Benefits and costs of earwig ( Forficula auricularia ) family life

The evolution of parental care and family group formation critically depends on offspring survival benefits and parental fecundity costs of care under given ecological conditions. Investigations of the functional significance of care in insect species that exhibit facultative parental care have been relatively rare but may be of particular interest for better understanding of benefit and cost schedules at an early evolutionary stage. In this study, aspects of benefits and costs of care were addressed in the sub-social European earwig (Forficula auricularia; Dermaptera: Forficulidae) by manipulating the presence of tending mothers and brood size in a fully crossed experimental design. Larvae growing in broods tended by their mother or of reduced size showed a higher survival probability than larvae growing in untended or large broods, as predicted if maternal care is beneficial and shaped by a trade-off between number and quality of offspring. Analysis of patterns of food consumption and developmental time further suggested that the benefit of maternal attendance is mediated by the maternal provisioning of food, while the quality-quantity trade-off seemed to be driven by sibling rivalry. Further, tending mothers delayed the production of a second clutch, indicating a potential cost of care in terms of lifetime fecundity. This study experimentally shows benefits and potential costs of maternal care and family group formation in the European earwig. More detailed behavioural experiments will be required to fully understand how behavioural interactions among family members mediate these reproductive outcome

Differential effects of offspring condition-dependent signals on maternal care regulation in the European earwig

Parent-offspring conflict theory predicts the evolution of offspring solicitation signals that can influence the amount and/or the duration of parental investment. Short-term effects of offspring solicitation signals on parental food provisioning have been widely demonstrated, but persistent effects of offspring signals on the maintenance of parental care have been rarely studied. Also, the relation between the amount of care provided to the brood and how it is distributed among individual offspring within a brood is not well enough understood. Here, we investigated in the European earwig (Forficula auricularia) the effects of offspring condition-dependent chemical signals on the maintenance of maternal care among broods and the distribution of maternal food within broods. Mothers were isolated from their brood for 3days and continuously exposed to chemical signals extracted from broods of experimentally manipulated nutritional state. After re-introducing mothers to their brood, a range of maternal behaviours were quantified. We found that earwig mothers groomed their offspring significantly more after exposure to chemical extract from high-food brood in comparison with mothers exposed to extract from low-food brood, which in turn displayed significantly more aggressive behaviour. Furthermore, we manipulated offspring individual nutritional condition within the brood to evaluate the effect of offspring state on the within-brood food distribution. Within broods, poorly fed individuals received significantly more food than well-fed individuals, probably due to scramble competition. These results show that earwig nymphs express multi-component condition-dependent signals and behaviours differentially affecting maternal care provisioned to the brood and the distribution of care within brood

Kin-selected siblicide and cannibalism in the European earwig

Aggression levels among individuals can severely increase under high density or shortage of crucial resources, sometimes resulting in individuals killing conspecifics. This is not uncommon in family groups of diverse taxa, where the dependent offspring compete for the limited resources provided by their parents. Killing a nest mate can relax the level of competition, and cannibalism provides a direct nutritional benefit. However, nest-mate killing bears the risk of reducing indirect fitness if the victim is related (i.e., siblicide), imposing selection on kin-recognition abilities. Based on this hypothesis, we predicted that first instar juveniles (nymphs) of the European earwig (Forficula auricularia) kill and cannibalize unrelated nest mates earlier and more often than related nest mates and that cannibalism has a direct nutritional benefit in terms of survival. We tested these predictions experimentally by establishing related and unrelated pairs of nymphs and recorded survival, aggregation behavior, and cannibalistic outcomes in the absence of alternative food sources. In order to obtain expected survival probabilities of victims and survivors in the absence of any interaction, we simulated virtual nymph pairs based on survival data of singly held control individuals. As predicted, victims lived for less time and survivor for longer than expected from the simulated survival distributions, demonstrating nest-mate killing and cannibalism. Furthermore, unrelated individuals were killed significantly earlier and were more often cannibalized than related individuals. The survival patterns of victims and survivors were quantitatively consistent with the expectations of Hamilton's rule. Our study shows that earwig nymphs recognize kin and adjust their nest-mate killing and cannibalistic behavior as predicted under the hypothesis of kin-selected siblicide and cannibalis

