Paternity alone does not predict long-term investment in juveniles by male baboons

Adult male chacma baboons (Papio hamadryas ursinus) form preferential associations, or friendships, with particular lactating females. Males exhibit high levels of affiliative contact with their friends’ infants and defend them from potentially infanticidal attacks (Palombit et al. 1997). Little is known about males’ associations with juveniles once they have passed the period of infanticidal risk. We conducted an observational, experimental, and genetic study of adult male and juvenile chacma baboons in the Moremi Reserve, Botswana. We identified preferential associations between males and juveniles and used behavioral data and a playback experiment to explore whether those associations have potential fitness benefits for juveniles. We determined whether males preferentially invest in care of their own offspring. We also determined how often males invest in care of their former friends’ offspring. The majority of juveniles exhibited preferential associations with one or two males, who had almost always been their mother’s friend during infancy. However, in only a subset of these relationships was the male the actual father, in part because many fathers died or disappeared before their offspring were weaned. Male caretakers intervened on behalf of their juvenile associates in social conflicts more often than they intervened on behalf of unconnected juveniles, and they did not appear to differentiate between genetic offspring and unrelated associates. Playbacks of juveniles’ distress calls elicited a stronger response from their caretakers than from control males. Chacma males may provide care to unrelated offspring of former friends because the costs associated with such care are low compared with the potentially high fitness costs of refusing aid to a juvenile who is a possible offspring.     

More than friends? Behavioural and genetic aspects of heterosexual associations in wild chacma baboons

In mammals, fathers are facultative caretakers, and male care is expected to evolve only if it is directed towards related young. Yet, in several promiscuous primate societies, males seem to care for infants despite a presumably low paternity confidence. In cercopithecines, cohesive associations (‘friendships’) between a lactating female and an adult male are frequent and provide the female and her infant with protection against various sources of aggression, including infanticide. However, the benefits gained by males through such relationships remain unclear, in part, because the relatedness between males and their protected infants has rarely been examined. Moreover, little is known about the nature of the cues underlying kin discrimination by males in societies where females mate polyandrously. In this study, we combine behavioural and genetic data from wild chacma baboons (Papio ursinus) in Namibia to investigate (1) whether males are related to their friend’s infant and (2) whether similarity between the major histocompatibility complex (MHC) genotype of males and infants (potentially perceived through odour phenotype) favours the establishment of friendships. We first show that males share close genetic ties with their friend’s infants, most often by having sired the infant. Secondly, we find that male–infant MHC (Class II–DRB) similarity, in contrast to paternity, does not predict male–infant associations. Overall, our results clarify the nature of the evolutionary benefits gained by males in these heterosexual associations, which can be considered as true paternal care. However, the proximate mechanisms underlying paternity recognition remain to be identified.     

“Friendships” between new mothers and adult males: adaptive benefits and determinants in wild baboons (<Emphasis Type="Italic">Papio cynocephalus</Emphasis>)

Close associations between adult males and lactating females and their dependent infants are not commonly described in non-monogamous mammals. However, such associations [sometimes called “friendships” (Smuts 1985)] are regularly observed in several primate species in which females mate with multiple males during the fertile period. The absence of mating exclusivity among “friends” suggests that males should invest little in infant care, raising questions about the adaptive significance of friendship bonds. Using data from genetic paternity analyses, patterns of behavior, and long-term demographic and reproductive records, we evaluated the extent to which friendships in four multi-male, multi-female yellow baboon (Papio cynocephalus) groups in Amboseli, Kenya represent joint parental care of offspring or male mating effort. We found evidence that mothers and infants benefited directly from friendships; friendships provided mother–infant dyads protection from harassment from other adult and immature females. In addition, nearly half of all male friends were the genetic fathers of offspring and had been observed mating with mothers during the days of most likely conception for those offspring. In contrast, nearly all friends who were not fathers were also not observed to consort with the mother during the days of most likely conception, suggesting that friendships between mothers and non-fathers did not result from paternity confusion. Finally, we found no evidence that prior friendship increased a male’s chances of mating with a female in future reproductive cycles. Our results suggest that, for many male–female pairs at Amboseli, friendships represented a form of biparental care of offspring. Males in the remaining friendship dyads may be trading protection of infants in exchange for some resources or services not yet identified. Our study is the first to find evidence that female primates gain social benefits from their early associations with adult males. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of male sexual harassment on female time budgets,feeding behavior,and metabolic rates in a tropical livebearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>)

