Invasive Herbivores Research Articles

Heat waves affect an invasive herbivore and its parasitoid differentially with impacts beyond the first generation

AbstractHeat waves are becoming more frequent with climate change, and their impact on lower and higher trophic levels can differ. Higher trophic levels, such as predators and parasitoids, are predicted to be more strongly affected by heat waves than herbivores because they may have smaller thermal windows and because of altered ecological interactions with lower trophic levels. We tested the effects of heat waves of varying intensity (36–42°C), imposed for 4 h during five consecutive days on the egg stage of an invasive herbivore, the brown marmorated stink bug (Halyomorpha halys), and on various developmental stages of its parasitoid,Trissolcus japonicus.Halyomorpha halyseggs had lower hatching success and fewer nymphs emerging with increasing intensity of heat stress, while exposure during the egg and larval stages ofT. japonicusdid not decrease adult emergence rates. Heat waves increased developmental time of parasitoids and decreased adult emergence when pupal stages were exposed. Adult parasitoids experiencing heat stress had lower survival, but their parasitism success overall was not affected. In addition, we detected transgenerational effects, whereas emergence success and the number of adult parasitoids emerging in the second generation declined when the parental generation was exposed to 42°C. Contrary to expectations, the results suggest thatH. halysmay be more sensitive to heat stress thanT. japonicus, at least during early development. Our findings indicate that biological control services provided byT. japonicuscan be maintained in the face of increasing heat wave events and point to the importance of following populations beyond the first generation to better understand the long‐term consequences of heat stress.

Host Range Expansion by the Invasive Herbivore Corythucha marmorata (Uhler, 1878) is not Caused by Better Quality of New Hosts

Phytophagous insects may become serious pests of crops when introduced into a new place. Better nutritional quality and lower toxicity of new host plants and escape from natural enemies can enhance survival. The chrysanthemum lace bug, Corythucha marmorata (Hemiptera: Tingidae), is native to North America, where it exploits mainly goldenrod and its relatives (Asteraceae). It was accidentally introduced into Japan by about 2000. Since then, many reports of injury to sweet potato (Convolvulaceae) and eggplant (Solanaceae) by this species have been published. Here, we tested larval performance on goldenrod, sweet potato, eggplant, and three other known or potential host plants to investigate why the lace bug began to exploit the new host plants. Survival to adult stage was nil on eggplant, extremely low on blue daze (Convolvulaceae) and crown daisy (Asteraceae), moderate on sweet potato, and ca. 80% on goldenrod and sunflower. Developmental time was shorter and adults grew larger on goldenrod and sunflower than on the other plants. These results show that plant nutritional or toxicological qualities are not major factors that facilitate host range expansion of C. marmorata in Japan. Possible factors are discussed.

The dynamic trophic niche of an island bird of prey.

Optimal foraging theory predicts an inverse relationship between the availability of preferred prey and niche width in animals. Moreover, when individuals within a population have identical prey preferences and preferred prey is scarce, a nested pattern of trophic niche is expected if opportunistic and selective individuals can be identified. Here, we examined intraspecific variation in the trophic niche of a resident population of striated caracara (Phalcoboenus australis) on Isla de los Estados (Staten Island), Argentina, using pellet and stable isotope analyses. While this raptor specializes on seabird prey, we assessed this population's potential to forage on terrestrial prey, especially invasive herbivores as carrion, when seabirds are less accessible. We found that the isotopic niche of this species varies with season, age, breeding status, and, to a lesser extent, year. Our results were in general consistent with classic predictions of the optimal foraging theory, but we also explore other possible explanations for the observed pattern. Isotopic niche was broader for groups identified a priori as opportunistic (i.e., nonbreeding adults during the breeding season and the whole population during the nonbreeding season) than it was for individuals identified a priori as selective. Results suggested that terrestrial input was relatively low, and invasive mammals accounted for no more than 5% of the input. The seasonal pulse of rockhopper penguins likely interacts with caracara's reproductive status by constraining the spatial scale on which individuals forage. Niche expansion in spatially flexible individuals did not reflect an increase in terrestrial prey input; rather, it may be driven by a greater variation in the types of marine prey items consumed.

