Social information use helps animals to rapidly acquire biologically relevant information, but it can increase competition or be erroneous. The "costly information' hypothesis proposes that animals especially rely on social information in high-risk contexts such as under predation pressure. However, whether predation drives the evolution of social learning remains enigmatic. Here we use artificial selection lines of Trinidadian guppies (Poecilia reticulata) that had been selected for predation survival for 3 generations (and control lines that were visually and olfactory exposed to predation but in a secured enclosure) to test how predation selects for social information use. In our study, social information use was assessed in a low-risk foraging context, which allows us to isolate evolved differences from acute responses under high-risk. Individuals observed a demonstrator on a screen solving a color association task, correctly or incorrectly, before attempting the task themselves. We show that individuals from both treatments learned the task, but only animals from the control lines relied on the use of social information during the process, while animals from the predation lines prioritized private information. Our results challenge the assumptions that social information use has adaptive benefits under increased predation risk. On the contrary, we propose that under high-predation certain social learning mechanisms may incur excessive costs due to, for instance, extensive acquisition time. We suggest that in a high-predation environment, disregarding social information and relying on personal information can be an adaptive strategy.

In animals, mature sperm are stored in males before ejaculation and sometimes in females before fertilization. Sperm storage provides evolutionary advantages; however, storage can also cause sperm deterioration owing to post-meiotic sperm senescence (PMSS). Yet, the extent of such deterioration, the mechanisms driving it and its fitness consequences remain poorly understood. We perform a meta-analysis across human (115 studies) and non-human animals (56 studies, 30 species) to understand the impacts of in vivo sperm storage. In men, we find that storage via sexual abstinence increases sperm oxidative stress and DNA damage, while reducing sperm viability and motility. In non-human animals, sperm storage in males or females reduces sperm performance, including fertilization success and embryo quality. In men, the method used for sampling individuals—and in other animals, the sex-specific duration of storage—modulates observed outcomes. Our results highlight the mechanisms of sperm deterioration during storage and its consequences for pre- and post-fertilization outcomes. These have important implications for fertility clinics, sperm selection, captive breeding and understanding evolved adaptations that mitigate stored-sperm deterioration. Importantly, our results can help to optimize the timing of ejaculate collection and storage.

A father's age at conception is predicted to affect not only his own fertility but also his offspring's fitness. Offspring born to old fathers are assumed to be less fit than those of young fathers. However, under low mating rates, paternal age might be confounded with the duration for which mature sperm are stored in fathers prior to ejaculation. Studies that disentangle the confounding paternal effects of sperm storage duration from those of age on offspring are lacking. We use Drosophila melanogaster to test the separate and interactive effects of paternal age and sexual rest on offspring fitness. As expected, old fathers produce fewer offspring than young fathers; however, paternal age does not influence the survival or age-dependent reproductive success of sons or daughters. Instead, a long duration of paternal sexual rest negatively impacts the reproductive success of the conceived sons. Furthermore, daughters of low reproductive quality selectively disappear with age, but sons do not, highlighting that demographic processes can further modulate paternal age effects. Overall, we highlight that paternal age effects might not be as pervasive as previously assumed and suggest that paternal sexual rest might be more important in influencing offspring phenotypes.

Predation is a major evolutionary driver of life history and morphology. However, whether these traits evolve directly via predation or indirect effects is largely unresolved. We used artificial selection to experimentally test the impact of adult predation on the evolution of life history and morphology in guppies (Poecilia reticulata). We found that, compared to control fish, predation-selected fish produced larger offspring and larger broods early in life. However, other life history parameters, such as interbrood interval and total number of offspring, showed no response. We also found that predation selected for smaller and lighter females and for shorter tails and gonopodia in males, with no effect on body coloration. Our results show that while several traits evolve fast under selection on adult predation, several "classic" predation-dependent traits seem unaffected by predation selection. By comparing our experimental results to those from natural populations, we can disentangle the contribution of direct and indirect effects on trait evolution under predation pressure.

