Cooperative partner choice in multi-level male dolphin alliances.

Investigations into cooperative partner choice should consider both potential and realised partners, allowing for the comparison of traits across all those available. Male bottlenose dolphins form persisting multi-level alliances. Second-order alliances of 4-14 males are the core social unit, within which 2-3 males form first-order alliances to sequester females during consortships. We compared social bond strength, relatedness and age similarity of potential and realised partners of individual males in two age periods: (i) adolescence, when second-order alliances are formed from all available associates, and (ii) adulthood, when first-order allies are selected from within second-order alliances. Social bond strength during adolescence predicted second-order alliance membership in adulthood. Moreover, males preferred same-aged or older males as second-order allies. Within second-order alliances, non-mating season social bond strength predicted first-order partner preferences during mating season consortships. Relatedness did not influence partner choice on either alliance level. There is thus a striking resemblance between male dolphins, chimpanzees and humans, where closely bonded non-relatives engage in higher-level, polyadic cooperative acts. To that end, our study extends the scope of taxa in which social bonds rather than kinship explain cooperation, providing the first evidence that such traits might have evolved independently in marine and terrestrial realms.

Seasonal flow dynamics exacerbate overlap between artisanal fisheries and imperiled Ganges River dolphins.

Here we quantify the effects of artisanal fisheries on the ecology of a small cetacean, the Ganges River dolphin (Platanista gangetica gangetica, GRD), in a large river system of Nepal. We examine the size-classes of fisheries' catches, behavioural changes in GRD in response to fishing activities, and diel overlap between GRD and fishing activity. We observed high human exploitation rates (> 60% of the total catch per effort) of GRD-preferred prey sizes, indicating risks of high resource competition and dietary overlap, especially during the low water season when resource availability is reduced. Competitive interactions in the feeding niches during the low water season, plus temporal overlap between the peak exploitation and critical life-history events (e.g., reproduction), likely have ecological consequences. Furthermore, we detected 48% (95% CI 43-52%) increase in the chance of behavioural changes among dolphins exposed to anthropopressure (fishing activity), risking social behaviour impairment in exposed dolphins. The higher diel overlap and increased diel coefficient as the surveys progressed towards the monsoon season suggest temporal shifts in GRD socio-behavioural states and seasonal effects on resource partitioning, respectively. This work identifies drivers of small cetaceans-fisheries interactions and their consequences, and can be used to help reduce biologically significant fishing impacts on small cetaceans. Mitigation strategies, together with river sanctuary and distanced-based approaches, should be urgently included in a framework of ecosystem-based management.

Potential Welfare Impacts of Chase and Capture of Small Cetaceans during Drive Hunts in Japan.

Drive hunts are a method to herd, capture and kill small cetaceans (whales and dolphins) in coastal waters of some countries including Japan and the Faroe Islands. In Japan, these methods are often associated with the acquisition of live dolphins for international marine parks and aquaria. During the hunts, dolphins are herded by a flotilla of fishing vessels and loud underwater noise created by fishermen banging hammers on metal poles. The prolonged and strenuous chase and use of sound barriers to herd, capture, and restrain the dolphins can result in acute stress and injury. The authors review physiological and behavioral data pertaining to chase, encirclement, and live capture of dolphins and draw comparisons between chase and capture data for marine and terrestrial species. This analysis raises substantial welfare concerns associated with the hunts and acquisition of dolphins from such capture operations. The authors assert that this data detailing the negative impacts of chase, herding and handling (capture) of small cetaceans renders these hunts inherently inhumane and should inform policy relating to the collection and management of dolphins in the wild.

Swimming features in captive odontocetes: Indicative of animals' emotional state?

