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1.
Numerous hypotheses suggest that natural enemies can influence the dynamics of biological invasions. Here, we use a group of 12 related native, invasive, and naturalized vines to test the relative importance of resistance and tolerance to herbivory in promoting biological invasions. In a field experiment in Long Island, New York, we excluded mammal and insect herbivores and examined plant growth and foliar damage over two growing seasons. This novel approach allowed us to compare the relative damage from mammal and insect herbivores and whether damage rates were related to invasion. In a greenhouse experiment, we simulated herbivory through clipping and measured growth response. After two seasons of excluding herbivores, there was no difference in relative growth rates among invasive, naturalized, and native woody vines, and all vines were susceptible to damage from mammal and insect herbivores. Thus, differential attack by herbivores and plant resistance to herbivory did not explain invasion success of these species. In the field, where damage rates were high, none of the vines were able to fully compensate for damage from mammals. However, in the greenhouse, we found that invasive vines were more tolerant of simulated herbivory than native and naturalized relatives. Our results indicate that invasive vines are not escaping herbivory in the novel range, rather they are persisting despite high rates of herbivore damage in the field. While most studies of invasive plants and natural enemies have focused on resistance, this work suggests that tolerance may also play a large role in facilitating invasions.  相似文献   

2.
Release from natural enemies may favor invasive plants evolving traits associated with reduced herbivore‐resistance and faster‐growth in introduced ranges. Given a genetic trade‐off between resistance and tolerance, invasive plants could also become more tolerant to herbivory than conspecifics in the native range. We conducted a field common garden study in the native range of Sapium sebiferum using seeds from native Chinese populations and invasive North American populations to compare their growth and herbivory resistance. We also performed a cage‐pot experiment to compare their resistance and tolerance to Bikasha collaris beetles that are specialist feeders on S. sebiferum trees in China. Results of the common garden study showed that Sapium seedlings of invasive populations relative to native populations were more frequently attacked by native herbivores. Growth and leaf damage were significantly higher for invasive populations than for native populations. Growth of invasive populations was not significantly affected by insecticide spray, but insecticide spray benefited that of native populations. In the bioassay trial, beetles preferentially consumed leaf tissue of invasive populations compared to native populations when beetles had a choice between them. Regression of percent leaf damage on biomass showed that invasive populations tolerated herbivory more effectively than native populations. Our results suggest that S. sebiferum from the introduced range had lower resistance but higher tolerance to specialist herbivores. Both defense strategies could have evolved as a response to the escape from natural enemies in the introduced range.  相似文献   

3.
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.  相似文献   

4.
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.  相似文献   

5.
During introduction, invasive plants can be released from specialist herbivores, but may retain generalist herbivores and encounter novel enemies. For fast-growing invasive plants, tolerance of herbivory via compensatory regrowth may be an important defense against generalist herbivory, but it is unclear whether tolerance responses are specifically induced by different herbivores and whether specificity differs among native and invasive plant populations. We conducted a greenhouse experiment to examine the variation among native and invasive populations of Chinese tallow tree, Triadica sebifera, in their specificity of tolerance responses to herbivores by exposing plants to herbivory from either one of two generalist caterpillars occurring in the introduced range of Triadica. Simultaneously, we measured the specificity of another defensive trait, extrafloral nectar (EFN) production, to detect potential tradeoffs between resistance and tolerance of herbivores. Invasive populations had higher aboveground biomass tolerance than native populations, and responded non-specifically to either herbivore, while native populations had significantly different and specific aboveground biomass responses to the two herbivores. Both caterpillar species similarly induced EFN in native and invasive populations. Plant tolerance and EFN were positively correlated or had no relationship and biomass in control and herbivore-damaged plants was positively correlated, suggesting little costs of tolerance. Relationships among these vegetative traits depended on herbivore type, suggesting that some defense traits may have positive associations with growth-related processes that are differently induced by herbivores. Importantly, loss of specificity in invasive populations indicates subtle evolutionary changes in defenses in invasive plants that may relate to and enhance their invasive success.  相似文献   

