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.     

Mating effort and female receptivity: how do male guppies decide when to invest in sex?

Males vary in the degree to which they invest in mating. Several factors can explain this variation, including differences in males’ individual condition and the fact that males allocate their energy depending on the context they face in each mating attempt. Particularly, female quality affects male reproductive success. Here, we studied whether male guppies (Poecilia reticulata) strategically allocated more mating effort, in terms of mating behaviour and male–male competition, when they were matched with a receptive (R) female than a non-receptive one. In accordance with our prediction, we found that males increased their mating behaviour when they were with a receptive female. Even though male guppies can inseminate non-receptive females, we only found high levels of courtship between males that were with a receptive female rather than a non-receptive one. Although there was little affect of female receptivity on male–male competition, we found that males chased and interrupted courtships more with receptive females than with non-receptive females regardless of odour. Finally, we also studied whether the sexual pheromone produced by receptive female guppies is a cue that males use in order to increase their mating effort. We found that males were more attracted to a female when they perceived the sexual pheromone, but only increased their mating and aggressive behaviours when females showed receptive behaviour. This strategic increase in mating effort could result in higher male reproductive success because mating attempts towards receptive females are likely to be less costly and males could have a greater probability of fertilisation.     

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.     

Selected polyandry: female choice and inter-sexual conflict in a small nocturnal solitary primate (<Emphasis Type="Italic">Microcebus murinus</Emphasis>)

Sex-specific interests over the maximization of reproductive success lead to an inter-sexual conflict over the optimal mating system in a species. Traditionally, the outcome of this inter-sexual conflict has been studied from the male perspective but it also depends on female mating strategies, such as manipulating the temporal distribution of sexual activity, advertisement, and mate choice. We used a small nocturnal primate, the gray mouse lemur (Microcebus murinus) to determine the relative importance of female mating strategies on the outcome of this conflict in a species where females are solitary during their activity period. We studied their mating behavior over three consecutive annual mating seasons and determined the genetic relationships among more than 300 study animals to quantify individual reproductive success. We found that most females were receptive asynchronously. Females did not exhibit any obvious direct mate choice, probably due to a highly male-biased operational sex ratio and the corresponding costs of choosiness. However, females exercised indirect choice for multiple matings. They mated with 1–7 males up to 11 times during their single night of receptivity. As a result, mixed paternity was common but heavier males sired more offspring, meaning that indirect female choice for superior males cannot be excluded. Females exhibited a mixed mating strategy, avoiding costly direct mate choice but still counteracting male efforts to monopolize mating, successfully increasing genetic variability among offspring. Thus, females had a major influence on the outcome of the inter-sexual conflict despite male monopolization attempts.Communicated by J. Setchell     

Male mate choice favors more colorful females in the gift-giving cabbage butterfly

While the phenomenon of male mate choice has attracted considerable attention in the last two decades, whether this sexual selection mechanism could drive the evolution of female ornaments remains poorly understood. Here, we used experimental manipulation of female wing coloration to investigate male mate choice in Pieris rapae, a gift-giving butterfly. Further, we tested whether males’ nutritional status influenced their mating preferences by subjecting larvae to short periods of starvation. We found that males showed significantly more mating approaches toward control females with more colorful wings (higher pteridine content), and that this preference was strongest in low-nutrition males. Additionally, a study of field-collected females revealed that pteridine-based wing coloration was positively correlated with female egg load, which suggests such ornaments may signal female quality. Pteridine-based ornaments are widespread in nature, however their potential as honest signals in male mate choice remains largely unexplored. This work furthers our understanding of how male mate choice and female ornamentation may evolve in species whose mating systems include nutritional nuptial gifts.     

