Increasing host age does not have the expected negative effects on the fitness parameters of an egg parasitoid

Host age is an important determinant of host acceptance and suitability for egg parasitoids. As host embryonic development advances, the quality of resources available to the parasitoid offspring typically declines, usually resulting in reduced acceptance levels by foraging females and lower offspring fitness. We examined the ability of the parasitoid Telenomus podisi Ashmead (Hymenoptera: Scelionidae) to parasitize and develop in Podisus maculiventris (Say) (Hemiptera: Pentatomidae) eggs of different ages. In laboratory experiments, we measured the effect of host age (6, 24, 48, 72, 96, or 120 h old) on parasitism rate and offspring fitness parameters such as survival, development time, sex ratio, and size. Contrary to our expectations, parasitism rate did not differ between host age treatments, nor did sex ratio allocation, offspring size, or the fecundity of newly emerged female offspring. However, parasitoid offspring had a longer development time with increasing host age. This trend was stronger for males than for females, which we suggest could reduce the degree of protandry among offspring emerging from older host eggs, thus increasing the rate of virginity upon leaving the emergence patch and resulting in more frequent off‐patch mating by female offspring in nature. Overall, our results suggest that all stages of P. maculiventris embryonic development are suitable for acceptance and development of T. podisi. Unlike most species of egg parasitoids, T. podisi has evolved mechanisms to utilize host resources, regardless of host developmental stage, with relatively minor fitness consequences.     

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.     

Patterns of parasitoid host utilization and development across a range of temperatures: implications for biological control of an invasive forest pest

Although climate change frequently has been linked to observed shifts in the distributions or phenologies of species, little is known about the potential effects of varying temperatures on parasitoids and their relationships with hosts. Using the egg parasitoid Oobius agrili (Hymenoptera: Encyrtidae) we examined host utilization patterns of this species across a range of temperatures (20–35 °C) to explore how changing climate could affect the interaction with its host—the emerald ash borer (EAB) (Coleoptera: Buprestidae), a serious invasive forest pest that has killed tens of millions of ash (Fraxinus spp.) trees in North America. Results from our study showed that the window of host susceptibility to O. agrili parasitism declined significantly from 14.8 to 2.6 days in an inverse second-order relationship with increasing exposure temperatures from 20 to 35 °C. In contrast, parasitoid host attack rate changed in a bell-shaped second-order relationship—i.e., increased with temperatures from 20 to 25 °C, but decreased at about the same rate when temperatures increased from 30 to 35 °C. This range of temperatures also significantly affected the development and mortality of immature parasitoids with 35 °C resulting in 100 % mortality. There was little mortality (0–4.5 %) and no significant differences in the percentage (20.9–34.9 %) of immature O. agrili that entered diapause (as mature larvae) at 20, 25, and 30 °C. However, there were significant differences in the time event of adult wasp emergence within this temperature range. The median time for 50 % of immature O. agrili emerging as adults at 20, 25, and 30 °C were 38, 18, and 17 days after parental wasp oviposition, respectively. Together these findings indicate that the non-linear and unequal temperature effects on these host utilization parameters are likely to result in differential host parasitism rates, and thus could reduce the efficacy of this parasitoid in suppressing host populations due to climate change (global warming and extreme heat).     

Genotype‐by‐genotype specificity remains robust to average temperature variation in an aphid/endosymbiont/parasitoid system

Genotype‐by‐genotype interactions demonstrate the existence of variation upon which selection acts in host–parasite systems at respective resistance and infection loci. These interactions can potentially be modified by environmental factors, which would entail that different genotypes are selected under different environmental conditions. In the current study, we checked for a G × G × E interaction in the context of average temperature and the genotypes of asexual lines of the endoparasitoid wasp Lysiphlebus fabarum and isolates of Hamiltonella defensa, a protective secondary endosymbiont of the wasp's host, the black bean aphid Aphis fabae. We exposed genetically identical aphids harbouring different isolates of H. defensa to three asexual lines of the parasitoid and measured parasitism success under three different temperatures (15, 22 and 29 °C). Although there was clear evidence for increased susceptibility to parasitoids at the highest average temperature and a strong G × G interaction between the host's symbionts and the parasitoids, no modifying effect of temperature, that is, no significant G × G × E interaction, was detected. This robustness of the observed specificity suggests that the relative fitness of different parasitoid genotypes on hosts protected by particular symbionts remains uncomplicated by spatial or temporal variation in temperature, which should facilitate biological control strategies.     

