Intensity Of Sperm Competition Research Articles

Sperm competition intensity shapes divergence in both sperm morphology and reproductive genes across murine rodents.

It remains unclear how variation in the intensity of sperm competition shapes phenotypic and molecular evolution across clades. Mice and rats in the subfamily Murinae are a rapid radiation exhibiting incredible diversity in sperm morphology and production. We combined phenotypic and genomic data to perform phylogenetic comparisons of male reproductive traits and genes across 78 murine species. We identified several shifts towards smaller relative testes mass, presumably reflecting reduced sperm competition. Several sperm traits were associated with relative testes mass, suggesting that mating system evolution selects for convergent suites of traits related to sperm competitive ability. We predicted that sperm competition would also drive more rapid molecular divergence in species with large testes. Contrary to this, we found that many spermatogenesis genes evolved more rapidly in species with smaller relative testes mass due to relaxed purifying selection. While some reproductive genes evolved rapidly under recurrent positive selection, relaxed selection played a greater role in underlying rapid evolution in small testes species. Our work demonstrates that postcopulatory sexual selection can impose strong purifying selection shaping the evolution of male reproduction, and that broad patterns of molecular evolution may help identify genes that contribute to male fertility.

Sperm competition intensity shapes divergence in both sperm morphology and reproductive genes across murine rodents.

It remains unclear how variation in the intensity of sperm competition shapes phenotypic and molecular evolution across clades. Mice and rats in the subfamily Murinae are a rapid radiation exhibiting incredible diversity in sperm morphology and production. We combined phenotypic and genomic data to perform phylogenetic comparisons of male reproductive traits and genes across 78 murine species. We identified several shifts towards smaller relative testes mass, presumably reflecting reduced sperm competition. Several sperm traits were associated with relative testes mass, suggesting that mating system evolution selects for convergent suites of traits related to sperm competitive ability. We predicted that sperm competition would also drive more rapid molecular divergence in species with large testes. Contrary to this, we found that many spermatogenesis genes evolved more rapidly in species with smaller relative testes mass due to relaxed purifying selection. While some reproductive genes evolved rapidly under recurrent positive selection, relaxed selection played a greater role in underlying rapid evolution in small testes species. Our work demonstrates that postcopulatory sexual selection can impose strong purifying selection shaping the evolution of male reproduction, and that broad patterns of molecular evolution may help identify genes that contribute to male fertility.

The transfer of male cuticular hydrocarbons provides a reliable cue of the risk and intensity of sperm competition in decorated crickets.

Theoretically, males should increase their ejaculate expenditure when the probability of sperm competition occurring (or risk) is high but decrease ejaculate expenditure as the number of competing ejaculates (or intensity) increases. Here we examine whether male decorated crickets (Gryllodes sigillatus) use cuticular hydrocarbons (CHCs) transferred to females by rival males at mating to assess the risk and intensity of sperm competition and adjust their ejaculate accordingly. Unmated females and those perfumed with CHCs extracted from one, three, or five males could be distinguished chemically, providing a reliable cue of the risk and intensity of sperm competition. In agreement with theory, males mating with these females increased sperm number with the risk of sperm competition and decreased sperm number with the intensity of sperm competition. Similarly, as the risk of sperm competition increased, males produced a larger and more attractive spermatophylax (an important non-sperm component of the ejaculate) but these traits did not vary with the intensity of sperm competition. Our results therefore demonstrate that both sperm and non-sperm components of the male ejaculate respond to the risk and intensity of sperm competition in different ways and that CHCs provide males with an important cue to strategically tailor their ejaculate.

Accessory gland size increases with sperm competition intensity in Cataglyphis desert ants

In many species, females have multiple mates, whose sperm compete for paternity. Males may subsequently invest in the increased production of sperm and/or seminal fluid. The latter is a complex mixture of proteins, peptides, and other compounds generated by the accessory glands (AGs) and is transferred to females along with a male’s sperm. Seminal fluid is known to be a key determinant of competitive outcomes among sperm, and its production may trade off with that of sperm. We show that AG size—a proxy for seminal fluid production—has a positive and phylogenetically robust correlation with both sperm competition intensity and sperm production in nine species of Cataglyphis desert ants. These results indicate a lack of trade-off between sperm production and seminal fluid production. They underscore that sperm competition may strongly shape sperm traits and could drive reproductive performance in eusocial hymenopterans.

