Use of population genetic data to infer oviposition behaviour: species-specific patterns in four oak gallwasps (Hymenoptera: Cynipidae)

Many species of oak gallwasp (Hymenoptera: Cynipidae: Cynipini) induce galls containing more than one larva (multilocular galls) on their host plant. To date, it has remained unclear whether multilocular galls result solely from clustered oviposition by a single female, or include the aggregated offspring of several females (multiple founding). We have developed a novel maximum-likelihood approach for use with population genetic data that estimates the number and genotypes of parents contributing to offspring from each gall. We apply this method to allozyme data from multiple populations of four oak gallwasps whose asexual generations develop in multilocular galls (Andricus coriarius, A. lucidus, A. panteli and A. seckendorffi). We find strong evidence for multiple founding in all four species, and show the data to be compatible with multiple founding rather than founding by a single foundress mated with multiple males. The extent of multiple founding differs among species: in A. lucidus and A. seckendorffi most galls are induced by a single female, whereas in A. coriarius and A. panteli over half of the galls sampled were multiple founded. We suggest that variation in levels of multiple founding may be due to consistent ecological differences between the four species.     

Sex ratio in relation to season and host plant quality in a monogenous stem-galling midge (Diptera: Cecidomyiidae)

Abstract.  1. The sex determination mechanism in gall midges is little understood, although it is known that the females of several species primarily or exclusively produce unisexual broods throughout their lifetime.
2. The gall midge Izeniola obesula Dorchin is a multivoltine species, inducing multi-chambered stem galls on the salt-marsh plant Suaeda monoica . Each gall contains 5–70 individuals, all being the progeny of a single female. Sampling of more than 450 galls, from which adult midges were reared, suggested that I .  obesula exhibits strict monogeny, resulting in galls that contain either all female or all male progeny.
3. Characterisation of the growth pattern of young S .  monoica shoots revealed that shoots in apical positions grew more rapidly than shoots in more basal positions. Galls that were induced on such shoots were larger and yielded more adult midges.
4. No difference in the site of gall induction was found between male and female galls, with galls of either sex being mainly induced on more rapidly growing shoots. It is concluded that I .  obesula females cannot control the sex of their progeny, and that both female-producers and male-producers strive to maximise their reproductive success by choosing the faster-growing shoots for gall induction.
5. Female galls were larger and more abundant than male galls at almost all times. The sex ratio among galls fluctuated throughout the year, ranging from 4:1 in spring to 1:1 in winter. The skewed sex ratio among galls possibly results from greater mortality rates among male galls than among female galls, due to either primary or secondary factors. Alternatively, it is possible that the number or fitness of male-producers in the population is reduced relative to female-producers.     

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.     

Pre-ovulation control of hatchling sex ratio in the Seychelles warbler

Females of some bird species have a high degree of control over the sex ratio of their offspring at laying. Although several mechanisms have been put forward to explain how females might control the sex of their eggs, virtually nothing is known. As females are the heterogametic sex in birds, adjustment of the clutch sex ratio could arise either by pre- or post-ovulation control mechanisms. The Seychelles warbler (Acrocephalus sechellensis) exhibits extreme adaptive egg sex ratio bias. Typically, warblers produce only single-egg clutches, but by translocating pairs to vacant habitat of very high quality, most females were induced to produce two-egg clutches. Overall, females skewed clutch sex ratios strongly towards daughters (86.6%). This bias was evident in the first egg, but critically, also in the second eggs laid a day apart, even when all absent, unhatched, or unsexed second eggs were assumed to be male. Although a bias in the first egg may arise through either pre- or post-ovulation mechanisms, the skew observed in second eggs could only arise through pre-ovulation control. Post-ovulation adjustment may also contribute to skewed hatchling sex ratios, but as sex-biased release of gametes is likely to be a more efficient process of control, pre-ovulation mechanisms may be the sole means of adjustment in this species. High fitness differentials between sons and daughters, as apparent in the Seychelles warblers, may be necessary for primary sex ratio adjustment to evolve.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

