Bottlenose dolphin (Tursiops truncatus) social structure in the Shannon Estuary,Ireland, is distinguished by age‐ and area‐related associations

Social and network analyses that incorporate information on individuals within a population enhance our knowledge of complex species. In this study, the social structure of wild bottlenose dolphins in the Shannon Estuary, Ireland, was analyzed by examining the dynamics of the whole population and then of individuals classed by sex, age, and area. One hundred and twenty‐one dolphins were identified during 522 sightings between 2012 and 2015. The mean half‐weight association index (HWI) of the population was 0.07 ± 0.02. The highest HWIs for individuals of known sex were for female‐male pairs. Mean within‐class HWI was significantly higher than mean between‐class HWI for both age and area classes. Ordinations and sociograms were used to visualize social networks. Permutation tests revealed nonrandom associations for the population overall and both between and within classes. Temporal analyses showed associations persisting for >1,000 d. The whole population's best fit model was for two levels of casual acquaintances. Movement analyses demonstrated the use of the inner estuary by only 25% of the population revealing a potential community division by area. The difference between mean HWI when socializing (0.09 ± 0.03) compared to foraging (0.06 ± 0.03) was significant. These results highlight the importance of localized research, reflecting the complexity found in bottlenose dolphin societies globally.     

Long‐term social structure of a resident community of Atlantic spotted dolphins,Stenella Frontalis,in the Bahamas 1991–2002

Long‐term social structure data on small delphinids is lacking for most species except the bottlenose dolphin. This study describes the long‐term social structure of one community of Atlantic spotted dolphins, Stenella frontalis, divided into three social clusters. Data from 12 yr were analyzed using SOCPROG 2.3. Coefficients of association (CoA) were calculated using the half‐weight index. The overall mean community CoA ranged from 0.09 to 0.12. Temporal analyses and mantel tests revealed significant differences between sex class associations due to high male‐male CoA (0.12–0.23) compared to female‐female and mixed sex CoA (0.08–0.10). Female associations were strongly influenced by reproductive status, calf care, and social familiarity, but not by age class. Male associations were strongly influenced by age, access to females, and alliance formation. Males showed two levels of alliance formation, long‐term first order pairs/trios (CoA 0.70–1.00) and shorter‐term second order alliances between two or more first order alliances (CoA 0.45–0.69), and a possible third level during interspecies interactions. Mating strategies, sex, and cluster formation shaped the social structure in this spotted dolphin community. Similar to many bottlenose dolphin studies, long‐term affiliations for spotted dolphins were correlated with age, sex, and reproductive status.     

Complex Social Structure of an Endangered Population of Bottlenose Dolphins (Tursiops truncatus) in the Aeolian Archipelago (Italy)

We investigated social structure and association patterns for a small population of Mediterranean bottlenose dolphins, Tursiops truncatus, inhabiting the Aeolian Archipelago (southern Italy). Specifically we evaluate the role of sex and age composition, residency patterns and interaction with trammel nets on this social organization. Association data for 23 regularly sighted individuals were obtained from summer photoidentification surveys collected from 2005–2012. Using a combined cluster and social network analysis approach, we found associations between dolphins were hierarchically structured, where two mixed-sex social units were subdivided into smaller temporarily dynamic groups. We found non-random and long-term preferred associations in the population; however, the degree of social cohesion, residence pattern and interaction with trammel nets differed considerably between the two social units. Six of eight females occurred in the more resident social unit-1; in addition, social unit-1 individuals had significantly stronger associations, higher preferred associates, lived in larger groups and occurred less frequently with trammel nets. Nine of eleven males were clustered in social unit-2 and five of these males, interacting with trammel nets, formed small groups and preferred associations. We propose that female and male groups associate in the study area during the breeding season and that some males choose to interact with reproductive females forming a distinct but interrelated social unit. Other males may be associating in a larger fission-fusion network, which consists of dolphins that appear to temporarily join the network from the coastal population. We cannot exclude that some males specialized in trammel net foraging, suggesting that this foraging technique may favor a solitary lifestyle. Large group sizes and high degree of social cohesion for females could be an indication of greater protection and more efficiency in detecting, deterring or repelling anthropogenic pressures. Most likely dolphins'' social organization depends on a combination of socio-ecological, demographic and anthropogenic factors.     

