Feeding preferences of Melanoplus femurrubrum grasshoppers on native and exotic grasses: behavioral and molecular approaches

Generalist insect herbivores, such as grasshoppers, may either avoid feeding on exotic plants, potentially enabling these plants to become invasive in the introduced range, or insects may incorporate exotic plants into their diet, contributing to the biotic resistance of native communities and potentially preventing plant invasions. Accurate determination of insect diet preferences with regard to native and exotic plants can be challenging, but this information is critical for understanding the interaction between native herbivores and exotic plants, and ultimately the mechanisms underlying plant invasions. To address this, we combined behavioral and molecular approaches to accurately compare food consumption of the polyphagous red‐legged grasshopper, Melanoplus femurrubrum (De Geer) (Orthoptera: Acrididae), on native [Andropogon gerardii Vitman and Bouteloua curtipendula (Michx.) Torr.] and exotic, potentially invasive grasses [Miscanthus sinensis Andersson and Bothriochloa ischaemum (L.) Keng] (all Poaceae). We found that M. femurrubrum grasshoppers demonstrated strong feeding preferences toward exotic grasses in experiments with intact plants under both field and greenhouse conditions, but they showed no preference in experiments with clipped leaves. Additionally, we sampled the gut contents of M. femurrubrum collected in the field and identified the ingested plant species based on DNA sequences for the non‐coding region of the chloroplast trnL (UAA) gene. We found that exotic plants were prevalent in the gut contents of grasshoppers collected at study sites in Ohio and Maryland, USA. These results suggest that the generalist herbivore M. femurrubrum does not avoid feeding on exotic grasses with which they do not share coevolutionary history. In addition, by demonstrating greater food consumption of exotic plants, these grasshoppers potentially provide biotic resistance should these grasses escape cultivation and become invasive in the introduced range.     

Increased resistance to generalist herbivores in invasive populations of the California poppy (Eschscholzia californica)

Escape from enemies in the native range is often assumed to contribute to the successful invasion of exotic species. Following optimal defence theory, which assumes a trade‐off between herbivore resistance and plant growth, some have predicted that the success of invasive species could be the result of the evolution of lower resistance to herbivores and increased allocation of resources to growth and reproduction. Lack of evidence for ubiquitous costs of producing plant toxins, and the recognition that invasive species may escape specialist, but not generalist enemies, has led to a new prediction: invasive species may escape ecological trade‐offs associated with specialist herbivores, and evolve increased, rather than decreased, production of defensive compounds that are effective at deterring generalist herbivores in the introduced range. We tested the performance of two generalist lepidopteran herbivores, Trichoplusia ni and Orgyia vetusta, when raised on diets of native and invasive populations of the California poppy, Eschscholzia californica. Pupae of T. ni were significantly larger when reared on native populations. Similarly, caterpillars of O. vetusta performed significantly better when raised on native populations, indicating that invasive populations of the California poppy are more resistant to herbivores than native populations. The chance of successful establishment of some non‐indigenous plant species may be increased by retaining resistance to generalist herbivores, and in some cases, invasive species may be able to escape ecological trade‐offs in their new range and evolve, as we observed, even greater resistance to generalist herbivores than native plants.     

Grasshopper Herbivory Affects Native Plant Diversity and Abundance in a Grassland Dominated by the Exotic Grass Agropyron cristatum

The indirect effects of native generalist insect herbivores on interactions between exotic and native grassland plants have received limited attention. Crested wheatgrass ( Agropyron cristatum ) is the most common exotic rangeland grass in western North America. Crested wheatgrass communities are resistant to colonization by native plant species and have strong competitive effects on native species, imposing problems for the restoration of native grasslands. Grasshoppers are generalist herbivores that are often abundant in Crested wheatgrass–dominated sites in the northern Great Plains. We conducted two experiments in a Crested wheatgrass–dominated grassland in western North Dakota to test the hypothesis that grasshopper herbivory influences local Crested wheatgrass community composition by impeding native seedlings. Grasshopper herbivory negatively affected the species richness, abundance, and Shannon diversity of native plants in 3 of 4 years. Although additional research is needed to determine if grasshoppers actively select native plants, the effects of grasshopper herbivory may be an important consideration in the restoration of Crested wheatgrass areas. Our findings illustrate the importance of understanding the impact of native generalist invertebrate herbivores on the relationships between exotic and native plants.     

