Social play in wild brown bears of varying age-sex class

Social play behaviour is a well-described phenomenon, almost ubiquitous among mammals. Despite its prevalence, social play takes several forms and may vary in function across species. For solitary species, the function of play outside of the family group remains unclear. Here, we describe the motor patterns of play among non-littermate wild brown bears Ursus arctos of different age-sex class. Play was documented during a time of abundant food availability in three different scenarios: play among non-littermate subadults, play among non-littermate cubs, and play among a ‘group’ of bears of different age and sex class. We used a previously described behavioural ethogram to recognise play. Play followed typical motor patterns and postures expressed by bears during play-fighting: relaxed face, puckered-lip, ears partially flattened to crescent, wrestling, jaw gaping, play-biting, paw-swatting, and lunging. No vocalisations were conducted during play bouts. Older bears displayed ‘self-handicapping’ and ‘role-reversal’ in the play postures they selected when playing with younger bears, suggesting that tactics vary according to age class and dominance ranking. Playing likely allows for the evaluation of conspecifics in a non-aggressive way during times of reduced competition and could also relieve stress in complex social situations.     

Behavioural differences between male and female replacement reproductives in Kalotermes flavicollis (Isoptera, Kalotermitidae)

Summary: An ethogram type study was performed on representative colonies of Kalotermes flavicollis to investigate the behavioural repertoires of replacement king and queen. Our observations suggested a sex-based behavioural specialisation in the two reproductives, affecting colony organisation and equilibrium. Interesting differences were detected in overall activity of the two reproductives, with the male showing the greatest frequency of movements in the nest, vibratory acts, and the highest rate and range of social contacts. In addition to his importance for periodical insemination of the queen, the king might also play a basic role in the social structure and dynamic development of the colony, as a "mediator" in social interactions between the queen and the other colony members.     

Playing it cool: Characterizing social play,bout termination,and candidate play signals of juvenile and infant Tibetan macaques(Macaca thibetana)

Play behaviors and signals during playful interactions with juvenile conspecifics are important for both the social and cognitive development of young animals.The social organization of a species can also influence juvenile social play. We examined the relationships among play behaviors, candidate play signals, and play bout termination in Tibetan macaques(Macaca thibetana) during juvenile and infant social play to characterize the species play style. As Tibetan macaques are despotic and live in groups with strict linear dominance hierarchies and infrequent reconciliation, we predicted that play would be at risk of misinterpretation by both the individuals engaged in the play bout and by those watching, possibly leading to injury of the players. Animals living in such societies might need to frequently and clearly signal playful intent to play partners and other group members to avoid aggressive outcomes. We gathered video data on 21 individually-identified juvenile and infant macaques(one month to five years of age)from the Valley of the Wild Monkeys, Mt. Huangshan,China. We used all-occurrence sampling to record play behaviors and candidate play signals based on an ethogram. We predicted that play groups would use multiple candidate play signals in a variety of contexts and in association with the number of audience members in proximity to the players and play bout length. In the 283 playful interactions we scored,juvenile and infant macaques used multiple body and facial candidate play signals. Our data showed that juvenile and infant Tibetan macaques use a versatile repertoire of play behaviors and signals to sustain play.     

Relationship between Serum Testosterone, Dominance and Mating Success in Père David's Deer Stags