Differences in sperm morphology between Alouatta palliata and Alouatta pigra are consistent with the intensity of sperm competition in each species.

The intensity of sperm competition, in which sperm compete within the female reproductive tract to reach and fertilize her eggs, varies in species with different mating systems. Sperm competition is more intense in species where males cannot monopolize access to reproductive females and females mate with multiple males. In this scenario, a morphological change that increases the ability of sperm to reach and fertilize eggs should rapidly spread in the population, leading to sperm morphological differences between closely related species. Differences in sperm morphology have been reported among primate species with different mating systems. However, due to the inherent logistical and ethical difficulties to sample sperm from wild primates, the extent of variation in sperm morphology within species and among closely related species remains understudied. Here, we compared sperm morphological traits from two sister howler monkey species (Alouatta palliata and Alouatta pigra) that have different mating systems to investigate the effect of sperm competition on sperm morphological traits. We predicted that sperm from A. palliata, where females have more opportunities to mate with multiple males, would show differences in traits associated with increase sperm competitiveness compared to A. pigra where females mostly mate with the central male. We used linear mixed models to determine species differences in sperm morphology, controlling for individual variation. We found that midpieces and heads in A. palliata sperm were on average 26.2% and 11.0% longer, respectively, than those of A. pigra. Differences in these traits are important for sperm speed and hydrodynamic movement in other species and can affect fertilization success. This study provides empirical evidence of sperm morphological traits that evolved through sexual selection in sister primate species with different mating systems.

Group size and mating system predict sex differences in vocal fundamental frequency in anthropoid primates

Vocalizations differ substantially between the sexes in many primates, and low-frequency male vocalizations may be favored by sexual selection because they intimidate rivals and/or attract mates. Sexual dimorphism in fundamental frequency may be more pronounced in species with more intense male mating competition and in those with large group size, where social knowledge is limited and efficient judgment of potential mates and competitors is crucial. These non-mutually exclusive explanations have not been tested simultaneously across primate species. In a sample of vocalizations (n = 1914 recordings) across 37 anthropoid species, we investigated whether fundamental frequency dimorphism evolved in association with increased intensity of mating competition (H1), large group size (H2), multilevel social organization (H3), a trade-off against the intensity of sperm competition (H4), and/or poor acoustic habitats (H5), controlling for phylogeny and body size dimorphism. We show that fundamental frequency dimorphism increased in evolutionary transitions towards larger group size and polygyny. Findings suggest that low-frequency male vocalizations in primates may have been driven by selection to win mating opportunities by avoiding costly fights and may be more important in larger groups, where limited social knowledge affords advantages to rapid assessment of status and threat potential via conspicuous secondary sexual characteristics.

Social Environment Influences the Temporal Dynamics of Sneak-Spawning in a Fish with Alternative Reproductive Tactics.

AbstractSeveral predictions of sperm competition theory are not well supported empirically. One potential reason is that most current theory and empirical research ignore how the social environment influence the temporal dynamics of mating. We propose that understanding these dynamics is key to understanding sexual selection and improving the predictive power of theory. To demonstrate the importance of these dynamics, we quantify how males' social role, interactions among males, and current social environment influence the timing of mating in Symphodus ocellatus, a species with three alternative male reproductive tactics. Nesting males spawn synchronously with females; sneakers and satellites sneak-spawn with some time delay. Satellites also cooperate with nesting males. We found that satellites have shorter sneak-spawning delays than sneakers, a benefit of their cooperation with nesting males. Sneak-spawning delays decreased with increasing nest activity for sneakers but not for satellites, suggesting that sneakers may benefit from increased sperm competition intensity. Current sperm competition models ignore this potential benefit, which may be why the prediction that males should decrease investment when sperm competition involves more than two males is not well supported. Our study provides insight into mechanisms that drive variation in the timing of spawning, which could explain mismatches between theoretical and empirical results.

