Post-copulatory mate guarding by males of the demselfly Hetaerina vulnerata Selys (Odonata: Calopterygidae)

Males of the calopterygid damselfly Hetaerina vulnerata remain with their mates after copulating with them. The species exhibits two unusual features of post-copulatory mate guarding. First, a male will often leave his territory to accompany a female in tandem on a search for oviposition sites elsewhere. Second, a male will perch near his ovipositing female even though she completely submerges when egg-laying and cannot be captured and mated by another male while she is underwater. These activities carry two potential costs: (1) a male may miss other receptive females while guarding one mate and (2) he may lose his territory to an interloper while he is absent. These costs were low, however, because territorial males secured only one mating per 3.6 days on average. Moreover, 23 times out of 26, territorial males reclaimed their plots quickly after being away for 30–60 min. The gain from postcopulatory guarding came from being present to recapture a female should she fly up from the water after rejecting an oviposition site. There was a 40% chance that a female would leave one site to search for another during an oviposition bout. If the male were not present, his mate would be captured and mated by another individual (no female ever selected an oviposition site without being carried to it by a male). Her new partner would fertilize the remaining eggs in the female's clutch (if sperm precedence occurs in this species). The total number of eggs fertilized by a male will be affected by how well he prevents any one mate from copulating again before she lays her entire clutch and the total number of receptive females he captures. The variation in the degree of mate guarding by male odonates seems to be the evolutionary outcome of differences in fitness gains derived from these two competing activities in different ecological settings.     

LIFETIME REPRODUCTIVE SUCCESS AND THE OPPORTUNITY FOR SELECTION IN A NONTERRITORIAL DAMSELFLY (ODONATA: COENAGRIONIDAE)

Major components of male and female lifetime reproductive success (LRS) were quantified for a damselfly that exhibits “scramble competition” for mates. The opportunity for selection on male reproduction was potentially 2.9 times that for females. Differential fertility/clutch and survivorship each accounted for about half of the total variation in female reproductive success. Variation in fertilization efficiency accounted for 7% of the total opportunity for selection on males. Although differences in survivorship and mating efficiency each contributed to about a third of the total opportunity for selection on male reproduction, both components appeared to be influenced by random factors. Survivorship was age-independent, and the mating distributions among males with equal mating opportunities were indistinguishable from those expected if matings were random with respect to male phenotype. Because the proportion of the standarized variance (I) in LRS that was attributed to sexual selection depended on the way the selective episodes were defined, the sample of individuals included in the partitioning analysis, and the degree of sexual selection on mated males that could be detected, my results caution against drawing conclusions about the dynamics of sexual selection on populations based on a superficial comparison of I values.     

Selection of male traits during and after copulation in the seedbug Lygaeus simulans (Heteroptera, Lygaeidae)

In order to evaluate selection of male morphological traits during copulation, a laboratory experiment was performed with the promiscuous seedbug Lygaeus simulans. Three male traits suspected as putative targets of selection were measured: weight, fluctuating asymmetry of a measure on the forewings, and length of a conspicuous genital structure, the processus gonopori. As fitness measures we considered total fecundity (number of fertilized eggs), insemination and fertilization success (established if a female laid fertilized eggs after copulation), and the interval between copulation and oviposition. Eighty-four males were allowed a single copulation with one virgin female each. Out of 67 copulations, 27 (40.2%) resulted in fertilized eggs and the oviposition latency ranged from 6 to 26 days. Regressions of male traits on the fitness measures showed significant phenotypical selection of two male traits: (1) males of average weight are more likely to achieve fertilization and (2) the oviposition latency was shorter for males with lower asymmetry. The copulation-oviposition interval may be especially important for male fertilization success because Lygaeus males perform copulatory mate guarding and the last male copulating with a female fertilizes most of the eggs. No selection of the genitalic trait was detected.     

