Neighbour tolerance,not suppression,provides competitive advantage to non‐native plants

High competitive ability has often been invoked as a key determinant of invasion success and ecological impacts of non‐native plants. Yet our understanding of the strategies that non‐natives use to gain competitive dominance remains limited. Particularly, it remains unknown whether the two non‐mutually exclusive competitive strategies, neighbour suppression and neighbour tolerance, are equally important for the competitive advantage of non‐native plants. Here, we analyse data from 192 peer‐reviewed studies on pairwise plant competition within a Bayesian multilevel meta‐analytic framework and show that non‐native plants outperform their native counterparts due to high tolerance of competition, as opposed to strong suppressive ability. Competitive tolerance ability of non‐native plants was driven by neighbour's origin and was expressed in response to a heterospecific native but not heterospecific non‐native neighbour. In contrast to natives, non‐native species were not more suppressed by hetero‐ vs. conspecific neighbours, which was partially due to higher intensity of intraspecific competition among non‐natives. Heterogeneity in the data was primarily associated with methodological differences among studies and not with phylogenetic relatedness among species. Altogether, our synthesis demonstrates that non‐native plants are competitively distinct from native plants and challenges the common notion that neighbour suppression is the primary strategy for plant invasion success.     

Native generalist herbivores promote invasion of a chemically defended seaweed via refuge‐mediated apparent competition

Refuge‐mediated apparent competition was recently suggested as a mechanism that enables plant invasions. The refuge characteristics of introduced plants are predicted to enhance impacts of generalist herbivores on native competitors and thereby result in an increased abundance of the invader. However, this prediction has so far not been experimentally verified. This study tested if the invasion of a chemically defended seaweed is promoted by native generalist herbivores via refuge‐mediated apparent competition. The invader was shown to offer herbivores a significantly better refuge against fish predation compared with native seaweeds. Furthermore, in an experimental community, the presence of herbivores decreased the performance of neighbouring native seaweeds, but increased growth and relative abundance of the invader. These results provides the first experimental evidence that native generalist herbivores can shift a community towards a dominance of a well‐defended invader, inferior to native species in direct competitive interactions, by means of refuge‐mediated apparent competition.     

Exotic grasses and feces deposition by an exotic herbivore combine to reduce the relative abundance of native forbs

Increased resource availability can facilitate establishment of exotic plant species, especially when coincident with propagule supply. Following establishment, increased resource availability may also facilitate the spread of exotic plant species if it enhances their competitive abilities relative to native species. Exotic Canada geese (Branta canadensis) introduce both exotic grass seed and nutrients to an endangered plant community on the Gulf Islands of southwestern British Columbia, Canada. I used greenhouse experiments to assess the competitive advantage of the exotic grasses relative to native and exotic forbs in this community and to test the impacts of nutrient addition from goose feces on competitive outcomes. I grew experimental communities varying in their proportion of forbs versus exotic grasses, and added goose feces as a nutrient source. I found that both native and exotic forbs produced significantly more biomass in competition with conspecifics than in competition with the grasses, and that the proportional abundance of two out of three native forbs was lowest in the combined presence of exotic grasses and nutrient addition. In a second experiment, I found that in monoculture all species of forbs and grasses showed equal growth responses to nutrients. The exotic species did not convert additional nutrients into additional biomass at a higher rate, but did germinate earlier and grow larger than the native species regardless of nutrient availability. This suggests that the exotic species may have achieved their competitive advantage partly by pre-empting resources in community mixtures. Small and late-germinating native forbs may be particularly vulnerable to competitive suppression from exotic grasses and forbs and may be at an even greater disadvantage if their competitors are benefiting from early access to additional nutrients. In combination, the input of exotic propagules and additional nutrients by nesting geese may compromise efforts to maintain native community composition in this system.     

