Impact of acidic deposition onEncelia farinosa gray (Compositae: Asteraceae) and feeding preferences ofTrirhabda geminata horn (Coleoptera: Chrysomelidae)

Container grownEncelia farinosa were exposed to three 3-hr episodes of acidic fog (pH 2.5) typical of events in southern California. Adults and larvae of the specialist leaf-feeding herbivore,Trirhabda geminata, preferred to feed on the acidic-treated foliage compared to control fogged (pH 6.3–6.5) foliage. Previous feeding damage on the plants did not affect feeding preference. The acidic-fogged foliage was significantly higher in total nitrogen and soluble protein but not different from control-treated tissue in water content. Stress on native populations of this drought-deciduous shrub caused by atmospheric pollutants may also result in altered feeding ecology of the beetle.     

Fate of the chromene encecalin in the interaction ofEncelia farinosa and its specialized herbivoreTrirhabda geminata

Leaf beetles of the speciesTrirhabda geminata are specialized herbivores that are able to feed on the chemically well-protected foliage of the desert sunflowerEncelia farinosa, which contains the insecticidal chromene derivative encecalin. Chemical analysis of the beetles and their fecal excretions indicated that encecalin is present only in the alimentary canal and is not absorbed across the gut membrane, as previously shown for other herbivorous insects (e.g., the Egyptian armyworm,Spodoptera littoralis) that are susceptible to this chromene derivative. Further differences betweenT. geminata and nonadapted insects were observed with regard to the metabolism of encecalin. Whereas the encecalin-resistent leaf beetles metabolize encecalin mainly to encecalol by reduction of the acetyl group, susceptible insects, such as larvae ofS. littoralis, metabolize encecalin mainly by exoxidation of the 3,4 double bond, which creates a powerful alkylating agent and is responsible for the toxicity of encecalin. Reductive rather than oxidative metabolism of encecalin therefore seems important for the resistance ofT. geminata against the chemical defense of their host plantE. farinosa.     

Interactions Among Insect Herbivore Guilds: Influence of Thrips Bud Injury on Foliar Chemistry and Suitability to Gypsy Moths

This study investigated the consequences of early season bud herbivory on host-plant phytochemistry and subsequent effects on a later mid-season leaf-feeding herbivore, to test the hypothesis that temporally segregated interguild interactions could affect herbivore success through plant-mediated responses. Our system consisted of American bass wood, Tilia americana, a bud-feeding thrips species, Thrips calcaratus, and the folivorous gypsy moth, Lymantria dispar. The impact of thrips bud-feeding on American basswood foliar chemistry and subsequent effects on gypsy moth larval preference and performance were measured. Foliar total nonstructural carbohydrates increased and phenolic levels decreased in response to bud injury, which affected larval feeding preference. In a two-choice test, gypsy moth larvae preferred leaf discs with high carbohydrate and low phenolic levels. The effects on larval performance depended on the extent of prior bud injury and were correlated with carbohydrate concentrations. In an early season assay, larval performance was lowest on moderately bud-damaged tissue, which also had the lowest total nonstructural carbohydrates. In a mid-season assay, larval performance and carbohydrate concentrations were highest in severely bud-damaged foliage. Foliar phenolics were highest in severely bud-damaged tissue in the early season assay, and in moderately damaged tissue in the mid-season assay. Gypsy moth performance was not correlated with foliar phenolic levels. Secondary (reflushed) foliage had higher carbohydrate levels than did primary (original) foliage, which correlated with increased larval performance. This study illustrates that bud-feeding herbivores can alter the phytochemistry and subsequent suitability of host-plant foliage for later folivores. The implications of these results to interactions between feeding guilds, community structure, and forest health are discussed.     

