Parasitoids follow herbivorous insects to a novel host plant,generalist predators less so

The ‘enemy‐free space’ hypothesis predicts that herbivorous insects can escape their natural enemies by switching to a novel host plant, with consequences for the evolution of host plant specialisation. However, if natural enemies follow herbivores to their novel host plants, enemy‐free space may only be temporary. We tested this by studying the colonisation of the introduced tree Eucalyptus grandis (Hill) Maiden (Myrtaceae) by insects in Brazil, where various species of herbivores have added eucalyptus to their host plant range, which consists of native myrtaceous species such as guava. Some herbivores, for example, Thyrinteina leucoceraea Ringe (Lepidoptera: Geometridae), cause outbreaks in eucalyptus plantations but not on guava, possibly because eucalyptus offers enemy‐free space. We sampled herbivores (mainly Lepidoptera species) and natural enemies on eucalyptus and guava and assessed parasitism of Lepidoptera larvae on both host plant species during ca. 2 years. Overall, predators were encountered more frequently on guava than on eucalyptus. In contrast, parasitoids were encountered equally and parasitism rates of Lepidoptera larvae were similar on both host plants. This indicates that herbivores may escape some enemies by moving to a novel host plant. However, this escape may be temporary and may vary with time. We argue that studying temporal and spatial patterns of enemy‐free space and the response of natural enemies to host use changes of their herbivorous prey is essential for understanding the role of natural enemies in the evolution of host plant use by herbivorous arthropods.     

Searching behaviour of an omnivorous predator for novel and native host plants of its herbivores: a study on arthropod colonization of eucalyptus in Brazil

Adaptation to novel host plants is a much‐studied process in arthropod herbivores, but not in their predators. This is surprising, considering the attention that has been given to the role of predators in host range expansion in herbivores; the enemy‐free space hypothesis suggests that plants may be included in the host range of herbivores because of lower predation and parasitism rates on the novel host plants. This effect can only be important if natural enemies do not follow their prey to the novel host plant, at least not immediately, thus allowing the herbivores to adapt to the novel host plant. Hence, depending on the speed with which natural enemies follow their prey to a new host plant, enemy‐free space on novel host plants may only exist for a limited period. This situation may presently be occurring in a system consisting of the herbivorous moth Thyrinteina arnobia Stoll (Lepidoptera: Geometridae) that attacks various species of Myrtaceae, such as guava (Psidium guajava L.) and jaboticaba (Myrciaria spp.), in Brazil. Since the introduction of eucalyptus (Myrtaceae) species into this country some 100 years ago, the moth has included this plant species in its host range and frequently causes outbreaks, a phenomenon that does not occur on the native host plant species. This suggests that the natural enemies that attack the herbivore on native species are not very effective on the novel host. We tested this hypothesis by studying the searching behaviour of one of the natural enemies, the omnivorous predatory bug Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae). When offered a choice between plants of the two species, the predators (originally collected in eucalyptus plantations) preferred guava to eucalyptus when both plant species were clean, infested with herbivores, or damaged by herbivores but with herbivores removed prior to the experiments. The bugs preferred herbivore‐damaged to clean guava, and showed a slight preference for damaged to clean eucalyptus. These results may explain the lack of impact of predatory arthropods on herbivore populations on eucalyptus and suggests that eucalyptus may offer an enemy‐free space for herbivores.     

How predation by Podisus nigrispinus is influenced by developmental stage and density of its prey Alabama argillacea

The functional response of a predator to the density of its prey is affected by several factors, including the prey's developmental stage. This study evaluated the functional response of Podisus nigrispinus (Dallas) (Hemiptera: Heteroptera: Pentatomidae) females to fourth instars and pupae of Alabama argillacea (Hübner) (Lepidoptera: Noctuidae), an important pest of cotton (Gossypium hirsutum L., Malvaceae) in Brazil. The prey were exposed to the predator for 12 and 24 h, and in densities of 1, 6, 12, 18, 24, and 30 items per predator female. The predation data were subjected to polynomial regression logistic analysis to determine the type of functional response. Holling and Rogers' equations were used to estimate parameters such as attack rate and handling time. Podisus nigrispinus females showed functional response types II and III by preying on larvae and pupae, respectively. The attack rate and handling time did not differ between the 12 and 24 h exposure times. Predation rate was higher at higher larval and pupal densities; predation was highest at a density of 30 prey items per female, and it was similar at 18 and 24 prey per predator. Understanding the interaction of predators and their food resources helps to optimize biological control strategies. It also helps the decision‐making and the improvement of release techniques of P. nigrispinus in the field.     

