Phoresy in the field: natural occurrence of Trichogramma egg parasitoids on butterflies and moths

Phoretic insects utilize other animals to disperse to new environments. We recently discovered how egg parasitoids use an exciting phoretic strategy to reach egg-laying sites of their butterfly hosts. In the laboratory, female Trichogramma wasps detect and mount mated female cabbage white butterflies that emit an anti-aphrodisiac pheromone. Hardly any information exists about the natural occurrence of phoresy in wasps of this genus. Therefore, we monitored the presence of phoretic Trichogramma wasps on lepidopteran hosts in the field. Only female wasps were found at low prevalence on six lepidopteran species. Wasps were mostly found on female hosts and mainly on abundant solitary host species. This is the first report of phoretic Trichogramma wasps on butterflies in nature. We suggest that phoresy is only one of several strategies used by these polyphagous egg parasitoids. The evolution of phoresy is discussed in relation to the nutritional ecology of egg parasitoids.     

Mothers modify eggs into shields to protect offspring from parasitism

Eggs are an immobile, vulnerable stage of development and their success often depends on the oviposition decisions of the mother. Studies show that female animals, and sometimes males, may invest parental resources in order to increase the survival of their offspring. Here, we describe a unique form of parental investment in offspring survival. The seed beetle Mimosestes amicus may lay eggs singly, or may cover eggs with additional egg(s). This egg stacking serves to significantly reduce the mortality of the protected egg from parasitism by the parasitic wasp, Uscana semifumipennis. The smaller top eggs serve only as protective shields; they are inviable, and wasps that develop in them suffer negative fitness consequences. Further, we found egg stacking to be inducible; M. amicus increase the number of stacks they lay when parasitoids are present. However, stacking invokes a cost. When wasps are absent, beetles lay more single eggs, and produce more offspring, highlighting the adaptive value of this extraordinary example of behavioural plasticity in parental investment.     

The dynamics of egg production, oviposition and resorption in a parasitoid wasp

1. The extent to which parasitoid wasps are limited by their egg supply is very important in understanding their reproductive strategies. Egg reserves are dynamic, with most wasps maturing new eggs throughout their life (synovigeny) and many species resorbing eggs that are not used in oviposition. We investigated the extent to which a parasitoid modulates its egg reserves in the light of its experience in finding hosts.
2. The egg dynamics of the Encyrtid Wasp Leptomastix dactylopii , a solitary parasitoid of mealybugs, were studied in the laboratory. This species is synovigenic and practises egg resorption.
3. We allowed newly emerged wasps to experience one of four environments of increasing value in terms of reproductive opportunities. We proposed that wasps that experienced good quality environments would maintain more mature eggs ready for oviposition. Dissection of wasps subject to different periods of host deprivation after the experimental treatment failed to confirm the hypothesis: egg load was independent of experience.
4. We also proposed that any adjustment of egg supply to make up for eggs oviposited would be effected through a reduction in egg resorption. Instead, we found that the wasp quickly made up for eggs oviposited by increased egg production.     

Egg deposition patterns of fig pollinating wasps: implications for studies on the stability of the mutualism

1. Fig pollinating wasps (Agaonidae) enter Ficus inflorescences (figs), oviposit in some of the flowers, and pollinate in the process. Each larva completes its development within a single flower. In most cases, an inflorescence entered by a wasp will represent its only egg‐laying site. The mechanisms that prevent pollinating wasps from exploiting all the flowers inside a fig are not understood. In this study, hypotheses about flower use by pollinating fig wasps were tested by investigating egg deposition patterns in three species. 2. Either one or three wasps were introduced into figs. The figs were then harvested. Serial sections allowed assessment of the presence or absence of a wasp egg in a sample of flowers in each fig. The overall proportion of flowers with eggs and the spatial distribution of eggs were then compared in single wasp figs and three foundress figs. 3. In all species, the proportion of flowers with a wasp egg increased with foundress number but less than three‐fold. 4. In all species, at least in single foundress figs, flowers near the fig cavity were more likely to receive a wasp egg than were flowers near the fig wall. 5. In two species, when the number of foundresses was multiplied by three, there was an increase in the use of flowers near the fig wall, while in the third species, the increase was spread evenly among flowers. 6. Factors affecting wasp egg deposition patterns and the potential of investigating such patterns for studying the stability of the mutualism are discussed.     

