The developmental strategies and related profitability of an idiobiont ectoparasitoid Sclerodermus pupariae vary with host size

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Co‐foundress confinement elicits kinship effects in a naturally sub‐social parasitoid

Kinship among interacting individuals is often associated with sociality and also with sex ratio effects. Parasitoids in the bethylid genus Goniozus are sub‐social, with single foundress females exhibiting post‐ovipositional maternal care via short‐term aggressive host and brood defence against conspecific females. Due to local mate competition (LMC) and broods normally being produced by a single foundress, sex ratios are female‐biased. Contests between adult females are, however, not normally fatal, and aggression is reduced when competing females are kin, raising the possibility of multi‐foundress reproduction on some hosts. Here, we screen for further life‐history effects of kinship by varying the numbers and relatedness of foundresses confined together with a host resource and also by varying the size of host. We confined groups of 1–8 Goniozus nephantidis females together with a host for 5+ days. Multi‐foundress groups were either all siblings or all nonsiblings. Our chief expectations included that competition for resources would be more intense among larger foundress groups but diminished by both larger host size and closer foundress relatedness, affecting both foundress mortality and reproductive output. From classical LMC theory, we expected that offspring group sex ratios would be less female‐biased when there were more foundresses, and from extended LMC theory, we expected that sex ratios would be more female‐biased when foundresses were close kin. We found that confinement led to the death of some females (11% overall) but only when host resources were most limiting. Mortality of foundresses was less common when foundresses were siblings. Developmental mortality among offspring was considerably higher in multi‐foundress clutches but was unaffected by foundress relatedness. Groups of sibling foundresses collectively produced similar numbers of offspring to nonsibling groups. There was little advantage for individual females to reproduce in multi‐foundress groups: single foundresses suppressed even the largest hosts presented and had the highest per capita production of adult offspring. Despite single foundress reproduction being the norm, G. nephantidis females in multi‐foundress groups appear to attune sex allocation according to both foundress number and foundress relatedness: broods produced by sibling foundresses had sex ratios similar to broods produced by single foundresses (ca. 11% males), whereas the sex ratios of broods produced by nonsibling females were approximately 20% higher and broadly increased with foundress number. We conclude that relatedness and host size may combine to reduce selection against communal reproduction on hosts and that, unlike other studied parasitoids, G. nephantidis sex ratios conform to predictions of both classical and extended LMC theories.     

Putting your sons in the right place: the spatial distribution of fig wasp offspring inside figs

Abstract. 1. Pollinating fig wasps (Hymenoptera, Agaonidae) display sex ratio adjustment, producing less female‐biased combined sex ratios as the number of ovipositing females (foundresses) inside a fig increases. Because males have low mobility, the oviposition sites (galled ovules) chosen by each foundress are likely to have consequences for the mating structure of wasp populations within the figs. 2. In this study, the spatial location of male and female progeny of the pollinating fig wasp Liporrhopalum tentacularis developing within figs of its host plant Ficus montana was examined to investigate two questions: (i) are male and/or female wasp offspring clustered together or interspersed? and (ii) is their distribution affected by whether one or two foundresses are present? Microsatellite markers were used to identify the progeny of different foundresses in dual‐foundress figs. 3. More offspring developed in the central part of the figs, compared with the ostiolar and basal parts, irrespective of foundress number. Neither male nor female wasp offspring were clustered within a fig. 4. The sons of the second foundress to enter a fig were positioned at similar minimum distances to both sibling and non‐sibling females, whereas the sons of the first foundress were closer to their sibling females than to non‐sibling females. If male wasps mate predominantly with females in adjacent galls, then the positioning of sons by the second foundresses is beneficial for them both in terms of reduced sibling mating and because they are provided with ready access to the female progeny of the first foundress.     

