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Coevolution
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{{Short description|Two or more species influencing each other's evolution}} {{Good article}} [[File:Dasyscolia ciliata.jpg|thumb|upright=1.5|The pollinating wasp ''[[Dasyscolia ciliata]]'' in [[pseudocopulation]] with a flower of ''[[Ophrys speculum]]''<ref name=Pijl/>]] 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 as [[gene-culture coevolution]]. [[Charles Darwin]] mentioned evolutionary interactions between [[flowering plant]]s and [[insect]]s in ''[[On the Origin of Species]]'' (1859). Although he did not use the word coevolution, he suggested how plants and insects could evolve through reciprocal evolutionary changes. Naturalists in the late 1800s studied other examples of how interactions among species could result in reciprocal evolutionary change. Beginning in the 1940s, plant pathologists developed breeding programs that were examples of human-induced coevolution. Development of new crop plant varieties that were resistant to some diseases favored rapid evolution in pathogen populations to overcome those plant defenses. That, in turn, required the development of yet new resistant crop plant varieties, producing an ongoing cycle of reciprocal evolution in crop plants and diseases that continues to this day. Coevolution as a major topic for study in nature expanded rapidly from the 1960s, when Daniel H. Janzen showed coevolution between [[acacia]]s and ants (see below) and [[Paul R. Ehrlich]] and [[Peter H. Raven]] suggested how [[Escape and radiate coevolution|coevolution between plants and butterflies]] may have contributed to the diversification of species in both groups. The theoretical underpinnings of coevolution are now well-developed (e.g., the geographic mosaic theory of coevolution), and demonstrate that coevolution can play an important role in driving major evolutionary transitions such as the evolution of sexual reproduction or shifts in [[ploidy]].<ref name="itct">{{cite book |last=Nuismer |first=Scott |date=2017|title=Introduction to Coevolutionary Theory |url=https://www.macmillanlearning.com/Catalog/product/introductiontocoevolutionarytheory-firstedition-nuismer/valueoptions |location=New York |publisher=W.F. Freeman |page=395 |isbn=978-1-319-10619-5 |access-date=2019-05-02 |archive-url=https://web.archive.org/web/20190502204606/https://www.macmillanlearning.com/Catalog/product/introductiontocoevolutionarytheory-firstedition-nuismer/valueoptions |archive-date=2019-05-02 |url-status=dead}}</ref><ref name="Thompson, John N">{{Cite book |title=Relentless evolution |last=Thompson |first=John N. |isbn=978-0-226-01861-4 |location=Chicago |oclc=808684836 |date=2013-04-15}}</ref> More recently, it has also been demonstrated that coevolution can influence the structure and function of ecological communities, the evolution of groups of mutualists such as plants and their pollinators, and the dynamics of infectious disease.<ref name="itct" /><ref>{{Cite journal |last1=Guimarães |first1=Paulo R. |last2=Pires |first2=Mathias M. |last3=Jordano |first3=Pedro |last4=Bascompte|first4=Jordi |last5=Thompson |first5=John N. |date=October 2017 |title=Indirect effects drive coevolution in mutualistic networks |journal=Nature |volume=550 |issue=7677 |pages=511–514 |doi=10.1038/nature24273 |pmid=29045396 |bibcode=2017Natur.550..511G |s2cid=205261069}}</ref> Each party in a coevolutionary relationship exerts [[evolutionary pressure|selective pressures]] on the other, thereby affecting each other's evolution. Coevolution includes many forms of [[mutualism (biology)|mutualism]], [[host–parasite coevolution|host-parasite]], and [[predation|predator-prey]] relationships between species, as well as [[intraspecific competition|competition within]] or [[interspecific competition|between species]]. In many cases, the selective pressures drive an [[evolutionary arms race]] between the species involved. '''Pairwise''' or '''specific coevolution''', between exactly two species, is not the only possibility; in '''multi-species coevolution''', which is sometimes called '''guild''' or '''diffuse coevolution''', several to many species may evolve a trait or a group of traits in reciprocity with a set of traits in another species, as has happened between the flowering plants and [[pollinator|pollinating]] insects such as [[bee]]s, [[fly|flies]], and [[beetle]]s. There are a suite of specific hypotheses on the mechanisms by which groups of species coevolve with each other.<ref name="Thompson, John N. 2005">{{Cite book|last=Thompson |first=John N. |title=The geographic mosaic of coevolution |date=2005|publisher=University of Chicago Press |isbn=978-0-226-11869-7 |location=Chicago |oclc=646854337}}</ref> Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as [[computer science]], [[sociology]], and [[astronomy]].
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