Effects of quantitative and qualitative differences in volatiles from host‐ and non‐host‐infested maize on the attraction of the larval parasitoid Cotesia kariyai

Cotesia kariyai Watanabe (Hymenoptera: Braconidae) is a specialist larval parasitoid of Mythimna separata Walker (Lepidoptera: Noctuidae). Cotesia kariyai wasps use herbivore‐induced plant volatiles (HIPVs) to locate hosts. However, complex natural habitats are full of volatiles released by both herbivorous host‐ and non‐host‐infested plants at various levels of intensity. Therefore, the presence of non‐hosts may affect parasitoid decisions while foraging. Here, the host‐finding efficiency of naive C. kariyai from HIPVs influenced by host‐ and non‐host‐infested maize [Zea mays L. (Poaceae)] plants was investigated with a four‐arm olfactometer. Ostrinia furnacalis Guenée (Lepidoptera: Crambidae) was selected as a non‐host species. One unit (1 U) of host‐ or non‐host‐infested plant was prepared by infesting a potted plant with five host or seven non‐host larvae. In two‐choice bioassays, host‐infested plants fed upon by different numbers of larvae, and various units of host‐ and non‐host‐infested plants (infestation units; 1 U, 2 U, and 3 U) were arranged to examine the effects of differences in volatile quantity and quality on the olfactory responses of C. kariyai with the assumption that volatile quantity and quality changes with differences in numbers of insects and plants. Cotesia kariyai was found to perceive quantitative differences in volatiles from host‐infested plants, preferring larger quantities of volatiles from larger numbers of larvae or plants. Also, the parasitoids discriminated between healthy plants, host‐infested plants, and non‐host‐infested plants by recognising volatiles released from those plants. Cotesia kariyai showed a reduced preference for host‐induced volatiles, when larger numbers of non‐host‐infested plants were present. Therefore, quantitative and qualitative differences in volatiles from host‐ and non‐host‐infested plants appear to affect the decision of C. kariyai during host‐habitat searching in multiple tritrophic systems.     

Olfactory cues explain differential attraction of the striped rice stem borer to different varieties of rice plant

In Iranian rice fields, different varieties of rice are cultivated which are differentially impacted by females of the rice stem borer, Chilo suppressalis. To elucidate the role odours may play in their host plant finding behaviour, female Ch. suppressalis were exposed to four varieties of rice plants and their volatiles in a four‐arm olfactometer. In whole plant tests, Ch. suppressalis were significantly attracted to the variety previously characterized as most susceptible, least attracted to one characterized as semisusceptible, and showed no attraction to those varieties characterized as semi‐ and highly resistant. Tests using headspace volatile extracts yielded similar results in the case of the most susceptible variety, but showed no attraction to the semisusceptible and highly resistant varieties, and low attraction to the semiresistant variety. Subsequent analysis of the volatile composition identified a panel of 27 components, some of which were either unique to, or abundantly present in, particular varieties, and may explain the observed variation in their attractiveness. Our findings show that rice plant volatiles can play a role in the host selection behaviour of this pest species, and we suggest compounds which may be important to this process and the future application of volatiles in rice pest management programs.     

寄主利它素对稻螟赤眼蜂寄生行为的影响

对稻螟赤眼蜂Trichogrammajaponicum(Ashmead)在寄主定位过程中所利用的寄主利它素的存在部位、作用方式及其应用效果作了初步研究。生测结果表明 :寄主利它素存在于三化螟Tryporyzaincertu las(Walker)卵块盖毛和成虫鳞片中 ,其中以盖毛中利它素含量最高 ,雌蛾鳞片次之 ,雄蛾鳞片含量最少。结果还表明盖毛在卵块上的完整性对稻螟赤眼蜂的寄主搜索行为有明显影响。室内应用效果表明利它素可以使稻螟赤眼蜂对寄主卵寄生率提高 1 0 %以上。     

Role of volatiles emitted by host and non-host plants in the foraging behaviour of Dentichasmias busseolae, a pupal parasitoid of the spotted stemborer Chilo partellus

