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第1行： − 此词条由Danny生产 − 此词条暂由彩云小译翻译，翻译字数共3648，未经人工整理和审校，带来阅读不便，请见谅。 − {{Short description|Two or more species influencing each other's evolution}}+ − {{Good article}}+ − + − + − 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 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. − 1859年，查尔斯·达尔文在他的著作''物种起源''中提到了被子植物和昆虫之间的进化互动。尽管他没有使用共同进化这个词，但他提出了植物和昆虫是如何通过相互的进化变化而进化的。19世纪晚期的博物学家研究了物种间的交互如何导致彼此演变的其他例子。从20世纪40年代开始的由植物病理学家开发的育种程序就是人类诱导共同进化的例子。培育能够抵抗某些疾病作物的新品种有利于病原体种群的快速进化以克服作物的这些抵御。这反过来又需要开发新的抗性作物品种，这样就造成了在作物和疾病之间的一个持续共同演化的循环；如是的循环一直持续到了今天。+ − 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 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.+ − 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.+ − 共同进化最初是一个生物学概念，但研究人员已将其应用于计算机科学、社会学和天文学等领域。+ − + − + − Coevolution is the [[evolution]] of two or more [[species]] which reciprocally affect each other, sometimes creating a [[Mutualism (biology)|mutualistic relationship]] between the species. Such relationships can be of many different types.<ref>{{cite book |title=Coevolution |author=[[Douglas J. Futuyma|Futuyma, D. J.]] and M. Slatkin (editors) |year=1983 |publisher=[[Sinauer Associates]] |isbn=978-0-87893-228-3 |pages=whole book}}</ref><ref>{{cite book |title=The Coevolutionary Process |author=Thompson, J. N.|year=1994 |publisher=University of Chicago Press |isbn=978-0-226-79759-5 |pages=whole book}}</ref>+ + − Coevolution is the evolution of two or more species which reciprocally affect each other, sometimes creating a mutualistic relationship between the species. Such relationships can be of many different types.+ − 共同进化是两个或两个以上物种相互影响，有时在物种之间创造一种互惠关系的进化。这样的关系可以有许多不同的类型。+ − + − ===Flowering plants===+ − = = 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. He first mentioned coevolution as a possibility in On the Origin of Species, and developed the concept further in Fertilisation of Orchids (1862).+ + − 在化石记录中，花朵相对突然地出现和多样化，创造了被查尔斯 · 达尔文描述为“令人憎恶的神秘”的花朵如何如此迅速地进化; 他考虑共同进化是否可以作为解释。他第一次提到共同进化的可能性是在20世纪物种起源，并在《兰花施肥》(1862)中进一步发展了这个概念。+ − ====Insects and insect-pollinated flowers====+ − {{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|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 insect-pollinated (entomophilous) flowers are conspicuously coadapted with insects to ensure pollination and in return to reward the pollinators 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.+ − 现代昆虫传粉的花朵与昆虫显著地共生，以确保授粉，并以花蜜和花粉回报授粉者。这两个群体已经共同进化了超过1亿年，创造了一个复杂的互动网络。要么它们一起进化，要么在某些后期阶段，它们一起进化，很可能是通过预适应，变得相互适应。+ − Several highly successful [[insect]] groups—especially the [[Hymenoptera]] (wasps, bees and ants) and [[Lepidoptera]] (butterflies and moths) as well as many types of [[Diptera]] (flies) and [[Coleoptera]] (beetles)—evolved in conjunction with [[flowering plant]]s during the [[Cretaceous]] (145 to 66 million years ago). The earliest bees, important pollinators today, appeared in the early Cretaceous.<ref name=Bristol>{{cite web |title=Coevolution of angiosperms and insects |url=http://palaeo.gly.bris.ac.uk/Palaeofiles/Angiosperms/coevolution.htm |publisher=University of Bristol Palaeobiology Research Group |access-date=16 January 2017 |archive-url=https://web.archive.org/web/20161220033247/http://palaeo.gly.bris.ac.uk/Palaeofiles/Angiosperms/coevolution.htm |archive-date=20 December 2016 |url-status=dead }}</ref> A group of wasps [[sister clade|sister]] to the bees evolved at the same time as flowering plants, as did the Lepidoptera.<ref name=Bristol/> Further, all the major [[clade]]s of bees first appeared between the middle and late Cretaceous, simultaneously with the adaptive radiation of the [[eudicots]] (three quarters of all angiosperms), and at the time when the angiosperms became the world's dominant plants on land.<ref name="CardinalDanforth2013">{{cite journal |last1=Cardinal |first1=Sophie |last2=Danforth |first2=Bryan N. |title=Bees diversified in the age of eudicots |journal=Proceedings of the Royal Society B |date=2013 |doi=10.1098/rspb.2012.2686 |volume=280 |issue=1755 |pages=20122686 |pmid=23363629 |pmc=3574388}}</ref>+ − Several highly successful insect groups—especially the Hymenoptera (wasps, bees and ants) and Lepidoptera (butterflies and moths) as well as many types of Diptera (flies) and Coleoptera (beetles)—evolved in conjunction with flowering plants during the Cretaceous (145 to 66 million years ago). The earliest bees, important pollinators today, appeared in the early Cretaceous. A group of wasps sister to the bees evolved at the same time as flowering plants, as did the Lepidoptera. Further, all the major clades of bees first appeared between the middle and late Cretaceous, simultaneously with the adaptive radiation of the eudicots (three quarters of all angiosperms), and at the time when the angiosperms became the world's dominant plants on land.+ − 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 some orchids mimic females of particular insects, deceiving males into pseudocopulation.+ + + − 至少有3个方面的花似乎是被子植物和昆虫共同进化的，因为它们涉及到这些有机体之间的交流。