Factors Influencing Realized Sex Ratios in Fig Wasps: Double Oviposition and Larval Mortalities

Pollinator fig wasps (Agaonidae) are a model system for studies of sex ratio evolution. They lay their eggs in galled ovules within figs. Only one adult emerges from each gall, suggesting that only one egg is always laid per ovule, but if double oviposition occurs then the assumption that adult (realised) sex ratios of fig wasps are representative of primary sex ratios may be violated. Many galls also fail to produce any wasps. If they initially contained eggs then differential mortality rates may also modify realized sex ratios. We investigated whether Kradibia (= Liporrhopalum) tentacularis foundresses in Ficus montana figs avoid laying in ovules that already contain eggs. Comparisons of oviposition frequencies and wasp emergence frequencies showed that most galls that failed to produce wasps will have had eggs laid in them, but few occupied ovules contained two eggs. Realised sex ratios therefore do not necessarily reflect primary sex ratios in this species, but double oviposition is not responsible.     

The structure of cynipid oak galls: patterns in the evolution of an extended phenotype

Galls are highly specialized plant tissues whose development is induced by another organism. The most complex and diverse galls are those induced on oak trees by gallwasps (Hymenoptera: Cynipidae: Cynipini), each species inducing a characteristic gall structure. Debate continues over the possible adaptive significance of gall structural traits; some protect the gall inducer from attack by natural enemies, although the adaptive significance of others remains undemonstrated. Several gall traits are shared by groups of oak gallwasp species. It remains unknown whether shared traits represent (i) limited divergence from a shared ancestral gall form, or (ii) multiple cases of independent evolution. Here we map gall character states onto a molecular phylogeny of the oak cynipid genus Andricus, and demonstrate three features of the evolution of gall structure: (i) closely related species generally induce galls of similar structure; (ii) despite this general pattern, closely related species can induce markedly different galls; and (iii) several gall traits (the presence of many larval chambers in a single gall structure, surface resins, surface spines and internal air spaces) of demonstrated or suggested adaptive value to the gallwasp have evolved repeatedly. We discuss these results in the light of existing hypotheses on the adaptive significance of gall structure.     

Fungal endophytes which invade insect galls: insect pathogens,benign saprophytes,or fungal inquilines?

Fungi are frequently found within insect galls. However, the origin of these fungi, whether they are acting as pathogens, saprophytes invading already dead galls, or fungal inquilines which invade the gall but kill the gall maker by indirect means, is rarely investigated. A pathogenic role for these fungi is usually inferred but never tested. I chose the following leaf-galling-insect/host-plant pairs (1) a cynipid which forms two-chambered galls on the veins of Oregon white oak, (2) a cynipid which forms single-chambered galls on California coast live oak, and (3) an aphid which forms galls on narrowleaf cottonwood leaves. All pairs were reported to have fungi associated with dead insects inside the gall. These fungi were cultured and identified. For the two cynipids, all fungi found inside the galls were also present in the leaves as fungal endophytes. The cottonwood leaves examined did not harbor fungal endophytes. For the cynipid on Oregon white oak, the fungal endophyte grows from the leaf into the gall and infects all gall tissue but does not directly kill the gall maker. The insect dies as a result of the gall tissue dying from fungal infection. Therefore, the fungus acts as an inquiline. Approximately 12.5% of these galls die as a result of invasion by the fungal endophyte.     

Evidence for Strategic Sex Allocation in the Guppy (Poecilia reticulata): Brood Sex Ratio and Size Predict Subsequent Adult Male Mating Success

Sex allocation theory predicts that females should produce more sons when the reproductive success of sons is expected to be high, whereas they should produce more daughters, not daughters when the reproductive success of sons is expected to be low. The guppy (Poecilia reticulata) is a live‐bearing fish, and female guppies are known to produce broods with biased sex ratios. In this study, we examined the relationship between brood sex ratio and reproductive success of sons and daughters, to determine whether female guppies benefit from producing broods with biased sex ratios. We found that sons in male‐biased broods had greater mating success at maturity than sons in female‐biased broods when brood sizes were larger. On the other hand, the reproductive output of daughters was not significantly affected by brood sizes and sex ratios. Our results suggest that female guppies benefit from producing large, male‐biased brood when the reproductive success of sons is expected to be high.     

