Play in adult bonobos (Pan paniscus): modality and potential meaning

The aim of this study was to thoroughly investigate social play and its modalities among adult bonobos. We evaluated how play intensity varies according to the sex-class combination of the playmates and we also performed an analysis on social locomotor-rotational movements (L-R play) and contact interactions (C play). Rough and gentle play sessions were performed with comparable frequencies by male-female and female-female adult dyads, with play signals unlikely when the playmates strongly differed in age and in rank position. L-R play rates did not differ according to the sex-combination of the players; in contrast, C play sessions were particularly frequent among females. Play faces (play signals) were significantly higher during C play than L-R play sessions, thus suggesting that playmates assess reciprocally yet safely their relationships by using facial displays to avoid any kind of misunderstanding. Play was positively correlated with grooming and contact sitting interactions, suggesting that it may be used as a social enhancer. Finally, we found no correlation between both play contexts (L-R and C play) and age, size and rank differences of the players. In conclusion, we suggest that bonobos with their egalitarian society, peculiar social structure, and playful tendency represent an attractive testing subject to examine empirically many emerging hypotheses on adult play behavior.     

Play behavior in immature moor macaques (Macaca maura) and Japanese macaques (Macaca fuscata)

Play is widespread across mammalian taxa, but species strongly vary in the ways they play. In less despotic primate species (i.e., with less steep dominance hierarchies, less severe conflicts, and more reconciliation), play has been described as being more frequent, cooperative, and freely expressed. To study the link between social play and dominance style, we compared play behavior in free-ranging infants, juveniles and subadults of more despotic Japanese macaques (Macaca fuscata, N = 24) and less despotic moor macaques (Macaca maura, N = 17). We found interspecific differences in play behavior that corresponded with the contrasting dominance styles of the study species, largely confirming our predictions. In particular, moor macaques spent a larger proportion of time in solitary and social play than Japanese macaques, while Japanese macaques spent a larger proportion of time in grooming interactions. In moor macaques, play sessions included more players, a larger variety of play behaviors, greater play face rates, a greater proportion of time in contact play, and a higher rate of reciprocal play-biting than in Japanese macaques. Aggressive escalations were not common, but more frequent in Japanese macaques. Finally, a higher frequency of play faces during play sessions predicted the occurrence of more reciprocal play-bites, but not the proportion of time spent in contact play behaviors. Additional studies on other groups and species will allow a better understanding of the link between dominance style and social play.     

Social play behavior in infant Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in Qinling Mountains, China

We describe the development of social play behavior and assess factors influencing the development of play in infant Sichuan snub-nosed monkeys (Rhinopithecus roxellana). Infant snub-nosed monkeys began to exhibit social play at 3 months of age, when they spent an average 0.89% of time engaging in this behavior (range: 0.7-1.12%). At 6 months of age, there was a significant increase in the proportion of time spent in social play, averaging 9.78% of observation time (range: 4.92-17.08%). However, from 7 to 9 months of age during the winter, social play decreased gradually before rising again from 10 months of age in the spring. Play behavior in infant snub-nosed monkeys is influenced by environmental temperature. Males were observed to play more than females, although further data on this are required. Social rank did not influence the social play of wild Sichuan snub-nosed monkey infants.     

Pattern of play behavior in infant (age 1 to 12 months) white‐headed langurs in limestone forests,southwest China

Play behavior is a significant trait of immature nonhuman primates (hereafter primates) that plays an important role in sensory, locomotor, socio‐cognitive, and developmental processes. It has been suggested that the function of play is to practice and improve motor skills related to foraging, avoiding predators, attracting mates, raising offspring, and strengthening the skills needed for the formation and maintenance of social bonds. From September 2009 to August 2010, we investigated the play behavior of 1–12 month‐old infant white‐headed langurs (Trachypithecus leucocephalus) a Critically Endangered primate species endemic to China. We carried out this study in the Guangxi Chongzuo White‐headed Langur National Nature Reserve, and recorded 4421 play bouts and 1302 min of play engaged in by seven infants. We found that infants of different ages exhibited different patterns of play behavior. Specifically, nonsocial play behaviors appeared at one month of age, social play behaviors at two months, and an expanded repertoire of social and non‐social play behaviors at three months of age. The frequency and duration of nonsocial play peaked at five months of age and then decreased, while social play gradually increased with age. Nonsocial play did not differ between the sexes, whereas social play showed sex specificity, with a higher frequency and longer duration of social play in male infants compared to female infants. In addition, male and female white‐headed langur infants appeared to prefer individuals of same sex as social playmates, but no obvious choice preference for a specific individual. In conclusion, we provide the first report of play behavior in a population of wild Critically Endangered white‐headed langurs. We suggest that age‐ and sex‐specific differences in play behavior of infants form the bases for age and sex‐based differences in the social interactions of adult langurs.     

