Editing Pollinator (section)

==Types of pollinators==
=== Insects ===
{{main|Entomophily}}

==== Bees ====
[[File:Lipotriches sp..jpg|thumb|left|''[[Lipotriches]]'' sp. bee pollinating flowers]]
The most recognized pollinators are the various species of [[bee]]s,<ref>{{cite journal | vauthors = Klein AM, Vaissière BE, Cane JH, Steffan-Dewenter I, Cunningham SA, Kremen C, Tscharntke T | title = Importance of pollinators in changing landscapes for world crops | journal = Proceedings. Biological Sciences | volume = 274 | issue = 1608 | pages = 303–313 | date = February 2007 | pmid = 17164193 | pmc = 1702377 | doi = 10.1098/rspb.2006.3721 }}</ref> which are plainly adapted to pollination. Bees typically are fuzzy and carry an [[electrostatic]] charge. Both features help pollen grains adhere to their bodies, but they also have specialized pollen-carrying structures; in most bees, this takes the form of a structure known as the [[Scopa (biology)|scopa]], which is on the hind legs of most bees, and/or the lower abdomen (e.g., of [[Megachilidae|megachilid]] bees), made up of thick, plumose [[seta]]e. [[Honey bee]]s, [[bumblebee]]s, and their relatives do not have a scopa, but the hind leg is modified into a structure called the corbicula (also known as the "[[pollen basket]]"). Most bees gather [[nectar]], a concentrated energy source, and pollen, which is high [[protein]] food, to nurture their young, and transfer some among the flowers as they are working.<ref>{{cite book| vauthors = Westbrook FE, Bergman PW, Wearne RA |title=Pollination and the Honey Bee|date=1975|publisher=U.S. Government Printing Office|location=Washington D.C.|url=https://books.google.com/books?id=gWwvAAAAYAAJ&q=bee+pollination|language=en}}</ref> [[Euglossini|Euglossine]] bees pollinate orchids, but these are male bees collecting floral scents rather than females gathering nectar or pollen. Female orchid bees act as pollinators, but of flowers other than orchids. [[Eusocial]] bees such as honey bees need an abundant and steady [[pollen source]] to multiply.

[[File:PrunusCerasifera0.jpg|thumb|upright|[[Honey bee]] pollinating a plum tree. Bees are the most effective insect pollinators.]]
Honey bees travel from flower to flower, collecting nectar (later converted to [[honey]]), and pollen grains. The bee collects the pollen by rubbing against the anthers. The pollen collects on the hind legs, in a structure referred to as a "pollen basket". As the bee flies from flower to flower, some of the pollen grains are transferred onto the stigma of other flowers. Nectar provides the energy for bee [[nutrition]]; pollen provides the [[protein]]. When bees are rearing large quantities of [[brood (honey bee)|brood]] (beekeepers say hives are "building"), bees deliberately gather pollen to meet the nutritional needs of the brood.

Good pollination management seeks to have bees in a "building" state during the bloom period of the crop, thus requiring them to gather pollen, and making them more efficient pollinators. Thus, the management techniques of a [[beekeeper]] providing pollination services are different from, and to some extent in tension with, those of a beekeeper who is trying to produce honey. Millions of [[Beehive (beekeeping)|hives]] of honey bees are [[contract]]ed out as pollinators by [[beekeeping|beekeepers]], and honey bees are by far the most important commercial pollinating agents, but many other kinds of pollinators, from blue bottle flies, to bumblebees, [[orchard mason bee]]s, and [[Alfalfa leafcutter bee|leaf cutter bees]] are cultured and sold for [[pollination management|managed pollination]].

