Introduced plants of the invasive Solidago gigantea (Asteraceae) are larger and grow denser than conspecifics in the native range

Introduced plant species that became successful invaders appear often more vigorous and taller than their conspecifics in the native range. Reasons postulated to explain this better performance in the introduced range include more favourable environmental conditions and release from natural enemies and pathogens. According to the Evolution of Increased Competitive Ability hypothesis (EICA hypothesis) there is a trade‐off between investment into defence against herbivores and pathogens, and investment into a stronger competitive ability. In this study, we conducted field surveys to investigate whether populations of the invasive perennial Solidago gigantea Ait (Asteraceae) differ with respect to growth and size in the native and introduced range, respectively. We assessed size and morphological variation of 46 populations in the native North American range and 45 populations in the introduced European range. Despite considerable variation between populations within continents, there were pronounced differences between continents. The average population size, density and total plant biomass were larger in European than in American populations. Climatic differences and latitude explained only a small proportion of the total variation between the two continents. The results show that introduced plants can be very distinct in their growth form and size from conspecifics in the native range. The apparently better performance of this invasive species in Europe may be the result of changed selection pressures, as implied by the EICA hypothesis.     

Increased competitive ability and herbivory tolerance in the invasive plant <Emphasis Type="Italic">Sapium sebiferum</Emphasis>

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.     

Different gardens, different results: native and introduced populations exhibit contrasting phenotypes across common gardens

Invasive plants may respond through adaptive evolution and/or phenotypic plasticity to new environmental conditions where they are introduced. Although many studies have focused on evolution of invaders particularly in the context of testing the evolution of increased competitive ability (EICA) hypothesis, few consistent patterns have emerged. Many tests of the EICA hypothesis have been performed in only one environment; such assessments may be misleading if plants that perform one way at a particular site respond differently across sites. Single common garden tests ignore the potential for important contributions of both genetic and environmental factors to affect plant phenotype. Using a widespread invader in North America, Cynoglossum officinale, we established reciprocal common gardens in the native range (Europe) and introduced range (North America) to assess genetically based differences in size, fecundity, flowering phenology and threshold flowering size between native and introduced genotypes as well as the magnitude of plasticity in these traits. In addition, we grew plants at three nutrient levels in a pot experiment in one garden to test for plasticity across a different set of conditions. We did not find significant genetically based differences between native and introduced populations in the traits we measured; in our experiments, introduced populations of C. officinale were larger and more fecund, but only in common garden experiments in the native range. We found substantial population-level plasticity for size, fecundity and date of first flowering, with plants performing better in a garden in Germany than in Montana. Differentiation of native populations in the magnitude of plasticity was much stronger than that of introduced populations, suggesting an important role for founder effects. We did not detect evidence of an evolutionary change in threshold flowering size. Our study demonstrates that detecting genetically based differences in traits may require measuring plant responses to more than one environment.     

Response to enemies in the invasive plant Lythrum salicaria is genetically determined

Background and Aims

The enemy release hypothesis assumes that invasive plants lose their co-evolved natural enemies during introduction into the new range. This study tested, as proposed by the evolution of increased competitive ability (EICA) hypothesis, whether escape from enemies results in a decrease in defence ability in plants from the invaded range. Two straightforward aspects of the EICA are examined: (1) if invasives have lost their enemies and their defence, they should be more negatively affected by their full natural pre-invasion herbivore spectrum than their native conspecifics; and (2) the genetic basis of evolutionary change in response to enemy release in the invasive range has not been taken sufficiently into account.

Methods

Lythrum salicaria (purple loosestrife) from several populations in its native (Europe) and invasive range (North America) was exposed to all above-ground herbivores in replicated natural populations in the native range. The experiment was performed both with plants raised from field-collected seeds as well as with offspring of these where maternal effects were removed.

Key Results

Absolute and relative leaf damage was higher for introduced than for native plants. Despite having smaller height growth rate, invasive plants attained a much larger final size than natives irrespective of damage, indicating large tolerance rather than effective defence. Origin effects on response to herbivory and growth were stronger in second-generation plants, suggesting that invasive potential through enemy release has a genetic basis.

Conclusions

The findings support two predictions of the EICA hypothesis – a genetically determined difference between native and invasive plants in plant vigour and response to enemies – and point to the importance of experiments that control for maternal effects and include the entire spectrum of native range enemies.     