Host gall size and oviposition success by the parasitoid Eurytoma gigantea

Abstract. 1. Eurytoma gigantea Walsh is a specialist parasitoid of the tephritid gallmaker Eurosta solidaginis (Fitch).
2. In the natural environment the incidence of parasitism by Eurytoma is greater in small galls than in large ones.
3. Laboratory experiments demonstrated that small galls are not more frequently discovered; however, oviposition attempts on small galls were more likely to be successful.
4. Eurytoma spends much time probing galls too big to penetrate; this leads to a decrease in foraging efficiency when many large galls are present.
5. The chance of successfully penetrating a gall depends on the thickness of the gall wall and the length of the parasitoid's ovipositor.
6. A simulation model was constructed which shows that a gallmak-er's chance of being parasitized depends on gall size, the number of parasitoids that discover the gall, and their ovipositor lengths.     

Hatching sex ratio and sex specific chick mortality in common terns Sterna hirundo

Bias in sex ratios at hatching and sex specific post hatching mortality in size dimorphic species has been frequently detected, and is usually skewed towards the production and survival of the smaller sex. Since common terns Sterna hirundo show a limited sexual size dimorphism, with males being only about 1–6% larger than females in a few measurements, we would expect to find small or no differences in production and survival of sons and daughters. To test this prediction, we carried out a 2-year observational study on sex ratio variation in common terns at hatching and on sex specific post hatching mortality. Sons and daughters hatched from eggs of similar volume. Post hatching mortality was heavily influenced by hatching sequence. In addition, we detected a sex specific mortality bias towards sons. Overall, hatching sex ratio and sex specific mortality resulted in fledging sex ratios 8% biased towards females. Thus, other reasons than body size may be influencing the costs of rearing sons. Son mortality was not homogeneous between brood sizes, but greater for two-chick broods. Since adults rearing two-chick broods were younger, lighter and bred consistently later than those rearing three-chick broods, it is suggested that lower capacity of two-chick brood parents adversely affected offspring survival of sons. Though not significantly, two-chick broods tended to be female biased at hatching, perhaps to counteract the greater male-biased nestling mortality. Thus, population bias in secondary sex ratio is not limited to strongly size dimorphic species, but species with a slight sexual size dimorphism can also show sex ratio bias through a combination of differential production and mortality of sons and daughters.     

ASSOCIATION BETWEEN SEX RATIO DISTORTION AND SEXUALLY ANTAGONISTIC FITNESS CONSEQUENCES OF FEMALE CHOICE

Genetic variation can be beneficial to one sex yet harmful when expressed in the other—a condition referred to as sexual antagonism. Because X chromosomes are transmitted from fathers to daughters, and sexually antagonistic fitness variation is predicted to often be X-linked, mates of relatively low-fitness males might produce high-fitness daughters whereas mates of high-fitness males produce low-fitness daughters. Such fitness consequences have been predicted to influence the evolution of female mating biases and the offspring sex ratio. Females might evolve to prefer mates that provide good genes for daughters or might adjust offspring sex ratios in favor of the sex with the highest relative fitness. We test these possibilities in a laboratory-adapted population of Drosophila melanogaster , and find that females preferentially mate with males carrying genes that are deleterious for daughters. Preferred males produce equal numbers of sons and daughters, whereas unpreferred males produce female-biased sex ratios. As a consequence, mean offspring fitness of unpreferred males is higher than offspring fitness of preferred males. This observation has several interesting implications for sexual selection and the maintenance of population genetic variation for fitness.     

