Ontogenetic changes and the stability of rhesus monkey dominance relationships

Dominance relationships were studied in a rhesus monkey group during five consecutive years. The group consisted of eight stable matriarchies and an adult male class which was replaced at the start, and again at the midpoint, of the study. Immature males were selectively harvested to maintain a sex ratio typical of natural troops. Maximum group size during the study was 77 animals.Dominance relationships were remarkably stable, with only 4.4% of dyads failing to show unidirectional relationships. Despite this stability, a linear ranking of all group members was not possible. Male dominance relationships with other males were among the most stable, following the fighting which ensued on male introductions. Male introductions did not disrupt female dominance relationships.Adult female dominance relationships were also quite stable, but immature females slowly achieved dominance over older sisters and females subordinate to their mothers. Such reversals were the result of processes lasting over many months. Many dominance assertions occurred prior to puberty but a significant number occurred following sexual maturity. Maturing females did not reverse dominance relationships according to any particular hierarchial order and, as a consequence, many were subordinate to animals that were dominated by others that they dominated.Although there was an alpha male that was dominant to all animals in the group, adult females dominated most adult males. Adult males, however, often reciprocated aggression directed at them. They almost invariably threatened or countercharged aggressive immature animals regardless of matriarchial membership. Adult males dominated some adult and most young females, even in families containing matriarchs and adult females to which the adult males always submitted.The dominance relationships of young males were similar to those of their sisters, until puberty. Young males did not necessarily bypass adult males that their mothers outranked, and often failed to win against adult females that their mothers dominated. Adolescent female aggression against females is seldom interfered with by adult males, and females may actively aid one another against males. In contrast, the aggression of young males often elicits interference by adult males, and young males often become the targets of redirected aggression in the group. As a consequence, whereas young females rise in rank to positions adjacent to their mothers, adolescent males often suffer losses to animals that they had dominated as juveniles.     

Food competition in captive female sooty mangabeys (Cercocebus torquatus atys)

We studied the social and foraging behavior of two captive groups of sooty mangabeys under two different spatial food situations. These food conditions were clumped (food was placed in a box) and dispersed (food was dispersed over the entire enclosure). In each group five adult females and two adult males were observed. As a criterion for food competition, individual differences in the relative food intake were used. Adult female mangabeys had a linear, stable, and unidirectional dominance hierarchy. Access to food was rank dependent among females only under clumped food distribution, as current models of the evolution of primate social systems predict. However, feeding success appeared to be mediated not by female but by male agonistic behavior toward females. High-ranking females received relatively less aggression from males and could, therefore, stay and feed longer in the feeding area. Male tolerance of higher-ranking females seems to mediate female feeding success under restricted food resources. The establishment of a special relationship with a high-ranking male might, therefore, be a strategy to get better access to food. This study demonstrates that female competition for access to food should not be analyzed separately from male influences on females and suggests that a more integral role of males in socioecological models of the evolution of primate social systems should be considered.     

Activity and ranging patterns of Colombian woolly monkeys in north-western Amazonia

In this study, I revise three aspects of the socioecology of woolly monkeys (genus Lagothrix) that might give us a better understanding of the patterns found in this species: (1) the association between temporal variation in fruit abundance and diet, activity, and ranging patterns; (2) the individual trade-offs associated with living in small or large groups, and (3) the relationship between social dominance and foraging success. Using behavioral and ecological data collected during 3 years in Tinigua Park, Colombia, I found that woolly monkeys tend to avoid open-degraded forests, where fruit production is generally lower than it is in mature forests. Diet and activity budgets were highly associated with temporal patterns of fruit production. Daily path length was positively correlated with group size and monthly fruit abundance, and negatively correlated with habitat quality. I found differences in activity budgets and the diet preferences of different age/sex classes. For example, adult males rest more and juveniles play more than other classes. Juveniles and adult females without infants look for arthropods more often than adult males and females with young infants, who showed the highest frequencies of fruit feeding. Dominant adult males were not consistently the most efficient foragers on fruits according to two different indexes. Most of these results are consistent with the expectations from strong intra-group competition for resources. However, females with infants received benefits during feeding similar to those of dominant adult males, which may be mediated by differential aggression from males to other group members (juveniles and females without infants).     