Meta‐analysis reveals evolution in invasive plant species but little support for Evolution of Increased Competitive Ability (EICA)

Ecological explanations for the success and persistence of invasive species vastly outnumber evolutionary hypotheses, yet evolution is a fundamental process in the success of any species. The Evolution of Increased Competitive Ability (EICA) hypothesis (Blossey and Nötzold 1995) proposes that evolutionary change in response to release from coevolved herbivores is responsible for the success of many invasive plant species. Studies that evaluate this hypothesis have used different approaches to test whether invasive populations allocate fewer resources to defense and more to growth and competitive ability than do source populations, with mixed results. We conducted a meta‐analysis of experimental tests of evolutionary change in the context of EICA. In contrast to previous reviews, there was no support across invasive species for EICA's predictions regarding defense or competitive ability, although invasive populations were more productive than conspecific native populations under noncompetitive conditions. We found broad support for genetically based changes in defense and competitive plant traits after introduction into new ranges, but not in the manner suggested by EICA. This review suggests that evolution occurs as a result of plant introduction and population expansion in invasive plant species, and may contribute to the invasiveness and persistence of some introduced species.     

The evolution of chemical defenses along invasion routes: Harmonia axyridis Pallas (Coccinellidae: Coleoptera) as a case study

The evolution of increased competitive ability (EICA) hypothesis (Blossey & Nötzold, 1995 ) postulates that escaping from coevolved enemies increases invaders fitness by energy reallocation from defenses and immunity to growth and reproduction. In this context, we evaluated the evidence of evolutionary change in invasive populations of Harmonia axyridis Pallas (Coccinellidae: Coleoptera). We measured egg defenses—cocktail of hydrocarbons on the egg's surface flagging egg toxicity and the concentration of the main alkaloid harmonine—in individuals from three populations along the invasion route (Japan: native, United States: introduced more than 30 years ago, South Africa: introduced in the early 2000s) in a common garden experiment. Our results support the EICA hypothesis: We found changes along the invasion route in the profiles of the hydrocarbons coating the eggs’ surface and a decrease in the concentration of harmonine in eggs from the most recent invasive South African population compared to the long established in the United States and the native Japanese ones.     

Palatability to a generalist herbivore,defence and growth of invasive and native <Emphasis Type="Italic">Senecio</Emphasis> species: testing the evolution of increased competitive ability hypothesis

This paper tests the prediction that introduced plants may become successful invaders because they experience evolutionary changes in growth and defence in their new range [evolution of increased competitive ability hypothesis (EICA)]. Interspecific and intraspecific binary feeding choices were offered to the snail Helix aspersa. The choices were between: (1) plants of the invasive Senecio inaequidens and Senecio pterophorus derived from populations in the introduced range (Europe) and plants of three indigenous species (Senecio jacobea, Senecio vulgaris and Senecio malacitanus) from populations in Europe; (2) plants of the invasive S. inaequidens and S. pterophorus from populations in the introduced range (Europe) and from populations in the native range (South Africa). We did not find a clear pattern of preference for indigenous or alien species of Senecio. However, we found that European invasive populations of S. inaequidens and S. pterophorus were less palatable than South African native populations. Moreover, in contrast to the predictions of the EICA hypothesis, the invasive genotypes of both species also showed a higher total concentration of pyrrolizidine alkaloids, and in the case of S. inaequidens we also found higher growth than in native genotypes. Our results are discussed with respect to the refinement of the EICA hypothesis that takes into account the difference between specialist and generalist herbivores and between qualitative and quantitative defences. We conclude that invasive populations of S. inaequidens and S. pterophorus are less palatable than native populations, suggesting that genetic differentiation associated with founding may occur and contribute to the plants’ invasion success by selecting the best-defended genotypes in the introduced range.     

Testing Predictions of the ‘Evolution of Increased Competitive Ability’ Hypothesis for an Invasive Crucifer

The successful spread of invasive plants may result from an evolutionary shift in resource allocation from defence to growth due to release from enemies, as proposed by the ‘evolution of increased competitive ability’ hypothesis (EICA). The crucifer Lepidium draba was used to test this hypothesis, measuring growth and levels of glucosinolates and myrosinase of leaves as constitutive defence parameters. Individuals from 21 populations of the native (Europe) and the invasive range (North-America) were grown under common greenhouse conditions. According to the EICA hypothesis it was predicted that plants from the invasive range might show stronger growth and have lower levels of defence as a result of selection favouring such genotypes. There was significant variation between populations in shoot, root, total biomass, and number of ramets of 3-month-old plants but no difference due to origin from both continents. The main glucosinolate p-hydroxybenzyl glucosinolate was significantly higher in seedlings of the invasive range while myrosinase activity was higher in old plants of the invasive range. Therefore, the EICA hypothesis does not hold, however, alternatively there is evidence for selection favouring stronger defence in the invasive range. The binary defence system of this crucifer is discussed with respect to the degree of specialisation of potential herbivores.     