Captive welfare studies in odontocete species have been mostly conducted on bottlenose dolphins (Tursiops truncatus) while the welfare of many other species' -including endangered species- remains poorly studied. More research is needed to find and validate potential indicators of welfare for each species and even for each group. Since captive odontocetes spend most of their time swimming, their swimming features are interesting to study in relation to their welfare state. We first analysed the circular swimming direction bias in three groups of captive odontocetes (Yangtze finless porpoises: Neophocaena asiaeorientalis asiaeorientalis; East-Asian finless porpoises: N. a. sunameri; and bottlenose dolphins, Tursiops truncatus). Second, we studied the effect of environmental and social factors (i.e., time of the day, delay to training, enrichment, potential perturbation, social grouping, public presence and housing pool) on circular swimming, fast swimming, group swimming, synchronous swimming and contact swimming in the three groups. Yangtze finless porpoises exhibited a clockwise swimming bias while East-Asian finless porpoises and bottlenose dolphins swam significantly more in the counter-clockwise direction. Each studied factor significantly impacted the animals' swimming behaviour slightly differently depending on the group. However, some patterns were common for the three groups: animals seemed to be more active in the morning than at noon and in the afternoon, and enrichment seemed to decrease circular swimming, fast swimming and social swimming (i.e., synchronous, contact and group swimming), while potential perturbations (e.g., pool cleaning, noise) seemed to increase it. In addition, behaviour differed for Yangtze finless porpoises and bottlenose dolphins right before the training or when other animals were being trained, suggesting an anticipation of this event or an excited/frustrated state in this context. Social separation also impacted these animals' swimming behaviour with less group swimming but more circular swimming, synchronous swimming and fast swimming when separated. The housing pool had an impact on bottlenose dolphins' behaviour with more circular swimming, more fast swimming and less group swimming when having access to a larger space. The effect of the presence of public was unclear and requires further investigation. From our results, we propose that circular swimming, synchronous swimming and contact swimming could be useful to monitor animals' emotional state, but that additional parameters should be added (e.g., swimming speed) since these behaviours can be expressed both in quiet and relaxed contexts and in stressful ones. In addition, fast swimming can be a useful indicator of stress for porpoises but might be more ambiguous for bottlenose dolphins that engage in intense social play bouts for instance. Finally, group swimming might be a good behaviour to monitor when wanting to investigate reactions to various conditions or events that can potentially be stressful. We suggest that further research should be conducted on other groups of odontocetes to validate our findings.

Acoustic recordings of rough-toothed dolphin (Steno bredanensis) offshore Eastern Sicily (Mediterranean Sea).

Rough-toothed dolphin's abundance and distribution is largely unknown worldwide and evaluation of its conservation status in the Mediterranean Sea is necessary. A rough-toothed dolphin was sighted offshore Eastern Sicily (Mediterranean Sea) in July 2017 and acoustic data were acquired in the same area of Watkins, Tyack, Moore, and Notarbartolo di Sciara [(1987). Mar. Mamm. Sci. 3, 78-82]. An automatic detection algorithm was developed to identify the echolocation clicks recorded within both datasets and a recurrent inter-click interval value was identified during the new encounter. Distinctive whistle classes were also identified with similar contour shapes within both datasets.

Global vs local modularity for network community detection.

Community structures are ubiquitous in various complex networks, implying that the networks commonly be composed of groups of nodes with more internal links and less external links. As an important topic in network theory, community detection is of importance for understanding the structure and function of the networks. Optimizing statistical measures for community structures is one of most popular strategies for community detection in complex networks. In the paper, by using a type of self-loop rescaling strategy, we introduced a set of global modularity functions and a set of local modularity functions for community detection in networks, which are optimized by a kind of the self-consistent method. We carefully compared and analyzed the behaviors of the modularity-based methods in community detection, and confirmed the superiority of the local modularity for detecting community structures on large-size and heterogeneous networks. The local modularity can more quickly eliminate the first-type limit of modularity, and can eliminate or alleviate the second-type limit of modularity in networks, because of the use of the local information in networks. Moreover, we tested the methods in real networks. Finally, we expect the research can provide useful insight into the problem of community detection in complex networks.

Selective reactions to different killer whale call categories in two delphinid species.

