Editing Pollination (section)

== Methods ==

Pollination may be biotic or abiotic. Biotic pollination relies on living [[pollinator]]s to move the [[pollen]] from one flower to another. Abiotic pollination relies on wind, water or even rain. Adding natural habitat areas into farm systems generally improves pollination, as farms that are closer to natural habitat have higher crop yield because they are visited by more pollinators.<ref>{{Cite web |title=About Pollinators |url=https://www.pollinator.org/pollinators |access-date=2023-11-10 |website=Pollinator.org |language=en-US}}</ref> 

=== Biotic pollination===
[[File:Hummingbird in ggp 7.jpg|thumb|[[Hummingbird]]s typically feed on red flowers]]
[[File:Melissodes desponsa, f, face, Maine, Du Clos 2015-12-01-17.37 (24469964305).jpg|thumb|A bee (''[[Melissodes desponsus]]'') covered in pollen]]
{{main|Pollinator}}

About 80% of [[Flowering plant|angiosperms]] rely on biotic pollination.<ref>{{Cite journal |vauthors=Ackerman JD |date=2000-03-01 |title=Abiotic pollen and pollination: Ecological, functional, and evolutionary perspectives |journal=Plant Systematics and Evolution |volume=222 |issue=1–4 |pages=167–185 |doi=10.1007/BF00984101 |bibcode=2000PSyEv.222..167A |s2cid=36015720}}</ref> (also called pollen vectors): organisms that carry or move the pollen grains from the [[anther]] of one flower to the receptive part of the carpel or pistil (stigma) of another.<ref name="crop_Type">{{Cite web | title=Types of Pollination, Pollinators and Terminology | work=CropsReview.Com | access-date=2015-10-20 | url=http://www.cropsreview.com/types-of-pollination.html}}</ref> Between 100,000 and 200,000 species of animal act as pollinators of the world's 250,000 species of flowering plant.<ref name="Abrol">{{cite book | last = Abrol | first = Dharam P. | title = Pollination Biology | name-list-style = vanc |year=2012 |chapter=Non Bee Pollinators-Plant Interaction |volume=Chapter 9 |pages=265–310 |doi=10.1007/978-94-007-1942-2_9|isbn=978-94-007-1941-5 }}</ref> The majority of these pollinators are [[insect]]s, but about 1,500 species of birds and mammals visit flowers and may transfer pollen between them. Besides birds and bats which are the most frequent visitors, these include monkeys, lemurs, squirrels, rodents and possums.<ref name="Abrol" />

[[Entomophily]], pollination by [[insect]]s, often occurs on plants that have developed colored petals and a strong [[Floral scent|scent]] to attract insects such as bees, wasps, and occasionally ants ([[Hymenoptera]]), [[beetle]]s ([[Coleoptera]]), moths and butterflies ([[Lepidoptera]]), and flies ([[Diptera]]). The existence of insect pollination dates back to the [[dinosaur]] era.<ref name="scie_Firs">{{Cite web | title=First ever record of insect pollination from 100 million years ago | work=ScienceDaily | access-date=2015-10-20 | url=https://www.sciencedaily.com/releases/2012/05/120514153113.htm }}</ref>