Lifetime reproductive success,selection on lifespan,and multiple sexual ornaments in male European barn swallows

Natural and sexual selection arise when individual fitness varies according to focal traits. Extra‐pair paternities (EPPs) can affect the intensity of selection by influencing variance in fitness among individuals. Studies of selection require that individual fitness is estimated using proxies of lifetime reproductive success (LRS). However, estimating LRS is difficult in large, open populations where EPPs cause reallocation of biological paternity. Here, we used extensive field sampling to estimate LRS in a population of barn swallows (Hirundo rustica) to estimate selection on lifespan and ornamental traits of males. We found selection on lifespan mediated both by within‐ and extra‐pair fertilization success and selection on tail length mediated by within‐ but not extra‐pair fertilization success. In addition, we found selection on tail white spots via extra‐pair fertilization success after controlling for selection on other traits. These results were not confounded by factors that hamper studies of LRS, including nonexhaustive sampling of offspring and biased sampling of males. Hence, natural and sexual selection mediated by LRS operates on lifespan, tail length, and size of the tail white spots in barn swallows.     

Brood conspicuousness and clutch viability in male‐caring assassin bugs (Rhinocoris tristis)

Abstract 1. Conspicuousness to mates can bring benefits to both males (increased mating success) and females (reduced search costs), but also brings costs (e.g. increased predation and parasitism). Assassin bugs, Rhinocoris tristis, lay egg clutches either on exposed stems or hidden under leaves. Males guard eggs against parasitoids. Guarding males are attractive to females who add subsequent clutches to the brood. This is an excellent opportunity to study the effects of conspicuousness on the fitness of males and females. 2. Using viable eggs in a multi‐clutch brood as a correlate of fitness, the present study examined whether laying eggs on stems affected (1) female fitness, through exposure to parasitism and cannibalism, and (2) male fitness, through attracting further females. 3. Stem broods were more parasitised. However, males on stems accumulated more mates and more eggs, a net benefit even accounting for parasitism. The eggs gained from being on a stem were cannibalised. By contrast, higher mortality on stems suggests that females should gain by ovipositing on leaves. To the extent that egg viability represents fitness, male and female interests may therefore differ. This suggests a potential for sexual conflict that may affect other species with male care. 4. Despite higher costs, females actually initiated more broods, and subsequently added bigger clutches to broods, on stems than under leaves. This suggests either that viable eggs do not reflect fitness, or that females laid in unfavourable locations. The key is now to address lifetime fitness, since unmeasured factors may affect offspring viability post‐hatching, and to investigate who controls the location of oviposition in R. tristis.     

Sexual selection in the monogamous barn swallow (Hirundo rustica). II. Mechanisms of sexual selection

Mechanisms of sexual selection in the monogamous, sexually dimorphic barn swallow (Hirundo rustica) were studied during a seven-year period. First, the sex ratio of reproducing adults was male-biased, and mated males had significantly longer tail ornaments than unmated males. Secondly, some of the unmated individuals later committed infanticide and became mated with the mother of the killed brood. Fathers of killed broods had significantly shorter tails than other males, and there was a tendency for infanticidal males to have longer tail ornaments than other unmated males. Thirdly, long-tailed male barn swallows were more successful in acquiring extra-pair copulations than other males, and females involved in extra-pair copulations, as compared to females not involved in such copulations, had mates with shorter tail ornaments. Fourthly, male barn swallows having long tails as compared to short-tailed males acquired mates in better body condition. Females mated to long-tailed males reproduced earlier, laid more eggs and were more likely to have two clutches than were females mated to short-tailed males. Finally, females mated to long-tailed males put more effort into reproduction than did other females, as evidenced by their relatively larger contribution to feeding of offspring. Thus, at least five different components of sexual selection affected male reproductive success. Selection arising from differential success during extra-pair copulations, differential reproductive success and differential male reproductive effort thus accounted for most of the selection on tail ornaments in male barn swallows.     

Unpredictable offspring survivorship in the damselfly, Megaloprepus coerulatus, shapes parental behavior, constrains sexual selection, and challenges traditional fitness estimates

Evolutionary biologists typically assume that the number of eggs fertilized or developing embryos produced is correlated with an individual's fitness. Using microsatellite markers, we document for the first time estimates of realized fitness quantified as the number of offspring surviving to adulthood in an insect under field conditions. In a territorial damselfly whose males defend tree hole oviposition sites, patterns of offspring survivorship could not be anticipated by adults. Fewer than half of the parents contributing eggs to a larval habitat realized any reproductive success from their investment. The best fitness correlate was the span over which eggs in a clutch hatched. Among parents, female fecundity and male fertilization success were poor predictors of realized fitness. Although body size was correlated with female clutch size and male mating success, larger parents did not realize greater fitness than smaller ones. The uncoupling of traditional fitness surrogates from realized fitness provides strong empirical evidence that selection at the larval stage constrains selection on mated adults.     

