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Introduced species

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"Alien species" redirects here. For life on planets other than Earth, seeExtraterrestrial life.
CattleBos primigenius taurusintroduced but not naturalized worldwide
Sweet clover(Melilotus sp.), introduced andnaturalizedin theAmericasfromEuropeas a forage and cover crop

Anintroduced species,alien species,exotic species,adventive species,immigrant species,foreign species,non-indigenous species,ornon-native speciesis aspeciesliving outside itsnativedistributionalrange,but which has arrived there byhumanactivity, directly or indirectly, and either deliberately or accidentally. Non-native species can have various effects on the local ecosystem. Introduced species that become established and spread beyond the place of introduction are considerednaturalized.The process of human-caused introduction is distinguished frombiological colonization,in which species spread to new areas through "natural" (non-human) means such as storms andrafting.The Latin expressionneobiotacaptures the characteristic that these species arenewbiotato their environment in terms of establishedbiological network(e.g.food web) relationships. Neobiota can further be divided intoneozoa(also: neozoons, sing. neozoon, i.e. animals) andneophyta(plants).

The impact of introduced species is highly variable. Some have a substantial negative effect on a local ecosystem (in which case they are also classified more specifically as aninvasive species), while other introduced species may have little or no negative impact (no invasiveness), and integrate well into the ecosystem they have been introduced to. Some species have been introduced intentionally to combat pests. They are calledbiocontrolsand may be regarded as beneficial as an alternative to pesticides in agriculture for example. In some instances the potential for being beneficial or detrimental in the long run remains unknown.[1][2][3]The effects of introduced species on natural environments have gained much scrutiny from scientists, governments, farmers and others.

Terminology

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The formal definition of an introduced species from theUnited States Environmental Protection Agencyis "A species that has been intentionally or inadvertently brought into a region or area. Also called an exotic or non-native species".[4][5]

In the broadest and most widely used sense, an introduced species is synonymous with "non-native" and therefore applies as well to most garden and farm organisms; these adequately fit the basic definition given above. However, some sources add to that basic definition "and are now reproducing in the wild",[6]which means that species growing in a garden, farm, or house may not meet the criteria unless they escape and persist.

Subset descriptions

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See also:Glossary of invasion biology terms

There are many terms associated with introduced species that represent subsets of introduced species, and the terminology associated with introduced species is now in flux for various reasons. Examples of these terms are "invasive", "acclimatized", "adventive", "naturalized", and "immigrant" species.

The term "invasive" is used to describe introduced species that cause ecological, economic, or other damage to the area in which they were introduced.

Acclimatized species are introduced species that have changed physically and/or behaviorally in order to adjust to their new environment. Acclimatized species are not necessarily optimally adjusted to their new environment and may just be physically/behaviorally sufficient for the new environment.

Adventive species are often considered synonymous with "introduced species", but this term is sometimes applied exclusively to introduced species that are not permanently established.[7]

Naturalized speciesare often introduced species that do not need human help to reproduce and maintain their population in an area outside their native range (no longer adventive), but that also applies to populations migrating and establishing in a novel environment (e.g.: inEurope,house sparrowsare well established since earlyIron Agethough they originated fromAsia).

Immigrant species are species that travel, sometimes by themselves, but often with human help, between two habitats. Invasiveness is not a requirement.[8]

Invasive species

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Introduction of a species outside its native range is all that is required to be qualified as an "introduced species". Such species might be termednaturalized,"established", or "wild non-native species". If they further spread beyond the place of introduction and cause damage to nearby species, they are called "invasive species".The transition from introduction, to establishment and to invasion has been described in the context of plants.[9]Introduced species are essentially "non-native" species. Invasive species are those introduced species that spread widely or quickly and cause harm, be that to the environment,[10]human health, other valued resources, or the economy. There have been calls from scientists to consider a species "invasive" only in terms of their spread and reproduction rather than the harm they may cause.[11]

According to a practical definition, an invasive species is one that has been introduced and become a pest in its new location, spreading (invading) by natural means. The term is used to imply both a sense of urgency and actual or potential harm. For example, U.S. Executive Order 13112 (1999) defines "invasive species" as "an alien species whose introduction does or is likely to cause economic or environmental harm or harm to human health".[12]The biological definition of invasive species, on the other hand, makes no reference to the harm they may cause, only to the fact that they spread beyond the area of original introduction.