We conducted an experiment in the Beijing Milu Park to study the social behavior of male Père David's deer, and related social behavior to social position and serum testosterone level of the stags during rut. We classified the stags into three rank classes according to their rutting behavior: ‘harem master’, ‘challenger’ and ‘bachelor’. We monitored the behaviors of four ‘harem masters’, five ‘challengers’ and five ‘bachelors’, and analyzed serum testosterone levels in blood samples of those 14 stags using radioimmunoassay. We defined the effectiveness value, E = A/T, to assess the effectiveness of herding or mating attempts made by stags (‘T’ represents the frequency of herding or mating attempts made by a stag and ‘A’ represents the frequency of herding or mating attempts accepted by hinds). We found that: (1) the ‘harem masters’ and the ‘challengers’ displayed more frequent rut and locomotive behaviors but fewer ingestion behaviors than the ‘bachelors’; (2) serum testosterone levels in the ‘harem masters’ and the ‘challengers’ were higher than that in the ‘bachelors’; (3) effectiveness value of herding attempts differed significantly between the three types of stags, being highest in the ‘harem masters’ and the lowest in the ‘bachelors’; and (4) effectiveness value of mating attempts was significantly greater for the ‘harem masters’ than for the ‘challengers’. We conclude that: (1) reproductive behavior of the Père David's deer stags is strongly associated with social rank; (2) social roles of Père David's deer stags during the rut are related to the testosterone secretion; and (3) rank class affects the mating opportunity of the stags.     

Demographic and social constraints on male chimpanzee behaviour

Male chimpanzees, Pan troglodytes, are well known for affiliating and cooperating in a variety of behavioural contexts. Prior field research indicates that maternal kinship does not affect patterns of affiliation and cooperation by males in the same social group. Two questions remain unclear from this finding. First, why do male chimpanzees not bias their behaviour towards maternal kin? Second, what factors account for who affiliates and cooperates with whom? We conducted behavioural observations of an unusually large community of chimpanzees at Ngogo, Kibale National Park, Uganda, to test the hypothesis that demographic constraints limit the number of maternal kin with whom male chimpanzees can cooperate, and thereby lead them to form selective bonds with nonkin of similar age and status. Results indicated that male age and rank are significantly associated with four measures of social behaviour. Members of the same age class and individuals close in rank were more likely to affiliate and cooperate than males that belonged to different age and rank classes. Additional analyses replicate earlier findings and show that males who affiliated and cooperated were not closely related through the maternal line, as assayed by mtDNA haplotype sharing. These results add to our growing understanding of the important role demographic and social constraints play in animal behaviour.     

‘Stick with your own kind,or hang with the locals?’ Implications of shoaling strategy for tropical reef fish on a range‐expansion frontline

Range shifts of tropical marine species to temperate latitudes are predicted to increase as a consequence of climate change. To date, the research focus on climate‐mediated range shifts has been predominately dealt with the physiological capacity of tropical species to cope with the thermal challenges imposed by temperate latitudes. Behavioural traits of individuals in the novel temperate environment have not previously been investigated, however, they are also likely to play a key role in determining the establishment success of individual species at the range‐expansion forefront. The aim of this study was to investigate the effect of shoaling strategy on the performance of juvenile tropical reef fishes that recruit annually to temperate waters off the south east coast of Australia. Specifically, we compared body‐size distributions and the seasonal decline in abundance through time of juvenile tropical fishes that shoaled with native temperate species (‘mixed’ shoals) to those that shoaled only with conspecifics (as would be the case in their tropical range). We found that shoaling with temperate native species benefitted juvenile tropical reef fishes, with individuals in ‘mixed’ shoals attaining larger body‐sizes over the season than those in ‘tropical‐only’ shoals. This benefit in terms of population body‐size distributions was accompanied by greater social cohesion of ‘mixed’ shoals across the season. Our results highlight the impact that sociality and behavioural plasticity are likely to play in determining the impact on native fish communities of climate‐induced range expansion of coral reef fishes.     

Rank relations of juvenile and subadult natal males of Barbary macaques (Macaca sylvanus) at Affenberg Salem

Rank relations of more than 100 juvenile and subadult natal Barbary macaque males were analyzed. Hierarchical relations among individuals of the same age were established early during the first year of life. With few exceptions concerning infants from very high-ranking genealogies, males dominated female peers regardless of maternal rank. Males started to outrank females from older cohorts during the second year of life and completed the process of rank reversal with adult females at 5-6 years of age. An age-graded dominance pattern existed among males from different birth cohorts. Only 3 rank reversals between males from different cohorts were observed. Rank reversals among males of the same birth cohort occurred more frequently. Rank position of a male among his male peers was influenced by birth order, by maternal rank, and by the presence of juvenile brothers. Most males without juvenile brothers had low positions, regardless of maternal rank. Males born late in the birth season were also low-ranking, even when juvenile brothers were present. There was no cohort where ranking among males was determined by maternal rank alone, as is the case in rhesus monkeys and Japanese macaques. Adult/subadult male carriers had no noticeable effect on rank positions of 'their' infants. It is suggested that a weaker influence of Barbary macaque mothers on rank of their sons is related to very early integration of male infants in male social/play groups.     

