Sex allocation and interactions between relatives in the bean beetle, Callosobruchus maculatus

When a small number of females contribute offspring to a discrete mating group, sex allocation (Local Mate Competition: LMC) theory predicts that females should bias their offspring sex ratio towards daughters, which avoids the fitness costs of their sons competing with each other. Conversely, when a large number of females contribute offspring to a patch, they are expected to invest equally in sons and daughters. Furthermore, sex ratios of species that regularly experience variable foundress numbers are closer to those predicted by LMC theory than species that encounter less variable foundress number scenarios. Due to their patterns of resource use, female Callosobruchus maculatus are likely to experience a broad range of foundress number scenarios. We carried out three experiments to test whether female C. maculatus adjust their sex ratios in response to foundress number and two other indicators of LMC: ovipositing on pre-parasitised patches and ovipositing with sisters. We did not find any evidence of the predicted sex ratio adjustment, but we did find evidence of kin biased behaviour.     

Dispersal and mating structure of a parasitoid with a female-biased sex ratio: Implications for theory

Summary Species of parasitic Hymenoptera that manifest female-biased sex ratios and whose offspring mate only with the offspring of the natal patch are assumed to have evolved biased sex ratios because of Local Mate Competition (LMC). Off-patch matings, i.e. outcrossing, are inconsistent with the conditions favouring biased sex ratios because they foster a mating structure approaching panmixia. Such a mating structure favours parents who invest equally in daughters and sons, assuming the production of each sex is of equal cost.Pachycrepoideus vindemiae (Rondani) is a solitary pupal parasitoid of patchily distributed frugivorousDrosophila, whose offspring manifest a female-biased sex ratio. Thus this species appears to manifest a population structure and progeny sex ratio consistent with LMC. However, preliminary observations and subsequent greenhouse experiments suggest that the males participate in off-patch matings and that this propensity is unlikely to be an experimental artefact. FemaleP. vindemiae dispersed from patches in which either the males were lacking (12% of the emigrant females), both resident (sibling) and immigrant males were present (23% of the females), only immigrant males were present (14% of the females), or their opportunity to mate could not be determined (14% of the females). Of the 12% that emigrated from a patch lacking males, an estimated 7% mated at an oviposition site and 5% remained unmated, presumably because they arrived at an oviposition site that lacked males before they were dissected to determine whether they were inseminated. Thus the degree of bias in the sex ratios of the progeny (18% males), coupled with the suggested outcrossing potential from the experiments (26–37%), is inconsistent with the assumptions of LMC or variants of it, i.e. asynchronous brood maturation. Thus the explanation for a biased sex ratio in the offspring ofP. vindemiae remains a conundrum. More importantly,P. vindemiae does not appear to be an isolated example.     

Mating system and sex allocation in the gregarious parasitoid Cotesia glomerata

The gregarious parasitoid Cotesia glomerata (L.) is often presumed to possess the characteristic attributes of a species that manifests local mate competition (LMC), as it commonly produces female-biased broods. However, our field surveys of sex ratio and laboratory observations of adult behaviour showed that this species is subject to partial local mate competition caused by natal dispersal. On average, 30% of males left their natal patch before mating, with the proportion of dispersing males increasing with an increase in the patch's sex ratio (i.e. proportion of males). Over 50% of females left their natal patch before mating, and only 27.5% of females mated with males emerging from the same natal patch. Although females showed no preference between males that were and were not their siblings, broods from females that mated with siblings had a significantly higher mean brood sex ratio (0.56) than broods from females that mated with nonsiblings (0.39). Furthermore, brood sex ratios increased as inbreeding was intensified over four generations. A field population of this wasp had a mean brood sex ratio of 0.35 over 3 years, which conformed well to the evolutionarily stable strategy sex ratio (r=0.34) predicted by Taylor's partial sibmating model for haplodiploid species. These results suggest that the sex allocation strategy of C. glomerata is based on both partial local mate competition in males and inbreeding avoidance in females. In turn, this mating system plays a role in the evolution of natal dispersal behaviour in this species.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Factors influencing brood sex ratios in polyembryonic Hymenoptera

