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1.
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   

2.
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   

3.
Biological invasions change native plant communities, but theory predicting whether introductions create naturalized or invasive species is lacking. Focusing on either plant traits or interactions of introduced plants with native biota creates unreliable results, and improvements may require integration of trait- and interaction-based approaches. To assess the importance of plant traits and herbivory on invasiveness, we incorporated herbivore effects in comparisons of growth and phenology of invasive Phragmites australis and its native congener P. australis subsp. americanus. Our results were influenced by venue (field or common garden), with extended life span and optimized leaf-age structure of introduced P. australis indicating greater potential for resource capture. Attack by introduced gallflies affected expression of plant traits, but we found no consistent effect of aphid attack. Origin did not affect leaf emergence or stem height, but preferential gallfly attack stunted native P. australis and delayed senescence. Greater resource capture and lower attack by nonnative herbivores could give introduced P. australis an advantage over the native subspecies. Our results demonstrating the importance of plant traits as well as their modification by interactions with natural enemies questions whether the outcome of plant introductions can be predicted.  相似文献   

4.
Studies of herbivory and its consequences on the growth of native and exotic plants could help elucidate some processes involved in plant invasions. Introduced species are likely to experience reduced herbivory in their new range due to the absence of specialist enemies and, thus, may have higher benefits if they reduce the investment in resistance and increase their compensatory capacity. In order to evaluate the role of herbivory in disturbed areas within the Patagonian steppe, we quantified and compared the leaf levels of herbivory of four native and five exotic species and recorded the associated insect fauna. We also performed greenhouse experiments in which we simulated herbivory in order to evaluate the compensatory capacity of native and exotic species under different herbivory levels that resembled naturally occurring damage. Natural herbivory levels in the field were similar between the studied exotic and native plants. Field observations confirmed that they both shared some herbivore insects, most of which are generalists. In the greenhouse experiments, both exotic and native plants fully compensated for herbivory. Our results suggest that the studied exotic plants are not released from herbivory in the Patagonian steppe but are able to fully compensate for it. The capacity to recover from herbivory coupled with other potential adaptations, such as a better performance under disturbance and greater competitive ability than that of the native species, may represent some of the mechanisms responsible for the success of plant invasion in the Patagonian steppe.  相似文献   

5.
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   

6.
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   

7.
Ecological release from herbivory due to chemical novelty is commonly predicted to facilitate biological invasions by plants, but has not been tested on a community scale. We used metabolomics based on mass spectrometry molecular networks to assess the novelty of foliar secondary chemistry of 15 invasive plant species compared to 46 native species at a site in eastern North America. Locally, invasive species were more chemically distinctive than natives. Among the 15 invasive species, the more chemically distinct were less preferred by insect herbivores and less browsed by deer. Finally, an assessment of invasion frequency in 2,505 forest plots in the Atlantic coastal plain revealed that, regionally, invasive species that were less preferred by insect herbivores, less browsed by white‐tailed deer, and chemically distinct relative to the native plant community occurred more frequently in survey plots. Our results suggest that chemically mediated release from herbivores contributes to many successful invasions.  相似文献   

8.
The popularly cited enemy release hypothesis, which states that non-native species are released from population control by their enemies, has not been adequately tested in plants. Many empirical studies have compared damage to native versus non-native invaders only in the invaded range, which can lead to erroneous conclusions regarding enemy release. Biogeographical studies that have compared natural enemies in native and introduced ranges have typically focused on a small area of the plants’ distributions in each range, only one plant species, and/or only one guild of natural enemies. To test enemy release, we first surveyed both pathogens and herbivores in multiple populations in both the native and naturalized ranges of three commonly co-occurring perennial bunchgrasses introduced to the United States from Europe. We then compared our field results to the number of fungal pathogens that have been documented on each species from published host-pathogen data compilations. Consistent with enemy release, our field survey showed less herbivory and denser populations in the naturalized range, but there was no evidence of release from pathogens. In contrast, the published host-pathogen data compilations produced evidence of enemy release from pathogens. The difference in results produced by the two approaches highlights the need for multiple approaches to testing mechanisms of invasions by introduced species, which can enable well supported theory to inform sound management practices.  相似文献   

9.
Invasive species may be released from consumption by their native herbivores in novel habitats and thereby experience higher fitness relative to native species. However, few studies have examined release from herbivory as a mechanism of invasion in oceanic island systems, which have experienced particularly high loss of native species due to the invasion of non-native animal and plant species. We surveyed putative defensive traits and leaf damage rates in 19 pairs of taxonomically related invasive and native species in Hawaii, representing a broad taxonomic diversity. Leaf damage by insects and pathogens was monitored in both wet and dry seasons. We found that native species had higher leaf damage rates than invasive species, but only during the dry season. However, damage rates across native and invasive species averaged only 2% of leaf area. Native species generally displayed high levels of structural defense (leaf toughness and leaf thickness, but not leaf trichome density) while native and invasive species displayed similar levels of chemical defenses (total phenolics). A defense index, which integrated all putative defense traits, was significantly higher for native species, suggesting that native species may allocate fewer resources to growth and reproduction than do invasive species. Thus, our data support the idea that invasive species allocate fewer resources to defense traits, allowing them to outperform native species through increased growth and reproduction. While strong impacts of herbivores on invasion are not supported by the low damage rates we observed on mature plants, population-level studies that monitor how herbivores influence recruitment, mortality, and competitive outcomes are needed to accurately address how herbivores influence invasion in Hawaii.  相似文献   