Being a generalist herbivore in a diverse world: how do diets from different grasslands influence food plant selection and fitness of the grasshopper Chorthippus parallelus?

1. Generalist insect herbivores occupy a variety of habitats that differ in food plant composition. Dietary mixing has been proposed as a possibility for generalists to overcome nutritional deficiencies of single plant species, but only a few studies have investigated herbivore feeding and fitness for diets that resemble natural scenarios. We studied feeding behaviour, survival, and reproduction of the generalist grasshopper Chorthippus parallelus raised on food plants of four typical habitats. 2. Grasshopper diet consisted of grasses (92.5%), legumes (6.7%) and, in small quantities, other forbs (0.8%). Diet selection differed between the four food plant mixtures, and depended on grasshopper sex and developmental stage. There was no correlation between the relative abundance of plant species in the field and the fraction of these species in the grasshopper diet. 3. Grasshoppers survived on average for 40.4 ± 1.0 days before maturity, grew 106.8 mg until maturity moult, and females laid 4.1 ± 0.4 egg pods, each of which contained 8.5 ± 0.4 eggs. However, despite the differences in feeding behaviour, grasshopper fitness was the same in all of the four food plant mixtures. While the digestibility of ingested food was similar in the four different treatments, indices indicated differences in the conversion efficiency to body mass. 4. Our results show that C. parallelus is a plastic feeder with no fixed preferences in diet composition. The results emphasise that generalist herbivores can counteract putative quality deficiencies of single food plants by selective dietary mixing.     

The enemy release and EICA hypothesis revisited: incorporating the fundamental difference between specialist and generalist herbivores

The success of invasive plants has been attributed to their escape from natural enemies and subsequent evolutionary change in allocation from defence to growth and reproduction. In common garden experiments with Senecio jacobaea, a noxious invasive weed almost worldwide, the invasive populations from North America, Australia, and New Zealand did indeed allocate more resources to vegetative and reproductive biomass. However, invasive plants did not show a complete change in allocation from defence to growth and reproduction. Protection against generalist herbivores increased in invasive populations and pyrrolizidine alkaloids, their main anti‐herbivore compounds, did not decline in invasive populations but were higher overall compared with native populations. In contrast, invasive plants lost additional protection against specialist herbivores adapted to pyrrolizidine alkaloids. Hence, the absence of specialist herbivores in invasive populations resulted in the evolution of lower protection against specialists and increased growth and reproduction, but also allowed a shift towards higher protection against generalist herbivores.     

Biotic resistance to plant invasions? Native herbivores prefer non‐native plants

In contrast to expectations of the enemy release hypothesis, but consistent with the notion of biotic resistance, we found that native generalist crayfishes preferred exotic over native freshwater plants by a 3 : 1 ratio when plants were paired by taxonomic relatedness. Native crayfishes also preferred exotic over native plants when tested across 57 native and 15 exotic plants found growing sympatrically at 11 sites throughout the southeastern USA. Exotic grass carp that share little evolutionary history with most of these plants exhibited no preference for native vs. exotic species. Analyses of three terrestrial data sets showed similar patterns, with native herbivores generally preferring exotic plants, while exotic herbivores rarely exhibited a preference. Thus, exotic plants may escape their coevolved herbivores only to be preferentially consumed by the native generalist herbivores in their new ranges, suggesting that native herbivores may provide biotic resistance to plant invasions.     

