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.     

Mating or ovipositing? A crucial decision in the life history of the cabbage aphid parasitoid Diaeretiella rapae (M’Intosh)

1. The reproductive fitness of a parasitoid depends on its mating and ovipositing success. Virgin haplodiploid females can reproduce, but produce only males, and may diminish fitness by producing more male offspring than required. Therefore, females must decide on whether to mate or oviposit first. 2. This study was conducted to assess the mating versus ovipositing decision and its impact on the reproductive fitness of Diaeretiella rapae (Hymenoptera: Aphididae), an endoparasitoid of the cabbage aphid Brevicoryne brassicae (Hemiptera: Aphididae). 3. When newly emerged females were given a choice between mating and ovipositing, about 62% of D. rapae females preferred to mate before ovipositing. Those females who oviposited before mating parasitised only 10% of the available aphids. After mating, females superparasitised their hosts with fertilised eggs, which resulted in a highly female‐biased sex ratio in the offspring. 4. Mating success was very high (91%) in the presence of hosts (cabbage aphid nymphs) compared with that in the absence of aphids. However, mating success was not influenced by the quality (size) of the hosts present in the mating arena, despite a parasitoid preference for larger hosts during oviposition. The time between pairing and mating was also shorter in the presence of host aphids. The mean number of aphids parasitised and the parasitism rate were significantly greater after mating.     

Oviposition decisions in an endoparasitoid under self-superparasitism conditions

Superparasitism is a widespread phenomenon. Having accepted superparasitism, mated female parasitoids must decide on the sex of each egg they subsequently lay into the same host. Theory predicts that this decision is either based on host quality, when more male eggs are laid in hosts that are already parasitized because they are perceived to be of poorer quality; or more eggs are laid of the sex that is most likely to be a strong larval competitor, i.e. generally females.Anastatus disparis is a facultative endoparasitic egg parasitoid. We used ‘artificial’ hosts to explore outcomes of decision making by A. disparis during superparasitism under a manipulated absence of larval competition. When only one egg was laid it was always female. As the number of eggs laid increased, so more of them were male. This supports the theory that oviposition decisions are based on host quality; more male eggs were laid in hosts that were already parasitized and thus of poorer quality.In a second experiment, eggs were exposed to parasitoids for different periods of time. Half the eggs were dissected to determine the number of parasitoid eggs that had been laid. The remaining eggs were incubated and the number and sex of offspring that ultimately emerged, following larval competition, were recorded. Under superparasitism conditions fierce larval competition ensued; only one offspring survived and they were predominantly female.In conclusion, oviposition decisions by female A. disparis accepting self-superparasitism were made based on host quality.     

Parasitoid size as a function of host sex: potential for different sex allocation strategies

Parasitoid females are known to preferentially allocate female eggs to hosts with the higher resource value, usually leading to oviposition of female eggs in larger hosts and male eggs in smaller hosts. For koinobiont parasitoids, if male and female hosts are of equal size at time of oviposition, but differ in size in later developmental stages, the sex of the host could be used to indicate future resource value. Using parasitoids of the braconid genus Asobara, which are larval parasitoids of Drosophila, it is shown that parasitoids emerging from female hosts are larger than those from male hosts. Given this difference in resource value, ovipositing females should preferentially allocate female eggs to female hosts. An alternative strategy would be to decrease the difference in resource value between male and female hosts by castrating male hosts. The primary sex ratio of A. tabida in their two main host species does not differ between male and female hosts. In contrast to A. tabida, A. citri is known to partially castrate male hosts, but this does not decrease the size difference between male and female hosts. As in A. tabida, there is no difference in sex allocation to male and female hosts in A. citri. Despite the clear difference between the resource value of male and female hosts, these parasitoid species do not seem to make optimal use of this difference. They may not be able to discriminate between host sexes or, alternatively, there is a presently unknown fitness disadvantage to ovipositing in female hosts.     

Host selection by virgin and inseminated females of the parasitic wasp, Dinarmus basalis (Pteromalidae, Hymenoptera)

1. Fitness is related to reproduction and survival. There apparently exists a negative correlation between the numbers of male and female offspring. There also exists a trade-off between survival and reproduction. This paper investigates optimal decisions with the reproduction and survival trade-off in host selection by wasps.
2. Whereas inseminated female wasps could manipulate the sex of their offspring, virgin females produced only male offspring. I surveyed behavioural differences and the consequences of oviposition by inseminated and virgin females of a solitary parasitic wasp in host choice situations.
3. Two host types were available at the same time to both inseminated and virgin female wasps: one (a 17-day-old host in one bean) presenting difficulties for the laying of eggs, but more benefits for the offspring and the other (five 12- or 13-day-old hosts in one bean) easier for the female wasp for laying of eggs but less beneficial for the offspring.
4. Inseminated female wasps chose more 17-day-old hosts than 12-day-old hosts, but more 13-day-old hosts than 17-day-old hosts in each pair-wise choice. Virgin females chose the smaller hosts in both situations.
5. Virgin females, having greater longevity than inseminated females, laid larger numbers of eggs than the inseminated females during their lifetime by adopting an energy-saving host choice that had little effect on male offspring fitness.     