The structure of a bottlenose dolphin society is coupled to a unique foraging cooperation with artisanal fishermen

Diverse and localized foraging behaviours have been reported in isolated populations of many animal species around the world. In Laguna, southern Brazil, a subset of resident bottlenose dolphins (Tursiops truncatus) uses a foraging tactic involving cooperative interactions with local, beach-casting fishermen. We used individual photo-identification data to assess whether cooperative and non-cooperative dolphins were socially segregated. The social structure of the population was found to be a fission-fusion system with few non-random associations, typical for this species. However, association values were greater among cooperative dolphins than among non-cooperative dolphins or between dolphins from different foraging classes. Furthermore, the dolphin social network was divided into three modules, clustering individuals that shared or lacked the cooperative foraging tactic. Space-use patterns were not sufficient to explain this partitioning, indicating a behavioural factor. The segregation of dolphins using different foraging tactics could result from foraging behaviour driving social structure, while the closer association between dolphins engaged in the cooperation could facilitate the transmission and learning of this behavioural trait from conspecifics. This unique case of a dolphin-human interaction represents a valuable opportunity to explore hypotheses on the role of social learning in wild cetaceans.     

THE SOCIAL STRUCTURE OF BOTTLENOSE DOLPHINS, TURSIOPS TRUNCATUS, IN THE BAHAMAS

The social structure of coastal ecotype bottlenose dolphins, Tursiops truncatus , is largely unknown as they inhabit regions far from shore. This study reports on a community of bottlenose dolphins ≥ 27 km from Grand Bahama Island (May-September, 1993–2002). Resident and non-resident dolphins occurred in the area. Some dolphins traveled over 320 km between communities; others showed long- term site fidelity up to 17 yr. Average group size was 3–5, and was significantly larger with calves present and significantly smaller when traveling. The half-weight index was used to determine coefficients of association (COA) for individuals of known sex annually and for pooled years. Permutation tests revealed non-random associations and presence of preferred/avoided companions in all data sets. Annual COAs were low: female-female χ= 0.31, male-male χ= 0.30, and mixed-sex χ= 0.26. Mother-calf associations showed the highest values. Some males formed strong, long-term bonds. Female COAs fluctuated with reproductive status. Using pooled data, COAs were lower and the same basic trends were evident. However, strong associations seen in the annual data were not evident in pooled data. Bottlenose dolphins that inhabit offshore, shallow water show many of the same social structure characteristics as in well-studied coastal populations.     

Cannibalism as an interacting phenotype: precannibalistic aggression is influenced by social partners in the endangered Socorro Isopod (Thermosphaeroma thermophilum)

Models for the evolution of cannibalism highlight the importance of asymmetries between individuals in initiating cannibalistic attacks. Studies may include measures of body size but typically group individuals into size/age classes or compare populations. Such broad comparisons may obscure the details of interactions that ultimately determine how socially contingent characteristics evolve. We propose that understanding cannibalism is facilitated by using an interacting phenotypes perspective that includes the influences of the phenotype of a social partner on the behaviour of a focal individual and focuses on variation in individual pairwise interactions. We investigated how relative body size, a composite trait between a focal individual and its social partner, and the sex of the partners influenced precannibalistic aggression in the endangered Socorro isopod, Thermosphaeroma thermophilum. We also investigated whether differences in mating interest among males and females influenced cannibalism in mixed sex pairs. We studied these questions in three populations that differ markedly in range of body size and opportunities for interactions among individuals. We found that relative body size influences the probability of and latency to attack. We observed differences in the likelihood of and latency to attack based on both an individual's sex and the sex of its partner but found no evidence of sexual conflict. The instigation of precannibalistic aggression in these isopods is therefore a property of both an individual and its social partner. Our results suggest that interacting phenotype models would be improved by incorporating a new conditional ψ, which describes the strength of a social partner's influence on focal behaviour.     