Some argue that "invasive" is a loaded word and harm is difficult to define.[6]

From a regulatory perspective, it is neither desirable nor practical to list as undesirable or outright ban all non-native species (although theState of Hawaiihas adopted an approach that comes close to this). Regulations require a definitional distinction between non-natives that are deemed especially onerous and all others. Introduced "pest" species, that are officially listed as invasive, best fit the definition of an invasive species. Early detection and rapid response is the most effective strategy for regulating a pest species and reducing economic and environmental impacts of an introduction.[13]Management of invasion pathways are on the forefront of eliminating unwanted invasive species this would include preliminary steps; educating the public, cooperation from industries and government resources.[14]

InGreat Britain,theWildlife and Countryside Act 1981prevents the introduction of any animal not naturally occurring in the wild or any of a list of both animals or plants introduced previously and proved to be invasive.

Nature of introductions

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Bydefinition,a species is considered "introduced" when its transport into an area outside of its native range is human mediated. Introductions by humans can be described as either intentional or accidental. Intentional introductions have been motivated by individuals or groups who either (1) believe that the newly introduced species will be in some way beneficial to humans in its new location or, (2) species are introduced intentionally but with no regard to the potential impact. Unintentional or accidental introductions are most often a byproduct of human movements and are thus unbound to human motivations. Subsequent range expansion of introduced species may or may not involve human activity.

WheatTriticumintroduced worldwide from its place of origin (Mesopotamia)

Intentional introductions

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Species that humans intentionally transport to new regions can subsequently become successfully established in two ways. In the first case, organisms are purposely released for establishment in the wild. It is sometimes difficult to predict whether a species will become established upon release, and if not initially successful, humans have made repeated introductions to improve the probability that the species will survive and eventually reproduce in the wild. In these cases, it is clear that the introduction is directly facilitated by human desires.

Malesilver pheasant

In the second case, species intentionally transported into a new region may escape from captive or cultivated populations and subsequently establish independent breeding populations. Escaped organisms are included in this category because their initial transport to a new region is human motivated.

The widespread phenomena of intentional introduction has also been described asbiological globalization.

Positive Introductions

Although most introduced species have negative impacts on the ecosystems they enter into, there are still some species that have affected the ecosystem in a positive way. For example, in New Hampshire invasive plants can provide some benefits to some species. Invasive species such as autumn olive, oriental bittersweet, and honeysuckle produce fruit that is used by a handful of fruit-eating bird species.[15]The invasive plants can also be a source of pollen and nectar for many insects, such as bees. These invasive plants were able to help their ecosystem thriving, and increase the native animal's chances of survival. Several introduced exotic trees served as nest sites for resident waterbird species in Udaipur city, India.[16]

Motivations for intentional introductions

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Economic
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Perhaps the most common motivation for introducing a species into a new place is that of economic gain. Non-native species can become such a common part of an environment, culture, and even diet that little thought is given to their geographic origin. For example,soybeans,kiwi fruit,wheat,honey bees,and alllivestockexcept theAmerican bisonand theturkeyare non-native species to North America. Collectively, non-native crops and livestock account for 98% of US food.[17]These and other benefits from non-natives are so vast that, according to the Congressional Research Service, they probably exceed the costs.[18]

Other examples of species introduced for the purposes of benefitingagriculture,aquacultureor other economic activities are widespread.[19]Eurasian carpwas first introduced to the United States as a potential food source. Theapple snailwas released in Southeast Asia with the intent that it be used as a protein source, and subsequently to places like Hawaii to establish a food industry. In Alaska,foxeswere introduced to many islands to create new populations for the fur trade. About twenty species of African and Europeandung beetleshave established themselves in Australia after deliberate introduction by theAustralian Dung Beetle Projectin an effort to reduce the impact of livestock manure. The timber industry promoted the introduction of Monterey pine (Pinus radiata) from California to Australia and New Zealand as a commercial timber crop. These examples represent only a small subsample of species that have been moved by humans for economic interests.

