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第1行： − ===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]] − {{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 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。+ − + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
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{{Short description|Two or more species influencing each other's evolution}}
{{Good article}}
[[File:Dasyscolia ciliata.jpg|thumb|upright=1.5|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}}
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.
[[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]]
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。
==Mutualism==
{{Main|Mutualism (biology)}}
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===
===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.
[[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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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。