Social runaway : fisherian elaboration (or reduction) of socially selected traits via indirect genetic effects

NWB was funded by fellowships from the UK Natural Environment Research Council [NE/G014906/1 and NE/L011255/1].Our understanding of the evolutionary stability of socially‐selected traits is dominated by sexual selection models originating with R. A. Fisher, in which genetic covariance arising through assortative mating can trigger exponential, runaway trait evolution. To examine whether non‐reproductive, socially‐selected traits experience similar dynamics—social runaway—when assortative mating does not automatically generate a covariance, we modelled the evolution of socially‐selected badge and donation phenotypes incorporating indirect genetic effects (IGEs) arising from the social environment. We establish a social runaway criterion based on the interaction coefficient, ψ, which describes social effects on badge and donation traits. Our models make several predictions. (1) IGEs can drive the original evolution of altruistic interactions that depend on receiver badges. (2) Donation traits are more likely to be susceptible to IGEs than badge traits. (3) Runaway dynamics in non‐sexual, social contexts can occur in the absence of a genetic covariance. (4) Traits elaborated by social runaway are more likely to involve reciprocal, but non‐symmetrical, social plasticity. Models incorporating plasticity to the social environment via IGEs illustrate conditions favouring social runaway, describe a mechanism underlying the origins of costly traits such as altruism, and support a fundamental role for phenotypic plasticity in rapid social evolution.PostprintPeer reviewe

Olfactory attractiveness of flowering plants to the parasitoid Microplitis mediator: potential implications for biological control

In agricultural landscapes, the lack of floral nectar can be a major difficulty for nectar feeding parasitoids. This problem can be reduced by the addition of suitable wildflowers. To date, flowers have mainly been studied in terms of effects on parasitoid fitness, not taking into account the essential role of flower attractiveness for foraging parasitoids. This study experimentally tested the olfactory attractiveness of five wildflowers (bishop's weed, cornflower, buckwheat, candytuft, and oregano) to the parasitoid Microplitis mediator (Haliday) (Hymenoptera: Braconidae). We conducted choice experiments in a Y-tube olfactometer to test the attractiveness of flowers against air, and relative attractiveness in paired choice tests. Our results showed that all the flowers were highly attractive and that in paired choice tests cornflower and candytuft were equally attractive and more attractive than buckwheat. These results indicate that M. mediator has evolved innate preferences that could be effectively exploited in biological contro

Maternal care provides antifungal protection to eggs in the European earwig

Many insects raise their offspring on organic substrates or in the soil where microorganisms are abundant. Microbes may pose a serious threat to offspring development and survival by either decomposing food resources or directly infecting the offspring. Selection to cope with these effects may favor social defenses, for example, through forms of parental care that can limit or eliminate these threats to offspring fitness. In this study, we experimentally tested if maternal egg attendance in the European earwig Forficula auricularia has a function as a social defense against mold infection of eggs by manipulating exposure of eggs to mold spores and the presence of the mother in a fully factorial design. Furthermore, we investigated the potential roles of egg grooming behavior and maternal transfer of chemicals as underlying mechanisms. As predicted, the beneficial effect of egg attendance on hatching success was significantly enhanced when eggs were exposed to the mold. Females significantly increased their egg grooming duration in response to mold exposure of her eggs, and the quantity of chemicals (identified as hydrocarbons) was maintained among attended eggs but decreased substantially among unattended eggs. Maternal transfer of chemicals was confirmed in extractions of glass beads that were mingled into attended or unattended clutches. This study shows that maternal egg attendance in the European earwig has a social defense function protecting offspring against mold infection. The maternal egg grooming behavior seems to be key for this effect, probably through both the mechanical removal of spores and the continued application of chemical substances on the egg surfac

