Male–male competition and alternative male mating tactics influence female behavior and fertility in Japanese medaka (Oryzias latipes)

Intense male–male competition driven by high male density during mating can result in the evolution of alternative mating tactics that increase male fertilization success. The effects of alternative male mating tactics on females can range from increased fertilization and genetic benefits to decreased fertilization and loss of paternal care. However, the influence of male competitive behavior and alternative mating tactics on female behavior and reproductive success has seldom been addressed. In this work, I investigated the occurrence of alternative male mating tactics and their potential influence on female behavior and fertilization success in Japanese medaka (Oryzias latipes). Groups of one, two, or four males competed for access to a female in a repeated-measures experiment. Male density had a significant influence on female reproductive output as a result of a change in competitive mode from contest to scramble competition that coincided with more disruption during mating when more than one male attempted to mate. By contrast, sneaking during mating was beneficial to males, as more than one male sired offspring in most spawnings involving sneaker males. These results suggest that there may be conflict between males and females over mating, such that females are detrimentally affected by the occurrence of alternative mating tactics, whereas males may benefit from sneak mating. The occurrence of conflict between the sexes can be related to ecological factors, such as male density, which cause behavioral change in both males and females.     

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.     

Female choice in a promiscuous wild guinea pig,the yellow-toothed cavy (<Emphasis Type="Italic">Galea musteloides</Emphasis>)

Promiscuity is traditionally considered to increase only male reproductive success but, more recently, female benefits are also assumed to be the driving force for promiscuous mating. The yellow-toothed cavy (Galea musteloides) is characterised by an extremely high degree of multiple paternity (>80%), and females have greater offspring viability after mating multiple males. However, so far it is not clear whether or not it is the female's decision that leads to mating with more than one male. To elucidate the female's role in bringing about promiscuity, female yellow-toothed cavies were given the choice between four different males each, in a mate-choice apparatus that simultaneously prevented monopolisation and harassment of the females by the males. In 10 of the 12 choice tests, mating occurred. Nine of these ten females actively sought copulations with more than one male, and their mating behaviour was displayed in a way that might have favoured the mixing of sperm. At the same time, they significantly preferred heavier and more frequently courting males. These results show that female yellow-toothed cavies are actively involved in mating with more than one male. Thus, the present study is the first to show that in a species in which females produce more viable offspring as a consequence of polyandrous mating, these females are indeed motivated to actively bring about promiscuity. Nevertheless, females are selective in the choice of their mates. Thus, both sperm competition and direct female choice seem to be important for the increased offspring viability due to promiscuous mating.     

Fitness benefits of multiple mating versus female mate choice in the cellar spider (Pholcus phalangioides)

Recent studies have demonstrated that mating with multiple males can be beneficial for females and her offspring even if males contribute nothing but sperm. This was mainly established for species in which sperm from several males mix in the reproductive tract of the female, thus allowing sperm competition and/or female sperm choice. However, in species with last male sperm precedence, female re-mating decides against the previous male by strongly limiting his reproductive success. We tested the effect of female re-mating behaviour using the cellar spider Pholcus phalangioides, which shows strong last males sperm precedence and moderate levels of polyandry under natural situations. We predicted that females prevented from remating even though they are receptive would show reduced reproductive success compared to females that accept two copulations and females that reject a second male, since the latter two treatments were allowed to behave according to their decisions. However, if the number of matings per se had an effect on oviposition or on offspring performance, double-mated females should perform better compared to both treatments of once-mated females. We measured female fecundity and fertility over a period of 140 days, comparable to the species' natural reproductive peak season. Two thousand one hundred and fifty-two offspring from 67 first egg sacs were reared under two feeding levels. We registered development time and survival, and measured offspring adult size and mass. We found a positive effect of double mating, as in this treatment, oviposition probability was higher compared to the other treatments. Interestingly, adult female offspring of the DM treatment that were raised under low food level had a higher condition index compared to those from FS and RM, but development time, size and mass at adulthood were not affected by mating treatment. Female choice only seemed to affect hatching latency of the offspring. Overall, the main predictor of female reproductive output and success was female body size.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