Play Behavior in Juvenile Golden Lion Tamarins (Callitrichidae: Primates): Organization in Relation to Costs

Play is a complex behavior that is widespread among vertebrates. Despite the potential benefits for the development of social, cognitive and motor skills, play behavior has costs: energy expenditure, and the risks of injury and predation. As the fitness benefits of play are presumably to be gained as adults, we expect young animals to minimize its immediate costs. Here, we describe play behavior in golden lion tamarins (GLTs) (Leontopithecus rosalia) and test cost-minimizing hypotheses. We collected data on play behavior and adult vigilance in nine groups (four wild and five supplemented, the latter deriving from a reintroduction) of wild-living lion tamarins (16 infants) in Brazil. Results showed that play occupied 3.8% of the activity budget, occurred during the hottest times of the day, and was characterized by being mainly social, involving partners of different ages, and sometimes different species. We found no differences in playing time between wild and supplemented groups. The lion tamarins played less in ‘dangerous’ substrates (canopy branches and forest floor), and more in ‘safe’ substrates (large branches and vine tangles), and favored playing in the center of the group over the periphery. The reproductive animals were vigilant during 78% of the time the infants were playing, and were mostly oriented away from the playing animals. The observed organization of play behavior suggests that juvenile GLTs minimize the risk of predation and accidental injuries, but do not seem to minimize energy expenditure and the risks of social injuries. The social-cognitive benefits acquired from experience of older partners during play may overcome the costs of the latter behavior, and because little time was spent in play, the energetic costs may be inconsequential.     

Social rank and habitat use in willow tit groups

One view depiets dominance and subordination as roles with equal fitness played as parts of a mixed behavioural strategy, but subordination can also be maintained as a transient strategy of ‘hopeful dominants’. Resource access in groups of non-kin willow tits, Parus montanus, was related to social rank. Dominants excluded subordinates from the preferred microhabitat. Fitness in terms of survival was correlated with access to the preferred foraging sites, indicating that a mixed-strategy explanation is inadequate. Social rank was linked to age, and first-year subordinates behaved as predicted for hopeful dominants. They were not ‘herded’ by dominant group mates because subordinates actively maintained contact with dominants. One possible reason for subordinates joining winter groups might be the opportunity to reduce dominants' pay-off from aggression.     

Does the environmental background (intensive v. outdoor systems) influence the behaviour of piglets at weaning?

Under intensive pig husbandry, outdoor systems offer a more complex physical and social environment compared with indoor systems (farrowing sheds). As the rearing environment affects behavioural development, it can, therefore, influence behavioural responses of pigs to stressful environments in later stages of production. We tested how the rearing environment influenced behavioural responses to a novel arena test in piglets on the day that they were weaned and mixed into large groups. We recorded video footage and compared the behavioural responses of 30 outdoor-raised and 30 farrowing shed-raised piglets tested in an experimental arena and sequentially exposed to four challenges (each for 5 min) on the day of weaning. Quantitative and qualitative behavioural measures were recorded using time budgets and scoring demeanour or ‘qualitative behavioural expression’ (using Qualitative Behavioural Assessment (QBA)). When held in isolation (challenge 1), both groups were scored as more ‘scared/worried’, while outdoor-raised piglets spent more time eating and jumping against the arena walls. Both groups interacted with a plastic ball (challenge 2: exposure to a novel object) during which they were scored as more ‘playful/curious’ than other challenges. When a food bowl was introduced (challenge 3), farrowing shed-raised piglets were more interested in playing with the food bowl itself, whereas outdoor-raised piglets spent more time eating the feed. Finally, there were no significant differences in social behaviour (challenge 4: introduction of another piglet) between the two groups in terms of the latency to contact each other, amount of time recorded engaged in aggressive/non-aggressive social interactions or QBA scores. Although piglets spent 30% of their time interacting with the other piglet, and half of this time (47%) was engaged in negative interactions (pushing, biting), the levels of aggression were not different between the two groups. Overall, outdoor-raised piglets ate more and were scored as more ‘calm/passive’, whereas farrowing shed-raised piglets spent more time investigating their environment and were scored as more ‘playful/inquisitive’. In conclusion, we did not find differences in behaviour between outdoor-raised and farrowing shed-raised piglets that would highlight welfare issues. The differences found in this study may reflect conflicting affective states, with responses to confinement, neophobia and motivation for exploration evident.     