首先，花朵通过气味与它们的传粉者交流; 昆虫利用这种气味来确定一朵花离它有多远，接近它，并确定在哪里落地，最后在哪里觅食。其次，花朵吸引昆虫的条纹图案导致花蜜和花粉的奖赏，以及蓝色和紫外线等颜色，它们的眼睛是敏感的; 相反，鸟类传粉的花朵往往是红色或橙色的。第三，像某些兰花这样的花朵模仿某些昆虫的雌性，欺骗雄性进入拟交配。+ − The [[yucca]], ''Yucca whipplei'', is pollinated exclusively by ''Tegeticula maculata'', a [[yucca moth]] that depends on the yucca for survival.<ref>{{cite journal |title=Pollination Partnerships Fact Sheet |journal=Flora of North America |year=2004 |first=Claire |last=Hemingway |pages=1–2 |url=http://www.fna.org/files/imported/Outreach/FNAfs_yucca.pdf |access-date=2011-02-18 |quote=Yucca and Yucca Moth}}</ref> The moth eats the seeds of the plant, while gathering pollen. The pollen has evolved to become very sticky, and remains on the mouth parts when the moth moves to the next flower. The yucca provides a place for the moth to lay its eggs, deep within the flower away from potential predators.<ref>{{cite journal |doi=10.1073/pnas.96.16.9178 |title=Forty million years of mutualism: Evidence for Eocene origin of the yucca-yucca moth association |journal=Proc. Natl. Acad. Sci. USA |date=August 1999 |first=Olle |last=Pellmyr |pmid=10430916 |author2=James Leebens-Mack |volume=96 |issue=16 |pmc=17753 |pages=9178–9183 |bibcode=1999PNAS...96.9178P|doi-access=free }}</ref>+ − The yucca, Yucca whipplei, is pollinated exclusively by Tegeticula maculata, a yucca moth that depends on the yucca for survival. The moth eats the seeds of the plant, while gathering pollen. The pollen has evolved to become very sticky, and remains on the mouth parts when the moth moves to the next flower. The yucca provides a place for the moth to lay its eggs, deep within the flower away from potential predators.+ − 亚卡的丝兰，只有斑点豆斑蛾才能为其授粉，斑点豆斑蛾是一种依靠丝兰生存的丝兰蛾。蛾子在采集花粉的同时吃植物的种子。花粉已经进化得非常粘，当飞蛾移动到下一朵花时，花粉仍然留在口腔部分。丝兰为蛾子提供了一个产卵的地方，在花的深处，远离潜在的捕食者。+ − ====Birds and bird-pollinated flowers====+ − {{Further|Ornithophily}} − [[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>+ − Hummingbirds and ornithophilous (bird-pollinated) flowers have evolved a 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.+ − 蜂鸟和喜鸟类(鸟类传粉)的花进化出了一种互惠的关系。这些花的花蜜适合鸟类的饮食，它们的颜色适合鸟类的视觉，它们的形状适合鸟的喙。人们还发现，这些花的开放时间与蜂鸟的繁殖季节相吻合。与昆虫传粉密切相关的物种相比，喜鸟类植物的花部特征差异很大。这些花也往往比昆虫授粉的同类更华丽、复杂和艳丽。人们普遍认为，植物首先与昆虫形成共同进化关系，喜鸟类的物种在后期分化。没有多少科学证据支持这种分歧的相反的例子: 从鸟类学到昆虫授粉。喜鸟类物种花器官表型的多样性和蜜蜂传粉物种花器官表型的相对一致性可以归因于传粉者偏好的转变方向。+ + − Flowers have converged to take advantage of similar birds.<ref name="Brown">{{cite journal |title=Convergence, Competition, and Mimicry in a Temperate Community of Hummingbird-Pollinated Flowers|author1=Brown James H. |author2=Kodric-Brown Astrid |s2cid=53604204 |journal=Ecology |year=1979 |volume=60 |issue=5 |pages=1022–1035 |doi=10.2307/1936870|jstor=1936870}}</ref> Flowers compete for pollinators, and adaptations reduce unfavourable effects of this competition. The fact that birds can fly during inclement weather makes them more efficient pollinators where bees and other insects would be inactive. Ornithophily may have arisen for this reason in isolated environments with poor insect colonization or areas with plants which flower in the winter.<ref name="Brown"/><ref>{{cite journal |last1=Cronk |first1=Quentin |last2=Ojeda |first2=Isidro |title=Bird-pollinated flowers in an evolutionary and molecular context |journal=Journal of Experimental Botany |date=2008 |volume=59 |issue=4 |pages=715–727 |doi=10.1093/jxb/ern009|pmid=18326865|doi-access=free }}</ref> Bird-pollinated flowers usually have higher volumes of nectar and higher sugar production than those pollinated by insects.<ref name="Stiles">{{cite journal |title=Geographical Aspects of Bird Flower Coevolution, with Particular Reference to Central America |author=Stiles, F. Gary |journal=Annals of the Missouri Botanical Garden |year=1981 |volume=68 |issue=2 |pages=323–351 |doi=10.2307/2398801|jstor=2398801|url=https://www.biodiversitylibrary.org/part/38387 }}</ref>+ − This meets the birds' high energy requirements, the most important determinants of flower choice.<ref name="Stiles"/> In ''[[Mimulus]]'', an increase in red pigment in petals and flower nectar volume noticeably reduces the proportion of pollination by bees as opposed to hummingbirds; while greater flower surface area increases bee pollination. Therefore, red pigments in the flowers of ''Mimulus cardinalis'' may function primarily to discourage bee visitation.<ref>{{cite journal |last1=Schemske |first1=Douglas W. |last2=Bradshaw |first2=H.D. |title=Pollinator preference and the evolution of floral traits in monkeyflowers (''Mimulus'') |journal=Proceedings of the National Academy of Sciences |date=1999 |volume=96 |issue=21 |pages=11910–11915 |doi=10.1073/pnas.96.21.11910|pmid=10518550 |bibcode=1999PNAS...9611910S |pmc=18386|doi-access=free }}</ref> In ''[[Penstemon]]'', flower traits that discourage bee pollination may be more influential on the flowers' evolutionary change than 'pro-bird' adaptations, but adaptation 'towards' birds and 'away' from bees can happen simultaneously.