Social Affiliations within Sperm Whale (Physeter macrocephalus) Groups

We examined patterns of affiliation within groups of sperm whales ( Physeter macrocephalus ), particularly concentrating on how short-term spatio–temporal associations reflect long-term relationships. Female and immature sperm whales live in stable, and partially matrilineal, social units. Two or more social units may move together for periods of several days, forming a cohesive group of about 20 animals. We observed that sperm whales in the eastern tropical Pacific quite consistently associated with members of their own social unit more than they did with other animals in their group with whom they did not share a long-term relationship. There was little evidence for preferred, or avoided, affiliations within social units, except in two large and relatively unstable units. In two well-studied groups, individuals did not show consistently favoured positions in the foraging rank relative to other members of their social unit. These results indicate the importance of long-term relationships to female and immature sperm whales, but suggest that relationships are quite homogeneous within social units.     

Behavior and social structure of the sperm whales of Dominica,West Indies

There is substantial geographic variation in the behavior and social structure of sperm whales worldwide. The population in the Eastern Caribbean is thought to be isolated from other areas in the North Atlantic. We describe the behavior and social structure of the sperm whales identified off Dominica during an eight year study (2005–2012; 92% of photographic identifications) with supplementary data collected from seven other organizations dating as far back as 1981. A total of 419 individuals were identified. Resighting rates (42% of individuals between years) and encounter rates with sperm whale groups (mean = 80.4% of days at sea) among this population were both comparatively high. Group sizes were small (7–9 individuals) and were comprised of just one social unit (mean = 6.76 individuals, SD = 2.80). We described 17 units which have been reidentified off Dominica across 2–27 yr. Mature males are seen regularly off Dominica, but residency in the area lasts only a few days to a few weeks. Males were reidentified across years spanning up to a decade. Management of this population within the multinational Wider Caribbean Region will require governments to work towards international agreements governing sperm whales as a cross‐border species of concern.     

An assessment of sperm whale occurrence and social structure off São Miguel Island,Azores using fluke and dorsal identification photographs

Female and immature sperm whales form stable social units in tropical to subtropical waters. One such area is the Azores archipelago, where details of their year-round occurrence and social organization are not well known. We used year-round sightings data collected from whale watching vessels to assess sperm whale occurrence and social structure off São Miguel Island, Azores from 2010 to 2017. Individuals were photo-identified by their flukes (n = 393) and dorsal pigmentation marks were examined to explore their use in assisting with reidentifications. Of all cataloged whales, 78.8% were sufficiently distinctive to be reidentified from dorsal pigmentation patterns. Associations between individuals were analyzed to determine social structure and delineate social units. We identified 12 units comprising 2–13 members that had stable, and perhaps preferred associations for periods up to the eight years of the study. Preferences between some pairs of units may exist, but more research is required to better understand dynamics within and between units. This local scale study is an important contributor to our knowledge of geographic variation on a global scale. We recommend making use of all available information from the entire archipelago, to further increase our understanding of sperm social organization in the Azores.     

Calf development and mother-calf spatial relationships in Southern right whales

Changes in spatial relationships between mother and calf right whales (Eubalaena australis) from birth to the separation of yearling calves from their mothers were observed. During the first few weeks of a calf's life, mother and calf were within close proximity over 90% of the time, and mothers were responsible for maintenance of contact with their infants. Later calves strayed farther and initiated many more leaves and approaches than their quiescent mothers. The mothers still maintained contact with their infants. Just before migration away from the area, pair members resumed close proximity and leaves and approaches by both diminished. Yearling calves, returning to the area with their mothers after six months, stayed close to their mothers and few leaves and approaches by either pair member were recorded. The yearling calves were responsible for maintaining contact as the mothers left them more than approached them. This behaviour on the part of mothers probably contributed to weaning of yearlings and separation after a few weeks in the area.     

