Applying competition theory to invasion: resource impacts indicate invasion mechanisms in California shrublands

Despite widespread work documenting invasion, it remains a challenge to determine invasion mechanisms and incorporate them into invasive species management. Competition theory presents a strong model for evaluating the role of resource reduction and requirements in invasion. Additionally, alternative models suggest fluctuations in resources, niche differences, or non-resource priority effects are key factors determining invasion success. We propose a comparative framework that incorporates resource impacts of native and invasive species, performance in controlled invasion trials, and long-term natural invasion patterns to elucidate relative importance of these invasion mechanisms. To demonstrate this framework, we established monocultures of two representative native and two invasive plant species in Southern California’s coastal sage scrub (CSS), measured resource impacts (i.e., R*), and conducted invasion trials to test whether resource impacts predicted invasion success. We then related experimental results to field invasion patterns. Compared to exotic herbaceous species, native shrubs were associated with greater resource depletion of key resources: light, soil water (at multiple depths), and soil inorganic nitrogen (particularly at depth). In invasion trials, natives resisted invasion by the exotics, as resource depletion measures would predict. However, these results did not follow long-term natural invasion patterns indicating that these exotic species invade areas once dominated by native shrubland. Applying our results to the invasion framework, we conclude that disturbance, or a similar mechanism causing resources to fluctuate, is needed for exotics to invade CSS habitats. This resource-based comparative analysis of invasion mechanisms can point out important processes and help suggest effective management actions.     

Regional analysis of the impacts of climate change on cheatgrass invasion shows potential risk and opportunity

Interactions between climate change and non-native invasive species may combine to increase invasion risk to native ecosystems. Changing climate creates risk as new terrain becomes climatically suitable for invasion. However, climate change may also create opportunities for ecosystem restoration on invaded lands that become climatically unsuitable for invasive species. Here, I develop a bioclimatic envelope model for cheatgrass ( Bromus tectorum ), a non-native invasive grass in the western US, based on its invaded distribution. The bioclimatic envelope model is based on the Mahalanobis distance using the climate variables that best constrain the species' distribution. Of the precipitation and temperature variables measured, the best predictors of cheatgrass are summer, annual, and spring precipitation, followed by winter temperature. I perform a sensitivity analysis on potential cheatgrass distributions using the projections of 10 commonly used atmosphere–ocean general circulation models (AOGCMs) for 2100. The AOGCM projections for precipitation vary considerably, increasing uncertainty in the assessment of invasion risk. Decreased precipitation, particularly in the summer, causes an expansion of suitable land area by up to 45%, elevating invasion risk in parts of Montana, Wyoming, Utah, and Colorado. Conversely, increased precipitation reduces habitat by as much as 70%, decreasing invasion risk. The strong influence of precipitation conditions on this species' distribution suggests that relying on temperature change alone to project future change in plant distributions may be inadequate. A sensitivity analysis provides a framework for identifying key climate variables that may limit invasion, and for assessing invasion risk and restoration opportunities with climate change.     

The Disturbed Resource-Flux Invasion Matrix: A New Framework for Patterns of Plant Invasion

Attempts to classify certain habitats as vulnerable to invasion or plant traits as invasive have met with limited success and applicability. Clearly, not all plant invaders are able to exploit all habitats and not all habitats are equally susceptible to invasion. Here we argue that it is critical for a successful model for invasions to incorporate both environmental and species traits and present just such a framework. Although disturbance has been targeted as a crucial event which renders habitats vulnerable to invasion, disturbances are often integral parts of ecosystems (e.g. floods, tree-falls, fire, etc.) and are not always associated with invasion events. We argue that disturbances that are associated with invasions alter historical patterns of turnover, or flux, of resources in an ecosystem. Given this perspective on the relationship between invasions and disturbances, and the need to integrate species traits with those of invaded ecosystems, we have developed an approach to characterize plant invasion patterns that we call the Disturbed Resource-Flux Invasion Matrix or DRIM. This is a 16-cell matrix that classifies habitats by the quality of changes in physical and chemical resource flux either increasing or decreasing flux relative to historical patterns. Within each matrix cell, it is then possible to apply basic ecological principles to target species traits that can facilitate successful invasion of habitats experiencing that particular kind of disturbance. We present examples from the literature of how habitats and species can be classified according to the DRIM, and demonstrate the application of this theoretical model.     