6.
The evolution of increased competitive ability (EICA) hypothesis predicts that release from natural enemies in the introduced range favors exotic plants evolving to have greater competitive ability and lower herbivore resistance than conspecifics from the native range. We tested the EICA hypothesis in a common garden experiment with Sapium sebiferum in which seedlings from native (China) and invasive (USA) populations were grown in all pairwise combinations in the native range (China) in the presence of herbivores. When paired seedlings were from the same continent, shoot mass and leaf damage per seedling were significantly greater for plants from invasive populations than those from native populations. Despite more damage from herbivores, plants from invasive populations still outperformed those from native populations when they were grown together. Increased competitive ability and higher herbivory damage of invasive populations relative to native populations of S. sebiferum support the EICA hypothesis. Regression of biomass against percent leaf damage showed that plants from invasive populations tolerated herbivory more effectively than those from native populations. The results of this study suggest that S. sebiferum has become a faster-growing, less herbivore-resistant, and more herbivore-tolerant plant in the introduced range. This implies that increased competitive ability of exotic plants may be associated with evolutionary changes in both resistance and tolerance to herbivory in the introduced range. Understanding these evolutionary changes has important implications for biological control strategies targeted at problematic invaders.  相似文献   

7.
Rapid post-introduction evolution has been found in many invasive plant species, and includes changes in defence (resistance and tolerance) and competitive ability traits. Here, we explored the post-introduction evolution of a trade-off between resistance to and tolerance of herbivory, which has received little attention. In a common garden experiment in a native range, nine invasive and 16 native populations of Brassica nigra were compared for growth and defence traits. Invasive populations had higher resistance to, but lower tolerance of, herbivore damage than native populations. Invasive populations survived better and produced more seeds than native ones when released from herbivores; but fitness was equivalent between the regions under ambient herbivory. The invasive populations grew taller, and produced more biomass and lighter seeds than natives, irrespective of insecticide treatment. In addition to supporting the idea of post-introduction rapid evolution of plant traits, our results also contribute to an emerging pattern of both increasing resistance and growth in invasive populations, contrary to the predictions of earlier theories of resistance-growth trade-offs.  相似文献   

8.
Tolerance, the degree to which plant fitness is affected by herbivory, is associated with invasiveness and biological control of introduced plant species. It is important to know the evolutionary changes in tolerance of invasive species after introduction in order to understand the mechanisms of biological invasions and assess the feasibility of biological control. While many studies have explored the evolutionary changes in resistance of invasive species, little has been done to address tolerance. We hypothesized that compared with plants from native populations, plants from invasive populations may increase growth and decrease tolerance to herbivory in response to enemy release in introduced ranges. To test this hypothesis, we compared the differences in growth and tolerance to simulated herbivory between plants from invasive and native populations of Chromolaena odorata, a noxious invader of the tropics and subtropics, at two nutrient levels. Surprisingly, flower number, total biomass (except at high nutrient), and relative increase in height were not significantly different between ranges. Also, plants from invasive populations did not decrease tolerance to herbivory at both nutrient levels. The invader from both ranges compensated fully in reproduction after 50?% of total leaf area had been damaged, and achieved substantial regrowth after complete shoot damage. This strong tolerance to damage was associated with increased resource allocation to reproductive structures and with mobilization of storage reserves in roots. The innately strong tolerance may facilitate invasion success of C. odorata and decrease the efficacy of leaf-feeding biocontrol agents. Our study highlights the need for further research on biogeographical differences in tolerance and their role in the invasiveness of exotic plants and biological control.  相似文献   

9.
Several hypotheses have been proposed to explain the defense strategies of invasive plants in new ranges. In the absence of specialist herbivores, it is believed that invasive plants may allocate fewer resources to resistance and more to growth and reproduction, thus increasing tolerance to damage in the invasive genotypes. In order to test these predictions, we compared both performance (growth and reproduction) and defense strategies (tolerance and resistance) of two populations of Taraxacum officinale, one from the native range in the French Alps, and one from the introduced range in the Chilean Andes. Individuals from the introduced population demonstrated improved reproductive traits relative to those from the native population, although there was no discernible difference in biomass accumulation. Additionally, reduced tolerance was evident in the case of the former; whereas fitness traits of native plants were unaffected by damage, invasive plants reduced growth and seed output by 25 and 30% respectively following damage treatments. Increases in levels of phenols and anthocyanins, produced as a defense response to herbivory, were observed in introduced plants. Our results suggest that reallocation of resources to reproduction may be an important factor favouring invasive success of T. officinale in Chile, and that a higher investment in chemical resistance traits in this population may also be a factor in this regard.  相似文献   

