Allelopathy and plant invasions: traditional,congeneric, and bio-geographical approaches

A relatively small subset of exotic plant species competitively exclude their neighbors in invaded “recipient” communities but coexist with neighbors in their native habitat. Allelopathy has been argued as one of the mechanisms by which such exotics may become successful invaders. Three approaches have been used to examine allelopathy as a mechanism for invasion. The traditional approach examines exotic invasives in the same way that other native plants also suspected of allelopathic activities are studied. In this approach dose, fate, and replenishment of chemicals can provide powerful evidence for allelopathic processes. The bio-geographical approach often does not provide as much mechanistic evidence for allelopathy, but comparing the allelopathic effects of exotic invasives on species from their native and invaded communities yields stronger evidence than the traditional approach for whether or not allelopathy actually contributes to invasive success. The congeneric, or phylogenetic, approach involves comparative studies of exotic species with natives in the same genus or that are as closely related as possible. Congeneric approaches are limited in inference and have been used to study the role of natural enemies in exotic invasion, but this approach has not been widely used to study allelopathy and invasion. We discuss these three approaches and present a data set for congeneric Lantana and Prosopis to illustrate how the congeneric approach can be used, and use Centaurea maculosa and (±)-catechin to demonstrate experimentally how traditional and bio-geographic approaches can be integrated to shed light on allelopathy in exotic plant invasions.     

Biogeographical variation in community response to root allelochemistry: novel weapons and exotic invasion

Centaurea diffusa is one of the most destructive invasive weeds in the western USA and allelopathy appears to contribute to its invasiveness ( Callaway & Aschehoug 2000 ). Here we identify a chemical from the root exudates of C. diffusa, 8‐hydroxyquinoline, not previously reported as a natural product, and find that it varies biogeographically in its natural concentration and its effect as an allelochemical. 8‐Hydroxyquinoline is at least three times more concentrated in C. diffusa‐invaded North American soils than in this weed's native Eurasian soils and has stronger phytotoxic effects on grass species from North America than on grass species from Eurasia. Furthermore, experimental communities built from North American plant species are far more susceptible to invasion by C. diffusa than communities built from Eurasian species, regardless of the biogeographical origin of the soil biota. Sterilization of North American soils suppressed C. diffusa more than sterilization of Eurasian soils, indicating that North American soil biota may also promote invasion by C. diffusa. Eurasian plants and soil microbes may have evolved natural resistance to 8‐hydroxyquinoline while North American plants have not, suggesting a remarkable potential for evolutionary compatibility and homeostasis among plants within natural communities and a mechanism by which exotic weeds destroy these communities.     

Allelopathy of a native grassland community as a potential mechanism of resistance against invasion by introduced plants

Successful plant invasions depend, at least partly, on interactions between introduced plants and native plant communities. While allelopathic effects of introduced invaders on native resident species have received much attention, the reverse, i.e. allelopathic effects of native residents on introduced plants, have been largely neglected. Therefore, we tested whether allelopathy of native plant communities decreases their invasibility to introduced plant species. In addition, we tested among the introduced species whether the invasive ones are more tolerant to allelopathy of native plant communities than the non-invasive ones. To test these hypotheses, we grew nine pairs of related (congeneric or confamilial) invasive and non-invasive introduced plant species (i.e. 18 species) in the presence or absence of a native grassland community, which consisted of three common forbs and three common grasses, with or without activated carbon in the soil. Activated carbon reduced the survival percentage and growth of introduced plants in the absence of the native plant community. However, its net effect on the introduced plants was neutral or even slightly positive in the presence of the native community. This might suggest that the native plant community imposed allelopathic effects on the introduced plants, and that these effects were neutralized or reduced by activated carbon. The invasive and non-invasive introduced plants, however, did not differ in their tolerance to such allelopathic effects of the native plant community. Thus, although allelopathy of native plant communities might increase their resistance against introduced plants, there was no evidence that tolerance to allelopathy of native plant communities contributes to the degree of invasiveness of introduced plants.     

化感——外来入侵植物的“Novel Weapons”

综述了与外来植物入侵相关的理论假说。并主要介绍了从化感角度提出的“N ove lW eapons”假说(NW假说)。某些外来入侵植物能产生一些化学物质。这些物质可以发挥较强的化感作用,或者成为植物和土壤微生物之间相互作用的调节者。由于不同区域的植物群落共同进化的轨迹不同,被入侵群落的植物对这些化学物质缺乏适应性,因此,这些物质成为了外来种入侵的利器。而对外来种发源地的群落来说,由于长期的共存关系,这些物质对其伴生植物的影响相对较小。NW假说为化感作为外来植物入侵的一种机制奠定了理论基础,为外来入侵植物的防治提供了理论依据,但作为一种较新的理论假说,还需要更多的实验研究加以支持。     

<Emphasis Type="Italic">Acroptilon repens</Emphasis>, an Asian invader,has stronger competitive effects on species from America than species from its native range

