Male sexual attractiveness predicts differential ovulatory shifts in female extra-pair attraction and male mate retention

Because ancestral women faced trade-offs in choosing mates, they may have evolved to pursue a dual-mating strategy in which they secured investment through one partner and obtained good genes through others. The dual-mating theory predicts that women will display greater interest in extra-pair sex near ovulation, especially if they are mated to a primary male partner who is low in sexual attractiveness. Forty-three normally ovulating women rated their partner's sexual attractiveness and separately reported their own desires and their partner's mate retention behaviors at high and low fertility (confirmed using luteinizing hormone tests). In the high-fertility session relative to the low, women who assessed their partners as being lower in sexual attractiveness reported greater extra-pair desires and more expressed love and attention from their male partners. Women's desire for their own partners did not differ significantly between high and low-fertility sessions.     

Ovulatory Shifts in Women��s Attractions to Primary Partners and Other Men: Further Evidence of the Importance of Primary Partner Sexual Attractiveness

Previous research has documented shifts in women’s attractions to their romantic partner and to men other than their partner across the ovulation cycle, contingent on the degree to which her partner displays hypothesized indicators of high-fitness genes. The current study set out to replicate and extend this finding. Forty-one couples in which the woman was naturally cycling participated. Female partners reported their feelings of in-pair attraction and extra-pair attraction on two occasions, once on a low-fertility day of the cycle and once on a high-fertility day of the cycle just prior to ovulation. Ovulation was confirmed using luteinizing hormone tests. We collected two measures of male partner sexual attractiveness. First, the women in the study rated their partner’s sexual attractiveness. Second, we photographed the partners and had the photos independently rated for attractiveness. Shifts in women’s in-pair attractions across the cycle were significantly moderated by women’s ratings of partner sexual attractiveness, such that the less sexually attractive women rated their partner, the less in-pair attraction they reported at high fertility compared with low fertility (partial r = .37, p _dir = .01). Shifts in women’s extra-pair attractions across the cycle were significantly moderated by third-party ratings of partner attractiveness, such that the less attractive the partner was, the more extra-pair attraction women reported at high relative to low fertility (partial r = −.33, p _dir = .03). In line with previous findings, we found support for the hypothesis that the degree to which a woman’s romantic partner displays indicators of high-fitness genes affects women’s attractions to their own partner and other men at high fertility.     

Hormonal influences on women's extra-pair sexual interests: The moderating impact of partner attractiveness

Based on the idea that women are especially attracted to ancestral markers of male genetic quality when conceptive in their cycle, scholars have conjectured that increases in women's extra-pair sexual interests during the conceptive phase of the cycle are moderated by their primary partners' sexual attractiveness. Multiple studies have examined this prediction, with largely supportive but mixed results. The current study is the first to examine whether hormonal influences—thought to mediate cycle shifts—on women's extra-pair sexual interests are moderated by partner attractiveness. 213 naturally cycling, romantically involved women (181 attending multiple sessions) participated in up to four sessions over about a month. Estrogen and progesterone levels were measured multiple times across women's cycles. Male partner attractiveness moderated associations between progesterone levels (though not estrogen levels) and women's extra-pair sexual interests. A negative association between progesterone levels and extra-pair sexual interests, a composite measure, was observed for women with relatively unattractive partners. For women with relatively attractive partners, no clear association emerged. The interaction between progesterone and partner attractiveness was robust for women's interest in extra-pair sex, a component preregistered as exploratory. The interaction effect was also significant for the absolute intensity of women's extra-pair attraction (a component item referenced in the preregistration) but was non-significant for the frequency of women's extra-pair attraction relative to typical days (a composite component specified in the preregistration). These findings inform theoretical understandings of women's sexuality.     

