A recently acquired host plant provides an oligophagous insect herbivore with enemy-free space

Enemy-free space (EFS) is a potentially important factor affecting host plant use by phytophagous insects. Yet only a few field studies have demonstrated that natural enemy activity is the sole mechanism underlying use of novel host plants by herbivorous insects. This may be due to the fact that in earlier studies, both herbivores and natural enemies had the opportunity to adapt to the new host plant. Here we studied the possibility that EFS underlies the recently recorded increase in Phthorimaea operculella densities on tomato plants in a few areas within its geographical range. Through field experiments in Ethiopia, we show that all three conditions proposed by Berdegue et al. to demonstrate EFS are fulfilled. First, a significantly higher proportion of larvae survive on caged than on exposed potato plants, showing that natural enemies are an important mortality factor on the original host, potato. Second, larval survival was significantly higher on exposed tomato than potato plants, implying greater protection for the herbivore from its natural enemies on tomato than on potato plants. Thus tomato plants provide P. operculella with an EFS. Finally, larval survival was significantly higher on caged potato than on caged tomato plants at the preblossom stage, indicating that, in the absence of natural enemies, there is a fitness cost when larvae feed on the sub-optimal tomato plants. Fulfillment of this third condition points to the importance of natural enemy activity relative to that of other unidentified factors, such as food quality and competition. An intensive field survey provides further support for this conclusion.     

Use of an exotic host plant reduces viral burden in a native insect herbivore

Incorporation of exotic plants into the diets of native herbivores is a common phenomenon, influencing interactions with natural enemies and providing insight into the tritrophic costs and benefits of dietary expansion. We evaluated how use of an exotic plant, Plantago lanceolata, impacted immune performance, development and susceptibility to pathogen infection in the neotropical herbivore Anartia jatrophae (Lepidoptera: Nymphalidae). Caterpillars were reared on P. lanceolata or a native plant, Bacopa monnieri, and experimentally infected with a pathogenic virus, Junonia coenia densovirus. We found that virus-challenged herbivores exhibited higher survival rates and lower viral burdens when reared on P. lanceolata compared to B. monnieri, though immune performance and development time were largely similar on the two plants. These findings reveal that use of an exotic plant can impact the vulnerability of a native herbivore to pathogen infection, suggesting diet-mediated protection against disease as a potential mechanism facilitating the incorporation of novel resources.     

A plant pathogen causes extensive mortality in an invasive insect herbivore

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Complex interactions between a plant pathogen and insect parasitoid via the shared vector-host: consequences for host plant infection

Plant viruses modify the development of their aphid vectors by inducing physiological changes in the shared host plant. The performance of hymenopterous parasitoids exploiting these aphids can also be modified by the presence of the plant pathogen. We used laboratory and glasshouse microcosms containing beans (Vicia faba) as the host plant to examine the interactions between a plant virus (pea enation mosaic virus; PEMV) and a hymenopterous parasitoid (Aphidius ervi) that share the aphid vector/host Acyrthosiphon pisum. Neither PEMV-infection of V. faba, nor the carriage of PEMV virions by A. pisum, affected the growth or morphology of the aphid, or the oviposition behaviour and development of A. ervi. The presence of developing Aphidius ervi larvae within Acyrthosiphon pisum did not affect the ability of the aphids to transmit PEMV. However, by reducing their longevity, parasitism ultimately decreased the time viruliferous aphids were able to inoculate plants. In terms of virus dispersal, parasitized aphids exhibited more movement around experimental arenas than unparasitized controls, causing a slight increase in the proportion of beans infected with PEMV. Exposure to adult Aphidius ervi caused Acyrthosiphon pisum to rapidly drop off bean plants and disperse to new hosts, resulting in considerably higher plant infection rates (70%) than that seen in control arenas (25%). The results of this investigation demonstrate that when parasitoids are added to a plant-pathogen-vector system, benefits to the host plant due to reduced herbivore infestation must be balanced against the consequences of parasitoid-induced aphid dispersal and a subsequent increase in the level of plant infection.     

