Herbivory by an introduced Asian weevil negatively affects population growth of an invasive Brazilian shrub in Florida

The enemy release hypothesis (ERH) is often cited to explain why some plants successfully invade natural communities while others do not. This hypothesis maintains that plant populations are regulated by coevolved enemies in their native range but are relieved of this pressure where their enemies have not been co-introduced. Some studies have shown that invasive plants sustain lower levels of herbivore damage when compared to native species, but how damage affects fitness and population dynamics remains unclear. We used a system of co-occurring native and invasive Eugenia congeners in south Florida (USA) to experimentally test the ERH, addressing deficiencies in our understanding of the role of natural enemies in plant invasion at the population level. Insecticide was used to experimentally exclude insect herbivores from invasive Eugenia uniflora and its native co-occurring congeners in the field for two years. Herbivore damage, plant growth, survival, and population growth rates for the three species were then compared for control and insecticide-treated plants. Our results contradict the ERH, indicating that E. uniflora sustains more herbivore damage than its native congeners and that this damage negatively impacts stem height, survival, and population growth. In addition, most damage to E. uniflora, a native of Brazil, is carried out by Myllocerus undatus, a recently introduced weevil from Sri Lanka, and M. undatus attacks a significantly greater proportion of E. uniflora leaves than those of its native congeners. This interaction is particularly interesting because M. undatus and E. uniflora share no coevolutionary history, having arisen on two separate continents and come into contact on a third. Our study is the first to document negative population-level effects for an invasive plant as a result of the introduction of a novel herbivore. Such inhibitory interactions are likely to become more prevalent as suites of previously noninteracting species continue to accumulate and new communities assemble worldwide.     

Plant invasions, generalist herbivores, and novel defense weapons

One commonly accepted mechanism for biological invasions is that species, after introduction to a new region, leave behind their natural enemies and therefore increase in distribution and abundance. However, which enemies are escaped remains unclear. Escape from specialist invertebrate herbivores has been examined in detail, but despite the profound effects of generalist herbivores in natural communities their potential to control invasive species is poorly understood. We carried out parallel laboratory feeding bioassays with generalist invertebrate herbivores from the native (Europe) and from the introduced (North America) range using native and nonnative tetraploid populations of the invasive spotted knapweed, Centaurea stoebe. We found that the growth of North American generalist herbivores was far lower when feeding on C. stoebe than the growth of European generalists. In contrast, North American and European generalists grew equally well on European and North American tetraploid C. stoebe plants, lending no support for an evolutionary change in resistance of North American tetraploid C. stoebe populations against generalist herbivores. These results suggest that biogeographical differences in the response of generalist herbivores to novel plant species have the potential to affect plant invasions.     

Combining optimal defense theory and the evolutionary dilemma model to refine predictions regarding plant invasion

Optimal defense theory posits that plants with limited resources deploy chemical defenses based on the fitness value of different tissues and their probability of attack. However, what constitutes optimal defense depends on the identity of the herbivores involved in the interaction. Generalists, which are not tightly coevolved with their many host plants, are typically deterred by chemical defenses, while coevolved specialists are often attracted to these same chemicals. This imposes an "evolutionary dilemma" in which generalists and specialists exert opposing selection on plant investment in defense, thereby stabilizing defenses at intermediate levels. We used the natural shift in herbivore community composition that typifies many plant invasions to test a novel, combined prediction of optimal defense theory and the evolutionary dilemma model: that the within-plant distribution of defenses reflects both the value of different tissues (i.e., young vs. old leaves) and the relative importance of specialist and generalist herbivores in the community. Using populations of Verbascum thapsus exposed to ambient herbivory in its native range (where specialist and generalist chewing herbivores are prevalent) and its introduced range (where only generalist chewing herbivores are prevalent), we illustrate significant differences in the way iridoid glycosides are distributed among young and old leaves. Importantly, high-quality young leaves are 6.5x more highly defended than old leaves in the introduced range, but only 2x more highly defended in the native range. Additionally, defense levels are tracked by patterns of chewing damage, with damage restricted mostly to low-quality old leaves in the introduced range, but not the native range. Given that whole-plant investment in defense does not differ between ranges, introduced mullein may achieve increased fitness simply by optimizing its within-plant distribution of defense in the absence of certain specialist herbivores.     