Life-history traits of non-native fishes in Iberian watersheds across several invasion stages: a first approach

Freshwater ecosystems are seriously imperiled by the spread of non-native fishes thus establishing profiles of their life-history characteristics is an emerging tool for developing conservation and management strategies. We did a first approach to determine characteristics of successful and failed non-native fishes in a Mediterranean-climate area, the Iberian Peninsula, for three stages of the invasion process: establishment, spread and integration. Using general linear models, we established which characteristics are most important for success at each invasion stage. Prior invasion success was a good predictor for all the stages of the invasion process. Biological variables relevant for more than one invasion stage were maximum adult size and size of native range. Despite these common variables, all models produced a different set of variables important for a successful invasion, demonstrating that successful invaders have a combination of biological traits that may favor success at all invasion stages. However, some differences were found in relation to published studies on fish invasions in other Mediterranean-climate areas, suggesting that characteristics of the recipient ecosystem are as relevant as the characteristics of the invading species.     

Twenty‐five years of plant community dynamics and invasion in New Zealand tussock grasslands

Understanding how plant communities respond to plant invasions is important both for understanding community structure and for predicting future ecosystem change. In a system undergoing intense plant invasion for 25 years, we investigated patterns of community change at a regional scale. Specifically, we sought to quantify how tussock grassland plant community structure had changed and whether changes were related to increases in plant invasion. Frequency data for all vascular plants were recorded on 124, permanent transects in tussock grasslands across the lower eastern South Island of New Zealand measured three times over a period of 25 years. Multivariate analyses of species richness were used to describe spatial and temporal patterns in the vegetation. Linear mixed‐effects models were used to relate temporal changes in community structure to the level and rate of invasion of three dominant invasive species in the genus Hieracium while accounting for relationships with other biotic and abiotic variables. There was a strong compositional gradient from exotic‐ to native‐dominated plant communities that correlated with increasing elevation. Over the 25 years, small‐scale species richness significantly decreased and then increased again; however, these changes differed in different plant communities. Exotic species frequency consistently increased on some transects and consistently declined on others. Species richness changes were correlated with the level of Hieracium invasion and abiotic factors, although the relationship with Hieracium changed from negative to positive over time. Compositional changes were not related to measured predictors. Our results suggest that observed broad‐scale fluctuations in species richness and community composition dynamics were not driven by Hieracium invasion. Given the relatively minor changes in community composition over time, we conclude that there is no evidence for widespread degradation of these grasslands over the last 25 years. However, because of continuing weed invasion, particularly at lower elevations, impacts may emerge in the longer term.     

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 nitrogen cycling by modifying the soil nitrifying community

Plant invasions have dramatic aboveground effects on plant community composition, but their belowground effects remain largely uncharacterized. Soil microorganisms directly interact with plants and mediate many nutrient transformations in soil. We hypothesized that belowground changes to the soil microbial community provide a mechanistic link between exotic plant invasion and changes to ecosystem nutrient cycling. To examine this possible link, monocultures and mixtures of exotic and native species were maintained for 4 years in a California grassland. Gross rates of nitrogen (N) mineralization and nitrification were quantified with ¹⁵N pool dilution and soil microbial communities were characterized with DNA‐based methods. Exotic grasses doubled gross nitrification rates, in part by increasing the abundance and changing the composition of ammonia‐oxidizing bacteria in soil. These changes may translate into altered ecosystem N budgets after invasion. Altered soil microbial communities and their resulting effects on ecosystem processes may be an invisible legacy of exotic plant invasions.     

