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.     

Invasion in space and time: non-native species richness and relative abundance respond to interannual variation in productivity and diversity

Ecologists have long sought to understand the relationships among species diversity, community productivity and invasion by non‐native species. Here, four long‐term observational datasets were analyzed using repeated measures statistics to determine how plant species richness and community resource capture (i.e. productivity) influenced invasion. Multiple factors influenced the results, including the metric used to quantify invasion, interannual variation and spatial scale. Native richness was positively correlated with non‐native richness, but was usually negatively correlated with non‐native abundance, and these patterns were stronger at the larger spatial scale. Logistic regressions indicated that the probability of invasion was reduced both within and following years with high productivity, except at the desert grassland site where high productivity was associated with increased invasion. Our analysis suggests that while non‐natives were most likely to establish in species rich communities, their success was diminished by high resource capture by the resident community.     

How invader traits interact with resident communities and resource availability to determine invasion success

Competition for limited resources is considered a key factor controlling invasion success. Resource availability can be viewed in either the long or short‐term. Long‐term availability depends on the baseline nutrient availability in the ecosystem and how those conditions shape the ecological community. Short‐term resource availability fluctuates with disturbances that alter nutrient availability and/or the density and composition of the ecological community. We investigated how species’ traits interact with short and long‐term resource availability to determine the outcome of invasions. We manipulated long‐term baseline resource availability, disturbance intensity, disturbance frequency, and propagule pressure in a fully factorial design using protist microcosms. Our results show that short and long‐term resource availability and the direct mortality from disturbance interact with the traits of resident community members and traits of invaders to determine community invasibility. While competitively dominant invaders with slow growth rates may suffer rather than benefit from short‐term resource fluctuations, quickly growing but competitively inferior invaders can benefit from both the resource fluctuations and the heterogeneity in community composition created by disturbance. Our findings empirically synthesize two explanations for invasion success, namely short‐term resource fluctuations and long‐term resource availability, and highlight the importance of considering traits of invaders and residents, such as growth rate and competitive ability, in the context of productivity and disturbance gradients. This species’ traits approach could resolve idiosyncratic results from natural systems undergoing disturbance and invasion that do not follow patterns predicted by traditional invasion frameworks.     

Can limiting similarity increase invasion resistance? A meta‐analysis of experimental studies

Synthesis We used meta‐analyses to examine experimental evidence that functional similarity between invaders and resident communities reduces invasion. We synthesized evidence from studies that experimentally added seed to resident communities in which the functional group composition had been manipulated. We found communities containing functionally similar resident species reduced invasion of forb but not grass invaders. However, experimental design dramatically influenced the results – with evidence for limiting similarity only found in artificially assembled communities, and not when studies used functional group removal from more ‘natural communities’. We suggest that functional group similarity plays a limited role in biotic resistance in established communities. The principle of limiting similarity suggests that species must be functionally different to coexist; based on the assumption that inter‐specific competition should be greatest between functionally similar species. There has been controversy over the generality of this assembly rule for plant communities with some studies finding evidence for limiting similarity and others not. One approach to testing this is to examine the ‘invasion’ success of species into communities in which the functional group composition has been manipulated. Using a meta‐analysis approach, we examined the generality of limiting similarity for plant communities based on published experimental studies. We asked – is establishment of an invading species less successful if it belongs to a functional group that is already present in the community compared to a community in which that functional group is absent? We explored separately colonisation (i.e. germination, establishment or seedling survival) and performance (i.e. biomass, cover or growth) of different functional groups (forbs and grasses) and experimental designs (removal experiments of more or less natural communities and synthetic‐assemblage experiments). We found that communities containing functionally similar resident species did reduce invader colonisation and performance of forb invaders, but did not reduce colonisation or performance of grass invaders. Evidence in support of limiting similarity was only detected in synthetic‐assemblage experiments and not when studies used functional group removal from ‘natural’ communities. Functional similarity is an important aspect of biotic resistance for forb invaders, but was only found in artificial communities. This has implications for restoration ecology especially when communities are built de novo. However, we suggest that limiting similarity plays a limited role in biotic resistance, because no evidence was detected in established communities.     

