Individual and social behavioral responses to injury in wild toque macaques (Macaca Sinica)

Toque macaques (Macaca sinica),inhabiting natural forest at Polonnaruwa, Sri Lanka, are frequently injured in fights with conspecifics. The behavior of known individuals when they were injured was compared to that after they had recovered their health. Thus, injured animals rested and alloand autogroomed more, but they foraged less and initiated fewer aggressive episodes. They spent most time being sedentary in the safety of arboreal refuges and reduced acrobatic movements by locomoting more often terrestrially. Other group members showed no special tolerance (or altruism) toward injury victims during the costly and highly competitive activity of foraging for food. In fact, some injured animals received more aggression, or lost dominance rank, and thereby had their competitive abilities further impaired. Care for the injured was manifest mostly by grooming and wound cleaning. All hair in the area surrounding a wound, as well as dirt, scabs, and fly larvae, were removed, and saliva was applied by licking the wound (wounds so treated healed with no obvious signs of infection). (1) Injured macaques sought and received significantly more grooming (owing to wound care); (2) the amount so received increased with the severity of the injury; and (3) the initiative of other group members often compensated for a victim’s inability to solicit care. Juvenile males were especially attentive to injured adult males, suggesting that they were investing in a social bond with these adults, which might reciprocate altruism toward their juvenile caregivers in the future. Injured juvenile females received most care from their mothers.     

Spatial structure of hierarchical groups: testing for processes of aggregation,clustering, and spatial centrality in crayfish (Orconectes rusticus)

Competing group members tend to arrange in a social order that governs who will likely submit to whom. In many species the spatial distribution of individuals often reflects social status: dominants tend to occupy central locations while subordinates are often found along the group's periphery. This article explores the emergence of spatial consequences as a result of social rank differentiation. Rather than orienting centripetally, the movements of crayfish (Orconectes rusticus) primarily indicated a tendency to remain close to arena walls. Spatial locations were affected by the location of group members; but, rather than actively aggregating or clustering, individuals maintained a minimum distance. Previously established social rank did not affect spatial distributions. High population densities in the field are likely attributed to habitat constraints, rather than any social or centripetal tendencies of individual crayfish.     

Psychosocial and hormonal aspects of hierarchy formation in groups of male squirrel monkeys

Accounts of primate social dominance hierarchies often imply that the achievement of superior status is a “goal”, akin to a valued resource or commodity, and that hierarchies emerge in multimale groups from prolonged competitive conflicts over social status. This possibility is not consistent with our observations of five newly formed triads of adult male squirrel monkeys (Saimiri sciureus). Stable linear hierarchies based on clear asymmetries in the direction of intermale agonism and genital displays were established quickly, with virtually no reciprocal fighting, and in the absence of rank-related differences in plasma cortisol or testosterone. Although affiliative social overtures were initiated more often by high-ranking and middle-ranking males, affiliative overtures were directed equally often toward all members of each group. From the outset of the study all males, regardless of rank, spent an average of 33% of their time huddling in affiliative contact with male cagemates. These results suggest that in newly formed groups of adult male squirrel monkeys, social hierarchies reflect an expedient convention that reduces conflict and facilitates the formation of small cohesive groups. © 1994 Wiley-Liss, Inc.     

Decision-making Processes in Group Departures of Cattle

To keep social cohesiveness, group-living animals have to reach consensus decisions through recruitment processes. This implies that decision-making depends on the behaviours and social relationships of several group members at different stages of movements. We tested these assumptions in a group of fifteen 18-mo-old Charolais heifers ( Bos taurus ) at pasture, in which two observers continuously videotaped social interactions and group departures after resting periods. These departures were preceded by a phase of preparation characterized by an increase in activity. The number of heifers participating to a movement increased with the number of group members oriented in the direction of the movement before departure. The first moving animal also recruited a higher number of mates when it had a greater number of close neighbours, the first individuals to follow being mainly its preferential partners. Coercive interactions such as pressing behaviours were observed within the 5 min preceding or following departure. After departure, the numbers of walks and restarts of the first two movers were still operative in recruiting others. The frequency of pauses of the first mover was significantly higher when it was not followed, meaning that it adjusted its behaviour to that of other group members. Decision-making was distributed among group members, with any individual being liable to move first. The behaviour of cows and their spatial distribution before departure, at departure and after departure significantly affected the number of participants in the movement, demonstrating that decision-making was time-distributed in the studied cattle group.     