<ref>{{cite journal |last1=Castellanos|first1=M. C. |last2=Wilson |first2=P. |last3=Thomson |first3=J.D. |title='Anti-bee' and 'pro-bird' changes during the evolution of hummingbird pollination in Penstemon flowers |journal=Journal of Evolutionary Biology |date=2005 |volume=17 |issue=4 |pages=876–885 |doi=10.1111/j.1420-9101.2004.00729.x |pmid=15271088|doi-access=free }}</ref> However, some flowers such as ''[[Heliconia angusta]]'' appear not to be as specifically ornithophilous as had been supposed: the species is occasionally (151 visits in 120 hours of observation) visited by ''[[Trigona]]'' stingless bees. These bees are largely pollen robbers in this case, but may also serve as pollinators.<ref>{{cite journal |last1=Stein |first1=Katharina |last2=Hensen |first2=Isabell |title=Potential Pollinators and Robbers: A Study of the Floral Visitors of Heliconia Angusta (Heliconiaceae) And Their Behaviour |journal=Journal of Pollination Ecology |date=2011 |volume=4 |issue=6 |pages=39–47|doi=10.26786/1920-7603(2011)7|doi-access=free }}</ref> − Flowers have converged to take advantage of similar birds. Flowers compete for pollinators, and adaptations reduce unfavourable effects of this competition. The fact that birds can fly during inclement weather makes them more efficient pollinators where bees and other insects would be inactive. Ornithophily may have arisen for this reason in isolated environments with poor insect colonization or areas with plants which flower in the winter. Bird-pollinated flowers usually have higher volumes of nectar and higher sugar production than those pollinated by insects.+ − This meets the birds' high energy requirements, the most important determinants of flower choice. In Mimulus, an increase in red pigment in petals and flower nectar volume noticeably reduces the proportion of pollination by bees as opposed to hummingbirds; while greater flower surface area increases bee pollination. Therefore, red pigments in the flowers of Mimulus cardinalis may function primarily to discourage bee visitation. In Penstemon, flower traits that discourage bee pollination may be more influential on the flowers' evolutionary change than 'pro-bird' adaptations, but adaptation 'towards' birds and 'away' from bees can happen simultaneously. However, some flowers such as Heliconia angusta appear not to be as specifically ornithophilous as had been supposed: the species is occasionally (151 visits in 120 hours of observation) visited by Trigona stingless bees. These bees are largely pollen robbers in this case, but may also serve as pollinators. − 花朵聚集在一起，利用同类鸟类的优势。花朵争夺传粉者，适应性减少了这种竞争的不利影响。鸟类可以在恶劣天气下飞行，这一事实使它们在蜜蜂和其他昆虫不活跃的地方成为更有效的授粉者。由于这个原因，鸟食现象可能出现在孤立的环境中，这些环境中的昆虫定居能力很差，或者在冬天有植物开花的地方。鸟类传粉的花朵通常比昆虫传粉的花朵有更多的花蜜和更高的糖分产量。这符合鸟类的高能量需求，最重要的决定因素花的选择。在蜜环菌中，花瓣中红色素的增加和花蜜体积的增加明显减少了蜜蜂授粉的比例，而蜂鸟则相反; 同时花朵表面积的增加增加了蜜蜂授粉。因此，红雀花中的红色色素可能主要起到抑制蜜蜂拜访的作用。在 Penstemon，阻碍蜜蜂授粉的花朵特征可能比“支持鸟类”的适应性对花朵进化变化的影响更大，但是“适应”鸟类和“远离”蜜蜂的适应性变化可以同时发生。然而，一些花，如棉铃虫似乎并不像人们想象的那样特别喜好鸟类: 这个物种偶尔(在120小时的观察中造访了151次)被无刺的 Trigona 蜜蜂采访。在这种情况下，这些蜜蜂大部分是花粉盗窃者，但也可能充当传粉者。+ − Following their respective breeding seasons, several species of hummingbirds occur at the same locations in [[North America]], and several hummingbird flowers bloom simultaneously in these habitats. These flowers have [[convergent evolution|converged]] to a common [[morphology (biology)|morphology]] and color because these are effective at attracting the birds. Different lengths and curvatures of the [[petal#Corolla|corolla]] tubes can affect the efficiency of extraction in hummingbird species in relation to differences in bill morphology. Tubular flowers force a bird to orient its bill in a particular way when probing the flower, especially when the bill and corolla are both curved. This allows the plant to place [[pollen]] on a certain part of the bird's body, permitting a variety of morphological [[co-adaptation]]s.<ref name="Stiles"/>+ + − Following their respective breeding seasons, several species of hummingbirds occur at the same locations in North America, and several hummingbird flowers bloom simultaneously in these habitats. These flowers have converged to a common morphology and color because these are effective at attracting the birds. Different lengths and curvatures of the corolla tubes can affect the efficiency of extraction in hummingbird species in relation to differences in bill morphology. Tubular flowers force a bird to orient its bill in a particular way when probing the flower, especially when the bill and corolla are both curved. This allows the plant to place pollen on a certain part of the bird's body, permitting a variety of morphological co-adaptations.+ − 在各自的繁殖季节之后，几种蜂鸟出现在北美的同一地点，几种蜂鸟的花同时在这些栖息地开放。这些花汇聚到一个共同的形态和颜色，因为这些是有效地吸引鸟类。蜂鸟喙形态的不同决定了花冠管的长度和曲率对蜂鸟提取效率的影响。管状花朵使得鸟类在探测花朵时，尤其是当喙和花冠都是弯曲的时候，它们会以一种特殊的方式来确定喙的方位。这允许植物把花粉放在鸟身体的某一部分，允许各种形态上的协同适应。+ − Ornithophilous flowers need to be conspicuous to birds.<ref name="Stiles"/> Birds have their greatest spectral sensitivity and finest hue discrimination at the red end of the [[visual spectrum]],<ref name="Stiles"/> so red is particularly conspicuous to them. Hummingbirds may also be able to see ultraviolet "colors". The prevalence of ultraviolet patterns and nectar guides in nectar-poor entomophilous (insect-pollinated) flowers warns the bird to avoid these flowers.