Infanticide in seasonally breeding multimale groups of Hanuman langurs (Presbytis entellus) in Ramnagar (South Nepal)

In the seasonally breeding langur (Presbytis entellus) population of Ramnagar, South Nepal, where multimale groups prevail, 25 attacks on 11 infants (including one actual killing) by seven adult males were witnessed in five groups by six observers between 1990 and 1996. Circumstantial evidence also indicates three additional attempts at infanticide and in seven additional cases infanticide was presumed or likely. Infanticide presumably accounted for 30.8–62.5% of infant mortality in the first 2 years of life. Most attackers (91.4%) were residents of the infants' group and had immigrated after the infants had been born (75.0%) or conceived (25.0%). Thus, they were not related to the victims. The interbirth interval was shortened if an infant died either prior to September of its 1st year of life (mean = 1.2 years), or its 2nd year (mean = 2.0) and even its 3rd year (mean = 2.4). All attackers remained in the group at least until the next mating season; high-ranking males maintained their dominance rank and lower-ranking males rose in rank. Since rank and mating success were correlated and rank and reproductive success might be correlated, all attackers had a good chance of siring the next infant of the victims' mothers and could thus have benefited by their action. Infanticide seems to be a male reproductive strategy at Ramnagar. Infanticide has never before been reported among seasonally breeding langurs living at such low densities. This is also the first detailed report of infanticide as a male reproductive strategy in a seasonally breeding primate population. Received: 19 December 1996 / Accepted after revision: 7 June 1997     

Female bank voles (Clethrionomys glareolus) prefer dominant males; but what if there is no choice?

Both intra-sexual competition between males and female mate choice have been found to affect mating behaviour in rodents. We studied female choice in the bank vole (Clethrionomys glareolus) under circumstances where male-male competition was excluded and the female could interact and mate freely with the males. Mating behaviour was observed in two situations: (1) the female encountered two males with a clear dominance relationship; and (2) the two males were equal in their social status. In all tests where a female in postpartum oestrus had a choice between males of different social rank she mated with the dominant one. When choosing between an even pair of males there was no difference in the frequency of lordotic responses, mounts or intromissions the female exhibited with either male before mating with one of them. The mean ejaculation latency was significantly longer in the tests with an even pair of males than in the tests where male hierarchy was clear. These results show that bank vole females are able to discriminate males according to their social status and strongly prefer dominant males as mating partners. However, when the females were presented with two equal males, they seemed to be unable to make a choice. The ability to choose the mating partner when the males are clearly different may be an important direct benefit for the female in terms of time saved during mating and thus decreased risk of predation and infanticide. Received: 16 March 1995/Accepted after revision: 11 February 1996     

Does female aggression prevent polygyny? An experiment with pied flycatchers (Ficedula hypoleuca)

The female aggression hypothesis states that resident females may be able to prevent polygyny by behaving aggressively towards intruding females. A critical test of the hypothesis is to provide prospecting females with a choice between displaying mated males some of which have initial mates with artificially reduced levels of aggressiveness. Here we present a mate choice experiment on pied flycatchers Ficedula hypoleuca. The species is a cavity nester, and resident females were prevented from behaving aggressively by enclosing them within their own nestboxes: narrowing the entrance hole so that they could not escape but could still let their head out and have some contact with their mate. This treament had only a minor influence on male behaviour. We studied whether the experimental males were better able to attract a new female than a control group of mated males. Four predictions from the female aggression hypothesis were supported. (1) Mating success of control males was positively related to the distance between their primary and secondary territory. (2) For experimental males, mating success was unrelated to interterritorial distance. (3) Experimental males had higher mating success than control males when the interterritorial distance was short but (4) not when it was long. Experimental males had much lower mating success than unmated males, as would be expected if prospecting females are able to discover male mating status from cues other than visits by primary females to their mates' secondary nest sites. Received: 5 January 1998 / Accepted after revision: 30 December 1998     

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     

Coalitionary mate guarding by male chimpanzees at Ngogo, Kibale National Park, Uganda

