Correlates of Intergroup Transfer in Male Grey-cheeked Mangabeys

We studied factors influencing intergroup transfer in male mangabeys (Lophocebus albigena) inhabiting 7 social groups in Kibale National Park over a 2-year period. The sample consisted of 40 males including 36 that we captured and marked during the study. Intergroup transfers are movements between groups that culminate in either long-term (dispersal) or short-term (visits) residency by males in the new groups. Both dispersers and visitors had a greater tendency to move into groups that contained higher numbers of estrous females than their prior group. Using averages of weekly group counts, we found a significant positive relationship between the number of estrous females in a group and the number of adult males in a group, but not between the number of nonestrous females and the number of adult males in the 7 study groups. There is no evidence that dispersal events were released by aggression. For a sample of males followed 2 mo before emigration, aggression did not increase just before emigration. Results suggest that spatiotemporal availability of estrous females is a major proximate factor influencing intergroup transfer in mangabeys. The results also suggest that dispersing males are more sensitive to relative numbers of estrous females than to measures of female availability such as operational sex ratio, socionomic ratio, and the number of excess females that take into account the potential for male-male competition.     

Causes of Female Emigration in the Group-Living Cichlid Fish Neolamprologus multifasciatus

Neolamprologus multifasciatus is a small cichlid breeding in empty gastropod shells that forms complex family groups with several reproductive males and females, presumably due to dispersal constraints. Nevertheless, some exchange of females between groups has been demonstrated. We investigated experimentally emigration patterns of female group members to elucidate which factors influence the decision of adult females either to stay or to emigrate into another territory. Four females resident in a large (13 shells) male territory were given the opportunity to stay there or to emigrate to another large (13 shells) or a small (four shells) male territory. In six replicates we found that (1) the most aggressive females had the largest subterritories (most shells); (2) invariably, the two females with the fewest shells emigrated; (3) both the emigrating and the staying females had more shells after emigration than before. Apparently, females explore their options in neighboring territories and decide accordingly whether to stay or emigrate.     

Social organization of white‐headed langurs (Trachypithecus leucocephalus) in the Nongguan Karst Hills,Guangxi, China

The number of males per group is the most variable aspect of primate social organization and is often related to the monopolizability of females, which is mainly determined by the number of females per group and their reproductive synchrony. Colobines show both inter‐specific and intra‐specific variations in the number of males per group. Compared with other colobine species, little is known about the social organization of white‐headed langur (Trachypithecus leucocephalus), despite its endangered status and unusual limestone habitat. As a part of a long‐term study of the white‐headed langurs in the Nongguan Karst Hills, Guangxi, China, we quantitatively investigated their social organization by analyzing census data from 1998 to 2003. The population censuses revealed that the predominant social organization of bisexual groups was the one‐male group, similar to a previous report on this species and many other Asian colobines. In such groups, one adult male associated with 5.1 adult females, 0.1 sub‐adult males, 2.6 juveniles and 2.9 infants on average, with a mean group size of 11.7 individuals. In addition, three multi‐male groups were recorded, consisting of 2–3 adult males, 1–5 adult females, 0–2 sub‐adult males, 0–7 juveniles and 0–2 infants. They did not contain more adult females than the one‐male groups and were unstable in group membership. The langurs outside bisexual groups were organized into small nonreproductive groups or lived as solitaries. The nonreproductive groups averaged 1.3 adult males, 1.3 sub‐adult males and 2.6 juveniles. Juvenile females were present in such groups on 52.4% of all occasions. As predicted by the monopolization model, the prevalence of the one‐male pattern in this species may mainly be attributed to the small number of females in the group. The possible reasons for the occurrence of multi‐male groups and the presence of juvenile females in nonreproductive groups are also discussed. Am. J. Primatol. 71:206–213, 2009. © 2008 Wiley‐Liss, Inc.     