Invasive plant population and herbivore identity affect latex induction

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Exotic Lonicera species both escape and resist specialist and generalist herbivores in the introduced range in North America

The enemy release hypothesis predicts that invasive plant species may benefit from a lack of top-down control by co-evolved herbivores, particularly specialists, in their new range. However, to benefit from enemy escape, invasive plants must also escape or resist specialist or generalist herbivores that attack related species in the introduced range. We compared insect herbivore damage on the exotic shrub, Lonicera maackii, the native congener Lonicera reticulata, and the native confamilial Viburnum prunifolium in North America. We also compared the laboratory preference and performance of a North American honeysuckle specialist sawfly (Zaraea inflata) and the performance of a widespread generalist caterpillar (Spodoptera frugiperda) on cut foliage from native and exotic Lonicera species. L. maackii received significantly lower amounts of foliar herbivory than L. reticulata across three seasons, while damage levels observed on V. prunifolium for two seasons was generally intermediate between L. reticulata and L. maackii. The specialist sawfly damaged L. reticulata heavily, but was not detected on L. maackii in the field. There were few statistical differences in the performance of sawfly larvae on L. reticulata and L. maackii, but the sawfly achieved higher pupal masses on L. reticulata than on L. maackii, and they strongly preferred L. reticulata over L. maackii when given a choice. The sawfly was unable to complete development on native L. sempervirens and non-native L. japonica. In contrast, the generalist caterpillar performed similarly on all Lonicera species. While L. maackii experienced little herbivory in the field compared to native relatives in the same habitat, laboratory assays indicate L. maackii appears to be a suitable host that escapes selection by the specialist, but L. japonica and L. sempervirens are highly resistant to it. These findings indicate that both enemy escape and resistance (to a specialist, but not a generalist herbivore) may contribute to the success of exotic Lonicera species.     

Effects of generalist herbivory on resistance and resource allocation by the invasive plant,Phytolacca americana

Successful invasions by exotic plants are often attributed to a loss of co‐evolved specialists and a re‐allocation of resources from defense to growth and reproduction. However, invasive plants are rarely completely released from insect herbivory because they are frequently attacked by generalists in their introduced ranges. The novel generalist community may also affect the invasive plant's defensive strategies and resource allocation. Here, we tested this hypothesis using American pokeweed (Phytolacca americana L.), a species that has become invasive in China, which is native to North America. We examined resistance, tolerance, growth and reproduction of plant populations from both China and the USA when plants were exposed to natural generalist herbivores in China. We found that leaf damage was greater for invasive populations than for native populations, indicating that plants from invasive ranges had lower resistance to herbivory than those from native ranges. A regression of the percentage of leaf damage against mass showed that there was no significant difference in tolerance between invasive and native populations, even though the shoot, root, fruit and total mass were larger for invasive populations than for native populations. These results suggest that generalist herbivores are important drivers mediating the defensive strategies and resource allocation of the invasive American pokeweed.     

Specificity of herbivore‐induced responses in an invasive species,Alternanthera philoxeroides (alligator weed)

Herbivory‐induced responses in plants can both negatively affect subsequently colonizing herbivores and mitigate the effect of herbivory on the host. However, it is still less known whether plants exhibit specific responses to specialist and generalist herbivores in non‐secondary metabolite traits and how specificity to specialists and generalists differs between invasive and native plant populations. We exposed an invasive plant, Alternanthera philoxeroides, to Agasicles hygrophila (Coleoptera, Chrysomelidae; specialist), Spodoptera litura (Lepidoptera, Noctuidae; generalist), manual clipping, or application of exogenous jasmonic acid and examined both the specificity of elicitation in traits of fitness (e.g., aboveground biomass), morphology (e.g., root:shoot ratio), and chemistry (e.g., C/N ratio and lignin), and specificity of effect on the subsequent performance of A. hygrophila and S. litura. Then, we assessed variation of the specificity between invasive and native populations (USA and Argentina, respectively). The results showed S. litura induced higher branching intensity and specific leaf area but lower C/N ratio than A. hygrophila, whereas A. hygrophila induced higher trichome density than S. litura. The negative effect of induction on subsequent larval growth was greater for S. litura than for A. hygrophila. Invasive populations had a weaker response to S. litura than to A. hygrophila in triterpenoid saponins and C/N ratio, while native populations responded similarly to these two herbivores. The specific effect on the two herbivores feeding on induced plants did not vary between invasive and native populations. Overall, we demonstrate specificity of elicitation to specialist and generalist herbivores in non‐secondary metabolite traits, and that the generalist is more susceptible to induction than the specialist. Furthermore, chemical responses specific to specialist and generalist herbivores only exist in the invasive populations, consistent with an evolutionary change in specificity in the invasive populations.     