Sexual harassment by males has the potential to affect almost any aspect of female behavior and life history. Using Atlantic mollies (Poecilia mexicana) as a model—a species in which males do not court but almost constantly try to forcefully mate with females—we asked whether and how male harassment influences (a) females’ time budgets and (b) feeding rates (e.g., through frequent flight from male approaches), and (c) whether metabolic rates are increased as a response to stress. Field observations in a natural P. mexicana population revealed that males (average feeding rate 15%) spent far less time feeding than females (60%), and clearly traded off frequent pursuit of females with foraging. Most importantly, females’ feeding times were dramatically reduced when being pursued by a harassing male. Also in standardized lab experiments, females spent significantly less time feeding when accompanied by a male as compared to being in the presence of another female. This effect was also observed when partner fish (male or female) were presented only visually, but could not interact physically with the focal female. It seems, therefore, that females increase vigilance when a harassing male is around, which keeps them from feeding even before males actually approach them. Based on the latter result, we asked whether a stress-induced increase in metabolic rates would be discernible. We measured oxygen consumption and gill ventilation frequencies (opercular rates) of females in different social contexts (alone, with another female, or a male). The predicted, strong body mass dependency of both physiological parameters was uncovered, but no evidence for an effect of social context was detected. We argue that male harassment represents such a constant (but non-lethal) stressor for poeciliid females that their metabolic stress responses have adapted to this through habituation.     

Vigilance in wild Thomas's langurs (Presbytis thomasi ): the importance of infanticide risk

This study examines vigilance as a behavioural indicator of the importance of infanticide risk by comparing the infanticide avoidance hypothesis with the predation avoidance and mate defence hypotheses for wild Thomas's langurs (Presbytis thomasi) in Sumatra. We found that all individuals were more vigilant in situations of high predation risk, i.e. lower in the trees and in the absence of neighbours. Females were also more vigilant on the periphery of the group. However, there were variations in vigilance levels that could not be accounted for by the predation avoidance hypothesis. Males without infants showed higher levels of vigilance in areas of home range overlap than in non-overlap areas during the early phase of their tenure, strongly suggesting mate defence. In these areas of home range overlap where Thomas's langur groups can interact, males may attack females and infants, and so the infanticide risk for males and females with infants is likely to be high in these areas. Only females with infants, but not males with infants or females without infants, showed higher vigilance levels in overlap areas than in non-overlap areas; in addition, in overlap areas, females with an infant were more vigilant than females without an infant, while this was not the case in non-overlap areas. Both females and males with infants were more vigilant high in the trees than at medium heights in overlap areas but not elsewhere. These findings can only be explained by the infanticide avoidance hypothesis. In contrast to predator attacks, infanticidal male attacks come from high in the canopy, and only occur in overlap areas. There was a significant sex difference in vigilance, but males were only more vigilant than females without an infant, and not more vigilant than females with an infant. We conclude that vigilance varied mainly in relation to the risk of predation and infanticide. Mate competition only played a role for males during the early phase of their tenure. Predation risk seems to offer the best explanation for vigilance for all individuals in the absence of infants. Both predation risk and infanticide risk played a role for females and males with infants. Received: 4 April 1998 / Accepted after revision: 6 September 1998     

Social monogamy and the threat of infanticide in larger mammals

The evolution of social monogamy in larger mammals is difficult to explain because males usually do not invest much in direct offspring care and might achieve greater fitness by deserting a pregnant female to reproduce with additional females elsewhere. It has been hypothesized that socially monogamous males remain with the female year-round to protect their offspring from infanticide by new immigrant males. We investigated this idea by analyzing all cases of infant loss in a wild population of white-handed gibbons (Hylobates lar; Primates), in which most groups were socially monogamous and some polyandrous (137.5 group years). We examined the influence of (a) male intruder pressure on male immigration rates and (b) the presence of a new male in the group on infant loss. We found no relation between intruder pressure and male immigration rates. Infant loss was lowest (4.5%) for stable monogamy (probable father stayed from conception through infancy) and intermediate (25.0%; p = 0.166) for stable polyandry. If a new male immigrated after conception, however, the infant was lost in all cases (p < 0.01) independent of the presumed father’s presence. Overall, 83.3% of infant losses were associated with the presence of a presumably unrelated male. Although the sample size is small, our results provide the first true support for the idea that the risk of infanticide is an important factor in the evolution of social monogamy in larger mammals.     