Fresh and Dry Weight Relations Are Predictors of Cycas micronesica Seed Age

Cycas micronesica is a foundation species in several Micronesian islands and its seeds have been a historical source of starch for the island residents. The species has become endangered by invasive specialist insect herbivores and conservationists struggle with the inability to estimate the age of observed seeds. To inform this agenda, we evaluated numerous Cycas micronesica seed traits to determine if any exhibited a relationship with age and a substantial change in absolute value. Of the 30 direct and derived seed traits that we evaluated, most of them were non-linear and exhibited minimal change after about 12 months in age. The only traits that emerged as unambiguous estimators of age were the quotients derived as gametophyte fresh weight/total seed fresh weight and sarcotesta dry weight/sclerotesta dry weight. These two simple metrics can be used to accurately estimate seed age for this arborescent cycad species.

Tritrophic interaction between the Mexican sunflower, the aphid Aphis gossypii and natural enemies in a greenhouse experiment

The establishment of invasive plants negatively affects natural environments. Invasive herbivores that attack weeds can be used as a form of biological control, but natural enemies of herbivores must be associated with this interaction to prevent the invasive phytophagous from become a local pest. We performed a greenhouse experiment to evaluate how the cotton aphid, Aphis gossypii, a ok and invasive herbivore, affects the performance of the weed Tithonia diversifolia, the Mexican sunflower. We also examined the relationship between the aphid and local natural enemies. Seedlings of T. diversifolia were divided in two groups: one infested by the aphid and another not infested. After 22 days, we assessed the relationship between aphid abundance and the presence of natural enemies (Coccinelidae and Aphidius platensis) on infested plants, and compared the vegetative performance of the two seedling groups. Both natural enemies were positively related to high aphid density on infested plants. Plants infested by the aphid presented foliar necrosis and senescence, and a reduction of around 50% in leaf number, foliar area, shoot length and shoot, root and total plant weight compared to non-infested plants. These results indicate potential biological control of Mexican sunflower seedlings by the cotton aphid, and control of this aphid by the studied natural enemies.

Facilitation between invasive herbivores: hemlock woolly adelgid increases gypsy moth preference for and performance on eastern hemlock

1. Interactions between invertebrate herbivores with different feeding modes are common on long‐lived woody plants. In cases where one herbivore facilitates the success of another, the consequences for their shared host plant may be severe. Eastern hemlock (Tsuga canadensis), a canopy‐dominant conifer native to the eastern U.S., is currently threatened with extirpation by the invasive stylet‐feeding hemlock woolly adelgid (Adelges tsugae). The effect of adelgid on invasive hemlock‐feeding folivores remains unknown.2. This study evaluated the impact of feeding by hemlock woolly adelgid on gypsy moth (Lymantria dispar) larval preference for, and performance on, eastern hemlock. To assess preference, 245 field‐grown hemlocks were surveyed for gypsy moth herbivory damage and laboratory paired‐choice bioassays were conducted. To assess performance, gypsy moth larvae were reared to pupation on adelgid‐infested or uninfested hemlock foliage, and pupal weight, proportional weight gain, and larval period were analysed.3. Adelgid‐infested hemlocks experienced more gypsy moth herbivory than did uninfested control trees, and laboratory tests confirmed that gypsy moth larvae preferentially feed on adelgid‐infested hemlock foliage. Gypsy moth larvae reared to pupation on adelgid‐infested foliage gained more weight than larvae reared on uninfested control foliage.4. These results suggest that the synergistic effect of adelgid and gypsy moth poses an additional threat to eastern hemlock that may increase extirpation risk and ecological impact throughout most of its range.

