Size‐dependent sex allocation in a simultaneous hermaphrodite parasite

Most models of sex allocation distinguish between sequential and simultaneous hermaphrodites, although an intermediate sexual pattern, size‐dependent sex allocation, is widespread in plants. Here we investigated sex allocation in a simultaneous hermaphrodite animal, the tapeworm Schistocephalus solidus, in which adult size is highly variable. Sex allocation was determined using stereological techniques, which allow measuring somatic and reproductive tissues in a common currency, namely volume. We investigated the relationships between individual volume and allocation to different reproductive tissues using an allometric model. One measure of female allocation, yolk gland volume, increased more than proportionally with individual volume. This is in contrast to the measure of male allocation, testis volume, which showed a strong tendency to increase less than proportionally with individual volume. Together these patterns led to sex allocation being strongly related to individual volume, with large individuals being more biased towards female allocation. We discuss these findings in the light of current ideas about size‐dependent sex allocation in, primarily, plants and try to extend them to simultaneous hermaphrodite animals.     

Frequency‐dependent two‐sex models: a new approach to sex ratio evolution with multiple maternal conditions

Mothers that experience different individual or environmental conditions may produce different proportions of male to female offspring. The Trivers‐Willard hypothesis, for instance, suggests that mothers with different qualities (size, health, etc.) will use different sex ratios if maternal quality differentially affects sex‐specific reproductive success. Condition‐dependent, or facultative, sex ratio strategies like these allow multiple sex ratios to coexist within a population. They also create complex population structure due to the presence of multiple maternal conditions. As a result, modeling facultative sex ratio evolution requires not only sex ratio strategies with multiple components, but also two‐sex population models with explicit stage structure. To this end, we combine nonlinear, frequency‐dependent matrix models and multidimensional adaptive dynamics to create a new framework for studying sex ratio evolution. We illustrate the applications of this framework with two case studies where the sex ratios depend one of two possible maternal conditions (age or quality). In these cases, we identify evolutionarily singular sex ratio strategies, find instances where one maternal condition produces exclusively male or female offspring, and show that sex ratio biases depend on the relative reproductive value ratios for each sex.     

Brood sex ratio variation in a cooperatively breeding bird

In cooperatively breeding species, the fitness consequences of producing sons or daughters depend upon the fitness impacts of positive (repayment hypothesis) and negative (local competition hypothesis) social interactions among relatives. In this study, we examine brood sex allocation in relation to the predictions of both the repayment and the local competition hypotheses in the cooperatively breeding long-tailed tit Aegithalos caudatus. At the population level, we found that annual brood sex ratio was negatively related to the number of male survivors across years, as predicted by the local competition hypothesis. At an individual level, in contrast to predictions of the repayment hypothesis, there was no evidence for facultative control of brood sex ratio. However, immigrant females produced a greater proportion of sons than resident females, a result consistent with both hypotheses. We conclude that female long-tailed tits make adaptive decisions about brood sex allocation.     

Partial local mate competition in the wasp Trichogramma euproctidis: the role of emergence sex ratio on female mating behaviour

1. Parasitic wasps with structured populations are generally assumed to follow the local mate competition (LMC) model: females lay only the minimal number of sons necessary to inseminate all daughters in the emergence patch, and increase this number when faced with additional broods from unrelated females. After emergence, daughters mate with local males before dispersing for host location and oviposition. The main predictions from the model have been verified for many species. 2. Conflicting evidence exists on the status of the egg parasitoids Trichogramma regarding their on‐patch versus off‐patch mating. Although the life‐history traits of several species indicate that mating must occur on the emergence patch, recent data suggest that mating could occur outside the natal patch. 3. In this study, we measured the level of off‐patch mating in the egg parasitoid Trichogramma euproctidis using two isofemale lines in a greenhouse experiment. The impact of the sex ratio on the level of off‐patch mating was also tested. 4. The overall off‐patch mating proportion was 40.5% with a range between 0 and 85.7%, and was influenced by the sex ratio on the emergence patch: the more males available at emergence, the less off‐patch mating occurring. 5. The mating structure of this species can be described as partial LMC.     