Evaluating and minimising social stress in the care of captive bottlenose dolphins (Tursiops aduncus)

Stress can increase an organism’s susceptibility to disease. Thus, managing stress and its causes are important elements of captive care. Social factors such as changes in group dynamics, competition over resources, and unstable dominance hierarchies are potential stressors for highly social animals such as bottlenose dolphins (Tursiops sp). We present three case studies of mortality and illness in captive bottlenose dolphins and suggest that stress, resulting from social instability and ensuing aggressive interactions, is likely to have played a role in these health consequences. Stress is implicated by blood profiles, loss of appetite, and gastric ulcers, and social problems and instability are evident in the quantitative analysis of individual activity levels and association patterns. This is a unique study on marine mammals in that it demonstrates a correlation between quantitative behavioral indices and physiological measures of stress and health. Recommendations are made for the management of captive dolphins including regular quantitative assessment of behavior and associations and maintenance of appropriate groupings of age and sex classes. Behavioral records can be an important early indicator of health problems and may also serve as a useful tool for recognizing potentially stressful social changes and circumstances. Zoo Biol 21:5–26, 2002. © 2002 Wiley‐Liss, Inc.     

Reserve Size,Conspecific Density,and Translocation Success for Black Rhinoceros

Abstract: Fighting and accidental injury commonly cause black rhinoceros (rhino; Diceros bicornis) death after release. Smaller reserves and higher conspecific density after release (release density) might increase a rhino's encounter rate with hazards like fenced boundaries and conspecifics. We conducted a science-by-management experiment on the influence of reserve size and release density on rates of movement, association, and injury and death amongst 39 black rhinos during the first 100 days after their release into 4 Namibian and 8 South African reserves ranging in size from 670 ha to 45,000 ha. Association rates were negatively related to reserve size and positively correlated with release density. There was also a negative relationship between the proportion of the reserve traversed by individual rhinos and reserve size. In reserves ≥18,000 ha association rates were consistently zero but became elevated in reserves ≤11,500 ha and at release densities ≤9 km²/rhino. Daily displacement did not increase with increasing reserve size >8,500 ha but in smaller reserves daily displacements indicated higher encounter rates by released rhinos with fenced boundaries. Three rhinos received fight-related injuries requiring intervention and 2 of 4 deaths were fight-related. All injuries and 3 deaths occurred in reserves ≤11,500 ha. Model selection based on Akaike's second-order Information Criterion indicated that the parameter release density alone best explained mortality risk. Traditionally considered risk factors, rhino sex, age, and presence of resident conspecifics, were superseded by the risk posed by releases into smaller reserves. Reserves ≤11,500 ha and release densities ≤9 km²/rhino pose an increasing risk to rhino survivorship and so larger reserves and lower densities than these should be favored as release sites.     

The social organization of giraffes in Niger

The social organization of the last population of giraffes ( Giraffa camelopardalis ) in West Africa was studied between October 1996 and December 1997. Population size increased from 51 to 63 individuals during the study period. Groups were larger during the rainy season (mean group size 9.4) than during the dry season (mean group size 6; U = 4131; P < 0.01). Giraffes did not show strong preferential associations. Group types observed were similar to those expected on the assumption that associations are independent of sex and age. Inter-individual associations were low (mean simple ratio value = 0.1), with an individual associating with only half of its partners of the previous day. Young males interacted more frequently than other classes did. Agonistic interactions occurred mostly among males. Contact interactions (i.e. non-agonistic or sexual) occurred between both sexes and most age classes. Links between local communities, domestic animals, fauna and the environment suggest that environment and development cannot be dissociated in the sustainable exploitation of natural resources.     