Egg parasitoid exploitation of plant volatiles induced by single or concurrent attack of a zoophytophagous predator and an invasive phytophagous pest

Zoophytophagous insect predators can induce physiological responses in plants by activating defence signalling pathways, but whether plants can respond to facultative phytophagy by recruiting natural enemies remains to be investigated. In Y-tube olfactometer bioassays, using a system including a Vicia faba plant, the zoophytophagous predator Podisus maculiventris and the egg parasitoid Telenomus podisi, we first demonstrated that T. podisi females are attracted by broad bean plants damaged by feeding activity of P. maculiventris and on which host egg masses had been laid, while they are not attracted by undamaged plants or plants damaged by feeding activity alone. In a second experiment, we evaluated the impact of the invasive phytophagous pest Halyomorpha halys on this plant volatile-mediated tritrophic communication. Results showed that the invasive herbivorous adults do not induce plants to recruit the native egg parasitoid, but they can disrupt the local infochemical network. In fact, T. podisi females are not attracted by volatiles emitted by plants damaged by H. halys feeding alone or combined with oviposition activity, nor are they attracted by plants concurrently infested by P. maculiventris and H. halys, indicating the specificity in the parasitoid response and the ability of the invasive herbivore in interrupting the semiochemical communication between plants and native egg parasitoids. To the best of our knowledge, this is the first study showing that zoophytophagous predator attacks induce indirect plant defences similarly to those defence strategies adopted by plants as a consequence of single or concurrent infestations of herbivorous insects.

Anatomy of the stridulation apparatus of the beech leaf‐mining weevil and characterization of, and behavioral responses to, stridulation sounds

AbstractWe investigated auditory signals and morphology of the stridulatory apparatus of the European beech leaf‐mining weevil, Orchestes fagi L. (Coleoptera: Curculionidae), an invasive herbivore now established in Nova Scotia, Canada, to determine their potential for enhancing survey tools to monitor the spread of the species in Canada. We recorded and described sounds produced by adult O. fagi, analyzed the morphology of the stridulatory mechanism for intersexual differences and asymmetry, and examined behavioral responses elicited in conspecifics by playback of stridulation recordings. Adult O. fagi produced sounds under three conditions: male in distress, female in distress, and male in the presence of female. Female distress chirps lasted significantly longer than male distress chirps and male chirps in the presence of females, but peak frequencies and mean number of chirps per s did not differ significantly among the three groups. Morphology of the stridulation structures in male and female O. fagi was compared using scanning electron microscopy. Orchestes fagi have an elytro‐tergal file‐ and scraper‐type sound production apparatus, through which sound is produced upon anterior motion of the abdomen. Female O. fagi have a ‘pars stridens’ that is longer and has more ridges than males. Width and number of ridges per length of pars stridens did not differ between the sexes. Evidence of asymmetry was found in male pars stridens, with the right side being longer than the left. Playback of recorded sounds to adult weevils suggests female O. fagi were repelled by sounds produced by distressed males.

Herbivore range expansion triggers adaptation in a subsequently-associated third trophic level species and shared microbial symbionts

Invasive species may change the life history strategies, distribution, genetic configuration and trophic interactions of native species. The diamondback moth, Plutella xylostella L., is an invasive herbivore attacking cultivated and wild brassica plants worldwide. Here we present phylogeographic analyses of P. xylostella and one of its major parasitoids, Cotesia vestalis, using mitochondrial markers, revealing the genetic diversity and evolutionary history of these two species. We find evidence that C. vestalis originated in Southwest China, then adapted to P. xylostella as a new host by ecological sorting as P. xylostella expanded its geographic range into this region. Associated with the expansion of P. xylostella, Wolbachia symbionts were introduced into local populations of the parasitoid through horizontal transfer from its newly associated host. Insights into the evolutionary history and phylogeographic system of the herbivore and its parasitoid provide an important basis for better understanding the impacts of biological invasion on genetic configuration of local species.