Social play does not reduce aggression in wild meerkats

Of the numerous hypotheses advanced to explain the adaptive significance of play, several assert that social play increases social harmony, cementing alliances and reducing aggression between group members or littermates. These hypotheses are frequently cited, but their validity remains unknown. We examined the relation between social play and aggression in juvenile meerkats, Suricata suricatta, living in a wild population in the southern Kalahari Desert. We tested the hypothesis that social play reduces aggression, by examining rates of play, play partner choices, the structure of social play and rates of aggressive interactions during foraging. We found no relation between frequency of play and level of aggression, either between individuals or during the course of development. Pups that played together frequently were just as aggressive towards one another as pairs of pups that played infrequently, and play interactions had no subsequent effect on the likelihood of aggression. In contrast, aggressive interactions during foraging inhibited the subsequent likelihood of play, and high levels of aggression during foraging changed the structure of social play, with victimized pups avoiding play wrestling. We conclude that social play does not reduce aggression in young meerkats. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Boldness and Information Use in Three‐Spined Sticklebacks

In foraging groups, individuals may utilise information from their social environment to aid decision making when choosing where to search for food. Little work has looked at the costs or benefits of behavioural differences, such as consistent individual variation in boldness, with respect to learning ability. Here, we investigate the response of three‐spined stickleback (Gasterosteus aculeatus) to ‘social cues’, ‘local enhancement’ and ‘public information’ during foraging tasks. Our results confirm previous work suggesting that this species responds to social cues and local enhancement but not public information. Variation in boldness did not affect the use of different types of information. However, time taken to make a choice and reach a patch varied between fish with different levels of boldness. Contrary to expectation, shy fish were the more variable individuals, having a greater range of reaction times when responding to the tasks. This suggests that individual behavioural differences still play a role when utilising information obtained from the environment and may influence the relative benefits that could result in different contexts.     

Winter survival and breeding success of dominant and subordinate Willow Tits Parus montanus

We studied the effect of winter rank on survival rate and reproductive success in Willow Tits Parus montanus, a resident passerine living in dominance-structured flocks during the nonbreeding season, in 6 years. Winter survival was dependent on both the birds' age and rank. Adults survived better than first-year birds, and within first-year males, dominants survived better than subordinates. In other sex and age classes, rank did not contribute to survival. Although first-year males were in excess among nonbreeders, no connection existed between breeding status and rank. Female rank did not explain the variation in the start of laying, clutch size, number of fledglings or recruit production. We conclude that social status in Willow Tits affects individual fitness mainly through rank-dependent survival. Acquiring a high rank position seems to be most important for first-year birds, especially first-year males.     

