Corresponding author: Luke J. Potgieter ( lukepotgieter2@gmail.com ) Academic editor: John Ross Wilson
© 2020 Luke J. Potgieter, Marc W. Cadotte.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Potgieter LJ, Cadotte MW (2020) The application of selected invasion frameworks to urban ecosystems. In: Wilson JR, Bacher S, Daehler CC, Groom QJ, Kumschick S, Lockwood JL, Robinson TB, Zengeya TA, Richardson DM. NeoBiota 62: 365-386. https://doi.org/10.3897/neobiota.62.50661
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Urbanization is a major driver of global change. Profound human-mediated changes to urban environments have provided increased opportunities for species to invade. The desire to understand and manage biological invasions has led to an upsurge in frameworks describing the mechanisms underpinning the invasion process and the ecological and socio-economic impacts of invading taxa. This paper assesses the applicability of three commonly used invasion frameworks to urban ecosystems. The first framework describes the mechanisms leading to invasion; the second and third frameworks assess individual species, and their associated environmental and socio-economic impacts, respectively.
In urban areas, the relative effectiveness of the barriers to invasion is diminished (to varying degrees) allowing a greater proportion of species to move through each subsequent invasion stage, i.e. “the urban effect” on invasion. Impact classification schemes inadequately circumscribe the full suite of impacts (negative and positive) associated with invasions in urban areas. We suggest ways of modifying these frameworks to improve their applicability to understanding and managing urban invasions.
Biological invasions, framework, impacts, invasive alien plants, urban ecosystems
Urbanization is now a major agent of change for social, economic, and ecological systems (
Basic understanding of invasion and management frameworks needs to be evaluated in urban ecosystems for several reasons. Firstly, urban areas are directly shaped by human activities that can transport and foster alien species. Secondly, the beliefs, priorities, and concerns of human populations in urban areas directly impact the creation and implementation of management policies for invasive alien species. Many frameworks exist to classify biological invasions, for example, by pathways of dispersal (
We selected three invasion frameworks: the Unified Framework for Biological Invasions (
The frequently employed and commonly cited Unified Framework for Biological Invasions proposed by
The unified framework for biological invasions by
The EICAT scheme classifies alien taxa in terms of the magnitude of their impacts on the environment. It relies on published evidence of impacts via an exhaustive literature search to identify all published literature on the impacts of each alien taxon under assessment. Each impact record for an alien taxon is classified into one of five sequential categories in ascending order of impact, from “Minimal Concern” to “Massive” depending on the level of biological organization (individual, population, community or ecosystem) impacted. This includes the mechanisms (and magnitude) of impact for each taxon, a confidence score for each record, and additional information including the spatial location at which the impact is realized, and which native species are impacted.
Based on the capability approach from welfare economics, the SEICAT scheme presents a system for ranking and comparing the negative impacts of alien taxa on human well-being. The process also relies on published evidence to classify alien taxa based on changes in human activities that result from their impacts. By focusing on changes in people’s activities, SEICAT captures impacts of introduced species on human well-being that systems based on monetary values cannot. The scheme defines eight categories into which alien taxa can be classified according to the magnitude of changes in people’s realized activities. This classification is analogous to the EICAT and IUCN Red List schemes (
Urban environments result in novel ecological patterns and processes, and dynamics (
Through sustained human facilitation in cities, intensive cultivation and repeated introductions of many alien species, biotic and abiotic manipulation, humanity’s preference for traits associated with high reproductivity (e.g. large showy flowers, colourful fruits), greater number and diversity of pathways and vectors that can facilitate the movement of alien plants, and increased levels of human-mediated habitat disturbance, collectively increase the likelihood of establishment and spread of alien plant species in urban areas.