Biotic interactions limit species richness in an alpine plant community,especially under experimental warming

The determinants of local species richness in plant communities have been the subject of much debate. Is species richness the result of stochastic events such as dispersal processes, or do local environmental filters sort species into communities according to their ecological niches? Recent studies suggest that these two processes simultaneously limit species richness, although their relative importance may vary in space and time. Understanding the limiting factors for species richness is especially important in light of the ongoing global warming, as new species establish in resident plant communities as a result of climate‐driven migration. We examined the relative importance of dispersal and environmental filtering during seedling recruitment and plant establishment in an alpine plant community subjected to seed addition and long‐term experimental warming. Seed addition increased species richness during the seedling recruitment stage, but this initial increase was cancelled out by a corresponding decrease in species richness during plant establishment, suggesting that environmental filters limit local species richness in the long term. While initial recruitment success of the sown species was related to both abiotic and biotic factors, long‐term establishment was controlled mainly by biotic factors, indicating an increase in the relative importance of biotic interactions once plants have germinated in a microhabitat with favourable abiotic conditions. The relative importance of biotic interactions also seemed to increase with experimental warming, suggesting that increased competition within the resident vegetation may decrease community invasibility as the climate warms.     

Differential invasion success of salmonids in southern Chile: patterns and hypotheses

Biological invasions create complex ecological and societal issues worldwide. Most of the knowledge about invasions comes only from successful invaders, but less is known about which processes determine the differential success of invasions. In this review, we develop a framework to identify the main dimensions driving the success and failure of invaders, including human influences, characteristics of the invader, and biotic interactions. We apply this framework by contrasting hypotheses and available evidence to explain variability in invasion success for 12 salmonids introduced to Chile. The success of Oncorhynchus mykiss and Salmo trutta seems to be influenced by a context-specific combination of their phenotypic plasticity, low ecosystem resistance, and propagule pressure. These well-established invaders may limit the success of subsequently introduced salmonids, with the possible exception of O. tshawytscha, which has a short freshwater residency and limited spatial overlap with trout. Although propagule pressure is high for O. kisutch and S. salar due to their intensive use in aquaculture, their lack of success in Chile may be explained by environmental resistance, including earlier spawning times than in their native ranges, and interactions with previously established and resident Rainbow Trout. Other salmonids have also failed to establish, and they exhibit a suite of ecological traits, environmental resistance, and limited propagule pressure that are variably associated with their lack of success. Collectively, understanding how the various drivers of invasion success interact may explain the differential success of invaders and provide key guidance for managing both positive and negative outcomes associated with their presence.     

Methods to test the interactive effects of drought and plant invasion on ecosystem structure and function using complementary common garden and field experiments

Abiotic global change drivers affect ecosystem structure and function, but how they interact with biotic factors such as invasive plants is understudied. Such interactions may be additive, synergistic, or offsetting, and difficult to predict. We present methods to test the individual and interactive effects of drought and plant invasion on native ecosystems. We coupled a factorial common garden experiment containing resident communities exposed to drought (imposed with rainout shelters) and invasion with a field experiment where the invader was removed from sites spanning a natural soil moisture gradient. We detail treatments and their effects on abiotic conditions, including soil moisture, light, temperature, and humidity, which shape community and ecosystem responses. Ambient precipitation during the garden experiment exceeded historic norms despite severe drought in prior years. Soil moisture was 48% lower in drought than ambient plots, but the invader largely offset drought effects. Additionally, temperature and light were lower and humidity higher in invaded plots. Field sites spanned up to a 10‐fold range in soil moisture and up to a 2.5‐fold range in light availability. Invaded and resident vegetation did not differentially mediate soil moisture, unlike in the garden experiment. Herbicide effectively removed invaded and resident vegetation, with removal having site‐specific effects on soil moisture and light availability. However, light was generally higher in invader‐removal than control plots, whereas resident removal had less effect on light, similar to the garden experiment. Invasion mitigated a constellation of abiotic conditions associated with drought stress in the garden experiment. In the field, where other factors co‐varied, these patterns did not emerge. Still, neither experiment suggested that drought and invasion will have synergistic negative effects on ecosystems, although invasion can limit light availability. Coupling factorial garden experiments with field experiments across environmental gradients will be effective for predicting how multiple stressors interact in natural systems.     

