Why do good hunters have higher reproductive success?

Anecdotal evidence from many hunter-gatherer societies suggests that successful hunters experience higher prestige and greater reproductive success. Detailed quantitative data on these patterns are now available for five widely dispersed cases (Ache, Hadza, !Kung, Lamalera, and Meriam) and indicate that better hunters exhibit higher age-corrected reproductive success than other men in their social group. Leading explanations to account for this pattern are: (1) direct provisioning of hunters’ wives and offspring, (2) dyadic reciprocity, (3) indirect reciprocity, (4) costly signaling, and (5) phenotypic correlation. I examine the qualitative and quantitative evidence bearing on these explanations and conclude that although none can be definitively rejected, extensive and apparently unconditional sharing of large game somewhat weakens the first three explanations. The costly signaling explanation has support in some cases, although the exact nature of the benefits gained from mating or allying with or deferring to better hunters needs further study. For comments on earlier drafts, I thank Monique Borgerhoff Mulder, Mike Gurven, Ray Hames, Kristen Hawkes, Kim Hill, Robert Kelly, Frank Marlowe, John Patton, and Polly Wiessner. Rebecca Bliege Bird and Douglas W. Bird invited me to collaborate in the Meriam research and (along with Del Passi of Mer) collected the data on Meriam demography. Geoff Kushnick and Matt Wimmer ably assisted with coding and statistical analysis of these data. Eric Alden Smith (PhD 1980, Cornell University) is a professor of anthropology at the University of Washington, Seattle. His research interests include the links between production and reproduction, the ecology and evolution of collective action, and politics in small-scale societies. He has conducted fieldwork among Inuit on Hudson Bay, Meriam in Torres Strait, and Mardu Aborigines in the Australian Western Desert.     

Greater opportunities for sexual selection in male than in female obligate brood parasitic birds

Females are expected to have evolved to be more discriminatory in mate choice than males as a result of greater reproductive investment into larger gametes (eggs vs. sperm). In turn, males are predicted to be more promiscuous than females, showing both a larger variance in the number of mates and a greater increase in reproductive success with more mates, yielding more intense sexual selection on males vs. females (Bateman's Paradigm). However, sex differences in costly parental care strategies can either reinforce or counteract the initial asymmetry in reproductive investment, which may be one cause for some studies failing to conform with predictions of Bateman's Paradigm. For example, in many bird species with small female‐biased initial investment but extensive biparental care, both sexes should be subject to similar strengths of sexual selection because males and females are similarly restricted in their ability to pursue additional mates. Unlike 99% of avian species, however, obligate brood parasitic birds lack any parental care in either sex, predicting a conformation to Bateman's Paradigm. Here we use microsatellite genotyping to demonstrate that in brood parasitic brown‐headed cowbirds (Molothrus ater), per capita annual reproductive success increases with the number of mates in males, but not in females. Furthermore, also as predicted, the variance of the number of mates and offspring is greater in males than in females. Thus, contrary to previous findings in this species, our results conform to predictions of the Bateman's Paradigm for taxa without parental care.     

The hierarchy of virtue: mutualism, altruism and signaling in Martu women's cooperative hunting

Cooperative hunting is often assumed to be mutualistic, maintained through returns to scale, where, by working together, foragers can gain higher per capita return rates or harvest sizes than they can by hunting alone. We test this hypothesis among Martu hunters and find that cooperation only provides increased returns to poorer hunters while disadvantaging better hunters. Even so, better hunters still cooperate as frequently as poorer hunters. We ask whether better hunters are advantaged in secondary sharing distributions or whether they bias their partner choice to kin or household members. We find that better hunters are not more likely to pair up with kin and they do not gain consumption benefits from acquiring more. They share a greater proportion of their harvest than poorer hunters: no matter how much one produces — better hunter, worse hunter, cooperator, solitary hunter — all eat the same amount in the end. Such a result suggests the hypothesis that cooperation might be a costly signal of commitment to the public interest on the part of better hunters, which generates trust among camp members and facilitates strong social networks, particularly among women, who cooperate more than men. While some foragers may benefit through cooperation from returns to scale or risk reduction, others may benefit more through signaling commitment and generating trust.     

