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.     

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.     

Response of two parasitoid species (Hymenoptera: Braconidae,Figitidae) to tephritid host and host food substrate cues

The Neotropical‐native figitid Aganaspis pelleranoi (Brèthes) and the Asian braconid Diachasmimorpha longicaudata (Ashmead) are two parasitoids of Tephritidae fruit flies with long and recent, respectively, evolutionary histories in the Neotropics. Both species experienced a recent range of overlap. In Argentina, A. pelleranoi is a potential species in biological control programs against the pestiferous tephritid species, Anastrepha fraterculus (Wiedemann) and Ceratitis capitata (Wiedemann), whereas D. longicaudata is already used in open‐field releases against Medfly in central‐western Argentina. To characterize the host‐foraging strategies of A. pelleranoi and D. longicaudata, olfactometer experiments were conducted comparing responses to C. capitata and A. fraterculus larvae, in two kinds of food substrate: fruit and artificial larval medium. To control the possible influence of host larvae used for parasitoid rearing on olfactory response, two strains of both parasitoid species, reared on both tephrtid species, were studied. Volatiles directly emanating either from A. fraterculus or C. capitata larvae may be detected by both A. pelleranoi and D. longicaudata, although chemical stimuli originating from the combination of host larvae and the habitat of the host were preferred. However, olfactory cues associated with host larvae probably play a relevant role in host searching behaviour of A. pelleranoi, whereas for D. longicaudata, the host‐habitat olfactory stimuli would be highly essential in short‐range host location. The strain of the parasitoids did not affect host search ability on the two tephritid species evaluated. These evidences are relevant for mass production of both parasitoids and their impact following open‐field augmentative releases.     

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.     

Host selection by Ibalia leucospoides based on temporal variations of volatiles from the hosts’ fungal symbiont

Insect parasitoids locate hosts via reliable and predictable cues such as volatile emissions from hosts and/or host plants. For insects that depend on mutualistic organisms, such as many wood‐boring insects, symbiont‐derived semiochemicals may represent a source of such cues to be exploited by natural enemies. Ultimately, exploitation of these signals may increase fitness by optimizing foraging efficiency. Female parasitoids of Ibalia leucospoides use volatiles from the fungal symbiont Amylostereum areolatum of their host Sirex noctlio to find concealed host eggs and young larvae within the xylem. We hypothesize that the temporal pattern of fungal emissions may indicate not only the presence of host larvae but also be used as a cue that indicates host suitability and age. Such information would allow female parasitoids to discern more efficiently between hosts within ovipositor reach from those already buried too deep into the xylem and out of reach. In this context, we assessed the behaviour of I. leucospoides females to volatiles of A. areolatum in a ‘Y’‐tube olfactometer at regular intervals over 30 days. We concurrently examined the fungal volatiles by headspace sampling through solid‐phase microextraction (SPME) followed by gas chromatography mass spectrometry (GC‐MS). We observed that females were attracted to volatiles produced by two‐week‐old fungal cultures, a period that matches when older larvae are still within ovipositor reach. Four chemical compounds were detected: ethanol, acetone, acetaldehyde and the sesquiterpene 2,2,8‐trimethyltricyclo[6.2.2.01,6]dodec‐5‐ene, with each compounds’ relative abundance changing over time. Results are discussed in the context of parasitoids fitness. Future studies involving electrophysiology, different collection techniques and further behavioural assays will help in identifying the compounds that convey temporal information to female parasitoids and have the potential for being used in integrated pest management programmes.     

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.     