Species diversity reduces invasion success in pathogen‐regulated communities

The loss of natural enemies is thought to explain why certain invasive species are so spectacularly successful in their introduced range. However, if losing natural enemies leads to unregulated population growth, this implies that native species are themselves normally subject to natural enemy regulation. One possible widespread mechanism of natural enemy regulation is negative soil feedbacks, in which resident species growing on home soils are disadvantaged because of a build‐up of species‐specific soil pathogens. Here we construct simple models in which pathogens cause resident species to suffer reduced competitive ability on home soils and consider the consequences of such pathogen regulation for potential invading species. We show that the probability of successful invasion and its timescale depend strongly on the competitive ability of the invader on resident soils, but are unaffected by whether or not the invader also suffers reduced competitive ability on home soils (i.e. pathogen regulation). This is because, at the start of an invasion, the invader is rare and hence mostly encounters resident soils. However, the lack of pathogen regulation does allow the invader to achieve an unusually high population density. We also show that increasing resident species diversity in a pathogen‐regulated community increases invasion resistance by reducing the frequency of home‐site encounters. Diverse communities are more resistant to invasion than monocultures of the component species: they preclude a greater range of potential invaders, slow the timescale of invasion and reduce invader population size. Thus, widespread pathogen regulation of resident species is a potential explanation for the empirical observation that diverse communities are more invasion resistant.     

Life after establishment: factors structuring the success of a mountain invader away from disturbed roadsides

Climate change and increased anthropogenic activity may both alter the current ranges of non-native plant species in mountainous areas, and could result in increased success of such species at higher elevations in the future. However, the course that management should take is often unclear due to a lack of information about the dynamics of how successful mountain invaders spread away from roadsides. The goals of this study were to determine if patterns of growth of a successful mountain invader, Linaria dalmatica (L.) Mill., (as measured by species cover) were: (1) similar to those of establishment (as measured by probability of occurrence), and (2) structured by the extant plant community. Study sites were established throughout the current elevation range of L. dalmatica in the Greater Yellowstone Ecosystem, and cover of the species was measured along with several vegetative community characteristics. Elevation influenced probability of occurrence (i.e., chance of establishment) for L. dalmatica, but not cover (which represents growth after establishment). L. dalmatica cover was negatively associated with several vegetative community characteristics which did not appear to be influenced by the presence of L. dalmatica. These results suggest that L. dalmatica establishment may be limited by climate, but that spread of established populations away from roadsides is most influenced by properties of the vegetative community. They further suggest that the resident vegetative communities structure the abundance of this invader, and that to limit spread of this species in mountainous areas, disturbance to the existing vegetative communities should be minimized.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

The impact of interactions on invasion and colonization resistance in microbial communities

In human microbiota, the prevention or promotion of invasions can be crucial to human health. Invasion outcomes, in turn, are impacted by the composition of resident communities and interactions of resident members with the invader. Here we study how interactions influence invasion outcomes in microbial communities, when interactions are primarily mediated by chemicals that are released into or consumed from the environment. We use a previously developed dynamic model which explicitly includes species abundances and the concentrations of chemicals that mediate species interaction. Using this model, we assessed how species interactions impact invasion by simulating a new species being introduced into an existing resident community. We classified invasion outcomes as resistance, augmentation, displacement, or disruption depending on whether the richness of the resident community was maintained or decreased and whether the invader was maintained in the community or went extinct. We found that as the number of invaders introduced into the resident community increased, disruption rather than augmentation became more prevalent. With more facilitation of the invader by the resident community, resistance outcomes were replaced by displacement and augmentation. By contrast, with more facilitation among residents, displacement outcomes shifted to resistance. When facilitation of the resident community by the invader was eliminated, the majority of augmentation outcomes turned into displacement, while when inhibition of residents by invaders was eliminated, invasion outcomes were largely unaffected. Our results suggest that a better understanding of interactions within resident communities and between residents and invaders is crucial to predicting the success of invasions into microbial communities.     