Male genital length and mating status differentially affect mating behaviour in an earwig

Phenotypic variation in male genitalia may affect copulation behaviour, which can have important fitness consequences for males. Male genitalia commonly possess traits that increase male control over copulation, but in species where females control mating, a poor functional understanding often prevents insight into the processes responsible for such effects. Here, I investigate the effect of male genital length on copulation behaviour in the earwig Euborellia brunneri, where both sexes exhibit extremely elongated genitalia that correspond in shape. This model system is particularly suitable because pairs mate repeatedly and females can limit both the number and duration of copulations. I used both virgin and mated males and females in a double-mating design because longer male genitalia confer benefits in sperm competition. Consistent with a greater predicted male mating effort in mated females, the duration of individual copulations increased, but this traded off against mating frequency as cumulative mating duration remained unchanged. In contrast, male genital length increased both individual and cumulative mating duration, regardless of mating status. This difference suggests that, while males may modify copulation duration in response to mating status, females facultatively adjust mating frequency to prevent mating excessively or express preferences for increased male genital length. Notably, this study demonstrates that male genital phenotypes that are successful in sperm competition also enjoy female-mediated mating benefits.     

Influence of alternate reproductive tactics and pre- and postcopulatory sexual selection on paternity and offspring performance in a lizard

Sexual selection theory predicts different optima for multiple mating in males and females. We used mating experiments and genetic paternity testing to disentangle pre- and postcopulatory mechanisms of sexual selection and alternate reproductive tactics in the highly promiscuous lizard Eulamprus heatwolei. Both sexes mated multiply: 30–60 % of clutches were sired by two to four fathers, depending on the experiment. Larger males sired more offspring when we allowed male contest competition: 52 % of large males but only 14 % of small males sired at least one offspring. In the absence of male contest competition, females mated promiscuously and there was no large male advantage: 80 % of large males and 90 % of small males sired at least one offspring, and there was no evidence for last-male precedence. Multiple mating did not yield obvious direct or indirect benefits to females. E. heatwolei represents a complex system in which males attempt to improve their fertility success by limiting rivals from access to females and through adopting alternate reproductive tactics. Conversely, females exhibit no obvious precopulatory mate choice but may influence fitness through postcopulatory means by either promoting sperm competition or through cryptic female choice. Our results support the hypothesis that female multiple mating in nonavian reptiles is best explained by the combined effect of mate encounter frequency and high benefits to males but low costs to females.     

Factors affecting male and female reproductive success in a chipmunk (Tamias sibiricus) with a scramble competition mating system

While sexual selection in mammals with female defense systems has historically received considerable attention, much less is known about the factors affecting reproductive success in mammalian species with scramble competition mating systems. Using mark–recapture techniques and DNA microsatellite loci to assign parentage, we examined the mating system and factors affecting the variation of the annual reproductive success in an introduced population of the Siberian chipmunk, Tamias sibiricus, a small, monomorphic, solitary squirrel. Our analyses showed that breeding females were spatially dispersed, setting the stage for a scramble competition mating system. Male reproductive success was positively associated with the size of the home range. The strength of sexual selection on this behavioral trait was very strong, equaling previous estimates for morphological traits in female defense mating systems. These findings suggest that a behavioral trait, space use, strongly influences the annual reproductive success in males, which is consistent with expectations in a scramble competition mating system. In both sexes, reproductive success was influenced by habitat, with twice as many juveniles produced in semi-open than in closed habitat, possibly due to differences in food availability between the two habitats.     

