Number Of Foundresses Research Articles

Exploring the Biology of Quasi-Social Idiobiont Parasitoids in the Genus Sclerodermus (Hymenoptera: Bethylidae)

Species in the genus Sclerodermus are among the most socially complex parasitoids, unlike most parasitoids, which are solitary and do not provide care after laying eggs. In Sclerodermus, groups of females paralyse their host, lay eggs on it, and work together to care for the brood (a quasi-social form of reproduction). This research, through database analysis and meta-analysis, covers the biology of the genus, which has 80 species, though only 24 have been studied in detail. It describes their morphology and behaviour, focusing on offspring production, developmental time, and the factors influencing these, such as kinship and the number of females tending the brood. The materials and methods used provide a comprehensive approach to data collection and analysis, drawing on diverse sources, rigorous classification, and advanced statistical techniques. This approach revealed that Sclerodermus species display a high degree of consistency in their responses to temperature, host size, and foundress number.

The Effect of Ficus semicordata Fig Quality on the Sex Ratio of Its Pollinating Wasp Ceratosolen gravelyi

The interaction between fig wasps and their host fig trees (Ficus spp.) is a striking example of an obligate pollination mutualism. Male and female fig wasps are confined within their natal patch instead of panmictic; under this circumstance, mating only occurs between individuals of the same patch. This is known as a local mate competition (LMC). It pays foundresses to invest mainly in daughters and to only produce enough sons to ensure that all female offspring can be fertilized, but in nature, pollinating fig wasps may face many problems with host quality, such as limitation of oviposition sites and the nutrition deficiency of the host fig. The sex ratio of wasps can determine the stability of fig–fig wasp mutualistic system and, thus, the stability of other species associated with it. In this study, we controlled the quality of host figs in three ways. The results showed that the host fig age can influence the sex ratio of pollinator offspring, while the foundress numbers and the presence of pollen have no significant effect on it. A compelling explanation for this result is that the sex-dependent mortality occurs. This is a novel finding of how host quality influences the interaction of fig and fig wasps, which can also help us understand the evolution and stability mechanism of this symbiotic system.

Who holds the reins? Context-dependent resource allocation in the mutualism between fig trees and their fig wasp pollinators.

Mutualisms are consumer-resource interactions, in which goods and services are exchanged. Biological market theory states that exchanges should be regulated by both partners. However, most studies on mutualisms are one-sided, focusing on the control exercised by host organisms on their symbionts. In the brood-site pollination mutualism between fig trees and their symbiont wasp pollinators, galled flowers are development sites for pollinator larvae and are exchanged for pollination services. We determined if pollinator galls influenced resource allocation to fig inflorescences called syconia and considered feedbacks from the host tree. We experimentally produced syconia containing only seeds (S), only pollinator galls (G) or seeds and galls (SG) with varying number of introduced female pollinator wasps, i.e., foundress wasps. Biomass allocation to syconia was affected by foundress numbers and treatment groups; SG treatments received highest biomass allocation at low foundress numbers, and both G and SG treatments at high foundress numbers. Seeds are important determinants of allocation at low foundress numbers; galls are likely more influential at high foundress numbers. Most allocation in the G and SG treatment was to the syconium wall, likely as protection from parasitoids and temperature/humidity fluctuations. Dry mass of individual seeds and wasps (except at low foundress numbers) was unchanged between treatment groups, indicating seeds and wasps regulate resource flow into them, with lower flow into galls containing the smaller males compared to females commensurate with sexual dimorphism. We demonstrate the importance of considering the direct role of symbionts in accessing resources and controlling exchanges within mutualisms.

Local ecological factors, not interference competition, drive the foundress number of two species of fig wasp sharing Ficus septica figs.