Copidosoma sp. is a polyembryonic encyrtid wasp which parasitizes isolated hosts. Most broods of this wasp are unisexual, but some contain both sexes and the secondary sex ratio of these is usually highly female biased. The overall population secondary sex ratio is female biased. Walter and Clarke (1992) argue that because the majority of individuals must mate outside the natal patch, the bias in the population secondary sex ratio contradicts predictions made by Hamilton's (1967) theory of local mate competition (LMC). We suggest that the primary sex ratio is unbiased and that Walter and Clarke's results do not cast doubt on LMC. Instead these results imply that ovipositing females make a combined clutch size and sex ratio decision influencing whether individuals developing from a particular brood will outbreed or largely inbreed; for each case the predictions of LMC theory are not violated. If this interpretation is correct, what is of interest is the basis on which this decision is made rather than the population secondary sex ratio. We show that host encounter rate influences the proportions of mixed and single sex broods laid by Copidosoma floridanum, a related polyembryonic parasitoid. Among single-sex broods the primary sex ratio is female biased, but our results are in agreement with LMC theory since offspring developing from these broods will probably mate with siblings from adjacent hosts. We consider the egg load of females to be of major influence on oviposition behaviour, and that the mating structure of parasitoid offspring, potentially differential costs of male and female broods and the natural distributions of hosts both at oviposition and eclosion, require further study.     

对叶榕传粉小蜂性比率的调节和稳定

传粉榕小蜂呈现偏雌的性比率,单双倍体性别决定系统、局域配偶竞争和近交效应被认为是调节偏雌性比率的3个主要机制。通过研究影响对叶榕传粉小蜂性比率的因素,结果表明传粉榕小蜂的偏雌性比率随局域配偶竞争强度的降低而增加;受母代雌蜂交配次数的影响,随着母代雌蜂交配次数的增加,子代的偏雌性比率逐渐降低,这一结果首次揭示了传粉榕小蜂的交配制次数对性比率的影响,并在个体水平上定量了性比率变异与雌蜂交配频次的关系。传粉小蜂的性比率与共生的非传粉小蜂的关系,非传粉小蜂的介入直接减少了传粉小蜂的数量,甚至对传粉小蜂的种群有显著影响,结果发现非传粉小蜂对传粉小蜂雌雄性的分配比率没有显著影响,传粉榕小蜂仍能正常地进行繁殖。传粉与非传粉者小蜂之间作用关系的确定,可为进一步理解两者的稳定共生的机制提供科学证据。     

Alternative mating tactics and extreme male dimorphism in fig wasps

The dimorphisms in morphology and behaviour of male fig wasps are among the most extreme in the animal kingdom, and offer excellent opportunities to test the predictions of certain sexual selection models. Winged males resemble their conspecific females closely, but wingless males are so divergent in form that they have repeatedly been classified into different taxa. Wingless males mate within their natal fig fruits, whereas winged males disperse to mate. Individual species may have winged males, wingless males or both morphs. A key hypothesis proposes that sexual selection on male mating opportunities favours winged males in species with small broods and wingless males in species with large broods. Using data from 114 species in 33 genera, we show that both simple and formal comparative analyses support the correlated evolution of large brood size and male winglessness. Theoretical models further predict that, in male dimorphic species, the proportion of winged males should equal (in cases without local mate competition) or exceed (in cases with local mate competition) the proportion of females developing in fig fruits without wingless males. These predictions are met by eight out of nine male dimorphic species studied. Taken together, the patterns across all species, and between different male dimorphic species, strongly support sexual selection on mating opportunities as the major determinant of male morph ratios in fig wasps.     

Competition and opportunity shape the reproductive tactics of males in the ant Cardiocondyla obscurior

Context-dependent adjustment of mating tactics can drastically increase the mating success of behaviourally flexible animals. We used the ant Cardiocondyla obscurior as a model system to study adaptive adjustment of male mating tactics. This species shows a male diphenism of wingless fighter males and peaceful winged males. Whereas the wingless males stay and exclusively mate in the maternal colony, the mating behaviour of winged males is plastic. They copulate with female sexuals in their natal nests early in life but later disperse in search for sexuals outside. In this study, we observed the nest-leaving behaviour of winged males under different conditions and found that they adaptively adjust the timing of their dispersal to the availability of mating partners, as well as the presence, and even the type of competitors in their natal nests. In colonies with virgin female queens winged males stayed longest when they were the only male in the nest. They left earlier when mating partners were not available or when other males were present. In the presence of wingless, locally mating fighter males, winged males dispersed earlier than in the presence of docile, winged competitors. This suggests that C. obscurior males are capable of estimating their local breeding chances and adaptively adjust their dispersal behaviour in both an opportunistic and a risk-sensitive way, thus showing hitherto unknown behavioural plasticity in social insect males.     