When there is no escape: the effects of natural enemies on native, invasive, and noninvasive plants

An important question in the study of biological invasions is the degree to which successful invasion can be explained by release from control by natural enemies. Natural enemies dominate explanations of two alternate phenomena: that most introduced plants fail to establish viable populations (biotic resistance hypothesis) and that some introduced plants become noxious invaders (natural enemies hypothesis). We used a suite of 18 phylogenetically related native and nonnative clovers (Trifolium and Medicago) and the foliar pathogens and invertebrate herbivores that attack them to answer two questions. Do native species suffer greater attack by natural enemies relative to introduced species at the same site? Are some introduced species excluded from native plant communities because they are susceptible to local natural enemies? We address these questions using three lines of evidence: (1) the frequency of attack and composition of fungal pathogens and herbivores for each clover species in four years of common garden experiments, as well as susceptibility to inoculation with a common pathogen; (2) the degree of leaf damage suffered by each species in common garden experiments; and (3) fitness effects estimated using correlative approaches and pathogen removal experiments. Introduced species showed no evidence of escape from pathogens, being equivalent to native species as a group in terms of infection levels, susceptibility, disease prevalence, disease severity (with more severe damage on introduced species in one year), the influence of disease on mortality, and the effect of fungicide treatment on mortality and biomass. In contrast, invertebrate herbivores caused more damage on native species in two years, although the influence of herbivore attack on mortality did not differ between native and introduced species. Within introduced species, the predictions of the biotic resistance hypothesis were not supported: the most invasive species showed greater infection, greater prevalence and severity of disease, greater prevalence of herbivory, and greater effects of fungicide on biomass and were indistinguishable from noninvasive introduced species in all other respects. Therefore, although herbivores preferred native over introduced species, escape from pest pressure cannot be used to explain why some introduced clovers are common invaders in coastal prairie while others are not.     

Direct and indirect effects of CO2, nitrogen, and community diversity on plant-enemy interactions

Resource abundance and plant diversity are two predominant factors hypothesized to influence the amount of damage plants receive from natural enemies. Many impacts of these environmental variables on plant damage are likely indirect and result because both resource availability and diversity can influence plant traits associated with attractiveness to herbivores or susceptibility to pathogens. We used a long-term, manipulative field experiment to investigate how carbon dioxide (CO2) enrichment, nitrogen (N) fertilization, and plant community diversity affect plant traits and the amount of herbivore and pathogen damage experienced by the common prairie legume Lespedeza capitata. We detected little evidence that CO2 or N affected plant traits; however, plants growing in high-diversity treatments (polycultures) were taller, were less pubescent, and produced thinner leaves (higher specific leaf area). Interestingly, we also detected little evidence that CO2 or N affect damage. Plants growing in polycultures compared to monocultures, however, experienced a fivefold increase in damage from generalist herbivores, 64% less damage from specialist herbivores, and 91% less damage from pathogens. Moreover, within diversity treatments, damage by generalist herbivores was negatively correlated with pubescence and often was positively correlated with plant height, while damage by specialist herbivores typically was positively correlated with pubescence and negatively associated with height. These patterns are consistent with changes in plant traits driving differences in herbivory between diversity treatments. In contrast, changes in measured plant traits did not explain the difference in disease incidence between monocultures and polycultures. In summary, our data provide little evidence that CO2 or N supply alter damage from natural enemies. By contrast, plants grown in monocultures experienced greater specialist herbivore and pathogen damage but less generalist herbivore damage than plants grown in diverse communities. Part of this diversity effect was mediated by changes in plant traits, many of which likely are plastic responses to diversity treatments, but some of which may be the result of evolutionary changes in response to these long-term experimental manipulations.     

Evaluating the impacts of multiple generalist fungal pathogens on temperate tree seedling survival