The ability to competitively suppress native species is key to successful invasion. Since invasions involve an increase in abundance or dominance of a species in its non-native range, competitive effects might be expected to be stronger in the non-native range of an invader; however, there have been few comparisons of the competitive effects of invasive plants on species from invaded ranges versus species from native ranges. We compared the competitive and allelopathic effects of Acroptilon repens on native North American species to effects on related species from the native range of Acroptilon in Uzbekistan. We also compared the competitive interactions among these North American and Eurasian species, in the absence of Acroptilon, examining the hypothesis that particular regional species pools may show differences in competitive ability. The results showed that Acroptilon had stronger competitive effects against native North American species than against species native to Uzbekistan. There was no difference in the competitive effects among Eurasians and North Americans. The effects of leachates collected from Acroptilon roots were weak but more negative on species from North America than on species from Uzbekistan. Our results suggest that inherently stronger competitive and allelopathic effects of Acroptilon on North American plants than on plants from its native range may contribute to its invasive success.     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

Competitive ability,not tolerance,may explain success of invasive plants over natives

When entering a new community, introduced species leave behind members of their native community while simultaneously forming novel biotic interactions. Escape from enemies during the process of introduction has long been hypothesized to drive the increased performance of invasive species. However, recent studies and quantitative syntheses find that invaders often receive similar, or even more, damage from enemies than do native species. Therefore, invasives may be those more tolerant to enemy damage, or those able to maintain competitive ability in light of enemy damage. Here, we investigate whether tolerance and competitive ability could contribute to invasive plant success. We determined whether invasive plants were more competitive than native or noninvasive exotic species in both the presence and absence of simulated herbivory. We found competition and herbivory additively reduced individual performance, and affected the performance of native, invasive, and noninvasive exotic species’ to the same degree. However, invasives exerted stronger competitive effects on an abundant native species (Elymus canadensis) in both the presence and absence of herbivory. Therefore, while invasive species responded similarly to competition and simulated herbivory, their competitive effects on natives may contribute to their success in their introduced range.     

Phylogenetic isolation increases plant success despite increasing susceptibility to generalist herbivores

Aim Theory suggests that introduced species that are phylogenetically distant from their recipient communities should be more successful than closely related introduced species because they can exploit open niches and escape enemies in their new range, i.e. Darwin’s Naturalization Hypothesis. Alternatively, it has also been hypothesized that closely related invaders might be more successful than novel invaders because they are pre‐adapted to conditions in their new range; a paradox coined Darwin’s Naturalization Conundrum. To date, these hypotheses have been tested primarily at the regional scale, not within local plant communities where introduced species colonize, compete and encounter herbivores. Location Global. Methods and Results We used community phylogenetics to analyse data from 49 published experiments to examine the importance of phylogenetic relatedness and generalist herbivory on native and exotic plant success at the community level. Plants that were categorized as ‘invasive’ were indeed less related to the recipient community than ‘non‐pest’ exotic plants. Distantly related exotic plants were also more abundant than closely related species. Phylogenetic relatedness predicted herbivore impact, but in a way that was opposite to predictions, as herbivores had stronger, not lesser, impacts on distantly related plants. Importantly, these same patterns generally held for native plants, as distantly related native plants were more abundant and more susceptible to herbivores than closely related species, ultimately resulting in herbivores suppressing community‐level phylogenetic diversity. Main conclusions Distantly related plants were more locally successful despite experiencing stronger control by generalist herbivores, a finding that was robust across native and exotic species. To our knowledge, this is the first evidence that phylogenetic matching influences the local success of both native and exotic species and that herbivores can influence community phylodiversity. Phylogenetic relatedness explained a relatively small portion of the variance in the data even after taking herbivory into account, however, suggesting that phylogenetic matching works in combination with other factors to influence community assembly.     

Testing for allelopathy in invasive plants: it all depends on the substrate!

Invasive plants can affect native plants through competition or allelopathy, and researchers often use pot experiments as a tool to measure the strength of these interactions. Recently, such pot experiments provided inconsistent estimates of the impact and allelopathic potential of invasive knotweed, one of the world’s most successful plant invaders. We suspected that the inconsistencies may be explained by the use of different substrates in different experiments. To test this, we conducted an experiment in which knotweed competed pairwise with five common native European species in several different substrates: two compost-based potting substrates and two natural soils, with or without extra fertilizer added. To test for allelopathy, we added activated carbon to half of the pots. We found that knotweed was generally much more successful, and there was much more evidence for its allelopathy, when tested in artificial potting substrates than in natural soils. Furthermore, addition of extra fertilizer decreased the dominance of knotweed and changed patterns of allelopathy. The physicochemical properties of potting soil, such as lower bulk density, higher pore space, permeability and nitrogen content may better allow rhizomes to penetrate and/or allelochemicals to be produced and diffused. If artificial substrates generally exaggerate dominance and allelopathy also in other invasive plants, then many previous studies may have overestimated the potential impact of invaders, and the results of these experiments should be interpreted with caution. To avoid misleading results, experiments that test the competitive or allelopathic impact of invasive plants should be done with natural soils, preferably from the targeted habitats.