Introduced alien species must pass through a series of barriers before naturalizing in their new environment (
The perennial herbaceous vine Vincetoxicum rossicum (Kleopow) Barbar. (Asclepiadaceae; syn. Cynanchum rossicum), also known as dog-strangling vine, is a prolific invader in the city of Toronto, Canada. Below we describe how V. rossicum overcame the barriers to invasion defined by Geographic barrier Vincetoxicum rossicum was directly imported into the city of Toronto in the late 1800s from the Ukraine and planted as an ornamental and for erosion stabilization in several locations in the city ( Captivity barrier There was a significant lag period before V. rossicum became a species of concern, but by the 1980s, it had become widespread and abundant in Toronto, especially in urban woodlots and meadows. Its ornamental use, likely facilitated by the nursery trade, provided opportunities for its escape from captivity ( Survival barrier Gardeners allowed V. rossicum to escape its natural enemies, resulting in much improved fitness. It demonstrates wide environmental tolerance to variations in light intensity and soil moisture ( Reproduction barrier Seeds of V. rossicum are often polyembryonic, giving rise to two, three and (rarely) four seedlings ( Dispersal barrier Vincetoxicum rossicum possesses feathery pappus-covered seeds that are easily carried by wind, and with cars and trains creating air currents, seeds can travel long distances along linear corridors ( Environmental barrier Vincetoxicum rossicum successfully establishes and survives across a range of disturbance regimes, and improved growth has been shown in more disturbed habitats ( |
Impacts
Having successfully overcome these barriers, V. rossicum negatively impacts native biodiversity – it reduces the diversity of plant and other trophic levels by excluding species with certain traits ( |
Figure B1. An urban site in Toronto, Canada, where the invasive Vincetoxicum rossicum forms a monoculture in open and understory habitats (photo credit: LJ Potgieter). |
The barriers to invasion proposed by Blackburn and colleagues (2011) in the context of urban areas, and the underlying mechanisms driving invasion in urban areas. *Indicates potential mechanisms strengthening the barriers to successful establishment of alien plants in urban areas.
Barriers to invasion | Urban effect on barriers | Facilitating mechanisms | References |
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Geographic | Significant opportunities for alien species introduction | • Urban areas are often the first entry point for newly introduced alien species. |
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• Cities serve as transportation and trade hubs, and are highly interconnected, thus increasing the movement of species globally. | |||
Captivity or cultivation | Significant opportunities to escape captivity / not relevant | • Horticultural activities result in a major pathway for the repeated importation of many alien plants into urban areas (e.g. gardens which serve as regular sources of alien plant propagules). |
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• Small urban settlements act as launching sites for plant invasions into natural areas. | |||
• Alien species introduced unintentionally by humans directly into the new environment. | |||
Survival | Significant potential for survival opportunities, unlike neighbouring, less disturbed areas | • Humans manipulate abiotic and biotic conditions to improve the survivorship of alien plant species. For example, reducing competition, increasing nutrient and water input, and altering soil pH. |
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• Urban heat island effect can provide suitable conditions for more heat-tolerant alien plants. | |||
• Selective breeding and dissemination of ‘urban suitable’ genotypes. | |||
• Intra- and inter-specific hybridization can create novel, potentially invasive genotypes. | |||
• *Repeated introductions of the same species at a location (propagule pressure) can increase the likelihood of successful establishment. | |||
Reproduction | Potential opportunities for reproductive success for some alien plants | • Habitat fragmentation selects for species with high seed production. |
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• Selective breeding or intraspecific hybridization of cultivars can increase reproductive success. | |||
• Longer growing season and earlier flowering and seeding for alien plants in response to urban climates. | |||
• Cultivar selection for desirable traits can inadvertently result in greater fecundity. | |||
• *Habitat fragmentation reduces the size and increases the isolation of urban plant populations, increasing extinction risk and reducing pollination. | |||
Dispersal | Potential for dispersal opportunities unlike neighbouring less disturbed areas | • Alien plants which possess traits conducive to effective dispersal through prominent urban dispersal pathways are more likely to proliferate, such as wind dispersal by vehicle traffic. |
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• *Many alien plants do not possess the appropriate suite of traits required for efficient dispersal in urban areas. | |||
• *Increased propagule pressure can enable alien species to overcome urban dispersal barriers despite poor dispersal abilities. | |||
Environmental | Environmental conditions provide potential opportunities for alien plant spread | • High level of heterogeneity and disturbance results in frequent colonization opportunities for alien plants. |
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• *Habitat fragmentation can limit the spread from established alien plant populations. | |||
• *High levels of disturbance can also act as a barrier to the establishment of some plant species. |
Urban areas as hotspots of globalization provide significant opportunities for alien species’ introduction
Human activities are progressively weakening biogeographical barriers to dispersal (
Urban areas have more alien taxa in captivity and cultivation, and so greater propagule pressure.