Male spotted bowerbirds preferentially choose,arrange and proffer objects that are good predictors of mating success

Females from a wide range of species have been shown to prefer males producing exaggerated, elaborate sexual displays. The question of whether males can adapt their behaviour in order to accentuate these preferred aspects has received less attention. Male bowerbirds provide an excellent system in which to address this question. Males build and decorate structures--bowers--which females use to assess males. Males exhibiting high numbers of particular decorations on their bowers receive increased mating success. Males can choose what objects to place on bowers and where to put them. Therefore, their behaviour can directly influence their mating success. I studied a population of spotted bowerbirds Chlamydera maculata in central Queensland, Australia. I show, using two sets of observations and a choice experiment, that males exhibit strong and consistent preferences for certain objects. The male's preferences have an adaptive value, targeting objects that predict his mating success. Such behaviour operates at three levels. Fundamentally, object selection and acquisition is biased in favour of certain objects for use as decorations. These decorations are then placed prominently on the bower, within the avenue. Finally, males actively proffer influential decorations, used as props, to visiting females, ensuring that crucial components of the sexual display cannot be ignored.     

Polyandrous mating in treetops: how male competition and female choice interact to determine an unusual carnivore mating system

The diversity of mammalian mating systems is primarily shaped by sex-specific reproductive strategies. In the present study, we explored determinants and consequences of a unique mating system exhibited by fossas (Cryptoprocta ferox), the largest Malagasy carnivore, where females mate polyandrously on traditional mating trees, and males exhibit intrasexual size dimorphism. Males face both contest and scramble competition, and inter-sexual size dimorphism can be pronounced, but its magnitude depends on the male morph. Using a continuous behavioral observation of six estrous females over 4 years, we investigated correlates of male contest competition and female choice based on 316 copulations. Furthermore, we assessed correlates of male scramble competition based on testes size and movement data obtained from GPS tracking. We found that females dominated males regardless of their smaller size and that females actively solicited copulations. Heavy males had highest mating success during the female’s peak mating activity, but were discriminated against afterwards. Female choice and male–male competition thus converged to generate a mating advantage for heavier males. Our results suggest that females actively seek polyandrous matings, presumably for indirect genetic benefits. Since body mass is the major determinant of male mating success and is at the same time dependent on the degree of sociality and associated hunting mode of the respective male morph, a male’s feeding ecology is likely to influence its reproductive tactic. A combination of benefits from female polyandry and the consequences of different subsistence strategies may thus ultimately explain this unusual mating system.     

The significance of hotspots to lekking topi antelopes (<Emphasis Type="Italic">Damaliscus lunatus</Emphasis>)

Controversy has surrounded the question of why lek-breeding has evolved in certain ungulate species. Can the behavior be explained simply by males mapping onto a female distribution that is determined by factors unrelated to mating? Or are leks created because estrous females distinguish between males and favor males who cluster? Here I address these questions by looking at spatial distribution in lekking topi antelopes (Damaliscus lunatus). Contrary to the predictions of a model assuming male clustering in the zone of maximum female range overlap, territories were highly clustered also within this zone, and lek size correlated positively with population density. In support of models derived from the ideal free distribution of males onto female dispersion, leks were in areas with high female density during the rut. However, models not taking into account both individual variation in male quality and female mate preferences failed to explain the extreme male clumping also within high density areas, which was revealed by a strongly male-biased sex-ratio on leks. Additional support for the female preference-based model came from the finding that estrous females concentrated onto leks. Female preference for clustered males may develop if males initially follow an ideal free distribution of unequal competitors with high quality males slightly clustered at density hotspots; positive feedback between female benefits of preference for clustered males and male benefits of clustering could lead to contraction of the territorial network and lek behavior. Thus only the female preference-based model correctly predicted a negative correlation between male mating rate and resource density.     

Strategic male signalling effort in a desert-dwelling fish

Males often use elaborate courtship displays to attract females for mating. Much attention, in this regard, has been focused on trying to understand the causes and consequences of signal variation among males. Far less, by contrast, is known about within-individual variation in signal expression and, in particular, the extent to which males may be able to strategically adjust their signalling output to try to maximise their reproductive returns. Here, we experimentally investigated male courtship effort in a fish, the Australian desert goby, Chlamydogobius eremius. When offered a simultaneous choice between a large and a small female, male gobies spent significantly more time associating with, and courting, the former, probably because larger females are also more fecund. Male signalling patterns were also investigated under a sequential choice scenario, with females presented one at a time. When first offered a female, male courtship was not affected by female size. However, males adjusted their courtship effort towards a second female depending on the size of the female encountered previously. In particular, males that were first offered a large female significantly reduced their courtship effort when presented with a subsequent, smaller, female. Our findings suggest that males may be able to respond adaptively to differences in female quality, and strategically adjust their signalling effort accordingly.     