In poor families, mothers' milk is richer for daughters than sons: A test of Trivers-Willard hypothesis in agropastoral settlements in Northern Kenya

The Trivers–Willard hypothesis predicts the unequal parental investment between daughters and sons, depending on maternal condition and offspring reproductive potential. Specifically, in polygynous populations where males have higher reproductive variance than females, it predicts that mothers in good condition will invest more in sons, whereas mothers in poor condition will invest more in daughters. Previous studies testing this hypothesis focused on behavioral investment, whereas few examined biological investment. This study investigates the Trivers–Willard hypothesis on both behavioral and biological parental investment by examining breastfeeding frequencies and breast milk fat concentrations. Data from exclusively breastfeeding mothers in Northern Kenya were used to test hypotheses: Economically sufficient mothers will breastfeed sons more frequently than daughters, whereas poor mothers will breastfeed daughters more frequently than sons, and economically sufficient mothers will produce breast milk with higher fat concentration for sons than daughters, whereas poor mothers will produce breast milk with higher fat concentration for daughters than sons. Linear regression models were applied, using breastfeeding frequency or log‐transformed milk fat as the dependent variable, and offspring's sex (son = 1/daughter = 0), socioeconomic status (higher = 1/lower = 0), and the sex‐wealth interaction as the predictors, controlling for covariates. Our results only supported the milk fat hypothesis: infant's sex and socioeconomic status interacted (P = 0.014, n = 72) in their relation with milk fat concentration. The model estimated that economically sufficient mothers produced richer milk for sons than daughters (2.8 vs. 0.6 gm/dl) while poor mothers produced richer milk for daughters than sons (2.6 vs. 2.3 gm/dl). Further research on milk constituents in relation to offspring's sex is warranted. Am J Phys Anthropol , 2012. © 2012 Wiley Periodicals, Inc.     

Intralocus sexual conflict,adaptive sex allocation,and the heritability of fitness

Intralocus sexual conflict arises when selection favours alternative fitness optima in males and females. Unresolved conflict can create negative between‐sex genetic correlations for fitness, such that high‐fitness parents produce high‐fitness progeny of their same sex, but low‐fitness progeny of the opposite sex. This cost of sexual conflict could be mitigated if high‐fitness parents bias sex allocation to produce more offspring of their same sex. Previous studies of the brown anole lizard (Anolis sagrei) show that viability selection on body size is sexually antagonistic, favouring large males and smaller females. However, sexual conflict over body size may be partially mitigated by adaptive sex allocation: large males sire more sons than daughters, whereas small males sire more daughters than sons. We explored the evolutionary implications of these phenomena by assessing the additive genetic (co)variance of fitness within and between sexes in a wild population. We measured two components of fitness: viability of adults over the breeding season, and the number of their progeny that survived to sexual maturity, which includes components of parental reproductive success and offspring viability (RS^V). Viability of parents was not correlated with adult viability of their sons or daughters. RS^V was positively correlated between sires and their offspring, but not between dams and their offspring. Neither component of fitness was significantly heritable, and neither exhibited negative between‐sex genetic correlations that would indicate unresolved sexual conflict. Rather, our results are more consistent with predictions regarding adaptive sex allocation in that, as the number of sons produced by a sire increased, the adult viability of his male progeny increased.     