Feeding and locomotor rhythm of the ornamental cichlid fish Amatitlania sp. in different social groups

The aim of this study was to evaluate the feeding activity and the rhythm of daily locomotor activity of the convict cichlid (Amatitlania sp.) kept in different social groups under a self-feeding system. A total of 120 animals was distributed among six repetitions of four social groups, as follows: group 1 with one male and one female per tank; group 2 with three males and three females per tank; group 3 with six males per tank; and group 4 with six females per tank. Feeding activity (FA) and locomotor activity (LA) were evaluated using photoelectric presence-sensors connected to automatic feeders. The fish were fed a commercial extruded diet (46% crude protein and 3600 kcal kg⁻¹ of digestible energy). Animal growth was evaluated for all groups. After 30 days of experimentation, the fish stabilized their demands by adjusting their consumption. Amatitlania sp. showed predominantly diurnal FA and LA. All groups showed a peak of activity when the light was turned on and when it was turned off. In summary, FA and LA of Amatitlania sp. are predominantly diurnal and independent of social group. Pairs and groups of males and females together consume less food in relation to groups of one sex or the other due to reproductive behaviour. On the other hand, groups of only males or females consume more food because they lack reproductive stimuli and thus prioritize growth. These results may support good feeding management practices for this ornamental cichlid. Studies relating feeding behaviour with different social groups are of great importance for determining effective feeding strategies for this species in captivity. Thus, such a study assists in a more efficient production of Amatitlania sp.     

Female-mediated causes and consequences of status change in a social fish

In highly social species, dominant individuals often monopolize reproduction, resulting in reproductive investment that is status dependent. Yet, for subordinates, who typically invest less in reproduction, social status can change and opportunities to ascend to dominant social positions are presented suddenly, requiring abrupt changes in behaviour and physiology. In this study, we examined male reproductive anatomy, physiology and behaviour following experimental manipulations of social status in the cooperatively breeding cichlid fish, Neolamprologus pulcher. This unusual fish species lives in permanent social groups composed of a dominant breeding pair and 1-20 subordinates that form a linear social dominance hierarchy. By removing male breeders, we created 18 breeding vacancies and thus provided an opportunity for subordinate males to ascend in status. Dominant females play an important role in regulating status change, as males successfully ascended to breeder status only when they were slightly larger than the female breeder in their social group. Ascending males rapidly assumed behavioural dominance, demonstrated elevated gonadal investment and androgen concentrations compared with males remaining socially subordinate. Interestingly, to increase gonadal investment ascending males appeared to temporarily restrain somatic growth. These results highlight the complex interactions between social status, reproductive physiology and group dynamics, and underscore a convergent pattern of reproductive investment among highly social, cooperative species.     

Agonistic Interactions of Juvenile Savanna Baboons

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

The Social Organization of the Cape Ground Squirrel (Xerus inauris; Rodentia: Sciuridae)

The Cape ground squirrel (Xerus inauris) of southern Africa is a tropical species that does not hibernate. Field observations using scan and all-occurrence sampling revealed that this species was highly social. Female Cape ground squirrels formed social units of related females and their subadult young, as is typical for other ground squirrels. Female social groups were usually composed of 2–3 adult females and 2–3 subadults of either sex. Members of these female social groups shared sleeping burrows and feeding ranges. Female social groups did not cooperatively defend their feeding ranges from adjacent groups in other burrow clusters. Interactions within female social groups were highly amicable, and no dominance hierarchy was evident. Males in this species also lived in groups. These all-male bands of up to 19 individuals lived almost independently from female groups. The entire male band shared one home range, although ephemeral sub-bands were formed daily. The composition and size of these sub-bands changed daily. Interactions among males, which were largely amicable, included allogrooming and sleeping together. Analysis of interactions within the band indicated a stable, linear, dominance hierarchy among males. Dispersal in this species appeared to be male biased as is typical of other ground-dwelling squirrels, with males dispersing at reproductive maturity. Males joining male bands were thus dispersers and were not likely to be closely related. Sociality in the Cape ground squirrel may be summarized as highly social female kin clusters and associated social non-kin bands of males.     

Serum testosterone, male dominance, and aggression in captive groups of vervet monkeys (Cercopithecus aethiops sabaeus)

The relationship of serum testosterone concentration to male dominance rank and frequency of aggression was investigated in stable vervet monkey social groups, each containing two or three adult males, several adult females, and their offspring. Dominance relationships were determined by noting an animal's success in intermale aggressive encounters. A striking finding was the marked within-subject variation in testosterone concentration: 5- to 10-fold fluctuations were often observed on successive days. When all 15 groups were considered together, testosterone concentration was unrelated to dominance rank. Although mean testosterone concentration for all dominant males was higher than the mean for all subordinate males, this difference was not significant. In a subset of 4 groups, the rate of aggression initiated was significantly correlated with same-day testosterone in dominant but not in subordinate males.     

