Resource choice in orchard populations of Drosophila

Three capture-recapture experiments were performed on natural populations of Drosophila from orchards in California. Two of the experiments investigated choice between known breeding substrates: oranges v. grapefruits and figs v. peaches. The third involved choice between citrus baits in a citrus orchard and both banana and citrus baits in an adjacent almond orchard. Drosophila pseudoobscura showed no fidelity to the bait or area of initial capture. However, a statistically significant but low level of resource fidelity was displayed by D. melanogaster/D. simulans. An index of differential choice is used to compare our results to those of previous studies. Possible biological bases of choice behaviour are discussed.     

Female mate choice across mating stages and between sequential mates in flour beetles

Few studies have examined how female premating choice correlates with the outcome of copulatory and post-copulatory processes. It has been shown that polyandrous Tribolium castaneum females discriminate among males before mating based on olfactory cues, and also exert cryptic choice during mating through several mechanisms. This study tested whether a male's relative attractiveness predicted his insemination success during copulation. Bioassays with male olfactory cues were used to rank two males as more and less attractive to females; each female was then mated to either her more attractive male followed by less attractive male, or vice versa. Dissections immediately after second copulations revealed a significantly higher percent of successful inseminations for females that remated with more attractive males compared with those that remated with less attractive males. These results indicate that cryptic female choice during copulation reinforces precopulatory female choice in T. castaneum, and suggest that females could use cryptic choice to trade up to more attractive males, possibly gaining better phenotypic or genetic quality of sires.     

Female goats use courtship display as an honest indicator of male quality

Due to the differential cost of reproduction in promiscuous species, like domesticated goat(s) (Capra hircus), it is expected that females should mate with higher quality males, while males should mate with a greater number of females. Females may use conspicuous secondary sexual characteristics of males such as courtship display to distinguish among high and low quality males. Testosterone (T) controls a large suite of secondary sexual characteristics and variation in T concentrations may account of differences in courtship rates. Two studies were conducted to examine the relationship between T concentrations and courtship rate and its role in mammalian female mate choice. Experiment 1 utilized bucks (intact males) and Experiment 2 utilized T-replaced wethers (castrated pre-pubertally). During the first year of Experiment 2, T-replaced wethers received either vehicle control (CON), 25 mg or 100 mg testosterone propionate (TP). During the second year of Experiment 2, T-replaced wethers were treated with either 5 mg, 15 mg or 25 mg TP. For all experiments, mean courtship rates and circulating T concentrations were measured, as well as female preference for males displaying different courtship frequencies. T concentrations and courtship rate were positively correlated for bucks and estrous females preferred high courting bucks. Males receiving 25 mg and 100 mg TP courted females at a similar rate, but both were significantly higher than courtship rates of the CON wethers. Courtship rates of the 25 mg, 15 mg and 5 mg TP-treated males were all significantly different. Females did not show a preference between the 100 mg or 25 mg TP-treated wethers, however both were preferred in comparison to the CON wethers. Females did not show preference between the 25 mg or 15 mg TP-treated wethers, however both were preferred in comparison to the 5 mg TP-treated wethers. Taken together, these studies suggest that courtship rate is T-dependent. Further, females can use courtship rate to distinguish among males.     

Stronger convex (stabilizing) selection on homologous sexual display traits in females than in males: a multipopulation comparison in Drosophila serrata

Mutual mate choice for homologous sexual display traits has been demonstrated in several recent studies yet little attention has been given to quantitative comparison of the strength and form of mate preferences between the sexes. Such comparisons may provide important insight into the evolution of mate choice for honest signals. In particular, because females generally provide the majority of resources for initial offspring development, female displays may trade-off with fecundity, causing preference evolution to differ between the sexes. Recent theory suggests that adaptive male preferences for honest displays in females are possible under certain conditions and may result in preferences that are convex (i.e., stabilizing) in form. We compared sexual selection on a suite of contact pheromones arising from mutual mate choice using nine separate geographic populations of Drosophila serrata. We show that the convex selection is stronger on females than on males overall in these populations, and that convex selection is the predominate form of nonlinear selection on females but not males.     