<ref name="Stiles"/> Each of the two subfamilies of hummingbirds, the [[Phaethornithinae]] (hermits) and the [[Trochilinae]], has evolved in conjunction with a particular set of flowers. Most Phaethornithinae species are associated with large [[monocotyledon]]ous herbs, while the Trochilinae prefer [[dicotyledon]]ous plant species.<ref name="Stiles"/>+ − <!-- could extend examples of mutualism indefinitely - might mention fish/anemone [[cleaning symbiosis]] etc. − − − Ornithophilous flowers need to be conspicuous to birds. Birds have their greatest spectral sensitivity and finest hue discrimination at the red end of the visual spectrum, so red is particularly conspicuous to them. Hummingbirds may also be able to see ultraviolet "colors". The prevalence of ultraviolet patterns and nectar guides in nectar-poor entomophilous (insect-pollinated) flowers warns the bird to avoid these flowers. Each of the two subfamilies of hummingbirds, the Phaethornithinae (hermits) and the Trochilinae, has evolved in conjunction with a particular set of flowers. Most Phaethornithinae species are associated with large monocotyledonous herbs, while the Trochilinae prefer dicotyledonous plant species.+ − + − 喜鸟类的花对鸟类来说需要引人注目。鸟类在视觉光谱的红色一端拥有最大的光谱敏感度和最好的色调辨别能力，所以红色对它们来说特别显眼。蜂鸟也可以看到紫外线的“颜色”。在缺少花蜜的昆虫传粉的花朵中，紫外线图案和花蜜向导的流行警告鸟类要避开这些花。蜂鸟的两个亚科——隐士蜂鸟亚科(phaeethornithinae，hermits)和热带蜂鸟亚科(Trochilinae，Trochilinae)的每一个亚科都与一组特定的花一起进化。大多数种类与大型单子叶植物有关，而金线菊亚科更喜欢双子叶植物。
无编辑摘要
===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/>|链接=Special:FilePath/Ficus_plant.jpg]]
[[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]]
{{Main|Reproductive coevolution in Ficus}}
{{Evolutionary biology}}
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 syconiums, 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.
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"/>|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===
= = = 相思蚂蚁和金合欢 = =
{{Main|Pseudomyrmex ferruginea}}
The [[acacia ant]] (''Pseudomyrmex ferruginea'') is an obligate plant ant that protects at least five species of "Acacia" (''[[Vachellia]]''){{efn|The acacia ant protects at least 5 species of "Acacia", now all renamed to ''Vachellia'': ''[[Vachellia chiapensis|V. chiapensis]]'', ''[[Vachellia collinsii|V. collinsii]]'', ''[[Vachellia cornigera|V. cornigera]]'', ''[[Vachellia hindsii|V. hindsii]]'', and ''[[Vachellia sphaerocephala|V. sphaerocephala]]''.}} from preying insects and from other plants competing for sunlight, and the tree provides nourishment and shelter for the ant and its larvae.<ref name="Hölldobler-532">{{cite book |last1=Hölldobler |first1=Bert |last2=Wilson |first2=Edward O. |title=The ants |publisher=Harvard University Press |year=1990 |url=https://archive.org/details/ants0000hlld |url-access=registration |isbn=978-0-674-04075-5 |pages=[https://archive.org/details/ants0000hlld/page/532 532]–533}}</ref><ref>{{cite web|last=National Geographic|title=Acacia Ant Video|url=http://video.nationalgeographic.com/video/player/animals/bugs-animals/ants-and-termites/ant_acaciatree.html|url-status=dead|archive-url=https://web.archive.org/web/20071107085438/http://video.nationalgeographic.com/video/player/animals/bugs-animals/ants-and-termites/ant_acaciatree.html|archive-date=2007-11-07}}</ref> Such mutualism is not automatic: other ant species exploit trees without reciprocating, following different [[evolutionary strategy|evolutionary strategies]]. These cheater ants impose important host costs via damage to tree reproductive organs, though their net effect on host fitness is not necessarily negative and, thus, becomes difficult to forecast.<ref>{{cite journal |doi=10.1073/pnas.1006872107 |vauthors=Palmer TM, Doak DF, Stanton ML, Bronstein JL, Kiers ET, Young TP, Goheen JR, Pringle RM |year=2010 |title=Synergy of multiple partners, including freeloaders, increases host fitness in a multispecies mutualism |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=107 |issue=40 |pages=17234–9 |pmid=20855614 |pmc=2951420 |bibcode=2010PNAS..10717234P|doi-access=free }}</ref><ref>{{cite journal |title=Kinship and incompatibility between colonies of the acacia ant ''Pseudomyrex ferruginea'' |journal=Behavioral Ecology and Sociobiology |first=Alex |last=Mintzer |author2=Vinson, S.B. |volume=17 |issue=1 |pages=75–78 |doi=10.1007/bf00299432 |jstor=4599807 |year=1985|s2cid=9538185 }}</ref>
The acacia ant (Pseudomyrmex ferruginea) is an obligate plant ant that protects at least five species of "Acacia" (Vachellia) from preying insects and from other plants competing for sunlight, and the tree provides nourishment and shelter for the ant and its larvae. Such mutualism is not automatic: other ant species exploit trees without reciprocating, following different evolutionary strategies. These cheater ants impose important host costs via damage to tree reproductive organs, though their net effect on host fitness is not necessarily negative and, thus, becomes difficult to forecast.
