Why do men marry and why do they stray?

Humans are quite unusual compared to other great apes in that reproduction typically takes place within long-term, iteroparous pairings--social arrangements that have been culturally reified as the institution of marriage. With respect to male behaviour, explanations of marriage fall into two major schools of thought. One holds that marriage facilitates a sexual division of labour and paternal investment, both important to the rearing of offspring that are born helpless and remain dependent for remarkably long periods (provisioning model). And the other suggests that the main benefits which men receive from entering into marriage derive from monopolizing access to women's fertility (mating effort model). In this paper, we explore extramarital sexual relationships and the conditions under which they occur as a means of testing predictions derived from these two models. Using data on men's extramarital sexual relationships among Tsimane forager-horticulturists in lowland Bolivia, we tested whether infidelity was more common when men had less of an opportunity to invest in their children or when they risked losing less fertility. We found that Tsimane men appear to be biasing the timing of their affairs to when they are younger and have fewer children, supporting the provisioning model.     

The male's dilemma: Increased offspring production is more paternity to steal

Summary Large potential effects of male care on the number of offspring females successfully raise are not sufficient to select for caring males because of the pervasive importance of mating competition. Males face a version of the social dilemma, in which increased production increases the pay-off for theft. Models of the allocation of male effort partitioned between caring for babies and competing for paternity show that the optimal allocation to care is very low under a wide range of conditions. Like sex allocation where the alternatives are male versus female function or sons versus daughters, the pay-offs to one alternative are always strongly frequency dependent. Because that alternative (male function, sons, male mating effort) pays so well when rare, it cannot remain rare under most conditions. Here we consider the consequences of partitioning mating effort into mate guarding and all other forms of mating conflict. If a male gets all his partner's conceptions while guarding, gaining them at a constant rate, there are two possible regions of stability. The evolutionarily stable strategy (ESS) depends on a parameter scaling the decisiveness of (non-guarding) mating conflict. When marginal returns from conflict decrease with scale, almost all effort goes into guarding. When marginal returns increase, the ESS devotes all effort to mating. Even when the potential effect of care is large, male equilibrium strategies allocate little effort to it. We also report the results of computer simulations showing that care increases if gains from guarding saturate quickly, so that a male is assured of the paternity of most of his partner's offspring with little guarding, and consequently the pool of unguarded conceptions open to competion shrinks sharply. But even when the male's dilemma is very much reduced, it still substantially limits the allocation to care. The results of both computer simulations and mathematical analysis converge with other lines of evidence that mating has much stronger effects than parenting in shaping male strategies.     

Parental care in captive siamangs (Hylobates syndactylus)

Siamangs exhibit paternal care to the extent that the male of a monogamous unit carries its infant offspring beginning late in the infant's first year of life. Field studies have documented this but without behavioral detail. It has been hypothesized that the transfer is facilitated by a desertion of the infant by the female. An infant siamang born in captivity at the Washington Park Zoo, Portland, Oregon, was observed through its first year of life. The infant transfer to the male was documented in captivity and was associated with a high rate of infant-initiated contact with the male and a high rate of infant retrieval by the female. The study concludes that the infant probably played a key role in facilitating the transfer to the male.     

Male brood care without paternity increases mating success

We investigate under which conditions we can expect the evolutionof costly male care for unrelated offspring, when the benefitof such care is in the form of increased mating success. Thisapplies to male helping behavior that cannot be explained aspaternal care because the male's own offspring does not benefitfrom his behavior. Our model shows that caring for others' offspringcan be a stable strategy for males, if a male that does not"help" loses mating opportunities, for example if females discriminateagainst non-helping males as mating partners. This is possiblewhen females are polyandrous. Increasing population densitydecreases the parameter region where male care is stable. Malecare is also more likely to be stable when male mortality rateis higher than that of females. We discuss the results withspecial reference to the golden egg bug Phyllomorpha laciniata,where females lay eggs on conspecifics, often on males beforemating. Males therefore carry mostly unrelated eggs. We investigatehow oviposition rate and female mating rate influences whenegg carrying is an evolutionary stable strategy. We concludethat in the golden egg bug, male egg carrying could be explainedas a form of mating investment.     