The rise in the use ofgenetically modified organismshas added another potential economic advantage to introducing new/modified species into different environments. Companies such asMonsantothat earn much of their profit through the selling of genetically modified seeds has added to the controversy surrounding introduced species. The effect of genetically modified organisms varies from organism to organism and is still being researched today, however, the rise of genetically modified organisms has added complexity to the conversations surrounding introduced species.

Human enjoyment
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Introductions have also been important in supporting recreation activities or otherwise increasing human enjoyment. Numerous fish andgame animalshave been introduced for the purposes of sport fishing and hunting. The introduced amphibian (Ambystoma tigrinum) that threatens the endemic California salamander (A. californiense) was introduced to California as a source of bait for fishermen.[20]Pet animalshave also been frequently transported into new areas by humans, and their escapes have resulted in several introductions, such as feralcats,[21]parrots,[22]andpond slider.[23]

Lophura nycthemera(silver pheasant), a native of East Asia, has been introduced into parts ofEuropefor ornamental reasons.

Many plants have been introduced with the intent of aesthetically improving public recreation areas or private properties. The introducedNorway maplefor example occupies a prominent status in many of Canada's parks.[24]The transport ofornamental plantsforlandscapinguse has and continues to be a source of many introductions. Some of these species have escaped horticultural control and become invasive. Notable examples includewater hyacinth,salt cedar,andpurple loosestrife.

In other cases, species have been translocated for reasons of "cultural nostalgia", which refers to instances in which humans who have migrated to new regions have intentionally brought with them familiar organisms. Famous examples include the introduction ofcommon starlingsto North America by the AmericanEugene Schieffelin,a lover of the works of Shakespeare and the chairman of theAmerican Acclimatization Society,who, it is rumoured, wanted to introduce all of the birds mentioned in Shakespeare's plays into the United States. He deliberately released eighty starlings into Central Park in New York City in 1890, and another forty in 1891.

Yet another prominent example of an introduced species that became invasive is theEuropean rabbit in Australia.Thomas Austin,a British landowner, had rabbits released on his estate inVictoriabecause he missed hunting them. A more recent example is the introduction of thecommon wall lizard(Podarcis muralis)to North America by aCincinnatiboy, George Rau, around 1950 after a family vacation toItaly.[25]

Addressing environmental problems
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Intentional introductions have also been undertaken with the aim of ameliorating environmental problems. A number of fast spreading plants such askudzuhave been introduced as a means oferosioncontrol. Other species have been introduced asbiological controlagents to controlinvasive species.This involves the purposeful introduction of anatural enemyof the target species with the intention of reducing its numbers or controlling its spread.

A special case of introduction is the reintroduction of a species that has become locally endangered or extinct, done in the interests of conservation.[26]Examples of successful reintroductions include wolves toYellowstone National Parkin the U.S., and thered kiteto parts of England and Scotland. Introductions or translocations of species have also been proposed in the interest ofgenetic conservation,which advocates the introduction of new individuals into genetically depauperate populations of endangered or threatened species.[27]

Unintentional introductions

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Unintentional introductions occur when species are transported by human vectors. Increasing rates of human travel are providing accelerating opportunities for species to be accidentally transported into areas in which they are not considered native. For example, three species ofrat(the black, Norway and Polynesian) have spread to most of the world as hitchhikers on ships, andarachnidssuch asscorpionsand exoticspidersare sometimes transported to areas far beyond their native range by riding in shipments of tropical fruit. This was seen during the introduction ofSteatoda nobilis(Noble false widow) worldwide throughbananashipments.[28]

Further there are numerous examples of marine organisms being transported inballast water,among them the invasivecomb jellyMnemiopsis leidyi,the dangerous bacteriumVibrio cholerae,or the foulingzebra mussel.The Mediterranean and Black Seas, with their high volume shipping from exotic sources, are most impacted by this problem.[29]Busy harbors are all potential hotspots as well: over 200 species have been introduced to theSan Francisco Bayin this manner making it the most heavily invaded estuary in the world.[30]

There is also the accidental release of theAfricanized honey bees(AHB), known colloquially as "killer bees" ) orAfricanized beeto Brazil in 1957 and theAsian carpto the United States. The insect commonly known as thebrown marmorated stink bug(Halyomorpha halys) was introduced accidentally in Pennsylvania. Another form of unintentional introductions is when an intentionally introduced plant carries a parasite or herbivore with it. Some become invasive, for example, theoleander aphid,accidentally introduced with the ornamental plant,oleander.

