The parasitoid wasp, Ooencyrtus kuvanae (Howard) (Hymenoptera: Encyrtidae), is a natural enemy of the spongy moth, a significant forest pest in North America. We investigated the oviposition behavior of O. kuvanae females on spongy moth egg masses by (i) presenting female parasitoids with a single spongy moth egg mass that was replaced every day, 2nd day, 4th day, 8th day, or 16th day (which is the total length of the oviposition period) and (ii) presenting female parasitoids with 1, 2, 4, or 8 egg masses at a time. Offspring developmental length ranged from 18 to 24 days. On average, male offspring exhibited faster developmental times, emerging approximately 1 day ahead of females. The amount of time that adult females spent on an egg mass affected the number of parasitized eggs. Specifically, more offspring emerged in the 4-, 8-, and 16-day treatments than in scenarios involving daily or every second-day egg mass replacement. The percentage of male offspring decreased as the number of egg masses presented to females increased. Interestingly, the total number of female offspring remained constant, but the number of male offspring decreased with an increase in the number of egg masses and time spent by the parent within a patch. The observed sexual dimorphism in development time, the influence of resource availability on offspring sex ratios, and flexible oviposition patterns illustrate the adaptability of O. kuvanae in response to varying conditions. These insights have implications for our understanding of parasitoid-host interactions and their potential role in biological control strategies.

The relative per-flower production of ovules and pollen varies broadly with angiosperm mating systems, with outcrossing types commonly producing more pollen grains per ovule than selfing types. The evolutionary causes of this variation are contentious, especially the relevance of pollination risk. Resolution of this debate may have been hampered by its focus on pollen:ovule (P:O) ratios rather than on the evolution of pollen and ovule numbers per se.
Using published mean ovule and pollen counts, we analyzed associations with the proportion of removed pollen that reaches stigmas (pollen-transfer efficiency) and differences between pollinator-dependent and autogamous forms within and among species. Analyses involved Bayesian methods that simultaneously considered variation in pollen and ovule numbers and accounted for phylogenetic relatedness. We also assessed the utility of P:O ratios as mating-system proxies and their association with female outcrossing rates.
Median pollen number declined consistently with pollen-transfer efficiency among species, whereas median ovule number did not. Similarly, in both intraspecific and interspecific analyses, pollinator-dependent plants produced more pollen than autogamous plants, whereas ovule production did not differ statistically. Distributions of P:O ratios overlapped extensively for self-incompatible and self-compatible species and for different mating-system classes, and P:O ratios correlated weakly with outcrossing rate.
Our findings demonstrate that pollinator dependence and pollination efficiency commonly influence the evolution of pollen number per flower but have more limited effects on ovule number. P:O ratios provide ambiguous, possibly misleading, information about mating systems, especially when compared among clades.

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.

A surprising result emerging from the theory of sex allocation is that the optimal sex ratio is predicted to be completely independent of the rate of dispersal. This striking invariance result has stimulated a huge amount of theoretical and empirical attention in the social evolution literature. However, this sex-allocation invariant has been derived under the assumption that an individual\'s dispersal behaviour is not modulated by population density. Here, we investigate how density-dependent dispersal shapes patterns of sex allocation in a viscous-population setting. Specifically, we find that if individuals are able to adjust their dispersal behaviour according to local population density, then they are favoured to do so, and this drives the evolution of female-biased sex allocation. This result obtains because, whereas under density-independent dispersal, population viscosity is associated not only with higher relatedness-which promotes female bias-but also with higher kin competition-which inhibits female bias-under density-dependent dispersal, the kin-competition consequences of a female-biased sex ratio are entirely abolished. We derive analytical results for the full range of group sizes and costs of dispersal, under haploid, diploid and haplodiploid modes of inheritance. These results show that population viscosity promotes female-biased sex ratios in the context of density-dependent dispersal.