Effects of size and fluctuating asymmetry on field fitness of the parasitoid Trichogramma carverae (Hymenoptera: Trichogrammatidae)

1. The effects of body size and asymmetry in morphological traits on field fitness were studied in the parasitoid Trichogramma carverae .
2. Significant fluctuating asymmetry was detected in four bristle counts and two wing measurements made on forewings. There was no evidence for directional asymmetry in any of the traits. An estimate of field fitness was obtained in grapevines by collecting released wasps arriving at oviposition sites consisting of egg rafts of the tortricid Epiphyas postvittana .
3. Comparisons of ovipositing and emergence samples indicated that wasps at oviposition sites were relatively larger and more variable in their size distribution. A non-parametric analysis of the relationship between fitness and size indicated that extremely large wasps had the highest fitness and suggested that small as well as large wasps may have had a fitness advantage.
4. For asymmetry, the only trait showing an association with field fitness was wing length. Wasps with a low length asymmetry were more likely to be collected at oviposition sites, although fitness curves indicated that wasps had a similar fitness once an intermediate length asymmetry was exceeded.
5. Mother–daughter comparisons for wasps from a genetically heterogeneous stock provided no evidence that size measures or asymmetries were heritable when wasps were reared on a factitious host.
6. These findings have implications for improving parasitism rates in inundative releases.     

A new biological method to control Anobium punctatum,by using the parasitoid wasp Spathius exarator

Biological pest control by means of beneficial organisms is for long part of agriculture and Integrated Pest Management (IPM). A new and efficient strategy to control the most common timber pest species in churches and museums, the furniture beetle Anobium punctatum (De Geer) (Coleoptera: Anobiidae), is based on the parasitoid wasp species Spathius exarator (Linnaeus) (Hymenoptera: Braconidae). Once this braconid wasp detects its host species beneath the surface, it pierces the wood with its ovipositor to lay one single egg onto the beetle larva. After hatching, the wasp larva feeds on the beetle larva thereby killing it. Afterwards, it pupates and emerges through a self-gnawed hole as an adult wasp. The tiny, 0.5-mm-wide exit hole can easily be distinguished from the 1- to 2-mm-wide exit hole of A. punctatum. Laboratory tests revealed that female wasps have an average life span of 85 days and produce a total of 24 offspring, when nutrition is provided. Between 2012 and 2019, braconid wasps were introduced into 54 different A. punctatum infested buildings. Treatment success was monitored by examining exit holes of new beetles and wasps thereby calculating corresponding parasitism rates. After the first year of treatment, parasitism rates were significantly higher with a mean value of 0.15 when compared to untreated objects with a natural parasitism and a mean value of 0.08 (n = 54). Following treatment of three objects over a period of eight years, parasitism rates continuously increased from 0.02 up to 0.31. In a church organ, which was treated during this period, the monitoring revealed a complete suppression of pest activity. These data prove that this biological method of pest control is an efficient, sustainable and non-toxic option to manage the common furniture beetle.     

