Influence of egg load and host size on host-feeding behaviour of the parasitoid Aphytis lingnanensis

Abstract.

• 1 Direct behavioural assays were used to investigate the influences of host size and parasitoid egg load on the decision to host feed versus oviposit made by the parasitoid Aphytis lingnanensis Compere. Egg load was manipulated without concurrent influences on the history of host contact by exploiting size-related variation in fecundity and by holding parasitoids at different temperatures to vary the rate of oocyte maturation.
• 2 Host feeding comprised a series of feeding bouts, separated by renewed probing of the scale insect body. Successive feeding bouts were progressively shorter, suggesting that hosts represent ‘patches’ yielding resources at a decelerating rate.
• 3 Parasitoids were significantly more likely to host feed on smaller hosts and oviposit on larger hosts.
• 4 Neither egg load nor the treatment variables (parasitoid size and holding temperature) exerted significant influences on the decision to host feed versus oviposit on second instar (low quality) hosts.
• 5 The failure to observe an effect of egg load on host-feeding decisions was not simply a reflection of the parasitoids being entirely insensitive to egg load; significant effects of egg load on parasitoid search intensity and clutch size decisions were observed.
• 6 Parasitoids developing on second instar (low quality) hosts experienced high levels of mortality late during development and yielded very small adults.
• 7 The discord between these experimental results and predictions regarding the importance of egg load underscores the need for additional work on the proximate basis for host-feeding decisions and the nutritional ecology of insect parasitoids.

     

Age-dependent clutch size in a koinobiont parasitoid

Abstract.  1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.
2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals.
3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.
4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time.
5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.
6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.
7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

Pre‐Patch Experience Affects the Egg Distribution Pattern in a Polyembryonic Parasitoid of Moth Egg Batches

According to foraging theory, female parasitoids should alter their host choice in response to cues that indicate a limitation of resources. We tested whether females of the polyembryonic parasitoid Ageniaspis fuscicollis (Hymenoptera: Encyrtidae), which attack egg batches of small ermine moths (Lepidoptera: Yponomeutidae), would alter their host acceptance pattern in response to different pre‐patch experience. We kept females of the parasitoid prior to a patch visit under different conditions, which should indicate different levels of competition for hosts. With increased competition as pre‐patch experience, females laid more eggs per host egg and self‐superparasitized more often, and the resultant egg distributions showed a trend from more regular distributions to increasingly Poisson and aggregated distributions. Consequently, females with a pre‐patch experience that would indicate low competition for hosts had the most even egg distributions. We conclude that pre‐patch experience of competitors may lead to a significant change of mutual interference patterns in egg‐laying A. fuscicollis wasps.     

Innate Host Habitat Preference in the Parasitoid Diachasmimorpha longicaudata: Functional Significance and Modifications through Learning

Parasitoids searching for polyphagous herbivores can find their hosts in a variety of habitats. Under this scenario, chemical cues from the host habitat (not related to the host) represent poor indicators of host location. Hence, it is unlikely that naïve females show a strong response to host habitat cues, which would become important only if the parasitoids learn to associate such cues to the host presence. This concept does not consider that habitats can vary in profitability or host nutritional quality, which according to the optimal foraging theory and the preference-performance hypothesis (respectively) could shape the way in which parasitoids make use of chemical cues from the host habitat. We assessed innate preference in the fruit fly parasitoid Diachasmimorpha longicaudata among chemical cues from four host habitats (apple, fig, orange and peach) using a Y-tube olfactometer. Contrary to what was predicted, we found a hierarchic pattern of preference. The parasitism rate realized on these fruit species and the weight of the host correlates positively, to some extent, with the preference pattern, whereas preference did not correlate with survival and fecundity of the progeny. As expected for a parasitoid foraging for generalist hosts, habitat preference changed markedly depending on their previous experience and the abundance of hosts. These findings suggest that the pattern of preference for host habitats is attributable to differences in encounter rate and host quality. Host habitat preference seems to be, however, quite plastic and easily modified according to the information obtained during foraging.     