Factors influencing mate guarding and territory defence in the stitchbird (hihi)

Socially monogamous male birds are predicted to maximise their reproductive success by pursuing extra-pair copulations (EPCs) while engaging in anti-cuckoldry behaviour such as mate guarding. In the stitchbird, Notiomystis cincta, high levels of forced EPCs and a high proportion of nestlings resulting from extra- pair fertilisations lead to the prediction that males of this species should exhibit intense paternity guarding behaviours. While studying an isolated stitchbird population on Tiritiri Matangi Island New Zealand (3636'S, 17453'E), I collected daily behavioural data throughout the breeding season from 15 males in 2000/01 and 27 males in 2001/02. In this study, male stitchbirds demonstrated clear paternity guarding by exhibiting: (1) an increased likelihood of being close to their mate during her fertile period, (2) an increased initiation of mate contact during her fertile period, (3) switching from site-specific territorial defence during the pre-fertile period to defending an area centring on the their female partners location during her fertile period, and (4) an increased following of the female to communal feeding sites outside the territory during her fertile period. For polygynous males, mate guarding and territorial defence were conditional on which of their females was fertile. Additional evidence supporting the hypothesis that mate guarding in this species is a form of paternity assurance, rather than protection from harassment, is that males protected their partner from harassment by other stitchbird males but did not intervene when females were harassed by male bellbirds, Anthornis melanura. While mate-guarding intensity in many species is conditional on the stage of female fertility, male stitchbirds also modified their behaviour depending on the location of the female and the rate of intrusions by extra-pair males. Resident males adopted a best-of-a-bad-job tactic when they were unable to locate their female by defending an area around her last known location. Furthermore, when the rate of intrusions by extra-pair males increased they traded-off the area they could defend within their territory against their ability to guard the female. Territory takeovers were uncommon, but when they did occur older males displaced younger males and healthy birds displaced sick ones. Contrary to the prevailing view that mate guarding is a male response to female infidelity, male stitchbirds appear to use mate guarding primarily to prevent paternity losses from forced EPCs. Future assessments of mate guarding function should consider the possibility that mate guarding involves a combination of conflict and co-operation between the sexes.     

Changes in women's sexual interests and their partners' mate-retention tactics across the menstrual cycle: evidence for shifting conflicts of interest

Because ancestral women could have obtained genetic benefits through extra-pair sex only near ovulation, but paid costs of extra-pair sex throughout the cycle, one might expect selection to have shaped female interest in partners, other than primary partners, to be greater near ovulation than during the luteal phase. Because men would have paid heavier costs if their partners had extra-pair sex near ovulation, one might also expect selection to have shaped males' efforts to track their primary partners' whereabouts to be increased near ovulation, relative to the luteal phase. Women filled out questionnaires about their sexual interests and their partners' mate-retention tactics twice: once within 5 days before a lutenizing hormone surge and once during the luteal phase. Results showed that: (i) women reported greater sexual interest in, and fantasy about, non-primary partners near ovulation than during the luteal phase; (ii) women did not report significantly greater sexual interest in, and fantasy about, primary partners near ovulation; (iii) women reported that their primary partners were both more attentive and more proprietary near ovulation.     

Adaptive mating strategies and the problem of mate retention

"Adaptations" are evolved solutions to the problems posed by survival and reproduction. The evolutionary psychologists believe that as well as the physical adaptations so the adaptations in human mind evolved, called "strategies". The "mating strategies" are adaptive solutions to successful mating. The mating strategies, designed to preserve access to a mate by preventing encroachment of intrasexual rivals and by preventing a mate from defecting from the mateship for a prospective better partner, are called "mate guarding strategies". The previous research found that humans do use a wide variety of behavioural tactics of mate guarding, ranging from vigilance to violence. Our research group explores the type and the intensity of behavioural tactics of mate guarding used in several contexts. Presently, the link between the woman's fertility status across her menstrual cycle and the man's mate guarding is examined. Discussing the preliminary results, a more intensive man's mate guarding of his partner around the ovulation when her fertility peaks may be assumed. These outcomes could be explained as an adaptive prevention to shift in woman's preferences to increase her extra-pair sexual attempts and following to a possible genetic cuckoldry at that most fertile time.     

