Invasive species


The term as most often used applies to introduced species that adversely affect the habitats and bioregions they invade economically, environmentally, or ecologically. Such species may be either plants or animals and may disrupt by dominating a region, wilderness areas, particular habitats, or wildland–urban interface land from loss of natural controls (such as predators or herbivores). This includes plant species labeled as exotic pest plants and invasive exotics growing in native plant communities.[3][4][5][6] The European Union defines "Invasive Alien Species" as those that are, firstly, outside their natural distribution area, and secondly, threaten biological diversity.[7][8] The term is also used by land managers, botanists, researchers, horticulturalists, conservationists, and the public for noxious weeds.[9]

The term "invasive" is often poorly defined or very subjective[6] and some broaden the term to include indigenous or "native" species, that have colonized natural areas[6] - for example deer considered by some to be overpopulating their native zones and adjacent suburban gardens in the Northeastern and Pacific Coast regions of the United States.[10]

The definition of "native" is also sometimes controversial. For example, the ancestors of Equus ferus (modern horses) evolved in North America and radiated to Eurasia before becoming locally extinct. Upon returning to North America in 1493 during their hominid-assisted migration, it is debatable as to whether they were native or exotic to the continent of their evolutionary ancestors.[11]

Notable examples of invasive plant species include the kudzu vine, Andean pampas grass, and yellow starthistle. Animal examples include the New Zealand mud snail, feral pigs, European rabbits, grey squirrels, domestic cats, carp and ferrets.[12][13][14]

Invasion of long-established ecosystems by organisms from distant bio-regions is a natural phenomenon, but has been accelerated massively by humans, from their earliest migrations through to the age of discovery, and now international trade.


Scientists include species and ecosystem factors among the mechanisms that, when combined, establish invasiveness in a newly introduced species.

Species based mechanisms


Researchers disagree about the usefulness of traits as invasiveness markers. One study found that of a list of invasive and noninvasive species, 86% of the invasive species could be identified from the traits alone.[15] Another study found invasive species tended to have only a small subset of the presumed traits and that many similar traits were found in noninvasive species, requiring other explanations.[15][16][17] Common invasive species traits include the following:

  • Fast growth
  • Rapid reproduction
  • High dispersal ability
  • Phenotype plasticity (the ability to alter growth form to suit current conditions)
  • Tolerance of a wide range of environmental conditions (Ecological competence)
  • Ability to live off of a wide range of food types (generalist)
  • Association with humans[18]
  • Prior successful invasions[19]








Ecosystem-based mechanisms

In ecosystems, the amount of available resources and the extent to which those resources are used by organisms determines the effects of additional species on the ecosystem. In stable ecosystems, equilibrium exists in the use of available resources. These mechanisms describe a situation in which the ecosystem has suffered a disturbance, which changes the fundamental nature of the ecosystem.[31]


Every species occupies a niche in its native ecosystem; some species fill large and varied roles, while others are highly specialized. Some invading species fill niches that are not used by native species, and they also can create new niches.[33] An example of this type can be found within the Lampropholis delicata species of skink.


One interesting finding in studies of invasive species has shown that introduced populations have great potential for rapid adaptation and this is used to explain how so many introduced species are able to establish and become invasive in new environments. When bottlenecks and founder effects cause a great decrease in the population size and may constrict genetic variation,[35] the individuals begin to show additive variance as opposed to epistatic variance. This conversion can actually lead to increased variance in the founding populations which then allows for rapid adaptive evolution.[36] Following invasion events, selection may initially act on the capacity to disperse as well as physiological tolerance to the new stressors in the environment. Adaptation then proceeds to respond to the selective pressures of the new environment. These responses would most likely be due to temperature and climate change, or the presence of native species whether it be predator or prey.[37] Adaptations include changes in morphology, physiology, phenology, and plasticity.



