Female dispersal,social organization,and infanticide in langurs: Are they linked to human disturbance?

Female dispersal in gregarious animals can involve the desertion of a site, desertion of a social group, or both. Group desertion may be related to inbreeding avoidance. Group fidelity may result from cooperation among females in a group. Site fidelity will be more likely when food can be monopolized and when the population density is close to habitat saturation. The degree of habitat saturation was approximated with a measure of human disturbance. The influence of these various factors on the incidence of female dispersal was investigated for langur populations using data from the literature. The results suggest that female dispersal in langurs involved site desertion, not group desertion. The incidence of female dispersal may affect the social organization of langurs. I propose that when females do not disperse, male takeovers prevail, whereas in populations where female dispersal regularly occurs bisexual groups are disbanded or new groups are formed, a process I call female split-merger. Male takeover is thought to occur when site fidelity is high, female split-merger when site fidelity is low. These processes were indeed found to occur in these circumstances. The dispersal of females might prevent infanticide, whereas male takeover might promote it. Indeed, in studies with male takeover, more infants fell victim to infanticide than in studies with female split-merger. Therefore, female dispersal in langurs is an effective female counterstrategy to infanticide. The factor that had the most profound effect on female dispersal, social organization, and infanticide was habitat saturation. Habitat saturation was measured as the degree of human disturbance, and its influence on the behavior of langurs is probably of relatively recent date. This may lead to an evolutionary transient situation and may explain the discrepancy between current socioecological theories and the behavior of langurs in populations lacking female dispersal. Am. J. Primatol. 44:235–254, 1998. © 1998 Wiley-Liss, Inc.     

Social and seasonal influences on reproductive biology in male moustached tamarins (Saguinus mystax)

In several primate taxa there is evidence that the social and physical environment can exert a significant effect on reproductive behavior and biology. In this paper we examine social and physiological factors influencing group composition and reproduction in free-ranging moustached tamarin monkeys (Saguinus mystax mystax). This species is characterized by cooperative care of the young and a breeding system that includes both polyandrous and polygyandrous matings. Body measurements collected on adult males residing in multimale groups indicate marked within-group differences in testes volume. In 12 of 17 groups examined, testes volume of at least two resident adult males differed by 21–174%. Among these males, testes volume was not correlated with either body weight or adult age class. We also examine whether factors such as time of year had an effect on reproductive condition. An analysis of body measurements of 128 adult male and 127 adult female moustached tamarins, wild-trapped and released in northeastern Peru, indicates cyclic changes in genital size. For males, mean monthly testes volume in July (712 mm³) was twice that recorded in June (351 mm³). Females exhibited a similar pattern. Although endocrine information on intra- and intersexual social effects on fertility are unavailable for S. mystax, given the high degree of social cooperation and lack of overt aggression among adult male group members, we offer the possibility that resident male moustached tamarins compete for access to the groups' lone breeding female through socially induced reproductive suppression and sperm competition. © 1996 Wiley-Liss, Inc.     

Male reproductive strategies in new world primates

Patterns of three variables of reproductive strategies in male New World primates are examined: (i) how males obtain access to potential mates; (ii) how males obtain actual mating opportunities; and (iii) how males affect infant survival and female reproductive success. Male opportunities to associate with females, whether by remaining in their natal groups, dispersing and forming new groups, or dispersing and taking over or joining established groups, are strongly influenced by local population densities and correlate with female reproductive rates and the extent of female reproductive seasonality and synchrony. Differences in male mating success are affected by female accessibility, whether male-male and male-female relationships are hierarchical or egalitarian, and whether female reproduction is seasonally restricted. Patterns of male behavior toward infants, characterized as active assistance, overt interference, or benign tolerance, appear to co-vary with differences in the degree to which males can affect female reproductive rates. These qualitative analyses suggest that the reproductive strategies of male New World primates can be classified along a continuum ranging from conservative to daring depending on whether female reproductive rates are relatively slow or fast and whether reproduction is strongly or weakly linked to seasonal ecological variables. Males adopt the conservative strategy of staying in their natal groups, forfeiting exclusive mating opportunities, and treating infants with tolerance when female reproduction is constrained by ecological factors. Conversely, males adopt the more daring strategy of dispersing and competing when potential payoffs through their ability to affect female reproduction are high.     

