High multiple paternity and low last-male sperm precedence in a hermaphroditic planarian flatworm: consequences for reciprocity patterns

It is difficult to predict a priori how mating success translates into fertilization success in simultaneous hermaphrodites with internal fertilization. Whereas insemination decisions will be determined by male interests, fertilization will depend on female interests, possibly leading to discrepancies between insemination and fertilization patterns. The planarian flatworm Schmidtea polychroa, a simultaneous hermaphrodite in which mating partners trade sperm was studied. Sperm can be stored for months yet individuals mate frequently. Using microsatellites, maternity and paternity data were obtained from 748 offspring produced in six groups of 10 individuals during four weeks. Adults produced young from four mates on average. Reciprocal fertilization between two mates was found in only 41 out of 110 registered mate combinations, which is clearly less than what is predicted from insemination patterns. Multiple paternity was high: > 80% of all cocoons had two to five fathers for only three to five offspring per cocoon. Because animals were collected from a natural population, 28% of all hatchlings were sired by unknown sperm donors in the field, despite a 10-day period of acclimatization and within-group mating. This percentage decreased only moderately throughout the experiment, showing that sperm can be stored and used for at least a month, despite frequent mating and sperm digestion. The immediate paternity a sperm donor could expect to obtain was only about 25%. Male reproductive success increased linearly with the number of female partners, providing support for Bateman's principle in hermaphrodites. Our results suggest that hermaphrodites do not trade fertilizations when trading sperm during insemination, lending support to the view that such conditional sperm exchange is driven by exchange of resources.     

Precopulatory stabbing, hypodermic injections and unilateral copulations in a hermaphroditic sea slug

Reciprocity constitutes the prevalent mating mechanism among simultaneous hermaphrodites. Yet, when copulations in the female role confer fitness costs through male manipulation, it becomes advantageous sometimes to mate unilaterally in the male role only. In the sea slug Siphopteron quadrispinosum, acting males stab their partner with a bipartite penis, which not only hypodermically injects prostate fluids, but also apparently mechanically enforces unilateral male matings. Despite a pronounced male mating drive in both partners, unilaterality ensued when one slug stabbed more rapidly than its partner. The acting male may thus avoid the costs inflicted by traumatic injections and penial spines. While future studies need to elucidate the net fitness consequences of stabbing, our behavioural evidence is in line with the hypothesis that mating in S. quadrispinosum represents conflicting rather than complementary mating interests between mates.     

Sex allocation in a simultaneously hermaphroditic marine shrimp

Two fundamental questions dealing with simultaneous hermaphrodites are how resources are optimally allocated to the male and female function and what conditions determine shifts in optimal sex allocation with age or size. In this study, I explored multiple factors that theoretically affect fitness gain curves (that depict the relationship between sex-specific investment and fitness gains) to predict and test the overall and size-dependent sex allocation in a simultaneously hermaphroditic brooding shrimp with an early male phase. In Lysmata wurdemanni, sperm competition is absent as hermaphrodites reproducing in the female role invariably mated only once with a single other shrimp. Shrimps acting as females preferred small over large shrimps as male mating partners, male mating ability was greater for small compared to large hermaphrodites, and adolescent males were predominant in the population during the breeding season. In addition, brooding constraints were not severe and varied linearly with body size whereas the ability to acquire resources increased markedly with body size. Using sex allocation theory as a framework, the findings above permitted to infer the shape of the male and female fitness gain curves for the hermaphrodites. The absence of sperm competition and the almost unconstrained brooding capacity imply that both curves saturate, however the male curve levels off much more quickly than the female curve with increasing level of investment. In turn, the predominance of adolescent males in the population implies that the absolute gain of the female curve is greater than that of the male curve. Last, the size-dependent female preference and male mating ability of hermaphrodites determines that the absolute gain of the male curve is greater for small than for large hermaphrodites. Taking into consideration the inferred shape of the fitness gain curves, two predictions with respect to the optimal sex allocation were formulated. First, overall sex allocation should be female biased; it permits hermaphrodites to profit from the female function that provides a greater fitness return than the male function. Second, sex allocation should be size-dependent with smaller hermaphrodites allocating more than proportionally resources to male reproduction than larger ones. This size-dependent sex allocation permits hermaphrodites to profit from male mating opportunities that are the greatest at small body sizes. Size-dependent sex allocation is also expected because the male fitness gain curve decelerates more quickly than the female gain curve and experiments indicated that resources are greater for large than small hermaphrodites. These two predictions were tested when determining the sex allocation of hermaphrodites by dissecting their gonad and quantifying ovaries versus testes mass. Supporting the predictions above, hermaphrodites allocated, on average, 118 times more to the female than to the male gonad and the proportion of resources devoted to male function was higher in small than in large hermaphrodites. A trade-off between male and female allocation is assumed by theory but no negative correlation between male and female reproductive investment was observed. In L. wurdemanni, the relationship between sex-specific investment and fitness changes during ontogeny in a way that is consistent with an adjustment of sex allocation to improve size-specific reproductive success.     

