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Coextinction and cothreatened refer to the phenomena of the loss or decline of a host
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
resulting in the loss or endangerment of an other species that depends on it, potentially leading to cascading effects across
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it ...
s. The term originated by the authors Stork and Lyal (1993) and was originally used to explain the extinction of
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
following the loss of their specific hosts. The term is now used to describe the loss of any interacting species, including competition with their counterpart, and specialist
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
s with their food source. Coextinction is especially common when a
keystone species A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintaini ...
goes extinct.


Causes

The most frequently cited example is that of the extinct passenger pigeon and its parasitic
bird lice A bird louse is any chewing louse (small, biting insects) of order Phthiraptera which parasitizes warm-blooded animals, especially birds. Bird lice may feed on feathers, skin, or blood. They have no wings, and their biting mouth parts distin ...
''
Columbicola extinctus ''Columbicola extinctus'', also known as the passenger pigeon chewing louse, is an extant species of phtilopterid louse that was once believed to have been extinct with its only known host, the passenger pigeon, prior to its rediscovery livin ...
'' and ''Campanulotes defectus''. Recently, ''C. extinctus'' was rediscovered on the
band-tailed pigeon The band-tailed pigeon (''Patagioenas fasciata'') is a medium-sized bird of the Americas. Its closest relatives are the Chilean pigeon and the ring-tailed pigeon, which form a clade of ''Patagioenas'' with a terminal tail band and iridescent p ...
, and ''C. defectus'' was found to be a likely case of misidentification of the existing ''
Campanulotes flavus ''Campanulotes'' is a genus of lice in the disputed, probably paraphyletic, family Philopteridae, the chewing lice, or in the family Goniodidae. It is mostly a genus of parasites on birds. Some species in the genus may have been cases of co ...
''. However, even though the passenger pigeon louse was rediscovered, coextinctions of other parasites, even on the passenger pigeon, may have occurred. Several louse species—such as ''
Rallicola extinctus ''Rallicola extinctus'' is an extinct species of phtilopterid louse.Palma, R. L. (1999): Amendments and additions to the 1982 list of chewing lice (Insecta: Phthiraptera) from birds in New Zealand. ''Notornis'' 46(3): 373–387PDF fulltext This ...
'', a huia parasite—probably became extinct together with their hosts. Recent studies have suggested that up to 50% of species may go extinct in the next 50 years. This is in part due to coextinction; for example the loss of tropical butterfly species from Singapore is attributed to the loss of their specific larval host plants. To see how possible future cases of coextinction would play out, researchers have made models to show probabilistic relationships between affiliate and host extinctions across co-evolved inter-specific systems. The subjects are pollinating
Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extendi ...
Wasps and Ficus, primate parasites, (Pneumocystis
Fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
,
Nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
, and
Lice Louse ( : lice) is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a resul ...
) and their hosts, parasitic mites and lice and their avian hosts, butterflies and their larval host plants, and ant butterflies and their host ants. For all but the most host-specific affiliate groups (e.g., primate Pneumocystis fungi and primates), affiliate extinction levels may be modest at low levels of host extinction but can be expected to rise quickly as host extinctions increase to levels predicted in the near future. This curvilinear relationship between host and affiliate extinction levels may also explain, in part, why so few coextinction events have been documented to date. Investigations have been carried out into coextinction risk among the rich Psyllid fauna
Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to arou ...
– Psylloidea inhabiting acacias (Fabaceae-Mimosoideae: Acacia) in central eastern New South Wales, Australia. The results, suggest that ''A. ausfeldii'' hosts one specialist psyllid species, Acizzia, and that ''A. gordonii'' hosts one specialist psyllid, Acizzia. Both psyllid species may be threatened at the same level of their host species with coextinction. Interaction patterns can be used to anticipate the consequences of phylogenetic effects. By using a system of methodical observations, scientists can use the phylogenetic relationships of species to predict the number of interactions they exhibit in more than one-third of the networks, and the identity of the species with which they interact in about half of the networks. Consequentially, simulated extinction events tend to trigger coextinction cascades of related species. This results in a non-random pruning of the evolutionary tree. In a 2004 paper in ''
Science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence ...
'',
ecologist Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
Lian Pin Koh Lian Pin Koh (born 1976 in Singapore) is a Singaporean conservation scientist. He is the Kwan Im Thong Hood Cho Temple Professor of Conservation, Vice Dean of Research at thFaculty of Science Director of thCentre for Nature-based Climate Solutio ...
and colleagues discuss coextinction, stating
"Species coextinction is a manifestation of the interconnectedness of organisms in complex
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
s. The loss of species through coextinction represents the loss of irreplaceable
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary and
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
ary history. In view of the global extinction crisis, it is imperative that coextinction be the focus of future research to understand the intricate processes of species extinctions. While coextinction may not be the most important cause of species extinctions, it is certainly an insidious one." (Koh ''et al.'' 2004)
Koh ''et al.'' also define coendangered as
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
"likely to go extinct if their currently endangered hosts ..become extinct." One example is the near extinction of the genus '' Hibiscadelphus'' as a consequence of the disappearance of several of the
Hawaiian honeycreeper Hawaiian honeycreepers are a group of small, passerine birds endemic to Hawaii. They are closely related to the rosefinches in the genus ''Carpodacus'', but many species have evolved features unlike those present in any other finch. Their great ...
s, its
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the m ...
s. There are several instances of
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s and scavengers dying out following the disappearance of species which represented their source of food: for example, the coextinction of the Haast's eagle with the moa. Coextinction may also occur on a local level: for example, the decline in the red ant ''
Myrmica sabuleti ''Myrmica sabuleti'' is a species of ant in the genus ''Myrmica''. The species is indigenous to Europe, and most colonies are polygynous. Caterpillars of the large blue butterfly (''Phengaris arion'') parasitically prey on this ant. The caterp ...
'' in southern
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe ...
, caused by habitat loss, resulted in the local extinction of the
large blue The large blue (''Phengaris arion'') is a species of butterfly in the family Lycaenidae. The species was first defined in 1758 and first recorded in Britain in 1795. In 1979 the species became mostly extinct in Britain but has been successfully ...
butterfly, which is dependent on the ant as a host for the larvae. In this case the ant avoided local extinction, and the butterfly has been reintroduced. Another example of species going through coextinction is the rhinoceros stomach bot fly (''Gyrostigma rhinocerontis'') and its host species the endangered
black rhinoceros The black rhinoceros, black rhino or hook-lipped rhinoceros (''Diceros bicornis'') is a species of rhinoceros, native to eastern and southern Africa including Angola, Botswana, Kenya, Malawi, Mozambique, Namibia, South Africa, Eswatini, Tanzania ...
and
white rhinoceros The white rhinoceros, white rhino or square-lipped rhinoceros (''Ceratotherium simum'') is the largest extant species of rhinoceros. It has a wide mouth used for grazing and is the most social of all rhino species. The white rhinoceros consists ...
(''Diceros bicornis'' and ''Ceratotherium simum''). The fly's larvae mature in a rhinoceros's stomach lining, having entered the body via the digestive tract, and so are dependent on rhinoceros species to reproduce.