Use of an exotic host plant affects mate choice in an insect herbivore

The colonization of exotic plants by herbivorous insects has provided opportunities for investigating causes and consequences of the evolution of niche breadth. The butterfly Lycaeides melissa utilizes exotic alfalfa, Medicago sativa, which is a relatively poor larval resource, and previous studies have found that caterpillars that consume M. sativa develop into smaller and less fecund adults. Here we investigate the effect of smaller female body size on male mate preference, a previously unexplored consequence of novel host use. Smaller females, which developed on the exotic host, were less likely to be visited by males. This result was confirmed with a second set of choice tests involving females reared on a single plant species, thus ruling out host-specific confounding factors. We suggest that an effect on mate choice be considered part of the complex suite of factors determining persistence of herbivorous insects following colonization of new habitats or resources.     

A plant pathogen modulates the effects of secondary metabolites on the performance and immune function of an insect herbivore

Host plant chemical composition critically shapes the performance of insect herbivores feeding on them. Some insects have become specialized on plant secondary metabolites, and even use them to their own advantage such as defense against predators. However, infection by plant pathogens can seriously alter the interaction between herbivores and their host plants. We tested whether the effects of the plant secondary metabolites, iridoid glycosides (IGs), on the performance and immune response of an insect herbivore are modulated by a plant pathogen. We used the IG‐specialized Glanville fritillary butterfly Melitaea cinxia, its host plant Plantago lanceolata, and the naturally occurring plant pathogen, powdery mildew Podosphaera plantaginis, as model system. Pre‐diapause larvae were fed on P. lanceolata host plants selected to contain either high or low IGs, in the presence or absence of powdery mildew. Larval performance was measured by growth rate, survival until diapause, and by investment in immunity. We assessed immunity after a bacterial challenge in terms of phenoloxidase (PO) activity and the expression of seven pre‐selected insect immune genes (qPCR). We found that the beneficial effects of constitutive leaf IGs, that improved larval growth, were significantly reduced by mildew infection. Moreover, mildew presence downregulated one component of larval immune response (PO activity), suggesting a physiological cost of investment in immunity under suboptimal conditions. Yet, feeding on mildew‐infected leaves caused an upregulation of two immune genes, lysozyme and prophenoloxidase. Our findings indicate that a plant pathogen can significantly modulate the effects of secondary metabolites on the growth of an insect herbivore. Furthermore, we show that a plant pathogen can induce contrasting effects on insect immune function. We suspect that the activation of the immune system toward a plant pathogen infection may be maladaptive, but the actual infectivity on the larvae should be tested.     

Plant polyploidy and host expansion in an insect herbivore

Polyploidization has played an essential role in the diversification of seed plants and often has profound effects on plant physiology and morphology. Yet, little is known about how plant polyploidization has shaped the ecology and evolution of interactions between phytophagous insects and their hosts. Polyploidization could either facilitate or impede colonization of new hosts. Greya politella (Lepidoptera: Prodoxidae) is highly specialized on plants in the genus Lithophragma (Saxifragaceae) throughout most of its geographic range. In central Idaho, some populations have shifted to the related Heuchera grossulariifolia, a plant that has repeatedly undergone autopolyploidization. Previous studies have shown that populations feeding natively on H. grossulariifolia prefer tetraploids to diploids in naturally mixed stands. We investigated whether this difference is caused by an inherent preference for tetraploids, or if the preference in present Heuchera-feeding populations has evolved over time. Moths from a strictly Lithophragma-feeding population were tested for preference of diploid or tetraploid H. grossulariifolia, using a combination of field experiments and caged choice trials. In all trials, attack rates on these non-hosts were very low, with no significant difference between ploidies. In addition, there was little evidence that females manipulated their clutch sizes when ovipositing into different plant species or ploidy levels. Hence, the local shift from Lithophragma to Heuchera in central Idaho is not due to failure of the moths to discriminate between these plant species. Furthermore, the higher attack rates on tetraploids in native H. grossulariifolia-feeding populations cannot be caused by a higher initial preference for these plants, but must instead be a result of differences in plant phenology and/or selection acting on local populations.     

