Male crickets in poor condition engage in less same-sex sexual behavior.

Same-sex sexual behavior (SSB) is widespread among animals and is often treated as an evolutionary anomaly or mistake. An alternative view is that SSB occurs because individuals have broader or more permissive "mating filters." A broader filter means directing courtship toward anything that resembles a potential mate, while a narrower filter means having stricter criteria about when to court. Broader filters help ensure animals do not miss out on mating opportunities. However, it may be advantageous for individuals to narrow their mating filters under some conditions, such as when the costs of courtship increase because courting individuals are in poor condition. We examined whether mating filters, and the expression of SSB, is condition dependent in the Pacific field cricket (Teleogryllus oceanicus). When males were in poor condition (because they were reared on a poor diet), they courted other males less. However, condition did not influence courtship toward females. This finding provides evidence that animals can decrease their expression of SSB by flexibly narrowing their mating filters when in poor condition. Our results demonstrate the benefits of a mating filters approach to studying SSB-rather than being a mistake or an anomaly in need of special explanation, SSB may be better viewed as a consequence of flexible, condition-dependent changes in mating behavior. Such an approach can be applied to other contexts where behavior is directed toward unintended targets. We emphasize that our results, like other studies of SSB in nonhuman animals, tell us nothing about sexual identities in humans.

Meta-analytical evidence that males prefer virgin females.

Males are often predicted to prefer virgin over non-virgin females because of the reduced risk of sperm competition. Does this prediction hold across studies? Our systematic meta-analysis of 138 studies, mainly conducted in invertebrates, confirms that males generally prefer virgin females. However, males preferred virgin females even in species with last male sperm precedence, suggesting that sperm competition alone does not drive male preferences. Furthermore, our results suggest that males may reject mated females even when no alternative exists. Preference for virgins is unlikely to influence female reproductive success since virginity cannot be selected for, but strong preference for virgin females could swamp or reinforce selection on other traits. Our results add to growing evidence that males are not indiscriminate in mating. However, given the unexplained heterogeneity in effect sizes, we urge caution in assuming that males will prefer virgins and recommend considering the natural context of mating decisions.

Rethinking same-sex sexual behaviour: male field crickets have broad mating filters.

Same-sex sexual behaviour (SSB) occurs in many animals and is often treated as an anomaly requiring special explanation. One common explanation for SSB is mistaken identity. However, animals make similar 'mistakes' in other contexts-such as attempting to mate with immature individuals or inanimate objects. Framing such behaviours as 'mistakes' risks misinterpreting how animals make flexible mating decisions. Here we make a case for an alternative approach to thinking about SSB by instead considering an individual's mating filter. A broad filter means directing courtship toward anything that resembles a potential mate, whilst a narrow filter means only courting with receptive targets. We illustrate this approach by examining the mating filters of male Pacific field crickets (Teleogryllus oceanicus). We find that males engage in SSB but also misdirect courtship toward juveniles (but not plastic crickets). This finding suggests that SSB is not an anomaly and is better considered alongside other misdirected behaviours. We argue that by viewing misdirected behaviours through the lens of mating filters rather than as 'mistakes' we can build a more nuanced understanding of reproductive behaviour and begin to determine when having a broader mating filter can be advantageous.

Sex-specific associations between life-history traits and a novel reproductive polymorphism in the Pacific field cricket.

Associations between heritable polymorphisms and life-history traits, such as development time or reproductive investment, may play an underappreciated role in maintaining polymorphic systems. This is because selection acting on a particular morph could be bolstered or disrupted by correlated changes in life history or vice versa. In a Hawaiian population of the Pacific field cricket (Teleogryllus oceanicus), a novel mutation (flatwing) on the X-chromosome is responsible for a heritable polymorphism in male wing structure. We used laboratory cricket colonies fixed for male wing morph to investigate whether males and females bearing the flatwing or normal-wing (wild-type) allele differed in their life-history traits. We found that flatwing males developed faster and had heavier testes than normal-wings, whereas flatwing homozygous females developed slower and had lighter reproductive tissues than normal-wing homozygous females. Our results advance our understanding of the evolution of polymorphisms by demonstrating that the genetic change responsible for a reproductive polymorphism can also have consequences for fundamental life-history traits in both males and females.

Females adjust maternal hormone concentration in eggs according to male condition in a burying beetle.