Group structure,within-group conflict and reproductive tactics in cooperatively breeding Galápagos mockingbirds,Nesomimus parvulus

Reproductive conflict within groups can be an important feature of cooperative breeding systems, especially when more than one individual of a sex breeds within a social group. Relationships between group structure, dominance, within-group conflict and reproductive tactics of cooperatively breeding Galápagos mockingbirds were examined on Isla Genovesa. Territorial groups of 2–24 adults included up to three breeding females, with 42% of the groups containing more than one (plural groups); females in most plural groups nested separately. Territory size increased with group size, but the area available per pair in plural groups was smaller than in singular groups (groups with only one breeding pair). Most pairings were monogamous, and males usually outnumbered females; high-ranking males obtained mates more frequently than subordinate males. In 3 relatively dry years, but not in a wet El Niño year, subordinate pairs in plural groups fledged fewer young than dominant pairs or pairs breeding in singular groups. Interference by dominant breeders, often leading to abandonment of nests by subordinate pairs, appears to account for these differences: through nest disruption in drier years, dominant individuals may reduce the cost of sharing their territories and increase the chances of recruiting helpers. Dominant males in plural groups may also father young through extra-pair copulations with subordinate females. Despite costs imposed by within-group conflict, subordinate breeders have higher long-term reproductive success than birds that defer breeding. Plural group structure is maintained because unpredictable climatic variation favours opportunistic breeding by subordinates.     

Rank,relationships, and responses to intruders among adult male vervet monkeys

We examined the influences of dyadic relationships among captive adult male vervet monkeys (Cercopithecus aethiops sabaeus) on behavior directed toward caged “intruder” males placed inside subjects' enclosures. Subjects were all 9 adult male residents from three stable social groups, each of which contained 3 adult males, at least 3 adult females, and their immature offspring. Every male was observed in two 3-hour sessions, each time with one of the 2 other adult males from his group. Observation sessions consisted of six consecutive 30-min stages in which group composition and the presence of the intruder were manipulated. All groups exhibited a stable, linear male dominance hierarchy prior to and throughout the study. In each group, there was one pair of males, when together, in which each member exhibited higher rates of intruder-directed approach and aggressive behaviors than when either animal was paired with the third male of his social group. Such pairs were also distinguished by high levels of within-pair agonistic interactions. The higher-ranking member of each dyad was the most aggressive male toward the intruder in his social group, although only one of these animals was the dominant male of his group. Mutual facilitation of aggression against intruding males is interpreted as cooperative behavior benefitting both males by increasing the likelihood of repelling a potential competitor for resident females. Such cooperation provides further evidence in nonhuman primates for cohesive male-male dyads between animals whose social interactions are characterized by agonism. © 1993 Wiley-Liss, Inc.     

White Flank Spots Signal Feeding Dominance in Female Diamond Firetails,Stagonopleura guttata

Plumage colour can be used as an honest signal to convey health and status, which has traditionally been examined in the sexual selection context of choosy females and elaborate males. We use a model avian system to study the role of plumage coloration in a social context such as inter‐ and intrasexual competition over food resources. The diamond firetail (Stagonopleura guttata) is an endemic Australian finch: females have more white flank spots than males, and white spot number was correlated with cell‐mediated immune response in females. We use two experimental designs to test the role of white flank spots for feeding dominance and dominance discrimination in a group‐living bird. The results from two‐choice trials and from single‐arena trials showed that female ornamentation was consistently important in social food contests, and males consistently responded to female spot number. Females with higher spot number fed first, in trials with males and/or females. Also, females preferred to feed next to test birds with low spot number, but males showed no preference for feeding next to birds with few or many spots. Finally, latency to feed was predicted by spot number: both males and females had longer latency to feed if test birds had more spots than the focal birds. We conclude that female, but not male, ornamentation was important for inter‐ and intrasexual food competition. This is one of the very few studies to show that the same plumage ornament can have a different function between the sexes as a signal of social status. Moreover, this study shows that white plumage can be a signal of dominance.     

Dominance and feeding competition in captive rhesus monkeys

The feeding behavior of 16 adult female rhesus monkeys living in three captive social groups was observed. Estimates of relative food intake, feeding rate, and location of feeding in relation to food sources were compared between females of different dominance ranks. Higher-ranking females had greater access to feeding sites and were supplanted or threatened less frequently while feeding than subordinates. However, no consistent differences in estimates of total intake were found between females of high and females of low rank. The effects of dominance on feeding behavior were most pronounced in the group receiving the least food relative to estimates of overall group nutritional requirements. Higher-ranking females, both over the long term and during the study period, tended to produce more surviving offspring. The effects of dominance on reproductive performance appeared to be less related to food intake than to competitive and aggressive interactions, potentially resulting in higher levels of stress for subordinates.     