Behavioral adaptations increase the value of enemy-free space for Heliothis subflexa,a specialist herbivore

We investigated the importance of specialized behaviors in the use of enemy-free space by comparing the host-use behavior of two closely related moths, Heliothis subflexa Guenee and H. virescens Fabricius. Heliothis subflexa is a specialist on plants in the genus Physalis, whereas H. virescens is an extreme generalist, feeding on plants in at least 14 families. Heliothis subflexa uses the inflated calyx surrounding Physalis fruits as enemy-free space, and field rates of parasitism for H. subflexa on Physalis are much lower than for H. virescens on tobacco and cotton, common hosts found in the same habitat as Physalis. If Physalis, architecture were solely responsible for H. subflexa's low rates of parasitism on Physalis, we predicted that H. virescens larvae experimentally induced to feed on Physalis would experience parasitism rates similar to those of H. subflexa. We found, however, that specialized host-use and host-acceptance behaviors are integral to the use of enemy-free space on Physalis and strongly augment the effects of the structural refuge. In laboratory assays, we found considerable differences between the larval behavior of the specialist. H. subflexa, and the generalist, H. virescens, and these contributed to H. subflexa's superior use of enemy-free space on Physalis. We tested the importance of these behavioral differences in the field by comparing parasitism of H. virescens on Physalis, H. virescens on tobacco, and H. subflexa on Physalis by Cardiochiles nigriceps Vierick, a specialist braconid parasitoid. For H. virescens, a threefold decrease in parasitism occurred when feeding on Physalis (mean parasitism +/- SEM = 13 +/- 4%) rather than tobacco (43 +/- 4%), a difference we attribute to the structural refuge provided by Physalis. However, parasitism of H. virescens on Physalis was more than ten times as great as that of H. subflexa on Pliv.salis (1 +/- 4%), supporting the hypothesis that specialized behaviors have a substantial impact on use of Physalis as enemy-free space. Behavioral adaptations may be central to the use of enemy-free space by phytophagous insects and may act as an important selective force in the evolution of dietary specialization.     

A fungal root symbiont modifies plant resistance to an insect herbivore

Vesicular-arbuscular mycorrhizal (VAM) fungi are common root-colonizing symbionts that affect nutrient uptake by plants and can alter plant susceptibility to herbivores. I conducted a factorial experiment to test the hypotheses that colonization by VAM fungi (1) improves soybean (Glycine max) tolerance to grazing by folivorous Mexican bean beetle (Epilachna varivestis), and (2) indirectly affects herbivores by increasing host resistance. Soybean seedlings were inoculated with the VAM fungus Glomus etunicatum or VAM-free filtrate and fertilized with high-[P] or low-[P] fertilizer. After plants had grown for 7 weeks first-instar beetle larvae were placed on bagged leaves. Growth of soybean was little affected by grazing larvae, and no effects of treatments on tolerance of soybeans to herbivores were evident. Colonization by VAM fungus doubled the size of phosphorus-stressed plants but these plants were still half the size of plants given adequate phosphorus. High-[P] fertilizer increased levels of phosphorus and soluble carbohydrates, and decreased levels of soluble proteins in leaves of grazed plants. Colonization of grazed plants by VAM fungus had no significant effect on plant soluble carbohydrates, but increased concentration of phosphorus and decreased levels of proteins in phosphorus-stressed plants to concentrations similar to those of plants given adequate phosphorus. Mexican bean beetle mass at pupation, pupation rate, and survival to eclosion were greatest for beetles reared on phosphorus-stressed, VAM-colonized plants, refuting the hypothesis that VAM colonization improves host plant resistance. VAM colonization indirectly affected performance of Mexician bean beetle larvae by improving growth and nutrition of the host plant. Received: 28 February 1997 / Accepted: 23 June 1997     