Development of the gregarious ectoparasitoid Nasonia vitripennis using five species of necrophagous flies as hosts and at various developmental temperatures

The utility of five species of necrophagous flies (Diptera) as pupal hosts for Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) was examined by comparing incidences of parasitism, fecundity, and several features of wasp development at three rearing temperatures. Species differences in host suitability were evident in all life history features examined, with the highest incidences of parasitism, largest clutches and adult body sizes, and shortest periods of development occurring when the sarcophagid Sarcophaga bullata Parker served as hosts, regardless of temperature in which the wasps developed. Puparia of the calliphorids Lucilia illustris Meigen, Phormia regina Meigen, and Protophormia terraenovae Robineau‐Desvoidy were also accepted as hosts by the female parasitoids, albeit not equally so, and each yielded large, female‐biased broods. By contrast, pupae of the calliphorid Chrysomya rufifacies (Macquart) were not well suited to serve as an oviposition site or support the development of N. vitripennis. When successful parasitism did occur on any host species, duration of parasitoid development increased, adult body sizes were truncated, male‐biased sex ratios were produced, and mortality from egg hatch to adult emergence elevated with increasing rearing temperature. Unlike with the four other fly species, C. rufifacies did not yield any adult parasitoids when the rearing temperature was 35 °C. The results argue that developmental data determined for this wasp derived from a single host species is not sufficient for applying to all scenarios in which wasp development is necessary to estimate a postmortem interval or periods of insect activity.     

Performance of Sclerodermus brevicornis,a parasitoid of invasive longhorn beetles,when reared on rice moth larvae

Biological control efficiency can be improved by developing effective mass‐rearing systems to produce large numbers of high‐quality parasitoids. This study explored an alternative host for rearing Sclerodermus brevicornis (Kieffer) (Hymenoptera: Bethylidae), a potential biocontrol agent for the suppression of exotic and invasive wood‐boring longhorn beetle (Coleoptera: Cerambycidae) populations in the European agroforestry ecosystems. We tested larvae of the rice moth, Corcyra cephalonica Stainton (Lepidoptera: Pyralidae), as host for the parasitoid. We quantified the probability and timing of host attack and parasitism as well as reproductive success, offspring production, and the characteristics of adult offspring. As S. brevicornis is a quasi‐social species (multiple females, communally produced offspring broods), we also explored the effects of varying the number of females to which individual hosts were presented, with the aim of determining the optimal female‐to‐host ratio. As time to host attack can be a limiting factor in S. brevicornis rearing protocols, we tested the use of adult females of another bethylid species, Goniozus legneri Gordh, to paralyse C. cephalonica larvae prior to presentation. We identified the conditions within our experiment that maximized offspring production per host and offspring production per adult female parasitoid. We found that C. cephalonica is suitable as a factitious host and, as it is considerably more straightforward for laboratory rearing than cerambycid species, it is a good candidate for adoption by future S. brevicornis mass‐rearing and release programmes.     

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.     

Spatial distribution patterns of three fig wasps on Ficus semicordata: How non-pollinators affect pollinator’s sex ratio

Sex ratio theory is one of the most productive fields in research on evolutionary biology. Pollinating fig wasps, due to their particular natural life history, are considered to be a valuable model for the study of sex ratio evolution. A great deal of research concerning the factors that affect pollinator fig wasp (Agaonidae) progeny sex ratio has been done, and at present three main factors (haplodiploidy, local mate competition and inbreeding) are found to be important at the population level. However, there still exists variation between empirical data and model predictions. Another factor to which little thought has been given before is the effect of non-pollinating fig wasps (NPFWs) which parasitize in the larvae gall of pollinator thus kill pollinators and exploit the fig/fig pollinator mutualistic systems. In this study, we focus on why and how non-pollinating fig wasps distort pollinator fig wasp’s original sex ratio. Through controlling the number of ovipositing foundresses inside a fig, combined with the observation of ovipositing behavior and sequence, we studied three species of wasp in the figs of a dioecious fig Ficus semicordata including the pollinator Ceratosolen gravely and NPFWs Platyneura cunia, Sycoscapter trifemmensis in tropical area of Xishuangbanna from September to December 2009. First, we observed the timing of oviposition of all fig wasps utilizing F. semicordata and found differences when compared to previous studies. Such as P. cunia is the fourth rather then the secondary fig wasps to oviposit on the syconia approximately 10 days after the pollinator. S. trifemmensis oviposits much earlier than previously thought, 14–32 days after the pollinators. We examined the spatial location of male and female progeny of the pollinator. We found foundresses of pollinator prefer to use innermost ovules first. Only at high offspring numbers were the outer ovules used. More male pollinator offspring were developed near the fig cavity, while female pollinator offspring were more evenly distributed among ovule layers. As pollinator offspring numbers increased, this phenomenon became more pronounced. This pattern of segregation of male larvae gall in inner ovules and female larvae gall in outer ovules suggests that female offspring might be more vulnerable to attack by parasitic wasps that oviposit from outside the syconium. Experiments later demonstrated that NPFWs are restricted by their ovipositor length and they prefer to or can only lay their eggs into ovules near the fig wall. Then we examined the spatial location of NPFWs and compared this with the spatial location of male/female progeny of pollinator. NPFWs had a high probability of parasitizing female pollinator larvae. Thus, NPFWs have a substantial effect on the sex ratio of the pollinator, as parasitism risk decreases towards the center of the syconium, where inner ovules provide enemy-free space for most of male pollinator offspring. Partial correlation analyse shows that sex ratio of pollinator progeny has a positive relationship with the number of NPFWs. We suggest that the resulting gradient in offspring viability between male and female contributes to selection on pollinators’ for a less femalebiased sex ratio. When the affect of NPFWs was excluded, the pollinator sex ratio was not in good agreement with local mate competition theory, although it was still female-biased. In addition, the average number of offspring per foundress decreased with increasing foundress number, but pollinator sex ratio was positively related to brood size. Thus, pollinator females do not appear to adjust their sex ratio to foundress density directly, but use brood size and foundress density simultaneously as cues to assess potential LMC.     