Males as infant protectors in Hanuman langurs (Presbytis entellus) living in multimale groups – defence pattern, paternity and sexual behaviour

Hypotheses about the evolution of gregariousness and social organisation in primates are based on ecological explanations as well as on social factors such as conspecific threat (especially infanticide by males). The social explanation fits well with the conditions found in strepsirrhine primates and furthermore explains why infanticide in anthropoid primates living in one-male groups mainly occurs when the resident male (protector) is replaced. However, whether it likewise fits to the conditions in multimale groups will depend on the role of resident males as infant protectors, which has rarely been examined. We investigated long-term data of wild Hanuman langurs (Presbytis entellus) inhabiting a forest near the village of Ramnagar (southern Nepal). Twenty-two eye-witnessed attacks on infants by males were analysed in connection with male residency, paternity (DNA analyses) and sexual behaviour. Adult males played a major role in infant defence (65%). Only the genetic father or males who had been residents when the infant was conceived were observed to protect infants. Males who immigrated after a female had conceived may later attack her infant and were never observed to defend it. lt seems that the males took only copulations with potentially fertile females but not with pregnant females as clues for paternity. In the light of these results it seems likely that the risk of infanticide is an important determinant in female-male associations even in anthropoid primate multimale groups. Received: 22 December 1998 / Received in revised form: 30 April 1999 / Accepted: 1 May 1999     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Independent effects of male and female density on sexual harassment, female fitness, and male competition for mates in the western mosquitofish Gambusia affinis

Operational sex ratio (the ratio of sexually active males to fertilizable females) has a major influence on male competition for mates and male–female interactions. The contributions of male and female density per se to mating system dynamics, however, are rarely examined, and the fitness consequences are often inferred rather than quantified. Male mosquitofish (Gambusia affinis) compete aggressively and frequently harass females for copulations, a behavior thought to reduce female fitness. Female fitness can also be reduced by increases in female density, which may affect food availability, cannibalism rates, and chemical interactions between females. I manipulated male and female densities of G. affinis to measure their effects on male–male aggression, male harassment toward females, and female fitness. I found that males chased rivals more often and attempted fewer copulations when female density decreased, but surprisingly male density had no significant effect on the frequency of these male behaviors. In contrast, males’ agonistic displays toward other males increased with male density, but display behavior was unaffected by female density. These results suggest that male and female density do not always contribute equally or at all to the patterns of behavior we observe. Female fitness declined as female density increased, the opposite pattern expected if male harassment is costly to females. This suggests that a strong, negative effect of female density overwhelmed any potential costs of male harassment. Sources of female density dependence and the consequences of changes in male and female density to patterns of male behavior are discussed.     

Innate responses to male sexual harassment in female mosquitofish

Male mosquitofish are very persistent in their sexual activity and harass any female they encounter. Gravid females pay a large tribute to this intense male sexual activity in terms of reduced foraging efficiency. Previous observations have demonstrated that gravid females, when chased by a male, dilute male harassment by moving closer to other females to form shoals. They also approach other males to promote male competition, and when males differ in size, they preferentially target large males, whose harassment is less intense. In this study, we tested whether the modulation of females’ social preferences in response to male harassment is innate or learned. We tested social preference in three groups of females that differed in experience of sexual harassment and in the factors affecting it. Females of the first group were reared without any sexual experience, and pregnancy was induced through artificial insemination. The second group was composed of naive females kept singly with a male; these females experienced sexual harassment but were prevented from experiencing the effects of male–male competition and shoaling on the amount of male sexual harassment. In the third group (controls), females were reared in multi-male, multi-female groups and could experience the modulating effects of social interactions on sexual harassment. When exposed to a harassing male, females of the three groups immediately reduced their distance from another female, approached a group of males or moved toward the larger of two available males. Moreover, the results for these three groups of females were similar to those obtained in wild-caught females that were tested in the same three tests in a previous study (Dadda et al. An. Behav., 70:463–471, 2005). This suggests that the strategies adopted by females in response to male sexual harassment do not need to be learned through specific experience of the social contexts.     