Population responses of hymenopteran parasitoids to the emerald ash borer (Coleoptera: Buprestidae) in recently invaded areas in north central United States

Populations of hymenopteran parasitoids associated with larval stages of the invasive emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) were surveyed in 2009 and 2010 in the recently invaded areas in north central United States (Michigan), where two introduced EAB larval parasitoids, Tetrastichus planipennisi Yang and Spathius agrili Yang were released for classical biological control. Results from two years of field surveys showed that several hymenopteran parasitoids have become associated with EAB in Michigan. Among these parasitoids, the gregarious species T. planipennisi was the most abundant, accounting for 93% of all parasitoid individuals collected in 2009 (immediately after field release) and 58% in 2010 (a year later after field releases). Low levels (1–5%) of parasitism of EAB larvae by T. planipennisi were consistently detected at survey sites in both years. Separately, the abundance of the native parasitoid, Atanycolus spp., increased sharply, resulting in an average parasitism rate of EAB larvae from <0.5% in 2009 to 19% in 2010. Other parasitoids such as Phasgonophora sulcata Westwood, Spathius spp., Balcha indica Mani & Kaul, Eupelmus sp., and Eurytomus sp. were much less abundant than T. planipennisi and Atanycolus spp., and each caused <1% parasitism. Besides hymenopteran parasitoids, woodpeckers consumed 32–42% of the immature EAB stages present at our study sites, while undetermined biotic factors (such as microbial disease and host tree resistance) caused 10–22% mortality of observed EAB larvae. Relevance of these findings to the potential for biological control of EAB in the invaded areas of North America is discussed.     

Egg load dynamics and the risk of egg and time limitation experienced by an aphid parasitoid in the field

Insect parasitoids and herbivores must balance the risk of egg limitation and time limitation in order to maximize reproductive success. Egg and time limitation are mediated by oviposition and egg maturation rates as well as by starvation risk and other determinants of adult lifespan. Here, we assessed egg load and nutritional state in the soybean aphid parasitoid Binodoxys communis under field conditions to estimate its risk of becoming either egg‐ or time‐limited. The majority of female B. communis showed no signs of egg limitation. Experimental field manipulations of B. communis females suggested that an average of 4–8 eggs were matured per hour over the course of a day. Regardless, egg loads remained constant over the course of the day at approximately 80 eggs, suggesting that egg maturation compensates for oviposition. This is the first case of such “egg load buffering” documented for a parasitoid in the field. Despite this buffering, egg loads dropped slightly with increasing host (aphid) density. This suggests that egg limitation could occur at very high host densities as experienced in outbreak years in some locations in the Midwestern USA. Biochemical analyses of sugar profiles showed that parasitoids fed upon sugar in the field at a remarkably high rate. Time limitation through starvation thus seems to be very low and aphid honeydew is most likely a source of dietary sugar for these parasitoids. This latter supposition is supported by the fact that body sugar levels increase with host (aphid) density. Together, these results suggest that fecundity of B. communis benefits from both dynamic egg maturation strategies and sugar‐feeding.     

Strong specificity in the interaction between parasitoids and symbiont‐protected hosts

Coevolution between hosts and parasites may promote the maintenance of genetic variation in both antagonists by negative frequency‐dependence if the host–parasite interaction is genotype‐specific. Here we tested for specificity in the interaction between parasitoids (Lysiphlebus fabarum) and aphid hosts (Aphis fabae) that are protected by a heritable defensive endosymbiont, the γ‐proteobacterium Hamiltonella defensa. Previous studies reported a lack of genotype specificity between unprotected aphids and parasitoids, but suggested that symbiont‐conferred resistance might exhibit a higher degree of specificity. Indeed, in addition to ample variation in host resistance as well as parasitoid infectivity, we found a strong aphid clone‐by‐parasitoid line interaction on the rates of successful parasitism. This genotype specificity appears to be mediated by H. defensa, highlighting the important role that endosymbionts can play in host–parasite coevolution.     