Consequences of differing competitive abilities between juvenile and adult plants

The competitive ability of perennial plants can change with life-stage, but whether these changes have fitness consequences is unknown. We present a simple model of two components of fitness, mortality and flowering rates, for two grassland species with very different patterns of competitive ability and life-stage. Achillea millefolium seedlings are poor competitors while the adults are good competitors. In contrast, Solidago missouriensis seedlings and adults have similar competitive ability. Models of the two species show that the overall effects of competition on growth are more important than interspecific differences in competitive ability in determining mortality and flowering rates, though the higher seedling competitive ability of S. missouriensis relative to A. millefolium seedlings does result in slightly lower mortality and higher flowering rates for the former species. Simulations where both average competitive ability and relative seedling and adult competitive ability are varied predict that dominant species with high overall competitive ability should experience no advantage or disadvantage from varying competitive ability through development. When overall competitive ability is moderate, the relative costs and benefits of differential competitive abilities among adults and seedlings are variable. High seedling competitive ability relative to adult competitive ability should be favored among species with low overall competitive ability. We predict that communities with high intensity of competition should have a high frequency of species with high seedling competitive ability, while communities with lower intensity of competition should have species with a wide range of relative seedling and adult competitive ability.     

Disturbance governs dominance of an invasive forb in a temporary wetland

Dominance of invasive species is often assumed to be due to a superior ability to acquire resources. However, dominance in plant communities can arise through multiple interacting mechanisms, including disturbance. Inter-specific competition can be strongly affected by abiotic conditions, which can determine the outcome of competitive interactions. We evaluated competition and disturbance as mechanisms governing dominance of Phyla canescens (hereafter lippia), an invasive perennial forb from South America, in Paspalum distichum (perennial grass, hereafter water couch) meadows in floodplain wetlands of eastern Australia. Water couch meadows (in the study area) are listed under the Ramsar Convention due to their significance as habitat for migratory waterbirds. In the field, we monitored patterns of vegetation boundaries between the two species in response to flooding. Under controlled glasshouse conditions, we explored competitive interactions between the native water couch and lippia subject to different soil moisture/inundation regimes. We did this using a pairwise factorial glasshouse experiment that manipulated neighbor density (9 treatments) and soil moisture/inundation (4 treatments). In the field trial, inundation increased the cover of water couch. Under more controlled conditions, the invader had a competitive effect on the native species only under dry soil conditions, and was strongly inhibited by inundation. This suggests that dry conditions favor the growth of the invader and wetter (more historical) conditions favor the native grass. In this system, invader dominance is governed by altered disturbance regimes which give the invader a competitive advantage over the native species.     

Shift in competitive ability mediated by soil biota in an invasive plant

Understanding the shifts in competitive ability and its driving forces is key to predict the future of plant invasion. Changes in the competition environment and soil biota are two selective forces that impose remarkable influences on competitive ability. By far, evidence of the interactive effects of competition environment and soil biota on competitive ability of invasive species is rare. Here, we investigated their interactive effects using an invasive perennial vine, Mikania micrantha. The competitive performance of seven M. micrantha populations varying in their conspecific and heterospecific abundance were monitored in a greenhouse experiment, by manipulating soil biota (live and sterilized) and competition conditions (competition‐free, intraspecific, and interspecific competition). Our results showed that with increasing conspecific abundance and decreasing heterospecific abundance, (1) M. micrantha increased intraspecific competition tolerance and intra‐ vs. interspecific competitive ability but decreased interspecific competition tolerance; (2) M. micrantha increased tolerance of the negative soil biota effect; and (3) interspecific competition tolerance of M. micrantha was increasingly suppressed by the presence of soil biota, but intraspecific competition tolerance was less affected. These results highlight the importance of the soil biota effect on the evolution of competitive ability during the invasion process. To better control M. micrantha invasion, our results imply that introduction of competition‐tolerant native plants that align with conservation priorities may be effective where M. micrantha populations are long‐established and inferior in inter‐ vs. intraspecific competitive ability, whereas eradication may be effective where populations are newly invaded and fast‐growing.     

Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (±)-catechin

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the “novel weapons” hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the “evolution of increased competitive ability” hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (±)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial Heterogeneity in Soil Microbes Alters Outcomes of Plant Competition

Plant species vary greatly in their responsiveness to nutritional soil mutualists, such as mycorrhizal fungi and rhizobia, and this responsiveness is associated with a trade-off in allocation to root structures for resource uptake. As a result, the outcome of plant competition can change with the density of mutualists, with microbe-responsive plant species having high competitive ability when mutualists are abundant and non-responsive plants having high competitive ability with low densities of mutualists. When responsive plant species also allow mutualists to grow to greater densities, changes in mutualist density can generate a positive feedback, reinforcing an initial advantage to either plant type. We study a model of mutualist-mediated competition to understand outcomes of plant-plant interactions within a patchy environment. We find that a microbe-responsive plant can exclude a non-responsive plant from some initial conditions, but it must do so across the landscape including in the microbe-free areas where it is a poorer competitor. Otherwise, the non-responsive plant will persist in both mutualist-free and mutualist-rich regions. We apply our general findings to two different biological scenarios: invasion of a non-responsive plant into an established microbe-responsive native population, and successional replacement of non-responders by microbe-responsive species. We find that resistance to invasion is greatest when seed dispersal by the native plant is modest and dispersal by the invader is greater. Nonetheless, a native plant that relies on microbial mutualists for competitive dominance may be particularly vulnerable to invasion because any disturbance that temporarily reduces its density or that of the mutualist creates a window for a non-responsive invader to establish dominance. We further find that the positive feedbacks from associations with beneficial soil microbes create resistance to successional turnover. Our theoretical results constitute an important first step toward developing a general understanding of the interplay between mutualism and competition in patchy landscapes, and generate qualitative predictions that may be tested in future empirical studies.     

Conundrums of competitive ability in plants: what to measure?

A survey of recent literature indicates that competitive ability in plants has been measured, in most studies, only in terms of the relative intensity of size suppression experienced by competitors within one growing season. Far fewer studies have recorded relative success in terms of survival and even fewer studies have recorded fecundity under competition. Differences in size suppression are usually assumed to reflect differences in relative abilities to deny resources to competitors. However, most previous studies have failed to control or account for other sources of variation in the size suppression that plants experience under competition, i.e. variation between mixtures in the resource supply/demand ratio (approach to carrying capacity), or variation in the degree of niche overlap between competitors, or variation in the intensity of concurrent facilitative interactions between competitors. For future studies, much greater caution is required in recognizing these inherent limitations of traditional measures of competitive ability and, hence, guarding against unfounded conclusions or predictions about potential for competitive success that are based on these measures. There is also a significant challenge for future studies to adopt empirical approaches for minimizing these limitations. Some initial recommendations are considered here based on an emerging view of competitive ability measured in terms of traits associated with all three conventional components of Darwinian fitness, i.e. not just growth (plant size) but also survival and fecundity allocation (offspring production per unit plant size per unit time). According to this model, differences in competitive ability imply differences in the ability, despite intense competition (i.e. low resource supply/demand ratio), to recruit offspring into the next generation and thereby limit offspring recruitment by other plants. The important traits of competitive ability, therefore, are not only those that allow a plant to deny resources to competitors, suppress their sizes and hence, maximize the plant's own size, but also those traits that allow the plant to withstand suppression from competition enough to persist, both as an individual (through survival) and across generations (through descendants).     

EVOLUTION OF BROAD AND SPECIFIC COMPETITIVE ABILITY IN NOVEL VERSUS FAMILIAR ENVIRONMENTS IN DROSOPHILA SPECIES

We used nine pairs of competing Drosophila melanogaster and Drosophila simulans populations to test three hypotheses. (1) Weaker competitors undergo greater evolutionary increases in competitive ability, compared with stronger ones. (2) Increased competitive ability against a specific competitor population causes a correlated increase in competitive ability against other competitor populations. (3) In a novel environment, adaptation to the abiotic environment contributes more to competitive ability than adaptation to the competitor population. After 11 generations of competition, initially weaker competitor populations showed relatively greater increases in competitive ability. Broad and specific competitive abilities, the latter being specific to a particular competitor population, were positively correlated in both familiar and novel environments. Adaptation to the abiotic environment seemed to be a more important component of competitive ability in the novel environments. We conclude that in geographically structured species, biotic and abiotic factors affecting the evolution of competitive ability may interact to help create a mosaic of outcomes that can affect the evolutionary dynamics of the interaction over the range of the competing species.     