Fighting and Mating Between Groups in a Cooperatively Breeding Mammal, the Banded Mongoose

Many cooperatively breeding animals actively defend a territory containing resources such as food and shelter, which are essential for reproduction. Some observations, however, indicate that conflicts between groups are often triggered by the attempts of males or females, or both, to gain extra‐group copulations. We studied interactions between 12 groups of banded mongooses (Mungos mungo) in Uganda to test whether the frequency of inter‐group encounters was linked to the reproductive status of females, and conducted an experiment to examine the responses of individuals to mongooses from other groups. The rate at which focal groups fought with other groups was higher when its females were in estrous, suggesting that many fights take place over access to mates. Both males and estrous females were instrumental in instigating encounters with rival groups, and extra‐group copulation in the midst of a fight was observed on three occasions. We experimentally simulated encounters with foreign individuals by presenting each of six groups with cages containing a male and a female from a rival group. Subordinate males were the first to investigate these ‘intruders’, spent more time around the intruders' cages, and were more aggressive to the intruders than either dominant males or females. Subordinate males directed more attention and aggression towards the male intruder than the female intruder. We suggest that male banded mongooses actively seek extra‐group copulations in pursuit of paternity, while females may actively seek extra‐group copulations as a way of reducing inbreeding depression.     

Male and female range use in a group of white‐bellied spider monkeys (Ateles belzebuth) in Yasuní National Park,Ecuador

Spider monkeys (Ateles sp.) live in a flexible fission–fusion social system in which members of a social group are not in constant association, but instead form smaller subgroups of varying size and composition. Patterns of range use in spider monkeys have been described as sex‐segregated, with males and females often ranging separately, females utilizing core areas that encompass only a fraction of the entire community range, and males using much larger portions of the community range that overlap considerably with the core areas of females and other males. Males are also reported to use the boundary areas of community home ranges more often than females. Spider monkeys thus seem to parallel the “male‐bonded” patterns of ranging and association found among some groups of chimpanzees. Over several years of research on one group of spider monkeys (Ateles belzebuth) in Yasuní National Park, Ecuador, we characterized the ranging patterns of adult males and females and evaluated the extent to which they conform to previously reported patterns. In contrast to ranging patterns seen at several other spider monkey sites, the ranges of our study females overlapped considerably, with little evidence of exclusive use of particular areas by individual monkeys. Average male and female home range size was comparable, and males and females were similar in their use of boundary areas. These ranging patterns are similar to those of “bisexually bonded” groups of chimpanzees in West Africa. We suggest that the less sex‐segregated ranging patterns seen in this particular group of spider monkeys may be owing to a history of human disturbance in the area and to lower genetic relatedness between males, highlighting the potential for flexibility some aspects of the spider monkeys' fission–fusion social system. Am. J. Primatol. 72:129–141, 2010. © 2009 Wiley‐Liss, Inc.     

Unraveling the dispersal patterns and the social drivers of natal emigration of a cooperative breeding mammal,the golden lion tamarin

The study of the social drivers of animal dispersal is key to understanding the evolution of social systems. Among the social drivers of natal emigration, the conspecific attraction, aggressive eviction, and reduced social integration hypotheses predict that sexually mature individuals who receive more aggressive behavior and are engaged in less affiliative interactions are more likely to disperse. Few reports have explored these proximate factors affecting emigration in cooperatively breeding species, particularly of Neotropical primates. In this study, we investigated the dispersal patterns and tested the social drivers of natal emigration in the golden lion tamarin (Leontopithecus rosalia) — an endangered species inhabiting Atlantic rainforests fragments in Brazil. We used behavioral and demographic data collected during 7 years from 68 groups of tamarins inhabiting 20 forest fragments. Our analyses from the 160 dispersing individuals showed that dispersal success is higher for males and for those engaged in parallel dispersal, but that males and females use different strategies to enhance their dispersal success, males immigrate into established groups while females form new groups. We did not find high levels of agonistic behavior among group members before natal emigration. Instead we found that conspecific attraction drives natal emigration in both sexes, while additionally the low level of affiliative interactions within the natal group triggers male emigration. We discuss natal emigration in the broader perspective of the cooperative breeding system and the implications of these findings for the conservation of the species.     

