Sexual Selection on male cuticular hydrocarbons via male–male competition and female choice

Traditional views of sexual selection assumed that male–male competition and female mate choice work in harmony, selecting upon the same traits in the same direction. However, we now know that this is not always the case and that these two mechanisms often impose conflicting selection on male sexual traits. Cuticular hydrocarbons (CHCs) have been shown to be linked to both social dominance and male attractiveness in several insect species. However, although several studies have estimated the strength and form of sexual selection imposed on male CHCs by female mate choice, none have established whether these chemical traits are also subject to sexual selection via male–male competition. Using a multivariate selection analysis, we estimate and compare sexual selection exerted by male–male competition and female mate choice on male CHC composition in the broad‐horned flour beetle Gnatocerus cornutus. We show that male–male competition exerts strong linear selection on both overall CHC abundance and body size in males, while female mate choice exerts a mixture of linear and nonlinear selection, targeting not just the overall amount of CHCs expressed but the relative abundance of specific hydrocarbons as well. We discuss the potential implications of this antagonistic selection with regard to male reproductive success.     

Sexual selection in a lekking bird: the relative opportunity for selection by female choice and male competition

Leks are classic models for studies of sexual selection due to extreme variance in male reproductive success, but the relative influence of intrasexual competition and female mate choice in creating this skew is debatable. In the lekking lance-tailed manakin (Chiroxiphia lanceolata), these selective episodes are temporally separated into intrasexual competition for alpha status and female mate choice among alpha males that rarely interact. Variance in reproductive success between status classes of adult males (alpha versus non-alpha) can therefore be attributed to male-male competition whereas that within status largely reflects female mate choice. This provides an excellent opportunity for quantifying the relative contribution of each of these mechanisms of sexual selection to the overall opportunity for sexual selection on males (I males). To calculate variance in actual reproductive success, we assigned genetic paternity to 92.3% of 447 chicks sampled in seven years. Reproduction by non-alphas was rare and apparently reflected status misclassifications or opportunistic copulations en route to attaining alpha status rather than alternative mating strategies. On average 31% (range 7-44%, n=6 years) of the total I males was due to variance in reproductive success between alphas and non-alphas. Similarly, in a cohort of same-aged males followed for six years, 44-58% of the total I males was attributed to variance between males of different status. Thus, both intrasexual competition for status and female mate choice among lekking alpha males contribute substantially to the potential for sexual selection in this species.     

The effects of female choice and male-male competition on the mating success of male guppies

This study examined the effects of female choice and male-male competition on male mating success in the guppy, Poecilia reticulata. The fraction of a male's displays that elicited a sexual response from a female was used as a measure of relative female preference for that male. Male mating success was measured by scoring paternity of sons, using colour pattern as a genetic marker. Female preference had a significant effect on male mating success. Male-male competition was relatively uncommon, and did not involve direct threats or combat. Data on male-male competition did not significantly predict mating success. Males from different experimental groups with the same colour pattern had similar mating success. This suggests that female choice could have been based on colour pattern. However, there was no obvious relationship of particular colour pattern elements to mating success or female preference. The males with the same colour pattern were related and most were reared together, so other genetic or environmental factors could have led to similar mating success.     

Male-male competition and female choice in brown trout

In some salmonid species, the females have been assumed to choose their mates on the size of the male's adipose fin. This hypothesis was tested in a stream water aquarium, in which 19 brown trout, Salmo trutta, females were allowed to choose between two males of the same body size but with different adipose fin sizes. The two males were separated from each other in cages. After the female had started to prepare her nest close to one of them, the males were released and allowed to fight each other for the opportunity to spawn. Out of 19 females, 14 prepared a nest closest to the male with the larger adipose fin. However, only six of the 14 females spawned with this male. Males that spawned were more dominant (i.e. were more likely to win fights). When the female spawned with the male she chose, he was less aggressive towards her than when she spawned with the other male. There were no significant differences in the plasma levels of testosterone (T) and 11-ketotestosterone (11-KT) between the chosen males and those not chosen. However, the dominant males had significantly higher plasma levels of T and 11-KT both before and after the experiment. The results support the view that female brown trout exhibit mate choice, but their choice is overruled by male-male competition. Copyright 1999 The Association for the Study of Animal Behaviour.     