Multiple paternity and mate competition in non-selfing,monogamous, egg-trading hermaphrodites

In animals in which the two sexes invest relatively similar amounts of resources in their young, the number of mates is expected to affect male and female reproductive success similarly and gender conflicts on the number of mates may not arise. Correspondingly, in non-selfing, simultaneous hermaphrodites with long-term monogamy, the two partners are expected to alternate repeatedly their sexual roles and invest similarly in their offspring. Therefore, the gender conflict on the number of mating partners should not arise. However, when >2 conspecifics are present, hermaphrodites are known to plastically adjust their behavior and sex allocation and compete for mating repeatedly in the male role. We tested whether this leads to multiple paternities of single egg clutches in experimental replicates of small and large groups of non-selfing, egg-trading, behaviorally monogamous polychaete worms (Ophryotrocha diadema) by using neutral genetic markers to estimate paternity. Multiply fertilized egg cocoons were common in these worms; two or more individuals succeeded in fertilizing the same egg cocoon and mate competition increased with group size. Multiply fertilized egg cocoons had a higher proportion of eggs developing into mature worms than singly fertilized egg cocoons. Possibly singly fertilized cocoons had a lower fertilization rate owing to low sperm counts and aflagellate sperm.     

Size-assortative mating in the earthworm Eisenia fetida (Oligochaeta, Lumbricidae)

In many species of simultaneous hermaphrodites, body size correlates with fecundity, and larger partners are preferred to small ones. Since sperm exchange is usually reciprocal, small individuals may be rejected by larger partners resulting in size-assortative mating. We studied the mating patterns in a natural population of the simultaneous hermaphroditic earthworm Eisenia fetida (Oligochaeta, Lumbricidae). We found that size-assortative mating processes existed, with variance in body weight within pairs lower than between pairs in mating earthworms. This non-random mating pattern probably reveals the existence of mate selection in this species, which lives at elevated densities with high availability of potential mates.     

Sperm transfer is affected by mating history in the simultaneously hermaphroditic snail Lymnaea stagnalis

Abstract. Males are predicted to strategically allocate sperm across mating partners in order to maximize their chances of paternity. This requires that males have the ability to detect aspects of their partner's mating history or the number of potential mates. We investigated whether simultaneous hermaphrodites mating in the male role strategically adjust sperm transfer depending on rearing conditions. The pond snail Lymnaea stagnalis (Basommatophora) is known to donate sperm repeatedly to different partners during a breeding season and store received sperm for >3 months. The rearing conditions of the donor as well as the recipient affect the amount of sperm transferred. Sperm donors raised in isolation transfer more sperm than those raised in groups. Furthermore, isolated sperm donors transfer less sperm to partners that were raised in groups than to those raised in isolation, i.e., virgins. These findings suggest that snails raised in isolation shift their sex allocation toward the male function and indicate that they can somehow assess the mating status of their partner.     

Sperm trading and sex roles in the hermaphroditic opisthobranch sea slug Navanax inermis : eager females or opportunistic males?

