Cues to paternity: Do partner fidelity and offspring resemblance predict daughter-directed sexual aversions?

Despite the profound influence of relatedness on mating and cooperative behavior in humans, the cues men use to assess paternity and guide offspring-directed behavior have yet to be fully resolved. According to leading theories of kin detection, kinship cues should influence both sexual and altruistic motivations because of fitness consequences associated with inbreeding and welfare tradeoff decisions, respectively. Prior work with paternity assessment, however, has generally evaluated candidate cues solely by demonstrating associations with altruism. Here we (i) replicate past work that found effects of phenotypic resemblance and perceived partner fidelity on offspring investment; and (ii) evaluate whether both phenotypic resemblance and perceived partner fidelity meet the more stringent criteria suggested by theory—that is, whether they also predict inbreeding aversions. We report on two studies, one from a population-based sample of Finnish fathers (N?=?390), the other from a Mechanical Turk sample (N?=?700), and furnish evidence in strong support of perceived partner fidelity as a cue to paternity. Support for resemblance as a cue to paternity was decidedly weaker. We discuss a non-kin-based role that resemblance might play in altruistic decision-making, consider whether men might use additional kinship cues to meet the computational challenges associated with paternity assessment, and provide suggestions for future research.     

Is there a link between paternity concern and female genital cutting in West Africa?

Here we explore the relationship between female genital cutting (FGC), sexual behaviour, and marriage opportunities in five West African countries. Using large demographic datasets (n 72,438 women, 12,704 men, 10,695 couples) we explore key (but untested) assumptions of an evolutionary proposal that FGC persists because it provides evolutionary fitness benefits for men by reducing non-paternity rates. We identify and test three assumptions implicit in this proposal. We test whether cut women have reduced extra-pair sex before or within marriage; whether FGC is associated with a younger age at marriage as an indication of partner preference; and whether individual and group-level indicators of paternity concern are associated with a stronger preference for marriage to women with FGC.Our results show that FGC status does not affect the odds of women engaging in several indicators of premarital sex, however women with FGC have significantly lower odds of having more than one lifetime sexual partner. We also show that women with FGC get married at a younger age which supports the argument that FGC status influences women's marriage opportunities, even when it does not restrict sexual activity. Finally, we find that in population groups where reported sexual activity and perceived risk of women's extra-pair sex is high, men have higher odds of marrying a first wife with FGC. Together, these results indicate that paternity certainty may be one of several factors contributing to the persistence of FGC in this sample, and that group-level sexual norms are key to maintaining the practice of FGC through the marriage market.     

Nuptial gift consumption influences female remating in a scorpionfly: male or female control of mating rate?

In the scorpionfly Panorpa cognata, males provide females with saliva secretions as nuptial food gifts. Consequently, females derive material benefits and possibly also genetic benefits from multiple matings. Females therefore generally should have a high motivation to remate. Males, on the other hand, do not share this interest, which will generate a sexual conflict over remating interval, possibly leading to male adaptations that prevent females from remating with other males. In this study, I found that mated females were less prone to copulate than virgin females, despite female benefits of multiple matings. Further, I found that the remating interval was significantly longer if the first copulation was long compared to shorter matings. This effect does not entirely depend on copulation duration per se, but on the amount of saliva, that a female is consuming during copulation. These results suggest a mating-induced refractory period and can be interpreted as male manipulation of female remating behaviour mediated through substances in the nuptial gift. Alternatively, receiving large nuptial gifts may decrease the prospective direct fitness benefits from further copulations, and thus change optimal female remating rate. Furthermore, gift size has been shown to correlate with male nutritional condition, which may be an indicator of male genetic quality. Females may therefore benefit indirectly by not remating following copulations involving large saliva gifts. In this scenario, female remating interval would be an effect of cryptic female choice.     

Does female mortality drive male semelparity in dasyurid marsupials?

In some members of the marsupial families Didelphidae and Dasyuridae, males are semelparous, that is, they live for only one mating season. Semelparity is proposed to be the result of the high energy demands of competing for matings with many females during a short breeding season. We argue that high adult female mortality rates between mating and weaning of the offspring selects for a 'bethedging' mating strategy in males. We tested this hypothesis in a well-studied field population of Antechinus agilis by estimating the number of females a male needs to mate with in order to have a high chance of siring at least one offspring that survives to the next breeding season. Our hypothesis predicts that species in which males are semelparous should have higher female mortality rates than species in which males are iteroparous. The limited available data for dasyurid marsupials support this prediction.     