Bateman gradients in field and laboratory studies: a cautionary tale

Since tools of molecular genetics became readily available,our understanding of bird mating systems has undergone a revolution.The majority of passerine species investigated are sociallymonogamous, but have been shown to be genetically polygamous.Data sets from natural populations of juncos suggest that multiplemating by females results in a sexual selection gradient assteep for females as for males (a result that does not supportBateman's predictions). However, in males, fitness is enhanceddirectly through fertilization success with multiple matings;in females fitness benefits may be enhanced immediately throughdirect access to food, protection against predators, or otherresources received from males, or they may be delayed throughimprovement in offspring quality (e.g., through good genes,or greater genetic compatibility between the female and theextra-pair male). But a steep sexual selection gradient forfemales can be difficult to interpret. If all females copulatewith multiple partners that are equally likely to fertilizeeggs, then females that produce larger clutch sizes, for anyreason, will appear to have copulated with more males. Thatis, multiple sires have a higher probability of detection inlarger clutches than in smaller ones, giving the impressionthat females that mate with multiple males increase their reproductivesuccess. Yet, in most studies in which there is a correlationbetween number of offspring produced by females and number ofextra-pair males, causation has not been clearly establishedand other factors may explain the results. Additional complicationsin understanding male and female reproductive strategies are:(1) Molecular studies cannot detect extra-pair copulations thatdid not result in fertilizations; yet if a female acquires foodor other resources from extra-pair males, such extra-pair matingsmay have significant effects on female fitness. Thus, molecularstudies provide only a conservative estimate of the number ofextra-pair copulations or "mates" that a female has. (2) Clutchsize affects the probability that any given male will be successfulin fertilizing a female's eggs. Specifically, at any given point,a male's chances of fertilizing at least one egg in the female'sclutch will be greater as clutch size increases. We predictthat in avian species with small clutch sizes, males may beselected to be choosy and avoid extra-pair copulations, whilefemales should be selected to be less discriminating. Moreover,if extra-pair males provide resources that increase female fitness,the females should seek extra-pair copulations, whether or notthe males are likely to fertilize any of her eggs. Laboratory studies with insects have yielded clearer evidenceof the causal relationship between multiple mating and increasedfemale fitness. We review studies on a tenebrionid beetle inwhich female fecundity increases directly with number of mates.In these experiments, the nutritive value of the spermatophoresdoes not fully explain the increase in female reproductive success.     

Females avoid manipulative males and live longer

Abstract Female mate choice has been demonstrated in a wide variety of species and is now accepted as an important factor in sexual selection. One of the remaining questions, however, is why females prefer specific males. Do females or their offspring benefit from their choice? Or do females choose mates to minimize costs of mating? Here we show that, in the ovoviviparous cockroach Nauphoeta cinerea, where sexual selection has been well documented, females chose mates to avoid costly male manipulation. Females were partnered with preferred or nonpreferred mates, and fitness of the females measured. We found that females lived longer when they mated with preferred males. Female lifespan depended on the rate at which offspring developed from egg to parturition: slower development led to longer life. We manipulated the male pheromone and showed that the component of the pheromone blend that makes males attractive to females also delayed parturition. Thus, like other aspects of sexual conflict in this species, offspring development and thereby the mother's lifespan depended on exposure of females to specific components of the male pheromone. Males benefit from manipulating offspring development because females with accelerated parturition remained unreceptive whereas females with slower developing offspring readily remated after giving birth to their offspring. Our results suggest a hormone‐like role for the male pheromone in N. cinerea and provide the first direct evidence of mate choice to avoid male manipulation. This study shows that dominant males may not be preferred males if they are manipulating females, why multiple components with contrasting effects can exist in a sexual signal, and emphasizes the complex fitness relationships that can arise in species with sexual conflict.     