Male and female Brandt’s voles have higher reproductive success when they have more mating partners regardless of population density

Abstract Although population density and body mass are known to alter the mating strategies of individuals, their impacts on reproductive success have not been fully investigated. We examined genetic mating system variation (estimated number of mating partners), reproductive success (offspring number), and sperm competition intensity (sires per litter) in semi-natural populations of Brandt’s voles (Lasiopodomys brandtii) under low-, medium- and high-densities. We used 12 microsatellite genetic markers and parentage assignment of 3516 voles, with the aim of identifying the estimated number of mating partners and offspring produced by founder individuals. We provided strong evidence that individuals with more mating partners experienced increased reproductive success in all density groups, as measured by the production of more offspring, compared with individuals that had fewer mating partners. Further, we found that large males had more mating partners (especially in high-population density enclosures) and also produced more offspring relative to smaller males. In high-density (HD) enclosures, the average reproductive success per female vole was lower (a negative density-dependent effect), but voles had more mating partners, more sires per litter, but lower paternity skew compared to low-density enclosures. Our results suggests that Brandt’s voles increased reproductive success through increasing mating partners under HD stress which may have significant implications in offsetting the negative density-dependency effect on population growth.

Male position in a sexual network reflects mating role and body size

Several species are characterised by male mating polymorphisms, which are often associated with sperm competition for which some phenotypes show specialisation. With high-resolution behavioural data, we used network analysis to quantify sperm competition intensity; the probability of overlap of the ejaculates of different males among competing male European bitterling (Rhodeus amarus) in a large experimental mesocosm. Implementing Bayesian inference with informative priors, we modelled sperm competition intensity among males adopting two alternative mating roles to understand how sperm competition intensity varied between roles as a function of body size. We demonstrated that larger males performing a “guarder” role experienced lower sperm competition intensity than smaller males. In contrast, for “sneaker males”, larger males experienced greater sperm competition intensity. We discuss these findings in the context of mating system evolution and the reproductive biology of bitterling.

Sperm competition increases sperm production and quality in Cataglyphis desert ants.

Sperm competition is a pervasive evolutionary force that shapes sperm traits to maximize fertilization success. Indeed, it has been shown to increase sperm production in both vertebrates and invertebrates. However, sperm production is energetically costly, which may result in trade-offs among sperm traits. In eusocial hymenopterans, such as ants, mating dynamics impose unique selective pressures on ejaculate. Males are sperm limited: they enter adulthood with a fixed amount of sperm that will not be renewed. We explored whether sperm competition intensity was associated with sperm quantity and quality (i.e. sperm viability and DNA fragmentation) in nine Cataglyphis desert ants. Our results provide phylogenetically robust evidence that sperm competition is positively correlated with sperm production and sperm viability. However, it was unrelated to sperm DNA integrity, indicating the absence of a trade-off involving this trait. These findings underscore that sperm competition may strongly mould sperm traits and drive reproductive performance in eusocial Hymenoptera.

How Female-Female Competition Affects Male-Male Competition: Insights into Postcopulatory Sexual Selection from Socially Polyandrous Species.

AbstractSexual selection is a major driver of trait variation, and the intensity of male competition for mating opportunities has been linked with sperm size across diverse taxa. Mating competition among females may also shape the evolution of sperm traits, but the effect of the interplay between female-female competition and male-male competition on sperm morphology is not well understood. We evaluated variation in sperm morphology in two species with socially polyandrous mating systems, in which females compete to mate with multiple males. Northern jacanas (Jacana spinosa) and wattled jacanas (J. jacana) vary in their degree of social polyandry and sexual dimorphism, suggesting species differences in the intensity of sexual selection. We compared mean and variance in sperm head, midpiece, and tail length between species and breeding stages because these measures have been associated with the intensity of sperm competition. We found that the species with greater polyandry, northern jacana, has sperm with longer midpieces and tails as well as marginally lower intraejaculate variation in tail length. Intraejaculate variation was also significantly lower in copulating males than in incubating males, suggesting flexibility in sperm production as males cycle between breeding stages. Our results indicate that stronger female-female competition for mating opportunities may also shape more intense male-male competition by selecting for longer and less variable sperm traits. These findings extend frameworks developed in socially monogamous species to reveal that sperm competition may be an important evolutionary force layered atop female-female competition for mates.

Loss of Avian Intromittent Organs as a Sperm Competition Strategy: A Race to Be Last.

