Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

The effects of male–male contests and female eavesdropping on female mate choice and male mating success in the jumping spider, <Emphasis Type="Italic">Thiania bhamoensis</Emphasis> (Araneae: Salticidae)

In experiments that comprised of three phases (fight, choice, and mating) under “seen” and “unseen” conditions, we examined the effects of the outcomes of male–male contests and female eavesdropping on female mate choice and male mating success in the fighting spider, Thiania bhamoensis (Salticidae). The results revealed female eavesdropping on agonistic interactions. Females that had watched an aggressive interaction showed no distinctive preference for the winner over the loser, but they preferred the loser when they had not observed a fight. Winners, however, achieved a greater mating success than did losers during the mating phase. Gaining access to females was based on the insistence of the winners in courtship in terms of the number of quivers, rather than on the fighting behavior of the males. Hence, the outcome of male–male contests may not be an important determinant of a male’s mating success in T. bhamoensis. Instead, courtship display plays an important role in determining the success of male mating in this species. This study also suggests that female mate preference may not be a good indicator of eventual female mate choice and male mating success. Thus, a causal relationship between female mate preference and male mating success cannot be inferred. Joanna P. Y. Chan, Pei Rong Lau, and Ai Jie Tham contributed equally.     

Phenotypic correlates and survival consequences of male mating success in lek-mating greater prairie-chickens (<Emphasis Type="Italic">Tympanuchus cupido</Emphasis>)

Female choice and male–male aggression are two modes of sexual selection that can lead to elaboration of male morphological and behavioral traits. In lek-mating species, male mating success is often strongly skewed, and it is puzzling why variation in male traits is still observed given directional female choice. If male traits correlated with reproductive success are honest signals of male quality, there may be survival costs associated with the expression of those traits. In this study, we examined whether morphological, behavioral, and territorial traits are correlated with male mating success and survival in the lek-mating greater prairie-chicken (Tympanuchus cupido). We introduce a novel application of multinomial discrete choice models for analysis of female mate choice behavior. We found that behavioral and territorial attributes showed 6.5 times more variability among males than morphological traits. Both display and aggressive behaviors were strong predictors of male mating success, suggesting that both female choice and male–male aggression were important in determining mating success among male greater prairie-chickens. Moreover, annual survival of male prairie-chickens was independent of mating success and male traits. Females appear to be choosing males based on behavioral traits where large variation exists between males (coefficient of variation >30%). Behavioral traits were the most important factor in determining mating success of male prairie-chickens, but the mechanism underlying this relationship is unknown. In the future, experimental manipulations of male hormones or parasite loads could bridge the proximate mechanisms and ultimate consequences of factors mediating male mating success in lek-mating grouse.     

Bushcricket song as a clue for spermatophore size?

Bushcricket males of Poecilimon zimmeri transfer large and protein-rich spermatophores during mating, which females directly ingest. There is correlational evidence that heavier males transfer larger nuptial gifts. In no-choice mating trials, females mated randomly with respect to male’s body weight. In contrast, in two-choice mating trials, female bushcrickets exhibit clear choice for the heavier male. This heavier male advantage was also found in pre-mating choice during phonotaxis. With manipulated mute males, females mated at random with regard to body weight of the competitors. The number of physical encounters between a female and males was low in all tests with a single male (no choice) and greater in choice-tests with two competing males. The latencies to mate also differed significantly between treatments. The time mating pairs spent in precopula was short in experiments where the males could hear rivals and significantly longer in all other tests using either a single male or mute males. Thus, acoustic signalling in male bushcrickets seems to signal male body weight. A preference for heavier males may reflect a female’s preference for a larger spermatophore and therefore a greater direct benefit.     

