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separated biogeographic frequency descriptions from 'plant adaptations' section into new section. added references. clarified in intro that ornithophily need not be 'coevolutionary' (ie not all bird pollinators are necessarily under selection by plants)
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[[File:Phaethornis longirostris.jpg|thumb|Hummingbird ''[[Phaethornis longirostris]]'' on an ''[[Etlingera]]'' inflorescence]]
[[File:Phaethornis longirostris.jpg|thumb|Hummingbird ''[[Phaethornis longirostris]]'' on an ''[[Etlingera]]'' inflorescence]]
'''Ornithophily''' or '''bird pollination''' is the [[pollination]] of flowering plants by [[bird]]s. This [[coevolution]]ary association is derived from insect pollination ([[entomophily]]) and is particularly well developed in some parts of the world, especially in the tropics and on some island chains.<ref>{{cite journal|author1=Valido, Alfredo |author2=Dupont, Yoko L. |author3=Olesen, Jens M. |lastauthoramp=yes |year=2004 |title= Bird–flower interactions in the Macaronesian islands |journal=J. Biogeogr. |volume=31|pages=1945–1953|url=http://www.biosci.ohio-state.edu/~awolfe/class/Biogeography/Readings/Culver.pdf |doi=10.1111/j.1365-2699.2004.01116.x|issue=12}}</ref> The association involves several distinctive plant adaptations forming a "[[pollination syndrome]]". The plants typically have colourful, often red, flowers with long tubular structures holding ample nectar and orientations of the stamen and stigma that ensure contact with the pollinator. Birds involved in ornithophily tend to be specialist [[nectarivore]]s with brushy tongues, long bills, capable of hovering flight or are light enough to perch on the flower structures.
'''Ornithophily''' or '''bird pollination''' is the [[pollination]] of flowering plants by [[bird]]s. This sometimes (but not always) [[coevolution]]ary association is derived from insect pollination ([[entomophily]]) and is particularly well developed in some parts of the world, especially in the tropics, Southern Africa, and on some island chains.<ref>{{cite journal|author1=Valido, Alfredo |author2=Dupont, Yoko L. |author3=Olesen, Jens M. |lastauthoramp=yes |year=2004 |title= Bird–flower interactions in the Macaronesian islands |journal=J. Biogeogr. |volume=31|pages=1945–1953|url=http://www.biosci.ohio-state.edu/~awolfe/class/Biogeography/Readings/Culver.pdf |doi=10.1111/j.1365-2699.2004.01116.x|issue=12}}</ref> The association involves several distinctive plant adaptations forming a "[[pollination syndrome]]". The plants typically have colourful, often red, flowers with long tubular structures holding ample nectar and orientations of the stamen and stigma that ensure contact with the pollinator. Birds involved in ornithophily tend to be specialist [[nectarivore]]s with brushy tongues and long bills, that are either capable of hovering flight or light enough to perch on the flower structures.