Recent studies have challenged assumptions about the classic fig-fig wasp pollination mutualism model, suggesting that further investigation into the receptive phase of fig development is needed. This study assessed the pollination mechanisms of Ficus septica in southern Taiwan and identified two species of wasps as the primary pollinators. Machine learning was used to identify and rank the factors that explain the relative abundance of these wasps. The two wasp species showed the highest level of cohabitation ever reported in the literature, with three-quarters of the figs containing multiple foundresses. The study also reported re-emerged foundresses and a 10% ratio of pollinated figs without foundresses. Local factors, such as the sampling period and tree identity, were the best predictors of the presence and number of each foundress species, with fig size also affecting the number of foundresses. The study highlights the variability in pollinator abundance between figs, crops, and trees. It also shows that the local environment of the trees and the availability of figs are crucial factors in determining which figs the pollinator wasps choose. These findings challenge assumptions about the classic mutualism model and suggest that long-term surveys are needed to estimate the relative contributions of each partner and provide data for evolutionary and ecological models. This study also provides valuable insights into the factors that affect the abundance and interactions of pollinator wasps during the receptive phase of fig development, with implications for understanding the behaviour of pollinating wasps and advancing our knowledge of population dynamics in Ficus species.

Individual- and group-level sex ratios under local mate competition: consequences of infanticide and reproductive dominance.

Extremely female-biased sex ratios of parasitoid wasps in multiple-foundress groups challenges evolutionary theory which predicts diminishing bias as foundress numbers increase. Recent theory based on foundress cooperation has achieved qualitative rather than quantitative success in explaining bias among parasitoids in the genus Sclerodermus. Here, we develop an explanation, expanding the theory of local mate competition, based on the observation that male production seems dominated by some foundresses within groups. Two sex ratio effects arise from such reproductive dominance: an immediate effect via suppression of male production, and a long-term evolutionary response to reproductive skew. We analyze the outcome of these effects at the individual and group level, the latter being more readily observable. Three model scenarios are analyzed: (1) random killing of developing sons in a group by all foundresses, without reproductive skew, (2) the development of reproductive dominance by some foundresses after sex allocation decisions by all foundresses have been implemented, and (3) reproductive dominance within foundress groups before sex allocation decisions are implemented. The 3 scenarios have subtly different implications for sex ratio evolution, with Models 2 and 3 being novel additions to theory, showing how reproductive dominance can alter the outcome of sex ratio evolution. All models match observations in their outcomes better than other recently proposed theory, but Models 2 and 3 are closest to observations in their underlying assumptions. Further, Model 2 shows that differential offspring mortality after parental investment can influence the primary sex ratio even when random with respect to parental and offspring characters, but targeted at entire clutches. The novel models are solved for both diploid and haplodiploid genetic systems, and confirmed with simulations. Overall, these models provide a feasible explanation for the extremely female-biased sex ratios produced by multi-foundress groups and expand the scope of local mate competition theory to consider reproductive dominance.

Testing the local mate competition rule in a quasi-gregarious parasitoid with facultative superparasitism

Abstract As an important domain of evolutionary ecology, sex allocation theory well explains the evolution of investment into female versus male offspring. Local mate competition (LMC) is a good predictor of sex allocation, where the optimal sex ratio becomes less female-biased and asymptotically approaches 0.5 as the number of foundresses increases. Parasitoid wasps, with haplodiploid sex determination, offer excellent opportunities to test how organisms manipulate their offspring sex ratio in response to environmental variation, and many species have been proved to allocate sex according to predictions under LMC. When hosts are spatially clustered, as in gregarious species, the mating systems of quasi-gregarious parasitoids meet the essential assumptions (female mating before dispersal) of LMC. However, inconsistent with predictions, in the quasi-gregarious species Anastatus disparis (Hymenoptera: Eupelmidae), a strongly female-biased eclosion sex ratio (0.156 ± 0.018 to 0.185 ± 0.016) was observed as the number of females laying eggs in a patch increased. Superparasitism, in which 44.7% of parasitized hosts contained more than one egg but only one adult emerged from each host, was common in A. disparis. However, the egg sex ratio was determined by microsatellites and likely fit the predictions of LMC theory. Male-biased offspring mortality arising from superparasitism during development likely contributes to the shift from the primary sex ratio predicted under LMC to the observed female-biased eclosion sex ratio. Inconsistent with results in gregarious parasitoids, the role of superparasitism in driving sex ratio shifts in quasi-gregarious parasitoids should be incorporated into LMC-based predictions of sex ratios.