Male moths undertake both pre- and in-copulation mate choice based on female age and weight

Variation in reproductive potential usually occurs among individuals of both sexes; for example, some individuals may carry more gametes or be able to continue to generate more gametes in their reproductive life than others. Therefore, to maximize their reproductive success, both sexes are expected to show adaptations for mate choice. However, most authors concentrated on how females choose their mates with the belief that females invest more in reproduction than males, and males are generally eager to pair with any female. In this paper, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), with special emphasis on male pre- and in-copulation mate choice in relation to the reproductive quality and re-copulation potential of females. We show that in E. kuehniella, younger and heavier females have significantly higher reproductive value than older and lighter ones, and male sperm supply significantly decreases over successive copulations. Males exercise pre-copulation mate choice by selecting females with higher reproductive potential for copulation and in-copulation mate choice by allocating more sperm to females with higher reproductive potential. However, high-quality females are more likely to re-copulate than low-quality ones, and allocation of more sperm to females by males does not increase female reproductive outputs. It is suggested that the allocation of more sperm to high-quality females functions to increase sperm competitiveness against rivals.     

Allocation of nuptial gifts in tree crickets changes with both male and female diet

Male tree crickets, Oecanthus nigricornis, offer a nuptial gift to females during mating in the form of a secretion from a dorsal metanotal gland. I examined the effects of male and female nutrient limitation on allocation of the gift. Males were fed ¹⁴C radiolabeled amino acids, placed onto high- or low-quality diets and then mated with females also maintained on high- or low-quality diets. Female acquisition of radiolabel increased, and residual radiolabel within the metanotal gland decreased, over time verifying that gift size correlates with duration of courtship feeding. High-diet females produced greater egg numbers and allocated proportionally more of the male-derived amino acids to their ovaries. As predicted, duration of courtship feeding was greater for males on high-quality diets and females on low-quality diets. Amino acid transfer, measured as the proportion of total radiolabel transferred to the female, showed a significant interaction between male and female diet. High-diet females acquired available radiolabel more rapidly from high-diet males, but low-diet females acquired available radiolabel more rapidly from low-diet males. The causes of these differences between feeding duration and nutrient allocation are discussed.     

Audience effect alters male but not female mating preferences

Males often face strong mating competition by neighboring males in their social environment. A recent study by Plath et al. (Anim Behav 75:21–29, 2008a) has demonstrated that the visual presence of a male competitor (i.e., an audience male) affects the expression of male mating preferences in a poeciliid fish (Poecilia mexicana) with a weaker expression of mating preferences when an audience male observed the focal male. This may be a tactic to reduce sperm competition, since surrounding males likely share intrinsic preferences for female traits or copy mate choice decisions. Here, we examined the hypothesis that a same-sex audience would affect female mate preferences less than male mating preferences. Our hypothesis was based on the assumptions that (1) competition for mates in a fashion that would be comparable in strength to sperm competition or overt male–male aggression is absent among Poecilia females, and (2) P. mexicana females typically form female-biased shoals, such that almost any female mate choice in nature occurs in front of a female audience. Poecilia females (P. mexicana, surface and cave form, and the closely related gynogenetic Poecilia formosa) were given a choice between a large and a small male, and the tests were repeated while a conspecific, a heterospecific, or no audience female (control) was presented. Females spent more time in the neutral zone and, thus, less time near the males during the second part of a trial when an audience was presented, but—consistent with predictions—females showed only slightly weaker expression of mate preferences during the second part of the tests. This decline was not specific to the treatment involving an audience and was significantly weaker than the effect seen in the male sex.     