Extended leaf phenology may drive plant invasion through direct and apparent competition

Invasive plants can inflict great harm, yet drivers of successful invasion remain unclear. Many invaders of North American deciduous forests exhibit extended leaf phenology (ELP), or longer growing season relative to natives. ELP may grant invaders competitive advantages, but we argue that ELP more potently drives invasion in the presence of herbivores. ELP invaders can support herbivores by lessening starvation during winter; consequently, native plants may suffer when attacked later through apparent competition. As modeled here, even short ELP can promote competitive success of invaders, and apparent competition sharply enhances ELP invader dominance. In ‘partial enemy escape’ scenarios, a less palatable ELP invader nearly excludes a preferred native where an invader without ELP could not. Together, ELP and apparent competition enhance invasion even when biotic resistance should suppress it, i.e. when the invader competes weakly or provides preferred forage. Thus, ELP‐apparent competition interactions grant invaders considerable success while challenging core tenets of invasion ecology.     

A trait-based approach across the native and invaded range to understand plant invasiveness and community impact

Alien plant species invasiveness and impact on diversity (i.e. species richness and composition) can be driven by the altered competitive interactions experienced by the invader in its invaded range compared to its native range. Trait-based competition effects on invasiveness can be mediated through size-asymmetric competition, i.e. a trait suit of the invader that drives competitive dominance, and through ‘niche differences', i.e. trait differentiation and thus minimized competition between invader and the invaded community. In terms of invasion impact, size-asymmetric competition is expected to result in competitive exclusion of co-occurring subordinate species, whereas ‘niche differences' might result in competitive exclusion of the most functionally similar co-occurring species. Although observational work does not allow the full disentanglement of both trait-based effects, it does allow to verify the occurrence of expected theoretical trait patters. In this study, we explored the trait-based competition effects on invasiveness and diversity impact for Rosa rugosa in both its invaded range in Belgium and its native range in Japan, based on seven functional traits across 100 vegetation plots. Following the predictions for enhanced invasiveness, we found much lower functional overlap between R. rugosa and the co-occurring species in the invaded range compared to the native range. This likely also explains the absence of diversity impact in its native range. Despite the absence of changes in species richness in the invaded range, the invader did strongly impact species composition of invaded communities. This impact occurred through strong shade tolerance responses, suggesting size-asymmetric competition effects and cover loss of co-occurring dominant species, next to exclusion of co-occurring species most functionally similar to the invader; suggesting niche difference effects. In conclusion, this case-study illustrates how exploring functional trait patterns across a species native and invaded range can help in understanding how trait-based competition processes can affect invasiveness and community impact.     

Are alien plants more competitive than their native conspecifics? A test using Hypericum perforatum L.

The evolution of increased competitive ability hypothesis predicts that introduced plants that are long liberated from their natural enemies may lose costly herbivore defense, enabling them to reallocate resources previously spent on defense to traits that increase competitive superiority. We tested this prediction by comparing the competitive ability of native St John's wort ( Hypericum perforatum) from Europe with introduced St John's wort from central North America where plants have long grown free of specialist herbivores, and introduced plants from western North America where plants have been subjected to over 57 years of biological control. Plants were grown in a greenhouse with and without competition with Italian ryegrass ( Lolium multiflorum). St John's wort from the introduced range were not better interspecific competitors than plants from the native range. The magnitude of the effect of ryegrass on St John's wort was similar for introduced and native genotypes. Furthermore, introduced plants were not uniformly larger than natives; rather, within each region of origin there was a high variability in size between populations. Competition with ryegrass reduced the growth of St John's wort by >90%. In contrast, St John's wort reduced ryegrass growth <10%. These results do not support the contention that plants from the introduced range evolve greater competitive ability in the absence of natural enemies.     