Social grooming in Tonkean macaques (Macaca tonkeana)

The distribution, form, and contexts of occurrence of social grooming were studied in two captive groups of Tonkean macaques (Macaca tonkeana), using interaction-dependent sampling. The social events surrounding grooming had little influence on its form, the participants' behavior being shaped mainly by physical constraints. Adult females were most often involved in grooming interactions. Grooming between adult females appeared more intimate than that between adult males and females. Kinship and dominance had no effect on the form or distribution of social grooming among adult females. It is concluded that social systems that are characterized by mild dominance relations allow individuals the freedom to interact in the way and with whom they wish.     

定量评估豢养长江江豚个体间的优势关系

2000年1—4月和2000年4月—2002年4月,分别连续3个月和2年观察了中国科学院水生生物研究所白鱀豚馆6头长江江豚（3雌3雄,8个不同组合群）个体间发生的3种交互关系,即接近—逃开、接近—接触和接近—尾鳍击打。在累计8,162min的观察时间内,共观察到这些交互关系1,685次。本研究将江豚个体间发生的这3种交互关系作为个体间优势关系的指示,并分别给交互双方赋予不同的分值,采用优势比分矩阵方法定量分析了江豚个体之间的优势关系。结果表明江豚群体内存在优势关系。同性别个体之间,优势关系与年龄和饲养时间有关,年龄大的个体比年龄小的个体具有优势,饲养时间短的个体比饲养时间长的个体具有优势。但是,异性成年个体之间,优势关系通常是可变的。     

Effects of age and sex on intra-group spacing behaviour in juvenile savannah baboons,Papio cynocephalus cynocephalus

Twenty juvenile members of known genealogies in two baboon groups were studied over a 16-month period to evaluate a number of predictions about juvenile spacing behaviour based on the natural history of savannah baboons. Young juveniles (1–2·5 years old) approached more frequently and spent more time in proximity to other group members than did old juveniles (3–5·5 years old). In particular, young juveniles associated more closely with their mothers, particular adult males (possible fathers) and age-peers than did old juveniles. Approaches of young juveniles towards unrelated, high-ranking adults were more likely to occur during feeding than were those of old juveniles. Also, following such an approach, young juveniles were more likely than old juveniles to begin feeding immediately. The overall rates of feeding of old juveniles were depressed when they were in proximity to unrelated, high-ranking adults, whereas the feeding rates of young juveniles were not. Juvenile males approached adult males more often than did juvenile females. Juvenile females approached unrelated adult females more often than did juvenile males. Sex differences also existed in juveniles' choices of unrelated adult female neighbours. Juvenile females associated most often with lactating females, whereas juvenile males associated primarily with cycling females. During group resting, juvenile females approached adult females from higher-ranking matrilines more often than they approached adult females from lower-ranking matrilines. Juvenile males did not exhibit this attraction. Also, among old juveniles, females associated closely with their mothers, whereas males did not. Taken together, the results support the hypotheses that juvenile baboons associate with group members in ways that (1) enhance the probability of surviving an early period of high mortality, (2) create opportunities for social learning of sex-typical behaviours/skills, and, for females, (3) facilitate acquisition of familial dominance status.     

Agonistic Interactions and Matrifocal Dominance Rank of Wild Bonobos (Pan paniscus) at Wamba

I studied dominance relations in a wild group of bonobos at Wamba, Democratic Republic of Congo. Although agonistic interactions between males occurred frequently, most of them consisted only of display, and physical attacks were infrequent. Dominance rank order seemed to exist among males, but its linearity is unclear. Dominant males rarely disturbed copulatory behavior by subordinate males. However, high-ranking males usually stayed in the central position of the mixed party and, so, would have more chance of access to estrous females. Among females, older individuals tended to be dominant over younger individuals. However, agonistic interactions between females occurred rather infrequently, and most consisted of displacement without any overt aggressive behavior. Dominance between males and females is unclear, but females tended to have priority of access to food. The close social status between males and females may be related to the prolonged estrus of females and their close aggregation during ranging. Existence of a male's mother in the group and her dominance status among females seemed to influence his dominance rank among males. Young adult males whose mothers were alive in the group tended to have high status. In some cases, change in dominance between high-ranking males was preceded by a corresponding change in dominance between their mothers. As the dominance status of females is similar to that of males, mothers may be able to support their sons to achieve high status, stay in the center of the mixed party, and so have greater access to females, which may maximize the number of descendants of the mothers.     

