Social relations in family-groups of wood miceApodemus sylvaticus under laboratory and enclosure conditions

Two family-groups of wood miceApodemus sylvaticus (Linnaeus, 1758) were observed: one in the laboratory (terrarium 100 × 60 × 60 cm) and the other under enclosure conditions (outdoor enclosure 200 × 100 × 100 cm). Three consecutive periods in the social relations of the family-groups were described: the linear hierarchy, the multidirectional social relations, and the variable dominance-subordination relations. The linear social hierarchy was reflected in the subordination behaviour of family members in relation to the dominant male, the father of the family. During the period of coexistence of a pair of parents with two litters of juveniles, both in the laboratory and under enclosure conditions, amicable relations between individuals occurred. Advanced forms of social care (paternal care, allomaternal lactation, transportation and licking by relatives) were also observed. In the laboratory, agonistic interactions were lacking during the linear hierarchy period, but the attainment of sexual maturity by individuals of the first two litters, accompanied by an increase in the number of mice, resulted in aggression between mature males and competition for dominance. Further, under increasingly crowded conditions, agonistic behaviour prevailed over other types of social interaction, and the survival rate of juveniles decreased. Under enclosure conditions a seasonal variation in agonistic interaction between mature individuals was recorded, similar to that observed in the wild. Despite the systematic increase in population density, no attacks by mature males on juveniles were observed.     

Response to social challenge in young bonnet (Macaca radiata) and pigtail (Macaca nemestrina) macaques is related to early maternal experiences

Previous experience affects how young primates respond to challenging social situations. The present retrospective study looked at one aspect of early experience, the quality of the mother-infant relationship, to determine its relationship to young bonnet and pigtail macaques' responses to two social challenges: temporary maternal separation at 5-6 months and permanent transfer to an unfamiliar peer group at 16-17 months. Relationship quality was measured quantitatively on 30 macaque mother-infant pairs with the Relationship Quality Index (RQI), the ratio of relative affiliation to relative agonism as previously applied to capuchin monkeys. Infants with high RQI values had amicable mother-infant relationships and infants with low RQI values had agonistic mother-infant relationships. Young monkeys with amicable and agonistic relationships showed consistent differences in behavior before, during, and after each social challenge, supporting the hypothesis that juveniles from amicable mother-infant relationships based on the RQI coped more effectively with social challenges than did juveniles from agonistic mother-infant relationships. Results suggest 1) characteristic amicability or agonism in early mother-offspring macaque relationships has the potential to influence offspring behavior in tense social contexts and 2) the RQI is useful as one of a coordinated suite of methods for studying the development of social skills.     

Inter-individual relationships in proboscis monkeys: a preliminary comparison with other non-human primates

This is the first report on inter-individual relationships within a one-male group of proboscis monkeys (Nasalis larvatus) based on detailed identification of individuals. From May 2005 to 2006, focal and ad libitum data of agonistic and grooming behaviour were collected in a forest along the Menanggul River, Sabah, Malaysia. During the study period, we collected over 1,968 h of focal data on the adult male and 1,539 h of focal data on the six females. Their social interactions, including agonistic and grooming behaviour, appeared to follow typical patterns reported for other colobines: the incidence of social interaction within groups is low. Of 39 agonistic events, 26 were displacement from sleeping places along the river, 6 were the α male threatening other monkeys to mediate quarrels between females and between females and juveniles, and 7 were displacement from feeding places. Although the agonistic behaviour matrix based on the 33 intra-group agonistic events (excluding events between adults and juveniles and between adults and infants) was indicative of non-significant linearity, there were some specific dominated individuals within the group of proboscis monkeys. Nonetheless, grooming behaviour among adult females within a group were not affected by the dominance hierarchy. This study also conducted initial comparisons of grooming patterns among proboscis monkeys and other primate species. On the basis of comparison of their grooming networks, similar grooming patterns among both-sex-disperse and male-philopatric/female-disperse species were detected. Because adult females in these species migrate to groups repeatedly, it may be difficult to establish the firm grooming exchange relationship for particular individuals within groups, unlike in female-philopatric/male-disperse species. However, grooming distribution patterns within groups among primate species were difficult to explain solely on the basis of their dispersal patterns. Newly immigrated females in some species including proboscis monkeys are eager to have social interactions with senior group members to improve their social position.     

Male and female agonistic and affiliative relationships in a social group of farm cats (Felis catus L.)