Body size preferences in the pot-bellied seahorse <Emphasis Type="Italic">Hippocampus abdominalis:</Emphasis> choosy males and indiscriminate females

Male seahorses (genus Hippocampus) provide all post-fertilization parental care, yet despite high levels of paternal investment, these species have long been thought to have conventional sex roles, with female mate choice and male–male competition. Recent studies of the pot-bellied seahorse (Hippocampus abdominalis) have shown that sex-role reversal occurs in high-density female-biased populations, indicating that male mating preferences may lead to sexual selection on females in this species. Egg size, egg number, and offspring size all correlate positively with female body size in Hippocampus, and by choosing large mating partners, male seahorses may increase their reproductive success. While male brood size is also positively correlated with body size, small H. abdominalis males can carry exceptionally large broods, suggesting that the fecundity benefits of female preference for large partners may be limited. We investigated the importance of body size in reproductive decisions of H. abdominalis, presenting focal individuals of both sexes with potential mating partners of different sizes. Mating preferences were quantified in terms of time spent courting each potential partner. Male seahorses were highly active throughout the mate-choice trials and showed a clear behavioral preference for large partners, while females showed significantly lower levels of activity and equivocal mating preferences. The strong male preferences for large females demonstrated here suggest that sexual selection may act strongly on female body size in wild populations of H. abdominalis, consistent with predictions on the importance of female body size for reproductive output in this species. An erratum to this article can be found at     

Providing parental care entails variable mating opportunity costs for male Temminck’s stints

Males of many species theoretically face a fitness tradeoff between mating and parental effort, but quantification of this is rare. We estimated the magnitude of the mating opportunity cost paid by incubating male Temminck’s stints (Calidris temminckii), taking advantage of uniparental care provided by both sexes in this species. “Incubating males” provide all care for an early clutch, limiting subsequent mating possibilities. “Non-incubating” males include males that failed to obtain, lost to predation, or actively avoided incubating clutches. These males were free to pursue extrapair copulations and to mate with females laying later clutches, which females usually incubate themselves. Male incubation classes did not differ in measures of quality, and many individuals changed classes between years, suggesting the use of conditional reproductive tactics. However, specialist non-incubators may also exist. Using microsatellites to assign parentage, we compare males’ total fertilizations and the subset “free of care” fertilizations between incubation classes. Incubators were more likely to gain at least one fertilization per season and averaged one more per season than non-incubators. However, successful non-incubators were more likely to gain “free of care” fertilizations, averaging two more than successful incubators. The relative success of male incubation classes also changed with local sex ratios. With higher female proportions, non-incubators gained disproportionately more offspring, suggesting that the use of tactics should be partly determined by the availability of potentially incubating females. Overall, we estimate the opportunity cost of incubating to be 13–25 % of the potential annual reproductive output.     

Multiple sexual pigments,assortative social pairing,and genetic paternity in the yellow warbler (Setophaga petechia)

In socially monogamous species, extra-pair paternity may increase the reproductive success of highly ornamented males, mediating the evolution of sexual ornaments. However, ornaments may also attract social mates, and a tradeoff between extra-pair paternity (EPP) and within-pair paternity (WPP) may complicate mating strategies. Further, in many socially monogamous species, females are also ornamented, and the relationship between female ornamentation and patterns of EPP has been neglected. We investigated the patterns of genetic paternity with respect to carotenoid- and melanin-based pigmentation in yellow warblers (Setophaga petechia) of both sexes. We asked whether males face tradeoffs between EPP and WPP, how paternity patterns relate to carotenoid- versus melanin-based pigmentation, and whether less EPP occurs in broods when males and females are assortatively paired. Males faced a tradeoff between EPP and WPP. Moreover, non-additive relationships existed between paternity patterns and the two pigment types in both sexes. Males with high melanin coverage but dull carotenoid pigmentation achieved EPP but lost WPP, whereas males with high levels of both pigment types had high WPP but gained little EPP. A parallel pattern occurred in females. Warblers paired assortatively by pigmentation and EPP was less common in broods when the sexes were assortatively paired by carotenoid pigmentation. Results suggest that the most colorful birds obtain high quality social mates and advance reproductive success through WPP, show that correlations can arise between female ornamentation and patterns of EPP, and also uniquely suggest that social pairing patterns may influence extra-pair mating strategies.     