Nepotistic colony fission in dense colony aggregations of an Australian paper wasp

Social insects are highly diverse in their social structures, aside from the consistent presence of reproductive castes. Among social insects, the Australian paper wasp Ropalidia plebeiana constructs extremely dense colony aggregations consisting of hundreds of colonies within a few square meters; however, little is known about the aggregation structures. We genetically analyzed the colony and population structure of R. plebeiana, and concomitant variations in colony sex ratios. In spring, the foundress (candidate queen) group started their colonies on a single old comb from the previous season, subsequently dividing these old combs via relatedness-based comb-cutting. Female philopatry, a prerequisite condition of Local Resource Competition (LRC), was confirmed. The colony sex ratio of reproductive individuals (male and female offspring for the next generation) became slightly male-biased in larger colonies, as predicted under LRC. However, the number of foundresses was positively associated with the number of reproductive individuals, suggesting that Local Resource Enhancement (LRE) also operates. Although the population structure appears to meet the prerequisites of LRC, the sex ratio appears to be modulated by factors other than LRC. Rather, through LRE, the availability of female helpers at the founding stage is likely to mitigate the sex ratios predicted under LRC.

How many cooperators are too many? Foundress number, reproduction and sex ratio in a quasi‐social parasitoid

AbstractIn the parasitoid genusSclerodermus, multiple foundresses produce and care for communal broods on large hosts, which can lead to greater reproductive success for group members than attempting to reproduce alone.We explore the consequences of foundress group size on the benefits of cooperative brooding and on brood sex ratios by providing groups of 10–55 foundresses with a single host and no alternative reproductive options.Within this range, increasing foundress group size leads to increasingly common failure in brood production and diminishedper capitasuccess. Group production of adult offspring declines once foundress number reaches around 25.Brood failure is usually at the early developmental stages, and current evidence suggests that there may be competition among foundresses for oviposition sites, possibly involving reproductive dominance and ovicide, which also delays initial brood production. Once broods become established, their rate of development is enhanced by large foundress numbers.The sex ratios of broods are very strongly female biased, irrespective of the foundress number. As this bias is not easily explained by standard models of local mate competition or by a recent model of local resource enhancement, we suggest an explanation based on control of sex allocation by a minority of dominant foundresses, which monopolise the production of adult males.

Co-foundress confinement elicits kinship effects in a naturally sub-social parasitoid.

Kinship among interacting individuals is often associated with sociality and also with sex ratio effects. Parasitoids in the bethylid genus Goniozus are sub-social, with single foundress females exhibiting post-ovipositional maternal care via short-term aggressive host and brood defence against conspecific females. Due to local mate competition (LMC) and broods normally being produced by a single foundress, sex ratios are female-biased. Contests between adult females are, however, not normally fatal, and aggression is reduced when competing females are kin, raising the possibility of multi-foundress reproduction on some hosts. Here, we screen for further life-history effects of kinship by varying the numbers and relatedness of foundresses confined together with a host resource and also by varying the size of host. We confined groups of 1-8 Goniozus nephantidis females together with a host for 5+days. Multi-foundress groups were either all siblings or all nonsiblings. Our chief expectations included that competition for resources would be more intense among larger foundress groups but diminished by both larger host size and closer foundress relatedness, affecting both foundress mortality and reproductive output. From classical LMC theory, we expected that offspring group sex ratios would be less female-biased when there were more foundresses, and from extended LMC theory, we expected that sex ratios would be more female-biased when foundresses were close kin. We found that confinement led to the death of some females (11% overall) but only when host resources were most limiting. Mortality of foundresses was less common when foundresses were siblings. Developmental mortality among offspring was considerably higher in multi-foundress clutches but was unaffected by foundress relatedness. Groups of sibling foundresses collectively produced similar numbers of offspring to nonsibling groups. There was little advantage for individual females to reproduce in multi-foundress groups: single foundresses suppressed even the largest hosts presented and had the highest per capita production of adult offspring. Despite single foundress reproduction being the norm, G.nephantidis females in multi-foundress groups appear to attune sex allocation according to both foundress number and foundress relatedness: broods produced by sibling foundresses had sex ratios similar to broods produced by single foundresses (ca. 11% males), whereas the sex ratios of broods produced by nonsibling females were approximately 20% higher and broadly increased with foundress number. We conclude that relatedness and host size may combine to reduce selection against communal reproduction on hosts and that, unlike other studied parasitoids, G.nephantidis sex ratios conform to predictions of both classical and extended LMC theories.

