Risk-prone hunting by chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) increases during periods of high diet quality

Most studies suggest that during times of nutritional stress, an animal faced with two foraging choices should follow a risk-prone strategy, choosing the option with highest payoff variance. This “scarcity/risk” hypothesis was developed to account for the foraging patterns of small animals with high metabolic rates susceptible to the threat of starvation. In this paper, we propose that animals should also be risk-prone when their diet quality is particularly high, far exceeding that which is needed to survive. Under these circumstances, the costs of experiencing a low or negative payoff can easily be recouped. We suggest that large-bodied omnivores are most likely to adopt this “abundance/risk” strategy. We investigate this question among wild chimpanzees (Pan troglodytes) that choose between a risk-averse strategy of feeding on plant material and a risk-prone strategy of hunting red colobus monkeys. Using 14 years of data on the Kanyawara chimpanzees of Kibale National Park, Uganda, we find strong evidence that chimpanzees follow the “abundance/risk” strategy. Both hunting rate (hunts/100 observation hours) and the probability of hunting upon encountering red colobus monkeys were positively correlated with seasonal consumption of ripe drupe fruits, a class of preferred food associated with elevated reproductive performance by females. Critically, these results remained statistically significant after controlling for the potentially confounding effects of male chimpanzee party size and the presence of sexually receptive females. These findings suggest that the relationship between risk-sensitive foraging and diet quality depends upon the daily probability of starvation, the number of alternative foraging strategies, and the degree to which diet quality satisfies an animal’s nutritional requirements.     

Reciprocity and trades in wild West African chimpanzees

Why do animals help other individuals and provide benefits to the recipient, sometimes at personal cost? In this study, we aim to determine if some of the helpful behaviors observed in a group of wild chimpanzees (Taï chimpanzee group, Côte d'Ivoire, West Africa) are exchanged among individuals resulting in a net benefit for both participants. We adopted an inclusive view of exchanges by considering that all commodities (i.e., social behaviors as grooming, sex, support, as well as resources, such as meat) can be exchanged. This would result in “market” type social interactions in which debts of one commodity can be met by giving the same or other commodities. We investigated whether both in-kind exchanges and trades of commodities occurred. The Taï chimpanzees reciprocated the amount of grooming they received and were more likely to give support in agonistic conflicts to those who also supported them. In addition, they traded support for meat and meat for sex. Both male and female chimpanzees exchanged many different commodities, which seemed to result in long-term balanced relationships. Our results suggest that wild chimpanzees rely on other group members to obtain many of the important commodities they need.     

Why chimpanzee (Pan troglodytes schweinfurthii) mothers are less gregarious than nonmothers and males: the infant safety hypothesis

Socialization of young is an important component of maternal care in social mammals. It is therefore perplexing that female chimpanzees with dependent offspring spend more time alone than females without dependent offspring, and than males. We propose that chimpanzee mothers are less gregarious than nonmothers and males to reduce the risk of injury that aggressive males pose to their offspring. We predict that mothers will associate less with males, associate with fewer males, and reduce mother−infant proximity in the presence of males, and that these effects will reduce with increasing offspring age. We test whether the pattern of gregariousness and mother−offspring proximity support these predictions in the Kanyawara chimpanzee community in Uganda. The probability that a female was found in the presence of males was lower for mothers than nonmothers and increased with offspring age. The probability that a female was found with other females did not differ between mothers and nonmothers. Mother-to-offspring distance was higher when a mother was in an all-female group than in a mixed-sex group and increased with her offspring's age. Mother-to-offspring distance was greatest when there were relatively low numbers of males and relatively high numbers of females in a subgroup. We propose that mothers avoid grouping with males because of the vulnerability of their young, and that the presence of males in a subgroup increases a female's protectiveness of her young. We discuss the implications of our findings and the relevance of fission−fusion group formation to chimpanzee mothers.     

