Sex ratio, mating behaviour and Wolbachia infections in a sheetweb spider

The solitary sheetweb spider Pityohyphantes phrygianus has a female-biased primary sex ratio. Earlier studies have indicated that female post-copulatory position may influence the degree of bias, but the adaptive significance of the departure from a Fisherian 1 : 1 sex ratio is not known. In this study we show that there is a significant relationship between abdominal position and offspring sex ratio in undisturbed females. Moreover, female mating behaviour (and thus sex ratio) is correlated with female size, but this relationship may shift from negative to positive depending on the presence of the maternally acquired endosymbiotic bacterium Wolbachia in both sexes. A survey of wild-caught individuals shows that the incidence of Wolbachia infection in natural populations is high. This indicates that Wolbachia may be a significant factor influencing female post-copulatory behaviour and sex ratio in natural populations. We discuss various adaptive reasons for sex-ratio bias and behavioural plasticity in females.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 181–186.     

Body-size effect on egg size in eublepharid geckos (Squamata: Eublepharidae), lizards with invariant clutch size: negative allometry for egg size in ectotherms is not universal

Within a single clutch, smaller species of ectotherms generally lay a smaller number of relatively larger eggs than do larger species. Many hypotheses explaining both the interspecific negative allometry in egg size and egg size–number trade-off postulate the existence of an upper limit to the egg size of larger species. Specifically, in lizards, large eggs of large species could have too long a duration of incubation, or they could be too large to pass through the pelvic opening, which is presumably constrained mechanically in larger species. Alternatively, negative allometry could be a result of limits affecting eggs of smaller species. Under the latter concept, hatchling size in smaller species may be close to the lower limit imposed by ecological interactions or physiological processes, and therefore smaller species have to invest in relatively larger offspring. Contrary to these lower limit hypotheses, explanations based on the existence of an upper limit always predict negative egg-size allometry even in animals with invariant clutch size, in which naturally there is no egg size–number trade-off. We studied egg-size allometry in lizards of the family Eublepharidae, a monophyletic group of primitive geckos with large variance in body size and an invariant number of two eggs per clutch. We found an isometric relationship between egg and female size that does not support the upper limit hypotheses.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 527–532.     

A sex-specific size–number tradeoff in clonal broods

Polyembryonic parasitoids producing single-sex broods of clonal offspring provide an unusually clear window into the classic tradeoff between the number and size of offspring. We conducted a laboratory study of the encyrtid parasitoid Copidosoma bakeri parasitizing the noctuid Agrotis ipsilon to examine the way that size and number of offspring tradeoff in broods of each sex and to determine how the fit between host and parasitoid brood is achieved. We found that brood mass (wasp body mass ×brood size) was proportional to host mass, independent of brood sex, indicating a tight fit between brood and host and ensuring a size–number tradeoff. By correcting brood size and body mass of each brood for host mass, we demonstrated the expected inverse relationship between wasp variables. We postulated that the wasp brood might achieve the fit to the host by (1) adjusting brood size based on information available early in host development before and during division of the embryo, (2) manipulating host size late in host development after completion of embryo division, or (3) simply adjusting individual wasp mass to fill the host. We evaluated host responses to parasitism – and correlations between brood size and host growth early and late in development – for broods of each sex. The data are consistent with adjustment of brood size to the amount of host growth early in host development and with manipulation of host mass late in host development. The tight link between host mass and brood mass also suggests a final adjustment by parasitoid growth to achieve complete filling. Within the tight fit, female broods were smaller but contained larger individuals than male broods. The sex-specific balance point of the tradeoff and sex differences in balancing mechanisms and responses to host size suggest different selection pressures on each sex requiring future investigation.     

Superparasitism and host size effects in Oomyzus sokolowskii, a parasitoid of diamondback moth

Many aspects of a parasitoid's biology may be affected by its host. Host size, for example, could affect parasitoid fitness, especially in gregarious parasitoids, in which the resource is used by multiple siblings. Oomyzus sokolowskii (Kurdjumov) (Hymenoptera: Eulophidae) is a gregarious larval–pupal endoparasitoid of the diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), a major pest of crucifers worldwide, and is able to superparasitize the host. This study focuses on the hypothesis that because resource availability is higher in larger hosts, parasitoids developing in larger hosts will fare better. However, superparasitized hosts are expected to yield larger numbers of parasitoid offspring of smaller body size. Results showed that superparasitism increased the number of parasitoid offspring produced per host and increased offspring longevity, but decreased offspring body size. However, developmental time and sex ratio of parasitoid offspring was similar among hosts parasitized once, twice, or three times. Regardless of superparasitism, parasitoids emerging from larger hosts that were fed honey solution lived longer than similarly fed progeny from smaller hosts (36.4 vs. 22.1 days). The results partially support the hypothesis that Oomyzus gained fitness from an increase in host size; moreover, superparasitism seems advantageous for Oomyzus due to increased offspring numbers and longevity.     