The sea slug Navanax inermis (Gastropoda, Opisthobranchia) is a well‐known example of a simultaneous hermaphrodite in which mating partners trade sperm. According to previous work by others, sperm trading follows from a general preference for the female role when the expected variance in reproductive success in the male role is higher and the risk of failure therefore larger. However, this view contradicts theoretical and empirical studies of other systems, which predict a general preference for the male role; sperm trading is assumed to follow from the fact that individuals benefit from sperm receipt as a nutritional compensation for sperm investment. In this study, we investigate the behaviour of N. inermis in more detail. In addition to observations of regular pairs, we also paired individuals with a partner that had been isolated for 33 days in order to induce changes in sex‐role preference in the non‐isolated partner. We also collected all clutches produced throughout the study to check for signs of infertility as a consequence of allosperm depletion. Fertility of field‐collected and isolated individuals suggested that sperm depletion occurs under natural conditions and may be caused by a lack of partners. Although this argues in favour of female preference, low mating rates both reduce variance in male reproductive success and remove the intent to use sperm for nutritional purposes, thus eliminating the conditions under which both hypotheses are intended to operate. The observational data indicate that animals are eager to mate as males, particularly at the beginning of a mating session. Intromissions lasted longer when a simultaneous intromission was received from the partner. Increases in intromission were recorded in non‐isolated individuals with partners which were previously isolated and therefore more attractive as females. This response would not have been expected were the female role the preferred one. A summary of the arguments concerning preference for either gender suggests some preference for the male role, but indicates that sexual preferences may actually change or become neutral within an individual, even in the course of a mating session. Overall, our results clearly confirm previously published observations of sperm trading in N. inermis . © 2003 The Linnean Society of London. Biological Journal of the Linnean Society, 2003, 78 , 105?116.     

Size-dependent sex allocation and sexual selection in Aplysia kurodai, a hermaphrodite with nonreciprocal mating

Abstract. For simultaneous hermaphrodites, a male-to-female shift in sex allocation with growth, and weak sexual selection on the male function, is predicted by many theories, although empirical data for both predictions are insufficient for internally fertilizing hermaphrodites with nonreciprocal mating. To address these issues, I studied mating and egg-laying behavior of the sea hare, Aplysia kurodai (Gastropoda: Opisthobranchia) in the laboratory. Both frequency and duration of egg laying increased with body weight, indicating that fecundity increased with weight. On the other hand, frequency and duration of mating as males did not increase with body weight, suggesting that sperm usage was independent of weight. Therefore, sex allocation shifted from male to female functions with growth. The lack of a relationship between body weight and mating activities as males also suggests that there was no "female" choice for large partners. However, the frequency of mating as females increased with body weight, suggesting "male" choice for large partners. This "male" choice is further supported by the presence of size-assortative mating and a longer duration of mating when the female partner was large. In addition, the variance in mating frequency as females was larger than that as males. As a whole, the mating behavior in A. kurodai can be summarized as choosy as males and unchoosy as females, the opposite of the patterns known in most gonochoric and hermaphroditic animals.     

The role of male age, sperm age and mating history on fecundity and fertilization success in the hide beetle

Models of age-related mate choice predict female preference for older males as they have proven survival ability. However, these models rarely address differences in sperm age and male mating history when evaluating the potential benefits to females from older partners. We used a novel experimental design to assess simultaneously the relative importance of these three parameters in the hide beetle, Dermestes maculatus. In a two-part experiment we first explored age-related male mating success and subsequently examined the consequences of male age, sperm age and male mating history on female fecundity and fertilization success. In a competitive mating environment, intermediate-age males gained significantly higher mating success than younger or older males. To test the consequences for females of aged-related male mating success, a second set of females were mated to males varying in age (young, intermediate-age and old), in numbers of matings and in timing of the most recent mating. We found that male age had a significant impact on female fecundity and fertilization success. Females mated to intermediate-age males laid more eggs and attained consistently higher levels of fertilization success than females with young and old mates. A male's previous mating history determined his current reproductive effort; virgin males spent longer in copula than males with prior mating opportunities. However, differences in copulation duration did not translate into increased fecundity or fertilization success. There was also little evidence to suggest that fertilization success was dependent on the age of a male's sperm. The experiment highlights the potential direct benefits accrued by females through mating with particular aged males. Such benefits are largely ignored by traditional viability models of age-related male mating success.     