A dominance hierarchy based on the outcome of agonistic encounters was found among male and female domestic cats. A female dominated over some males. The dominance concept is also discussed in terms of social bonding. The relationships among adult females were amicable, whereas adult males showed reciprocal tolerance. The flow of affiliative behaviour was directed mainly from females to one male of the group. The analysis of marking behaviour showed that this male sprayed urine and rubbed the perioral and cheek regions of the face on the objects of the environment at a higher rate than the other members of the group. Nevertheless, rubbing the perioral and cheek regions of the face on objects was not correlated to dominance rank, possibly because it has some function in social communication other than territorial defence against strangers. No relationships have been found between claw scratching, rolling on the ground and social rank, or between the former and other marking behaviour. It is concluded that claw scratching and rolling were not utilised to mark territory.     

Development of Social Behaviour in Four Litters of Dogs (Canis familiaris)

The development of social behaviour in 4 litters of dogs was observed without interfering with the puppies from birth to 8 weeks of age. Direct and continuous observation was combined with video recording. Three of the litters were observed during one session of 2 h once a week, and the fourth litter during one session of 40 min twice a week. Social interactions were divided into 1) investigation of litter mates (licking, sniffing or investigating orally), 2) social play, 3) and interactions in which agonistic elements (dominance postures, threats, bites or submission) were displayed. The different forms of social interactions appeared for the first time when the puppies were between 14 and 21 days of age. Social investigation appeared first and was followed by play and agonistic interactions. From week 5, differences between the puppies in the tendency to initiate social play and agonistic interactions emerged. Generally, within the litters individual differences were consistent over weeks 6-8 (positive correlations between weeks), whereas the tendency in the puppies during these weeks were negatively correlated with those of week 3 (play) or weeks 3 and 4 (agonistic interactions), indicating a rebound effect for both play and agonistic behaviour. No significant correlations, however, were found for social investigation. More often than expected males played or engaged in agonistic with other males, whereas these behaviours occurred less often than expected between females. Both males and females, however, preferred male partners for agonistic interactions. No sex differences were found in the direction of social investigation. Agonistic behaviour was often responded to by play and play was often responded to by agonistic behaviour, and the results indicated that before 8 weeks of age differences in social behaviour between the puppies were already established.     

Social behaviour of juvenile red sea urchins, Strongylocentrotus franciscanus (Agassiz)

This study examined social behaviour of juvenile red sea urchins, Strongylocentrotus franciscanus (Agassiz). At four coastal locations in British Columbia, Canada, distributions of juvenile sea urchins were examined in relation to adults. One-third of all juveniles were found underneath an adult “spine canopy”, another third outside the spine canopy but close to an adult, and the remainder were found apart from adults. Juveniles associated with adults were contagiously distributed with each other. In the laboratory, juveniles chose adults over other target locations. They selected adults in particular locations over others, and in one case showed selection among adults; but they did not select between fed and non-fed adults, nor between adults that had been “protective” and “non-protective” of juveniles in the field. In time-lapse film studies, starved juveniles showed little change in social behaviour when food was introduced, but showed a dramatic change in behaviour when predators were active. We suggest that juvenile red sea urchins are found under the spine canopies of adults as a result of juvenile behaviour; and that this interaction functions to protect juveniles from predation. The strong gregarious nature of juveniles among themselves may have evolved in a different selective situation as a defence against predation and to ensure reproductive success.     

The Ontogeny of Social Behaviour in Four Conilurine Rodents (Muridae) of Australia

This paper describes the ontogeny of social behaviour in 4 closely related rodents from diverse habitats. These rodents, Notomys alexis, Pseudomys albocinereus, P. shortridgei and P. desertor exemplify 4 different social organizations. The young were observed daily under semi-natural conditions until weaning, and subsequently at intervals of one week until sexual maturity. Relationships between the young and their families were determined from the relative frequencies of attracting (amicable) and repelling (agonistic) behaviour. The social behaviour of the young is correlated with the social behaviour of adults. Social interactions experienced from birth throughout sub-adulthood appear to prepare these species for their adult social life.     

Play in Harbour seals (Phoca vitulina)

Harbour seals in a breeding colony at Miquelon exhibited 13 distinct types of play behaviour, 80% of which was solitary locomotor play. Most play was displayed by yearlings and juveniles, though a significant amount of adult play occurred as well. Mothers and pups rarely played. Yearlings played socially as often as alone, but seals of other age classes played by themselves most often. Only social play resembled adult activities. It is suggested that the function of mammalian play has been obscured by its similarity to adult behaviour.     