Host-specific mortality driven by natural enemies is a widely discussed mechanism for explaining plant diversity. In principle, populations of plant species can be regulated by distinct host-specific natural enemies that have weak or nonexistent effects on heterospecific competitors, preventing any single species from becoming dominant and thus promoting diversity. Two of the first steps in exploring the role of natural enemies in diversity regulation are to (1) identify potential enemies and (2) evaluate their levels of host specificity by determining if interactions between any one host and its enemy have equivalent survival impacts on co-occurring host species. We developed a bioinformatics framework to evaluate impacts of potential pathogens on seedling survival, for both single and multiple infections. Importantly, we consider scenarios not only if there are specialist pathogens for each plant, but also when generalist pathogens have differential effects on multiple host species, and when co-infection has species-specific effects. We then applied this analytical framework to a field experiment using molecular techniques to detect potential fungal pathogens on co-occurring tree seedling hosts. Combinatorial complexity created by 160 plant-fungus interactions was reduced to eight combinations that affect seedling survival. Potential fungal pathogens had broad host ranges, but seedling species were each regulated by different combinations of fungi or by generalist fungi that had differential effects on multiple plant species. Soil moisture can have the potential to shift the nature of the interactions in some plant-fungal combinations from neutral to detrimental. Reassessing the assumption of single-enemy-single-host interactions broadens the mechanisms through which natural enemies can influence plant diversity.     

Selecting for Tolerance against Pathogens and Herbivores to Enhance Success of Reintroduction and Translocation

Abstract: Some species have insufficient defenses against climate change, emerging infectious diseases, and non‐native species because they have not been exposed to these factors over their evolutionary history, and this can decrease their likelihood of persistence. Captive breeding programs are sometimes used to reintroduce individuals back into the wild; however, successful captive breeding and reintroduction can be difficult because species or populations often cannot coexist with non‐native pathogens and herbivores without artificial selection. In captive breeding programs, breeders can select for host defenses that prevent or reduce pathogen or herbivore burden (i.e., resistance) or traits that limit the effects of parasitism or herbivory on host fitness (i.e., tolerance). We propose that selection for host tolerance may enhance the success of reintroduction or translocation because tolerant hosts generally have neutral effects on introduced pathogens and herbivores. The release of resistant hosts would have detrimental effects on their natural enemies, promoting rapid evolution to circumvent the host resistance that may reduce the long‐term probability of persistence of the reintroduced or translocated species. We examined 2 case studies, one on the pathogenic amphibian chytrid fungus ( Batrachochytrium dendrobatidis [Bd]) and the other on the herbivorous cactus moth ( Cactoblastis cactorum) in the United States, where it is not native. In each case study, we provide recommendations for how captive breeders and managers could go about selecting for host tolerance. Selecting for tolerance may offer a promising tool to rescue hosts species from invasive natural enemies as well as new natural enemies associated with climate change‐induced range shifts.     

On the risk of extinction of a wild plant species through spillover of a biological control agent: Analysis of an ecosystem compartment model

Invasive plant species can be controlled by introducing natural enemies (insect herbivores) from their native range. However, such introduction entails the risk that the introduced herbivores attack indigenous plant species in the area of introduction. Here, we study the effect of spillover of a herbivore from a managed ecosystem compartment (agriculture) to a natural compartment (non-managed) and vice versa. In the natural compartment, an indigenous plant species is attacked by the introduced herbivores, whereas another indigenous plant species, which competes with the first, is not attacked. The combination of competition and herbivory may result in extinction of the attacked wild plant species. Using a modelling approach, we determine model parameters that characterize the risk of extinction for a wild plant species. Risk factors include: (1) a high attack rate of the herbivores on the wild non-target species, (2) niche overlap expressed as strong competition between the attacked non-target species and its competitor(s), and (3) factors favouring large spillover from the managed ecosystem compartment to the natural compartment; these include (3a) a high dispersal ability, and (3b) a moderate attack rate of the introduced herbivore on the target species, enabling large resident populations of the insect herbivore in the managed compartment. The analysis thus indicates that a high attack rate on the target species, which is a selection criterion for biocontrol agents with respect to their effectiveness, also mitigates risks resulting from spillover and non-target effects. While total eradication of an invasive plant species is not possible in the one-compartment-one-plant-one-herbivore system, natural enemy spillover from a natural to a managed compartment can make the invasive weed go extinct.     

Spillover of Agriculturally Subsidized Predators as a Potential Threat to Native Insect Herbivores in Fragmented Landscapes

Abstract:  Habitat loss and fragmentation can have strong negative impacts on populations of some native species. Spillover of generalist natural enemies from the surrounding landscape matrix is one mechanism potentially generating such effects, yet this has been rarely studied in insects. We examined the influence of habitat conversion to agriculture on the abundance and potential effects of predatory coccinellid beetles on native insect herbivores within 12 grassland remnants in central Nebraska (U.S.A.). Results of sweep sampling revealed that coccinellids were three to six times more abundant at native grassland sites embedded within cropland-dominated landscapes compared with control sites in grassland-dominated landscapes over the 3 years of the study. Exclusion experiments further demonstrated that predation intensity was strongly related to coccinellid abundances across sites and that coccinellids can dramatically reduce densities of a native aphid herbivore. In contrast to studies of specialized insect parasitoids, which have generally found reduced enemy pressure in fragmented landscapes, our results suggest that native herbivores may in some cases experience increased consumer pressure in landscapes with increasing habitat loss because of spillover of generalist predators from surrounding cropland habitats.     