Adaptiveness of sex ratio control by the pupal parasitoid Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size

Adaptiveness of sex ratio control by the solitary parasitoid wasp Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size was studied, by examining whether differential effects of host size on the fitness of resulting wasps are to be found between males and females. The offspring sex ratio (male ratio) decreased with increasing host size. Larger hosts yielded larger wasps. Male larvae were less efficient in consuming larger hosts than female larvae. No significant interaction in development time was found between parasitoid sex and host size. Larger female wasps lived longer than smaller females, while longevity of male wasps did not increase with increasing wasp size. Smaller males were able to mate either with small or with large females, while larger males failed to mate with small females. Larger female wasps had a greater number of ovarioles and mature eggs at any one time than smaller females, although the number of eggs produced per host-feeding was not influenced by female wasps. Thus, the differential effect of host size on the fitness of males and females exists in I. naranyae. The basic assumption of the host-size model was therefore satisfied, demonstrating that sex ratio control by I. naranyae in response to host size is adaptive.     

A field study of size–fitness relationships in the parasitoid Asobara tabida

1. Many evolutionary models of parasitoid behaviour assume a positive correlation between size and fitness. In this paper we study the size–fitness relationship in the laboratory and in the field using females of the solitary parasitoid Asobara tabida (Hymenoptera: Braconidae).
2. In the laboratory, fecundity, fat reserves and longevity without food were positively correlated with size.
3. Release–recapture experiments in the field showed that dispersal diminishes fat reserves. Dispersal ability is size-dependent: larger females, with larger fat reserves, disperse over larger distances than smaller females.
4. The form of the relationship between size and fitness in the field was estimated in two ways: one based on a comparison of the size distribution of released and recaptured females; the other based on the egg load and fat reserves of wild-caught females. Both showed an accelerating increase of fitness with size.
5. The majority of females appeared to be time-limited. Therefore, the increase in fitness with size is predominantly due to a larger dispersal ability and not to a higher egg load.     

Female morphology,web design,and the potential for multiple mating in Nephila clavipes: do fat‐bottomed girls make the spider world go round?

In animal species where females mate with multiple males, female mating success might be expected to covary with aspects of female morphology, such as size or shape. Spiders are especially interesting in this regard, as the females of several spider groups weave intricate webs that often accommodate multiple male spiders, all of whom are potential mates. Because web design is likely to be dependent on female size/shape, we use multivariate methods to assess the relationships among female morphology, web design, and reproductive ecology over a range of body sizes in the orb-weaving spider Nephila clavipes . Of the measured variables, only abdomen size explained a significant amount of the variation in number of males on a web, and this relationship holds even after statistically accounting for body size. Because abdomen size is an indicator of body condition in spiders, we suggest that condition is likely to be an important factor relating to potential mating success in female spiders. We found no evidence for an association between web design and number of males on a web, although our data indicate that larger females build webs that are both larger and further from the ground than those of smaller females.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87 , 95−102.     

Sexual dimorphism and the differential mortality model: is behaviour related to survival?

There are numerous hypotheses to explain the evolution of sexual dimorphism in spiders. One of the most controversial is the differential mortality model (DMM) which proposes that differing rates of (adult) male and female mortality can result in a skewed operational sex ratio and lead to the evolution of small males. This hypothesis has been examined using a comparative approach which assumes that the behaviour of males and females could be used as a surrogate measure of mortality. We tested this assumption using two model species, Hogna helluo and Pardosa milvina (Araneae: Lycosidae) that differ in the degree of sexual dimorphism both in terms of body size and level of activity. Our data demonstrate that differences in male and female behaviour are not predictive of differences in mortality. Rather, as in other organisms, mortality is a complex phenomenon dependent on activity as well as size. These data call into question the methods previously used to test the DMM and suggest that understanding sexual size dimorphism (SSD) in spiders will require evaluation of historical constraints as well as how size currently influences fitness in each sex.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 97−103.     

SEX-RATIO MANIPULATION IN RESPONSE TO HOST SIZE BY THE PARASITOID WASP SPALANGIA CAMERONI: A LABORATORY STUDY

The prediction of Charnov et al.'s (1981) host-size model that there should be a negative relationship between host size and wasp sex ratio (proportion sons) was supported for Spalangia cameroni, a solitary parasitoid wasp. The relationship was shown to be a result of offspring sex manipulation by females in response to host size rather than a result of differential mortality of the sexes. A major assumption of the host-size model is that host size has a greater effect on the ultimate reproductive success of emerging female wasps than of males. This assumption was not supported. Host size had a positive effect on the size of both male and female S. cameroni. However, neither host size nor wasp size affected longevity, production of offspring by females, or ability of males to compete for mates. Host size may differentially affect the reproductive success of female and male wasps through effects on other aspects of reproductive success. Tests of the assumptions of offspring sex-ratio manipulation hypotheses are scarce but critical, not only for parasitoid wasps, but also for other organisms.     