The risk of predation is often invoked as an important factor influencing the evolution of social organization in cetaceans, but little direct information is available about how these aquatic mammals respond to predators or other perceived threats. We used controlled playback experiments to examine the behavioral responses of short-finned pilot whales (Globicephala macrorhynchus) off Cape Hatteras, NC, USA, and Risso's dolphins (Grampus griseus) off the coast of Southern California, USA, to the calls of a potential predator, mammal-eating killer whales. We transmitted calls of mammal-eating killer whales, conspecifics and baleen whales to 10 pilot whales and four Risso's dolphins equipped with multi-sensor archival acoustic recording tags (DTAGs). Only playbacks of killer whale calls resulted in significant changes in tagged animal heading. The strong responses observed in both species occurred only following exposure to a subset of killer whale calls, all of which contained multiple non-linear properties. This finding suggests that these structural features of killer whale calls convey information about predatory risk to pilot whales and Risso's dolphins. The observed responses differed between the two species; pilot whales approached the sound source while Risso's dolphins fled following playbacks. These divergent responses likely reflect differences in anti-predator response mediated by the social structure of the two species.

Whale and dolphin behavioural responses to dead conspecifics.

The scientific study of death across animal taxa-comparative thanatology-investigates how animals respond behaviourally, physiologically and psychologically to dead conspecifics, and the processes behind such responses. Several species of cetaceans have been long known to care for, attend to, be aroused by, or show interest in dead or dying individuals. We investigated patterns and variation in cetacean responses to dead conspecifics across cetacean taxa based on a comprehensive literature review. We analysed 78 records reported between 1970 and 2016, involving 20 of the 88 extant cetacean species. We adopted a weighted comparative approach to take observation effort into account and found that odontocetes (toothed cetaceans) were much more likely than mysticetes (baleen whales) to attend to dead conspecifics. Dolphins (Delphinidae) had the greatest occurrence of attentive behaviour (92.3% of all records), with a weighed attendance index 18 times greater than the average of all other cetacean families. Two dolphin genera, Sousa and Tursiops, constituted 55.1% of all cetacean records (N=43) and showed the highest incidence of attentive behaviour. Results of analyses intended to investigate the reasons behind these differences suggested that encephalisation may be an important predictor, consistent with the "social brain" hypothesis. Among attending individuals or groups of known sex (N=28), the majority (75.0%) were adult females with dead calves or juveniles (possibly their own offspring, with exceptions), consistent with the strong mother-calf bond, or, in a few cases, with the bond between mothers and other females in the group. The remaining records (25.0%) involved males either showing sexual interest in a dead adult or subadult, or carrying a dead calf in the presence of females. Because an inanimate individual is potentially rescuable, responses to dead conspecifics-especially by females-can be explained at least in part by attempts to revive and protect, having a clear adaptive value. In some cases such responses are followed by apparently maladaptive behaviour such as the long-term carrying of, or standing by, a decomposed carcass, similar to observations of certain terrestrial mammals. Among the possible explanations for the observed cetacean behavioural responses to dead conspecifics are strong attachment resulting in a difficulty of "letting go"-possibly related to grieving-or perhaps individuals failing to recognise or accept that an offspring or companion has died. Our current understanding is challenged by small sample size, incomplete descriptions, and lack of information on the physiology and neural processes underpinning the observed behaviour. We provide research recommendations that would improve such understanding.

Risso's dolphins plan foraging dives.

Humans remember the past and use that information to plan future actions. Lab experiments that test memory for the location of food show that animals have a similar capability to act in anticipation of future needs, but less work has been done on animals foraging in the wild. We hypothesized that planning abilities are critical and common in breath-hold divers who adjust each dive to forage on prey varying in quality, location and predictability within constraints of limited oxygen availability. We equipped Risso's dolphins with sound-and-motion recording tags to reveal where they focus their attention through their externally observable echolocation and how they fine tune search strategies in response to expected and observed prey distribution. The information from the dolphins was integrated with synoptic prey data obtained from echosounders on an underwater vehicle. At the start of the dives, whales adjusted their echolocation inspection ranges in ways that suggest planning to forage at a particular depth. Once entering a productive prey layer, dolphins reduced their search range comparable to the scale of patches within the layer, suggesting that they were using echolocation to select prey within the patch. On ascent, their search range increased, indicating that they decided to stop foraging within that layer and started searching for prey in shallower layers. Information about prey, learned throughout the dive, was used to plan foraging in the next dive. Our results demonstrate that planning for future dives is modulated by spatial memory derived from multi-modal prey sampling (echoic, visual and capture) during earlier dives.