Sex differences in the movement patterns of free-ranging chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>): foraging and border checking

Most social primates live in cohesive groups, so travel paths inevitably reflect compromise: decision processes of individuals are obscured. The fission–fusion social organisation of the chimpanzee, however, allows an individual's movements to be investigated independently. We followed 15 chimpanzees (eight male and seven female) through the relatively flat forest of Budongo, Uganda, plotting the path of each individual over periods of 1–3 days. Chimpanzee movement was parsed into phases ending with halts of more than 20 min, during which individuals fed, rested or engaged in social activities. Males, lactating or pregnant females and sexually receptive females all travelled similar average distances between halts, at similar speeds and along similarly direct beeline paths. Compared to lactating or pregnant females, males did travel for a significantly longer time each day and halted more often, but the most striking sex differences appeared in the organisation of movement phases into a day's path. After a halt, males tended to continue in the same direction as before. Lactating or pregnant females showed no such strategy and often retraced the preceding phase, returning to previously visited food patches. We suggest that female chimpanzee movements approximate an optimal solution to feeding requirements, whereas the paths of males allow integration of foraging with territorial defence. The ‘continually moving forwards’ strategy of males enables them to monitor their territory boundaries—border checking—whilst foraging, generally avoiding the explicit boundary patrols observed at other chimpanzee study sites.     

Association patterns among wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) reflect sex differences in cooperation

Theory predicts that frequent dyadic association should promote cooperation through kin selection or social tolerance. Here we test the hypothesis that sex differences in the strength and stability of association preferences among free-ranging chimpanzees conform to sex differences in cooperative behavior. Using long-term data from the Kanyawara chimpanzee (Pan troglodytes schweinfurthii) community (Kibale National Park, Uganda), we calculated indices of intra-sexual dyadic association over a 10-year period. We found that (1) male–male dyads had significantly stronger association indices than female–female dyads, (2) the pattern of association preferences in both sexes changed little over the entire study period, and (3) when comparing periods with different alpha males, changes in association strength were more frequent among males. These results demonstrate that both the strength and stability of association patterns are important components of social relationships. Male chimpanzees, which are characterized by frequent cooperation, had association preferences that were both strong and stable, suggesting that forming long-term bonds is an important dominance strategy. However, the fact that male association patterns were sensitive to upheaval in the male dominance hierarchy suggests that males also take advantage of a changing social climate when choosing association partners. By contrast, the overall strength of female associations was relatively weak. Female association preferences were equally stable as males’; however, this reflected a dyad’s tendency to be found in the same party rather than to associate closely within that party. Therefore, in this community, female association patterns appear to be more a consequence of individual ranging behavior rather than a correlate of cooperation.     

Social context-dependent immune gene expression in bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)

Social insects are prone to attack by parasites as they provide numerous resources of food and brood, homeostatic nest conditions and a high density of individuals, enhancing the transmission of parasites. The defence of social insects might occur at different levels, the individual and the group. Individual defence occurs in part via the innate immune system resulting in the expression of antimicrobial substances. Group level defences, summarised as ‘social immunity’, represent a suite of behavioural and organisational features. Here, all effects contributing to social immunity except for the social context were removed from bumblebee (Bombus terrestris) workers, kept either in groups or solitarily. The gene expression of six effector molecules of the immune system was monitored in both groups and in controls from the same source colonies. The social treatment has a highly significant effect on immune gene expression, with groups exhibiting higher levels of two antimicrobial peptides (AMPs) and two lysozymes. Phenoloxidase is affected at the regulatory level, with a strong upregulation of its suppressor Spn27A in groups suggesting a trade-off with antimicrobial activity. AMPs are strongly upregulated in groups, whereas lysozymes are strongly downregulated in solitary treatments suggesting another trade-off. Clearly, social immunity impacts elements of individual immunity.     

The Contribution of Long-Term Research at Gombe National Park to Chimpanzee Conservation

Abstract:  Long-term research projects can provide important conservation benefits, not only through research specifically focused on conservation problems, but also from various incidental benefits, such as increased intensity of monitoring and building support for the protection of an area. At Gombe National Park, Tanzania, long-term research has provided at least four distinct benefits to wildlife conservation. (1) Jane Goodall's groundbreaking discoveries of chimpanzee (Pan troglodytes) tool use, hunting, and complex social relationships in what was then a game reserve drew attention to the area and created support for upgrading Gombe to national park status in 1968. (2) The highly publicized findings have earned Gombe and Tanzania the attention of a worldwide public that includes tourists and donors that provide financial support for Gombe, other parks in Tanzania, and chimpanzee conservation in general. (3) Crucial information on social structure and habitat use has been gathered that is essential for effective conservation of chimpanzees at Gombe and elsewhere. (4) A clear picture of Gombe's chimpanzee population over the past 40 years has been determined, and this has helped identify the greatest threats to the viability of this population, namely disease and habitat loss outside the park. These threats are severe and because of the small size of the population it is extremely vulnerable. Research at Gombe has led to the establishment of conservation education and development projects around Gombe, which are needed to build local support for the park and its chimpanzees, but saving these famous chimpanzees will take a larger integrated effort on the part of park managers, researchers, and the local community with financial help from international donors.     