Cooperative mate guarding by males is unusual in mammals and birds, largely because fertilizations are non-shareable. Chimpanzees live in fission-fusion communities that have cores of philopatric males who cooperate in inter-group aggression and in defending access to the females in their community. Male contest mating competition is restrained within communities, but single high-ranking males sometimes try to mate guard estrous females. Data from an unusually large chimpanzee commmunity at Ngogo, Kibale National Park, Uganda, that contains more males than any previously studied community show new variation in chimpanzee mate-guarding behavior. Contrary to expectation given the large number of males, mate guarding was as common as, or more common than, at other sites, and males other than the alpha male guarded more often. More strikingly, pairs or trios of top-ranking males sometimes engaged in cooperative aggression to prevent estrous females from mating with other males, but tolerated each other's mating activities. Both single males and coalitions mostly guarded periovulatory females. Mate-guarding coalitions were previously unknown in chimpanzees. Coalitions occurred in large mating parties, seemingly because these often contained too many males for single males to maintain exclusive access to estrous females. Coalition members gained higher shares of copulations than they could have expected from solo mate guarding, and suffered lower per capita costs of guarding (as inferred from aggression rates). Two males who most often participated in coalitions formed two-male coalitions at about the point where the number of males present made it unlikely that either could get 50% or more of total copulations on his own, and formed trios when this value dropped below 33%. Kin selection could be a factor in cooperation among male chimpanzees, but coalition members were not necessarily close relatives and the apparent structure of payoffs fit that of mutualism. Furthermore, reliance of male chimpanzees on support from allies to maintain high rank could have led to trading of mating exclusivity for support against mating competitors. Received: 28 May 1997 / Accepted after revision: 16 May 1998     

“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.     

Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits <Emphasis Type="Italic">Remiz pendulinus</Emphasis>

Melanin-based ornaments are often involved in signaling aggression and dominance, and their role in sexual selection is increasingly recognized. We investigated the functions of a melanin-based plumage ornament (facial ‘mask’) in male Eurasian penduline tits Remiz pendulinus in the contexts of male–male aggression, mating success, and parental care. The penduline tit is a passerine bird with a unique mating system in which both sexes may mate with several mates in a breeding season, and one (or both) parent deserts the clutch. Our study revealed that mask size of males is more likely an honest signal used by females in their mate choice decisions than a trait involved in male–male competition. First, mask size increased with both age and body condition, indicating that the mask may signal male quality. Second, males with larger masks paired more quickly and had more mates over the breeding season than males with smaller masks. Third, we found no evidence that male mask size signals male–male aggression or dominance during competitive encounters. The increased mating success of large-masked males, however, did not translate into higher reproductive success, as nestling survival decreased with mask size. Therefore, we conclude that there is either no directional selection on male mask size or males with larger masks receive indirect, long-term benefits.     

Post-copulatory aggression toward their mates by males of the rove beetle Leistotrophus versicolor (Coleoptera: Staphylinidae)

Summary Rove beetles (Leistotrophus versicolor) forage and mate at dung and carrion in the riparian forest of northwestern Costa Rica. After copulating, males often launch a post-copulatory attack against their recent partner. We test four hypotheses on the adaptive value of male behavior: (1) The sperm competition hypothesis proposes that the behavior may be the functional equivalent of mate-guarding, (2) the sperm-transfer signal hypothesis states that males bite their mates after copulating to signal that they have successfully passed sperm, (3) the feeding competition hypothesis argues that male aggression toward mates occurs to drive away competitors for fly prey, and (4) the redirected aggression hypothesis is that male attacks after mating occur when threatened males redirect their aggression onto their partners. Only the sperm competition hypothesis withstands testing. As required by this hypothesis, females are usually receptive while at dung, and will mate with more than one male in a morning. In addition, males are more likely to attack a mate when they have fought earlier in the day with other males, an indicator of the presence of rival males and the risk of sperm competition. Contrary to the sperm-transfer signal hypothesis (2), biting of mates does not occur after nearly 40% of all copulations; it seems unlikely that mating males so often fail to transfer sperm. Whether males have fed or not prior to mating has no effect on the probability of post-copulatory attack, a result that contradicts the food competition hypothesis (3). Finally, the occurrence of attacks by males on females in the absence of an immediate threat from a rival argues against the redirected aggression hypothesis (4).     