Insect pollinators such as [[honey bee]]s (''Apis'' spp.),<ref>{{cite journal | vauthors = Hill PS, Wells PH, Wells H | title = Spontaneous flower constancy and learning in honey bees as a function of colour | journal = Animal Behaviour | volume = 54 | issue = 3 | pages = 615–27 | date = September 1997 | pmid = 9299046 | doi = 10.1006/anbe.1996.0467 | s2cid = 24674731 }}</ref>
[[bumblebee]]s (''Bombus'' spp.),<ref>{{cite journal | vauthors = Stout JC, Allen JA, Goulson D | title = The influence of relative plant density and floral morphological complexity on the behaviour of bumblebees | journal = Oecologia | volume = 117 | issue = 4 | pages = 543–550 | date = December 1998 | pmid = 28307680 | doi = 10.1007/s004420050691 | bibcode = 1998Oecol.117..543S | s2cid = 5829708 }}</ref><ref>{{cite journal | vauthors = Chittka L, Gumbert A, Kunze J | year=1997 | title=Foraging dynamics of bumble bees: correlates of movement within and between plant species | journal=Behavioral Ecology | volume=8 | issue=3 | pages=239–249 | doi=10.1093/beheco/8.3.239}}</ref> and [[Butterfly|butterflies]] (e.g., ''Thymelicus flavus'')<ref>{{cite journal | vauthors = Goulson D, Ollerton J, Sluman C | year=1997 | title=Foraging strategies in the small skipper butterfly, ''Thymelicus flavus'': when to switch? | journal=Animal Behaviour | volume=53 | issue= 5| pages=1009–1016 | doi=10.1006/anbe.1996.0390| s2cid=620334 }}</ref> have been observed to engage in [[flower constancy]], which means they are more likely to transfer pollen to other conspecific plants.<ref>{{cite book | vauthors = Harder LD, Williams NM, Jordan CY, Nelson WA | chapter = The effects of Floral design and display on pollinator economics and pollen dispersal | pages = 297–317 | veditors = Chittka L, Thomson JD | title =  Cognitive Ecology of Pollination: Animal Behavior and Floral Evolution | date = 2001 | publisher = Cambridge University Press }}</ref> This can be beneficial for the pollinators, as flower constancy prevents the loss of pollen during interspecific flights and pollinators from clogging stigmas with pollen of other flower species. It also improves the probability that the pollinator will find productive flowers easily accessible and recognisable by familiar clues.<ref>{{cite journal | vauthors = Chittka L, Thomson JD, Waser NM | year=1999 | title=Flower constancy, insect psychology, and plant evolution | journal=Naturwissenschaften | volume=86 | issue= 8| pages=361–377 | doi=10.1007/s001140050636| bibcode=1999NW.....86..361C | s2cid=27377784 }}</ref> The primary insect pollinators are [[hymenoptera]]ns, mostly [[bee]]s, but also including [[sawfly|sawflies]], [[ant]]s, and many species of wasps.<ref>{{cite journal | vauthors = Sühs RB, Somavilla A, Köhler A, Putzke J |url=http://www.ufrgs.br/seerbio/ojs/index.php/rbb/article/view/1123 |title=Pollen vector wasps (Hymenoptera, Vespidae) of ''Schinus terebinthifolius'' Raddi (Anacardiaceae) |year=2009 |pages=138–143  |journal=Brazilian Journal of Biosciences |volume=7 |issue=2}}</ref>

Many flowers attract pollinators by odor. For example, [[orchid bee]] species such as ''[[Euglossa cordata]]'' are attracted to orchids this way, and it has been suggested that some orchid species intoxicate bees during visits which can last up to 90 minutes.<ref>{{cite journal | vauthors = Dressler RL | title = Pollination by Euglossine Bees | journal = Evolution; International Journal of Organic Evolution | volume = 22 | issue = 1 | pages = 202–210 | date = March 1968 | pmid = 28564982 | doi = 10.2307/2406664 | jstor = 2406664 }}</ref> However, in general, plants that rely on pollen vectors tend to be adapted to their particular type of vector, for example day-pollinated species tend to be brightly coloured and have little odor, but if they are pollinated largely by birds or specialist mammals, they tend to be larger and have larger nectar rewards than species that are strictly insect-pollinated. Night-blooming flowers have little color, but are often very aromatic. Plants with vertebrate pollinators also tend to spread their rewards over longer periods, having long flowering seasons; their specialist pollinators would be likely to starve if the pollination season were too short.<ref name=":Potts1995">{{cite web | last1 = Potts | first1 = Brad | last2 = Gore | first2 = Peter | name-list-style = vanc | title = Reproductive Biology and Controlled Pollination of Eucalyptus | publisher = School of Plant Science, University of Tasmania | date = 1995 | url = https://eprints.utas.edu.au/7447/1/Potts_and_Gore_reproductive_biology_manual.pdf }}</ref>