肿腿蜂类寄生蜂室内控害效能评价——以松脊吉丁肿腿蜂为例

为了开发利用天敌资源防治害虫,科学而客观地评价肿腿蜂对蛀干害虫的控害作用,研究了松脊吉丁肿腿蜂(Sclerodermus sp.)对松褐天牛3龄幼虫的功能反应,建立了肿腿蜂类寄生蜂室内控害效能评价体系,并证明了其可行性。该评价体系包括供试虫源标准、寄生蜂生物学与寄生行为学观察、寄生蜂对寄主的功能反应和寻找寄主效应以及寄生蜂的忠岐指数(Y)4个部分。研究结果表明:松脊吉丁肿腿蜂具有较强的控害潜力,单头雌成虫能防治3—4头松褐天牛3龄幼虫,21d内最多可致死9.07头寄主幼虫;寄主密度对单头寄生蜂寄生作用功能反应的测定结果与对寄生作用寻找效应的测定结果以及对产卵量影响的测定结果一致;综合评价该蜂对松褐天牛3龄幼虫控制效能,释放比为1∶1(即1头肿腿蜂雌蜂:1头3龄松褐天牛幼虫)时最高,其忠岐指数Y为39.63,这可能是肿腿蜂类天敌特有的抚幼习性所致,结论也与寄生蜂生物学特性及寄生行为学观察结果相同。研究结论证明该评价体系是稳定可靠的。评价体系在利用天敌昆虫生物防治害虫时为筛选最佳寄生性天敌种类提供了可行的方法和依据。     

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.     

Transfer of a chromosomal <Emphasis Type="Italic">Maverick</Emphasis> to endogenous bracovirus in a parasitoid wasp

Bracoviruses are used by parasitoid wasps to allow development of their progeny within the body of lepidopteran hosts. In parasitoid wasps, the bracovirus exists as a provirus, integrated in a wasp chromosome. Viral replication occurs in wasp ovaries and leads to formation of particles containing dsDNA circles (segments) that are injected into the host body during wasp oviposition. We identified a large DNA transposon Maverick in a parasitoid wasp bracovirus. Closely related elements are present in parasitoid wasp genomes indicating that the element in CcBV corresponds to the insertion of an endogenous wasp Maverick in CcBV provirus. The presence of the Maverick in a bracovirus genome suggests the possibility of transposon transfers from parasitoids to lepidoptera via bracoviruses.     