Social facilitation of exploration in mosquitofish (<Emphasis Type="Italic">Gambusia holbrooki</Emphasis>)

The social environment can exert a powerful influence on the expression of an individual’s behaviour patterns. For example, social facilitation occurs where individuals are more likely to express a given behaviour, or express it a greater rate, in the presence of conspecifics. Social facilitation is partly driven by an individual’s perception of risk, which is a function both of the size of its social group and the information that it gathers relating to predator activity and risk. Here I tested the effects of social group size (one, two, four, eight or 16 fish) and the presence of ‘social’, ‘predation’ or ‘neutral’ chemical cues (derived respectively from live conspecifics, injured conspecifics or a blank water control) on the exploratory behaviour of juvenile mosquitofish (Gambusia holbrooki) in a novel environment. Focal fish in larger groups explored a greater proportion of the arena during the course of the experiment, demonstrating social facilitation of exploration. After 4 h in the arena, focal fish in all group sizes showed significantly reduced swimming activity, suggesting that the initial faster swimming activity of fish on entry to the arena may be in response to the motivation to explore. The presence of predation cues in the environment had the effect of reducing exploratory behaviour across groups in a novel environment, whereas social cues had no effect on exploratory behaviour. Taken as a whole, these results suggest that there is a high degree of context dependency in the expression of exploratory behaviour, with a strong influence of both the presence of conspecifics and cues relating to potential danger.     

Family insurance: kin selection and cooperative breeding in a solitary primate (Microcebus murinus)

Lactation imposes substantial physiological costs on mothers and should therefore not be directed towards foreign offspring. Such allonursing, however, is common in mammal species that share roosts. Hypotheses to explain allonursing among such plural breeders include misdirected parental care, milk evacuation, brood parasitism, reciprocity, and kin selection. The necessary behavioral data, in combination with data on kinship and kin recognition, have rarely been available to distinguish among these explanations, however. In this study, we provide evidence for cooperative nursing and adoption by plural-breeding females in a nocturnal primate, the gray mouse lemur (Microcebus murinus), in which females forage solitarily during the night, but form day-time sleeping groups with one to two other females. We observed 34 resident females in an 8 ha study area in Kirindy Forest, Madagascar, over three consecutive annual breeding seasons and determined genetic relationships among all members of this population. Five sleeping groups of adult females were filmed inside their roosts during one breeding season after females gave birth. The composition of groups changed substantially across years, but they always consisted of close maternal relatives. All females within a group gave birth to one to three infants. They regularly transferred only their own offspring among roosting sites, demonstrating an ability to discriminate between their own and other’s offspring, but they regularly groomed and nursed related offspring other than their own and adopted related dependent young after their mother’s death. Kin selection may therefore be the main selective force behind cooperative breeding among these closely related females with a high mortality risk, providing each of them with family insurance.     

Coalitionary mate guarding by male chimpanzees at Ngogo, Kibale National Park, Uganda

Cooperative mate guarding by males is unusual in mammals and birds, largely because fertilizations are non-shareable. Chimpanzees live in fission-fusion communities that have cores of philopatric males who cooperate in inter-group aggression and in defending access to the females in their community. Male contest mating competition is restrained within communities, but single high-ranking males sometimes try to mate guard estrous females. Data from an unusually large chimpanzee commmunity at Ngogo, Kibale National Park, Uganda, that contains more males than any previously studied community show new variation in chimpanzee mate-guarding behavior. Contrary to expectation given the large number of males, mate guarding was as common as, or more common than, at other sites, and males other than the alpha male guarded more often. More strikingly, pairs or trios of top-ranking males sometimes engaged in cooperative aggression to prevent estrous females from mating with other males, but tolerated each other's mating activities. Both single males and coalitions mostly guarded periovulatory females. Mate-guarding coalitions were previously unknown in chimpanzees. Coalitions occurred in large mating parties, seemingly because these often contained too many males for single males to maintain exclusive access to estrous females. Coalition members gained higher shares of copulations than they could have expected from solo mate guarding, and suffered lower per capita costs of guarding (as inferred from aggression rates). Two males who most often participated in coalitions formed two-male coalitions at about the point where the number of males present made it unlikely that either could get 50% or more of total copulations on his own, and formed trios when this value dropped below 33%. Kin selection could be a factor in cooperation among male chimpanzees, but coalition members were not necessarily close relatives and the apparent structure of payoffs fit that of mutualism. Furthermore, reliance of male chimpanzees on support from allies to maintain high rank could have led to trading of mating exclusivity for support against mating competitors. Received: 28 May 1997 / Accepted after revision: 16 May 1998     