Agonistic Behaviour and Development of Territoriality During Ontogeny of the Sea Anemone Dwelling Crab Allopetrolisthes Spinifrons (H. Milne Edwards, 1837) (Decapoda: Anomura: Porcellanidae)

Allopetrolisthes spinifrons is an ectosymbiotic crab of the sea anemone Phymactis clematis. As a consequence of low host abundance, these represent a scarce and limited resource for the crab. Additionally, the relatively small size of the sea anemone host suggests that few symbiotic crabs can cohabit on one host individual, forcing crabs to adopt a territorial behaviour. In order to examine the potential presence and ontogenetic development of territoriality, the agonistic behaviour between crabs of various ontogenetic stages (adults, juveniles, and recruits) was studied in the laboratory. Laboratory experiments demonstrated that adult or juvenile crabs aggressively defended their sea anemone hosts against adult or juvenile intruders, respectively, but both adult and juvenile crabs tolerated recruits. Adult crabs behaved indifferently towards juvenile crabs, sometimes tolerating them, sometimes expelling them. Recruits never showed agonistic behaviour among themselves. The agonistic interactions observed in the laboratory and the uniform population distribution pattern on sea anemones recently described for A. spinifrons indicate that this species exhibits territorial behaviour, which develops during ontogeny. Territoriality in this species and other symbiotic decapods may function as a density-dependent mechanism of population regulation, being mediated by the availability of hosts. Resource monopolisation behaviours may be common among other symbiotic and free-living marine invertebrates inhabiting discrete habitats that represent a limiting resource.     

Agonistic Behaviour and Development of Territoriality During Ontogeny of the Sea Anemone Dwelling Crab Allopetrolisthes Spinifrons (H. Milne Edwards, 1837) (Decapoda: Anomura: Porcellanidae)

Allopetrolisthes spinifrons is an ectosymbiotic crab of the sea anemone Phymactis clematis . As a consequence of low host abundance, these represent a scarce and limited resource for the crab. Additionally, the relatively small size of the sea anemone host suggests that few symbiotic crabs can cohabit on one host individual, forcing crabs to adopt a territorial behaviour. In order to examine the potential presence and ontogenetic development of territoriality, the agonistic behaviour between crabs of various ontogenetic stages (adults, juveniles, and recruits) was studied in the laboratory. Laboratory experiments demonstrated that adult or juvenile crabs aggressively defended their sea anemone hosts against adult or juvenile intruders, respectively, but both adult and juvenile crabs tolerated recruits. Adult crabs behaved indifferently towards juvenile crabs, sometimes tolerating them, sometimes expelling them. Recruits never showed agonistic behaviour among themselves. The agonistic interactions observed in the laboratory and the uniform population distribution pattern on sea anemones recently described for A . spinifrons indicate that this species exhibits territorial behaviour, which develops during ontogeny. Territoriality in this species and other symbiotic decapods may function as a density-dependent mechanism of population regulation, being mediated by the availability of hosts. Resource monopolisation behaviours may be common among other symbiotic and free-living marine invertebrates inhabiting discrete habitats that represent a limiting resource.     

Play as a reinforcer for maze-learning in juvenile rats

The reinforcing value of play for juvenile rats was examined, relative to that of other social experiences. Successive experimental groups chose, in a T-maze, between normal companions and companions whose social behaviour had been rendered abnormal by physical confinement or by amphetamine or chlorpromazine treatment. Both these drugs inhibited play, as did physical restriction, but they had opposing effects on other social behaviours, so that the choice presented was between social experiences with and without play. Young rats learnt these social discriminations as easily as a food/no food discrimination in the same apparatus, developing a preference for the playing partner. Non-playing partners indulging in large amounts of amicable social behaviour were chosen more frequently than unsociable animals.     

Social and reproductive behaviour of critically endangered northern white rhinoceros in a zoological garden

Northern and southern white rhinos have poor reproduction in captivity and social interactions between them, especially increased agonistic behaviour, are believed to be one of the possible reasons. Northern white rhino is currently on the brink of extinction with less than ten animals surviving. We studied the social behaviour of northern white rhinos in zoological garden and investigated the effects of separation of the oldest, wild-born female from the herd on the social behaviour of other group members. After the separation, the numbers of agonistic and the play interactions between the animals significantly increased, no change was found in cohesive behaviour. Our results suggest that a composition of white rhino groups has a significant influence on social interactions between the animals and that better knowledge of proper composition of their groups in captivity in terms of age, sex and wild or zoo origin might improve animals’ well-being and also increase a chance for reproduction.     