When mothers make sons sexy: maternal effects contribute to the increased sexual attractiveness of extra-pair offspring

Quality differences between offspring sired by the social and by an extra-pair partner are usually assumed to have a genetic basis, reflecting genetic benefits of female extra-pair mate choice. In the zebra finch (Taeniopygia guttata), we identified a colour ornament that is under sexual selection and appears to have a heritable basis. Hence, by engaging in extra-pair copulations with highly ornamented males, females could, in theory, obtain genes for increased offspring attractiveness. Indeed, sons sired by extra-pair partners had larger ornaments, seemingly supporting the genetic benefit hypothesis. Yet, when comparing ornament size of the social and extra-pair partners, there was no difference. Hence, the observed differences most likely had an environmental basis, mediated, for example, via differential maternal investment of resources into the eggs fertilized by extra-pair and social partners. Such maternal effects may (at least partly) be mediated by egg size, which we found to be associated with mean ornament expression in sons. Our results are consistent with the idea that maternal effects can shape sexual selection by altering the genotype-phenotype relationship for ornamentation. They also caution against automatically attributing greater offspring attractiveness or viability to an extra-pair mate's superior genetic quality, as without controlling for differential maternal investment we may significantly overestimate the role of genetic benefits in the evolution of extra-pair mating behaviour.     

Explicit experimental evidence for the effectiveness of proximity as mate-guarding behaviour in reducing extra-pair fertilization in the Seychelles warbler

Extra-pair copulations (EPCs; copulations outside the pair bond) are widespread in birds and may result in extra-pair fertilizations (EPFs). To increase reproductive success, males should not only seek to gain EPFs, but also prevent their own females from gaining EPFs. Although males could reduce the number of EPCs by their mates, this does not necessarily mean that they reduce the number of EPFs; indeed several studies have found no association between EPCs and EPFs. Male Seychelles warblers (Acrocephalus sechellensis) follow their partner closely during the period when the pair female is most receptive (fertile period). We show that males that guarded their mates more closely were less likely to have extra-pair young in their nest. This study on the Seychelles warbler is the first to provide explicit experimental evidence that mate guarding is effective in reducing EPFs. First, in territories where free-living males were induced to stop mate guarding during the pair female's fertile period, extra-pair parentage was higher than in the control group. Second, in the experimental group, the probability of having an extra-pair nestling in the nest was positively associated with the number of days during the fertile period for which mate guarding was artificially stopped. Thus, male mate guarding was effective in reducing the risk of cuckoldry.     

Changes in Sleep Time and Sleep Quality across the Ovulatory Cycle as a Function of Fertility and Partner Attractiveness

Research suggests that near ovulation women tend to consume fewer calories and engage in more physical activity; they are judged to be more attractive, express greater preferences for masculine and symmetrical men, and experience increases in sexual desire for men other than their primary partners. Some of these cycle phase shifts are moderated by partner attractiveness and interpreted as strategic responses to women''s current reproductive context. The present study investigated changes in sleep across the ovulatory cycle, based on the hypothesis that changes in sleep may reflect ancestral strategic shifts of time and energy toward reproductive activities. Participants completed a 32-day daily diary in which they recorded their sleep time and quality for each day, yielding over 1,000 observations of sleep time and quality. Results indicated that, when the probability of conception was high, women partnered with less attractive men slept more, while women with more attractive partners slept less.     

Human mate guarding

Long-term committed mating is a fundamental strategy in the human repertoire. Successful enactment of this strategy requires solving two related adaptive problems--fending off potential mate poachers and preventing a mates from defecting. Mate guarding adaptations evolved to solve these persistent problems. Those who failed in mate guarding risked suffering substantial reproductive costs ranging from genetic cuckoldry to reputational damage to the entire loss of a mate. Because the precise nature of the adaptive problems confronted differed historically for the sexes, men and women evolved corresponding differences in the underlying psychology of mate guarding. Men's mate guarding, relative to that of women's, is strongly triggered as a consequence of being mated to young and physically attractive women, being confronted by interested rivals who have superior economic resources or prospects, and having a mate who displays signs of sexual involvement with a rival. Women's mate guarding, relative to that of men's, is triggered as a consequence of being mated to men high in income and status striving, rivals who are more physically attractive, and having a partner who shows signs of emotional involvement with another woman. Behavioral output of mate guarding adaptations range from vigilance to violence.     