Body size and fitness relation in male and female Diaeretiella rapae

A strong relationship exists between body size and fitness in parasitoids. However, it is unclear whether the relationship is symmetric or asymmetric in males and females. The present study investigated the body size and fitness relationship in Diaeretiella rapae emerged from small and large nymphs of cabbage aphid Brevicoryne brassicae. A positive relationship existed between the size of the aphid host and growth of parasitoid larva developing in it. The fitness gain in males and females was not proportionate to their body size gain. Females mated with larger males produced 10?% more female offspring than females mated with smaller males. However, females that developed in large hosts produced 62?% more offspring (total male and female) than the females emerged from smaller hosts. The findings suggest that the number of offspring and the progeny sex ratio were affected by the body size of both male and female D. rapae.     

Offspring size and survival in the frog Rana latastei: from among-population to within-clutch variation

Egg size is considered to be a major maternal effect for offspring in oviparous organisms. It has profound consequences on fitness, and differences in egg size are viewed as plastic responses to environmental variability. However, it is difficult to identify the effect of egg size per se because egg size can covary with genetic features of the mother and with other nongenetic factors. We analysed the relationship between offspring starting size (i.e. a proxy of egg size) and larval survival in the frog Rana latastei . We analysed this relationship: (1) among five populations at different altitudes; (2) among clutches laid from different females; and (3) among siblings within clutches, to evaluate the effect of starting size. We observed differences among populations for offspring size, but starting size was not related to altitude or genetic diversity. Mortality was higher in populations and families with small average starting size; however, among siblings, the relationship between starting size and mortality was not verified. The relationship observed among clutches may therefore be caused by covariation between egg size and other effects. This suggests that the covariation between egg size and other effects can result in apparent relationships between egg size and fitness-related traits. Proximate and ultimate factors can cause the phenotypic variation of hatchlings in the wild, and key traits can be related to this variation, but the underlying causes require further investigation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 845–853.     

Guppies control offspring size at birth in response to differences in population sex ratio

Females that invest adaptively in their offspring are predicted to channel more resources to the sex that will be at an advantage in the prevailing environmental conditions. Here, we report, for the first time, that female Trinidadian guppy, Poecilia reticulata, respond in reproductively distinct ways when faced with differences in operational sex ratio. We show that females assigned to a female‐biased sex ratio produce larger male offspring than females in an environment in which males predominate. Given the link between size at birth and fitness, and the marked reproductive skew in this species, larger male offspring are expected to have reproductive advantages in guppy populations with an excess of females. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 414–419.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

Gestation temperature affects sexual phenotype, morphology, locomotor performance, and growth of neonatal brown forest skinks, Sphenomorphus indicus

We maintained pregnant Sphenomorphus indicus under four thermal conditions for the whole gestation period to assess the effects of gestation temperature on offspring phenotypes. Parturition occurred between late June and early August, with females at high body temperatures giving birth earlier than those maintained at low body temperatures. Litter size, litter mass, and postpartum body mass did not differ among treatments, and females with relatively higher fecundity produced smaller offspring. Females gave birth to predominantly female offspring (85.7% of the 14 sexed offspring were females) at 24 °C and to predominantly male offspring (76.5% of the 17 sexed offspring were males) at 28 °C. Females with the opportunity to regulate body temperature produced a mix of sexes that did not differ from equality. Offspring produced in different treatments differed in head size, hind-limb length, and tympanum length, but not in snout-vent length, tail length, body mass, fore-limb length, and eye length. Offspring produced at 28 °C were not only smaller in head size, but also shorter in hind-limb length and tympanum length than those offspring produced at lower temperatures. Offspring produced at 28 °C performed more poorly in the racetrack and grew more slowly than offspring produced in the other three treatments. Taken together, our results show that S. indicus might be a temperature-dependent sex determination species and that offspring phenotypes are impaired at high gestation temperatures but maximized at relatively low gestation temperatures.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 453–463.     

Primary sex-ratio and differential progeny survivorship in solitary haplo-diploid parasitoids

ABSTRACT.

