Fitness effects of sex ratio response to host quality and size in the parasitoid wasp Spalangia cameroni

The parasitoid wasp Spalangia cameroni oviposited a greaterproportion of daughters in stable fly pupae than in house flypupae, even when I controlled for stable flies being smallerthan house flies. Sex ratio manipulation in response to hostquality has been modeled as being adaptive through an effectof host quality on the size and hence offspring production ofdaughters. 5. cameronis response to host species may insteadbe adaptive through an effect on larval survivorship, the developmenttime of daughters, and the size of sons. There was greater survivalof daughters than sons on stable flies. Controlling for hostsize, I found that development time of daughters was about 2%less on stable flies than on house flies. The decrease in developmenttime corresponds to a 2% increase in fitness as estimated byr, the intrinsic rate of increase, and is equivalent to abouta 9% increase in offspring production. Sons were about 2% largerfrom house flies than stable flies, which may increase offspringproduction by up to 3%. Host species had no consistent effecton size of daughters or development time of sons. In additionto the response to host species, mothers oviposited a greaterproportion of daughters in larger stable fly hosts. Whetherthis behavior is adaptive is unclear. Although offspring werelarger when they developed on larger stable flies, the rateof increase was less for daughters dian for sons. Effects ofstable fly size on offspring development time were negligible.     

Using sex ratios to estimate what limits reproduction in parasitoids

The reproductive success of a foraging parasitoid may be limited by the number of eggs that she produces and/or the number of hosts that she can locate. Despite the significance for population dynamics and numerous areas of behavioural ecology, the relative importance of these factors remains an issue of contention. Attempts to resolve this controversy have been hindered because estimating the importance of factors limiting reproduction in the field can be extremely laborious and time consuming. We show how sex ratio data can be used as a relatively easy method to indirectly estimate the relative importance of the factors limiting reproduction. Sex ratio data from 48 samples of eight species suggest that: (a) the extent of host or egg limitation in a species varies between site collected and time of year; and (b) on average, species are at an intermediate position on the egg/host limitation continuum, with a bias towards host limitation.     

Sex ratio manipulation in response to host size in the parasitoid wasp Spalangia cameroni: is it adaptive?

Many species of parasitoid wasps produce a greater proportionof sons in small than in large hosts. As described by the host-sizemodel, natural selection is becoming a standard explanationfor the evolution of this phenomenon. We examined a criticalassumption of the host-size model, that host size has a morepositive effect on female than on male reproductive success.In laboratory experiments with the parasitoid wasp Spalangiacameroni, females that developed on larger hosts contained moreeggs at emergence. However, more eggs did not translate intomore offspring, at high or low density and regardless of whethera female had to burrow to reach hosts. The size of host on whicha female developed was also unrelated to her longevity, regardlessof the presence or absence of hosts. The size of host on whicha male developed had no effect on his sperm production or abilityto inseminate females, regardless of whether insemination abilitywas measured by the amount of sperm transferred to a female,by the proportion of a male's mates that produced any daughters,or by the proportion of daughters that a male's mates produced.Thus, despite data on multiple measures of fitness under a rangeof conditions, sex ratio manipulation in response to host sizein S. cameroni does not appear to be adaptive, and another explanationis needed.     

How two different host species influence the performance of a gregarious parasitoid: host size is not equal to host quality

1. Hyssopus pallidus Askew (Hymenoptera, Eulophidae) is a gregarious ectoparasitoid of the two tortricid moths species Cydia molesta Busck and C. pomonella L. (Lepidoptera, Tortricidae). It paralyses and parasitizes different larval instars of both species inside the apple fruit, which leads to the death of the caterpillar. 2. We assessed the influence of host species characteristics and host food on the performance of the parasitoid female in terms of clutch size decisions and fitness of the F(1) generation. 3. A comparison of clutch size revealed that female parasitoids deposited similar numbers of eggs on the comparatively smaller C. molesta hosts as on the larger C. pomonella hosts. The number of parasitoid offspring produced per weight unit of host larva was significantly higher in C. molesta than in C. pomonella, which is contrary to the general prediction that smaller hosts yield less parasitoid offspring. However, the sex ratio was not influenced by host species that differed considerably in size. 4. Despite the fact that less host resources were available per parasitoid larva feeding on C. molesta caterpillars, the mean weight of emerging female wasps was higher in the parasitoids reared on C. molesta. Furthermore, longevity of these female wasps was neither influenced by host species nor by the food their host had consumed. In addition we did not find a positive relationship between adult female weight and longevity. 5. Parasitoid females proved to be able to assess accurately the nutritional quality of an encountered host and adjust clutch size accordingly. These findings indicate that host size is not equal to host quality. Thus host size is not the only parameter to explain the nutritional quality of a given host and to predict fitness gain in the subsequent generation.     