In many animals, sperm are stored for extended periods either in the reproductive tracts of males before ejaculation, or of females after copulation. Sperm storage reduces the risk of sperm limitation in both sexes and avoids the costs of female re-mating. However, sperm storage can lead to post-meiotic sperm senescence, i.e. within-sperm-age-dependent deterioration, potentially impacting conceived offspring and lowering male and female fitness. Yet, the extent and magnitude of such deterioration and the variables modulating it during sperm storage are not well understood. Using a meta-analysis across humans (115 studies) and non-human animals (56 studies from 30 species), we investigate how in-vivo sperm storage affects sperm quality, fertilisation success, and offspring quality. In humans, sperm storage leads to greater sperm oxidative stress and DNA damage, and reduces sperm viability and motility. In other animals, sperm performance and embryo quality decline. We identify the duration of sperm storage, the design used for sampling individuals, and the sex of the individual storing sperm as potentially important moderators of the effects of sperm storage. These findings have key biomedical implications, including optimising the timing of ejaculation and fertilisation in fertility clinics or captive breeding programs. Overall, our results reveal the mechanisms that cause post-meiotic sperm senescence, the fitness consequences of sperm storage, and provide evolutionary insights into sex-specific adaptations that potentially mitigate the detrimental effects of sperm storage. ### Competing Interest Statement The authors have declared no competing interest. All data associated with our study, and the code used to analyse this data, is available at OSF with the DOI: 10.17605/OSF.IO/JXV7Z.

Mate availability and social information can influence mating behaviour in both males and females. Social information obtained from conspecifics can influence mate choice, particularly shown by studies on mate choice copying. However, the role of directly observing conspecific mating on mating behaviour has been less explored. As such, whether conspecifics are copulating or not could inform ‘observers’ about the availability of mates, or even stimulate observers to mate themselves. Using Drosophila melanogaster, we tested whether exposure to the visual cue of a mating pair would increase the mating propensity of an observer pair (i.e., voyeur). We followed a factorial design where a male-female pair (voyeur flies) were placed together with or without visual access to another pair of flies (who were either mating or not mating). We found no evidence that mating latency or duration of mating were affected by whether voyeurs had visual access to a mating or non-mating pair. These results could be due to biological factors (e.g., use of other non-visual cues by flies to acquire information related to sex), or methodological limitations of our study (e.g., flies unable to watch other pairs). Generally, our results suggest that fruit flies do not use visual cues from conspecifics mating to adjust their own mating latency or mating duration.

Male reproductive traits such as ejaculate size and quality, are expected to decline with advancing age due to senescence. It is however unclear whether this expectation is upheld across taxa. We perform a meta-analysis on 379 studies, to quantify the effects of advancing male age on ejaculate traits across 157 species of non-human animals. Contrary to predictions, we find no consistent pattern of age-dependent changes in ejaculate traits. This result partly reflects methodological limitations, such as studies sampling a low proportion of adult lifespan, or the inability of meta-analytical approaches to document non-linear ageing trajectories of ejaculate traits; which could potentially lead to an underestimation of senescence. Yet, we find taxon-specific differences in patterns of ejaculate senescence. For instance, older males produce less motile and slower sperm in ray-finned fishes, but larger ejaculates in insects, compared to younger males. Notably, lab rodents show senescence in most ejaculate traits measured. Our study challenges the notion of universal reproductive senescence, highlighting the need for controlled methodologies and a more nuanced understanding of reproductive senescence, cognisant of taxon-specific biology, experimental design, selection pressures, and life-history.

Fight outcomes often affect male fitness by determining their access to mates. Thus, "winner-loser" effects, where winners often win their next contest while losers tend to lose, can influence how males allocate resources toward pre- and postcopulatory traits. We experimentally manipulated the winning/losing experiences of pairs of size-matched male Gambusia holbrooki for 1 day, 1 week, or 3 weeks to test whether prior winning/losing experiences differentially affect the plasticity of male investment into either mating effort (precopulatory) or ejaculates (postcopulatory). When winner/loser pairs directly competed for a female, winners had better precopulatory outcomes than losers for three of the four traits we measured: mating attempts, successful attempts, and time spent with the female (but not aggression). However, winners and losers did not differ in either total sperm counts or sperm velocity. Interestingly, absolute male size, an important predictor of fighting success, mediated winner-loser effects on how long males then spent near a female. Compared with losers, smaller winners spent more time with the female than did larger winners, suggesting that how males respond to prior social experiences is size dependent. We discuss the general importance of controlling for inherent male condition when comparing male investment into condition-dependent traits.