How do introduction characteristics influence the invasion success of Mediterranean alien plants?

Invasive plant species are becoming increasingly widespread following accelerated anthropogenic activity in the Mediterranean region. Humans have played a central role in the expansion process, and it is important to incorporate such considerations into management plans. Using generalized linear models, our first aim was to describe how the invasion success of 862 prominent alien plant species on Mediterranean islands is related to characteristics of the introduction process: introduction frequency, date and region of origin, range size and purpose of import. The importance of each was measured by the numbers of species present and their average invasiveness. The main findings were: (a) accidental imports and ornamentals accounted for a high proportion of all aliens, although neither group had particularly high average invasiveness; (b) introduction frequency had a comparatively modest influence, with the most commonly-introduced species naturalized only three times more widely than those rarely-introduced; (c) rates of species introduction appear to have increased dramatically in the last century, although aliens which have been present in the region for more than 200 years were most widespread, indicating that it may be centuries before some species fill their potential range; (d) there were small tendencies for successful invaders to originate in the Neotropics or in regions with Mediterranean climate biomes and to have large range sizes. Our second aim was to determine whether the number or average invasiveness of species introduced via a given pathway had the most influence on the overall probability of invasion on a given island. An elasticity analysis suggested that the number of species was substantially the best predictor of the two. This finding arises largely because invasion events are rare and remain unpredictable, and has significant implications for assessing invasion risk. We discuss how substantial sources of error and intrinsic variability in invasiveness within species groups limit the potential for developing accurate risk models.     

A functional trait perspective on plant invasion

Background and Aims

Global environmental change will affect non-native plant invasions, with profound potential impacts on native plant populations, communities and ecosystems. In this context, we review plant functional traits, particularly those that drive invader abundance (invasiveness) and impacts, as well as the integration of these traits across multiple ecological scales, and as a basis for restoration and management.

Scope

We review the concepts and terminology surrounding functional traits and how functional traits influence processes at the individual level. We explore how phenotypic plasticity may lead to rapid evolution of novel traits facilitating invasiveness in changing environments and then ‘scale up’ to evaluate the relative importance of demographic traits and their links to invasion rates. We then suggest a functional trait framework for assessing per capita effects and, ultimately, impacts of invasive plants on plant communities and ecosystems. Lastly, we focus on the role of functional trait-based approaches in invasive species management and restoration in the context of rapid, global environmental change.

Conclusions

To understand how the abundance and impacts of invasive plants will respond to rapid environmental changes it is essential to link trait-based responses of invaders to changes in community and ecosystem properties. To do so requires a comprehensive effort that considers dynamic environmental controls and a targeted approach to understand key functional traits driving both invader abundance and impacts. If we are to predict future invasions, manage those at hand and use restoration technology to mitigate invasive species impacts, future research must focus on functional traits that promote invasiveness and invader impacts under changing conditions, and integrate major factors driving invasions from individual to ecosystem levels.     

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.     

The perfect storm: Weed invasion and intense storms in tropical forests

Climate change scientists predict an increased intensity of storms (cyclones, hurricanes and typhoons) in the future. Intense storms facilitate plant invasion by increasing resource availability, reducing competition and increasing opportunities for propagule dispersal. We document here the state of current understanding about the response of invasive plant species to intense storms and suggest that the structure and function of forests in storm‐prone regions may be much altered in the future as a result of weed invasion. Intense storms provide a large spatial and temporal window of opportunity for invasion and empirical research demonstrates growth and recruitment rates of invasive species increase following such events, and they spread readily. In particular, lianas and woody invasive species that are shade tolerant and recruit from the seedling layer may constitute the greatest threat to tropical forests following storm events. Forests persisting in fragmented landscapes will be exposed to some of the most severe consequences of intense storms and subsequent weed invasion. In storm‐prone regions, forests of the future are likely to experience a decrease in diversity of native species and homogenization of communities at landscape and regional scales, slower rates of forest succession, increasing degradation of forest fragments and ultimately a decrease in ecosystem function.     