Population and leaf-level variation of iridoid glycosides in the invasive weed Verbascum thapsus L. (common mullein): implications for herbivory by generalist insects

Plant–insect interactions, which are strongly mediated by chemical defenses, have the potential to shape invasion dynamics. Despite this, few studies have quantified natural variation in key defensive compounds of invasive plant populations, or how those defenses relate to levels of herbivory. Here, we evaluated variation in the iridoid glycosides aucubin and catalpol in rosette plants of naturally occurring, introduced populations of the North American invader, Verbascum thapsus L. (common mullein; Scrophulariaceae). We examined two scales that are likely to structure interactions with insect herbivores—among populations and within plant tissues (i.e., between young and old leaves). We additionally estimated the severity of damage incurred at these scales due to insect chewing herbivores (predominantly grasshoppers and caterpillars), and evaluated the relationship between iridoid glycoside content and leaf damage. We found significant variation in iridoid glycoside concentrations among populations and between young and old leaves, with levels of herbivory strongly tracking leaf-level investment in defense. Specifically, across populations, young leaves were highly defended by iridoids (averaging 6.5× the concentration present in old leaves, and containing higher proportions of the potentially more toxic iridoid, catalpol) and suffered only minimal damage from generalist herbivores. In contrast, old leaves were significantly less defended and accordingly more substantially utilized. These findings reveal that quantitative variation in iridoid glycosides is a key feature explaining patterns of herbivory in an introduced plant. In particular, these data support the hypothesis that defenses limit the ability of generalists to feed on mullein’s well-defended young leaves, resulting in minimal losses of high-quality tissue, and increasing performance of this introduced species.     

Herbivores on a dominant understory shrub increase local plant diversity in rain forest communities

Indirect effects of trophic interactions on biodiversity can be large and common, even in complex communities. Previous experiments with dominant understory Piper shrubs in a Costa Rican rain forest revealed that increases in herbivore densities on these shrubs caused widespread seedling mortality as a result of herbivores moving from Piper to seedlings of many different plant genera. We tested components of the Janzen-Connell hypothesis by conducting focused studies on the effects of specialist and generalist Piper herbivores on local seedling diversity. Whereas specialist herbivores are predicted to increase mortality to neighboring seedlings that are closely related to the source plant, true generalists moving from source plants may cause density-dependent mortality of many species, and possibly increase richness if new species replace abundant species that have been thinned by herbivores. Therefore, we hypothesized that seedling richness would be greater in understory control plots created in patches of Piper that had normal densities of generalist herbivores compared to plots from which we removed generalist herbivores manually from all Piper shrubs. After 15 months, generalist-herbivore-removal plots had > 40% fewer seedlings, > 40% fewer species, and 40% greater seedling evenness, on average, than control plots with generalist herbivores intact. Using a complementary approach in unmanipulated plots in four forests, we used path analysis to test for a positive association between seedling diversity and herbivore damage on Piper species. In unmanipulated plots, for both generalist and specialist herbivores, our data were significant fits to the causal model that Piper herbivores decrease evenness and increase plant species richness, corroborating the experimental results. Because herbivores changed how individuals were apportioned among the species and families present (lower evenness), one interpretation of these associations between herbivores on Piper shrubs and local seedling richness is that high seedling mortality in dominant families allowed the colonization or survival of less common species. If interspecific or apparent competition allowed for a relative increase in species richness, then the Janzen-Connell hypothesis may extend its predictions to generalist seedling predators. We speculate that apparent competition may explain some of the deviations from neutral model predictions, especially at small scales.     