AbstractThe loss of the intromittent organ (IO) in the majority of birds remains unexplained. Here, I propose that IO loss results from sperm competition in the context of the unique avian sperm storage system. The first stage of fertilization is the movement of sperm through the vagina from the site of ejaculation to the sperm storage tubules (SSTs) at the uterovaginal junction. In a second stage, sperm are released from the SSTs and move through the uterus to the site of fertilization in the infundibulum. Last-male fertilization precedence can occur if sperm that arrive later at the uterovaginal junction occupy uterus-side SSTs, which then have a head start in the race to the infundibulum with each ovulation. Under such "last-in, first-out" conditions, there is strong sperm competition to arrive later at the SSTs. Consequently, the optimal male strategy would be placement of the ejaculate at the cloacal opening to allow any other sperm in the vagina to reach the SSTs first. Cloacal placement is effectively achieved by loss of the IO. The evolution of altricial development in Neoaves, the largest clade that lacks IOs, created conditions that favor IO loss. Specifically, the smaller clutch sizes and hatching asynchrony of altricial birds increase the intensity of sperm competition for fertilization of early eggs in the laying sequence and thus the selective advantage of later arrival at the SSTs. The rarity of IO loss among all animals suggests that the complex mechanism of avian fertilization creates unique conditions for sperm competition.

Does diet influence ejaculate expenditure under experimentally altered risk of sperm competition in guppies?

According to the sperm competition risk (SCR) model, males should increase their investment in ejaculates when there is a high chance that their sperm will compete for fertilization with those from another male. However, as ejaculates are energetically costly to produce, the extent to which males can adjust ejaculate investment may depend on the availability of metabolic resources. We tested these ideas using the guppy, Poecilia reticulata, a livebearing fish known to exhibit strong condition dependence in ejaculate traits and among the highest recorded polyandry rates in any vertebrate. Specifically, we manipulated SCR by visually exposing mature males to females either in the presence (high SCR) or absence (low SCR) of a rival male over 1 month, which comprises a full spermatogenic cycle. To test for condition dependence in male response to SCR, both groups were further subdivided into two diet quantity treatments (high and low), enabling us to test fixed and interactive effects of SCR and diet treatment on ejaculate expenditure (sperm production, sperm length and sperm velocity). Contrary to our predictions, we found no evidence that males increase ejaculate expenditure under high SCR, and indeed report the opposite pattern (higher expenditure under low SCR) for sperm velocity. Furthermore, we found no evidence that dietary restriction (when administered to mature adult males over 1 month) affected any ejaculate traits, leading us to suspect that ejaculates may only be susceptible to dietary restriction effects during a critical ontogenetic stage of development. We discuss these unexpected findings in the context of guppy biology, which may render the species less likely to exhibit long-term adjustments in ejaculates when perceiving short-term changes in SCR. Moreover, the high polyandry rates in this species may mean that the visual presence of a rival is a cue of altered sperm competition intensity, rather than risk.

Sperm competition intensity affects sperm precedence patterns in a polyandrous gift-giving spider.

Sperm competition drives traits that enhance fertilization success. The amount of sperm transferred relative to competitors is key for attaining paternity. Female reproductive morphology and male mating order may also influence fertilization, however the outcome for sperm precedence under intense sperm competition remains poorly understood. In the polyandrous spider Pisaura mirabilis, males offer nuptial gifts which prolong copulation and increase sperm transfer, factors proposed to alter sperm precedence patterns under strong sperm competition. First, we assessed the degree of female polyandry by genotyping wild broods. A conservative analysis identified up to four sires, with a mean of two sires per brood, consistent with an optimal mating female rate. Then we asked whether intense sperm competition shifts sperm precedence patterns from first male priority, as expected from female morphology, to last male advantage. We varied sexual selection intensity experimentally and determined competitive fertilization outcome by genotyping broods. In double matings, one male monopolised paternity regardless of mating order. A mating order effect with first male priority was revealed when females were mated to four males, however this effect disappeared when females were mated to six males, probably due to increased sperm mixing. The proportion of males that successfully sired offspring drastically decreased with the number of competitors. Longer copulations translated into higher paternity shares independently of mating order, reinforcing the advantage of traits that prolong copulation duration under intense competition, such as the nuptial gift. Sperm competition intensity enhances the impact of competitive sexual traits and imposes multiple effects on paternity.