In birds and other vertebrates, there is good evidence that females adjust the allocation of hormones in their eggs in response to prenatal environmental conditions, such as food availability or male phenotype, with profound consequences for life history traits of offspring. In insects, there is also evidence that females deposit juvenile hormones (JH) and ecdysteroids (ESH) in their eggs, hormones that play a key role in regulating offspring growth and metamorphosis. However, it is unclear whether females adjust their hormonal deposition in eggs in response to prenatal environmental conditions. Here we address this gap by conducting an experiment on the burying beetle Nicrophorus vespilloides, in which we manipulated the presence of the male parent and the size of the carcass used for breeding at the time of laying. We also tested for effects of the condition (i.e., body mass) of the parents. We then recorded subsequent effects on JH and ESH concentrations in the eggs. We found no evidence for an effect of these prenatal environmental conditions (male presence and carcass size) on hormonal concentration in the eggs. However, we found that females reduced their deposition of JH when mated with heavier males. This finding is consistent with negative differential allocation of maternal hormones in response to variation in the body mass of the male parent. We encourage further work to investigate the role of maternally derived hormones in insect eggs.

Increased allocation to reproduction reduces future competitive ability in a burying beetle.

The existence of a trade-off between current and future reproduction is a fundamental prediction of life history theory. Support for this prediction comes from brood size manipulations, showing that caring for enlarged broods often reduces the parent's future survival or fecundity. However, in many species, individuals must invest in competing for the resources required for future reproduction. Thus, a neglected aspect of this trade-off is that increased allocation to current reproduction may reduce an individual's future competitive ability. We tested this prediction in the burying beetle, Nicrophorus vespilloides, a species where parents care for their offspring and where there is fierce competition for resources used for breeding. We manipulated reproductive effort by providing females with either a small brood of 10 larvae or a large brood of 40 larvae and compared the ability of these females, and virgin females that had no prior access to a carcass, to compete for a second carcass against a virgin competitor. We found that increased allocation to current reproduction reduced future competitive ability, as females that had cared for a small brood were more successful when competing for a second carcass against a virgin competitor than females that had cared for a large brood. In addition, the costs of reproduction were offset by the benefits of feeding from the carcass during an initial breeding attempt, as females that had cared for a small brood were better competitors than virgin females that had no prior access to a carcass, whilst females that had cared for a large brood were similar in competitive ability to virgin females. Our results add to our understanding of the trade-off between current and future reproduction by showing that this trade-off can manifest through differences in future competitive ability and that direct benefits of reproduction can offset some of these costs.

Effects of variation in resource acquisition during different stages of the life cycle on life-history traits and trade-offs in a burying beetle.

Individual variation in resource acquisition should have consequences for life-history traits and trade-offs between them because such variation determines how many resources can be allocated to different life-history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life-history traits and trade-offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade-off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life-history traits but limited effects on trade-offs. This suggests that in species that acquire a fixed-sized resource at the onset of breeding, the size of this resource has larger effects on life-history trade-offs than resources acquired at earlier stages.

Interplay between age-based competitive asymmetries within the brood and direct competition between inbred and outbred offspring in a burying beetle.

Theory suggests that intraspecific competition associated with direct competition between inbred and outbred individuals should be an important determinant of the severity of inbreeding depression. The reason is that, if outbred individuals are stronger competitors than inbred ones, direct competition should have a disproportionate effect on the fitness of inbred individuals. However, an individual's competitive ability is not only determined by its inbreeding status but also by competitive asymmetries that are independent of an individual's inbreeding status. When this is the case, such competitive asymmetries may shape the outcome of direct competition between inbred and outbred individuals. Here, we investigate the interface between age-based competitive asymmetries within broods and direct competition between inbred and outbred offspring in the burying beetle Nicrophorus vespilloides. We found that inbred offspring had lower survival than outbred ones confirming that there was inbreeding depression. Furthermore, seniors (older larvae) grew to a larger size and had higher survival than juniors (younger larvae), confirming that there were age-based competitive asymmetries. Nevertheless, there was no evidence that direct competition between inbred and outbred larvae exacerbated inbreeding depression, no evidence that inbreeding depression was more severe in juniors and no evidence that inbred juniors suffered disproportionately due to competition from outbred seniors. Our results suggest that direct competition between inbred and outbred individuals does not necessarily exacerbate inbreeding depression and that inbred individuals are not always more sensitive to poor and stressful conditions than outbred ones.

Effects of Offspring and Parental Inbreeding on Parent-Offspring Communication.

There is mounting evidence that inbreeding can have complex effects on social interactions among inbred and outbred individuals. Here, we investigate effects of offspring and maternal inbreeding on parent-offspring communication in the burying beetle Nicrophorus vespilloides. We find effects of the interaction between offspring and maternal inbreeding on maternal behavior. Outbred females provided more direct care toward inbred larvae, while inbred females provided similar levels of direct care toward inbred and outbred larvae. Furthermore, we find direct and indirect effects of offspring inbreeding on offspring begging and maternal behavior, respectively. Inbred larvae spent less time begging than outbred larvae, and (outbred) females provided more direct care and less indirect care toward inbred larvae. Finally, we find effects of the interaction between offspring and maternal inbreeding on larval body mass. Inbred and outbred offspring grew to a similar size when the female was outbred, while inbred offspring were of a smaller size when the female was inbred. Our results suggest that outbred females provided more care toward inbred offspring to compensate for their poor genetic quality. Our study advances our understanding of inbreeding by showing that inbreeding can have direct effects on the behavior of inbred individuals and indirect effects on the behavior of outbred individuals and that indirect effects on outbred individuals may in turn influence the fitness of inbred individuals.