金合欢蚂蚁(相思树蚁)是一种专性植物蚂蚁，它能保护至少5种金合欢树(Vachellia)免受捕食昆虫和其他植物争夺阳光的伤害，并为蚂蚁及其幼虫提供营养和庇护。这种互利共生并不是自然而然的: 其他蚂蚁种类遵循不同的进化策略，不作回报地利用树木。这些骗子蚂蚁通过破坏树木的生殖器官给寄主造成重大损失，但它们对寄主适合度的净影响并不一定是负面的，因此难以预测。
==Mutualism==
==Hosts and parasites==
{{Main|Mutualism (biology)}}
{{Main|Host–parasite coevolution}}
===Parasites and sexually reproducing hosts===
[[Host–parasite coevolution]] is the coevolution of a [[host (biology)|host]] and a [[parasite]].<ref name="Woolhouse">{{cite journal |doi=10.1038/ng1202-569 |last1=Woolhouse |first1=M. E. J. |last2=Webster |first2=J. P. |last3=Domingo |first3=E. |last4=Charlesworth|first4=B. |last5=Levin |first5=B. R. |title=Biological and biomedical implications of the coevolution of pathogens and their hosts |journal=[[Nature Genetics]] |date=December 2002 |pmid=12457190 |volume=32 |issue=4 |pages=569–77 |url=http://www.era.lib.ed.ac.uk/bitstream/1842/689/2/Charlesworth_Woolhouse.pdf|hdl=1842/689 |s2cid=33145462 |hdl-access=free }}</ref> A general characteristic of many viruses, as [[obligate parasite]]s, is that they coevolved alongside their respective hosts. Correlated mutations between the two species enter them into an evolution arms race. Whichever organism, host or parasite, that cannot keep up with the other will be eliminated from their habitat, as the species with the higher average population fitness survives. This race is known as the [[Red Queen hypothesis]].<ref>{{cite journal |author=Van Valen, L. |date=1973 |title=A New Evolutionary Law |journal=Evolutionary Theory |volume=1 |pages=1–30}} cited in: [http://pespmc1.vub.ac.be/REDQUEEN.html The Red Queen Principle]</ref> The Red Queen hypothesis predicts that sexual reproduction allows a host to stay just ahead of its parasite, similar to the [[Red Queen's race]] in ''[[Through the Looking-Glass]]'': "it takes all the running ''you'' can do, to keep in the same place".<ref>{{cite book |last=Carroll |first=Lewis |author-link=Lewis Carroll |orig-year=1871 |title=Through the Looking-glass: And what Alice Found There |url=https://books.google.com/books?id=cJJZAAAAYAAJ |publisher=Macmillan |date=1875 |page=42 |quote=it takes all the running ''you'' can do, to keep in the same place.}}</ref> The host reproduces sexually, producing some offspring with immunity over its parasite, which then evolves in response.<ref>{{cite journal |doi=10.1038/srep10004 |last=Rabajante |first=J. |display-authors=etal |title=Red Queen dynamics in multi-host and multi-parasite interaction system |journal=[[Scientific Reports]] |year=2015 |volume=5 |pages=10004 |pmid=25899168 |pmc=4405699|bibcode=2015NatSR...510004R}}</ref>
Host–parasite coevolution is the coevolution of a host and a parasite. A general characteristic of many viruses, as obligate parasites, is that they coevolved alongside their respective hosts. Correlated mutations between the two species enter them into an evolution arms race. Whichever organism, host or parasite, that cannot keep up with the other will be eliminated from their habitat, as the species with the higher average population fitness survives. This race is known as the Red Queen hypothesis. cited in: The Red Queen Principle The Red Queen hypothesis predicts that sexual reproduction allows a host to stay just ahead of its parasite, similar to the Red Queen's race in Through the Looking-Glass: "it takes all the running you can do, to keep in the same place". The host reproduces sexually, producing some offspring with immunity over its parasite, which then evolves in response.