Broadband ship noise and its potential impacts on Indo-Pacific humpback dolphins: Implications for conservation and management.

Ship noise pollution has raised considerable concerns among regulatory agencies and cetacean researchers worldwide. There is an urgent need to quantify ship noise in coastal areas and assess its potential biological impacts. In this study, underwater broadband noise from commercial ships in a critical habitat of Indo-Pacific humpback dolphins was recorded and analyzed. Data analysis indicated that the ship noise caused by the investigated commercial ships with an average length of 134 ± 81 m, traveling at 18.8 ± 2.5 km/h [mean ± standard deviation (SD), n = 21] comprises mid-to-high components with frequencies approaching and exceeding 100 kHz, and the ship noise could be sensed auditorily by Indo-Pacific humpback dolphins within most of their sensitive frequency range. The contributions of ship noise to ambient noise were highest in two third-octave bands with center frequencies of 8 and 50 kHz, which are within the sensitive hearing range of Indo-Pacific humpback dolphins and overlap the frequency of sounds that are biologically significant to the dolphins. It is estimated that ship noise in these third-octave bands can be auditorily sensed by and potentially affect the dolphins within 2290 ± 1172 m and 848 ± 358 m (mean ± SD, n = 21), respectively.

Acoustic occurrence detection of a newly recorded Indo-Pacific humpback dolphin population in waters southwest of Hainan Island, China.

In 2014, Indo-Pacific humpback dolphins were recorded for the first time in waters southwest of Hainan Island, China. In this paper, the temporal occurrence of Indo-Pacific humpback dolphins in this region was detected by stationary passive acoustic monitoring. During the 130-day observation period (from January to July 2016), 1969 click trains produced by Indo-Pacific humpback dolphins were identified, and 262 ten-minute recording bins contained echolocation click trains of dolphins, of which 70.9% were at night and 29.1% were during the day. A diurnal rhythm with a nighttime peak in acoustic detections was found. Passive acoustic detections indicated that the Indo-Pacific humpback dolphins frequently occurred in this area and were detected mainly at night. This information may be relevant to conservation efforts for these dolphins in the near future.

A simulation of temperature influence on echolocation click beams of the Indo-Pacific humpback dolphin (Sousa chinensis).

A finite element method was used to investigate the temperature influence on sound beams of the Indo-Pacific humpback dolphin. The numerical models of a dolphin, which originated from previous computed tomography (CT) scanning and physical measurement results, were used to investigate sound beam patterns of the dolphin in temperatures from 21 °C to 39 °C, in increments of 2 °C. The -3 dB beam widths across the temperatures ranged from 9.3° to 12.6°, and main beam angle ranged from 4.7° to 7.2° for these temperatures. The subsequent simulation suggested that the dolphin's sound beam patterns, side lobes in particular, were influenced by temperature.

Multi-modal sexual displays in Australian humpback dolphins.

Sexual displays enriched by object carrying serve to increase individual male fitness, yet are uncommon phenomena in the animal kingdom. While they have been documented in a variety of taxa, primarily birds, they are rare outside non-human mammals. Here, we document marine sponge presenting associated with visual and acoustic posturing found in several, geographically widespread populations of Australian humpback dolphins (Sousa sahulensis) over ten years of observation. Only adult males presented marine sponges, typically doing so in the presence of sexually mature females, although social groups predominantly consisted of mixed age and sex classes. Male humpback dolphins appear to be using sponges for signalling purposes in multi-modal sexual displays. Further, based on limited behavioural and genetic data, we hypothesise that pairs of adult male Sousa form at least temporary coalitions or alliances. The use of objects in sexual displays by non-human mammals is rare and, moreover, cooperation between males in the pursuit of an indivisible resource is an evolutionary hurdle relatively few species have overcome. These findings suggest a hitherto unrecognised level of social complexity in humpback dolphins.

Categorizing click trains to increase taxonomic precision in echolocation click loggers.