Shared or unshared consensus decision in macaques?

Members of a social group have to make collective decisions in order to synchronise their activities. In a shared consensus decision, all group members can take part in the decision whereas in an unshared consensus decision, one individual, usually a dominant member of the group, takes the decision for the rest of the group. It has been suggested that the type of decision-making of a species could be influenced by its social style. To investigate this further, we studied collective movements in two species with opposed social systems, the Tonkean macaque (Macaca tonkeana) and the rhesus macaque (Macaca mulatta). From our results, it appears that the decision to move is the result of the choices and actions of several individuals in both groups. However, this consensus decision involved nearly all group members in Tonkean macaques whereas dominant and old individuals took a prominent role in rhesus macaques. Thus, we suggest that Tonkean macaques display equally shared consensus decisions to move, whereas in the same context rhesus macaque exhibit partially shared consensus decisions. Such a difference in making a collective decision might be linked to the different social systems of the two studied species.     

Social Structure Affects Initiations of Group Movements but Not Recruitment Success in Japanese Macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>)

Research on collective movements has often focused on the sociodemographic parameters explaining the success of some individuals as leaders or initiators of collective movements. Several of these studies have shown the influence of social structure, through kinship and affiliative relationships, on the organization of collective movements. However, these studies have been conducted on semi-free-ranging groups of macaques that were not faced with a natural environment and its constraints. In the socially intolerant rhesus macaque (Macaca mulatta) the success of an initiator correlates with its hierarchical rank, the most dominant individuals being the most successful. We investigated the collective movements of another socially intolerant macaque species, Japanese macaques, in the wild, to assess whether the social structure was still a determinant factor under natural conditions. In line with previous studies of macaques, we found that social structure drove the organization of collective movements. More dominant individuals initiated more collective movements. However, dominance did not affect the success of an initiation, i.e., the number of individuals joining. In addition, kinship strongly constrained the associations observed in females during collective movements. These results reflect the social structure of Japanese macaques, in which strong power asymmetry and kinship relationships constrain the majority of interactions between individuals within the group. Moreover, these results are similar to those observed in semi-free-ranging rhesus macaques and support the hypothesis of an effect of social determinants on collective movements of primates even under natural conditions.     

Social correlates of variation in urinary cortisol in wild male bonobos (Pan paniscus)

Cortisol excretion in males of group living species is often associated with social rank and competition for oestrous females. Rank-related patterns of cortisol levels can be used to study mechanisms of rank maintenance and costs associated with mate competition. Bonobos (Pan paniscus) are interesting because males form a linear dominance hierarchy but are not dominant over females and therefore aggressive male-male competition over access to females alone is not considered to be a successful reproductive strategy. In this study on social correlates of urinary cortisol in wild male bonobos, we investigated the relationship between cortisol levels and several aspects of mate competition, including male rank, aggression rates, and association time with oestrous females. We found that cortisol levels correlated positively with dominance rank when oestrous females were present, but not when they were absent. This result is consistent with the idea that aggressive behaviour plays a minor role in maintenance of high rank. While aggression received from males and females explained within-individual variation in cortisol levels, it was the time spent in association with oestrous females that best explained between-individual variation in male cortisol levels. The observed increase in male cortisol may be associated with spatial proximity to oestrous females and could result from anticipated aggression from other group members, reduced feeding time in the males, or a combination of both.     