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.     

Olfactory responses of Anaphes nitens (Hymenoptera,Mymaridae) to host and habitat cues

The eucalyptus weevil, Gonipterus spp. Schoenherr, 1833 (Coleoptera, Curculionidae) is considered a major pest of eucalyptus plantations. In regions where control is achieved, success is usually brought by the action of the solitary egg parasitoid Anaphes nitens (Girault, 1928) (Hymenoptera, Mymaridae). Research was conducted to ascertain which cues might mediate female wasp host location and selection. In Petri dish arenas, females were attracted to Gonipterus platensis Marelli, 1927 egg capsules, to G. platensis mated female faeces and to leaves of Eucalyptus globulus Labillardière, 1799. Gas chromatography–mass spectrometry electroantennographic detection analysis was conducted using extracts obtained from leaves of E. globulus, from G. platensis egg capsules, as well as from adults of both sexes and their faeces, in order to detect and identify compounds perceived by the wasp's olfactory system. The parasitoids were shown to detect a wide range of compounds emitted by each one of these sources, and for 31 compounds, antennal response was confirmed by dose-response tests. Further behavioural trials were conducted in Petri dishes in order to decode the effect, on parasitoid behaviour, of selected compounds emitted by E. globulus and of the pheromones, emitted by the weevils on parasitoid behaviour. Attraction was observed to two compounds emitted by E. globulus, namely 1,8-cineole and γ-terpinene, and to the main component of the male sex/aggregation pheromone, cis-verbenol. To our knowledge, this is the first report of attraction of a parasitoid from the family Mymaridae to a component of its host's sexual/aggregation pheromone. Similarly, to other egg parasitoid species, A. nitens females are likely to use the host plant volatiles as long-range host location cues and to adopt the ‘infochemical detour’ strategy in order to get in the vicinity of their hosts.     

Behavioural and body mass changes before egg laying in the Barn Owl: cues for clutch size determination?

To investigate laying decision and clutch size determination in indeterminate layers, we analysed in-nest activity (nest presence, and copulation, prey deliveries, and entrance frequencies) and female body mass change, as well as their relation to clutch size variation in five Barn Owl pairs (Tyto alba) nesting in eastern France. Body mass of the female and behaviour [copulation frequency, entrance frequency, and prey delivery to the nest by the male (in number and mass)] were monitored using an automated weighing system and a video camera. There was a consistent change of behaviour and foraging activity among pairs ca. 18 days before laying indicating that the females may be tied to the nest at this time. Barn Owls being indeterminate layers have their clutch size determined at the oviposition of the first egg of the clutch. Window correlation analyses between the clutch size and the female body mass gain indicate that the clutch size might be determined no later than a few days before the laying of the first egg. Our results suggest that female Barn Owls may use the pre-laying period to determine the clutch size using cues such as the male food deliveries (a proxy for male quality).     

Attractive blue‐green egg coloration and cuckoo−host coevolution

Recent evidence suggests that blue‐green coloration of bird eggshells may be related to female and/or egg phenotypic quality, and that such colour may affect parental effort and therefore the nutritional environment of developing nestlings. Here we suggest that these relationships and the signal function of eggshell coloration would affect the outcome of coevolution between avian brood parasites and their hosts in at least three different non‐exclusive evolutionary pathways. First, by laying blue‐green coloured eggs, cuckoo females may exploit possible sensory biases of their hosts, constraining the evolution of parasitic egg recognition, and thus avoid rejection. Second, because of the relatively high costs of laying blue eggs, cuckoo females may be limited in their ability to mimic costly blue‐green eggs of their hosts because cuckoo females lay many more eggs than their hosts. Furthermore, costs associated with foreign egg recognition errors would be relatively higher for hosts laying blue eggs. Third, cuckoos may use coloration of host eggs for selecting individuals or specific hosts of appropriate phenotypic quality (i.e. parental abilities). We here explored some predictions emerging from the above scenarios and found partial support for two of them by studying egg coloration of European cuckoos (Cuculus canorus) and that of their 25 main hosts, as well as parasitism and rejection rate of hosts. Cuckoo hosts parasitized with more blue, green, and ultraviolet cuckoo eggs, or those laying more blue‐green eggs, were more prone to accept experimental parasitism with artificial cuckoo eggs. In addition, coloration of cuckoo eggs is more variable when parasitizing hosts laying bluer‐greener eggs, even after controlling for the effect of host egg coloration (i.e. degree of egg matching). Globally, our results are consistent with the proposed hypothesis that host egg traits that are related to phenotypic quality of hosts, such as egg coloration, may have important implications for the coevolutionary interaction between hosts and brood parasites. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 154–168.     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