Behaviour of a specialist parasitoid, Cotesia melitaearum: from individual behaviour to metapopulation processes

Abstract. 1. Foraging behaviour and movement within and among host patches of the specialist parasitoid wasp Cotesia melitaearum (Braconidae) attacking the larvae of Melitaea cinxia (Nymphalidae) were studied in the field and in the laboratory.
2. In the field, female wasps aggregated in large host groups in the autumn and caused positive spatial density-dependent parasitism in the field. Wasps stayed longer with groups of pre-diapause caterpillars than with post-diapause caterpillars, but attacked them less frequently.
3. In the laboratory, wasps attacked larger larvae more readily than smaller larvae. Also in the laboratory, wasps exposed to larvae outside their protective webs showed differences in the rates at which they attacked larvae fed different diets, implicating host plant-derived chemicals as proximate cues for foraging wasps.
4. Mark–recapture studies indicated that there was a low rate of successful movement of wasps among groups of young larvae within a habitat patch in the autumn and no successful movement of wasps across non-habitat. In contrast, wasps moved frequently among groups of late-instar caterpillars in the spring.
5. Host caterpillars of different ages responded very differently to wasp attacks. Pre-diapause larvae remained in groups and used collective head-jerking behaviour to defend themselves, whereas post-diapause larvae dispersed away from the group immediately after being attacked.
6. Population and metapopulation level dynamics of the host–parasitoid interaction are discussed in light of these observations of the behaviour of individual wasps.     

Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on Banisteriopsis malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system

In the Brazilian savanna many plant species bear regular associations with patrolling ants that are aggressive towards insect herbivores. However, not only ants but also several species of predatory wasps are attracted to plants due to the extrafloral nectaries (EFNs). Such wasps feed on both herbivores and plant exudates. In this study we describe the foraging behavior of the social Polistinae wasp Brachygastra lecheguana in the extrafloral nectaried shrub Banisteriopsis malifolia, and investigated the influence of patrolling ants Camponotus blandus on the activity of the wasp. Brachygastra lecheguana fed on the endophytic larvae of Anthonomus (Curculionidae) beetles that developed inside flower buds. The wasp lacerated the bud layers to reach the beetle larvae located at the bud core. The wasp visits to Ba. malifolia were statistically related to the abundance of flower buds and beetles. Ant exclusion experiments revealed that the hunting behavior of B. lecheguana on beetles was not related to the absence of C. blandus. However we found that wasps spent more time consuming extrafloral nectar on branches where ants were excluded. This is the first study reporting extrafloral nectar consumption by B. lecheguana, as well as the predation on herbivores in natural areas. In cerrado vegetation, ants benefit the plant by reducing insect herbivores, and our study provides evidence that the B. lecheguana – Ba. malifolia system represents a potential interaction where the wasp may also benefit the host plant. The value of this wasp species as a plant‐guard is discussed.     

Analysis of Tiphia parasitoids preovipositional behaviors and of their scarab host defensive responses

Tiphia vernalis Rohwer and Tiphia popilliavora Rohwer were introduced as biocontrol agents against Japanese beetles (Popillia japonica Newman) and oriental beetles (Anomala orientalis Waterhouse). Studies have shown that under field conditions, T. vernalis parasitize Japanese beetles more often than oriental beetles. This study was done to understand how tiphiid wasps handle the two different host species and the influence of host defensive behaviors on the oviposition process of tiphiid wasps. The preovipositional behaviors performed by Tiphia wasps included: stinging, examining, moving soil, kneading, host-feeding, and host scraping. The frequency, sequence, and total time spent on each behavior before oviposition were scored and compared between two host species. The sequence and frequency of preovipositional behaviors performed by both Tiphia wasps did not show a difference between the two host species. However, female T. vernalis spent significantly longer time trying to sting oriental beetles than Japanese beetles in order to paralyze them. The time T. popilliavora spent on prestinging behaviors did not show a difference between Japanese and oriental beetles. The defensive behaviors performed by Japanese and oriental beetle grubs included: vigorous movements, rubbing their abdomen or head against the wasp’s abdomen, and biting at the attacking wasp. The frequency and total time spent on each defensive behavior was scored and compared between two host species. Overwintered, third instar oriental beetle grubs spent significantly longer time on defensive behaviors when they were attacked by T. vernalis which likely cause wasps to spend longer time trying to sting oriental beetle grubs. The active host resistance gained through behavioral defenses could make oriental beetle grubs less susceptible to T. vernalis attack than Japanese beetle grubs, especially under field conditions. Younger grubs attacked by T. popilliavora did not exhibit these differences.     