The role of volatiles from stemborer host and non‐host plants in the host‐finding process of Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae) a pupal parasitoid of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) was studied. The non‐host plant, molasses grass (Melinis minutiflora Beauv. (Poaceae)), is reported to produce some volatile compounds known to be attractive to some parasitoid species. The studies were conducted to explore the possibility of intercropping stemborer host plants with molasses grass in order to enhance the foraging activity of D. busseolae in such a diversified agro‐ecosystem. Olfactometric bioassays showed that volatiles from the host plants maize, Zea mays L., and sorghum, Sorghum bicolor (L.) (Poaceae), were attractive to the parasitoid. Infested host plants were the most attractive. Volatiles from molasses grass were repellent to the parasitoid. Further tests showed that volatiles from infested and uninfested host plants alone were preferred over those from infested and uninfested host plants combined with the non‐host plant, molasses grass. In dual choice tests, the parasitoid did not discriminate between volatiles from maize infested by either of the two herbivore species, C. partellus or Busseola fusca Fuller (Lepidoptera: Noctuidae). Volatiles from sorghum infested by C. partellus were preferred over those from C. partellus‐infested maize. The study showed that the pupal parasitoid D. busseolae uses plant volatiles during foraging, with those from the plant–herbivore complex being the most attractive. The fact that volatiles from molasses grass were deterrent to the parasitoid suggested that intercropping maize or sorghum with molasses grass was not likely to enhance the foraging behaviour of D. busseolae. Volatiles from the molasses grass may hinder D. busseolae's host location efficiency.     

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.     

Trichobaris weevils distinguish amongst toxic host plants by sensing volatiles that do not affect larval performance

Herbivorous insects use plant metabolites to inform their host plant selection for oviposition. These host‐selection behaviours are often consistent with the preference–performance hypothesis; females oviposit on hosts that maximize the performance of their offspring. However, the metabolites used for these oviposition choices and those responsible for differences in offspring performance remain unknown for ecologically relevant interactions. Here, we examined the host‐selection behaviours of two sympatric weevils, the Datura (Trichobaris compacta) and tobacco (T. mucorea) weevils in field and glasshouse experiments with transgenic host plants specifically altered in different components of their secondary metabolism. Adult females of both species strongly preferred to feed on D. wrightii rather than on N. attenuata leaves, but T. mucorea preferred to oviposit on N. attenuata, while T. compacta oviposited only on D. wrightii. These oviposition behaviours increased offspring performance: T. compacta larvae only survived in D. wrightii stems and T. mucorea larvae survived better in N. attenuata than in D. wrightii stems. Choice assays with nicotine‐free, JA‐impaired, and sesquiterpene‐over‐produced isogenic N. attenuata plants revealed that although half of the T. compacta larvae survived in nicotine‐free N. attenuata lines, nicotine did not influence the oviposition behaviours of both the nicotine‐adapted and nicotine‐sensitive species. JA‐induced sesquiterpene volatiles are key compounds influencing T. mucorea females’ oviposition choices, but these sesquiterpenes had no effect on larval performance. We conclude that adult females are able to choose the best host plant for their offspring and use chemicals different from those that influence larval performance to inform their oviposition decisions.     

The Response Specificity of <Emphasis Type="Italic">Trichogramma</Emphasis> Egg Parasitoids towards Infochemicals during Host Location

Parasitoids are confronted with many different infochemicals of their hosts and food plants during host selection. Here, we investigated the effect of kairomones from the adult host Pieris brassicae and of cues present on Brussels sprout plants infested by P. brassicae eggs on the behavioral response of the egg parasitoid Trichogramma evanescens. Additionally, we tested whether the parasitoid’s acceptance of P. brassicae eggs changes with different host ages. The wasps did not discriminate between olfactory cues from mated and virgin females or between mated females and males of P. brassicae. T. evanescens randomly climbed on the butterflies, showing a phoretic behavior without any preference for a certain sex. The parasitoid was arrested on leaf parts next to 1-day-old host egg masses. This arrestment might be due to cues deposited during oviposition. The wasps parasitized host eggs up to 3 days old equally well. Our results were compared with former studies on responses by T .brassicae showing that T. evanescens makes less use of infochemicals from P. brassicae than T. brassicae.     

Histories of host shifts and cospeciation among free‐living parasitoids of Rhagoletis flies

Host shifts by specialist insects can lead to reproductive isolation between insect populations that use different hosts, promoting diversification. When both a phytophagous insect and its ancestrally associated parasitoid shift to the same novel host plant, they may cospeciate. However, because adult parasitoids are free living, they can also colonize novel host insects and diversify independent of their ancestral host insect. Although shifts of parasitoids to new insect hosts have been documented in ecological time, the long‐term importance of such shifts to parasitoid diversity has not been evaluated. We used a genus of flies with a history of speciation via host shifting (Rhagoletis [Diptera: Tephritidae]) and three associated hymenopteran parasitoid genera (Diachasma, Coptera and Utetes) to examine cophylogenetic relationships between parasitoids and their host insects. We inferred phylogenies of Rhagoletis, Diachasma, Coptera and Utetes and used distance‐based cophylogenetic methods (ParaFit and PACo) to assess congruence between fly and parasitoid trees. We used an event‐based method with a free‐living parasitoid cost model to reconstruct cophylogenetic histories of each parasitoid genus and Rhagoletis. We found that the current species diversity and host–parasitoid associations between the Rhagoletis flies and parasitoids are the primary result of ancient cospeciation events. Parasitoid shifts to ancestrally unrelated hosts primarily occur near the branch tips, suggesting that host shifts contribute to recent parasitoid species diversity but that these lineages may not persist over longer time periods. Our analyses also stress the importance of biologically informed cost models when investigating the coevolutionary histories of hosts and free‐living parasitoids.     