Effects of group composition on agonistic behavior of captive pigtail macaques,Macaca nemestrina

We surveyed agonistic behaviors of 20 captive groups of pigtail macaques (Macaca nemestrina) housed under identical spatial conditions. Fifteen groups contained one male each; the other five groups contained no adult males. Groups included six to twelve adult females, some of which had infants with them. We found no relationship between social density of groups and incidence of agonistic behavior, but significantly more contact aggression (grab, hit, push, bite) and noncontact aggression (chase, open-mouth “threat,” bark vocalization) occurred among females in groups containing no males than in those containing one male each. Apparently, males played an important role in the inhibition of intragroup conflict. We also found that females in groups containing males exhibited less noncontact aggression if infants were present than if no infants resided in their groups. Thus, competition of females over infants must not have been an important constituent of intragroup conflict under the conditions of this survey.     

Development of Social Structure in a Captive Colony of the Common Vampire Bat,Desmodus rotundus

The development of social structures of the vampire bat (Desmodus rotundus) was studied in small groups in a captive colony over a period of 30 months. The membership of individuals in the main group and the development of group formation were evaluated by the coherence index. The main group of the vampire colony consisted of several adult females, their infants and one harem male. All members of the main group showed relatively high coherence indices, while the coherence index of adult males, except the harem male, was 0. The females and young in the main group formed a very stable group in comparison to the adult males. The young males left the main group at 20–27 months of age and formed temporary bachelor groups. The harem males changed during the study in the period from 1 to 20 months. They exhibited territorial behaviour and made vigorous attempts to approach the main group within the bounds of their roosting areas, whilst also keeping out intruders.     

Natal emigration by both sexes in the La Pacifica population of mantled howlers: when do some stay?

We have reported previously that all male and female mantled howlers emigrate from natal groups at Hacienda La Pacifica, Costa Rica. In the years since that report, a small number of juveniles have stayed in the natal group without experiencing a solitary phase. Here, we present a post hoc analysis on juvenile emigration in six groups of howlers under observation for varying amounts of time between 1972 and 2005. Our records revealed 139 juveniles for whom emigration status was certain, and 125 of these did emigrate. There was a significant association between presence of mother and emigration: juveniles without mothers were more likely to remain in their natal group (chi(1)(2) = 53.1, P<.0001). The mean age of emigration for all juveniles (n = 125) was 2.47 years (SD = 0.9, range = 1.5-6.5). There was no difference in age of emigration by adult male composition (one-male, multi-male, both), but juveniles of unknown sex emigrated younger than either known males or females (F(2,116) = 4.4, P<.02). For emigrating juveniles of known sex (n = 99), both males and females without mothers left at a later age than those with mothers (F(1,95) = 6.5, P<.02). Although philopatry or delayed emigration occurs in a few motherless animals, most males and females do emigrate from their natal groups at ages consistent with those reported for other species of howlers.     

Secondary transfer of adult mantled howlers (<Emphasis Type="Italic">Alouatta palliata</Emphasis>) on Hacienda La Pacifica,Costa Rica: 1975–2009

Natal emigration by male and female mantled howlers (Alouatta palliata), and subsequent immigration into breeding groups, is well documented for the free-ranging population on Hacienda La Pacifica, Costa Rica, but secondary transfer was considered rare (Glander in Int J Primatol 3:415–436, 1992). Population surveys in 1998 and 2006 caused us to question our assumptions and to re-evaluate our long-term data set from a post hoc perspective. We first identified all animals observed or captured as adults in more than one non-natal group anywhere in the population. We then systematically analyzed joining or leaving by adults in seven groups tracked for various times from 1975 to 2005 for patterns suggesting secondary transfer. Fourteen adults (nine females, five males) were found in two different non-natal groups as adults. In addition, one male and one female that became dominant and reproduced in their natal group later transferred to a second group, and one female was known to be a tertiary transfer. Data from the seven tracked social groups indicate that 35% of all the males and 29% of all the females were potential secondary transfers. In these groups, males leaving or joining was not associated with group size or absolute number of females. Females leaving or joining was not associated with group size or absolute number of males, but females left groups with more females and joined groups with fewer females. Both sexes left groups with unfavorable sex ratios for their sex and joined groups with sex ratios more favorable for their sex. Since a favorable sex ratio is associated with reproductive success in other howler populations, this suggests secondary transfer as a reproductive strategy. Other factors could also influence secondary transfer.     

