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{{Short description|Two or more species influencing each other's evolution}}

{{Short description|Two or more species influencing each other's evolution}}

{{Good article}}

{{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/>]]

[[File:Dasyscolia ciliata.jpg|thumb|upright=1.5|授粉的黄蜂 ''Dasyscolia ciliata'' 在与''Ophrys speculum''花进行拟交配<ref name=Pijl/>|链接=Special:FilePath/Dasyscolia_ciliata.jpg]]

{{Evolutionary biology}}

{{Evolutionary biology}}

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]].

在生物学中, 当两个或多个物种通过自然选择过程相互影响彼此各自的演化时，就会发生'''共同演化'''。该词语有时用于同一物种中存在相互影响和演化的两个特征，例如基因和文化的共同演化。

 

 

 

 

 

 

[[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.

[[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.

Charles Darwin mentioned evolutionary interactions between flowering plants and insects 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.

Charles Darwin mentioned evolutionary interactions between flowering plants and insects 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 after the middle 1960s, when Daniel H. Janzen showed coevolution between acacias 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, 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|language=en|volume=550|issue=7677|pages=511–514|doi=10.1038/nature24273|pmid=29045396|issn=1476-4687|bibcode=2017Natur.550..511G|s2cid=205261069}}</ref>

Coevolution as a major topic for study in nature expanded rapidly after the middle 1960s, when Daniel H. Janzen showed coevolution between acacias 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. 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[[Javascript:;|展示]] of infectious disease.

Coevolution as a major topic for study in nature expanded rapidly after the middle 1960s, when Daniel H. Janzen showed coevolution between acacias and ants (see below) and Paul R. Ehrlich and Peter H. Raven suggested how 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. 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.

Coevolution as a major topic for study in nature expanded rapidly after the middle 1960s, when Daniel H. Janzen showed coevolution between acacias and ants (see below) and Paul R. Ehrlich and Peter H. Raven suggested how 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. 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.

共同进化作为自然界研究的一个主要课题，在20世纪60年代中期之后迅速扩大，当时丹尼尔 · h · 詹森(Daniel h. Janzen)展示了金合欢和蚂蚁之间的共同进化(见下文) ，保罗 · r · 埃利希(Paul r. Ehrlich)和彼得 · h · 雷文(Peter h. Raven)提出，植物和蝴蝶之间的共同进化可能促进了两个群体的物种多样化。共同进化的理论基础现在已经很成熟了(例如，共同进化的地理镶嵌理论) ，并且证明了共同进化在推动主要的进化转变中扮演着重要的角色，例如有性生殖的演化或者倍性的变化。最近，共同进化也被证明可以影响生态群落的结构和功能，相互作用者群体的进化，例如植物和它们的传粉者，以及传染病的动态。

共同演化作为自然界研究的一个主要课题，在20世纪60年代中期之后迅速扩大，丹尼尔·H·詹森（Daniel H. Janzen）展示了金合欢和蚂蚁之间的共同演化（见下文），保罗·R·埃利希（Paul R. Ehrlich）和彼得·H·雷文（Peter H. Raven）提出植物和蝴蝶之间的共同演化可能促进了两个群体的物种多样化。如今共同进化的理论基础已经颇为成熟（例如共同进化的地理镶嵌理论），而且向我们表明了共同演化在推动主要的进化转变中扮演着重要的角色，例如有性生殖的演化或者倍性的变化。<ref name="Thompson, John N">{{Cite book|title=Relentless evolution|last=Thompson, John N.|isbn=978-0-226-01861-4|location=Chicago|oclc=808684836|date = 2013-04-15}}</ref><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>最近，共同进化也被证实可以影响生态群落的结构和功能以及共生群体的演化，例如植物和它们的传粉者，以及传染病的动态过程。<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|language=en|volume=550|issue=7677|pages=511–514|doi=10.1038/nature24273|pmid=29045396|issn=1476-4687|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|title=The geographic mosaic of coevolution|last=Thompson, John N.|date=2005|publisher=University of Chicago Press|isbn=978-0-226-11869-7|location=Chicago|oclc=646854337}}</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.