Intrasexual competition underlies sexual selection on male breeding coloration in the orangethroat darter,Etheostoma spectabile

Elaborate, sexually dimorphic traits are widely thought to evolve under sexual selection through female preference, male–male competition, or both. The orangethroat darter (Etheostoma spectabile) is a sexually dichromatic fish in which females exhibit no preferences for male size or coloration. We tested whether these traits affect individual reproductive success in E. spectabile when multiple males are allowed to freely compete for a female. The quality and quantity of male coloration were associated with greater success in maintaining access to the female and in spawning as the primary male (first male to participate). On the other hand, sneaking behavior showed little correlation with coloration. Male breeding coloration in E. spectabile may therefore demonstrate how intrasexual competition can be a predominant factor underlying the evolution of male ornaments.     

Male competition reverses female preference for male chemical cues

Females must choose among potential mates with different phenotypes in a variety of social contexts. Many male traits are inherent and unchanging, but others are labile to social context. Competition, for example, can cause physiological changes that reflect recent wins and losses that fluctuate throughout time. We may expect females to respond differently to males depending on the outcome of their most recent fight. In Bolitotherus cornutus (forked fungus beetles), males compete for access to females, but copulation requires female cooperation. In this study, we use behavioral trials to determine whether females use chemical cues to differentiate between males and whether the outcome of recent male competition alters female preference. We measured female association time with chemical cues of two size‐matched males both before and after male–male competition. Females in our study preferred to associate with future losers before males interacted, but changed their preference for realized winners following male competitive interactions. Our study provides the first evidence of change in female preference based solely on the outcome of male–male competition.     

Influence of Adult Size on Mate Choice in the Solitary and Gregarious Parasitoids, Cotesia marginiventris and Cotesia flavipes

Female and male mate choice in relation to adult size were examined for the solitary and gregarious parasitoids, Cotesia marginiventris (Cresson) and Cotesia flavipes Cameron, respectively. In addition, male precopulatory behaviors were observed for evidence of male competition or a large-male advantage in mate acquisition. Male parasitoids are not known to offer female mates direct benefits, but females that mate high quality males may obtain indirect benefits, such as offspring that are more successful in obtaining mates. Female choice experiments for C. marginiventris found that large males approached females first more frequently than small males, and that females mated large males significantly more often than small males. Male choice experiments for C. marginiventris did not demonstrate a male preference for female size. In contrast, female choice experiments with C. flavipes found that females mated equally with large or small males, while male choice experiments showed that males attempted copulation and mated more frequently with smaller females. Male competition was not observed in the gregarious species C. flavipes, but competition in this gregarious parasitoid could be moderated by dispersal.     

Dynamics of male residence and female oestrus during a breeding season of blue monkeys in the Kakamega Forest, Kenya

Blue monkeys (Cercopithecus mitis stuhlmanni) are seasonal breeders with a dynamic mating system in which typical one‐male social units are regularly disrupted and replaced by multi‐male ones. The number of males in the group is correlated with the number of oestrous females. We used observations of male presence and female oestrus on individual days during a 6‐month period to assess whether the presence of multiple males in a group stimulates female oestrus or whether oestrous females attract multiple males to the group. We confirmed prior observations with our finding that the number of males in a group was significantly correlated (r_s = 0.435, P < 0.0001) with the number of oestrous females across 126 observation days. A transition matrix did not show an obvious relationship between day‐to‐day changes in the numbers of oestrous females and males. However, cross‐correlation analysis provided stronger support for the idea that the number of oestrous females attracts males to a group than for the idea that the influx of strange males stimulates oestrus in female blue monkeys. Autocorrelation analysis showed that while female oestrus appeared to show a high degree of synchrony, as expected in a seasonal breeder, there was no evidence that the number of males accompanying a group of females influenced the likelihood of other males joining or leaving the group. Overall, our results confirm that female oestrous behaviour stimulates changes in male residence patterns. However, other observations suggest that changes in male residence may also stimulate female oestrus in some circumstances.     