Some flowers have specialized mechanisms to [[pollination trap|trap pollinators]] to increase effectiveness,<ref name=":Potts1995" /> attach pollen to specific body parts (as happens in many orchid and ''[[Asclepias]]'' species<ref>Ollerton, J. & S. Liede. 1997. Pollination systems in the Asclepiadaceae: a survey and preliminary analysis. Biological Journal of the Linnean Society (1997), 62: 593–610.</ref>), or require specialized behaviors or morphology in order to extract pollen or nectar. One such syndrome is "[[buzz pollination]]" (or "sonication"), where a bee must vibrate at a certain frequency in order to cause pollen to be released from the [[anthers]].<ref>{{Cite journal|title = Reproductive biology of six Brazilian Myrtaceae: - is there a syndrome associated with buzz-pollination?|journal = New Zealand Journal of Botany|date = 2009-12-01|issn = 0028-825X|pages = 355–365|volume = 47|issue = 4|doi = 10.1080/0028825x.2009.9672712|first1 = Adriana De O.|last1 = Fidalgo|first2 = Astrid De M. P.|last2 = Kleinert|doi-access = | bibcode=2009NZJB...47..355F }}</ref>

In [[zoophily]], pollination is performed by vertebrates such as [[bird]]s and [[bat]]s, particularly, [[hummingbird]]s, [[sunbird]]s, [[spiderhunter]]s, [[honeyeater]]s, and [[fruit bat]]s. [[Ornithophily]] or bird pollination is the pollination of flowering plants by birds. [[Chiropterophily]] or bat pollination is the pollination of flowering plants by bats. Plants adapted to use bats or moths as pollinators typically have white petals, strong [[Floral scent|scent]] and flower at night, whereas plants that use birds as pollinators tend to produce copious nectar and have red petals.<ref>{{cite journal | vauthors = Rodríguez-Gironés MA, Santamaría L | title = Why are so many bird flowers red? | journal = PLOS Biology | volume = 2 | issue = 10 | pages = e350 | date = October 2004 | pmid = 15486585 | pmc = 521733 | doi = 10.1371/journal.pbio.0020350 | doi-access = free }}</ref>
[[File:Bee Leg with Pollen Bulb.jpg|thumb|Hind leg of a honey bee with pollen pellet stuck on the pollen basket or corbicula. When the worker bee is collecting pollen, their legs make the transfer of pollen from the inner basitarsal combs to the outer pollen basket (shown in figure).]] 

Mammals are not generally thought of as pollinators, but some rodents, bats and marsupials are significant pollinators and some even specialise in such activities. In South Africa certain species of ''[[Protea]]'' (in particular ''Protea humiflora'', ''P. amplexicaulis'', ''P. subulifolia'', ''P. decurrens'' and ''P. cordata'') are adapted to pollination by rodents (particularly [[Cape Spiny Mouse]], ''Acomys subspinosus'')<ref>{{cite journal | vauthors = Wiens D, Rourke JP, Casper BB, Rickart EA, LaPine TR, Peterson CJ, Channing A | title = Nonflying Mammal Pollination of Southern African Proteas. | journal = Annals of the Missouri Botanical Garden | volume = 70 | issue = 1 | date = 1983 | doi = 10.2307/2399006 | jstor = 2399006 | url = https://www.biodiversitylibrary.org/part/31073 }}</ref> and [[elephant shrew]]s (''Elephantulus'' species).<ref>{{cite journal | vauthors = Fleming PA, Nicolson SW | title = Arthropod fauna of mammal-pollinated Protea humiflora: ants as an attractant for insectivore pollinators? | journal = African Entomology | date = March 2003 | volume = 11 | issue = 1 | pages = 9–14 | url = https://www.ingentaconnect.com/content/sabinet/ento/2003/00000011/00000001/art00002 }}</ref> The flowers are borne near the ground, are yeasty smelling, not colourful, and sunbirds reject the nectar with its high [[xylose]] content. The mice apparently can digest the xylose and they eat large quantities of the pollen.<ref>{{cite web |url=http://www.proteaatlas.org.za/p52prhumi.htm |title=Who is pollinating Pr humiflora |url-status=live|archive-url=https://web.archive.org/web/20130219070835/http://protea.worldonline.co.za/p52prhumi.htm |archive-date=2013-02-19 |first1= Trish|last1 = Fleming|first2= Sue|last2=Nicholson | name-list-style = vanc  }}</ref> In Australia pollination by flying, gliding and earthbound mammals has been demonstrated.<ref>{{cite journal | vauthors = Goldingay RL, Carthew SM, Whelan RJ |title=The Importance of Non-Flying Mammals in Pollination |journal= Oikos |volume= 61| issue =  1 |date=May 1991 |pages= 79–87 |jstor=3545409|doi=10.2307/3545409 |bibcode=1991Oikos..61...79G }}</ref>