Rapid Herbivore-Induced Changes in Mountain Birch Phenolics and Nutritive Compounds and Their Effects on Performance of the Major Defoliator, Epirrita autumnata

Insect damage changes plant physiology and chemistry, and such changes may influence the performance of herbivores. We introduced larvae of the autumnal moth (Epirrita autumnataBorkh.) on individual branches of its main host plant, mountain birch (Betula pubescens ssp. czerepanovii (Orlova) Hämet-Ahti) to examine rapid-induced plant responses, which may affect subsequent larval development. We measured systemic responses to herbivory by analyzing chemistry, photosynthesis, and leaf growth, as well as effects on larval growth and feeding, in undamaged branches of damaged and control trees. Larvae reared on leaves from intact branches of the herbivore-damaged trees grew faster than those reared on leaves of control trees, indicating systemic-induced susceptibility. Herbivore damage did not lead to systemic changes in levels of primary nutrients or phenolic compounds. The analyses of photosynthetic activity and individual hydrolyzable tannins revealed a reversal of leaf physiology-herbivore defense patterns. On control trees, consumption by E. autumnata larvae was positively correlated with photosynthetic activity; on damaged trees, this correlation was reversed, with consumption being negatively correlated with photosynthetic activity. A similar pattern was found in the relationship between monogalloylglucose, the most abundant hydrolyzable tannin of mountain birch, and leaf consumption. Among the control trees, consumption was positively correlated with concentrations of monogalloylglucose, whereas among herbivore-damaged trees, this correlation was reversed and became negative. Our results suggest that herbivore performance is related to both concentrations of phenolic compounds and photosynthetic activity in leaves. This linkage between herbivore performance, leaf chemistry, and physiology was sensitive to induced plant responses caused by slight herbivore damage.     

Inhibition ofCampoletis sonorensis parasitism ofHeliothis zea and of parasitoid development by 2-tridecanone-mediated insect resistance of wild tomato

Field populations ofHeliothis spp. were sampled for levels of naturally occurring larval parasitism on six tomato lines varying in levels of 2-tridecanone-mediated resistance toManduca sexta (L.) andLeptinotarsa decemlineata (Say). Second and third instars were parasitized byCampoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae) andCotesia (=Apantales)marginiventris (Cresson) (Hymenoptera: Braconidae) in 1984 through 1986 and byMicropletis croceipes (Cresson) (Hymenoptera: Braconidae) in 1986. Differences in parasitism by individual and multiple species among host plants were not demonstrated. However, levels of parasitism were low and variable among replicates. Total larval parasitism averaged across all plant lines was less than 6% in 1984 and 1986 and approximately 11% in 1985. In laboratory cage studies,C. sonorensis parasitized fewerH. zea larvae on tomato foliage with high levels of 2-tridecanone than on foliage with low levels. RearingH. zea on diet containing 2-tridecanone and 2-undecanone did not alter incidence of parasitism byC, sonorensis; nor did rearing parasitizedH. zea larvae on chemically treated host diets precondition the parasitoid to higher or lower mortality when transferred to foliage as a substrate for cocoon spinning, regardless of the foliage genotype. However, parasitoid survival during cocoon spinning on foliage varied significantly among plant lines in a manner corresponding to the level of 2-tridecanone-mediated resistance of the foliage. Parasitoid mortality was greatest on highly resistant foliage and lowest on susceptible foliage.     

Tolerance to Thiarubrines from Ambrosia chamissonis by the Rare Moth Lasionycta wyatti

The larvae of a rare noctuid moth, Lasionycta wyatti, were found to be associated with the stems and roots of Ambrosia chamissonis, a species found along sandy beaches in western North America. High-performance liquid chromatographic (HPLC) analyses of L. wyatti larvae and their frass revealed significant amounts of thiarubrines, secondary compounds characteristic of this host plant. Artificial diet studies showed that thiarubrines were well tolerated by these larvae. These same diets, however, were toxic to larvae of Manduca sexta and inhibited larval growth of Spodoptera litura. Even in the presence of simulated sunlight, artificial diets containing thiarubrines did not affect larvae of L. wyatti. However, dietary incorporation of thiophenes, natural UV-activated breakdown products of thiarubrines, was deleterious to larvae of L. wyatti.     