Each party in a coevolutionary relationship exerts selective pressures on the other, thereby affecting each other's evolution. Coevolution includes many forms of mutualism, host-parasite, and predator-prey relationships between species, as well as competition within or 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 pollinating insects such as bees, flies, and beetles. There are a suite of specific hypotheses on the mechanisms by which groups of species coevolve with each other.

Each party in a coevolutionary relationship exerts selective pressures on the other, thereby affecting each other's evolution. Coevolution includes many forms of mutualism, host-parasite, and predator-prey relationships between species, as well as competition within or 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 pollinating insects such as bees, flies, and beetles. There are a suite of specific hypotheses on the mechanisms by which groups of species coevolve with each other.

共同进化关系中的每一方都向对方施加选择压力，从而影响彼此的进化。共同进化包括多种形式的互利共生、宿主-寄生、物种间的捕食-被捕食关系以及物种内部或物种间的竞争。在许多情况下，选择压力驱动了相关物种之间进化的军备竞赛。在多物种共同进化中，有时被称为群体共同进化或分散共同进化，几个到多个物种可能进化出一个特征或一组特征，这些特征与另一个物种的一系列特征相互作用，就像被子植物与蜜蜂、苍蝇和甲虫等传粉昆虫之间发生的情况一样。关于物种群之间共同进化的机制，有一套具体的假说。

共同演化关系中的每一方都会向对方施加选择压，从而影响彼此的演化。共同演化包括各种形式的互利共生、宿主-寄生、物种间的捕食-被捕食关系以及物种内部或物种间的竞争。在许多情况下，选择压驱动了相关物种之间进化的较量。在特定两个物种之间中，'''两两'''和'''单独'''的共同演化的存在并不是唯一的可能；在'''多物种共同演化'''，有时被称为散协同演化中，几个到多个物种可能进化出同一个特征或同一组特征，这些特征与另一个物种的一系列特征相互作用，就像被子植物与蜜蜂、苍蝇和甲虫等传粉昆虫之间发生的情况那样。关于物种群之间共同进化的机制，有一套具体的假说。<ref name="Thompson, John N. 2005">{{Cite book|title=The geographic mosaic of coevolution|last=Thompson, John N.|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]].

Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as [[computer science]], [[sociology]], and [[astronomy]].

Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as computer science, sociology, and astronomy.

Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as computer science, sociology, and astronomy.

==Mutualism==

==Mutualism==

= = 被子植物 = =  

= = 被子植物 = =  

Flowers appeared and diversified relatively suddenly in the fossil record, creating what [[Charles Darwin]] described as the "abominable mystery" of how they had evolved so quickly; he considered whether coevolution could be the explanation.<ref name="CardinalDanforth2013"/><ref>{{cite journal |author=Friedman, W. E. |date=January 2009 |title=The meaning of Darwin's 'abominable mystery' |journal=Am. J. Bot. |volume=96 |issue=1 |pages=5–21 |doi=10.3732/ajb.0800150 |url=http://www.amjbot.org/content/96/1/5.full |pmid=21628174}}</ref> He first mentioned coevolution as a possibility in ''[[On the Origin of Species]]'', and developed the concept further in ''[[Fertilisation of Orchids]]'' (1862).<ref name=t24>{{cite book |first=John N. |last=Thompson |title=The coevolutionary process |publisher=[[University of Chicago Press]] |location=Chicago |year=1994 |isbn=978-0-226-79760-1 |url=https://books.google.com/books?id=AyXPQzEwqPIC&q=Wallace+%22creation+by+law%22+Angr%C3%A6cum&pg=PA27 |access-date=2009-07-27}}</ref><ref name=origins94>{{cite book |last=Darwin |first=Charles |year=1859 |title=On the Origin of Species |edition=1st |location=London |publisher=John Murray |url=http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text&pageseq=1 |access-date=2009-02-07}}</ref><ref name=orchids1>{{cite book |last=Darwin |first=Charles |year=1877 |title=On the various contrivances by which British and foreign orchids are fertilised by insects, and on the good effects of intercrossing |location=London |publisher=John Murray |edition=2nd |url=http://darwin-online.org.uk/content/frameset?itemID=F801&viewtype=text&pageseq=1 |access-date=2009-07-27}}</ref>