Reptile pollinators are known, but they form a minority in most ecological situations. They are most frequent and most ecologically significant in island systems, where insect and sometimes also bird populations may be unstable and less species-rich. Adaptation to a lack of animal food and of predation pressure, might therefore favour reptiles becoming more herbivorous and more inclined to feed on pollen and nectar.<ref name="OJMVA">{{cite journal | vauthors = Olesen JM, Valido A | title = Lizards as pollinators and seed dispersers: an island phenomenon. | journal = Trends in Ecology & Evolution | date = April 2003 | volume = 18 | issue = 4 | pages = 177–81 | doi = 10.1016/S0169-5347(03)00004-1 | bibcode = 2003TEcoE..18..177O }}</ref> Most species of lizards in the families that seem to be significant in pollination seem to carry pollen only incidentally, especially the larger species such as [[Varanidae]]  and [[Iguanidae]], but especially several species of the [[Gekkonidae]] are active pollinators, and so is at least one species of the [[Lacertidae]], ''[[Podarcis lilfordi]]'', which pollinates various species, but in particular is the major pollinator of ''[[Euphorbia dendroides]]'' on various Mediterranean islands.<ref name="GAH">{{cite journal | last = Godínez-Álvarez | first = Héctor | name-list-style = vanc | title = Pollination and seed dispersal by lizards. | journal = Revista Chilena de Historia Natural | volume = 77 | pages = 569–577 | date = 2004 | issue = 3 | doi = 10.4067/S0716-078X2004000300015 | doi-access = free }}</ref>

{{Excerpt|Pollinator|Other invertebrates}}

=== Abiotic pollination===
Abiotic pollination uses nonliving methods such as wind and water to move pollen from one [[flower]] to another. This allows the plant to spend energy directly on pollen rather than on attracting pollinators with flowers and [[nectar]]. Pollination by wind is more common amongst abiotic pollination.