Iridoid glycosides and host-plant specificity in larvae of the buckeye butterfly,Junonia coenia (Nymphalidae)

Larvae of the buckeye,Junonia coenia (Nymphalidae) feed primarily on plants in four families: Scrophulariaceae, Plantaginaceae, Verbenaceae, and Acanthaceae. These plant families have in common the presence of a group of plant secondary compounds, the iridoid glycosides. Larvae were reared on three plant species and two artificial diets, one with and one without iridoid glycosides.Larvae grew poorly and had low survivorship on the artificial diet without iridoid glycosides, while growth and survival on the artificial diet with iridoid glycosides was comparable to that on plants. Choice tests using artificial diets with and without iridoid glycosides showed that larvae: (1) chose diets with iridoid glycosides (in the form of a crude extract or pure compound) over a diet without; (2) showed no preference between the diet with the crude extract and that with pure iridoid glycoside, and (3) preferred the artificial diet with ground leaves of the host plant,Plantago lanceolata, over the diet with pure iridoid glycosides. The artificial diet that larvae had been reared on prior to these tests had no effect on subsequent larval preferences in the choice tests.     

Quantitative Variability of Direct Chemical Defense in Primary and Secondary Leaves of Lima Bean (<Emphasis Type="Italic">Phaseolus lunatus</Emphasis>) and Consequences for a Natural Herbivore

Ontogenetic variability in chemical plant defenses against herbivores is a common phenomenon, but the effects of this variability on herbivore–plant interactions are little understood. In a previous study on lima bean (Phaseolus lunatus), we found a trade-off between cyanogenesis, a direct defense, and the release of herbivore-induced volatile organic compounds (VOCs; mainly functioning as an indirect defense). Moreover, the expression of these two defenses could change during plant ontogeny. The present study aimed at elucidating whether such ontogenetic changes in plant defense can affect herbivore–plant interactions. We quantified feeding rates of a natural insect herbivore, the Mexican bean beetle (Epilachna varivestis), on primary and secondary leaves of individual lima bean plants. These insects strongly preferred low cyanogenic primary leaves over high cyanogenic secondary leaves. Although weakly defended by cyanogenesis, lima beans’ primary leaves showed protein concentrations and photosynthetic activities that did not differ significantly from secondary leaves at the time of analysis. Based on our findings, we suggest that lima beans’ long-lived primary leaves function as efficient source organs, even beyond the stage of seedlings. This hypothesis may explain why primary leaves express a strong indirect defense by the release of herbivore induced VOCs.     

Similarity of Cuticular Lipids Between a Caterpillar and Its Host Plant: A Way to Make Prey Undetectable for Predatory Ants?

Ithomiine butterflies (Nymphalidae) have long-lived, aposematic, chemically protected adults. However, little is known about the defense mechanisms in larvae and other juvenile stages. We showed that larvae Mechanitis polymnia are defended from ants by a chemical similarity between their cuticular lipids and those of the host plant, Solanum tabacifolium (Solanaceae). This is a novel defense mechanism in phytophagous insects. A field survey during one season showed that larval survivorship was up to 80%, which is high when compared with other juvenile stages. In a laboratory bioassay, live larvae on their host plant were not attacked by the predatory ant Camponotus crassus (Formicidae). Two experiments showed that the similarity between the cuticular lipids of M. polymnia and S. tabacifolium protected the larvae from C. crassus: (a) when the caterpillar was switched from a host plant to a non-host plant, the predation rate increased, and (b) when a palatable larva (Spodoptera frugiperda, Noctuidae) was coated with the cuticular lipids of M. polymnia and placed on S. tabacifolium leaves, it no longer experienced a high predation rate. This defensive mechanism can be defined as chemical camouflage, and may have a double adaptive advantage, namely, protection against predation and a reduction in the cost of sequestering toxic compounds from the host plant.     