Other species of bees differ in various details of their behavior and pollen-gathering habits, and honey bees are not native to the [[Western Hemisphere]]; all pollination of native plants in the Americas and Australia historically has been performed by various native bees. It has also been found that non-native plants may have positive effects on native bee pollinators while also influencing their foraging patterns and bee–plant networks.<ref>{{Cite journal |last1=Seitz |first1=Nicola |last2=vanEngelsdorp |first2=Dennis |last3=Leonhardt |first3=Sara D. |date=2020 |title=Are native and non-native pollinator friendly plants equally valuable for native wild bee communities? |journal=Ecology and Evolution |language=en |volume=10 |issue=23 |pages=12838–12850 |doi=10.1002/ece3.6826 |issn=2045-7758 |pmc=7713930 |pmid=33304497|bibcode=2020EcoEv..1012838S }}</ref>

==== Butterflies and moths ====
[[File:Australian painted lady feeding closeup.jpg|thumb|upright|left|An [[Australian painted lady]] feeding on nectar]]
[[Lepidoptera]] ([[butterfly|butterflies]] and [[moth]]s) may also pollinate to various degrees.<ref>{{cite web | title = Butterfly Pollination | work = Celebrating Butterflies | publisher = U.S. Forestry Service | url = http://www.fs.fed.us/wildflowers/pollinators/butterflies.shtml | archive-url = https://web.archive.org/web/20110723083314/http://www.fs.fed.us/wildflowers/pollinators/butterflies.shtml| archive-date=23 July 2011}}</ref> They are not major pollinators of [[food crop]]s, but various moths are important pollinators of other commercial crops such as [[tobacco]]. Pollination by certain moths may be important, however, or even crucial, for some wildflowers mutually adapted to specialist pollinators. Spectacular examples include orchids such as ''[[Angraecum sesquipedale]]'', dependent on a particular [[Sphingidae|hawk moth]], [[Xanthopan morganii|Morgan's sphinx]]. ''[[Yucca]]'' species provide other examples, being fertilised in elaborate ecological interactions with particular species of [[Prodoxidae|yucca moths]].

==== Flies ====
Many [[Bombyliidae|bee flies]], and some [[Horse-fly|Tabanidae]] and [[Nemestrinidae]] are particularly adapted to pollinating [[fynbos]] and [[Karoo]] plants with narrow, deep [[Petal|corolla tubes]], such as ''[[Lapeirousia]]'' species. Part of the adaptation takes the form of remarkably long probosces. This also applies to empidine dance flies ([[Empidinae]]) that visit a wide range of flowering plants, some species of which can pollinate the [[Geranium sylvaticum|woodland geranium]] ([[Geranium sylvaticum|''Geranium sylvaticum'' L.]]) as effectively as [[bee]]s.<ref>{{cite journal | vauthors = Lefebvre V, Daugeron C, Villemant C, Fontaine C | title = Empidine dance flies pollinate the woodland geranium as effectively as bees | journal = Biology Letters | volume = 15 | issue = 7 | pages = 20190230 | date = July 2019 | pmid = 31362609 | pmc = 6684995 | doi = 10.1098/rsbl.2019.0230 }}</ref>

[[File:Goudoogdaas zijaanzicht 2009 08 23.png|thumb|left|[[Horse-fly|Tabanid]] fly on a [[thistle]] flower]]
Carrion flies and flesh flies in families such as [[Calliphoridae]] and [[Flesh-fly|Sarcophagidae]] are important for some species of plants whose flowers [[Carrion flower|exude a fetid odor]]. The plants' ecological strategy varies; several species of ''[[Stapelia]]'', for example, attract carrion flies that futilely lay their eggs on the flower, where their larvae promptly starve for lack of [[carrion]]. Other species do decay rapidly after ripening, and offer the visiting insects large masses of food, as well as pollen and sometimes seed to carry off when they leave.