==Plant adaptations==
==Plant adaptations==
[[File:Colibri-thalassinus-001-edit.jpg|thumb|left|A [[lesser violetear]]]] [[File:ErythrinaFlower.jpg|thumb|Inflorescences of ''[[Butea]]'' allow birds to perch on the stalk<ref>{{cite journal|author=Cotton, Peter A. |year=2001 |title= The Behavior and Interactions of Birds Visiting ''Erythrina fusca'' Flowers in the Colombian Amazon|journal= Biotropica |volume=33 |issue=4 |pages=662–669 |doi=10.1646/0006-3606(2001)033[0662:tbaiob]2.0.co;2}}</ref><ref>{{cite journal|author=Tandon R|author2=Shivanna, K. R.|author3=Mohan Ram, HY|last-author-amp=yes |year=2003 |title= Reproductive Biology of ''Butea monosperma'' (Fabaceae)|journal= Annals of Botany|volume=92|pages=715–723 |doi=10.1093/aob/mcg193|pmid=14500327|issue=5 |pmc=4244857}}</ref>]]
[[File:Colibri-thalassinus-001-edit.jpg|thumb|left|A [[lesser violetear]]]] [[File:ErythrinaFlower.jpg|thumb|Inflorescences of ''[[Butea]]'' allow birds to perch on the stalk<ref>{{cite journal|author=Cotton, Peter A. |year=2001 |title= The Behavior and Interactions of Birds Visiting ''Erythrina fusca'' Flowers in the Colombian Amazon|journal= Biotropica |volume=33 |issue=4 |pages=662–669 |doi=10.1646/0006-3606(2001)033[0662:tbaiob]2.0.co;2}}</ref><ref>{{cite journal|author=Tandon R|author2=Shivanna, K. R.|author3=Mohan Ram, HY|last-author-amp=yes |year=2003 |title= Reproductive Biology of ''Butea monosperma'' (Fabaceae)|journal= Annals of Botany|volume=92|pages=715–723 |doi=10.1093/aob/mcg193|pmid=14500327|issue=5 |pmc=4244857}}</ref>]]
Bird pollination is considered as a costly strategy for plants and it evolves only where there are particular benefits for the plant.<ref>{{cite journal|author=Stiles, Gary F.|year= 1978 |title= Ecological and Evolutionary Implications of Bird Pollination |journal=American Zoologist |volume= 18 |issue=4 |pages=715–727 |doi=10.1093/icb/18.4.715}}</ref> High altitude ecosystems that lack insect pollinators, those in dry regions or isolated islands tend to favour the evolution of ornithophily in plants.<ref name=cronk/>
Bird pollination is considered as a costly strategy for plants and it evolves only where there are particular benefits for the plant.<ref>{{cite journal|author=Stiles, Gary F.|year= 1978 |title= Ecological and Evolutionary Implications of Bird Pollination |journal=American Zoologist |volume= 18 |issue=4 |pages=715–727 |doi=10.1093/icb/18.4.715}}</ref> High altitude ecosystems that lack insect pollinators, those in dry regions or isolated islands tend to favour the evolution of ornithophily.<ref name=cronk/>


Plants adaptations can be grouped into mechanisms that attract birds, those that exclude insects,<ref>{{cite journal|year=2004|title='Anti-bee' and 'pro-bird' changes during the evolution of hummingbird pollination in ''Penstemon'' flowers|journal=Journal of Evolutionary Biology |volume=17|issue=4|pages=876–85|url=https://www.csun.edu/~hcbio028/JEBCastellanos.pdf|vauthors=Castellanos MC, Wilson P, Thomson JD |doi=10.1111/j.1420-9101.2004.00729.x|pmid=15271088}}</ref> protect against nectar theft and pollination mechanisms in the strict sense.<ref name=cronk>{{cite journal|author1=Cronk, Quentin |author2=Isidro Ojeda |lastauthoramp=yes |year=2008 |title= Bird-pollinated flowers in an evolutionary and molecular context |journal=J. Exp. Bot. |volume=59 |pages=715–727 |doi=10.1093/jxb/ern009|pmid=18326865|issue=4}}</ref> The ovules of bird flowers also tend to have adaptations that protect them from damage.<ref>{{cite journal|title=The Protection of the Ovules in Flowering Plants|first=Verne|jstor=2405331|last=Grant|journal=Evolution |volume= 4|issue=3|year= 1950 |pages=179–201|doi=10.2307/2405331}}</ref>
Plant adaptations for ornithophily can be grouped primarily into those that attract and facilitate pollen transfer by birds, and those that exclude other groups, primarily insects<ref>{{cite journal|year=2004|title='Anti-bee' and 'pro-bird' changes during the evolution of hummingbird pollination in ''Penstemon'' flowers|journal=Journal of Evolutionary Biology |volume=17|issue=4|pages=876–85|url=https://www.csun.edu/~hcbio028/JEBCastellanos.pdf|vauthors=Castellanos MC, Wilson P, Thomson JD |doi=10.1111/j.1420-9101.2004.00729.x|pmid=15271088}}</ref>, protecting against 'theft' of [[Nectar robbing|nectar]] and pollen.<ref name=cronk>{{cite journal|author1=Cronk, Quentin |author2=Isidro Ojeda |lastauthoramp=yes |year=2008 |title= Bird-pollinated flowers in an evolutionary and molecular context |journal=J. Exp. Bot. |volume=59 |pages=715–727 |doi=10.1093/jxb/ern009|pmid=18326865|issue=4}}</ref> The ovules of bird flowers also tend to have adaptations that protect them from damage during vigorous foraging by hard bird bills.<ref>{{cite journal|title=The Protection of the Ovules in Flowering Plants|first=Verne|jstor=2405331|last=Grant|journal=Evolution |volume= 4|issue=3|year= 1950 |pages=179–201|doi=10.2307/2405331}}</ref>