Gall wasps and their parasitoids in cork oak fragmented forests

Abstract.  1. This paper explores the potential effects of host-plant fragmentation on cork oak gall wasp populations (Cynipidae, Hymenoptera) and on their predators, lethal inquilines, and parasitoids. To address this objective, galls were collected across a gradient of cork oak ( Quercus suber ) forest fragmentation in the East Pyrenees (Albera, Spain), and they were incubated to obtain the parasitism rates.
2. Two hypotheses were tested: (1) Host-plant fragmentation may induce a decline in gall wasp populations because of area and isolation effects on local extinction and dispersal; as a consequence of that, parasitoids may decline even more strongly in fragmented habitats than their prey. (2) Host-plant fragmentation may cause a decline in gall wasp parasitoid populations that, in turn, can lead to an ecological release in their prey populations.
3. Among the eight cork oak gall wasps sampled in the study area of Albera, the gall abundances of three species ( Callirhytis glandium , Callirhytis rufescens , and Andricus hispanicus ) were significantly related to forest fragmentation. The overall abundance of gall wasps was affected by a radius of ≈ 890 m surrounding landscape, presenting constant abundances with forest loss until forest cover is reduced at ≈ 40%; below that value the abundance increased rapidly. Three inquilines and 23 parasitoids species were recorded after gall incubation. In 25 cases, species of inquilines and parasitoids were newly recorded for the corresponding host in the Iberian peninsula.
4. Although the overall parasitism rate was high (1.1), it was uncorrelated with fragmentation and with overall cynipid abundance. These results indicate that host-plant fragmentation was correlated with higher abundance of gall wasps, whereas the parasitism rate could not explain this hyper-abundance in small forest fragments.     

Sex ratio and maternal investment in the multivoltine large carpenter bee Xylocopa sulcatipes (Apoideh: Anthophoridae)

Abstract. 1. Females of the multivoltine carpenter bee Xylocopa sulcutipes (Maa) (Hymenoptera: Anthophoridae) usually excavate a straight tunnel in dead twigs and mass provision a linear array of up to ten brood cells with pollen and nectar. An egg is deposited upon each food mass within one cell.
2. Female offspring generally receive a higher provisioning mass (0.180 ± 0.048 g) than males, a significant difference ( P > 0.001). There are, however, male larvae that receive as much food or more as their sisters or female larvae reared in another nest.
3. There is a close positive association between the size of a mother and the weight of provisions for individual daughters, but not for sons.
4. Female offspring are positioned in the innermost brood cells (Gositions 1, 2 and 3). The sex ratio of the outer cells is either significantly male biased (positions 4–6) or skewed towards males (positions 8 and 9). Positions 7 and 10 are in equilibrium.
5. Solitary females produce a significantly female biased sex ratio ( P < 0.01). Sex ratio in social nests is skewed toward females, but not significantly so ( P < 0.2). There is no significant difference between the sex ratio of solitary and social nests ( P = 0.361). The population sex ratio (pooled sex ratio of all broods produced) is significantly female biased ( P = 0.003).
6. Females kept in the laboratory produced female biased sex ratios whilst unmated females produced all-male broods indicating that insemination and ovarian development are not causally related.
7. The expected sex ratio (ESR) under equal investment, calculated as 1/CR (CR = mean male provision weight/mean female provision weight), is 137.5:117.5 (males:females), and differs significantly from that observed, 104:151 (males:females) ( P < 0.001). The 'Local Resource Enhlancement' hypothesis best explains the female biased sex ratio found in X.sulcatipes and its maintenance in the population.     

The parasitoid community of <Emphasis Type="Italic">Andricus quercuscalifornicus</Emphasis> and its association with gall size,phenology, and location

Plant galls are preyed upon by a diverse group of parasitoids and inquilines, which utilize the gall, often at the cost of the gall inducer. This community of insects has been poorly described for most cynipid-induced galls on oaks in North America, despite the diversity of these galls. This study describes the natural history of a common oak apple gall (Andricus quercuscalifornicus [Cynipidae]) and its parasitoid and inquiline community. We surveyed the abundance and phenology of members of the insect community emerging from 1234 oak apple galls collected in California’s Central Valley and found that composition of the insect community varied with galls of different size, phenology, and location. The gall maker, A. quercuscalifornicus, most often reached maturity in larger galls that developed later in the season. The parasitoid Torymus californicus [Torymidae] was associated with smaller galls, and galls that developed late in the summer. The most common parasitoid, Baryscapus gigas [Eulophidae], was more abundant in galls that developed late in the summer, though the percentage of galls attacked remained constant throughout the season. A lepidopteran inquiline of the gall (Cydia latiferreana [Tortricidae] and its hymenopteran parasitoid (Bassus nucicola [Braconidae]) were associated with galls that developed early in the summer. Parasitoids and inquilines, in general, had a longer emergence period and diapause within the gall than the gall-inducer. The association of different parasite species with galls of different size and phenology suggests that different parasite species utilize galls with slight differences in traits.     