Female competition and its evolutionary consequences in mammals

Following Darwin's original insights regarding sexual selection, studies of intrasexual competition have mainly focused on male competition for mates; by contrast, female reproductive competition has received less attention. Here, we review evidence that female mammals compete for both resources and mates in order to secure reproductive benefits. We describe how females compete for resources such as food, nest sites, and protection by means of dominance relationships, territoriality and inter‐group aggression, and by inhibiting the reproduction of other females. We also describe evidence that female mammals compete for mates and consider the ultimate causes of such behaviour, including competition for access to resources provided by mates, sperm limitation and prevention of future resource competition. Our review reveals female competition to be a potentially widespread and significant evolutionary selection pressure among mammals, particularly competition for resources among social species for which most evidence is currently available. We report that female competition is associated with many diverse adaptations, from overtly aggressive behaviour, weaponry, and conspicuous sexual signals to subtle and often complex social behaviour involving olfactory signalling, alliance formation, altruism and spite, and even cases where individuals appear to inhibit their own reproduction. Overall, despite some obvious parallels with male phenotypic traits favoured under sexual selection, it appears that fundamental differences in the reproductive strategies of the sexes (ultimately related to parental investment) commonly lead to contrasting competitive goals and adaptations. Because female adaptations for intrasexual competition are often less conspicuous than those of males, they are generally more challenging to study. In particular, since females often employ competitive strategies that directly influence not only the number but also the quality (survival and reproductive success) of their own offspring, as well as the relative reproductive success of others, a multigenerational view ideally is required to quantify the full extent of variation in female fitness resulting from intrasexual competition. Nonetheless, current evidence indicates that the reproductive success of female mammals can also be highly variable over shorter time scales, with significant reproductive skew related to competitive ability. Whether we choose to describe the outcome of female reproductive competition (competition for mates, for mates controlling resources, or for resources per se) as sexual selection depends on how sexual selection is defined. Considering sexual selection strictly as resulting from differential mating or fertilisation success, the role of female competition for the sperm of preferred (or competitively successful) males appears particularly worthy of more detailed investigation. Broader definitions of sexual selection have recently been proposed to encompass the impact on reproduction of competition for resources other than mates. Although the merits of such definitions are a matter of ongoing debate, our review highlights that understanding the evolutionary causes and consequences of female reproductive competition indeed requires a broader perspective than has traditionally been assumed. We conclude that future research in this field offers much exciting potential to address new and fundamentally important questions relating to social and mating‐system evolution.     

Morphological and genetic predictors of parental care in the North American barn swallow Hirundo rustica erythrogaster

Sexually dimorphic traits often signal the fitness benefits an individual can provide to potential mates. In species with altricial young, these signals may also predict the level of parental care an individual is expected to provide to shared offspring. In this study, we tested three hypotheses that traditionally relate sexually dimorphic traits to parental care in two populations of North American barn swallows Hirundo rustica erythrogaster. The good parent hypothesis predicts a positive relationship between an individual's ornamentation and his or her care whereas the differential allocation (more care given by individuals when paired to high quality mates) and reproductive compensation (more care given by individuals when paired to low quality mates) hypotheses predict that an individual's level of parental investment is relative to the quality of their mate. Male and female North American barn swallows have colorful ventral feathers and elongated tail streamers, but there is evidence that ventral color, not tail streamer length, predicts measures of seasonal reproductive success. Accounting for the positive correlation between within‐pair feeding rates and other potentially confounding variables in all of our models, we found no support for the good parent hypothesis because in both males and females, traits shown to be under sexual selection did not predict feeding rates in either sex. However, our data reveal that male coloration, and not streamer length, predicted a female's provisioning rate to shared offspring (females fed more when paired with darker individuals) in two separate populations, supporting the differential allocation, but not the reproductive compensation hypothesis. Because genetic traits have also been shown to affect parental investment, we evaluated this variable as well and found that a male's paternity did not have significant effects on either male or female feeding rates. Overall, our results suggest that females do not pair with darker males in order to gain direct benefits in terms of his expected levels of parental care to shared offspring, but do themselves invest greater levels of care when paired to darker males. Further, our results are consistent with previous studies which suggest that ventral feather color, not streamer length, is a target of sexual selection in North American populations of barn swallow because females invested more in their offspring when paired to darker mates.     