Production of male flowers does not decrease with plant size in insect‐pollinated Sagittaria trifolia,contrary to predictions of size‐dependent sex allocation

Abstract It has been proposed that relative allocation to female function increases with plant size in animal‐pollinated species. Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size‐dependent sex allocation (SDS), throwing doubt on the generalization of SDS. Plant size, phenotypic gender, and flower production were measured in experimental populations of an aquatic, insect‐pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities. The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors. In the high‐density population, 48% of ramets were male without female flowers, but in the low‐density population all ramets were monoecious. We observed allometric growth in reproductive allocation with ramet size, as evident in biomass of reproductive structures and number of flowers. However, within both populations female and male flower production were isometric with ramet size, in contrast to an allometric growth in femaleness as predicted by SDS. Phenotypic gender was not related to ramet size in either population. The results indicated that large plants may increase both female and male function even in animal‐pollinated plants, pointing towards further studies to test the hypothesis of size‐dependent sex allocation using different allocation currencies.     

Budding dispersal and the sex ratio

There is much interest in understanding how population demography impacts upon social evolution. Here, we consider the impact of rate and pattern of dispersal upon a classic social evolutionary trait--the sex ratio. We recover existing analytical results for individual dispersal, and we extend these to allow for budding dispersal. In particular, while a cancelling of relatedness and kin competition effects means that the sex ratio is unaffected by the rate of individual dispersal, we find that a decoupling of relatedness and kin competition means that budding dispersal favours increasingly female-biased sex ratios. More generally, our analysis illustrates the relative ease with which biological problems involving class structure can be solved using a kin selection approach to social evolution theory.     

The influence of developmental mortality on optimal sex allocation under local mate competition

In panmictic populations, optimal sex allocation is, under theassumptions of Fisher's model, not influenced by the probabilityof offspring developmental mortality, or by differences in mortalitybetween the sexes. In contrast, when mating opportunities areconfined to siblings, developmental mortality can influenceoptimal sex allocation. Many animal species have both localmating and developmental mortality. We show that when developmentalmortality is random for individual offspring, optimal sex allocationis influenced by mortality among males but not among females.Male mortality increases the allocation to males, but this shouldnever be male biased, even under extreme male mortality. Thisresult applies both when mothers are able to control the sexof individual offspring precisely, and when sex is allocatedwith binomial probability. The influence of mortality becomesprogressively larger when the variance of the distribution ofmortality over clutches diminishes. The reduction in fitnessis greater than the proportion of mortality, especially at smallclutch sizes, and mortality reduces the advantage of producingprecise sex ratios, and of local mate competition in general.     

Spatiotemporal fluctuations in natural selection acting on the gall‐parasitic aphid Tetraneura sorini

The measurement of the selection gradient is crucial for understanding the magnitude of selection acting directly on a trait and predicting the evolutionary trajectory of that trait. This study evaluated the selection gradient acting on the morphology of the gall‐parasitic aphid Tetraneura sorini during the galling process and compared the strength among populations. Gall formers (first instars) frequently fight with conspecifics or heterospecifics for usurping incipient galls using their well‐developed hind legs. First instars that successfully acquired galls were found within galls, whereas those that failed were found dead on leaf surfaces. Selection gradients were estimated using logistic stepwise regression and partial least square (PLS) regression. Calculated selection differentials indicated that first instars that secured galls were larger in body size than failed individuals through all populations. However, selection gradients on weapon traits varied largely among populations or among years in the same population. We confirmed microevolutionary changes in the relationship between traits, which accorded with the expectation from changes in the selection gradients. When gall formers were transferred onto developing buds individually, individuals that successfully induced galls had smaller body size than failed individuals. Available evidence suggests that the selection gradient on body size becomes higher with an increasing proportion of T. sorini in the Tetraneura species community. Thus, we concluded that more intense fighting with conspecifics leads to stronger selective pressure on body size, but that selective pressure for each trait is variable depending on differences in the tactics and species composition among populations.     

Impact of competition on sex allocation by Trichogramma

Hymenopteran parasitoids change their sex ratio following different factors. One of these factors is the exploitation of a host patch by several females. The Local Mate Competition (LMC) model ( Hamilton, 1967 ) states that when there are many foundresses on a patch, they should lay a higher sex ratio. The impact of both intra‐ and interspecific competition on sex allocation was measured in two egg parasitoids, Trichogramma minutum Riley and T. pintoi Voegele (Hymenoptera: Trichogrammatidae), by comparing the sex ratio deposited by females exploiting host patches alone and in groups. Results showed that the sex ratio deposited by both species was higher when in groups, as predicted by the LMC model. When the sex ratio produced was compared between females either alone or in interspecific groups, T. minutum females deposited the same sex ratio, while T. pintoi produced more sons when in interspecific groups than when alone. These results are discussed following their natural habitat and their discrimination abilities.     