Nectar feeding increases exploratory behaviour in the aphid parasitoid Diaeretiella rapae (McIntosh)

Feeding on floral nectar has multiple positive effects on parasitic wasps, including increased longevity and fecundity, and in addition, nectar feeding can also alter parasitoid behaviour. To advance understanding of the effects of nectar feeding on Diaeretiella rapae (McIntosh) [Hymenoptera: Braconidae], the activities of 1‐day‐old female D. rapae with or without a prior buckwheat (Fagopyrum esculentum) nectar meal were quantified. Nectar increased searching time of D. rapae by a factor of 40 compared with individuals provided with water only and reduced the time spent stationary. The number of attacks to aphids by nectar‐fed parasitoids was not significantly (P = 0.06) higher than that of unfed D. rapae, suggesting that the conditions of the experiment facilitated host finding by ‘quiet’ parasitoids. Nevertheless, nectar feeding modified the behaviour of D. rapae in a way that parasitoids were more explorative and less inactive. This represents one additional mechanism through which nectar feeding impacts parasitoid biology and suggests that a synergy between increased host searching, increased longevity and increased fecundity should lead to an enhancement of biocontrol when D. rapae females have access to nectar in the field.     

An improved method for monitoring parasitism and establishment of Oobius agrili (Hymenoptera: Encyrtidae), an egg parasitoid introduced for biological control of the emerald ash borer (Coleoptera: Buprestidae) in North America

Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) is a solitary egg parasitoid that has been released in the United States since 2007 for biocontrol of the invasive emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). Field and laboratory trials with ash logs infested with EAB eggs were conducted in Michigan between 2009 and 2010 to improve methods for monitoring the establishment of O. agrili. Naturally occurring EAB eggs were collected in both parasitoid-release and control (non-parasitoid-release) plots to compare with the EAB egg-sentinel log (ESL) technique. In three parasitoid-release plots, >50% of ESLs had O. agrili-parasitized eggs ranging from 3.9% to 48.2% egg parasitism after one week of field exposure. No EAB eggs were attacked by O. agrili on the ESLs deployed in control plots. In the laboratory, 100% of ESLs exposed to O. agrili inside rearing jars for one week had parasitized-eggs (68.5% egg parasitism). Deployment of ESLs detected low levels of parasitism by O. agrili in all three ash stands where O. agrili was released in previous years. In contrast, collection of naturally occurring EAB eggs detected the parasitism in only one of these three parasitoid-release ash stands. No parasitism was detected in control ash stands with either method. These findings indicate that populations of O. agrili released in previous years had successfully overwintered and established in the released ash stands by 2010, but had not yet dispersed to the control stands.     

Development of methods for the field evaluation of Oobius agrili (Hymenoptera: Encyrtidae) in North America,a newly introduced egg parasitoid of the emerald ash borer (Coleoptera: Buprestidae)

A field study was conducted in forested plots near Lansing, Michigan in 2008 and 2009 to evaluate the newly introduced egg parasitoid Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) for control of the invasive emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). To measure parasitism by O. agrili, laboratory-reared “sentinel EAB eggs” were deployed under bark flaps on trunks of selected ash trees in both parasitoid-release and non-release control plots. In addition, naturally occurring EAB eggs were collected in both parasitoid-release and control plots to measure parasitism. While no parasitism was detected with either sentinel or naturally occurring EAB eggs in control plots in either 2008 or 2009, a low level of parasitism by O. agrili was detected in the parasitoid-release plots in both artificially deployed sentinel eggs (?1%) and field-collected, naturally occurring eggs (1.1–4.2%) in both years. In addition to losses due to parasitism by O. agrili, a large proportion (37–52%) of the field-deployed sentinel eggs disappeared, possibly due to predators such as ants, in both parasitoid-release and control plots. While no statistical differences in parasitism by O. agrili were detected between parasitoid release and control plots, other sources of egg mortality such as disappearance due to predation on eggs, varied significantly across study sites in both 2008 and 2009. The relevance of these findings to future release and evaluation strategies for O. agrili for biological control of the invasive emerald ash borer in the US is discussed.     