Flowers appeared and diversified relatively suddenly in the fossil record, creating what [[Charles Darwin]] described as the "abominable mystery" of how they had evolved so quickly; he considered whether coevolution could be the explanation.<ref name="CardinalDanforth2013"/><ref>{{cite journal |author=Friedman, W. E. |date=January 2009 |title=The meaning of Darwin's 'abominable mystery' |journal=Am. J. Bot. |volume=96 |issue=1 |pages=5–21 |doi=10.3732/ajb.0800150 |url=http://www.amjbot.org/content/96/1/5.full |pmid=21628174}}</ref> He first mentioned coevolution as a possibility in ''[[On the Origin of Species]]'', and developed the concept further in ''[[Fertilisation of Orchids]]'' (1862).<ref name="t24">{{cite book |first=John N. |last=Thompson |title=The coevolutionary process |publisher=[[University of Chicago Press]] |location=Chicago |year=1994 |isbn=978-0-226-79760-1 |url=https://books.google.com/books?id=AyXPQzEwqPIC&q=Wallace+%22creation+by+law%22+Angr%C3%A6cum&pg=PA27 |access-date=2009-07-27}}</ref><ref name="origins94">{{cite book |last=Darwin |first=Charles |year=1859 |title=On the Origin of Species |edition=1st |location=London |publisher=John Murray |url=http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text&pageseq=1 |access-date=2009-02-07}}</ref><ref name="orchids1">{{cite book |last=Darwin |first=Charles |year=1877 |title=On the various contrivances by which British and foreign orchids are fertilised by insects, and on the good effects of intercrossing |location=London |publisher=John Murray |edition=2nd |url=http://darwin-online.org.uk/content/frameset?itemID=F801&viewtype=text&pageseq=1 |access-date=2009-07-27}}</ref>

Flowers appeared and diversified relatively suddenly in the fossil record, creating what Charles Darwin described as the "abominable mystery" of how they had evolved so quickly; he considered whether coevolution could be the explanation. He first mentioned coevolution as a possibility in On the Origin of Species, and developed the concept further in Fertilisation of Orchids (1862).

Flowers appeared and diversified relatively suddenly in the fossil record, creating what Charles Darwin described as the "abominable mystery" of how they had evolved so quickly; he considered whether coevolution could be the explanation. He first mentioned coevolution as a possibility in On the Origin of Species, and developed the concept further in Fertilisation of Orchids (1862).

====Insects and insect-pollinated flowers====

====Insects and insect-pollinated flowers====

{{Further|Entomophily}}

{{Further|Entomophily}}

[[File:Apis mellifera - Melilotus albus - Keila.jpg|thumb|upright|[[Honey bee]] taking a reward of [[nectar]] and collecting pollen in its [[pollen basket]]s from [[Melilotus albus|white melilot]] flowers]]

[[File:Apis mellifera - Melilotus albus - Keila.jpg|thumb|upright|[[Honey bee]] taking a reward of [[nectar]] and collecting pollen in its [[pollen basket]]s from [[Melilotus albus|white melilot]] flowers|链接=Special:FilePath/Apis_mellifera_-_Melilotus_albus_-_Keila.jpg]]