Role of volatile semiochemicals in host location by the egg parasitoid Anagrus breviphragma

Recent investigations conducted on several tritrophic systems have demonstrated that egg parasitoids, when searching for host eggs, may exploit plant synomones that have been induced as a consequence of host oviposition. In this article we show that, in a system characterized by host eggs embedded in the plant tissue, naïve females of the egg parasitoid Anagrus breviphragma Soyka (Hymenoptera: Mymaridae) responded in a Y‐tube olfactometer to volatiles from leaves of Carex riparia Curtis (Cyperaceae) containing eggs of one of its hosts, Cicadella viridis (L.) (Hemiptera: Cicadellidae). The wasp did not respond to host eggs or to clean leaves from non‐infested plants compared with clean air, whereas it showed a strong preference for the olfactometer arm containing volatiles of leaves with embedded host eggs, compared with the arm containing volatiles of leaves from a non‐infested plant or host eggs extracted from the plant. When the eggs were removed from an infested leaf, the parasitoid preference was observed only if eggs were added aside, suggesting a synergistic effect of a local plant synomone and an egg kairomone. The parasitoid also responded to clean leaves from an egg‐infested plant when compared with leaves from a non‐infested plant, indicating a systemic effect of volatile induction.     

Associative learning of host presence in non‐host environments influences parasitoid foraging

1. Parasitoids are known to utilise learning of herbivore‐induced plant volatiles (HIPVs) when foraging for their herbivorous host. In natural situations these hosts share food plants with other, non‐suitable herbivores (non‐hosts). Simultaneous infestation of plants by hosts and non‐hosts has been found to result in induction of HIPVs that differ from host‐infested plants. Each non‐host herbivore may have different effects on HIPVs when sharing the food plant with hosts, and thus parasitoids may learn that plants with a specific non‐host herbivore also contain the host. 2. This study investigated the adaptive nature of learning by a foraging parasitoid that had acquired oviposition experience on a plant infested with both hosts and different non‐hosts in the laboratory and in semi‐field experiments. 3. In two‐choice preference tests, the parasitoid Cotesia glomerata shifted its preference towards HIPVs of a plant–host–non‐host complex previously associated with an oviposition experience. It could, indeed, learn that the presence of its host is associated with HIPVs induced by simultaneous feeding of its host Pieris brassicae and either the non‐host caterpillar Mamestra brassicae or the non‐host aphid Myzus persicae. However, the learned preference found in the laboratory did not translate into parasitisation preferences for hosts accompanying non‐host caterpillars or aphids in a semi‐field situation. 4. This paper discusses the importance of learning in parasitoid foraging, and debates why observed learned preferences for HIPVs in the laboratory may cancel out under some field experimental conditions.     

Behavioral responses of the parasitoid Psyllaephagus pistaciae (Hymenoptera: Encyrtidae) to host plant volatiles and honeydew

The behavioral responses of the parasitoid Psyllaephagus pistaciae, the major biocontrol agent of the common pistachio psylla, Agonoscena pistaciae, to volatiles emanating from its host plant and host honeydew, were examined using a four‐arm airflow olfactometer. In addition, the arrestment behavior of this parasitoid on clean and honeydew‐treated leaves of the pistachio, Pistacia vera, was monitored. The infested pistachio leaves were the most favored source of the volatile attracting the parasitoids. The parasitoid clearly distinguished and responded to infochemicals emitted by psyllid honeydew but at a lower level than to the volatiles from infested host plants. However, the searching time, locomotory behavior, antennal drumming and ovipositor probing were all affected when they encountered honeydew‐contaminated zones on pistachio leaves. These findings suggest that the psyllid honeydew releases kairomones that stimulate the parasitoids to greater searching activity, as well as providing a directional cue. The intensive searching activities in the presence of the volatiles tested were very similar to responses by the parasitoid females when encountering patches treated with psyllid honeydew. Such behavior could retain the parasitoid in a favorable area, thereby increasing the probability of additional host encounters.     

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.     