Impaired sperm quality,delayed mating but no costs for offspring fitness in crickets winning a fight

The outcome of male–male contest competition is known to affect male mating success and is believed to confer fitness benefits to females through preference for dominant males. However, by mating with contest winners, females can incur significant costs spanning from decreased fecundity to negative effects on offspring. Hence, identifying costs and benefits of male dominance on female fitness is crucial to unravel the potential for a conflict of interests between the sexes. Here, we investigated males' pre‐ and post‐copulatory reproductive investment and its effect on female fitness after a single contest a using the field cricket Gryllus bimaculatus. We allowed males to fight and immediately measured their mating behaviour, sperm quality and offspring viability. We found that males experiencing a fight, independently of the outcome, delayed matings, but their courtship effort was not affected. However, winners produced sperm of lower quality (viability) compared to losers and to males that did not experience fighting. Results suggest a trade‐off in resource allocation between pre‐ and post‐mating episodes of sexual selection. Despite lower ejaculate quality, we found no fitness costs (fecundity and viability of offspring) for females mated to winners. Overall, our findings highlight the importance of considering fighting ability when assessing male reproductive success, as winners may be impaired in their competitiveness at a post‐mating level.     

Mate-search efficiency can determine the evolution of separate sexes and the stability of hermaphroditism in animals

Limited availability of mating partners has been proposed as an explanation for the occurrence of simultaneous hermaphroditism in animals with pair mating. When low population density or low mobility of a species limits the number of potential mates, simultaneous hermaphrodites may have a selective advantage because, first, they are able to adjust the allocation of resources between male and female functions in order to maximize fitness; second, in a hermaphroditic population the likelihood of meeting a partner is higher because all individuals are potential mates; and, third, in the absence of mating partners, many simultaneously hermaphroditic animals have the option of reproducing through self-fertilization. Recognizing that mate availability is central to the existing theory of hermaphroditism in animals, it is important to examine the effects of mate search on predictions of the stability of hermaphroditism. Many hermaphroditic animals can increase the number of potential mates they contact by active searching. However, since mate search has costs in terms of time and energy, the increased number of potential mates will be traded off against the amount of resources that can be allocated to the production of gametes. We explore the consequences of this trade-off to the evolution of mating strategies and to the selective advantage of self-fertilization. We show that in low and moderate population densities, poor mate-search efficiency and high costs of searching stabilize hermaphroditism and bias sex allocation toward female function. In addition, in very low population densities, there is strong selective advantage for self-fertilization, but this advantage decreases considerably in species with high mate-search efficiency. Most important, however, we present a novel evolutionary prediction: when mate search is efficient, disruptive frequency-dependent selection on time allocation to mate search leads to the evolution of searching and nonsearching phenotypes and, ultimately, to the evolution of males and females.     

Sperm trading in a hermaphroditic flatworm: reluctant fathers and sexy mothers

When matings are frequent and received sperm are digested, hermaphrodites should trade sperm when mating. We investigated sperm trading in the flatworm Schmidtea (Dugesia) polychroa and manipulated mating interests to investigate its possible causes. In 106 mating pairs consisting of nonisolated individuals, no sperm donation in either direction (35%) and reciprocal exchange (38%) were more common than expected by chance, whereas unilateral transfer (27%) was less frequent, confirming sperm trading. The amount of sperm donated depended on the availability of self-sperm, not on the amount received. Animals with more allosperm from previous matings had more self-sperm and consequently donated more. This suggests that sperm digestion boosts sperm production. In a second experiment, 'mixed-interest' pairs consisting of a nonisolated (N) and an isolated individual (I), NxI, were compared with IxI and NxN pairs. Whereas IxI pairs were eager and NxN reluctant to mate, NxI pairs showed an intermediate likelihood of mating. Whereas NxN pairs traded sperm, the other two groups did not. The change in behaviour in N individuals in the NxI treatment suggests precopulatory assessment and mating in relation to phenotypic mate quality. Isolated individuals are attractive, presumably because they donate large sperm clumps unconditionally and contain fewer allosperm, implying reduced sperm competition. The reduced reluctance in N individuals to mate with, and to inseminate, previously isolated partners suggests that female quality is an important factor in male sperm donation decisions. Hence, S. polychroa may be choosier than previously assumed. Copyright 2000 The Association for the Study of Animal Behaviour.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Determinants of female fecundity in a simultaneous hermaphrodite: the role of polyandry and food availability