Vocal repertoires and insights into social structure of sperm whales (Physeter macrocephalus) in Mauritius,southwestern Indian Ocean

Sperm whales communicate using codas (stereotyped click sequences). Females and juveniles live in long-term social units, and units with similar coda repertoires share vocal clan membership. Vocal clans exhibit culturally defined differences in their multilevel social structure. Here, we aimed to identify different social units among sperm whales in Mauritius in the southwestern Indian Ocean, and to describe and compare their coda repertoires to investigate the presence of different vocal clans. We conducted six boat-based surveys between 2008 and 2013, during which 101 different individuals were photo-identified. Analysis of associations between 22 resighted individuals divided them into four candidate social units, but a lack of resightings impedes solid delineation of social units. Based on number and rhythm of clicks, at least 24 discrete coda types were detected among 4,767 analyzed codas using two different classification methods. Comparison of coda repertoires recorded from seven sperm whale groupings revealed the possible existence of two sympatric vocal clans, but the size differences of recorded repertoires warrant caution of interpretation. To further evaluate social structuring and the presence of different vocal clans in this region, future surveys should aim at obtaining long-term sighting and acoustic data, and cover a more extensive area.     

Kinship of long-term associates in the highly social sperm whale

The evolution of stable social groups can be promoted by both indirect and direct fitness benefits. Sperm whales (Physeter macrocephalus) are highly social, with a hierarchical social structure based around core groups of adult females and subadults, a rare level of complexity among mammals. We combined long-term satellite tracking (ranging from 11 to 607 days) of 51 individual sperm whales with genetic kinship analysis to assess the pattern of kin associations within and among coherent social units. Unlike findings for other species with similar social structure, we find no consistent correlation between kinship and association apart from close associations between two pairs of first-order relatives. A third pair of first-order relatives did not associate, and overall, the mean relatedness was the same within as among social groups. However, social behaviour can also be promoted by ecological factors such as resource dispersion. We assessed putative foraging behaviour during travel from the satellite-tracking data, which suggested that prey resources were dispersed and unpredictable, a condition that could promote living in groups.     

Long-term social structure of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited by around 216 pilot whales, which are present all year round, and nothing is known about their social structure. The aim of this study is to analyse the inter-individual association patterns within this pilot whales community to (1) provide an insight on their long-term social system and (2) to assess the relationship between sexes within this social system. Between 1999 and 2006, 23,004 km was sampled in the Strait of Gibraltar, and 4,887 images of dorsal fins of pilot whales were taken from 226 groups. The sex of 56 of the individuals could be determined genetically. The strength of the behavioural relationships between dyads of individuals was calculated, and the temporal aspects of the social structure were evaluated, showing in a non-random social structure made by constant companions. The preferred associations between individuals consisted in associations of males–females. Eight long-term units could be found with different degrees of association rates. Consequently, we propose that, in the Strait, the pilot whales exhibit a hierarchical social system composed of a population encompassing several clans of pilot whales each containing several pods. Pods will then be formed by several line units, similar to killer whale matrilineal units.     

Occurrence and social structure of Baird's beaked whales,Berardius bairdii,in the Commander Islands,Russia

The social structure of Baird's beaked whales is completely unstudied, and it is unknown if either females or males form long‐term social associations or occur in stable groups. In this paper we summarize our observations of individually identified animals over the span of 6 yr to provide insight on their long‐term social structure. We have identified 122 whales, with 28 of them encountered three times or more and thus included in the analysis of social structure. We found that the whales exhibited nonrandom patterns of social associations with some individuals preferentially associating with each other. Whales with more scarred skin had higher maximum association coefficients, which indicates that older animals and/or males were more inclined to form stable associations. Cluster analysis with a modularity test for gregariousness divided the whales into four clusters. Whales from the same clusters did not always occur together, but some individuals retained stable associations over several years. The strength of social relationships decayed over periods of months, with between‐year relationships showing little deviation from what would be expected if association was random. Generally these findings do not correspond to a stable society with fixed groups but instead suggest a fission‐fusion society with some stable alliances.     

Dominance rank predicts social network position across developmental stages in rhesus monkeys

Social network analysis is increasingly common in studying complex interactions among individuals. Across a range of primates, high-ranking adults are generally more socially connected, which results in better fitness outcomes. However, it still remains unclear whether this relationship between social network position and dominance rank emerges in infancy and whether, in species with a social transmission of dominance rank, social network positions are driven by the presence of the mother. To fill this gap, we first explored whether dominance ranks were related to social network position, measured via eigenvector centrality, in infants, juveniles, and adults in a troop of semi-free-ranging rhesus macaques (Macaca mulatta). We then examined relationships between dominance rank and eigenvector centrality in a peer-only group of yearlings who were reared with their mothers in either a rich, socially complex environment of multigenerational (MG) kin support or a unigenerational group of mothers and their infants from birth through 8 months. In Experiment 1, we found that mother's network position predicted offspring network position and that dominants across all age categories were more central in affiliative networks (social contact, social grooming, and social play). Experiment 2 showed that high-ranking yearlings in a peer-only group were more central only in the social contact network. Moreover, yearlings reared in a socially complex environment of MG kin support were more central. Our findings suggest that the relationship between dominance rank and social network position begins early in life, and that complex early social environments can promote later social competency. Our data add to the growing body of evidence that the presence/absence of the mother and kin influence how dominance rank affects social network position. These findings have important implications for the role of caregivers in the social status of developing primates, which ultimately ties to health and fitness outcomes.     