The role of play objects and object play in human cognitive evolution and innovation

In this contribution, we address a major puzzle in the evolution of human material culture: If maturing individuals just learn their parental generation's material culture, then what is the origin of key innovations as documented in the archeological record? We approach this question by coupling a life‐history model of the costs and benefits of experimentation with a niche‐construction perspective. Niche‐construction theory suggests that the behavior of organisms and their modification of the world around them have important evolutionary ramifications by altering developmental settings and selection pressures. Part of Homo sapiens' niche is the active provisioning of children with play objects — sometimes functional miniatures of adult tools — and the encouragement of object play, such as playful knapping with stones. Our model suggests that salient material culture innovation may occur or be primed in a late childhood or adolescence sweet spot when cognitive and physical abilities are sufficiently mature but before the full onset of the concerns and costs associated with reproduction. We evaluate the model against a series of archeological cases and make suggestions for future research.     

Intentionality and Expertise: Learning from Observations of Children at Play in Multilingual, Multicultural Contexts

This article draws from three ethnographic studies of children playing sociodramatically in multilingual, multicultural contexts. Countering a deficit perspective that focuses on what children from nondominant cultures do not know, we use the concept of syncretism to illuminate children's expertise and intentionality as they blend knowledge from multiple worlds to reinvent cultural practices and create new spaces for learning. Findings encourage educators to learn from the syncretic nature of children's teaching and learning through play.     

Play and social dynamics in a captive herd of gelada baboons (Theropithecus gelada)

Here, we provide quantitative data concerning adult and immature play distribution in geladas discussing the results in the light of their social dynamics. Sex differences in immature play does not seem to reflect sex differences in adult role; in fact, frequency, modality (C-play and LR-play), and motivation (measured by play initiations) to play did not vary according to the different sex-class combinations. The occurrence of adult-adult play suggests that geladas are characterized by an apparent more than actual rigidity in their social relationships and this is particularly evident in females which show peculiar traits in the use of social play. Specifically, adult females played with other adult females as much as with immature ones, thus suggesting their high motivation to engage in such interactions. This adult play behavioural pattern is typical of species showing relaxed/fluid more than despotic/rigid social relationships. Such assumption is also supported by the positive correlation observed between play and affinitive behaviours (grooming and agonistic support) found within each age-category (adult females and immature subjects). This evidence, together with the lack of correlation between aggressive contacts and social play, suggests that geladas use play for social assessment purposes and/or to increase their social affiliation levels.     

Early play may predict later dominance relationships in yellow-bellied marmots (Marmota flaviventris)

Play has been defined as apparently functionless behaviour, yet since play is costly, models of adaptive evolution predict that it should have some beneficial function (or functions) that outweigh its costs. We provide strong evidence for a long-standing, but poorly supported hypothesis: that early social play is practice for later dominance relationships. We calculated the relative dominance rank by observing the directional outcome of playful interactions in juvenile and yearling yellow-bellied marmots (Marmota flaviventris) and found that these rank relationships were correlated with later dominance ranks calculated from agonistic interactions, however, the strength of this relationship attenuated over time. While play may have multiple functions, one of them may be to establish later dominance relationships in a minimally costly way.     