Sexual dimorphism and the mating ecology of polar bears (Ursus maritimus) at Svalbard

We assessed the role of size, mass, and age in mating and non-mating polar bears (Ursus maritimus) at Svalbard, Norway, during the spring breeding season. The ratio of male to female mass, in male-female pairs, ranged from 1.00 to 3.02 ([`(x)] = 1.99 \overline x = 1.99 ) indicating that mating males were larger than mating females but with substantial variation. Paired males were older than unpaired males and male mass was related to age. However, males paired with females were not significantly different in body mass from those males caught alone. Wounds and scars resulting from fights between males began at about 6 years of age and peaked at about 17 and 20 years of age, respectively. The frequency of broken canines in males, presumably due to increased male-male conflicts, increased with age but showed little increase in females. The wide range of male size in male-female pairs and the age-related signs of injury suggest that male polar bears engage in both scramble competition and contest competition for access to breeding females. The mating system of polar bears is variable but is best described as female defense polygyny or serial monogamy.     

Sex in the dark: determinants and consequences of mixed male mating tactics in<Emphasis Type="Italic"> Microcebus murinus</Emphasis>, a small solitary nocturnal primate

In mammals with solitary females, the potential for males to monopolize matings is relatively low, and scramble competition polygyny is presumed to be the predominant mating system. However, combinations of male traits and mating tactics within this type of polygyny have been described. The main aim of our study was to identify the relative importance of, and interactions among, potential determinants of contrasting male reproductive tactics, and to determine their consequences for male reproductive success in a small solitary nocturnal Malagasy primate, the gray mouse lemur (Microcebus murinus). We studied their mating behavior over three consecutive annual mating seasons. In addition, we determined the genetic relationships among more than 300 study animals to quantify the reproductive success of individual males. We found that, with a given relatively low overall monopolization potential, successful male mouse lemurs roamed extensively in search of mates, had superior finding ability and mated as early as possible. However, contest competition was important too, as temporary monopolization was also possible. Males exhibited different mating tactics, and heavier males had a higher reproductive success, although most litters had mixed paternities. Switching between tactics depended on short-term local variation in monopolization potential determined by a pronounced dynamic in fertilization probability, number of alternative mating opportunities, and the operational sex ratio. This study also revealed that the dynamics of these determinants, as well as the mutual interactions between them, necessitate a detailed knowledge of the mating behavior of a species to infer the impact of determinants of alternative mating tactics.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by S. AlbertsThis revised version was published online in August 2004 with corrections to Figure 2.     

Extra-pair copulations in a lek: the secondary mating system of monogamous razorbills

Summary I studied the mating strategies of razorbills Alca torda, a monogamous colonial seabird, during 1987–1989 in Wales, U.K. The outstanding feature of the study population's mating system was the performance of most copulations in mating arenas outside the colony. Females visited the arenas and sometimes accepted extra-pair copulations (EPCs), and males aggressively competed for EPCs. During the fertilizable period of most females, the arenas resembled leks because they contained no resources except mates, and males contributed only sperm to extra-pair females. Typical of leks, the operational sex ratio was strongly male-biased, and male mating success was highly skewed. To examine the potential for sexual selection to operate in this system, I correlated male EPC success with variables that could be associated with male-male competition and/or female choice. The frequencies of three behaviors (interference of copulation attempts, fights won against other males, and EPC attempts), accounted for 62% of the variance in EPC success and were strongly intercorrelated, with interference explaining most of the variance in a multivariate test. EPC success was not correlated with body size, age, paired status or either of two estimates of the time spent in the arena. The hypothesis that male EPC success was determined by active female choice was tested by examining the relative success (EPCs per attempt) of males. Relative success was not correlated with any of the three behaviors associated with absolute success, or with age, body size or attendance, suggesting that male-male competition, mainly in the form of copulation interference, is the principal correlate of EPC success. These findings in a monogamous species illustrate that EPC is a secondary mating system of razorbills in which sexual selection operates. The existence of this system outside the colony raises the possibility that lekking for EPCs may also occur within the nesting territories of other monogamous, colonial species but is hidden by competition for nests and breeding partners.     