Swim Speed,Behavior, and Movement of North Atlantic Right Whales (Eubalaena glacialis) in Coastal Waters of Northeastern Florida,USA

In a portion of the coastal waters of northeastern Florida, North Atlantic right whales (Eubalaena glacialis) occur close to shore from December through March. These waters are included within the designated critical habitat for right whales. Data on swim speed, behavior, and direction of movement – with photo-identification of individual whales – were gathered by a volunteer sighting network working alongside experienced scientists and supplemented by aerial observations. In seven years (2001–2007), 109 tracking periods or “follows” were conducted on right whales during 600 hours of observation from shore-based observers. The whales were categorized as mother-calf pairs, singles and non-mother-calf pairs, and groups of 3 or more individuals. Sample size and amount of information obtained was largest for mother-calf pairs. Swim speeds varied within and across observation periods, individuals, and categories. One category, singles and non mother-calf pairs, was significantly different from the other two – and had the largest variability and the fastest swim speeds. Median swim speed for all categories was 1.3 km/h (0.7 kn), with examples that suggest swim speeds differ between within-habitat movement and migration-mode travel. Within-habitat right whales often travel back-and-forth in a north-south, along-coast, direction, which may cause an individual to pass by a given point on several occasions, potentially increasing anthropogenic risk exposure (e.g., vessel collision, fishing gear entanglement, harassment). At times, mothers and calves engaged in lengthy stationary periods (up to 7.5 h) that included rest, nursing, and play. These mother-calf interactions have implications for communication, learning, and survival. Overall, these behaviors are relevant to population status, distribution, calving success, correlation to environmental parameters, survey efficacy, and human-impacts mitigation. These observations contribute important parameters to conservation biology, predictive modeling, and management. However, while we often search for predictions, patterns, and means, the message here is also about variability and the behavioral characteristics of individual whales.     

The social organization of resident-type killer whales (Orcinus orca) in Avacha Gulf,Northwest Pacific,as revealed through association patterns and acoustic similarity

Northeast Pacific resident-type killer whales (Orcinus orca) are known to form stable associations based on kinship between maternal relatives (matrilines) with a system of vocal dialects thought to reflect kinship relationships. We analyzed association patterns and acoustic similarity to study the social organization of killer whales in Avacha Gulf (Kamchatka, Russia), in the Northwest Pacific. The resident-type killer whales of Avacha Gulf formed temporally stable units that included maternal relatives with no dispersal observed. Acoustically, the killer whale community of Avacha Gulf was characterized by a system of dialects comparable to the communities of Northeast Pacific resident-type killer whales. Different units rarely associated with each other and these associations were nonrandom. Associations at different spatial levels did not always coincide with each other and with the patterns of acoustic similarity. Associations between units could change quickly irrespective of kinship relationships. The vocal dialect of a unit, which is more stable than the association patterns between units, might better reflect the overall kinship relationships. The stability and frequency of associations between units depended on the number of mature males in a unit, which could contribute to differences in the speed of change in vocal dialects and association patterns.     

Individual vocal production in a sperm whale (Physeter macrocephalus) social unit

The vocal repertoires of group‐living animals may communicate individual or group identity. Female and juvenile sperm whales live in long‐term social units that can be assigned to vocal clans based on the pattern of clicks in coda vocalizations. An unusual set of circumstances allowed us to record the vocalizations of photo‐identified individuals within a single social unit over a 41 d period. Using click interpulse intervals, we were able to assign codas to individuals and investigate coda production at the individual level within a social unit for the first time. Adult females in the unit vocalized at approximately equal rates. A calf and juvenile, both male, vocalized less often than the adult females. Repertoires were indistinguishable for all unit members apart from a mother and her calf, which possessed significantly different repertoires—even from one another. We suggest that similarity among the coda repertoires of most unit members indicates a function in advertising unit identity. In contrast, the distinctive repertoires of the calf and its mother may facilitate reunions between these whales. We hypothesize that sperm whales may be able to vary their vocal repertoires as their reproductive status alters the trade‐off between the benefits of individual and group identification.     