Effects of light availability on host plant chemistry and the consequences for behavior and growth of an insect herbivore

We examined how light availability influenced the defensive chemistry of tomato (Lycopersicon esculentum: Solanaceae). Tomato plants were grown either in full sunlight or under shade cloth rated at 73%. Leaves from plants grown in full sunlight were tougher, had higher concentrations of allelochemicals (chlorogenic acid, rutin and tomatine), and had less protein than leaves from plants grown in shade. We determined how these differences in host plant quality due to light availability affected the behavior and growth of a Solanaceae specialist, Manduca sexta. Both in the greenhouse and in the field, caterpillars on shade-grown plants grew heavier in a shorter amount of time than those on plants that had previously been grown in full sunlight. In contrast, the effects of previous light availability to plants on caterpillar behavior appeared to be minor.To further investigate how light availability to plants influenced herbivore growth, we examined the effects of leaf-powder diets made from tomato leaves of different ages (new, intermediate, or mature) grown in full sunlight or shade on caterpillar performance. Caterpillars fed diets made from plants grown in shade consumed less but grew faster than larvae fed diets made from tomato plants grown in full sunlight. Caterpillars fed diets made from new leaves grew larger in less time than caterpillars fed diets made from intermediate aged leaves. Caterpillars did not survive on the mature leaf powder diets. There were plant-light treatment by larval thermal regime interactions. For example, at 26:15 °C , plant-light treatment had no effect on stadium duration, but at 21:10 °C, stadium duration was prolonged with the full sunlight-new leaf diet compared with the shaded-new leaf diet. In a second diet experiment, we examined the interactive effects of protein and some tomato allelochemicals (rutin, chlorogenic acid and tomatine) on the performance of caterpillars. There were food quality by thermal regime interactions. For instance, at 26:15 °C , neither protein nor allelochemical concentration influenced stadium duration, whereas at 21:10 °C, stadium duration was prolonged with the low protein-high allelochemical diet, which simulated full sunlight leaves. In sum, light availability to plants affected defensive chemistry and protein concentration. The difference in food quality was great enough to influence the growth of a specialist insect herbivore, but the effects were temperature-dependent.     

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     

Trade-offs underlying polyphagy in a facultative ant-tended florivorous butterfly: the role of host plant quality and enemy-free space

The underlying mechanisms mediating the use of multiple host plants were investigated in Parrhasius polibetes (Lycaenidae), a florivorous and facultative myrmecophilous butterfly. Plant traits such as presence of ant–treehopper associations as a source of enemy-free space, flower bud dimensions, toughness, thickness, trichomes, and the corresponding performance and wear of P. polibetes mandibles were examined for three natural hosts: Schefflera vinosa (Araliaceae), Pyrostegia venusta (Bignoniaceae) and Luehea grandiflora (Malvaceae). Parasitism levels of larvae found on the three hosts were also determined. Almost all Luehea had ant–treehopper associations, and all larvae found on this host were non-parasitized. Parasitism was low in larvae found on Schefflera, half of which hosted ant–treehopper associations. No ant–treehopper association was found on Pyrostegia, where parasitism was significantly higher compared to other hosts. In the laboratory, P. polibetes performed well on Schefflera, followed by Pyrostegia. No larvae survived when fed with Luehea. Flower buds of Luehea were thicker and tougher than those of Schefflera and Pyrostegia. Indeed, mandibles of larvae reared on Luehea showed substantial wear, whereas those reared either on Schefflera or Pyrostegia presented no significant damage. Additionally, we suggest that co-occurrence with ant–treehopper associations on a plant provides parasitoid-free space for P. polibetes larvae. Our results support the hypothesis that ecological trade-offs among host plants (i.e., food quality and enemy-free space) promote polyphagy in natural populations of P. polibetes. Host morphological traits seem to play a relevant role in P. polibetes performance. To our knowledge, this is the first report showing the costs of polyphagy in a myrmecophilous butterfly.     