10.
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   

11.
The invasion success of exotic plants is often attributed to escape from natural enemies in their introduced ranges and subsequent evolutionary change in resource allocation from defense to growth and reproduction. We tested this idea by comparing resistance, tolerance, and growth between native (China) and invasive (US) populations of kudzu (Peuraria montana var. lobata) exposed to natural herbivores in the native range. The percentage of foliar damage was much higher in invasive populations than in native populations, indicating that plants from invasive populations had lower resistance to herbivory. Regression of total mass on percentage of foliar damage showed no significant differences in tolerance to herbivory between native and invasive populations. However, stem diameter and mass were significantly greater in invasive populations than in native populations. Our results may suggest geographic variation in herbivory damage and plant growth among kudzu native and invasive populations, but the role of herbivores influencing kudzu invasion requires further investigation.  相似文献   

12.
Herbivory has long been recognized as a significant driver of plant population dynamics, yet its effects along environmental gradients are unclear. Understanding how weather modulates plant–insect interactions can be particularly important for predicting the consequences of exotic insect invasions, and an explicit consideration of weather may help explain why the impact can vary greatly across space and time. We surveyed two native prickly pear cactus species (genus Opuntia) in the Florida panhandle, USA, and their specialist insect herbivores (the invasive South American cactus moth, Cactoblastis cactorum, and three native insect species) for five years across six sites. We used generalized linear mixed models to assess the impact of herbivory and weather on plant relative growth rate (RGR) and sexual reproduction, and we used Fisher's exact test to estimate the impact of herbivory on survival. Weather variables (precipitation and temperature) were consistently significant predictors of vital rate variation for both cactus species, in contrast to the limited and varied impacts of insect herbivory. Weather only significantly influenced the impact of herbivory on Opuntia humifusa fruit production. The relationships of RGR and fruit production with precipitation suggest that precipitation serves as a cue in determining the trade‐off in the allocation of resources to growth or fruit production. The presence of the native bug explained vital rate variation for both cactus species, whereas the invasive moth explained variation only for Ostricta. Despite the inconsistent effect of herbivory across vital rates and cactus species, almost half of Ostricta plants declined in size, and the invasive insect negatively affected RGR and fruit production. Given that fruit production was strongly size‐dependent, this suggests that Ostricta populations at the locations surveyed are transitioning to a size distribution of predominantly smaller sizes and with reduced sexual reproduction potential.  相似文献   

13.
Meyer G  Clare R  Weber E 《Oecologia》2005,144(2):299-307
The mechanisms that allow introduced plants to become invasive are poorly understood. Here, we present a test of the evolution of increased competitive ability hypothesis, which holds that because specialized natural enemies may be absent from the introduced range, exotic plants may evolve to invest less in anti-herbivore defenses and thereby gain a competitive advantage over native plants. We grew Solidago gigantea plants derived from both the native range (North America) and the invasive range (Europe) in a common garden in the native range for 2 years. Half the plants were treated with insecticide to protect them from insect herbivores and the other half were exposed to insects that colonized the garden from nearby fields. Insect herbivore biomass was significantly higher on European plants than US plants in the first year but not the second. European plants were more heavily attacked by pathogens in both years of the study. When exposed to insect herbivores, US plants produced more seed than European plants, but when plants were protected from herbivores, seed production was equivalent between US plants and European plants. The presence of insect herbivores suppressed seed production of European plants much more than that of US plants, even though the level of herbivory experienced by European and US plants was similar in the second year, suggesting that the ability to tolerate herbivory was diminished in European plants. These results partially support the EICA hypothesis: plants from the introduced range were more susceptible to some natural enemies and benefited more from insect removal than plants from the native range. The prediction that European plants would perform better than US plants in the absence of insect herbivores was not supported. Electronic Supplementary Material Supplementary material is available for this article at  相似文献   

14.
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   

15.
Interactions among herbivores or between herbivores and other plant natural enemies, such as fungal pathogens, range from competition to facilitation. Moreover, the outcome of these interactions depends on the ecological context where they occur. In this study we examined the effects of clipping, as a surrogate of herbivory by ungulates, on the damage caused by two types of natural enemies (herbivorous insects and foliar fungal pathogens) on bilberry, Vaccinium myrtillus, in combination with nitrogen (N) fertilization representing current N atmospheric deposition. To examine whether the responses of both these natural enemies were mediated by changes in the plant, we estimated the effects of the treatments on bilberry growth and branching and on chlorophyll content as proxy of N content in leaves. Clipping increased the proportion of leaves damaged by herbivorous insects regardless of whether it was combined with N fertilization or not in 2008. In 2007 and 2009 repeated damage to the shrub also facilitated insect herbivory but only under N applications. Regarding fungal infestation incidence, clipping decreased the proportion of infected leaves in all the years considered but only in fertilized plots. Our results suggest that vertebrate herbivores facilitate insect herbivory and reduce fungal infestation but that these effects are dependent on nutritional conditions. Moreover, we found a negative residual correlation between insect herbivory and fungal infestation on bilberry leaves. Therefore, interactions between insect herbivores and fungal pathogens could be implicated in the final outcome of interactions between browsing ungulates and both bilberry natural enemies.  相似文献   