What makes a nest-building male successful? Male behavior and female care in penduline tits

Why do females increase parental effort when caring for theoffspring of attractive males? First, attractive males may bepoor fathers so that their females are compelled to increasetheir own contribution in order to fledge some young (the partner-compensationhypothesis). Second, females mated to attractive males may bewilling to increase their parental effort to reap high indirectbenefits for their offspring, and in turn males can decreasetheir own contribution (the differential allocation hypothesis[DAH]). We investigated these hypotheses in the penduline titRemiz pendulinus, a small passerine bird that has sequentialpolygamy by both sexes and strict uniparental care either bythe male or the female. We focused on two sexually selectedmale traits: nest size and nest-building behavior. We show thatmale care is unrelated to nest-building behavior, whereas femalesare more likely to care for the offspring of those males thatspend more time nest building. Females also more likely carefor the offspring of males that build large nests. Consequently,the reproductive success of males increases with nest size andnest-building behavior. Our results are consistent with theDAH and suggest that nest-building behavior and nest size areunder postmating sexual selection in penduline tits.     

The Bruce effect revisited: is pregnancy termination in female rodents an adaptation to ensure breeding success after male turnover in low densities?

Animal-mediated seed dispersal (zoochory) is considered to be an important mechanism regulating biological processes at larger spatial scales. To date, intra-specific variation in seed disperser quality within seed-dispersing animals has not been studied. Here, I employed seed feeding trials to quantify individual differences in disperser quality within the common carp (Cyprinus carpio) using seeds of two aquatic plants: unbranched bur-reed (Sparganium emersum, Sparganiaceae) and arrowhead (Sagittaria sagittifolia, Alismataceae). I found substantial variation among carp individuals in their propensity to ingest seeds and their ability to digest them, resulting in up to 31-fold differences in the probability of seed dispersal. In addition, there were significant differences in the time that seeds are retained in their digestive systems, generating a twofold difference in the maximum distance over which they can potentially disperse seeds. I propose that seed-eating animal species consist of individuals that display continuous variation in disperser quality, with at one end of the continuum individuals that are likely to eat seeds, pass them unharmed through their digestive tract and transport them over large distances to new locations (i.e. high-quality seed dispersers) and at the other end individuals that rarely eat seeds, destroy most of the ones they ingest and transport the few surviving seeds over relatively short distances (low-quality seed dispersers). Although individual differences in seed dispersal quality could be the result of a variety of factors, these results underline the ecological and evolutionary potential of such variation for both plants and animals.     

Do female spotless starlings Sturnus unicolor adjust maternal investment according to male attractiveness?

In birds, female egg allocation patterns have a strong influence in offspring development and differential investment in egg size and composition has been shown to respond to male attractiveness. In this study we experimentally manipulated the perceived attractiveness of male starlings Sturnus unicolor by increasing the amount of green material in some nests (a male courtship display in this species). We predicted that, if female investment before laying is related to male attractiveness, experimental females would increase their reproductive investment in response to the addition of plants in their nests when compared to control females. We found that our manipulation caused variations in female reproductive investment in a way that seems to influence offspring quantity but not offspring quality: Females laid larger clutch sizes but not larger eggs when green plant material was added. However, yolk androgens contents were not related to the experimental manipulation. Contrary to expectations, females breeding in experimental nests laid eggs with smaller amounts of eggshell pigments. Interestingly, we found that eggs laid later in the sequence had higher testosterone levels and showed more intense egg colouration than eggs laid earlier in the sequence. These differences at the within-clutch level suggest that selection has favoured compensatory strategies for hatching asynchrony. Alternatively, since nest sabotages by other females are most common at the beginning of laying, this could be seen as female strategy to minimise losses due to nest sabotages. As far as we know, this is the first study to show that an external egg characteristic such as blue-green colouration reflects yolk androgen concentration.     

Do male ornaments signal immunity in the common yellowthroat?

Male ornaments may advertise genetic benefits to females choosingmates. These benefits may come in the form of genes for resistanceto parasites and disease. Thus, females that prefer more ornamentedmales as mates may receive genes for enhanced immune systemfunction for their offspring. The common yellowthroat (Geothlypistrichas) is the only species to date in which a male plumageornament (size of the black facial mask) is known to be relatedto extrapair mating, and extrapair young are more immunocompetentthan their within-pair half-sibs (at least in colder years).To investigate whether male mask size signals superior immunefunction, we examined male ornaments (mask and bib size andcolor) in relation to measures of overall health (hematocrit)and immune system function (plasma immunoglobulin G [IgG] concentrationand cutaneous immune activity). We also investigated the rolethat testosterone may play in mediating the relationship betweenornaments and immunity. Male mask size was correlated positivelywith IgG level, suggesting that male mask size may indicatehumoral immunity. However, mask size was correlated negativelywith hematocrit and cutaneous immune activity (our measure ofnestling immunocompetence). Bib size and color were not relatedto these indices of immune function or health. Plasma testosteronelevel was neither related to immune function or health nor tothe expression of male ornaments. These results suggest thatthere might be a trade-off between immune system components,as well as between immunity and the production or advertisementof male ornaments.     