Sex Allocation by a Parasitoid Wasp (Hymenoptera: Ichneumonidae) to Different Host Species: A Question for the Mechanism of Host Size Estimation

Sex allocation by the polyphagous solitary pupal parasitoid wasp Pimpla luctuosa Smith to a small host species, Galleria mellonella (L.), and a large host species, Mamestra brassicae L., was investigated to test whether female wasps responded to hosts of different sizes across different host species. In the experiments, both host species were presented to each test female wasp. Primary and secondary sex ratio experiments revealed that female wasps laid more female eggs in larger pupae of each host species, indicating that female wasps recognized size differences within host species. The wasp sex ratio (male ratio) from M. brassicae, however, was much higher than that expected on the basis of the sex ratio curve from different-sized G. mellonella. Larger hosts of each host species yielded larger wasps, indicating that the host size estimation by female wasps across different host species was incomplete or was not simple. These results suggested that P. luctuosa evaluated host size not only by physical measures such as dimension but also by other unknown measures. A possible explanation for the adaptiveness of different sex ratio responses by Pimpla luctuosa to different host species was discussed.     

Sex allocation in haplodiploids is mediated by egg size: evidence in the spider mite Tetranychus urticae Koch

Haplodiploid species display extraordinary sex ratios. However, a differential investment in male and female offspring might also be achieved by a differential provisioning of eggs, as observed in birds and lizards. We investigated this hypothesis in the haplodiploid spider mite Tetranychus urticae, which displays highly female-biased sex ratios. We show that egg size significantly determines not only larval size, juvenile survival and adult size, but also fertilization probability, as in marine invertebrates with external fertilization, so that female (fertilized) eggs are significantly larger than male (unfertilized) eggs. Moreover, females with on average larger eggs before fertilization produce a more female-biased sex ratio afterwards. Egg size thus mediates sex-specific egg provisioning, sex and offspring sex ratio. Finally, sex-specific egg provisioning has another major consequence: male eggs produced by mated mothers are smaller than male eggs produced by virgins, and this size difference persists in adults. Virgin females might thus have a (male) fitness advantage over mated females.     

Sex ratio, sex-specific chick mortality and sexual size dimorphism in birds

It has been suggested that sexual size dimorphism (SSD) may influence sex ratios at different life stages. Higher energy requirements during growth associated with larger body size could lead to a greater mortality of the larger sex and ultimately to an overproduction of the smaller sex. To explore the associations between SSD and hatching and fledging sex ratio we performed a species-level analysis and a phylogenetically controlled analysis, based on 83 bird species. Overall, there was a significant inverse relationship between the degree of SSD and the proportion of males at hatching and fledging. Sex-specific mortality related to SSD showed a weak but persistent negative tendency, suggesting a mortality bias towards the larger sex. These results suggest that changes in relation to SSD may take place mainly at the conception stage, but could be adjusted during growth. However, conclusions should be treated cautiously as these relationships weaken when additional variables are considered.     

Sex-determining and floating translocation complexes in Viscutn cruciatum Sieber ex Boiss.(Viscaceae) in southern Spain. Some evolutionary and ecological comments

As in other dioecious species of Viscum, males of V.cruciatum always produce large rings of chromosomes during meiosis due to the existence of permanent translocation heterozygosity. Only rings often (and very rarely 12) chromosomes have been found in this species, probably a relative of V.album in the Mediterranean area. Females form bivalentS. Another two floating translocations–not sex-associated and affecting different chromosomes–have been observed in both male and female plants in some populations. These produce meiotic configurations previously unrecorded in Viscum. This floating heterozygosity seems to be correlated with the aptitude of V.cruciatum to develop on different hosts, especially Prunus. All studied populations showed an excess of female plants, perhaps partly due to facultative agamospermy.     

Effects of host age and cocoon position on attack rate by the hyperparasitoid,Eurytoma sp. [Hym.: Eurytomidae], on cocoons of the parasitoid,Cotesia (=Apanteles) glomerata [Hym.: Braconidae]

Host age preference of the hyperparasitoid,Eurytoma sp., attacking the cocoons of the gregarious parasitoid wasp,Cotesia (=Apanteles) glomerata L., was investigated in the field and laboratory. Under laboratory conditions of 20°C and L16D8 photoperiod.Eurytoma sp. parasitized cocoons of all ages, laying 7 to 10 eggs per cluster during a 24 h period. Field-collected cocoons also indicated that the host was parasitized regardless of its developmental stage. However, the mortality ofEurytoma sp. laid in cocoons on the day before host emergence was as high as 60%. Furthermore, progeny sex ratio (proportion males) reached 0.708 in eggs laid in the oldest cocoon clusters, whereas that for younger cocoons was strongly female-biased. Together, these facts suggest that older hosts are less suitable forEurytoma sp. than are younger ones, even though there was no significant decreasing tendency in the number of parasitized cocoons per cluster. In addition, the effect of cocoon position within a cluster was apparent, outer cocoons being more easily parasitized than inner ones. TheEurytoma sp. female oviposited at random on the free surface ofC. glomerata cocoons.