Most bird pollinated flowers are red and have a lot of nectar. They also tend to be unscented.<ref>{{cite journal|author=Knudsen, Jette T., Tollsten, Lars, Groth, Inga|author2=Bergström, Gunnar|author3=Raguso, Robert A. |lastauthoramp=yes |year=2004 |title= Trends in floral scent chemistry in pollination syndromes: floral scent composition in hummingbird-pollinated taxa|journal=Botanical Journal of the Linnean Society |volume=146 |issue=2|pages=191–199|doi=10.1111/j.1095-8339.2004.00329.x}}</ref> Flowers with generalist pollinators tend to have dilute nectar but those that have specialist pollinators such as hummingbirds or sunbirds tend to have more concentrated nectar.<ref>{{cite journal|author1=Johnson, Steven D |author2=Susan W Nicolson |lastauthoramp=yes |year=2008 |title= Evolutionary associations between nectar properties and specificity in bird pollination systems |journal=Biol. Lett.|volume=4|issue=1|pages=49–52|doi=10.1098/rsbl.2007.0496|pmid=17999944|pmc=2412932}}</ref><ref>{{cite journal|vauthors=Rodríguez-Gironés MA, Santamaría L |year=2004|title= Why Are So Many Bird Flowers Red? |journal=PLoS Biology|pmid=15486585 |volume=2|issue=10|pmc=521733|page=e350|doi=10.1371/journal.pbio.0020350}}</ref> The nectar of ornithophilous flowers vary in the sugar composition, with hexoses being high in passerine pollinated species while those that are insect pollinated tend to be sucrose rich. Hummingbird pollinated flowers however tend to be sucrose rich.<ref>{{cite journal|author=Dupont, YL|author2=Hansen, DM|author3=Rasmussen, JT|author4=Olesen, JM|last-author-amp=yes |year=2004 |title=Evolutionary changes in nectar sugar composition associated with switches between bird and insect pollination: the Canarian bird-flower element revisited |journal=Functional Ecology |volume=18|issue=5|pages=670–676|doi=10.1111/j.0269-8463.2004.00891.x}}</ref> Many plants of the family [[Loranthaceae]] have explosive flowers that shower pollen on a bird that forages near it. They are associated mainly with flowerpeckers in the [[Dicaeidae]] family.<ref>{{cite journal|author=Feehan, J.|year=1985 |title= Explosive flower-opening in ornithophily: A study of pollination mechanisms in some Central African Loranthaceae |journal=Botanical Journal of the Linnean Society |volume=90|pages=129–144|doi=10.1111/j.1095-8339.1985.tb02205.x|issue=2}}</ref> In Australia, some species of ''[[Banksia]]'' have flowers that open in response to bird actions thereby reducing the wastage of pollen.<ref>{{cite journal|author=Ramsey, MW |year=1988 |title= Floret Opening in ''Banksia menziesii'' R.Br.; The Importance of Nectarivorous Birds |journal=Australian Journal of Botany |volume=36 |issue=2 |pages=225–232 |doi=10.1071/BT9880225}}</ref> In tropical dry forests in southern India, ornithophilous flowers were found to bloom mainly in the hot dry season.<ref>{{cite journal| title= Reproductive Phenology of a Tropical Dry Forest in Mudumalai, Southern India| author1=Murali, K.S.| author2=Sukumar, R.| journal=Journal of Ecology| volume=82| issue=4| year=1994| pages=759–767|jstor=2261441| doi=10.2307/2261441}}</ref> As many as 129 species of North American plants have evolved ornithophilous associations.<ref>{{cite journal|author=Grant, V |year= 1994 |title= Historical development of ornithophily in the western North American flora |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=91|issue=22|pages=10407–10411 |doi=10.1073/pnas.91.22.10407|pmid=7937964|pmc=45029}}</ref> Nearly a fourth of the 900 species of the genus ''Salvia'' are bird pollinated in the South African region.<ref>{{cite journal|author1=Wester, Petra |author2=Regine Claßen-Bockhoff |year=2006 |title= Bird pollination in South African Salvia species |journal=Flora – Morphology, Distribution, Functional Ecology of Plants |volume=201|issue=5|pages=396–406|doi=10.1016/j.flora.2005.07.016}}</ref> Tropical China and the adjacent Indochinese countries harbor relatively few bird-pollinated flowers, among them is ''[[Rhodoleia championii]]'', a member of the [[Hamamelidaceae]] family, which at any one site can be visited and pollinated by up to seven species of nectar-foraging birds, including [[Japanese white-eye]]s (''Zosterops japonicus'', Zosteropidae) and [[fork-tailed sunbird]]s (''Aethopyga christinae'', Nectariniidae).<ref>{{cite journal|author=Gu |year=2010 |title= Passerine pollination of ''Rhodoleia championii'' (Hamamelidaceae) in subtropical China |journal=Biotropica |volume=42 |pages=336–341 |doi=10.1111/j.1744-7429.2009.00585.x |issue=3 |last2=Luo|first2=Zhonglai|last3=Zhang|first3=Dianxiang|last4=Renner|first4=Susanne S.|display-authors=etal}}</ref> ''[[Calceolaria uniflora]]'', a species of Scrophularaceae from South America has a special fleshy appendage on the lower lip of the flower which is rich in sugar. This is fed on by the [[Least seedsnipe|least seedsnipe]] (''Thinocorus rumicivorus'') and in the process they brush pollen onto their head and transfer them to other flowers.<ref>{{Cite journal|last=Sérsic|first=A. N.|last2=Cocucci|first2=A. A.|date=1996|title=A Remarkable Case of Ornithophily in Calceolaria : Food Bodies as Rewards for a Non-nectarivorous Bird*|journal=Botanica Acta|language=en|volume=109|issue=2|pages=172–176|doi=10.1111/j.1438-8677.1996.tb00558.x}}</ref>
Most bird pollinated flowers are red and have a lot of nectar. They also tend to be unscented.<ref>{{cite journal|author=Knudsen, Jette T., Tollsten, Lars, Groth, Inga|author2=Bergström, Gunnar|author3=Raguso, Robert A. |lastauthoramp=yes |year=2004 |title= Trends in floral scent chemistry in pollination syndromes: floral scent composition in hummingbird-pollinated taxa|journal=Botanical Journal of the Linnean Society |volume=146 |issue=2|pages=191–199|doi=10.1111/j.1095-8339.2004.00329.x}}</ref> Flowers with generalist pollinators tend to have dilute nectar but those that have specialist pollinators such as hummingbirds or sunbirds tend to have more concentrated nectar.<ref>{{cite journal|author1=Johnson, Steven D |author2=Susan W Nicolson |lastauthoramp=yes |year=2008 |title= Evolutionary associations between nectar properties and specificity in bird pollination systems |journal=Biol. Lett.|volume=4|issue=1|pages=49–52|doi=10.1098/rsbl.2007.0496|pmid=17999944|pmc=2412932}}</ref><ref>{{cite journal|vauthors=Rodríguez-Gironés MA, Santamaría L |year=2004|title= Why Are So Many Bird Flowers Red? |journal=PLoS Biology|pmid=15486585 |volume=2|issue=10|pmc=521733|page=e350|doi=10.1371/journal.pbio.0020350}}</ref> The nectar of ornithophilous flowers vary in the sugar composition, with hexoses being high in passerine pollinated species while those that are insect pollinated tend to be sucrose rich. Hummingbird pollinated flowers however tend to be sucrose rich.<ref>{{cite journal|author=Dupont, YL|author2=Hansen, DM|author3=Rasmussen, JT|author4=Olesen, JM|last-author-amp=yes |year=2004 |title=Evolutionary changes in nectar sugar composition associated with switches between bird and insect pollination: the Canarian bird-flower element revisited |journal=Functional Ecology |volume=18|issue=5|pages=670–676|doi=10.1111/j.0269-8463.2004.00891.x}}</ref>