Female choice and male competition in longear sunfish

Mating systems with spatial clumping of nests or display sitestheoretically can result from female preference for clumpedmales or from competitive interactions among males. We testedpredictions derived from these two types of models against observationsof nest dispersion, mating success, and alternative male matingtactics in a naturally occurring population of central longearsunfish (Lepomis megalotis megalotis). The data were consistentwith models for male-initiated spatial clumping of nest sites;no evidence of female preference for clumped nests was found.We observed that large males had greater mating success thansmall males and that solitarily nesting males were larger thancolonial males; the most successful males were solitary nesters.Mating success of large males did not increase with colony size.Furthermore, colonial males were often cuckolded by neighbors,whereas solitary males were not cuckolded. We propose that colonialnesting is maintained in longear sunfish because group nestingallows subordinate males to obtain access to females.     

Opposing selection on a sexually dimorphic trait through female choice and male competition in a water boatman

Female choice and male-male competition are traditionally considered to act in concert, with male competition facilitating female choice. This situation would enforce the strength of directional selection, which could reduce genetic variation and thus the benefits of choice. Here I show that in a water boatman, Sigara falleni, the direction of selection through female choice and male competition vary among traits under laboratory conditions. The two forces were mutually enforcive in acting on body size but exerted opposing selection on a sexually selected trait, male foreleg pala size. Female choice favored large palae, whereas male competition favored smaller palae, suggesting that large palae are costly in competition. This conflicting selection through female choice and male competition could be one of the forces that contribute to the maintenance of genetic variation in sexually selected traits.     

Sexual selection in pinyon jays I: female choice and male-male competition

Although pinyon jays, Gymnorhinus cyanocephalus, mate monogamously and male parental investment is high, the mechanisms of sexual selection occur in this species. Experiments were performed using captive birds to determine which male traits are favoured by female choice and which are favoured by inter-male competition. Females mated preferentially with males that were brighter in colour and had larger testes. Females did not prefer large males, but inter-male competition favoured males with larger bills and heavier bodies. Male dominance was not the primary determinant of male mating succes, but it probably conferred an advantage.     

Sexual selection in pinyon jays II: male choice and female-female competition

Experiments were performed on a group of captive pinyon jays, Gymnorhinus cyanocephalus, to determine which female traits are favoured by female-female competition and which are favoured by male choice of mates. Brightness of malar feathers and body weight were weakly associated with female dominance. Female rank, weight and bill depth were related to success in pairing. Competition and choice favoured specific traits less strongly in females than in males.     

Size-assortative mating, male choice and female choice in the curculionid beetle Diaprepes abbreviatus

In the beetle Diaprepes abbreviatus (L.) females are larger on average than males, as indicated by elytra length. Size-assortative matings were observed in wild populations in Florida and in laboratory mating experiments. We tested three mechanisms for this size-assortative mating: (1) mate availability; (2) mating constraints; and (3) mate choice. We found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females; (2) large males successfully competing for large females, leaving small males to mate with small females; and (3) females accepting large males as mates more readily than small males. Males increased their reproductive success by mating with larger, more fecund females. They transferred protein to females during mating. Copyright 1999 The Association for the Study of Animal Behaviour.     

Sperm competition and female procurement of male resources

This study investigates changes in sexual motivation over the duration of a partnership in a population sample stratified by age. The results replicate and extend the findings of a previous study that was based on a sample of college students. In the samples of 30- and 45-year-olds, male sexual motivation remains constant regardless of the duration of the partnership. Female sexual motivation matches male sexual motivation in the first years of the partnership and then steadily decreases. In the sample of 60-year-olds, male sexual motivation always exceeds female sexual motivation, and both are little affected by duration of the partnership. This pattern is clearly evident for some measures of sexual motivation and less so or not at all for others. Interpretations of the current results from social constructivism or from mainstream psychology are difficult to conceive. The results seem more intelligible from an evolutionary perspective as reflections of evolved design for sexual motivation, fine-tuned to the different conditions governing the reproductive success of males and females. In this view male sexual motivation promotes a constant frequency of copulation in order to guard against cuckoldry. Female sexual motivation, in contrast, promotes copulation to solve the adaptive problem of procuring male resources by establishing and maintaining a pair bond. Dr. Dietrich Klusman is a psychologist at the Institute of Medical Psychology, University Clinic Hamburg-Eppendorf, Germany. His research focus is sexual motivation.     