植物功能性状与外来植物入侵

揭示影响外来植物入侵性的功能性状及其生态机制是入侵植物生态学的核心任务之一。本文综述了植物功能性状与外来植物入侵性的研究进展, 通过分析植物功能性状对外来植物入侵的贡献以及外来植物的不同入侵阶段对其功能性状的需求, 探讨植物功能性状与外来植物入侵的相关性及其入侵机理。迄今研究较多的影响外来植物入侵性的功能性状主要包括形态性状、生长性状、生理性状、繁殖性状、种子性状、克隆性状、表型可塑性和遗传变异等。这些功能性状对外来植物入侵的贡献随着入侵阶段的不同而变化。在传播到达阶段, 种子性状对入侵具有重要影响; 在定居建群阶段, 与植物抗逆性和适应性相关的生理性状和繁殖性状发挥主要作用; 在扩散入侵阶段, 克隆性状和影响植物竞争能力的生理性状对植物成功入侵具有重要贡献。由于植物入侵性是其功能性状和环境因素互作的结果, 且功能性状的作用随环境因素和入侵阶段不同而异, 因此, 结合外来植物入侵阶段, 并考虑功能性状与环境因子的互作, 是入侵生物学中植物功能性状研究的发展趋势。     

Assessing the level of plant invasion: A multi-scale approach based on vegetation plots

Recent studies highlight the importance of selecting the appropriate scale and indices of invasion level for evaluating the abundance and impact of alien plants. Our survey considers the use of vegetation plot databases compared with floristic checklists to address invasion patterns regarding alien–native relationships across vegetation types by means of a multi-scale approach. We analysed the alien–native richness relationship in 1077 vegetation plots from the Basque Country (N. Spain) at ecosystem level and phytosociological class and alliance levels. According to our results, the alien species richness (Alo)–native species richness (Nat) relationship is variable and depends not only on the scale but also on the vegetation type. In contrast with other multi-scale approaches, no negative correlation has been detected at any studied level. The strong correlation existing between plot number and cumulative Alo and cumulative Nat highlights the constraints of using checklists to generalize invasion patterns. Our results demonstrate that the combined use of both relative alien species richness and relative alien species cover facilitates the understanding of invasion patterns across plant communities at different scales. In addition to climate, disturbance and propagule pressure, habitat type proved to be an important filter for alien species, capable of explaining such patterns.     

Guild‐specific shifts in visitation rates of frugivores with habitat loss and plant invasion

Habitat loss and plant invasions are two major drivers of global change in subtropical and tropical ecosystems. Both lead to a loss of biodiversity and alter species interactions, which may imperil vital ecosystem processes such as seed dispersal by frugivores. Reponses of frugivores to disturbance are often linked to their specialization on certain habitats or resources. Yet, it is poorly understood how habitat loss and plant invasion structure interactions between plants and different habitat or feeding guilds. Here we investigated whether visitation rates of frugivores change guild‐specifically with increasing habitat loss and invasion level in a heterogeneous subtropical landscape. In 756 h of observations, we recorded 1446 plant–frugivore interactions among 18 plant species and 42 avian frugivore species. Visitation rates of forest specialists decreased with increasing habitat loss, but not with changes in invasion level. In contrast forest generalists and forest visitors were unaffected by either driver. Similarly, obligate frugivores that overall showed a generalized fruit choice were unaffected by habitat loss and changes in invasion level. Contrary, visitation rates of specialized partial and opportunistic frugivores decreased with higher invasion level. Importantly, the negative effect of plant invasion on partial frugivores was more pronounced as habitat loss in the same study site increased, indicating a synergistic effect of the two drivers. The implications of our study are twofold: first, frugivores respond guild‐specifically to habitat loss and plant invasion. Thereby forest dependency is mainly related to habitat loss, and degree of frugivory mainly related to plant invasion. Forest generalists and obligate frugivores in turn may play a key‐role for forest regeneration in disturbed forest landscapes. Second, particularly frugivores with a specialized fruit choice may be threatened by synergistic effects between habitat loss and plant invasion.     

