更改

===被子植物===

===被子植物===

Flowers appeared and diversified relatively suddenly in the fossil record, creating what [[Charles Darwin]] described as the "abominable mystery" of how they had evolved so quickly; he considered whether coevolution could be the explanation.<ref>{{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).

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

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

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。<ref name="CardinalDanforth2013" />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。<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>

花朵在化石记录中比较突发地出现和多样化，创造了被查尔斯 · 达尔文描述为如此迅速进化的“令人憎恶的神秘”; 他考虑是否可以以共同演化作为解释。<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>他可能在''物种起源''中首次提到了共同演化，并在''兰花的传粉''(1862)中进一步发展了这个概念。<ref name="CardinalDanforth2013" /><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>

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

====昆虫和昆虫传粉的花朵====

{{Further|Entomophily}}

{{Further|Entomophily}}

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

[[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 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.

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.

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 plant]]s 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 clade|sister]] to the bees evolved at the same time as flowering plants, as did the Lepidoptera. 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.

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.

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。

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）与被子植物共同演化。最早的蜜蜂，今天重要的传粉者，出现在白垩纪早期。<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>一群与蜜蜂相近亲的黄蜂与被子植物同时演化，鳞翅目也是如此。<ref name="Bristol" />此外，所有主要的蜜蜂分支种系均首次出现在白垩纪中到晚期间，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） ，正当被子植物成为世界上陆地的主要植物时。<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>

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

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

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

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

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.<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.

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.

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.

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

====鸟类和鸟类传粉的花朵====

{{Further|Ornithophily}}

{{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]]

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

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

[[Hummingbird]]s and ornithophilous (bird-pollinated) flowers have evolved a [[mutualism (biology)|mutualistic]] relationship. The flowers have [[nectar]] suited to the birds' diet, their color suits the birds' vision and their shape fits that of the birds' bills. The blooming times of the flowers have also been found to coincide with hummingbirds' breeding seasons. The floral characteristics of ornithophilous plants vary greatly among each other compared to closely related insect-pollinated species. These flowers also tend to be more ornate, complex, and showy than their insect pollinated counterparts. It is generally agreed that plants formed coevolutionary relationships with insects first, and ornithophilous species diverged at a later time. There is not much scientific support for instances of the reverse of this divergence: from ornithophily to insect pollination. The diversity in floral phenotype in ornithophilous species, and the relative consistency observed in bee-pollinated species can be attributed to the direction of the shift in pollinator preference.

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.

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。

蜂鸟和喜鸟类（通过鸟类传粉）的花演化出了一种互惠的关系。这些花的花蜜合于鸟类的饮食，颜色亦合于鸟类的视觉，形状则合于鸟的喙。这些花的开放时间还被发现与蜂鸟的繁殖季节相吻合。同与昆虫传粉密切相关的植物相比，喜鸟类植物的花部特征差异很大。这些花也会较昆虫授粉的同类更华丽、复杂和艳丽。被普遍认为的是，植物首先与昆虫形成共同演化关系，喜鸟类植物在后期分化。从鸟类学到昆虫授粉，并没有多少科学依据支持此分歧中相反的例子。喜鸟类植物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。<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>

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>

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>

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.

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.

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.

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。

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。<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>

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 [[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"/>