Relationship between predation by Podisus nigrispinus and developmental phase and density of its prey,Plutella xylostella

This study evaluated the predation by Podisus nigrispinus (Dallas) (Hemiptera: Pentatomidae) at various densities of larvae and pupae of the pest Plutella xylostella (L.) (Lepidoptera: Plutellidae). We tested predator behavior of female P. nigrispinus at six experimental densities (1, 5, 10, 15, 20, or 25 prey items in a 1‐l transparent plastic container, replicated 15 times for each density) of both the fourth instar and pupae of P. xylostella. The number of prey consumed was monitored every 15 min for 12 h and was subsequently monitored at 24 h. Podisus nigrispinus females were weighed before and after the experiments to determine the effect of different densities of prey on their weight gain. Female predators had a Type‐II functional response, with attack rate estimated at 1.387 and 0.260 and a handling time of 0.091 and 0.183 h^?1 for larvae and pupae, respectively. Podisus nigrispinus consumed on average 10.9 larvae or 5.5 pupae in 24 h. Despite the similarity of the response type, P. nigrispinus preferred to feed on larvae, rather than on pupae.     

Predatory behaviour of Podisus nigrispinus (Heteroptera: Pentatomidae) on different densities of Anticarsia gemmatalis (Lepidoptera: Noctuidae) larvae

The functional response of the predatory bug Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae) feeding on its prey, Anticarsia gemmatalis Hübner (Lepidoptera: Noctuidae), was studied in a greenhouse compartment. Each cage enclosed three soybean plants plus two, four, six, eight, 10, 12 or 14 prey larvae. One adult predator was released and kept inside the cages for 24h. The predation rate of adult male P. nigrispinus was highest at densities of eight or more A. gemmatalis larvae with a handling time of 5.76h and an attack rate of 0.68h^?1. Adult females had higher predation rates on plants with 10 or more A. gemmatalis larvae, with a handling time of 3.84h and an attack rate of 0.65h^?1. The maximum number of larvae consumed by males and females of this predator were 4.1 and 6.0 per day, respectively, in groups of three plants. The results suggest that P. nigrispinus may be used in biological control programmes against A. gemmatalis in soybean fields.     

Carbon dioxide‐ and temperature‐mediated changes in plant defensive compounds alter food utilization of herbivores

Although the impact of elevated carbon dioxide and rising temperature on plants and animals has been extensively documented recently, only limited understanding exists regarding their combined effects. The objective of this research was to address the consequences of using combinations of elevated CO₂ and elevated temperature on a plant's defensive chemistry, and subsequent utilization of the plant as insect food. Our results indicated that elevated CO₂ and increased temperature, for the most part, act independently on the production of defensive compounds in broccoli leaves (Brassica oleracea L. var. italica). CO₂ concentrations had significant effects on the foliar water content, total phenolic compounds, polyphenol oxidase and trypsin inhibitor concentrations. The herbivore Spodoptera litura (Fabricius; Lepidoptera: Noctuidae) responded to changes in the plant secondary chemistry, with larvae consuming more plant materials that had been exposed to elevated CO₂. The food utilization efficiencies of second‐instar larvae were more sensitive to CO₂‐treated foliage than those of the third‐ and fourth‐instar larvae. Temperature did exert a significant effect on food utilization (ECD) by the larvae. Our study will provide important information in future predictions on plant–insect interactions as a result of climate change. The study also demonstrated that since various larval stages might respond differently to climate change, this possibility needs to be considered in future forecasting and monitoring.     