Agonistic Interactions of Juvenile Savanna Baboons

19 juvenile members of known genealogies in two wild baboon groups were studied over a 16-month period to compare the ontogeny of agonistic experience and dominance relations for males and females. Juveniles of all age-sex classes were disproportionately likely to receive aggression from and submit to adult males per unit of time spent in proximity. This pattern intensified with increasing juvenile age. With age, juvenile females more often submitted to unrelated adult females from higher-ranking families, whereas this was not true for juvenile males. All juveniles received aggression from older group members more often during feeding than was expected by chance. High rates of agonistic interaction with unrelated adult females accounted for old juvenile females (3–5.5 years-old) interacting agonistically more frequently than male age peers and young juveniles of either sex (1–2.5 years-old). Adult females were also more aggressive toward females among young juveniles, suggesting that adult females target females among juveniles for aggression and resistance to rank reversal. Within juvenile age groups, males dominated all females and all younger males, irrespective of maternal dominance status. Dominance relations among female age-peers were generally isomorphic with relations among their mothers. No juvenile targeted any older male for rank reversal. Males targeted all older females, whereas females typically targeted only older females from families lower-ranking than their own. The strong sexual dimorphism in adult body size in baboons may explain why juvenile males' dominance relations with peers and adult females are not structured along lines of family membership as is true for the less dimorphic macaques. Acquisition of higher agonistic status probably allows juveniles of both sexes to increase their success in within-group feeding competition during late stages of juvenility, which, in turn, could affect important life-history traits such as age at menarche and adult body size.     

Playmate relationships among individuals of the Japanese monkey troop in arashiyama

Observations of play behavior were made on a troop of Japanese monkeys for five months. The troop consisted of 125 animals during the study period. Only 104 animals were observed playing with the troop members while the other 21 animals were never observed playing with other individuals. Two-member play was the most frequent. On the average, a monkey played with 20.7 individuals. A total of 6,068 play bouts were observed. The frequency of play appeared to be affected by age, sex, and degree of relatedness. One-year-old infant males played most with other members and the frequency of play decreased with age. Between monkeys whose disparity of age was less than two years, 5,763 bouts (95.0% of the total) were observed. Moreover, among sameaged monkeys who comprised 10.6% of the possible pair combinations, 2,739 play bouts (45.1%) were observed. Juvenile males played with same-sexed peers more than with opposite-sexed peers, whereas older juvenile females appeared to play with infants of both sexes. Individuals who were related and similarly-ranked tended to play together. There was no apparent preference for animals to play with the offspring of the highest-ranking female. Dominance rank of infnats and juveniles was primarily affected by rank of their mothers and to a lesser extent by play partners. Dominance rank of older juvenile males is more likely to be affected by play partners than females. It may be a critical time for males when they leave their natal troop and join a new troop. The timing of troop shifting by males seemed to be affected by the presence or absence of play-mates. For male Japanese monkeys, play is very important in developing social bonds. Play may act to perpetuate social bonds, enhance the chance of survival, and may contribute to their future reproductive success.     

Social interactions of juvenile Collared peccaries, Tayassu tajacu (Mammalia: Artiodactyla)

Is there a predictable relationship between species-typical behavioural development and social organization in mammals? Earlier work on canids and rodents suggested that adult sociality is associated with a developmental schedule in which juveniles showed, among themselves: (1) frequent social play; (2) frequent amicable contact; (3) frequent olfactory contact; (4) a delay in the emergence of agonistic behaviour. In the highly social Collared peccary (Tayassu tajacu) the behaviour of juveniles conformed only partially to this scheme. Juveniles engaged in social play with each other at an early age, but showed no other observable amicable or olfactory contact until eight to twelve weeks of age and showed no delay in the emergence of agonistic behaviour among themselves. Adults, in contrast, were conspicuously tolerant of juveniles, and 86% of juvenile social interactions in the first year took place with adults.     

Juvenile social play and yearling behavior and reproductive success in female Belding’s ground squirrels