Sexual conflict and cryptic female choice in the black field cricket, Teleogryllus commodus

The prevalence and evolutionary consequences of cryptic female choice (CFC) remain highly controversial, not least because the processes underlying its expression are often concealed within the female reproductive tract. However, even when female discrimination is relatively easy to observe, as in numerous insect species with externally attached spermatophores, it is often difficult to demonstrate directional CFC for certain male phenotypes over others. Using a biological assay to separate male crickets into attractive or unattractive categories, we demonstrate that females strongly discriminate against unattractive males by removing their spermatophores before insemination can be completed. This results in significantly more sperm being transferred by attractive males than unattractive males. Males respond to CFC by mate guarding females after copulation, which increases the spermatophore retention of both attractive and unattractive males. Interestingly, unattractive males who suffered earlier interruption of sperm transfer benefited more from mate guarding, and they guarded females more vigilantly than attractive males. Our results suggest that postcopulatory mate guarding has evolved via sexual conflict over insemination times rather than through genetic benefits of biasing paternity toward vigorous males, as has been previously suggested.     

Female mate choice and male–male competition in Tonkean macaques: Who decides?

The theory of sexual selection predicts that females should be discriminatory in the choice of sexual partners. Females can express their choice in two ways. In direct mate choice, they show preferences for certain partners. In indirect mate choice, they select partners by displaying sexually attractive traits, thus eliciting contest competition between males. We focused on a primate species in which females advertise the timing of their ovulation and studied the balance between these two choice strategies. We tested predictions related to three hypotheses about direct and indirect female choice, namely the best‐male, graded‐signal and weak‐selectivity hypotheses. We investigated the sexual and agonistic interactions occurring during oestrous periods in five captive groups of Tonkean macaques (Macaca tonkeana). The results showed that dominant males used mate guarding to monopolise sexual access to parous females that were in the fertile stage of their reproductive cycle, while lower‐ranking males monitored only nulliparous females. The distribution of sexual presentations indicated that females accepted different types of partners, supporting the weak‐selectivity hypothesis regarding direct mate choice. The analysis of behavioural sequences revealed that mate‐guarding males used mild coercive behaviours to prevent females from mating with other males at conception time. The distribution of mounts showed that females mainly mated with dominant males, which leads us to argue that the best‐male hypothesis provides the most parsimonious explanation regarding indirect mate choice in Tonkean macaques. At the individual level, it may be concluded that male competitive strategies prevented females from exercising direct mate choice. At the evolutionary level, however, female sexual advertising and thus indirect choice promoted competition between males. The outcome is that indirect mate choice appears more important than direct mate choice in female Tonkean macaques.     

Sexual selection,germline mutation rate and sperm competition

Background  

An important component of sexual selection arises because females obtain viability benefits for their offspring from their mate choice. Females choosing extra-pair fertilization generally favor males with exaggerated secondary sexual characters, and extra-pair paternity increases the variance in male reproductive success. Furthermore, females are assumed to benefit from 'good genes' from extra-pair sires. How additive genetic variance in such viability genes is maintained despite strong directional selection remains an evolutionary enigma. We propose that sexual selection is associated with elevated mutation rates, changing the balance between mutation and selection, thereby increasing variance in fitness and hence the benefits to be obtained from good genes sexual selection. Two hypotheses may account for such elevated mutation: (1) Increased sperm production associated with sperm competition may increase mutation rate. (2) Mutator alleles increase mutation rates that are revealed by the expression of condition-dependent secondary sexual characters used by choosy females during their mate choice. M Petrie has independently developed the idea that mutator alleles may account for the maintenance of genetic variation in viability despite strong directional selection.     