Vocalizations by Alaskan moose: female incitation of male aggression

Evidence of female fomentation of male–male aggression as a mechanism of mate choice is rare, especially in mammals. Female choice of mates in polygynous species may be masked by intense male competition or by males attempting to restrict female choice. We studied protest moans of female Alaskan moose Alces alces gigas in interior Alaska, USA, from 1987 to 1990, to determine if moans incited male–male aggression. Alaskan moose exhibit a mating system in which one dominant male (the harem master) herds, defends, courts, and attempts to mate with females in his harem. Protest moans were given by females only in response to courtship. We hypothesized that if protest moans were related to females reducing harassment and exercising mate choice, females should give protest moans more frequently when courted by small males and less often when courted by large males, and that rates of male–male aggression would be elevated following protest moans. Harems were composed of one large male, with a mean of 4.4 females (median = 3 females); 10% of 132 harems included ≥10 females. The temporal pattern of protest moans from late August through November was associated with, but tended to lag behind, mating behavior. The rate of protest moans given by females decreased with increasing size of males courting them. Male–male aggression was significantly less during periods without protest moans than during periods in which protest moans occurred. These results indicate that female moose gave protest moans to reduce harassment by smaller males, and assure a mating opportunity with the most dominant male. Such a subtle mechanism of indirect mate choice by females may occur in other vertebrates in which choice is limited by a mating system in which male–male combat and male dominance over females reduces opportunities for female choice. The importance of female choice may be undervalued in studies of sexual selection in mammals.     

Mating effort and female receptivity: how do male guppies decide when to invest in sex?

Males vary in the degree to which they invest in mating. Several factors can explain this variation, including differences in males’ individual condition and the fact that males allocate their energy depending on the context they face in each mating attempt. Particularly, female quality affects male reproductive success. Here, we studied whether male guppies (Poecilia reticulata) strategically allocated more mating effort, in terms of mating behaviour and male–male competition, when they were matched with a receptive (R) female than a non-receptive one. In accordance with our prediction, we found that males increased their mating behaviour when they were with a receptive female. Even though male guppies can inseminate non-receptive females, we only found high levels of courtship between males that were with a receptive female rather than a non-receptive one. Although there was little affect of female receptivity on male–male competition, we found that males chased and interrupted courtships more with receptive females than with non-receptive females regardless of odour. Finally, we also studied whether the sexual pheromone produced by receptive female guppies is a cue that males use in order to increase their mating effort. We found that males were more attracted to a female when they perceived the sexual pheromone, but only increased their mating and aggressive behaviours when females showed receptive behaviour. This strategic increase in mating effort could result in higher male reproductive success because mating attempts towards receptive females are likely to be less costly and males could have a greater probability of fertilisation.     

Song length variation serves multiple functions in the white-crowned sparrow

A wide variety of hypotheses has been proposed to explain the structural diversity in bird song repertoires. Song diversity is frequently described in terms of song “types” or within-type “variants.” Male Puget Sound white-crowned sparrows, Zonotrichia leucophrys pugetensis, produce variants of their single adult song type by altering the number of repetitions of syllables in the terminal trill. We tested whether variation in trill length correlated with distance to the receiver and with signaling context as predicted by the eavesdropping avoidance hypothesis and the strategic signaling hypothesis. In accordance with the eavesdropping avoidance hypothesis, males sang variants with shorter trills, and sang quieter and less frequently when near their mate during the incubation phase than during spontaneous singing while unpaired. Males also sang variants with short trills, but at a high rate and variable amplitude when within 10 m of an opponent during close male–male territorial interactions. In agreement with the strategic signaling hypothesis, males decreased trill length immediately before chasing an opponent, but did not change length consistently prior to flight. We conclude that the occurrence of short quiet songs sung near the mate agrees with predictions of the eavesdropping avoidance hypothesis, while short songs sung near other males are best explained by the strategic signaling hypothesis. Trill length variation may be a conventional signal of aggressive intentions in male–male contests stabilized by receiver-imposed retaliation costs.     

Vocal interactions in common nightingales (<Emphasis Type="Italic">Luscinia megarhynchos</Emphasis>): males take it easy after pairing

Seasonal patterns of bird song have been studied intensively with a focus on individual males. However, little is known about seasonal patterns of singing during vocal interactions between males. Vocal interactions have been shown to be important in sexual selection as males may signal aspects of motivation or quality. Here, we investigated in nightingales (Luscinia megarhynchos) whether a male’s behaviour in vocal interactions at different stages of the breeding season is influenced by its mating status. We examined how males that differ in their subsequent mating success respond to a non-interactive, nocturnal playback presented during the period of mate attraction and subsequently during the egg-laying period. We found that mated males overlapped fewer songs and had a lower song rate during the egg-laying period compared to their responses during the mate-attraction period, whereas unpaired males did not vary in their responses between the two periods. Our results suggest that mating status is a key factor affecting singing behaviour in vocal interactions and that a time-specific singing pattern like song overlapping is used flexibly during vocal interactions. Because song overlapping is thought to be a signal of aggression in male–male vocal interactions, it seems that males vary the level of aggression in vocal interactions according to their mating status and to the stage in the breeding season.     