Context-dependent transmission of a generalist plant pathogen: host species and pathogen strain mediate insect vector competence

The specificity of pathogen–vector–host interactions is an important element of disease epidemiology. For generalist pathogens, different pathogen strains, vector species, or host species may all contribute to variability in disease incidence. One such pathogen is Xylella fastidiosa Wells et al., a xylem-limited bacterium that infects dozens of crop, ornamental, and native plants in the USA. This pathogen also has a diverse vector complex and multiple biologically distinct strains. We studied the implications of diversity in this pathogen–vector–host system, by quantifying variability in transmission efficiency of different X. fastidiosa strains (isolates from almond and grape genetic groups) for different host plants (grape, almond, and alfalfa) by two of the most important vectors in California: glassy-winged sharpshooter [ Homalodisca vitripennis (Germar)] and green sharpshooter ( Draeculacephala minerva Ball) (both Hemiptera: Cicadellidae). Transmission of isolates of the almond strain by H. vitripennis did not differ significantly, whereas transmission varied significantly among isolates from the grape strain (15–90%). Host plant species did not affect H. vitripennis transmission. Conversely, D. minerva efficiency was mediated by both host plant species and pathogen strain. No acquisition of an almond isolate occurred regardless of plant type (0/122), whereas acquisition of a grape isolate from alfalfa was 10-fold higher than from grape or almond plants. These results suggest that pathogen, vector, and host diversity impose contingencies on the transmission ecology of this complex disease system. Studies aimed at the development of management strategies for X. fastidiosa diseases should consider the complexity of these interactions as they relate to disease spread.     

Transgenerational acclimatization in an herbivore–host plant relationship

Twenty years ago, scientists began to recognize that parental effects are one of the most important influences on progeny phenotype. Consequently, it was postulated that herbivorous insects could produce progeny that are acclimatized to the host plant experienced by the parents to improve progeny fitness, because host plants vary greatly in quality and quantity, and can thus provide important cues about the resources encountered by the next generation. However, despite the possible profound implications for our understanding of host-use evolution of herbivores, host-race formation and sympatric speciation, intense research has been unable to verify transgenerational acclimatization in herbivore–host plant relationships. We reared Coenonympha pamphilus larvae in the parental generation (P) on high- and low-quality host plants, and reared the offspring (F₁) of both treatments again on high- and low-quality plants. We tested not only for maternal effects, as most previous studies, but also for paternal effects. Our results show that parents experiencing predictive cues on their host plant can indeed adjust progeny''s phenotype to anticipated host plant quality. Maternal effects affected female and male offspring, whereas paternal effects affected only male progeny. We here verify, for the first time to our knowledge, the long postulated transgenerational acclimatization in an herbivore–host plant interaction.     

Geography is more important than host plant use for the population genetic structure of a generalist insect herbivore

Population divergence can occur due to mechanisms associated with geographic isolation and/or due to selection associated with different ecological niches. Much of the evidence for selection‐driven speciation has come from studies of specialist insect herbivores that use different host plant species; however, the influence of host plant use on population divergence of generalist herbivores remains poorly understood. We tested how diet breadth, host plant species and geographic distance influence population divergence of the fall webworm (Hyphantria cunea; FW). FW is a broadly distributed, extreme generalist herbivore consisting of two morphotypes that have been argued to represent two different species: black‐headed and red‐headed. We characterized the differentiation of FW populations at two geographic scales. We first analysed the influence of host plant and geographic distance on genetic divergence across a broad continental scale for both colour types. We further analysed the influence of host plant, diet breadth and geographic distance on divergence at a finer geographic scale focusing on red‐headed FW in Colorado. We found clear genetic and morphological distinction between red‐ and black‐headed FW, and Colorado FW formed a genetic cluster distinct from other locations. Although both geographic distance and host plant use were correlated with genetic distance, geographic distance accounted for up to 3× more variation in genetic distance than did host plant use. As a rare study investigating the genetic structure of a widespread generalist herbivore over a broad geographic range (up to 3,000 km), our study supports a strong role for geographic isolation in divergence in this system.     