Invasibility is strongly influenced by propagule pressure (
The effect of propagule pressure on invasibility is also apparent in smaller urban settlements which can act as launching sites for plant invasions into rural (
The captivity and cultivation barrier might also be skipped entirely by alien species which are introduced unintentionally by humans (e.g. as a contaminant (stowaway) of a commodity) (
Human intervention improves the survivorship of alien species
Inherent features of the urban landscape could serve to select for species that are able to persist under more stressful conditions or rapidly take advantage of the resource-rich conditions. For example, higher temperatures associated with urban areas (i.e. urban heat island effect) might select for the establishment of alien plants preadapted to warmer conditions than the natural environment provides (
Below the species’ level, genotypes proving successful in urban areas are selectively bred (e.g. for flowering, architecture, foliage and for disease-resistance traits) and are widely disseminated through plant exchanges and the horticultural trade (regulated and unregulated) (
While the survival barrier in urban areas might be high for some species, the failure of individuals or populations to survive is not just a consequence of the environment, for subsequent human-mediated introductions of the same species at that location could succeed (
Urbanization can enhance reproductive success of some alien plants
Urban habitats are highly fragmented, thus reducing the size and increasing the isolation of urban plant populations, resulting in a decline in pollinator services and ultimately lowering reproductive success of plants (
While selective breeding or intraspecific hybridization of cultivars can reduce invasive potential (
The complex network of dispersal pathways and vectors in urban areas facilitates the movement of some alien plants.
Urban areas comprise a complex network of pathways and vectors that can facilitate plant movement within the urban matrix and into surrounding natural areas (
Alien plant species with fleshy fruits are more likely to expand their range in urban environments because of their ability to utilize bird dispersal (
Urban areas provide environmental conditions favourable for the spread of some alien plants
In urban areas, abiotic conditions such as climate, land use, pollution, and nutrient loads are dramatically altered through human intervention (
Habitat fragmentation greatly increases the amount of edge habitat, which can increase the susceptibility of vegetation patches to disturbances (
The influence of management
While the relative importance of the barriers to reproduction, dispersal, and environment for determining invasion success can be greater in urban areas compared to natural areas (Table
A taxon with a high ecological impact in a rural or natural environment will not necessarily have the same impact in an urban environment, or impact at the same spatial and temporal scales. Generally, the magnitude of an alien species’ impact (which can be highly variable and context-dependent,
Combining the ICAT Schemes might achieve a more comprehensive assessment of the negative ecological and socio-economic impacts from an invasion. Both schemes explicitly focus on deleterious impacts and do not set out to weigh deleterious against beneficial impacts to determine the net value of an introduction of an alien taxon (cf.
Robust and comparable data on the impacts of most alien species are still lacking, and in many cases, uncertainties in impact assessments remain significant (
With ongoing dissemination of alien plant propagules, there are likely to be significant time-lags before any impact is realized (
Urban areas present increased economic and social opportunities for people compared to rural or natural areas (
With an abundance of invasion frameworks and a growing body of literature exploring the many facets of invasions in urban areas, it is important to consider whether current invasion frameworks apply to urban systems.
Biological invasions represent a complex societal issue. Consequently, impact assessments and subsequent management decisions should include input from a wide range of stakeholders to elucidate the positive and negative effects of invasive alien plants in urban areas (e.g.
Urban areas have a complex network of dispersal pathways and vectors that can facilitate the movement of alien plants. Existing pathway frameworks (e.g.
Urbanization provides insights into how species will respond and interact under future global change scenarios (
This paper emerged from a workshop on ‘Frameworks used in Invasion Science’ hosted by the DSI-NRF Centre of Excellence for Invasion Biology in Stellenbosch, South Africa, 11–13 November 2019, that was supported by the National Research Foundation of South Africa and Stellenbosch University. Funding was provided by the Connaught Global Challenges Award, the Office of the Vice-President International, the School of Graduate Studies and the HKU-U of T Strategic Partnership Fund at the University of Toronto, and the Office of the Vice-Principal Research at the University of Toronto Scarborough. MWC wishes to further acknowledge funding from the Natural Sciences and Engineering Research Council of Canada (#386151).