Traits of invaded ecosystems

In 1958, Charles S. Elton[39] claimed that ecosystems with higher species diversity were less subject to invasive species because of fewer available niches. Other ecologists later pointed to highly diverse, but heavily invaded ecosystems and argued that ecosystems with high species diversity were more susceptible to invasion.[40]


Invasion was more likely in ecosystems that were similar to the one in which the potential invader evolved.[43] Island ecosystems may be more prone to invasion because their species faced few strong competitors and predators, or because their distance from colonizing species populations makes them more likely to have "open" niches.[44] An example of this phenomenon was the decimation of native bird populations on Guam by the invasive brown tree snake.[45] Conversely, invaded ecosystems may lack the natural competitors and predators that check invasives' growth in their native ecosystems.

Invaded ecosystems may have experienced disturbance, typically human-induced.[43] Such a disturbance may give invasive species a chance to establish themselves with less competition from natives less able to adapt to a disturbed ecosystem.[20]



An early human vector occurred when prehistoric humans introduced the Pacific rat (Rattus exulans) to Polynesia.[47]


The arrival of invasive propagules to a new site is a function of the site's invasibility.[48]




Within the aquatic environment


Many marine organisms have the capacity to attach themselves to vessel hulls. Therefore, these organisms are easily transported from one body of water to another and are a significant risk factor for a biological invasion event.[53] Unfortunately, controlling for vessel hull fouling is voluntary and there are no regulations currently in place to manage hull fouling. However, the governments of California and New Zealand have announced more stringent control for vessel hull fouling within their respective jurisdictions.[54]

The other main vector for the transport of non-native aquatic species is ballast water. Ballast water taken up at sea and released in port by transoceanic vessels is the largest vector for non-native aquatic species invasions.[55][56] In fact, it is estimated that 10,000 different species, many of which are non-indigenous, are transported via ballast water each day.[57] Many of these species are considered harmful and can negatively affect their new environment. For example, freshwater zebra mussels, native to the Black, Caspian and Azov seas, most likely reached the Great Lakes via ballast water from a transoceanic vessel.[58] Zebra mussels outcompete other native organisms for oxygen and food, such as algae. Although the zebra mussel invasion was first noted in 1988, and a mitigation plan was successfully implemented shortly thereafter, the plan had a serious flaw or loophole, whereby ships loaded with cargo when they reached the Seaway were not tested because their ballast water tanks were empty. However, even in an empty ballast tank, there remains a puddle of water filled with organisms that could be released at the next port (when the tank is filled with water after unloading the cargo, the ship takes on ballast water which mixes with the puddles and then everything including the living organisms in the puddles is discharged at the next port).[59] Current regulations for the Great Lakes rely on ‘salinity shock’ to kill freshwater organisms left in ballast tanks.[60]

Even though ballast water regulations are in place to protect against potentially invasive species, there exists a loophole for organisms in the 10-50 micron size class. For organisms between 10 and 50 microns, such as certain types of phytoplankton, current regulations allow less than 10 cells per milliliter be present in discharge from treatment systems.[61] The discharge gets released when a ship takes on cargo at a port so the discharged water is not necessarily the same as the receiving body of water. Since many species of phytoplankton are less than 10 microns in size and reproduce asexually, only one cell released into the environment could exponentially grow into many thousands of cells over a short amount of time. This loophole could have detrimental effects to the environment. For example, some species in the genus Pseudo-nitzschia are smaller than 10 microns in width and contain domoic acid, a neurotoxin. If toxic Pseudo-nitzschia spp. are alive in ballast discharge and get released into their “new environment” they could cause domoic acid poisoning in shellfish, marine mammals and birds.[62] Fortunately, human deaths related to domoic acid poisoning have been prevented because of stringent monitoring programs that arose after a domoic acid outbreak in Canada in 1987.[62] Ballast water regulations need to be more rigorous to prevent future ramifications associated with the potential release of toxic and invasive phytoplankton.

