The decision to take risks in the presence of a predator involves complex trade-offs between immediate survival and future reproduction. Individuals may gain fitness advantages if they are able to optimally alter their risk-taking strategies depending on the differential costs and benefits of risky behaviours across contexts. Male fiddler crabs (Austruca mjoebergi) exhibited a higher propensity to take risks in the presence of a female compared with conspecifics that were not presented with a female during both mating and nonmating periods. Contrary to predictions, however, risk-taking behaviour did not differ between mating and nonmating periods.

Spatial and temporal variation in environmental factors and the social setting can help to maintain genetic variation in sexually selected traits if it affects the strength of directional selection. A key social parameter which affects the intensity of, and sometimes predicts the response to, mating competition is the operational sex ratio (OSR; ratio of receptive males to females). How the OSR affects selection for specific male traits is poorly understood. It is also unclear how sexual selection is affected by interactions between the OSR and environmental factors, such as habitat complexity, that alter key male-female interactions such as mate encounter rates. Here, we experimentally manipulated the OSR and habitat complexity and quantified sexual selection on male mosquitofish (Gambusia holbrooki) by directly measuring male reproductive success (i.e. paternity). We show that despite a more equitable sharing of paternity (i.e. higher levels of multiple paternity) under a male-biased OSR, selection on focal male traits was unaffected by the OSR or habitat complexity. Instead, sexual selection consistently, and significantly, favoured smaller bodied males, males with higher genome wide heterozygosity (based on >3,000 SNP markers) and males with a relatively long gonopodium (intromittent organ). Our results show that sexual selection on male body size, relative genital size and heterozygosity in this system is consistent across environments that vary in ecological parameters that are expected to influence mate encounter rates.

Sexual selection is a cornerstone of evolutionary theory, but measuring it has proved surprisingly difficult and controversial. Various proxy measures-e.g., the Bateman gradient and the opportunity for sexual selection-are widely used in empirical studies. However, we do not know how reliably these measures predict the strength of sexual selection across natural systems, and most perform poorly in theoretical worst-case scenarios. Here we provide a rigorous comparison of eight commonly used indexes of sexual selection. We simulated 500 biologically plausible mating systems, based on the templates of five well-studied species that cover a diverse range of reproductive life histories. We compared putative indexes to the actual strength of premating sexual selection, measured as the strength of selection on a simulated "mating trait." This method sidesteps a key weakness of empirical studies, which lack an appropriate yardstick against which proxy measures can be assessed. Our model predicts that, far from being useless, the best proxy measures reliably track the strength of sexual selection across biologically realistic scenarios. The maximum intensity of precopulatory sexual selection s'(max) (the Jones index) outperformed all other indexes and was highly correlated with the strength of sexual selection. In contrast, the Bateman gradient and the opportunity for sexual selection were poor predictors of sexual selection, despite their continuing popularity.

Environmental variation can maintain genetic variation in sexually selected traits if it affects the strength of directional selection. Specifically, environmental variation in sex-specific mortality will change the operational sex ratio (OSR), which predicts the intensity of mating competition. How the OSR affects selection for specific male traits is poorly understood; and it is unknown how often sexual selection is affected by interactions between the OSR and environmental factors that alter social variables such as mate encounter rates. Here, we experimentally manipulated the OSR and habitat complexity and quantified sexual selection on male mosquitofish ( Gambusia holbrooki ). In G. holbrooki there is high within-population variation in male size, which may exist because of a tradeoff between the ability to sneak copulate (favouring small males) and monopolize females (favouring large males). The success of each tactic is predicted to depend on the OSR, encounter rates and the ability to stealthily approach conspecifics. We show that, despite greater sharing of paternity under a male-biased OSR, neither the opportunity for selection, nor selection on male traits was affected by the OSR or habitat complexity. Instead, sexual selection consistently favored smaller males with high genetic heterozygosity (using >3000 SNP markers), and a relatively long gonopodium (intromittent organ).