Partial bivoltinism in a gregarious endoparasitoid: larval diapause as influenced by season and sharing a host

The yearly timing of the life cycle of a parasitoid is a key element of its life‐history strategy. I examine here factors influencing the expression of partial bivoltinism in Tetrastichus julis Walker (Hymenoptera: Eulophidae), a specialist parasitoid introduced to North America to attack its univoltine host, the cereal leaf beetle, Oulema melanopus (L.) (Coleoptera: Chrysomelidae). The varying tendency was assessed of individuals of this gregarious larval parasitoid to either emerge as adults in the same summer they mature, or to enter diapause to emerge the following year. Parasitized hosts were obtained by rearing cereal leaf beetles collected as mature larvae from grain fields in northern Utah (western USA) throughout the growing seasons in 2013 and 2014. Cocoons spun by these beetles were held to determine patterns over the spring and summer in the tendency of the parasitoid to forgo larval diapause. A high percentage (nearly 90% in 2013) of parasitoid individuals were found to forgo diapause and emerge in the same summer from earliest maturing hosts. This percentage rapidly declined to 20% or less of individuals forgoing diapause and emerging from cocoons as the summer advanced. The percentage of parasitoid individuals forgoing diapause increased significantly at a given time of season (early or late) as the number of conspecifics with which an individual shared a host larva increased. These results may reflect a trade‐off for individual parasitoids in which greater success in finding – and ovipositing in – host larvae the following spring vs. in summer, is countered by reduced survivorship in diapausing over the winter vs. emerging in the same summer in which the parasitoid matures. Expression of partial bivoltinism of T. julis, as affected strongly by both season and within‐host density, results in high rates of parasitism of cereal leaf beetles both early and late in the season.     

The roles of foraging environment,host species,and host diet for a generalist pupal parasitoid

Even for parasitoids with a wide host range, not all host species are equally suitable, and host quality often depends on the plant the host feeds on. We compared oviposition choice and offspring performance of a generalist pupal parasitoid, Pteromalus apum (Retzius) (Hymenoptera: Pteromalidae), on two congeneric hosts reared on two plant species under field and laboratory conditions. The plants contain defensive iridoid glycosides that are sequestered by the hosts. Sequestration at the pupal stage differed little between host species and, although the concentrations of iridoid glycosides in the two plant species differ, there was no effect of diet on the sequestration by host pupae. The rate of successful parasitism differed between host species, depending on the conditions they were presented in. In the field, where plant‐associated cues are present, the parasitoid used Melitaea cinxia (L.) over Melitaea athalia (Rottemburg) (Lepidoptera: Nymphalidae), whereas more M. athalia were parasitised in simplified laboratory conditions. In the field, brood size, which is partially determined by rate of superparasitism, depended on both host and plant species. There was little variation in other aspects of offspring performance related to host or plant species, indicating that the two host plants are of equal quality for the hosts, and the hosts are of equal quality for the parasitoids. Corresponding to this, we found no evidence for associative learning by the parasitoid based on their natal host, so with respect to these host species they are truly generalist in their foraging behaviour.     

Varying degree of physiological integration among host instars and their endoparasitoid affects stress‐induced mortality

In natural populations of insect herbivores, genetic differentiation is likely to occur due to variation in host plant utilization and selection by the local community of organisms with which they interact. In parasitoids, engaging in intimate associations with their host during immature development, local variation may exist in host quality for parasitoid development. We compared the development of a gregarious endoparasitoid, Cotesia glomerata L. (Hymenoptera: Braconidae), collected in The Netherlands, in three strains and three caterpillar instars (L1–L3) of its main host, Pieris brassicae L. (Lepidoptera: Pieridae). Hosts had been collected in The Netherlands and France, and were reared in the laboratory for one generation. We also used an established Dutch laboratory strain that had not been exposed to parasitoids for at least 24 generations. Parasitoid survival to adulthood was inversely correlated with host instar at parasitism. Adult parasitoid body mass was largest when hosts were parasitized as L1 and smallest when hosts were parasitized as L3, whereas egg‐to‐adult development time was quickest on L3 hosts and slowest on L1 hosts. Higher survival and faster development of C. glomerata on French L2 hosts also showed that there is variation in host‐instar‐related suitability. Many L2 and most L3 caterpillars that were parasitized exhibited signs of pathogen infection and perished within a few days of parasitism, whereas this never happened when hosts were parasitized as L1 or in non‐parasitized control caterpillars. Our results reveal that, irrespective of the host strain, L1 hosts are optimally synchronized with C. glomerata development. By contrast, the high precocious mortality of L3 larvae may be due to stress‐induced regulation by the parasitoid in order to ‘force’ its developmental program into synchrony with the developing parasitoid larvae. Our results underscore a potentially important role played by pathogens in mediating herbivore–parasitoid interactions that are host‐instar‐dependent in their expression.     