= = = 寄生虫和有性生殖的宿主 = = = 宿主-寄生虫的共同进化是宿主和寄生虫的共同进化。许多病毒作为专性寄生虫的一个普遍特征是它们与各自的宿主共同进化。这两个物种之间的相关突变使它们进入了进化的军备竞赛。无论是哪种生物、宿主或寄生物，如果不能跟上其他生物的步伐，就会从它们的栖息地消失，因为平均适合度较高的物种会幸存下来。这一种族被称为红皇后假说。红皇后原则红皇后假说预测有性生殖可以让寄主在寄生虫之前保持领先，就像爱丽丝镜中奇遇的红皇后比赛一样: “你可以尽你所能地跑，保持在同一个地方。”。宿主进行有性繁殖，产生一些对寄生虫具有免疫力的后代，然后进化为应对措施。
===Flowering plants===
The parasite–host relationship probably drove the prevalence of sexual reproduction over the more efficient asexual reproduction. It seems that when a parasite infects a host, sexual reproduction affords a better chance of developing resistance (through variation in the next generation), giving sexual reproduction variability for fitness not seen in the asexual reproduction, which produces another generation of the organism susceptible to infection by the same parasite.<ref>{{cite web |title=Sexual reproduction works thanks to ever-evolving host, parasite relationships |website=PhysOrg |url=https://phys.org/news/2011-07-sexual-reproduction-ever-evolving-host-parasite.html |date=7 July 2011}}</ref><ref>{{cite journal |author1=Morran, L.T. |author2=Schmidt, O.G. |author3=Gelarden, I.A. |author4=Parrish, R.C. II |author5= Lively, C.M. |title=Running with the Red Queen: Host-Parasite Coevolution Selects for Biparental Sex |journal=Science |volume=333 |issue=6039 |pages=216–8 |date=8 July 2011 |id=Science.1206360 |bibcode=2011Sci...333..216M |doi=10.1126/science.1206360 |pmid=21737739 |pmc=3402160}}</ref><ref>{{cite encyclopedia |author=Hogan, C. Michael |date=2010 |url=https://editors.eol.org/eoearth/wiki/Virus |title=Virus |encyclopedia=Encyclopedia of Earth |editor=Cutler Cleveland |editor2=Sidney Draggan}}</ref> Coevolution between host and parasite may accordingly be responsible for much of the genetic diversity seen in normal populations, including blood-plasma polymorphism, protein polymorphism, and histocompatibility systems.<ref>{{cite journal |author1=Anderson, R. |author2=May, R. |date=October 1982 |title=Coevolution of hosts and parasites |journal=Parasitology |volume=85 |issue=2 |pages=411–426 |doi=10.1017/S0031182000055360 |pmid=6755367}}</ref>
The parasite–host relationship probably drove the prevalence of sexual reproduction over the more efficient asexual reproduction. It seems that when a parasite infects a host, sexual reproduction affords a better chance of developing resistance (through variation in the next generation), giving sexual reproduction variability for fitness not seen in the asexual reproduction, which produces another generation of the organism susceptible to infection by the same parasite. Coevolution between host and parasite may accordingly be responsible for much of the genetic diversity seen in normal populations, including blood-plasma polymorphism, protein polymorphism, and histocompatibility systems.
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/>|链接=Special:FilePath/Reed_warbler_cuckoo.jpg]]
===Brood parasites===
{{Main|Brood parasitism}}
[[Brood parasite|Brood parasitism]] demonstrates close coevolution of host and parasite, for example in some [[cuckoo]]s. These birds do not make their own nests, but lay their eggs in nests of other species, ejecting or killing the eggs and young of the host and thus having a strong negative impact on the host's reproductive fitness. Their eggs are camouflaged as eggs of their hosts, implying that hosts can distinguish their own eggs from those of intruders and are in an evolutionary arms race with the cuckoo between camouflage and recognition. Cuckoos are counter-adapted to host defences with features such as thickened eggshells, shorter incubation (so their young hatch first), and flat backs adapted to lift eggs out of the nest.<ref name=Weiblen>{{cite web |last1=Weiblen |first1=George D. |title=Interspecific Coevolution |url=http://geo.cbs.umn.edu/Weiblen2003.pdf |publisher=Macmillan |date=May 2003}}</ref><ref>{{cite journal |last1=Rothstein |first1=S.I |year=1990 |title=A model system for coevolution: avian brood parasitism |journal=Annual Review of Ecology and Systematics |volume=21 |pages=481–508 |doi=10.1146/annurev.ecolsys.21.1.481}}</ref><ref>{{Cite book|title=Cuckoo : cheating by nature|last=Davies, N. B. (Nicholas B.), 1952-|others=McCallum, James (Wildlife artist)|date=7 April 2015|isbn=978-1-62040-952-7|edition=First U.S.|location=New York, NY|oclc=881092849}}</ref>
Brood parasitism demonstrates close coevolution of host and parasite, for example in some cuckoos. These birds do not make their own nests, but lay their eggs in nests of other species, ejecting or killing the eggs and young of the host and thus having a strong negative impact on the host's reproductive fitness. Their eggs are camouflaged as eggs of their hosts, implying that hosts can distinguish their own eggs from those of intruders and are in an evolutionary arms race with the cuckoo between camouflage and recognition. Cuckoos are counter-adapted to host defences with features such as thickened eggshells, shorter incubation (so their young hatch first), and flat backs adapted to lift eggs out of the nest.
