A new level of complexity in the male alliance networks of Indian Ocean bottlenose dolphins (Tursiops sp.)

Male bottlenose dolphins in Shark Bay, Western Australia form two levels of alliances; two to three males cooperate to herd individual females and teams of greater than three males compete with other groups for females. Previous observation suggested two alliance tactics: small four to six member teams of relatives that formed stable pairs or trios and unrelated males in a large 14-member second-order alliance that had labile trio formation. Here, we present evidence for a third level of alliance formation, a continuum of second-order alliance sizes and no relationship between first-order alliance stability and second-order alliance size. These findings challenge the 'two alliance tactics' hypothesis and add to the evidence that Shark Bay male bottlenose dolphins engage in alliance formation that likely places considerable demands on their social cognition.     

Complex social structure, alliance stability and mating access in a bottlenose dolphin 'super-alliance'

Large brain size in mammals has been related to the number and complexity of social relationships, particularly social alliances within groups. The largest within-group male alliance known outside of humans is found in a social network (> 400) of Indian Ocean bottlenose dolphins (Tursiops aduncus) in Shark Bay Western Australia. Members of this dolphin 'super-alliance' cooperate against other alliances over access to females. Males within the super-alliance form temporary trios and occasionally pairs in order to consort with individual females. The frequent switching of alliance partners suggests that social relationships among males within the super-alliance might be relatively simple and based on an equivalence rule', thereby allowing dolphins to form large alliances without taxing their 'social intelligence'. The equivalence model predicts that the 14 males in the super-alliance should not exhibit differences in alliance stability or partner preferences. However, data from 100 consortships do not support the equivalence hypothesis. The 14 males exhibited striking differences in alliance stability and partner preferences suggesting that the super-alliance has a complex internal structure. Further, within the super-alliance, alliance stability correlates with consortship rate, suggesting that differentiated relationships within the super-alliance are based on competition for access to females.     

Alliance membership and kinship in wild male bottlenose dolphins (Tursiops aduncus) of southeastern Australia

Bottlenose dolphins are one of only a few mammalian taxa where the males are known to cooperate within their social group in order to maintain mating access to single females against other males. Male bonds in bottlenose dolphins have been hypothesized as evolving through kinship and associated inclusive fitness effects. In this study we tested whether individually identified male bottlenose dolphins preferentially associate and form alliances with kin in a small coastal resident population of southeastern Australia using a combination of behavioural data, genetic sexing, sequences of the mitochondrial DNA control region and nuclear microsatellite markers. Males generally associated significantly more often than expected with one to three other males, with whom they jointly herded females for mating. Associations and alliance membership were not associated with either maternal kinship or genetic relatedness. The majority of male pairs within alliances were randomly related, although high relatedness values were found between males of different alliances in the resident population. These findings indicate that mechanisms other than kin selection may be foremost in the development and maintenance of cooperation between male bottlenose dolphins.     

Synchrony,leadership, and association in male Indo-pacific bottlenose dolphins (Tursiops aduncus)

Male Indo-pacific bottlenose dolphins in Shark Bay, Western Australia, have converged with humans in the formation of nested male alliances and the use of synchrony in alliance behavior. Further, the strength of association among allied male dolphins varies and the stability of alliances correlates with the rate that males consort with estrus females (and is thus a possible indicator of dominance). To examine the possibility that synchrony reflects alliance association strength and dominance relationships, we analyzed videotapes from focal follows of two groups of males that reflect the range of alliance size and the strength of association between individuals in the population. We examined two variables: leadership during synchronous behaviors, based on which animal in a synchronously surfacing pair surfaced first, and the degree of synchrony, based on temporal differences in synchronous surfacing. We predicted that closer associates would exhibit a greater degree of synchrony and that one dolphin in a dyad would consistently lead. Contrary to our predictions, the degree of synchrony was inversely related to strength of association within alliances. This surprising result suggests that individuals with less secure bonds may strive more to achieve synchrony. We found no evidence of leadership during synchronous surfacing or between synchrony and other behavioral variables. Proximate mechanisms for synchronous behavior, such as entrainment and mutual motor imitation (“the mirror game” paradigm), may inhibit leadership in this context. Our results show that synchrony during surfacing is not a useful behavior to examine for dominance relationships in wild dolphins but it may be a useful tool to examine variation in alliance relationships.     