Female‐biased dispersal in a bat with a female‐defence mating strategy

The ultimate causes for predominant male‐biased dispersal (MBD) in mammals and female‐biased dispersal (FBD) in birds are still subject to much debate. Studying exceptions to general patterns of dispersal, for example, FBD in mammals, provides a valuable opportunity to test the validity of proposed evolutionary pressures. We used long‐term behavioural and genetic data on individually banded Proboscis bats (Rhynchonycteris naso) to show that this species is one of the rare mammalian exceptions with FBD. Our results suggest that all females disperse from their natal colonies prior to first reproduction and that a substantial proportion of males are philopatric and reproduce in their natal colonies, although male immigration has also been detected. The age of females at first conception falls below the tenure of males, suggesting that females disperse to avoid father–daughter inbreeding. Male philopatry in this species is intriguing because Proboscis bats do not share the usual mammalian correlates (i.e. resource‐defence polygyny and/or kin cooperation) of male philopatry. They have a mating strategy based on female defence, where local mate competition between male kin is supposedly severe and should prevent the evolution of male philopatry. However, in contrast to immigrant males, philopatric males may profit from acquaintance with the natal foraging grounds and may be able to attain dominance easier and/or earlier in life. Our results on Proboscis bats lent additional support to the importance of inbreeding avoidance in shaping sex‐biased dispersal patterns and suggest that resource defence by males or kin cooperation cannot fully explain the evolution of male philopatry in mammals.     

Range use of the forest chimpanzees of Kibale: Implications for the understanding of chimpanzee social organization

We contrast the range use patterns of male and female chimpanzees (Pan troglodytes) based on repeated sightings over three years of 19 individuals in the Kanyawara community of Kibale Forest Reserve, Uganda. Depending on how home range size was calculated, male chimpanzees used an area that was 1.5 to 2 times greater than that of females. There was no difference between the sexes in whether their home ranges were used in a clumped or uniform fashion. However, males were more likely to be seen in boundary areas than females. These results are discussed in light of previously proposed models of chimpanzee social organization. It is concluded that the scenario in which females have smaller core areas within the defended home range of the males is most strongly supported by the range use patterns observed in Kibale chimpanzees. © 1993 Wiley-Liss, Inc.     

Aspects of Male Competition in <Emphasis Type="Italic">Colobus vellerosus</Emphasis>: Preliminary Data on Male and Female Loud Calling,and Infant Deaths After a Takeover

Male Colobus vellerosus are the main participants in intergroup encounters, and lead incursions in neighboring groups during which they attack infants. Extragroup copulations, all-male groups, and male takeover occur in the species. Here, we provide additional information on behaviors associated with male reproductive competition in Colobus vellerosus. We examined 1 resident male loud calling and participation in intergroup encounters in relation to a takeover. We also report a second case of takeover that led to the death of the former resident male and the death of 2 male infants, presumably as a result of aggression from the all-male group. The new resident male wounded the third infant of the group, which apparently died after its mother abandoned it. During the period characterized by the attacks on the infant and after its disappearance, females initiated and participated in loud call bouts with the new resident male. We examine the possible functions of female loud calling, and suggest that in this context, it might force the resident male to call along to indicate his presence.     

Intergroup encounters in wild white-faced capuchins (Cebus capucinus)

Wrangham (1980) hypothesized that knowledge of the nature of intergroup encounters is crucial to understanding primate social relationships and social organization. I studied a single social group of wild white-faced capuchins over a period of 26 months and observed 44 encounters between social groups during 3703 hr of observation. All intergroup encounters consisted of predominantly hostile social interactions. However, nonaggressive interactions between males of different social groups occurred in a few cases. Adult males were the sole participants in 39 encounters and the primary participants in all 44 encounters. The alpha male was the most frequent participant. High-ranking females participated aggressively in five encounters, and low-ranking females never participated. There was no stable intergroup dominance hierarchy. I hypothesize that the need for male-male cooperation in intergroup aggression is an important factor influencing the quality of intragroup male-male relationships. Behavior during intergroup encounters is consistent with the idea that intergroup behavior is related to male reproductive strategies, but inconsistent with the idea that intergroup aggression is related to female defense of resources. The possibility that males are “hired guns” (Wrangham, 1980) cannot be ruled out.     