巢寄生证明了宿主和寄生虫的密切共同进化，例如在一些杜鹃中。这些鸟不自己筑巢，而是在其他物种的巢中产卵，排出或杀死寄主的卵和幼鸟，从而对寄主的生殖适应性产生严重的负面影响。它们的卵伪装成它们寄主的卵，这意味着寄主能够区分自己的卵和入侵者的卵，并且处于一种进化的军备竞赛中，杜鹃介于伪装和识别之间。杜鹃与寄主相反，具有加厚的蛋壳、较短的孵化期(所以它们的幼鸟先孵化)以及适于将蛋提出巢外的平背等特征。
===Antagonistic coevolution===
===Antagonistic coevolution===
= = = 对抗性共同进化 = =
Antagonistic coevolution is seen in the [[harvester ant]] species ''[[Pogonomyrmex barbatus]]'' and ''[[Pogonomyrmex rugosus]]'', in a relationship both parasitic and mutualistic. The queens are unable to produce worker ants by mating with their own species. Only by crossbreeding can they produce workers. The winged females act as parasites for the males of the other species as their sperm will only produce sterile hybrids. But because the colonies are fully dependent on these hybrids to survive, it is also mutualistic. While there is no genetic exchange between the species, they are unable to evolve in a direction where they become too genetically different as this would make crossbreeding impossible.<ref name="Herrmann Cahan pp. 20141771–20141771">{{cite journal |last1=Herrmann |first1=M. |last2=Cahan |first2=S. H. |title=Inter-genomic sexual conflict drives antagonistic coevolution in harvester ants |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1797 |date=29 October 2014 |doi=10.1098/rspb.2014.1771 |pmid=25355474 |pages=20141771 |pmc=4240986}}</ref>
Antagonistic coevolution is seen in the harvester ant species Pogonomyrmex barbatus and Pogonomyrmex rugosus, in a relationship both parasitic and mutualistic. The queens are unable to produce worker ants by mating with their own species. Only by crossbreeding can they produce workers. The winged females act as parasites for the males of the other species as their sperm will only produce sterile hybrids. But because the colonies are fully dependent on these hybrids to survive, it is also mutualistic. While there is no genetic exchange between the species, they are unable to evolve in a direction where they become too genetically different as this would make crossbreeding impossible.
拮抗性的共同进化在收获蚂蚁种类 Pogonomyrmex barbatus 和 Pogonomyrmex rugosus 中可以看到，它们之间既有寄生关系也有互惠关系。蚁后无法通过与同类交配来繁殖工蚁。只有通过杂交，他们才能生产工人。有翅膀的雌性像寄生虫一样为其他物种的雄性服务，因为它们的精子只会产生不育的杂种。但由于殖民地完全依赖这些杂交种生存，这也是互惠互利的。虽然两个物种之间没有基因交换，但它们无法朝着基因差异太大的方向进化，因为这将使杂交繁殖变得不可能。
==Predators and prey==
[[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}}
[[Predator]]s and prey interact and coevolve: the predator to catch the prey more effectively, the prey to escape. The coevolution of the two mutually imposes [[selective pressure]]s. These often lead to an [[evolutionary arms race]] between prey and predator, resulting in [[anti-predator adaptation]]s.<ref>{{cite web|title=Predator-Prey Relationships|url=https://necsi.edu/predator-prey-relationships|publisher=New England Complex Systems Institute|access-date=17 January 2017}}</ref>
Predators and prey interact and coevolve: the predator to catch the prey more effectively, the prey to escape. The coevolution of the two mutually imposes selective pressures. These often lead to an evolutionary arms race between prey and predator, resulting in anti-predator adaptations.
捕食者和猎物相互作用并共同进化: 捕食者更有效地捕捉猎物，猎物逃跑。两者的共同进化相互施加选择压力。这往往导致进化的军备竞赛之间的猎物和捕食者，导致反捕食适应。
The same applies to [[herbivore]]s, animals that eat plants, and the plants that they eat. [[Paul R. Ehrlich]] and [[Peter H. Raven]] in 1964 proposed the theory of [[escape and radiate coevolution]] to describe the evolutionary diversification of plants and butterflies.<ref>{{cite journal |last1=Ehrlich |first1=Paul R. |author1-link=Paul R. Ehrlich |last2=Raven |first2=Peter H. |author2-link= Peter H. Raven |year=1964 |title=Butterflies and Plants: A Study in Coevolution |journal=Evolution |volume=18 |issue=4 |pages=586–608 |doi=10.2307/2406212 |jstor=2406212}}</ref> In the [[Rocky Mountains]], [[red squirrel]]s and [[crossbill]]s (seed-eating birds) compete for seeds of the [[lodgepole pine]]. The squirrels get at pine seeds by gnawing through the cone scales, whereas the crossbills get at the seeds by extracting them with their unusual crossed mandibles. In areas where there are squirrels, the lodgepole's cones are heavier, and have fewer seeds and thinner scales, making it more difficult for squirrels to get at the seeds. Conversely, where there are crossbills but no squirrels, the cones are lighter in construction, but have thicker scales, making it more difficult for crossbills to get at the seeds. The lodgepole's cones are in an evolutionary arms race with the two kinds of herbivore.<ref name="Berkeley">{{cite web |title=Coevolution |url=https://evolution.berkeley.edu/evolibrary/article/evo_33 |publisher=University of California Berkeley |access-date=17 January 2017}} and the two following pages of the web article.</ref>
The same applies to herbivores, animals that eat plants, and the plants that they eat. Paul R. Ehrlich and Peter H. Raven in 1964 proposed the theory of escape and radiate coevolution to describe the evolutionary diversification of plants and butterflies. In the Rocky Mountains, red squirrels and crossbills (seed-eating birds) compete for seeds of the lodgepole pine. The squirrels get at pine seeds by gnawing through the cone scales, whereas the crossbills get at the seeds by extracting them with their unusual crossed mandibles. In areas where there are squirrels, the lodgepole's cones are heavier, and have fewer seeds and thinner scales, making it more difficult for squirrels to get at the seeds. Conversely, where there are crossbills but no squirrels, the cones are lighter in construction, but have thicker scales, making it more difficult for crossbills to get at the seeds. The lodgepole's cones are in an evolutionary arms race with the two kinds of herbivore. and the two following pages of the web article.