Haplo-diploid sex determination in the parasitoid wasp, Nasonia vitripennis (Walker), allows females to adjust their brood sex ratios. Females influence whether ova are fertilized, producing diploid females, or remain unfertilized, producing haploid males. Females appear to adjust their brood sex ratios to minimize \'local mate competition,\' i.e., competition among sons for mates. Because mating occurs between siblings, females may optimize mating opportunities for their offspring by producing only enough sons to inseminate daughters when ovipositing alone, and producing more sons when superparasitism is likely. Although widely accepted, this hypothesis makes no assumptions about gamete limitation in either sex. Because sperm are used to produce daughters, repeated oviposition could reduce sperm supplies, causing females to produce more sons. In contrast, if egg-limited females produce smaller broods, they might use fewer sperm, making sperm limitation less likely. To investigate whether repeated oviposition and female fertility influence gamete limitation within females, we created two treatments of six mated female wasps, which each received a series of six hosts at intervals of 24 or 48 h. All females produced at least one mixed-sex brood (63 total broods; 3,696 offspring). As expected, if females became sperm-limited, in both treatments, brood sex ratios became increasingly male-biased with increasing host number. Interhost interval did not affect brood size, total offspring number, or sex ratio, indicating females did not become egg limited. Our results support earlier studies showing sperm depletion affects sex allocation in N. vitripennis¸ and could limit adaptive sex ratio manipulation in these parasitoid wasps.

Sex allocation theory in simultaneous hermaphrodites predicts that optimal sex allocation is influenced by local sperm competition, which occurs when related sperm compete to fertilize a given set of eggs. Different factors, including the mating strategy and the ability to self-fertilize, are predicted to affect local sperm competition and hence the optimal SA. Moreover, since the local sperm competition experienced by an individual can vary temporally and spatially, this can favour the evolution of sex allocation plasticity. Here, using seven species of the free-living flatworm genus Macrostomum, we document interspecific variation in sex allocation, but neither their mating strategy nor their ability to self-fertilize significantly predicted sex allocation among these species. Since we also found interspecific variation in sex allocation plasticity, we further estimated standardized effect sizes for plasticity in response to (i) the presence of mating partners (i.e. in isolation vs. with partners) and (ii) the strength of local sperm competition (i.e. in small vs. large groups). We found that self-fertilization predicted sex allocation plasticity with respect to the presence of mating partners, with plasticity being lower for self-fertilizing species. Finally, we showed that interspecific variation in sex allocation is higher than intraspecific variation due to sex allocation plasticity. Our study suggests that both sex allocation and sex allocation plasticity are evolutionarily labile, with self-fertilization predicting the latter in Macrostomum.

Synthetic gene drive constructs are being developed to control disease vectors, invasive species, and other pest species. In a well-mixed random mating population a sufficiently strong gene drive is expected to eliminate a target population, but it is not clear whether the same is true when spatial processes play a role. In species with an appropriate biology it is possible that drive-induced reductions in density might lead to increased inbreeding, reducing the efficacy of drive, eventually leading to suppression rather than elimination, regardless of how strong the drive is. To investigate this question we analyse a series of explicitly solvable stochastic models considering a range of scenarios for the relative timing of mating, reproduction, and dispersal and analyse the impact of two different types of gene drive, a Driving Y chromosome and a homing construct targeting an essential gene. We find in all cases a sufficiently strong Driving Y will go to fixation and the population will be eliminated, except in the one life history scenario (reproduction and mating in patches followed by dispersal) where low density leads to increased inbreeding, in which case the population persists indefinitely, tending to either a stable equilibrium or a limit cycle. These dynamics arise because Driving Y males have reduced mating success, particularly at low densities, due to having fewer sisters to mate with. Increased inbreeding at low densities can also prevent a homing construct from eliminating a population. For both types of drive, if there is strong inbreeding depression, then the population cannot be rescued by inbreeding and it is eliminated. These results highlight the potentially critical role that low-density-induced inbreeding and inbreeding depression (and, by extension, other sources of Allee effects) can have on the eventual impact of a gene drive on a target population.

