An invasive species is a species that is not native to a specific location (an introduced species), and that has a tendency to spread to a degree believed to cause damage to the environment, human economy or human health.
The criteria for invasive species has been controversial, as
widely divergent perceptions exist among researchers as well as concerns
with the subjectivity of the term "invasive".
Several alternate usages of the term have been proposed. The term as
most often used applies to introduced species (also called
"non-indigenous" or "non-native") that adversely affect the habitats and bioregions
they invade economically, environmentally, or ecologically. Such
invasive 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 non-native invasive plant species labeled as exotic pest plants and invasive exotics growing in native plant communities. It has been used in this sense by government organizations as well as conservation groups such as the International Union for Conservation of Nature (IUCN) and the California Native Plant Society.
The European Union defines "Invasive Alien Species" as those that are,
firstly, outside their natural distribution area, and secondly, threaten
biological diversity.
The term is also used by land managers, botanists, researchers, horticulturalists, conservationists, and the public for noxious weeds. The kudzu vine (Pueraria lobata), Andean pampas grass (Cortaderia jubata), and yellow starthistle (Centaurea solstitialis) are examples. An alternate usage broadens the term to include indigenous or "native" species along with non-native species, that have colonized natural areas (p. 136). Deer are an example, considered to be overpopulating their native zones and adjacent suburban gardens, by some in the Northeastern and Pacific Coast regions of the United States. Sometimes the term is used to describe a non-native or introduced species that has become widespread (p. 136). However, not every introduced species has adverse effects on the environment. A nonadverse example is the common goldfish (Carassius auratus), which is found throughout the United States, but rarely achieves high densities (p. 136). Notable examples of invasive species include European rabbits, grey squirrels, domestic cats, carp and ferrets.
Dispersal and subsequent proliferation of species is not solely an anthropogenic phenomenon. There are many mechanisms by which species from all Kingdoms have been able to travel across continents in short periods of time such as via floating rafts, or on wind currents. Charles Darwin, a British naturalist, performed many experiments to better understand long distance seed dispersal, and was able to germinate seeds from insect frass, faeces of waterfowl,
dirt clods on the feet of birds, all of which may have traveled
significant distances under their own power, or be blown off course by
thousands of miles.
Invasion of long-established ecosystems by organisms from distant
bio-regions is a natural phenomenon, which has likely been accelerated
via hominid-assisted migration although this has not been adequately
directly measured.
The definition of "native" is controversial in that there is no
way to precisely determine nativity. 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.
Causes
Scientists
include species and ecosystem factors among the mechanisms that, when
combined, establish invasiveness in a newly introduced species.
Species based mechanisms
While
all species compete to survive, invasive species appear to have
specific traits or specific combinations of traits that allow them to
outcompete native species.
In some cases, the competition is about rates of growth and
reproduction. In other cases, species interact with each other more
directly.
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.
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. 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
- Prior successful invasions
Typically, an introduced species must survive at low population densities before it becomes invasive in a new location.
At low population densities, it can be difficult for the introduced
species to reproduce and maintain itself in a new location, so a species
might reach a location multiple times before it becomes established.
Repeated patterns of human movement, such as ships sailing to and from
ports or cars driving up and down highways offer repeated opportunities
for establishment (also known as a high propagule pressure).
An introduced species might become invasive if it can outcompete native species for resources such as nutrients, light, physical space, water, or food. If these species evolved under great competition or predation, then the new environment may host fewer able competitors, allowing the invader to proliferate quickly. Ecosystems which are being used to their fullest capacity by native species can be modeled as zero-sum systems in which any gain for the invader is a loss for the native. However, such unilateral competitive superiority (and extinction of native species with increased populations of the invader) is not the rule.
Invasive species often coexist with native species for an extended
time, and gradually, the superior competitive ability of an invasive
species becomes apparent as its population grows larger and denser and
it adapts to its new location.
An invasive species might be able to use resources that were
previously unavailable to native species, such as deep water sources
accessed by a long taproot, or an ability to live on previously uninhabited soil types. For example, barbed goatgrass (Aegilops triuncialis) was introduced to California on serpentine soils, which have low water-retention, low nutrient levels, a high magnesium/calcium ratio, and possible heavy metal
toxicity. Plant populations on these soils tend to show low density,
but goatgrass can form dense stands on these soils and crowd out native
species that have adapted poorly to serpentine soils.