Long-term monitoring of the introduced emerald ash borer (Coleoptera: Buprestidae) egg parasitoid,Oobius agrili (Hymenoptera: Encyrtidae), in Michigan,USA and evaluation of a newly developed monitoring technique

Emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a serious invasive pest of ash trees (Fraxinus spp.) in North America from China. The egg parasitoid Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) was introduced from China as a biological control agent for this pest in Michigan and throughout the infested area of the United States. A critical component of any biological control program is post-release monitoring and evaluation; however, because of the small size and cryptic nature of O. agrili, evaluation of its impact is difficult. We compared two methods for measuring parasitism of emerald ash borer eggs: (1) timed visual searches of bark on standing ash trees and (2) bark collection, sifting, and sorting. Both methods were carried out in paired parasitoid-release and control plots, the visual search method over a six-year period (2008–2013) and the more recently developed bark-collection and sifting method for 2 years (2012–2013). The visual search method found parasitism in release plots remained low (0.7–4.2%) in samples taken from 2008 to 2012 and reached 10.6% in 2013. In comparison, the bark-sifting method found that rates of egg parasitism were considerably higher in release plots, 21.8% and 18.9% for samples taken in 2012 and 2013, respectively. These findings indicate that the population-level impact of O. agrili is increasing and may be an important source of mortality for EAB populations. We recommend the bark-collection and sifting method as the more effective method to recover parasitoids and estimate parasitism rates of O. agrili.     

Herbivore egg deposition induces tea leaves to arrest the egg‐larval parasitoid Ascogaster reticulata

Plants are able to activate direct and indirect defences against egg deposition by herbivorous insects. A known indirect defence is the production of synomones to help egg‐ and egg‐larval parasitoids to locate their hosts. The wasp Ascogaster reticulata Watanabe (Hymenoptera: Braconidae) is a solitary egg‐larval parasitoid of the moth Adoxophyes honmai Yasuda (Lepidoptera: Tortricidae), which lays eggs and feeds as caterpillars on the leaves of the tea plant Camellia sinensis (L.) Kuntze (Theaceae). Here, we studied whether or not oviposition by A. honmai induces tea plants to produce synomones that help the parasitoid to locate its host. An olfactometer bioassay suggested that synomones produced by the infested plants did not attract the parasitoid over a short range. However, a contact bioassay showed that tea leaves were induced to arrest the parasitoid 24 h after egg deposition and remained induced until the host‐egg masses were no more attractive to the parasitoids. Wing scales and deposits of adult moths and the contents of the egg masses did not induce the tea leaves to arrest the parasitoid, but the contents of the female moth's reproductive system did. Synomone induction was systemic: uninfested leaves in the vicinity of egg‐laden leaves also arrested the parasitoid.     

Effect of adult feeding and timing of host exposure on the fertility and longevity of the parasitoid Anaphes flavipes

The parasitoid Anaphes flavipes (Foerster) (Hymenoptera: Mymaridae) is a gregarious egg parasitoid which is widely used in biological control against important crop pest beetles of the genus Oulema (Coleoptera: Chrysomelidae). Here, we present the first experimental examination of the influence of adult feeding and timing of host exposure on the longevity and fertility of this parasitoid. We confirmed a positive effect of adult feeding on longevity of both sexes. Fed parasitoids lived 3× longer than unfed ones. On the other hand, adult feeding and feeding time had no effect on female fertility. The number of hatched offspring was not increased by adult feeding, which suggests that the parasitoid emerges with already mature ovaries (proovigenic type). However, the fertility of fed females was strongly influenced by the timing of host egg exposure. By providing distinct groups of parasitoids with host eggs at different times, we were able to show lower fertility of fed females that had been offered host eggs more than 24 h after hatching. Our results thus show that the parasitoid's fertility is determined by her age at the time of parasitization rather than by feeding.     

Food web associations and effect of trophic resources and environmental factors on parasitoids expanding their host range into non‐native hosts