Oviposition pattern of the predator Podisus nigrispinus (Heteroptera: Pentatomidae) under different temperatures

The daily reproductive rate of Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae) fed with Alabama argillacea (Hübner) (Lepidoptera: Noctuidae) larvae was studied at constant temperatures of 20, 23, 25, 28, 30 and 33±0.2°C, relative humidity of 60±10% and photoperiod of L:D 14:10. Daily reproductive rate of P. nigripinus was affected by age of this predator. Each P. nigrispinus female laid 5.3 (20°C) to 19.9 eggs/day (28°C) which developed into 4.3–16.5 nymphs, respectively. Highest daily reproductive rate of P. nigrispinus was recorded at 28 and 30°C for 5–30-day-old females. This predator showed higher daily reproductive rate than its prey A. argillacea at 25°C. It was also able to reproduce at temperatures from 20 to 33°C with maximum daily reproductive rate between 25 and 30°C. These results are important for optimizing mass rearing of P. nigrispinus in the laboratory.     

Caterpillar‐induced plant volatiles attract conspecific herbivores and a generalist predator

Plants release volatiles in response to caterpillar feeding that attracts natural enemies of the herbivores, a tritrophic interaction which has been considered to be an indirect plant defence against herbivores. On the other hand, the caterpillar‐induced plant volatiles have been reported to either repel or attract conspecific adult herbivores. This work was undertaken to investigate the response of both herbivores and natural enemies to caterpillar‐induced plant volatiles in apple orchards. We sampled volatile compounds emitted from uninfested apple trees, and apple trees infested with generalist herbivore the pandemis leafroller moth, Pandemis pyrusana (Lepidoptera, Tortricidae) larvae using headspace collection and analysed by gas chromatography/mass spectrometry. Infested apple trees uniquely release six compounds (benzyl alcohol, phenylacetonitrile, phenylacetaldehyde, 2‐phenylethanol, indole and (E)‐nerolidol). These compounds were tested on two species of herbivores and one predator in apple orchards. Binary blends of phenylacetonitrile + acetic acid or 2‐phenylethanol + acetic acid attracted a large number of conspecific male and female adult herbivores. The response of pandemis leafroller to herbivore‐induced plant volatiles (HIPVs) was so pronounced that over one thousand and seven hundred conspecific male and female adult herbivores were caught in traps baited with HIPVs in three‐day trapping period. In addition, significantly higher number of male and female obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera, Tortricidae), was caught in traps baited a binary blend of 2‐phenylethanol + acetic acid, or a ternary blend contains 2‐phenylethanol and phenylacetonitrile + acetic acid. This result challenges the current paradigm hypothesized that HIPVs repel herbivores and question the indirect defensive function proposed for these compounds. On the other hand, a ternary blend of phenylacetonitrile and 2‐phenylethanol + acetic acid attracted the largest numbers of the general predator, the common green lacewing, Chrysoperla plorabunda. To our knowledge, this is the first record of the direct attraction of conspecific adult herbivores as well as a predator to the caterpillar‐induced plant volatiles in the field.     

Effects of carotenoid sequestration on a caterpillar's cryptic coloration and susceptibility to predation

Carotenoids are used for many functions by animals, including combining with other pigments to produce aposematic and cryptic coloration. Carotenoids in combination with blue pigments are responsible for green coloration in many caterpillars, and thus carotenoid sequestration may reduce their contrast against a green foliage background. We tested the hypothesis that carotenoid sequestration reduces contrast and enhances survival by rearing Trichoplusia ni Hübner (Lepidoptera: Noctuidae) on Brassica oleracea L. var. Acephala (Brassicaceae) leaves and exposing them to predators. We found that carotenoids derived from the host plant are partially excreted, along with chlorophyll, but also sequestered in hemolymph. Larvae that were given plants that provided carotenoids showed less contrast against their host plants within 1 day compared to larvae that were not provided with carotenoids. Last, both short‐term field observations and laboratory trials of larvae caged with predatory Podisus maculiventris Say (Hemiptera: Pentatomidae) nymphs showed that survival of carotenoid‐sequestering larvae was higher compared to larvae that did not sequester. These results suggest that carotenoid sequestration may be an important adaptive strategy that reduces susceptibility to natural enemies that hunt by sight. Further research that examines the mechanisms by which carotenoids are absorbed and modified will lend insights into the evolution of carotenoids functioning as passive defensive compounds.     