Modern [[entomophily|insect-pollinated (entomophilous) flowers]] are conspicuously coadapted with insects to ensure pollination and in return to reward the [[pollinator]]s with nectar and pollen. The two groups have coevolved for over 100 million years, creating a complex network of interactions. Either they evolved together, or at some later stages they came together, likely with pre-adaptations, and became mutually adapted.<ref name=Lunau>{{cite journal |last1=Lunau |first1=Klaus |title=Adaptive radiation and coevolution — pollination biology case studies |journal=Organisms Diversity & Evolution |date=2004 |volume=4 |issue=3 |pages=207–224 |doi=10.1016/j.ode.2004.02.002 }}</ref><ref>{{cite book |author=Pollan, Michael |title=The Botany of Desire: A Plant's-eye View of the World |publisher=Bloomsbury |isbn=978-0-7475-6300-6 |title-link=The Botany of Desire |year=2003}}</ref>

Modern [[entomophily|insect-pollinated (entomophilous) flowers]] are conspicuously coadapted with insects to ensure pollination and in return to reward the [[pollinator]]s with nectar and pollen. The two groups have coevolved for over 100 million years, creating a complex network of interactions. Either they evolved together, or at some later stages they came together, likely with pre-adaptations, and became mutually adapted.<ref name=Lunau>{{cite journal |last1=Lunau |first1=Klaus |title=Adaptive radiation and coevolution — pollination biology case studies |journal=Organisms Diversity & Evolution |date=2004 |volume=4 |issue=3 |pages=207–224 |doi=10.1016/j.ode.2004.02.002 }}</ref><ref>{{cite book |author=Pollan, Michael |title=The Botany of Desire: A Plant's-eye View of the World |publisher=Bloomsbury |isbn=978-0-7475-6300-6 |title-link=The Botany of Desire |year=2003}}</ref>

一些非常成功的昆虫群体---- 尤其是膜翅目(黄蜂、蜜蜂和蚂蚁)和鳞翅目(蝴蝶和飞蛾)以及许多种双翅目(苍蝇)和鞘翅目(甲虫)---- 在白垩纪(1.45亿至6.6亿年前)与被子植物共同进化。最早的蜜蜂，今天重要的传粉者，出现在白垩纪早期。一群蜜蜂的姐妹黄蜂与被子植物同时进化，鳞翅目也是如此。此外，所有主要的蜜蜂群首次出现在白垩纪中期和晚期之间，同时出现的是真根植物的辐射适应(占所有被子植物的四分之三) ，当时被子植物成为世界上陆地上的主要植物。

一些非常成功的昆虫群体---- 尤其是膜翅目(黄蜂、蜜蜂和蚂蚁)和鳞翅目(蝴蝶和飞蛾)以及许多种双翅目(苍蝇)和鞘翅目(甲虫)---- 在白垩纪(1.45亿至6.6亿年前)与被子植物共同进化。最早的蜜蜂，今天重要的传粉者，出现在白垩纪早期。一群蜜蜂的姐妹黄蜂与被子植物同时进化，鳞翅目也是如此。此外，所有主要的蜜蜂群首次出现在白垩纪中期和晚期之间，同时出现的是真根植物的辐射适应(占所有被子植物的四分之三) ，当时被子植物成为世界上陆地上的主要植物。

At least three aspects of flowers appear to have coevolved between flowering plants and insects, because they involve communication between these organisms. Firstly, flowers communicate with their pollinators by scent; insects use this scent to determine how far away a flower is, to approach it, and to identify where to land and finally to feed. Secondly, flowers attract insects with patterns of stripes leading to the rewards of nectar and pollen, and colours such as blue and ultraviolet, to which their eyes are sensitive; in contrast, bird-pollinated flowers tend to be red or orange. Thirdly, flowers such as [[Ophrys|some orchids]] mimic females of particular insects, deceiving males into [[pseudocopulation]].<ref name=Bristol/><ref name=Pijl>{{cite book |first1=Leendert |last1=van der Pijl |first2=Calaway H. |last2=Dodson |title=Orchid Flowers: Their Pollination and Evolution |chapter-url=https://archive.org/details/orchidflowersthe0000pijl |chapter-url-access=registration |chapter=Chapter 11: Mimicry and Deception |publisher=[[University of Miami]] Press |location=Coral Gables |year=1966 |pages=[https://archive.org/details/orchidflowersthe0000pijl/page/129 129–141] |isbn=978-0-87024-069-0}}</ref>