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     

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.     

Host acceptance and sex allocation of Nasonia wasps in response to conspecifics and heterospecifics

Species recognition is an important aspect of an organism''s biology. Here, we consider how parasitoid wasps vary their reproductive decisions when their offspring face intra- and interspecific competition for resources and mates. We use host acceptance and sex ratio behaviour to test whether female Nasonia vitripennis and Nasonia longicornis discriminate between conspecifics and heterospecifics when ovipositing. We tested pairs of conspecific or heterospecific females ovipositing either simultaneously or sequentially on a single host, using strains varying in their recent history of sympatry. Both N. vitripennis and N. longicornis rejected parasitized hosts more often than unparasitized hosts, although females were more likely to superparasitize their own species in the sequential treatment. However, sex ratio behaviour did not vary, suggesting similar responses towards conspecifics and heterospecifics. This contrasts with theory predicting that heterospecifics should not influence sex ratios as their offspring do not influence local mate competition, where conspecifics would. These non-adaptive sex ratios reinforce the lack of adaptive kin discrimination in N. vitripennis and suggest a behavioural constraint. Discrimination between closely related species is therefore context dependent in Nasonia. We suggest that isolating mechanisms associated with the speciation process have influenced behaviour to a greater extent than selection on sex ratios.     

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.     

Sex allocation of three solitary ectoparasitic wasp species on bean weevil larvae: Sex ratio change with host quality and local mate competition

Sex ratio manipulation by ovipositing females was surveyed in 3 solitary ectoparastic wasp species,Dinarmus basalis (Pteromalidae),Anisopteromalus calanrae (Pteromalidae), andHeterospilus prosopidis (Braconidae), that parasitize azuki bean weevil (Callosobruchus chinensis (L) (Coleoptera: Buruchidae)) larvae within azuki beans (Vigna angularis). Variables were local mate competition (LMC) and host quality (HQ). We used host age as a measure of host quality (from 9-to 16-day-old hosts), changed the number of ovipositing females to control the level of local mate competition (1 female and 10 females), and examined oviposition patterns of the wasps. The offspring sex ratios (proportion of females) of the 3 wasp species respond qualitatively same to HQ and LMC. The common qualitative tendency among the 3 species is an increase of sex ratios increase with host age. In the process of changing the sex ratio (9–13-day-old) 3 wasp species respond only to HQ. In the hosts that end development in size (14–16-day-old) wasps respond to LMC. The response of sex ratio change to LMC in the old host ageclasses are different among the 3 species. In the situation that there exists LMC (10 females) sex ratios are the same among the 3 wasps. However, the sex ratios in no LMC (single female) are heterogeneous among the 3 wasps.     

The influence of previous host age on current host acceptance in Trichogramma

Trichogramma principium Sug. & Sor. females were sequentially offered two portions of the grain moth (Sitotroga cerealella Oliv.) eggs, either young (1-day old) or old (eggs that had developed 6 days at a temperature of 20 °C). The probability of host acceptance depended not only on current host age, but also on the age of the previously offered host. Particularly, Trichogramma females more often oviposited in old host eggs when previously offered young eggs (35–45% of Trichogramma females laid eggs) compared to females which were sequentially offered two portions of old eggs (15–20% of Trichogramma females laid eggs). In other words, parasitization by Trichogramma was stable even when transferred from young (preferred) to old (usually rejected) eggs. Dissections showed that refusing females had significantly more mature eggs than ovipositing females, independent of host age. Among ovipositing females, wasps provided with young hosts had fewer mature ovarial eggs than wasps provided with old hosts. Supposedly, Trichogramma females offered old hosts require a higher motivation to oviposit and have a correspondingly higher egg load than females offered young (preferred) hosts.     

Egg allocation of the autoparasitoid encarsia tricolor at different relative densities of the primary host (Trialeurodes vaporariorum) and two secondary hosts (Encarsia formosa and E. tricolor)

Parasitism, offspring sex ratio and superparasitism of the facultative autoparasitoid Encarsia tricolor Foërster (Hymenoptera: Aphelinidae) when given access to arenae with different proportions of the primary host (Trialeurodes vaporariorum (Westwood)) and two species of secondary hosts (E. tricolor and Encarsia formosa Gahan) were studied.Parasitism and offspring sex ratio were not affected by female age in the range 3–10 days old. When the secondary hosts were young E. tricolor pupae, eggs were mostly laid on primary hosts, so the offspring sex ratio was more female-biased than expected, and secondary hosts were not superparasitized at all. When the secondary hosts were fully grown E. formosa larvae, superparasitism was small and offspring sex ratio was more male-biased than expected. E. tricolor females were able to discriminate between hosts previously parasitized by themselves and non-parasitized hosts.     