Sex-reversed correlation between stress levels and dominance rank in a captive non-breeder flock of crows

Group living has both benefits and costs to individuals; benefits include efficient acquisition of resources, and costs include stress from social conflicts among group members. Such social challenges result in hierarchical dominance ranking among group members as a solution to avoid escalating conflict that causes different levels of basal stress between individuals at different ranks. Stress-associated glucocorticoid (corticosterone in rodents and birds; CORT) levels are known to correlate with dominance rank in diverse taxa and to covary with various social factors, such as sex and dominance maintenance styles. Although there is much evidence for sex differences in the basal levels of CORT in various species, the correlation of sex differences in basal CORT with dominance rank is poorly understood. We investigated the correlation between CORT metabolites (CM) in the droppings and social factors, including rank and sex, in a captive non-breeder group of crows. In this group, all the single males dominated all the single females, and dominance ranks were stable among single males but relatively unstable among single females. CM levels and rank were significantly correlated in a sex-reversed fashion: males at higher rank (i.e., more dominant) had higher CM, whereas females at higher rank exhibited lower CM. This is the first evidence of sex-reversed patterns of CM–rank correlation in birds. The results suggest that different mechanisms of stress–dominance relationships operate on the sexes in non-breeder crow aggregations; in males, stress is associated with the cost of aggressive displays, whereas females experience subordination stress due to males' overt aggression.     

Social Dominance Hierarchies and Phenotypic Correlates of Dominance in Captive Groups of the Grey‐cheeked Fulvetta Alcippe morrisonia in Taiwan

Dominance hierarchies usually form quickly among avian foraging groups because they are beneficial to most individuals by reducing conflict. Several characteristics that correlate with dominance rank have been identified in birds, but most of these conclusions rely on studies of temperate species. Hence, we studied whether captive group members of a subtropical species, grey‐cheeked fulvetta Alcippe morrisonia, form social dominance hierarchies when competing for food during the non‐breeding season. We also investigated whether sex, age, body condition and fat score were related to an individual's dominance rank which was established by counting aggressive interactions in six captive groups of nine individuals each. In all groups, linear dominance hierarchies were formed whereby yearlings dominated over adult birds, and individuals with a better body condition were also more dominant, while sex and fat score had no discernable effect. Male yearlings had significantly higher body masses and body condition indices than male adults, while female yearlings had significantly higher body masses, body condition indices and fat scores than female adults. However, there were no significant differences between male and female yearlings or adults for any of these variables. We suggest possible reasons for the dominance of yearlings, such as captive conditions or the higher body weight of yearlings.     

秦岭羚牛群体分离与重组的变化

对4只佩戴无线电颈圈的羚牛进行直接观察的结果表明,羚牛群体的结构和组成是经常发生变化的。在羚牛各集群类型中,家群的稳定性最高,社群次之,混合群较差。虽然家群具有较高的稳定性,但是除母牛及其幼仔外,家群的其它成员也经常变化,羚牛的亚成体也不是构成家群的稳定成员。繁殖是羚牛集群类型变化的主要原因之一。在繁殖期中健壮的成年雄性个体因寻找配偶离开原群,在各群间游荡,成为羚牛群中最不稳定的成员。亚成体在其母亲产下幼仔后可能会中断与母牛间的关系,进入混合群和社群。羚牛反捕食策略和护幼行为也会影响羚牛群的分群和重组。     

Relatedness of matrilines,dispersing males and social groups in long-tailed macaques (Macaca fascicularis).

Genealogical relatedness is thought to be an important causal factor in the evolution of cooperation. We inferred relatedness on the basis of 11 blood protein markers using the Queller and Goodnight index of relatedness in a macaque population with long-term demographic records. This estimate reflected independently determined pedigree relationships in our data set. Mean relatedness among all members of a social group was 0.10 but much higher levels of relatedness (0.30-0.47) were found among the members of matrilineal families with a high or intermediate social rank. Groups of dispersing males that had been born into the same social group were sometimes closely related (0.43 and 0.58), but they could also be less related (0.08). We found that the pattern of distribution of relatedness was associated with gene flow and differential reproduction in males, rather than with group fission and the presence of geographical barriers.     

Sex differences in leadership during group movement in mantled howler monkeys (Alouatta palliata)

Benefits of group life depend in large part on whether animals remain cohesive, which often requires collective decisions about where and when to move. During a group movement, the leader may be considered as the individual occupying the vanguard position of the group progression, when its movement evokes following by other group members. In nondespotic societies, individuals with greater incentives to move frequently are leaders. During 15 months of observations (1,712 contact hours), we investigated two mantled howler monkey (Alouatta palliata) groups at La Flor de Catemaco (Los Tuxtlas, Mexico) to examine whether sex and female reproductive state influenced leadership likelihood in two contexts: movements toward feeding trees; movements associated with loud calls, a group-defense behavior used by males of this genus. Females led and occupied forward positions during group movements toward feeding trees more often than adult males. Adult females led these movements more frequently when they were gestating than when they were lactating or cycling. There were no differences between sexes in the leadership of group movements associated with loud calls. Leadership by gestating females is perhaps the result of their higher nutritional/energetic needs when compared with cycling females, and of their greater mobility when compared with lactating females carrying dependent offspring. Female leadership during movements toward feeding trees may be a mechanism to optimize access to food resources in mantled howler monkeys.     