Egg load is a cue for offspring sex ratio adjustment in a fig-pollinating wasp with male-eggs-first sex allocation.

Fig-pollinating wasps (Agaonidae) only reproduce within fig treeinflorescences (figs). Agaonid offspring sex ratios are usually female-biased and often concur with local mate competition theory (LMC). LMC predicts less female-bias when several foundresses reproduce in a fig due to reduced relatedness among intra-sexually competing male offspring. Clutch size, the offspring produced by each foundress, is a strong predictor of agaonid sex ratios and correlates negatively with foundress number. However, clutch size variation can result from several processes including egg load (eggs within a foundress), competition among foundresses and oviposition site limitation, each of which can be used as a sex allocation cue. We introduced into individual Ficus racemosa figs single Ceratosolen fusciceps foundresses and allowed each to oviposit from zero to five hours thus variably reducing their eggs-loads and then introduced each wasp individually into a second fig. Offspring sex ratio (proportion males) in second figs correlated negatively with clutch size, with males produced even in very small clutches. Ceratosolen fusciceps lay mainly male eggs first and then female eggs. Our results demonstrate that foundresses do not generally lay or attempt to lay a 'fixed' number of males, but do 'reset to zero' their sex allocation strategy on entering a second fig. With decreasing clutch size, gall failure increased, probably due to reduced pollen. We conclude that C.fusciceps foundresses can use their own egg loads as a cue to facultatively adjust their offspring sex ratios and that foundresses may also produce more 'insurance' males when they can predict increasing rates of offspring mortality.

Foundress numbers and the timing of selective events during interactions between figs and fig wasps

In intimate mutualisms between hosts and symbionts, selection can act repeatedly over the development times of the interacting individuals. Although much is now known about the overall ecological conditions that favor the evolution of mutualism, a current challenge is to understand how natural selection acts on the number and kinds of partners to shape the evolution and stability of these interactions. Using the obligate fig-fig wasp mutualism, our experiments showed that the proportion of figs developed to maturity increased quickly to 1.0 as the number of foundresses increased, regardless of whether the foundresses carried pollen. Selection against pollen-free wasps did not occur at this early stage in fig development. Within figs that developed, the proportion of galls producing adult wasps remained high as the number of pollen-carrying foundresses increases. In contrast, the proportion of galls producing adult wasps decreased as the number of pollen-free foundresses increased. Viable seed production increased as the number or proportion of pollen-carrying foundresses increased, but the average number of wasp offspring per pollen-carrying foundress was highest when she was the sole foundress. These results show that figs and their pollinator wasps differ in how fitness effects are distributed throughout the development of the interaction and depend on the number and proportion of pollen-carrying foundresses contributing to the interaction. These results suggest that temporal fluctuations in the local number and proportion of pollen-carrying wasps available to enter figs are likely to have strong but different effects on the figs and the wasps.

Differential deployment of sanctioning mechanisms by male and female host trees in a gynodioecious fig-wasp mutualism.