The benefits of social capital: close social bonds among female baboons enhance offspring survival

Sociality has evolved in many animal taxa, but primates are unusual because they establish highly differentiated bonds with other group members. Such bonds are particularly pronounced among females in species like baboons, with female philopatry and male dispersal. These relationships seem to confer a number of short-term benefits on females, and sociality enhances infant survival in some populations. However, the long-term consequences of social bonds among adult females have not been well established. Here we provide the first direct evidence that social relationships among female baboons convey fitness benefits. In a group of free-ranging baboons, Papio cynocephalus ursinus, the offspring of females who formed strong social bonds with other females lived significantly longer than the offspring of females who formed weaker social bonds. These survival benefits were independent of maternal dominance rank and number of kin and extended into offspring adulthood. In particular, females who formed stronger bonds with their mothers and adult daughters experienced higher offspring survival rates than females who formed weaker bonds. For females lacking mothers or adult daughters, offspring survival was closely linked to bonds between maternal sisters. These results parallel those from human studies, which show that greater social integration is generally associated with reduced mortality and better physical and mental health, particularly for women.     

Anthropogenic injuries disrupt social associations of common bottlenose dolphins (Tursiops truncatus) in Sarasota Bay,Florida

Social connectivity is important for measuring the fitness of common bottlenose dolphins (Tursiops truncatus). While interactions in fission-fusion societies vary between individuals, studies show that repeated interactions enhance reproduction and foraging success. Injuries that potentially remove an individual from its association network may disrupt these interactions. Using data from the long-term resident dolphin community in Sarasota Bay, Florida, we investigated how anthropogenic injuries affect the dolphins' social associations by examining the differences before and after injury to individuals. We examined group size, strength, eigenvector centrality, clustering coefficient, and number of triangles and analyzed whether the animal's sex, age class, type of injury, or human intervention affected these values. We found that while group size did not change, injured dolphins had fewer preferred associates (HWI > 0.14) and were found in more fluid groups immediately after injury, but started returning to normal association levels after 2 years. This initial decrease in connectivity was not related to the age, sex, type of injury, or intervention. Despite the fluidity in individual associations, the strongest bonds remained stable, those between mothers and calves and those between male alliance partners. These findings provide some of the first information relating injuries and social networks for animals.     

Adaptation of sperm whales to open-boat whalers: rapid social learning on a large scale?

Animals can mitigate human threats, but how do they do this, and how fast can they adapt? Hunting sperm whales was a major nineteenth century industry. Analysis of data from digitized logbooks of American whalers in the North Pacific found that the rate at which whalers succeeded in harpooning (‘striking’) sighted whales fell by about 58% over the first few years of exploitation in a region. This decline cannot be explained by the earliest whalers being more competent, as their strike rates outside the North Pacific, where whaling had a longer history, were not elevated. The initial killing of particularly vulnerable individuals would not have produced the observed rapid decline in strike rate. It appears that whales swiftly learned effective defensive behaviour. Sperm whales live in kin-based social units. Our models show that social learning, in which naive social units, when confronted by whalers, learned defensive measures from grouped social units with experience, could lead to the documented rapid decline in strike rate. This rapid, large-scale adoption of new behaviour enlarges our concept of the spatio-temporal dynamics of non-human culture.     

Whale killers: Prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia

Reports of killer whales (Orcinus orca) preying on large whales have been relatively rare, and the ecological significance of these attacks is controversial. Here we report on numerous observations of killer whales preying on neonate humpback whales (Megaptera novaeangliae) off Western Australia (WA) based on reports we compiled and our own observations. Attacking killer whales included at least 19 individuals from three stable social groupings in a highly connected local population; 22 separate attacks with known outcomes resulted in at least 14 (64%) kills of humpback calves. We satellite‐tagged an adult female killer whale and followed her group on the water for 20.3 h over six separate days. During that time, they attacked eight humpback calves, and from the seven known outcomes, at least three calves (43%) were killed. Overall, our observations suggest that humpback calves are a predictable, plentiful, and readily taken prey source for killer whales and scavenging sharks off WA for at least 5 mo/yr. Humpback “escorts” vigorously assisted mothers in protecting their calves from attacking killer whales (and a white shark, Carcharodon carcharias). This expands the purported role of escorts in humpback whale social interactions, although it is not clear how this behavior is adaptive for the escorts.     

