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The Fisher-inspired, arbitrary intersexual selection models of Lande (1981) and Kirkpatrick (1982), including both stable and unstable equilibrium conditions, provide the appropriate null model for the evolution of traits and preferences by intersexual selection. Like the Hardy-Weinberg equilibrium, the Lande-Kirkpatrick (LK) mechanism arises as an intrinsic consequence of genetic variation in trait and preference in the absence of other evolutionary forces. The LK mechanism is equivalent to other intersexual selection mechanisms in the absence of additional selection on preference and with additional trait-viability and preference-viability correlations equal to zero. The LK null model predicts the evolution of arbitrary display traits that are neither honest nor dishonest, indicate nothing other than mating availability, and lack any meaning or design other than their potential to correspond to mating preferences. The current standard for demonstrating an arbitrary trait is impossible to meet because it requires proof of the null hypothesis. The LK null model makes distinct predictions about the evolvability of traits and preferences. Examples of recent intersexual selection research document the confirmationist pitfalls of lacking a null model. Incorporation of the LK null into intersexual selection will contribute to serious examination of the extent to which natural selection on preferences shapes signals.

Females of many species show strong directional preferences for costly male signals, which serve as indicators of male quality. Leader preference, a directional preference for males producing their signals ahead of their neighbors', is unusual in that there are no inherent costs to producing the attractive signal. Nonetheless, interacting males compete to produce leading signals, with the winner gaining a large fitness advantage. Here, we test whether competition to produce leading calls is sufficient to make leadership an honest indicator of male quality in the absence of costs in Neoconocephalus katydids. We include one species with leader preference and two without to determine whether any correlations between signal and quality arose before or after the evolution of leader preference. We found no correlations between leadership and male quality in any species tested, suggesting that the production of leading calls does not serve as an honest indicator of quality. This is likely the ancestral state for this system. The ability to produce leading calls in N ensiger, the species with leader preference, was not correlated with costly call traits. However, the ability to produce leading calls was highly repeatable, allowing for the possibility that it encodes meaningful information about males. The inter-individual variation in the ability to produce leading calls is likely caused by neural properties within the central pattern generator; this variation is likely due to both genetic and environmental factors. Though we found no evidence that females gain a fitness benefit by mating with leading males, we cannot exclude the possibility without tests of female fitness.

1. The indicator theory of sexual selection suggests that mating displays honestly signal aspects of fitness. While rarely studied, kinematic (locomotor) performance is an excellent candidate for an honest indicator, as mating displays of many animals include rapid or extended locomotion that may be physiologically correlated with performance traits that impact survival. 2. We investigate the indicator value of display locomotion of wild-caught male guppies, Poecilia reticulata, by examining relationships between mating display kinematic traits, anti-predator kinematic traits, and survival during a subsequent staged encounter with a natural predator, the pike cichlid Crenicichla alta. 3. We first compared guppy display kinematics with subsequent survival, and found that display body angle and angular speed positively predicted survival. We next compared anti-predator kinematic and tactical traits with survival, to identify traits that might link mating displays to survival. We measured anti-predator traits in two tests, first in response to a standardized stimulus (fast start test), and second in response to the live predator (encounter test). Guppy fast start speed and encounter speed, time in refuges, and approach distance (response distance) all positively predicted survival, while encounter swim duration negatively predicted survival. These data provided our final hypothesis, that these particular anti-predator traits would be correlated with mating display kinematics. However, we detected only one of eight predicted correlations, a negative relationship between display body angle and encounter swim duration that may reflect an energy trade-off. 4. We conclude that courtship locomotor performance can be an honest survival indicator in guppies, and that the mechanism linking courtship to survival merits further study. These results suggest that courtship locomotion may contribute to viability impacts on the evolution of animal mate choice, and support others in suggesting that these traits may reward greater attention in sexual and natural selection studies.