Senescence, the deterioration of organismal function with advancing age, is a puzzling biological phenomenon. While actuarial senescence ( i.e. , age-dependent increases in mortality rates) is well described across some taxa, reproductive senescence ( i.e. age- dependent declines in reproduction) is less understood, especially in males, with mixed patterns reported across studies. To examine the evidence for male reproductive senescence, we investigated how advancing male age affects ejaculate traits across non-human animals via a meta-analysis yielding 1814 effect sizes from 379 studies. We found no evidence for a general pattern of reproductive senescence. Instead, we found high heterogeneity for how reproduction changes with male age across animals. Some of this heterogeneity (>10%) was associated with biological factors. For example, there were taxonomical differences for some ejaculate traits — sperm motility declined with male age in lab rodents and fish, whereas ejaculate size improved with male age in bulls, fish, and insects. Some methodological factors were also important in explaining this heterogeneity: studies sampling a larger proportion of a species’ lifespan were more likely to detect senescence in ejaculate traits, emphasising the need to examine the full life cycle of species to document senescence. Contrary to predictions, we reveal that the evidence for senescence in ejaculate traits is sporadic. Our findings will help generate novel hypotheses and identify more effective methodological approaches for studying male reproductive senescence.

Spatial cognitive abilities allow individuals to remember the location of food patches, predator hide-outs, or shelters. Animals typically incorporate learnt spatial information or use external environmental cues to navigate their surroundings. A spectacular example of how some fishes move is through aerial jumping. For instance, fish that are trapped within isolated pools, cut off from the main body of water during dry periods, may jump over obstacles and direct their jumps to return to safe locations. However, what information such re-orientation behaviour during jumping is based on remains enigmatic. Here we combine a lab and field experiment to test if guppies ( Poecilia reticulata ) incorporate learnt spatial information and external environmental cues (visual and auditory) to determine where to jump. In a spatial memory assay we found that guppies were more likely to jump towards deeper areas, hence incorporating past spatial information to jump to safety. In a matched vs. mismatched spatial cue experiment in the field, we found that animals only showed directed jumping when visual and auditory cues matched. We show that in unfamiliar entrapments guppies direct their jumps by combining visual and auditory cues, while in familiar entrapments they use a cognitive map. We hence conclude that jumping behaviour is a goal-directed behaviour, guided by different sources of information and involving important spatial cognitive skills.

Inbreeding depression, the reduced fitness of the offspring of related individuals, can affect males and females differently. Although a comprehensive theoretical framework describing the causes of sex-specific inbreeding depression is lacking, empirical evidence suggests that often one sex tends to be more vulnerable than the other. However, the generality, direction, and degree of sex-specific difference in inbreeding depression remains enigmatic as studies on this topic have reported conflicting results. Here, we conduct a meta-analysis to test for sex-specific differences in the magnitude of inbreeding depression. We synthetised 321 effect sizes of experimental studies across 47 species and found a small difference in inbreeding depression between the sexes: females suffered slightly higher inbreeding depression than males. Furthermore, a higher inbreeding coefficient was correlated with higher inbreeding depression. However, there was a large amount of heterogeneity that remained unexplained, even when considering different factors that could affect inbreeding between the sexes, such as sexual size dimorphism, heterogamety, the type of trait measured and whether animals were tested in a stressful environment. As such, we highlight the need to further explore inbreeding depression across different species to determine the occurrence and causes of sex differences to increase our understanding of the evolutionary consequences of sex-specific inbreeding depression.