Elevational and structural shifts in the treeline of an oceanic island (Tenerife, Canary Islands) in the context of global warming

Global warming is changing the structure and elevational limits of treelines around the world. This could become a threat particularly on islands, where usually high mountain ecosystems occupy small areas. Tenerife, with a maximum elevation of 3715 m, is an excellent example of this. In this subtropical island, the treeline composed by endemic pine forests is expected to go up in elevation due to global warming, invading the summit scrub ecosystem. However, there is a lack of knowledge about shifts and trends of the island treeline from a multi-temporal perspective and how invasive herbivores are influencing these changes. In this study, we evaluated the past and current state of the Pinus canariensis treeline on the southern areas of Tenerife, where natural forests persist, and an increase in temperature due to global warming has been already detected in the last decades. For that purpose, we counted the number of pine trees in aerial photographs for the years 1963, 1987 and 2016 and performed generalized additive models to evaluate the role of the different macro-variables involved in the regeneration processes. Complementarily, we performed ten transects to evaluate current forest structure and the influence of invasive herbivores (rabbits and mouflons) from 1600 m to the upper limits (2400 m). Our results reveal an increase in tree density and slow but consistent advance of the treeline in this part of the island during the last 53 years. Interestingly, positive relationships were found between number of trees and temperature. On the contrary, negative correlation was detected between seedlings and saplings and herbivores, a factor that is influencing the forests structure at all elevation levels. Our results show the importance of the herbivore control to ensure a healthy forest structure that allow an adequate migratory capacity of the species with the global warming.

Do native and invasive herbivores have an effect on Brassica rapa pollination?

Mutualistic (e.g. pollination) and antagonistic (e.g. herbivory) plant-insect interactions shape levels of plant fitness and can have interactive effects. By using experimental plots of Brassica rapa plants infested with generalist (Mamestra brassicae) and specialised (Pieris brassicae) native herbivores and with a generalist invasive (Spodoptera littoralis) herbivore, we estimated both pollen movement among treatments and the visiting behaviour of honeybees versus other wild pollinators. Overall, we found that herbivory has weak effects on plant pollen export, either in terms of inter-treatment movements or of dispersion distance. Plants infested with the native specialised herbivore tend to export less pollen to other plants with the same treatment. Other wild pollinators preferentially visit non-infested plants that differ from those of honeybees, which showed no preferences. Honeybees and other wild pollinators also showed different behaviours on plants infested with different herbivores, with the former tending to avoid revisiting the same treatment and the latter showing no avoidance behaviour. When taking into account the whole pollinator community, i.e. the interactive effects of honeybees and other wild pollinators, we found an increased avoidance of plants infested by the native specialised herbivore and a decreased avoidance of plants infested by the invasive herbivore. Taken together, our results suggest that herbivory may have an effect on B. rapa pollination, but this effect depends on the relative abundance of honeybees and other wild pollinators.

Conifer responses to a stylet‐feeding invasive herbivore and induction with methyl jasmonate: impact on the expression of induced defences and a native folivore

Abstract Trees attacked by multiple herbivores need to defend themselves against dynamic biotic challenges; appropriate responses to one stressor can elicit hormonal responses that are antagonistic to another. Hemlock (Tsuga canadensis) infestation by hemlock woolly adelgid (HWA; Adelges tsugae) results in the accumulation of the defensive hormone salicylic acid. We explored the potential for HWA infestation to interfere with anti‐folivore‐induced defence signalling and its implications for a native folivore (hemlock looper; Lambdina fiscellaria). Hemlocks were infested with HWA and/or sprayed with methyl jasmonate; foliar defences were analyzed and foliage quality for looper larvae was assessed. Both treatments activated foliar defensive traits, including a HWA‐mediated increase in peroxidase activity and an accumulation of cell wall‐bound phenolics and lignin, as well as a methyl jasmonate‐mediated increase in lipoxygenase activity. The two treatments had an additive effect on other defensive traits and both treatments negatively affected looper performance. These results suggest that salicylic acid and jasmonic acid are not strictly antagonistic in conifers and that both have a role in anti‐folivore defence signalling. The present study illustrates the need for a better understanding of hormone signalling, cross‐talk and induced responses in conifers.

The influence of warming on the biogeographic and phylogenetic dependence of herbivore-plant interactions.