[[Hoverfly|Hoverflies]] are important pollinators of flowering plants worldwide.<ref name="Larson 2001 439–465">{{cite journal | vauthors = Larson BM, Kevan PG, Inouye DW |title=Flies and flowers: taxonomic diversity of anthophiles and pollinators. |journal=Canadian Entomologist |year=2001 |volume=133 |issue=4  |pages=439–465 |doi=10.4039/ent133439-4|s2cid=55767580 }}</ref> Often hoverflies are considered to be the second most important pollinators after wild bees.<ref name="Larson 2001 439–465"/> Although hoverflies as a whole are generally considered to be nonselective pollinators, some species have more specialized relationships. The orchid species ''Epipactis veratrifolia'' mimics alarm [[pheromones]] of aphids to attract hover flies for pollination.<ref>{{cite journal | vauthors = Stökl J, Brodmann J, Dafni A, Ayasse M, Hansson BS | title = Smells like aphids: orchid flowers mimic aphid alarm pheromones to attract hoverflies for pollination | journal = Proceedings. Biological Sciences | volume = 278 | issue = 1709 | pages = 1216–1222 | date = April 2011 | pmid = 20943694 | pmc = 3049078 | doi = 10.1098/rspb.2010.1770 }}</ref> Another plant, the [[slipper orchid]] in southwest China, also achieves pollination by deceit by exploiting the [[innate]] yellow colour preference of syrphids.<ref>{{cite journal | vauthors = Shi J, Luo YB, Bernhardt P, Ran JC, Liu ZJ, Zhou Q | title = Pollination by deceit in Paphiopedilum barbigerum (Orchidaceae): a staminode exploits the innate colour preferences of hoverflies (Syrphidae) | journal = Plant Biology | volume = 11 | issue = 1 | pages = 17–28 | date = January 2009 | pmid = 19121110 | doi = 10.1111/j.1438-8677.2008.00120.x | bibcode = 2009PlBio..11...17S }}</ref>

Some male [[Dacini|dacine fruit flies]] are exclusive pollinators of some wild ''[[Bulbophyllum]]'' orchids that lack nectar and have a specific chemical attractant and reward (methyl eugenol, raspberry ketone or zingerone) present in their floral fragrances.<ref>{{cite journal | vauthors = Tan KH, Nishida R, Toong YC | date = 2002 | title = ''Bulbophyllum cheiri'' floral synomone lures fruit flies to perform pollination. | journal = Journal of Chemical Ecology | volume = 28 | issue = 6 | pages = 1161–1172 | doi = 10.1023/A:1016277500007 | pmid = 12184394 | s2cid = 36621985 }}</ref><ref>{{cite journal | vauthors = Tan K, Nishida R | title = Synomone or kairomone?-Bulbophyllum apertum flower releases raspberry ketone to attract Bactrocera fruit flies. | journal = Journal of Chemical Ecology | date = 2005 | volume = 31 | issue = 3 | pages = 497–507 | doi = 10.1007/s10886-005-2023-8| pmid = 15898497 | bibcode = 2005JCEco..31..497K | s2cid = 39173699 }}</ref><ref>{{cite journal | vauthors = Tan KH, Nishida R | title = Zingerone in the floral synomone of ''Bulbophyllum baileyi'' (Orchidaceae) attracts ''Bactrocera'' fruit flies during pollination. | journal = Biochemical Systematics and Ecology | date = June 2007 | volume = 35 | issue = 6 | pages = 334–341 | doi = 10.1016/j.bse.2007.01.013 | bibcode = 2007BioSE..35..334T }}</ref>

Some flies, especially [[Anthomyiidae]], [[Empididae]] and [[Muscidae]], may be the main pollinators at higher elevations of mountains,<ref>{{cite journal |vauthors=Lefebvre V, Fontaine C, Villemant C, Daugeron C |date=November 2014 |title=Are empidine dance flies major flower visitors in alpine environments? A case study in the Alps, France |journal=Biology Letters |volume=10 |issue=11 |pages=20140742 |doi=10.1098/rsbl.2014.0742 |pmc=4261866 |pmid=25376804}}</ref><ref>{{cite journal |vauthors=Lefebvre V, Villemant C, Fontaine C, Daugeron C |date=March 2018 |title=Altitudinal, temporal and trophic partitioning of flower-visitors in Alpine communities |journal=Scientific Reports |volume=8 |issue=1 |pages=4706 |bibcode=2018NatSR...8.4706L |doi=10.1038/s41598-018-23210-y |pmc=5856740 |pmid=29549294}}</ref> whereas [[bumblebee]] species are typically the only other pollinators in alpine regions at timberline and beyond.