Secondary invasion: When invasion success is contingent on other invaders altering the properties of recipient ecosystems

Positive interactions between exotic species may increase ecosystem‐level impacts and potentially facilitate the entry and spread of other exotic species. Invader‐facilitated invasion success—”secondary invasion”—is a key conceptual aspect of the well‐known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of “secondary invasion” and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term “secondary invasion” has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to “invader‐facilitated invasion,” which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting “secondary invasion” in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology.     

Plant invasion alters trait composition and diversity across habitats

Increased globalization has accelerated the movement of species around the world. Many of these nonnative species have the potential to profoundly alter ecosystems. The mechanisms underpinning this impact are often poorly understood, and traits are often overlooked when trying to understand and predict the impacts of species invasions on communities. We conducted an observational field experiment in Canada's first National Urban Park, where we collected trait data for seven different functional traits (height, stem width, specific leaf area, leaf percent nitrogen, and leaf percent carbon) across an abundance gradient of the invasive Vincetoxicum rossicum in open meadow and understory habitats. We assessed invasion impacts on communities, and associated mechanisms, by examining three complementary functional trait measures: community‐weighted mean, range of trait values, and species’ distances to the invader in trait space. We found that V. rossicum invasion significantly altered the functional structure of herbaceous plant communities. In both habitats V. rossicum changed the community‐weighted means, causing invaded communities to become increasingly similar in their functional structure. In addition, V. rossicum also reduced the trait ranges for a majority of traits indicating that species are being deterministically excluded in invaded communities. Further, we observed different trends in the meadow and understory habitats: In the understory, resident species that were more similar to V. rossicum in multivariate trait space were excluded more, however this was not the case in the meadow habitat. This suggests that V. rossicum alters communities uniquely in each habitat, in part by creating a filter in which only certain resident species are able to persist. This filtering process causes a nonrandom reduction in species' abundances, which in turn would be expected to alter how the invaded ecosystems function. Using trait‐based frameworks leads to better understanding and prediction of invasion impacts. This novel framework can also be used in restoration practices to understand how invasion impacts communities and to reassemble communities after invasive species management.     

Biotic resistance buffers the effects of nutrient enrichment on the success of a highly invasive aquatic plant

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Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Variation in the impact of non‐native seaweeds along gradients of habitat degradation: a meta‐analysis and an experimental test

Biological invasions are acknowledged among the main drivers of global changes in biodiversity. Despite compelling evidence of species interactions being strongly regulated by environmental conditions, there is a dearth of studies investigating how the effects of non‐native species vary among areas exposed to different anthropogenic pressures. Focusing on marine macroalgae, we performed a meta‐analysis to test whether and how the direction and magnitude of their effects on resident communities and species varies in relation to cumulative anthropogenic impact levels. The relationship between human impact levels and non‐native species impact intensity emerged only for a reduced subset of the response variables examined. Yet, there was a trend for the effects of non‐native species on community biomass and abundance and on species abundance to become less negative at heavily impacted sites. By contrast, the magnitude of negative effects of seaweed on community evenness tended to increase with human impact levels. The hypothesis of decreasing severity of invader’ impacts along a gradient of habitat degradation was also tested experimentally at a regional scale by comparing the effects of the removal of non‐native alga, Caulerpa cylindracea, on resident assemblages among rocky reefs exposed to different anthropogenic pressures. Assemblages at urban and pristine site did not differ when invaded, but did so when C. cylindracea was removed. Our results suggest that, despite the generally weak relationship between human impacts levels and non‐native species impacts, more negative impacts can be expected in less stressful environments (i.e. less degraded or pristine sites), where competitive interactions are presumably the driving force structuring resident communities. Implementing strategies for controlling the establishment of non‐native seaweeds should be, thus, considered a priority for preserving biodiversity in relatively pristine areas. On the other hand, control of invaders at degraded sites could be warranted to lessen their role as propagule sources. Synthesis Local anthropogenic stressors that severely alter biotic and abiotic conditions may underpin context‐dependency in the impacts of biological invasions. We used a meta‐analysis and an experimental test to examine the relationship between cumulative human impacts and ecological impact of non‐native seaweeds on resident assemblages. Our results suggest that more negative impacts of non‐native seaweeds on the abundance and biomass of resident assemblages can be expected in less degraded or pristine sites. Possibly, stronger impacts prevail at pristine sites, where assemblages are mainly structured by biotic interactions. Hence, management efforts should be mostly directed to prevent the establishment and spread of non‐native seaweeds in pristine areas. On the other hand, weak, but positive effects of seaweeds at the most degraded sites add to the ongoing debate on the role of non‐native species in rehabilitation plans.     