Does body contact contribute towards repairing relationships?: The association between flipper-rubbing and aggressive behavior in captive bottlenose dolphins

Bottlenose dolphins exhibit complex social affiliations that may be shaped by interactions among individuals. Affiliative body contact among dolphins may repair deteriorated relationships or reduce tension within the group following aggressive interactions. We investigated the time-series association between one type of contact behavior (flipper-rubbing) and aggression by continuous observation of three captive bottlenose dolphins. For all three dolphin pairs, the elapsed time to aggressive events was significantly greater following flipper-rubbing. In two dolphin pairs comprised of a young male and an adult female, one-zero score of inter-opponent flipper-rubbing was higher for 10 min following aggressive bouts (post-AG periods) than for the same length of control (Ctrl) periods. For all three focal pairs, one-zero score of third-party rubbing was higher for post-AG than Ctrl periods. Neither the direction of rubbing nor the identity of the partner that approached prior to rubbing showed any significant tendencies. Flipper-rubbing may contribute to restore friendly relationships between former opponents or reduce conflicts in at least juvenile-adult female associations. Our results also give preliminary suggestions of the functions of third-party flipper-rubbing among bottlenose dolphins, including tension easing by the third party, or displacement as a result of aggressive interactions.     

Blood mercury concentrations in common bottlenose dolphins from the Indian River Lagoon,Florida: Patterns of social distribution

Social network analysis has been shown to be effective in studying the social structure of cetacean populations. Common bottlenose dolphins (Tursiops truncatus) inhabiting the Indian River Lagoon (IRL), Florida, have among the highest concentrations of total mercury (THg) in blood reported worldwide. The purpose of this study was to examine the relationship between THg concentrations in IRL dolphins and their social affiliations. Whole blood samples from 98 dolphins with photo‐identification sighting histories were collected between 2003–2007 and 2010–2012. Dolphins were categorized into approximate tertiles of low (mean 199.7 μg/L), medium (mean 366.8 μg/L), and high (mean 990.5 μg/L) THg exposure. Social associations between individual dolphins were defined by the proportion of sightings documented with another known individual. Social network measures of individuals and associations between dyads were examined to determine differences among THg categories. Strong social affiliations of individuals within the highest category of THg were found (P = 0.04), suggesting shared exposures among dolphins foraging in specific areas of the estuary. Network measures of strength and affinity were significantly higher in the highest exposure category. This report used social network analysis as a novel way to examine patterns of exposure to an environmental contaminant in a cetacean population.     

Replacement dolphins? Social restructuring of a resident pod of Atlantic bottlenose dolphins,Tursiops truncatus,after two major hurricanes

Environmental variations can influence the structure of ecological communities that in turn alter the grouping and association patterns of social communities. This study compares the social structure of bottlenose dolphins in the Bahamas before and after two major hurricanes. Approximately 30% of regularly seen individuals disappeared after the hurricanes, with an equal number of immigrants arriving afterwards. The primary goal of this study was to quantitatively describe social structure changes occurring after this large‐scale emigration (or death) and subsequent immigration of individuals using the social analysis program, SOCPROG 2.3. The pre‐hurricane results revealed one community with association patterns that were consistent with previous work on this population as well as other well‐documented populations. Post‐hurricane associations revealed that the community split into two distinct units, whose members associated highly within, but rarely between units. Association patterns varied between units. Immigrants assimilated well into the population, especially males. Over half of the post‐hurricane associations involved immigrants, the majority between residents and immigrants, and primarily involving immigrant males. The costs/benefits of choosing to associate with an immigrant individual differ between males and females and may have been the driving force for the changes in social structure that occurred.     