Male Participation in Nest Building in the Dung Beetle Scarabaeus catenatus (Coleoptera: Scarabaeidae): Mating Effort Versus Paternal Effort

The dung beetle, Scarabaeus catenatus, shows not only the rolling but also the tunneling tactic for nest building with bisexual cooperation. Sex roles, however, differed between the tactics. In rolling, the male took the initiative like that of ball-roller species: he rolled a dung ball away and buried it. In tunneling, in contrast, the male usually had a secondary role like that of tunnelers: he was less active in burrow excavation and provisioning. Regardless of the tactics, male participation did not increase female reproductive output measured by the number or size of brood balls in the field, but seemed to function as mate guarding against conspecific males. This suggests that, in both tactics, the male S. catenatus invests primarily in mating effort compared with paternal effort. The relative importance of mating effort in male participation seems to hold true in other dung beetles, irrespective of whether they are ball-roller or tunneler species. In addition, the male mating strategy of S. catenatus is compared with that of other ball-rollers.     

SPERMATOPHORE SIZE IN BUSHCRICKETS: COMPARATIVE EVIDENCE FOR NUPTIAL GIFTS AS A SPERM PROTECTION DEVICE

During courtship and copulation, males of many insect species provide the female with a nuptial gift of a prey item or synthesized material. These gifts may be explained as a form of paternal investment by increasing female reproductive output, or in terms of mating effort by increasing male fertilization success. These explanations, while not mutually exclusive, are controversial. While experimental studies examine the maintenance of nuptial gifts in single species, comparative studies are required to indicate more general evolutionary trends. Male bushcrickets provide females with a nuptial gift, a spermatophylax, which is transferred to females at mating along with the sperm-containing ampulla. Analysis of comparative data of 28 species of bushcrickets (Orthoptera: Tettigoniidae), reveals that male spermatophore size (spermatophylax and ampulla weight) is positively correlated with female refractory period, which, in turn, correlates with male fertilization success. Moreover, gift size (the spermatophylax) covaries with ejaculate size (the ampulla), which is consistent with the hypothesis that it serves as a sperm protection device. In contrast, there is no significant correlation between any measure of female fecundity and male spermatophylax size. This indicates that the variation in spermatophore size among bushcrickets is better explained by a mating-effort function than a paternal investment function.     

The function of nuptial feeding in insects: a review of empirical studies

Nuptial feeding encompasses any form of nutrient transfer from the male to the female during or directly after courtship and/or copulation. In insects, nuptial gifts may take the form of food captured or collected by the male, parts, or even the whole of the male's body, or glandular products of the male such as salivary secretions, external glandular secretions, the spermatophore and substances in the ejaculate. Over the past decade, there has been considerable debate over the current function of nuptial feeding in insects. This debate has centred on the issue of whether nuptial gifts function as paternal investment (i.e. function to increase the fitness and/or number of the gift-giving male's own offspring) or as mating effort (i.e. function to attract females, facilitate coupling, and/or to maximize ejaculate transfer), although the two hypotheses are not mutually exclusive. In the present article, evidence for the potential of nuptial gifts to function as either paternal investment, mating effort, or both is reviewed for each form of nuptial feeding in each insect taxon for which sufficient data are available. Empirical evidence suggests that many diverse forms of nuptial feeding in different insect taxa function, at least in part, as mating effort. For example, nuptial prey and salivary masses in the Mecoptera, regurgitated food in Drosophila (Diptera), hind-wing feeding in Cyphoderris (Orthoptera) and the secretion of the male's cephalic gland in Neopyrochroa (Coleoptera) and Zorotypus (Zoraptera) appear to function to entice females to copulate and/or to facilitate coupling. Nuptial prey and salivary masses in the Mecoptera also appear to function to maximize ejaculate transfer (which is also a form of mating effort), as do nuptial prey in Empis (Diptera), external glandular secretions in Oecanthus and Allonemobius (Orthoptera) and the spermatophylax in gryllids and tettigoniids (Orthoptera). Large spermatophores in, for example, the Lepidoptera and Coleoptera, also appear to be maintained by selection on the male to maximize ejaculate transfer and thereby counter the effects of sperm competition. In contrast to the large amount of evidence in support of the mating effort hypothesis, there is a relative lack of good evidence to support the paternal investment hypothesis. Certain studies have demonstrated an increase in the weight and/or number of eggs laid as a result of the receipt of larger gifts, or a greater number of gifts, in tettigoniids, gryllids, acridids, mantids, bruchid beetles, drosophilids and lepidopterans. However, virtually all of these studies (with the possible exception of studies of the spermatophylax in tettigoniids) have failed to control adequately for hormonal substances in the ejaculate that are known to affect female reproductive output. Furthermore, in at least four tettigoniids (but not in the case of two species), three lepidopterans, a drosophilid and probably also bruchid beetles and bittacids, evidence suggests that the male has a low probability of fertilising the eggs that stand to benefit from his nuptial gift nutrients. Therefore, the hypothesis that paternal investment might account for the function of nuptial gifts in general is not supported.