Evolutionary experience and the phylogenetic relationships of plants have both been proposed to influence herbivore–plant interactions and plant invasion success. However, the direction and magnitude of these effects, and how such patterns are altered with increasing temperature, are rarely studied. Through laboratory functional response experiments, we tested whether the per capita feeding efficiency of an invasive generalist herbivore, the golden apple snail, Pomacea canaliculata, is dependent on the biogeographic origin and phylogenetic relatedness of host plants, and how increasing temperature alters these dependencies. The feeding efficiency of the herbivore was highest on plant species with which it had no shared evolutionary history, that is, novel plants. Further, among evolutionarily familiar plants, snail feeding efficiency was higher on those species more closely related to the novel plants. However, these biogeographic dependencies became less pronounced with increasing temperature, whereas the phylogenetic dependence was unaffected. Collectively, our findings indicate that the susceptibility of plants to this invasive herbivore is mediated by both biogeographic origin and phylogenetic relatedness. We hypothesize that warming erodes the influence of evolutionary exposure, thereby altering herbivore–plant interactions and perhaps the invasion success of plants.

Exploring the relationships between marine protected areas and invasive fish in the world's most invaded sea.

Marine Protected Areas (MPAs) are an essential tool for marine biodiversity conservation. Yet, their effectiveness in protecting marine ecosystems from global stressors is debated. Biological invasions are a major driver of global change, causing biodiversity loss and altering ecosystem functioning. Here, we explored the relationships between MPAs and alien/native range-expanding fishes in the Mediterranean Sea, the world's most invaded sea. We surveyed fish and benthic communities in nine MPAs and adjacent unprotected areas across six countries. In the South and Eastern Mediterranean MPAs, the biomass of alien and native range-expanding fishes often exceeded 50% of the total fish biomass. Conversely, in the North and Western Mediterranean, alien fishes were absent. A negative relationship was found between native and alien species richness. Average and minimum sea surface temperature (SST) over six consecutive years were positively correlated with the total biomass of alien species; no alien fishes were recorded below 20.5°C average SST and 13.8°C minimum SST. We also found a negative relationship between alien fishes' biomass and the distance from the Suez Canal, which is the main pathway for the introduction of alien fish in the Mediterranean Sea. The biomass of alien and native range-expanding fishes was found to be higher in the South and Eastern Mediterranean MPAs than in adjacent unprotected areas. The association of barrens (rocky reefs deprived of vegetation) and invasive herbivores was observed at all eastern sites, regardless of protection status. Currently, the level of fishing pressure exerted on alien and native range-expanding fishes seems to be the most influential factor determining the lower biomass of invasive fishes at unprotected sites compared to MPAs. Our findings suggest that complementary management actions, such as species-targeted removals, should be taken in MPAs to effectively control invasive fish populations.

Associational susceptibility of a native shrub induced by context‐dependent attraction of an invasive herbivore

AbstractIndirect interactions among native and invasive species are notoriously difficult to predict. Here, we apply theory from trait‐mediated indirect effects and plant–insect interactions to explain the outcomes of multiple invader interactions. The present study investigates the roles of herbivore preference and plant ontogeny in mediating associational effects among a southern California native shrub, Atriplex canescens, an invasive annual forb, Brassica tournefortii, and an invasive stink bug, Bagrada hilaris. Ba. hilaris have been observed to form dense aggregations on A. canescens in late spring following senescence of Br. tournefortii, with subsequent increased mortality of A. canescens. A manipulative experiment found Ba. hilaris recruited to A. canescens in approximately 70 times greater numbers when neighbored by Br. tournefortii than when alone. Ba. hilaris nymph production, while minimized experimentally, occurred only on Br. tournefortii, suggesting A. canescens is not of sufficient quality for Ba. hilaris reproduction. In greenhouse preference trials, Ba. hilaris exhibited an overall preference for Br. tournefortii over A. canescens; however, the magnitude of preference depended on ontogenetic stage of Br. tournefortii. Overall, our results suggest significant potential for associational susceptibility mediated by a combination of Ba. hilaris preference for reproductive Br. tournefortii, substantial aggregation of Ba. hilaris onto Br. tournefortii, and a marked decline in Ba. hilaris preference for Br. tournefortii with advancing ontogeny—triggering spillover onto neighboring plants. Our results demonstrate an important role for multiple invaders and phenologically driven trait changes in mediating associational susceptibility.