10.
Although induced defenses are widespread in nature, and a potentially important strategy used by invasive plants, it is unclear how induced defenses vary among populations and whether the intensity and duration of induced defenses depends on herbivore type. For invasive plants, low herbivore loads in their introduced ranges can lead to differences in herbivore defense compared to their native ranges, but we currently know little about how induced defenses vary among native and invasive populations. We conducted a greenhouse experiment to examine variation in one type of induced defense, extrafloral nectar (EFN) production, among native and invasive populations of Chinese tallow tree, Triadica sebifera. We experimentally manipulated herbivory from an exotic generalist scale insect, a native generalist caterpillar, both herbivores, or neither and then examined EFN production by Triadica. Damage from leaf-chewing caterpillars resulted in strongly induced EFN in both native and invasive populations while damage from phloem-feeding scales did not. Extrafloral nectar production and dissolved solute content peaked 4 days after caterpillar herbivory for both native and invasive populations. Number and proportion of leaves producing EFN, EFN volume and concentration of dissolved solutes were similar among native and invasive populations. These results suggest that selection for indirect defenses may be different than selection for other defenses in the introduced ranges of invasive plants, as constitutive and induced EFN production is retained in invasive populations.  相似文献   

11.
The Enemy Release hypothesis holds that exotic plants may have an advantage over native plants because their specialized natural enemies are absent. We tested this hypothesis by measuring leaf damage and plant abundance for naturally-occurring plants in prairies, and by removing natural enemies in an enemy exclusion experiment. We classified plants as invasive exotic, noninvasive exotic, or native, to determine if their degree of invasiveness influenced their relationships with natural enemies. Our field surveys showed that invasive exotic plants generally had significantly lower levels of foliar damage than native species while there was no consistent pattern for noninvasive exotics compared to natives. The relationship between damage and abundance was different for exotic and native plants: foliar damage decreased with increasing abundance for exotic plants while the trend was positive for native plants. While these results from the field surveys supported the Enemy Release Hypothesis, the enemy exclusion experiment did not. There was no relationship between a species?? status as exotic or native and its degree of release from herbivory. Pastinaca sativa, the invasive exotic in this experiment, experienced gains in leaf area and vegetative biomass when treated with pesticides, indicating substantial herbivore pressure in the introduced range. These results show that foliar damage may not accurately predict the amount of herbivore pressure that plants actually experience, and that the Enemy Release hypothesis is not sufficient to explain the invasiveness of P. sativa in prairies.  相似文献   

12.
Trade-offs associated with maintaining herbivory resistance and herbivory tolerance are frequently inferred in plant life histories. Invasive success for many non-native plants is often attributed to novel resistance that repels native herbivores. Chinese tallow tree (Sapium sebiferum, Euphorbiaceae) is a non-native invader that threatens ecosystems throughout the southeastern United States, including imperiled coastal prairie regions. Low herbivore loads due to costly resistance are generally assumed to give Sapium a competitive advantage over native plants. We predicted that if Sapium experienced higher damage levels it would show significant decreases in growth and reduced ability to compete with native prairie vegetation. We conducted full-factorial, paired greenhouse and field experiments designed to assess the effects of simulated leaf herbivory on Sapium growth in the presence of prairie vegetation at different levels of light and nitrogen. Contrary to our expectations, neither low-intensity, chronic defoliation nor high-intensity, acute defoliation negatively affected Sapium seedlings in any resource combination in either experiment. These studies reveal that Sapium possesses considerable phenotypic plasticity, and herbivory tolerance is a newly appreciated trait that likely contributes to its invasive potential.  相似文献   