Life-history strategies are diverse. While understanding this diversity is a fundamental aim of evolutionary biology and biodemography, life-history data for some traits—in particular, age-dependent reproductive investment—are biased towards females. While other authors have highlighted this sex skew, the general scale of this bias has not been quantified and its impact on our understanding of evolutionary ecology has not been discussed. This review summarizes why the sexes can evolve different life-history strategies. The scale of the sex skew is then discussed and its magnitude compared between taxonomic groups, laboratory and field studies, and through time. We discuss the consequences of this sex skew for evolutionary and ecological research. In particular, this sex bias means that we cannot test some core evolutionary theory. Additionally, this skew could obscure or drive trends in data and hinder our ability to develop effective conservation strategies. We finally highlight some ways through which this skew could be addressed to help us better understand broad patterns in life-history strategies.

Cognitive and sensory abilities are vital in affecting survival under predation risk, leading to selection on brain anatomy. However, how exactly predation and brain evolution are linked has not yet been resolved, as current empirical evidence is inconclusive. This may be due to predation pressure having different effects across life stages and/or due to confounding factors in ecological comparisons of predation pressure. Here, we used adult guppies ( Poecilia reticulata ) to experimentally test how direct predation during adulthood would impact the relative brain size and brain anatomy of surviving individuals to examine if predators selectively remove individuals with specific brain morphology. To this end, we compared fish surviving predation to control fish, which were exposed to visual and olfactory predator cues but could not be predated on. We found that predation impacted the relative size of female brains. However, this effect was dependent on body size, as larger female survivors showed relatively larger brains, while smaller survivors showed relatively smaller brains when compared to control females. We found no differences in male relative brain size between survivors and controls, nor for any specific relative brain region sizes for either sex. Our results corroborate the important, yet complex, role of predation as an important driver of variation in brain size.

Animals are usually expected to avoid mating with relatives (kin avoidance) as incestuous mating can lead to the expression of inbreeding depression. Yet, theoretical models predict that unbiased mating with regards to kinship should be common, and that under some conditions, the inclusive fitness benefits associated with inbreeding can even lead to a preference for mating with kin. This mismatch between empirical and theoretical expectations generates uncertainty as to the prevalence of inbreeding avoidance in animals. Here, we synthesized 677 effect sizes from 139 experimental studies of mate choice for kin versus non-kin in diploid animals, representing 40 years of research, using a meta-analytical approach. Our meta-analysis revealed little support for the widely held view that animals avoid mating with kin, despite clear evidence of publication bias. Instead, unbiased mating with regards to kinship appears widespread across animals and experimental conditions. The significance of a variety of moderators was explored using meta-regressions, revealing that the degree of relatedness and prior experience with kin explained some variation in the effect sizes. Yet, we found no difference in kin avoidance between males and females, choice and no-choice experiments, mated and virgin animals or between humans and animals. Our findings highlight the need to rethink the widely held view that inbreeding avoidance is a given in experimental studies.

The outcomes of fights often affect the fitness of males by determining their access to mates. ‘Winner-loser’ effects, where winners often win their next contest, but losers tend to lose, can therefore influence how males allocate resources towards traits under pre- and post-copulatory sexual selection. We experimentally manipulated the winning/losing experiences of size-matched male Gambusia holbrooki for either a day, a week or three weeks to test whether prior winning/losing experiences differentially affect the plasticity of male investment into either mating effort (pre-copulatory) or ejaculates (post-copulatory). Winners had better pre-copulatory outcomes than losers for three of the four traits we measured: number of mating attempts, number of successful attempts, and time spent with the female. Winners also produced faster sperm than losers, but there was no difference in total sperm counts. Interestingly, absolute male size, an important predictor of fighting success, mediated the effect of winning or losing on how long males then spent near a female. Smaller winners spent more time with the female than did larger winners, suggesting that how males respond to prior social experiences is size-dependent. We discuss the general importance of controlling for inherent male condition when comparing male investment into condition-dependent traits.

