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.     

Traits linked with species invasiveness and community invasibility vary with time,stage and indicator of invasion in a long‐term grassland experiment

Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.     

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.     

Intraspecific genotypic richness and relatedness predict the invasibility of microbial communities

Biological invasions can lead to extinction events in resident communities and compromise ecosystem functioning. We tested the effect of two widespread biodiversity measurements, genotypic richness and genotypic dissimilarity on community invasibility. We manipulated the genetic structure of bacterial communities (Pseudomonas fluorescens) and submitted them to invasion by Serratia liquefaciens. We show that the two diversity measures impact on invasibility via distinct and additive mechanisms. Genotypic dissimilarity of the resident communities linearly increased productivity and in parallel decreased invasion success, indicating that high dissimilarity prevents invasion through niche pre-emption. By contrast, genotypic richness exerted a hump-shaped effect on invasion and was linked to the production of toxins antagonistic to the invader. This effect peaked at intermediate richness, suggesting that high richness levels may increase invasibility. Invasibility could be well predicted by the combination of these two mechanisms, documenting that both genotypic richness and dissimilarity need to be considered, if we are to understand the biotic properties determining the susceptibility of ecosystems to biological invasions.     

Distinct invasion strategies operating within a natural annual plant system

Alien plant species are known to have a wide range of impacts on recipient communities, from resident species’ exclusions to coexistence with resident species. It remains unclear; however, if this variety of impacts is due to different invader strategies, features of recipient communities or both. To test this, we examined multiple plant invasions of a single ecosystem in southwestern Australia. We used extensive community data to calculate pairwise segregation between target alien species and many co‐occurring species. We related segregation to species’ positions along community trait hierarchies and identified at least two distinct invasion strategies: ‘exploiters’ which occupy high positions along key trait hierarchies and reduce local native species diversity (particularly in nutrient‐enriched situations), and ‘coexisters’ who occupy intermediate trait positions and have no discernable impact on native diversity. We conclude that trait hierarchies, linked to measures of competition, can provide valuable insights about the processes driving different invasion outcomes.     

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.     

Siderophores drive invasion dynamics in bacterial communities through their dual role as public good versus public bad

Microbial invasions can compromise ecosystem services and spur dysbiosis and disease in hosts. Nevertheless, the mechanisms determining invasion outcomes often remain unclear. Here, we examine the role of iron-scavenging siderophores in driving invasions of Pseudomonas aeruginosa into resident communities of environmental pseudomonads. Siderophores can be ‘public goods’ by delivering iron to individuals possessing matching receptors; but they can also be ‘public bads’ by withholding iron from competitors lacking these receptors. Accordingly, siderophores should either promote or impede invasion, depending on their effects on invader and resident growth. Using supernatant feeding and invasion assays, we show that invasion success indeed increased when the invader could use its siderophores to inhibit (public bad) rather than stimulate (public good) resident growth. Conversely, invasion success decreased the more the invader was inhibited by the residents’ siderophores. Our findings identify siderophores as a major driver of invasion dynamics in bacterial communities under iron-limited conditions.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

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.     

Every Coin Has a Back Side: Invasion by Limnohabitans planktonicus Promotes the Maintenance of Species Diversity in Bacterial Communities

One of the earliest challenges for ecologists has been to study the impact of invasive species on microbial communities. Although bacteria are fundamental in biological processes, current knowledge on invasion effects by aquatic non-pathogenic bacteria is still limited. Using pure cultures of diverse planktonic bacteria as model organisms at two different carbon concentration levels, we tested the response of an assembled community to the invasion by Limnohabitans planktonicus, an opportunistic bacterium, successful in freshwaters. The invader, introduced at the early stationary growth phase of the resident community, caused a strong decrement of the abundance of the dominant species. This was due to competition for nutrients and a potential allelopathic interaction. Simultaneously, resident species formerly unable to successfully compete within the community, thus potentially exposed to competitive exclusion, increased their abundances. The overall result of the invasion was preservation of species diversity, the higher the lower was the substrate content available. Our study provides new insights into bacterial invasions, offering an alternative interpretation of invasions for community ecology.     