The physiological effects of social status in the cooperatively breeding cichlid Neolamprologus pulcher

The physiological effects of social rank were examined in three different experiments with Neolamprologus pulcher a cooperatively breeding cichlid, endemic to Lake Tanganyika, East Africa. The effects of rank on physiology between pairs of dominant and subordinate size‐matched fish (experiment 1) and among groups of four size‐matched fish (experiment 2) were examined. A third experiment mimicked the natural social structure in the wild; pairs were observed with other group members including breeders. The effect of social position was investigated on growth rates, liver concentrations of adenosine triphospate (ATP), lipids, proteins, creatine phosphate (CrP), glucose and glycogen as well as plasma cortisol. In naturalistic group settings, dominants displayed higher levels of liver protein and plasma cortisol. In the absence of breeders, dominant individuals (of helper pairs) had higher liver glycogen levels and dominant fish (held in groups of four) grew most. These results support previous cooperatively breeding mammal studies and suggest that dominant individuals experience higher cortisol levels as well as higher growth rates.     

Alloparental behavior in a captive group of spider monkeys (Ateles geoffroyi) at the Auckland zoo

I studied alloparental behavior in a captive group of spider monkeys at the Auckland Zoo using seven infants as focal subjects and assessed the effects of age, sex, and reproductive status of alloparents on patterns of infant-other interaction. Adult males initiated interactions with infants most often, followed by adult females. Immature individuals interacted with infants infrequently. Infants themselves initiated contact with adult males more often than with other members of the group. Alloparental behavior in spider monkeys differs from that in most other species in that the infant is an active rather than a passive participant in alloparental interactions. I discuss the patterns of infant-other interaction in relation to the social structure and dispersal patterns of Ateles.     

How many for dinner? Recruitment and monitoring by glances in capuchins

Group members present physical and physiological differences according to their age, sex or social status, which could generate motivation differences among individuals during travel. In spite of these divergences of interest among individuals, the group succeeds more often than not in making a collective decision about departure time and which direction to take. To reach a consensus decision, animals should exchange information relating to characteristics of group movement through different communication channels. The main purpose of this study is to understand the function of behaviour patterns displayed during movements of white-faced capuchins (Cebus capucinus). We designed experiments in which we provoked collective movements involving a binary choice. During experiments, a video camera recorded the behaviour of each capuchin, which enabled us to determine which individuals displayed a behavioural pattern during travel and how this behaviour influenced the other group members. We found that looking backwards seemed to permit the recruitment of group mates during collective movement. This behaviour also seemed to allow the quantification of the number of followers, since the emitter modified its locomotion speed according to this number. In this preliminary study, we showed that visual behaviour was used to recruit and monitor group mates during collective movements and provided information on mechanisms involved in maintaining cohesion and coordination among group members during travel.     

Personality and Affinities Play a Key Role in the Organisation of Collective Movements in a Group of Domestic Horses

Understanding how groups of individuals with different motives come to daily decisions about the exploitation of their environment is a key question in animal behaviour. While interindividual differences are often seen only as a threat to group cohesion, growing evidence shows that they may to some extent facilitate effective collective action. Recent studies suggest that personality differences influence how individuals are attracted to conspecifics and affect their behaviour as an initiator or a follower. However, most of the existing studies are limited to a few taxa, mainly social fish and arthropods. Horses are social herbivores that live in long‐lasting groups and show identifiable personality differences between individuals. We studied a group of 38 individuals living in a 30‐ha hilly pasture. Over 200 h, we sought to identify how far individual differences such as personality and affinity distribution affect the dynamic of their collective movements. First, we report that individuals distribute their relationships according to similar personality and hierarchical rank. This is the first study that demonstrates a positive assortment between unrelated individuals according to personality in a mammal species. Second, we measured individual propensity to initiate and found that bold individuals initiated more often than shy individuals. However, their success in terms of number of followers and joining duration did not depend on their individual characteristics. Moreover, joining process is influenced by social network, with preferred partners following each other and bolder individuals being located more often at the front of the movement. Our results illustrate the importance of taking into account interindividual behavioural differences in studies of social behaviours.     