Passive acoustic monitoring is an efficient way to study acoustically active animals but species identification remains a major challenge. C-PODs are popular logging devices that automatically detect odontocete echolocation clicks. However, the accompanying analysis software does not distinguish between delphinid species. Click train features logged by C-PODs were compared to frequency spectra from adjacently deployed continuous recorders. A generalized additive model was then used to categorize C-POD click trains into three groups: broadband click trains, produced by bottlenose dolphin (Tursiops truncatus) or common dolphin (Delphinus delphis), frequency-banded click trains, produced by Risso's (Grampus griseus) or white beaked dolphins (Lagenorhynchus albirostris), and unknown click trains. Incorrect categorization rates for broadband and frequency banded clicks were 0.02 (SD 0.01), but only 30% of the click trains met the categorization threshold. To increase the proportion of categorized click trains, model predictions were pooled within acoustic encounters and a likelihood ratio threshold was used to categorize encounters. This increased the proportion of the click trains meeting either the broadband or frequency banded categorization threshold to 98%. Predicted species distribution at the 30 study sites matched well to visual sighting records from the region.

Geographic variation in Risso's dolphin echolocation click spectra.

Discrimination of bioacoustic signals to the species or population level is critical for using passive acoustic monitoring to study cetacean ecology. Risso's dolphins off southern California have distinctive peaks and notches in their echolocation clicks, but it was unknown whether Risso's dolphins from other geographic areas have similarly distinctive click spectra and whether populations are acoustically distinct. This study investigates using clicks for species and population identification by characterizing the spectral structure of Risso's dolphin echolocation clicks recorded over wide-ranging geographic regions including the U.S. waters of the North Atlantic Ocean, Gulf of Mexico, and North Pacific Ocean; and international waters of the Eastern Tropical Pacific. All recordings with Risso's dolphin clicks exhibited the spectral peak and notch pattern described off southern California, indicating the presence of peak banding patterns is useful for species discrimination. Geographic regions were a significant explanatory factor for variability in the frequencies of click spectral peaks, with relatively higher frequency peaks and notches found off Hawaii compared to California waters and off the southeast U.S. compared to the Gulf of Mexico. In the North Atlantic Ocean, a latitudinal cline in frequencies was evident. Potential causes of acoustic variation within and among acoustic encounters are evaluated.

Limited capacity of working memory in unihemispheric random walks implies conceivable slow dispersal.

Phenomenologically inspired by dolphins' unihemispheric sleep, we introduce a minimal model for random walks with physiological memory. The physiological memory consists of long-term memory which includes unconscious implicit memory and conscious explicit memory, and working memory which serves as a multi-component system for integrating, manipulating and managing short-term storage. The model assumes that the sleeping state allows retrievals of episodic objects merely from the episodic buffer where these memory objects are invoked corresponding to the ambient objects and are thus object-oriented, together with intermittent but increasing use of implicit memory in which decisions are unconsciously picked up from historical time series. The process of memory decay and forgetting is constructed in the episodic buffer. The walker's risk attitude, as a product of physiological heuristics according to the performance of objected-oriented decisions, is imposed on implicit memory. The analytical results of unihemispheric random walks with the mixture of object-oriented and time-oriented memory, as well as the long-time behavior which tends to the use of implicit memory, are provided, indicating the common sense that a conservative risk attitude is inclinable to slow movement.

Responses to familiar and unfamiliar objects by belugas (Delphinapterus leucas), bottlenose dolphins (Tursiops truncatus), and Pacific white-sided dolphins (Lagenorhynchus obliquidens).

Previous research with bottlenose dolphins (Tursiops truncatus) demonstrated their ability to discriminate between familiar and unfamiliar stimuli. Dolphins gazed longer at unfamiliar stimuli. The current study attempted to extend this original research by examining the responses of three species of cetaceans to objects that differed in familiarity. Eleven belugas from two facilities, five bottlenose dolphins and five Pacific white-sided dolphins housed at one facility were presented different objects in a free-swim scenario. The results indicated that the animals gazed the longest at unfamiliar objects, but these gaze durations did not significantly differ from gaze durations when viewing familiar objects. Rather, the animals gazed longer at unfamiliar objects when compared to the apparatus alone. Species differences emerged with longer gaze durations exhibited by belugas and bottlenose dolphins and significantly shorter gaze durations for Pacific white-sided dolphins. It is likely that the animals categorized objects into familiar and unfamiliar categories, but the free-swim paradigm in naturalistic social groupings did not elicit clear responses. Rather this procedure emphasized the importance of attention and individual preferences when investigating familiar and unfamiliar objects, which has implications for cognitive research and enrichment use.