A focus on novel, confirmatory, and statistically significant results leads to substantial bias in the scientific literature. One type of bias, known as "p-hacking," occurs when researchers collect or select data or statistical analyses until nonsignificant results become significant. Here, we use text-mining to demonstrate that p-hacking is widespread throughout science. We then illustrate how one can test for p-hacking when performing a meta-analysis and show that, while p-hacking is probably common, its effect seems to be weak relative to the real effect sizes being measured. This result suggests that p-hacking probably does not drastically alter scientific consensuses drawn from meta-analyses.

Generally, sex-specific mortality is not expected to affect optimal patterns of sex allocation. Several authors have, however, made verbal arguments that this is not true if juvenile mortality is sex specific during the period of parental care. Here, we provide formal mathematical models exploring the effect of such mortality on optimal sex allocation. We confirm the prediction that biased production of the sex with higher mortality during care is favoured. Crucially, however, this is only true when juvenile mortality in the period of parental care frees up resources for their current/future siblings (i.e. the saved investment is transferable). Furthermore, we show that although optimal sex allocation is consistent with the theory of equal investment (as asserted by previous authors), thinking in terms of equal investment is not readily feasible in some scenarios. We also show that differences in early mortality overcome biased sex allocation such that the sex ratio at independence is generally, but not always, biased in the opposite direction from that at birth. Our models should prove useful to empiricists investigating the effect of sex-specific juvenile mortality and antagonistic sibling interactions on sex allocation.

Mothers vary in their effects on their offspring, but studies of variation in maternal effects rarely ask whether differences between mothers are consistent for sons and daughters. Here, we analysed maternal effects in the mosquitofish Gambusia holbrooki for development time and adult size of sons and daughters, and a primary male sexual character (gonopodium length). We found substantial maternal effects on all traits, most notably for gonopodium length. There were significant correlations within each sex for maternal effects on different traits, indicative of trade-offs between development rate and adult size. By contrast, there was no evidence of any consistency in maternal effects on sons and daughters. This suggests that the evolution of maternal effects will follow independent trajectories dependent on sex-specific selection on offspring. Importantly, failure to recognize the sex-specific nature of maternal effects in this population would have substantially underestimated the extent of their variation between mothers.

In many taxa, temporary nutritional shortage early in development can favour compensatory strategies that include elevated growth (‘compensatory growth’) and/or extension of the usual period of development (‘catch-up growth’) once conditions improve. The net gains from each strategy depend on the extent to which larger body size increases fitness relative to associated costs (e.g. long-term effects on adult performance, or a greater risk of juvenile mortality). These costs and benefits are likely to differ between the sexes due to sex-specific selection. We documented the responses of male and female mosquitofish (Gambusia holbrooki) to 3 weeks of low food availability (7–28 days old) that restricted subsequent growth and morphology compared to control fish continuously reared on a high food diet (N = 635 fish total). Neither sex elevated their growth rate immediately after being returned to a normal diet compared to control fish. When measured over the entire period until maturation, however, females showed compensatory growth. Males did not. Both sexes also exhibited catch-up growth but the delay until maturation was significantly longer for males. Despite early growth restriction, both sexes eventually matured at almost the same size as control fish, although males had a significantly smaller gonopodium (a sexually selected trait) than that of control males. Reasons for these sex differences are discussed.

Biological signaling usually occurs in complex environments, yet signals are most often studied in controlled experiments that strip away this complexity. Male fiddler crabs possess one enlarged claw that is waved during courtship displays, and females preferentially respond to larger claws and faster waves. Fiddler crab vision is evolutionarily specialized to their predominantly level mudflat habitats, although some populations inhabit topographically complex environments. Here, we investigated how the elevation of signaling males relative to receiving females affects attractiveness. Experiments with robotic crabs revealed a strong female aversion to males signaling from atop small (>2cm) mud mounds. This aversion entirely masked previously documented strong preferences for large claws and faster waving. Our results suggest that variation in signaling environment might substantially weaken selection on males, thereby helping to maintain genetic variation in courtship traits.