Mating behavior of <Emphasis Type="Italic">Abdopus aculeatus</Emphasis> (d’Orbigny 1834) (Cephalopoda: Octopodidae) in the wild

The mating system of Abdopus aculeatus incorporates sneaker matings, mate guarding, sex-specific body patterns, frequent copulations, and male–male competition for mates, making it more similar to that of aggregating decapod cephalopods than any previously known octopus social system. Large male–female A. aculeatus occupy ‘Adjacent’ (G_A) dens and copulate frequently in mate-guarding situations over successive days. Nearby individuals copulate in ‘Temporary guarding’ (G_T) and ‘Transient’ (T; non-guarding) situations, the latter of which can involve ‘Sneaker’ (S) mating. In a focal animal study of these octopuses in the wild (Sulawesi, Indonesia) we addressed the hypotheses that they demonstrate: (1) precopulatory mate choice, (2) differential copulation rates by individuals employing different mating tactics, and (3) distant sex identification. We quantified daily copulation rates of A. aculeatus of reproductive size as well as aspects of copulation duration, display, mate-competition, and mate rejection. Mating tactic correlated with daily copulation rates. ♂G_A spent significantly more time copulating than did ♂T, while ♀G_A spent more than twice as much time per day in copula than did other females. Sneaker copulations lasted longer than those by males adopting other tactics. Mate-guarding was an effective and important tactic used by males to temporarily monopolize mating with apparently non-selective females. Males demonstrated clear pre-copulatory mate choice by guarding and mating repeatedly with large females (typically ♀G_A). While foraging alone away from the den, ♂G procured ‘Transient’ copulations with unguarded females. However, mate-guarding reduced the amount of time ♂G were alone and may impede their ability to seek out new mates. Low-copulation rates by ♀T, the smallest female tactic on average, may reflect this trade-off between mate preference and mate-searching by males, or non-receptivity of some females. A male-typical body pattern (black and white stripes) appeared to facilitate distant sex identification. Although mating and aggression were often initiated before contact between individuals, same-sex copulations and intense male–female aggression were rare. By contrast frequent male–female copulations and intense male–male aggression were consistent behavioral components of mating in A. aculeatus at these sites. Because the behavioral and ecological characters conducive to this complex system are not exclusive to A. aculeatus, it is possible that other octopuses exhibit some or all of these behaviors. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Prior encounters with the opposite sex affect male and female mating behavior in a wolf spider (Araneae,Lycosidae)

Mate availability can vary widely in nature depending upon population density and sex ratio and can affect the ability of individuals to be selective in mate choice. We tested the effects of prior encounters with the opposite sex (i.e., exposure to the opposite sex either with or without mating) on subsequent mating behavior in two experiments that manipulated mate availability for both males and females in the wolf spider, Hogna helluo. The probability of mating in the experimental trial depended upon whether the prior encounter involved mating or not, and males and females responded in opposite directions. Exposure without mating resulted in a higher subsequent frequency of mating for females and a lower subsequent frequency of mating for males, while prior mating experience resulted in a lower frequency of female remating and a higher frequency of male remating. Prior exposure without mating did not affect female aggression. However, mated females engaged in precopulatory cannibalism more frequently than virgins. Mated males escaped postcopulatory cannibalism more frequently than virgins. Our results show that males respond to exposure without mating in the expected manner. However, prior mating (1 week earlier) had unexpected effects on males, which may be due to mated males being of higher quality. There were little or no effects of the size of the prior exposure individual or mate on subsequent mating behaviors. Further research is needed to determine why different species use different degrees of prior information in mate choice.     