Development of Asian parasitoids in larvae Of Drosophila Suzukii feeding on blueberry and artificial diet

In just a few years, the Asian fly Drosophila suzukii has invaded several continents and has become a very serious pest of many fruit crops worldwide. Current control methods rely on chemical insecticides or expensive and labour‐intensive cultural practices. Classical biological control through the introduction of Asian parasitoids that have co‐evolved with the pest may provide a sustainable solution on condition that they are sufficiently specific to avoid non‐target effects on local biodiversity. Here, we present the first study on the development of three larval parasitoids from China and Japan, the Braconidae Asobara japonica and the Figitidae Leptopilina japonica and Ganaspis sp., on D. suzukii. The Asian parasitoids were compared with Leptopilina heterotoma, a common parasitoid of several Drosophilidae worldwide. The three Asian species were successfully reared on D. suzukii larvae in both, blueberry and artificial diet, in contrast to L. heterotoma whose eggs and larvae were encapsulated by the host larvae. All parasitoids were able to oviposit one day after emergence. Asobara japonica laid as many eggs in larvae feeding in blueberry as in artificial diet, whereas L. heterotoma oviposited more in larvae on the artificial diet and the Asian Figitidae oviposited more in larvae feeding on blueberry. Ganaspis sp. laid very few eggs in larvae in the artificial diet, suggesting that it may be specialized in Drosophila species living in fresh fruits. These data will be used for the development of a host range testing to assess the suitability of Asian parasitoids as biological control agents in invaded regions.     

Volatiles from non‐host aromatic plants repel tea green leafhopper Empoasca vitis

The tea green leafhopper, Empoasca vitis Göthe (Hemiptera: Cicadellidae), is a serious pest of tea crops in China. The effectiveness of five aromatic non‐host plants, Corymbia citriodora (Hook.) (Myrtaceae), Ocimum basilicum L. (Lamiaceae), Lavandula pinnata L. (Lamiaceae), Ruta graveolens L. (Rutaceae), and Rosmarinus officinalis L. (Lamiaceae), was investigated to determine their ability to suppress E. vitis on tea plants. Volatile organic compounds derived from L. pinnata, R. officinalis, and C. citriodora were found to repel leafhoppers and to mask the host's odors. Intercropping L. pinnata and C. citriodora with tea plants significantly reduced the E. vitis population levels in the tea plantation associated with higher population densities of generalist predators. The volatile compounds from the five non‐host plants were collected by headspace absorption under field conditions, and the 10 major components were identified. Qualitative and quantitative differences were found among the five odor profiles. Moreover, the emission dynamics of non‐host volatiles were monitored. Non‐host volatile emissions showed two peak periods, one in the spring and one in the autumn period. These peaks were almost consistent with the population dynamics of E. vitis. Our findings suggested that this newer approach to ecologically based, sustainable pest management implemented via intercropping with non‐host aromatic plants such as L. pinnata and C. citriodora offers a promising alternative to chemical control of the leafhopper population.     

Fitness of Bt‐resistant cabbage loopers on Bt cotton plants

Development of resistance to the insecticidal toxins from Bacillus thuringiensis (Bt) in insects is the major threat to the continued success of transgenic Bt crops in agriculture. The fitness of Bt‐resistant insects on Bt and non‐Bt plants is a key parameter that determines the development of Bt resistance in insect populations. In this study, a comprehensive analysis of the fitness of Bt‐resistant Trichoplusia ni strains on Bt cotton leaves was conducted. The Bt‐resistant T. ni strains carried two genetically independent mechanisms of resistance to Bt toxins Cry1Ac and Cry2Ab. The effects of the two resistance mechanisms, individually and in combination, on the fitness of the T. ni strains on conventional non‐Bt cotton and on transgenic Bt cotton leaves expressing a single‐toxin Cry1Ac (Bollgard I) or two Bt toxins Cry1Ac and Cry2Ab (Bollgard II) were examined. The presence of Bt toxins in plants reduced the fitness of resistant insects, indicated by decreased net reproductive rate (R₀) and intrinsic rate of increase (r). The reduction in fitness in resistant T. ni on Bollgard II leaves was greater than that on Bollgard I leaves. A 12.4‐day asynchrony of adult emergence between the susceptible T. ni grown on non‐Bt cotton leaves and the dual‐toxin‐resistant T. ni on Bollgard II leaves was observed. Therefore, multitoxin Bt plants not only reduce the probability for T. ni to develop resistance but also strongly reduce the fitness of resistant insects feeding on the plants.     

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.     