Leaf biomechanical properties as mechanisms of resistance to black cutworm (Agrotis ipsilon) among Poa species

Biomechanical properties can be important parameters in resistance of plants to herbivorous insects. As plants age, however, there can be dramatic changes in physical defenses that can then influence their susceptibility to insect herbivores. We measured changes in leaf biomechanical properties during ontogeny of Poa species and the relationship of these changes to the development of a generalist herbivore, the black cutworm, Agrotis ipsilon Hufnagel (Lepidoptera: Noctuidae), was investigated. Larvae were reared on two representative age classes, i.e., young (<60 days after planting) and old (>1 year after planting), of foliage in laboratory assays. Foliage generally reaches a peak fracture force between 80 and 109 days after planting depending on grass type. Foliage from old plants was significantly tougher than that of young plants, and black cutworm larvae reared on foliage from young plants gained significantly (ca. four times) more weight than those fed on foliage from old Poa plants. In addition, fracture force has a negative relationship with black cutworm development. Plant fiber, particularly neutral detergent fiber accounted for 65 and 46% of the variation in fracture force and larval development, respectively. These results provide additional insight into how plant ontogeny influences physical defenses to an insect herbivore in a grass system. Likewise, this is seemingly the first study to suggest a mechanism for host plant resistance to black cutworm. Plant fiber may be a useful trait to explore in plant improvement programs in which black cutworm is a primary pest (e.g., managed turfgrass).     

Plant silicon effects on insect feeding dynamics are influenced by plant nitrogen availability

Although there is growing evidence that silicon (Si)‐based plant defenses effectively reduce both the palatability and digestibility of leaves, and thus impact nutrient assimilation by insect herbivores, much less is known about how this is affected by extrinsic and intrinsic factors. For example, do herbivores exhibit compensatory feeding on poor‐quality diets with Si or are Si defenses less effective in agroecosystems where high N availability increases plant quality? To investigate the interactive effects of N and Si on insect feeding, we conducted insect performance and compensatory feeding bioassays using maize, Zea mays L. (Poaceae), and the true armyworm, Pseudeletia unipuncta Haworth (Lepidoptera: Noctuidae). In the performance assay, the addition of Si alone resulted in increased larval mortality compared with the controls, likely because early instars with poorly developed mandibles could not feed effectively. However, larvae fed on plants treated with both Si and N survived better than on plants treated with Si only, although pupal mass did not differ between treatments. In our compensatory assay, Si addition reduced maize consumption, but increased both armyworm approximate digestibility and N assimilation efficiency, suggesting that enhanced post‐ingestion feeding physiology, rather than compensatory food intake, could have accounted for the lack of Si effects on pupal weight. Overall, our results demonstrate that, similar to other chemical and mechanical defenses, the effectiveness of plant Si defense is influenced by plant nutrient status and consumer compensatory ability.     

Enemy release but no evolutionary loss of defence in a plant invasion: an inter-continental reciprocal transplant experiment