In some insect nursery pollination mutualisms, plant hosts impose net costs to uncooperative "cheater" symbionts. These "sanctions" promote mutualism stability but their precise adaptive nature remains unclear. In fig-wasp mutualisms host trees (Ficus spp.) are only pollinated by female agaonid wasps whose larvae only use galled fig flowers as food. In actively pollinated systems, if wasps fail to pollinate, sanctions can result via fig abortion, killing all wasp offspring, or by increased offspring mortality within un-aborted figs. These sanctions result from selective investment to pollinated inflorescences, a mechanism present in almost all angiosperms. To more fully understand how selective investment functions as sanctions requires the measurement of variation in their costs and benefits to both hosts and symbionts. Gynodioecious fig-tree-fig-wasp mutualisms are particularly suitable for this because pollen and wasps are produced only in the figs of "male" trees and seeds only in the figs of "female" trees. Male and female trees thus incur different net costs of pollen absence, and costs of sanctions to pollen-free "cheater" wasps only occur in male trees. We used the actively pollinated host tree Ficus hispida and introduced into male and female figs either 1, 3, 5, 7, or 9 all pollen-laden "cooperative" (P+) or all pollen-free "cheater" (P-) wasps. Abortion in both male and female trees was highest in P- figs, with P- fig abortion higher in females (~90%) than in males (~40%). Fig abortion was negatively associated with foundress number mainly in P+ figs; in P- figs abortion was only weakly associated with the number of "cheater" wasps, especially in female figs. In un-aborted male figs, wasp offspring mortality was higher in P- figs than in P+ figs, and in P- figs correlated positively with foundress (cheater) number. Increased offspring mortality was biased against female wasp offspring and likely resulted from reduced larval nutrition in unpollinated flowers. Variation in selective investment to P- figs thus reflects costs and benefits of pollen absence/presence to hosts, variation that translates directly to net costs to cheater wasps.

Evolutionary stable sex ratios with non‐facultative male‐eggs first sex allocation in fig wasps

Sex allocation theory has long generated insights into the nature of natural selection. Classical models have elucidated causal phenomena such as local mate competition and inbreeding on the degree of female bias exhibited by various invertebrates. Typically, these models assume mothers facultatively adjust sex allocation using predictive cues of future offspring mating conditions. Here we relax this assumption by developing a sex allocation model for haplodiploid mothers experiencing local mate competition that lay a fixed number of male eggs first. Female egg number is determined by remaining oviposition sites or remaining eggs of the mother, depending on which is exhausted first. Our model includes parameters for variation in foundress number, patch size, fecundity and offspring mortality that allow us to generate secondary sex ratio predictions based on specific parameterizations for natural populations. Simulations show that: 1) in line with classical models, factors that increase sib‐mating result in mothers laying relatively more female eggs; 2) high offspring mortality leads to relatively more males as fertilization insurance; 3) unlike classical model predictions, sub‐optimal predictions, such as more males than females are possible. In addition, our model provides the first quantitative predictions for the expected number of males and females in a patch where typically only one mother utilizes a given patch. We parameterized the model with data obtained from seven species of southern African fig wasps to predict expected means and variances for numbers of male and female offspring for typical numbers of mothers utilizing a patch. These predictions were compared to secondary sex ratio data from single foundress patches, the most commonly encountered situation for these species. Our predictions matched both the observed number and variance of male and female offspring with a high degree of accuracy suggesting that facultative adjustment is not required to produce evolutionary stable sex ratios.

Offspring sex ratio shifts of the solitary parasitoid wasp, Trichopria drosophilae (Hymenoptera: Diapriidae), under local mate competition

Localmate competition (LMC) models predict a female-biased offspring sex ratio when a single foundress oviposits alone in a patch and an increasing proportion of sons with increasing foundress number. We tested whether the solitary pupal parasitoid, Trichopria drosophilae (Hymenoptera: Diapriidae), adjusted offspring sex ratio with foundress number when parasitizing Drosophila melanogaster pupae. Mean number of female offspring was higher than that of males, with a male proportion of 26 ± 16% when only one foundress oviposited. However, male proportion reached 58 ± 26%, 48 ± 22%, and 51 ± 19% in three-, five and seven-foundress cohorts. That the male proportion of offspring increased with foundress number is consistent with LMC models.

Differential gene expression is not required for facultative sex allocation: a transcriptome analysis of brain tissue in the parasitoid wasp Nasonia vitripennis.