<Emphasis Type="Italic">Barbarea vulgaris</Emphasis> Glucosinolate Phenotypes Differentially Affect Performance and Preference of Two Different Species of Lepidopteran Herbivores

The composition of secondary metabolites and the nutritional value of a plant both determine herbivore preference and performance. The genetically determined glucosinolate pattern of Barbarea vulgaris can be dominated by either glucobarbarin (BAR-type) or by gluconasturtiin (NAS-type). Because of the structural differences, these glucosinolates may have different effects on herbivores. We compared the two Barbarea chemotypes with regards to the preference and performance of two lepidopteran herbivores, using Mamestra brassicae as a generalist and Pieris rapae as a specialist. The generalist and specialist herbivores did not prefer either chemotype for oviposition. However, larvae of the generalist M. brassicae preferred to feed and performed best on NAS-type plants. On NAS-type plants, 100% of the M. brassicae larvae survived while growing exponentially, whereas on BAR-type plants, M. brassicae larvae showed little growth and a mortality of 37.5%. In contrast to M. brassicae, the larval preference and performance of the specialist P. rapae was unaffected by plant chemotype. Total levels of glucosinolates, water soluble sugars, and amino acids of B. vulgaris could not explain the poor preference and performance of M. brassicae on BAR-type plants. Our results suggest that difference in glucosinolate chemical structure is responsible for the differential effects of the B. vulgaris chemotypes on the generalist herbivore. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of iridoid glycoside content on oviposition host plant choice and parasitism in a specialist herbivore

The Glanville fritillary butterfly Melitaea cinxia feeds upon two host plant species in Å land, Finland, Plantago lanceolataand Veronica spicata, both of which produce iridoid glycosides. Iridoids are known to deter feeding or decrease the growth rate of many generalist insect herbivores, but they often act as oviposition cues to specialist butterflies and are feeding stimulants to their larvae. In this study, two iridoid glycosides (aucubin and catalpol) were analyzed by micellar electrokinetic capillary chromatography. We measured the spatial and temporal variation of iridoid glycosides in natural populations of the host plants of M. cinxia. We also analyzed the aucubin and catalpol content in plants in relation to their use by ovipositing females, and in relation to the incidence of parasitism of M. cinxia larvae in natural populations. The mean concentrations of aucubin and catalpol were higher in P. lanceolata than in V. spicata, and catalpol concentrations were higher than aucubin concentrations in both host species. Plantago lanceolata individuals that were used for oviposition by M. cinxia had higher aucubin concentrations than random plants and neighboring plants. Additionally, oviposition and random plants had higher catalpol concentrations than neighboring plants, indicating that ovipositing females select for high iridoid glycoside plants or that oviposition induces iridoid glycoside production in P. lanceolata. Parasitism by the specialist parasitoid wasp Cotesia melitaearum occurred most frequently in larval groups that were feeding on plants with low concentrations of catalpol, irrespective of year, population, and host plant species. Therefore, parasitoids appear to avoid or perform poorly in host larvae with high catalpol content.     

Induction of Systemic Acquired Resistance in Cotton Foliage Does Not Adversely Affect the Performance of an Entomopathogen

Baculoviral efficacy against lepidopteran larvae is substantially impacted by the host plant. Here, we characterized how baculoviral pathogenicity to cotton-fed Heliothis virescens larvae is affected by induction of systemic acquired resistance (SAR). Numerous studies have shown that SAR induced by the plant elicitor benzo-(1,2,3)-thiadiazole-7-carbothioic acid S-methyl ester (BTH) can protect against plant pathogens, but reports on the impacts of SAR on chewing herbivores or on natural enemies of herbivores are few. We found that BTH application significantly increased foliar peroxidase activity, condensed tannin levels, and total phenolic levels but did not alter dihydroxyphenolic levels. Consumption of BTH-treated foliage did not influence H. virescens pupal weight or larval mortality by the microbial control agent Autographa californica multiple nucleopolyhedrovirus any more than did consumption of untreated foliage. Thus, activation of SAR, although it did not protect the plant against a chewing herbivore, also did not reduce the effect of a natural enemy on a herbivore, indicating that SAR and microbial control agents may be compatible components of integrated pest management.     