Reduced competitive ability in an invasive plant

One explanation for successful plant invaders is that they evolved to be more competitive. An intuitive prediction of this Evolution of Increased Competitive Ability (EICA) hypothesis never previously tested is that invasive populations should outcompete their native ‘ancestors’ in a common environment. We tested this idea in a diallel competition experiment with Alliaria petiolata where offspring from native and invasive populations were grown alone or in all pairwise combinations. While without competition, there were no differences between native and invasive populations, native populations outperformed invasive ones when competing against each other. Our results contradict the EICA hypothesis and we conclude that it does not not hold for Alliaria petiolata. Instead, we formulate a new ERCA (Evolutionary Reduced Competitive Ability) hypothesis: if there is less competition in the invasive range and competitive ability involves traits that have a fitness cost, then selection might act against it, thereby reducing intraspecific interactions too.     

For or against: the importance of variation in growth rate for testing the EICA hypothesis

The evolution of increased competitive ability (EICA) hypothesis proposes that invasive species evolve decreased defense and increased competitive ability following natural enemy release. Previous studies have found evidence both for and against EICA. The resource-enemy release hypothesis (R-ERH) suggests that fast-growing species may experience stronger enemy release than slow-growing species. On the basis of R-ERH, the prediction of EICA will be held true for slow-growing genotypes, i.e., the slow-growing genotypes from the introduced range will be less resistant to herbivory and grow faster than those from the home range; while the EICA will not be held for fast-growing genotypes, i.e., there will be no significant differences in growth and defense traits between the introduced and native fast-growing genotypes. We tested these predictions preliminarily using five populations of the invasive plant Alternanthera philoxeroides. This species has two varieties in its home range, which showed a distinct growth-defense strategy: the northern A. p. var. acutifolia (Apa) had higher growth rate but lower resistance, while the southern A. p. var. obtusifolia (Apo) had lower growth rate but higher resistance level. Our results suggest that the EICA hypothesis is consistent with the slow-growing Apo, but not with the fast-growing Apa. We suggest that evolutionary changes in growth or resistance following enemy release are influenced by variation in growth rate within an invasive alien plant. These findings have important implications for the EICA hypothesis, and may partially explain why previous studies have found evidence both for and against EICA.     

Increased reproductive capacity and physical defense but decreased tannin content in an invasive plant

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.     

Invasive Mahonia plants outgrow their native relatives

Invasive populations often grow more vigorously than conspecific populations in the native range. This has frequently been attributed to evolutionary changes resulting either from founder effects, or from natural selection owing to enemy release. Another mechanism contributing to evolutionary change has largely been neglected in the past: Many invasive plant species do actually descend from cultivated plants and were therefore subject to breeding, including hybridization and artificial selection. In a common garden experiment, we compared invasive Central European populations of the ornamental shrub, Mahonia, with native populations of its putative parental species, Mahonia aquifolium and M. repens, from North America. We hypothesized that plants of invasive populations show increased growth and retained high levels of heritable variation in phenotypic traits. Indeed, invasive Mahonia plants grew larger in terms of stem length, number of leaves and above-ground biomass than either of the two native species, which did not differ significantly from each other. Since there are no hints on release of invasive Mahonia populations from natural enemies, it is likely that hybridization and subsequent selection by breeders have lead to an evolutionary increase of plant vigour in the introduced range. Further on, heritable variation was not consistently reduced in invasive populations compared with populations of the two native species. We suggest that interspecific hybridization among the Mahonia species has counteracted the harmful effects of genetic bottlenecks often associated with species introductions. Based on this case study, we conclude that, more attention has to be paid on the role of plant breeding when assessing the mechanisms behind successful plant invasions in future.     