Benefits of play behavior have been described for individuals during the juvenile period; however, it is less clear whether benefits of juvenile play commonly extend beyond the period of juvenile development. I evaluated possible associations between juvenile social play and yearling maternal territorial behavior and reproductive success in female Belding’s ground squirrels (Urocitellus beldingi). Higher rates of juvenile social play were associated with greater intensity of territorial aggressive and vigilant behavior during gestation but not later phases of the yearling reproductive cycle, suggesting that both juvenile play and later experience may influence maternal territorial behavior in yearling females. Juvenile social play was also a reliable predictor of whether a yearling female successfully weaned a litter, raising the possibility that juvenile play behavior influences yearling reproductive success via effects on maternal territorial behavior in female U. beldingi. Other factors were also related to reproductive success in yearling females. Body mass at the beginning of the yearling reproductive period was not correlated with juvenile social play but was associated with the likelihood of successfully weaning a litter. Yearling females whose mother was still alive were more likely to wean a litter than those whose mother was not alive. Mothers and yearling daughters tended to have maternal territories near each other, and mothers with a surviving yearling daughter expressed aggressive behavior at elevated rates and directed aggression toward intruders on the daughter’s territory. Overall, results of the study suggest that juvenile social play is among the variables that influence maternal territorial behavior and reproductive success in yearling female U. beldingi, and are consistent with the idea that juvenile play has benefits beyond the juvenile period for female U. beldingi.     

Unexplained sexual segregation in polygamous ungulates: a defense of an ontogenetic approach

Two processes are generally mixed when considering ungulate sexual segregation: the social segregation which appears to be a rule in polygamous ungulate species and the spatial segregation which is facultative. Early in life, there is evidence that males and females exhibit different levels of activity and patterns of interaction which could lead to behavioural incompatibility. It is proposed that juvenile females, differently motivated than males to interact socially, may soon avoid the pseudo-sexual and agonistic male behavioural components. as body and behavioural dimorphism increase with age, the behavioural incompatibility would be intensified. Since adults often interact agonistically with or avoid them, juveniles will attend to interact with same-sex peers of similar status. Because social experiences are occurring since early stages of development, i.e., during sensitive periods, a social auto-segregation by sex/age classes is expected. Then population would be structured in coherent socio-spatial sets of compatible individuals, sharing similar constraints and properties. Around the birth period, females become asocial promoting a temporal social segregation. Costs linked to gestation and lactation may make females more dependent on water and rich food resources. Female with offspring are also likely to select areas where the risks of predation are reduced, sometimes at detriment to energy intake, while other animals exhibit opportunistic feeding behaviour, resulting in seasonally ecological segregation. Body growth channels a lot of energy for males which are expected to use the best feeding areas. However, indirect competition for the same resources may result in a spatial segregation between the sexes. Sex- and age-specific activity budgets are suspected to favour social and spatial segregations. The social segregation among males and females use exclusive ranges. Whether spatial segregation results in ecological segregation would however depend on habitat heterogeneity, population density, local ecological conditions and the biological cycle periods. It is suggested that spatial segregation will occur not only between the sexes but also among males. If populations are structured in coherent socio-spatial units, spatial segregation among individuals of age and sex classes may result from indirect interactions between sets of individuals (of same age and sex class), that may be translated as an indirect competition between individuals. Studies on ethogenesis are considered as fundamental as they allow to know on what basis social and spatial segregations are set up. In particular, studies on behavioural development could highlight how social segregation between adults may result from a process that originates much earlier in the life of animals. By considering the role of behaviour, we defend the idea that social and spatial segregations may be logically distinguished.     

Social Dominance among Male Meadow Voles is Inversely Related to Reproductive Success

Intrasexual selection can occur through direct aggressive interactions between males for access to females. We tested the relationship between social dominance and male reproductive success among meadow voles, Microtus pennsylvanicus. Dominance ranks of wild‐caught males were determined using neutral arena trials, with the winner of two of three trials considered dominant. These males were then released into field enclosures and allowed to visit females housed in nestboxes for 8 wk, and males’ home range sizes were determined using weekly grid trapping. Male reproductive success was determined using molecular paternity analysis (six microsatellite primers) for all pups born during the field experiment. Males with higher dominance ranks had larger home ranges. However, male dominance rank was not predictive of the number of total visits to females’ nestboxes or the number of visits to each male's most frequently visited nestbox. Males that made more visits to nestboxes sired more litters. Males that had higher dominance ranks sired fewer litters. These results suggest that there is a reproductive disadvantage to having higher dominance rank among male meadow voles.