Influence of cnicin,a sesquiterpene lactone ofCentaurea maculosa (Asteraceae), on specialist and generalist insect herbivores

The sesquiterpene lactone cnicin was extracted fromCentaurea maculosa andCentaurea vallesiaca. We examined its effects on the ovipositional response and larval development of generalist and specialist insect herbivores associated withC. maculosa. For the oviposition trials, three plant species (C. maculosa, Achillea millefolium, andCichorium intybus), half of which were sprayed with 3% of cnicin, were exposed to the specialist mothsStenodes straminea, Agapeta zoegana, andPterolonche inspersa in field cages. All three species significantly preferredC. maculosa to other plants andP. inspersa significantly preferred cnicin-sprayed plants to untreated plants for oviposition. Tested over all species, cnicin significantly increased the number of eggs laid on a given plant. A larval diet test examined the toxicity of cnicin for larvae of the generalist noctuid mothSpodoptera littoralis. Cnicin concentrations of 3% and 6% were lethal and 1% and 0.5% seriously inhibited growth and development. The larvae of theC. maculosa specialistStenodes straminea survived at 6% cnicin, but none of the pupae hatched.Agapeta zoegana was able to survive at 1% and 3% cnicin. Both specialists had difficulties with the artificial diet, but weight increase and survival was not further reduced when cnicin was present compared with on the control diet. In conclusion, cnicin influenced host recognition by the specialist species, and larvae of the generalist did not survive on natural levels of cnicin. Growth and survival of the specialist were not influenced by cnicin but were considerably hampered on artificial diet.     

The Role of Fresh <Emphasis Type="Italic">versus</Emphasis> Old Leaf Damage in the Attraction of Parasitic Wasps to Herbivore-Induced Maize Volatiles

The odor produced by a plant under herbivore attack is often used by parasitic wasps to locate hosts. Any type of surface damage commonly causes plant leaves to release so-called green leaf volatiles, whereas blends of inducible compounds are more specific for herbivore attack and can vary considerably among plant genotypes. We compared the responses of naïve and experienced parasitoids of the species Cotesia marginiventris and Microplitis rufiventris to volatiles from maize leaves with fresh damage (mainly green leaf volatiles) vs. old damage (mainly terpenoids) in a six-arm olfactometer. These braconid wasps are both solitary endoparasitoids of lepidopteran larvae, but differ in geographical origin and host range. In choice experiments with odor blends from maize plants with fresh damage vs. blends from plants with old damage, inexperienced C. marginiventris showed a preference for the volatiles from freshly damaged leaves. No such preference was observed for inexperienced M. rufiventris. After an oviposition experience in hosts feeding on maize plants, C. marginiventris females were more attracted by a mixture of volatiles from fresh and old damage. Apparently, C. marginiventris has an innate preference for the odor of freshly damaged leaves, and this preference shifts in favor of a blend containing a mixture of green leaf volatiles plus terpenoids, after experiencing the latter blend in association with hosts. M. rufiventris responded poorly after experience and preferred fresh damage odors. Possibly, after associative learning, this species uses cues that are more directly related with the host presence, such as volatiles from host feces, which were not present in the odor sources offered in the olfactometer. The results demonstrate the complexity of the use of plant volatiles by parasitoids and show that different parasitoid species have evolved different strategies to exploit these signals.     