Some adult [[mosquito]]es, if they feed on nectar, may act as pollinators; ''Aedes communis'', a species found in North America, is known to pollinate ''[[Platanthera obtusata]]'', commonly referred as the blunt-leaved orchid.<ref>{{cite web |date=8 July 2015 |title=Year of Pollination: Mosquitoes as Pollinators |url=https://awkwardbotany.com/2015/07/08/year-of-pollination-mosquitoes-as-pollinators/ |access-date=28 July 2017 |website=awkward botany}}</ref><ref>{{cite web |title=''Aedes communis'': The Pollinating Mosquito |url=https://www.fs.fed.us/wildflowers/pollinators/pollinator-of-the-month/aedes_communis.shtml |access-date=28 July 2017 |website=[[United States Forest Service]] |vauthors=Statman-Weil Z}}</ref>

Biting midges ([[Ceratopogonidae]]) pollinate ''[[Theobroma cacao]]'' (Malvaceae), whose flowers have pollen inaccessible to larger pollinators.<ref>{{cite journal |last1=Arnold |first1=S.E.J. |last2=Forbes |first2=S.J. |last3=Hall |first3=D.R. |last4=Farman |first4=D.I. |last5=Bridgemohan |first5=P. |last6=Spinelli |first6=G.R. |last7=Bray |first7=D.P. |last8=Perry |first8=G.B. |last9=Grey |first9=L. |last10=Belmain |first10=S.R. |last11=Stevenson |first11=P.C. |year=2019 |title=Floral Odors and the Interaction between Pollinating Ceratopogonid Midges and Cacao |journal=Journal of Chemical Ecology |volume=45 |issue=10 |pages=869–878 |bibcode=2019JCEco..45..869A |doi=10.1007/s10886-019-01118-9 |pmid=31741191 |s2cid=208086796 |doi-access=free}}</ref>

==== Other insects ====
[[File:Unidentified Scoliidae foraging 2012 02 26 3034s.jpg|thumb|A [[Scoliid wasp]] (''Scolia chrysotricha'') foraging]]
Many insects other than bees accomplish pollination by visiting flowers for nectar or pollen, or commonly both. Many do so adventitiously, but the most important pollinators are specialists for at least parts of their life cycles for at least certain functions.

Prominent among Hymenoptera other than bees are wasps, especially [[Crabronidae]], [[Cuckoo wasp|Chrysididae]], [[Ichneumonidae]], [[Sphecidae]] and [[Vespidae]].<ref name=":03">{{Cite journal |last=Borchardt |first=Kate E. |last2=Holthaus |first2=Danielle |last3=Soto Méndez |first3=Paola A. |last4=Toth |first4=Amy L. |date=2024 |title=Debunking wasp pollination: Wasps are comparable to bees in terms of plant interactions, body pollen and single‐visit pollen deposition |url=https://resjournals.onlinelibrary.wiley.com/doi/10.1111/een.13329 |journal=Ecological Entomology |language=en |volume=49 |issue=4 |pages=569–584 |doi=10.1111/een.13329 |issn=0307-6946|doi-access=free }}</ref> Some wasps are comparable to or even superior to some bees as pollinators.<ref name=":03" /> The term "[[pollen wasp]]s", in particular, is widely applied to the Masarinae, a [[Taxonomic rank|subfamily]] of the Vespidae; they are remarkable among [[hunting wasp|solitary wasp]]s in that they specialise in gathering pollen for feeding their larvae, carried internally and regurgitated into a mud chamber prior to oviposition. Also, males of many species of bees and wasps, though they do not gather pollen, rely on flowers as sources of energy (in the form of nectar) and also as territories for meeting fertile females that visit the flowers.

[[Beetle]]s of species that specialise in eating pollen, nectar, or flowers themselves, may be important cross-pollinators of some plants such as members of the [[Araceae]] and [[Zamiaceae]], that produce prodigious amounts of pollen. Others, for example the [[Hopliini]], specialise on flowers of [[Asteraceae]] and [[Aizoaceae]].