The benefits of social capital: close social bonds among female baboons enhance offspring survival

Sociality has evolved in many animal taxa, but primates are unusual because they establish highly differentiated bonds with other group members. Such bonds are particularly pronounced among females in species like baboons, with female philopatry and male dispersal. These relationships seem to confer a number of short-term benefits on females, and sociality enhances infant survival in some populations. However, the long-term consequences of social bonds among adult females have not been well established. Here we provide the first direct evidence that social relationships among female baboons convey fitness benefits. In a group of free-ranging baboons, Papio cynocephalus ursinus, the offspring of females who formed strong social bonds with other females lived significantly longer than the offspring of females who formed weaker social bonds. These survival benefits were independent of maternal dominance rank and number of kin and extended into offspring adulthood. In particular, females who formed stronger bonds with their mothers and adult daughters experienced higher offspring survival rates than females who formed weaker bonds. For females lacking mothers or adult daughters, offspring survival was closely linked to bonds between maternal sisters. These results parallel those from human studies, which show that greater social integration is generally associated with reduced mortality and better physical and mental health, particularly for women.     

Consensus decision making in animals

Individual animals routinely face decisions that are crucial to their fitness. In social species, however, many of these decisions need to be made jointly with other group members because the group will split apart unless a consensus is reached. Here, we review empirical and theoretical studies of consensus decision making, and place them in a coherent framework. In particular, we classify consensus decisions according to the degree to which they involve conflict of interest between group members, and whether they involve either local or global communication; we ask, for different categories of consensus decision, who makes the decision, what are the underlying mechanisms, and what are the functional consequences. We conclude that consensus decision making is common in non-human animals, and that cooperation between group members in the decision-making process is likely to be the norm, even when the decision involves significant conflict of interest.     

Multilevel selection and the social transmission of behavior

Many evolutionary models assume that behaviors are caused directly by genes. An implication is that behavioral uniformity should be found only in groups that are genetically uniform. Yet, the members of human social groups often behave in a uniform fashion, despite the fact that they are genetically diverse. Behavioral uniformity can occur through a variety of psychological mechanisms and social processes, such as imitation, consensus decision making, or the imposition of social norms. We present a series of models in which genes code for social transmission rules, which in turn govern the behaviors that are adopted. Transmission rules can evolve in randomly formed groups that concentrate phenotypic variation at the between-group level, favoring the evolution of altruistic behaviors and other group-advantageous traits. In addition, a direct bias toward adopting altruistic behaviors can evolve. Our models begin to show how group selection can be a strong force in human evolution, despite the absence of extreme genetic variation among groups.     

Decision-Making in Przewalski Horses (Equus ferus przewalskii) is Driven by the Ecological Contexts of Collective Movements

We addressed decision-making processes in the collective movements of two groups of Przewalski horses ( Equus ferus przewalskii ) living in a semi free-ranging population. We investigated whether different patterns of group movement are related to certain ecological contexts (habitat use and group activity) and analysed the possible decision-making processes involved. We found two distinct patterns; 'single-bout' and 'multiple-bout' movements occurred in both study groups. The movements were defined by the occurrence of collective stops between bouts and differed by their duration, distance covered and ecological context. For both movements, we found that a preliminary period involving several horses occurred before departure. In single-bout movements, all group members rapidly joined the first moving horse, independently of the preliminary period. In multiple-bout movements, however, the joining process was longer; in particular when the number of decision-makers and their pre-departure behaviour before departure increased. Multiple-bout movements were more often used by horses to switch habitats and activities. This observation demonstrates that the horses need more time to resolve motivational conflicts before these departures. We conclude that decision-making in Przewalski horses is based on a shared consensus process driven by ecological determinants.