Contest versus scramble competition for mates: The composition and spatial structure of a population of gray mouse lemurs (Microcebus murinus) in North-west Madagascar

The modes of intrasexual competition interacting in many dispersed societies of nocturnal solitary foragers are still poorly understood. In this study we investigate the spatial structure within a free-living population of gray mouse lemurs (Microcebus murinus) in order to test for the first time the predictions from two contrasting models of male intrasexual competition on the population level. The contest competition model predicts an uneven distribution of the sexes in a population nucleus with a female biased sex ratio in the center and a male biased sex ratio in the periphery. In contrast the scramble competition model predicts males and females being distributed evenly throughout their habitat with a constant sex ratio. Nine capture/recapture periods within three consecutive mating seasons revealed a continuous male biased sex ratio in the adult population with even trapping rates for the sexes. The male biased sex ratio could either be explained with postnatal female biased mortality or with a male biased natal sex ratio. This male biased sex ratio was apparent in all parts of the study site, indicating that the population was not subdivided into a female biased core and a male biased periphery. Furthermore, the majority of adult males have been captured at the same site as or in vicinity to females. Consequently, a large proportion of males had spatial access to females during the mating season. No signs of monopolization of females by certain dominant males could be detected. These data support the predictions from the scramble competition model and the concept of a promiscuous mating system for this species.     

Sex ratios under asymmetrical local mate competition: theory and a test with parasitoid wasps

Sex ratio theory allows unparalleled opportunities for testing how well animal behavior can be predicted by evolutionary theory. For example, Hamilton's theory of local mate competition (LMC) is well understood and can explain variation in sex allocation across numerous species. This allows more specific predictions to be developed and tested. Here we extend LMC theory to a situation that will be common in a range of species: asymmetrical LMC. Asymmetrical LMC occurs when females lay eggs on a patch asynchronously and male offspring do not disperse, leading to relatively weaker LMC for males emerging from later broods. Varying levels of LMC then lead to varying optimal sex ratios for females, depending on when and where they oviposit. We confirm the assumptions of our theory using the wasp Nasonia vitripennis and then test our predictions. We show that females adjust their offspring sex ratios in the directions predicted, laying different sex ratios on different hosts within a patch. Specifically, there was a less female-biased sex ratio when ovipositing on an unparasitized host if another host on the patch had previously been parasitized and a less female-biased sex ratio on parasitized hosts if females also oviposited on an unparasitized host.     

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.     

钝叶榕(Ficus curtipes)非传粉小蜂交配行为

榕树及其传粉榕小蜂繁殖上相互依存,被认为是生物界中协同进化时间最悠久,相互关系最密切的一对生物;在大多数榕树种类的隐头花序内,除了传粉榕小蜂外,还共存着多种非传粉小蜂,它们的繁殖行为直接影响着榕树和传粉榕小蜂的繁殖和互惠稳定。钝叶榕(Ficus curtipes Corner),是一种雌雄同株的绞杀性榕树。研究在西双版纳热带植物园里共收集钝叶榕100个隐头果内的榕小蜂,获得9493号标本;其中,包括1种传粉小蜂和5种非传粉小蜂,钝叶榕传粉小蜂Eupristina sp.占总数的4466%,杨氏榕树金小蜂Diaziella yangi 占46.13%,而其它4种非传粉小蜂（Lipothymus sp., Sycobia sp., Philotrypesis sp.和Sycoscopter sp.）仅占9.20%。前3种榕小蜂雌蜂进到果腔产卵,其余3种在果外产卵。观测23个钝叶榕榕果出蜂情况发现,6种榕小蜂在钝叶榕隐头花序内遵循严格的羽化出蜂顺序,首先是Sycobia sp.,次之是Lipothymus sp.,再次之是杨氏榕树金小蜂,最后是钝叶榕传粉小蜂、Philotrypesis sp.和Sycoscopter sp.。5种非传粉小蜂的交配场所与雄蜂翅型无关,雄蜂有翅型的杨氏榕树金小蜂大部分交配在果内完成,而且它们的雄蜂为争夺交配机会存在激烈的打斗行为;雄蜂无翅型的Lipothymus sp.有部分雄蜂爬出隐头果,在果壁和附近的叶片背面交配;雄蜂有翅型的Sycobia sp.,其所有交配都发生在果外;Philotrypesis sp.和Sycoscopter sp. 雄蜂均无翅,它们的交配全发生在果内。局域配偶竞争使榕小蜂性比偏雌,杨氏榕树金小蜂雄蜂虽然有翅,但大部分交配发生在榕果内,这将影响其最佳的性比率。因此,依赖雄蜂翅型并不能很好地预测榕小蜂的交配场所和性比率。     