Invasive species might alter their environment by releasing chemical compounds, modifying abiotic factors, or affecting the behaviour of herbivores, creating a positive or negative impact on other species. Some species, like Kalanchoe daigremontana, produce allelopathic compounds,
that might have an inhibitory effect on competing species, and
influence some soil processes like carbon and nitrogen mineralization. Other species like Stapelia gigantea facilitates the recruitment of seedlings of other species in arid environments by providing appropriate microclimatic conditions and preventing herbivory in early stages of development.
Other examples are Centaurea solstitialis (yellow starthistle) and Centaurea diffusa (diffuse knapweed). These Eastern European noxious weeds have spread through the western and West Coast states. Experiments show that 8-hydroxyquinoline, a chemical produced at the root of C. diffusa,
has a negative effect only on plants that have not co-evolved with it.
Such co-evolved native plants have also evolved defenses. C. diffusa and C. solstitialis
do not appear in their native habitats to be overwhelmingly successful
competitors. Success or lack of success in one habitat does not
necessarily imply success in others. Conversely, examining habitats in
which a species is less successful can reveal novel weapons to defeat
invasiveness.
Changes in fire regimens are another form of facilitation. Bromus tectorum,
originally from Eurasia, is highly fire-adapted. It not only spreads
rapidly after burning but also increases the frequency and intensity
(heat) of fires by providing large amounts of dry detritus
during the fire season in western North America. In areas where it is
widespread, it has altered the local fire regimen so much that native
plants cannot survive the frequent fires, allowing B. tectorum to further extend and maintain dominance in its introduced range.
Ecological facilitation also occurs where one species physically modifies a habitat in ways that are advantageous to other species. For example, zebra mussels increase habitat complexity on lake floors, providing crevices in which invertebrates live. This increase in complexity, together with the nutrition provided by the waste products of mussel filter-feeding, increases the density and diversity of benthic invertebrate communities.
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.
When changes such as a forest fire occur, normal succession favors native grasses and forbs.
An introduced species that can spread faster than natives can use
resources that would have been available to native species, squeezing
them out. Nitrogen and phosphorus are often the limiting factors in these situations.
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. An example of this type can be found within the Lampropholis delicata species of skink.
Ecosystem changes can alter species' distributions. For example, edge effects describe what happens when part of an ecosystem is disturbed as when land is cleared for agriculture.
The boundary between remaining undisturbed habitat and the newly
cleared land itself forms a distinct habitat, creating new winners and
losers and possibly hosting species that would not thrive outside the
boundary habitat.
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,
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.
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. Adaptations include changes in morphology, physiology, phenology, and plasticity.
Rapid adaptive evolution in these species leads to offspring that
have higher fitness and are better suited for their environment.
Intraspecific phenotypic plasticity, pre-adaptation and
post-introduction evolution are all major factors in adaptive evolution.
Plasticity in populations allows room for changes to better suit the
individual in its environment. This is key in adaptive evolution because
the main goal is how to best be suited to the ecosystem that the
species has been introduced. The ability to accomplish this as quickly
as possible will lead to a population with a very high fitness.
Pre-adaptations and evolution after the initial introduction also play a
role in the success of the introduced species. If the species has
adapted to a similar ecosystem or contains traits that happen to be well
suited to the area that it is introduced, it is more likely to fare
better in the new environment. This, in addition to evolution that takes
place after introduction, all determine if the species will be able to
become established in the new ecosystem and if it will reproduce and
thrive.
Ecology
Traits of invaded ecosystems
In 1958, Charles S. Elton 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.
This debate hinged on the spatial scale
at which invasion studies were performed, and the issue of how
diversity affects susceptibility remained unresolved as of 2011.
Small-scale studies tended to show a negative relationship between diversity
and invasion, while large-scale studies tended to show the reverse. The
latter result may be a side-effect of invasives' ability to capitalize
on increased resource availability and weaker species interactions that
are more common when larger samples are considered.
Invasion was more likely in ecosystems that were similar to the one in which the potential invader evolved. 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. An example of this phenomenon was the decimation of native bird populations on Guam by the invasive brown tree snake.
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.
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.