Evolutionary increases in defense during a biological invasion

Invasive plants generally escape from specialist herbivores of their native ranges but may experience serious damage from generalists. As a result, invasive plants may evolve increased resistance to generalists and tolerance to damage. To test these hypotheses, we carried out a common garden experiment comparing 15 invasive populations with 13 native populations of Chromolaena odorata, including putative source populations identified with molecular methods and binary choice feeding experiments using three generalist herbivores. Plants from invasive populations of C. odorata had both higher resistance to three generalists and higher tolerance to simulated herbivory (shoot removal) than plants from native populations. The higher resistance of plants from invasive populations was associated with higher leaf C content and densities of leaf trichomes and glandular scales, and lower leaf N and water contents. Growth costs were detected for tolerance but not for resistance, and plants from invasive populations of C. odorata showed lower growth costs of tolerance. Our results suggest that invasive plants may evolve to increase both resistance to generalists and tolerance to damage in introduced ranges, especially when the defense traits have low or no fitness costs. Greater defenses in invasive populations may facilitate invasion by C. odorata by reducing generalist impacts and increasing compensatory growth after damage has occurred.     

Herbivore phenology can predict response to changes in plant quality by livestock grazing

Livestock grazing can have a strong impact on herbivore abundance, distribution and community. However, not all species of herbivores respond the same way to livestock grazing, and we still have a poor understanding of the underlying mechanisms driving these differential responses. Here, we investigate the effect of light intensity cattle grazing on the abundance of two grasshoppers (Euchorthippus cheui and E. unicolor) that co-occur in the same grasslands and feed on the same food plant (the dominant grass Leymus chinensis). The two grasshopper species differ in phenology so that their peak abundances are separated into early- and late-growing seasons. We used an exclosure experiment to monitor grasshopper abundance and food quality in the field under grazed and ungrazed conditions, and performed feeding trials to examine grasshopper preference for grazed or ungrazed food plants in the laboratory. We found that the nitrogen content of L. chinensis leaves continuously declined in the ungrazed areas, but was significantly enhanced by cattle grazing over the growing season. Cattle grazing facilitated the early-season grasshopper E. cheui, whereas it suppressed the late-season grasshopper E. unicolor. Moreover, feeding trials showed that E. cheui preferred L. chinensis from grazed plots, while E. unicolor preferred the leaves from ungrazed plots. We conclude that livestock grazing has opposite effects on the two grasshopper species, and that these effects may be driven by grazing-induced changes in plant nutrient content and the unique nutritional niches of the grasshoppers. These results suggest that insects that belong to the same guild can have opposite nutrient requirements, related to their distinct phenologies, and that this can ultimately affect their response to cattle grazing. Our results show that phenology may link insect physiological needs to local resource availabilities, and should be given more attention in future work on interactions between large herbivores and insects.     