What happens in the pith stays in the pith: tissue‐localized defense responses facilitate chemical niche differentiation between two spatially separated herbivores

Herbivore attack is known to elicit systemic defense responses that spread throughout the host plant and influence the performance of other herbivores. While these plant‐mediated indirect competitive interactions are well described, and the co‐existence of herbivores from different feeding guilds is common, the mechanisms of co‐existence are poorly understood. In both field and glasshouse experiments with a native tobacco, Nicotiana attenuata, we found no evidence of negative interactions when plants were simultaneously attacked by two spatially separated herbivores: a leaf chewer Manduca sexta and a stem borer Trichobaris mucorea. T. mucorea attack elicited jasmonic acid (JA) and jasmonoyl‐l ‐isoleucine bursts in the pith of attacked stems similar to those that occur in leaves when M. sexta attacks N. attenuata leaves. Pith chlorogenic acid (CGA) levels increased 1000‐fold to levels 6‐fold higher than leaf levels after T. mucorea attack; these increases in pith CGA levels, which did not occur in M. sexta‐attacked leaves, required JA signaling. With plants silenced in CGA biosynthesis (irHQT plants), CGA, as well as other caffeic acid conjugates, was demonstrated in both glasshouse and field experiments to function as a direct defense protecting piths against T. mucorea attack, but not against leaf chewers or sucking insects. T. mucorea attack does not systemically activate JA signaling in leaves, while M. sexta leaf‐attack transiently induces detectable but minor pith JA levels that are dwarfed by local responses. We conclude that tissue‐localized defense responses allow tissue‐specialized herbivores to share the same host and occupy different chemical defense niches in the same hostplant.     

Generalism Versus Specialism: Responses of Diadegma mollipla (Holmgren) and Diadegma semiclausum (Hellen), to the Host Shift of the Diamondback Moth (Plutella xylostella L.) to Peas

A population of the diamondback moth Plutella xylostella (L.) (Lepidoptera: Plutellidae) (DBM) was recently found to infest sugar snap- and snowpeas in the Rift Valley in Kenya, causing heavy damage. The influence of this host shift on host location preferences of two parasitoids was investigated: The indigenous Diadegma mollipla (Holmgren) regarded as a relative generalist, and Diadegma semiclausum(Hellen), regarded as highly specific to DBM. The attractiveness of different odour sources was compared for the two parasitoid species using a Y-tube olfactometer using naïve females. D. mollipla was not significantly attracted to any cabbage related odours but showed a significant preference for the DBM infested pea plant when tested against clean air. D. semiclausum was highly attracted to the undamaged cabbage plant and odours related to cabbage. On the other hand, peas infested with DBM, showed no attractiveness to this parasitoid. The results showed that specialisation of D. semiclausum is mediated by host plant signals, associated with crucifers, which are not encountered in DBM feeding on peas. For D. mollipla,although a frequent parasitoid on DBM in crucifers, volatiles emitted by these plants might not be used as primary cues for host location. This species may respond largely to chemicals yet unknown and associated with a variety of plant-herbivore interactions.     

Diadegma insulare development is altered by Plutella xylostella reared on water‐stressed host plants

Natural enemies of herbivores function in a multitrophic context, and their performance is directly or indirectly influenced by herbivores and their host plants. Very little is known about tritrophic interactions between host plants, pests and their parasitoids, particularly when the host plants are under any stress. Herbivores and their natural enemies’ response to plants under stress are diverse and variable. Therefore, in this study we investigated how diamondback moth, Plutella xylostella (L.), reared on water‐stressed host plants (Brassica napus L. and Sinapis alba L.) influenced the development of its larval parasitoid, Diadegma insulare (Cresson). No significant differences were observed in development of P. xylostella when reared on water‐stressed host plants. However, all results indicated that water stress had a strong effect on developmental parameters of D. insulare. Development of D. insulare was delayed when the parasitoid fed on P. xylostella, reared on stressed host plants. Egg to adult development of D. insulare was faster on non‐stressed B. napus than non‐stressed S. alba followed by stressed B. napus and S. alba. Female parasitoids were heavier on non‐stressed host plants than stressed counterparts. Furthermore, the parasitoid lived significantly longer on stressed B. napus. However, body size was not affected by water treatment. Most host plant parameters measured were significantly lower for water‐stressed than non‐stressed treatments. Results suggest that development of this important and effective P. xylostella parasitoid was influenced by both water stress and host plant species.     