Oviposition site preference and developmental performance of a gall‐inducing leafhopper on galled and non‐galled host plants

Oviposition preferences of herbivorous insects affect offspring performance. Both positive and negative links between oviposition preference and offspring performance have been reported for many species. A gall‐inducing leafhopper, Cicadulina bipunctata Melichar (Hemiptera: Cicadellidae), feeds on various Poaceae plants and induces galls of enhanced nutritional value for their offspring. Although gall induction by C. bipunctata improves nymphal performance, the oviposition preference of females between galled and non‐galled host plants is still unclear. In this paper, the nymphal performance and oviposition and feeding‐site preference of C. bipunctata were investigated using galled wheat, Triticum aestivum L., and non‐galled barley, Hordeum vulgare L., as host plants. The survival rate of C. bipunctata on wheat was significantly higher than on barley. In the choice test, significantly more eggs were laid into barley, whereas the number of eggs deposited on both hosts was not significantly different in the no‐choice test. The number of settling individuals per leaf area was not significantly different between wheat and barley, suggesting no clear preference for oviposition between these plants. Experience as a nymph with a growing host did not affect oviposition preference as adult female. The inconsistent correspondence between offspring performance and oviposition preference of C. bipunctata may reflect the high mobility of nymphs and/or differences in leaf area between host plants. The results indicate that the previous finding that oviposition preference and offspring performance are not always positively correlated in herbivorous insects is applicable to gall‐inducing insects.     

Estimating costs of aphid resistance to parasitoids conferred by a protective strain of the bacterial endosymbiont Regiella insecticola

Heritable bacterial endosymbionts are common in aphids (Hemiptera: Aphididae), and they can influence ecologically important traits of their hosts. It is generally assumed that their persistence in a population is dependent on a balance between the costs and benefits they confer. A good example is Hamiltonella defensa Moran et al., a facultative symbiont that provides a benefit by strongly increasing aphid resistance to parasitoid wasps, but becomes costly to the host in the absence of parasitoids. Regiella insecticola Moran et al. is another common symbiont of aphids and generally does not influence resistance to parasitoids. In the green peach aphid, Myzus persicae (Sulzer), however, one strain (R5.15) was discovered that behaves like H. defensa in that it provides strong protection against parasitoid wasps. Here we compare R5.15‐infected and uninfected lines of three M. persicae clones to test whether this protective symbiont is costly as well, i.e., whether it has any negative effects on aphid life‐history traits. Furthermore, we transferred R5.15 to two other aphid species, the pea aphid, Acyrthosiphon pisum (Harris), and the black bean aphid, Aphis fabae Scopoli, where this strain is also protective against parasitoids and where we could compare its effects with those of additional, non‐protective strains of R. insecticola. Negative effects of R5.15 on host survival and lifetime reproduction were limited and frequently non‐significant, and these effects were comparable or in one case weaker than those of R. insecticola strains that are not protective against parasitoid wasps. Unless the benefit of protection is counteracted by detrimental effects on traits that were not considered in this study, R. insecticola strain R5.15 should have a high potential to spread in aphid populations.     

Herbivore species identity rather than diversity of the non‐host community determines foraging behaviour of the parasitoid wasp Cotesia glomerata

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species‐rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore‐induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non‐suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non‐host herbivores influence the behaviour of parasitoids, we created environments with different levels of non‐host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non‐host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non‐host species diversity nor non‐host identity influenced the preference of the parasitoid for herbivore‐infested plants. However, after landing on the plant, non‐host species identity did affect parasitoid behaviour, whereas non‐host diversity did not. One of the non‐host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non‐host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non‐host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.     