Pollinating fig wasp Ceratosolen solmsi adjusts the offspring sex ratio to other foundresses

Abstract Local mate competition theory predicts that offspring sex ratio in pollinating fig wasps is female‐biased when there is only one foundress, and increased foundress density results in increased offspring sex ratio. Information of other foundresses and clutch size have been suggested to be the main proximate explanations for sex ratio adjustment under local mate competition. Our focus was to show the mechanism of sex ratio adjustment in a pollinating fig wasp, Ceratosolen solmsi Mayr, an obligate pollinator of the functionally dioecious fig, Ficus hispida Linn., with controlled experiments in the field. First, we obtained offspring from one pollinator and offspring at different oviposition sequences, and found that offspring sex ratio decreased with clutch size, and pollinators produced most of their male offspring at the start of bouts, followed by mostly females. Second, we found that offspring sex ratio increased with foundress density, and pollinators did adjust their offspring sex ratio to other females in the oviposition patches. We suggest that when oviposition sites are not limited, pollinators will mainly adjust their offspring sex ratio to other foundresses independent of clutch size changes, whereas adjusting clutch size may be used to adjust sex ratio when oviposition sites are limited.     

Unifying cornerstones of sexual selection: operational sex ratio,Bateman gradient and the scope for competitive investment

What explains variation in the strength of sexual selection across species, populations or differences between the sexes? Here, we show that unifying two well‐known lines of thinking provides the necessary conceptual framework to account for variation in sexual selection. The Bateman gradient and the operational sex ratio (OSR) are incomplete in complementary ways: the former describes the fitness gain per mating and the latter the potential difficulty of achieving it. We combine this insight with an analysis of the scope for sexually selected traits to spread despite naturally selected costs. We explain why the OSR sometimes does not affect the strength of sexual selection. An explanation of sexual selection becomes more logical when a long ‘dry time’ (‘time out’, recovery after mating due to e.g. parental care) is understood to reduce the expected time to the next mating when in the mating pool (i.e. available to mate again). This implies weaker selection to shorten the wait. An integrative view of sexual selection combines an understanding of the origin of OSR biases with how they are reflected in the Bateman gradient, and how this can produce selection for mate acquisition traits despite naturally selected costs.     

Facultative adjustment of the offspring sex ratio and male attractiveness: a systematic review and meta‐analysis

Females can benefit from mate choice for male traits (e.g. sexual ornaments or body condition) that reliably signal the effect that mating will have on mean offspring fitness. These male‐derived benefits can be due to material and/or genetic effects. The latter include an increase in the attractiveness, hence likely mating success, of sons. Females can potentially enhance any sex‐biased benefits of mating with certain males by adjusting the offspring sex ratio depending on their mate's phenotype. One hypothesis is that females should produce mainly sons when mating with more attractive or higher quality males. Here we perform a meta‐analysis of the empirical literature that has accumulated to test this hypothesis. The mean effect size was small (r = 0.064–0.095; i.e. explaining <1% of variation in offspring sex ratios) but statistically significant in the predicted direction. It was, however, not robust to correction for an apparent publication bias towards significantly positive results. We also examined the strength of the relationship using different indices of male attractiveness/quality that have been invoked by researchers (ornaments, behavioural displays, female preference scores, body condition, male age, body size, and whether a male is a within‐pair or extra‐pair mate). Only ornamentation and body size significantly predicted the proportion of sons produced. We obtained similar results regardless of whether we ran a standard random‐effects meta‐analysis, or a multi‐level, Bayesian model that included a correction for phylogenetic non‐independence. A moderate proportion of the variance in effect sizes (51.6–56.2%) was due to variation that was not attributable to sampling error (i.e. sample size). Much of this non‐sampling error variance was not attributable to phylogenetic effects or high repeatability of effect sizes among species. It was approximately equally attributable to differences (occurring for unknown reasons) in effect sizes among and within studies (25.3, 22.9% of the total variance). There were no significant effects of year of publication or two aspects of study design (experimental/observational or field/laboratory) on reported effect sizes. We discuss various practical reasons and theoretical arguments as to why small effect sizes should be expected, and why there might be relatively high variation among studies. Currently, there are no species where replicated, experimental studies show that mothers adjust the offspring sex ratio in response to a generally preferred male phenotype. Ultimately, we need more experimental studies that test directly whether females produce more sons when mated to relatively more attractive males, and that provide the requisite evidence that their sons have higher mean fitness than their daughters.     