Inbred burying beetles suffer fitness costs from making poor decisions.

There is growing interest in how environmental conditions, such as resource availability, can modify the severity of inbreeding depression. However, little is known about whether inbreeding depression is also associated with differences in individual decision-making. For example, decisions about how many offspring to produce are often based upon the prevailing environmental conditions, such as resource availability, and getting these decisions wrong may have important fitness consequences for both parents and offspring. We tested for effects of inbreeding on individual decision-making in the burying beetle Nicrophorus vespilloides, which uses the size of a carrion resource to make decisions about number of offspring. Both inbred and outbred females adjusted their initial decisions about number of eggs to lay based on carcass size. However, when we forced individuals to update this initial decision by providing them with a different-sized carcass partway through reproduction, inbred females failed to update their decision about how many larvae to cull. Consequently, inbred females reared too many larvae, resulting in negative fitness consequences in the form of smaller offspring and reduced female post-reproductive condition. Our study provides novel insights into the effects of inbreeding by showing that poor decision-making by inbred individuals can negatively affect fitness.

Intraspecific Competition and Inbreeding Depression: Increased Competitive Effort by Inbred Males Is Costly to Outbred Opponents.

A recent theoretical model suggests that intraspecific competition is an important determinant of the severity of inbreeding depression. The reason for this is that intraspecific competition is density dependent, leading to a stronger negative effect on inbred individuals if they are weaker competitors than outbred ones. In support of this prediction, previous empirical work shows that inbred individuals are weaker competitors than outbred ones and that intraspecific competition often exacerbates inbreeding depression. Here, we report an experiment on the burying beetle Nicrophorus vespilloides, in which we recorded the outcome of competition over a small vertebrate carcass between an inbred or outbred male resident caring for a brood and a size-matched inbred or outbred male intruder. We found that inbred males were more successful as intruders in taking over a carcass from a male resident and were injured more frequently as either residents or intruders. Furthermore, inbred males gained less mass during the breeding attempt and had a shorter adult life span than outbred males. Finally, successful resident males reared a substantially smaller brood comprised of lighter larvae when the intruder was inbred than when it was outbred. Our results shows that inbred males increased their competitive effort, thus contradicting previous work suggesting that inbred males are weaker competitors. Furthermore, our results shows that inbred intruders impose a greater cost to resident males, suggesting that outbred individuals can suffer fitness costs as a result of competition with inbred ones.

No evidence of sibling cooperation in the absence of parental care in Nicrophorus vespilloides.

Interactions among siblings fall on a continuum with competition and cooperation at opposite ends of the spectrum. Prior work on the burying beetle Nicrophorus vespilloides suggests that parental care shifts the balance between competition and cooperation by masking a density-dependent shift from cooperation to competition. However, these results should be interpreted with caution because they were based on correlational evidence for an association between larval density at dispersal and mean larval mass at dispersal. Here, we test for a causal effect of the initial density of larvae in a brood on the larvae's subsequent performance in the absence of care. We find no effect of the initial larval density on mean larval mass. Thus, our results provide no evidence for sibling cooperation in the absence of care in this species. However, using larval density at dispersal as a predictor of mean larval mass at dispersal, there was a significant correlation between larval density and mean larval mass. Our study highlights the importance of using experimental designs that exclude confounding effects due to shared environmental conditions that otherwise could be misinterpreted as evidence for sibling cooperation.

Waist-to-height ratio is more predictive of years of life lost than body mass index.

OBJECTIVE: Our aim was to compare the effect of central obesity (measured by waist-to-height ratio, WHtR) and total obesity (measured by body mass index, BMI) on life expectancy expressed as years of life lost (YLL), using data on British adults. METHODS: A Cox proportional hazards model was applied to data from the prospective Health and Lifestyle Survey (HALS) and the cross sectional Health Survey for England (HSE). The number of years of life lost (YLL) at three ages (30, 50, 70 years) was found by comparing the life expectancies of obese lives with those of lives at optimum levels of BMI and WHtR. RESULTS: Mortality risk associated with BMI in the British HALS survey was similar to that found in US studies. However, WHtR was a better predictor of mortality risk. For the first time, YLL have been quantified for different values of WHtR. This has been done for both sexes separately and for three representative ages. CONCLUSION: This study supports the simple message "Keep your waist circumference to less than half your height". The use of WHtR in public health screening, with appropriate action, could help add years to life.