Herbivory on invasive exotic plants and their non-invasive relatives

The Enemy Release Hypothesis links exotic plant success to escape from enemies such as herbivores and pathogens. Recent work has shown that exotic plants that more fully escape herbivores and pathogens are more likely to become highly invasive, compared to plants with higher enemy loads in their novel ranges. We predicted that highly invasive plants from the Asteraceae and the Brassicaceae would be less acceptable, in laboratory no-choice feeding trials, to the generalist herbivore the American grasshopper, Schistocerca americana. We also compared herbivory on invasive and non-invasive plants from the genus Centaurea in no-choice feeding trials using the red-legged grasshopper Melanoplus femurrubrum and in a common garden in the field. In accordance with our predictions, highly invasive plants were fed on less by grasshoppers in the laboratory. They also received less damage in the field, suggesting that they contain feeding deterrents that render them less acceptable to generalist herbivores than non-invasive plants.     

Relationships among leaf damage, natural enemy release, and abundance in exotic and native prairie plants

The Enemy Release hypothesis holds that exotic plants may have an advantage over native plants because their specialized natural enemies are absent. We tested this hypothesis by measuring leaf damage and plant abundance for naturally-occurring plants in prairies, and by removing natural enemies in an enemy exclusion experiment. We classified plants as invasive exotic, noninvasive exotic, or native, to determine if their degree of invasiveness influenced their relationships with natural enemies. Our field surveys showed that invasive exotic plants generally had significantly lower levels of foliar damage than native species while there was no consistent pattern for noninvasive exotics compared to natives. The relationship between damage and abundance was different for exotic and native plants: foliar damage decreased with increasing abundance for exotic plants while the trend was positive for native plants. While these results from the field surveys supported the Enemy Release Hypothesis, the enemy exclusion experiment did not. There was no relationship between a species?? status as exotic or native and its degree of release from herbivory. Pastinaca sativa, the invasive exotic in this experiment, experienced gains in leaf area and vegetative biomass when treated with pesticides, indicating substantial herbivore pressure in the introduced range. These results show that foliar damage may not accurately predict the amount of herbivore pressure that plants actually experience, and that the Enemy Release hypothesis is not sufficient to explain the invasiveness of P. sativa in prairies.     

Soil fungal community changes in response to long-term fire cessation and N fertilization in tallgrass prairie

In grasslands, fire management and fertilization are established drivers of plant community change, but associated soil fungal responses are less well defined. We predicted that soil fungal communities would change seasonally, that decades of fire cessation and nitrogen (N) fertilization would alter fungal distributions, and that plant and fungal community change would be correlated. Surface soils were sampled monthly for 1 y from a 30-y fire by fertilization experiment to evaluate fungal community dynamics and assess correlation with plant community heterogeneity. ITS gene community composition was seasonally stable, excepting increased arbuscular mycorrhizal fungal summer abundance in the burned, fertilized treatment. Long-term treatments affected soil fungal and plant communities, with correlated heterogeneity patterns. Despite woody encroachment in the fire cessation treatment, soil fungal communities did not resemble those of forests. This study provides evidence supporting the strength of feedbacks between fungal and plant community change in response to long-term grassland fire and N management changes.     

Persistence of native and exotic plants 10 years after prairie reconstruction

Prairie reconstructions are a critical component of preservation of the imperiled tallgrass prairie ecosystem in the Midwestern United States. Sustainability of this endeavor depends on establishment of persistent cover of planted native species and resistance to noxious weeds. The goal of this study was to understand the influence of early reconstruction practices on long‐term outcomes. Twelve replicates of three planting methods (dormant‐season broadcast, growing‐season broadcast, and growing‐season drill) and three seed mix richness levels (10, 20, or 34 species), fully crossed in a completely randomized design were planted in 2005 on nine former agricultural fields located in Iowa and Minnesota. Cover by species was estimated in 2005–2007, 2010, and 2015. In 2015, cover of planted species, native nonplanted species, and exotic species were similar to those recorded in 2010. Cover of the noxious weed Cirsium arvense had also declined by an average of 49% without herbicide from a peak in 2007 to low stable levels from 2010 to 2015. Richness of planted forbs, on the other hand, were still increasing in high‐richness broadcast treatments (e.g. 17–59% increase 2010–1015 in Minnesota). Two results in 2015 are reasons for concern: cover of planted species is only slightly over 50% in both Minnesota and Iowa, though with forbs still increasing, this may improve; and the cool‐season exotic grasses Poa pratensis and Bromus inermis are increasing at both Minnesota and Iowa sites. Control of these invasive grasses will be necessary, but care will be needed to avoid negative impacts of control methods on natives.     