Plant chemical defenses and escape from natural enemies have been postulated to select for dietary specialization in herbivorous insects. In field and laboratory bioassays, we evaluated the effectiveness of intact and chemically modified larval shield defenses of the generalist Chelymorpha alternans and the specialists Acromis sparsa and Stolas plagiata (Chrysomelidae: Cassidinae) against three natural predators, using larvae reared on two morning glory (Convolvulaceae) species. We assessed whether: (1) specialists were better defended than generalists when both were fed and assayed on the same plant; (2) larval shield defenses were chemical, physical, or both; and (3) specialists exploit chemistry better than generalists. Live specialist larvae survived at higher rates than did generalists in predator bioassays with the bug Montina nigripes (Reduviidae), but there were no differences among groups against two species of Azteca ants (Hymenoptera: Dolichoderinae). Solvent leaching by H₂O or MeOH significantly reduced shield efficacy for all species compared to larvae with intact shields. In contrast, freshly killed specialist larvae exhibited significantly lower capture rates and frequencies than the generalists. Although solvent leaching significantly reduced overall shield efficacy for freshly killed larvae of all species, the pattern of leaching effects differed between specialists and generalists, with H₂O-leaching having a greater impact on the specialists. The overall vulnerability of the generalists appears due to lower chemical protection, which is ameliorated by increased escape behaviors, suggesting a selective trade-off between these defensive components. These experiments indicate that shield defenses are essential for larval survival and that specialists are superior at exploiting plant compounds residing in the aqueous fraction. Our results support the hypothesis that diet-specialized herbivorous insects have more effective defenses than generalists when both feed on the same plant due to the differential ability to exploit defensive precursors obtained from the host. The evolution of dietary specialization may therefore confer the advantage of enhanced enemy-free space.     

Effects of prey availability, facultative plant feeding, and plant defenses on a generalist insect predator

This study examined the effects of feeding interval, access to host plants (thus, a source of sap), and plant defenses on the predatory insect, Podisus maculiventris Say (Hemiptera: Pentatomidae). The experiment consisted of a 2 × 2 design with two feeding intervals (1 day or 5 days) and predators living on either tomato plants or plastic plants. Females fed every day had greater body weights and egg hatch rates than females fed every five days. Females on tomato plants lived longer than females on plastic plants. However, access to plants did not alleviate the effects of low prey level on predator weight or reproductive output. In a second experiment, third instar nymphs were placed on either tomato plants or plastic plants for four days to examine the effects of tomato trichome defenses on these predators. Nymphs on tomato plants experienced 50% mortality compared to 15% mortality for nymphs on plastic plants. Some nymphs living on tomato plants were trapped by the hairy trichomes of the plant; others had gummed up legs from the exudates of the plants’ glandular trichomes, which inhibited their movement and ability to feed on prey. Although predators appeared to benefit from feeding on tomato plants, their ability to live on the plants was negatively affected by the defensive features of the plants. The potential effects of trichome defenses on predator survival and population dynamics must be considered when evaluating the benefits of plants on insect predator life histories and efficacy as biological control agents.     

Development and reproduction of a predatory stinkbug, Podisus nigrispinus in relation to two different prey types and environmental conditions

Abiotic factors, prey availability, prey type, and plant host actsimultaneously under field conditions. Their combined effects werepartially investigated for the predatory stinkbugPodisus nigrispinus(Dallas) (Heteroptera: Pentatomidae) preying on tomato leafminer (TLM)caterpillars Tuta absoluta (Meyrick)(Lepidoptera: Gelechiidae) (a preyfeeding on a plant host), and compared to yellow mealworm (YMW)Tenebrio molitor (Coleoptera: Tenebrionidae)pupae (a prey not feeding ona plant host). These experiments were carried out in the laboratory and inan open-sided greenhouse. In general, the environmental variationsbetween laboratory and open-sided greenhouse did not induce differencesin life history characteristics forP. nigrispinus feeding on the sameprey type. However, development and reproductionof P. nigrispinuswas greatly affected by prey type. Nymphal developmental times weresimilar between prey types under laboratory conditions; but were longer(and produced adults of lower body weight) when predators in theopen-sided greenhouse were fed on TLM compared with YMW. Mean mortalityduring the second instar was higher on both prey types in the open-sidedgreenhouse than in the laboratory. Adult emergencewas 38.5 and 50% inthe laboratory, and 32.5 and 48.6%in the open-sided greenhouse for bugsfed on TLM and YMW, respectively. Reproductive characteristics weresimilar for the different environmental conditions when the predator fed onthe same prey, but offspring production was greater for females fed on YMWcompared to females fed on TLM. These results suggest thatP. nigrispinus can adjust its performance accordingto the type of prey available, independently from environmentalvariations. It has the ability to survive on alternative prey,enabling it to maintain a presence in thetomato ecosystem until a primary pest species arrives.     