Testing population-level mechanisms of invasion by a mobile vertebrate: a simple conceptual framework for salmonids in streams

Invasion biology suffers from a lack of the ability to predict the outcome of particular invasions because of reliance on verbal models and lack of rigorous experimental data at the appropriate scale. More progress is likely to be made by considering invasions as population-level phenomena and initially focusing on specific taxa or particular categories of invasions. To this end, we propose a simple conceptual framework to motivate studies of invasion by salmonids (salmon, trout, grayling, and whitefish) in streams that emphasizes population-level mechanisms affecting native species and promoting spread by the invader. Specifically, the only direct mechanisms by which the abundance of the native species can decline are through biotic interactions which cause decreased reproductive rates or survival at specific life stages, net emigration, debilitating or fatal diseases introduced by the invader, or a combination of these factors. Conversely, abundance of the invader must increase by local reproduction, high survival, net immigration, or a combination of these factors. Review of existing salmonid invasion literature suggests that future studies could be improved by using manipulative field experiments at a spatial and temporal scale appropriate to address population-level processes, characterizing how movement affects the establishment and spread of an invader, and including abiotic context in experimental designs. Using the example of brook trout (Salvelinus fontinalis) invasion into streams containing native Colorado River cutthroat trout (Oncorhynchus clarki pleuriticus) in the central Rocky Mountains (USA), we demonstrate how the framework can be used to design a manipulative field experiment to test for population-level mechanisms causing ecological effects and promoting invasion success. Experiments of this type will give invasion ecologists a useful example of how a taxon-specific invasion framework can improve the ability to predict ecological effects, and provide fishery biologists with the quantitative foundation necessary to better manage stream salmonid invasions.     

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.     

Can Invasive Species Facilitate Native Species? Evidence of How, When, and Why These Impacts Occur

Although the predatory and competitive impacts of biological invasions are well documented, facilitation of native species by non-indigenous species is frequently overlooked. A search through recent ecological literature found that facilitative interactions between invasive and native species occur in a wide range of habitats, can have cascading effects across trophic levels, can re-structure communities, and can lead to evolutionary changes. These are critical findings that, until now, have been mostly absent from analyses of ecological impacts of biological invasions. Here I present evidence for several mechanisms that exemplify how exotic species can facilitate native species. These mechanisms include habitat modification, trophic subsidy, pollination, competitive release, and predatory release. Habitat modification is the most frequently documented mechanism, reflecting the drastic changes generated by the invasion of functionally novel habitat engineers. Further, I predict that facilitative impacts on native species will be most likely when invasive species provide a limiting resource, increase habitat complexity, functionally replace a native species, or ameliorate predation or competition. Finally, three types of facilitation (novel, substitutive, and indirect) define why exotic species can lead to facilitation of native species. It is evident that understanding biological invasions at the community and ecosystem levels will be more accurate if we integrate facilitative interactions into future ecological research. Since facilitative impacts of biological invasions can occur with native endangered species, and can have wide-ranging impacts, these results also have important implications for management, eradication, and restoration.Contribution Number 2293, Bodega Marine Laboratory, University of California at Davis.     

Soil biota composition and the performance of a noxious weed across its invaded range

The success of invasive plant species is driven, in part, by feedback with soil ecosystems. Yet, how variation in belowground communities across latitudinal gradients affects invader distributions remains poorly understood. To determine the effect of soil communities on the performance of the noxious weed Cirsium arvense across its invaded range, we grew seedlings for 40 days in soils collected across a 699 km linear distance from both inside and outside established populations. We also described the mesofaunal and bacterial communities across all soil samples. We found that C. arvense typically performed better when grown in soils sourced from northern populations than from southern locations where it has a longer invasion history. We also found evidence that C. arvense performed best in soils sourced from outside invaded patches, although this was not consistent across all sites. The bacterial community showed a significant increase in the magnitude of compositional change in invaded sites at higher latitudes, while the mesofaunal community showed the opposite pattern. Bacterial community composition was significantly correlated with C. arvense performance, although mesofaunal community composition was not. Our results demonstrate that the interactions between an invasive plant and associated soil communities change across the invaded range, and the bacterial community in particular may affect variation in plant performance. Observed patterns may be caused by C.arvense presence and time since invasion allowing for an accumulation of species‐specific pathogens in southern soils, while the naïveté of northern soils to invasion results in a more responsive bacterial community. Although these interactions are difficult to predict, such effects could possibly facilitate the establishment of this exotic species to novel locations.     