Good genes and old age: Do old mates provide superior genes?

It has been suggested that female preference for older mates in species without parental care has evolved in response to an alleged higher genetic quality of older individuals. This is based on the widespread assumption that viability selection produces older individuals that are genetically superior to younger individuals. In contrast, we propose that the oldest individuals rarely are genetically superior. Quantitative genetic models of life history evolution indicate that young to intermediately aged individuals are likely to possess the highest breeding values of fitness. This conclusion is based on four arguments: 1) Viability selection on older individuals may decrease or at least not substantially increase breeding values of fitness, because there may exist negative genetic correlations between late-age and early-age life history parameters, 2) Fertility selection is likely to raise the fitness of gametes produced by young individuals more than those produced by old individuals, because the covariance between fertility and fitness often decreases with age, 3) The history of selection on their parents makes younger individuals more fit than older individuals, 4) Germ-line mutations, which are generally deleterious, significantly decrease the breeding value of fitness of an individual throughout its lifespan, especially in males. Therefore, females that mate with the oldest males in a population are doing so for reasons other than to obtain offspring of high genetic quality.     

Male choice generates stabilizing sexual selection on a female fecundity correlate

We know very little about male mating preferences and how they influence the evolution of female traits. Theory predicts that males may benefit from choosing females on the basis of traits that indicate their fecundity. Here, we explore sexual selection generated by male choice on two components of female body size (wing length and body mass) in Drosophila serrata. Using a dietary manipulation to alter female size and 828 male mate choice trials, we analysed linear and nonlinear sexual selection gradients on female mass and wing length. In contrast to theoretical expectations and prevailing empirical data, males exerted stabilizing rather than directional sexual selection on female body mass, a correlate of fecundity. Sexual selection was detected only among females with access to standard resource levels as an adult, with no evidence for sexual selection among resource-depleted females. Thus the mating success of females with the same body mass differed depending upon their access to resources as an adult. This suggests that males in this species may rely on signal traits to assess body mass rather than assessing it directly. Stabilizing rather than directional sexual selection on body mass together with recent evidence for stabilizing sexual selection on candidate signal traits in this species suggests that females may trade-off resources allocated to reproduction and sexual signalling.     

Correlates of male mating success in the lekking great snipe (Gallinago media): results from a four-year study

To investigate behavioral or morphological traits importantas mate choice cues, we measured selection differentials (s)as the covariances between each trait and male mating success,and directional selection gradients (J3) from multiple linearregression of the standardized traits on male mating success.Data from two leks in four consecutive years were included,and the annual data were analyzed separately. The main findingsare: (1) the distribution of male mating success proved to beless skewed than those found in many other lekking species,(2) only a few traits yielded significant selection gradients,(3) the importance of age on male mating success changed acrossyears, (4) females may use traits with a high variance as matechoice cues, and (5) individual males achieved similar matingsuccesses between years. Attendance and age were the traitsmost consistently correlated with male mating success, but notraits showed significant selection gradients in all years.Our results indicate that variable sexual selection pressuresexisted between years, but the high correlation found betweenthe mating success of individual males in successive seasonsalso indicates that permanent differences in male traits areimportant. Key words: lek, mate choice, sexual selection.     

Reinforcement shapes clines in female mate discrimination in Drosophila subquinaria

Reinforcement of species boundaries may alter mate recognition in a way that also affects patterns of mate preference among conspecific populations. In the fly Drosophila subquinaria, females sympatric with the closely related species D. recens reject mating with heterospecific males as well as with conspecific males from allopatric populations. Here, we assess geographic variation in behavioral isolation within and among populations of D. subquinaria and use cline theory to understand patterns of selection on reinforced discrimination and its consequences for sexual isolation within species. We find that selection has fixed rejection of D. recens males in sympatry, while significant genetic variation in this behavior occurs within allopatric populations. In conspecific matings sexual isolation is also asymmetric and stronger in populations that are sympatric with D. recens. The clines in behavioral discrimination within and between species are similar in shape and are maintained by strong selection in the face of gene flow, and we show that some of their genetic basis may be either shared or linked. Thus, while reinforcement can drive extremely strong phenotypic divergence, the long‐term consequences for incipient speciation depend on gene flow, genetic linkage of discrimination traits, and the cost of these behaviors in allopatry.     