16.
We explored consequences of spatial and temporal heterogeneity in herbivory on the survival, growth, and reproduction of the Californian native dune thistle, Cirsium occidentale, in coastal and inland sites, for 2 years. We assessed the relative impacts of insect and mammalian herbivores and compared the relative importance of herbivory in coastal and inland habitats and among locations with different microclimates across a coastal dune. Effects of insect and mammalian herbivores were tested with a combination of insecticidal spray and cage exclusion treatments in a factorial experiment at the coastal site. Mammalian herbivores strongly affected the population dynamics of C. occidentale in both years, and their effects were augmented by fungal infection (1991), herbivory by stem-borers (1990) and, to a lesser extent, by insect seed predators in both years. Mammals caused most plant deaths, but the mammal species responsible differed among sites. Rabbit herbivory altered the vegetative growth of coastal thistles and significantly modified other key aspects of Cirsium demography, including growth rate and timing of reproduction. Small, uncaged plants grazed by rabbits took at least 1 year longer to mature than did caged plants. Larvae of Pyrausta subsequalis were the only insects that killed established plants. In 1990 and 1991, the numbers of insects damaging seed heads before dispersal were low, but were sufficient to cause receptacle and seed damage. The number of mature, undamaged seeds (and percent successful seed production) was reduced significantly only for heads infested by fungi near the ocean in 1991: the fungus occurred in 37% of heads and caused a 77% reduction in mature seeds. Received: 21 October 1996 / Accepted: 27 March 1997  相似文献   

17.
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   

18.
Plants are frequently attacked by both above- and belowground arthropod herbivores. Nevertheless, studies rarely consider root and shoot herbivory in conjunction. Here we provide evidence that the root-feeding insect Agriotes lineatus reduces the performance of the foliage feeding insect Spodoptera exigua on cotton plants. In a bioassay, S. exigua larvae were allowed to feed on either undamaged plants, or on plants that had previously been exposed to root herbivory, foliar herbivory, or a combination of both. Previous root herbivory reduced the relative growth rates as well as the food consumption of S. exigua by more than 50% in comparison to larvae feeding on the undamaged controls. We found no effects in the opposite direction, as aboveground herbivory by S. exigua did not affect the relative growth rates of root-feeding A. lineatus . Remarkably, neither did the treatment with foliar herbivory affect the food consumption and relative growth rate of S. exigua in the bioassay. However, this treatment did result in a significant change in the distribution of S. exigua feeding. Plants that had been pre-exposed to foliar herbivory suffered significantly less damage on their young terminal leaves. While plant growth and foliar nitrogen levels were not affected by any of the treatments, we did find significant differences between treatments with respect to the level and distribution of plant defensive chemicals (terpenoids). Exposure to root herbivores resulted in an increase in terpenoid levels in both roots as well as in mature and immature foliage. Foliar damage, on the other hand, resulted in high terpenoid levels in young, terminal leaves only. Our results show that root-feeding herbivores may change the level and distribution of plant defenses aboveground. Our data suggest that the reported interactions between below- and aboveground insect herbivores are mediated by induced changes in plant secondary chemistry.  相似文献   

19.
Exotic plant invasion can have dramatic impacts on native plants making restoration of native vegetation at invaded sites challenging. Though invasives may be superior competitors, it is possible their dominance could be enhanced by insect herbivores if native plants are preferred food sources. Insect herbivory can regulate plant populations, but little is known of its effects in restoration settings. There is a need to better understand relationships between insect herbivores and invasive plants with regard to their combined potential for impacting native plant establishment and restoration success. The objective of this study was to assess impacts of grasshopper herbivory and the invasive grass Bromus tectorum (cheatgrass) on mortality and growth of 17 native plant species used in restoration of critical sagebrush steppe ecosystems. Field and greenhouse experiments were conducted using moderate densities of a common, generalist pest grasshopper (Melanoplus bivittatus). Grasshoppers had stronger and more consistent impacts on native restoration plants in field and greenhouse studies than cheatgrass. After 6 weeks in the greenhouse, grasshoppers were associated with 36% mortality over all native restoration species compared to 2% when grasshoppers were absent. Herbivory was also associated with an approximately 50% decrease in native plant biomass. However, effects varied among species. Artemisia tridentata, Chrysothamnus viscidiflorus, and Coreopsis tinctoria were among the most negatively impacted, while Oenothera pallida, Pascopyrum smithii, and Leymus cinerus were unaffected. These findings suggest restoration species could be selected to more effectively establish and persist within cheatgrass infestations, particularly when grasshopper populations are forecasted to be high.  相似文献   

20.
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   

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