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     

Mate quality,not aggressive spillover,explains sexual cannibalism in a size-dimorphic spider

Sexual cannibalism particularly before mating is costly for the male victim but also for the female aggressor if she risks remaining unmated. The aggressive spillover hypothesis explains the persistence of this behavior as a maladaptive side effect of positive selection on aggressiveness in a foraging context. The hypothesis predicts that the occurrence of sexual cannibalism is explained by female aggressiveness but is not related to male phenotype or behavioral type. An alternative hypothesis invokes sexual selection and makes the opposite prediction namely that sexual cannibalism is an expression of female choice and should hence mainly target males of low quality. We tested the above hypotheses on a sexually dimorphic nephilid spider Nephilengys livida, known for male monopolization of females via genital damage, female genital plugging, and mate guarding, by staging mating trials during which we recorded mating behaviors and occurrences of pre- and postcopulatory cannibalism. We did not restrict assessment of aggressiveness to the mating and foraging context but also included aggression against same sex conspecifics. To assess female personalities, i.e., consistent individual differences in behavior including aggressiveness, we repeatedly tested them for intra-sex aggression, voracity towards prey, locomotory activity, and boldness. Females exhibited consistent differences in intra-sex aggressiveness, latency to attack prey, and boldness. Aggressive females had shorter latencies to attack prey and were more active than non-aggressive ones. In contrast to the predictions of the aggressive spillover hypothesis, females that were aggressive towards prey and towards other females were not more likely to attack a male than non-aggressive females. In support of the mate choice hypothesis, less aggressive males were more likely attacked and cannibalized than more aggressive ones. This hints at sexual selection for aggressiveness in males and raises the question of mechanisms that maintain variation in male aggressiveness.     

Sex-specific reproductive behaviours and paternity in free-ranging Barbary macaques (Macaca sylvanus)

In a wide variety of species, male reproductive success is determined by contest for access to females. Among multi-male primate groups, however, factors in addition to male competitive ability may also influence paternity outcome, although their exact nature and force is still largely unclear. Here, we have investigated in a group of free-ranging Barbary macaques whether paternity is determined on the pre- or postcopulatory level and how male competitive ability and female direct mate choice during the female fertile phase are related to male reproductive success. Behavioural observations were combined with faecal hormone analysis for timing of the fertile phase (13 cycles, 8 females) and genetic paternity analysis (n = 12). During the fertile phase, complete monopolisation of females did not occur. Females were consorted for only 49% of observation time, and all females had ejaculatory copulations with several males. Thus, in all cases, paternity was determined on the postcopulatory level. More than 80% of infants were sired by high-ranking males, and this reproductive skew was related to both, male competitive ability and female direct mate choice as high-ranking males spent more time in consort with females than low-ranking males, and females solicited copulations mainly from dominant males. As most ejaculatory copulations were female-initiated, female direct mate choice appeared to have the highest impact on male reproductive success. However, female preference was not directly translated into paternity, as fathers were not preferred over non-fathers in terms of solicitation, consortship and mating behaviour. Collectively, our data show that in the Barbary macaque, both sexes significantly influence male mating success, but that sperm of several males generally compete within the female reproductive tract and that therefore paternity is determined by mechanisms operating at the postcopulatory level.     

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     

Sexual coercion by male chimpanzees shows that female choice may be more apparent than real