Attractiveness of Michigan native plants to arthropod natural enemies and herbivores

The use of plants to provide nectar and pollen resources to natural enemies through habitat management is a growing focus of conservation biological control. Current guidelines frequently recommend use of annual plants exotic to the management area, but native perennial plants are likely to provide similar resources and may have several advantages over exotics. We compared a set of 43 native Michigan perennial plants and 5 frequently recommended exotic annual plants for their attractiveness to natural enemies and herbivores for 2 yr. Plant species differed significantly in their attractiveness to natural enemies. In year 1, the exotic annual plants outperformed many of the newly established native perennial plants. In year 2, however, many native perennial plants attracted higher numbers of natural enemies than exotic plants. In year 2, we compared each flowering plant against the background vegetation (grass) for their attractiveness to natural enemies and herbivores. Screening individual plant species allowed rapid assessment of attractiveness to natural enemies. We identified 24 native perennial plants that attracted high numbers of natural enemies with promise for habitat management. Among the most attractive are Eupatorium perfoliatum L., Monarda punctata L., Silphium perfoliatum L., Potentilla fruticosa auct. non L., Coreopsis lanceolata L., Spiraea alba Duroi, Agastache nepetoides (L.) Kuntze, Anemone canadensis L., and Angelica atropurpurea L. Subsets of these plants can now be tested to develop a community of native plant species that attracts diverse natural enemy taxa and provides nectar and pollen throughout the growing season.     

The role of natural enemies in the expression and evolution of mixed mating in hermaphroditic plants and animals

Although a large portion of plant and animal species exhibit intermediate levels of outcrossing, the factors that maintain this wealth of variation are not well understood. Natural enemies are one relatively understudied ecological factor that may influence the evolutionary stability of mixed mating. In this paper, we aim for a conceptual unification of the role of enemies in mating system expression and evolution in both hermaphroditic animals and plants. We review current theory and detail the potential effects of enemies on fundamental mating system parameters. In doing so, we identify situations in which consideration of enemies alters expectations about the stability of mixed mating. Generally, we find that inclusion of the enemy dimension may broaden conditions in which mixed mating systems are evolutionarily stable. Finally, we highlight avenues ripe for future theoretical and empirical work that will advance our understanding of enemies in the expression and evolution of mixed mating in their hosts/victims, including examination of feedback cycles between victims and enemies and quantification of mating system-related parameters in victim populations in the presence and absence of enemies.     

Post-dispersal seed mortality of exotic and native species: Effects of fungal pathogens and seed predators

The invasive behaviour of exotic species is assumed to be due to the reduced impact of enemies on their performance, along with other possible mechanisms. I studied whether the seeds of exotics (6 species) are less impacted by seed predators and seed fungal pathogens than the seeds of their related natives (5 species). I also explored whether the co-occurrence of related natives and the time since introduction increased the percentage of lost seeds in exotics. Seeds were either left unprotected during a period of seven months or treated with fungicide, protected by seed predator exclosures or subjected to both treatments. Both treatments improved seed survival rate. Fungicide treatment had more positive effect on seeds of native than of exotic species but the fungicide-by-origin interaction was insignificant. When exotic species only were considered, fungicide had neutral effect on survival of their seeds, irrespective of the co-occurrence of related natives in the vegetation. Time since introduction was shown not to influence the proportion of seeds lost due to fungi or seed predators. Though the results of this study did not support enemy release as a possible mechanism causing the invasiveness of exotic species, it identified fungal pathogens as an enemy group with possibly differential impacts on native and exotic seeds, which thus deserves attention in future studies.     