Transgenic GNA Expressing Potato Plants Augment the Beneficial Biocontrol of Lacanobia Oleracea (Lepidoptera: Noctuidae) by the parasitoid Eulophus Pennicornis (Hymenoptera; Eulophidae)

The effect of expressing the gene encoding snowdrop lectin (Galanthus nivalis agglutinin, GNA) in transgenic potato plants, on parasitism of the phytophagous insect pest Lacanobia oleracea by the gregarious ectoparasitoid Eulophus pennicornis, was investigated in glasshouse trials. Expression of GNA (approx. 1.0% total soluble protein) by transgenic plants significantly reduced the level of pest damage, thus confirming previous studies. Furthermore, the presence of the parasitoid significantly reduced the levels of damage incurred either by the transgenic or control plants when compared to those plants grown in the absence of the parasitoid. For the GNA expressing plants the presence of the parasitoid resulted in further reductions (ca. 21%) in the level of damage caused by the pest species. The ability of the wasp to parasitise and subsequently develop on the pest larvae was not altered by the presence of GNA in the diet of the host. E. pennicornis progeny that developed on L. oleracea reared on GNA expressing plants showed no significant alteration in fecundity when compared with wasps that had developed on hosts fed on control potato plants, although mean size and longevity of female parasitoids was significantly reduced. The number of F ₂ progeny produced by parasitoids derived from hosts fed on GNA-expressing plants was not significantly different to those produced by parasitoids from hosts fed control plants. Results from the present study demonstrate that the use of transgenic plants expressing insecticidal proteins can be compatible with the deployment of beneficial insects and that the two factors may interact in a positive manner.     

A large-scale rearing method for Peristenus digoneutis (Hymenoptera: Braconidae), a biological control agent of Lygus lineolaris (Hemiptera: Miridae)

Releases of Peristenus digoneutis against Lygus spp. in North America have been conducted for many years; however, no published procedures for mass production of the biological control agent were available. A laboratory rearing method was developed using Lygus lineolaris as the host to enhance establishment efforts and provide large numbers of wasps for inundative releases into high value fruit crops. Experiments were conducted to determine optimum host:parasitoid density and rearing temperature. The effects of nymph:wasp ratios and temperature on parasitism and wasp survival showed a 20:1 ratio at 20°C provided high parasitism (256 parasitized nymphs/wasp over lifetime) and excellent wasp survival of 27 days. Experiments on diapause-inducing conditions for P. digoneutis demonstrated that fluctuating temperatures of 23°C (day) and <16°C (night) and corresponding photo phases of 16 h light, for rearing parasitized nymphs, produced 100% diapausing parasitoids whereas non-diapausing parasitoids were only produced at more than 16 h light. Furthermore, parasitized Lygus nymphs need to be transferred to short day conditions no later than 10 days after parasitism to produce diapausing parasitoids. Critical life stages for exposure to conditions inducing diapause, the egg, first and second instar parasitoid larva, occurred from 0 to 10 days at 24°C constant temperature. Increased time in cold storage reduced the number of days to first emergence of parasitoids from diapausing cocoons when transferred to warm temperatures. The optimum storage time for diapausing P. digoneutis is between 25 and 44 weeks, depending upon the length of time that cocoons remain at warm conditions prior to chilling.     

Mass‐rearing optimization of the parasitoid Psyttalia lounsburyi for biological control of the olive fruit fly

The olive fruit fly, Bactrocera oleae (Tephritidae), is a direct pest of olives that has invaded the Mediterranean Region and California. Psyttalia lounsburyi (Braconidae), a larval parasitoid from Africa, has been approved for release in the USA as a classical biological agent. However, it has been difficult to rear the parasitoid in the laboratory because it is multivoltine, and the host develops only in fresh olives, which are not available for most of the year. A method to rear the parasitoid on the factitious host, Mediterranean fruit fly (Ceratitis capitata) was developed, but it was not very efficient for producing large numbers of parasitoids needed for release. We developed a number of ways to improve the efficiency of rearing, including the frequency and duration of exposure for oviposition, optimizing the density of adult parasitoids, host age, as well as methods to quickly standardize the number of larvae exposed and to count emerging adult parasitoids. We significantly improved the number of progeny produced per female and the sex ratio of progeny. Thanks to these improvements, we produced in 2017 over 119,000 adults and shipped over 53,900 for release in California.     