Strong Competition Does Not Always Predict Play Asymmetry: The Case of South American Sea Lions (Otaria flavescens)

Although play fighting has been studied for over a century in both human and non‐human animals, quantitative data on marine mammals are still scarce. Here, we investigated play fighting in South American sea lions (Otaria flavescens), one of the most sexually dimorphic species with an extreme polygynous mating system, high levels of both intra‐ and inter‐sexual competition. All these features make South American sea lions a good model species to test some predictions on play fighting. Our results indicate play is restricted to juveniles, being inhibited among adults, and as to be expected in a species that shows a high degree of sexual dimorphism, it is mainly expressed in males. Even though playful interactions were punctuated by competitive behaviours, animals played in a highly symmetric way and were able to adjust their competitive playful interactions in a flexible manner and so reduce the risk of escalation to a minimum level. They were highly selective in their choice of playmates by limiting the number of players per session and playing more with age‐matched companions and friends. All these factors taken together are probably at the basis of the low risk of escalation recorded during the study. This result is predictive of a high ability and motivation of these animals to engage in play behaviour which can have a possible role not only in the acquisition of dominance status, but also in establishing and maintaining social relationships, an unexpected role in a so highly competitive species.     

Social play in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): Implications for natural social systems and interindividual relationships

This study compares adult play behavior in the two Pan species in order to test the effects of phylogenetic closeness and the nature of social systems on play distribution. The social play (both with fertile and immature subjects) performed by adults did not differ between the two species. In contrast, in bonobos, play levels among fertile subjects were higher than in chimpanzees. Findings regarding levels of undecided conflicts (more frequent in bonobos) and formal submission displays (lacking in bonobos) confirm, in the two colonies under study, that bonobos exhibit "egalitarianism" more than chimpanzees. Some authors emphasized the importance of play-fighting for social assessment when relationships among individuals are not codified and structured according to rank-rules. Indeed, adult bonobos played more roughly than chimpanzees. Moreover, adult bonobos displayed the full play-face at a high frequency especially during rough play sessions, whereas in chimpanzees, the frequency of play signals was not affected by roughness of play. The frequency of social play among bonobo females was higher than in any other sex combinations, whereas no difference was found for chimpanzees. As a matter of fact, social play can be viewed as a balance between cooperation and competition. Among bonobo females, characterized by social competence and affiliation, social play might enhance their behavioral flexibility and increase their socially symmetrical relationships which, after all, are the basis for their egalitarian society.     

Sex‐specific recombination rates and allele frequencies affect the invasion of sexually antagonistic variation on autosomes

The introduction and persistence of novel, sexually antagonistic alleles can depend upon factors that differ between males and females. Understanding the conditions for invasion in a two‐locus model can elucidate these processes. For instance, selection can act differently upon the sexes, or sex linkage can facilitate the invasion of genetic variation with opposing fitness effects between the sexes. Two factors that deserve further attention are recombination rates and allele frequencies – both of which can vary substantially between the sexes. We find that sex‐specific recombination rates in a two‐locus diploid model can affect the invasion outcome of sexually antagonistic alleles and that the sex‐averaged recombination rate is not necessarily sufficient to predict invasion. We confirm that the range of permissible recombination rates is smaller in the sex benefitting from invasion and larger in the sex harmed by invasion. However, within the invasion space, male recombination rate can be greater than, equal to or less than female recombination rate in order for a male‐benefit, female‐detriment allele to invade (and similarly for a female‐benefit, male‐detriment allele). We further show that a novel, sexually antagonistic allele that is also associated with a lowered recombination rate can invade more easily when present in the double heterozygote genotype. Finally, we find that sexual dimorphism in resident allele frequencies can impact the invasion of new sexually antagonistic alleles at a second locus. Our results suggest that accounting for sex‐specific recombination rates and allele frequencies can determine the difference between invasion and non‐invasion of novel, sexually antagonistic alleles in a two‐locus model.     

Physical and activity changes associated with pregnancy in captive lowland gorillas (Gorilla gorilla gorilla)

The physical appearance and the behavior of five lowland gorilla females at two zoos were recorded before and during 14 pregnancies. Whereas only the quality of changes associated with pregnancy had been described previously, in this study the behavior of the females was recorded quantitatively and evaluated statistically. All females at Stuttgart showed distention of the abdomen three to four months before delivery and development of the breasts a few weeks before parturition. Milk was produced by the third month in one case, and in the sixth month or later in the others, sometimes after delivery. The female at Frankfurt stopped social play seven days after conception; the Stuttgart females, in general, stopped, in the first weeks of pregnancy. Social activity and locomotion decreased considerably within the first three months in all females and were significantly different between the pregnant and the nonpregnant state in all the Stuttgart females. In this study, as in others in the literature, behavioral changes were much more reliable signs of pregnancy than were physical changes. Thus the quantitative results confirm those obtained solely by qualitative observation.