Independent effects of male and female density on sexual harassment, female fitness, and male competition for mates in the western mosquitofish Gambusia affinis

Operational sex ratio (the ratio of sexually active males to fertilizable females) has a major influence on male competition for mates and male–female interactions. The contributions of male and female density per se to mating system dynamics, however, are rarely examined, and the fitness consequences are often inferred rather than quantified. Male mosquitofish (Gambusia affinis) compete aggressively and frequently harass females for copulations, a behavior thought to reduce female fitness. Female fitness can also be reduced by increases in female density, which may affect food availability, cannibalism rates, and chemical interactions between females. I manipulated male and female densities of G. affinis to measure their effects on male–male aggression, male harassment toward females, and female fitness. I found that males chased rivals more often and attempted fewer copulations when female density decreased, but surprisingly male density had no significant effect on the frequency of these male behaviors. In contrast, males’ agonistic displays toward other males increased with male density, but display behavior was unaffected by female density. These results suggest that male and female density do not always contribute equally or at all to the patterns of behavior we observe. Female fitness declined as female density increased, the opposite pattern expected if male harassment is costly to females. This suggests that a strong, negative effect of female density overwhelmed any potential costs of male harassment. Sources of female density dependence and the consequences of changes in male and female density to patterns of male behavior are discussed.     

Proximal traits and mechanisms for biasing paternity in the red flour beetle Tribolium castaneum (Coleoptera: Tenebrionidae)

A comprehensive understanding of sexual selection requires knowledge of the traits and mechanisms responsible for increasing a male’s paternity share (proportion of progeny sired) relative to that of other males mating with the same female. In this study we manipulated by starvation the expression of traits that might influence male paternity share in Tribolium castaneum. We then conducted experiments to examine how male starvation affects male performance during sequential episodes of sexual selection from mating to progeny production, and investigated female control over specific stages by using live vs dead females. Comparison of starved vs fed males revealed that T. castaneum females have control over spermatophore transfer during mating, as live females rejected inseminations by starved (“low quality”) males. None of the measured male copulatory behaviors (leg-rubbing frequency, asymmetry, and percent of time spent rubbing) affected the probability of successful insemination, but the last two were positively associated with male paternity share. Spermatophore positioning within the female reproductive tract was not affected by male treatment (starved/fed), by female treatment (live/dead), or by male copulatory behaviors. Starvation, however, had a dramatic effect on male reproductive physiology, decreasing both accessory gland size and total number of sperms transferred (but not sperm viability in seminal vesicles). In addition, females who mated to starved males stored fewer sperms in their spermathecae, which, together with decreased ejaculate size, may explain the reduced paternity share of starved males compared to fed males. This study elucidates some cryptic mechanisms influencing male reproductive success and aids our understanding of trait evolution through sexual selection.     

Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits <Emphasis Type="Italic">Remiz pendulinus</Emphasis>

Melanin-based ornaments are often involved in signaling aggression and dominance, and their role in sexual selection is increasingly recognized. We investigated the functions of a melanin-based plumage ornament (facial ‘mask’) in male Eurasian penduline tits Remiz pendulinus in the contexts of male–male aggression, mating success, and parental care. The penduline tit is a passerine bird with a unique mating system in which both sexes may mate with several mates in a breeding season, and one (or both) parent deserts the clutch. Our study revealed that mask size of males is more likely an honest signal used by females in their mate choice decisions than a trait involved in male–male competition. First, mask size increased with both age and body condition, indicating that the mask may signal male quality. Second, males with larger masks paired more quickly and had more mates over the breeding season than males with smaller masks. Third, we found no evidence that male mask size signals male–male aggression or dominance during competitive encounters. The increased mating success of large-masked males, however, did not translate into higher reproductive success, as nestling survival decreased with mask size. Therefore, we conclude that there is either no directional selection on male mask size or males with larger masks receive indirect, long-term benefits.     