Errors in egg-laying by female Common Cuckoo Cuculus canorus in nests of its common host

Dozens of studies have documented that brood parasites are well adapted to a brood parasitic lifestyle but not all parasitism events are successful. Co-evolution between brood parasites and their hosts is a dynamic process so it is reasonable to expect that a female brood parasite may commit errors during egg deposition by laying her eggs outside the laying period of the host, with consequent impacts on her fitness. Using an extensive dataset from a long-term study, we evaluated egg-laying patterns and errors related to the timing of egg-laying in the Common Cuckoo Cuculus canorus (hereafter ‘Cuckoo’). Specifically, we tested whether the Cuckoo avoids laying before or on the day of host clutch initiation to reduce the risk of rejection of parasitic eggs, whether laying errors will be more frequent in periods with a lack of active host nests, and whether the laying errors will be more frequent in periods with intense Cuckoo parasitism and a consequent lack of suitable host nests. We found that about one-third of Cuckoo eggs were laid on the host clutch initiation day or 1 day before, and the percentage of Cuckoo eggs laid decreased thereafter. Surprisingly, the probability of Cuckoo egg acceptance by the hosts was not affected by the egg-laying stage of the host clutch. Errors in the timing of egg-laying with fatal consequences (i.e. those precluding Cuckoo hatching because of laying in incubated or deserted clutches) were recorded in about 5% of cases. Only laying date of a Cuckoo egg had a significant effect on the probability of errors, which increased during the breeding season. This may be related to the higher number of deserted and incubated host nests at the site at the end of the breeding season. Errors in egg-laying may be attributed to young and inexperienced females but also impaired body condition or intraspecific competition may cause this behaviour. Future studies, which will test these possible explanations, will help to understand better the mechanism of co-evolutionary arms races and differences between host specialist and generalist brood parasites in various host–parasite systems.     

Potential for climate effects on the size-structure of host–parasitoid indirect interaction networks

Communities of insect herbivores are thought to be structured mainly by indirect processes mediated by shared natural enemies, such as apparent competition. In host–parasitoid interaction networks, overlap in natural enemy communities between any pair of host species depends on the realized niches of parasitoids, which ultimately depend on the foraging decisions of individuals. Optimal foraging theory predicts that egg-limited parasitoid females should reject small hosts in favour of future opportunities to oviposit in larger hosts, while time-limited parasitoids are expected to optimize oviposition rate regardless of host size. The degree to which parasitoids are time- or egg-limited depends in part on weather conditions, as this determines the proportion of an individual''s lifespan that is available to foraging. Using a 10-year time series of monthly quantitative host–parasitoid webs, we present evidence for host-size-based electivity and sex allocation in the common secondary parasitoid Asaphes vulgaris. We argue that this electivity leads to body-size-dependent asymmetry in apparent competition among hosts and we discuss how changing weather patterns, as a result of climate change, may impact foraging behaviour and thereby the size-structure and dynamics of host–parasitoid indirect interaction networks.     