Different plants have also developed specific adaptations for bird pollination. Many plants of the family [[Loranthaceae]] have explosive flowers that shower pollen on a bird that forages near it. They are associated mainly with flowerpeckers in the [[Dicaeidae]] family.<ref>{{cite journal|author=Feehan, J.|year=1985 |title= Explosive flower-opening in ornithophily: A study of pollination mechanisms in some Central African Loranthaceae |journal=Botanical Journal of the Linnean Society |volume=90|pages=129–144|doi=10.1111/j.1095-8339.1985.tb02205.x|issue=2}}</ref> In Australia, some species of ''[[Banksia]]'' have flowers that open in response to bird actions thereby reducing the wastage of pollen.<ref>{{cite journal|author=Ramsey, MW |year=1988 |title= Floret Opening in ''Banksia menziesii'' R.Br.; The Importance of Nectarivorous Birds |journal=Australian Journal of Botany |volume=36 |issue=2 |pages=225–232 |doi=10.1071/BT9880225}}</ref> In tropical dry forests in southern India, ornithophilous flowers were found to bloom mainly in the hot dry season.<ref>{{cite journal| title= Reproductive Phenology of a Tropical Dry Forest in Mudumalai, Southern India| author1=Murali, K.S.| author2=Sukumar, R.| journal=Journal of Ecology| volume=82| issue=4| year=1994| pages=759–767|jstor=2261441| doi=10.2307/2261441}}</ref> ''[[Calceolaria uniflora]]'', a species of Scrophularaceae from South America, has a special fleshy appendage on the lower lip of the flower that is rich in sugar. This is fed on by the [[least seedsnipe]] (''Thinocorus rumicivorus'') and in the process the birds brush pollen onto their head and transfer them to other flowers.<ref>{{Cite journal|last=Sérsic|first=A. N.|last2=Cocucci|first2=A. A.|date=1996|title=A Remarkable Case of Ornithophily in Calceolaria : Food Bodies as Rewards for a Non-nectarivorous Bird*|journal=Botanica Acta|language=en|volume=109|issue=2|pages=172–176|doi=10.1111/j.1438-8677.1996.tb00558.x}}</ref>
⚫
The rat's tail babiana (''[[Babiana ringens]]'') is a species of plant that produces a strong stalk within the inflorescence that serves as a perch for the [[malachite sunbird]] as it visits the flower.<ref name=cronk/> [[Heliconias]] have special sticky threads that help in the adhesion of pollen to smooth structures such as the bill of a hummingbird.<ref>{{cite journal|author1=Rose, Marie-Jeanette |author2=Barthlott, Wilhelm |lastauthoramp=yes |year=1995|title= Pollen-connecting threads in Heliconia (Heliconiaceae)|journal=Plant Systematics and Evolution|volume= 195|issue=1|pages=61–65|doi=10.1007/BF00982315}}</ref> Some African orchids of the genus ''[[Disa (plant)|Disa]]'' have [[Pollinium|pollinaria]] that stick to the feet of visiting sunbirds.<ref>{{cite journal|title=Transfer of pollinaria on birds' feet: a new pollination system in orchids|journal=Plant Systematics and Evolution |year=2004|volume=244|issue=3|pages=181–188| doi = 10.1007/s00606-003-0106-y |vauthors=Johnson SD, Brown M }}</ref>