Whole-transcriptome technologies have been widely used in behavioural genetics to identify genes associated with the performance of a behaviour and provide clues to its mechanistic basis. Here, we consider the genetic basis of sex allocation behaviour in the parasitoid wasp Nasonia vitripennis. Female Nasonia facultatively vary their offspring sex ratio in line with Hamilton's theory of local mate competition (LMC). A single female or ‘foundress’ laying eggs on a patch will lay just enough sons to fertilize her daughters. As the number of ‘foundresses’ laying eggs on a patch increases (and LMC declines), females produce increasingly male-biased sex ratios. Phenotypic studies have revealed the cues females use to estimate the level of LMC their sons will experience, but our understanding of the genetics underlying sex allocation is limited. Here, we exposed females to three foundress number conditions, i.e. three LMC conditions, and allowed them to oviposit. mRNA was extracted from only the heads of these females to target the brain tissue. The subsequent RNA-seq experiment confirmed that differential gene expression is not associated with the response to sex allocation cues and that we must instead turn to the underlying neuroscience to reveal the underpinnings of this impressive behavioural plasticity.

Spatial variation in pollinator gall failure within figs of the gynodioecious Ficus hirta

Figs, the inflorescences of Ficus species (Moraceae), contain numerous uni-ovulate flowers. Male trees of gynodioecious Ficus have figs that support development of pollinator fig wasp offspring (Agaonidae) and rarely produce seeds. Pollinator larvae develop inside galled ovules that expand rapidly after eggs are laid to fill the available space. Galls that fail to support successful larval development can be abundant and failures may influence oviposition behavior and modify realized offspring sex ratios. We examined pollinator reproductive success in figs of the Asian Ficus hirta where we had allowed entry by either one or two foundresses and prevented attack by parasitoids. At the developmental stage when adult offspring were about to emerge from their galls, we recorded where in the figs their galls were located, the distributions of sons and daughters in the galls and whether galls that developed closest to the periphery of the figs were more likely to fail. Foundress number had an effect on gall location, but not total offspring numbers. No spatial variation in the distribution of male and female adult offspring was detected. Overall, over 25% of the galled ovaries failed to support offspring development, and failure rates were independent of foundress number. More peripheral galls were more likely to fail in figs entered by two foundresses. Gall location in gynodioecious figs is determined largely by the extent to which their basal pedicels expand after galling. Competition for nutrients between galls, with those developing shorter pedicels being at a disadvantage, may explain the results. If pedicel length is related to timing of oviposition, then pollinator eggs laid later are less likely to survive.

Molecular markers reveal reproductive strategies of non‐pollinating fig wasps

1. Fig wasps have proved extremely useful study organisms for testing how reproductive decisions evolve in response to population structure. In particular, they provide textbook examples of how natural selection can favour female‐biased offspring sex ratios, lethal combat for mates and dimorphic mating strategies.2. However, previous work has been challenged, because supposedly single species have been discovered to be a number of cryptic species. Consequently, new studies are required to determine population structure and reproductive decisions of individuals unambiguously assigned to species.3. Microsatellites were used to determine species identity and reproductive patterns in three non‐pollinating Sycoscapter species associated with the same fig species. Foundress number was typically one to five and most figs contained more than one Sycoscapter species. Foundresses produced very small clutches of about one to four offspring, but one foundress may lay eggs in several figs.4. Overall, the data were a poor match to theoretical predictions of solitary male clutches and gregarious clutches with n − 1 females. However, sex ratios were male‐biased in solitary clutches and female‐biased in gregarious ones.5. At the brood level (all wasps in a fig), a decrease in sex ratio with increasing brood size was only significant in one species, and sex ratio was unrelated to foundress number. In addition, figs with more foundresses contain more wasp offspring.6. Finally, 10–22% of females developed in patches without males. As males are wingless, these females disperse unmated and are constrained to produce only sons from unfertilised eggs.