Predator Presence Moves <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> Larvae to Distraction

Displacement of herbivorous insects by the presence of predators on whole plants has rarely been studied. By semi-continuous observations of an externally feeding insect herbivore and a predator, we show how the mere presence of the predator, Geocoris lubra Kirkaldy (Hemiptera: Geocoridae), on a plant can have a strong influence on the movement and behaviors of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae. The presence of predators, as opposed to mortality by predators, influenced the proportion of larvae feeding, resting and implementing avoidance activities. In addition, the proportion of time individual larvae allocated to feeding, resting and dropping off plants was affected when predators were present with and without contact between the two. Predators do more than just reduce numbers of herbivores; they influence feeding, displacement and subsequently the distribution of plant damage.     

The exploitation of an ant-defended host plant by a shelter-building herbivore

Larvae of a Polyhymno species (Lepidoptera: Gelechiidae) feed on the ant-defended acacia, Acacia cornigera, in the tropical lowlands of Veracruz, Mexico. Polyhymno larvae construct sealed shelters by silking together the pinna or pinnules of acacia leaves. Although larval density and larval survival are higher on acacias not occupied by ants, shelters serve as a partial refuge from the ant Pseudomyrmex ferruginea (Hymenoptera: Formicidae), which defends A. cornigera plants; thus, shelters provide Polyhymno larvae access to an ant-defended host plant. P. ferruginea ants act as the primary antiherbivore defense of A. cornigera plants, which lack the chemical and mechanical defenses of non-ant-defended acacias. Thus, defeating the ant defense of A. cornigera provides Polyhymno larvae access to an otherwise poorly defended host plant. Damage caused by Polyhymno larval feeding reaches levels which can kill A. cornigera plants. Received: 6 June 1996 / Accepted: 6 September 1996     

Interactions of Bacillus thuringiensis bioinsecticides and the predatory stink bug Podisus nigrispinus to control Plutella xylostella

The diamondback moth (DBM), Plutella xylostella (L.), is a major pest of brassica crops worldwide. Control of this pest is difficult because it rapidly develops resistance to synthetic and biological insecticides and because of the effects of insecticides on its natural enemies. Podisus nigrispinus (Dallas) is a predator that feeds on its prey, as well as on the host plants of its prey, and is an important biological control agent of DBMs. The aim of this study was to determine the susceptibility of P. xylostella larvae to two bioinsecticides: the HD1 strain of Bacillus thuringiensis (B. thuringiensis var. kurstaki) and the commercial product Agree^® (B. thuringiensis var. aizawai CG 91). In addition, the impact of these bioinsecticides on the P. nigrispinus consumption of DBM larvae and phytophagy was evaluated. Both the HD1 strain and Agree^® caused 100% mortality in P. xylostella larvae. P. nigrispinus nymphs fed only with kale leaves (Brassica oleracea var. acephala) sprayed with water, the HD1 strain, or Agree^® did not complete their nymphal development. When prey was also available, P. nigiripinus fed on kale leaves to obtain water. Both nymphs and adults of P. nigrispinus consumed greater numbers of DBM larvae, and fed less on kale leaves, when sprayed with the HD1 strain or Agree^®. These results suggest a positive interaction of B. thuringiensis‐based products and the predator P. nigrispinus in the control of P. xylostella larvae.     

Dissemination of the biocontrol agent Vairimorpha necatrix by the spined soldier bug, Podisus maculiventris