⚫
The rat's tail babiana (''[[Babiana ringens]]'') produces a strong stalk within the inflorescence that serves as a perch for the [[malachite sunbird]] as it visits the flower.<ref>{{Cite web|url=https://phys.org/news/2011-09-bird-pollinated-sex.html|title=Bird pollinated plant mixes it up when it comes to sex|website=phys.org|language=en-us|access-date=2019-03-20}}</ref> [[Heliconias]] have special sticky threads that help in the adhesion of pollen to smooth structures such as the bill of a hummingbird.<ref>{{cite journal|author1=Rose, Marie-Jeanette |author2=Barthlott, Wilhelm |lastauthoramp=yes |year=1995|title= Pollen-connecting threads in Heliconia (Heliconiaceae)|journal=Plant Systematics and Evolution|volume= 195|issue=1|pages=61–65|doi=10.1007/BF00982315}}</ref> Some African orchids of the genus ''[[Disa (plant)|Disa]]'' have [[Pollinium|pollinaria]] that stick to the feet of visiting sunbirds.<ref>{{cite journal|title=Transfer of pollinaria on birds' feet: a new pollination system in orchids|journal=Plant Systematics and Evolution |year=2004|volume=244|issue=3|pages=181–188| doi = 10.1007/s00606-003-0106-y |vauthors=Johnson SD, Brown M }}</ref>
Plants need to protect against nectar being taken by non-pollinators. These agents are classified into nectar robbers, which may destroy the flower, for example cut the flower at the base to obtain nectar and nectar thieves that obtain nectar without pollinating the flower.<ref>{{cite journal|author1=Valdivia, Carlos E. |author2=Paulina L. González-Gómez |year=2006|title= A trade-off between the amount and distance of pollen dispersal triggered by the mixed foraging behaviour of Sephanoides sephaniodes (Trochilidae) on Lapageria rosea (Philesiaceae) |journal=Acta Oecologica |volume=29|issue=3|pages=324–327|url=http://captura.uchile.cl/dspace/bitstream/2250/2772/1/Valdivia%20CE-Trade.pdf |doi=10.1016/j.actao.2005.12.005}}</ref>