Why male orangutans do not kill infants

Infanticide is widespread among mammals, is particularly common in primates, and has been shown to be an adaptive male strategy under certain conditions. Although no infanticides in wild orangutans have been reported to date, several authors have suggested that infanticide has been an important selection pressure influencing orangutan behavior and the evolution of orangutan social systems. In this paper, we critically assess this suggestion. We begin by investigating whether wild orangutans have been studied for a sufficiently long period that we might reasonably expect to have detected infanticide if it occurs. We consider whether orangutan females exhibit counterstrategies typically employed by other mammalian females. We also assess the hypothesis that orangutan females form special bonds with particular “protector males” to guard against infanticide. Lastly, we discuss socioecological reasons why orangutan males may not benefit from infanticide. We conclude that there is limited evidence for female counterstrategies and little support for the protector male hypothesis. Aspects of orangutan paternity certainty, lactational amenorrhea, and ranging behavior may explain why infanticide is not a strategy regularly employed by orangutan males on Sumatra or Borneo.     

Interactions in male baboons: the importance of both males’ testosterone

To date, research on testosterone and behavior has focused on individuals, even when studying social behaviors that necessarily involve multiple participants. Here, we explore male responses to other males of different dominance ranks and testosterone levels in a population of wild baboons. In chacma baboons (Papio hamadryas ursinus) of the Okavango Delta, a male’s testosterone is related to his rank trajectory and, therefore, the threat he poses to other males. To examine the effects of testosterone and rank on male–male interactions, we used playback experiments to measure how a target male responded to the simulated approach of another male, scoring responses by whether or not the subject moved away from the speaker in the first minute. High testosterone subjects did not move away from the speaker more often than low testosterone subjects, but high testosterone callers elicited a move more often than low testosterone callers. When the combined testosterone of the subject and caller was high, moves were most common. The rank relationship between subject and caller did not predict moves, but the effect of combined testosterone on moving was most pronounced in adjacently ranked males. Adjacently ranked, high testosterone males are the most likely to be competing for each others’ rank, and our experiments on these dyads elicited the most moves. Both behavioral and experimental observations indicate that testosterone may be more important than the rank relationship in predicting the outcome of male–male interactions. Furthermore, combined information on the testosterone of both males was the best predictor of results, highlighting the utility of dyadic analyses when relating testosterone to behavior.     

Female mate choice in the non-territorial lizard Ameiva plei (Teiidae)

Mate choice by females has been documented in a variety of taxa. Female mate choice in species lacking male resource control or paternal care might occur if preferred males provide protection from harassment. Female mate choice was investigated in a natural population of the non-territorial lizard Ameiva plei (Teiidae). Consort pairs were allowed to form naturally. Consort males were significantly larger than non-consort males. After removal of consort males, the “abandoned” female's reaction to the first male who approached her was recorded. Females rejected all small males. Female preference for large males was significantly higher than preference for small males. Large males may be better equipped to guard the females from harassment and behavior of large males is less harassing than behavior of small males, thereby affording the female increased foraging time. Received: 21 June 1996 / Accepted after revision: 28 December 1996     

Vocalizations by Alaskan moose: female incitation of male aggression

Evidence of female fomentation of male–male aggression as a mechanism of mate choice is rare, especially in mammals. Female choice of mates in polygynous species may be masked by intense male competition or by males attempting to restrict female choice. We studied protest moans of female Alaskan moose Alces alces gigas in interior Alaska, USA, from 1987 to 1990, to determine if moans incited male–male aggression. Alaskan moose exhibit a mating system in which one dominant male (the harem master) herds, defends, courts, and attempts to mate with females in his harem. Protest moans were given by females only in response to courtship. We hypothesized that if protest moans were related to females reducing harassment and exercising mate choice, females should give protest moans more frequently when courted by small males and less often when courted by large males, and that rates of male–male aggression would be elevated following protest moans. Harems were composed of one large male, with a mean of 4.4 females (median = 3 females); 10% of 132 harems included ≥10 females. The temporal pattern of protest moans from late August through November was associated with, but tended to lag behind, mating behavior. The rate of protest moans given by females decreased with increasing size of males courting them. Male–male aggression was significantly less during periods without protest moans than during periods in which protest moans occurred. These results indicate that female moose gave protest moans to reduce harassment by smaller males, and assure a mating opportunity with the most dominant male. Such a subtle mechanism of indirect mate choice by females may occur in other vertebrates in which choice is limited by a mating system in which male–male combat and male dominance over females reduces opportunities for female choice. The importance of female choice may be undervalued in studies of sexual selection in mammals.     