Facilitation promotes invasions in plant‐associated microbial communities

While several studies have established a positive correlation between community diversity and invasion resistance, it is less clear how species interactions within resident communities shape this process. Here, we experimentally tested how antagonistic and facilitative pairwise interactions within resident model microbial communities predict invasion by the plant–pathogenic bacterium Ralstonia solanacearum. We found that facilitative resident community interactions promoted and antagonistic interactions suppressed invasions both in the lab and in the tomato plant rhizosphere. Crucially, pairwise interactions reliably explained observed invasion outcomes also in multispecies communities, and mechanistically, this was linked to direct inhibition of the invader by antagonistic communities (antibiosis), and to a lesser degree by resource competition between members of the resident community and the invader. Together, our findings suggest that the type and strength of pairwise interactions can reliably predict the outcome of invasions in more complex multispecies communities.     

Strengthening Invasion Filters to Reassemble Native Plant Communities: Soil Resources and Phenological Overlap

Preventing invasion by exotic species is one of the key goals of restoration, and community assembly theory provides testable predictions about native community attributes that will best resist invasion. For instance, resource availability and biotic interactions may represent “filters” that limit the success of potential invaders. Communities are predicted to resist invasion when they contain native species that are functionally similar to potential invaders; where phenology may be a key functional trait. Nutrient reduction is another common strategy for reducing invasion following native species restoration, because soil nitrogen (N) enrichment often facilitates invasion. Here, we focus on restoring the herbaceous community associated with coastal sage scrub vegetation in Southern California; these communities are often highly invaded, especially by exotic annual grasses that are notoriously challenging for restoration. We created experimental plant communities composed of the same 20 native species, but manipulated functional group abundance (according to growth form, phenology, and N‐fixation capacity) and soil N availability. We fertilized to increase N, and added carbon to reduce N via microbial N immobilization. We found that N reduction decreased exotic cover, and the most successful seed mix for reducing exotic abundance varied depending on the invader functional type. For instance, exotic annual grasses were least abundant when the native community was dominated by early active forbs, which matched the phenology of the exotic annual grasses. Our findings show that nutrient availability and the timing of biotic interactions are key filters that can be manipulated in restoration to prevent invasion and maximize native species recovery.     

Is biotic resistance enhanced by natural variation in diversity?

Theories linking diversity to ecosystem function have been challenged by the widespread observation of more exotic species in more diverse native communities. Few studies have addressed the underlying processes by dissecting how biotic resistance to new invaders may be shaped by the same environmental influences that determine diversity and other community properties. In grasslands with heterogeneous soils, we added invaders and removed competitors to analyze the causes of invasion resistance. Abiotic resistance was measured using invader success in the absence of the resident community. Biotic resistance was measured as the reduction in invader success in the presence of the resident community. Invaders were most successful where biotic resistance was lowest and abiotic resistance was highest, confirming the dominant role of biotic resistance. Contrary to theory, though, biotic resistance was highest where both species richness and functional diversity were lowest. In the multivariate framework of a structural equation model, biotic resistance was independent of community diversity, and was highest where fertile soils led to high community biomass. Seed predation slightly augmented biotic resistance without qualitatively changing the results. Soil‐related genotypic variation in the invader also did not affect the results. We conclude that in natural systems, diversity may be correlated with invasibility and yet have little effect on biotic resistance to invasion. More generally, the environmental causes of variation in diversity should be considered when examining the potential functional consequences of diversity.     