Classical sexual selection theory assumes that the reproductive success of females is primarily limited by the resources available for egg production rather than by the number of mating partners. However, there is now accumulating evidence that multiple mating can entail fitness costs or benefits for females. In this study we investigated the effect of polyandry (i.e., the mating with different mating partners) and food availability on the reproductive output of the female sex function in an outcrossing simultaneous hermaphrodite, the free-living flatworm Macrostomum lignano. We exposed virgin worms to different group sizes, a treatment that has previously been shown to affect the level of polyandry in this species. Moreover, we manipulated the food availability throughout the subsequent egg laying period, during which the worms were kept in isolation. The number of offspring produced was used as an estimate of female fecundity. We found that food availability, but not group size, had a significant effect on female fecundity. Additionally, female fecundity was positively correlated with the number of stored sperm in the female sperm-storage organ at the time of isolation, but it was not correlated with body or ovary size of the worms. Our results suggest that female fecundity in M. lignano is primarily determined by the resources available for egg production, and not by the level of polyandry, confirming classic sexual selection theory for simultaneous hermaphrodites.     

Reproductive behavior,egg trading,and correlates of male mating success in the simultaneous hermaphrodite,Serranus tabacarius

Synopsis Serranus tabacarius (Serranidae), the tobaccofish, is a simultaneous hermaphrodite which belongs to a group of seabasses that exhibit a wide variety of social and mating systems. The reproductive behavior of tobaccofish is similar to other hermaphroditic seabasses, with individuals assuming sex-specific spawning behaviors that allow for the assignment of male and female roles in a mating sequence. Virtually all matings involved pairs of individuals, although streaking, an alternative male mating tactic, was observed once. Pairs engage in egg trading, where individuals divide their daily clutch into a series of sequentially released parcels and take turns releasing eggs for their partner to fertilize. Individuals mate over a late afternoon spawning period with a number of partners sequentially. Larger individuals have both more total matings and more spawning partners. Egg trading is not symmetrical, the number of male and female matings for an individual in a spawning sequence is often unequal. Overall, the ratio of male to female matings increases with individual size. Large individuals are socially dominant, chase conspecifics during the reproductive period, and are more likely to end a spawning bout with a partner immediately after mating in the male role. In addition, larger individuals are less likely to reciprocate female matings by a partner, either by only mating once (as a male) in a spawning bout or by mating consecutively as a male within a series of matings. Although larger individuals show this relative specialization in the male role, they maintain their simultaneous hermaphroditism and obtain a substantial percentage of their mating success through female function. Egg trading appears to reduce the opportunity for large individuals to specialize as pure males, and thus interacts with the environmental potential for polygamy in shaping the mating system and sex allocation pattern in this species.     

Costs of receipt and donation of ejaculates in a simultaneous hermaphrodite

Background  

Sexual conflicts between mating partners can strongly impact the evolutionary trajectories of species. This impact is determined by the balance between the costs and benefits of mating. However, due to sex-specific costs it is unclear how costs compare between males and females. Simultaneous hermaphrodites offer a unique opportunity to determine such costs, since both genders are expressed concurrently. By limiting copulation of focal individuals in pairs of pond snails (Lymnaea stagnalis) to either the male role or the female role, we were able to compare the fecundity of single sex individuals with paired hermaphrodites and non-copulants. Additionally, we examined the investment in sperm and seminal fluid of donors towards feminized snails and hermaphrodites.     