The ability of the spined soldier bug, Podisus maculiventris (Say) (Heteroptera: Pentatomidae), to disseminate infective forms of two lepidopteran pathogens, Vairimorpha necatrix (Kramer) (Microspora: Microsporidia) and Lacanobia oleracea granulovirus (LoGV) was investigated. Individual female P. maculiventris that had fed on Lacanobia oleracea L. (Lepidoptera: Noctuidae) larvae, infected with V. necatrix, excreted approximately 6 × 10⁸V. necatrix spores during the subsequent 7 days. Excreted spores were fed to L. oleracea larvae, causing 100% mortality, indicating that the spores remained viable after passing through the gut of the predator. Podisus maculiventris that had fed on V. necatrix or LoGV‐infected larvae were allowed to defecate on the foliage of tomato plants, prior to the infestation of the plants with L. oleracea or Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) larvae. This proved to be an effective way of infecting the pest larvae with the pathogens, particularly when five predatory bugs were used per plant. After 20 days, the number of S. littoralis and L. oleracea surviving on the plants was reduced by 75% and 61%, respectively. Female P. maculiventris maintained on V. necatrix‐infected prey showed reduced egg production and longevity, whilst those fed on LoGV‐infected prey showed only reduced egg production. The potential for P. maculiventris to disseminate insect pathogens is discussed in the context of improved biological control of lepidopteran pests.     

Detrimental effects of plant compounds on a polyembryonic parasitoid are mediated through its highly polyphagous herbivore host

Plant defensive compounds can have sometimes severe deleterious effects on both herbivores and their natural enemies. Iridoid glycosides (IGs) are defensive compounds that are well established as deterrent to several generalist herbivores and generalist predators. Trichoplusia ni Hübner (Lepidoptera: Noctuidae) is exceptional among generalist herbivores for its ability to tolerate and thrive when feeding upon IG‐producing plant species; however, it is not known whether the compounds themselves have a harmful effect on T. ni and whether the effects in turn affect its oligophagous endoparasitoid Copidosoma floridanum Ashmead (Hymenoptera: Encyrtidae). To examine these effects, a semi‐purified extract of the IG‐containing plant Plantago lanceolata L. (Plantaginaceae), containing the IGs aucubin and catalpol, was added to artificial diets at 0, 1, 5, or 10% diet dry weight. These diets were fed to both C. floridanum‐parasitized and unparasitized T. ni. Diets higher in IGs tended to be more toxic to both parasitized and unparasitized larvae: host larvae that did survive were slightly smaller and took longer to develop on higher IG diets and total clutch size and survival of the parasitoid C. floridanum were greatly reduced as the host's dietary intake of IGs increased. Only small amounts of aucubin were detected inside the T. ni hemocoel, suggesting that the negative effect of these compounds on C. floridanum is due to nutritional quality of the host being reduced rather than direct toxic effects of the compounds.     

Insect oviposition preference between Epichloë‐symbiotic and Epichloë‐free grasses does not necessarily reflect larval performance

Variation in plant communities is likely to modulate the feeding and oviposition behavior of herbivorous insects, and plant‐associated microbes are largely ignored in this context. Here, we take into account that insects feeding on grasses commonly encounter systemic and vertically transmitted (via seeds) fungal Epichloë endophytes, which are regarded as defensive grass mutualists. Defensive mutualism is primarily attributable to alkaloids of fungal origin. To study the effects of Epichloë on insect behavior and performance, we selected wild tall fescue (Festuca arundinacea) and red fescue (Festuca rubra) as grass–endophyte models. The plants used either harbored the systemic endophyte (E+) or were endophyte‐free (E?). As a model herbivore, we selected the Coenonympha hero butterfly feeding on grasses as larvae. We examined both oviposition and feeding preferences of the herbivore as well as larval performance in relation to the presence of Epichloë endophytes in the plants. Our findings did not clearly support the female's oviposition preference to reflect the performance of her offspring. First, the preference responses depended greatly on the grass–endophyte symbiotum. In F. arundinacea, C. hero females preferred E+ individuals in oviposition‐choice tests, whereas in F. rubra, the endophytes may decrease exploitation, as both C. hero adults and larvae preferred E? grasses. Second, the endophytes had no effect on larval performance. Overall, F. arundinacea was an inferior host for C. hero larvae. However, the attraction of C. hero females to E+ may not be maladaptive if these plants constitute a favorable oviposition substrate for reasons other than the plants' nutritional quality. For example, rougher surface of E+ plant may physically facilitate the attachment of eggs, or the plants offer greater protection from natural enemies. Our results highlight the importance of considering the preference of herbivorous insects in studies involving the endophyte‐symbiotic grasses as host plants.