Interactions of the ant Crematogaster scutellaris with the fig/fig wasp mutualism

Abstract. 1. A classical example of specialised pollination mutualisms is the relationship between fig trees and their pollinating wasps, in which each partner depends completely on the other for its reproduction; however the fig/fig wasp association is also the target of a great diversity of other species, ranging from specialised parasites to opportunistic foragers, among them ants.
2. The ant community and the sources of ant attraction observed on the Mediterranean fig tree Ficus carica were characterised.
3. A guild of ants attracted by homopterans tended on the plant was distinguished from a second guild composed of two co-dominant ant species ( Crematogaster scutellaris and Pheidole pallidula ) that prey mostly on pollinating wasps, abundant during certain parts of the fig cycle.
4. Foraging workers of C. scutellaris search for prey on the fig inflorescence (syconium), capturing pollinating wasps mostly at the peak of wasp emergence and at a rate estimated to reach 600 prey per day for an entire tree.
5. Detailed study of the predatory sequences displayed under experimental conditions showed that ant workers captured 100% of the pollinating wasps offered, while they captured only 5.5% of the parasitoid wasp specific to the pollinator. The respective impacts of the interaction on ants and on the figs are discussed, as well as several behavioural traits of predation by the dominant ant on pollinators.     

Host-feeding and acceptance by a parasitic wasp (Hymenoptera: Ichneumonidae) as influenced by egg load and experience in a patch

Optimal host selection models based on dynamic programming predict that the physiological state of a foraging insect, i.e. egg load, energy reserves etc., influences behavioral decisions. To test this prediction, the effect of physiological state on host acceptance of the ectoparasitic wasp Agrothereutes lanceolatus was investigated. Female wasps in plastic cups (regarded as patches) were presented with hosts, and their responses to the hosts were continuously observed. After observations, the wasps were dissected and the number of mature and immature eggs they carried were counted. The results showed that behavioral decisions by the female wasps were influenced by mature egg load, but not by wasp size or immature egg load. Hence the wasps with higher egg loads were more likely to oviposit. The number of hosts previously encountered in a patch (i.e. wasp experience) also had an independent effect on females' host acceptance, indicating that female informational state was updated during foraging in that patch. Female wasps host-fed only when mature egg load approached zero. Concurrent host-feeding was not observed. Parasitoid survival was almost zero when parasitoid eggs were transferred onto hosts that were fed upon, indicating that concurrent host-feeding could cause a high degree of offspring mortality. These three results supported the assumption and prediction of optimal host-feeding models. Parasitoid host selection and host-feeding are discussed in the context of recent models.     

Rho 1 participates in parasitoid wasp eggs maturation and host cellular immunity inhibition

Endoparasitoid wasps introduce venom into their host insects during the egg-laying stage. Venom proteins play various roles in the host physiology, development, immunity, and behavior manipulation and regulation. In this study, we identified a venom protein, MmRho1, a small guanine nucleotide-binding protein derived from ovary in the endoparasitoid wasp Microplitis mediator and found that knockdown of its expression by RNA interference caused down-regulation of vitellogenin and juvenile hormone, egg production, and cocoons formation in the female wasps. We demonstrated that MmRho1 entered the cotton bollworm's (host) hemocytes and suppressed cellular immune responses after parasitism using immunofluorescence staining. Furthermore, wasp MmRho1 interacted with the cotton bollworm's actin cytoskeleton rearrangement regulator diaphanous by yeast 2-hybrid and glutathione s-transferase pull-down. In conclusion, this study indicates that MmRho1 plays dual roles in wasp development and the suppression of the host insect cellular immune responses.     

