In a review published in New Phytologist (2007), Kathleen Theoharides and Jeffrey Dukes examine four stages of invasion as they relate to alien (i.e. introduced or non-indigenous) plant species. In part one we discussed transport and colonization, in which species must survive being transported long distances and then take root and reach maturity in an unfamiliar location. Alien plant species don’t become invasive until they have reached the last two stages: establishment and landscape spread. Removal of the species upon reaching these stages is no easy task. Luckily, introduced species have a few barriers to overcome before this point.
An established population is one that is self-sustaining and expanding. Environmental conditions may be a limiting factor, as they were during colonization, but the main constraints at this stage are “biotic filters.” Theoharides and Dukes define these as “barriers to invasion created by the actions or presence of living organisms.” Through competition for various resources, as well as herbivory and disease, neighboring organisms affect the survival, growth, and reproduction of introduced plant species. Thus, “traits that enhance competitive performance, reduce niche overlap between [introduced species] and natives, or increase enemy resistance may be most important during establishment… Other advantageous traits include secondary chemical compounds that deter herbivores, ‘novel weapons’ such as root exudates that negatively impact other plants, fast growth, and high fecundity.”
Plants compete for light, moisture, and soil nutrients, as well as for pollinators and seed dispersers. Competition inhibits the establishment of invaders when neighboring plants consume available resources more efficiently. Introduced plants risk being outcompeted by plants that are of the same functional type (plants that are “morphologically, phenologically, and physiologically similar”). They also risk competition by a single dominant species (or group of similar species) or by “an assemblage of species with different traits.” As a general rule, plant communities with greater diversity are more resistant to invasion.Two hypotheses postulate the success of some plant invaders in establishing themselves: the enemy release hypothesis and the evolution of increased competitive ability hypothesis. In the first hypothesis, plant species – having been removed from their native habitat – are freed from their natural enemies and are thus able to allocate more resources to growth and reproduction. The second hypothesis states that, in light of “reduced enemy pressure,” introduced species quickly evolve to allocate resources “from enemy defense to faster growth.” Escape from herbivory and diseases, however, is likely not the only factor in the success of invaders, and much still depends on the competitiveness of the plant and the availability of key resources.
After introduced plants become established, a lag phase generally occurs before landscape spread. This can be a result of a lack of genetic variation, a dearth of suitable habitat, unfavorable environmental conditions, or some combination of the three. New introductions may occur, and the population may continue to adapt and expand. Suitable habitat may be made available, and environmental conditions may shift. In time, landscape spread becomes a possibility.
Landscape spread occurs when multiple populations of a species are connected via long-distance dispersal. At this “metacommunity scale,” populations of an introduced plant species interact across a large area, with each population in a different stage of colonization and establishment. This means that transport, colonization, and establishment are all at play during the landscape spread stage.Dispersal ability and habitat connectivity are key factors in determining the success of an introduced plant species during landscape spread. Long-distance dispersal can occur via wind, water, or animals. Species that depend on animals to spread their seeds rely on specific animals to be present. The seeds of Prunus serotina (black cherry), for example, are dispersed by birds. So, landscape spread is reliant on birds and “roosting trees” where the birds can perch and defacate the seeds. In many cases, “humans also play a large role in intraregional dispersal.”
Habitats vary across the landscape due to a combination of numerous geological and biological processes. The disturbance regime – “the frequency, spatial extent, severity, and intensity of killing events over time” – also helps determine landscape patterns. Natural disturbances, such as fire, weather, and natural disasters, are differentiated from disturbances caused by human activity. Large scale development and disturbance of natural areas by humans disrupts the natural disturbance regime and alters historical landscape patterns. As the authors write, “alterations of the disturbance regime that increase resource availability or change landscape patterns can promote non-indigenous plant species spread by creating favorable patches for colonization and establishment.”
Fragmented landscapes consisting of small patches of natural areas dispersed among large areas of human development are particularly prone to invasion by introduced plant species for many reasons, including increased influx of propagules and a high degree of edge effects (habitat edges have environmental conditions that are generally more prone to invasion than habitat interiors).
Habitat patches can be connected via corridors. It is through these corridors that dispersal can occur between populations in a metacommunity. Corridors connect populations of both introduced and native plant species. However, “native plants often require wide undisturbed corridors of intact habitat, while [introduced plant species] may disperse best through strips of human-disturbed habitat or ‘disturbance corridors.'” The environmental conditions in disturbance corridors and the presence of dispersal agents (including humans and domesticated animals) help facilitate the connectivity of populations of introduced plant species and promote the colonization and establishment of new populations.
In their abstract, Theoharides and Dukes write, “both research and management programs may benefit from employing multiscale and stage approaches to studying and controlling invasion.” With their conclusion they provide a list of potential management strategies for each stage, and they advise employing “natural filters in order to prevent invasion succees.” Examples include reducing habitat fragmentation and edge effects, promoting intact native communities, reducing human disturbances, promoting natural disturbance regimes, and minimizing disturbance corridors.