Volatiles emitted by different cotton varieties damaged by feeding beet armyworm larvae

Volatile compounds elicited by insect herbivore feeding damage in five cotton cultivars and one naturalized cotton variety were examined by allowing beet armyworm larvae to feed overnight on leaves and collecting volatiles from the plants in situ. Of 23 compounds identified from larval damaged leaves, terpenes and lipoxygenase-hydroperoxide lyase-derived volatiles predominated. No pronounced differences in the levels of volatile emission were noted from leaves of undamaged plants of the different varieties. However, average volatile emission from damaged leaves of the naturalized variety was almost sevenfold higher than from damaged leaves of the commercial cultivars. This was despite the fact that larvae preferred feeding on the leaves of commercial cultivars over those of the naturalized variety in choice tests.     

Influence of foliar glucosinolates in oilseed rape and mustard on feeding and growth of the bertha armyworm,Mamestra configurata Walker

The relationship between host plant glucosinolate profile and feeding and growth of the Bertha armyworm,Mamestra configurata Walker was investigated using eight cultivated rape and mustard varieties. Mean larval weights of neonates reared on intact rosette-stage plants were significantly different on the different species in the orderBrassica juncea <Sinapis alba <B. napus <B. campestris. WhileB. juncea was least preferred,S. alba was significantly more attractive to neonate larvae in choice tests. Relative consumption and growth rates of fourth-instar larvae were also reduced onB. juncea foliage. Other differences were dependent on the plant growth stage. Neonate preference was not correlated to total glucosinolate levels, but rather to the concentrations of isothiocyanate-releasing glucosinolates. However, the relationship between consumption and glucosinolate levels was inconsistent. Relative growth rate was negatively correlated to total glucosinolate content for stage 3 and 4 foliage—mainly due to the concentration of isothiocyanatereleasing glucosinolates. The relative importance of isothiocyanate-releasing glucosinolates was verified by rearing neonates on meridic diets containing equimolar concentrations of sinigrin, its metabolite, allyl isothiocyanate, and indole-3-carbinol, metabolite of 3-indolylmethyl glucosinolate. Sinigrin and allyl isothiocyanate in the diet produced virtually identical negative weight vs. concentration regression lines. No such dose-response effect was observed with indole-3-carbinol. The data suggest that foliar isothiocyanate-releasing glucosinolates may provide some degree of plant protection from polyphagous insects.     

Processing of iridoid glycoside antirrinoside fromMaurandya antirrhiniflora (Scrophulariaceae) byMeris paradoxa (Geometridae) andLepipolys species (Noctuidae)

The iridoid glycoside antirrinoside was found to be sequestered by highly aposematic larvae of the geometrid mothMeris paradoxa and two noctuid mothLepipolys species feeding onMaurandya antirrhiniflora (Scrophulariaceae), a natural food plant from southern Arizona. The antirrinoside content of leaves and petioles being consumed, eariy-instar larvae, late-instar larvae, larval frass, regurgitant, reflex-bleeding emission (Meris paradoxa), cocoons, pupae, meconium emitted upon eclosion, and adult moths was determined. Larvae, other than the earliest instars, did not excrete antirrinoside in the frass, but sequestered it in amounts of 3–11 % of the dry weight. Small amounts of antirrinoside remained in various pupal or cocoon parts and some was emitted in the meconium upon eclosion. The total antirrinoside accounted for was, however, considerably below that expected based upon the remarkably high 20% content of the leaves and petioles being consumed. The adult cryptic moths of both species contained little or no antirrinoside. This is the first report of a natural food plant and larval stages forM. paradoxa and a previously undescribedLepipolys species. It is also the first report of antirrinoside sequestration and utilization by insects.Chemistry of the Scrophulariaceae. 20. Paper 19: Stermitz, F.R., Ianiro, T.T., Robinson, R.D., and Gardner, D.R. 1991.J. Nat. Prod. 54:626–628.     