Nutritional ecology of the interaction between larvae of the gregarious ectoparasitoid, Muscidifurax raptorellus (Hymenoptera: Pteromalidae), and their pupal host, Musca domestica (Diptera: Muscidae)

In this study we examined the relationship between clutch size and parasitoid development of Muscidifurax raptorellus (Hymenoptera: Pteromalidae), a gregarious idiobiont attacking pupae of the housefly, Musca domestica (Diptera: Muscidae). Host quality was controlled in the experiments by presenting female parasitoids with hosts of similar size and age. This is the first study to monitor the development of a gregarious idiobiont parasitoid throughout the course of parasitism. Most female wasps laid clutches of one to four eggs per host, although some hosts contained eight or more parasitoid larvae. In both sexes, parasitoids completed development more rapidly, but emerging adult wasp size decreased as parasitoid load increased. Furthermore, the size variability of eclosing parasitoid siblings of the same sex increased with clutch size. Irrespective of clutch size, parasitoids began feeding and growing rapidly soon after eclosion from the egg and this continued until pupation. However, parasitoids in hosts containing five or more parasitoid larvae pupated one day earlier than hosts containing one to four larvae. The results are discussed in relation to adaptive patterns of host utilization by gregarious idiobiont and koinobiont parasitoids.     

Egg and time limitation mediate an egg protection strategy

The number of mature eggs remaining in the ovaries and the time left for oviposition determine the reproductive decisions of the hyperdiverse guild of insects that require discrete and potentially limiting resources for oviposition (such as seeds, fruits or other insects). A female may run out of eggs before all available oviposition sites are used (egg limitation), or die before using all of her eggs (time limitation). Females are predicted to change clutch size depending on whether eggs or time is the limiting resource. We extend this framework and ask whether the same constraints influence a strategy in which females modify eggs into protective shields. In response to egg parasitism cues, female seed beetles (Mimosestes amicus) lay eggs in vertical groups of 2–4, modifying the top 1–3 eggs into shields in order to protect the bottom egg from attack by parasitoids. We made contrasting predictions of how egg and time limitation would influence egg size and the incidence and level of egg protection. By varying access to seed pods, we manipulated the number of remaining eggs a female had at the time she received a parasitism cue. Although egg size was not affected, our results confirm that egg‐limited females protected fewer eggs and time‐limited females protected more eggs. Female body size explained the number of eggs in a stack rather than host deprivation or the timing of parasitoid exposure. Our results clearly show that host availability relative to female age influences the incidence of egg protection in M. amicus. Furthermore, our study represents a novel use of life history theory to explain patterns in an unusual but compelling defensive behaviour.     

Adaptive decision making or differential mortality: what causes offspring emergence in a gregarious parasitoid?

Hosts represent a limited resource for the developing offspring of parasitic insects laying eggs in or on spatially discrete resources like fruits, seeds, or other insects. The quality of hosts differs with respect to the value and amount of resources they provide for the feeding larvae. Accordingly, the size of a clutch of eggs laid on a given host should be a function of host quality, because severe competition between developing larvae can lead to increased mortality and/or decreased size of the offspring, both causing a fitness loss for the offspring and the mother. Therefore, females should be selected for the ability to estimate host quality and to adjust their clutch size accordingly. Using the parasitic wasp Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) this study investigated the respective contribution of developmental mortality of offspring vs. the clutch size decision of the mother as a determinant of final offspring emergence per host. In addition, taking offspring size into account, the study examined the fitness consequences of female oviposition decisions. Developmental mortality was very low in all quality classes of hosts except previously frozen and thus dead host pupae. Females laid reduced clutch sizes on dead, previously parasitized, and smaller hosts. In contrast to offspring number, offspring size did not differ between host qualities. We conclude that females are able to sense the quality of a host and adjust the number of eggs they lay to mitigate larval competition.     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non‐significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

Cuckoo females preferentially use specific habitats when searching for host nests