Social personality traits in chimpanzees: temporal stability and structure of behaviourally assessed personality traits in three captive populations

Animals of many species show consistency in behaviour across time and contexts that differs from other individuals' behaviour in the same population. Such ‘personality’ affects fitness and has therefore become an increasingly relevant research topic in biology. However, consistent variation in social behaviour is understudied. In socially living species, behaviour occurs in a social environment and social interactions have a significant influence on individual fitness. This study addressed personality in social behaviour of 75 captive chimpanzees in three zoos by coding observed behaviour. Fifteen behavioural variables were significantly repeatable (range 0.21–0.93) in at least two of the three zoos. The behaviours showed considerable long-term stability across 3 years, which did not differ from the short-term repeatability. The repeatable behaviours were then analysed with factor analyses. They formed five independent factors, three of which consisted of social traits and were labelled ‘sociability’, ‘positive affect’ and ‘equitability’. The two non-social behaviour factors were labelled ‘anxiety’ and ‘activity’. The factor scores were analysed for sex and population differences. Males had higher factor scores in all traits except ‘sociability’. The factor scores differed also between the zoos, implying considerable external effects in trait expression. The results show that chimpanzees show personality in a broad range of social and non-social behaviours. The study highlights the importance of assessing personality in the social behaviour, especially in cohesive social species, as only then can we understand the consequences of personality in socially living species.     

Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana

According to mate choice models, a female should prefer males with traits that are reliable indicators of genetic quality which the sire can pass on to their progeny. However, good genes may depend on the social environment, and female choice for good genes should be context dependent. The side-blotched lizard, Uta stansburiana, exhibits genetically based throat colors (orange, blue, or yellow) that could be used as a sexually selected signal since they reliably predict the genetic quality of mates. The frequencies of male and female morphs cycle between years, and both male and female morphs have an advantage when rare; thus genetic quality will depend on morph frequency. A female should choose a sire that maximizes the reproductive success of both male and female progeny. We examine a game theoretical model that predicts female mate choice as a function of morph frequency and population density. The model predicts the following flexible mate choice rule: both female morphs should prefer rare males in ’boom years’ of the female cycle (e.g., ’rarest-of-N rule’), but prefer orange males in ’crash years’ of the female cycle (’orange-male rule’). Cues from the current social environment should be used by females to choose a mate that maximizes the future reproductive success of progeny, given the social environment of the next generation. We predict that the cue is the density of aggressive orange females. In the side-blotched lizard, cycling mate choice games and context-dependent mate choice are predicted to maintain genetic variation in the presence of choice for good genes. Received: 8 March 2000 / Revised: 26 August 2000 / Accepted: 4 September 2000     

To fission or to fusion: effects of community size on wild chimpanzee (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) social organisation

To balance advantages and disadvantages of group living, some species have fission-fusion social systems in which members of the same group form frequently changing subgroups. This allows flexible responses of group size to external conditions while at the same time retaining group stability. In chimpanzees, subgroup (party) size and composition depend mainly on the presence of receptive females, food availability and the activity of the party. Here we analyse the extent to which fission-fusion parameters are influenced by changes of demographic variables like community size and composition. Data were collected from a habituated West African chimpanzee community (Taï forest, Côte dIvoire) over 10 years, during which total community size decreased from 51 to 21, and the number of adult males decreased from 9 to 2. Taï chimpanzees are highly gregarious, as they spend more than 80% of their time with unrelated conspecifics. With decreasing community size, party size, party duration and male-female association increased. Neither activity nor the presence of receptive females or feeding competition could explain the observed changes in grouping patterns. Thus, the decrease in community size led to an increase of party cohesion and also enhanced cohesiveness between the sexes, while general sociality remained unchanged. Therefore, our data support the notion that small communities are more cohesive and have a less flexible fission-fusion system.Communicated by D. Watts     