Do pheromones reveal male immunocompetence?

Pheromones function not only as mate attractors, but they may also relay important information to prospective mates. It has been shown that vertebrates can distinguish, via olfactory mechanisms, major histocompatibility complex types in their prospective mates. However, whether pheromones can transmit information about immunocompetence is unknown. Here, we show that female mealworm beetles (Tenebrio molitor) prefer pheromones from males with better immunocompetence, indicated by a faster encapsulation rate against a novel antigen, and higher levels of phenoloxidase in haemolymph. Thus, the present study indicates that pheromones could transmit information about males' parasite resistance ability and may work as a reliable sexual ornament for female choice.     

Why do males emerge before females? protandry as a mating strategy in male and female butterflies

Summary The reproductive strategy of butterfly males can be defined as being to maximize the number of females mated. We have earlier shown that, if the eclosion period of females is regarded as given, males should emerge before females to achieve maximal reproductive success. However, females may also be considered to have a reproductive strategy with respect to the issue when to emerge. In this paper we assume that females are selected to minimize the time spent unmated (to minimize prereproductive death), and analyze when females should optimally emerge in relation to males to achieve this end. We show that there is no conflict between the sexes with respect to the timing of eclosion when the length of the eclosion period is approximately equal for males and females. Thus, protandry should be considered a reproductive strategy of both males and females.     

A paternity advantage for speedy males? Sperm precedence patterns and female re-mating frequencies in a sexually cannibalistic praying mantid

Scramble competition polygyny is expected when females and/or resources are widely dispersed and not easily monopolized by males, or when there is an abundance of mates during an extremely restricted reproductive period. Additional factors such as first male sperm precedence or low female re-mating rate might further explain the propensity of males to engage in scramble competition. The sexually cannibalistic praying mantid Pseudomantis albofimbriata exhibits a polygynous mating system, where females exist in low-density populations and male competition manifests as the race to find females rather than as direct physical fighting. Here, we aim to determine whether there is a paternity advantage for the first-male to mate and/or a low frequency of female re-mating. First, we determined sperm precedence patterns in P. albofimbriata using the sterile male technique. Second, we tested the likelihood of female re-mating in P. albofimbriata by comparing the close-range approach behaviour and frequency of successful mating attempts for males when paired with virgin as opposed to recently mated females, and by comparing the frequency of long-distance male attraction between virgin and mated females. We found no paternity advantage for the first male to mate, rather a second male advantage. Although mated females were not rejected by males when approached from close-range, they were chemically unattractive to males searching from a distance. Since initial mate attraction in many praying mantids, including P. albofimbriata, is mediated via long-distance chemical communication, we believe the latter result is more ecologically relevant and therefore more important. These results suggest that the relatively low frequency of female re-mating observed in P. albofimbriata may be an additional factor driving scramble competition in this system.     

Why do multiple traits determine mating success? Differential use in female choice and male competition in a water boatman

Mating success is often determined by multiple traits, but why this occurs is largely unknown. Much attention has been paid to female preferences for multiple traits, but surprisingly few researchers have addressed the possibility that multiple traits are important because they serve different functions in female choice and male-male competition. Differential trait function could result from a conflict of interest between the sexes or from constraints forcing the sexes to pay attention to different traits. I show that traits determined at distinct life-history stages differ in their importance in female choice and male-male competition in a water boatman Sigara falleni. Juvenile conditions determined body and foreleg pala size and were the main determinants of mating success under female choice, whereas adult conditions determined body mass and influenced mating success when male competition was included. This differential use of condition-dependent traits under the two selection regimes appeared to arise partly from a conflict between the sexes, since the two selection forces (female choice and male competition) conflict for selection on pala size, and partly from constraints, as females appeared unable to assess adult condition.     

Do males choose their mates in the lekking moth Achroia grisella? Influence of female body mass and male reproductive status on male mate choice

Lekking males aggregate to attract females and contribute solely to egg fertilization, without any further parental care. Evolutionary theory therefore predicts them to be nonchoosy toward their mates, because any lost mating opportunities would outweigh the benefits associated with such preferences. Nevertheless, due to time costs, the production of energetically costly sexual displays, and potential sperm limitation, the mating effort of lekking males is often considerable. These factors, combined with the fact that many females of varying quality are likely to visit leks, could favor the evolution of male mate preferences. Here, we show that males of the lekking lesser wax moth, Achroia grisella, were indeed more likely to mate with heavier females in choice experiments, even at their virgin mating (i.e., when their reproductive resources have not yet been depleted by previous matings). This differential female mating success could not be attributed to female behavior as heavy and light females showed similar motivation to mate (i.e., latency to approach the males) and time to copulate. Males seem to benefit from mating with heavier females, as fecundity positively correlated with female mass. This new empirical evidence shows that male mate choice may have been underestimated in lekking species.     