Adaptive seasonal trend in brood sex ratio: test in two sister species with contrasting breeding systems

Evolutionary theory predicts adaptive adjustment in offspring sex ratio by females. Seasonal change in sex ratio is one possibility, tested here in two sister species, the Common sandpiper and the Spotted sandpiper Actitis hypoleucos and A. macularia. In the monogamous Common sandpiper, males are the most competitive sex. In each of 3 years, there was a change from mainly sons in early clutches to mainly daughters in late clutches. This seasonal adjustment of clutch sex ratio took place within the female before the eggs were laid, not by differential egg or chick survival. The sex of all eggs laid in the clutches used here was determined molecularly from chick blood taken at the time of hatching. The Spotted sandpiper in contrast is polyandrous, with partly reversed sex roles. There was no seasonal trend from sons to daughters in this species. When tested together, the two species differed significantly as predicted by the hypothesis of adaptive sex ratio adjustment by females.     

The protein content of tissues in cynipid galls (Hymenoptera: Cynipidae): Similarities between cynipid galls and seeds

Cynipid galls are examples of induced plant development, where the gall inducer is in control of cell differentiation and morphogenesis of a new plant organ. This study concentrates on the tissues of the larval chamber common to all cynipid galls. The protein content of the inner gall tissue was compared to that of non‐gall plant tissues. We investigated three oak and two rose galls and their respective host plants. Total protein signatures of inner gall tissues were different from those of non‐gall plant tissues, and among the five galls. N‐terminal sequences were obtained for two abundant proteins from the inner gall tissues of D. spinosa and A. quercuscalicis, which were common to all galls, at 62 and 43 kDa. Database queries suggest the 62 kDa protein to be homologous to a protein disulphide isomerase (PDI), and the 43 kDa protein to be homologous to NAD‐dependent formate dehydrogenase (FDH). A naturally biotinylated protein was detected at 33 kDa during Western analyses with streptavidin. Western analyses revealed the presence of the biotinylated protein and PDI in the inner gall tissues of all five galls, while FDH was only detected in A. quercuscalicis and A. fecundator. PDI was also common to all non‐gall tissues, while FDH was not detected in non‐gall tissues, and the biotinylated protein was only detected in seeds. The proteins identified in the inner gall tissue suggest that (a) inner gall tissues in some galls are under respiratory stress, and (b) cynipid gall formation might involve the ectopic expression of seed‐specific proteins.     

Condition Dependent Effects on Sex Allocation and Reproductive Effort in Sequential Hermaphrodites

Theory predicts the optimal timing of sex change will be the age or size at which half of an individual''s expected fitness comes through reproduction as a male and half through reproduction as a female. In this way, sex allocation across the lifetime of a sequential hermaphrodite parallels the sex allocation of an outbreeding species exhibiting a 1∶1 ratio of sons to daughters. However, the expectation of a 1∶1 sex ratio is sensitive to variation in individual condition. If individuals within a population vary in condition, high-condition individuals are predicted to make increased allocations to the sex with the higher variance in reproductive success. An oft-cited example of this effect is seen in red deer, Cervus elaphus, in which mothers of high condition are more likely to produce sons, while those in low condition are more likely to produce daughters. Here, we show that individual condition is predicted to similarly affect the pattern of sex allocation, and thus the allocation of reproductive effort, in sequential hermaphrodites. High-condition sex-changers are expected to obtain more than half of their fitness in the high-payoff second sex and, as a result, are expected to reduce the allocation of reproductive effort in the initial sex. While the sex ratio in populations of sequential hermaphrodites is always skewed towards an excess of the initial sex, condition dependence is predicted to increase this effect.