Warming neutralizes host-specific competitive advantages between a native and invasive herbivore

Although native-invasive species interactions have become a common mechanism shaping ecosystems, whether these interactions shift under warming remains unclear. To investigate how warming may affect native and invasive species separately and together (intraspecific and interspecific competition, respectively) and whether any warming impact is resource dependent, we examined the performance of two competing herbivores (native Pieris canidia and invasive P. rapae) on two common host plants under three temperature settings (control, 3 °C, and 6 °C warming using environmental chambers). The results revealed that warming benefited the development and growth of both Pieris under intraspecific competition, but the benefits were host-plant dependent. Notably, the native or invasive Pieris gained an advantage from interspecific competition (host-plant dependent), but warming neutralized the competitive advantages of either Pieris species. These findings raise the possibility that warming-induced shifts in competitive status may become a crucial mechanism shaping ecosystems worldwide, because most ecosystems are challenged by species invasion and warming. Moreover, this study revealed a discrepancy in species thermal performance between intra- and interspecific competition. Therefore, to predict native-invasive species competition under warming, current thermal performance applications should use species performance curves derived from interspecific rather than intraspecific competition studies (although the latter is more readily available).

Chronic impacts of invasive herbivores on a foundational forest species: a whole‐tree perspective

Forests make up a large portion of terrestrial plant biomass, and the long-lived woody plants that dominate them possess an array of traits that deter consumption by forest pests. Although often extremely effective against native consumers, invasive species that avoid or overcome these defenses can wreak havoc on trees and surrounding ecosystems. This is especially true when multiple invasive species co-occur, since interactions between invasive herbivores may yield non-additive effects on the host. While the threat posed by invasive forest pests is well known, long-term field experiments are necessary to explore these consumer-host interactions at appropriate spatial and temporal scales. Moreover, it is important to measure multiple variables to get a "whole-plant" picture of their combined impact. We report the results of a 4-yr field experiment addressing the individual and combined impacts of two invasive herbivores, the hemlock woolly adelgid (Adelges tsugae) and elongate hemlock scale (Fiorinia externa), on native eastern hemlock (Tsuga canadensis) in southern New England. In 2011, we planted 200 hemlock saplings into a temperate forest understory and experimentally manipulated the presence/absence of both herbivore species; in 2015, we harvested the 88 remaining saplings and assessed plant physiology, growth, and resource allocation. Adelgids strongly affected hemlock growth: infested saplings had lower above/belowground biomass ratios, more needle loss, and produced fewer new needles than control saplings. Hemlock scale did not alter plant biomass allocation or growth, and its co-occurrence did not alter the impact of adelgid. While both adelgid and scale impacted the concentrations of primary metabolites, adelgid effects were more pronounced. Adelgid feeding simultaneously increased free amino acids local to feeding sites and a ~30% reduction in starch. The cumulative impact of adelgid-induced needle loss, manipulation of nitrogen pools, and the loss of stored resources likely accelerates host decline through disruption of homeostatic source-sink dynamics occurring at the whole-plant level. Our research stresses the importance of considering long-term impacts to predict how plants will cope with contemporary pressures experienced in disturbed forests.

Impact of an invasive herbivore and human trampling on lichen-rich dry grasslands: Soil-dependent response of multiple taxa