Male takeover,infanticide, and female countertactics in white-headed leaf monkeys (<Emphasis Type="Italic">Trachypithecus leucocephalus</Emphasis>)

Infanticide by males is common in mammalian species such as primates in which lactation lasts much longer than gestation. It frequently occurs in one-male groups following male takeovers and is likely a male reproductive strategy. Reported female countertactics include abrupt weaning of infants, dispersal, or paternity confusion. Here, we estimated costs of female countertactics in terms of weaning ages and interbirth intervals. We observed a population of white-headed leaf monkeys (Trachypithecus leucocephalus) in Nongguan Nature Reserve, China (1995–2006) mainly composed of one-male groups. Takeovers (N = 11) coincided with the peak conception period. Detailed data are presented for five takeovers (34 females, 29 infants, and 47 group-years) leading to six infant disappearances (42.9% of infant mortality). All presumed infanticides were in accordance with the sexual selection hypothesis. Following a takeover, females without infants or with old infants stayed with the new males, incurring no or low costs (via abrupt weaning). Females with young infants dispersing with the old males also experienced low costs. High costs (due to infant loss) were incurred by pregnant females and those with young infants who stayed with the new males indicating that paternity was not confused. Costs in terms of long interbirth intervals were also high for females leaving with the old males to later join the new males, despite infant survival. Female countertactics reflected female philopatry mediated by infant age. Presumably due to the seasonal timing of takeovers, most countertactics seemed successful given that 32.3% of females apparently incurred no costs and 41.2% incurred only low costs.     

Adoption or infanticide: options of replacement males in the European starling

The behaviour of a male bird towards a potential mate and her clutch may depend both on his expected paternity and on the likelihood that she will produce a replacement clutch if he commits infanticide. In this study we evaluate the choices made by replacement male European starlings Sturnus vulgaris. By removing males before and during laying, we induced other males, mainly neighbours, to mate with the reproductively active females. When the original male was removed before laying, a new male adopted the subsequent clutch in 14 out of 15 cases. When ten females were widowed during their laying period, replacement males never adopted their clutches. The paternity of replacement males was a function of when they replaced the former male. When replacement occurred more than 3 days before egglaying, the new male fathered nearly all offspring; when it occurred the day before laying, the new male still fathered more than every second young. When the original male was removed during his mate’s laying period, in five out of ten cases a replacement male committed infanticide by throwing out the eggs, but this only occurred in one out of 15 cases when removal took place before laying. The evidence for infanticide actually being committed by the replacement male was circumstantial. Four out of six of the females affected by apparent infanticide produced replacement clutches in which the male presumably had higher paternity than in the original clutch. In all cases, the male adopted the replacement clutch. In five cases when the original male was removed during laying, the neighbours neither adopted the brood nor committed infanticide, although they sometimes were seen courting the widowed female and copulating with her. These cases occurred later during laying than those were males comitted infanticide. The time from infanticide to the laying of the replacement clutch tended to increase as infanticide was committed later in the laying sequence. We conclude that strategies of potential replacement males are influenced by their expected paternity in the current brood and the probability that the female will produce an early replacement clutch. Received: 10 March 1995 / Accepted after revision: 28 October 1995     

O brother,where art thou? The varying influence of older siblings in rank acquisition by female baboons