Changes in Defense of an Alien Plant Ambrosia artemisiifolia before and after the Invasion of a Native Specialist Enemy Ophraella communa

The evolution of increased competitive ability hypothesis (EICA) predicts that when alien plants are free from their natural enemies they evolve lower allocation to defense in order to achieve a higher growth rate. If this hypothesis is true, the converse implication would be that the defense against herbivory could be restored if a natural enemy also becomes present in the introduced range. We tested this scenario in the case of Ambrosia artemisiifolia (common ragweed) – a species that invaded Japan from North America. We collected seeds from five North American populations, three populations in enemy free areas of Japan and four populations in Japan where the specialist herbivore Ophraella communa naturalized recently. Using plants grown in a common garden in Japan, we compared performance of O. communa with a bioassay experiment. Consistent with the EICA hypothesis, invasive Japanese populations of A. artemisiifolia exhibited a weakened defense against the specialist herbivores and higher growth rate than native populations. Conversely, in locations where the herbivore O. communa appeared during the past decade, populations of A. artemisiifolia exhibited stronger defensive capabilities. These results strengthen the case for EICA and suggest that defense levels of alien populations can be recuperated rapidly after the native specialist becomes present in the introduced range. Our study implies that the plant defense is evolutionary labile depending on plant-herbivore interactions.     

No evidence for an ‘evolution of increased competitive ability’ for the invasive Lepidium draba

The ‘evolution of increased competitive ability’ (EICA) hypothesis states that reduced herbivory in the introduced range causes an evolutionary shift in resource allocation from herbivore defense to growth. Therefore, according to EICA, introduced genotypes are expected to grow more vigorously than conspecific native genotypes when cultivated under common standardized conditions. The EICA hypothesis also assumes that herbivores will perform better on introduced genotypes compared to native genotypes, because they are less well defended. However, selection for either defense or growth will depend on the type of defense (quantitative or qualitative) employed by the plant, and whether the plant is released from generalist or specialist herbivores. The predictions of the EICA hypothesis might be reversed if a plant experiences increased generalist herbivore pressure in the introduced range, and therefore invests more in qualitative defense. We examined this idea with the invasive perennial mustard, Lepidium draba. We grew a total of 16 populations of L. draba from both its native European and introduced western US ranges under common conditions in a greenhouse. We also tested for differences in plant resistance to the specialist herbivore, Psylliodes wrasei, by conducting a leaf disc feeding bioassay with native and introduced L. draba genotypes. Furthermore, we quantified the generalist herbivore load on L. draba in both ranges in order to assess the selection pressure for increased qualitative defense. Contrary to the original EICA prediction, all plant traits (biomass, number of shoots, length and diameter of longest leaf) tended to be greater for the native, rather than introduced L. draba genotypes. There was no significant difference in the proportion of leaf area consumed by the specialist herbivore between native and introduced genotypes. The generalist herbivore load on L. draba was significantly greater in the introduced range. Our data suggest that the EICA hypothesis does not explain the invasion success of L. draba in the US. Instead, we propose that the reduced vigor of introduced genotypes may be due to selection for increased defense against generalist herbivores in the introduced range.     

Invasive aliens and sampling bias

Two hypotheses have been proposed to explain the observation of increased vigour of invasive alien plants in their nonindigenous ranges: phenotypic plasticity, and the post‐invasion evolution of increased competitive ability (EICA). Here I specify how a general pattern of increased vigour may result from sampling bias. Ignoring failed invasions can account for the illusion of increased vigour over a broad range of assumptions. Plasticity and EICA need not be viewed as explanations for a general pattern of increased vigour even if they are the mechanisms underlying every occurrence.     

Selection on herbivory resistance and growth rate in an invasive plant

The evolution of increased competitive ability (EICA) hypothesis proposes that invasive species evolve decreased defense and increased competitive ability following natural enemy release. Previous tests of EICA examined the result of evolution by comparing individuals from home and introduced ranges, but no previous study of this hypothesis has examined the process of evolution by analyzing patterns of selection. On the basis of EICA, there should be selection for competitive ability without herbivores and selection for defense with herbivores. Selection on competitive ability should be stronger for genotypes accustomed to herbivores (home range genotypes), and selection on defense should be stronger for genotypes unaccustomed to herbivores (introduced range genotypes). Using a field experiment, we tested these hypotheses for the invasive plant Melaleuca quinquenervia. There was a negative genetic correlation between resistance and growth, indicating a trade-off. However, selection for stem elongation (an indicator of competitive ability) was always positive, and selection on resistance was always negative and did not depend on genotype source or the presence of herbivores. The patterns of selection found in this study contrast with predictions from EICA and accurately predict the lack of evolutionary change in growth and resistance following the introduction of this species from Australia to Florida.     