Studies of sexual communication typically focus on the design and information content of a signal of interest, but the timing of signal production relative to nearby competitors can be crucial. Male fiddler crabs, Uca mjoebergi, court females with a stereotyped claw-waving display, and males are often observed waving in synchrony with nearby claw-waving males. Using female mate preference experiments with robots that imitate male claw waves, we found evidence that females are more attracted to males whose waves immediately precede a synchronous group of waves (leaders); females also favoured males that waved in opposite phase to a synchronous group (alternators). By contrast, males whose waves lagged behind a group of synchronous wavers (laggards) were no more attractive. We discuss a simple sensory process that could explain how this female preference arises. Our results agree with past findings suggesting that synchrony in fiddler crabs occurs as an epiphenomenon of adaptive male responses to female preferences.

Very few studies have presented compelling evidence for adaptive shifts in female mating preferences. Additionally, there is almost no data on how males adjust courtship effort when female mating preferences change predictably. How should males respond? Should a currently more attractive male increase his courtship effort because he has, for now, a better chance of reproducing? Or should he maintain/lower his courtship effort and conserve energy because he already has an edge on the competition? We experimentally measured female mating preferences and male courtship effort (i.e., male mating preferences) in the fiddler crab Uca mjoebergi. Using robotic males, we documented a consistent shift in female preferences for male claw size across each of 6 biweekly mating periods: females tested at the beginning of a mating period preferred large males, whereas those tested at the end preferred small males. This is one of the fastest-known temporal changes in the mean mating preference of a population and supports our prediction of an adaptive response due to time constraints on larval development. Males adjusted their courtship effort across the mating period in concert with the observed daily shift in the mean female mating preference. Interestingly, changes in courtship effort depended on male size. We interpret this shift as males increasing their courtship effort to take advantage of their current attractiveness to females. To our knowledge, this is the first study of the interactions between shifts in female and male mating preferences.

How should males respond to sexual rivals? We found evidence that male crickets increase their courtship calling effort when competing with a rival, but relax when their rival dies. Also, competing males died sooner than those that were free to court without a rival. These results highlight the importance of behavioral changes in courtship by males to life history strategies.Temporal variation in the strength of selection on male sexual traits is often attributable to changes in the social environment that alter the number of competitors. Selection could favor phenotypic plasticity in male investment into sexual traits if there are cues indicative of current and, possibly, future levels of mating competition. In many taxa, males court more intensely when rivals are present, but the extent to which phenotypically plastic responses differ predictably among males is less well studied. For example, will larger males show a greater or smaller change in courtship in response to the presence of rivals? In addition, the effects of any changes in courtship on key life-history traits have been understudied. In this study, we experimentally tested male crickets (Teleogryllus commodus) from 3 populations to determine how the presence or absence of a rival affects: 1) calling effort, 2) life span, and 3) whether male body size (correlated with dominance) influences any changes in life span or calling effort. Calling effort increased significantly with body size, mainly due to daily calling effort increasing with age and larger males living longer. Considering all males, there was no effect of rival presence on lifetime calling effort. However, within the rival present treatment, after correcting for age, the longer lived of 2 paired males called significantly more before his rival died than afterwards. This implies that there is a plastic shift in courtship effort. Finally, larger males lived significantly longer and, crucially, males housed with a rival had, on average, a significantly shorter life span.

Sperm competition and uncertainty of paternity hamper the evolution of male parental care. Thus, maternal care predominates in most taxa. What if males can, however, limit cuckoldry by guarding the eggs postmating? Here, we show that this provides a reason to reconsider an old and nowadays rather discredited hypothesis: that external fertilization is associated with male care because the parent who releases its gametes first can depart leaving the other in a "cruel bind," having to care for the offspring. In our model, protection of paternity provides an additional incentive for the male to stay associated with its young. When we then assume that offspring survive better if guarded, paternity protection proves enough to kick-start the evolution of male-only parental care from a scenario with no care. This fits with data from fishes, where male-only care is associated with external fertilization, whereas female-only care almost always evolves after an initial transition to internal fertilization. Our model unifies disparate hypotheses regarding parental care roles and provides support for the idea that care roles can be influenced by sex differences in selection to be physically close to the offspring, including selection that is initially not based on offspring survival.