Male-Biased Sex Ratios in Offspring of Attractive Males in the Guppy

The attractiveness hypothesis predicts that females produce offspring with male-biased sex ratios when they mate with attractive males because their male offspring will inherit the paternal sexual attractiveness and may have high reproductive success. In this study, we examined the effect of the attractiveness of the male guppy Poecilia reticulata in terms of the conspicuousness of its orange spot patterns, important criteria affecting female choice in this species, on the offspring sex ratios. We found that food-manipulation treatment altered the conspicuousness of the orange spot patterns in a full-sibling male pair. When females were presented to these males, they showed a greater mate preference for males having brighter orange spots than for those having duller orange spots. Subsequently, half of the females were mated with the preferred males and the remaining females were mated with the less preferred males. When the females exhibited a greater preference for their mates, their offspring sex ratios were more male biased. These results appear to be consistent with the prediction of the attractiveness hypothesis. In the guppy, as male sexual attractiveness is heritable, the male-biased sex ratios of the broods of attractive males may be adaptive.     

Explicit experimental evidence for the role of mate guarding in minimizing loss of paternity in the Seychelles warbler

Extra-pair copulations (EPCs) (copulations outside the pair bond) resulting in extra-pair fertilizations (EPFs) are widespread in birds. To increase reproductive success, males should not only seek EPCs, but also prevent their females from having EPFs. Male Seychelles warblers (Acrocephalus sechellensis) follow their partner closely during the period when these females are most receptive (fertile period). The Seychelles warbler is the first species to offer explicit experimental evidence that mate guarding functions as paternity guarding: in territories where free-living males were induced to stop mate guarding during the pair female''s fertile period, the rates of intrusions by other males and successful EPCs (male mounting female) were significantly higher than those observed in the control group and in the absence of mate guarding the frequency of successful EPCs increased significantly with local male density. Male warblers do not assure their paternity through frequent copulations to devalue any sperm from other males: males do not copulate with their partners immediately following a successful EPC obtained by their partners, the frequency of successful within-pair copulations does not increase with the frequency of successful EPCs and females initiate all successful copulations and are capable of resisting copulation attempts.     

The evolutionary psychology of physical attractiveness: Sexual selection and human morphology

Psychological evidence suggests that sex differences in morphology have been modified by sexual selection so as to attract mates (intersexual selection) or intimidate rivals (intrasexual selection). Women compete with each other for high quality husbands by advertising reproductive value in terms of the distribution of fat reserves and by exaggerating morphological indicators of youthfulness such as a small nose and small feet and pale, hairless skin. Men's physical appearance tends to communicate social dominance, which has the combined effects of intimidating reproductive rivals and attracting mates. In addition to their attractiveness and intimidatory effects, human secondary sexual characters also provide cues to hormonal status and phenotypic quality consistent with the good genes model of sexual selection (which includes parasite resistance). Low waist-hip ratio is sexually attractive in women and indicates a high estrogen/testosterone ratio (which favors reproductive function). Facial attractiveness provides honest cues to health and mate value. The permanently enlarged female breast appears to have evolved under the influence of both the good genes and the runaway selection mechanisms. The male beard is not obviously related to phenotypic quality and may have evolved through a process of runaway intersexual selection.     

