Sexual Selection and Variance in Reproductive Success

Influence du risque du a la presence de predateurs et a au jeune subit sur le taux d'accouplement et sur sa duree

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.

Previous work (Reynolds and Gross 1992) has demonstrated that mate choice enables female guppies, Poecilia reticulata , to obtain heritable benefits (‘good genes’) for their offspring. It is not yet known whether males also signal their functional fertility, that is, their viability as fathers, to female guppies. Our study tested the hypothesis that a male9s behaviour reveals characteristics of his ejaculate. We uncovered a strong correlation between display rate and sperm number in male guppies derived from two wild Trinidadian populations. This result indicates that information on the functional fertility of potential mates is available to females. However, our investigation also found a significant relationship between the frequency of sneaky mating and sperm number. Female guppies in wild populations, particularly those experiencing a high predation regime, are constantly subjected to sneaky mating attempts. As the males responsible for these sneaky matings have highest reserves of sperm they may have the capacity to undermine female choice. The implications of these findings for sperm competition are discussed.

A "Good-Sperm" Model Can Explain the Evolution of Costly Multiple Mating by Females

The Cost of Mating

Reproduction is costly and individual decisions about when and how much to invest in reproduction will relate to potential benefits in terms of offspring survivorship or mating success. For those organisms that can reproduce more than once, individuals should invest relatively more in reproduction when the potential benefits are high. If females gain directly or indirectly from choosing attractive, highly ornamented males, then it could be predicted that females mated to these males should invest more in reproduction. Here we report the first test of this prediction for a lekking species, peafowl (Pavo cristatus), in which males do not provide resources for offspring, and females prefer to mate with those males that possess the most elaborate trains. We find that peahens randomly mated to males that vary in the degree of ornament produce more eggs for those peacocks with more elaborate trains. We could find no evidence to support the possibility that this difference arises as a result of difference between males in their ability to fertilize eggs.

Female preference for males successful in male–male competition is generally assumed to result in mating with high quality males. Here I report results from an experiment disentangling the effects of intra– and intersexual selection in the sand goby, Pomatoschistus minutus , a marine fish that exhibits paternal care. I show that large males are successful in male–male competition, but contrary to what one would expect, dominants are not preferred by females and are not better at taking care of the eggs. Female preference, however, correlated with the subsequent hatching success of the eggs. Thus, female choice selects for good parenting. Hence, direct benefits in the form of superior paternal care can explain female choice in this species, supporting a good parent process of sexual selection. However, choosing on the outcome of male–male competition does not enable females to mate with the ‘best’ males.

Models of sexual selection predict that females use ornament size to evaluate male condition. It has also been suggested that ornament asymmetry provides females with accurate information about condition. To test these ideas we experimentally manipulated condition in the stalk-eyed fly, Cyrtodiopsis dalmanni, by varying the amount of food available to developing larvae. Males of this species have greatly exaggerated eyestalk length and females prefer to mate with males with wider eyespans. Our experiments show that male ornaments (eyestalks) display a disproportionate sensitivity to condition compared with the homologous character in females, and to non-sexual traits (wing dimensions). In contrast, in neither sex did asymmetry reflect condition either in sexual ornaments or in non-sexual traits. We conclude that ornament size is likely to play a far greater role in sexual selection as an indicator of individual condition than does asymmetry.

In cooperatively breeding superb fairy-wrens (Malurus cyaneus), all males contribute to the feeding and defence of young. Despite the expectation that such paternal care should be directed only to relatives, DNA fingerprinting revealed that most offspring (76%, 138 out of 181) were sired by extra-group males that contributed no care, and that almost all broods (95%, 38 out of 40) contained young sired by extra-group fathers. This is the highest known incidence of cuckoldry. This remarkable mating system is produced by female control of fertilization and consistent preference for certain extra-group male genotypes. This choice leads to the production of sons that also gain extra-group fertilizations. One constraint on the extra-pair mate choice of females is the level of parental assistance received from males. Males living in pairs contribute relatively more parental care and are more likely to gain paternity in one of their broods (85%, 11 out of 13) than dominant males in multi-male cooperative groups (30%, 8 out of 27). In groups helpers compensate for the lower parental assistance of dominant males, so the total feeding rate is similar between pairs and groups. This suggests that females allow males in pairs more paternity to ensure their assistance with parental care. Helpers provide an alternative source of paternal investment, and allow females to express unrestricted mate choice. Mating options for females in other species with female control of fertilization may also reflect a trade-off between acquiring the genes of high-quality males for their offspring and parental care of those offspring.