Another important factor to consider about marine invasive species is the role of environmental changes associated with climate change, such as an increase in ocean temperature. There have been multiple studies suggesting an increase in ocean temperature will cause range shifts in organisms,[63][64] which could have detrimental effects on the environment as new species interactions emerge. For example, Hua and Hwang proposed that organisms in a ballast tank of a ship traveling from the temperature zone through tropical waters can experience temperature fluctuations as much as 20 °C.[65] To further examine the effects of temperature on organisms transported on hulls or in ballast water, Lenz et al. (2018) carried out study where they conducted a double heat stress experiment. Their results suggest that heat challenges organisms face during transport may enhance the stress tolerance of species in their non-native range by selecting for genetically adapted genotypes that will survive a second applied heat stress, such as increased ocean temperature in the founder population.[66] Due to the complexity of climate change induced variations, it is difficult to predict the nature of temperature-based success of non-native species in-situ. Since some studies have suggested increased temperature tolerance of “hijackers” on ships’ hulls or in ballast water, it is necessary to develop more comprehensive fouling and ballast water management plans in an effort to prevent against future possible invasions as environmental conditions continue to change around the world.

Effects of wildfire and firefighting









Invasive species can change the functions of ecosystems. For example, invasive plants can alter the fire regime (cheatgrass, Bromus tectorum), nutrient cycling (smooth cordgrass Spartina alterniflora), and hydrology (Tamarix) in native ecosystems.[68] Invasive species that are closely related to rare native species have the potential to hybridize with the native species. Harmful effects of hybridization have led to a decline and even extinction of native species.[69][70] For example, hybridization with introduced cordgrass, Spartina alterniflora, threatens the existence of California cordgrass (Spartina foliosa) in San Francisco Bay.[71] Invasive species cause competition for native species and because of this 400 of the 958 endangered species under the Endangered Species Act are at risk.[72]


Primary geomorphological effects of invasive plants are bioconstruction and bioprotection. For example, Kudzu Pueraria montana, a vine native to Asia was widely introduced in the southeastern USA in the early 20th century to control soil erosion. While primary effects of invasive animals are bioturbation, bioerosion, and bioconstruction. For example, invasion of Chinese mitten crab Eriocheir sinensis have resulted in higher bioturbation and bioerosion rates.[73]


Some invaders cause negativity affect the economy of the local area. For example, in the Great Lakes Region the sea lamprey is an invasive species that acts as a predator. In its original habitat, the sea lamprey used co-evolution to act as a parasite without killing the host organism. However, in the Great Lakes Region, this co-evolutionary link is absent, so the sea lamprey acts as a predator and can consume up to 40 pounds of fish in its 12-18 month feeding period.[74] Sea lampreys prey on all types of large fish such as lake trout and salmon. The sea lampreys' destructive effects on large fish negatively affect the fishing industry and have helped cause the collapse of the population of some species.[74]

Economic opportunities

Some invasions offer potential commercial benefits. For instance, silver carp and common carp can be harvested for human food and exported to markets already familiar with the product, or processed into pet foods, or mink feed. Water hyacinth can be turned into fuel by methane digesters,[75] and other invasive plants can also be harvested and utilized as a source of bioenergy.[76]


Although most people focus on the negative effects of invasive and non-native species, they can actually be harmless or even beneficial in some cases. Ecosystems thrive because of biodiversity and some need non-native species in order to succeed. There are four major ways that non-natives can be very beneficial for an ecosystem. The first is that they can provide a suitable habitat or food source for other organisms. In areas where a native has become extinct or reached a point that it cannot be restored, non-native species can fill their role. A good example of this is the Tamarisk, a non-native woody plant, and the Southwestern Willow Flycatcher, an endangered bird. 75% of Southwestern Willow Flycatchers were found to nest in these plants and their success was the same as the flycatchers that had nested in native plants. The removal of Tamarisk would be detrimental to Southwestern Willow Flycatcher as their native nesting sites are unable to be restored.