How conception risk affects competition and cooperation with attractive women and men

We investigated competition and cooperation for resources across the menstrual cycle in the context of bargaining games. Although bargaining has been studied within an evolutionary framework, little attention has been paid specifically to the role of mating motives in economic behavior. To investigate how motives related to reproductive success affect bargaining, participants at high or low risk for conception or who were on oral contraceptives played ultimatum and dictator games with partners who varied in sex and facial attractiveness. In ultimatum games, women in the fertile phase were more competitive over resources with attractive women than with less attractive women. Intrasexual competition was not observed in dictator games. Women were more cooperative with attractive men than with less attractive men in both games, regardless of fertility status. Low fertility women were more cooperative with attractive members than with less attractive members of both sexes in both games. Results support the view that, during periods of high fertility, when women are most intrasexually competitive for mates, withholding resources from potential rivals would enable women to gain the means to enhance their attractiveness and weaken competitors' abilities to do the same at a time when relative advantages in appearance are most crucial to reproductive success. The lack of a fertility effect for cooperation with potential mates supports the view that displays of generosity accrue benefits for women across the cycle in their efforts to attract men who will invest in relationships.     

Criminal offending as part of an alternative reproductive strategy: investigating evolutionary hypotheses using Swedish total population data

Criminality is highly costly to victims and their relatives, but often also to offenders. From an evolutionary viewpoint, criminal behavior may persist despite adverse consequences by providing offenders with fitness benefits as part of a successful alternative mating strategy. Specifically, criminal behavior may have evolved as a reproductive strategy based on low parental investment reflected in low commitment in reproductive relationships. We linked data from nationwide total population registers in Sweden to test if criminality is associated with reproductive success. Further, we used several different measures related to monogamy to determine the relation between criminal behavior and alternative mating tactics. Convicted criminal offenders had more children than individuals never convicted of a criminal offense. Criminal offenders also had more reproductive partners, were less often married, more likely to get remarried if ever married, and had more often contracted a sexually transmitted disease than non-offenders. Importantly, the increased reproductive success of criminals was explained by a fertility increase from having children with several different partners. We conclude that criminality appears to be adaptive in a contemporary industrialized country, and that this association can be explained by antisocial behavior being part of an adaptive alternative reproductive strategy.     

Sensation seeking and men's face preferences

Findings from previous studies suggest that only men who are in good physical condition can afford to pursue high-risk activities and that men who engage in high-risk activities are considered particularly attractive by women. Here, we show that men's interest in high-sensation activities, a personality trait that is known to increase the likelihood of those individuals engaging in high-risk behaviors, is positively related to the strength of their preferences for femininity in women's faces (Studies 1–3) but is not related to the strength of their preferences for femininity in men's faces (Study 2). We discuss these findings as evidence for potentially adaptive condition-dependent mate preferences, whereby men who exhibit signals of high quality demonstrate particularly strong preferences for facial cues of reproductive and medical health in potential mates because they are more likely than lower-quality men to succeed in acquiring such partners.     