Evolutionary game theory as a framework for studying biological invasions

Although biological invasions pose serious threats to biodiversity, they also provide the opportunity to better understand interactions between the ecological and evolutionary processes structuring populations and communities. However, ecoevolutionary frameworks for studying species invasions are lacking. We propose using game theory and the concept of an evolutionarily stable strategy (ESS) as a conceptual framework for integrating the ecological and evolutionary dynamics of invasions. We suggest that the pathways by which a recipient community may have no ESS provide mechanistic hypotheses for how such communities may be vulnerable to invasion and how invaders can exploit these vulnerabilities. We distinguish among these pathways by formalizing the evolutionary contexts of the invader relative to the recipient community. We model both the ecological and the adaptive dynamics of the interacting species. We show how the ESS concept provides new mechanistic hypotheses for when invasions result in long- or short-term increases in biodiversity, species replacement, and subsequent evolutionary changes.     

Experimental test of the Invasional Meltdown Hypothesis: an exotic herbivore facilitates an exotic plant,but the plant does not reciprocally facilitate the herbivore

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Biodiversity and belowground interactions mediate community invasion resistance against a tall herb invader

Aims Species-rich plant communities are hypothesized to be more resistant against plant invasions because they use resources in a more efficient way. However, the relative contributions of aboveground competition and belowground interactions for invasion resistance are still poorly understood.Methods We compared the performance of Knautia arvensis transplants growing in plots differing in plant diversity both under full competition and with shoots of neighbors tied back to determine the relative strength of aboveground competition in suppressing this test invader without the confounding effect of shading. In addition, we assessed the effects of belowground competition and soil-borne pathogens on transplant performance.Important findings Both aboveground competition and plant species richness strongly and independently affected invader performance. Aboveground biomass, height, leaf mass per area and flowering of transplanted individuals of K. arvensis decreased with increasing species richness of the host community. Species-rich and species-poor communities both imposed equally strong aboveground competition on K. arvensis. However, belowground interactions (especially belowground root competition) had strong negative effects on transplant performance. In addition, the presence of grasses in a plant community further reduced the performance of K. arvensis. Our results suggest that belowground competition can render species-rich host communities more suppressive to newly arriving species, thus enhancing community invasion resistance.     

The importance of biological inertia in plant community resistance to invasion

Abstract. Insights into the ecology of historic invasions by introduced species can be gained by studying long‐term patterns of invasions by native species. In this paper, we review literature in palaeo‐ecology, forest‐stand simulation modelling, and historical studies of plant species invasions to illustrate the relevance of biological inertia in plant communities to invasion ecology. Resistance to invasion occurs in part because of environmental, demographic, and biotic factors influencing the arrival and establishment of invading species. We propose that biological inertia within the resident community is a fourth component of resistance to invasion, because of the lag time inherent in eliminating resident species and perhaps their traces after environmental conditions become suitable for invasion by immigrating species. Whether or not an introduced species invades can be conditioned by the presence of the pre‐existing community (and/or its legacy) in addition to the other biotic and abiotic factors.     

Invasions: the perspective of diverse plant communities

Exotic plant invasions represent a threat to natural and managed ecosystems. Understanding of the mechanisms that determine why a given species may invade a given ecosystem, or why some biomes and regions seem more prone to invasions, is limited. One potential reason for this lack of progress may lie in how few studies have addressed invasion mechanisms from the point of view of the invaded community. On the other hand, the renewed debate about the relationship between ecological diversity and ecosystem stability offers the opportunity to revisit existing theory and empirical evidence, and to attempt to investigate which characteristics of plant communities, including their diversity, contribute to their invasibility. Empirical studies have shown both positive and negative relationships between species diversity of resident plant communities and their invasibility by external species. Rather than attempting to build a larger collection of case studies, research now needs to address the mechanisms underlying these relationships. Previous knowledge about the mechanisms favouring invasion needs to be coupled with community theory to form the basis of these new investigations. Modern community theory offers hypotheses and techniques to analyse the invasibility of communities depending on their diversity and other factors, such as species’ life histories and environmental variability. The body of knowledge accumulated in invasion ecology suggests that the role of disturbances, in interaction with fertility, and the importance of interactions with other trophic levels, are specific factors for consideration. In addition, it is essential for future studies to explicitly tease apart the effects of species richness per se from the effects of other components of ecological diversity, such as functional diversity (the number of functional groups) and trophic diversity (the number of interactions among trophic levels).