Reproductive efficiency of the bethylid wasp Cephalonomia tarsalis: the influences of spatial structure and host density

The parasitoid wasp Cephalonomia tarsalis (Ashmead) (Hymenoptera: Bethylidae) is commonly present in stored product facilities. While beneficial, it does not provide a high 24 degree of biological pest control against its host, the saw-toothed beetle Oryzaephilus surinamensis (L.) (Coleoptera: Silvanidae). A candidate explanation for poor host population suppression is that adult females interfere with each other’s foraging and reproductive behavior. We used simple laboratory microcosms to evaluate such mutual interference in terms of its overall effects on offspring production. We varied the density of the hosts and also the spatial structure of the environment, via the extent of population sub-division and the provision of different substrates. Production of C. tarsalis offspring was positively influenced by host density and by the isolation of females. With incomplete sub-division within microcosms offspring production was, in contrast, low and even zero. The provision of corrugated paper as a substrate enhanced offspring production and partially mitigated the effects of mutual interference. We recommend simple improvements to mass rearing practice and identify promising areas for further behavioral and chemical studies towards a better understanding of the mechanisms of mutual interference

Detecting non-binomial sex allocation when developmental mortality operates

Optimal sex allocation theory is one of the most intricately developed areas of evolutionary ecology. Under a range of conditions, particularly under population sub-division, selection favours sex being allocated to offspring non-randomly, generating non-binomial variances of offspring group sex ratios. Detecting non-binomial sex allocation is complicated by stochastic developmental mortality, as offspring sex can often only be identified on maturity with the sex of non-maturing offspring remaining unknown. We show that current approaches for detecting non-binomiality have limited ability to detect non-binomial sex allocation when developmental mortality has occurred. We present a new procedure using an explicit model of sex allocation and mortality and develop a Bayesian model selection approach (available as an R package). We use the double and multiplicative binomial distributions to model over- and under-dispersed sex allocation and show how to calculate Bayes factors for comparing these alternative models to the null hypothesis of binomial sex allocation. The ability to detect non-binomial sex allocation is greatly increased, particularly in cases where mortality is common. The use of Bayesian methods allows for the quantification of the evidence in favour of each hypothesis, and our modelling approach provides an improved descriptive capability over existing approaches. We use a simulation study to situations where current methods fail, and we illustrate the approach in real scenarios using empirically obtained datasets on the sexual composition of groups of gregarious parasitoid wasps demonstrate substantial improvements in power for detecting non-binomial sex allocation in situations where current methods fail, and we illustrate the approach in real scenarios using empirically obtained datasets on the sexual composition of groups of gregarious parasitoid wasps

Objective and subjective components of resource value in lethal fights between male entomopathogenic nematodes

Males sometimes engage in fights over contested resources such as access to mates; in this case, fighting behaviour may be adjusted based on the value they place on the females. Resource value (RV) can have two components. First, males can assess the quality of females, which constitutes an objective assessment of RV. Second, internal state such as previous mating experience can also influence motivation to fight thus constituting a subjective assessment of RV. If mating opportunities are scarce and available females have a major impact on the lifetime reproductive success of males, then fighting can be fatal; in this situation it is uncertain whether males would adjust fighting behaviour based on RV. We found that both female quality, that is, virginity (objective component of RV) and male mating status (subjective component of RV) influenced fighting intensity between males of the entomopathogenic nematode Steinernema longicaudum which engage in lethal fights. Male nematodes were more likely to engage in fighting and fought longer and more frequently in the presence of virgin (high-quality) females than in the presence of mated (lower-quality) females. Male mating status was also found to influence fighting behaviour: mated males were the winners in staged fights between mated and virgin males. Mated males may have superior fighting ability (greater resource-holding potential), but RV asymmetries between mated and virgin males cannot be excluded. Males were more likely to win when they were resident, but we did not find a significant interaction effect between male mating and residency status

Sustenance and performance: nutritional reserves, longevity and contest outcomes of fed and starved adult parasitoid wasps