Movement and reproductive behavior of solitary male redtail guenons (Cercopithecus ascanius)

Adult male redtail guenons (Cercopithecus ascanius) live as resident males in heterosexual groups or as nonresident males outside groups. Data are reported here on movement and temporary intrusions by nonresident males in heterosexual groups. These data further elucidate the reproductive strategies of this class of individual. Observations of three radiocollared males provided data on the details of their movement relative to groups. Regular sampling of three heterosexual groups provided data on patterns of nonresident males' intrusions. During peak mating months, nonresident males formed loose affiliations with groups. The males affiliated with one group at a time, with each affiliation lasting several weeks. Nonresident male intrusions in groups were unevenly distributed and were not correlated with the number of potentially fertile females in each group. Estimates of male density indicate that there were about two nonresident males per group, which suggests an adult sex ratio of three females per male within the study area. Nonresident male redtail guenons apparently do not spread their reproductive efforts over several groups at once, but instead each male tends to direct his attention toward the group offering the best opportunity for reproductive payoffs. Factors that determine which groups they attempt to infiltrate may include the characteristics of resident males, female choice, or both, but the number of fertile females probably has no effect. The male low density and female-biased sex ratio among adults should facilitate resident males' attempts to monopolize fertilizations in groups; however, resident males were unable to prevent copulations by nonresident males even when only one nonresident was intruding in a group.     

Intra-group home ranges in a female-biased group of sex-changing fish

Simultaneous loss of multiple males from large, multi-male social groups of the coral reef fish, Anthias squamipinnis, induces an equivalent number of females to change sex. One-to-one replacement could be produced in large groups if the group was organized according to either of two hypotheses. The group could consist of an aggregation of single-male, multi-female units with the home ranges of particular females falling within the home ranges of each male, and with sex change controlled separately within each unit. Alternatively, females might range freely throughout the group, with sex change controlled by differential interaction rates among males and females. In the latter case, female home ranges would overlap substantially with the home ranges of several males. To choose between these hypotheses, the three-dimensional, intra-group home ranges of five males and five adult females were measured in a group containing five males and 32 adult females. All 10 fish had definable home ranges. Home ranges of females were larger than home ranges of males. All females ranged much farther outside of the home range of individual males than inside, and most females spent most of the observation time outside of the home range of any one male. These results effectively disproved the hypothesis of spatially-defined, single-male, multi-female units. Rather, large groups consist of females ranging freely over the discrete home ranges of several males.     

Group size and group composition of the mona monkey (Cercopithecus mona) on the Island of Grenada,West Indies

Cercopithecus, the genus of guenons, is the largest of the African primate genera, and yet more than half of the species belonging to this group have never been the focus of a long-term field study. In this paper, I present data on group size and composition for a previously unstudied population of guenons on the Caribbean island of Grenada. The mona monkey, Cercopithecus mona, was introduced to Grenada from Africa approximately 200–300 years ago. Two types of social groups were found for Cercopithecus mona on Grenada: all-male groups consisting of two to four individuals and bisexual groups containing 5–32 individuals. All-male groups of Grenada mona monkeys contained any combination of juveniles, subadults, and/or adults. All-male groups were a common occurrence on Grenada but have never been reported for African C. mona and have been reported only in two other forest Cercopithecusspecies. Bisexual groups appeared to consist of one adult male, one to six adult females, subadult females, and juveniles and infants of both sexes. Even though no more than one adult male was ever seen in each bisexual group of monas on Grenada, other males were heard giving copulation calls simultaneously with resident adult male loud calls, suggesting that other males occasionally infiltrate bisexual groups. Am. J. Primatol. 43:167–173, 1997. © 1997 Wiley-Liss, Inc.     