Local mate competition (LMC) favours female biased clutch sex ratios because it reduces competition between brothers and provides extra mating opportunities for sons. Fig wasps seem to fit LMC model assumptions and lay female-biased sex ratios as predicted. These female biased sex ratios increase fitness greatly. In line with predictions, their sex ratios become less female-biased as the number of mothers laying in the same fig increases. However, this variation results in comparatively small fitness benefits compared to just biased ratios and data suggest substantial mismatches with LMC theory. The mismatches are due to several factors. (1) Multiple foundresses typically lay too many daughters. (2) Single foundress sex ratios are explained by sequential oviposition and ladies-last models. (3) Mortality that typically exceeds 10% may decouple the link between primary sex ratios, the focus of model predictions, and secondary sex ratios of adult wasps that are counted by researchers. (4) Model assumptions are frequently violated: (a) clutch sizes are unequal, (b) oviposition may not be simultaneous (c) cryptic/multiple wasp species inhabit the same host, (d) foundress numbers are systematically undercounted, (e) inbreeding coefficient calculations are inaccurate, and (f) male wasps sometimes disperse. These data and calculations suggest that alternative explanations must be considered seriously. Substantial data show that wasps typically lay most of their male eggs first followed by mostly female eggs require a new approach. These \"slope\" strategies result in more accurate sex ratios that are automatically adjusted to foundress number, own and relative clutch sizes and to sequential clutches. This effect will alter sex ratios in all species once the egg capacity of a fig is crossed or when interference reduces clutch sizes. In addition to this passive response, the females of about half the studied species have a conditional response that reduces female bias under higher foundress numbers by laying more sons. Therefore, wasps seem to use a very simple strategy that increases their fitness. Natural selection could have optimized parameters of the slope strategy and possibly the existence of the slope strategy itself. Variation in the slope strategy that is the result of natural selection is adaptive. Research should therefore focus on quantifying variables of this slope strategy. Currently, it is unclear how much of the variation is adaptive as opposed to being coincidental by-products.

Hamilton\'s local mate competition theory provided an explanation for extraordinary female-biased sex ratios in a range of organisms. When mating takes place locally, in structured populations, a female-biased sex ratio is favored to reduce competition between related males, and to provide more mates for males. However, there are a number of wasp species in which the sex ratios appear to more female biased than predicted by Hamilton\'s theory. It has been hypothesized that the additional female bias in these wasp species results from cooperative interactions between females. We investigated theoretically the extent to which cooperation between related females can interact with local mate competition to favor even more female-biased sex ratios. We found that (i) cooperation between females can lead to sex ratios that are more female biased than predicted by local competition theory alone, and (ii) sex ratios can be more female biased when the cooperation occurs from offspring to mothers before dispersal, rather than cooperation between siblings after dispersal. Our models formally confirm the verbal predictions made in previous experimental studies, which could be applied to a range of organisms. Specifically, cooperation can help explain sex ratio biases in Sclerodermus and Melittobia wasps, although quantitative comparisons between predictions and data suggest that some additional factors may be operating.

Parker, Baker, and Smith provided the first robust theory explaining why anisogamy evolves in parallel in multicellular organisms. Anisogamy sets the stage for the emergence of separate sexes, and for another phenomenon with which Parker is associated: sperm competition. In outcrossing taxa with separate sexes, Fisher proposed that the sex ratio will tend towards unity in large, randomly mating populations due to a fitness advantage that accrues in individuals of the rarer sex. This creates a vast excess of sperm over that required to fertilize all available eggs, and intense competition as a result. However, small, inbred populations can experience selection for skewed sex ratios. This is widely appreciated in haplodiploid organisms, in which females can control the sex ratio behaviorally. In this review, we discuss recent research in nematodes that has characterized the mechanisms underlying highly skewed sex ratios in fully diploid systems. These include self-fertile hermaphroditism and the adaptive elimination of sperm competition factors, facultative parthenogenesis, non-Mendelian meiotic oddities involving the sex chromosomes, and environmental sex determination. By connecting sex ratio evolution and sperm biology in surprising ways, these phenomena link two \"seminal\" contributions of G. A. Parker.