Exogenous testosterone increases female aggression in the European starling (<Emphasis Type="Italic">Sturnus vulgaris</Emphasis>)

In the European starling, Sturnus vulgaris, optimal mating systems differ between males and females. Males gain from polygyny, whereas monogamy increases female fitness. The cost of polygyny to females lead to intense female–female competition, and it has previously been shown that the intensity of female aggression during the pre-breeding period can predict the realised mating system. The physiological regulation of such female aggression in starlings is not yet known. This study examines the role of testosterone in mediating aggressive behaviours involved in intra-specific reproductive competition in female starlings. Testosterone levels were experimentally elevated with testosterone implants in females during the pre-laying period. To simulate a situation in which an additional female tried to mate with the focal female’s mate, a caged female was presented close to a nest-site to which the male could attract a secondary female. Testosterone was significantly related to several behaviours involved in female–female interactions. Females with testosterone implants spent significantly more time close to the caged female and produced more song bouts than control females. In contrast, male behaviour was unrelated to the experimental status of the mate. Females mated to males that attracted a secondary female were less aggressive towards the caged female than those that remained monogamously mated. The effect of exogenous testosterone in this study indicates that androgens may mediate social behaviours in female starlings during the breeding season.     

A female preference for experienced males in the almond moth, Cadra cautella

Male mating status can affect female reproductive output if male ejaculate investment declines over consecutive matings. Accordingly, females are predicted to mate preferentially with virgin males. In mildly polyandrous lepidopterans, female fitness is less affected by reduced male investment than in more polyandrous species, and so the predictions for female mating preferences are less clear. We examined female mating preferences in the mildly polyandrous almond moth, Cadra cautella, in which ejaculate size does not affect female reproductive output. First, we allowed females to mate with virgin or once-mated males, in which the males were presented individually or simultaneously. We recorded the latency to mating and, in the case of the simultaneously presented trials, the identity of the successful, copulating male. We found that females mated more frequently with mated males (when simultaneously presented with both males), yet females did not differ in the time taken to initiate copulation with any male. We further examined if this mated male advantage was due to differential mate detection or locomotory behaviour of the male treatments. We tested the ability of virgin and mated males to locate a receptive female within a wind tunnel using long-distance pheromone cues and recorded their activity budget. We found no difference in the ability of mated or virgin males to locate or approach a receptive female, or in their activity levels. These data suggest a female preference for mated males in this species, a preference that may minimise other potential costs of mating.     

The downfall of mating: the effect of mate-carrying and flight muscle ratio on the escape ability of a pierid butterfly

In many species, the ability to evade predators is known to be periodically impaired by increased weight loads due to feeding and reproduction. Not only may extra weight reduce escape speed, but feeding and mating can also make the prey more noticeable to visually hunting predators. A number of butterfly species mate for hours, and if a mating couple is disturbed, one of the butterflies is responsible for flying, whereas its partner remains still. This study investigated the ability of male Pieris napi butterflies to fly while mating, with the prediction that mate carrying impairs flight ability compared to single flying males and that males with relatively high flight muscle ratios (FMR; male thorax mass/male + female body mass) will have better flight performance in copula. Our results clearly show that whereas single males always take off at steep angles and fly upwards, couples invariably have a negative take-off angle and rarely gain height. Moreover, landing height of the couples is positively associated with higher FMR. Hence, male flight ability when in copula is positively associated with a high relative thorax mass. Butterfly pairs may thus be at greater risk of predation as a consequence of their impaired flight ability, especially couples with critically low FMRs (<16%).     

The effect of perceived female parasite load on post-copulatory male choice in a sex-role-reversed pipefish

The last several decades of research in behavioral ecology have resulted in a deeper appreciation of post-mating processes and sexual conflict in sexual selection. One of the most controversial aspects of sexual selection is cryptic mate choice. Here, we take advantage of male pregnancy in a sex-role-reversed pipefish (Syngnathus typhle) to quantify cryptic choice based on perceived parasite load and other sources of variance in female fitness. Studies have shown that S. typhle males preferentially mate with females with lower parasite loads and that a male’s perception of female parasite load can be altered by tattooing females. We manipulated the apparent parasite load of females in controlled mating experiments to test the hypothesis that post-copulatory sexual selection is dependent on a male’s perception of female parasite load in pipefish. Our results provided no evidence for cryptic male choice based on perceived female parasite load. However, we found evidence that eggs from larger females were more likely to result in viable offspring than eggs from smaller females and that the first female to mate with a male transferred more eggs per copulation on average. Overall, our results show that potential for post-copulatory sexual selection does exist in pipefish, but the male’s perception of female parasite load does not play a major role in this process.     