The second way that non-native species can be beneficial is that they act as catalysts for restoration. This is because the presence of non-native species increases the heterogeneity and biodiversity in an ecosystem. This increase in heterogeneity can create microclimates in sparse and eroded ecosystems, which then promotes the growth and reestablishment of native species. Another benefit of non-native species is that they can act as a substitute for an existing ecosystem engineer. In many cases, non-native species can be introduced to fill a niche that had previously been occupied by a native species.

Many non-native species have similar characteristics and functions and can keep an ecosystem functioning properly without collapse. An example of this is the Aldabra giant tortoises, which were introduced on several small islands and have successfully taken over the roles of herbivore and seed disperser. The last benefit of non-native species is that they provided ecosystem services. There are many examples of this. The major one being pollinators. The American Honey bee was introduced in the rainforest to pollinate fragmented landscapes that native species cannot. Also, non-native species can function as biocontrol agents to limit the effects of invasive species. Such as the use of non-native species to control agricultural pests.[77]

Non-native species can have other benefits. Asian oysters, for example, filter water pollutants better than native oysters. They also grow faster and withstand disease better than natives. Biologists are currently considering releasing this mollusk in the Chesapeake Bay to help restore oyster stocks and remove pollution. A recent study by the Johns Hopkins School of Public Health found the Asian oyster could significantly benefit the bay's deteriorating water quality.[78] Additionally, some species have invaded an area so long ago that they have found their own beneficial niche in the environment, a term referred to as naturalisation. For example, L. leucozonium, shown by population genetic analysis to be an invasive species in North America,[79] has become an important pollinator of caneberry as well as cucurbit, apple trees, and blueberry bushes.[80]



In 2005 Chef Bun Lai of Miya's Sushi in New Haven, Connecticut created the first menu dedicated to the idea of using invasive species, during which time half the menus invasive species offerings were conceptual because invasive species were not yet commercially available.[81] Today, Miya’s offers a plethora of invasive species such as Chesapeake blue catfish, Florida lionfish, Kentucky silver carp, Georgia cannonball jellyfish, and invasive edible plants such as Connecticut Japanese knotweed and Autumn olive.[82][83][84][85][86]



Proponents of invasivorism argue that humans have the ability to eat away any species that it has an appetite for, pointing to the many animals which humans have been able to hunt to extinction—such as the Dodo bird, the Caribbean monk seal, and the passenger pigeon. Proponents of invasivorism also point to the success that Jamaica has had in significantly decreasing the population of lionfish by encouraging the consumption of the fish.[91]

In recent years, organizations including Reef Environmental Educational Foundation[92] and the Institute for Applied Ecology, among others, have published cookbooks and recipes that include invasive species as ingredients.[93][94][95]


Economic costs from invasive species can be separated into direct costs through production loss in agriculture and forestry, and management costs. Estimated damage and control cost of invasive species in the U.S. alone amount to more than $138 billion annually.[96] Economic losses can also occur through loss of recreational and tourism revenues.[97] When economic costs of invasions are calculated as production loss and management costs, they are low because they do not consider environmental damage; if monetary values were assigned to the extinction of species, loss in biodiversity, and loss of ecosystem services, costs from impacts of invasive species would drastically increase.[96] The following examples from different sectors of the economy demonstrate the impact of biological invasions.

It is often argued that the key to reducing the costs of invasive species damage and management is early detection and rapid response,[98] meaning that incurring an initial cost of searching for and finding an invasive species and quickly controlling it, while the population is small, is less expensive that managing the invasive population when it is widespread and already causing damage. However, an intense search for the invader is only important to reduce costs in cases where the invasive species is (1) not frequently reintroduced into the managed area and (2) cost effective to search for and find.[99]