Female dominance rank and behavior during artificial menstrual cycles in social groups of rhesus monkeys (Macaca mulatta)

We recently reported (Michael & Zumpe: American Journal of Primatology 15:157–170, 1988) evidence for the hypothesis that copulations by pairs of rhesus monkeys are linked more closely to the ovulatory phase of the female's menstrual cycle when a male has access to several females in different cycle phases rather than access to females in the same cycle phase. We have now used data from this earlier study to assess the role of female dominance. Artificial menstrual cycles were induced with hormone injections in two of four ovariectomized females in eight social groups (each consisting of one male and four females). The artificial cycles were either synchronized or offset by 7-day increments. There were differences in the behavioral interactions of the higher-ranking (dominant) and lower-ranking (subordinate) hormone-treated females in each group. The amount of male sexual activity received by dominant and subordinate females differed little, but dominant females initiated significantly more sexual activity (P<.0001) than did subordinate females. Dominant females were significantly more aggressive (P<.0001) toward other females and received more direct and redirected aggression from males. Males generally initiated and maintained more social contacts with dominant than with subordinate females, and dominant females were less active than subordinates in initiating and maintaining social interactions with males. The findings provide new evidence for both female mate competition and mate choice by males for higher-ranking females.     

Spatial and Social Sexual Segregation Patterns in Indo-Pacific Bottlenose Dolphins (Tursiops aduncus)

Sexual segregation seems to be common in bottlenose dolphins, whereby males and females live in different pods that mix mainly for mating. Male dolphins often use aggressive behaviour to mate with females, while females with calves may have different activity and dietary requirements to males and different susceptibility to predation. We investigated the degree of spatial and social sexual segregation in Indo-Pacific bottlenose dolphins (Tursiops aduncus) in a subtropical estuary in Australia. Based on surveys completed over three years, dolphin groups were mostly mixed-sex or female. Mixed-sex groups were found in larger groups in mostly deeper water, whereas, female groups were foraging across all water depths in smaller groups. Aggressive coercive behaviour by males towards females was high, occurring mainly in deeper water, at higher tides, and outside the breeding season. Habitat use by female dolphin groups suggests that shallow tributaries may provide a sanctuary from aggressive males, access to suitable prey items and density for mothers and their calves, or a combination of these factors.     

Female Coordination of Group Travel in Wild Propithecus and Eulemur

Coordination of primate group movements by individual group members is generally categorized as leadership behavior, which entails several steps: deciding where to move next, initiating travel, and leading a group between food, water sources, and rest sites. Presumably, leaders are able to influence their daily foraging efficiency and nutritional intake, which could influence an individual's feeding ecology and long-term reproductive success. Within anthropoid species, females lead group movements in most female-bonded groups, while males lead groups in most nonfemale-bonded groups. Group leadership has not been described for social prosimians, which are typically not female-bonded. We describe group movements in two nonfemale-bonded, lemurid species living in southeastern Madagascar, Propithecus diadema edwardsi and Eulemur fulvus rufus. Although several social lemurids exhibit female dominance Eulemur fulvus rufus does not, and evidence for female dominance is equivocal in Propithecus diadema edwardsi. Given the ecological stresses that females face during reproduction, we predict that females in these two species will implement alternative behavioral strategies such as group leadership in conjunction with, or in the absence of, dominance interactions to improve access to food. We found that females in both species initiated and led group movements significantly more often than males did. In groups with multiple females, one female was primarily responsible for initiating and leading group movements. We conclude that female nutritional needs may determine ranging behavior to a large extent in these prosimian species, at least during months of gestation and lactation.     

Effect of Dominance,Reproductive State,and Group Size on Body Mass in Macaca radiata

Body mass can impact reproductive performance in males and females. In nonhuman primates the relationship is often mediated by dominance. We measured body mass monthly in a provisioned group of bonnet macaques (Macaca radiata) living at a Hindu temple. We also measured body mass on 3 occasions in a wild population of bonnet macaques. In the temple group, females that reproduced lost body mass, while females that did not reproduce gained body mass. Mass loss among females occurred primarily while they were nursing. Adult males from the temple group lost mass during the mating season and gained it during the non-mating season. Subadult males experienced less seasonal variation in body mass. Body mass and changes in mass were not related to dominance rank in either the temple or the forest group. Furthermore, maternal dominance rank did not affect infant mass. Females from the smallest forest group weighed significantly less than females from the two larger forest groups, which suggests intergroup competition in the population. Body mass was not related to dominance rank in a straightforward manner but may indirectly affect reproductive performance. The pattern of body mass change suggests that the period of lactation is critical for females and endurance rivalry is an important form of competition among males.