Anti-predation behaviour of red colobus monkeys in the presence of chimpanzees

Predator-prey interactions are usually regarded as evolutionary “arms races”, but evidence is still scarce. We examined whether the anti-predation strategies of red colobus monkeys (Procolobus badius) are adapted to the hunting strategies of chimpanzees (Pan troglodytes) in the Taï National Park, Ivory Coast. Taï chimpanzees search for red colobus groups, approach them silently and hunt co-operatively. Our playback experiments and observations of natural encounters revealed that red colobus hid higher up the trees in positions where exposure to the forest floor is minimal and became silent, when chimpanzees were close. They moved away silently through the canopy, when chimpanzees were still at some distance. However, if a group of diana monkeys was nearby in the latter situation, red colobus sought their presence even if they had to move towards the chimpanzees. Chimpanzees refrained from hunting associated red colobus groups, probably because diana monkeys are excellent sentinels for predators approaching over the forest floor. Thus several elements of both the predator's and the prey's strategies correspond to each other. Finally, we compared the interactions between the two species in Taï and in Gombe, Tanzania. We suggest that the difference in size ratio between the two species at the two sites and adaptation of hunting techniques and of escape modes to different forest structures can explain why Gombe red colobus attack chimpanzees while Taï red colobus try to escape. We conclude that predator-prey interactions can indeed lead to evolutionary arms races, with the specific form of co-adaptations depending on environmental factors.     

Acquisition of fission–fusion social organization in a chimpanzee (Pan troglodytes troglodytes) community released into the wild

The social aspects of primate reintroduction are of primary importance to the success of release programs and need to be assessed through the study of changes in social behavior over time. This study reports on the development of social structure and organization in a community of 37 chimpanzees (Pan troglodytes troglodytes) released into the wild in the Conkouati Douli National Park, Republic of Congo. Analyses of post-release monitoring data collected over 10 years on association patterns between individuals show that during the years following individual releases, chimpanzees exhibited changing social structure and organization until they stabilized in one community living in a fission–fusion system. Social organization development was directly affected by several factors including community size and experience in the wild, while social structure was affected by individual characteristics: gender, pre-release history, and release history. Similarities between social structure and organization observed in the released community and wild chimpanzee communities demonstrate that the release program is a success from a social point of view.     

Urban wild meat consumption and trade in central Amazonia

The switch from hunting wild meat for home consumption to supplying more lucrative city markets in Amazonia can adversely affect some game species. Despite this, information on the amounts of wild meat eaten in Amazonian cities is still limited. We estimated wild meat consumption rates in 5 cities in the State of Amazonas in Brazil through 1046 door-to-door household interviews conducted from 2004 to 2012. With these data, we modeled the relationship between wild meat use and a selection of socioeconomic indices. We then scaled up our model to determine the amounts of wild meat likely to be consumed annually in the 62 urban centers in central Amazonia. A total of 80.3% of all interviewees reported consuming wild meat during an average of 29.3 (CI 11.6) days per year. Most wild meat was reported as bought in local markets (80.1%) or hunted by a family member (14.9%). Twenty-one taxa were cited as consumed, mostly mammals (71.6%), followed by reptiles (23.2%) and then birds (5.2%). The declared frequency of wild meat consumption was positively correlated with the proportion of rural population as well as with the per capita gross domestic product of the municipality (administrative divisions) where the cities were seated. We estimated that as much as 10,691 t of wild meat might be consumed annually in the 62 urban centers within central Amazonia, the equivalent of 6.49 kg per person per year. In monetary terms, this amounts to US$21.72 per person per year or US$35.1 million overall, the latter figure is comparable to fish and timber production in the region. Given this magnitude of wild meat trade in central Amazonia, it is fundamental to integrate this activity into the formal economy and actively develop policies that allow the trade of more resilient taxa and restrict trade in species sensitive to hunting.     