Male reproductive success increases with alliance size in Indo-Pacific bottlenose dolphins (Tursiops aduncus)

1. Determining the extent of variation in male mating strategies and reproductive success is necessary to understand the fitness benefits of social and cooperative behaviour. 2. This study assesses the reproductive success of male Indo-Pacific bottlenose dolphins in a small embayment population where different behavioural strategies of males have previously been identified. Parentage for 44 sampled calves was examined using 23 microsatellite loci and one mitochondrial DNA marker. Our candidate parent pool of 70 males and 64 females contained individuals sampled from both the embayment and adjacent coastal populations. 3. A moderate level of polygyny was detected in our sample. We assigned paternity of 23 calves to 12 males at the strict 95% confidence level and an additional nine calves to two males at the 80% confidence level. The majority (92%) of successful males were identified as residents to the embayment, and 46% of offspring were located within the same social group or community as their father. 4. Our results suggest that the size of alliances was the best predictor of reproductive success for males in this population, while the strength of association among allied males, alliance stability and male ranging patterns had little influence. In line with predictions for male alliances formed between unrelated individuals, we found that reproductive skew within alliances was not large. 5. Together, our genetic and behavioural analyses demonstrate that alliance formation between male dolphins is a successful strategy to enhance reproductive output.     

Male‐Biased Mate Competition in King Penguin Trio Parades

Darwin devised sexual selection theory to explain sexual dimorphisms. Further developments of the theory identified the operational sex‐ratio (OSR) as one of its cornerstones, and it was commonly admitted that an OSR biased toward one sex would lead to stronger selection pressures toward that sex. Recent theoretical developments have challenged this view and showed that the OSR alone does not determine the direction of sexual selection, more particularly in mutually ornamented species exhibiting high and similar parental investment by both sexes. These developments, however, focused on mutual intersexual selection, and little is known about intrasexual selection of both males and females in species exhibiting such characteristics. The first aim of our study was to test the relative involvement of males and females in same‐sex contest over mates in the king penguin, a species exhibiting mutual ornamentation of the sexes, high parental investment by both sexes, and a male‐biased OSR. We investigated the sex composition of trio parades, which are groups of three individuals that compete for mates during pair formation. We found that these trios consist of a female trailed by two fighting males in 19 of 20 cases; the 20th trio was all male. The second aim of our study was to investigate the existence of within‐sex differences in colour ornaments between individuals involved in such trios and individuals already paired. While limited sample sizes precluded detection of statistically significant differences between trios vs. pairs, reflectance measurements suggested that the beak spot of males in trios were more strongly ultraviolet than the beak spot of males in pairs. We concluded that intrasexual selection in our colony follows the typical pattern of mate competition observed in species in which sexual dimorphisms and OSR are male biased, and discussed the ultraviolet difference within the framework of the king penguins' colour perception.     

Speckled warblers break cooperative rules: absence of helping in a group-living member of the Pardalotidae

In the majority of cooperatively breeding bird species, subordinates help the dominant pair to provision the young, regardless of the origin of groups and relatedness between members. Within the family that includes speckled warblers, Chthonicola sagittata, there is considerable variation in social organization and parental behaviour, but the societies of other group-living species in this family are of kin, and subordinates provision the young of breeders. Speckled warblers differ in that offspring are never philopatric, and breeding groups comprise unrelated individuals. We used behavioural observations and DNA fingerprinting to examine genetic parentage and the mating system of the group-living speckled warbler in Canberra, Australia. Speckled warblers breed as pairs or in trios consisting of a female with two males. Alpha males are socially dominant to beta males and maintain dominance through aggression. Beta males never helped to feed nestlings or fledglings, even though they competed for copulations and in one case gained paternity of two of three young in a brood. The social system is therefore polyandrous but not cooperative. As well as competing for copulations with females on their own territories, beta males and single males (those that had lost their breeding partner) regularly undertook extraterritorial forays, most often when the neighbouring female was fertile. Failure by beta males to help even with broods containing their own offspring may be adaptive in this species and may result from increased opportunity for extrapair fertilizations or enhancement of prospects for future reproduction through the formation of alliances with females.     