13.
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.  相似文献   

14.
Tolerance and resistance are defence strategies evolved by plants to cope with damage due to herbivores. The introduction of exotic species to a new biogeographical range may alter the plant–herbivore interactions and induce selection pressures for new plant defence strategies with a modified resource allocation. To detect evolution in tolerance to herbivory in common ragweed, we compared 3 native (North America) and 3 introduced (France) populations, grown in a common garden environment. We explored the effect of leaf herbivory on plant vegetative and reproductive traits. Plants were defoliated by hand, simulating different degrees of insect grazing by removing 0%, 50% or 90% of each leaf blade. Total and shoot dry biomasses were not affected by increasing defoliation, whereas root dry biomass and root:shoot ratio decreased significantly for native and introduced populations. Furthermore, defoliation treatments did not affect any of the plant reproductive traits measured. Hence, common ragweed displayed an efficient reallocation of resources in shoot biomass at the expense of roots following defoliation, which allows the species to tolerate herbivory without obvious costs for fitness. We did not detect any difference in herbivory tolerance between introduced and native populations, but significant differences were found in reproduction with invasive populations producing more seeds than native populations. As a result, tolerance to herbivory has been maintained in the introduced plant populations. We discuss some implications of these preliminary results for biological control strategies dedicated to common ragweed.  相似文献   

15.
The European herb garlic mustard (Alliaria petiolata) is a serious invader of North American deciduous forests. One explanation for its success could be that in the absence of specialized herbivores, selection has favored less defended but more vigorous genotypes. This idea was addressed by comparing offspring from several native and introduced Alliaria populations with respect to their palatability to insect herbivores and their tolerance to simulated herbivory. Feeding rates of a specialist weevil from the native range were significantly greater on American plants, suggesting a loss of resistance in the introduced range. In contrast, there was significant population variation but no continent effect in the feeding rates of a generalist caterpillar. After simulated herbivory, A. petiolata showed a substantial regrowth capacity that involved changes in plant growth, architecture, and allocation. Removal of 75% leaf area or of all bolting stems reduced plant fitness to 81% and 58%, respectively, of the fitness of controls. There was no indication of a difference in tolerance between native and introduced Alliaria populations or of a trade-off between tolerance and resistance.  相似文献   

16.

Background and Aims

Invasive plants can be released from specialist herbivores and encounter novel generalists in their introduced ranges, leading to variation in defence among native and invasive populations. However, few studies have examined how constitutive and induced indirect defences change during plant invasion, especially during the juvenile stage.

Methods

Constitutive extrafloral nectar (EFN) production of native and invasive populations of juvenile tallow tree (Triadica sebifera) were compared, and leaf clipping, and damage by a native specialist (Noctuid) and two native generalist caterpillars (Noctuid and Limacodid) were used to examine inducible EFN production.

Key results

Plants from introduced populations had more leaves producing constitutive EFN than did native populations, but the content of soluble solids of EFN did not differ. Herbivores induced EFN production more than simulated herbivory. The specialist (Noctuid) induced more EFN than either generalist for native populations. The content of soluble solids in EFN was higher (2·1 times), with the specialist vs. the generalists causing the stronger response for native populations, but the specialist response was always comparable with the generalist responses for invasive populations.

Conclusions

These results suggest that constitutive and induced indirect defences are retained in juvenile plants of invasive populations even during plant establishment, perhaps due to generalist herbivory in the introduced range. However, responses specific to a specialist herbivore may be reduced in the introduced range where specialists are absent. This decreased defence may benefit specialist insects that are introduced for classical biological control of invasive plants.  相似文献   

17.
Intact tropical forests are generally considered to be resistant to invasions by exotic species, although the shrub Clidemia hirta (Melastomataceae) is highly invasive in tropical forests outside its native range. Release from natural enemies (e.g., herbivores and pathogens) contributes to C. hirta invasion success where native melastomes are absent, and here we examine the role of enemies when C. hirta co-occurs with native Melastomataceae species and associated herbivores and pathogens. We study 21 forest sites within agricultural landscapes in Sabah, Malaysian Borneo, recording herbivory rates in C. hirta and related native Melastoma spp. plants along two 100-m transects per site that varied in canopy cover. Overall, we found evidence of enemy release; C. hirta had significantly lower herbivory (median occurrence of herbivory per plant = 79% of leaves per plant; median intensity of herbivory per leaf = 6% of leaf area) than native melastomes (93% and 20%, respectively). Herbivory on C. hirta increased when closer to native Melastoma plants with high herbivory damage, and in more shaded locations, and was associated with fewer reproductive organs on C. hirta. This suggests host-sharing by specialist Melastomataceae herbivores is occurring and may explain why invasion success of C. hirta is lower on Borneo than at locations without related native species present. Thus, natural enemy populations may provide a “biological control service” to suppress invasions of exotic species (i.e., biotic resistance). However, lower herbivory pressures in more open canopy locations may make highly degraded forests within these landscapes more susceptible to invasion.  相似文献   