飞机草入侵种群与原产地种群生长性状的差异

飞机草(Chromolaena odorata)是我国热带地区危害严重的外来入侵植物,为揭示适应进化对其成功入侵的贡献,在同质种植园中,比较研究了飞机草10个入侵地种群与12个原产地种群生长性状的差异,为排除奠基者效应的可能影响,进一步比较了飞机草10个入侵地种群与其原产地可能的祖先种群间的差异。结果表明,飞机草10个入侵地种群的基茎、株高、分枝数、生物量和比叶面积均显著高于12个原产地种群;与可能的祖先种群相比,飞机草10个入侵种群的生物量、分枝数和比叶面积仍更高。这些结果表明,在长期的入侵过程中飞机草通过进化提高了资源向生长的分配,支持增强竞争能力的进化假说。     

Is the increased vigour of invasive weeds explained by a trade-off between growth and herbivore resistance?

Blossey and Nötzold (1995) recently hypothesised that the increased vigour of certain invasive plant species has been at the expense of defences against natural enemies. A prediction of their evolution of increased competitive ability (EICA) hypothesis is that invasive genotypes are relatively poorly defended. We tested this prediction with herbivore bioassays and with direct quantification of plant secondary metabolites comparing non-indigenous genotypes of Lythrum salicaria L. (purple loosestrife) with indigenous forms. The herbivore bioassays revealed no significant intra-specific variation in herbivore resistance between indigenous and non-indigenous hosts. The phenolic content of L. salicaria leaves was significantly higher in indigenous genotypes, as predicted by the EICA hypothesis. The average phenolic content of leaves (regardless of their origin) was, however, low, implying that the role of plant phenolics in purple loosestrife anti-herbivore defence is probably limited. It is suggested that the EICA hypothesis, as tested in the current study, does not explain the increased vigour of L. salicaria in non-indigenous habitats.     

Evolutionary responses of natives to introduced species: what do introductions tell us about natural communities?

Biological invasions dramatically affect the distribution, abundance and reproduction of many native species. Because of these ecological effects, exotic species can also influence the evolution of natives exposed to novel interactions with invaders. Evolutionary changes in natives in response to selection from exotics are usually overlooked, yet common responses include altered anti-predator defenses, changes in the spectrum of resources and habitats used, and other adaptations that allow native populations to persist in invaded areas. Whether a native population is capable of responding evolutionarily to selection from invaders will depend on the demographic impact of the invader, the genetic architecture and genetic variability of the native population and potentially the history of previous invasions. In some cases, natives will fail to evolve or otherwise adapt, and local or global extinction will result. In other cases, adaptive change in natives may diminish impacts of invaders and potentially promote coexistence between invaders and natives. Here, we review the evidence for evolutionary responses of native species to novel community members. We also discuss how the effects of introduced species may differ from those caused by natural range expansions of native species. Notably, introduced species may come from remote biotas with no previous evolutionary history with the native community. In addition, the rate of addition of introduced species into communities is much greater than all but the most extreme cases of historical biotic exchange. Understanding the evolutionary component of exotic/native species interactions is critical to recognizing the long-term impacts of biological invasions, and to understanding the role of evolutionary processes in the assembly and dynamics of natural communities.     

Effects of latitudinal variation on field and common garden comparisons between native and introduced groundsel (Senecio vulgaris) populations

纬度变异对欧洲千里光原产地与引入地种群野外和同质生物园比较的影响 野外和同质生物园比较经常用于研究入侵植物快速进化。当原产地或引入地的种群地理分布范围较广时，表型性状的纬度渐变可能会影响这种比较分析。我们检验了纬度渐变是否会影响欧洲千里光(Senecio vulgaris)引入地和原产地种群间的比较。欧洲千里光原产于欧洲，现已引入到中国东北和西南地区。我们对欧洲原产地和中国引入地的野外种群进行了调查，同时在瑞士开展了同质生物园实验，比较了中国引入地种群与欧洲原产地种群的株高、分枝数和花序数，以此衡量两地欧洲千里光种群在生长和繁殖方面的差异。野外调查表明，中国引入地种群的株高和花序数量显著大于欧洲原产地种群，这一结果与增强竞争力进化假说的预测一致。在同质生物园比较中，当考虑纬度渐变的作用时，中国种群的花序数量显著大于欧洲种群。分别将中国东北部、中国西南部两个地区的种群与欧洲种群进行比较，无论在野外还是同质生物园，与欧洲种群处于相似纬度的中国东北部种群表现出更高的株高和更多的花序。但位于更低纬度的中国西南部种群的繁殖表现并不强于原产地种群。因此我们的结果表明，当引入地种群地理分布较广时，无论野外还是同质生物园的比较都应该考虑表型性状纬度渐变的影响。