黑盾胡蜂单母建群习性的观察

通过引蜂筑巢研究黑盾胡蜂VespabicolorFabricius单母建群的生物学习性。结果表明 :母蜂在最初 3只子代工蜂羽化之前兼负蜂王和工蜂的职能 ,建成外包巢壁内具有 3 4个巢房的小巢 ,饲育的幼虫数不超过 1 7只 ,各虫态历期分别为卵期 5 5d ,幼虫期 1 2d,蛹期 1 2 5d ;母蜂日外勤飞翔次数与天气相关 ,晴天可达 42次 ,其中近 80 %为捕食。作者认为 ,黑盾胡蜂单母建群时期可能是其生活史中既脆弱又关键的阶段。     

Importance of plant size, distribution of egg masses, and weather conditions on egg parasitism of the European corn borer, Ostrinia nubilalis by Trichogramma ostriniae in sweet corn

Trichogramma ostriniae (Hym: Trichogrammatidae), an egg parasitoid of the European corn borer, Ostrinia nubilalis (Lep: Pyralidae), were released into sweet corn (Zea mays L.) fields to study the effects of weather, plant size and distribution of egg masses on egg parasitism by the wasp. Sentinel European corn borer eggs were stapled onto leaves located in the upper, middle and lower third of sweet corn plants 5 to 35 meters away from the wasp release point in either a radial or grid manner. Weather conditions and plant architecture were monitored during the experiments. Logistic regression was used to analyze the data. The results indicated that percentage of eggs parasitized was negatively related to an increase in leaf area as well as an increase in distance eggs were located from the point of release of wasps. Eggs distributed on plants at different directions from the release point received different levels of parasitism. Eggs that were stapled onto leaves in the upper third of a corn plant received much less parasitism than those on the middle and lower third of the plant. Higher mean temperature adversely affected the level of parasitism during hotter times of the season and conversely, lower temperatures (<17 °C ) reduced the egg parasitism during cooler times of the season. The longer the exposure of eggs to wasps, the higher the level of egg parasitism. However, the levels of egg parasitism for 2 day's exposure were almost the same as that for 3 day's exposure due to the limited longevity and egg-laying behavior of the wasp. These results suggest that inundative releases of T. ostriniae should be made every two to three days, with multiple release points per hectare. In addition, weather conditions and plant architecture, especially temperature, plant height and leaf area must be taken into consideration to optimize levels of parasitism.     

聚果榕传粉榕小蜂传粉与产卵之间的权衡

【目的】榕树(Ficus)依赖专性榕小蜂(Agaonidae)传粉,同时为传粉榕小蜂提供繁衍后代的场所,两者形成动植物间经典的协同进化关系。在雌花期果内,榕小蜂需在有限的存活时间内完成传粉和产卵,而传粉榕小蜂如何在传粉与产卵之间进行权衡仍然是悬而未解的问题。本研究旨在明确传粉榕小蜂——一种栉颚榕小蜂Ceratosolen sp.在雌雄同株的聚果榕Ficus racemosa雌花期果内的行为活动及繁殖模式。【方法】借助测微尺测量聚果榕榕果雌花花柱长度与传粉榕小蜂(Ceratosolen sp.)产卵器长度,通过显微视频记录传粉榕小蜂在雌花期果内搜索、传粉及产卵行为;结合单果控制性引蜂试验,测定不同阶段榕小蜂个体大小、孕卵量、携粉量,以及雄花期最终繁殖的榕小蜂后代和榕果种子数量。【结果】聚果榕雌花花柱长度存在树间变异,榕小蜂产卵器长度比绝大多数的雌花花柱长,说明该小蜂可以产卵于大部分的雌花子房里。通常个体大的榕小蜂孕卵量更多,但个体大小与携粉量之间相关性不显著。观察发现,榕小蜂进入雌花期榕果内,前6 h集中产卵,可产下孕卵量的95%,平均搜索用时27 s,产卵用时46 s,此期间传粉行为少见,花粉筐中携带花粉量亦无明显变化;榕小蜂进果后6-24 h,主要执行传粉,其行为主动,连贯高效,单次传粉用时平均为2 s,最终可传完携粉量的80%。控制引蜂试验也证实榕小蜂进入榕果内前6 h主要执行产卵繁殖后代,之后6-24 h主要执行传粉以繁殖榕树种子。【结论】在雌雄同株的聚果榕雌花期榕果内,榕小蜂先产卵、后传粉。本研究首次展示了传粉榕小蜂在聚果榕雌花期榕果内的产卵和传粉行为,并获得与行为相匹配的产卵量和传粉繁殖量,反映了具主动传粉行为的榕小蜂在传粉和产卵之间存在时间和数量上的权衡。     