Reproductive strategies of two forms of flightless males in a non‐pollinating fig wasp under partial local mate competition

1. The underlying basis of alternative male reproductive strategies is either genetic or environmental. Several non‐pollinating fig wasp lineages have dimorphic males, typically with winged males that disperse from natal figs to mate and flightless males that seek mating opportunities in natal figs. 2. Walkerella sp. from Ficus benjamina has dark and pale wingless males. Observations and experiments in Xishuangbanna, Southern China found that (i) the sex ratio of Walkerella sp. did not vary with foundress number or brood size. (ii) The frequency of dark males increased with brood size and foundress number and they were absent from figs with a single foundress. This produced a higher proportion of dark males at higher densities. (iii) Males of both morphs fought, but injuries to dark males were more frequent. (iv) Dark males were more likely to disperse away from their natal figs and (v) they were more resistant to dehydration. 3. Responses to selection are constrained by the genetic options available. Consequently, selection pressures acting on different lineages can produce similar outcomes that are achieved in different ways. Walkerella species lack winged males, but dark males display some of their features, dispersing from natal figs and displaying appropriate physiological and behavioural adaptations. However, dark males also displayed increased levels of damage from fighting – a feature unlikely to be shared with the winged males of other species.     

Courtship with two spoons—Anatomy and presumed function of the bizarre antennae of Cardiocondyla zoserka ant males

Mating in ants often occurs on the wing during nuptial flights or on the ground when scattered female sexuals attract males by pheromones. In both scenarios, there is little opportunity for males to engage in prolonged aggressive competition or elaborate courtship displays. Male morphology is therefore adapted to locating female sexuals and mating, and it lacks specific weapons or other traits associated with courtship. In contrast, sexuals of the ant genus Cardiocondyla typically mate in their natal nests. As a consequence, in many species winged males have been replaced by wingless fighter or territorial males, which kill or expel rival males with their strong mandibles and show complex mating behavior. However, no wingless males are known from Cardiocondyla zoserka from West Africa, and instead, winged males have evolved a bizarre secondary sexual trait: uniquely shaped antennae with spoon‐like tips that show heavily sculptured ventral surfaces with numerous invaginations. We here report on the courtship behavior of C. zoserka males and describe antennal glands with class 3 gland cells, which presumably secrete a close range sex pheromone. Antennal glands have not yet been found in males of other ant species, including a close relative of C. zoserka, suggesting that in ants with intranidal mating sexual selection can rapidly lead to highly divergent adaptations and the evolution of novel structures.     

Sex allocation of parasitic wasps: local mate competition,dispersal before mating and host quality variation

We constructed a sex allocation model for parasitic wasps to explain the wide variation in their sex ratio, considering the effects of local mate competition, partial dispersal of progeny before mating, and heterogeneity in host quality among patches. We conducted an experiment to compare with the predictions of our model. We considered the following situations. First, the hosts are distributed in discrete patches: a number of female wasps visit and oviposit in each patch. Second, all the progeny do not mate within the natal patch; some of them disperse to take part in population-wide random mating. We calculated ES sex ratios in cases where there are two kinds of patches: good ones and poor ones. We examined the dependency of ES sex ratios on several parameters, i.e., 1) the probability that a daughter mates in her natal patch, 2) the ratio of the female fitness of the good patch to that of the poor patch, 3) the proportion of poor patches, and 4) the number of foundresses in a patch. The result of our experiment showed the same tendency as the calculation in case where the LMC effect is high in each patch. We briefly discuss a possible selection pressure for dispersal of progeny, with special reference to the mating structure of parasitic wasps.     

Adaptive production of fighter males: queens of the ant Cardiocondyla adjust the sex ratio under local mate competition

Hamilton's concept of local mate competition (LMC) is the standard model to explain female-biased sex ratios in solitary Hymenoptera. In social Hymenoptera, however, LMC has remained controversial, mainly because manipulation of sex allocation by workers in response to relatedness asymmetries is an additional powerful mechanism of female bias. Furthermore, the predominant mating systems in the social insects are thought to make LMC unlikely. Nevertheless, several species exist in which dispersal of males is limited and mating occurs in the nest. Some of these species, such as the ant Cardiocondyla obscurior, have evolved dimorphic males, with one morph being specialized for dispersal and the other for fighting with nest-mate males over access to females. Such life history, combining sociality and alternative reproductive tactics in males, provides a unique opportunity to test the power of LMC as a selective force leading to female-biased sex ratios in social Hymenoptera. We show that, in concordance with LMC predictions, an experimental increase in queen number leads to a shift in sex allocation in favour of non-dispersing males, but does not influence the proportion of disperser males. Furthermore, we can assign this change in sex allocation at the colony level to the queens and rule out worker manipulation.     