Mating behavior and reproductive morphology predict macroevolution of sex allocation in hermaphroditic flatworms

BackgroundSex allocation is the distribution of resources to male or female reproduction. In hermaphrodites, this concerns an individual’s resource allocation to, for example, the production of male or female gametes. Macroevolutionary studies across hermaphroditic plants have revealed that the self-pollination rate and the pollination mode are strong predictors of sex allocation. Consequently, we expect similar factors such as the selfing rate and aspects of the reproductive biology, like the mating behaviour and the intensity of postcopulatory sexual selection, to predict sex allocation in hermaphroditic animals. However, comparative work on hermaphroditic animals is limited. Here, we study sex allocation in 120 species of the hermaphroditic free-living flatworm genus Macrostomum. We ask how hypodermic insemination, a convergently evolved mating behaviour where sperm are traumatically injected through the partner’s epidermis, affects the evolution of sex allocation. We also test the commonly-made assumption that investment into male and female reproduction should trade-off. Finally, we ask if morphological indicators of the intensity of postcopulatory sexual selection (female genital complexity, male copulatory organ length, and sperm length) can predict sex allocation.ResultsWe find that the repeated evolution of hypodermic insemination predicts a more female-biased sex allocation (i.e., a relative shift towards female allocation). Moreover, transcriptome-based estimates of heterozygosity reveal reduced heterozygosity in hypodermically mating species, indicating that this mating behavior is linked to increased selfing or biparental inbreeding. Therefore, hypodermic insemination could represent a selfing syndrome. Furthermore, across the genus, allocation to male and female gametes is negatively related, and larger species have a more female-biased sex allocation. Finally, increased female genital complexity, longer sperm, and a longer male copulatory organ predict a more male-biased sex allocation.ConclusionsSelfing syndromes have repeatedly originated in plants. Remarkably, this macroevolutionary pattern is replicated in Macrostomum flatworms and linked to repeated shifts in reproductive behavior. We also find a trade-off between male and female reproduction, a fundamental assumption of most theories of sex allocation. Beyond that, no theory predicts a more female-biased allocation in larger species, suggesting avenues for future work. Finally, morphological indicators of more intense postcopulatory sexual selection appear to predict more intense sperm competition.

Cannibalism and protective behavior of eggs in Arctic charr (Salvelinus alpinus).

From video recordings of spawning events, we quantified protective and cannibalistic behavior of Arctic charr occurring immediately after spawning. The number of fish cannibalizing on stray eggs was examined regarding (a) whether more than one male shed milt during the spawning event, that is, whether sperm competition occurred, (b) whether the sperm competition included few or many males, that is, the intensity of sperm competition, and (c) the density of fish at the spawning site. Response behavior toward egg cannibalism was also examined among females and dominant males in order to determine any parental investment toward protecting the eggs after spawning. Cannibalistic behavior was seen in almost 50% of the spawnings, and the multiple spawning events showed the highest numbers of fish cannibalizing on eggs. Both the number of males releasing milt and the number of fish approaching the spawning site were positively correlated with egg cannibalism. Sperm competition was, however, not a prerequisite for egg cannibalism. Although we also observed partial filial cannibalism, protective behavior of eggs was seen both among dominant males and females, suggesting that charr actually conduct parental care.

Evolution of nuptial gifts and its coevolutionary dynamics with male-like persistence traits of females for multiple mating

BackgroundMany male animals donate nutritive materials during courtship or mating to their female mates. Donation of large-sized gifts, though costly to prepare, can result in increased sperm transfer during mating and delayed remating of the females, resulting in higher paternity. Nuptial gifting sometimes causes severe female-female competition for obtaining gifts (i.e., sex-role reversal in mate competition) and selection on females to increase their mating rate, changing the intensity of sperm competition and the resultant paternity gains. We built a theoretical model to simulate such coevolutionary feedbacks between nuptial gift size (male trait) and propensity for multiple mating (female trait). Donation of nuptial gifts sometimes causes development of female persistence trait for gift acquisition. We also analyzed the causes and consequences of this type of traits, taking double receptacles for nutritious seminal gifts, which are known to occur in an insect group with a “female penis” (Neotrogla spp.), as an illustrative example.ResultsOur individual-based simulations demonstrated that female-female competition for male-derived nutrients always occur when the environment is oligotrophic and mating costs are low for females. However, a positive correlation between donated gift size and the resultant paternity gain was a requisite for the co-occurrence of large gifts and females’ competitive multiple mating for the gifts. When gift donation satisfied female demands and thus resulted in monandry, exaggeration of nuptial gift size also occurred under the assumption that the last male monopolizes paternity. The evolution of double slots for gift acquisition and digestion (female persistence trait) always occurred when males could not satisfy the demands of females for gifts. However, through coevolutionary reduction in male gift size, fixation of this trait in a population drastically reduced the average female fitness.ConclusionSperm usage patterns, which have rarely been examined for animals with nuptial gifts, can be a critical factor for determining the extent of exaggeration in nuptial gifting. Sex-role reversals in mate competition, as a result of donation of nuptial gifts from males to females, can involve the evolution of male-like, persistent traits in females that reduce population productivity, as is the case with persistence traits in males.