歪叶榕非传粉小蜂的繁殖策略及其对榕-蜂共生系统的影响

2004年8月至2005年8月在西双版纳热带植物园内,通过广泛收集歪叶榕榕小蜂标本、非传粉小蜂产卵行为学观察和阻止传粉者入果等实验方法,研究了我国西双版纳热带雨林下的一种榕树——歪叶榕Ficus cyrtophylla的榕小蜂群落组成结构、非传粉小蜂的繁殖策略以及它们对榕-蜂共生系统的影响。结果表明,歪叶榕中除了具有唯一传粉榕小蜂Blastophag sp.以外,还具有3种非传粉小蜂Platyneura sp.、Philotrypesis sp.和Sycoscapter sp.。在歪叶榕榕小蜂群落中,传粉榕小蜂占整个群落总数的92.21%,是群落的最主要组成者;主要的非传粉小蜂是Sycoscaptersp.,占5.78%; 其次是Philotrypesissp.,占1.84%,而Platyneurasp.仅占群落总数的0.17%。歪叶榕中的非传粉小蜂通过各自产卵时间和食性分化的策略来利用榕果中的资源繁殖后代。非传粉小蜂寄生使传粉榕小蜂的总数和其雌蜂数量都显著地降低,但是对传粉小蜂雄蜂数量没有显著影响,从而导致传粉榕小蜂的雄性性比显著地增加。这说明非传粉小蜂在选择寄居宿主时具有明显的倾向性,而且更多地将卵产于含有雌性传粉小蜂的瘿花之中。因此,非传粉小蜂通过减少雌性传粉小蜂的数量而降低了榕树的雄性适合度,从而在一定程度上对榕 蜂共生系统的稳定存在和发展产生了负面影响。     

Natural selection in the laboratory for a change in resistance byDrosophila melanogaster to the parasitoid waspAsobara tabida

The selection response of the polymorphic hostD. melanogaster (Meigen) to the braconid waspA. tabida (Nees) is addressed. Cages of flies with and without wasps were initiated with a population ofD. melanogaster that exhibited variation both in larval foraging behavior and in encapsulation ability. Encapsulation ability was measured as the proportion of parasitized larvae that produce a hardened capsule which encapsulates the wasp egg and ultimately kills the wasp larva. We determined whether the host population changed its encapsulation ability and/or its foraging behavior in response to the wasp. Both species were collected from a local orchard whereA. tabida is the only wasp known to parasitizeD. melanogaster larvae. The naturally occurring genetic polymorphism for rover and sitter larval foraging behavior inD. melanogaster is also found in this field population.A. tabida's vibrotactic search behavior enables it to detect rover more frequently than sitter larvae. Rover larvae move significantly more while feeding than do sitter larvae. In this field population, rover larvae also show higher encapsulation abilities than do sitter larvae. Six cage populations, three without wasps and three with wasps, each containing an equal mixture of rover and sitter flies, were established in the laboratory and maintained for 19 fly generations. Selection pressure in the laboratory was similar to that found in the field population from which the flies and wasps were derived. We found that larvae from cages with wasps developed a significantly higher frequency of encapsulation than those reared without wasps. We were, however, unable to detect a change in larval movement (rover or sitter behavior) in larvae from cages subject to selection from wasps compared to larvae from cages containing no wasps. This may have resulted from a balance between two selective forces, selection against rovers by the wasps' use of vibrotaxis, and selection for rovers resulting from their increased encapsulation abilities     