At least three aspects of flowers appear to have coevolved between flowering plants and insects, because they involve communication between these organisms. Firstly, flowers communicate with their pollinators by scent; insects use this scent to determine how far away a flower is, to approach it, and to identify where to land and finally to feed. Secondly, flowers attract insects with patterns of stripes leading to the rewards of nectar and pollen, and colours such as blue and ultraviolet, to which their eyes are sensitive; in contrast, bird-pollinated flowers tend to be red or orange. Thirdly, flowers such as [[Ophrys|some orchids]] mimic females of particular insects, deceiving males into [[pseudocopulation]].<ref name=Bristol/><ref name="Pijl">{{cite book |first1=Leendert |last1=van der Pijl |first2=Calaway H. |last2=Dodson |title=Orchid Flowers: Their Pollination and Evolution |chapter-url=https://archive.org/details/orchidflowersthe0000pijl |chapter-url-access=registration |chapter=Chapter 11: Mimicry and Deception |publisher=[[University of Miami]] Press |location=Coral Gables |year=1966 |pages=[https://archive.org/details/orchidflowersthe0000pijl/page/129 129–141] |isbn=978-0-87024-069-0}}</ref>

At least three aspects of flowers appear to have coevolved between flowering plants and insects, because they involve communication between these organisms. Firstly, flowers communicate with their pollinators by scent; insects use this scent to determine how far away a flower is, to approach it, and to identify where to land and finally to feed. Secondly, flowers attract insects with patterns of stripes leading to the rewards of nectar and pollen, and colours such as blue and ultraviolet, to which their eyes are sensitive; in contrast, bird-pollinated flowers tend to be red or orange. Thirdly, flowers such as some orchids mimic females of particular insects, deceiving males into pseudocopulation.

At least three aspects of flowers appear to have coevolved between flowering plants and insects, because they involve communication between these organisms. Firstly, flowers communicate with their pollinators by scent; insects use this scent to determine how far away a flower is, to approach it, and to identify where to land and finally to feed. Secondly, flowers attract insects with patterns of stripes leading to the rewards of nectar and pollen, and colours such as blue and ultraviolet, to which their eyes are sensitive; in contrast, bird-pollinated flowers tend to be red or orange. Thirdly, flowers such as some orchids mimic females of particular insects, deceiving males into pseudocopulation.

====Birds and bird-pollinated flowers====

====Birds and bird-pollinated flowers====

{{Further|Ornithophily}}

{{Further|Ornithophily}}

[[File:Purple-throated carib hummingbird feeding.jpg|thumb|left|[[Purple-throated carib]] feeding from and pollinating a flower]]

[[File:Purple-throated carib hummingbird feeding.jpg|thumb|left|[[Purple-throated carib]] feeding from and pollinating a flower|链接=Special:FilePath/Purple-throated_carib_hummingbird_feeding.jpg]]

[[Hummingbird]]s and ornithophilous (bird-pollinated) flowers have evolved a [[mutualism (biology)|mutualistic]] relationship. The flowers have [[nectar]] suited to the birds' diet, their color suits the birds' vision and their shape fits that of the birds' bills. The blooming times of the flowers have also been found to coincide with hummingbirds' breeding seasons. The floral characteristics of ornithophilous plants vary greatly among each other compared to closely related insect-pollinated species. These flowers also tend to be more ornate, complex, and showy than their insect pollinated counterparts. It is generally agreed that plants formed coevolutionary relationships with insects first, and ornithophilous species diverged at a later time. There is not much scientific support for instances of the reverse of this divergence: from ornithophily to insect pollination. The diversity in floral phenotype in ornithophilous species, and the relative consistency observed in bee-pollinated species can be attributed to the direction of the shift in pollinator preference.<ref>{{cite journal |last1=Kay |first1=Kathleen M.|last2=Reeves |first2=Patrick A. |last3=Olmstead |first3=Richard G. |last4=Schemske|first4=Douglas W. |s2cid=2991957|title=Rapid speciation and the evolution of hummingbird pollination in neotropical Costus subgenus Costus (Costaceae): evidence from nrDNA ITS and ETS sequences |journal=American Journal of Botany |date=2005 |volume=92 |issue=11|pages=1899–1910 |doi=10.3732/ajb.92.11.1899 |pmid=21646107|doi-access=free }}</ref>