Dyadic contests for possession of resources occur across a wide range of animal taxa, with contest outcome often being heavily influenced by the energetic reserves of the competitors. The majority of studied parasitoid wasps lack de novo lipogenesis, with adult lipid reserves being limited to those acquired throughout larval development. Carbohydrate- and lipid-rich adult diets can increase adult parasitoid lifespan and fecundity by potentially acting as a maintenance energy store. However, the effects of such diets on fat reserve compositions and some aspects of parasitoid behaviour contest outcome dynamics have not been examined. This study assesses the effect of a carbohydrate rich diet (honey) on the longevity, metabolic state and contest performance of Goniozus legneri, a bethylid wasp. The longevity of honey-fed diet adults was typically more than twice that of starved wasps. Compared to similarly-aged starved wasps, honey feeding increased concentrations of common haemolymph sugars and amino acids in a 3-day old and 7-day old wasps, and higher concentrations of stored lipid in7-day old wasps. However, nutritional status did not affect the outcome of dyadic contests over host possession when both contestants were either 3 days or 7 days old. While contest dynamics outcome may be unaffected by diet, it remains possible that an enhancing effect of feeding on contest ability is matched by an effect of higher value being placed on winning by starved wasps

Annual Review of Entomology

Parasitoids of the hymenopterous family Encyrtidae are one of the most important groups of natural enemies of soft scale insects and have been used extensively in biological control. We summarize existing knowledge of the biology, ecology, and behavior of these parasitoids and how it relates to biological control. Soft scale stage/size and phenology are important determinants of host range and host utilization, which are key aspects in understanding how control by these parasitoids is exerted. Furthermore, the nutritional ecology of encyrtids and their physiological interactions with their hosts affect soft scale insect population dynamics. Lastly, the interactions among encyrtids, heteronomous parasitoids, and ants shape parasitoid species complexes and consequently have a direct impact on the biological control of soft scale insects

Data from: Higher relatedness mitigates mortality in a nematode with lethal male fighting

According to kin selection theory, individuals show less aggression towards their relatives. Limited dispersal promotes interactions among relatives but also increases competition among them. The evolution of cooperation in viscous populations has been subject of mainly theoretical exploration. We investigated the influence of relatedness on aggression in males of entomopathogenic nematode Steinernema longicaudum that engage in lethal fighting. In a series of in vitro experiments, we found that both competitor male group size and relatedness influence male mortality rates. Higher relatedness led to progressively lower rates of male mortality. In experimentally infected insects, wherein large numbers of males and females interact, the proportion of dead and paralysed (= terminally injured) males was higher when infection was established by infective juveniles originating from a mixture of three lines than in those infected by a single line. The results collectively show that Steinernema longicaudum males recognize their kin and consequently male mortality rates are lower in groups consisting of more related males. Furthermore, this monotonic negative relationship between aggression and relatedness suggests that kin selection benefits are still substantial even under extreme competition. Our experiments also suggest that kin recognition in entomopathogenic nematodes has a genetic basis rather than being strictly based on environmental cues. We discuss our findings within the theoretical context of the evolution of altruistic/cooperative behaviour in structured populations

Sequences of sex allocation and mortality in clutches of Metaphycus parasitoids of soft scale insects and the prevalence of all-female broods

International audience1. In many gregarious or quasi-gregarious parasitoids that experience local mate competition, precise sex ratios with low variance are observed. Precise sex ratios can be achieved by laying male and female eggs in non-random sequences. 2. Developmental mortality can also alter sex ratios of emerging offspring, and subsequently influence sex ratio optima. 3. The present study investigates sex allocation by Metaphycus flavus Howard, M. luteolus Timberlake, and M. angustifrons Compere (Hymenoptera: Encyrtidae), endoparasitoids of soft scale insects, in the laboratory. 4. All three Metaphycus species had precise secondary sex ratios when parasitising brown soft scale, Coccus hesperidum, L. in the laboratory. Moreover, we documented that all three species lay fertilised (= female) eggs first followed by unfertilised (= male) eggs at the end of the oviposition bout. However, there were significant differences in sex allocation sequences among species. 5. Mortality rates of eggs allocated within an oviposition bout also varied considerably, indicating that there is a significant interspecific variation in sequence position-specific mortality. 6. Using a stochastic Monte Carlo simulation approach, we provide evidence that the incidence of all-female broods in these parasitoid wasps appears mainly due to developmental mortality and not due to decisions by the ovipositing female. In two species the prevalence of all-female broods was independent of clutch size, contrary to what is expected from theory. The influence of mortality on optimal sex allocation in these parasitoids is discussed