The extent to which active female mating preferences influence male reproductive success in mammals is unclear, particularly for promiscuously breeding species like chimpanzees (Pan troglodytes). Previous studies from multiple long-term study sites have shown that female chimpanzees mate more restrictively around ovulation, and this has been taken as evidence for female choice. However, none of these studies rigorously evaluated the alternative hypothesis that restrictive mating results not from unconstrained choice, but in response to coercive mate guarding, in which males use punishment and intimidation to reduce female promiscuity and promote their own mating interests. Nor did they consider evidence for the potential genetic or phenotypic benefits that females might be choosing. Using 11 years of data from the Kanyawara community in Kibale National Park, Uganda, we previously demonstrated that males achieve elevated mating success with those females toward whom they direct high levels of aggression. Here we extend those findings to show that even female copulatory approaches, which have previously been attributed to female choice, are correlated with male aggression. Specifically, individual females at our site initiated periovulatory copulations most frequently with the males who were most aggressive toward them throughout their cycles. Those males showed high rates of aggression toward females throughout estrus, despite achieving high copulation rates, demonstrating a continuing conflict of interest over the exclusivity of mating access. Because sexual coercion is potentially widespread in primates and other mammals, our results stress the importance of considering the influence of male aggression in studies of female choice.     

Forced dispersal of juvenile guanacos (<Emphasis Type="Italic">Lama guanicoe</Emphasis>): causes,variation, and fates of individuals dispersing at different times

We examined adult-juvenile conflict in the guanaco (Lama guanicoe). During spring, territorial males become increasingly aggressive toward all juveniles born the previous year and begin expelling them from family groups. In an apparent effort to reduce aggression, juveniles display submissive crouches when being observed, approached, or attacked by the territorial male. Therefore, we assessed the influence of juvenile submissive behavior on the timing of dispersal and also examined if dispersal time was related to survival and reproductive performance as adults. We also evaluated hypotheses regarding the evolution of juvenile mammalian dispersal in the context of if and how each may favor the forced dispersal of juvenile guanacos by territorial males. Juveniles generally dispersed in late spring and early summer, and a nearly equal proportion of females (n=46; 48%) and males (n=49; 52%) dispersed. More-submissive animals generally dispersed later than less-submissive animals. Juvenile sex and dispersal time were not related to survival. In contrast, juvenile sex and dispersal time were related to reproductive performance. The probability of reproducing was highest when juveniles dispersed early and decreased with increasing time in family groups prior to dispersal. The largest proportion of juveniles was forced to disperse during a 2-week interval following the peak of the breeding season. Competition for food resources is likely very intense at this juncture and territorial males may force older juveniles to disperse in order to divert food resources to younger neonates. Additionally, juveniles may be forced to disperse after territorial males mate their mothers to prevent lost mating opportunities, because females leave territories when their offspring disperse and possibly prior to mating with males. We conclude that the forced dispersal of juvenile guanacos by territorial males is ultimately driven by competition for food resources on territories. The timing of dispersal, however, may be tempered by the chronology of matings between territorial males and particular adult females, and/or genetic relatedness between territorial males and juveniles.     

Female mate choice,male migration,and the threat of infanticide in ringtailed lemurs

Summary Paternity determination by DNA fingerprinting is reported for a long-term study group of semi-free-ranging ringtailed lemurs (Lemur catta), together with behavioral data collected independently. In 1985, fraternal twin males unfamiliar and unrelated to the resident ringtailed lemurs were introduced to the forest enclosure. Every mature resident male attacked the immigrants frequently across the next 5 months, whereas no female ever did. All observed estrous females showed sexual proceptivity toward the' immigrant males; three solicited copulation exclusively from them. Each female repelled sons, matrilineal brothers, and other resident males from attempting to copulate. Over a 5-year period, four of five females always reproduced with distantly related or unrelated males (Fig. 3). Despite low dominance status throughout the case study, an immigrant sired the off-spring of each female that was proceptive toward only the immigrants, demonstrating that female choice can override male dominance relations to determine reproductive success among male ringtailed lemurs. In the birth season following the 1985–1986 immigration, each of four females targeted one or two particular adult males for consistent attack across the period of infant dependency, beginning days after parturition. Paternity determinations, colony records, and subsequent study of two groups allowed 66 cases of this mode of maternal aggression to be documented. In each, the targeted male had not fathered the protected infant, and almost invariably, he was unrelated to the infant's mother. New mothers attacked every male that immigrated following their infants' conceptions and a few familiar males with whom they had not been seen to copulate during the previous breeding season. Recent attempts by immigrant males to kill infants confirmed the anti-infanticidal function of maternal targeting of males. All results were interpreted together to advance a prospective model of the mating system of ringtailed lemurs. Female avoidance of incest has led to the evolution of natal male dispersal. Subsequently, males should prefer to transfer into groups containing few and/or status-vulnerable males. We predict that, by killing others' infants, males simultaneously increase chances for success in females' next reproductive efforts and terminate current fathers' reproductive eligibility in a group. Basic hypotheses that await testing are that (a) raising an infant through weaning reduces a female's chances for reproductive success the following year and (b) males that demonstrate the capacity to promote the survival of infant offspring are most attractive to females as mates.     