The quantitative genetic basis of sex ratio variation in Nasonia vitripennis: a QTL study

Our understanding of how natural selection should shape sex allocation is perhaps more developed than for any other trait. However, this understanding is not matched by our knowledge of the genetic basis of sex allocation. Here, we examine the genetic basis of sex ratio variation in the parasitoid wasp Nasonia vitripennis, a species well known for its response to local mate competition (LMC). We identified a quantitative trait locus (QTL) for sex ratio on chromosome 2 and three weaker QTL on chromosomes 3 and 5. We tested predictions that genes associated with sex ratio should be pleiotropic for other traits by seeing if sex ratio QTL co-occurred with clutch size QTL. We found one clutch size QTL on chromosome 1, and six weaker QTL across chromosomes 2, 3 and 5, with some overlap to regions associated with sex ratio. The results suggest rather limited scope for pleiotropy between these traits.     

Social situation,sperm competition and sex allocation in a simultaneous hermaphrodite parasite,the cestode Schistocephalus solidus

Evolutionary theory predicts an influence of mating group size on sex allocation in simultaneous hermaphrodites. We experimentally manipulated the social situation during reproduction in a simultaneous hermaphrodite parasite, the tapeworm Schistocephalus solidus, by placing worms as singles, pairs or triplets into an in vitro system that replaces the final host. We then determined the reproductive allocation patterns after 24 h (i.e. before the start of egg release) and after 72 h (i.e. around the peak of egg release rate) using stereology. After 24 h, sex allocation strongly depended on worm volume (which is determined in the second intermediate host), but was not significantly affected by the social situation experienced during reproduction. After 72 h, worms in groups had less vesicular sperm (i.e. sperm to be used in future inseminations) than singles. They also stored significantly more received sperm in their seminal receptacles than singles, suggesting that more sperm had been transferred in groups. Moreover, worms in triplets stored significantly more received sperm than worms in pairs, suggesting that they either mated more often and/or transferred more sperm per mating. This suggests a behavioural response to the increased risk of sperm competition in triplets. We further discuss the relative importance of sex allocation decisions at different life‐history stages.     

Can the extremely female‐biased sex ratio of the social spider mites be explained by Hamilton’s local mate competition model?

Abstract.  1. Extremely female-biased sex ratios are known in the social spider mite species, Stigmaeopsis longus and S. miscanthi . Whether Hamilton's local mate competition (LMC) theory can explain such sex ratios was investigated.
2. Significant changes of the progeny sex ratios in the direction predicted by the LMC model were found in both species when the foundress number changed. Therefore, LMC can partly explain the skewed sex ratios in these species.
3. When the foundress number increased, the progeny sex ratio was still female biased and significantly different from the prediction of the LMC model for haplodiploidy. Relatedness between foundresses could not fully explain the female-biased sex ratios. Therefore, these results suggest that there are factors other than LMC skewing the sex ratios of these species toward female.     

Inbreeding effects on progeny sex ratio and gender variation in the gynodioecious Silene vulgaris (Caryophyllaceae)

In gynodioecious species, sex expression is generally determined through cytoplasmic male sterility genes interacting with nuclear restorers of the male function. With dominant restorers, there may be an excess of females in the progeny of self-fertilized compared with cross-fertilized hermaphrodites. Moreover, the effect of inbreeding on late stages of the life cycle remains poorly explored. Here, we used hermaphrodites of the gynodioecious Silene vulgaris originating from three populations located in different valleys in the Alps to investigate the effects of two generations of self- and cross-fertilization on sex ratio and gender variation. We detected an increase in females in the progeny of selfed compared with outcrossed hermaphrodites and inbreeding depression for female and male fertility. Male fertility correlated positively with sex ratio differences between outbred and inbred progeny, suggesting that dominant restorers are likely to influence male fertility qualitatively and quantitatively in S. vulgaris. We argue that the excess of females in the progeny of selfed compared with outcrossed hermaphrodites and inbreeding depression for gamete production may contribute to the maintenance of females in gynodioecious populations of S. vulgaris because purging of the genetic load is less likely to occur.     