Burst pulses of Guiana dolphin (Sotalia guianensis) in southeastern Brazil.

Acoustic studies of Guiana dolphin have been focused on whistles, with little known about pulse signals in this species. This study characterized the temporal and spectral properties of Guiana dolphin burst pulses. Groups of 2 to 23 Guiana dolphins were recorded while feeding and socializing in shallow waters in Guanabara Bay, southeastern Brazil, in 2013 and 2014. Burst pulse analysis involved two steps: signal detection and acoustic parameter analysis. Eight variables were analyzed for 197 total burst pulses: number of clicks, burst pulse duration, interclick interval, click duration, peak frequency, center frequency, -3 dB bandwidth, and -10 dB bandwidth. Mean burst pulse duration was 108.6 ms [standard deviation (SD) = 91.3] with a mean of 168 clicks (SD = 137.3). Burst pulses had short interclick interval (0.7 ms, SD = 0.3) and mean click duration of 300 µs (SD = 100). Mean peak frequency and center frequency were 28 kHz (SD = 11.6) and 29 kHz (SD = 11.0), respectively. Mean -3 dB bandwidth was 15 kHz (SD = 7.5) and mean -10 dB bandwidth was 40.5 kHz (SD = 14.3). The quantitative characterization of Guiana dolphin burst pulses is an important step in describing the full acoustic repertoire of this species.

Automatic classification of whistles from coastal dolphins of the southern African subregion.

Passive acoustic monitoring (PAM) is commonly used to generate information on the distribution, abundance, and behavior of cetacean species. In African waters, the utilization of PAM lags behind most other continents. This study examines whether the whistles of three coastal delphinid species (Delphinus delphis, Tursiops truncatus, and Tursiops aduncus) commonly encountered in the southern African subregion can be readily distinguished using both statistical analysis of standard whistle parameters and the automated detection and classification software PAMGuard. A first account of whistles recorded from D. delphis from South Africa is included. Using PAMGuard, classification to species was high with an overall mean correct classification rate of 87.3%. Although lower, high rates of correct classification were also found (78.4%) when the two T. aduncus populations were included separately. Classification outcomes reflected patterns observed in standard whistle parameters. Such acoustic discrimination may be useful for confirmation of morphologically similar species in the field. Classification success was influenced by training and testing the classifier with data from different populations, highlighting the importance of locally collected acoustic data to inform classifiers. The small number of sampling populations may have inflated the classification success, therefore, classification trials using a greater number of species are recommended.

Social sounds produced by franciscana dolphins, Pontoporia blainvillei (Cetartiodactyla, Pontoporiidae).

Franciscana dolphin (Pontoporia blainvillei) whistles were documented for the first time during 2003-2013 in Babitonga Bay estuary, South Brazil, together with burst pulses. Recordings were made from small boats under good sea conditions, and recording equipment that allowed analysis of sounds up to 96 kHz. The recordings were made in the presence of 2-31 franciscana dolphins. During 23 h and 53 min, 90 whistles and 51 burst pulse series were recorded. Although Guiana dolphins (Sotalia guianensis) inhabit nearby waters, none were observed in the area during the recordings. The authors recorded ten types of whistles. The initial frequency varied between 1.6 and 94.6 kHz, and the final frequency varied between 0.7 and 94.5 kHz; the authors were not able to determine if dolphin whistles exceeded the 96 kHz recording limit of the authors' equipment, although that is likely, especially because some whistles showed harmonics. Whistle duration varied between 0.008 and 0.361 s. Burst pulses had initial frequencies between 69 and 82.1 kHz (77 ± 3.81). These results showed that P. blainvillei produces whistles and burst pulses, although they seem to be produced infrequently.