外来植物入侵机制研究进展与展望

从植物入侵的内因、外因、驱动力和入侵过程等方面综述了植物入侵机制的研究进展:(1)外来植物由于预适应性较强或由于杂交和多倍体化而具有较强的侵染力:(2)外来植物由于逃脱原生区域的自然天敌,或由于土著种竞争较弱、生物多样性较低、植食动物或疾病较少等导致的生物抵抗力较低,或因良好的资源可获得性及入侵植物的化感作用等造成新栖息地可侵人性的增加;(3)自然或人为引入外来植物及对新栖息地的扰动是植物入侵的驱动力;(4)外来植物可能通过基于事件的入侵、入侵崩溃和协同入侵等人侵过程机制实现其成功入侵,分析了各入侵机制对入侵植物防控、生态学和进化学研究意义以及待解决问题.结果表明,植物入侵机制高度复杂,不同的外来植物种、新柄息地类型及植物入侵的不同阶段,各入侵机制可能单独或共同作用,为针对性地设计植物入侵防控方案提供了理论支撑.最后对植物入侵机制理论及其在植物入侵预测、危险性分区和防控中的应用研究等进行了展望.     

Role of plant enemies in the forestry of indigenous vs. nonindigenous pines

Plantations of rapidly growing trees are becoming increasingly common because the high productivity can enhance local economies, support improvements in educational systems, and generally improve the quality of life in rural communities. Landowners frequently choose to plant nonindigenous species; one rationalization has been that silvicultural productivity is enhanced when trees are separated from their native herbivores and pathogens. The expectation of enemy reduction in nonindigenous species has theoretical and empirical support from studies of the enemy release hypothesis (ERH) in the context of invasion ecology, but its relevance to forestry has not been evaluated. We evaluated ERH in the productive forests of Galicia, Spain, where there has been a profusion of pine plantations, some with the indigenous Pinus pinaster, but increasingly with the nonindigenous P. radiata. Here, one of the most important pests of pines is the indigenous bark beetle, Tomicus piniperda. In support of ERH, attacks by T. piniperda were more than twice as great in stands of P. pinaster compared to P. radiata. This differential held across a range of tree ages and beetle abundance. However, this extension of ERH to forestry failed in the broader sense because beetle attacks, although fewer on P. radiata, reduced productivity of P. radiata more than that of P. pinaster (probably because more photosynthetic tissue is lost per beetle attack in P. radiata). Productivity of the nonindigenous pine was further reduced by the pathogen, Sphaeropsis sapinea, which infected up to 28% of P. radiata but was absent in P. pinaster. This was consistent with the forestry axiom (antithetical to ERH) that trees planted "off-site" are more susceptible to pathogens. Fungal infections were positively correlated with beetle attacks; apparently T. piniperda facilitates S. sapinea infections by creating wounds and by carrying fungal propagules. A globally important component in the diminution of indigenous flora has been the deliberate large-scale propagation of nonnative trees for silviculture. At least for Pinus forestry in Spain, reduced losses to pests did not rationalize the planting of nonindigenous trees. There would be value in further exploration of relations between invasion ecology and the forestry of nonindigenous trees.     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

Direct and interactive effects of enemies and mutualists on plant performance: a meta-analysis

Plants engage in multiple, simultaneous interactions with other species; some (enemies) reduce and others (mutualists) enhance plant performance. Moreover, effects of different species may not be independent of one another; for example, enemies may compete, reducing their negative impact on a plant. The magnitudes of positive and negative effects, as well as the frequency of interactive effects and whether they tend to enhance or depress plant performance, have never been comprehensively assessed across the many published studies on plant-enemy and plant-mutualist interactions. We performed a meta-analysis of experiments in which two enemies, two mutualists, or an enemy and a mutualist were manipulated factorially. Specifically, we performed a factorial meta-analysis using the log response ratio. We found that the magnitude of (negative) enemy effects was greater than that of (positive) mutualist effects in isolation, but in the presence of other species, the two effects were of comparable magnitude. Hence studies evaluating single-species effects of mutualists may underestimate the true effects found in natural settings, where multiple interactions are the norm and indirect effects are possible. Enemies did not on average influence the effects on plant performance of other enemies, nor did mutualists influence the effects of mutualists. However, these averages mask significant and large, but positive or negative, interactions in individual studies. In contrast, mutualists ameliorated the negative effects of enemies in a manner that benefited plants; this overall effect was driven by interactions between pathogens and belowground mutualists (bacteria and mycorrhizal fungi). The high frequency of significant interactive effects suggests a widespread potential for diffuse rather than pairwise coevolutionary interactions between plants and their enemies and mutualists. Pollinators and mycorrhizal fungi enhanced plant performance more than did bacterial mutualists. In the greenhouse (but not the field), pathogens reduced plant performance more than did herbivores, pathogens were more damaging to herbaceous than to woody plants, and herbivores were more damaging to crop than to non-crop plants (suggesting evolutionary change in plants or herbivores following crop domestication). We discuss how observed differences in effect size might be confounded with methodological differences among studies.     