Density‐dependent and density‐independent drivers of population change in Barton Springs salamanders

Understanding population change is essential for conservation of imperiled species, such as amphibians. Worldwide amphibian declines have provided an impetus for investigating their population dynamics, which can involve both extrinsic (density‐independent) and intrinsic (density‐dependent) drivers acting differentially across multiple life stages or age classes. In this study, we examined the population dynamics of the endangered Barton Springs Salamander (Eurycea sosorum) using data from a long‐term monitoring program. We were interested in understanding both the potential environmental drivers (density‐independent factors) and demographic factors (interactions among size classes, negative density dependence) to better inform conservation and management activities. We used data from three different monitoring regimes and multivariate autoregressive state‐space models to quantify environmental effects (seasonality, discharge, algae, and sediment cover), intraspecific interactions among three size classes, and intra‐class density dependence. Results from our primary data set revealed similar patterns among sites and size classes and were corroborated by our out‐of‐sample data. Cross‐correlation analysis showed juvenile abundance was most strongly correlated with a 9‐month lag in aquifer discharge, which we suspect is related to inputs of organic carbon into the aquifer. However, sedimentation limited juvenile abundance at the surface, emphasizing the importance of continued sediment management. Recruitment from juveniles to the sub‐adult size class was evident, but negative density‐dependent feedback ultimately regulated each size class. Negative density dependence may be an encouraging sign for the conservation of E. sosorum because populations that can reach carrying capacity are less likely to go extinct compared to unregulated populations far below their carrying capacity. However, periodic population declines coupled with apparent migration into the aquifer complicate assessments of species status. Although both density‐dependent and density‐independent drivers of population change are not always apparent in time series of animal populations, both have important implications for conservation and management of E. sosorum.     

母代雌蜂数、进果时间及非传粉小蜂对传粉榕小蜂后代数量及性比的影响

【目的】对性比的研究有助于我们理解自然界生物的选择压力及其所产生的原因和结果,榕树和榕小蜂之间的互惠共生关系以及生物学和生态学特性使其成为研究性比和局域配偶竞争模型（local mate competition）的理想材料。本研究旨在探明榕小蜂性比调节和进化机制。【方法】对分布于西双版纳地区的鸡嗉子榕Ficus semicordata进行了人工控制性放蜂实验。测定了母代雌蜂数量及其进果时间间隔、非传粉小蜂Sycoscapter trifemmensis数量对传粉榕小蜂Ceratosolen gravelyi后代数量（成蜂数量）和性比的影响,并分析了小蜂后代数量和性比的相关性。【结果】在榕果发育期一致的前提下,随着母代雌蜂数量的增加,每头雌蜂的平均后代数量明显下降(P<0.001),后代性比显著升高(P<0.001),后代数量和性比呈显著负相关(P<0.05);随着雌蜂进果间隔的延长,后代数量亦呈现下降趋势,且性比增大,放2头雌蜂和3头雌蜂的处理呈同样趋势,但差异均不显著(P=0.87; P=0.49),小蜂后代数量与性比无显著相关性(P=0.86)。此外,非传粉小蜂数量与传粉小蜂后代数量呈显著负相关(P<0.001),与传粉小蜂性比呈正相关(P<0.001),小蜂后代数量和性比同样呈现显著负相关(P<0.001)。【结论】本实验模拟了自然界中榕 蜂的相互作用,所得结果有助于我们理解自然状态下榕小蜂性比调节模式和机制,以及榕 蜂互利共生系统的进化机制。     

母体亲缘关系在杨氏榕树金小蜂后代性比调节中的作用

局域配偶竞争(local mate competition, LMC)理论问世以来在很多方面获得了发展,其中一个观点就是相互作用的繁殖母体之间的亲缘关系越近,后代性比越低,即产生更多的雌性后代。本研究以西双版纳广泛分布的钝叶榕Ficus curtipes上寄生的非传粉小蜂杨氏榕树金小蜂Diaziella yangi为研究对象,研究在同一榕果上产卵的繁殖雌蜂之间的亲缘关系对后代性比的影响。处理1：在同一榕果产卵的2头繁殖雌蜂来自蜂源1号树的同一个榕果;处理2：在同一榕果产卵的2头繁殖雌蜂来自蜂源2号树的同一个榕果,处理3：在同一榕果上产卵的2头繁殖蜂分别来自1、2号蜂源树。实验结果却显示出与理论预测值不一致。实验结果中,具有姐妹关系的2头繁殖雌蜂的2个处理的后代性比分别为0.195±0.028和0.189±0.043;亲缘关系较远的2头繁殖雌蜂的后代性比为0.240±0.030;不同处理的后代性比并没有显著差异。说明杨氏榕树金小蜂没有识别相互作用的繁殖雌蜂之间的亲缘关系以及据此调整后代性比的能力,这与其他大量研究 结果一致。