‘O father: where art thou?’— Paternity assessment in an open fission–fusion society of wild bottlenose dolphins (Tursiops sp.) in Shark Bay,Western Australia

Sexually mature male bottlenose dolphins in Shark Bay cooperate by pursuing distinct alliance strategies to monopolize females in reproductive condition. We present the results of a comprehensive study in a wild cetacean population to test whether male alliance membership is a prerequisite for reproductive success. We compared two methods for inferring paternity: both calculate a likelihood ratio, called the paternity index, between two opposing hypotheses, but they differ in the way that significance is applied to the data. The first method, a Bayesian approach commonly used in human paternity testing, appeared to be overly conservative for our data set, but would be less susceptible to assumptions if a larger number of microsatellite loci had been used. Using the second approach, the computer program cervus 2.0, we successfully assigned 11 paternities to nine males, and 17 paternities to 14 out of 139 sexually mature males at 95% and 80% confidence levels, respectively. It appears that being a member of a bottlenose dolphin alliance is not a prerequisite for paternity: two paternities were obtained by juvenile males (one at the 95%, the other at the 80% confidence level), suggesting that young males without alliance partners pursue different mating tactics to adults. Likelihood analyses showed that these two juvenile males were significantly more likely to be the true father of the offspring than to be their half-sibling (P < 0.05). Using paternity data at an 80% confidence level, we could show that reproductive success was significantly skewed within at least some stable first-order alliances (P < 0.01). Interestingly, there is powerful evidence that one mating was incestuous, with one calf apparently fathered by its mother's father (P < 0.01). Our study suggests that the reproductive success of both allied males, and of nonallied juveniles, needs to be incorporated into an adaptive framework that seeks to explain alliance formation in male bottlenose dolphins.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. I. Partner Number and Diversity and Grooming Reciprocity

Allogrooming serves many social functions in primates. Grooming can help individuals to service social relationships generally, sometimes reciprocally, and may be particularly important in the development and maintenance of alliances. However, time constraints limit the number of partners with whom one individual can groom enough to maintain cooperative relationships. As a result, the size of its grooming network may reach an asymptote as the size of its group increases, and it may distribute its grooming less equally among potential partners. Chimpanzees live in multimale, fission-fusion communities; males are philopatric, and commonly associate and groom with each other. Males form within-community alliances that influence dominance rank and access to mates, and allies groom with each other regularly; males also cooperate in aggression between communities. The chimpanzee community at Ngogo, in Kibale National Park, Uganda, is unusually large and has more males than any other known community. Field data show that adult Ngogo males groomed far more with other adult males than with females or with adolescent males, in contrast to a previous report (Ghiglieri, 1984). Adolescent males groomed adults much more than the reverse; males groomed and were groomed by females about equally. Individual males groomed mostly with a small number of other males. On average, males at Ngogo had only slightly more male grooming partners overall and had the same number of important partners as those of males in a much smaller community in the Mahale National Park, Tanzania, and they distributed their grooming less equitably. These results fit those expected if limits on available grooming time cause males to have a loyalty problem as the number of potential grooming and alliance partners increases. Despite differences in the extent and equitability of their grooming networks, males at both Ngogo and Mahale showed reciprocity in grooming. Grooming reciprocity has been demonstrated for captive chimpanzee males, but the Ngogo findings are the first demonstrations of reciprocity in wild communities.     

Alliances and reproductive success in Camargue stallions

A study of a herd of Camargue horses Equus caballus, showed that while the majority of high-ranking stallions held single-male harems, some sons of low-ranking mares, being low ranking themselves, formed alliances that could last a lifetime. The two stallions were each other's closest associate and preferential grooming partner. Alliances were based on coalitions in which either both partners confronted an intruder synchronously or the dominant of the pair tended the female(s) while the subordinate simultaneously displayed towards the rival. Alliance partners were of similar age but were not more closely related to each other than to other stallions in the herd. Long-term paternity data revealed that subordinates sired close to a quarter of the foals born into the alliance group, and significantly more foals than low-ranking stallions in the herd adopting a 'sneak'-mating strategy. The dominant appeared to benefit from the presence of his subordinate partner. Fights occurred all year round, and the subordinate stallion of each alliance pair fought outside competitors more than twice as often as the dominant. Forming short-term alliances before defending mares on their own may enhance long-term reproductive success for both partners. Other benefits to both partners include higher survivorship of their foals and increased access to proven reproductive mares. These results suggest that the relationship between alliance partners is based on mutualism, but several conditions for reciprocity seem to be fulfilled: the benefit to the dominant (assistance in fights), and the benefit to the subordinate (access to reproduction), are both costly to the other partner and delayed in time. Copyright 1999 The Association for the Study of Animal Behaviour.     