Males often possess traits, such as horns, claws, and tusks, which are used during male–male combat. Studies suggest that selection has fine tuned these weapons to improve their effectiveness and that the shape of these weapons provides cues for males to assess the strength of rivals. Similarly, females might also assess a male’s weaponry to determine his value as a mate. The largest weapon relative to body size is the major claw of male fiddler crabs (genus: Uca). Males use their claws both as a weapon and as a courtship signal, waving it to attract females. We used robotic crabs in two-choice experiments to test female mating preferences based on male claw shape in Uca mjoebergi. First, females did not prefer conspecific claws over any of three alternatively shaped stimuli matched for color and for the rate and pattern of waving. The alternative stimuli were two different heterospecific claws and a plain rectangular shape. Second, females significantly preferred the alternative stimuli when they were presented at a faster wave rate. We conclude that claw shape in U. mjoebergi has not evolved under sexual selection imposed by female mating preferences and is more likely to have evolved under selection for effectiveness during male–male competition.

Sex allocation theory explains why most species produce equal numbers of sons and daughters, and highlights situations that select for deviation from this norm. Past research has, however, heavily focused on situations with discrete generations. When temporally varying generational overlap affects future mate availability, models predict cyclical shifts in sex allocation, but these predictions have not yet been appropriately tested. Here we provide evidence that mosquitofish (Gambusia holbrooki) populations possess a suitable life history: some autumn-born females bred alongside their own offspring, while such overlap was rare or absent for spring-born females and for all males. Our analytic model of sex allocation for these populations produced a perfect rank-order correlation between observed birth sex ratio biases and theoretical predictions, with stronger biases observed as the extent of female generational overlap increased. This is the first robust evidence that sex allocation theory accounts for cases when mating opportunities vary predictably over time.

A poor start in life owing to a restricted diet can have readily detectable detrimental consequences for many adult life-history traits. However, some costs such as smaller adult body size are potentially eliminated when individuals modify their development. For example, male mosquitofish (Gambusia holbrooki) that have reduced early food intake undergo compensatory growth and delay maturation so that they eventually mature at the same size as males that develop normally. But do subtle effects of a poor start persist? Specifically, does a male's developmental history affect his subsequent attractiveness to females? Females prefer to associate with larger males but, controlling for body length, we show that females spent less time in association with males that underwent compensatory growth than with males that developed normally.

The action of sexual selection is highly variable among taxa. This creates challenges when trying to generalize (e.g. determine if a particular relationship exists based on its average strength, or if it varies in response to theoretically relevant factors). Consequently, accounting for moderating factors is likely to be crucial to explain differences in sexual selection among studies. In principle, given measures of key theoretical parameters we can predict the strength of sexual selection on different sexual signals, the benefits of mate choice, the extent of sex differences (e.g. in immune function or survival) and the likely life history trade-offs between investment into different sexual traits (e.g. sperm vs. courtship) or non-sexual traits (e.g. immune function, traits that increase longevity, parental care). How well does empirical data support theoretical expectations? First, we provide a short history of the use of meta-analysis in sexual selection studies. We present a table summarizing 94 meta-analyses that have asked questions about sexual selection or allied topics of interest to those studying sexual selection (e.g. the link between heterozygosity and fitness). Second, we list the main ways that meta-analysis has been used in sexual selection work and provide illustrative examples. Third, we provide practical advice to identify questions that are ripe for meta-analysis. We highlight 11 sexual selection topics where meta-analyses are needed (e.g. there are no meta-analyses testing game theory models of fighting contests). Finally, we discuss some general issues that will arise as the use of meta-analysis in sexual selection studies becomes more sophisticated.

Sexual selection is a major force behind the rapid evolution of male genital morphology among species. Most within-species studies have focused on sexual selection on male genital traits owing to events during or after copulation that increase a male's share of paternity. Very little attention has been given to whether genitalia are visual signals that cause males to vary in their attractiveness to females and are therefore under pre-copulatory sexual selection. Here we show that, on average, female eastern mosquitofish Gambusia holbrooki spent more time in association with males who received only a slight reduction in the length of the intromittent organ ('gonopodium') than males that received a greater reduction. This preference was, however, only expressed when females chose between two large males; for small males, there was no effect of genital size on female association time.