Plants need to protect against nectar and pollen being taken by non-pollinators<ref>{{Cite journal|last=Hargreaves|first=Anna L.|last2=Harder|first2=Lawrence D.|last3=Johnson|first3=Steven D.|date=2009|title=Consumptive emasculation: the ecological and evolutionary consequences of pollen theft|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-185X.2008.00074.x|journal=Biological Reviews|language=en|volume=84|issue=2|pages=259–276|doi=10.1111/j.1469-185X.2008.00074.x|issn=1469-185X}}</ref>. Such animals are sometimes classified as thieves, which simply remove resources without pollinating, and robbers, which damage the flower to access resources. Flowers specialized for pollination by long-billed birds may be especially vulnerable to theft<ref>{{Cite journal|last=Hargreaves|first=AL|last2=Harder|first2=LD|last3=Johnson|first3=SD|date=2012|title=Floral traits mediate the vulnerability of aloes to pollen theft and inefficient pollination by bees|url=https://academic.oup.com/aob/article/109/4/761/129355|journal=Annals of Botany|language=en|volume=109|issue=4|pages=761–772|doi=10.1093/aob/mcr324|issn=0305-7364|pmc=PMC3286288|pmid=22278414|via=}}</ref>. For example, some bees and birds that cannot reach down the long tubes of bird pollinated flowers simply pierce the flower at the base to obtain nectar, without pollinating<ref>{{cite journal|author1=Valdivia, Carlos E. |author2=Paulina L. González-Gómez |year=2006|title= A trade-off between the amount and distance of pollen dispersal triggered by the mixed foraging behaviour of Sephanoides sephaniodes (Trochilidae) on Lapageria rosea (Philesiaceae) |journal=Acta Oecologica |volume=29|issue=3|pages=324–327|url=http://captura.uchile.cl/dspace/bitstream/2250/2772/1/Valdivia%20CE-Trade.pdf |doi=10.1016/j.actao.2005.12.005}}</ref>.

== Frequency of Ornithophily ==
As many as 129 species of North American plants have evolved ornithophilous associations.<ref>{{cite journal|author=Grant, V |year= 1994 |title= Historical development of ornithophily in the western North American flora |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=91|issue=22|pages=10407–10411 |doi=10.1073/pnas.91.22.10407|pmid=7937964|pmc=45029}}</ref> Nearly a fourth of the 900 species of the genus ''Salvia'' are bird pollinated in the South African region.<ref>{{cite journal|author1=Wester, Petra |author2=Regine Claßen-Bockhoff |year=2006 |title= Bird pollination in South African Salvia species |journal=Flora – Morphology, Distribution, Functional Ecology of Plants |volume=201|issue=5|pages=396–406|doi=10.1016/j.flora.2005.07.016}}</ref> Tropical China and the adjacent Indochinese countries harbor relatively few bird-pollinated flowers, among them is ''[[Rhodoleia championii]]'', a member of the [[Hamamelidaceae]] family, which at any one site can be visited and pollinated by up to seven species of nectar-foraging birds, including [[Japanese white-eye]]s (''Zosterops japonicus'', Zosteropidae) and [[fork-tailed sunbird]]s (''Aethopyga christinae'', Nectariniidae).<ref>{{cite journal|author=Gu |year=2010 |title= Passerine pollination of ''Rhodoleia championii'' (Hamamelidaceae) in subtropical China |journal=Biotropica |volume=42 |pages=336–341 |doi=10.1111/j.1744-7429.2009.00585.x |issue=3 |last2=Luo|first2=Zhonglai|last3=Zhang|first3=Dianxiang|last4=Renner|first4=Susanne S.|display-authors=etal}}</ref>


==Bird adaptations==
==Bird adaptations==

Revision as of 19:40, 20 March 2019

Hummingbird Phaethornis longirostris on an Etlingera inflorescence

Ornithophily or bird pollination is the pollination of flowering plants by birds. This sometimes (but not always) coevolutionary association is derived from insect pollination (entomophily) and is particularly well developed in some parts of the world, especially in the tropics, Southern Africa, and on some island chains.[1] The association involves several distinctive plant adaptations forming a "pollination syndrome". The plants typically have colourful, often red, flowers with long tubular structures holding ample nectar and orientations of the stamen and stigma that ensure contact with the pollinator. Birds involved in ornithophily tend to be specialist nectarivores with brushy tongues and long bills, that are either capable of hovering flight or light enough to perch on the flower structures.