Herbivory and population dynamics of invasive and native <Emphasis Type="Italic">Lespedeza</Emphasis>

Some exotic plants are able to invade habitats and attain higher fitness than native species, even when the native species are closely related. One explanation for successful plant invasion is that exotic invasive plant species receive less herbivory or other enemy damage than native species, and this allows them to achieve rapid population growth. Despite many studies comparing herbivory and fitness of native and invasive congeners, none have quantified population growth rates. Here, we examined the contribution of herbivory to the population dynamics of the invasive species, Lespedeza cuneata, and its native congener, L. virginica, using an herbivory reduction experiment. We found that invasive L. cuneata experienced less herbivory than L. virginica. Further, in ambient conditions, the population growth rate of L. cuneata (λ = 20.4) was dramatically larger than L. virginica (λ = 1.7). Reducing herbivory significantly increased fitness of only the largest L. virginica plants, and this resulted in a small but significant increase in its population growth rate. Elasticity analysis showed that the growth rate of these species is most sensitive to changes in the seed production of small plants, a vital rate that is relatively unaffected by herbivory. In all, these species show dramatic differences in their population growth rates, and only 2% of that difference can be explained by their differences in herbivory incidence. Our results demonstrate that to understand the importance of consumers in explaining the relative success of invasive and native species, studies must determine how consumer effects on fitness components translate into population-level consequences. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Variations in reproductive strategies between one invasive population and two native populations of Pseudorasbora parva

Reproductive traits of one invasive population in Lake Fuxian and two native populations in Lakes Chao and Dongting for Pseudorasbora parva were investigated to determine the variations in their reproductive strategies associated with the change of environmental conditions. Compared with the two native populations,P.parva markedly experienced the protraction in their spawning time and the decrease in mortality for adult individuals in Lake Fuxian. The three populations all got their first sexual maturity at the same age,but their total lengths at maturity were quite different for those in Lake Chao were significantly lower than in Lakes Fuxian and Dongting. Additionally,in spite that no significant difference occurred in their relative fecundities for P.parva in Lakes Fuxian and Dongting,the population in Lake Chao showed the markedly higher relative fecundity than the other two populations. These results were analyzed with the variations in water temperature,nutrient status,fishing stress among the three lakes. The relative decline in fecundity for the invasive population,which conflicted with the enemy release hypothesis,was disscussed with the context of the preponderance of another invasive fish,Neosalanx taihuensis,in Lake Fuxian.     

Parasites of native and exotic freshwater fishes in south‐western Australia

In this study, 1429 fishes of 18 different species (12 native and six exotic) were sampled from 29 localities to compare the levels of parasitism between native and exotic fish species and to examine the relationship between environmental degradation and parasite diversity. Forty‐four putative species of parasites were found and most of these appear to be native parasites, which have not previously been described. Two parasite species, Lernaea cyprinacea and Ligula intestinalis, are probably introduced. Both were found on or in a range of native fish species, where they may cause severe disease. Levels of parasitism and parasite diversity were significantly greater in native fishes than in exotic species, and this may contribute to an enhanced demographic performance and competitive ability in invading exotics. Levels of parasitism and parasite diversity in native fishes were negatively related to habitat disturbance, in particular to a suite of factors that indicate increased human use of the river and surrounding environment. This was due principally to the absence in more disturbed habitats of a number of species of endoparasites with complex life cycles, involving transmission between different host species.     

Vegetation responses to different spatial patterns of soil disturbance in burned and unburned tallgrass prairie

Pocket gopher (Geomyidae) disturbances are created in spatiallypredictable patterns. This may influence resource heterogeneity and affectgrassland vegetation in a unique manner. We attempt to determine the extent towhich density and spatial pattern of soil disturbances influence tallgrassprairie plant community structure and determine how these disturbances interactwith fire. To investigate the effects of explicit disturbance patterns we createdsimulated pocket gopher burrows and mounds in various spatial patterns.Simulated burrows were drilled into the soil at different densities inreplicated plots of burned and unburned prairie. Separate plots of simulatedmounds were created in burned and unburned prairie at low, medium, or high mounddensities in clumped, uniform, or random spatial dispersions. In both burned and unburned plots, increased burrow density decreasedgraminoid biomass and increased forb biomass. Total-plant and graminoid biomasswere higher in burned than unburned plots while forb biomass was higher inunburned plots. Total-plant species richness was not significantly affected byburrow density or burning treatments, but graminoid species richness increasedin unburned plots and forb species richness increased in burned plots. Plant species richness was temporarily reduced directly on mounddisturbances compared to undisturbed prairie. Over time and at larger samplingscales, the interaction of fire and mound disturbance patterns significantlyaffected total-plant and graminoid species richness. The principal effect inburned and unburned prairie was decreased total-plant and graminoid speciesrichness with increased mound disturbance intensity. Although species richness at small patch scales was not increased by anyintensity of disturbance and species composition was not altered by theestablishment of a unique guild of disturbance colonizing plants, our studyrevealed that interactions between soil disturbances and fire alter the plantcommunity dominance structure of North American tallgrass prairie primarily viachanges to graminoids. Moreover, these effects become increasingly pronouncedover time and at larger spatial sampling scales.     