Early emergence and resource availability can competitively favour natives over a functionally similar invader

Invasive plant species can form dense populations across large tracts of land. Based on these observations of dominance, invaders are often described as competitively superior, despite little direct evidence of competitive interactions with natives. The few studies that have measured competitive interactions have tended to compare an invader to natives that are unlikely to be strong competitors because they are functionally different. In this study, we measured competitive interactions among an invasive grass and two Australian native grasses that are functionally similar and widely distributed. We conducted a pair-wise glasshouse experiment, where we manipulated both biotic factors (timing of establishment, neighbour identity and density) and abiotic factors (nutrients and timing of water supply). We found that the invader significantly suppressed the performance of the natives; but its suppression ability was contingent on resource levels, with pulsed water/low nutrients or continuous watering reducing its competitive effects. The native grasses were able to suppress the performance of the invader when given a 3-week head-start, suggesting the invader may be incapable of establishing unless it emerges first, including in its own understorey. These findings provide insight for restoration, as the competitive effect of a functionally similar invader may be reduced by altering abiotic and biotic conditions in favour of natives.     

Competition strength of two significant invasive species in coastal dunes

To investigate the effect of increased nutrient availability on competition amongst invasive and native plants, I measured changes in above and below ground biomass of Chrysanthemoides monilifera spp. rotundata (bitou bush) and Asparagus aethiopicus (asparagus fern) competing with two native species, Banksia integrifolia and Ficinia nodosa, under high- and low-nutrient regimes. Bitou bush, as a primary invader, was competitive under all conditions lowering the growth of native species in both high and low nutrients. Asparagus fern as a secondary invader, did not influence growth of native species but responded, like bitou bush, to high nutrients. Native species were generally negatively affected by increases in nutrients. With bitou bush soils often providing higher nutrients, the chance of secondary invasion by asparagus fern is more likely, although asparagus fern is unlikely to invade low nutrient soils quickly. The invasive species, therefore, differed in their competitive ability in these coastal dune communities.     

Differential effect of inter- and intraspecific competition on the performance of invasive and native Taraxacum species

Inter- and intraspecific competitive abilities are significant determinants of invasive success and the ecological impact of non-native plants. We tested two major hypotheses on the competitive ability of invasive species using invasive (Taraxacum officinale) and native (T. platycarpum) dandelions: differential interspecific competitive ability between invasive and native species and the kin recognition of invasive species. We collected seeds from two field sites where the two dandelion species occurred nearby. Plants were grown alone, with kin (plants from the same maternal genotype) or strangers (plants from different populations) of the same species, or with different species in a growth chamber, and the performance at the early developmental stage between species and treatments was compared. The invasive dandelions outcompeted the native dandelions when competing against each other, although no difference between species was detected without competition or with intraspecific competition. Populations of native species responded to interspecific competition differently. The effect of kinship on plant performance differed between the tested populations in both species. A population produced more biomass than the other populations when grown with a stranger, and this trend was manifested more in native species. Our results support the hypothesis that invasive plants have better competitive ability than native plants, which potentially contributes to the establishment and the range expansion of T. officinale in the introduced range. Although kin recognition is expected to evolve in invasive species, the competitive ability of populations rather than kinship seems to affect plant growth of invasive T. officinale under intraspecific competition.     