Brood size, sibling competition, and the cost of begging in great tits (Parus major)

Evolutionary theory of parent-offspring conflict explains begging displays of nestling birds as selfish attempts to influence parental food allocation. Models predict that this conflict may be resolved by honest signaling of offspring need to parents, or by competition among nestmates, leading to escalated begging scrambles. Although the former type of models has been qualitatively supported by experimental studies, the potential for a begging component driven by scramble competition cannot be excluded by the evidence. In a brood-size manipulation experiment with great tits, Parus major, we explored the scramble component in the begging activity of great tit nestlings by investigating the mechanisms of sibling competition in relation to brood size. While under full parental compensation, the feeding rate per nestling will remain constant over all brood sizes for both types of models; the scramble begging models alone predict an increase in begging intensity with brood size, if begging costs do not arise exclusively through predation. Great tit parents adjusted feeding rates to brood size and fed nestlings at similar rates and with similar prey sizes in all three brood-size categories. Despite full parental compensation, the begging and food solicitation activities increased with experimental brood size, whereas nestling body condition deteriorated. These findings support a scramble component in begging and suggest that the competition-induced costs of food solicitation behavior play an important role in the evolution of parent-offspring communicatio

Offspring social network structure predicts fitness in families.

addresses: Centre for Ecology and Conservation, Biosciences, College of Life and Environmental Sciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK. [email protected]: PMCID: PMC3497231types: Journal Article; Research Support, Non-U.S. Gov'tSocial structures such as families emerge as outcomes of behavioural interactions among individuals, and can evolve over time if families with particular types of social structures tend to leave more individuals in subsequent generations. The social behaviour of interacting individuals is typically analysed as a series of multiple dyadic (pair-wise) interactions, rather than a network of interactions among multiple individuals. However, in species where parents feed dependant young, interactions within families nearly always involve more than two individuals simultaneously. Such social networks of interactions at least partly reflect conflicts of interest over the provision of costly parental investment. Consequently, variation in family network structure reflects variation in how conflicts of interest are resolved among family members. Despite its importance in understanding the evolution of emergent properties of social organization such as family life and cooperation, nothing is currently known about how selection acts on the structure of social networks. Here, we show that the social network structure of broods of begging nestling great tits Parus major predicts fitness in families. Although selection at the level of the individual favours large nestlings, selection at the level of the kin-group primarily favours families that resolve conflicts most effectively

One clutch or two clutches? Fitness correlates of coexisting alternative female life-histories in the European earwig

Whether to reproduce once or multiple times (semelparity vs. iteroparity) is a major life-history decision that organisms have to take. Mode of parity is usually considered a species characteristic. However, recent models suggested that population properties or condition-dependent fitness payoffs could help to maintain both life-history tactics within populations. In arthropods, semelparity was also hypothesised to be a critical pre-adaptation for the evolution of maternal care, semelparous females being predicted to provide more care due to the absence of costs on future reproduction. The aim of this study was to characterize potential fitness payoffs and levels of maternal care in semel- and itero-parous females of the European earwig Forficula auricularia. Based on 15 traits measured in 494 females and their nymphs, our results revealed that iteroparous females laid their first clutch earlier, had more eggs in their first clutch, gained more weight during the 2weeks following hatching of the first clutch, but produced eggs that developed more slowly than semelparous females. Among iteroparous females, the sizes of first and second clutches were significantly and positively correlated, indicating no investment trade-off between reproductive events. Iteroparous females also provided more food than semelparous ones, a result contrasting with predictions that iteroparity is incompatible with the evolution of maternal care. Finally, a controlled breeding experiment reported full mating compatibility among offspring from females of the two modes of parity, confirming that both types of females belong to one single species. Overall, these results indicate that alternative modes of parity represent coexisting life-history tactics that are likely to be condition-dependent and associated with offspring development and specific levels of maternal care in earwig