Kinematics and energetics of foraging behavior in Rice's whales of the Gulf of Mexico.

Rorqual foraging behavior varies with species, prey type and foraging conditions, and can be a determining factor for their fitness. Little is known about the foraging ecology of Rice's whales (Balaenoptera ricei), an endangered species with a population of fewer than 100 individuals. Suction cup tags were attached to two Rice's whales to collect information on their diving kinematics and foraging behavior. The tagged whales primarily exhibited lunge-feeding near the sea bottom and to a lesser extent in the water-column and at the sea surface. During 6-10 min foraging dives, the whales typically circled their prey before executing one or two feeding lunges. Longer duration dives and dives with more feeding-lunges were followed by an increase in their breathing rate. The median lunge rate of one lunge per dive of both animals was much lower than expected based on comparative research on other lunge-feeding baleen whales, and may be associated with foraging on fish instead of krill or may be an indication of different foraging conditions. Both animals spent extended periods of the night near the sea surface, increasing the risk for ship strike. Furthermore, their circling before lunging may increase the risk for entanglement in bottom-longline fishing gear. Overall, these data show that Rice's whale foraging behavior differs from other lunge feeding rorqual species and may be a significant factor in shaping our understanding of their foraging ecology. Efforts to mitigate threats to Rice's whales will benefit from improved understanding of patterns in their habitat use and fine-scale ecology.

SoundScape learning: An automatic method for separating fish chorus in marine soundscapes.

Marine soundscapes provide the opportunity to non-invasively learn about, monitor, and conserve ecosystems. Some fishes produce sound in chorus, often in association with mating, and there is much to learn about fish choruses and the species producing them. Manually analyzing years of acoustic data is increasingly unfeasible, and is especially challenging with fish chorus, as multiple fish choruses can co-occur in time and frequency and can overlap with vessel noise and other transient sounds. This study proposes an unsupervised automated method, called SoundScape Learning (SSL), to separate fish chorus from soundscape using an integrated technique that makes use of randomized robust principal component analysis (RRPCA), unsupervised clustering, and a neural network. SSL was applied to 14 recording locations off southern and central California and was able to detect a single fish chorus of interest in 5.3 yrs of acoustically diverse soundscapes. Through application of SSL, the chorus of interest was found to be nocturnal, increased in intensity at sunset and sunrise, and was seasonally present from late Spring to late Fall. Further application of SSL will improve understanding of fish behavior, essential habitat, species distribution, and potential human and climate change impacts, and thus allow for protection of vulnerable fish species.

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.

An echosounder view on the potential effects of impulsive noise pollution on pelagic fish around windfarms in the North Sea.

Anthropogenic noise in the oceans is disturbing marine life. Among other groups, pelagic fish are likely to be affected by sound from human activities, but so far have received relatively little attention. Offshore wind farms have become numerous and will become even more abundant in the next decades. Wind farms can be interesting to pelagic fish due to food abundance or fisheries restrictions. At the same time, construction of wind farms involves high levels of anthropogenic noise, likely disturbing and/or deterring pelagic fish. Here, we investigated whether bottom-moored echosounders are a suitable tool for studying the effects of impulsive - intermittent, high-intensity - anthropogenic noise on pelagic fish around wind farms and we explored the possible nature of their responses. Three different wind farms along the Dutch and Belgian coast were examined, one with exposure to the passing by of an experimental seismic survey with a full-scale airgun array, one with pile driving activity in an adjacent wind farm construction site and one control site without exposure. Two bottom-moored echosounders were placed in each wind farm and recorded fish presence and behaviour before, during and after the exposures. The echosounders were successful in detecting variation in the number of fish schools and their behaviour. During the seismic survey exposure there were significantly fewer, but more cohesive, schools than before, whereas during pile driving fish swam shallower with more cohesive schools. However, the types and magnitudes of response patterns were also observed at the control site with no impulsive sound exposure. We therefore stress the need for thorough replication beyond single case studies, before we can conclude that impulsive sounds, from either seismic surveys or pile driving, are a disturbing factor for pelagic fish in otherwise attractive habitat around wind farms.