Observations on sex ratio and behavior of males in Xyleborinus saxesenii Ratzeburg (Scolytinae, Coleoptera)

Strongly female-biased sex ratios are typical for the fungalfeeding haplodiploid Xyleborini (Scolytinae, Coleoptera), and are a result of inbreeding and local mate competition (LMC). These ambrosia beetles are hardly ever found outside of trees, and thus male frequency and behavior have not been addressed in any empirical studies to date. In fact, for most species the males remain undescribed. Data on sex ratios and male behavior could, however, provide important insights into the Xyleborini's mating system and the evolution of inbreeding and LMC in general.In this study, I used in vitro rearing methods to obtain the first observational data on sex ratio, male production, male and female dispersal, and mating behavior in a xyleborine ambrosia beetle. Females of Xyleborinus saxesenii Ratzeburg produced between 0 and 3 sons per brood, and the absence of males was relatively independent of the number of daughters to be fertilized and the maternal brood sex ratio. Both conformed to a strict LMC strategy with a relatively precise and constant number of males. If males were present they eclosed just before the first females dispersed, and stayed in the gallery until all female offspring had matured. They constantly wandered through the gallery system, presumably in search of unfertilized females, and attempted to mate with larvae, other males, and females of all ages. Copulations, however, only occurred with immature females. From galleries with males, nearly all females dispersed fertilized. Only a few left the natal gallery without being fertilized, and subsequently went on to produce large and solely male broods. If broods were male-less, dispersing females always failed to found new galleries.     

Decision-making conditions for intra- or inter-nest mating of winged males in the male-dimorphic ant <Emphasis Type="Italic">Cardiocondyla minutior</Emphasis>

Only winged male and female ants generally mate through nuptial flight during the reproductive season. In the ants of Cardiocondyla, the males show wing dimorphism and their reproductive strategies differ depending on the differences in wing morphology. It has been suggested that wingless “ergatoid” males bearing very similar external morphologies to workers mate within natal nests, whereas winged males bearing typical ant male morphology disperse from their nests to mate. However, some behavioral observations suggest that the winged males of some Cardiocondyla ants such as C. obscurior and C. minutior may mate within natal nests before dispersion. We evaluated the factors affecting the mating behaviors of the winged males of C. minutior under laboratory conditions. We found that (1) the winged males remained and mated with virgin females in natal nests when either virgin winged females or the relatively mature pupae of winged females (i.e., at least 10 days) were present in the nest, (2) the winged males dispersed to adjacent nests with virgin winged females when only mated queens and the relatively young pupae of winged females (i.e., <9 days) were present in the nest, and (3) all winged males were accepted by the workers of non-natal nests irrespective of the distance from the natal nests in the field. Although most ergatoid males were accepted by the workers of close non-natal nests, they were all attacked and killed by the workers of distant non-natal nests. These results suggest that intra-nest mating and the dispersion of the winged males of C. minutior are facultatively determined by the condition of winged females (virginity and relative pupal age) in natal nests. Furthermore, our results suggest that winged males are likely to seek mating partners chemically and to mate with virgin winged females.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

INEXPLICABLY FEMALE‐BIASED SEX RATIOS IN MELITTOBIA WASPS

The sex ratio behavior of parasitoid wasps in the genus Melittobia is scandalous. In contrast to the prediction of Hamilton's local mate competition theory, and the behavior of numerous other species, their extremely female‐biased sex ratios (1–5% males) change little in response to the number of females that lay eggs on a patch. We examined the mating structure and fitness consequences of adjusting the sex ratio in M. australica and found that (1) the rate of inbreeding did not differ from that expected with random mating within each patch; (2) the fitness of females that produced less female‐biased sex ratios (10 or 20% males) was greater than that of females who produced the sex ratio normally observed in M. australica. These results suggest that neither assortative mating nor asymmetrical competition between males can explain the extreme sex ratios. More generally, the finding that the sex ratios produced by females led to a decrease in their fitness suggests that the existing theory fails to capture a key aspect of the natural history of Melittobia, and emphasizes the importance of examining the fitness consequences of different sex ratio strategies, not only whether observed sex ratios correlate with theoretical predictions.