Impact of grasshoppers and an invasive grass on establishment and initial growth of restoration plant species

Exotic plant invasion can have dramatic impacts on native plants making restoration of native vegetation at invaded sites challenging. Though invasives may be superior competitors, it is possible their dominance could be enhanced by insect herbivores if native plants are preferred food sources. Insect herbivory can regulate plant populations, but little is known of its effects in restoration settings. There is a need to better understand relationships between insect herbivores and invasive plants with regard to their combined potential for impacting native plant establishment and restoration success. The objective of this study was to assess impacts of grasshopper herbivory and the invasive grass Bromus tectorum (cheatgrass) on mortality and growth of 17 native plant species used in restoration of critical sagebrush steppe ecosystems. Field and greenhouse experiments were conducted using moderate densities of a common, generalist pest grasshopper (Melanoplus bivittatus). Grasshoppers had stronger and more consistent impacts on native restoration plants in field and greenhouse studies than cheatgrass. After 6 weeks in the greenhouse, grasshoppers were associated with 36% mortality over all native restoration species compared to 2% when grasshoppers were absent. Herbivory was also associated with an approximately 50% decrease in native plant biomass. However, effects varied among species. Artemisia tridentata, Chrysothamnus viscidiflorus, and Coreopsis tinctoria were among the most negatively impacted, while Oenothera pallida, Pascopyrum smithii, and Leymus cinerus were unaffected. These findings suggest restoration species could be selected to more effectively establish and persist within cheatgrass infestations, particularly when grasshopper populations are forecasted to be high.     

Phylogenetic isolation increases plant success despite increasing susceptibility to generalist herbivores

Aim Theory suggests that introduced species that are phylogenetically distant from their recipient communities should be more successful than closely related introduced species because they can exploit open niches and escape enemies in their new range, i.e. Darwin’s Naturalization Hypothesis. Alternatively, it has also been hypothesized that closely related invaders might be more successful than novel invaders because they are pre‐adapted to conditions in their new range; a paradox coined Darwin’s Naturalization Conundrum. To date, these hypotheses have been tested primarily at the regional scale, not within local plant communities where introduced species colonize, compete and encounter herbivores. Location Global. Methods and Results We used community phylogenetics to analyse data from 49 published experiments to examine the importance of phylogenetic relatedness and generalist herbivory on native and exotic plant success at the community level. Plants that were categorized as ‘invasive’ were indeed less related to the recipient community than ‘non‐pest’ exotic plants. Distantly related exotic plants were also more abundant than closely related species. Phylogenetic relatedness predicted herbivore impact, but in a way that was opposite to predictions, as herbivores had stronger, not lesser, impacts on distantly related plants. Importantly, these same patterns generally held for native plants, as distantly related native plants were more abundant and more susceptible to herbivores than closely related species, ultimately resulting in herbivores suppressing community‐level phylogenetic diversity. Main conclusions Distantly related plants were more locally successful despite experiencing stronger control by generalist herbivores, a finding that was robust across native and exotic species. To our knowledge, this is the first evidence that phylogenetic matching influences the local success of both native and exotic species and that herbivores can influence community phylodiversity. Phylogenetic relatedness explained a relatively small portion of the variance in the data even after taking herbivory into account, however, suggesting that phylogenetic matching works in combination with other factors to influence community assembly.     

Plant chemical defense against herbivores and pathogens: generalized defense or trade-offs?