Constraints on foraging success due to resource ecology limit colony productivity in social insects

The benefit of group living is a fundamental question in social evolution. For sociality to evolve, each individual must gain in terms of some fitness component by living in larger groups. However, in social insects, a decrease in per capita success in brood production has been observed in larger groups. While it has been proposed that this decrease could be outweighed by an increase in the predictability of success, a functional basis to this hypothesis has so far never been demonstrated. In this paper, using foraging economics as a functional proxy to colony productivity, we construct a model to explore how number of foragers in the colony interacts with the ecology of resources to influence per capita foraging success and its predictability. The results of the model show that there is no increase in per capita foraging success in larger colonies under most circumstances, though there is an increase in its predictability. We then test the model with empirical data on the foraging behavior of the primitively eusocial wasp, Ropalidia marginata. The consistency between the data and the model suggests that foraging economics could provide a robust functional basis in explaining the relationship between colony size and productivity.     

Bisexually bonded ranging in chimpanzees (Pan troglodytes verus)

Abstact While male mammals seek to maximize access to potential mates, females maximize feeding efficiency. Ranging patterns are therefore often sex specific. Sex-specific ranging patterns have also been reported for East African chimpanzees and a recent study on female ranging patterns concludes that social organization is best described by a male-bonded community model, where females occupy individual home ranges that are distributed within the boundaries of the male-defended range. In West African chimpanzees, however, such sex-specific ranging patterns have not been consistently observed and a bisexually bonded community model, where both sexes use the entire home range equally, has been suggested to best describe social organization. In this study we analyze 5 years of data on individual ranging patterns of chimpanzees in the Taï National Park (Côte dIvoire) to test specific predictions of the different models of chimpanzee social organization. We found that although males in Taï had slightly larger home ranges than females, all individual home ranges and core areas overlapped highly. Small individual home range size differences were entirely due to the use of peripheral areas and were correlated with female social dominance. These findings strongly support the bisexually bonded community model for Taï chimpanzees. Thus, we conclude that there are fundamental differences in the space-use patterns of East and West African chimpanzees and discuss possible factors leading to such differences.Communicated by D. Watts     

Between-group contest competition for food in a highly folivorous population of black and white colobus monkeys (Colobus guereza)

In theory, between-group contest (BGC) competition for food can greatly influence female social relationships and reproductive success in primates, but few studies have investigated whether such effects occur and, if so, under what ecological conditions. There is evidence that adult male black and white colobus monkeys (Colobus guereza, “guerezas”) defend the food their mates need against other groups, suggesting that BGC competition is important in this species. Using data on feeding, ranging, vegetation patterns, and intergroup encounters between six neighboring guereza groups, I provide evidence that the highly folivorous guerezas at Kanyawara, Kibale National Park, Uganda, engaged in BGC competition over unevenly dispersed, relatively high-quality feeding areas or “core areas”. Intergroup aggression was common, and groups’ home ranges overlapped. Groups were more likely to initiate high-level aggression if they encountered another group within or near their core area, and groups that initiated and won encounters often fed in the same areas in which losing groups had fed. Guerezas fed selectively on species with contagious (clumped) distributions and concentrated their feeding efforts in areas of the forest that contained the most food (core areas). Groups could be ranked in a linear dominance hierarchy, and group rank number was inversely related to the quantity and quality of food in groups’ core areas. This study not only provides good evidence that BGC competition occurs in primates but it also reinforces the idea that folivore food resources may be worth defending.     

Communal nesting is explained by subsequent mating rather than kinship or thermoregulation in the Siberian flying squirrel

Research on group living in animals is concentrated on highly social species, but studying less social species may hint at the factors possibly leading to the evolution of increased sociality. Thermoregulation is often thought to explain communal nesting in solitarily breeding mammals but also other factors may be involved. For example, it is observed that even solitary species may have cryptic kin cooperation. We studied factors affecting communal nesting in the Siberian flying squirrel. Flying squirrels breed solitarily but, similar to most other rodents, adults may sometimes huddle in groups. Communal nesting in flying squirrels was most frequent during winter and autumn, but also occurred during other seasons. This pattern was explained by the breeding season, which took place in the spring–summer, when communal nesting was less common. Neither monthly temperature, after accounting for breeding season, nor daily temperatures in winter explained communal nesting. Group size was small, two to three individuals. In most cases the group was a pair consisting of unrelated male and female, possibly indicating that group formation was related to mating behavior. This study contributes to the understanding of mammalian group formation in two major ways. First, our study contributes to the understanding of the role of relatedness in rodent group formation, demonstrating a case where close relatedness is not related to group formation. Second, our study indicates that in a solitarily breeding, rodent huddling may be more driven by other factors than temperature.