Post‐fire vegetation dynamics in nutrient‐enriched and non‐enriched sclerophyll woodland

Abstract Exotic plant invasions are a significant problem in urban bushland in Sydney, Australia. In low‐nutrient Hawkesbury Sandstone communities, invasive plants are often associated with urban run‐off and subsequent increases in soil nutrients, particularly phosphorus. Fire is an important aspect of community dynamics in Sydney vegetation, and is sometimes used in bush regeneration projects as a tool for weed control. This study addressed the question: ‘Are there differences in post‐fire resprouting and germination of native and exotic species in nutrient‐enriched communities, compared with communities not disturbed by nutrient enrichment?’ We found that in non‐enriched areas, few exotic species emerged, and those that did were unable to achieve the rapid growth that was seen in exotic plants in the nutrient‐enriched areas. Therefore, fire did not promote the invasion of exotic plants into areas that were not nutrient‐enriched. In nutrient‐enriched areas after fire, the diversity of native species was lower than in the non‐enriched areas. Some native species were able to survive and compete with the exotic species in terms of abundance, per cent cover and plant height. However, these successful species were a different suite of natives to those commonly found in the non‐enriched areas. We suggest that although fire can be a useful tool for short‐term removal of exotic plant biomass from nutrient‐enriched areas, it does not promote establishment of native species that were not already present.     

Biodiversity influences invasion success of a facultative epiphytic seaweed in a marine forest

The biotic resistance hypothesis predicts that more diverse communities should have greater resistance to invasions than species-poor communities. However for facultative and obligate epiphytic invaders a high native species richness, abundance and community complexity might provide more resources for the invader to thrive to. We conducted surveys across space and time to test for the influence of native algal species abundance and richness on the abundance of the invasive facultative epiphytic filamentous alga Lophocladia lallemandii in a Mediterranean Cystoseira balearica seaweed forest. By removing different functional groups of algae, we also tested whether these relationships were dependent on the complexity and abundance of the native algal community. When invasion was first detected, Lophocladia abundance was positively related to species richness, but the correlation became negative after two years of invasion. Similarly, a negative relationship was also observed across sites. The removal experiment revealed that more complex native communities were more heavily invaded, where also a positive relationship was found between native algal richness and Lophocladia, independently of the native algal abundance. Our observational and experimental data show that, at early stages of invasion, species-rich seaweed forests are not more resistant to invasion than species-poor communities. Higher richness of native algal species may increase resource availability (i.e. substrate) for invader establishment, thus facilitating invasion. After the initial invasion stage, native species richness decreases with time since invasion, suggesting negative impacts of invasive species on native biodiversity.     

Invasibility of experimental grassland communities: the role of earthworms, plant functional group identity and seed size

Invasions of natural communities by non‐indigenous species threaten native biodiversity and are currently rated as one of the most important global‐scale environmental problems. The mechanisms that make communities resistant to invasions and drive the establishment success of seedlings are essential both for management and for understanding community assembly and structure. Especially in grasslands, anecic earthworms are known to function as ecosystem engineers, however, their direct effects on plant community composition and on the invasibility of plant communities via plant seed burial, ingestion and digestion are poorly understood. In a greenhouse experiment we investigated the impact of Lumbricus terrestris, plant functional group identity and seed size of plant invader species and plant functional group of the established plant community on the number and biomass of plant invaders. We set up 120 microcosms comprising four plant community treatments, two earthworm treatments and three plant invader treatments containing three seed size classes. Earthworm performance was influenced by an interaction between plant functional group identity of the established plant community and that of invader species. The established plant community and invader seed size affected the number of invader plants significantly, while invader biomass was only affected by the established community. Since earthworm effects on the number and biomass of invader plants varied with seed size and plant functional group identity they probably play a key role in seedling establishment and plant community composition. Seeds and germinating seedlings in earthworm burrows may significantly contribute to earthworm nutrition, but this deserves further attention. Lumbricus terrestris likely behaves like a ‘farmer’ by collecting plant seeds which cannot directly be swallowed or digested. Presumably, these seeds are left in middens and become eatable after partial microbial decay. Increased earthworm numbers in more diverse plant communities likely contribute to the positive relationship between plant species diversity and resistance against invaders.     

Non-random recruitment of invader species in experimental grasslands

To assess potential effects of seed limitation, characteristics of invader species and characteristics of established plant communities on recruitment success, we conducted a split-plot experiment factorially combining three weeding treatments corresponding to increasing successional age (regular weeding相似文献