Vocal clans in sperm whales (Physeter macrocephalus)

Cultural transmission may be a significant source of variation in the behaviour of whales and dolphins, especially as regards their vocal signals. We studied variation in the vocal output of 'codas' by sperm whale social groups. Codas are patterns of clicks used by female sperm whales in social circumstances. The coda repertoires of all known social units (n = 18, each consisting of about 11 females and immatures with long-term relationships) and 61 out of 64 groups (about two social units moving together for periods of days) that were recorded in the South Pacific and Caribbean between 1985 and 2000 can be reliably allocated into six acoustic 'clans', five in the Pacific and one in the Caribbean. Clans have ranges that span thousands of kilometres, are sympatric, contain many thousands of whales and most probably result from cultural transmission of vocal patterns. Units seem to form groups preferentially with other units of their own clan. We suggest that this is a rare example of sympatric cultural variation on an oceanic scale. Culture may thus be a more important determinant of sperm whale population structure than genes or geography, a finding that has major implications for our understanding of the species' behavioural and population biology.     

Resightings of humpback whales in Hawaiian waters over spans of 10–32 years: Site fidelity,sex ratios,calving rates,female demographics,and the dynamics of social and behavioral roles of individuals

From a database of approximately 5,000 Hawaiian humpback whales identified photographically between 1976 and 2010, we extracted 71 males and 39 females having resighting spans of 10 or more years, from first to most recent sighting. Findings included: (1) the male‐biased sex ratio was like that found in breeding grounds worldwide; (2) the mean span for males of 20.7 yr (maximum = 32 yr) did not differ significantly from the mean of 19.8 yr (maximum = 29 yr) for females, but males were seen in significantly more years during their spans than were females; (3) the mean number of females seen with and without calf across 11 three‐year intervals from 1977 to 2009 did not differ significantly; (4) the calving rate for the 39 females was 0.48 and seven females produced two to eight calves over spans of 22–26 yr; (5) females attracted significantly more escorts in years without calf than in years with calf; (6) individuals showed great diversity in the social units they occupied over their sighting spans, but with the most frequently observed unit for both sexes being the trio of mother, calf, and escort. Males were also observed frequently in competitive groups centered about a female without calf.     

A killer whale social network is vulnerable to targeted removals

Individuals play various roles in maintaining social integrity of mammalian populations. However, many models developed for managing wildlife resources assume that all individuals are equal. Killer whales are social animals that rely on relationships within and among family groups for survival. In the northeastern Pacific, fish-eating, 'resident' killer whale populations are composed of matrilines from which offspring do not disperse. We analysed the influence of various individuals' age, sex and matrilineal affiliation on their position in a social network. Here, we show that some matrilines appeared to play more central roles than others in the network. Furthermore, juvenile whales, especially females, appeared to play a central role in maintaining network cohesion. These two key findings were supported subsequently by simulating removal of different individuals. The network was robust to random removals; however, simulations that mimicked historic live-captures from the northeastern Pacific were likely to break the network graph into isolated groups. This finding raises concern regarding targeted takes, such as live-capture or drive fisheries, of matrilineal cetaceans.     

How does social behavior differ among sperm whale clans?

When individuals primarily associate with and learn from those who behave similarly, society and culture become closely tied. Sperm whales (Physeter macrocephalus) exhibit multilevel social structure, the levels of which are differentiated in part by characteristic cultural behaviors. Sperm whales are organized into sympatric clans, with distinctive vocal repertoires that are socially learned. Other behaviors, such as movement patterns and foraging, also differ among clans. Here we ask whether the clan partition also includes divergences in social behavior. Off the Galápagos Islands, members of two clans differed consistently in diving synchrony, heterogeneity, and temporal stability of social relationships. While number of associates (indicated by social unit, group, and cluster sizes) were similar between clans, Regular clan members dived more synchronously and had more homogeneous relationships than the Plus‐One clan members. Plus‐One social units had generally longer associations than those of the Regular clan. Differences in surface‐time coordination and quality of social relationships are likely byproducts of the clan segregation, which could affect alloparental care giving, therefore scaling up to differential calf survival rates between clans. This new dimension of behavioral divergence between sperm whale clans indicates that sympatric, socio‐cultural entities of nonhumans can also display characteristic social behavior.