Adaptive female choice for middle-aged mates in a lekking sandfly

Most theoretical models of age-related mate choice predict that females should prefer older males because they have proven survival ability. An alternative view is that older males represent inferior mates because of negative genetic correlations between early and late fitness components, or because older males have traded off longevity against other fitness components, have accumulated deleterious germ-line mutations, or are less well adapted to current conditions than more recently born individuals. While numerous studies have reported female choice for older males, few have explicitly examined the fitness consequences of such a preference. We present evidence from a lekking sandfly, Lutzomyia longipalpis, showing that choosy females discriminate against older males and gain a fitness benefit from their choice. When permitted free choice from an aggregation consisting of males aged zero to two days (young), four to six days (middle-aged) and eight to ten days (old), females preferentially mated with middle-aged males, but all measures of female reproductive success were independent of male age. In contrast, when a second set of females was randomly assigned single virgin males of known age, the eggs of those paired to old mates exhibited lower hatching success than the eggs of females mated to young or middle-aged males. These results suggest that females avoid mating with older males because they represent poorer quality mates. Age-related differences in male quality may have a genetic basis, but could equally well arise through a phenotypic decline in sperm quality or sperm transfer ability with male age. The lack of evidence of female discrimination against older males from other studies may be because these did not explore the reproductive success of the full age range of males.     

Male chimpanzees exchange political support for mating opportunities

Male chimpanzees, Pan troglodytes, differ from males in most other mammalian taxa because they remain in their natal communities throughout their lives, form close bonds with one another, and cooperate in a range of activities [1]. However, males also compete fiercely for status within their groups [2,3], and high rank enhances male reproductive success [4,5]. Males rely partly on coalitions to achieve and maintain status [2,3,6-9], and shifts in male alliances can have dramatic political effects [2,3,6]. It is not known what benefits are obtained by low-ranking coalition partners. Here we report that the highest-ranking (alpha) male in one well-studied community of chimpanzees rewarded his allies by allowing them preferential access to mates.     

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.     

Divorce in the barn owl: securing a compatible or better mate entails the cost of re‐pairing with a less ornamented female mate

Two nonmutually exclusive hypotheses can explain why divorce is an adaptive strategy to improve reproductive success. Under the ‘better option hypothesis’, only one of the two partners initiates divorce to secure a higher‐quality partner and increases reproductive success after divorce. Under the ‘incompatibility hypothesis’, partners are incompatible and hence they may both increase reproductive success after divorce. In a long‐term study of the barn owl (Tyto alba), we address the question of whether one or the two partners derive fitness benefits by divorcing. Our results support the hypothesis that divorce is adaptive: after a poor reproductive season, at least one of the two divorcees increase breeding success up to the level of faithful pairs. By breeding more often together, faithful pairs improve coordination and thereby gain in their efficiency to produce successful fledglings. Males would divorce to obtain a compatible mate rather than a mate of higher quality: a heritable melanin‐based signal of female quality did not predict divorce (indicating that female absolute quality may not be the cause of divorce), but the new mate of divorced males was less melanic than their previous mate. This suggests that, at least for males, a cost of divorce may be to secure a lower‐quality but compatible mate. The better option hypothesis could not be formally rejected, as only one of the two divorcing partners commonly succeeded in obtaining a higher reproductive success after divorce. In conclusion, incompatible partners divorce to restore reproductive success, and by breeding more often together, faithful partners improve coordination.     

Is Divorce in Young Common Terns,Sterna hirundo,after Recruitment just a Question of Timing?

The probability of divorce in birds has been linked with age, breeding experience, reproductive output and synchrony in return. Here, we investigate the consequences of first breeding attempts in common terns for mating in the subsequent season. Nearly 20% of all first‐time breeders disappeared or skipped at least one season after recruitment. In 84 pairs, which consisted of at least one recruit and of which both partners returned to the colony, the divorce rate was 45%. We compared reproductive success, arrival dates, and asynchrony in arrival dates of pairs of the first breeding season against the second season, for both reunited and divorced pairs and males and females separately. First, in pairs of which both members came back to the colony, we found an increase of reproductive success most pronounced in males. In the second season reproductive success of divorced compared with reunited pairs was higher, as only divorced pairs significantly improved the number of fledglings, and again this relation was stronger in males. Secondly, females of reunited pairs arrived significantly earlier from the first to the second season and by far more days than their males. However, in divorced pairs former mates did not differ in the number of days they advance their arrivals. Finally, divorced males arrived on average 4 d earlier than their former mates, whereas divorced females arrived 5 d later compared with their former mates of the recruitment season. Contradictory to nearly all other divorce studies in birds so far, we found a clear fitness gain in divorced males. We suggest that the improvement in reproductive success of young males stems from a side‐effect of the birds’ quality and ability to reach the breeding site in appropriate time and earlier as potential competitors. In long‐lived bird species the heterogeneity among young individuals in the timing of arrival at the colony seems to explain why former recruit‐pairs reunite or split. For young males we suggest as best explanation of divorce that they profit from ‘pushing for an empty chair’, while females seem to profit from their choosiness and may actively decide between former and other mates.     