Social Relationships Among Male <Emphasis Type="Italic">Alouatta pigra</Emphasis>

We investigated male social relationships in 2 groups of black howlers (Alouatta pigra) in Palenque National Park, Mexico, over a 14-mo study characterized by frequent changes in male group membership. Both single males and pairs of males entered our focal groups. Single males tended to join groups, whereas pairs of males entering groups together successfully evicted resident males. The 19 male dyads across the 11 periods defined by changes in group membership were rarely in close proximity or interacting with one another. Nevertheless, 1 male formed significantly closer associations with resident females in 6 periods and achieved higher mating success during 4 of the periods when ≥1 female was sexually active. In the other 5 periods, no particular male maintained significantly closer associations with resident females, which may be a result of the rapid sequence of changes in male membership and corresponding social instability. Resident males participated frequently and consistently in mutual howling, but the resident male with the strongest female associations and highest mating success initiated howling bouts more frequently in all but 1 of the 6 socially stable periods. Though still preliminary, our findings suggest that variation in social relationships among male black howlers may be more related to their ability to establish strong relationships with females than to their relationships with one another.     

Male rank,reproductive behavior,and reproductive success in free-ranging rhesus macaques

Paternity assessment through DNA fingerprinting by synthetic oligonucleotide probes was applied to one birth cohort in a social group of free-ranging rhesus macaques (Macaca mulatta) on Cayo Santiago. The 11 group males and 9 males from other groups were observed mating with the females. Paternity was determined for 11 of the 15 infants. Male dominance rank was not associated with reproductive success. High-ranking resident males (N=5) sired 27% of the infants born during a one-year study. Four of the 11 infants of known paternity were sired by males of other social groups. The four infants of unknown paternity were sired either by males not observed mating with the females or the low-ranking male who was not fingerprinted. Male dominance rank was not associated with reproductive activity during conception cycles. These results suggest that the effect of rank on male reproductive success is not a predictable correlation, but a conditional probability.     

Impact of Male Takeovers on Infant Deaths,Births and Conceptions in Cebus capucinus at Santa Rosa,Costa Rica

Male takeovers are associated with infant wounding and death in 3 of 4 capuchin species. In this paper, I analyze the effects of male takeovers on infant mortality and the subsequent conceptions and interbirth intervals of their mothers over an 18-yr period and test predictions of the sexual selection model of infanticide for white-faced capuchins, (Cebus capucinus). Major findings are that infants are significantly more likely to die in the 3- and 12-mo periods following a takeover than in times of peace and that a female whose infant dies experiences a significantly shorter interbirth interval before her next infant is born than she would have had the former infant survived. In the vast majority of cases, the invading males become resident in the group and are present during the subsequent conceptions of the females in the group. However, overall conception rates do not rise significantly in the year after a takeover, there is no relationship between the age of the infant at death and the length of the mother's subsequent interbirth interval, and it is not yet clear if male infants are preferentially targeted by invading males. Most takeovers occur during the 6-mo dry season and most conceptions occur in the wet season, 3–6 mo later. My findings support the major predictions of the sexual selection model of infanticide in primates and demonstrate that male takeovers of social groups have substantial effects on infant survival and maternal parturition patterns in Cebus capucinus.     

What is driving male mate preference evolution in Jamaican field crickets?

Male mating preferences are often a neglected aspect of studies on sexual selection. Male mating preferences may evolve if they provide males with direct‐fitness benefits such as increased opportunity to fertilize more eggs or indirect‐fitness benefits such as enhanced offspring survival. We tested these ideas using Jamaican field crickets, Gryllus assimilis, previously shown to exhibit male mating preferences. We randomly mated males to either their preferred or non‐preferred potential mates and then asked whether mating treatment influenced egg oviposition or offspring viability. Preferred females were not significantly more fecund and did not produce more viable eggs or offspring than non‐preferred females. Male mate preferences were therefore inconsistent with both the direct‐ and indirect‐fitness benefits hypotheses under the conditions of our experiment. Our null results leave us with an open question about what is driving the evolution of mating preferences in male crickets. Future research should explore the whether the offspring of preferred females are more attractive, have stronger immune systems, and/or experience higher adult longevity.     