Septate endophyte colonization and host responses of grasses and forbs native to a tallgrass prairie

Native tallgrass prairies support distinct dark septate endophyte (DSE) communities exemplified by Periconia macrospinosa and Microdochium sp. that were recently identified as common root symbionts in this system. Since these DSE fungi were repeatedly isolated from grasses and forbs, we aimed to test their abilities to colonize different hosts. One Microdochium and three Periconia strains were screened for colonization and growth responses using five native grasses and six forbs in an in vitro system. Previously published data for an additional grass (Andropogon gerardii) were included and reanalyzed. Presence of indicative inter- and intracellular structures (melanized hyphae, microsclerotia, and chlamydospores) demonstrated that all plant species were colonized by the DSE isolates albeit to varying degrees. Microscopic observations suggested that, compared to forbs, grasses were colonized to a greater degree in vitro. Host biomass responses varied among the host species. In broad comparisons, more grass species than forbs tended to respond positively to colonization, whereas more forb species tended to be non-responsive. Based on the suspected differences in the levels of colonization, we predicted that tallgrass prairie grasses would support greater DSE colonization than forbs in the field. A survey of field-collected roots from 15 native species supported this hypothesis. Our study supports the “broad host range” of DSE fungi, although the differences in the rates of colonization in the laboratory and in the field suggest a greater compatibility between grasses and DSE fungi. Furthermore, host responses to DSE range from mutualism to parasitism, suggesting a genotype-level interplay between the fungi and their hosts that determines the outcome of this symbiosis.     

Soil microbial responses to experimental warming and clipping in a tallgrass prairie

Global surface temperature is predicted to increase by 1.4–5.8°C by the end of this century. However, the impacts of this projected warming on soil C balance and the C budget of terrestrial ecosystems are not clear. One major source of uncertainty stems from warming effects on soil microbes, which exert a dominant influence on the net C balance of terrestrial ecosystems by controlling organic matter decomposition and plant nutrient availability. We, therefore, conducted an experiment in a tallgrass prairie ecosystem at the Great Plain Apiaries (near Norman, OK) to study soil microbial responses to temperature elevation of about 2°C through artificial heating in clipped and unclipped field plots. While warming did not induce significant changes in net N mineralization, soil microbial biomass and respiration rate, it tended to reduce extractable inorganic N during the second and third warming years, likely through increasing plant uptake. In addition, microbial substrate utilization patterns and the profiles of microbial phospholipid fatty acids (PLFAs) showed that warming caused a shift in the soil microbial community structure in unclipped subplots, leading to the relative dominance of fungi as evidenced by the increased ratio of fungal to bacterial PLFAs. However, no warming effect on soil microbial community structure was found in clipped subplots where a similar scale of temperature increase occurred. Clipping also significantly reduced soil microbial biomass and respiration rate in both warmed and unwarmed plots. These results indicated that warming‐led enhancement of plant growth rather than the temperature increase itself may primarily regulate soil microbial response. Our observations show that warming may increase the relative contribution of fungi to the soil microbial community, suggesting that shifts in the microbial community structure may constitute a major mechanism underlying warming acclimatization of soil respiration.     

Predicting invasiveness in exotic species: do subtropical native and invasive exotic aquatic plants differ in their growth responses to macronutrients?