Inoculation of susceptible and resistant potato plants with the late blight pathogen Phytophthora infestans: effects on an aphid and its parasitoid

Plants are exposed to microbial pathogens as well as herbivorous insects and their natural enemies. Here, we examined the effects of inoculation of potato plants, Solanum tuberosum L. (Solanaceae), with the late blight pathogen Phytophthora infestans (Mont.) de Bary (Peronosporales: Pythiaceae) on an aphid species commonly infesting potato crops and one of the aphid's major parasitoids. We observed the peach‐potato aphid, Myzus persicae Sulzer (Hemiptera: Aphididae), and its natural enemy, the biocontrol agent Aphidius colemani Viereck (Hymenoptera: Braconidae), on potato either inoculated with water or P. infestans. Population growth of the aphid, parasitism rate of its natural enemy, and other insect life‐history traits were compared on several potato genotypes, the susceptible cultivar Désirée and genetically modified (GM) isogenic lines carrying genes conferring resistance to P. infestans. Effects of P. infestans inoculation on the intrinsic rate of aphid population increase and the performance of the parasitoid were only found on the susceptible cultivar. Insect traits were similar when comparing inoculated with non‐inoculated resistant GM genotypes. We also tested how GM‐plant characteristics such as location of gene insertion and number of R genes could influence non‐target insects by comparing insect performance among GM events. Different transformation events leading to different positions of R‐gene insertion in the genome influenced aphids either with or without P. infestans infection, whereas effects of position of R‐gene insertion on the parasitoid A. colemani were evident only in the presence of inoculation with P. infestans. We conclude that it is important to study different transformation events before continuing with further stages of risk assessment of this GM crop. This provides important information on the effects of plant resistance to a phytopathogen on non‐target insects at various trophic levels.     

Climate variation alters the synchrony of host–parasitoid interactions

Observed changes in mean temperature and increased frequency of extreme climate events have already impacted the distributions and phenologies of various organisms, including insects. Although some research has examined how parasitoids will respond to colder temperatures or experimental warming, we know relatively little about how increased variation in temperature and humidity could affect interactions between parasitoids and their hosts. Using a study system consisting of emerald ash borer (EAB), Agrilus planipennis, and its egg parasitoid Oobius agrili, we conducted environmentally controlled laboratory experiments to investigate how increased seasonal climate variation affected the synchrony of host–parasitoid interactions. We hypothesized that increased climate variation would lead to decreases in host and parasitoid survival, host fecundity, and percent parasitism (independent of host density), while also influencing percent diapause in parasitoids. EAB was reared in environmental chambers under four climate variation treatments (standard deviations in temperature of 1.24, 3.00, 3.60, and 4.79°C), while O. agrili experiments were conducted in the same environmental chambers using a 4 × 3 design (four climate variation treatments × 3 EAB egg densities). We found that EAB fecundity was negatively associated with temperature variation and that temperature variation altered the temporal egg laying distribution of EAB. Additionally, even moderate increases in temperature variation affected parasitoid emergence times, while decreasing percent parasitism and survival. Furthermore, percent diapause in parasitoids was positively associated with humidity variation. Our findings indicate that relatively small changes in the frequency and severity of extreme climate events have the potential to phenologically isolate emerging parasitoids from host eggs, which in the absence of alternative hosts could lead to localized extinctions. More broadly, these results indicate how climate change could affect various life history parameters in insects, and have implications for consumer–resource stability and biological control.     

Patch exploitation by non‐aggressive parasitoids under intra‐ and interspecific competition

The behavioral strategies evolved by insect parasitoids to optimize their foraging efforts have been the subject of many theoretical and empirical studies. However, the effects competition may have on these strategies, especially for species that do not engage in antagonistic behaviors, have received little attention. The objective of this study was to evaluate the effects of intraspecific and interspecific competition on patch exploitation strategies by two non‐aggressive species, Trichogramma pintoi Voegelé and Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), both generalist egg parasitoids. We analyzed the patch residence times of females, their patch‐leaving mechanisms, and the sex allocation of their progeny while foraging either alone, with an intraspecific competitor, or with an interspecific competitor. To some extent, each species responded differently to the presence of a competitor in the patch. Trichogramma pintoi females did not change their patch‐leaving mechanisms in response to competition and behaved as if under an exploitative competition regime, whereas T. minutum females did change their patch‐leaving mechanisms in response to competition and remained longer in the host patch than expected. Antennal rejection, and not oviposition, was the proximate behavioral mechanism underlying patch‐leaving decisions by both species. Neither species adjusted the sex allocation of their progeny in response to competition. These results indicate that the effects of competition differ even among closely related parasitoid species.     

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.     

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.