Female tactile cues maximize paternal behavior in prairie voles

Most monogamous male mammals display parental care, but the mechanism by which they become motivated to care for infants remains unknown. We used prairie voles (Microtus ochrogaster) as a model of biparental species to test the hypothesis that physical contact with a female enhances male parental responsiveness before the birth of offspring. We tested four groups of males: mated with physical contact with their mate, mated with only distal cues from their mate, mated with no cues, and unmated with distal cues from a female. The subjects were placed in an oversized cage divided into two compartments by a perforated barrier. At mid-gestation, the bedding and females were removed, and the males’ responses toward two infants were videotaped. Although most males behaved parentally, mated males exposed to their mate’s tactile or distal cues approached the young faster and were more likely to care for both infants than unmated males which received female distal cues. Males who had physical contact with their mate showed “kyphosis” (a crouching position over infants) and contacted infants more frequently than mated males which received no cues from their mate or unmated males which received female distal cues. Furthermore, the frequency of non-social behaviors was lowest in mated males that had physical contact with their mate. The data suggest that exposure to female distal cues after mating is sufficient to increase male parental attentiveness, but female somatosensory cues affect the male's tendency to remain in physical contact with infants.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

The evolution of female copulation calls in primates: a review and a new model

Female copulation calls are mating-associated vocalizations that occur in some species of Old World monkeys and apes. We argue that copulation calls have two immediate functions: to encourage mating attempts by other males and to increase mate guarding by the consort male. We hypothesize that female copulation calls have evolved under the selective pressures of risk of infanticide and sperm competition. When male mate guarding is effective, copulation calls allow females to concentrate paternity in dominant males and benefit from their protection against the risk of infanticide. When mate guarding is ineffective, copulation calls may bring genetic benefits to females through facilitation of sperm competition. We present a quantitative model in which interspecific variation in females' promiscuity predicts their tendency to use copulation calls in conjunction with mating. The model predicts that in species with little female promiscuity, copulation calls should be rare and exhibited only in association with mating with dominant males. In species in which females are highly promiscuous, copulation calls should be frequent and unrelated to male dominance rank. The limited data available to test the model support its main predictions as well as the predicted relation between copulation calls and male dominance rank.

Prior encounters with the opposite sex affect male and female mating behavior in a wolf spider (Araneae,Lycosidae)

Mate availability can vary widely in nature depending upon population density and sex ratio and can affect the ability of individuals to be selective in mate choice. We tested the effects of prior encounters with the opposite sex (i.e., exposure to the opposite sex either with or without mating) on subsequent mating behavior in two experiments that manipulated mate availability for both males and females in the wolf spider, Hogna helluo. The probability of mating in the experimental trial depended upon whether the prior encounter involved mating or not, and males and females responded in opposite directions. Exposure without mating resulted in a higher subsequent frequency of mating for females and a lower subsequent frequency of mating for males, while prior mating experience resulted in a lower frequency of female remating and a higher frequency of male remating. Prior exposure without mating did not affect female aggression. However, mated females engaged in precopulatory cannibalism more frequently than virgins. Mated males escaped postcopulatory cannibalism more frequently than virgins. Our results show that males respond to exposure without mating in the expected manner. However, prior mating (1 week earlier) had unexpected effects on males, which may be due to mated males being of higher quality. There were little or no effects of the size of the prior exposure individual or mate on subsequent mating behaviors. Further research is needed to determine why different species use different degrees of prior information in mate choice.     

The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.