Paternity exclusion by DNA fingerprinting,and mate guarding in the hooded seal Cystophora cristata

Hooded seal Cystophora cristata trios consist of an adult female, her pup, and an attending adult male. Using DNA fingerprinting, we excluded the possibility that the attending males within hooded seal trios were the fathers of the pups, proving that these hooded seals did not remain paired from one breeding season to the next. Behavioural observations of the trios after capture and release revealed that male hooded seals displace one another in attending nursing females. Mate guarding appears to be the preferred mating strategy available to male hooded seals given intense competition for females, a very brief nursing period, and oestrus occuring soon after weaning, but its effectiveness remains unclear.     

Chillingham Cattle: Dominance and Affinities and Access to Supplementary Food

A herd of cattle of natural sex ratio and age distribution, inhabiting a 134-ha park in northern England, was studied during supplementary feeding in 4 winters. Interactions could be summarised by conventional dominance hierarchies, more strictly linear and less stable among males than among females. Personal associations among individuals were not important, but affinities among social classes were, in determining the composition of feeding groups. Dominant males often fed in the same groups as dominant females. Dominant animals were less often seen to feed alone, implying that social dominance did not confer exclusive access to food. Cattle often fed in groups of two or three; certain combinations (notably those including two males, or one male and one female, or three calves) were stable, others unstable, notably combinations of females and calves, or of two or three females. This implies that females may defend resources more vigorously against other females than males do against other males.     

Multiple matings in the blue gourami, Trichogaster trichopterus (Pisces,Belontiidae)

The reproductive behaviour of the blue gourami, Trichogaster trichopterus, was examined in 15 bisexual pairs and 22 trios maintained in 75-litre aquaria for 23 days. During this time, 74 spawning episodes occurred among 26 different pairs and trios. Sixty-nine of these spawnings were directly observed. Both males and females were capable of mating repeatedly with the same or different partners with as little as 20 to 24 h separating successive spawnings. Females often mated with two males in the same afternoon, while only one male succeeded in mating with two females in one day. It is concluded that blue gouramis probably do not form pair bonds and that both sexes will mate with differet partners when the physical environment permits two or more males to maintain territories in the same aquarium.     

Whistle Convergence among Allied Male Bottlenose Dolphins (Delphinidae, Tursiops sp.)

Dolphins are adept at learning new vocalizations (whistles) throughout life, an ability thus far demonstrated in few nonhuman mammals. In dolphins, this ability is well documented in captivity but poorly studied in the wild, and little is known of its role in natural social behavior. This study documents the previously unknown phenomenon of whistle convergence among habituated free-living male bottlenose dolphins ( Tursiops sp.). Over a 4 yr study period, three male subjects formed an alliance, spending most of their time together and cooperating to herd females. Within individuals, whistle repertoires were more variable than expected based on previous studies, mostly performed with captive dolphins, but became less so during the course of the study. Among individuals, the distinctiveness of individual repertoires decreased such that the three males were virtually indistinguishable by the end of the study. Initially, some whistle types were shared. By the end of the study, the three males had formed a close alliance, and had all converged on one particular shared whistle form which they had rarely produced before forming the alliance. The results are discussed in terms of their implications for the prevailing 'signature whistle' hypothesis, as well as possible mechanisms and functional significance of whistle convergence among cooperating males.     

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.     

The use of low-frequency vocalizations in African elephant(Loxodonta africana) reproductive strategies

Fertility-advertisement calls in females are predicted to occur in nonmonogamous species where males and females are widely separated in space. In African elephants, low-frequency vocalizations have thus been suggested as a reproductive strategy used by fertile females to attract mates. This study examined the use of low-frequency vocalizations with respect to different phases of the estrous cycle in African elephants by simultaneously monitoring vocalizations, behavior, and hormonal profiles. Subjects were one male and six female African elephants housed at Disney's Animal Kingdom. No acoustically distinct vocalizations were restricted to the ovulatory follicular phase. However, overall rate of low-frequency vocalization as well as the rate of one acoustically distinct vocalization changed over the estrous cycle, with highest rates of calling related to the first period of follicular growth, or anovulatory follicular phase. Elevated rates of vocalization thus were not restricted to behavioral estrus and occurred much earlier in the estrous cycle than in most species that produce fertility-advertisement calls. Both herd composition and elephant identity also affected rates of vocalization. Vocalizations therefore may not be reliable signals of actual fertility. However, the increase in vocalizations in advance of estrus may attract males to the herd prior to ovulation, facilitating both male-male competition and female choice. Once present in the herd, males may then switch strategies to use more reliable chemical and visual cues to detect ovulating females.     