A general formulation of inclusive fitness is proposed which specifically accounts for competitive effects between relatives. As an example, for an asexual population in a homogeneous inelastic environment, such as is found in a stepping-stone model of dispersal, the inclusive fitness of a breeding female, under weak selection, is independent of her direct effect on the fitness of other individuals in the population. More precisely, suppose a female acts in a way that changes not only her own fitness but the fitness of several other females in the population who may be relatives (i.e. she changes the number of their offspring). I call these changes the direct effects of the action. There will also be indirect effects on the fitness of these and other females which come from the competitive impact of the extra offspring in the next generation. The principal result is that these indirect effects exactly cancel all the direct effects except the direct effect of the actor on herself.

Most recent models of the partitioning of reproduction attempt to explain patterns of skew on the assumption that dominant individuals have complete control over breeding opportunities within the group, but may nevertheless concede a share of direct reproduction to subordinates as an incentive to remain peacefully in the association. Although these models may be applicable to some animal societies, we argue that they fail to provide a comprehensive theory of skew. Instead, we suggest that subordinates may often be able to claim unsanctioned reproduction for themselves, but will be forced to exercise a degree of reproductive restraint lest they incite ejection by the dominant. Reproductive skew, in other words, may reflect the threat of ejection (inducing subordinate restraint) rather than the threat of subordinate departure (inducing reproductive concessions by dominants). We present a simple ESS model of reproductive skew under these circumstances, which demonstrates that a shift in emphasis from reproductive concessions by dominants to reproductive restraint on the part of subordinates, radically alters the predictions of skew models. High group productivity, high relatedness and (when group members are related) strong ecological constraints are all expected to lead to reduced skew (the opposite conclusions to those of previous, concession-based analyses). The reason is that these factors reduce the benefits (or increase the costs) of ejection to the dominant, who therefore does best to tolerate more subordinate reproduction.

Evolutionary theory predicts that differences in parental care patterns among species arose from interspecific differences in the costs and benefits of care for each sex. In Galilee St Peter9s fish, Sarotherodon galilaeus (Cichlidae), male care, female care and biparental care all occur in the same population. We exploit this unusual variability to isolate conditions favouring biparental versus uniparental mouth–brooding by males or females. We first review a game–theoretic model of parental care evolution, predictions of which we test experimentally in this paper. Manipulations of the operational sex ratio show that males and females desert their offspring more frequently when the costs of care are high (in terms of lost mating opportunities). Breeding trials with males of different sizes show that small fathers desert more frequently than large fathers. We attribute this to the associated difference in the fitness benefit of biparental care relative to female–only care. Our experimental results confirm that in St Peter9s fish the probability of caring is determined facultatively according to current conditions at each spawn. The experiments and model together suggest that interspecific variation in remating opportunities and clutch size may be responsible for differences in care patterns within the sub–family Tilapiini. Our results support the hypothesis that biparental mouth–brooding was the ancestral state of both male and female uniparental mouth–brooding in cichlid fishes.

Predation Risk Affects Mate Choice in a Gobiid Fish

On Wicken Fen and nearby watercourses eastern England, parasitism by cuckoos, Cuculus canorus, declined from 26 and 16 of reed warbler (Acrocephalus scirpaceus) nests in 1985 and 1986, respectively...

In Atlantic salmon, as in most salmonids, males can mature early in the life cycle, as small freshwater fish, termed parr, and/or undergo a sea migration before maturing as full–size adults. The alternative life histories are contingent on environmental and social circumstances, such as growth rate, territory quality or any other factor that affects the individual9s state. In order to model the choice of life history in this group of commercially valuable species, it is necessary to understand not only the relative contribution of the different male types to subsequent generations, but also to know the factors that affect reproductive success in each type. In this paper we present the results of a study designed to investigate the factors that affect the reproductive success of mature parr. We used highly polymorphic minisatellite DNA markers to analyse paternity in a series of mating experiments where the number and body size of parr were manipulated. The fraction of eggs fertilized by mature parr ranged from 26 to 40 per cent, with individual parr fertilizing up to 26 per cent of the eggs. A strong positive correlation was found between parr size and reproductive success. The relative success of parr decreased with increasing parr number. Data from this and other studies on variation in the timing and degree of parr reproductive success are discussed in relation to the evolution of male mating strategies and life history in salmonids.