Predation risk during early ontogeny can impact developmental trajectories and permanently alter adult phenotypes. Such phenotypic plasticity often leads to adaptive changes in traits involved in anti-predator responses. While plastic changes in cognition may increase survival, it remains unclear whether early predation experience shapes cognitive investment and drives developmental plasticity in cognitive abilities. Here, we show that predation risk during early ontogeny induces developmental plasticity in two cognitive domains. We reared female guppies (Poecilia reticulata) with and without predator cues and tested their adult cognitive abilities. We found that females reared under simulated predation took longer to learn a simple association task, yet outperformed animals reared without predation threat in a reversal learning task testing cognitive flexibility. These results show that predation pressure during ontogeny shapes adult cognitive abilities, which we argue is likely to be adaptive. Our study highlights the important role of predator-mediated developmental plasticity on cognitive investment in natural populations and the general role of plasticity in cognitive performance.

In many species, males exhibit phenotypic plasticity in sexually selected traits when exposed to social cues about the intensity of sexual competition. To date, however, few studies have tested how this plasticity affects male reproductive success. We initially tested whether male mosquitofish, Gambusia holbrooki (Poeciliidae), change their investment in traits under pre- and postcopulatory sexual selection depending on the social environment. For a full spermatogenesis cycle, focal males were exposed to visual and chemical cues of rivals that were either present (competitive treatment) or absent (control). Males from the competitive treatment had significantly slower-swimming sperm but did not differ in sperm count from control males. When two males competed for a female, competitive treatment males also made significantly fewer copulation attempts and courtship displays than control males. Further, paternity analysis of 708 offspring from 148 potential sires, testing whether these changes in reproductive traits affected male reproductive success, showed that males previously exposed to cues about the presence of rivals sired significantly fewer offspring when competing with a control male. We discuss several possible explanations for these unusual findings.

1.Phenotypic plasticity requires animals to acquire reliable environmental information. When multiple sources of information agree, cues should be perceived as reliable and induce a relatively strong response. Conversely, where stimuli conflict, animals must weigh the accuracy of the sources of information and responses should be reduced. 2.Availability of reliable information is often considered a limitation on plasticity, yet how animals integrate seemingly contradictory or incomplete information remains enigmatic, as empirical tests are scarce. 3.We tested how incomplete information determines phenotypic plasticity by simulating predation risk during early ontogeny of guppies (Poecilia reticulata). We exposed guppy fry to a combination of visual and/or olfactory cues of the predatory pike cichlid (Crenicichla alta), and monitored growth of the body and brain. After five weeks of exposure, guppies were returned to common no-risk conditions and their activity rates were monitored for four weeks post-treatment. 4.Visual predator exposure more strongly affected development; reducing body size of adult males and increasing brain size in females. However, there was little evidence for the hypothesised additive effect, with the combined treatment not inducing a larger effect than when only receiving olfactory or visual treatments. 5.While there was consistent individual variation in activity rates, this was unaffected by developmental risk and uncorrelated with the growth parameters. 6.Our results demonstrate the differential reliability of cues during development. Visual exposure to a predator was a highly reliable environmental cue, while environmental certainty was unaffected by combined stimuli.

Predation is a near ubiquitous factor of nature and a powerful selective force on prey. Moreover, it has recently emerged as an important driver in the evolution of brain anatomy, though population comparisons show ambiguous results with considerable unexplained variation. Here, we test the reproducibility of reduced predation on evolutionary trajectories of brain evolution. We make use of an introduction experiment, whereby guppies (Poecilia reticulata) from a single high predation stream were introduced to four low predation streams. After 8–9 years of natural selection in the wild and two generations of common garden conditions in the laboratory, we quantified brain anatomy. Relative brain region sizes did not differ between populations. However, we found a general increase and striking variation in relative brain size of introduced populations, which varied from no change to a 12.5% increase in relative brain weight, relative to the ancestral high predation population. We interpret this as evidence for non-parallel evolution, which implies a weak or inconsistent association of relative brain size with fitness in low predation sites. The evolution of brain anatomy appears sensitive to unknown environmental factors, or contingent on either chance events or historical legacies of environmental change.