Consequences

Coextinction can mean loss of biodiversity and diversification. Coextinctions can influence not only parasite and mutualist diversification but also their hosts. Arguably, parasites facilitate host diversification through sexual selection. That loss of parasites can reduce host diversification rates. Coextinction can also result in loss of evolutionary history. The extinction of related hosts can lead to the extinction of related parasites. The loss of history is likely to be greater than the loss expected, were species to go extinct at random. Furthermore, if coextinctions are clustered, it is more likely that coextinction can produce non-random trait loss. Species that are at risk of coextinction are expected to be larger because rare hosts tend to be larger and larger hosts have larger parasites. They can also be expected to have lengthy generation times or higher tropic positions. Coextinction can extend beyond biodiversity and has direct and indirect consequences from the communities of lost species. One main consequence of coextinction that goes beyond biodiversity is mutualism, by loss of food production with a decline in threatened pollinators. Losses of parasites can have negative impacts on humans or the species. In rare hosts, losses of specialist parasites can predispose hosts to infection by emergent parasites. Furthermore, relating to the consequences of removing specialist parasites from rare hosts, is the problem of where the parasites will go once their host is extinct. If the parasites are dependent on only those species than there are parasite species that are at risk of extinction through co-endangerment. On the other hand, if they are able to find and switch onto alternative hosts, those hosts can turn out to be humans. Either way, the loss of parasites by co extinction or the acquiring of new parasites by alternative hosts, proves to be a major issue. Coextinction can go beyond the decreased biodiversity, it can range into various biomes and link various ecosystems. A study conducted in
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
has shown that extinction of a
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
-associated fish species of average size would eventually result in the co-extinction of at least ten species of parasites.


Risks

The host Specificity and Life Cycle is a major factor in the risk of coextinction. Species of mutalists, parasites, and many free-living insects that have staged life cycles are more likely to be a victim of coextinction. This is due to the fact that these organisms may depend on multiple hosts throughout their lives in comparison to simple life cycled organisms. Also, if organisms are evolutionary flexible, then these organisms may escape extinction. The area with that has the greatest effect of coextinction is the tropics. There is a continued disappearance in the habitat, human intervention, and a great loss in vital
ecosystem services Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...
. This is threatening because the tropics contain 2/3 of the all known species but they aren't in a situation where they can be fully taken care of. Along with forest loss other risk factors include: coastal development,
overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term ap ...
of wildlife, and habitat conversion, that also affect human well-being. In an effort to find a stop to coextinction, researchers have found that the first step would be to conserve the host species in which other species are dependent on. These hosts serve as major components for their habitat and need them to survive. In deciding what host to protect, it is important to choose one that can benefit an array of other dependent species.


See also

*
Dodo The dodo (''Raphus cucullatus'') is an extinct flightless bird that was endemic to the island of Mauritius, which is east of Madagascar in the Indian Ocean. The dodo's closest genetic relative was the also-extinct Rodrigues solitaire. ...
and
tambalacoque ''Sideroxylon grandiflorum'', known as tambalacoque or dodo tree, is a long-lived mesocaul tree in the sapote family Sapotaceae, superfiially resembling the unrelated Plumeria, but the Dodo Tree's flowers and fruit are cauliflorous. endemic t ...
, for a supposed case of near-coextinction that turned out to be much more complex


References


External links


The Extinction Website
{{extinction Extinction