Plant adaptations

A lesser violetear
Inflorescences of Butea allow birds to perch on the stalk[2][3]

Bird pollination is considered as a costly strategy for plants and it evolves only where there are particular benefits for the plant.[4] High altitude ecosystems that lack insect pollinators, those in dry regions or isolated islands tend to favour the evolution of ornithophily.[5]

Plant adaptations for ornithophily can be grouped primarily into those that attract and facilitate pollen transfer by birds, and those that exclude other groups, primarily insects[6], protecting against 'theft' of nectar and pollen.[5] The ovules of bird flowers also tend to have adaptations that protect them from damage during vigorous foraging by hard bird bills.[7]

Most bird pollinated flowers are red and have a lot of nectar. They also tend to be unscented.[8] Flowers with generalist pollinators tend to have dilute nectar but those that have specialist pollinators such as hummingbirds or sunbirds tend to have more concentrated nectar.[9][10] The nectar of ornithophilous flowers vary in the sugar composition, with hexoses being high in passerine pollinated species while those that are insect pollinated tend to be sucrose rich. Hummingbird pollinated flowers however tend to be sucrose rich.[11]

Different plants have also developed specific adaptations for bird pollination. Many plants of the family Loranthaceae have explosive flowers that shower pollen on a bird that forages near it. They are associated mainly with flowerpeckers in the Dicaeidae family.[12] In Australia, some species of Banksia have flowers that open in response to bird actions thereby reducing the wastage of pollen.[13] In tropical dry forests in southern India, ornithophilous flowers were found to bloom mainly in the hot dry season.[14] Calceolaria uniflora, a species of Scrophularaceae from South America, has a special fleshy appendage on the lower lip of the flower that is rich in sugar. This is fed on by the least seedsnipe (Thinocorus rumicivorus) and in the process the birds brush pollen onto their head and transfer them to other flowers.[15]

The rat's tail babiana (Babiana ringens) produces a strong stalk within the inflorescence that serves as a perch for the malachite sunbird as it visits the flower.[16] Heliconias have special sticky threads that help in the adhesion of pollen to smooth structures such as the bill of a hummingbird.[17] Some African orchids of the genus Disa have pollinaria that stick to the feet of visiting sunbirds.[18]

Plants need to protect against nectar and pollen being taken by non-pollinators[19]. Such animals are sometimes classified as thieves, which simply remove resources without pollinating, and robbers, which damage the flower to access resources. Flowers specialized for pollination by long-billed birds may be especially vulnerable to theft[20]. For example, some bees and birds that cannot reach down the long tubes of bird pollinated flowers simply pierce the flower at the base to obtain nectar, without pollinating[21].

Frequency of Ornithophily

As many as 129 species of North American plants have evolved ornithophilous associations.[22] Nearly a fourth of the 900 species of the genus Salvia are bird pollinated in the South African region.[23] Tropical China and the adjacent Indochinese countries harbor relatively few bird-pollinated flowers, among them is Rhodoleia championii, a member of the Hamamelidaceae family, which at any one site can be visited and pollinated by up to seven species of nectar-foraging birds, including Japanese white-eyes (Zosterops japonicus, Zosteropidae) and fork-tailed sunbirds (Aethopyga christinae, Nectariniidae).[24]

Bird adaptations

Ruby-throated hummingbird (Archilochus colubris) at scarlet beebalm flowers (Monarda didyma)

The main families of specialized nectar feeding birds that are involved in ornithophily are the hummingbirds (Trochilidae), sunbirds (Nectariniidae), and the honey-eaters (Meliphagidae). Other important bird groups include those in the families the Icteridae, the honeycreepers (Thraupidae, Drepanidae), white-eyes (Zosteropidae) and the South African sugar-birds (Promeropidae). Birds may obtain nectar either by perching or by hovering with the latter mainly found in the hummingbirds and sunbirds. Within the hummingbirds, two kinds of foraging are noted with territorial "hermit" hummingbirds and the non-hermits which forage longer distances [5]

Hummingbirds have the ability to digest sucrose unlike many passerines that prefer hexoses (fructose and glucose). Starlings and their relatives will completely avoid sucrose.[25] Nectar feeding birds typically have a mechanism to quickly excrete excess water. They may have to drink four to five times their body mass of liquid during the day to obtain enough energy.[26] Hummingbirds are capable of excreting nitrogenous wastes as ammonia since they can afford more water loss than birds that feed on low-moisture food sources.[27][28] Hummingbirds and sunbirds also have special anatomical and physiological adaptations that allow them to quickly excrete excess water. Hummingbirds are also able to turn off their kidney function at night.[29]