Effects of mating history on ejaculate size, fecundity, longevity, and copulation duration in the ant-tended lycaenid butterfly, Jalmenus evagoras

The mating system of the Australian lycaenid butterfly, Jalmenus evagoras, is highly unusual compared to most other Lepidoptera. Characteristics of this system, which has been termed an ’explosive mating strategy,’ include the formation of an intensely competitive mating aggregation of males, a highly male biased operational sex ratio, a lack of discrimination and mate choice by both sexes, a high variance in male mating success, and female monogamy. In this study, we tested the hypothesis that multiple mating by males imposes physiological costs resulting in smaller spermatophores, and that this results in a fitness cost to females. We found that male J. evagoras transferred only 2.2% of their eclosion weight during their first mating, consistent with the hypothesis that males of monandrous species produce a relatively small investment. The wet weight of the ejaculate declined by an average of 27% at the second mating and the dry weight by 29%, and an intermating interval of 5–9 days was needed for the ejaculate to return to the size at the first mating, regardless of male size or age. Wet ejaculate mass increased proportionally with male size, though dry mass was proportionally larger in smaller males. Ejaculate mass tended to increase with male age at both first and second matings. Female characteristics, in general, did not affect ejaculate mass, although the wet weight of the ejaculate was positively associated with female weight at the second mating. Copulation duration increased from 2.4 h to approximately 3 h at the second mating, and to over 4 h at the third and fourth matings. Fecundity was positively correlated with female size but not with mating history, copulation duration, or any other characteristics measured for either males or females. Female longevity declined significantly as the number of times the male partner had previously mated increased. We conclude that despite the small male investment in ejaculate, the costs of multiple mating may nonetheless be significant, as indicated by the reduction in ejaculate mass, an increase in copulation duration, and reduction in female lifespan with increasing mating number. Received: 22 January 1999 / Received in revised form: 28 July 1999 / Accepted: 18 September 1999     

Audience effect alters male but not female mating preferences

Males often face strong mating competition by neighboring males in their social environment. A recent study by Plath et al. (Anim Behav 75:21–29, 2008a) has demonstrated that the visual presence of a male competitor (i.e., an audience male) affects the expression of male mating preferences in a poeciliid fish (Poecilia mexicana) with a weaker expression of mating preferences when an audience male observed the focal male. This may be a tactic to reduce sperm competition, since surrounding males likely share intrinsic preferences for female traits or copy mate choice decisions. Here, we examined the hypothesis that a same-sex audience would affect female mate preferences less than male mating preferences. Our hypothesis was based on the assumptions that (1) competition for mates in a fashion that would be comparable in strength to sperm competition or overt male–male aggression is absent among Poecilia females, and (2) P. mexicana females typically form female-biased shoals, such that almost any female mate choice in nature occurs in front of a female audience. Poecilia females (P. mexicana, surface and cave form, and the closely related gynogenetic Poecilia formosa) were given a choice between a large and a small male, and the tests were repeated while a conspecific, a heterospecific, or no audience female (control) was presented. Females spent more time in the neutral zone and, thus, less time near the males during the second part of a trial when an audience was presented, but—consistent with predictions—females showed only slightly weaker expression of mate preferences during the second part of the tests. This decline was not specific to the treatment involving an audience and was significantly weaker than the effect seen in the male sex.     

Pre- and post-mating sexual selection both favor large males in a rainbowfish

Sexual selection can act through female choice and male–male competition. Although both processes can act simultaneously, they are typically studied independently. Here, we adopt a more integrated approach to studying sexual selection by incorporating measures of both processes using the western rainbowfish Melanotaenia australis, a freshwater fish endemic to northwestern Australia. We assessed male–male competition and female choice separately while measuring the performance of individual males under both processes and used paternity analyses to estimate male reproductive success. We then related the performance of males during each of these stages to their phenotype, which was described using linear measures of size and color pattern traits, and spectrographic measures of the reflectance of color patches. We found that female choice favored relatively large males and that these preferences were consistent within individual females and repeatable between different females. Larger males were also more dominant in the competition trials and sired the majority of offspring produced when females spawned. There was little evidence to suggest that sexual selection acted on male color patterns either via female choice or male contest competition or during subsequent post-mating episodes of sexual selection. We conclude, therefore, that male–male competition and female choice act concordantly to favor relatively large males and that these patterns of mating success are reflected during post-mating episodes of sexual selection.