Remote analysis of biological invasion and the impact of enemy release

Escape from natural enemies is a widely held generalization for the success of exotic plants. We conducted a large-scale experiment in Hawaii (USA) to quantify impacts of ungulate removal on plant growth and performance, and to test whether elimination of an exotic generalist herbivore facilitated exotic success. Assessment of impacted and control sites before and after ungulate exclusion using airborne imaging spectroscopy and LiDAR, time series satellite observations, and ground-based field studies over nine years indicated that removal of generalist herbivores facilitated exotic success, but the abundance of native species was unchanged. Vegetation cover <1 m in height increased in ungulate-free areas from 48.7% +/- 1.5% to 74.3% +/- 1.8% over 8.4 years, corresponding to an annualized growth rate of lambda = 1.05 +/- 0.01 yr(-1) (median +/- SD). Most of the change was attributable to exotic plant species, which increased from 24.4% +/- 1.4% to 49.1% +/- 2.0%, (lambda = 1.08 +/- 0.01 yr(-1)). Native plants experienced no significant change in cover (23.0% +/- 1.3% to 24.2% +/- 1.8%, lambda = 1.01 +/- 0.01 yr(-1)). Time series of satellite phenology were indistinguishable between the treatment and a 3.0-km2 control site for four years prior to ungulate removal, but they diverged immediately following exclusion of ungulates. Comparison of monthly EVI means before and after ungulate exclusion and between the managed and control areas indicates that EVI strongly increased in the managed area after ungulate exclusion. Field studies and airborne analyses show that the dominant invader was Senecio madagascariensis, an invasive annual forb that increased from < 0.01% to 14.7% fractional cover in ungulate-free areas (lambda = 1.89 +/- 0.34 yr(-1)), but which was nearly absent from the control site. A combination of canopy LAI, water, and fractional cover were expressed in satellite EVI time series and indicate that the invaded region maintained greenness during drought conditions. These findings demonstrate that enemy release from generalist herbivores can facilitate exotic success and suggest a plausible mechanism by which invasion occurred. They also show how novel remote-sensing technology can be integrated with conservation and management to help address exotic plant invasions.     

Two herbivore-deterrent iridoid glycosides reduce the in-vitro growth of a specialist but not of a generalist pathogenic fungus of Plantago lanceolata L

Summary. Many secondary plant compounds are involved in defense against both insect herbivores and pathogens. Two secondary plant compounds of Plantago lanceolata, the iridoid glycosides catalpol and its precursor aucubin, are well known for their deterrent effects on generalist and non-adapted specialist insect herbivores. We tested the effects of these compounds on the in-vitro growth of a specialist and generalist fungal pathogen of this host species. Two chemical forms of these iridoids were tested. The glycosides and their aglycones, the products of enzymatic conversion by specific $/Beta$-glucosidase enzymes. The glycosides enhanced growth of both the specialist fungus Diaporthe adunca and the generalist fungus Fusarium moniliforme var. subglutinans. The positive effect of these glycosides on the generalist fungus is in sharp contrast with the generally negative effects of these glysosides on generalist insect herbivores. The aglycones of aucubin and catalpol reduced the growth of the specialist fungus D. adunca, but, contrary to expectation, enhanced the growth of the generalist fungus F. moniliforme var. subglutinans. Effects of aucubin on D. adunca were stronger than effects of catalpol. This was true both for the growth stimulating effects of the glycosides and for the fungitoxic effects of the aglycones. We therefore expect that the effects of these iridoids in P. lanceolata on the specialist fungus will strongly depend on the ratio between catalpol and its precursor aucubin and the chemical form (glycoside or aglycone) in which these compounds are encountered by the fungus during growth. Our results suggest that iridoid glycosides in P. lanceolata can be used as defense against both herbivores and pathogens, but that their effects are highly specific with respect to the natural enemy species that is encountered. Received 11 April 2002; accepted 9 August 2002     