[[Hummingbird]]s and ornithophilous (bird-pollinated) flowers have evolved a [[mutualism (biology)|mutualistic]] relationship. The flowers have [[nectar]] suited to the birds' diet, their color suits the birds' vision and their shape fits that of the birds' bills. The blooming times of the flowers have also been found to coincide with hummingbirds' breeding seasons. The floral characteristics of ornithophilous plants vary greatly among each other compared to closely related insect-pollinated species. These flowers also tend to be more ornate, complex, and showy than their insect pollinated counterparts. It is generally agreed that plants formed coevolutionary relationships with insects first, and ornithophilous species diverged at a later time. There is not much scientific support for instances of the reverse of this divergence: from ornithophily to insect pollination. The diversity in floral phenotype in ornithophilous species, and the relative consistency observed in bee-pollinated species can be attributed to the direction of the shift in pollinator preference.<ref>{{cite journal |last1=Kay |first1=Kathleen M.|last2=Reeves |first2=Patrick A. |last3=Olmstead |first3=Richard G. |last4=Schemske|first4=Douglas W. |s2cid=2991957|title=Rapid speciation and the evolution of hummingbird pollination in neotropical Costus subgenus Costus (Costaceae): evidence from nrDNA ITS and ETS sequences |journal=American Journal of Botany |date=2005 |volume=92 |issue=11|pages=1899–1910 |doi=10.3732/ajb.92.11.1899 |pmid=21646107|doi-access=free }}</ref>

===Fig reproduction and fig wasps===

===Fig reproduction and fig wasps===

[[File:Ficus plant.jpg|thumb|left|A [[Common fig|fig]] exposing its many tiny matured, seed-bearing [[gynoecia]]. These are pollinated by the fig wasp, ''[[Blastophaga psenes]]''. In the cultivated fig, there are also asexual varieties.<ref name=Suleman/>]]

[[File:Ficus plant.jpg|thumb|left|A [[Common fig|fig]] exposing its many tiny matured, seed-bearing [[gynoecia]]. These are pollinated by the fig wasp, ''[[Blastophaga psenes]]''. In the cultivated fig, there are also asexual varieties.<ref name=Suleman/>|链接=Special:FilePath/Ficus_plant.jpg]]

{{Main|Reproductive coevolution in Ficus}}

{{Main|Reproductive coevolution in Ficus}}

The genus ''[[Ficus]]'' is composed of 800 species of vines, shrubs, and trees, including the cultivated fig, defined by their [[syconium]]s, the fruit-like vessels that either hold female flowers or pollen on the inside. Each fig species has its own [[fig wasp]] which (in most cases) pollinates the fig, so a tight mutual dependence has evolved and persisted throughout the genus.<ref name=Suleman>{{cite journal |last1=Suleman |first1=Nazia |last2=Sait |first2=Steve |last3=Compton |first3=Stephen G. |title=Female figs as traps: Their impact on the dynamics of an experimental fig tree-pollinator-parasitoid community |journal=Acta Oecologica |volume=62 |year=2015 |pages=1–9 |doi=10.1016/j.actao.2014.11.001 |bibcode=2015AcO....62....1S|url=http://eprints.whiterose.ac.uk/85568/7/Female%20plants%20as%20traps%20paper%20%283%29.pdf }}</ref>