Functions and consequences of play behaviour in juvenile Belding's ground squirrels

We evaluated hypotheses suggesting that play behaviour in free-living juvenile Belding's ground squirrels, Spermophilus beldingi, helps to regulate energy balance and promotes development of motor skills. We also examined the possibility that play behaviour and motor development influence important early life events in S. beldingi. We regularly measured body fat of juveniles using nondestructive methods, observed their behaviour, conducted motor skills tests, monitored dispersal status of males, and evaluated weaning success of females during their first effort to reproduce as yearlings. Rates of feeding were greater among juveniles who engaged in both social and nonsocial play behaviour at low compared to high rates, suggesting that play does not help young animals expend excess energy that they consume. By contrast, body fat was greater among juveniles who engaged in social play at high compared to low rates, supporting the idea that energetic variables such as body fat limit the expression of play behaviour. Motor skills of juvenile S. beldingi improved throughout the period in which juveniles engaged in play behaviour. Improvement in motor skills was greater in juveniles who engaged in social play at high compared to low rates, suggesting a link between social play and motor development. Motor skill levels at the end of the play period were greater among males who dispersed by the end of the juvenile summer than among males still residing in their natal areas. Rates of social play and improvement in motor skills over the course of the play period were greater in females who successfully weaned a litter as yearling than in females who did not. These last two results raise the possibility that play behaviour and motor development during the play period influence important events in the early lives of S. beldingi, which may ultimately influence long-term reproductive success.     

Juvenile Richardson's Ground Squirrels (Spermophilus richardsonii) Manifest Both Littermate and Neighbour/Stranger Discrimination

Newborn ground squirrels (Spermophilus spp.) are reared in isolation of individuals belonging to other litters. At or near weaning, they broaden their social interactions to encompass conspecifics other than littermates. In laboratory studies, juvenile ground squirrels commonly discriminate littermates from non-littermates, but few studies have examined whether behavioural discrimination at that level persists under natural circumstances. Previous studies have also all but ignored the possibility of alternative levels of social discrimination augmenting or replacing the discrimination of littermates from non-littermates. To examine those possibilities, I staged interactions in a neutral arena between littermate, neighbour and non-neighbour (stranger) pairs of otherwise free-living juvenile Richardson's ground squirrels (Spermophilus richardsonii). Littermates interacted more frequently than other pair types and showed the highest proportion of cohesive behaviour and the least agonism. Strangers interacted less frequently than all other pair types, showed the fewest recognitive contacts, and were the least cohesive and most agonistic. Neighbours interacted less than littermates, but more than strangers, and showed intermediate levels of cohesion and agonism. Neighbours also engaged in more recognitive contacts than strangers, but did not have fewer recognitive contacts than littermates. Defecation rates also differed among interactant types: littermates left the fewest scats in the arena, neighbours deposited an intermediate number and strangers defecated the most. My results substantiate those of earlier laboratory studies suggesting that the discrimination of littermates from neighbouring juveniles persists in the natural context. These data also extend those earlier findings by demonstrating that juveniles come to discriminate neighbours from strangers. Comparisons of the behaviour of related vs. unrelated neighbours reveal that neighbour/stranger discrimination is contingent upon differential familiarity. Simultaneous social discrimination at the levels documented could promote kin-biased behaviour and cooperation among both related and unrelated neighbours.     

The diurnal patterns of behaviour of the wild Barbary macaque Macaca sylvanus

The daily routine of a population of wild Barbary macaques is described. The day journey is mainly on the ground with the animals taking to the trees only for sleeping, resting, avoiding predators and, during some seasons, for feeding. Two methods (based on individual activity records and on social interactions) were used to measure the diurnal distribution of behaviour. In the summer, feeding was bimodally distributed with peaks morning and afternoon. The initiation of social interactions peaked at the same time, owing to the frequent use of agonistic behaviour to maintain individual distance while feeding. The reduction in feeding at midday was accompanied by an increase in allogrooming and resting and in the proportion of animals in the trees. Peaks of friendly approaches and in the initiation of interactions involving unweaned monkeys (excluding play, maternal behaviour and agonistic behaviour) occurred on either side of the midday rest period and in the evening. In the winter, there was no midday rest period; grooming decreased throughout the day while feeding increased. Sexual behaviour was rare during the summer.     