Yet another unintentional pathway of introduction is during the delivery ofhumanitarian aidin the aftermath of natural disasters.[31][32]This occurred during relief efforts forHurricane MariainDominica,it was found that thecommon green iguana,theCuban tree frog,and potentially theVenezuela snouted tree frogwere introduced with the former two becoming established.[32]

Most accidentally or intentionally introduced species do not become invasive as the ones mentioned above. For instance, Some 179 coccinellid species have been introduced to the U.S. and Canada; about 27 of these non-native species have become established, and only a handful can be considered invasive, including the intentionally introducedHarmonia axyridis,multicoloredAsian lady beetle.[33]However the small percentage of introduced species that become invasive can produce profound ecological changes. In North America,Harmonia axyridishas become the most abundantlady beetleand probably accounts for more observations than all the nativelady beetlesput together.[34]

Introduced plants

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See also:Adventive plant
Thehorse chestnut,Aesculus hippocastanum,native toGreeceand theBalkan peninsula,has been introduced across most of Europe and parts of North America as an ornamental plant. Introduced to the United Kingdom in 1616,[35]thisneophytespecies has become widely distributed across the country. Though non-native, its leaves attract insects which serve as a food source for populations of native birds.[36]

Many non-native plants have been introduced into new territories, initially as eitherornamental plantsor forerosioncontrol, stock feed, or forestry. Whether an exotic will become aninvasive speciesis seldom understood in the beginning, and many non-native ornamentals languish in the trade for years before suddenlynaturalizingand becoming invasive. Studies have shown that introduced species display a greater likeliness of naturalizing when there is an appropriate environmental match, the plant species are short lived herbs or cultivate from seeds.[37]

Peaches,for example, originated in China, and have been carried to much of the populated world.Tomatoesare native to theAndes.Squash (pumpkins),maize(corn), and tobacco are native tothe Americas,but were introduced to theOld World.Many introduced species require continued human intervention to survive in the new environment. Others may become feral, but do not seriouslycompetewith natives, but simply increase the biodiversity of the area. One example would beDandelionsinNorth America,which have become an essential source of early season nectar for both native and introduced pollinators, and do not meaningfully compete with native grasses or flowers.

A very troublesome marine species in southern Europe is theseaweedCaulerpa taxifolia.Caulerpawas first observed in theMediterranean Seain 1984, off the coast ofMonaco.By 1997, it had covered some 50 km2.It has a strong potential to overgrow naturalbiotopes,and represents a major risk for sublittoralecosystems.The origin of the alga in the Mediterranean was thought to be either as a migration through theSuez Canalfrom the Red Sea, or as an accidental introduction from an aquarium. Another troublesome plant species is the terrestrial plantPhyla canescens,which was intentionally introduced into many countries in North America, Europe, and Africa as an ornamental plant.[38][39]This species has become invasive in Australia, where it threatens native rare plants and causes erosion and soil slumping around river banks.[40]It has also become invasive in France where it has been listed as an invasive plant species of concern in the Mediterranean region, where it can form monocultures that threaten critical conservation habitats.[41]

Japanese knotweedgrows profusely in many nations. Human beings introduced it into many places in the 19th century. It is a source ofresveratrol,a dietary supplement. It can grow in building foundations, threatening their stability, and spreads quite quickly. Mesquite (Prosopis juliflora) has spread very widely across the Indian subcontinent and is impacting a range of wild species including urban birds.[42]

Introduced animals

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ChickensGallus gallus domesticus,from Asia, introduced in the rest of the world