Males in sexually dimorphic species like baboons appear to have surprisingly little influence on the reproduction and dominance ranks of their female kin, even though they could potentially increase their fitness by helping their relatives improve their ranks. Male baboons are able to dominate females several years before they emigrate, but their presence has no effect on relatives’ dominance ranks, at least when female kin are present. As a result, females usually acquire ranks within their matriline, above their older sisters. We describe the process of rank acquisition among orphaned and non-orphaned juvenile and adolescent females in a group of free-ranging baboons. Orphaned females were significantly more likely than non-orphaned females to acquire unexpected ranks. Orphaned females with older sisters often acquired ranks within the matriline, but below their older sisters’. Orphaned females with older brothers were likely to rise in rank above their matriline. Females’ interventions on behalf of younger sisters always supported the existing female dominance hierarchy, while males’ interventions tended to act against it. Similarly, in playback experiments, females appeared to be willing to support their younger sister only in disputes with lower-ranking females. In contrast, males appeared to be willing to support their sister even in disputes with higher-ranking females. Fraternal support enables females to improve their dominance ranks, but only if their mothers have died. It remains a puzzle why males have so little influence on their female relatives’ ranks when female kin are present, and so much when they are absent.     

Predation risk,food deprivation and non-random mating by size in the stream water strider,Aquarius remigis

Summary Non-random mating by size (NRMS) plays a central role in the study of sexual selection and the evolution of mating systems. Theory suggests that NRMS should be influenced by conflicting demands (e.g., predation risk, hunger); few experimental studies, however, have addressed these effects. We used a factorial experiment to examine the influence of predatory green sunfish and food deprivation on NRMS in male and female stream water striders, Aquarius remigis. As predicted by theory, food deprivation decreased the large-male mating advantage. The influence of predation risk, however, went against existing theory; that is, risk increased the large male mating advantage. The degree of large-male mating advantage was negatively related to a measure of the rate of male harassment of females. A behavioral mechanism that can explain these patterns emphasizes the contrasting effects of different competing demands on male harassment rates, female resistance and the role of male size in overcoming female resistance. Females usually resist male mating attempts. Successful mating occurs when males overcome female resistance. If harassment rates (of females by males) are low, larger males have a mating advantage over smaller males perhaps because females resist heavily and thus only larger males can overcome female resistance. If, however, male harassment rates are very high, female resistance might be swamped; mating should then be more random with respect to male size. Food deprivation increases gerrid activity and thus increases harassment rates which should then reduce NRMS. In contrast, risk decreases gerrid activity, thus decreasing harassment rates and increasing NRMS. Females did not show significant NRMS. Females did, however, show a pattern of change in NRMS that is consistent with male choice for larger females. Correspondence to: A. Sih     

Sexual conflict in the snake den

Red-sided garter snakes (Thamnophis sirtalis parietalis) court and mate in spring, soon after they emerge from large communal overwintering dens in south-central Manitoba. Because of a massive bias in the operational sex ratio, every female attracts intense courtship from dozens to hundreds of males. We suggest that this courtship constitutes significant ”harassment,” because it delays the females’ dispersal from the den and hence increases their vulnerability to predation. Small females may face the greatest costs, because they are less able to escape from amorous males (who court all females, even juvenile animals). Our measurements show that males are stronger and faster than females. Experimental trials confirm that the locomotor ability of females (especially small females) is greatly reduced by the weight of a courting male. Arena trials show that intense courtship stimulates females to attempt to escape. Remarkably, some females that are too small to produce offspring may nonetheless copulate. This precocious sexual receptivity may benefit juvenile females because copulation renders them unattractive to males, and thus allows them to escape more easily from the den. Female ”tactics” to escape male harassment may explain other puzzling aspects of garter snake biology including size-assortative mating, temporal patterns in dispersal from the den, avoidance of communal dens by young-of-the-year snakes, and female mimicry. Hence, sexual conflict may have influenced important features of the mating system and behavioral ecology of these animals. Received: 8 May 2000 / Revised: 28 July 2000 / Accepted: 30 July 2000     

The evolution of female social relationships in nonhuman primates

Considerable interspecific variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current “ecological model”, predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social relationships among gregarious females are here characterized as Dispersal-Egalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained differences in the occurrence of formal submission signals, decided dominance relationships, coalitions and female philopatry. Group size and female rank generally affect female reproduction success as the model predicts, and studies of closely related species in different ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the effects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment. Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains several phenomena that the ecological model does not, is a “socioecological model” based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail. Received: 30 September 1996 / Accepted after revision: 20 July 1997