The genus ''[[Ficus]]'' is composed of 800 species of vines, shrubs, and trees, including the cultivated fig, defined by their [[syconium]]s, the fruit-like vessels that either hold female flowers or pollen on the inside. Each fig species has its own [[fig wasp]] which (in most cases) pollinates the fig, so a tight mutual dependence has evolved and persisted throughout the genus.<ref name=Suleman>{{cite journal |last1=Suleman |first1=Nazia |last2=Sait |first2=Steve |last3=Compton |first3=Stephen G. |title=Female figs as traps: Their impact on the dynamics of an experimental fig tree-pollinator-parasitoid community |journal=Acta Oecologica |volume=62 |year=2015 |pages=1–9 |doi=10.1016/j.actao.2014.11.001 |bibcode=2015AcO....62....1S|url=http://eprints.whiterose.ac.uk/85568/7/Female%20plants%20as%20traps%20paper%20%283%29.pdf }}</ref>

榕属植物由800种藤本植物、灌木和乔木组成，其中包括栽培的无花果。每一种榕树都有自己的榕小蜂(在大多数情况下)为榕小蜂授粉，所以这个属中形成了一种紧密的相互依赖关系。

榕属植物由800种藤本植物、灌木和乔木组成，其中包括栽培的无花果。每一种榕树都有自己的榕小蜂(在大多数情况下)为榕小蜂授粉，所以这个属中形成了一种紧密的相互依赖关系。

[[File:Ant - Pseudomyrmex species, on Bull Thorn Acacia (Acacia cornigera) with Beltian bodies, Caves Branch Jungle Lodge, Belmopan, Belize - 8505045055.jpg|thumb|right|''Pseudomyrmex'' ant on bull thorn acacia (''[[Vachellia cornigera]]'') with Beltian bodies that provide the ants with protein<ref name="Hölldobler-532"/>]]

[[File:Ant - Pseudomyrmex species, on Bull Thorn Acacia (Acacia cornigera) with Beltian bodies, Caves Branch Jungle Lodge, Belmopan, Belize - 8505045055.jpg|thumb|right|''Pseudomyrmex'' ant on bull thorn acacia (''[[Vachellia cornigera]]'') with Beltian bodies that provide the ants with protein<ref name="Hölldobler-532"/>|链接=Special:FilePath/Ant_-_Pseudomyrmex_species,_on_Bull_Thorn_Acacia_(Acacia_cornigera)_with_Beltian_bodies,_Caves_Branch_Jungle_Lodge,_Belmopan,_Belize_-_8505045055.jpg]]

===Acacia ants and acacias===

===Acacia ants and acacias===

寄生虫与宿主的关系可能导致了有性生殖的流行，而不是更有效率的无性生殖。看起来，当寄生虫感染宿主时，有性生殖提供了一个更好的机会来发展抗药性(通过下一代的变异) ，使得适应性的有性生殖变异性在无性生殖中看不到，这就产生了另一代的有机体，易受同一寄生虫的感染。宿主和寄生虫之间的共同进化可能相应地导致了正常人群中的许多遗传多样性，包括血浆多态性、蛋白多态性和组织相容性系统。

寄生虫与宿主的关系可能导致了有性生殖的流行，而不是更有效率的无性生殖。看起来，当寄生虫感染宿主时，有性生殖提供了一个更好的机会来发展抗药性(通过下一代的变异) ，使得适应性的有性生殖变异性在无性生殖中看不到，这就产生了另一代的有机体，易受同一寄生虫的感染。宿主和寄生虫之间的共同进化可能相应地导致了正常人群中的许多遗传多样性，包括血浆多态性、蛋白多态性和组织相容性系统。

[[File:Reed warbler cuckoo.jpg|upright|thumb|[[Brood parasite]]: [[Eurasian reed warbler]] raising a [[common cuckoo]]<ref name=Weiblen/>]]