Volatile chemicals from leaf litter are associated with invasiveness of a neotropical weed in Asia

Some invasive plant species appear to strongly suppress neighbors in their nonnative ranges but much less so in their native range. We found that in the field in its native range in Mexico, the presence of Ageratina adenophora, an aggressive Neotropical invader, was correlated with higher plant species richness than found in surrounding plant communities where this species was absent, suggesting facilitation. However, in two nonnative ranges, China and India, A. adenophora canopies were correlated with much lower species richness than the surrounding communities, suggesting inhibition. Volatile organic compound (VOC) signals may contribute to this striking biogeographical difference and the invasive success of A. adenophora. In controlled experiments volatiles from A. adenophora litter caused higher mortality of species native to India and China, but not of species native to Mexico. The effects of A. adenophora VOCs on seedling germination and growth did not differ between species from the native range and species from the nonnative ranges of the invader. Litter from A. adenophora plants from nonnative populations also produced VOCs that differed quantitatively in the concentrations of some chemicals than litter from native populations, but there were no chemicals unique to one region. Biogeographic differences in the concentrations of some volatile compounds between ranges suggest that A. adenophora may be experiencing selection on biochemical composition in its nonnative ranges.     

Acceleration of Exotic Plant Invasion in a Forested Ecosystem by a Generalist Herbivore

Abstract: The successful invasion of exotic plants is often attributed to the absence of coevolved enemies in the introduced range (i.e., the enemy release hypothesis). Nevertheless, several components of this hypothesis, including the role of generalist herbivores, remain relatively unexplored. We used repeated censuses of exclosures and paired controls to investigate the role of a generalist herbivore, white‐tailed deer (Odocoileus virginianus), in the invasion of 3 exotic plant species (Microstegium vimineum, Alliaria petiolata, and Berberis thunbergii) in eastern hemlock (Tsuga canadensis) forests in New Jersey and Pennsylvania (U.S.A.). This work was conducted in 10 eastern hemlock (T. canadensis) forests that spanned gradients in deer density and in the severity of canopy disturbance caused by an introduced insect pest, the hemlock woolly adelgid (Adelges tsugae). We used maximum likelihood estimation and information theoretics to quantify the strength of evidence for alternative models of the influence of deer density and its interaction with the severity of canopy disturbance on exotic plant abundance. Our results were consistent with the enemy release hypothesis in that exotic plants gained a competitive advantage in the presence of generalist herbivores in the introduced range. The abundance of all 3 exotic plants increased significantly more in the control plots than in the paired exclosures. For all species, the inclusion of canopy disturbance parameters resulted in models with substantially greater support than the deer density only models. Our results suggest that white‐tailed deer herbivory can accelerate the invasion of exotic plants and that canopy disturbance can interact with herbivory to magnify the impact. In addition, our results provide compelling evidence of nonlinear relationships between deer density and the impact of herbivory on exotic species abundance. These findings highlight the important role of herbivore density in determining impacts on plant abundance and provide evidence of the operation of multiple mechanisms in exotic plant invasion.     

Effects of soil biota from different ranges on Robinia invasion: acquiring mutualists and escaping pathogens

The net effects of soil biota on exotic invaders can be variable, in part, because net effects are produced by many interacting mutualists and antagonists. Here we compared mutualistic and antagonistic biota in soils collected in the native, expanded, and invasive range of the black locust tree, Robinia pseudoacacia. Robinia formed nodules in all soils with a broad phylogenetic range of N-fixing bacteria, and leaf N did not differ among the different sources of soil. This suggests that the global expansion of Robinia was not limited by the lack of appropriate mutualistic N-fixers. Arbuscular mycorrhizal fungi (AMF) from the native range stimulated stronger positive feedbacks than AMF from the expanded or invasive ranges, a biogeographic difference not described previously for invasive plants. Pythium taxa collected from soil in the native range were not more pathogenic than those from other ranges; however, feedbacks produced by the total soil biota were more negative from soils from the native range than from the other ranges, overriding the effects of AMF. This suggests that escape from other pathogens in the soil or the net negative effects of the whole soil community may contribute to superior performance in invaded regions. Our results suggest that important regional evolutionary relationships may occur among plants and soil biota, and that net effects of soil biota may affect invasion, but in ways that are not easily explained by studying isolated components of the soil biota.