Sublethal effect of brood parasitism on the grass-carrying wasp Isodontia mexicana

Abstract.  1. The nests of solitary, nest-provisioning wasps (Sphecidae and Crabronidae) are commonly attacked by brood parasites, including flies of the families Phoridae and Sarcophagidae. Larvae of the flies commonly kill the wasp offspring directly or starve it by consuming prey provided by the adult female wasps.
2. To determine whether brood parasitic flies can have a sublethal effect (i.e. reduced body size) on wasp offspring, nests of the wasp Isodontia mexicana were collected at two field sites in upstate New York, U.S.A. Nest diameter had no effect on the probability that a wasp cell would be inhabited by a brood parasite. Most offspring that developed in cells also containing phorids or sarcophagids managed to complete development and emerge as adults. Nevertheless, they had significantly smaller body size than conspecifics emerging from unparasitised cells in which the developing wasp did not have to compete for food (which in this species consists of tree crickets and katydids). Apparently, this is the first time that a sublethal effect of brood parasitism on offspring body size has been quantified for a solitary wasp species. Known effects of body size on the reproductive success of adult wasps suggest that sublethal consequences of the presence of brood parasites may have a significant effect on the fitness of adult I. mexicana .     

Relative investment in egg load and poison sac in fig wasps: Implications for physiological mechanisms underlying seed and wasp production in figs

Fig pollinating wasps and most non-pollinator wasps apply secretions from their poison sacs into oviposited flowers that appear necessary to the formation of the galls that their developing offspring consume. Thus, both eggs and poison sac secretions appear to be essential for wasp reproduction, but the relative investment in each is unknown. We measured relative investment in poison sac and egg production in pollinating and non-pollinating wasps associated with seven species of monoecious Panamanian figs representing both active and passive pollination syndromes. We then collected similar data for four fig hosts in China, where some wasp species in the genus Eupristina have lost the ability to pollinate (“cheaters”). All wasps examined possessed large poison sacs, and we found a strong positive correlation between poison sac size and absolute egg production. In the Panamanian species, the relative poison sac to egg investment was highest in the externally ovipositing non-pollinator wasps, followed by active pollinators, then by passive pollinators. Further, pollinator wasps of fig species with demonstrated host sanctions against “cheating” wasps showed higher investment in the poison sac than wasps of species without sanctions. In the Chinese samples, relative investment in the poison sac was indistinguishable between pollinators and “cheaters” associated with the same fig species. We suggest that higher relative investment in poison sac across fig wasp species reflects higher relative difficulty in initiating formation of galls and subsequently obtaining resources from the fig. We discuss the implications for the stability of the fig–wasp mutualism, and for the ability of non-pollinators to exploit this mutualism.     

Endogenous and exogenous factors affecting parasitism of gypsy moth egg masses by shape Ooencyrtus kuvanae

Factors affecting the orientation, reproduction, and sex ratio of the egg parasitoid Ooencyrtus kuvanae Howard were examined. Adult females were attracted to airborne volatiles from the egg mass and accessory gland of the primary host, the gypsy moth Lymantria dispar L. Visual cues also affected host selection. Background colors against which egg masses were placed affected oviposition preference. In the absence of egg masses, color variation did not affect wasp behavior. Light is required for parasitism by O. kuvanae. The age and density of both the host and parasitoid affected wasp reproduction and sex ratios. Older egg masses issued relatively fewer wasps and higher proportions of males than did young egg masses. Likewise, wasp reproduction and the proportion of females declined with wasp age. Larger egg masses produced more wasps and lower proportions of males than did smaller egg masses. The number of offspring per female, and the proportion of female offspring, were inversely related to wasp density. Implications to biological control of the gypsy moth and parasitoid ecology are discussed.