[[File:Reed warbler cuckoo.jpg|upright|thumb|[[Brood parasite]]: [[Eurasian reed warbler]] raising a [[common cuckoo]]<ref name=Weiblen/>|链接=Special:FilePath/Reed_warbler_cuckoo.jpg]]

===Brood parasites===

===Brood parasites===

==Predators and prey==

==Predators and prey==

[[File:Leopard kill - KNP - 001.jpg|thumb|left|Predator and prey: a [[leopard]] killing a [[bushbuck]]]]

[[File:Leopard kill - KNP - 001.jpg|thumb|left|Predator and prey: a [[leopard]] killing a [[bushbuck]]|链接=Special:FilePath/Leopard_kill_-_KNP_-_001.jpg]]

{{Main|Predation}}

{{Main|Predation}}

这同样适用于食草动物，吃植物的动物，以及它们吃的植物。1964年，Paul r. Ehrlich 和 Peter h. Raven 提出了逃逸辐射共同进化理论来描述植物和蝴蝶的进化多样性。在落基山脉，红松鼠和斑鸠(食种子的鸟)争夺海滩松的种子。松鼠通过啃咬松果鳞片来获取松子，而交喙则通过它们不寻常的交叉下颚来获取松子。在有松鼠的地方，海滩鱼的球果更重，种子更少，鳞片更薄，这使得松鼠更难获得种子。相反，如果有交喙，但没有松鼠，球果的结构较轻，但有较厚的鳞片，使交喙更难以获得种子。海滩上的锥形细胞与这两种食草动物进行着一场进化中的军备竞赛。以及接下来两页的网络文章。

这同样适用于食草动物，吃植物的动物，以及它们吃的植物。1964年，Paul r. Ehrlich 和 Peter h. Raven 提出了逃逸辐射共同进化理论来描述植物和蝴蝶的进化多样性。在落基山脉，红松鼠和斑鸠(食种子的鸟)争夺海滩松的种子。松鼠通过啃咬松果鳞片来获取松子，而交喙则通过它们不寻常的交叉下颚来获取松子。在有松鼠的地方，海滩鱼的球果更重，种子更少，鳞片更薄，这使得松鼠更难获得种子。相反，如果有交喙，但没有松鼠，球果的结构较轻，但有较厚的鳞片，使交喙更难以获得种子。海滩上的锥形细胞与这两种食草动物进行着一场进化中的军备竞赛。以及接下来两页的网络文章。

[[File:Drosophila.melanogaster.couple.2.jpg|thumb|upright|[[Sexual conflict]] has been studied in ''[[Drosophila melanogaster]]'' (shown mating, male on right).]]

[[File:Drosophila.melanogaster.couple.2.jpg|thumb|upright|[[Sexual conflict]] has been studied in ''[[Drosophila melanogaster]]'' (shown mating, male on right).|链接=Special:FilePath/Drosophila.melanogaster.couple.2.jpg]]

==Competition==

==Competition==

== Multispecies ==

== Multispecies ==

[[File:Amegilla cingulata on long tube of Acanthus ilicifolius flower.jpg|thumb|upright|Long-tongued bees and long-tubed flowers coevolved, whether pairwise or "diffusely" in groups known as guilds.<ref name=Juenger/>]]

[[File:Amegilla cingulata on long tube of Acanthus ilicifolius flower.jpg|thumb|upright|Long-tongued bees and long-tubed flowers coevolved, whether pairwise or "diffusely" in groups known as guilds.<ref name=Juenger/>|链接=Special:FilePath/Amegilla_cingulata_on_long_tube_of_Acanthus_ilicifolius_flower.jpg]]

thumb|upright|Long-tongued bees and long-tubed flowers coevolved, whether pairwise or "diffusely" in groups known as guilds.

thumb|upright|Long-tongued bees and long-tubed flowers coevolved, whether pairwise or "diffusely" in groups known as guilds.