Pre- and post-mating sexual selection both favor large males in a rainbowfish

Sexual selection can act through female choice and male–male competition. Although both processes can act simultaneously, they are typically studied independently. Here, we adopt a more integrated approach to studying sexual selection by incorporating measures of both processes using the western rainbowfish Melanotaenia australis, a freshwater fish endemic to northwestern Australia. We assessed male–male competition and female choice separately while measuring the performance of individual males under both processes and used paternity analyses to estimate male reproductive success. We then related the performance of males during each of these stages to their phenotype, which was described using linear measures of size and color pattern traits, and spectrographic measures of the reflectance of color patches. We found that female choice favored relatively large males and that these preferences were consistent within individual females and repeatable between different females. Larger males were also more dominant in the competition trials and sired the majority of offspring produced when females spawned. There was little evidence to suggest that sexual selection acted on male color patterns either via female choice or male contest competition or during subsequent post-mating episodes of sexual selection. We conclude, therefore, that male–male competition and female choice act concordantly to favor relatively large males and that these patterns of mating success are reflected during post-mating episodes of sexual selection.     

Body size and sexual selection in the koala

Sexual selection is often characterized by polygynous breeding systems, size dimorphism, and skewed operational sex ratios. Koalas are sexually dimorphic in multiple domains, yet are absent from the literature on sexual selection and the structure of their mating system is unclear. We provide the first documentation of the strength of sexual selection in koalas by using microsatellite markers to identify sires. We combine the genetic data with morphological data in order to assess the role of body size in regulating reproductive output. During our 4-year study, 37% of males were identified as possible sires. Males were significantly larger than females, with sires heavier than non-sires. Male body mass correlated with annual reproductive output, with Crow’s Index of Opportunity for Selection revealing that variation in male reproductive success was threefold higher than that of females. Since it appears that male koalas rarely engage in physical confrontations over access to females, size dimorphism could be based upon non-agonistic competition and/or female mate choice. We propose that size dimorphism in koalas evolved as a consequence of endurance rivalry promoting vocal sexual advertisements that attract females. We suggest that female choice is a key mediator of male reproductive output.     

Scramble competition selects for greater tailfin size in male red-spotted newts (Amphibia: Salamandridae)

Darwin predicted that scramble competition for access to reproductive females would result in sexual dimorphism of locomotory structures, but direct evidence for this is extremely rare. I examined the relationship between variation in tailfin size and mating success in a field and laboratory study of red-spotted newts, Notophthalmus viridescens. Over three breeding seasons, male tailfin size was positively correlated with variation in male amplexus frequency, and indirectly correlated with male insemination frequency. In a laboratory study, I confirmed that males' ability to capture females is affected by variation in tailfin size. This is the first study to show that naturally occurring variation in male locomotory structures affects male mating success. It corroborates the prediction that scramble competition leads to sexual selection on locomotory structures and, potentially, to dimorphism in these structures. Received: 16 April 1999 / Received in revised form: 16 May 1999 / Accepted: 12 June 1999     

No cost of male mating experience on female reproductive success in the almond moth, <Emphasis Type="BoldItalic">Cadra cautella</Emphasis> (Lepidoptera; Pyralidae)

Male copulation experience may have a profound impact on female reproductive success if male reproductive investment declines over consecutive copulations and if females are unlikely to re-mate. Male reproductive investment is particularly interesting in lepidopterans because males produce dimorphic sperm: a fertilizing (eupyrene) and a non-fertilising (apyrene) sperm. In two experiments, we explored the lifetime reproductive investment of male almond moths, Cadra cautella (also known as Ephestia cautella) and examined its influence on female reproductive success. In the almond moth, females re-mate infrequently and males transfer sperm in a spermatophore. Attached to the spermatophore is a large chitinous process, the function of which is unknown. One group of males were permitted consecutive copulations with virgin females and the amount of sperm and size of the spermatophore transferred were compared for all females. We found that the number of both eupyrene and apyrene sperm per ejaculate decreased with his increased mating frequency, while the size of the spermatophore process decreased dramatically after the male’s first copulation. In a second experiment, we allowed males to mate with females throughout their lives and then compared female fecundity and fertilisation success. We found no obvious decrease in female fecundity and fertilisation success with increased male copulation experience, despite the likely reduction in male gametic investment. We discuss potential explanations for the development of this enlarged and elaborate first spermatophore of male almond moths given that it confers no clear fitness advantage to females.     