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.     

Sociality, Mate Choice, and Timing of Mating in American Bison (Bison bison): Effects of Large Males

We studied group size, composition, and mating activities in American bison (Bison bison) during rut on the Delta Junction Bison Range in interior Alaska, USA, in 1996 and 1997. Our purpose was to determine the effects of large males (≥5 yr old) on mating and associated activities. Groups with large males were larger than those containing smaller males. Most groups of bison were mixed‐sex (90%), but large males occurred in only one‐half of all groups. Moreover, females in groups with large males were more likely to copulate than those in groups with smaller males, indicating a female preference for large males. Nevertheless, our results are consistent with large males seeking out adult females for mating rather than vice versa. Mating peaked in mid‐August during both the study years and was highly synchronous. Scent marking was coincident with mating, an outcome consistent with a hypothesis of such behavior triggering ovulation. Scent marking by large male bison occurred in both male–male and male–female contexts, but was associated most often with sexual activities. No differences in group size occurred with changes in weather or among vegetation types occupied by bison. Group size of bison, however, was larger with increasing distance from the forest edge, which likely was a response to predation risk in this predator‐rich environment.     

Male Influx, Infanticide, and Female Transfer in Macaca radiata radiata

In bonnet macaques, males usually disperse between groups and females remain philopatric, but researchers have reported female transfer. We report a rare case of male influx during the mating season in our bonnet macaque study group in the Anaimalai Hills. The density of bonnet macaques in the study region was unusually high. The study group had a single, crippled adult male with a long tenure and 5 adult females. During the mating season, adult females approached and mated with outgroup males, and then several males entered the group. The adult male left the group without any resistance. The incoming males mated with 3 receptive females, forcibly mated with 2 lactating females, and attacked and killed 2 infants. During the influx, 2 outgroup females joined the group. The data suggest that male influxes provide an opportunity for infanticide and female transfer, which can have important fitness consequences even in species in which they rarely occur.     

Mating promiscuity and reproductive tactics in female black and gold howler monkeys (Alouatta caraya) inhabiting an island on the Parana river,Argentina

In several primate species, females mate promiscuously and several adult males peacefully co‐reside in the same social group. We investigated female mating behavior in two neighboring multimale–multifemale groups of Alouatta caraya in northern Argentina (27°20′S–58°40′W). All adult individuals in each group were marked with identification anklets and ear tags, and followed for five consecutive full days per month during 20 consecutive months. We recorded 219 copulations for eight resident females in these two groups. Thirty‐two percent of matings involved extra‐group copulations and 68% were with resident males. During periods when females were likely to conceive and during periods when females were nonfertile (pregnancy and lactation), there were no significant differences in the average number of resident and nonresident males with which they copulated (G‐test: G_adj=0.1, df=3, P>0.05). In both of our study groups, adult males were tolerant of the mating activities between resident males and resident females, but acted aggressively and collectively (howling, border vigilance, and fighting) when extragroup males attempted to enter the group and mate with resident females. Given the frequency of extragroup matings, we examined the distance females traveled to engage in these copulations, time engaged in pre‐ and postcopulatory behavior, and the risk of injury during extragroup copulations. These costs were found to be relatively small. We suggest that female promiscuity is the prime driver or constraint on male reproductive opportunities in this species. Female promiscuity in A. caraya appears to represent a mixed mating strategy that may serve to increase opportunities for genetic diversity between a female's successive offspring as well as minimize the risk of infanticide by spreading paternity estimates across a larger number of adult males. Am. J. Primatol. 72:734–748, 2010. © 2010 Wiley‐Liss, Inc.     