Herbivore-induced responses in alfalfa (Medicago sativa)

The herbivore-induced response of alfalfa (Medicago sativa) was examined through assays with Spodoptera littoralislarvae and analyses of important secondary substances. In food preference experiments, larvae preferred young undamaged alfalfa plants over plants that had been damaged by feeding larvae 5 and 7 days earlier, while no difference in feeding preferences could be detected 1, 9, and 14 days after damage. This suggests a peak in the herbivore induced resistance of alfalfa approximately one week after initial damage. The induced resistance in young plants was also shown to be systemic, while older flowering plants failed to show increased resistance after defoliation. Larvae gained weight slower and had lower pupal mass when fed damaged alfalfa than when fed undamaged alfalfa. Levels of total saponins were increased in foliage of damaged alfalfa, and detailed analyses of specific saponin components revealed doubled concentrations of 3GlcA,28AraRhaXyl medicagenate (medicagenic acid bidesmoside) and 3GlcAGalRha soyasapogenol B (soyasaponin I). Levels of the flavonoid apigenin (as free aglycone) also were increased in herbivore damaged plants. The herbivore-induced response of alfalfa was significantly weaker than that of cotton: S. littoralis larvae given a choice of undamaged cotton and undamaged alfalfa preferred to feed on cotton, whereas preferences shifted towards alfalfa when plants were damaged.     

Role of Detoxification of Plant Secondary Compounds on Diet Breadth in a Mammalian Herbivore, Trichosurus vulpecula

Theory predicts that mammalian herbivores detoxify different classes of plant secondary compounds via separate metabolic pathways and that generalist herbivores maintain broad diet breadth to avoid overloading individual detoxification pathways. We tested the hypothesis that a generalist marsupial herbivore, the common brushtail possum, Trichosurus vulpecula, can maintain a higher intake of food when allowed to select from two diets containing different profiles of secondary compounds (phenolics and terpenes) than when given access to the diets individually. Diets consisted of a fruit and vegetable mash to which was added ground leaves of either Eucalyptus melliodora or E. radiata. E. melliodora and E. radiata differ in their concentrations and types of secondary compounds. Brushtail possums include these eucalypt species as part of their natural diet. We measured food consumption and detoxification metabolites of possums on these diets. Consistent with the hypothesis, animals presented with a choice of both diets consumed more food than animals given diets singly. One of the two indicators of detoxification, acid load in urine, differed significantly between diets while the other, glucuronic acid, did not. These results provide partial support for the hypothesis that diet breadth is governed by detoxification abilities.     

Evaluation of the Evolution of Increased Competitive Ability (EICA) Hypothesis: Loss of Defense Against Generalist but not Specialist Herbivores

The evolution of increased competitive ability (EICA) hypothesis predicts that invasive plant species may escape their specialized natural enemies in their introduced range and subsequently evolve with a decrease in investment in anti-herbivore chemical defenses relative to native conspecifics. We compared the chemical profile of 10 populations of US native and 20 populations of European invasive Solidago gigantea. To test for differences in inducibility between native and invasive populations, we measured secondary chemistry in both damaged and undamaged plants. We also performed bioassays with three specialist and two generalist insect herbivores from four different feeding guilds. There was no evidence that invasive populations had reduced concentrations of sesquiterpenes, diterpenes, or short-chain hydrocarbons (SCH), although significant variation among populations was detected. Sesquiterpene and diterpene concentrations were not influenced by damage to the host plant, whereas SCH concentrations were decreased by damage for both native and invasive plants. Performance of the three specialist insects was not affected by the continental origin of the host plant. However, larval mass of the generalist caterpillar Spodoptera exigua was 37% lower on native plants compared to invasive plants. The other generalist insect, a xylem-tapping spittlebug that occurs on both continents, performed equally well on native and invasive plants. These results offer partial support for the defense predictions of the EICA hypothesis: the better growth of Spodoptera caterpillars on European plants shows that some defenses have been lost in the introduced range, even though our measures of secondary chemistry did not detect differences between continents. Our results show significant variation in chemical defenses and herbivore performance across populations on both continents and emphasize the need for analysis across a broad spatial scale and the use of multiple herbivores.