Trophic interactions and environmental conditions determine the structure of food webs and the host expansion of parasitoids into novel insect hosts. In this study, we investigate plant–insect–parasitoid food web interactions, specifically the effect of trophic resources and environmental factors on the presence of the parasitoids expanding their host range after the invasion of Chrysodeixis chalcites (Esper) (Lepidoptera: Noctuidae). We also consider potential candidates for biological control of this non‐native pest. A survey of larval stages of Plusiinae (Lepidoptera: Noctuidae) and their larval parasitoids was conducted in field and vegetable greenhouse crops in 2009 and 2010 in various locations of Essex and Chatham‐Kent counties in Ontario, Canada. Twenty‐one plant–host insect–host parasitoid associations were observed among Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae), C. chalcites, and larval parasitoids in three trophic levels of interaction. Chrysodeixis chalcites, an old‐world species that had just arrived in the region, was the most common in our samples. The larval parasitoids Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae), Cotesia vanessae (Reinhard), Cotesia sp., Microplitis alaskensis (Ashmead), and Meteorus rubens (Nees) (all Hymenoptera: Braconidae) expanded their host range into C. chalcites changing the structure of the food web. Copidosoma floridanum (Ashmead) (Hymenoptera: Encyrtidae) was the most common parasitoid of T. ni that was not found in the invasive species. Plant species, host abundance, and agro‐ecosystem were the most common predictors for the presence of the parasitoids expanding their host range into C. chalcites. Our results indicate that C. sonorensis, C. vanessae, and C. floridanum should be evaluated for their potential use in biological control of C. chalcites and T. ni.     

Effects of seasonal variation in host quality and availability on parasitism by the egg parasitoid Telenomus coloradensis

Hemlock looper, Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae), is one of the most important defoliator in North American forests. The common egg parasitoid Telenomus coloradensis Crawford (Hymenoptera: Platygastridae) plays a significant role as a natural control agent, with parasitism levels in spring typically higher than in fall. The objectives of this study were to quantify changes in host acceptance and reproductive performance of the parasitoid in relation to (1) host egg fertilization in fall, (2) host diapause status, (3) host embryonic development in spring, and (4) host deprivation during summer. Our results indicate that T. coloradensis do not have the capacity to develop in unfertilized host eggs, whereas early‐diapausing eggs are more suitable for the parasitoid than post‐diapausing eggs. Furthermore, the host physiological suitability decreases with embryonic development in spring. Finally, a host deprivation period during the summer tends to negatively affect the parasitic activity of T. coloradensis. These laboratory results confirm previous hypotheses concerning T. coloradensis seasonal ecology and contribute to a better understanding of the effect of hemlock looper egg physiology and availability on the reproductive potential of T. coloradensis.     

Testing for host‐associated differentiation in two egg parasitoids of a forest herbivore

Several studies underline the importance of ecological barriers and differential selection in driving sympatric speciation. Host‐associated differentiation (HAD) has been proposed as one of the mechanisms leading to sympatric speciation. However, it is still unclear how common HAD is or which are the factors that could promote it. In particular, not much is known about HAD in predators and parasitoids of herbivorous insects. One of the characteristics postulated to pre‐dispose insects to HAD is parthenogenesis as it may favour adaptive responses to particular environments, amplifying selected gene complexes. In this study, we used amplified fragment length polymorphism (AFLP) markers to determine whether HAD is present in two parthenogenetic egg parasitoids attacking the same herbivore species – the pine processionary moth, Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera: Notodontidae) – on two host Pinus species. A total of 100 loci for 59 individuals sampled in four populations of Baryscapus servadeii (Domenichini) (Hymenoptera: Eulophidae), a specialist parasitoid, and 106 loci for 117 individuals sampled in six populations of Ooencyrtus pityocampae Mercet (Hymenoptera: Encyrtidae), a generalist parasitoid, were analysed. Levels of genetic differentiation were also assessed with an outlier analysis, checking for alleles associated to host plants. No evidence of HAD was detected in any of the two parasitoid species. We hypothesize that both the lack of strict parthenogenetic reproduction and the ectophagous nature of the insect host could explain the absence of HAD. The genetic variation observed in the generalist parasitoid responded to a pattern of local adaptation, whereas no relationship with either host or geography was found in the specialist parasitoid.     

Native parasitoids and recovery of Spathius agrili from areas of release against emerald ash borer in eastern Tennessee,USA

The parasitoid Spathius agrili Yang, introduced in the USA to suppress populations of the emerald ash borer (EAB), Agrilus planipennis Fairmaire, has been recovered at a release site for the first time in eastern Tennessee after a single year of releases. Other native parasitoids, including Spathius floridanus Ashmead, undetermined species of Spathius (possibly Spathius elegans Matthews and Spathius parvulus Matthews) and Atanycolus cappaerti Marsh & Strazanac, also known to be associated with EAB, were recovered. These recoveries represent the first documentation of these four species, including the introduced S. agrili, associated with EAB in the southern USA. Implications for biological control efforts against EAB are discussed.