Cooperation and conflict among dunnocks, Prunella modularis, in a variable mating system

The mating system of a population of 90 breeding dunnocks (or hedge sparrow, Prunella modularis) included monogamy, polygyny, polyandry and polygynandry. Monogamous males guarded females during their fertile period to prevent neighbouring males from copulating. The most intense guarding occurred where two (unrelated) males shared a territory. Here, the alpha male tried to prevent the beta male from copulating with the female. Beta males were seen to copulate in only half the cases. They were more likely to succeed when the alpha male found it difficult to guard the female closely because her range was large, the vegetation was dense or there were other females breeding synchronously on the same territory. Close guarding and chasing by males reduced the female's feeding rate and was correlated with unhatched eggs in the nest. Females attempted to escape the alpha male's attentions and actively encouraged the beta male to mate. Beta males only helped to feed the young if they copulated with the female. Nestlings fed by two males and a female got more food and weighed more than those fed by just one male and a female. Indirect evidence suggested that when beta males failed to copulate, they destroyed eggs or young chicks. Females laid larger clutches when two males mated with them as opposed to one, thus adapting their clutch size to the amount of parental care they expected. The results of natural removal experiments and matched comparisons of the same female in different mating systems support these conclusions. For females, selection favours cooperative polyandry, whereas for males if favours polygyny; the variable mating system may reflect the different outcomes of this sexual conflict.     

Sexual selection: harem size and the variance in male reproductive success

Sexual selection is potentially stronger than natural selection when the variance in male reproductive fitness exceeds all other components of fitness variance combined. However, measuring the variance in male reproductive fitness is difficult when nonmating males are absent, inconspicuous, or otherwise difficult to find. Omitting the nonmating males inflates estimates of average male reproductive success and diminishes the variance, leading to underestimates of the potential strength of sexual selection. We show that, in theory, the proportion of the total variance in male fitness owing to sexual selection is approximately equal to H, the mean harem size, as long as H is large and females are randomly distributed across mating males (i.e., Vharem=H). In this case, mean harem size not only provides an easy way to estimate the potential strength of sexual selection but also equals the opportunity for sexual selection, I(mates). In nature, however, females may be overdispersed with VharemH. We show that H+(k-1) is a good measure of the opportunity for sexual selection, where k is the ratio Vharem/H. A review of mating system data reveals that in nature the median ratio for Vharem/H is 1.04, but as H increases, females tend to become more aggregated across mating males with V(harem) two to three times larger than H.     

Sex‐specific selection and sexual size dimorphism in the waterstrider Aquarius remigis

We estimated selection on adult body size for two generations in two populations of Aquarius remigis, as part of a long‐term study of the adaptive significance of sexual size dimorphism (SSD). Net adult fitness was estimated from the following components: prereproductive survival, daily reproductive success (mating frequency or fecundity), and reproductive lifespan. Standardized selection gradients were estimated for total length and for thorax, abdomen, genital and mesofemur lengths. Although selection was generally weak and showed significant temporal and spatial heterogeneity, patterns were consistent with SSD. Prereproductive survival was strongly influenced by date of eclosion, but size (thorax and genital lengths in females; total and abdomen lengths in males) played a significant secondary role. Sexual selection favoured smaller males with longer external genitalia in one population. Net adult fitness was not significantly related to body size in females, but was negatively related to size (thorax and total length) in males.     

Precopulation Sexual Selection in Nysius huttoni White (Heteroptera: Lygaeidae) in Relation to Morphometric Traits

Nysius huttoni White is a polygamous bug, endemic to New Zealand, and an important pest of wheat and brassicas. This bug has a female-biased sexual size dimorphism but relative to body length, males have longer antennae, suggesting that the allometric scales of antennal–body relationships may be highly selective in sexual selection. Body weight and most morphometric traits measured have no effect on mating success of either sex. Males significantly preferred mating with females having thicker abdomens, more mature eggs, and longer ovipositors. This result suggests that males may select their mates on the basis of immediate reproductive benefit: fertilizing more eggs and ensuring better survival of these eggs. Males with large genital structures have mating advantages over those with small ones, suggesting that precopulation sexual selection in this species act on male genital traits rather than body weight and nonsexual traits. Finally, females significantly preferred males with greater slopes for the antennal-body relationship for mating. The allometry in the male antennal length may be an indicator of male reproductive fitness.     