In some birds such as white-eyes, the pollen dusted by the plants on the forehead of the birds may increase the wear of these feathers leading to increased moulting and replacement.[30]

Other associations

Several mite species (mainly in the genera Proctolaelaps, Tropicoseius and Rhinoseius, family Ascidae) have evolved a phoretic mode of life, climbing into the nostrils of hummingbirds that visit flowers and hitching a ride to other flowers where they can feed on the nectar. Hummingbird flower mites favour plants in the families of Heliconiaceae, Costaceae, Zingiberaceae, Amaryllidaceae, Rubiaceae, Apocynaceae, Bromeliaceae, Gesneriaceae, Lobeliaceae and Ericaceae, members of which are associated with hummingbirds.[31]

See also

References

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  2. ^ Cotton, Peter A. (2001). "The Behavior and Interactions of Birds Visiting Erythrina fusca Flowers in the Colombian Amazon". Biotropica. 33 (4): 662–669. doi:10.1646/0006-3606(2001)033[0662:tbaiob]2.0.co;2.
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  16. ^ "Bird pollinated plant mixes it up when it comes to sex". phys.org. Retrieved 2019-03-20.
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  18. ^ Johnson SD, Brown M (2004). "Transfer of pollinaria on birds' feet: a new pollination system in orchids". Plant Systematics and Evolution. 244 (3): 181–188. doi:10.1007/s00606-003-0106-y.
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  21. ^ Valdivia, Carlos E.; Paulina L. González-Gómez (2006). "A trade-off between the amount and distance of pollen dispersal triggered by the mixed foraging behaviour of Sephanoides sephaniodes (Trochilidae) on Lapageria rosea (Philesiaceae)" (PDF). Acta Oecologica. 29 (3): 324–327. doi:10.1016/j.actao.2005.12.005.
  22. ^ Grant, V (1994). "Historical development of ornithophily in the western North American flora". Proc. Natl. Acad. Sci. U.S.A. 91 (22): 10407–10411. doi:10.1073/pnas.91.22.10407. PMC 45029. PMID 7937964.
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  24. ^ Gu; Luo, Zhonglai; Zhang, Dianxiang; Renner, Susanne S.; et al. (2010). "Passerine pollination of Rhodoleia championii (Hamamelidaceae) in subtropical China". Biotropica. 42 (3): 336–341. doi:10.1111/j.1744-7429.2009.00585.x.
  25. ^ Fleming, P. A.; Xie, S.; Napier, K.; McWhorter, T. J.; Nicolson, S. W. (2008). "Nectar concentration affects sugar preferences in two Australian honeyeaters and a lorikeet" (PDF). Functional Ecology. 22 (4): 599–605. doi:10.1111/j.1365-2435.2008.01401.x. {{cite journal}}: Unknown parameter |lastauthoramp= ignored (|name-list-style= suggested) (help)
  26. ^ Fleming, P. A.; Nicolson, S. W. (2003). "Osmoregulation in an avian nectarivore, the whitebellied sunbird Nectarinia talatala: response to extremes of diet concentration". J. Exp. Biol. 206 (Pt 11): 1845–1854. doi:10.1242/jeb.00351. PMID 12728006. {{cite journal}}: Unknown parameter |lastauthoramp= ignored (|name-list-style= suggested) (help)
  27. ^ McWhorter, T. J.; Powers, D. R.; Martínez del Rio, C. (2003). "Are hummingbirds facultatively ammonotelic? Nitrogen excretion and requirements as a function of body size". Physiol. Biochem. Zool. 76 (5): 731–743. doi:10.1086/376917. PMID 14671720. {{cite journal}}: Unknown parameter |last-author-amp= ignored (|name-list-style= suggested) (help)
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  31. ^ Da Cruz, Denise Dias; Vanessa Holanda Righetti De Abreu; Monique Van Sluys (2007). "The Effect of Hummingbird Flower Mites on Nectar Availability of Two Sympatric Heliconia Species in a Brazilian Atlantic Forest". Annals of Botany. 100 (3): 581–588. doi:10.1093/aob/mcm135. PMC 2533618. PMID 17638712. {{cite journal}}: Unknown parameter |last-author-amp= ignored (|name-list-style= suggested) (help)

External links

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