We investigated whether plasticity in growth responses to nutrients could predict invasive potential in aquatic plants by measuring the effects of nutrients on growth of eight non-invasive native and six invasive exotic aquatic plant species. Nutrients were applied at two levels, approximating those found in urbanized and relatively undisturbed catchments, respectively. To identify systematic differences between invasive and non-invasive species, we compared the growth responses (total biomass, root:shoot allocation, and photosynthetic surface area) of native species with those of related invasive species after 13 weeks growth. The results were used to seek evidence of invasive potential among four recently naturalized species. There was evidence that invasive species tend to accumulate more biomass than native species ( P  = 0.0788). Root:shoot allocation did not differ between native and invasive plant species, nor was allocation affected by nutrient addition. However, the photosynthetic surface area of invasive species tended to increase with nutrients, whereas it did not among native species ( P  = 0.0658). Of the four recently naturalized species, Hydrocleys nymphoides showed the same nutrient-related plasticity in photosynthetic area displayed by known invasive species. Cyperus papyrus showed a strong reduction in photosynthetic area with increased nutrients. H. nymphoides and C. papyrus also accumulated more biomass than their native relatives. H. nymphoides possesses both of the traits we found to be associated with invasiveness, and should thus be regarded as likely to be invasive.     

Laboratory evaluation of predation rates of two native natural enemies on the exotic pest Bagrada hilaris

Laboratory studies were conducted to determine the consumption rates of two native predators found attacking the exotic invasive stink bug Bagrada hilaris (Burmeister) (Hempitera: Pentatomidae) in field plots in New Mexico, USA. Individual field‐collected adults of the spined soldier bug, Podisus maculiventris (Say) (Hempitera: Pentatomidae) and the soft‐winged flower beetle, Collops vittatus (Say) (Coleoptera: Melyridae), were provided daily with fixed numbers of different life stages of B. hilaris under controlled conditions. Consumption rates were recorded daily for ten consecutive days for a total of 20 adult P. maculiventris and 20 adult C. vittatus per prey life stage. For P. maculiventris, predation rates were obtained in relation to adult, third and fifth instar prey, and for C. vittatus for first, second and third instar prey. On average, predation on third and fifth instar B. hilaris nymphs by P. maculiventris was 0.6 ± 0.1 and 0.9 ± 0.1 per day respectively. Predation rates on adults were slightly higher (1.3 ± 0.1 per day), with female prey being consumed at a significantly higher rate than male prey when three mating pairs of B. hilaris were provided per day (0.8 ± 0.1 females per day vs. 0.5 ± 0.1 males per day). Collops vittatus adults provisioned daily with 20 first instar B. hilaris nymphs killed a mean total of 4.7 ± 0.4 and 9.3 ± 0.6 prey each day (for male and female beetles respectively), with only approximately half that number of prey being fully consumed. Partial consumption of prey by this species was also observed with second and third instar nymphs, but to a lesser degree. Female beetles consumed significantly more prey than did male beetles when fed first and third instar B. hilaris, but not when given second instar prey.     

Spatio-temporal variation in seed predation by a native weevil in the invasive Prunus serotina

Invasive species may escape the enemies from their native range (‘enemy release’), but they can also acquire new enemies in their introduced range, which will affect the invasion process. For the invasive tree species Prunus serotina, seed predation by the native weevil Furcipus rectirostris has been reported in forests in its introduced range. In this study, we quantified how common the infestation of P. serotina seeds by F. rectirostris is in a 4000 km² area in northern Belgium. Seeds were sampled on P. serotina trees in different habitats and in two years, i.e., with low and high P. serotina fruit production. Infestation was found throughout the study region, in 43 and 62% of the sampled trees in the two years of the study; the maximum infestation levels of infested seed samples were 50 and 69%. Overall, predation occurred in 4.4 and 10.8% of the sampled seeds. The level of infestation differed between habitats and years, and the number of fruits per raceme was inversely related to the infestation level. Notwithstanding the rather high incidence of F. rectirostris infestation in our study, the impact on P. serotina's invasiveness might remain low seeing the overall high seed production and dispersal capacity of the species.     