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).|链接=Special:FilePath/Drosophila.melanogaster.couple.2.jpg]]
==Competition==
{{Main|Intraspecific competition|Interspecific competition}}
Both [[intraspecific competition]], with features such as [[sexual conflict]]<ref>{{cite journal |doi=10.1098/rstb.2005.1785 |title=Sexual conflict over mating and fertilization: An overview |year=2006 |last1=Parker |first1=G. A. |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |volume=361 |issue=1466 |pages=235–59 |pmid=16612884 |pmc=1569603}}</ref> and [[sexual selection]],<ref name="UCL">{{cite web|title=Biol 2007 - Coevolution|url=https://www.ucl.ac.uk/~ucbhdjm/courses/b242/Coevol/Coevol.html|publisher=[[University College, London]]|access-date=19 January 2017}}</ref> and [[interspecific competition]], such as between predators, may be able to drive coevolution.<ref>{{cite journal |last1=Connell |first1=Joseph H. |s2cid=5576868 |title=Diversity and the Coevolution of Competitors, or the Ghost of Competition Past |journal=Oikos |date=October 1980 |volume=35 |issue=2 |pages=131–138 |doi=10.2307/3544421 |jstor=3544421}}</ref>
Both intraspecific competition, with features such as sexual conflict and sexual selection, and interspecific competition, such as between predators, may be able to drive coevolution.
无论是具有如性冲突和性选择等特征的种内竞争，还是具有如食肉动物之间的种间竞争，都可能推动共同进化。
== 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/>|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.
多种蜂和长舌蜜蜂共同进化，不论是成对的还是“广泛”进化的，都被称为公会。
The types of coevolution listed so far have been described as if they operated pairwise (also called specific coevolution), in which traits of one species have evolved in direct response to traits of a second species, and vice versa. This is not always the case. Another evolutionary mode arises where evolution is reciprocal, but is among a group of species rather than exactly two. This is variously called guild or diffuse coevolution. For instance, a trait in several species of [[flowering plant]], such as offering its [[nectar]] at the end of a long tube, can coevolve with a trait in one or several species of pollinating insects, such as a long proboscis. More generally, flowering plants are pollinated by insects from different families including [[bee]]s, [[fly|flies]], and [[beetle]]s, all of which form a broad [[guild (ecology)|guild]] of [[pollinator]]s which respond to the nectar or pollen produced by flowers.<ref name=Juenger>Juenger, Thomas, and [[Joy Bergelson]]. "Pairwise versus diffuse natural selection and the multiple herbivores of scarlet gilia, Ipomopsis aggregata." Evolution (1998): 1583–1592.</ref><ref>{{cite book |author1=Gullan, P. J. |author2=Cranston, P. S. |date=2010 |title=The Insects: An Outline of Entomology |url=https://archive.org/details/insectsoutlineen00pjgu |url-access=limited |publisher=Wiley |edition=4th |isbn=978-1-118-84615-5 |pages=[https://archive.org/details/insectsoutlineen00pjgu/page/n315 291]–293}}</ref><ref>{{cite journal |last1=Rader |first1=Romina |last2=Bartomeus |first2=Ignasi |display-authors=etal |title=Non-bee insects are important contributors to global crop pollination |journal=PNAS |date=2016 |volume=113 |issue=1 |doi=10.1073/pnas.1517092112 |pmid=26621730 |pmc=4711867 |pages=146–151 |bibcode=2016PNAS..113..146R|doi-access=free }}</ref>
[[File:Common clownfish curves dnsmpl.jpg|thumb|[[Ocellaris clownfish]] and [[Heteractis magnifica|Ritter's sea anemones]] live together in a [[mutualism (biology)|mutual]] service-service symbiosis, the fish driving off butterfly fish and the anemone's tentacles protecting the fish from predators.]]
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The types of coevolution listed so far have been described as if they operated pairwise (also called specific coevolution), in which traits of one species have evolved in direct response to traits of a second species, and vice versa. This is not always the case. Another evolutionary mode arises where evolution is reciprocal, but is among a group of species rather than exactly two. This is variously called guild or diffuse coevolution. For instance, a trait in several species of flowering plant, such as offering its nectar at the end of a long tube, can coevolve with a trait in one or several species of pollinating insects, such as a long proboscis. More generally, flowering plants are pollinated by insects from different families including bees, flies, and beetles, all of which form a broad guild of pollinators which respond to the nectar or pollen produced by flowers.Juenger, Thomas, and Joy Bergelson. "Pairwise versus diffuse natural selection and the multiple herbivores of scarlet gilia, Ipomopsis aggregata." Evolution (1998): 1583–1592.
到目前为止，所列出的共同进化类型被描述为是两两运作的(也称为特定共同进化) ，其中一个物种的特征是直接响应第二个物种的特征而进化的，反之亦然。但事实并非总是如此。另一种进化模式出现在进化是相互的，但是是在一组物种之间而不是两个物种之间。这被称为公会或漫反射共同进化。例如，几种开花植物的一个特征，例如在长管的末端提供花蜜，可以与一种或几种传粉昆虫的特征共同进化，例如长喙。更一般地说，被子植物是由来自不同科的昆虫授粉的，包括蜜蜂、苍蝇和甲虫，所有这些昆虫形成了一个广泛的授粉者协会，它们对花朵产生的花蜜或花粉作出反应。朱恩格、托马斯和乔伊 · 贝尔格森。成对自然选择与扩散自然选择以及猩红吉利亚的多种食草动物——绿芽菜(Ipomopsis aggregata)进化(1998) : 1583-1592。