Colony productivity of Polistes dominula (Christ) (Hymenoptera, Vespidae: Polistinae) wasps nesting on plants by haplometrosis: Regression analysis

The paper presents the results of analysis of the relationships of various colony characteristics in the resocial wasp Polistes dominula nesting on plants in the south of Ukraine (Kherson Province, the Black Sea Biosphere Reserve) in 2003–2007. The number of future foundresses and nest size at the end of the life cycle depend on the queen longevity and on the number of workers in the colony. The number of males reared in the colony is positively correlated with the nest size (the number of cells) and negatively correlated with the queen longevity. An increase in the share of the brood infested by the parasitoids Latibulus argiolus (Hymenoptera, Ichneumonidae) and Elasmus schmitti (Hymenoptera, Eulophidae) results in a smaller nest size and a smaller number of males reared.

Biology of two members of the Euwallacea fornicatus species complex (Coleoptera: Curculionidae: Scolytinae), recently invasive in the U.S.A., reared on an ambrosia beetle artificial diet

Abstract Recent molecular studies have found that the ambrosia beetle Euwallacea fornicatus Eichhoff (Coleoptera: Curculionidae: Scolytinae) is a complex of cryptic species, each carrying a different species of symbiotic fungus, in the genus Fusarium, which they farm within galleries inside woody hosts. Several of these beetle species have become invasive pests around the world for attacking and infecting healthy trees with their phytopathogenic fungal symbionts. Diet and rearing protocols were developed for two members of the E. fornicatus species complex, polyphagous shot hole borer (PSHB) and tea shot hole borer (TSHB), using sawdust from host trees, allowing collection of data on beetle biology, phenology and sex ratios. Adults developed within 22 days at 24 °C. Single PSHB or TSHB foundresses averaged 32.4 and 24.7 adult female offspring, respectively, and up to 57 and 68 female adults within 6–7 weeks. A strong predictor of the number of offspring in a colony was the number of entry holes. Average sex ratios (% male) for PSHB and TSHB, respectively, were 7.4% and 7.2%. Being haplodiploid, virgin PSHB foundresses were able to produce and mate with male offspring, then subsequently produce female offspring, confirming that they have arrhenotokous reproduction. A cold tolerance study found significant mortality rates among PSHB colonies exposed to −5° or −1 °C but not colonies exposed to 0°, 1° or 5 °C. Given Hamilton's local mate competition (LMC) theory, a number of LMC predictions were violated. PSHB sex ratios were not affected by the number of foundresses; approximately 14% of broods did not contain males; males did not usually eclose before females but eclosed around the same time (22–23 days); and PSHB males were found walking outside of their natal galleries on the trunk of a heavily infested tree in the field. Alternatives to LMC are considered, such as early forms of sociality (maternal care, cooperative brood care), local resource enhancement and kin selection.

Determination of the optimal parasitoid‐to‐host ratio for efficient mass‐rearing of the parasitoid, Sclerodermus pupariae (Hymenoptera: Bethylidae)

AbstractSclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae) is used as a potential biocontrol agent for several buprestid and cerambycid larvae. This study aimed to enhance the efficiency of mass‐rearing of this parasitoid by investigating the fitness gain of this bethylid wasp, including the proportion of successful parasitism and development, brood size, sex ratio, proportion of winged female offspring, body size and longevity of female offspring, under eight different maternal parasitoid density treatments using Thyestilla gebleri Faldermann as host in the laboratory. The results indicated that the foundress densities did not affect the parasitism or emergence rate of this parasitoid. Brood size of the parasitoids increased significantly when the number of maternal wasps ranged from one to four. However, further increases in foundress number did not affect the parasitoid brood size. The sex ratios of S. pupariae were always female‐biased. The proportions of male in the progeny colonies were <10% throughout all experimental treatments. The percentage of winged female progeny was not significantly influenced by the density of adult maternal parasitoids. Body sizes of parasitoids significantly declined with increasing maternal parasitoid densities. Although the parasitoid body size reduced when maternal wasp number was higher, it could be compromised by the relatively higher number of female offspring produced. Further, more than 70% of the parasitoids remained alive when they were stored at 12°C for four months throughout the experiments. These findings suggest that exposure of four female wasps to a single host larva would result in the highest fitness of S. pupariae. Our findings might provide a new approach to enhance the efficiency of mass‐rearing of this bethylid wasp.