[[Thrips]] pollinate plants such as elderflower ''[[Sambucus nigra]]'' (Adoxaceae)<ref>{{cite journal |last1=Scott-Brown |first1=A.S. |last2=Arnold |first2=S.E.J. |last3=Kite |first3=G.C. |last4=Farrell |first4=I.F. |last5=Collins |first5=D.W. |last6=Stevenson |first6=P.C. |title=Mechanisms in mutualisms: A chemically mediated thrips pollination strategy in common elder |journal=Planta |year=2019 |volume=250 |issue=1 |pages=367–379 |doi=10.1007/s00425-019-03176-5|pmid=31069523 |s2cid=253886497 |doi-access=free |bibcode=2019Plant.250..367S }}</ref> and pointleaf manzanita, ''[[Arctostaphylos pungens]]'' ([[Ericaceae]]).<ref>{{Cite journal |last=Eliyahu |first=Dorit |last2=McCall |first2=Andrew C. |last3=Lauck |first3=Marina |last4=Trakhtenbrot |first4=Ana |last5=Bronstein |first5=Judith L. |date=2015 |title=Minute pollinators: The role of thrips (Thysanoptera) as pollinators of pointleaf manzanita, Arctostaphylos pungens (Ericaceae) |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC4509684/ |journal=Journal of Pollination Ecology |volume=16 |pages=64–71 |issn=1920-7603 |pmc=4509684 |pmid=26207155}}</ref> [[Ant]]s also pollinate some kinds of flowers, but for the most part they are parasites, consuming nectar and/or pollen without conveying useful amounts of pollen to a stigma. Other insect orders are rarely pollinators, and then typically only incidentally (e.g., [[Hemiptera]] such as [[Anthocoridae]] and [[Miridae]]).

A strategy of great biological interest is that of sexual deception, where plants, generally [[Orchidaceae#Pollination|orchids, produce remarkably complex combinations of pheromonal attractants and physical mimicry]] that induce male [[Apidae|bees]] or [[Tiphiidae|wasps]] to attempt to mate with them, conveying [[Pollinium|pollinia]] in the process. Examples are known from all continents apart from [[Antarctica]], though Australia appears to be exceptionally rich in examples.<ref name=OrchSex>{{cite journal | vauthors = Mant JG, Schiestl FP, Peakall R, Weston PH | title = A phylogenetic study of pollinator conservatism among sexually deceptive orchids | journal = Evolution; International Journal of Organic Evolution | volume = 56 | issue = 5 | pages = 888–98 | date = May 2002 | pmid = 12093025 | doi = 10.1111/j.0014-3820.2002.tb01402.x | s2cid = 42724740 | doi-access = free }}</ref>

Whole groups of plants, such as certain [[fynbos]] ''[[Moraea]]'' and ''[[Erica (plant)|Erica]]'' species produce flowers on sticky [[Peduncle (botany)|peduncles]] or with sticky [[Petal|corolla]] tubes that only permit access to flying pollinators, whether bird, bat, or insect.

=== Other invertebrates ===
Experimental evidence has shown invertebrates (mostly small [[crustacean]]s<ref name=":1" />) acting as pollinators in underwater environments. Beds of [[seagrass]] have been shown to reproduce this way in the absence of currents. It is not yet known how important invertebrate pollinators might be for other species.<ref name=":0" /><ref>{{cite journal | vauthors = van Tussenbroek BI, Villamil N, Márquez-Guzmán J, Wong R, Monroy-Velázquez LV, Solis-Weiss V | title = Experimental evidence of pollination in marine flowers by invertebrate fauna | journal = Nature Communications | volume = 7 | issue = 1 | pages = 12980 | date = September 2016 | pmid = 27680661 | pmc = 5056424 | doi = 10.1038/ncomms12980 | bibcode = 2016NatCo...712980V }}</ref> Later, ''[[Idotea balthica]]'' was discovered to help ''[[Gracilaria]] gracilis'' reproduce – the first known case of an animal helping [[algae]] reproduce.<ref>{{cite news | vauthors = Roth A |title=Like Bees of the Seas, These Crustaceans Pollinate Seaweed |url=https://www.nytimes.com/2022/07/28/science/pollination-algae-crustacean.html |access-date=21 August 2022 |work=The New York Times |date=28 July 2022}}</ref><ref>{{cite journal | vauthors = Lavaut E, Guillemin ML, Colin S, Faure A, Coudret J, Destombe C, Valero M | title = Pollinators of the sea: A discovery of animal-mediated fertilization in seaweed | journal = Science | volume = 377 | issue = 6605 | pages = 528–530 | date = July 2022 | pmid = 35901149 | doi = 10.1126/science.abo6661 | bibcode = 2022Sci...377..528L | s2cid = 251159505 | url = https://hal.archives-ouvertes.fr/hal-03740649/file/Lavaut_Science_2022_version%20Hal.pdf }}</ref>