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.     

Host parasitoid interactions affecting reproduction and oviposition byBrachymeria intermedia [Hym.: Chalcididae]

A maximum number of 6 parasitoid progeny are produced byBrachymeria intermedia (Nees) females, in a 24 h period, even when more than 6 hosts are available. The number of progeny produced by ovipositing females kept at a high temperature or high light intensity is greater than that of females kept at low temperatures or low light intensity. Host size and age of female parasitoids affect the sex ratio or progeny. Parasitism of small pupal hosts results in the production of a greater proportion of male progeny. Increasing maternal age appears to increase the proportion of males.     

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.     

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.     

Effect of host Bactrocera dorsalis sex on yield and quality of the parasitoid Fopius arisanus

This study examines the effect of host Bactrocera dorsalis Hendel (Diptera: Tephritidae) sex on performance of the parasitoid Fopius arisanus Sonan (Hymenoptera: Braconidae) to enable use of a genetic sexing strain (GSS) to transfer this parasitoid to regions where B. dorsalis is not established with lower risk of accidental introduction of the host. Sex ratio and yield of F. arisanus did not significantly differ between the two sexes of a GSS, or between GSS and a standard colony strain. F1 parasitoids of individuals that emerged from the standard colony strain, male GSS hosts and female GSS hosts had similar hatching and parasitization rates on colony eggs. Survivorship of F1 parasitoids with parents from female hosts was significantly greater than for those from male hosts. Using GSS to improve the safety of parasitoid shipments is suggested to be viable, but there may be fitness effects of host sex on F1 parasitoid fitness.     

Factors affecting the parasitism rate and the number and sex ratio of offspring of Asecodes hispinarum Bouček (Hymenoptera: Eulophidae), a biological control agent of Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae)

To optimise the production of Asecodes hispinarum Bou?ek (Hymenoptera: Eulophidae), a parasitoid of coconut leaf beetle, Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae), some of the factors affecting rates of parasitism, number of offspring produced per host and sex ratio of A. hispinarum were investigated. The numbers and sex ratio of A. hispinarum offspring per host reduced significantly at extreme low humidity (30% relative humidity [RH]), but there was no significant effect on parasitism. Photoperiod had no significant effects on any of the life traits tested. A. hispinarum was able to reproduce via arrhenotoky, and while increasing the proportion of female parents increased the number of parasitoids produced, the proportion of female offspring decreased. Older females showed a lower rate of parasitism than young females, however, maternal age did not affect the number or the sex ratio of offspring. Increasing the number of hosts offered to a pair of parasitoids significantly increased the number of parasitised hosts but decreased the parasitism rate while the sex ratio of progeny was not affected. Present work showed that to maximise the production of female parasitoids, a parasitoid/host ratio of 1:1, using one-day old A. hispinarum at a female/male ratio of 3:1 and RH of at least 55% is recommended.     

Effects of Host Size and Parasite Burden on Sex Ratio in the Mosquito Parasite Octomyomermis muspratti

The ratio of Octomyomermis muspratti to the host mosquito at the time of exposure had little effect on the ratio of male to female parasites that resulted. However, the ratio of males to females increased as the number of parasites/host increased. Hosts with a single nematode produced fewer than 1% males in comparison with hosts with 8 parasites which produced about 40% males; hosts with 10 or more nematodes generally produced more male than female nematodes. Males of O. muspratti usually emerged before females because of the earlier death of multiply-infected mosquitoes. The size of the host at the time of invasion bad no significant influence on nematode sex ratios. Since mating is apparently necessary for reproduction in O. muspratti, the low male to female ratios that occur will be important in developing successful mass production techniques.     

A sex ratio theory of gregarious parasitoids

Summary A mathematical model is constructed to explain a density-dependent increase in the progeny sex ratios of gregarious parasitoids. In the model we considered non-cooperative game between females concerned with their own inclusive fitness. Equilibrium progeny sex ratios of the first and second females ovipositing on the same host are expressed in terms of the probability of double parasitism (p), the ratio of a male to a female in contribution to resource competition (α), the clutch size ratio between the two females (β), the crowding effect on female reproductive success (γ), and the inbreeding coefficient (f). Major predictions from the model are: 1) the progeny sex ratios of both the first and second females increase withp, 2) as β becomes smaller, the progeny sex ratios of the first females decrease, while those of the second females dramatically increases, 3) when a host is attacked by at most two wasps, the sex ratio of the total number of eggs laid on the host does not exceed 0.25. The effects of α and preferential death by female progeny in doubly parasitized hosts are considered as factors responsible for an excess number of males at emergence. Some possible modes of density-dependent increase in the sex ratios of the overall progeny populations is also discussed on the basis of the present model.