Context‐dependent induction of allelopathy in plants under competition

Some plants use allelopathy to compete against neighbouring plants, and the ability to induce allelopathic compound production in response to competition is hypothesized to be adaptive, as plants can save costs of metabolite production in the absence of competitors. However, whether plants induce allelopathy has rarely been explored so far. We studied the inducibility of polyacetylenes – putative allelopathic compounds in Solidago altissima – in response to competition. Polyacetylenes were found in natural soil surrounding S. altissima patches within the range of concentration known to inhibit competitor growth. Individual S. altissima plants with higher polyacetylene concentration in roots suppressed the growth of the competitor plants more, suggesting that root polyacetylene levels proximate plants’ allelopathic capacity. Competition induced polyacetylenes in a context‐dependent manner: Whereas introduced Japanese and Australian populations of S. altissima had higher constitutive concentration of polyacetylenes than the native North American populations, inducibility was observed only in Australian plants, where the population is still at an early stage of invasion. Also, induction became more prominent under nutrient depletion, where enhanced allelopathy may be particularly beneficial for suppressing a competitor's exploitative capacity. Finally, we found weak evidence for a tradeoff between constitutive and induced polyacetylenes. The observed patterns suggest that allelopathic plants could respond to competition by inducing allelochemical production, but the benefit of such plasticity may vary across time and space. Shifts in competitor communities in introduced range over time may shape plant's plastic responses to competition, while variation in resource availability may alter competitive environment to influence the degree to which plants induce allelopathy.     

Plant tolerance and resistance in food webs: community-level predictions and evolutionary implications

While evolutionary ecologists emphasize different ways in which plants can evolutionarily respond to herbivory, such as resistance or tolerance, community ecology has lagged in its understanding of how these different plant traits can influence interactions, abundance, composition, and diversity within more complex food webs. In this paper, we present a series of models comparing community level outcomes when plants either resist or tolerate herbivory. We show that resistance and tolerance can lead to very different outcomes. A particularly important result is that resistant species should often coexist locally with other, less resistant competitors, whereas tolerant species should not be able to coexist locally with less tolerant competitors, although priority effects allow them to coexist regionally. We also use these models to suggest some insights into the evolution of these traits within more complex communities. We emphasize how understanding the differential effects of plant tolerance and resistance in food webs provides greater appreciation of a variety of empirical patterns that heretofore have appeared enigmatic. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic variation in tolerance of competition and neighbour suppression in Arabidopsis thaliana

Intraspecific competitive interactions can profoundly influence phenotypic evolution. However, prior studies have rarely evaluated the evolutionary potential of the two components of competitive ability, tolerance of competition and suppression of neighbours. Here, we grow a set of 20 Arabidopsis thaliana recombinant inbred lines in three competitive treatments (noncompetitive, intra‐genotypic competition and inter‐genotypic competition) to examine if there is genetic variation for the components of competitive ability and whether neighbour relatedness has an effect on fitness. We find evidence for genetic variation in tolerance of competition and neighbour suppression and that these two competitive strategies are correlated, such that genotypes that tolerate competition will also strongly suppress neighbours. We further observe that the effect of neighbour relatedness on fitness of target individuals depends on neighbour identity, i.e. whether target individuals perform better when competing against self vs. nonself individuals depends on the genotypic identity of the nonself neighbour. The results are particularly relevant to evolutionary responses under multi‐level selection.     

The competition–dispersal trade‐off exists in forbs but not in graminoids: A case study from multispecies alpine grassland communities

Much theoretical evidence has demonstrated that a trade‐off between competitive and dispersal ability plays an important role in facilitating species coexistence. However, experimental evidence from natural communities is still rare. Here, we tested the competition–dispersal trade‐off hypothesis in an alpine grassland in the Tianshan Mountains, Xinjiang, China, by quantifying competitive and dispersal ability using a combination of 4 plant traits (seed mass, ramet mass, height, and dispersal mode). Our results show that the competition–dispersal trade‐off exists in the alpine grassland community and that this pattern was primarily demonstrated by forbs. The results suggest that most forb species are constrained to be either good competitors or good dispersers but not both, while there was no significant trade‐off between competitive and dispersal ability for most graminoids. This might occur because graminoids undergo clonal reproduction, which allows them to find more benign microenvironments, forage for nutrients across a large area and store resources in clonal structures, and they are thus not strictly limited by the particular resources at our study site. To the best of our knowledge, this is the first time the CD trade‐off has been tested for plants across the whole life cycle in a natural multispecies plant community, and more comprehensive studies are still needed to explore the underlying mechanisms and the linkage between the CD trade‐off and community composition.