Male horn length in some horned beetles shows a sigmoidal relationship with body size. This has often been considered as the reflection of alternative reproductive tactics of males based on body size. Large males should possess long horns to acquire females through fights with other males using their horns, whereas small males do not require long horns because they usually avoid intermale fights and adopt alternative tactics such as sneaking. This may lead to a prediction that horn length is a reliable indicator of the fighting ability of the male. We examined the effects of both male horn length and body size of Allomyrina dichotoma on the outcomes of escalated fights. Results indicate that male horn length was more important than body size in predicting the outcomes of fight, and this may support the hypothesis that the evolution of the horn dimorphism in male horned beetles is the result of different reproductive tactics.[PUBLICATION ABSTRACT]

The dynamics of sexual selection (competition for mates and mate choice) and social selection (competition for resources other than mates) are overviewed with an emphasis on the evolutionary elaboration and condition-dependent expression of associated physical, behavioral, and brain and cognitive traits. These traits are more variable than naturally selected traits, and their expression is more easily disrupted by inbreeding, exposure to parasites, poor nutrition, social stress, and exposure to man-made toxins. Multiple examples across a wide range of species are provided to illustrate this condition-dependence and to lay the foundation for understanding human vulnerability.

The sexual swellings of female primates have generated a great deal of interest in evolutionary biology. Two hypotheses recently proposed to elucidate their functional significance argue that maximal swelling size advertises either female fertility within a cycle or female quality across cycles. Published evidence favours the first hypothesis, and further indicates that larger swellings advertise higher fertility between cycles. If so, a male preference for large swellings might evolve, driving females to use swellings as quality indicators, as proposed by the second hypothesis. In this paper, we explore this possibility using a combination of empirical field data and mathematical modelling. We first test and find support for three key predictions of the female-quality hypothesis in wild chacma baboons (Papio ursinus): (i) inter-individual differences in swelling size are maintained across consecutive cycles, (ii) females in better condition have larger swellings and higher reproductive success, and (iii) males preferentially choose females with large swellings. We then develop an individual-based simulation model that indicates that females producing larger swellings can achieve higher mating success even when female-female competition is low and within-female variance in the trait is high. Taken together, our findings show that once sexual swellings have evolved as fertility signals, they might, in certain socio-sexual systems, be further selected to act as quality signals. These results, by reconciling two hypotheses, help to clarify the processes underlying sexual swelling evolution. More generally, our findings suggest that mate choice for direct benefits (fertility) can lead to indirect benefits (good genes).

Song is used as a signal in sexual selection in a wide range of taxa. In birds, males of many species continue to sing after pair formation. It has been suggested that a high song output after pair formation might serve to attract extrapair females and to minimise their own partner's interest in extra-pair copulations. A non-exclusive alternative function that has received only scant attention is that the amount of song might stimulate the own female's investment into eggs in a quantitative way. We address these hypotheses in a captive population of zebra finches, Taeniopygia guttata, by relating male undirected song output (i.e. non-courtship song) to male egg siring success and female reproductive investment in two different set-ups. When allowed to breed in aviaries, males with the highest song output were no more attractive than others to females in an analysis of 4,294 extra-pair courtships involving 164 different males, and they also did not sire more offspring (both trends were against the expectation). When breeding in cages with two different partners subsequently, females produced larger eggs with more orange yolks when paired to a male with a high song output. These findings suggest that singing activity in paired zebra finch males might primarily function to stimulate the partner and not to attract extra-pair females.

In various models of sexual selection mediated by the viability indicator (\"good genes\") mechanism, a sexually selected trait will truly reflect male quality if its expression is costly for the male. However, in long-lived species, the expression of a trait often increases with age while the genotype of the male remains unchanged. This fact may obscure the indicator mechanism. Hitherto, game theory models of honesty in sexual advertisement have not taken life-history effects into account, whereas life-history models of reproductive effort have only seldom considered the dependence of mating success on the actions of other individuals. Here, the two approaches are combined, and I examine whether honesty is maintained if males can divide their advertisement effort over their lifetime. The model shows that an increase in the expression of the sexually selected trait over several years is an evolutionarily stable strategy (ESS) under a wide range of situations, so that a correlated preference for old age can emerge through a viability indicator mechanism. Honesty in the strict sense is not preserved: an optimally behaving low-quality male will in some cases advertise more than a high-quality male of equal age, to the extent that the strongest advertisement found in the population can be associated with a low-quality male. Due to life-history trade-offs, however, honesty in an average sense holds true over the lifetime of individuals: \"cheater\" age classes will remain small enough, that a female will obtain a higher expected mate quality if she trusts in the trait as an indicator of viability.