• 1 A method of partitioning emergence sex-ratio to give estimates of primary sex-ratio and male and female offspring survival from oviposition to emergence in solitary haplo-diploid parasitoids is presented.
• 2 This method is applied to sex-ratio data from a larval parasitoid, Aphidius ervi Haliday, and a pupal parasitoid, Coccygomimus turionella L., parasitizing a range of host types.
• 3 There was no evidence of facultative control of primary sex-ratio in either species. Parasitoid emergence sex-ratios were similar for all host sizes attacked by C. turionella. In contrast, emergence sex-ratios of A.ervi showed a significant male bias in smaller hosts. This shift in emergence sex-ratio was attributable to differential progeny survival.
• 4 Pre-emergence mortality in both species was a function of host size, with few offspring surviving from small hosts. This suggests that host size may be an important component in the dynamics of host-parasitoid interactions.
• 5 The evolution of sex-ratio regulatory mechanisms in solitary haplo-diploid parasitoids is discussed in the context of parasitoid life-history. We suggest that there is a constraint to the evolution of the facultative control of primary sex-ratio in parasitoids attacking larval stages as a result of the uncertainty of future host resource acquisition rates.

     

Employing evolutionary theory to improve biological pest control: Causes of non-adaptive sex allocation behavior in parasitoid wasps and implications

Models based on sex allocation theory predict that when the fitness gains from larger size differ between male and female offspring, mothers should produce the sex that will offer the greatest investment return. Behavioral studies on parasitoid wasps have confirmed predictions of models, which additionally have practical implications because of their relevance in biological control. We investigated how a parasitoid attacking a scale insect matches theoretical model predictions in a 2-year field study. As predicted by Charnov's host quality model, mothers laid female eggs in hosts above a threshold size. This threshold was absolute, i.e. independent of the host size distribution, independently of the sampling site and date. Further laboratory assays confirmed field results for at least one parasitoid generation and, moreover, excluded the possibility that the observed behavior was a consequence of immature mortality. By comparing the characteristics of our system with others, we hypothesize that this short-term absolute threshold might be favored in polyphagous parasitoids that attack multivoltine hosts. We propose three measures to mitigate the negative implications of this sex allocation behavior in classical and augmentative biological control programs.     

寄主大小及寄生顺序对蝇蛹佣小蜂寄生策略的影响

寄主大小模型认为寄生蜂后代性比与寄主大小相关,寄生蜂倾向于在大寄主上产出更多雌性后代,在小寄主上产出更多雄性后代.探讨了以家蝇蛹为寄主时,蝇蛹佣小蜂后代产量和性比变化;单次寄生情况下,寄主大小及寄生顺序对寄生蜂后代性比等影响.结果表明,蝇蛹佣小蜂的产卵期为(8.93±3.34)d,单头雌蜂能产雌性后代(34.11±16.34)头和雄性后代(11.04±8.87)头,且雄性百分比为0.24±0.11.随成蜂日龄的增大,寄生蜂产生雄性后代的比率显著增加.蝇蛹佣小蜂在寄生家蝇蛹时,会优先选择寄生个体较大的蛹;在单次寄生的情况下,蝇蛹佣小蜂倾向于在较大的家蝇蛹内产出更多的雌性后代.     

Sexual size dimorphism and sex ratios in dragonflies (Odonata)

Sexual size dimorphism and biased sex ratios are common in animals. Rensch's rule states that sexual size dimorphism (SSD) would increase with body size in taxa where males are larger than females and decrease with body size in taxa where females are larger. We tested this trend in dragonflies (Odonata) by analysing body size of 21 species and found support for Rensch's rule. The increase in SSD with increasing size among species can be explained by sexual selection favouring large males. We also estimated the slope of the relationship between sex ratio and size ratio in populations of the 21 species. A negative slope would suggest that the larger sex suffers from high mortality in the larval stage, consistent with riskier foraging. The slope of this relationship was negative, but after correcting for phylogentic non-independence with independent contrasts the relationship was no longer statistically significant, perhaps because of phylogenic inertia or low sample size.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 507–513.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Brothers smell similar: variation in the sex pheromone of male European Beewolves Philanthus triangulum F. (Hymenoptera: Crabronidae) and its implications for inbreeding avoidance

Female choice is thought to increase the fitness returns of females. The complementary choice model states that the best mate depends on the particular genotype of a female. Aculeate Hymenoptera represent a special case of complementary female choice because males should be chosen on the basis of their allele at the sex determination locus. The prevalent sex determination mechanism in bees and wasps (single-locus complementary sex determination) requires that, to produce a daughter, diploid offspring are heterozygous at the sex determination locus. Otherwise, infertile diploid males result. Inevitably, the proportion of diploid males increases with the rate of inbreeding. In the European Beewolf, males scent mark territories to attract mates and the composition of the pheromone might provide a basis for female choice. One crucial prerequisite for females to be able to discriminate against brothers and avoid inbreeding is that the male sex pheromone varies with familial affiliation. This hypothesis was tested by analysing the pheromone of male progeny of eight mothers using gas chromatography and mass spectrometry. A significantly higher similarity was found among brothers than among unrelated individuals. Such a genetic component of a male sex pheromone has not yet been described from aculeate Hymenoptera.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 433–442.