18.
The novel associations between invasive plants and their natural enemies in the introduced range have recently received increasing attention; however, the effects of novel enemies on exotic plant performance and competition with native species remain poorly explored. Here, we tested the impact of herbivory by a native beetle, Cassida piperata, on the performance of the exotic species Alternanthera philoxeroides and competition with a native congener, Alternanthera sessilis, using common garden experiments in central China. We found A. philoxeroides was able to fully compensate for intense herbivory by C. piperata. Herbivory by C. piperata that released at the average density in this region had no impact on competition between the native and exotic plant species. Our results indicate that herbivory by novel enemies may not reduce exotic plant performance due to plant compensation. However, high tolerance to herbivory may not confer a competitive advantage for exotic species compared to less tolerant native competitors if the herbivore damage is below a certain threshold. Thus, it is necessary to assess the impact of novel enemies on exotic plant performance and competition with native plants along gradients of insect densities. This may lead to a better understanding of how best to exploit the role of native herbivores in facilitating or slowing plant invasions.  相似文献   

19.
Both the length of the growing season and the intensity of herbivory often vary along climatic gradients, which may result in divergent selection on plant phenology, and on resistance and tolerance to herbivory. In Sweden, the length of the growing season and the number of insect herbivore species feeding on the perennial herb Lythrum salicaria decrease from south to north. Previous common‐garden experiments have shown that northern L. salicaria populations develop aboveground shoots earlier in the summer and finish growth before southern populations do. We tested the hypotheses that resistance and tolerance to damage vary with latitude in L. salicaria and are positively related to the intensity of herbivory in natural populations. We quantified resistance and tolerance of populations sampled along a latitudinal gradient by scoring damage from natural herbivores and fitness in a common‐garden experiment in the field and by documenting oviposition and feeding preference by specialist leaf beetles in a glasshouse experiment. Plant resistance decreased with latitude of origin, whereas plant tolerance increased. Oviposition and feeding preference in the glasshouse and leaf damage in the common‐garden experiment were negatively related to damage in the source populations. The latitudinal variation in resistance was thus consistent with reduced selection from herbivores towards the northern range margin of L. salicaria. Variation in tolerance may be related to differences in the timing of damage in relation to the seasonal pattern of plant growth, as northern genotypes have developed further than southern have when herbivores emerge in early summer.  相似文献   

20.
Abstract Plant invasions create novel plant–insect interactions. The EICA (evolution of increased competitive ability) hypothesis proposes that invasive plants will reallocate resources from defense to growth and/or reproduction because they have escaped from their co‐evolved insect natural enemies. Testing multiple herbivory by monophagous and oligophagous herbivores and simultaneous measurement of various plant traits will provide new insights into the evolutionary change of invasive plants. In this context, we conducted a common garden experiment to compare plant growth and reproduction, chemical and physical defense, and plant responses to herbivory by different types of herbivores between invasive North American populations and native East Asian populations of mile‐a‐minute weed, Persicaria perfoliata. We found that invasive mile‐a‐minute exhibited lower biomass, flowered earlier and had greater reproductive output than plants from the native range. Compared with native populations, plants from invasive populations had lower tannin content, but exhibited higher prickle density on nodes and leaves. Thus our results partially support the EICA hypothesis. When exposed to the monophagous insect, Rhinoncomimus latipes and the oligophagous insects, Gallerucida grisescens and Smaragdina nigrifrons, more damage by herbivory was found on invasive plants than on natives. R. latipes, G. grisescens and S. nigrifrons had strong, moderate and weak impacts on the growth and reproduction of mile‐a‐minute, respectively. The results indicate that mile‐a‐minute may have evolved a higher reproductive capacity in the introduced range, and this along with a lack of oligophagous and monophagous herbivores in the new range may have contributed to its invasiveness in North America.  相似文献   

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