Sex differences in infant integration in a semifree-ranging group of Barbary macaques (Macaca sylvanus,L. 1758) at Salem,Germany

Sixteen multiparous Barbary macaque females with newborns were studied over a 16 month period within the context of their naturally formed group. Analysis of their social behavior revealed 1) triadic interactions involving focal females, their newborns, and other group members occurred mainly with other females; and 2) mothers with female newborns interacted mainly with females of their own matriline, while mothers with male newborns interacted mainly with nonmatriline females. Observed in two successive birth seasons, this pattern indicates that partners of maternal interactions chose each other according to the sex of the newborn. Measures of distance from the mother also reflected differences between infants of different sex. At about five months of age, female infants were observed close to their mothers significantly more often than males. This finding follows the pattern of a sex-specific infant socialization process which integrates female infants into the network of their matrilines and male infants into the broader group. This sex-specific integration pattern is interpreted as supporting female philopatry and male dispersal. © 1995 Wiley-Liss, Inc.     

Lactation and care for unrelated infants in forest-living ringtailed Lemurs

An adult female ringtailed lemur (Lemur catta) known not to have been pregnant showed spontaneous lactation in response to twin infants born to an unrelated female. The females had met only 7 months earlier, when they and two other unrelated adult females were released from separate locations in a forest enclosure to form a new social group. Three months after release, an adult male from an adjacent enclosure gained access to the new group for 1 day, the day of one female's estrus. No males had access to the females throughout the remainder of the breeding season. Within 2 weeks of the birth of the twins, one of the other adult females began carrying the infants frequently, typically one at a time. All three females were checked for lactation when the infants were two months old. Both their mother and the unrelated adult who had been carrying the infants were producing milk. The third adult female, who never carried either infant, had no milk. The third adult female, who never carried either infant, had no milk. This female, however, like the two maternal females, frequently attacked unfamiliar immigrating adult males when the males approached the infants. Potential implications of these observations concerning the social organization of ringtailed lemurs are discussed.     

Sex differences in play among western lowland gorilla (Gorilla gorilla gorilla) infants: implications for adult behavior and social structure

Mammalian play is believed to improve motor skills as well as facilitate the development of social relationships. Given the marked sexual dimorphism in gorilla body size and the role assumed by the male in protecting the group from conspecifics and predators, the motor-training hypothesis of play predicts that male infants should exhibit higher frequencies of social play than female infants, and that males should prefer to play with other males. Given that adult female gorillas are strongly attracted to adult breeding males and form only weak social bonds with unrelated adult females, the social-relationship hypothesis of play predicts that female infants should prefer to play with males. These hypotheses were tested in a 22-month study of 12 gorilla infants, aged between 0-5 years, living in three zoological parks in Chicago and Atlanta. Consistent with the hypotheses, male infants played more than female infants did, and both male and female infants preferred to play with males rather than with females. These findings suggest that sex differences in play in the great apes and other primates can be predicted by the characteristics of adult behavior and social structure above and beyond the patterns of sex-biased dispersal or coalition formation with same-sex kin.     

Evolution of an ornament,the dewlap,in females of the lizard genus Anolis

Male ornaments have been the subject of numerous studies on sexual selection and communication, although female ornaments have garnered substantially less study, even though female ornaments are well developed in some species. The factors that have propelled the evolution of elaborate ornaments in females are poorly understood but may include genetic correlations between the sexes, social selection, sensory drive or species recognition. We used simulation‐based comparative methods and a newly estimated phylogeny to test these four hypotheses to explain female ornamentation within the diverse neotropical lizard genus Anolis. We found support for the sensory drive hypothesis and the social selection hypothesis; the female dewlap was larger in species that use more arboreal habitats, as well as in species where the sexes were less dimorphic. We did not find support for the genetic correlation hypothesis or the species recognition hypothesis. We propose that the size of the female dewlap may evolve in response to sensory drive differentially affecting species in different habitats, as well as social selection such as male mate choice or intrasexual competition for territory among females. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 191–201.