Mate compatibility,parental allocation and fitness consequences of mate choice in the sand goby <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Recent theory and empirical work suggests that there may be variation among females in mate preferences that is adaptive. One of the possible mechanisms maintaining variability in preferences and preferred traits is that the benefits of mate choice may depend on compatibility with potential mating partners. We examined fitness consequences of mate choice in a species of fish, the sand goby, Pomatoschistus minutus with a special focus on mate compatibility. Females were given the opportunity to establish their mate preferences in a dichotomous mate choice experiment. This information was then applied by mating the focal or control female with either the preferred or the non-preferred male. The parental performance of the males of these four mating combinations was then measured. In a separate experiment, we assessed the female differential allocation by determining the residual gonad weight of spawned females as a measure of the proportion of eggs spawned. We also estimated the amount of filial cannibalism separately for both sexes. Our results show that preferred males provided benefits in the form of an increased number of hatching eggs. This benefit was the same when the male was mated with a focal or a control female. Hence, we found no support for benefits that depend on mate compatibility. Neither did we find support for the hypothesis that females would lay a different number of eggs depending on the male status. The results also indicate that male filial cannibalism has a strong role in determining hatching success in this species.     

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.     

Female habitat choice as a determinant of the reproductive success of the territorial male marine iguana (Amblyrhynchus cristatus)

Summary The reproductive behaviour of marine iguanas (Amblyrhynchus cristatus) was studied to answer the following questions: Which physical factors determine the number of females found on a male's territory? Do males select territories according to preferences for physical conditions or according to the number of females there? What causes the strong correlation between the number of females on a territory and the number of copulations achieved there? Females preferred as their resting places territories with the following characteristics, in order of importance: High number of shady/shelter places, low temperature of rocks at noon, proximity to the sea, sun-basking places in the afternoon. Other factors measured (probability of being wetted by spray, sun-basking places in the morning and at noon) showed no significant correlation with the number of females. Males, however, preferred territories further from the sea (within about 7 m), with high rock temperatures and a high number of basking places in the morning (in order of importance). Males primarily followed their site preferences and not female density in their choice of a territory. The site choice of females determined the number of copulations achieved on a territory, as females copulated preferentially on their resting places. A shift of females from one territory to another within one colony between two succeeding years was correlated with the change in relative weight of territory owners. This result was interpreted as an indication of female choice. Site preferences of males and females are discussed in relation to the animals' physiological requirements. Possible influences of ecological factors on the evolution of this mating system are considered.     

Mating patterns of woolly spider monkeys,Brachyteles arachnoides: implications for female choice

Summary Woolly spider monkeys show a promiscuous, polygynous mating system in which a receptive female mates with several males, often in rapid succession. Copulation is prolonged with an average duration of 4.1±1.5 (SD) min. Coitus consists of a stationary phase with the male in intromission, followed by stercotypic behaviours of the female which appear to cue and/or accompany male climax and ejaculation. Subadult and adult males show different association patterns with individual females. Subadult males form long-term consortships with particular females while adult males appear strongly attracted to a particular female only when the is receptive. These different behavior patterns of males are viewed as age-specific mating tactics. Males in a mating aggregation show little intermale aggression for sexual access to a receptive female. Large testis size in this species suggests that much intermale competition for reproductive success may be carried out at a postcopulatory level, perhaps by sperm competition. The copulatory pattern of woolly spider monkeys may function primarily as a mechanism of female choice, aiding a female in assessing the quality of males in hei mating aggregation while helping to ensure that maximal high quality spermatozoa will be available at the proper time for fertilization.     

Repeatable female preferences, mating order and mating success in the poeciliid fish, Heterandria formosa

Sexual selection via female choice can afford preferred males comparably higher mating success than those males that lack preferred traits. In addition, many models of sexual selection assume that both male traits and female preferences are heritable. In this study we test whether females of the poeciliid fish, Heterandria formosa, have repeatable pre-copulatory preferences for larger males. We also test whether female pre-copulatory preferences are always reliable indicators of male mating success. When given a choice between a large and a small male, females prefer larger males, and the repeatability of this preference is high. Although there are no overall differences in male mating success between large and small males, large males have a higher mating success when they are the first to mate than when they are the second to mate. Likewise, preferred males also have higher mating success when they are the first to mate than when they mate second. Therefore, the repeatable female preferences observed in this study only predict male mating success when the preferred male mates first. These results illustrate that even significantly repeatable female preferences do not translate into male mating success, which is an assumption of many examinations of the importance of female choice in sexual selection.