Plants are often attacked by multiple enemies, including pathogens and herbivores. While many plant secondary metabolites show specific effects toward either pathogens or herbivores, some can affect the performance of both these groups of natural enemies and are considered to be generalized defense compounds. We tested whether aucubin and catalpol, two iridoid glycosides present in ribwort plantain (Plantago lanceolata), confer in vivo resistance to both the generalist insect herbivore Spodoptera exigua and the biotrophic fungal pathogen Diaporthe adunca using plants from P. lanceolata lines that had been selected for high- and low-leaf iridoid glycoside concentrations for four generations. The lines differed approximately three-fold in the levels of these compounds. Plants from the high-selection line showed enhanced resistance to both S. exigua and D. adunca, as evidenced by a smaller lesion size and a lower fungal growth rate and spore production, and a lower larval growth rate and herbivory under both choice and no-choice conditions. Gravimetric analysis revealed that the iridoid glycosides acted as feeding deterrents to S. exigua, thereby reducing its food intake rate, rather than having post-ingestive toxic effects as predicted from in vitro effects of hydrolysis products. We suggest that the bitter taste of iridoid glycosides deters feeding by S. exigua, whereas the hydrolysis products formed after tissue damage following fungal infection mediate pathogen resistance. We conclude that iridoid glycosides in P. lanceolata can serve as broad-spectrum defenses and that selection for pathogen resistance could potentially result in increased resistance to generalist insect herbivores and vice versa, resulting in diffuse rather than pairwise coevolution.     

Tolerance and resistance of invasive and native Eupatorium species to generalist herbivore insects

Invasive plants are exotic species that escape control by native specialist enemies. However, exotic plants may still be attacked by locally occurring generalist enemies, which can influence the dynamics of biological invasions. If invasive plants have greater defensive (resistance and tolerance) capabilities than indigenous plants, they may experience less damage from native herbivores. In the present study, we tested this prediction using the invasive plant Eupatorium adenophorum and two native congeners under simulated defoliation and generalist herbivore insect (Helicoverpa armigera and Spodoptera litura) treatments. E. adenophorum was less susceptible and compensated more quickly to damages in biomass production from both treatments compared to its two congeners, exhibiting greater herbivore tolerance. This strong tolerance to damage was associated with greater resource allocation to aboveground structures, leading to a higher leaf area ratio and a lower root: crown mass ratio than those of its native congeners. E. adenophorum also displayed a higher resistance index (which integrates acid detergent fiber, nitrogen content, carbon/nitrogen ratio, leaf mass per area, toughness, and trichome density) than its two congeners. Thus, H. armigera and S. litura performed poorly on E. adenophorum, with less leaf damage, a lengthened insect developmental duration, and decreased pupating: molting ratios compared to those of the native congeners. Strong tolerance and resistance traits may facilitate the successful invasion of E. adenophorum in China and may decrease the efficacy of leaf-feeding biocontrol agents. Our results highlight both the need for further research on defensive traits and their role in the invasiveness and biological control of exotic plants, and suggest that biocontrol of E. adenophorum in China would require damage to the plant far in excess of current levels.     

Invasion of weeping lovegrass reduces native food and habitat resource of Eusphingonotus japonicus (Saussure)

To examine the relative importance and interactions of trophic and non-trophic effects of plant invasions on herbivores, we investigated how the invasion of weeping lovegrass Eragrostis curvula impacted Eusphingonotus japonicus, a food generalist grasshopper endemic to gravelly floodplains with sparse vegetation in Japan. Field observations and laboratory experiments showed that the grasshopper fed mainly on herbs endemic to the gravelly floodplains, which were negatively impacted by the alien grass. The alien grass was not consumed as food. Field censuses also showed that the abundance of the grasshopper was positively correlated with the coverage of endemic herbs in a plot where weeping lovegrass was not dominant. Dominance of the grass (i.e., habitat modification for E. japonicus) negatively impacted the grasshopper and weakened the positive relationship between the grasshopper and endemic herbs. These results suggest that an understanding of the relationship between non-trophic and trophic effects is essential to predict the impacts of plant invasions on herbivores.     