Males Benefit from Mating with Outbred Females in Drosophila littoralis: Male Choice for Female Genetic Quality?

The evolution and expression of mate choice behaviour in either sex depends on the sex‐specific combination of mating costs, benefits of choice and constraints on choice. If the benefits of choice are larger for one sex, we would expect that sex to be choosier, assuming that the mating costs and constraints on choice are equal between sexes. Because deliberate inbreeding is a powerful genetic method for experimental manipulation of the quality of study organisms, we tested the effects of both male and female inbreeding on egg and offspring production in Drosophila littoralis. Female inbreeding significantly reduced offspring production (mostly due to lower egg‐to‐adult viability), whereas male inbreeding did not affect offspring production (despite a slight effect of paternal inbreeding on egg‐to‐adult viability). As inbreeding depressed female quality more than male quality, the benefits of mate choice were larger for males than for females. In mate choice experiments, inbreeding did not affect male mating success (measured as a probability to be accepted as a mate in a large group), suggesting that females did not discriminate among inbred and outbred males. In contrast, female mating success was affected by inbreeding, with outbred females having higher mating success than inbred females. This result was not explained by lower activity of inbred females. Our results show that D. littoralis males benefit from mating with outbred females of high genetic quality and suggest adaptive male mate choice for female genetic quality in this species. Thus, patterns of mating success in mate choice trials mirrored the benefits of choice: the sex that benefited more from choice (i.e. males) was more choosy.     

Nestmateship and body size do not influence mate choice in males and females: A laboratory study of a primitively eusocial wasp Ropalidia marginata

We investigated the effect of nestmateship and body size on mate selection through a choice based assay in the primitively eusocial wasp Ropalidia marginata. A recent study has shown that male and female R. marginata mate with their nestmates and non-nestmates with equal probability if no choice is available. That study could also not detect any influence of body size on mating probability in the absence of choice. To confirm that the same results can be obtained even when the wasps have a choice, we offered a choice of two virgin partners either to a virgin test male or to a virgin test female and measured the probability that the test individual would mate with any particular partner based on nestmateship or body size. We show here that even when a choice is available, neither male nor female test wasps base their mate choice on the nestmateship or body size of the partner. We therefore suggest that the natural mating habit of these wasps is sufficiently promiscuous and not constrained by such factors as nestmateship and body size.     

Modelling the evolution of female choice strategies under inbreeding conditions

Recently, many mate choice studies have discussed the role of genetic compatibility and inbreeding for the evolution of mate choice. With population genetic simulations I compared the potential advantage of three different female choice strategies under inbreeding conditions. Females were assumed to benefit indirectly via a preference for (i) complementary males, (ii) males with few detrimental mutations, and (iii) non-inbred males. Probably related to the reduced inbreeding depression in offspring of choosy females, the choice-allele increased for all three strategies. However, the advantage of the strategies differed widely. Choice of males with fewer mutations provided a comparatively large advantage, choice of complementary males led to a reasonable advantage, and choice of non-inbred males only resulted in a minor advantage of female choice. My results show that complementary mate choice can be almost as beneficial as conventional good-genes choice of mates with lower genetic load. Compared to the two other mate choice strategies, choice of non-inbred males is less likely to contribute to the evolution of costly mate choice. The results of a recent study showing that female sticklebacks prefer males with a larger number of MHC-loci is thus unlikely to be related to an indirect benefit of choosing non-inbred males.     