Brown spider monkeys (Ateles hybridus): a model for differentiating the role of social networks and physical contact on parasite transmission dynamics

Elevated risk of disease transmission is considered a major cost of sociality, although empirical evidence supporting this idea remains scant. Variation in spatial cohesion and the occurrence of social interactions may have profound implications for patterns of interindividual parasite transmission. We used a social network approach to shed light on the importance of different aspects of group-living (i.e. within-group associations versus physical contact) on patterns of parasitism in a neotropical primate, the brown spider monkey (Ateles hybridus), which exhibits a high degree of fission–fusion subgrouping. We used daily subgroup composition records to create a ‘proximity’ network, and built a separate ‘contact’ network using social interactions involving physical contact. In the proximity network, connectivity between individuals was homogeneous, whereas the contact network highlighted high between-individual variation in the extent to which animals had physical contact with others, which correlated with an individual''s age and sex. The gastrointestinal parasite species richness of highly connected individuals was greater than that of less connected individuals in the contact network, but not in the proximity network. Our findings suggest that among brown spider monkeys, physical contact impacts the spread of several common parasites and supports the idea that pathogen transmission is one cost associated with social contact.     

Does Innovation Success Influence Social Interactions? An Experimental Test in House Sparrows

In group‐living animals, individuals may benefit from the presence of an innovative group‐mate because new resources made available by innovators can be exploited, for example by scrounging or social learning. As a consequence, it may pay off to take the group‐mates' problem‐solving abilities into account in social interactions such as aggression or spatial association, for example because dominance over an innovative group‐mate can increase scrounging success, while spatial proximity may increase the chance of both direct exploitation and social learning. In this study, we tested whether the individuals' innovation success influences their social interactions with group‐mates in small captive flocks of house sparrows (Passer domesticus). First, we measured the birds' actual problem‐solving success in individual food‐extracting tasks. Then, we manipulated their apparent problem‐solving success in one task (by allowing or not allowing them to open a feeder repeatedly) while a new, unfamiliar group‐member (focal individual) had the opportunity to witness their performance. After this manipulation, we observed the frequency and intensity of aggression and the frequency of spatial associations between the focal individuals and their manipulated flock‐mates. Although flock‐mates behaved according to their treatments during manipulations, their apparent problem‐solving success did not affect significantly the focal individuals' agonistic behaviour or spatial associations. These results do not support that sparrows take flock‐mates' problem‐solving abilities into account during social interactions. However, focal individuals attacked those flock‐mates more frequently that had higher actual problem‐solving success (not witnessed directly by the focal individuals), although aggression intensity and spatial association by the focal birds were unrelated to the flock‐mates' actual success. If this association between flock‐mates' actual innovativeness and focal individuals' aggression is not due to confounding effects, it may imply that house sparrows can use more subtle cues to assess the group‐mates' problem‐solving ability than direct observation of their performance in simple foraging tasks.     

The building-up of social relationships: behavioural types, social networks and cooperative breeding in a cichlid

Consistent individual differences in behavioural types may not only cause variation in life-history decisions, but may also affect the choice of social partners and sociality in general. Here, we tested whether and how behavioural type influences the establishment of social ties using the cooperatively breeding cichlid, Neolamprologus pulcher. In a habitat saturation experiment with individuals pre-tested for behavioural type, we first analysed whether behavioural type affected the likelihood of settlement (i.e. social status), group sizes, and the types of dominant and subordinate individuals accepted as group members. Corrected for effects of body size and sex, the behavioural type did not affect settlement. However, bold dominant males only accepted smaller females, and grouped with bold subordinates, while shy dominant males accepted larger females than themselves, and grouped with shy subordinates. Second, we analysed the relationships between behavioural type and the aggressiveness or affiliation social network. Behavioural type significantly affected the number and quality of connections within the two networks. We show that behavioural types affect group composition, social networks and status achieved, in interaction with body size. Thus, the interactions within groups may depend not only on age, size and sex, but also on the behavioural type of the individuals involved.     