Mating system of an exceptional primate, the olive colobus (Procolobus verus)

In the olive colobus (Procolobus verus), many groups have multiple males and the males have large testes. This indicates that even though this species lives in small groups, single males do not monopolize the groups. We investigated the strategies employed by males to secure their mating success, and sought to determine whether the lack of male monopolization was a result of female mating strategies, as indicated by the exaggerated sexual swellings of the females. Four study groups were monitored for demographic changes, and group composition was determined in six additional groups in Taï National Park, Ivory Coast, between 1994 and 1999. Social behavior was recorded by scan and focal sampling in the study groups. The almost permanent association of olive colobus with Diana monkeys (Cercopithecus diana) in effect provided males a resource at which they could expect females to visit and sometimes even permanently join them, as well as protection from predators. As alternative strategies for obtaining females, one male took over the group of another male and one male immigrated into a bisexual group. Within bi‐male groups, dominant males mated most frequently and males defended their groups during intergroup interactions. Lone females that visited groups or solitary males had a swelling more often than expected, and generally mated with the males they visited. Females had long receptive periods, several consecutive receptive cycles, and some overlap in receptive periods within groups. Females mated with extragroup males, and during infertile periods. We concluded that the males used the Diana monkeys for safety reasons and to obtain mating partners, and that female reproductive biology and behavior prevented the monopolization of groups of females by single males. Our data were inconclusive as regards the benefits to females of avoiding monopolization by males. Am. J. Primatol. 62:261–273, 2004. © 2004 Wiley‐Liss, Inc.     

Social Influences on Group Membership in <Emphasis Type="Italic">Propithecus verreauxi verreauxi</Emphasis>

The size and composition of a social group can influence the reproductive success of its members. I examined the hypothesis that residents actively try to manipulate group size and composition in Verreaux’s sifaka (Propithecus verreauxi verreauxi). I collected behavioral data from 21 individuals in 5 social groups from November 2000 to March 2002 in the Kirindy Forest (C.F.P.F.) of western Madagascar. I investigated the affiliative and agonistic behaviors of resident males and females toward other group members to determine whether residents attempt to manipulate the costs and benefits of group membership. I also examined 3 cases of immigration attempts and 1 case of emigration. As expected, group members facilitated the residency of subordinate members of the opposite sex. For example, females sought out the subordinate males, responded to their calls, and even groomed them more. However, intrasexual relationships were also important for establishing or maintaining residency. The data, in conjunction with the observations of immigration and emigration events, suggest that resident individuals of both sexes attempt to influence group size and composition. I suggest that single–male groups are not the norm in Propithecus verreauxi verreauxi because females encourage, and males do not discourage, multiple males to reside in the group to increase group stability and to reduce the risk of infanticide.     

Aggression with mixed age-sex groups of bolivian squirrel monkeys following single animal introductions and new group formations

The social organization of the Bolivian squirrel monkey (Saimiri boliviensis boliviensis) is thought to be sexually segregated, with males and females forming separate social groups during the nonbreeding season. To investigate the influence of this social order on patterns of aggression, controlled single animal introductions within established groups and establishment of new groups were studied in a systematic manner. Behavioral observations were made using an all-occurrences technique to sample all agonistic interactions. In study I, 4 animals of each age-sex class (adult males and females, juvenile males and females) were introduced one at a time into 4 different social groups composed of 1 adult male and 8 to 10 adult females. Behavioral observations were made prior to and after the introductions. Newly introduced adult males received significantly more contact aggression than other age classes. Only the adult females increased aggression after the introduction of new animals. In study II, new social groups were formed and behavioral observations were made following formation. One group was formed from 7 pairs of familiar females and an unfamiliar adult male. This group had a high frequency of aggression during the first half hour, with contact aggression rising to peaks at 3 and 5 h. The second group consisted of 10 familiar females, 2 pairs of familiar females and an unfamiliar male. There was a significant peak in contact aggression 3 h into the observation.