宁波滑蜥两性异形和雌性繁殖

蜥蜴的雌性繁殖特征对理解两性异形的进化原因起着重要作用。于2011年4月在安徽滁州采集宁波滑蜥(Scincella modesta),定量研究该种形态特征的两性异形和雌性繁殖特征,检验成体形态特征两性异形与雌性繁殖的相关性。研究共采集43条(17♀♀,26♂♂)宁波滑蜥,雄性和雌性个体的最大体长分别为47.4 mm和46.6 mm。雌雄两性在体长和头宽上没有差异,而在腹长和头长上差异显著,雄性有较大的头长,雌性有较大的腹长。宁波滑蜥年产单窝卵。窝卵数和窝卵重与雌体体长及腹长呈正相关,卵重与雌体体长无相关性。窝卵数及卵重的变异系数分别为0.20和0.12。卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数来增加繁殖输出。这些结果表明,宁波滑蜥是雌雄个体大小同形的两性异形模式,性选择使得雄性有着较大的头长,以具有较高的交配成功率,生育力选择使得雌性有着较大的腹长,以具有较大的生育力和繁殖输出。     

Random size‐assortative mating despite size‐dependent fecundity in a Neotropical amphibian with explosive reproduction

Sexual selection theory predicts that, when body size is correlated with fecundity, there should be fitness advantages for mate choice of the largest females. Moreover, because larger males are expected to monopolise the largest females, this should result in an assortative mating based on body size. Although such patterns could be expected in both explosive and prolonged breeders, non‐assortative mating should be more widespread in species under time constraints. However, patterns of sexual selection are largely unexplored in explosive breeding species, and contrasting patterns have been found previously. We expect that the active choice of partners may be particularly risky when the time period during which sexual partners are available is severely limited. Therefore, to avoid missing an entire reproductive act, males and females should pair irrespective of traits, such as body size. We tested this hypothesis by investigating the mating patterns of the Pacific horned toad, Ceratophrys stolzmanni, a short‐lived fossorial species inhabiting Neotropical dry forests. This species is particularly adequate to test our prediction because it reproduces explosively over the course of a single night per year. Although the number of eggs laid was proportional to the size of females, and individuals of both sexes showed variation in body size, there was no assortative mating based either on size, body condition or age of mates. Egg size was not influenced by either female size or clutch size. The larger body size of females compared to males is likely due to fecundity selection, that is, the selective pressure that enhances reproductive output. Although we cannot dismiss the possibility that individuals could select their partners based on other criteria than those related to size or age, the results fit well our prediction, showing that the explosive breeding makes improbable an active choice of partners in both sexes and therefore favours a random mating pattern.     

What is driving male mate preference evolution in Jamaican field crickets?

Male mating preferences are often a neglected aspect of studies on sexual selection. Male mating preferences may evolve if they provide males with direct‐fitness benefits such as increased opportunity to fertilize more eggs or indirect‐fitness benefits such as enhanced offspring survival. We tested these ideas using Jamaican field crickets, Gryllus assimilis, previously shown to exhibit male mating preferences. We randomly mated males to either their preferred or non‐preferred potential mates and then asked whether mating treatment influenced egg oviposition or offspring viability. Preferred females were not significantly more fecund and did not produce more viable eggs or offspring than non‐preferred females. Male mate preferences were therefore inconsistent with both the direct‐ and indirect‐fitness benefits hypotheses under the conditions of our experiment. Our null results leave us with an open question about what is driving the evolution of mating preferences in male crickets. Future research should explore the whether the offspring of preferred females are more attractive, have stronger immune systems, and/or experience higher adult longevity.     

Sex, mutation and fitness: asymmetric costs and routes to recovery through compensatory evolution

A wild-type population of Drosophila melanogaster was used to assess the impact of a known deleterious mutation, nub(1), when it had (1) evolved for up to 180 generations with the mutation or (2) recently had the same mutant allele introgressed into it. Relative to this benchmark, we observed much stronger initial fitness depression in males (-74%) than in females (-38%) and also relatively greater fitness recovery by evolved males (+55%) than females (+17%). Experimental assays revealed amelioration in both juvenile and adult fitness and suggested that the greater relative recovery of male fitness was from gains through sexual selection. These evolutionary changes in male fertility depended on pairing with their coevolved mates for both mate choice and post-copulatory components of sexual selection. Without replication at the population level, these results are used to motivate a general hypothesis rather than definitively test it: Differences in reproductive optima may generally skew mutational effects towards the more strongly sexually-selected sex due to genic capture and condition dependence.     