Association patterns and population dynamics of bottlenose dolphins in the Strait of Sicily (Central Mediterranean Sea): implication for management

An understanding of the population dynamics and social organization of cetaceans is essential to manage the influence of anthropogenic activities. In this study, the population size, site fidelity and social interactions of bottlenose dolphins in the Strait of Sicily (Italy) were investigated to provide recommendations for their conservation. Mark-recapture analysis was based on the encounter histories of 103 marked dolphins from 2004 to 2015. The POPAN formulation of the Jolly–Seber model in MARK software was used to estimate the size of the super-population. Site fidelity and social organization were estimated for individuals re-sighted ≥3 times. The estimated population size was 140 (SE = 15.75; 95% CI = 106–164). Dolphins had low site fidelity, and both adults and sub-adults move outside the study area. Females with calves used the area longer than other individuals. Based on our results, dolphins’ home range likely extended beyond the study area. The mean value of the Half-Weight Association Index was low and the preferred association was by casual acquaintance. However, we found a distinct aggregation of post-parturition females during the final 2 years of the study. Therefore, the pattern of association was apparently a response to an ecological requirement, which was the possibility to breed in high productivity waters. Whether these individuals are part of a larger pelagic population is unknown; however, we can conclude that the management of only coastal waters is insufficient for the conservation of dolphins in the Strait of Sicily.     

Bats are able to maintain long-term social relationships despite the high fission-fusion dynamics of their groups

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission-fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed long-term data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20,500 individual roosting observations over 5 years, we show that in two wild Bechstein's bat colonies with high fission-fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechstein's bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals.     

Analysis of Spix's disc-winged bat association patterns and roosting home ranges reveal a novel social structure among bats

Spix's disc-winged bats, Thyroptera tricolor, roost in young, rolled leaves of Heliconia or Calathea plants. In this paper, we examined how the combination of high habitat availability, low occupancy rate and short longevity of those roosts may affect the pattern of interactions among individuals in the population. We regularly censused a 5.69-ha study area in northeastern Costa Rica and examined patterns of association using mark-recapture data. Thyroptera tricolor formed behaviourally cohesive social groups of mixed sex, ranging in size from four to 14 individuals. Approximately 85% of dyads maintained associations over time periods of up to 100 days, and 40% of dyads maintained longer-term associations of at least 420 days across sex classes. Individuals within social groups did not always roost together, but they shared a small common roosting home range, which averaged just 0.19 ha. Members of different social groups rarely associated, although limited associations between members of select social groups in one subunit were observed. However, roosting home ranges of adjacent social groups often overlapped (up to 39% of home ranges, and up to 92% of the area of the smaller home range), and home range centres were situated less than 100 m apart. Thus, social groups rarely interacted but overlapped in space. The features of this social system are unique among bats and mammals in general, and point to groupings based on kinship or cooperation.     

Conflict and reconciliation behavior trends of the bottlenose dolphin (Tursiops truncatus)

Wild bottlenose dolphin (Tursiops truncatus) populations display societal structures characterized by numerous and frequent changes in group composition, complex social relationships, and high levels of cooperation, attributes also observed in human and nonhuman primate cultures. Maintaining social relationships under such elemental conditions can frequently create conflict—and the opportunity for reconciliation—among group members. The conflict and reconciliation behavior patterns of the chimpanzee (Pan troglodytes) have been studied extensively; trends are well‐documented in the wild and in captivity. Apparent cultural similarities have prompted several analogous studies on wild and captive bottlenose dolphins. This research attempted to corroborate previous efforts by analyzing the social behavior of seven captive bottlenose dolphins to determine the effects of sex and age on the frequency of conflict and reconciliation, and to investigate the incidence of consolatory behavior within the group. A total of 3,428 interactions involving focal animals, 414 of which were conflict episodes, were documented during 261 hr of videotaped observations. Although the sample size precluded meaningful statistical evaluation of the influence of sex on conflict and reconciliation, participant age was a significant determinant of conflict frequency. Conversely, age did not impact frequency of reconciliation, which only occurred after 18% of all conflict interactions. Little to no definitive evidence of consolation was apparent within the study group. While results partially support the findings of previous dolphin reconciliation research, extensive behavioral studies of wild populations should be conducted before generating broad comparisons between human terms and nonhuman behavioral interactions. Zoo Biol 29:567–585, 2010. © 2009 Wiley‐Liss, Inc.