Plant industry

Weeds reduce yield in agriculture, though they may provide essential nutrients. Some deep-rooted weeds can "mine" nutrients (see dynamic accumulator) from the subsoil and deposit them on the topsoil, while others provide habitat for beneficial insects or provide foods for pest species. Many weed species are accidental introductions that accompany seeds and imported plant material. Many introduced weeds in pastures compete with native forage plants, threaten young cattle (e.g., leafy spurge, Euphorbia esula) or are unpalatable because of thorns and spines (e.g., yellow starthistle). Forage loss from invasive weeds on pastures amounts to nearly US$1 billion in the U.S. alone.[96] A decline in pollinator services and loss of fruit production has been caused by honey bees infected by the invasive varroa mite. Introduced rats (Rattus rattus and R. norvegicus) have become serious pests on farms, destroying stored grains.[96] The introduction of leaf miner flies, including the American serpentine leaf miner, to California has also caused losses in California's floriculture industry, as the larvae of these invasive species feed on ornamental plants.[100]



Aquaculture is a very common vector of species introductions – mainly of species with economic potential (e.g., Oreochromis niloticus).[102]




Tourism and recreation

Invasive species can impact outdoor recreation, such as fishing, hunting, hiking, wildlife viewing, and water-based activities. They can damage a wide array of environmental services that are important to recreation, including, but not limited to, water quality and quantity, plant and animal diversity, and species abundance.[108] Eiswerth states, "very little research has been performed to estimate the corresponding economic losses at spatial scales such as regions, states, and watersheds". Eurasian watermilfoil (Myriophyllum spicatum) in parts of the US, fill lakes with plants complicating fishing and boating.[109] The very loud call of the introduced common coqui depresses real estate values in affected neighborhoods of Hawaii.[110]


Encroachment of humans into previously remote ecosystems has exposed exotic diseases such as HIV[96] to the wider population. Introduced birds (e.g. pigeons), rodents and insects (e.g. mosquito, flea, louse and tsetse fly pests) can serve as vectors and reservoirs of human afflictions. Throughout recorded history, epidemics of human diseases, such as malaria, yellow fever, typhus, and bubonic plague, spread via these vectors.[39] A recent example of an introduced disease is the spread of the West Nile virus, which killed humans, birds, mammals, and reptiles.[111] The introduced Chinese mitten crabs are carriers of Asian lung fluke.[58] Waterborne disease agents, such as cholera bacteria (Vibrio cholerae), and causative agents of harmful algal blooms are often transported via ballast water.[112] Invasive species and accompanying control efforts can have long term public health implications. For instance, pesticides applied to treat a particular pest species could pollute soil and surface water.[96]


Biotic invasion is considered one of the five top drivers for global biodiversity loss and is increasing because of tourism and globalization.[113] This may be particularly true in inadequately regulated fresh water systems, though quarantines and ballast water rules have improved the situation.[114]


Genetic pollution

Native species can be threatened with extinction[117] through the process of genetic pollution. Genetic pollution is unintentional hybridization and introgression, which leads to homogenization or replacement of local genotypes as a result of either a numerical or fitness advantage of the introduced species.[118] Genetic pollution occurs either through introduction or through habitat modification, where previously isolated species are brought into contact with the new genotypes. Invading species have been shown to adapt to their new environments in a remarkably short amount of time.[119] The population size of invading species may remain small for a number of years and then experience an explosion in population, a phenomenon known as "the lag effect".[120]

Hybrids resulting from invasive species interbreeding with native species can incorporate their genotypes into the gene pool over time through introgression. Similarly, in some instances a small invading population can threaten much larger native populations. For example, Spartina alterniflora was introduced in the San Francisco Bay and hybridized with native Spartina foliosa. The higher pollen count and male fitness of the invading species resulted in introgression that threatened the native populations due to lower pollen counts and lower viability of the native species.[121] Reduction in fitness is not always apparent from morphological observations alone. Some degree of gene flow is normal, and preserves constellations of genes and genotypes.[122][123] An example of this is the interbreeding of migrating coyotes with the red wolf, in areas of eastern North Carolina where the red wolf was reintroduced.[124] The end result was a decrease in stable breeding pairs of red wolf, which may further complicate the social stability of packs and reintroduction efforts.