==== By wind ====
[[File:Pollen from Dactylis glomerata.jpg|alt=Cat grass (Dactylis glomerata) spreading pollen by wind|thumb|Cat grass (''[[Dactylis glomerata]]'') spreading pollen by wind]]
Some 98% of abiotic pollination is [[anemophily]], i.e., pollination by wind. This probably arose from insect pollination (entomophily), most likely due to changes in the environment or the availability of pollinators.<ref>{{Cite book |url=https://books.google.com/books?id=3zfLBAAAQBAJ&pg=PA34 |title=Principles of Pollination Ecology |vauthors=Faegri K, Van der Pijl L |date=2013-10-22 |publisher=Elsevier |isbn=9781483293035 |page=34}}</ref><ref>{{cite journal |vauthors=Whitehead DR |date=March 1969 |title=Wind Pollination in the Angiosperms: Evolutionary and Environmental Considerations |journal=Evolution; International Journal of Organic Evolution |volume=23 |issue=1 |pages=28–35 |doi=10.2307/2406479 |jstor=2406479 |pmid=28562955}}</ref><ref>{{Cite journal |last1=Culley |first1=Theresa M. |last2=Weller |first2=Stephen G. |last3=Sakai |first3=Ann K. |name-list-style=vanc |date=2002-08-01 |title=The evolution of wind pollination in angiosperms |journal=Trends in Ecology & Evolution |volume=17 |issue=8 |pages=361–369 |doi=10.1016/S0169-5347(02)02540-5}}</ref> The transfer of pollen is more efficient than previously thought; wind pollinated plants have developed to have specific heights, in addition to specific floral, [[stamen]] and stigma positions that promote effective pollen dispersal and transfer.<ref>{{cite journal |vauthors=Friedman J, Barrett SC |date=June 2009 |title=Wind of change: new insights on the ecology and evolution of pollination and mating in wind-pollinated plants |journal=Annals of Botany |volume=103 |issue=9 |pages=1515–27 |doi=10.1093/aob/mcp035 |pmc=2701749 |pmid=19218583}}</ref>

==== By water ====
Pollination by water, [[hydrophily]], uses water to transport pollen, sometimes as whole anthers; these can travel across the surface of the water to carry dry pollen from one flower to another.<ref name=":10">{{Cite journal |last=Cox |first=Paul Alan |name-list-style=vanc |date=1988 |title=Hydrophilous Pollination |journal=Annual Review of Ecology and Systematics |volume=19 |pages=261–279 |doi=10.1146/annurev.es.19.110188.001401 |jstor=2097155}}</ref> In ''[[Vallisneria spiralis]]'', an unopened male flower floats to the surface of the water, and, upon reaching the surface, opens up and the fertile anthers project forward. The female flower, also floating, has its stigma protected from the water, while its [[sepal]]s are slightly depressed into the water, allowing the male flowers to tumble in.<ref name=":10" />

==== By rain ====

Rain pollination is used by a small percentage of plants. Heavy rain discourages insect pollination and damages unprotected flowers, but can itself disperse pollen of suitably adapted plants, such as ''[[Ranunculus flammula]]'', ''[[Narthecium ossifragum]]'', and ''[[Caltha palustris]]''.<ref name="pollin" /> In these plants, excess rain drains allowing the floating pollen to come in contact with the stigma.<ref name="pollin" /> In some orchids ombrophily occurs, and rain water splashes cause the anther cap to be removed, allowing for the pollen to be exposed. After exposure, raindrops causes the pollen to be shot upward, when the stipe pulls them back, and then fall into the cavity of the stigma. Thus, for the orchid ''[[Acampe rigida]]'', this allows the plant to self-pollinate, which is useful when biotic pollinators in the environment have decreased.<ref name="Fan2012">{{cite journal |vauthors=Fan XL, Barrett SC, Lin H, Chen LL, Zhou X, Gao JY |date=October 2012 |title=Rain pollination provides reproductive assurance in a deceptive orchid |url= |journal=Annals of Botany |volume=110 |issue=5 |pages=953–8 |doi=10.1093/aob/mcs165 |pmc=3448421 |pmid=22851311 |doi-access=free}}</ref>

==== Switching methods ====
It is possible for a plant to have varying pollination methods, including both biotic and abiotic pollination. The orchid ''[[Oeceoclades maculata]]'' uses both rain and butterflies, depending on its environmental conditions.<ref>{{Cite journal |last1=Aguiar |first1=João M. R. B. V. |last2=Pansarin |first2=Ludmila M. |last3=Ackerman |first3=James D. |last4=Pansarin |first4=Emerson R. |name-list-style=vanc |date=2012 |title=Biotic versus abiotic pollination in Oeceoclades maculata (Lindl.) Lindl. (Orchidaceae) |journal=Plant Species Biology |volume=27 |issue=1 |pages=86–95 |doi=10.1111/j.1442-1984.2011.00330.x}}</ref>