Egg mimicry is an important adaptation of common cuckoos, Cuculus canorus, against rejection of eggs by their respective hosts. A precondition for the maintenance of egg mimicry is that female cuckoos find hosts with a matching egg type. Experimental evidence indicated that habitat imprinting may be important for host selection. We tested whether the spacing and laying patterns of female cuckoos in the field are compatible with the supposed habitat-imprinting mechanism. We observed 16 females, with the help of radiotelemetry; of seven females, we observed directly 26 egg layings and 27 nest visits without laying. As expected if females were imprinted on different vegetation types, (1) the distribution of vegetation types differed between female home ranges, (2) female habitat use differed from average habitat availability within the egg-laying area (habitat preference), (3) females visited nests and deposited their eggs in the habitat they preferred, and (4) females laid their eggs consistently in a particular habitat type, irrespective of the host species. These results indicate that cuckoo females show habitat preference when searching for suitable host nests. Hence our data are compatible with the habitat-imprinting hypothesis, but owing to the habitat specificity of hosts the data cannot disprove a potential role of host specificity in cuckoo females.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Wasp behavior leads to uniform parasitism of a host available only a few hours per year

The parasitoid wasp, Hyposoter horticola, parasitizes a nearlyfixed fraction of its host butterfly larvae within a host metapopulationof 300–500 local populations in a 50 x 70-km area. Weshow, through laboratory observation, that the wasp lays eggsin fully developed larvae that have not yet hatched from theegg, constraining the period of host vulnerability to severalhours out of the host's one year lifecycle. The parasitoid achievesa persistent high rate of parasitism over the entire host rangedespite the extremely limited period of host vulnerability aswell as a high rate of host population extinctions and colonizationsof new habitat patches every year. It does this in part by beingextremely mobile. In addition, we show by using a field experimentand observation of marked wasps foraging for hosts in naturalpopulations, that the wasp finds virtually all host egg clustersin the weeks before the hosts become vulnerable to parasitism,and then later returns to parasitize them. By locating the hostsbefore their vulnerability, the wasp extends the time availablefor searching from hours to weeks. After parasitizing aboutone-third of the larvae in a host cluster the wasp stops, apparentlyleaving a mark that deters further parasitism by other individuals.The result of this novel combination of mobility and local foragingbehavior is a stable population size despite an unstable hostthat is vulnerable during about one thousandth of its lifecycle.     

Body size affects host defensive behavior and progeny fitness in a parasitoid wasp,Lysiphlebus fabarum

This study tested effects of maternal body size on foraging behavior and progeny development in a thelytokous population of Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae). Small and large wasps were reared from first and second instar hosts [black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae)], respectively, and each was provided with a patch (bean leaf disk) containing either 15 small (second instar) or 15 large (fourth instar) hosts for a 30‐min foraging period. Neither body size nor host size affected time allocation to various behaviors within a patch, but second instar aphids produced significantly more mummies than fourth instars. The preferred attack orientation was from the side of the aphid, suggesting wasps were sensitive to the risk of smearing with cornicle secretions. Few wasps developed in fourth instar hosts, suggesting later host instars were somewhat resistant to parasitism. Second instar hosts, the most suitable stage for L. fabarum development, relied more on defensive behavior, specifically kicking and secreting cornicle droplets. Large wasps were more likely to elicit a double cornicle secretion, indicating that aphids graded their response to the size of the attacker. Larger wasps were also more likely to be smeared with cornicle secretion, suggesting they were more vulnerable than small wasps. Although small wasps had smaller eggs than large wasps, there was no effect of maternal egg size on the size of progeny. However, daughters of small females emerged with larger egg loads than daughters of large mothers, and their eggs tended to be slightly smaller, although not significantly. Regression analysis revealed a positive correlation between maternal egg size and progeny developmental time for small and large wasps, and between maternal egg size and progeny egg load for small wasps. These results confirm maternal effects of body size in an aphid parasitoid, and reveal that vulnerability to host behavioral defenses is also body size dependent.