Dry grasslands are listed among the habitats of conservation concern in Europe. Here, based on a multitaxon approach including vascular plants, bryophytes and lichens, we explored the effects of small-scale disturbance on lichen-rich dry grassland vegetation by surveying 60 sites across the Po Plain (Northern Italy). In particular, we evaluated the impact of human trampling and fecal pellet deposition by the alien invasive herbivore Sylvilagus floridanus. We found a soil-dependent response of multiple taxa to the impact of the herbivore. For plants, beside a negative effect of trampling, the interaction between fecal pellet amount and soil pH indicates that the negative effect of the invasive herbivore is stronger on acidic soils. Bryophyte cover increased with increasing soil pH, annual rainfall and fecal pellet, while it was not affected by trampling. Lichen richness and cover decreased with increasing soil pH. The marginal interaction between soil pH and amount of fecal pellet indicates that the more negative effects on lichens may be expected on calcareous soils. Trampling did not affect lichen patterns and the rainfall gradient marginally affected lichen cover with a negative effect. Lichen species richness is also negatively affected by increasing vascular plant cover. The main implications of this study for improving conservation are: (1) conservation practices should be tailored to organism and substrate type; (2) bryophyte and lichen diversity patterns are influenced also by climatic conditions, suggesting that the impact on these organisms may be exacerbated by climate change; and (3) strict conservation, even through active exclusion of wild fauna, of the most species-rich sites should be recommended, even if previous literature and the negative plant cover-lichen richness relationship found in this study indicate that moderate mechanical disturbance could be a practical tool to enhance cryptogams.

Two invasive herbivores on a shared host: patterns and consequences of phytohormone induction.

Herbivore-induced changes in host quality mediate indirect interactions between herbivores. The nature of these indirect interactions can vary depending on the identity of herbivores involved, species-specific induction of defense-signaling pathways, and sequence of attack. However, our understanding of the role of these signaling pathways in the success of multiple exotic herbivores is less known. Eastern hemlock (Tsuga canadensis) is attacked by two invasive herbivores [elongate hemlock scale (EHS; Fiorinia externa) and hemlock woolly adelgid (HWA; Adelges tsugae)] throughout much of its range, but prior attack by EHS is known to deter HWA. The potential role of phytohormones in this interaction is poorly understood. We measured endogenous levels of phytohormones in eastern hemlock in response to attack by these invasive herbivores. We also used exogenous application of methyl jasmonate (MJ) and acibenzolar-S-methyl (ASM), a salicylic acid (SA) pathway elicitor, to test the hypothesis that defense-signaling phytohormones typically induced by herbivores could deter HWA. Resistance to adelgid attack was assessed using a behavioral assay. Adelgid feeding significantly elevated both abscisic acid (ABA) and SA in local tissues, while EHS feeding had no detectable effect on either phytohormone. HWA progrediens and sistens crawlers preferred to settle on ASM-treated foliage. In contrast, HWA crawlers actively avoided settlement on MJ-treated foliage. We suggest that induction of ABA- and SA-signaling pathways, in concert with defense-signaling interference, may aid HWA invasion success, and that defense-signaling interference, induced by exotic competitors, may mediate resistance of native hosts.

Resource Availability, Propagule Supply, and Effect of Nonnative Ungulate Herbivores onSenecio madagascariensisInvasion

Nonnative invasive herbivores can create complex biotic interactions by differentially feeding on native and nonnative invasive plant species. The herbivores may act as enemies of nonnative plants and prevent them from becoming invasive, or they may facilitate invasion by having a greater negative impact on native plants compared with nonnative plants. It is also possible that within the same ecosystem nonnative herbivores could either facilitate or inhibit invasion under different abiotic or biotic conditions. In this study we experimentally investigated how abiotic (soil nutrients) and biotic (propagule density) conditions influence the effect of invasive generalist herbivores on Senecio madagascariensis, an invasive plant species in Hawaiian dry forest plant communities. We used fenced exclosures to manipulate presence or absence of invasive ungulates (feral goats and sheep), and we used seed addition to manipulate propagule supply of S. madagascariensis. The experiment was replicated in a recently burned and an unburned site to examine how a resource pulse following fire may alter plant-herbivore interactions. There were very few seeds of S. madagascariensis in the seed rain of both sites, and recruitment was four times higher when seeds were experimentally added, suggesting that S. madagascariensis is dispersal limited in this area. Recruitment of S. madagascariensis was five times higher in the burned site compared to the unburned site, suggesting that increased resources promote recruitment. Recruitment was three times higher when herbivores were present compared to when they were excluded, but plants were much smaller when herbivores were present. We conclude that herbivores can alter S. madagascariensis recruitment, even during dry conditions, and that propagule availability influences where S. madagascariensis can become established.