In the eye (and ears) of the beholder: Receiver psychology and human signal design

Although the study of signals has been part of human behavioral ecology since the field's inception,¹ only recently has signaling theory become important to the evolutionary study of human behavior and culture.² Signaling theory's rise to prominence has been propelled mainly by applications of costly signaling theory,³ which has shed light on a wide variety of human behaviors ranging from hunting⁴ to religion.^5,6 Costly signaling rests on the idea that wasteful but highly visible traits and behaviors can be explained as honest indicators of underlying qualities that are otherwise difficult to detect. For example, a laborious hunting technique may serve as a display of skill on the part of the hunter, who may then be favorably perceived by potential mates and allies.⁴ The costs of the activity ensure that the signal is honest, since unskilled hunters will not be able to perform as well. Despite the usefulness of this perspective, many such studies begin by documenting a costly behavior that is then explained with reference to costly signaling theory. Because such behaviors are easy to detect, they may be overemphasized in the literature.⁷ Moreover, costly signaling theory by itself can explain neither all signals nor all aspects of signal design. In this review, we argue that a focus on the role that the psychology of the intended receiver plays in signal design can expand the scope of signaling theory as a promising avenue to explain human behavior.     

Reproductive correlates of plumage coloration of female Mountain Bluebirds

Many studies have shown that the plumage coloration of male birds can act as an honest signal of quality, indicating benefits that a female could gain from pairing with a specific male. In some species, females also display ornamental plumage, but less is known about the function and potential adaptive significance of female coloration because most research has focused on male coloration. Male Mountain Bluebirds (Sialia currucoides) display full body, ultraviolet (UV)‐blue plumage, whereas female plumage is more subdued, with blue color focused on the rump, wing, and tail. During the 2011 and 2012 breeding seasons (May–July) near Kamloops, BC, Canada, we examined coloration of the rump and tail of female Mountain Bluebirds to determine if their plumage could act as an indicator of direct reproductive benefits (e.g., enhanced parental care or reproductive success) to potential mates. We found no relationship between female plumage coloration and either provisioning rate or fledging success. However, female coloration varied with age, with after‐second‐year (ASY) females having brighter, more UV‐blue tail feathers than second‐year (SY) females. In addition, ASY females with brighter, more UV‐blue tails had larger clutches. We also observed positive assortative mating by tarsus length. Because previous work with other species suggests that female body size may be a good predictor of breeding success, males could potentially benefit from pairing with larger females. However, reproductive success did not vary with female size in our study. Although our evidence that structural plumage coloration of female Mountain Bluebirds is a signal of direct reproductive benefits for males (e.g., higher reproductive success) is limited, our results (i.e., ASY females with brighter tails than SY females, and ASY females with brighter tails having larger clutches) do suggest the potential for sexual selection to act on female coloration.     

Mutual mate choice in crayfish: large body size is selected by both sexes, virginity by males only

Mate choice is often assumed to be a prerogative of females because of their putatively larger reproductive investment than males. However, recent evidence suggests that spermatogenesis is far from being limitless and that males show a high selectivity towards their mates, thus maximizing their reproductive success. We investigated mutual mate choice in the crayfish Procambarus clarkii through two experiments. The first experiment explored the effects of body size, chelar size and chelar symmetry and social status of a potential partner. In the second experiment, we asked whether this species can discriminate between partners of the same body size but with different mating status. We used a binary choice test paradigm, in which two 'targets' with opposing characteristics were simultaneously presented to a test animal, the 'chooser'. The results showed that P. clarkii males are more selective than expected. Similar to the other sex, they were significantly attracted by targets with large body sizes, but not by individuals with larger and symmetric chelae or with a dominant status. An inter-sexual difference was found in the second experiment, in which only males seemed to select virgin potential mates. The several adaptive explanations for these preferences, still under debate, are finally discussed.     