The ontogeny of social play in a feral troop of vervet monkeys (Cercopithecus aethiops sabaeus): The function of early play

We describe the ontogeny of social play over the first 30 weeks of age in a troop of feral vervet monkeys (Cercopithecus aethiops sabaeus) in Barbados. Play time increased rapidly for the first 10 weeks but remained relatively constant thereafter. The form of play changed with infant age; bouts became more frequent but of shorter duration. Play time, bout frequency, and bout duration at a given age differed between infants; younger infants altered their play patterns to complement those of older infants. All infants played more within their own year class than with older juveniles; play time increased with decreasing age difference between the infant and the play partner. Infants terminated a higher proportion of their play bouts the greater the age difference between themselves and their play partners. Preferred play partners are therefore individuals of similar age that will be similar in size and have matched motivation to play and form of play. Neither play time nor proportion of bouts initiated or terminated was correlated with the social rank of the play partner, which suggests that play is not directed toward individuals that may be high-value alliance partners. Maternal intervention in play occurred primarily when infants were<10 weeks old. It was not correlated with the age or social rank of the mother or with the age or social rank of her infant's play partner. Infants played more and terminated a lower proportion of their play bouts in the absence of their mothers than in their presence. Our results are consistent with the hypothesis that the primary function of early play is to enhance physical fitness and to develop coordination and other fighting skills, with minimal risk of injury.     

The effect of familiarity on social interactions in meadow voles,Microtus pennsylvanicus: a laboratory and field study

The role of familiarity in affecting the outcome of social interactions among meadow voles was investigated in both a laboratory and a field experiment. In the laboratory, captive meadow voles, Microtus pennsylvanicus, were exposed to a conspecific's odour. The voles were then placed into two groups: familiar and unfamiliar. Familiar voles were individuals who underwent staged dyadic encounters with the conspecific to whose odours they had been exposed. Unfamiliar voles were individuals who underwent paired encounters with conspecifics to whose odours they had not been exposed. In the field experiment, familiar voles were neighbours that were trapped within each other's home ranges over two consecutive bi-weekly trapping sessions. Unfamiliar voles were individuals that were trapped on different trapping grids. The results of the laboratory and field studies were similar. Encounters between familiar females resulted in less agonistic behaviour and more amicable acts than encounters between unfamiliar females. In contrast, encounters between familiar males resulted in more agonistic behaviour than encounters between unfamiliar males. Familiarity did not affect the outcome of male-female interactions. These results are discussed in the framework of the social system of the meadow vole.     

Ontogeny of Dominance in Free-living Red Foxes

We established the ethogram of fox pups of free-living foxes after emergence from the den. We recorded the reactions of the pups during encounters to determine their social status and documented the relationship between social status and social play. The ethogram comprised 12 behavioural classes including 64 behavioural acts. ‘Moving’ and ‘exploration’ were the most observed classes with a frequency of occurrence higher than 20%. Most behavioural acts occurred at frequencies ranging from 0.5% to 5%. The intra-litter hierarchy was linear. Social play was the predominant type of social activity among pups, and induced the maintenance of dominance relationships among them. Adult—juvenile interactions (‘relation with an adult’ and ‘suckling’) could depend on the social rank of the juveniles. Juvenile-juvenile contacts allowing strong social bonds to develop between littermates (‘tactile communication’ and ‘comfort social behaviour licking and nibbling fur’) were not correlated with the social rank of the juveniles.     

The ontogeny of sex differences in the behavior of patas monkeys

Many studies of sex differences in primates have been based on small experimental groups of peers in which only a limited range of social behavior could be expressed. In addition, the first few months of life are often the focus of such studies, with relatively little attention paid to older juveniles. In this study, 11 male and 9 female juvenile patas monkeys, living in a captive social group with all age-sex classes available, were observed between 1 and 4 years of age. A subset of seven patas monkeys was also observed between birth and 1 year of age. Here, we report the development of sex differences in independence, play, grooming, positioning behavior, and aggression over the juvenile period. Juvenile male patas monkeys played more and in longer bouts than females, but wrestling (rough-and-tumble play) was not more common among males. There were few differences in behaviors directed to male and female juveniles by other group members. Distinct differences emerged only in the behaviors of the juveniles themselves, with females being more active participants in social and aggressive interactions than males. In general, sex differences in patas monkeys show a mixture of patterns, some of which are predictive of adult sex differences and some of which appear to be specific to the particular demands of the juvenile period in this species     

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.