The Effect of Female Quality on Male Ejaculatory Expenditure and Reproductive Success in a Praying Mantid

Strategic ejaculation is a behavioural strategy shown by many animals as a response to sperm competition and/or as a potential mechanism of cryptic male choice. Males invest more mating resources when the risk of sperm competition increases or they invest more in high quality females to maximize their reproductive output. We tested this hypothesis in the false garden mantid Pseudomantis albofimbriata, where females are capable of multiply mating and body condition is an indicator of potential reproductive fitness. We predicted male mantids would ejaculate strategically by allocating more sperm to high quality females. To determine if and how males alter their ejaculate in response to mate quality, we manipulated female food quantity so that females were either in good condition with many eggs (i.e. high quality) or poor condition with few eggs (i.e. low quality). Half of the females from each treatment were used in mating trials in which transferred sperm was counted before fertilisation occurred and the other half of females were used in mating trials where fertilisation occurred and ootheca mass and total eggs in the ootheca were recorded. Opposed to our predictions, the total number of sperm and the proportion of viable sperm transferred did not vary significantly between female treatments. Male reproductive success was entirely dependent on female quality/fecundity, rather than on the number of sperm transferred. These results suggest that female quality is not a major factor influencing postcopulatory male mating strategies in P. albofimbriata, and that sperm number has little effect on male reproductive success in a single mating scenario.     

Females of the Cellar Spider Discriminate Against Previous Mates

Mate choice for novel partners should evolve when remating with males of varying genetic quality provides females with fitness‐enhancing benefits. We investigated sequential mate choice for same or novel mating partners in females of the cellar spider Pholcus phalangioides (Pholcidae) to understand what drives female remating in this system. Pholcus phalangioides females are moderately polyandrous and show reluctance to remating, but double‐mated females benefit from a higher oviposition probability compared to single‐mated females. We exposed mated females to either their former (same male) or a novel mating partner and assessed mating success together with courtship and copulatory behaviours in both sexes. We found clear evidence for mate discrimination: females experienced three‐fold higher remating probabilities with novel males, being more often aggressive towards former males and accepting novel males faster in the second than in the first mating trial. The preference for novel males suggests that remating is driven by benefits derived from multiple partners. The low remating rates and the strong last male sperm precedence in this system suggest that mating with novel partners that represent alternative genotypes may be a means for selecting against a former mate of lower quality.     

Differences between the sexes in direction and duration of response to seeing a potential sex partner mate with another

We have shown previously that: (1) female Japanese quail, Coturnix japonica, increase and male Japanese quail decrease their tendencies to affiliate with potential sex partners after seeing them mate, and (2) in both sexes of quail, affiliative preferences and choice of a sex partner are highly correlated. Here we predict that because effects of a prior male's sperm on a second male's probability of fertilizing a female are relatively brief, a male's avoidance of whichever member of a pair of females he has seen mating should be transitory. Conversely, because female quail seek high-quality males as mates and quality is a relatively permanent characteristic, females' preferences between males should remain constant over time. We found, as predicted, that the durations of effects on affiliation of seeing a potential sex partner mate differed in male and female quail. Forty-eight hours after male quail saw a female mate, they no longer avoided her, whereas 48 h after female quail saw a male mate, his attractiveness remained enhanced. We conclude by suggesting that both the direction and the duration of responses of male and female Japanese quail to seeing a member of the other sex mating enhance the fitness of members of each sex. Copyright 2000 The Association for the Study of Animal Behaviour.     

The evolution of repeated mating under sexual conflict

In insects, repeated mating by females may have direct effects on female fecundity, fertility, and longevity. In addition, a female's remating rate affects her fitness through mortality costs of male harassment and ecological risks of mating such as predation. We analyse a model where these female fitness factors are put into their life-history context, and traded against each other, while accounting for limitations because of mate availability. We solve analytically for the condition when female multiple mating will evolve. We show that the probability that a female mates with a courting male decreases with increases in population density. The extent of conflict between the sexes thus automatically becomes larger at higher densities. However, because at higher densities females meet males at a higher rate, the resulting ESS female remating rate is independent of population density. The female remating probability is in conflict with male adaptations that increase male mating rate by persuading or forcing females to mate, and also in conflict with male adaptations for protecting the own sperm from being removed by future female mates. We show that the relative importance of these conflicts depends on population density.