Effects of site fidelity and breeding performance on mate retention in a short‐lived passerine,the tree swallow Thachycineta bicolor

We studied the relationship among re‐mating, site fidelity and breeding performance in the tree swallow Tachycineta bicolor using 16 y of data on reproductive biology in a population breeding in nest boxes near Ithaca, New York. Of 217 pairs for which both members survived the non‐breeding season, 76% mated with a new partner and 24% reunited with their previous mate. Pairs did not increase their breeding success by breeding together for more than one breeding season. Males produced fewer fledglings after breeding with a new partner, but females neither increased nor decreased their success when breeding with a new mate. Females who bred with a new partner were younger than females that reunited with their previous mates, and they were more likely to move to a different nest box. Males that bred with a new mate were of similar age to males that reunited, and they did not move more often. The probability of breeding with a new partner was better predicted by female age than by previous breeding success, suggesting that re‐mating was not strongly affected by past breeding performance. Because younger females change breeding sites more frequently than do older females and females that mated with a new partner were younger than females that reunited with their previous mates, we suggest that the tendency of tree swallows to change partners between years is a by‐product of lower site fidelity of younger females rather than a strategy for increasing breeding success.     

Ejaculate economics: an experimental test in a moth

Increasing evidence shows that spermatogenesis is costly. As a consequence, males should optimize the use of their sperm to maximize their reproductive outputs in their lifetime. However, experimental evidence on this prediction is largely lacking. Here, we examine how a male moth Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) responds to the presence of rivals or additional mates and how such response influences his lifetime reproductive fitness. We show that when rival males are present around a copulating pair, the male ejaculates more sperm to win a sperm competition battle but in such an environment he inseminates fewer females, sires fewer offspring and lives shorter. The opposite is the case when additional females are present during copulation. These findings reveal that elevated reproductive expenditure owing to sperm competition intensity is made at the expense of longevity and future reproduction.     

Male and female mate choice affects offspring quality in a sex-role-reversed pipefish.

Where both sexes invest substantially in offspring, both females and males should discriminate between potential partners when choosing mates. The degree of choosiness should relate to the costs of choice and to the potential benefits to be gained. We measured offspring quality from experimentally staged matings with preferred and non-preferred partners in a sex-role-reversed pipefish, Syngnathus typhle L. Here, a substantial male investment in offspring results in a lower potential reproductive rate in males than in females, and access to males limits female reproductive success rather than vice versa. Thus, males are choosier than females and females compete more intensely over mates than do males. Broods from preferred matings were superior at escaping predation, when either males or females were allowed to choose a partner. However, only 'choosing' females benefited in terms of faster-growing offspring. Our results have important implications for mate-choice research: here we show that even the more competitive and less choosy sex may contribute significantly to sexual selection through mate choice.     

No reduction of female sexual receptivity following mating in a stalk‐eyed fly,Cyrtodiopsis dalmanni (Diptera: Diopsidae)

The level of female sexual receptivity is an important component of male and female reproductive success. In many insects, mating itself causes a sharp decline in female receptivity. This can be a direct result of the physical act of mating, or because of actions of sperm or seminal fluid proteins. The degree to which males can decrease female receptivity will directly affect their reproductive success, by altering the chance that their sperm will be used in fertilizations in the interval before the female mates again. In this study, we investigated the effect of mating on female receptivity in the sexually dimorphic stalk‐eyed fly, Cyrtodiopsis dalmanni. Our results showed no evidence for mating‐induced reductions in female receptivity. In addition, we found that matings with males that differed in eyespan did not cause differences in the level of female receptivity. There was also no evidence that females remated sooner when presented with large eyespan males. These results are surprising, given the indirect benefits that females gain from matings with large eyespan males. Finally we demonstrate that males do not appear to discriminate between females on the basis of female mating status.