Hylobius abietis L. feeding on the novel host Pinus brutia Ten. increases emission of volatile organic compounds

Plants respond to feeding by herbivorous insects by producing volatile organic chemicals, which mediate interactions between herbivores and plants. Yet, few studies investigated whether such plant responses to herbivory differ between historical host and novel plants. Here, we investigated whether herbivory by the pine weevil Hylobius abietis causes a release of volatile organic chemicals from a novel tree Pinus brutia and compared the relative amounts of volatiles released from herbivore's historical hosts and P. brutia. We collected volatiles emitted from P. brutia seedlings that were either subjected to feeding by H. abietis or no feeding. Our results indicated that feeding increased emission of volatile compounds, composed of monoterpenes and sesquiterpenes, and that the emission was several fold higher in the damaged seedlings than in undamaged seedlings. In particular, emission of monoterpenes and sesquiterpenes increased by 4.4‐and 10‐fold in the damaged plants, respectively. Strikingly, individual monoterpenes and sesquiterpenes showed much greater dissimilarity between damaged and undamaged seedlings. Furthermore, several minor monoterpenes showed negative relationships with the weevil gnawed area. We discussed these results with the results of previous studies focused on historical host plants of H. abietis and hypothesized the ecological relevance and importance of our results pertaining relevance to the plant–herbivory interactions.     

Close-Range Host Searching Behavior of the Stemborer Parasitoids <Emphasis Type="Italic">Cotesia sesamiae</Emphasis> and <Emphasis Type="Italic">Dentichasmias busseolae</Emphasis>: Influence of a Non-Host Plant <Emphasis Type="Italic">Melinis minutiflora</Emphasis>

Studies were conducted on the host searching behavior of the larval parasitoid Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) and the pupal parasitoid Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae), both of which attack lepidopteran (Crambidae, Noctuidae) cereal stemborers. The behavior of D. busseolae was observed in a diversified habitat that consisted of stemborer host plants (maize, Zea mays L. and sorghum, Sorghum bicolor (L). Moench (Poaceae)) and a non-host plant (molasses grass, Melinis minutiflora Beauv. (Poaceae)), while C. sesamiae was observed separately on host plants and molasses grass. In previous olfactometer studies, C. sesamiae was attracted to molasses grass volatiles while hboxD. busseolae was repelled. The aim of the present study was to investigate the influence of molasses grass on close-range foraging behavior of the parasitoids in an arena that included infested and uninfested host plants. Dentichasmias busseolae strongly discriminated between host and non-host plants, with female wasps spending most of the time on infested host plants and least time on molasses grass. Likewise, C. sesamiae spent more time on uninfested and infested host plants than it did on molasses grass in single choice bioassays. While on infested plants, the wasps spent more time foraging on the stem, the site of damage, than on other areas of the plant. Overall, the results indicate that presence of the non-host plant does not hinder close range foraging activities of either parasitoid.     

Use of herbivore‐induced plant volatiles as search cues by Tiphia vernalis and Tiphia popilliavora to locate their below‐ground scarabaeid hosts

Japanese beetle, Popillia japonica Newman, and oriental beetle, Anomala orientalis (Waterhouse) (both Coleoptera: Scarabaeidae) are considered invasive species and have been reported as key pests of urban landscapes in the Northeastern USA. Tiphia vernalis Rohwer and Tiphia popilliavora Rohwer (Hymenoptera: Tiphiidae) were introduced as biocontrol agents against these beetles. These parasitic wasps burrow into the soil and search for grubs. When a host is found, the wasp attaches an egg in a location that is specific for the wasp species. It is unknown if these wasps can detect patches of concealed hosts from a distance above ground and what role, if any, herbivore‐induced plant volatiles play in their host location. This study evaluated the responses of female T. vernalis and T. popilliavora to grub‐infested and healthy plants in Y‐tube olfactometer bioassays. Also the effect of root herbivory on the composition of turfgrass (Poaceae) volatile profiles was investigated by collecting volatiles from healthy and grub‐infested grasses. Tiphia wasps were highly attracted to volatiles emitted by grub‐infested tall fescue (Festuca arundinacea Schreb.) and Kentucky bluegrass (Poa pratensis L.) over healthy grasses. In contrast, wasps did not exhibit a significant preference for grub‐infested perennial ryegrass (Lolium perenne L.) as compared with the control plants. The terpene levels emitted by grub‐infested Kentucky bluegrass and tall fescue were greater than that of control plants. Low levels of terpenes were observed for both test and control perennial ryegrass. The elevated levels of terpenes emitted by grub‐infested Kentucky bluegrass and tall fescue coincided with the attractiveness to the tiphiid wasps. Here, we provide evidence that plant exposure to root‐feeding insects P. japonica and A. orientalis resulted in an increase in terpenoid levels in turfgrasses, which strongly attracts their above‐ground parasitoids.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.