Exploring the effects of extreme polyandry on estimates of sexual selection and reproductive success

Abstract Multiple mating by females can dramatically alter selection on males by creating indirect interactions between rivals via sperm competition. Exactly how this behavior alters the relationship between male mating and fertilization success depends on multiple factors: re-mating frequency, sperm usage patterns, and mating assortment (the extent to which the most promiscuous individuals mate with each other). Here, we explore the role these elements play in determining sexual selection in a highly polygyandrous species, the squash bug Anasa tristis. Using replicated semi-natural enclosures, in which individuals were able to freely interact for a 2-week period, we tracked matings between individuals and subsequent female offspring production. Multiple mating was extremely common, resulting in very high levels of sperm competition intensity. However, network analysis revealed that the most promiscuous males mated with less polyandrous females, and therefore experienced lower levels of sperm competition. As a result, estimated male reproductive success increased with mating success, but this relationship varied according to the mode of sperm utilization with which it was calculated. Furthermore, females with more mating partners produced more offspring, suggesting they also benefit from mating multiply. Our findings highlight that polyandry has numerous and complex effects on sexual selection which may only be exposed when examined under naturalistic conditions.

Intraspecific variation in testis organisation and sperm head morphology of the delicate mouse (

The delicate mouse (Pseudomys delicatulus) ranges from Queensland to northern Western Australia. In this study the morphology of the cellular organisation of the testis and cauda epididymal spermatozoa are compared. Individuals from the mainland of the Northern Territory and Western Australia invariably have only 60–70% of the testes composed of seminiferous tubules with the interstitial tissue between the tubules containing abundant lipid rich Leydig cells, whereas the cauda epididymal sperm have highly polymorphic, often pear-shaped, heads and basally attached tails. In individuals from Queensland the seminiferous tubules make up approximately 80% of the testis, whereas in the cauda epididymides the sperm populations are generally less variable and have bilaterally flattened heads with the tail attached to the lower concave surface. These differences in the morphology of the delicate mouse testis and spermatozoa in these two geographic regions suggest differences in intensity of intermale sperm competition, with individuals from northern Western Australia and the mainland of the Northern Territory exhibiting monogamy whereas those from Queensland may exhibit some degree of intermale sperm competition and hence possibly have a polyandrous or promiscuous mating system. These findings support the suggestion that P. delicatulus, as currently recognised, contains at least one cryptic species.

Sexual selection and personality: Individual and group-level effects on mating behaviour in red junglefowl.

Despite increasing evidence of the importance of repeatable among-individual differences in behaviour (animal personality) in ecology and evolution, little remains known about the role of animal personalities in sexual selection. Here, we present an investigation of the hypothesis that the personalities of individuals and their sexual partners play a role in different episodes of sexual selection, and the extent to which these effects are modulated by the social environment. We first examined how two repeatable behaviours-exploration and boldness-are associated with pre- and postcopulatory sexual selection in male red junglefowl Gallus gallus, using replicate groups across three experimental sex ratio treatments. We further explored how the social environment modulates relationships between male personality and mating performance, and whether mating is assortative or disassortative with respect to exploration or boldness. Finally, we examined behavioural mechanisms linking personality with mating performance. Across all sex ratios, the fastest and slowest exploring males courted females proportionally less, and faster exploring males associated with females more and received more sexual solicitations. In female-biased groups, the fastest and slowest exploring males experienced the highest mating success and lowest sperm competition intensity. Faster exploring males also obtained more mates in female-biased groups when their competitors were, on average, slower exploring, and the proportion of matings obtained by fast-exploring males decreased with the proportion of fast-exploring males in a group, consistent with negative frequency-dependent sexual selection. While boldness did not predict mating performance, there was a tendency for individuals to mate disassortatively with respect to boldness. Collectively, our results suggest that male exploration can play a role in sexual selection, and that sexual selection on personality is complex and contingent on the social environment.