Invasive exotic diseases

History is rife with the spread of exotic diseases, such as the introduction of smallpox into the indigenous peoples of the Americas by the Spanish, where it obliterated entire populations of indigenous civilizations before they were ever even seen by Europeans.

Problematic exotic disease introductions in the past century or so include the chestnut blight which has almost eliminated the American chestnut tree from its forest habitat. Responses to increase the population of the American chestnut include creating blight resistant trees that can be reintroduced. This displays both the positive and negative aspects of introduced species.

Another example is the Dutch elm disease, which has severely reduced the American elm trees in forests and cities.


It has been argued that some invasive species "...could fill niches in degraded ecosystems and help restore native biodiversity...".[126]

Study and eradication

Stage Characteristic
0 Propagules residing in a donor region
I Traveling
II Introduced
III Localized and numerically rare
IVa Widespread but rare
IVb Localized but dominant
V Widespread and dominant

While the study of invasive species can be done within many subfields of biology, the majority of research on invasive organisms has been within the field of ecology and geography where the issue of biological invasions is especially important. Much of the study of invasive species has been influenced by Charles Elton's 1958 book The Ecology of Invasion by Animals and Plants which drew upon the limited amount of research done within disparate fields to create a generalized picture of biological invasions.[127][128] Studies on invasive species remained sparse until the 1990s when research in the field experienced a large amount of growth which continues to this day.[128] This research, which has largely consisted of field observational studies, has disproportionately been concerned with terrestrial plants.[128] The rapid growth of the field has driven a need to standardize the language used to describe invasive species and events. Despite this, little standard terminology exists within the study of invasive species which itself lacks any official designation but is commonly referred to as "Invasion ecology" or more generally "Invasion biology".[127][128] This lack of standard terminology is a significant problem, and has largely arisen due to the interdisciplinary nature of the field which borrows terms from numerous disciplines such as agriculture, zoology, and pathology, as well as due to studies on invasive species being commonly performed in isolation of one another.[127]



Introduced species on islands

Perhaps the best place to study problems associated with introduced species is on islands. Depending upon the isolation (how far an island is located from continental biotas), native island biological communities may be poorly adapted to the threat posed by exotic introductions. Often this can mean that no natural predator of an introduced species is present, and the non-native spreads uncontrollably into open or occupied niche.

An additional problem is that birds native to small islands may have become flightless because of the absence of predators prior to introductions and cannot readily escape the danger brought to them by introduced predators. The tendency of rails in particular to evolve flightless forms on islands making them vulnerable has led to the disproportionate number of extinctions in that family.

The field of island restoration has developed as a field of conservation biology and ecological restoration, a large part of which deals with the eradication of invasive species. A 2019 study suggests that if eradications of invasive animals were conducted on just 169 islands the survival prospects of 9.4% of the Earth’s most highly threatened terrestrial insular vertebrates would be improved.[130]



New Zealand

The first invasive species were the dogs and rats brought by Polynesian settlers around 1300.[131][132] Cats, brought later by Europeans, have had a devastating effect upon the native birdlife, particularly as many New Zealand birds are flightless. Rabbits, introduced as a food source by sailors in the 1800s, have become a severe nuisance to farmers, notably in the South Island. Common gorse, originally a hedge plant in Britain, was introduced to New Zealand for the same purpose but grows aggressively and threatens to obliterate native plants in much of the country and is hence routinely eradicated.

The native forests are heavily impacted by several species of deer from North America and Europe and the Australian brushtail possum. These exotic species have all thrived in the New Zealand environment.

South Georgia Island

In 2018, the South Georgia Island was declared free of invasive rodents after a multi-year extermination effort.[133][134]

See also



This article incorporates CC-BY-3.0 text from the reference[115]


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