Vectors
Non-native species have many vectors, including biogenic vectors, but most invasions are associated with human activity. Natural range
extensions are common in many species, but the rate and magnitude of
human-mediated extensions in these species tend to be much larger than
natural extensions, and humans typically carry specimens greater
distances than natural forces.
An early human vector occurred when prehistoric humans introduced the Pacific rat (Rattus exulans) to Polynesia.
Vectors include plants or seeds imported for horticulture. The pet trade moves animals across borders, where they can escape and become invasive. Organisms stow away on transport vehicles.
The arrival of invasive propagules to a new site is a function of the site's invasibility.
Species have also been introduced intentionally. For example, to feel more "at home," American colonists
formed "Acclimation Societies" that repeatedly imported birds that were
native to Europe to North America and other distant lands. In 2008, U.S. postal workers in Pennsylvania noticed noises coming from inside a box from Taiwan; the box contained more than two dozen live beetles. Agricultural Research Service entomologists identified them as rhinoceros beetle, hercules beetle, and king stag beetle.
Because these species were not native to the U.S., they could have
threatened native ecosystems. To prevent exotic species from becoming a
problem in the U.S., special handling and permits are required when
living materials are shipped from foreign countries. USDA programs such as Smuggling Interdiction and Trade Compliance (SITC) attempt to prevent exotic species outbreaks in America.
Many invasive species, once they are dominant in the area, are
essential to the ecosystem of that area. If they are removed from the
location it could be harmful to that area.
Economics plays a major role in exotic species introduction. High demand for the valuable Chinese mitten crab is one explanation for the possible intentional release of the species in foreign waters.
Within the Aquatic Environment
The
development of maritime trade has rapidly affected the way marine
organisms are transported within the ocean. Two ways marine organisms
are transported to new environments are hull fouling and ballast water
transport. In fact, Molnar et al. 2008 documented the pathways of
hundreds of marine invasive species and found that shipping was the
dominant mechanism for the transfer of invasive species.
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.
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.
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.
In fact, it is estimated that 10,000 different species, many of which
are non-indigenous, are transported via ballast water each day. Many of these species are considered harmful and can negatively impact 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. 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). Current regulations for the Great Lakes rely on ‘salinity shock’ to kill freshwater organisms left in ballast tanks.
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.
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.
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.
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,
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.
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.
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.
Impacts of wildfire and firefighting
Invasive species often exploit disturbances to an ecosystem (wildfires, roads, foot trails) to colonize an area. Large wildfires can sterilize soils, while adding a variety of nutrients.
In the resulting free-for-all, formerly entrenched species lose their
advantage, leaving more room for invasives. In such circumstances plants
that can regenerate from their roots have an advantage. Non-natives
with this ability can benefit from a low intensity fire burns that
removes surface vegetation, leaving natives that rely on seeds for propagation to find their niches occupied when their seeds finally sprout.
Wildfires often occur in remote areas, needing fire suppression
crews to travel through pristine forest to reach the site. The crews can
bring invasive seeds with them. If any of these stowaway seeds become
established, a thriving colony of invasives can erupt in as few as six
weeks, after which controlling the outbreak can need years of continued
attention to prevent further spread. Also, disturbing the soil surface,
such as cutting firebreaks, destroys native cover, exposes soil, and can
accelerate invasions. In suburban and wildland-urban interface areas, the vegetation clearance and brush removal ordinances of municipalities for defensible space can result in excessive removal of native shrubs and perennials that exposes the soil to more light and less competition for invasive plant species.
Fire suppression vehicles are often major culprits in such
outbreaks, as the vehicles are often driven on back roads overgrown with
invasive plant species. The undercarriage of the vehicle becomes a
prime vessel of transport. In response, on large fires, washing stations
"decontaminate" vehicles before engaging in suppression activities. Large wildfires attract firefighters from remote places, further increasing the potential for seed transport.
Effects
Ecological
Land
clearing and human habitation put significant pressure on local
species. Disturbed habitats are prone to invasions that can have adverse
effects on local ecosystems, changing ecosystem functions. A species of
wetland plant known as ʻaeʻae in Hawaii (the indigenous Bacopa monnieri) is regarded as a pest species in artificially manipulated water bird refuges because it quickly covers shallow mudflats established for endangered Hawaiian stilt (Himantopus mexicanus knudseni), making these undesirable feeding areas for the birds.