An integrative view of sexual selection in Tribolium flour beetles

Sexual selection is a major force driving the evolution of diverse reproductive traits. This evolutionary process is based on individual reproductive advantages that arise either through intrasexual competition or through intersexual choice and conflict. While classical studies of sexual selection focused mainly on differences in male mating success, more recent work has focused on the differences in paternity share that may arise through sperm competition or cryptic female choice whenever females mate with multiple males. Thus, an integrative view of sexual selection needs to encompass processes that occur not only before copulation (pre-mating), but also during copulation (peri-mating), as well as after copulation (post-mating), all of which can generate differences in reproductive success. By encompassing mechanisms of sexual selection across all of these sequential reproductive stages this review takes an integrative approach to sexual selection in Tribolium flour beetles (Coleoptera: Tenebrionidae), a particularly well-studied and economically important model organism. Tribolium flour beetles colonize patchily distributed grain stores, and juvenile and adult stages share the same food resources. Adults are highly promiscuous and female reproduction is distributed across an adult lifespan lasting approximately 1 year. While Tribolium males produce an aggregation pheromone that attracts both sexes, there appears to be little pre-mating discrimination among potential mates by either sex. However, recent work has revealed several peri-mating and post-mating mechanisms that determine how offspring paternity is apportioned among a female's mates. During mating, Tribolium females reject spermatophore transfer and limit sperm numbers transferred by males with low phenotypic quality. Although there is some conflicting evidence, male copulatory leg-rubbing appears to be associated with overcoming female resistance to insemination and does not influence a male's subsequent paternity share. Evidence suggests that Tribolium beetles have several possible post-mating mechanisms that they may use to bias paternity. Male sperm precedence has been extensively studied in Tribolium spp. and the related Tenebrio molitor, and several factors influencing male paternity share among a female's progeny have been identified. These include oviposition time, inter-mating interval, male strain/genotype, the mating regimen of a male's mother, male starvation, and tapeworm infection. Females exert muscular control over sperm storage, although there is no evidence to date that females use this to differentiate among mates. Females could also influence offspring paternity by re-mating with additional males, and T. castaneum females more readily accept spermatophores when they are re-mating with more attractive males. Additional work is needed to examine the possible roles played by both male and female accessory gland products in determining male paternity share. Sexual selection during pre-mating episodes may be reinforced or counteracted by peri- and post-copulatory selection, and antagonistic coevolution between the sexes may be played out across reproductive stages. In Tribolium, males' olfactory attractiveness is positively correlated with both insemination success and paternity share, suggesting consistent selection across different reproductive stages. Similar studies across sequential reproductive stages are needed in other taxa to provide a more integrative view of sexual selection.     

Sexual selection and its evolutionary consequences in female animals

For sexual selection to act on a given sex, there must exist variation in the reproductive success of that sex as a result of differential access to mates or fertilisations. The mechanisms and consequences of sexual selection acting on male animals are well documented, but research on sexual selection acting on females has only recently received attention. Controversy still exists over whether sexual selection acts on females in the traditional sense, and over whether to modify the existing definition of sexual selection (to include resource competition) or to invoke alternative mechanisms (usually social selection) to explain selection acting on females in connection with reproduction. However, substantial evidence exists of females bearing characters or exhibiting behaviours that result in differential reproductive success that are analogous to those attributed to sexual selection in males. Here we summarise the literature and provide substantial evidence of female intrasexual competition for access to mates, female intersexual signalling to potential mates, and postcopulatory mechanisms such as competition between eggs for access to sperm and cryptic male allocation. Our review makes clear that sexual selection acts on females and males in similar ways but sometimes to differing extents: the ceiling for the elaboration of costly traits may be lower in females than in males. We predict that current and future research on female sexual selection will provide increasing support for the parsimony and utility of the existing definition of sexual selection.     

Paternity estimated by isozyme variation in the marine sculpinAlcichthys alcicornis (Pisces: Cottidae) exhibiting copulation and paternal care

InAlcichthys alcicornis, a sculpin copulating in conjunction with paternal care, the genetic relation between males and egg masses they guard was investigated using polymorphism of an isozyme. At least 3 out of the 10 egg masses investigated were not genetically related with the guarding males. In these 3 cases, the female parents must have copulated with males other than the guarding males. It was hypothesized that males take care of non-kin eggs for the opportunity to fertilize the next clutch females will produce.