Most introduced species do not become invasive. Examples of introduced animals that have become invasive include thegypsy mothin easternNorth America,thezebra musselandalewifein theGreat Lakes,theCanada gooseandgray squirrelin Europe, thebeaverinTierra del Fuego,themuskratinEuropeandAsia,thecane toadandred foxinAustralia,nutriainNorth America,Eurasia,andAfrica,and thecommon brushtail possuminNew Zealand.InTaiwan,the success of introduced bird species was related to their native range size and body size; larger species with larger native range sizes were found to have larger introduced range sizes.[43]

One notoriously devastating introduced species is the small Indian mongoose (Urva auropunctata). Originating in a region encompassingIranandIndia,it was introduced to theWest IndiesandHawaiiin the late 1800s for pest control. Since then, it has thrived on prey unequipped to deal with its speed, nearly leading to the local extinction of a variety of species.[44]

In some cases, introduced animals may unintentionally promote the cause ofrewilding.[45]For example, escapedhorsesanddonkeysthat have gone feral inthe Americasmay play ecological roles similar to those of theequidsthat becameextinctthere at the end of thePleistocene.[46]

The exotic pet trade has also been a large source of introduced species. The species favored as pets have more general habitat requirements and larger distributions.[47]Therefore, as these pets escape or are released, unintentionally or intentionally, they are more likely to survive and establish non-native populations in the wild. Among the popular exotic pets that have become alien or invasive species are parrots, frogs, terrapins, and iguanas.

Most commonly introduced species

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Some species, such as theWestern honey bee,brown rat,house sparrow,ring-necked pheasant,andEuropean starling,have been introduced very widely. In addition there are someagriculturalandpetspecies that frequently becomeferal;these includerabbits,dogs,ducks,snakes,goats,fish,pigs,andcats.Manywater fleassuch asDaphnia,BosminaandBythotrepheshave introduced around the world, causing dramatic changes in native freshwater ecosystems.[48]

Genetics

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When a new species is introduced, the species could potentially breed with members of native species, producing hybrids. The effect of the creating of hybrids can range from having little effect, a negative effect, to having devastating effects on native species. Potential negative effects include hybrids that are less fit for their environment resulting in a population decrease. This was seen in theAtlantic Salmonpopulation when high levels of escape from Atlantic Salmon farms into the wild populations resulted in hybrids that had reduced survival.[49]Potential positive effects include adding to thegenetic diversityof the population which can increase theadaptationability of the population and increase the number of healthy individuals within a population. This was seen in the introduction ofguppiesinTrinidadto encouragepopulation growthand introduce newallelesinto the population. The results of this introduction included increased levels ofheterozygosityand a largerpopulation size.[50]Wide-spread introductions of non-native iguanas are causing devastating effects on nativeIguanapopulations in theCaribbeanLesser Antilles,as hybrids appear to have higher fitness than native iguanas, leading to competitive outcompetition and replacement.[51][52]Numerous populations have already become extinct and hybridization continues to reduce the number of native iguanas on multiple islands.

In plants, introduced species have been observed to undergo rapid evolutionary change to adapt to their new environments, with changes in plant height, size, leaf shape, dispersal ability, reproductive output, vegetative reproduction ability, level of dependence on themycorrhizal network,and level of phenotype plasticity appearing on timescales of decades to centuries.[53]

On a planetary body

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Main article:Interplanetary contamination

It has been hypothesized thatinvasive speciesofmicrobial lifecould contaminate aplanetary bodyafter the former is introduced by aspace probeorspacecraft,either deliberately or unintentionally.[54]It has also been hypothesized that the origin of life on earth is due to introductions of life from other planets billions of years ago, possibly by a sentient race. Projects have been proposed to introduce life to other lifeless buthabitable planetsin other star systems some time in the future. In preparation for this, projects have been proposed to see if anything is still alive from any of the feces left behind during the sixMoon landingsfrom 1969 to 1972.[55]

See also

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References

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Further reading

[edit]
  • Chris D. Thomas (2017).Inheritors of the Earth: How Nature Is Thriving in an Age of Extinction.PublicAffairs.ISBN978-1610397278.
[edit]
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