Multiple successive introductions of different non-native species
can have interactive effects; the introduction of a second non-native
species can enable the first invasive species to flourish. Examples of
this are the introductions of the amethyst gem clam (Gemma gemma) and the European green crab (Carcinus maenas). The gem clam was introduced into California's Bodega Harbor from the East Coast of the United States a century ago. It had been found in small quantities in the harbor but had never displaced the native clam species (Nutricola
spp.). In the mid-1990s, the introduction of the European green crab,
found to prey preferentially on the native clams, resulted in a decline
of the native clams and an increase of the introduced clam populations.
In the Waterberg
region of South Africa, cattle grazing over the past six centuries has
allowed invasive scrub and small trees to displace much of the original grassland, resulting in a massive reduction in forage
for native bovids and other grazers. Since the 1970s, large scale
efforts have been underway to reduce invasive species; partial success
has led to re-establishment of many species that had dwindled or left
the region. Examples of these species are giraffe, blue wildebeest, impala, kudu and white rhino.
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.
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. For example, hybridization with introduced cordgrass, Spartina alterniflora, threatens the existence of California cordgrass (Spartina foliosa) in San Francisco Bay. 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.
Geomorphological
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.
Economic
Some invaders cause negative benefits towards 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 non
existent, so the sea lamprey acts as a predator, and can consume up to
40 pounds of fish in its 12-18 month feeding period. Sea lampreys prey on all types of large fish such as lake trout and salmon.
The sea lampreys' destructive effects towards large fish negatively
affects the fishing industry and has helped collapse the population of
some economy dependent species.
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, and other invasive plants can also be harvested and utilized as a source of bioenergy.
Benefits
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 Flycatcher 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.
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.
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, has become an important pollinator of caneberry as well as cucurbit, apple trees, and blueberry bushes.
Invasivorism
Invasive species are flora and fauna whose introduction into a
habitat disrupts the native eco-system. In response, Invasivorism is a
movement that explores the idea of eating invasive species in order to
control, reduce, or eliminate their populations. Chefs from around the
world have begun seeking out and using invasive species as alternative
ingredients. Miya's of New Haven, Connecticut created the first invasive species menu in the world. Skeptics point out that once a foreign species has entrenched itself in a new place—such as the Indo-Pacific lionfish that has now virtually taken over the waters of the Western Atlantic, Caribbean and Gulf of Mexico—eradication
is almost impossible. Critics argue that encouraging consumption might
have the unintended effect of spreading harmful species even more
widely.
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.
Costs
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. Economic losses can also occur through loss of recreational and tourism revenues.
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. 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,
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.
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. 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.
Invasive plant pathogens and insect vectors for plant diseases
can also suppress agricultural yields and nursery stock. Citrus greening
is a bacterial disease vectored by the invasive Asian citrus psyllid (ACP). Because of the impacts of this disease on citrus crops, citrus is under quarantine and highly regulated in areas where ACP has been found.
Aquaculture
Aquaculture is a very common vector of species introductions – mainly of species with economic potential (e.g., Oreochromis niloticus).
Forestry
The unintentional introduction of forest pest species and plant pathogens can change forest ecology and damage the timber industry. Overall, forest ecosystems in the U.S. are widely invaded by exotic pests, plants, and pathogens.
The Asian long-horned beetle (Anoplophora glabripennis)
was first introduced into the U.S. in 1996, and was expected to infect
and damage millions of acres of hardwood trees. As of 2005 thirty
million dollars had been spent in attempts to eradicate this pest and
protect millions of trees in the affected regions. The woolly adelgid has inflicted damage on old-growth spruce, fir and hemlock forests and damages the Christmas tree industry. And the chestnut blight fungus (Cryphonectria parasitica) and Dutch elm disease (Ophiostoma novo-ulmi) are two plant pathogens with serious impacts on these two species, and forest health. Garlic mustard, Alliaria petiolata,
is one of the most problematic invasive plant species in eastern North
American forests. The characteristics of garlic mustard are slightly
different from those of the surrounding native plants, which results in a
highly successful species that is altering the composition and function
of the native communities it invades. When garlic mustard invades the understory
of a forest, it affects the growth rate of tree seedlings, which is
likely to alter forest regeneration of impact forest composition in the
future.