The effect of host developmental stage at parasitism on sex‐related size differentiation in a larval endoparasitoid

• 1 For their larval development, parasitoids depend on the quality and quantity of resources provided by a single host. Therefore, a close relationship is predicted between the size of the host at parasitism and the size of the emerging adult wasp. This relationship is less clear for koinobiont than for idiobiont parasitoids.
• 2 As size differentiation in host species exhibiting sexual size dimorphism (SSD) is likely to occur already during larval development, in koinobiont larval endoparasitoids the size of the emerging adult may also be constrained based on the sex of the host caterpillar.
• 3 Sex‐specific growth trajectories were compared in unparasitised Plutella xylostella caterpillars and in second and fourth instar hosts that were parasitised by the solitary larval koinobiont endoparasitoid Diadegma semiclausum. Both species exhibit SSD, where females are significantly larger than males.
• 4 Healthy female P. xylostella caterpillars developed significantly faster than their male conspecifics. Host regulation induced by D. semiclausum parasitism depended on the instar attacked. Parasitism in second‐instar caterpillars reduced growth compared to healthy unparasitised caterpillars, whereas parasitism in fourth‐instar caterpillars arrested development. The reduction in growth was most pronounced in hosts producing male D. semiclausum.
• 5 Parasitism itself had the largest impact on host growth. SSD in the parasitoid is mainly the result of differences in growth rate of the parasitoid–host complex producing male and female wasps and differences in exploitation of the host resources. Female wasps converted host biomass more efficiently into adult biomass than males.

     

The influence of temperature on size as an indicator of host quality for the development of a solitary koinobiont parasitoid

The influence of aphid size on the host quality assessment and progeny performance of aphidiine parasitoids was examined using the mealy plum aphid parasitoid, Aphidius transcaspicus Telenga (Hymenoptera: Braconidae) and the black bean aphid, Aphis fabae Scopoli (Homoptera: Aphididae), as a readily acceptable alternate host. Aphid size in relation to stage of development was manipulated by rearing synchronous aphid cohorts at either 15 or 30 °C. At 15 °C, 2nd instar aphids were approximately the same size as 4th instar aphids reared at 30 °C. Cohorts of 30 aphids from each instar, reared at each temperature, were exposed to parasitism by a single parasitoid female for a period of 5 h. Overall susceptibility to parasitism did not vary between aphid cohorts, but the parasitoid response to aphid size differed significantly between rearing temperatures for both progeny sex ratio (parent female assessment of host quality) and larval growth and development (host suitability for parasitoid development). For aphids reared at 15 °C, the proportion of female progeny and emerging adult size for the parasitoid increased linearly with aphid size at the time of attack, while development time remained constant. In contrast, for aphids reared at 30 °C, the proportion of female progeny, emerging adult size, and the development time of the parasitoid all declined with aphid size at the time of attack. The contrasting responses of the parasitoid to host size for aphids reared at the two temperatures suggest that host quality is only indirectly related to aphid size among aphidiine parasitoids. The possible effects of higher temperatures on nutritional stress, obligate endosymbionts, and future growth potential of the aphids are discussed as explanations for the variation in host quality for parasitoid development.     

茉莉酸甲酯诱导大戟三萜类代谢的研究

京大戟是多年生草本药用植物,入药部分是其干燥根,但可入药的京大戟资源由于生长缓慢以及环境污染的加剧而越发匮乏,因此解决大戟资源日益紧张的问题是当今药用植物资源开发与利用方向的重要课题。京大戟含有三萜类、二萜类、黄酮类等丰富的活性成分,一些常见药用植物的有效成分是三萜类化合物,其在抗病毒、抗肿瘤、免疫调节等方面具有很好的活性。对植物萜类物质代谢起重要作用的关键酶,如3-羟基,3-甲基戊二酰辅酶A还原酶(hmgr)、鲨烯合酶(sqs)、法尼基焦磷酸合酶(fps)的基因克隆及活性研究取得了进展和突破,但通过调控萜类物质代谢途径中关键酶基因的表达来诱导终产物合成的研究鲜有报道。通过研究大戟萜类物质代谢途径进而利用基因工程手段提升目的物质的产量来解决京大戟药源短缺问题具有重要意义。该研究以大戟愈伤组织为材料,使用茉莉酸甲酯分别按时间梯度和浓度梯度进行诱导,将诱导后的愈伤组织分为两部分:一部分提取其总RNA,以actin为内参基因进行反转录,实时定量RT-PCR分析大戟三萜类代谢途径中hmgr、sqs与fps基因的相对表达差异;另一部分用于提取其总三萜并使用分光光度法进行含量测定。实时定量RT-PCR分析结果表明,茉莉酸甲酯可诱导3个基因的表达,但其表达模式不一样。相应的京大戟愈伤组织中总三萜的含量明显提高,最高可较未处理样品增加27%。研究结果可为茉莉酸甲酯促进药用植物大戟三萜类物质积累的分子机制研究提供参考。     