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 efficacy of female choice in chimpanzees of the Taï Forest, Côte d’Ivoire

Male–male competition has historically been considered the major force driving sexual selection. However, female choice and inter-sexual conflict are increasingly recognized as important influences affecting differential mating and reproductive success. Many females exhibit preferences for particular males; however, male strategies may conflict with females’ ability to obtain their mate preferences. To influence paternity, females must affect both (1) whether or not sexual interactions occur, particularly during the periovulatory period (POP) and (2) the outcome of sexual interactions. This study focuses on the effectiveness of female choice in wild chimpanzees (Pan troglodytes verus). Over 2,600 h of data were collected on two habituated chimpanzee communities in the Taï National Park, Côte d’Ivoire. Female mate preferences were measured by quantifying proceptive and resistance behavior toward males in both the periovulatory period and non-POP phases of estrus. The efficacy of female preference was measured both (1) by measuring success rates of female proceptivity and resistance behaviors and (2) by determining how well measures of female mate preference (proceptivity and resistance rates) predict male mating success. Though male chimpanzees are clearly dominant to females, the results indicate that females could effectively resist male solicitations and, in most cases, unwanted copulations were averted. Both female proceptivity and resistance rates correlate (positively and inversely, respectively) with male mating success in POP. Outside POP, female proceptivity rates corresponded with male mating success, but resistance rates did not. Males (irrespective of rank) that were preferred by females obtained higher mating success compared to other males during the POP, suggesting that females were effective in their mate choice and that, despite clear male dominance, female choice influences paternity in wild chimpanzees.     

Experimental assessment of ecological and phenotypic factors affecting male mating success and polyandry in northern watersnakes, Nerodia sipedon

To resolve conflicting field observations regarding the action of sexual selection, we used breeding experiments and paternity analysis of the 927 resulting offspring to assess how male size, condition, tail length, genetic similarity to the female, and variation in operational sex ratio (OSR) affected male reproductive success and the incidence of polyandry in northern watersnakes (Nerodia sipedon). Only size affected male mating success. Large males were more successful, but only when male size varied substantially and competition among males was intense (i.e., male-biased OSR). The conditional nature of the size advantage may explain why studies of free-living watersnakes have produced inconsistent results regarding the relationship between male size and mating success. Size differences between males did not affect the proportion of offspring each male sired within multiply sired litters. We found positive size-assortative mating, but only when the OSR was female biased, suggesting that smaller males had improved access to females when competition among males was reduced, but that competition with larger males still restricted mating opportunities of small males to less preferred, smaller females. Most litters (58%) were multiply sired and larger females were more likely to produce multiply sired litters, similar to free-living watersnakes. There was no association between the incidence of multiple paternity and OSR, however, suggesting that polyandry is not simply a function of opportunity, with females passively waiting for males to court them.     

Quantifying male attractiveness and mating behaviour through phenotypic size manipulation in the Trinidadian guppy, Poecilia reticulata

Although many studies have examined the effects of male size on attractiveness and mating behaviour, few have taken genetic background into consideration. Phenotypic manipulation permits the experimental adjustment of morphological traits while keeping genetic background constant. Here, male guppies, Poecilia reticulata, an ideal model for this type of manipulation, were raised at different temperatures to produce sibling pairs that differed in size. These were then used to investigate male mating behaviour and male attractiveness, assessed through female mate choice, in relation to this size dimorphism. Further, male–male competition, which is intrinsic to male mating behaviour, is also likely to be affected by their size. Through the use of repeated measures analyses we demonstrate that females significantly prefer larger males and male size and competition significantly affect several aspects of male mating behaviour. Larger siblings perform more sneaky mating attempts and spend more time chasing females. The frequencies of both these behaviours increase with competition. While display frequency is unaffected by male size and competition, display duration and the amount of time spent attending females are reduced in the presence of competitors. This study highlights the use of phenotypic manipulation as a valuable tool for investigating behavioural interactions and confirms that both male size and competition are significant factors in the guppy mating system.     

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.