Diving apart together: call propagation in diving long-finned pilot whales.

Group-living animals must communicate to stay in contact. In long-finned pilot whales, there is a trade-off between the benefits of foraging individually at depth and the formation of tight social groups at the surface. Using theoretical modelling and empirical data of tagged pairs within a group, we examined the potential of pilot whale social calls to reach dispersed group members during foraging periods. Both theoretical predictions and empirical data of tag pairs showed a potential for communication between diving and non-diving group members over separation distances up to 385 m (empirical) and 1800 m (theoretical). These distances match or exceed pilot whale dive depths recorded across populations. Call characteristics and environmental characteristics were analysed to investigate determinants of call detectability. Longer calls with a higher sound pressure level (SPL) that were received in a quieter environment were more often detected than their shorter, lower SPL counterparts within a noisier environment. In a noisier environment, calls were louder and had a lower peak frequency, indicating mechanisms for coping with varying conditions. However, the vulnerability of pilot whales to anthropogenic noise is still of concern as the ability to cope with increasing background noise may be limited. Our study shows that combining propagation modelling and actual tag recordings provides new insights into the communicative potential for social calls in orientation and reunion with group members for deep-diving pilot whales.

Spatial avoidance to experimental increase of intermittent and continuous sound in two captive harbour porpoises.

The continuing rise in underwater sound levels in the oceans leads to disturbance of marine life. It is thought that one of the main impacts of sound exposure is the alteration of foraging behaviour of marine species, for example by deterring animals from a prey location, or by distracting them while they are trying to catch prey. So far, only limited knowledge is available on both mechanisms in the same species. The harbour porpoise (Phocoena phocoena) is a relatively small marine mammal that could quickly suffer fitness consequences from a reduction of foraging success. To investigate effects of anthropogenic sound on their foraging efficiency, we tested whether experimentally elevated sound levels would deter two captive harbour porpoises from a noisy pool into a quiet pool (Experiment 1) and reduce their prey-search performance, measured as prey-search time in the noisy pool (Experiment 2). Furthermore, we tested the influence of the temporal structure and amplitude of the sound on the avoidance response of both animals. Both individuals avoided the pool with elevated sound levels, but they did not show a change in search time for prey when trying to find a fish hidden in one of three cages. The combination of temporal structure and SPL caused variable patterns. When the sound was intermittent, increased SPL caused increased avoidance times. When the sound was continuous, avoidance was equal for all SPLs above a threshold of 100 dB re 1 µPa. Hence, we found no evidence for an effect of sound exposure on search efficiency, but sounds of different temporal patterns did cause spatial avoidance with distinct dose-response patterns.

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.

On growth and form of irregular coiled-shell of a terrestrial snail: Plectostoma concinnum (Fulton, 1901) (Mollusca: Caenogastropoda: Diplommatinidae).

The molluscan shell can be viewed as a petrified representation of the organism's ontogeny and thus can be used as a record of changes in form during growth. However, little empirical data is available on the actual growth and form of shells, as these are hard to quantify and examine simultaneously. To address these issues, we studied the growth and form of a land snail that has an irregularly coiled and heavily ornamented shell-Plectostoma concinnum. The growth data were collected in a natural growth experiment and the actual form changes of the aperture during shell ontogeny were quantified. We used an ontogeny axis that allows data of growth and form to be analysed simultaneously. Then, we examined the association between the growth and the form during three different whorl growing phases, namely, the regular coiled spire phase, the transitional constriction phase, and the distortedly-coiled tuba phase. In addition, we also explored the association between growth rate and the switching between whorl growing mode and rib growing mode. As a result, we show how the changes in the aperture ontogeny profiles in terms of aperture shape, size and growth trajectory, and the changes in growth rates, are associated with the different shell forms at different parts of the shell ontogeny. These associations suggest plausible constraints that underlie the three different shell ontogeny phases and the two different growth modes. We found that the mechanism behind the irregularly coiled-shell is the rotational changes of the animal's body and mantle edge with respect to the previously secreted shell. Overall, we propose that future study should focus on the role of the mantle and the columellar muscular system in the determination of shell form.