On ecological fitting, plant–insect associations, herbivore host shifts, and host plant selection

"Pests soon colonize plants that are cultivated extensively, plant species recruit different pest species in different regions, and associations of insects with plants are often more casual, fortuitous, and labile than those usually interpreted as coevolutionary." ( Strong 1979 , pp. 89)

"…  when a parasite arrives in a new habitat, it will feed on those species whose defense traits it can circumvent because of the abilities it carries at the time. Such a parasite cannot be distinguished from one that evolved the ability to circumvent a defense while in trophic contact with its host." ( Janzen 1980 , pp. 611)

"We believe that a reasonable null hypothesis  …  is that many associations between insects and plants can occur without much evolution…" (Rey, McCoy and Strong 1981, pp. 620)

"The main role of secondary plant substances in insect/host plant relationships is that they form the 'fingerprint'  …  by which the insect recognizes the plants  …  The recognition of a plant as host is unrelated to whether the plant and the insect have evolved together or whether they meet for the first time in their evolutionary history." ( Jermy 1984 , pp. 620)

     

The nutritional landscape of host plants for a specialist insect herbivore

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Adaptation in an insect host–plant pathogen interaction

Selection on parasites to adapt to local host populations may be direct or through other components of the system such as vectors or the food plant on which the parasite is ingested. To test for local adaptation of nucleopolyhedrovirus among island populations of western tent caterpillars, Malacosoma californicum pluviale, we compared virus isolates from three geographically distinct sites with different dominant host plants. Pathogenicity, speed of kill and virus production of each isolate were examined on the three food plants. Virus isolates from the two permanent host populations had the fastest speed of kill on the host plant from which they were isolated. This was not the case for a caterpillar population that goes extinct when populations are regionally low. Virus isolates on some plant species combined rapid speed of kill with high virus yield. Infection of hosts by mixed microparasite populations could facilitate local adaptation in response to differing food plant chemistry.     

Realised toxicity of plant defences to an insect herbivore depends more on insect dehydration than on energy reserves

1. Climate effects on plant defences, and subsequently insect herbivores, have received considerable attention. However, climate also directly affects the physiological condition of insects that may influence their ability to combat plant defences. 2. This study tested the effects of body fat and water content on the ability of mountain pine beetles, Dendroctonus ponderosae Hopkins, to tolerate monoterpenes, the primary defences of their host pine trees. To manipulate their physiological condition, we exposed beetles to one of three environmental treatments: cold (4 °C, 30% RH), humid (21 °C, 70% RH) or dry (21 °C, 30% RH). Beetles had the highest fat and water content in the cold treatment, followed by lower fat but the same water content in the humid treatment, and the lowest of both in the dry treatment. Following environmental treatments, beetles were exposed to 0, 312.5 or 625 ppm vapours of monoterpenes (−)-α-pinene or (R)-(+)-limonene. 3. Survival during both environmental and monoterpene treatments was highest for the cold treatment, less for the humid treatment, and lowest for the dry treatment. Fat and water content positively predicted survival. In response to increasing monoterpene concentration, survival, body condition and water content, but not fat, declined. 4. These results indicate that water content of insect herbivores was important for the realised toxicity of plant defences, presumably due to the process of excreting detoxified compounds. Energetic costs of detoxification were less evident. Dehydration of insects has not been widely considered in insect–plant interactions, but effects of climate change on humidity may become increasingly important for these interactions.