Rhizosphere microbiome functional diversity and pathogen invasion resistance build up during plant development

The rhizosphere microbiome is essential for plant growth and health, and numerous studies have attempted to link microbiome functionality to species and trait composition. However, to date little is known about the actual ecological processes shaping community composition, complicating attempts to steer microbiome functionality. Here, we assess the development of microbial life history and community-level species interaction patterns that emerge during plant development. We use microbial phenotyping to experimentally test the development of niche complementarity and life history traits linked to microbiome performance. We show that the rhizosphere microbiome assembles from pioneer assemblages of species with random resource overlap into high-density, functionally complementary climax communities at later stages. During plant growth, fast-growing species were further replaced by antagonistic and stress-tolerant ones. Using synthetic consortia isolated from different plant growth stages, we demonstrate that the high functional diversity of ‘climax’ microbiomes leads to a better resistance to bacterial pathogen invasion. By demonstrating that different life-history strategies prevail at different plant growth stages and that community-level processes may supersede the importance of single species, we provide a new toolbox to understand microbiome assembly and steer its functionality at a community level.     

Predicting ecosystem shifts requires new approaches that integrate the effects of climate change across entire systems

Most studies that forecast the ecological consequences of climate change target a single species and a single life stage. Depending on climatic impacts on other life stages and on interacting species, however, the results from simple experiments may not translate into accurate predictions of future ecological change. Research needs to move beyond simple experimental studies and environmental envelope projections for single species towards identifying where ecosystem change is likely to occur and the drivers for this change. For this to happen, we advocate research directions that (i) identify the critical species within the target ecosystem, and the life stage(s) most susceptible to changing conditions and (ii) the key interactions between these species and components of their broader ecosystem. A combined approach using macroecology, experimentally derived data and modelling that incorporates energy budgets in life cycle models may identify critical abiotic conditions that disproportionately alter important ecological processes under forecasted climates.     

Quantifying levels of biological invasion: towards the objective classification of invaded and invasible ecosystems

Biological invasions are a global phenomenon that threatens biodiversity, and few, if any, ecosystems are free from alien species. The outcome of human‐mediated introductions is affected by the invasiveness of species and invasibility of ecosystems, but research has primarily focused on defining, characterizing and identifying invasive species; ecosystem invasibility has received much less attention. A prerequisite for characterizing invasibility is the ability to compare levels of invasion across ecosystems. In this paper, we aim to identify the best way to quantify the level of invasion by nonnative animals and plants by reviewing the advantages and disadvantages of different metrics. We explore how interpretation and choice of these measures can depend on the objective of a study or management intervention. Based on our review, we recommend two invasion indices and illustrate their use by applying them to two case studies. Relative alien species richness and relative alien species abundance indicate the contribution that alien species make to a community. They are easy to measure, can be applied to various taxa, are independent of scale and are comparable across regions and ecosystems, and historical data are often available. The relationship between relative alien richness and abundance can indicate the presence of dominant alien species and the trajectory of invasion over time, and can highlight ecosystems and sites that are heavily invaded or especially susceptible to invasion. Splitting species into functional groups and examining invasion patterns of transformer species may be particularly instructive for gauging effects of alien invasion on ecosystem structure and function. Establishing standard, transparent ways to define and quantify invasion level will facilitate meaningful comparisons among studies, ecosystem types and regions. It is essential for progress in ecology and will help guide ecosystem restoration and management.