DIRECT AND INDIRECT SEXUAL SELECTION AND QUANTITATIVE GENETICS OF MALE TRAITS IN GUPPIES (POECILIA RETICULATA)

Abstract.— The ornamentation and displays on which sexual attractiveness and thus mating success are based may be complex and comprise several traits. Predicting the outcome of sexual selection on such complex phenotypes requires an understanding of both the direct operation of selection on each trait and the indirect consequences of selection operating directly on genetically correlated traits. Here we report the results of a quantitative genetic analysis of the ornamentation, sexual attractiveness, and mating success of male guppies (Poecilia reticulata). We analyze male ornamentation both from the point of view of single ornamental traits (e.g., the area of each color) and of composite measures of the way the entire pattern is likely to be perceived by females (e.g., the mean and contrast in chroma). We demonstrate that there is substantial additive genetic variation in almost all measures of male ornamentation and that much of this variation may be Y linked. Attractiveness and mating success are positively correlated at the phenotypic and genetic level. Orange area and chroma, the area of a male's tail, and the color contrast of his pattern overall are positively correlated with attractiveness and/or mating success at the phenotypic and genetic levels. Using attractiveness and mating success as measures of fitness, we estimate gradients of linear directional sexual selection operating on each male trait and use equations of multivariate evolutionary change to predict the response of male ornamentation to this sexual selection. From these analyses, we predict that indirect selection may have important effects on the evolution of male guppy color patterns.     

Male courtship attractiveness and paternity success in Photinus greeni fireflies

Although female mate choice and male sperm competition have separately attracted much attention, few studies have addressed how precopulatory and postcopulatory episodes of sexual selection might interact to drive the evolution of male traits. In Photinus fireflies, females preferentially respond to males based on their bioluminescent courtship signals, and females gain direct benefits through male nuptial gifts acquired during multiple matings over several nights. We experimentally manipulated matings of P. greeni fireflies to test the hypothesis that postcopulatory paternity success might be biased toward males that are more attractive during courtship interactions. We first measured male courtship attractiveness to individual females using field behavioral assays. Females were then assigned to two double-mating treatments: (1) least attractive second male-females were first mated with their most attractive male, followed by their least attractive male, or (2) most attractive second male-females mated with males in reverse order. Larval offspring produced by each female following these double matings were genotyped using random amplified polymorphic DNA (RAPD) markers, and male paternity was determined. Contrary to prediction, firefly males that were more attractive to females based on their bioluminescent courtship displays subsequently showed significantly lower paternity, reflecting possible male trade-offs or sexual conflict. Differences in male paternity were not related to male body condition, testes or accessory gland mass, or to variation in female spermathecal size. Additionally, this study suggests that changes in phenotypic selection gradients may occur during different reproductive stages. These results indicate that it is crucial for future studies on sexual selection in polyandrous species to integrate both precopulatory and postcopulatory episodes to fully understand the evolution of male traits.     

CRYPTIC FEMALE CHOICE: CRITERIA FOR ESTABLISHING FEMALE SPERM CHOICE

In this paper, I consider the criteria necessary to demonstrate the postcopulatory ability of females to favor the sperm of one conspecific male over another, that is, sperm choice. In practice it is difficult to distinguish between sperm competition and sperm choice, and sperm choice can be demonstrated only if the effects of sperm competition can be controlled. Few studies have used experimental protocols that do this, so evidence for sperm choice is limited. Moreover, in those studies in which sperm choice occurs, it does so to avoid incompatible genetic combinations and is therefore unlikely to result in directional sexual selection.     

INTERACTION-INDEPENDENT SEXUAL SELECTION AND THE MECHANISMS OF SEXUAL SELECTION

Darwin identified explicitly two types of sexual selection, male contests (combat and displays) and female choice, and he devoted the overwhelming majority of his examples to traits that influence the outcome of these interactions. Subsequent treatments of sexual selection have emphasized the importance of intra- and intersexual interactions as sources of sexual selection. However, many traits that are important determinants of mating success influence mating success without necessarily affecting the outcome of intra- and intersexual interactions. Here, I argue that traits can be subject to sexual selection even if they do not affect the outcome of intra- and intersexual interactions. I distinguish two types of sexual selection, interaction-independent and interaction-dependent selection, based on whether variance in mating success is the result of trait-dependent outcomes of interactions between conspecifics. I then use this distinction to construct a framework for classifying types of sexual selection that unifies and expands previously proposed frameworks. Finally, I outline several implications that the concept of interaction-independent sexual selection has for the general theory of sexual selection.