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Male contests for access to receptive females are thought to have selected for the larger male body size and conspicuous weaponry frequently observed in mammalian species. However, when females copulate with multiple males within an oestrus, male reproductive success is a function of both pre-and postcopulatory strategies. The relative importance of these overt and covert forms of sexual competition has rarely been assessed in wild populations. The Soay sheep mating system is characterized by male contests for mating opportunities and high female promiscuity. We find that greater horn length, body size and good condition each independently influence a male's ability to monopolize receptive females. For males with large horns at least, this behavioural success translates into greater siring success. Consistent with sperm-competition theory, we also find that larger testes are independently associated with both higher copulation rates and increased siring success. This advantage of larger testes emerges, and strengthens, as the number of oestrous females increases, as dominant males can no longer control access to them all. Our results thus provide direct quantitative evidence that male reproductive success in wild populations of mammals is dependent upon the relative magnitude of both overt contest competition and covert sperm competition.

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Evolutionary theories of ageing posit that increased reproductive investment occurs at the expense of physiological declines in later life. Males typically invest heavily in costly sexual ornaments and behaviour, but evidence that the expression of these traits can cause senescence is lacking. Long-lived houbara bustards (Chlamydotis undulata) engage in extravagant sexual displays to attract mates and here we show that males investing most in these displays experience a rapid senescent deterioration of spermatogenic function at a younger age. This effect is sufficiently large that the expected links between male 'showiness' and fertility reverse in later life, despite 'showy' males continuing to display at near maximal levels. We show that our results cannot be explained by the selective disappearance of competitive phenotypes and that they are instead consistent with an early vs. late life trade-off in male reproductive competence, highlighting the potential significance of sexual selection in explaining rates of ageing.

Background: Sleep is a biological enigma. Despite occupying much of an animal's life, and having been scrutinized by numerous experimental studies, there is still no consensus on its function. Similarly, no hypothesis has yet explained why species have evolved such marked variation in their sleep requirements (from 3 to 20 hours a day in mammals). One intriguing but untested idea is that sleep has evolved by playing an important role in protecting animals from parasitic infection. This theory stems, in part, from clinical observations of intimate physiological links between sleep and the immune system. Here, we test this hypothesis by conducting comparative analyses of mammalian sleep, immune system parameters, and parasitism.

Summary1. Testosterone (T) is a key mediator in the expression of numerous morphological and behavioural traits in mammals, but the factors underlying individual variation in circulating T levels are poorly understood. 2. The intimate structural integration of sperm and T production within the testes, alongside the dependency of sperm production on high levels of T, suggests that T requirements for spermatogenesis could be an important driver of individual differences in T. 3. To test this hypothesis, we examine how male capacity for sperm production (as indicated by their testes size) is associated with T levels in a feral population of Soay sheep, resident on St. Kilda, Scotland, during their rutting season. 4. We found a strong positive relationship between an individual's testes size (as measured before their seasonal enlargement) and the levels of circulating T during their rut, suggesting that T requirements for spermatogenesis has a prominent influence on the production of this androgen. 5. In contrast, body condition and competitive ability did not independently predict T levels, findings that are inconsistent with conventional 'condition-dependent' and 'challenge' hypotheses of T production. 6. This influence of male's capacity for sperm production on T appeared to be substantial enough to be biologically relevant, as testes size also predicted male aggression and mate-seeking behaviour. 7. Our results suggest that a male's inherent capacity for sperm and T production is tightly phenotypically integrated, with potential consequences for a wide range of other T-mediated reproductive traits.

In most animal species, males are predicted to compete for reproductive opportunities, while females are expected to choose between potential mates. However, when males' rate of reproduction is constrained, or females vary widely in 'quality', male mate choice is also predicted to occur. Such conditions exist in the promiscuous mating system of feral Soay sheep on St Kilda, Scotland, where a highly synchronized mating season, intense sperm competition and limitations on sperm production constrain males' potential reproductive rate, and females vary substantially in their ability to produce successful offspring. We show that, consistent with predictions, competitive rams focus their mating activity and siring success towards heavier females with higher inclusive fitness. To our knowledge, this is the first time that male mate choice has been identified and shown to lead to assortative patterns of parentage in a natural mammalian system, and occurs despite fierce male-male competition for mates. An additional consequence of assortative mating in this population is that lighter females experience a series of unstable consorts with less adept rams, and hence are mated by a greater number of males during their oestrus. We have thus also identified a novel male-driven mechanism that generates variation in female promiscuity, which suggests that the high levels of female promiscuity in this system are not part of an adaptive female tactic to intensify post-copulatory competition between males.

Understanding whether the sperm of older males has a diminished capacity to produce successful offspring is a key challenge in evolutionary biology. We investigate this issue using 10 years of reproductive data on captive long-lived houbara bustards (Chlamydotis undulata), where the use of artificial insemination techniques means parents can only influence offspring quality via their gametes. Here we show that paternal aging reduces both the likelihood that eggs hatch and the rate at which chicks grow, with older males producing the lightest offspring after the first month. Surprisingly, this cost of paternal aging on offspring development is of a similar scale to that associated with maternal aging. Fitting with predictions on germline aging, the sperm of immature males produce the fastest growing offspring. Our findings thus indicate that any good genes benefit that might be offered by older ‘proven' males will be eroded by aging of their germline DNA.