Native weeds and exotic plants: relationships to disturbance in mixed-grass prairie

Disturbance frequently is implicated in the spread of invasive exotic plants. Disturbances may be broadly categorized as endogenous (e.g., digging by fossorial animals) or exogenous (e.g., construction and maintenance of roads and trails), just as weedy species may be native or exotic in origin. The objective of this study was to characterize and compare exotic and native weedy plant occurrence in and near three classes of disturbance – digging by prairie dogs (an endogenous disturbance to which native plants have had the opportunity to adapt), paved or gravel roads (an exogenous disturbance without natural precedent), and constructed trails (an exogenous disturbance with a natural precedent in trails created by movement of large mammals) – in three geographically separate national park units. I used plant survey data from the North and South Units of Theodore Roosevelt National Park and Wind Cave National Park in the northern mixed-grass prairie of western North and South Dakota, USA, to characterize the distribution of weedy native and exotic plants with respect to the three disturbance classes as well as areas adjacent to them. There were differences both in the susceptibility of the disturbance classes to invasion and in the distributions of native weeds and exotic species among the disturbance classes. Both exotic and native weedy species richness were greatest in prairie dog towns and community composition there differed most from undisturbed areas. Exotic species were more likely to thrive near roadways, where native weedy species were infrequently encountered. Exotic species were more likely to have spread beyond the disturbed areas into native prairie than were weedy native species. The response of individual exotic plant species to the three types of disturbance was less consistent than that of native weedy species across the three park units. This revised version was published online in June 2006 with corrections to the Cover Date.     

Responses of morphological and physiological traits to herbivory by snails of three invasive and native submerged plants

3种入侵和本地沉水植物形态和生理性状对螺类牧食的响应 沉水植物水盾草(Cabomba caroliniana)已成为中国太湖流域的优势入侵水生植物。与外来物种的原产地环境相比,引入地新环境中存在的专食性天敌较少。外来物种可能会逃避其原产地环境中的天敌牧食,又因为它们的适口性相对较差,从而导致在引入地外来物种通常比本地物种遭受的牧食者影响更低(天敌逃逸假说)。本研究的目的是比较水盾草与共生的本地沉水植物对本地牧食者的响应。我们进行了一个中宇宙尺度实验,研究了水盾草和两种共生的本地沉水植物黑藻(Hydrilla verticillata)和穗花狐尾藻(Myriophyllum spicatum)对两种本地广食性腹足纲螺类萝卜螺(Radix swinhoei)和环棱螺(Sinotaia quadrata)的牧食响应。记录了它们的形态性状指标(总生物量、冠根比和相对生长率)和生理性状指标(叶片总非结构性碳、木质素和纤维素)。研究结果表明,环棱螺对3种沉水植物性状指标的影响较少。随着本地广食性螺类萝卜螺数量的增加,黑藻和穗花狐尾藻大部分植物性状发生了改变,而水盾草的植物性状表现出相对稳定的趋势。水盾草对萝卜螺的牧食更具抵抗力,这与天敌逃逸假说的假设一致。这一发现说明牧食性螺类促进了水盾草的入侵,这可能会改变沉水植物群落中的物种组成。     

Reciprocal interactions between native and introduced populations of common milkweed,Asclepias syriaca,and the specialist aphid,Aphis nerii

Following its introduction into Europe (EU), the common milkweed (Asclepias syriaca) has been free of most specialist herbivores that are present in its native North American (NA) range, except for the oleander aphid Aphis nerii. We compared EU and NA populations of A. nerii on EU and NA milkweed populations to test the hypothesis that plant–insect interactions differ on the two continents. First, we tested if herbivore performance is higher on EU plants than on NA plants, because the former have escaped most of their herbivores and have perhaps been selected for lower defence levels following introduction. Second, we compared two A. nerii lines (one from each continent) to test whether genotypic differences in the herbivore may influence species interactions in plant–herbivore communities in the context of species introductions. The NA population of A. nerii developed faster, had higher fecundity and attained higher population growth rates than the EU population. There was no overall significant continental difference in aphid resistance between the plants. However, milkweed plants from EU supported higher population growth rates and faster development of the NA line of A. nerii than plants from NA. In contrast, EU aphids showed similar (low) performance across plant populations from both continents. In a second experiment, we examined how chewing herbivores indirectly mediate interactions between milkweeds and aphids, and induced A. syriaca plants from each continent by monarch caterpillars (Danaus plexippus) to compare the resulting changes in plant quality on EU aphid performance. As specialist chewing herbivores of A. syriaca are only present in NA, we expected that plants from the two continents may affect aphid growth in different ways when they are challenged by a specialist chewing herbivore. Caterpillar induction decreased aphid developmental times on NA plants, but not on EU plants, whereas fecundity and population growth rates were unaffected by induction on both plant populations. The results show that genetic variation in the plants as well as in the herbivores can determine the outcome of plant–herbivore interactions.