The effect of multi‐species host density on superparasitism and sex ratio in a gregarious parasitoid

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Egg load is a cue for offspring sex ratio adjustment in a fig‐pollinating wasp with male‐eggs‐first sex allocation

Fig‐pollinating wasps (Agaonidae) only reproduce within fig tree inflorescences (figs). Agaonid offspring sex ratios are usually female‐biased and often concur with local mate competition theory (LMC). LMC predicts less female‐bias when several foundresses reproduce in a fig due to reduced relatedness among intra‐sexually competing male offspring. Clutch size, the offspring produced by each foundress, is a strong predictor of agaonid sex ratios and correlates negatively with foundress number. However, clutch size variation can result from several processes including egg load (eggs within a foundress), competition among foundresses and oviposition site limitation, each of which can be used as a sex allocation cue. We introduced into individual Ficus racemosa figs single Ceratosolen fusciceps foundresses and allowed each to oviposit from zero to five hours thus variably reducing their eggs‐loads and then introduced each wasp individually into a second fig. Offspring sex ratio (proportion males) in second figs correlated negatively with clutch size, with males produced even in very small clutches. Ceratosolen fusciceps lay mainly male eggs first and then female eggs. Our results demonstrate that foundresses do not generally lay or attempt to lay a ‘fixed’ number of males, but do ‘reset to zero’ their sex allocation strategy on entering a second fig. With decreasing clutch size, gall failure increased, probably due to reduced pollen. We conclude that C. fusciceps foundresses can use their own egg loads as a cue to facultatively adjust their offspring sex ratios and that foundresses may also produce more ‘insurance’ males when they can predict increasing rates of offspring mortality.     

Brood Guarding by an Adult Parasitoid Reduces Cannibalism of Parasitoid-Attacked Conspecifics by a Caterpillar Host

Adult female bethylid parasitoids (Hymenoptera: Bethylidae) commonly guard their offspring until they develop into later immature stages to protect them from competing parasitoids and predators. Another possible mortality factor is from caterpillar larvae that cannibalize parasitized conspecific larvae. This study determined the effect of brood guarding on the occurrence of host cannibalism by using a brood guarding bethylid parasitoid Goniozus legneri and a non-guarding braconid parasitoid Habrobracon gelechiae (Hymenoptera: Braconidae), both are gregarious ectoprasitoids that attack the obliquebanded leafroller Choristoneura rosaceana (Lepidoptera: Tortricidae). When a C. rosaceana larva parasitized by G. legneri was presented to a live conspecific larva, the frequency of cannibalism on the parasitized conspecific was lower in the presence than in the absence of a guarding female G. legneri. In contrast, when a C. rosaceana larva parasitized by H. gelechiae was presented to a live conspecific larva, the presence or absence of H. gelechiae did not affect cannibalism frequency. Cannibals did not gain a fitness advantage in terms of larval survival, developmental time, or body mass of developed pupae; therefore, cannibalism by the host larva may be a group defense against increased parasitism in subsequent population generations. We suggest that brood guarding by parasitoids can act as a broad defense strategy to protect offspring from the cannibalism of parasitized conspecifics by the host larvae.     

Nonsiblicidal behavior and the evolution of clutch size in bethylid wasps

Parent-offspring conflict over clutch size may lead to siblicidal behavior between juveniles. In parasitoid wasps, selection for siblicide in small broods is predicted to produce a dearth of gregarious broods with few eggs. Here we document the clutch size distribution in the Bethylidae, a large family of aculeate parasitoids. Small gregarious clutches are the most common. Further data suggest that the most common gregarious clutches in the parasitoid Hymenoptera as a whole contain only a few eggs. Across bethylid species, both clutch size and wasp size increase with host size. Within genera clutch size is more closely related to host size, but between genera or larger clades wasp size is more closely related to host size. The volume of the emerging wasp brood does not depend on whether a species lays single- or multiple-egg clutches once host size is taken into account. These data suggest that clutch size in bethylid wasps is best described by traditional optimality models and that siblicide plays little role in this and possibly other families. We propose several ecological reasons for the rarity of siblicide in bethylids: ectoparasitism, idiobiosis, and a suite of characteristics associated with high within-brood relatedness.