Parasites, male-male competition and female mate choice in the sand goby

Male sand goby Pomatoschistus minutus dominance in competition over nest sites was associated with higher body condition but not with the intensity of infection with any individual parasite of six different species, nor with an overall index combining relative levels of infection by all parasites. Body condition was not related to the intensity of infection with any individual parasite nor with the index of total relative parasite load. In trials in which females spawned, they showed a tendency to choose dominant over subordinate males as mates, but did not consistently choose less parasitized males. Variation in the relative size of the dorsal fins of males was detected, and this related to numbers of the ectoparasitic monogenean Gyrodactylus sp., suggesting that at least some infections have phenotypic effects that could allow females to detect and avoid the most heavily infected males.     

Cryptic male choice: sperm allocation strategies when female quality varies

We examined evolutionary stable sperm allocation and included stochastic variation in male mating frequency, not included in previous models examining sperm allocation strategies. We assumed sperm mixing and variation in female quality and used a genetic algorithm to analyse the evolution of male sperm allocation. Our results show that males should invest more sperm in initial copulations than in subsequent copulations as a male might fail to mate again. The inclusion of variation in female fecundity had no influence on the evolutionary stable sperm allocation strategy if males were unable to recognize female quality. If males were assumed to allocate sperm in response to female quality, the proportion of sperm allocated was positively correlated with female quality. Moreover, with increasing variance in female quality, males conserved more sperm for later copulations. Literature data on sperm allocation from diverse taxa show a good fit with the predictions given by our model.     

The mate choice brain: comparing gene profiles between female choice and male coercive poeciliids

Genes that mediate mate preferences potentially play a key role in promoting and maintaining biological diversity. In this study, we compare mate preference behavior in two related poeciliid fishes with contrasting behavioral phenotypes and relate these behavioral differences to gene profiles in the brain. Results reveal that one poeciliid fish, the Northern swordtail, exhibits robust mate preference as compared to the Western mosquitofish, which utilizes a coercive mating system. Female swordtails display no significant difference in association time between male- and female-exposure trials, whereas female mosquitofish spend significantly less time associating with males relative to females. Furthermore, the preference strength for large males is significantly lower in female mosquitofish relative to swordtails. We then examine expression of three candidate genes previously shown to be associated with mate preference behavior in female swordtails and linked to neural plasticity in other vertebrates: neuroserpin (NS), neuroligin-3 (NLG-3) and N-methyl-d-aspartate receptor (NMDA-R). Whole brain gene expression patterns reveal that two genes (NS and NLG-3) are positively associated with mate preference behavior in female swordtails, a pattern opposing that of the mosquitofish. In mosquitofish females, these genes are downregulated when females express biases toward males yet are elevated in association with total motor activity patterns under asocial conditions, suggesting that the presence of males in mosquitofish species may inhibit expression of these genes. Both gene expression and female behavioral responses to males exhibit opposing patterns between these species, suggesting that this genetic pathway may potentially act as a substrate for the evolution of mate preference behavior.     

Some factors in the choice of male or female sterilisation in Aberdeen

In a random sample of 84 men and 167 women, medical considerations, an intransigent attitude of one partner or a fear of adverse effects, were related to the decision on which partner should seek sterilisation. Only 19% of couples felt that they had a real choice between male and female sterilisation.     

Balancing sexual selection through opposing mate choice and male competition

Male–male competition and female mate choice act contemporaneously in the cockroach Nauphoeta cinerea and the social pheromone of males influences the outcome of both forms of sexual selection. We therefore examined the joint and separate effects of male–male competition and female mate choice to determine if the selective optima for the pheromone were the same or different. Dominant males in a newly established hierarchy mated more frequently, but not exclusively. Manipulations of the multi-component social pheromone produced by males of N. cinerea showed that both long- and close-range attraction of females by males were influenced by the quantity and composition of the pheromone. The most attractive composition, however, differed from that which was most likely to confer high status to males. Since the outcome of male–male competition can conflict with mating preferences exhibited by females, there is balancing sexual selection on the social pheromone of N. cinerea. Such balancing selection might act to maintain genetic variation in sexually selected traits. We suggest that the different forms of sexual selection conflict in N. cinerea because females prefer a blend different to that which is most effective in male–male competition in order to avoid mating with overly aggressive males.