Willingness to engage in casual sex

Sexually dimorphic mate selection strategies were examined in 200 university students reporting their willingness to engage in casual sexual encounters with hypothetical individuals of the opposite sex. Using a questionnaire format, the possibility of forming a long-term relationship was manipulated, while risk of disease, pregnancy, and detection was eliminated across all conditions. In addition, potential partners varied in level of attractiveness, and in personality and behavioral characteristics. As expected, men reported a greater anticipated willingness to engage in sexual intercourse across all conditions compared with women. The possibility of forming a long-term relationship elevated women’s, but not men’s, willingness for sexual intercourse. While a potential partner’s attractiveness had a significant positive overall effect on responses, reducing their relative attractiveness had a greater negative impact on men’s responses. Reference to the parental qualities of a potential partner significantly increased women’s, but not men’s, anticipated willingness for sexual intercourse. Describing a hypothetical partner as non-aggressive (safe) marginally increased women’s willingness (p<.09) and did not affect men’s responses. The wording of items relevant to this condition may have resulted in the potential partner sounding "wimpy" rather than nonaggressive, and this may have reduced the expected effect of this manipulation. The possibility that women may trade off personality and behavioral characteristics with attractiveness to a greater degree than men when assessing potential mates is considered.     

Extra-pair paternity and song characteristics in the willow warbler Phylloscopus trochilus

Complexity in bird song is often argued to be advantageous in processes of sexual selection, and numerous studies show that characteristics of song are associated with increased success in territory defence or mate attraction. Less evidence exists on the relationship between bird song characteristics and patterns of extra-pair paternity. We tested whether males that suffered from extra-pair paternity differed in song characteristics from males with no extra pair paternity in the willow warbler Phylloscopus trochilus . In the Scottish population that we studied, we found that 23.5% of young were not related to the social father, and that 47% of nests contained at least one extra pair young. Older males were less likely to suffer paternity loss. While song repertoire size was not related to loss of paternity, males with short songs suffered higher paternity loss than males with long songs. Although arrival date is a good correlate of social mate choice in this population, it was not related to extra-pair paternity. These results suggest that females use song length, or a trait correlated with this song characteristic, as a cue for choosing extra-pair partners in this species, or alternatively that variance in the success of mate guarding or female coercion is related to this song variable.     

The role of genetic constraints and social environment in explaining female extra-pair mating

Why do females of socially monogamous species engage in extra-pair copulations? This long-standing question remains a puzzle, because the benefits of female promiscuous behavior often do not seem to outweigh the costs. Genetic constraint models offer an answer by proposing that female promiscuity emerges through selection favoring alleles that are either beneficial for male reproductive success (intersexual pleiotropy hypothesis) or beneficial for female fecundity (intrasexual pleiotropy hypothesis). A previous quantitative genetic study on captive zebra finches, Taeniopygia guttata, reported support for the first, but not for the second hypothesis. Here, we re-examine both hypotheses based on data from lines selected for high and low male courtship rate. In contrast to previous conclusions, our new analyses clearly reject the hypothesis that male and female promiscuity are genetically homologous traits. We find some support for a positive genetic correlation between female promiscuity and fecundity. This study also shows that the behavioral outcome of extra-pair courtships primarily depends on individual-specific female preferences and not on the “attractiveness” of the social mate. In contrast, patterns of paternity are strongly influenced by the social partner and the pair bond, presumably reflecting variation in copulation behavior, fertility, or sperm competitiveness.     

Is egg carrying attractive? Mate choice in the golden egg bug (Coreidae,Heteroptera)

In several species of fish, females select males that are already guarding eggs in their nests. It is a matter of debate as to whether a female selects a good nest site for her offspring (natural selection) or a male for his attractiveness (sexual selection). The golden egg bug, Phyllomorpha laciniata Vill, resembles fish in the sense that mating males carry more eggs than single males, but in the bugs, female mate choice is decoupled from egg site choice. The sexual selection hypothesis predicts that if females select males using male egg load as a cue for male quality, they should not mate with a male when eggs are removed, regardless of his mating attempts. When individual females were enclosed with an egg-loaded male and an unloaded male, they mated equally often with both males, although the loaded males courted more. In addition, when only successful males were used, females mated equally often with the loaded male and the unloaded male irrespective of sex ratio. Male choice rather than female choice affected mating frequency when sex ratio was equal. Therefore, females do not select the male by the eggs he carries, but successful males may receive many eggs due to egg dumping by alien females while they mate or as a consequence of mate guarding.