering performance in baleen whales: larger whales outperform expectations.

Despite their enormous size, whales make their living as voracious predators. To catch their much smaller, more maneuverable prey, they have developed several unique locomotor strategies that require high energetic input, high mechanical power output and a surprising degree of agility. To better understand how body size affects maneuverability at the largest scale, we used bio-logging data, aerial photogrammetry and a high-throughput approach to quantify the maneuvering performance of seven species of free-swimming baleen whale. We found that as body size increases, absolute maneuvering performance decreases: larger whales use lower accelerations and perform slower pitch-changes, rolls and turns than smaller species. We also found that baleen whales exhibit positive allometry of maneuvering performance: relative to their body size, larger whales use higher accelerations, and perform faster pitch-changes, rolls and certain types of turns than smaller species. However, not all maneuvers were impacted by body size in the same way, and we found that larger whales behaviorally adjust for their decreased agility by using turns that they can perform more effectively. The positive allometry of maneuvering performance suggests that large whales have compensated for their increased body size by evolving more effective control surfaces and by preferentially selecting maneuvers that play to their strengths.

Friend or foe: Risso's dolphins eavesdrop on conspecific sounds to induce or avoid intra-specific interaction.

The detection and use of emitters' signals by unintended receivers, i.e., eavesdropping, represents an important and often low-cost way for animals to gather information from their environment. Acoustic eavesdropping can be a key driver in mediating intra- and interspecific interactions (e.g., cooperation, predator-prey systems), specifically in species such as cetaceans that use sound as a primary sensory modality. While most cetacean species produce context-specific sounds, little is known about the use of those sounds by potential conspecific eavesdroppers. We experimentally tested the hypothesis that a social cetacean, Risso's dolphin (Grampus griseus), is able to gather biologically relevant information by eavesdropping on conspecific sounds. We conducted playback experiments on free-ranging dolphins using three context-specific sounds stimuli and monitored their horizontal movement using visual or airborne focal follow observations. We broadcasted natural sequences of conspecific foraging sounds potentially providing an attractive dinner bell signal (n = 7), male social sounds simulating a risk of forthcoming agonistic interaction (n = 7) and female-calf social sounds representing no particularly threatening context (n = 7). We developed a quantitative movement response score and tested whether animals changed their direction of horizontal movement towards or away from the playback source. Dolphins approached the foraging and the social female-calf sounds whereas they avoided the social male sounds. Hence, by acoustically eavesdropping on conspecifics, dolphins can discriminate between social and behavioural contexts and anticipate potential threatening or beneficial situations. Eavesdropping and the ensuing classification of 'friend or foe' can thus shape intra-specific social interactions in cetaceans.

Risso's dolphins perform spin dives to target deep-dwelling prey.

Foraging decisions of deep-diving cetaceans can provide fundamental insight into food web dynamics of the deep pelagic ocean. Cetacean optimal foraging entails a tight balance between oxygen-conserving dive strategies and access to deep-dwelling prey of sufficient energetic reward. Risso's dolphins (Grampus griseus) displayed a thus far unknown dive strategy, which we termed the spin dive. Dives started with intense stroking and right-sided lateral rotation. This remarkable behaviour resulted in a rapid descent. By tracking the fine-scale foraging behaviour of seven tagged individuals, matched with prey layer recordings, we tested the hypothesis that spin dives are foraging dives targeting deep-dwelling prey. Hunting depth traced the diel movement of the deep scattering layer, a dense aggregation of prey, that resides deep during the day and near-surface at night. Individuals shifted their foraging strategy from deep spin dives to shallow non-spin dives around dusk. Spin dives were significantly faster, steeper and deeper than non-spin dives, effectively minimizing transit time to bountiful mesopelagic prey, and were focused on periods when the migratory prey might be easier to catch. Hence, whereas Risso's dolphins were mostly shallow, nocturnal foragers, their spin dives enabled extended and rewarding diurnal foraging on deep-dwelling prey.

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. SIGNIFICANCE STATEMENT: 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.

Evidence for distinct coastal and offshore communities of bottlenose dolphins in the north east Atlantic.

Bottlenose dolphin stock structure in the northeast Atlantic remains poorly understood. However, fine scale photo-id data have shown that populations can comprise multiple overlapping social communities. These social communities form structural elements of bottlenose dolphin (Tursiops truncatus) [corrected] populations, reflecting specific ecological and behavioural adaptations to local habitats. We investigated the social structure of bottlenose dolphins in the waters of northwest Ireland and present evidence for distinct inshore and offshore social communities. Individuals of the inshore community had a coastal distribution restricted to waters within 3 km from shore. These animals exhibited a cohesive, fission-fusion social organisation, with repeated resightings within the research area, within a larger coastal home range. The offshore community comprised one or more distinct groups, found significantly further offshore (>4 km) than the inshore animals. In addition, dorsal fin scarring patterns differed significantly between inshore and offshore communities with individuals of the offshore community having more distinctly marked dorsal fins. Specifically, almost half of the individuals in the offshore community (48%) had characteristic stereotyped damage to the tip of the dorsal fin, rarely recorded in the inshore community (7%). We propose that this characteristic is likely due to interactions with pelagic fisheries. Social segregation and scarring differences found here indicate that the distinct communities are likely to be spatially and behaviourally segregated. Together with recent genetic evidence of distinct offshore and coastal population structures, this provides evidence for bottlenose dolphin inshore/offshore community differentiation in the northeast Atlantic. We recommend that social communities should be considered as fundamental units for the management and conservation of bottlenose dolphins and their habitat specialisations.