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.
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. The very loud call of the introduced common coqui depresses real estate values in affected neighborhoods of Hawaii.
Health
Encroachment of humans into previously remote ecosystems has exposed exotic diseases such as HIV 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. A recent example of an introduced disease is the spread of the West Nile virus, which killed humans, birds, mammals, and reptiles. The introduced Chinese mitten crabs are carriers of Asian lung fluke. Waterborne disease agents, such as cholera bacteria (Vibrio cholerae), and causative agents of harmful algal blooms are often transported via ballast water. 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.
Biodiversity
Biotic invasion is considered one of the five top drivers for global biodiversity loss and is increasing because of tourism and globalization. This may be particularly true in inadequately regulated fresh water systems, though quarantines and ballast water rules have improved the situation.
Invasive species may drive local native species to extinction via competitive exclusion, niche displacement, or hybridisation
with related native species. Therefore, besides their economic
ramifications, alien invasions may result in extensive changes in the
structure, composition and global distribution of the biota of sites of
introduction, leading ultimately to the homogenisation of the world's
fauna and flora and the loss of biodiversity.
Nevertheless, it is difficult to unequivocally attribute extinctions to
a species invasion, and the few scientific studies that have done so
have been with animal taxa. Concern over the impacts of invasive species
on biodiversity must therefore consider the actual evidence (either
ecological or economic), in relation to the potential risk.
Genetic pollution
Native species can be threatened with extinction 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.
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.
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".
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. 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. 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.
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.
Diseases may also be vectored by invasive insects such as the Asian citrus psyllid and the bacterial disease citrus greening.
But in recent years some argue that some introduced species may have a positive ecological impact on an environment.
Study and eradication
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.
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. This research, which has largely consisted of field observational studies, has disproportionately been concerned with terrestrial plants.
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".
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.
In an attempt to avoid the ambiguous, subjective, and pejorative
vocabulary that so often accompanies discussion of invasive species even
in scientific papers, Colautti and MacIsaac proposed a new nomenclature
system based on biogeography rather than on taxa.
By discarding taxonomy, human health,
and economic factors, this model focused only on ecological factors.
The model evaluated individual populations rather than entire species.
It classified each population based on its success in that environment.
This model applied equally to indigenous and to introduced species, and
did not automatically categorize successful introductions as harmful.
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.
New Zealand
In New Zealand the largest commercial crop is Pinus radiata, the native Californian Monterey pine tree, which grows as well in New Zealand as in California. However, the native forests are also occupied by deer from North America and Europe.
They are exotic species and have thrived in the New Zealand
environment. The pines are seen as beneficial while the deer are
regarded as serious pests.
Common gorse, originally a hedge plant in Britain,
was introduced to New Zealand for the same purpose. Like the Monterey
pine, it has shown a favour to its new climate. It is, however, regarded
as a noxious plant that threatens to obliterate native plants in much
of the country and is hence routinely eradicated, though it can also
provide a nursery environment for native plants to reestablish
themselves.
Rabbits, introduced as a food source by sailors in the 1800s, have become a severe nuisance to farmers, notably in the South Island. The rabbit calicivirus
was illegally imported and released, but it had little lasting effect
upon the rabbit population other than to make it more resistant.
Cats, brought later by Europeans, have had a devastating effect
upon the native birdlife, particularly as many New Zealand birds are
flightless. Feral cats and dogs which were originally brought as pets
are also known to kill large numbers of birds. A recent (2006) study in
the South Island has shown that even domestic cats with a ready supply
of food from their owners may kill hundreds of birds in a year,
including natives.
Sparrows, which were brought to control insects upon the introduced grain crops, have displaced native birds as have rainbow lorikeets and cockatoos (both from Australia) which fly free around areas west of Auckland City such as the Waitakere Ranges.
Two notable varieties of spiders have also been introduced: the white tail spider and the redback spider. Both may have arrived inside shipments of fruit. Until then, the only spider (and the only venomous animal) dangerous to humans was the native katipo, which is very similar to the redback and interbreed with the more aggressive Australian variety.
South Georgia Island
In 2018, the South Georgia Island was declared free of invasive rodents after a multi-year extermination effort.