Intraspecific Variation in Parental Care May Reflect Variation in Parental Quality.

The existence of life-history trade-offs is a fundamental assumption of evolutionary biology and behavioural ecology, yet empirical studies have found mixed evidence for this. Such trade-offs are expected when individuals vary in how they allocate their limited resource budgets between different life-history functions (variation in resource allocation), but they may be masked when individuals vary in how many resources they have acquired that they can later allocate to life-history functions (variation in resource acquisition). We currently lack studies on the extent to which individual differences in behaviour reflect variation between individuals in resource acquisition and resource allocation.

Revisiting the ecology and evolution of burying beetle behavior (Staphylinidae: Silphinae).

Investigating fundamental processes in biology requires the ability to ground broad questions in species-specific natural history. This is particularly true in the study of behavior because an organism's experience of the environment will influence the expression of behavior and the opportunity for selection. Here, we provide a review of the natural history and behavior of burying beetles of the genus to provide the groundwork for comparative work that showcases their remarkable behavioral and ecological diversity.

Flexible females: nutritional state influences biparental cooperation in a burying beetle.

In species that provide biparental care, there is a sexual conflict between parents over how much each should contribute toward caring for their joint offspring. Theoretical models for the resolution of this conflict through behavioral negotiation between parents assume that parents cannot assess their partner's state directly but do so indirectly by monitoring their partner's contribution. Here, we test whether parents can assess their partner's state directly by investigating the effect of nutritional state on cooperation between parents in the burying beetle .

Access to resources buffers against effects of current reproduction on future ability to provide care in a burying beetle.

Studies investigating the trade-off between current and future reproduction often find that increased allocation to current reproduction is associated with a reduction in the number or quality of future offspring. In species that provide parental care, this effect on future offspring may be mediated through a reduced future ability to provide care. Here, we test this idea in the burying beetle , a species in which parents shift the cost of reproduction toward future offspring and provide elaborate parental care.

Having a better home range does not reduce the cost of reproduction in Soay sheep.

A cost of reproduction may not be observable in the presence of environmental or individual heterogeneity because they affect the resources available to individuals. Individual space use is critical in determining both the resources available to individuals and the exposure to factors that mediate the value of these resources (e.g.

Carry on caring: infected females maintain their parental care despite high mortality.

Parental care is a key component of an organism's reproductive strategy that is thought to trade-off with allocation toward immunity. Yet, it is unclear how caring parents respond to pathogens: do infected parents reduce care as a sickness behavior or simply from being ill or do they prioritize their offspring by maintaining high levels of care? To address this issue, we investigated the consequences of infection by the pathogen on mortality, time spent providing care, reproductive output, and expression of immune genes of female parents in the burying beetle . We compared untreated control females with infected females that were inoculated with live bacteria, immune-challenged females that were inoculated with heat-killed bacteria, and injured females that were injected with buffer.

Resource allocation is determined by both parents and offspring in a burying beetle.

Parents and offspring have different optima for the level of parental resource allocation and the timing of nutritional independence. Theoretical models assume that either parents or offspring control the allocation of resources within a brood; however, control may also be mutual. Here, we investigate whether the resolution of parent-offspring conflict is biased towards cues from either the parents' or the offspring's behaviour, or whether the conflict is under mutual control.

Effects of inbreeding on behavioural plasticity of parent-offspring interactions in a burying beetle.

Inbreeding depression is defined as a fitness decline in progeny resulting from mating between related individuals, the severity of which may vary across environmental conditions. Such inbreeding-by-environment interactions might reflect that inbred individuals have a lower capacity for adjusting their phenotype to match different environmental conditions better, as shown in prior studies on developmental plasticity. Behavioural plasticity is more flexible than developmental plasticity because it is reversible and relatively quick, but little is known about its sensitivity to inbreeding.

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.

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.

The resolution of conflict in families.

The emergence of family groups is associated with conflict over the allocation of food or other limited resources. Understanding the mechanisms mediating the resolution of such conflict is a major aim in behavioral ecology. Most empirical work on familial conflict has focused on birds.

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.

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.

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.

Biparental care is more than the sum of its parts: experimental evidence for synergistic effects on offspring fitness.

Despite an extensive body of theoretical and empirical literature on biparental cooperation, it is still unclear whether offspring fare equally, better or worse when receiving care by two parents versus a single parent. Some models predict that parents should withhold the amount of care they provide due to sexual conflict, thereby shifting as much of the workload as possible to their partner. This conflict should lead to offspring faring worse with two parents.

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.

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.

Differential effects of offspring and maternal inbreeding on egg laying and offspring performance in the burying beetle Nicrophorus vespilloides.

We investigate the effect of offspring and maternal inbreeding on maternal and offspring traits associated with early offspring fitness in the burying beetle Nicrophorus vespilloides. We conducted two experiments. In the first experiment, we manipulated maternal inbreeding only (keeping offspring outbred) by generating mothers that were outbred, moderately inbred or highly inbred.

Flexible parents: joint effects of handicapping and brood size manipulation on female parental care in Nicrophorus vespilloides.

Parental care is highly variable, reflecting that parents make flexible decisions in response to variation in the cost of care to themselves and the benefit to their offspring. Much of the evidence that parents respond to such variation derives from handicapping and brood size manipulations, the separate effects of which are well understood. However, little is known about their joint effects.

Females manipulate behavior of caring males via prenatal maternal effects.

In species with biparental care, there is sexual conflict as each parent is under selection to minimize its personal effort by shifting as much as possible of the workload over to the other parent. Most theoretical and empirical work on the resolution of this conflict has focused on strategies used by both parents, such as negotiation. However, because females produce the eggs, this might afford females with an ability to manipulate male behavior via maternal effects that alter offspring phenotypes.

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.

Accounting for female space sharing in St. Kilda Soay sheep (Ovis aries) results in little change in heritability estimates.

When estimating heritability in free-living populations, it is common practice to account for common environment effects, because of their potential to generate phenotypic covariance among relatives thereby biasing heritability estimates. In quantitative genetic studies of natural populations, however, philopatry, which results in relatives being clustered in space, is rarely accounted for. The two studies that have been carried out so far suggest absolute declines in heritability estimates of up to 43% when accounting for space sharing by relatives.

Maternal effects alter the severity of inbreeding depression in the offspring.

A maternal effect is a causal influence of the maternal phenotype on the offspring phenotype over and above any direct effects of genes. There is abundant evidence that maternal effects can have a major impact on offspring fitness. Yet, no previous study has investigated the potential role of maternal effects in influencing the severity of inbreeding depression in the offspring.

Effects of Prior Contest Experience and Contest Outcome on Female Reproductive Decisions and Offspring Fitness.

Winning or losing a prior contest can influence the outcome of future contests, but it might also alter subsequent reproductive decisions. For example, losers may increase their investment in the current breeding attempt if losing a contest indicates limited prospects for future breeding. Using the burying beetle Nicrophorus vespilloides, we tested whether females adjust their prehatching and posthatching reproductive effort after winning or losing a contest with a same-sex conspecific.