===Vertebrates===
[[File:Tacca chantrieri172799839.jpg|thumb|Tropical flowers like ''[[Tacca chantrieri]]'' are [[bat]]-pollinated.]]
[[File:Colibri thalassinus Curicancha 02.jpg|thumb|[[Green violetear]] with pollen on bill, [[Curi Cancha Wildlife Refuge]], Costa Rica]]
[[Bat]]s are important pollinators of some tropical flowers, visiting to take nectar.<ref>{{Cite journal | vauthors = Stewart AB, Dudash MR |date=1 January 2018 |title=Foraging strategies of generalist and specialist Old World nectar bats in response to temporally variable floral resources |journal=Biotropica |volume=50 |issue=1 |pages=98–105 |doi=10.1111/btp.12492|bibcode=2018Biotr..50...98S |s2cid=90515964 }}</ref> Birds, particularly [[hummingbird]]s, [[honeyeaters]] and [[sunbirds]] also accomplish much pollination, especially of deep-throated flowers. Other [[vertebrates]], such as [[kinkajou]]s, [[monkey]]s, [[lemur]]s, [[Phalangeriformes|possums]], [[rodent]]s, [[lizard]]s,<ref>{{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 = 177–181 | doi = 10.1016/S0169-5347(03)00004-1 | bibcode = 2003TEcoE..18..177O }}]</ref><ref>{{cite journal | vauthors = Baeckens S, Van Damme R | title = The island syndrome | journal = Current Biology | volume = 30 | issue = 8 | pages = R338–R339 | date = April 2020 | pmid = 32315628 | doi = 10.1016/j.cub.2020.03.029 | doi-access = free | bibcode = 2020CBio...30.R338B }}</ref> and [[Canidae|canids]]<ref>{{Cite journal |last1=Lai |first1=Sandra |last2=Léandri-Breton |first2=Don-Jean |last3=Lesaffre |first3=Adrien |last4=Samune |first4=Abdi |last5=Marino |first5=Jorgelina |last6=Sillero-Zubiri |first6=Claudio |date=2024 |title=Canids as pollinators? Nectar foraging by Ethiopian wolves may contribute to the pollination of Kniphofia foliosa |journal=Ecology |language=en |volume=105 |issue=12 |pages=e4470 |doi=10.1002/ecy.4470 |issn=1939-9170 |pmc=11610677 |pmid=39561987}}</ref> have been recorded pollinating some plants.

Humans can be pollinators, as many [[gardener]]s have discovered that they must [[hand pollination|hand pollinate]] garden [[vegetable]]s, whether because of [[pollinator decline]] or simply to keep a strain genetically pure. This can involve using a small brush or [[cotton]] swab to move pollen, or to simply tap or shake [[tomato]] blossoms to release the pollen for the [[self-pollinating]] flowers. Tomato blossoms are self-fertile, but (with the exception of potato-leaf varieties) have the pollen inside the [[anther]], and the flower requires shaking to release the pollen through [[Wiktionary: pore|pores]]. This can be done by wind, by humans, or by a [[Buzz pollination|sonicating]] bee (one that vibrates its wing muscles while perched on the flower), such as a bumblebee. Sonicating bees are extremely efficient pollinators of tomatoes, and colonies of bumblebees are quickly replacing humans as the primary pollinators for [[greenhouse]] tomatoes.