Weaponry,size, and sex ratio affect spatial distribution within small and large groups of the maritime earwig (Anisolabis maritima)

Dispersion patterns within a group can reveal important aspects about social interactions and sexual selection within a species. We examined the distribution patterns of the maritime earwig (Anisolabis maritima), an insect well suited for studies of aggression, sociality, and sexual selection since both sexes live in close proximity and possess weaponry in the form of sexually dimorphic pincers. To examine intra‐ and intersexual interactions within small groups, we conducted trials with three earwigs with limited access to shelters. In single‐sex trios, we found that both males and females exhibited strong size‐based intrasexual aggression, as larger individuals were less likely to be excluded from shelters; however, males were more likely to cohabitate than females. In mixed‐sex trios, we found that both males and females preferred smaller opposite‐sex partners, and cohabitation patterns indicate that both sex‐ and size‐based differences in aggression can influence overall spatial distribution. We also examined larger single‐sex and mixed‐sex groups of 18 earwigs to determine whether they had random, uniform, or clumped distributions. Similar to previous field observations, males tended to form aggregations, whereas females were distributed uniformly, a pattern indicative of territoriality. Mixed‐sex groups, on the other hand, were uniform during nocturnal periods of high activity but then become clumped after settling into more stable daytime positions. Overall, our results suggest that females have high levels of aggression regardless of the social context, whereas males alter their aggressive behavior in the presence of females.     

Genetic evidence for a family structure in stable social aggregations of the Australian lizard Egernia stokesii

In this study we used data from six unlinked microsatellite loci to examine stable aggregations of Egernia stokesii, from a population in the southern Flinders Ranges of South Australia. We show that these aggregations are comprised of breeding partners, their offspring from two or more cohorts, and related adults, providing the first genetic evidence of a family structure in any lizard species. Despite this high level of relatedness within aggregations, most breeding pairs were unrelated and partners were less closely related to each other than they were to other potential within-group partners. Where individuals dispersed, both sexes usually moved to social groups close to their natal group. Although both sexes showed natal philopatry, there was some evidence that females in groups were more related than males in groups. These data suggest that an active choice of unrelated partners and male-biased dispersal may be the mechanisms used by E. stokesii to avoid inbreeding within groups.     

Dolphin social intelligence: complex alliance relationships in bottlenose dolphins and a consideration of selective environments for extreme brain size evolution in mammals

Bottlenose dolphins in Shark Bay, Australia, live in a large, unbounded society with a fission-fusion grouping pattern. Potential cognitive demands include the need to develop social strategies involving the recognition of a large number of individuals and their relationships with others. Patterns of alliance affiliation among males may be more complex than are currently known for any non-human, with individuals participating in 2-3 levels of shifting alliances. Males mediate alliance relationships with gentle contact behaviours such as petting, but synchrony also plays an important role in affiliative interactions. In general, selection for social intelligence in the context of shifting alliances will depend on the extent to which there are strategic options and risk. Extreme brain size evolution may have occurred more than once in the toothed whales, reaching peaks in the dolphin family and the sperm whale. All three 'peaks' of large brain size evolution in mammals (odontocetes, humans and elephants) shared a common selective environment: extreme mutual dependence based on external threats from predators or conspecific groups. In this context, social competition, and consequently selection for greater cognitive abilities and large brain size, was intense.     

Social Organization of Lepilemur ruficaudatus

We characterize the spatial organization of red-tailed sportive lemurs (Lepilemur ruficaudatus) as a key aspect of their social organization and social system. Sportive lemurs are small (<1000 g), nocturnal and folivorous primates endemic to Madagascar. We studied a population of 57 individually-marked individuals in Kirindy Forest, western Madagascar, between 1995 and 2001. We radio-tracked 20 males and 26 females of the marked population to obtain detailed information on the size and location of their home ranges. Census data and morphometric measurements provided complementary data sets. Males and females occupied small (<1 ha) home-ranges. Long-term records from 9 individuals revealed home-range stability over several years. In 4 cases home ranges overlapped extensively with that of one member of the opposite sex; in 2 cases, a spatial association of 1 male and 2 females occurred. However, home ranges overlapped very little with neighboring individuals of both sexes. During the study period, spatially associated individuals used on average 5.6 sleeping trees within 117 days, but they spent on average only about every fourth night together. The data suggest that home ranges in red-tailed sportive lemurs are exclusively used by pairs or trios and that the modal social organization of red-tailed sportive lemurs is pair-living.