Male–male competition facilitates female choice in sticklebacks

In many species, secondary sexual characteristics are used in both male–male competition and in attracting females. This suggests that social control of deception could contribute to the maintenance of honest sexual signalling. In the three-spined stickleback Gasterosteus aculeatus, male red breeding coloration plays a dual role in sexual selection by functioning as both a threat signal in male–male competition and as a cue for female choice. To investigate whether male competition determines the level of signalling, the expression of red coloration and courtship activity were recorded both before and after male interactions. The results show that male competition influences signal expression by increasing the difference between males in signalling level. This in turn facilitates female choice and induces a preference for dominant males. Since a preference for dominant males may benefit females both directly and indirectly in this species with exclusively paternal care, male–male competition seems to increase the honesty of signalling and, thus, facilitates female choice in relation to male quality. This may increase the intensity of sexual selection and promote the evolution of breeding aggregations.     

Can too strong female choice deteriorate male ornamentation?

Competition for limited resources can have fundamental implications for population dynamics. However, the effects of resource depletion have rarely been discussed in the context of sexual selection, even though mate choice typically favours males who outperform others in securing access to some limited resource. Here, we develop a model to investigate the question of resource competition as a form of male-male competition in the context of male sexual displays. We phrase our model in terms of male bowerbirds either searching for or stealing resources (ornamental objects) valued by females, and compare the model findings with published studies of time allocation to various activities in different species of bowerbirds. The basic idea of the model, however, extends to cases where the resource is used less directly for the development of sexual ornamentation, such as males excluding others' access to food. We show that if males compete for resources used in sexual displays, intense female preference for high-quality displays can lead to poorer prospects for efficient choice by females. This is because males benefit from excluding others' access to resources used in displays, damaging the overall efficiency of resource use in the population, and the accuracy with which females can judge male ability to gain such resources. The evolution of female choice may therefore have a self-limiting nature when it poses a selection pressure on male resource acquisition.     

Why does inbreeding reduce male paternity? Effects on sexually selected traits

Mating with relatives has often been shown to negatively affect offspring fitness (inbreeding depression). There is considerable evidence for inbreeding depression due to effects on naturally selected traits, particularly those expressed early in life, but there is less evidence of it for sexually selected traits. This is surprising because sexually selected traits are expected to exhibit strong inbreeding depression. Here, we experimentally created inbred and outbred male mosquitofish (Gambusia holbrooki). Inbred males were the offspring of matings between full siblings. We then investigated how inbreeding influenced a number of sexually selected male traits, specifically: attractiveness, sperm number and velocity, as well as sperm competitiveness based on a male's share of paternity. We found no inbreeding depression for male attractiveness or sperm traits. There was, however, evidence that lower heterozygosity decreased paternity due to reduced sperm competitiveness. Our results add to the growing evidence that competitive interactions exacerbate the negative effects of the increased homozygosity that arises when there is inbreeding.     

Do all Blackcaps Sylvia atricapilla show winter site fidelity?

Fidelity to previous wintering areas (i.e. site fidelity) has important ecological and evolutionary implications. However, since the percentage of recaptures of ringed birds at the same wintering area in subsequent years does not allow the estimation of the proportion of birds alive that exhibit site fidelity, previous studies on different passerine species have failed to show the true extent of this site fidelity. Here we use a recent approach, based on the comparison of survival rate estimates from capture-recapture data at single field stations with recovery data from a much larger area. The idea is to determine the proportion of birds still alive that return to the area. The study was carried out on the Blackcap Sylvia atricapilla , comparing the capture-recapture data of two field stations (Pilas [SW] and Tiana [NE]) situated 1000 km apart in Spain with the complete winter recovery data of the Spanish Ringing Office. Totals of 1936 and 3976 Blackcaps were ringed at Pilas (1981–1986) and Tiana (1975–1986), respectively, and the numbers of recaptures in subsequent winters were 94 and 34. The pooled annual survival rate estimation based on the two sites is 0.40 (s.e. = 0.05). The annual survival rate estimated from the Spanish ringing recovery data was 0.48 (s.e. = 0.08), which is not significantly different. We can conclude, therefore, that migrant Blackcaps in Spain, if alive, tend to return year after year to the previous wintering site.