Larval feeding induced defensive responses in tobacco: comparison of two sibling species of <Emphasis Type="Italic">Helicoverpa</Emphasis> with different diet breadths

Plants respond differently to damage by different herbivorous insects. We speculated that sibling herbivorous species with different host ranges might also influence plant responses differently. Such differences may be associated with the diet breadth (specialization) of herbivores within a feeding guild, and the specialist may cause less intensive plant responses than the generalist. The tobacco Nicotinana tabacum L. is the common host plant of a generalist Helicoverpa armigera (Hübner) and a specialist H. assulta Guenée (Lepidoptera, Noctuidae). The induced responses of tobacco to feeding of these two noctuid herbivores and mechanical wounding were compared. The results showed that the feeding of the specialist H. assulta and the generalist H. armigera resulted in the same inducible defensive system, but response intensity of plants was different to these two species. Inductions of jasmonic acid (JA), lipoxygenase (LOX), and proteinase inhibitors (PIs) were not significantly different concerning these two species, but H. assulta caused the less intensive foliar polyphenol oxidase (PPO) increase, more intensive nicotine and peroxidase (POD) increases in tobacco than H. armigera. The defensive response of plant to herbivores with different diet breadth seems to be more complicated than we expected, and the specialist does not necessarily cause less intensive plant responses than the generalist.     

Invasive plants and their escape from root herbivory: a worldwide comparison of the root-feeding nematode communities of the dune grass <Emphasis Type="Italic">Ammophila arenaria</Emphasis> in natural and introduced ranges

Invasive plants generally have fewer aboveground pathogens and viruses in their introduced range than in their natural range, and they also have fewer pathogens than do similar plant species native to the introduced range. However, although plant abundance is strongly controlled by root herbivores and soil pathogens, there is very little knowledge on how invasive plants escape from belowground enemies. We therefore investigated if the general pattern for aboveground pathogens also applies to root-feeding nematodes and used the natural foredune grass Ammophila arenariaas a model. In the late 1800s, the European A. arenariawas introduced into southeast Australia (Tasmania), New Zealand, South Africa, and the west coast of the USA to be used for sand stabilization. In most of these regions, it has become a threat to native vegetation, because its excessive capacity to stabilize wind-blown sand has changed the geomorphology of coastal dunes. In stable dunes of most introduced regions, A. arenaria is more abundant and persists longer than in stabilized dunes of the natural range. We collected soil and root samples and used additional literature data to quantify the taxon richness of root-feeding nematodes on A.␣arenaria in its natural range and collected samples from the four major regions where it has been introduced. In most introduced regions A. arenaria did not have fewer root-feeding nematode taxa than the average number in its natural range, and native plant species did not have more nematode taxa than the introduced species. However, in the introduced range native plants had more feeding-specialist nematode taxa than A. arenaria and major feeding specialists (the sedentary endoparasitic cyst and root knot nematodes) were not found on A. arenaria in the southern hemisphere. We conclude that invasiveness of A. arenaria correlates with escape from feeding specialist nematodes, so that the pattern of escape from root-feeding nematodes is more alike escape from aboveground insect herbivores than escape from aboveground pathogens and viruses. In the natural range of A. arenaria, the number of specialist-feeding nematode taxa declines towards the margins. Growth experiments are needed to determine the relationship between nematode taxon diversity, abundance, and invasiveness of A. arenaria.     

Do seeds from invasive bromes experience less granivory than seeds from native congeners in the Great Basin Desert?

In part, the enemy release hypothesis of plant invasion posits that generalist herbivores in the non-native ranges of invasive plants will prefer native plants to exotic invaders. However, the extent to which this occurs in natural communities is unclear. Here, I examined the foraging preferences of an important guild of generalist herbivores—granivorous rodents—with respect to seeds from a suite of native and invasive Bromus (“brome”) species at five study sites distributed across?≈?80,000 km² of the Great Basin Desert, USA. By examining only congeners, I accounted for a potentially large source of interspecific variation (phylogenetic relatedness). In general, granivorous rodents removed seeds from native bromes at a 23% higher rate than seeds from invasive bromes, suggesting a preference for native species. This preference was not entirely explained by seed size, and patterns of seed removal were consistent across study sites. These findings suggest that invasive bromes in the Great Basin might experience less rodent granivory than native congeners, which is consistent with a key prediction derived from the enemy release hypothesis.