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A pollinator is an animal that moves
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
from the male
anther The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
of a
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
to the female stigma of a flower. This helps to bring about
fertilization Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...
of the
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the sporangium, megasporangium), ...
s in the flower by the male gametes from the pollen grains. Insects are the major pollinators of most plants, and insect pollinators include all families of
bees Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
and most families of aculeate wasps;
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s; many families of
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
; many
lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...
ns (both
butterflies Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
and
moths Moths are a group of insects that includes all members of the order Lepidoptera that are not butterflies. They were previously classified as suborder Heterocera, but the group is paraphyletic with respect to butterflies (suborder Rhopalocera) a ...
); and many families of
beetles Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
. Vertebrates, mainly bats and birds, but also some non-bat mammals (
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
s,
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
s,
possums Possum may refer to: Animals * Didelphimorphia, or (o)possums, an order of marsupials native to the Americas ** Didelphis, a genus of marsupials within Didelphimorphia *** Common opossum, native to Central and South America *** Virginia opossum, ...
,
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s) and some
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s pollinate certain plants. Among the pollinating birds are
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
s, honeyeaters and sunbirds with long beaks; they pollinate a number of deep-throated flowers. Humans may also carry out
artificial pollination Hand-pollination, also known as mechanical pollination, is a technique that can be used to pollinate plants when natural or open pollination is either undesirable or insufficient. Method This method of pollination is done by manually trans ...
. A pollinator is different from a
pollenizer A pollenizer (or polleniser), sometimes pollinizer (or polliniser, see spelling differences), is a plant that provides pollen. The word ''pollinator'' is often used when ''pollenizer'' is more precise. A pollinator is the biotic agent that move ...
, a plant that is a source of pollen for the
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
process.


Background

Plants fall into
pollination syndrome Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different Pollination, pollen vectors, which can be abiotic (wind and water) or biotic, such as birds, bees, flies, and so forth thro ...
s that reflect the type of pollinator being attracted. These are characteristics such as: overall flower size, the depth and width of the corolla, the color (including patterns called
nectar guides Nectar guides are markings or patterns seen in flowers of some angiosperm species, that guide pollinators to their rewards. These markings may appear as lines, spots, or "blotches". Such patterns are also known as "pollen guides" and "honey gui ...
that are visible only in
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
light), the
scent An odor (American English) or odour (English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is a smell or a scent caused by one or more volatilized ...
, amount of nectar, composition of nectar, etc. For example, birds visit red flowers with long, narrow tubes and much nectar, but are not as strongly attracted to wide flowers with little nectar and copious pollen, which are more attractive to beetles. When these characteristics are experimentally modified (altering colour, size, orientation), pollinator visitation may decline. Although non-bee pollinators have been seen to be less effective at depositing pollen than bee pollinators one study showed that non-bees made more visits than bees resulting in non-bees performing 38% of visits to crop flowers, outweighing the ineffectiveness of their ability to pollinate. It has recently been discovered that
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
s, which are not
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s, are also pollinated by insects. In 2016, researchers showed evidence of pollination occurring underwater, which was previously thought not to happen.


Types of pollinators


Insects


Bees

The most recognized pollinators are the various species of bees, which are plainly adapted to pollination. Bees typically are fuzzy and carry an
electrostatic Electrostatics is a branch of physics that studies slow-moving or stationary electric charges. Since classical times, it has been known that some materials, such as amber, attract lightweight particles after rubbing. The Greek word (), mean ...
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 (; ) is an Italian card game, and one of the three major national card games in Italy, the others being and . It is also popular in Argentina and Brazil, brought in by Italian diaspora, Italian immigrants, mostly in the variation. is also p ...
, which is on the hind legs of most bees, and/or the lower abdomen (e.g., of megachilid bees), made up of thick, plumose
seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...
e.
Honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the ...
s,
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
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 The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of the female of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive where it is used as food by the col ...
"). Most bees gather
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
, a concentrated energy source, and pollen, which is high
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
food, to nurture their young, and transfer some among the flowers as they are working. 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 Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
bees such as honey bees need an abundant and steady pollen source to multiply. Honey bees travel from flower to flower, collecting nectar (later converted to
honey Honey is a sweet and viscous substance made by several species of bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of pl ...
), 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 Nutrition is the biochemistry, biochemical and physiology, physiological process by which an organism uses food and water to support its life. The intake of these substances provides organisms with nutrients (divided into Macronutrient, macro- ...
; pollen provides the
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
. When bees are rearing large quantities of 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 A beekeeper is a person who keeps honey bees, a profession known as beekeeping. The term beekeeper refers to a person who keeps honey bees in beehives, boxes, or other receptacles. The beekeeper does not control the creatures. The beekeeper ow ...
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
hives Hives, also known as urticaria, is a kind of skin rash with red or flesh-colored, raised, itchy bumps. Hives may burn or sting. The patches of rash may appear on different body parts, with variable duration from minutes to days, and typically ...
of honey bees are
contract A contract is an agreement that specifies certain legally enforceable rights and obligations pertaining to two or more parties. A contract typically involves consent to transfer of goods, services, money, or promise to transfer any of thos ...
ed out as pollinators by 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 bees, and leaf cutter bees are cultured and sold for 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 The Western Hemisphere is the half of the planet Earth that lies west of the Prime Meridian (which crosses Greenwich, London, United Kingdom) and east of the 180th meridian.- The other half is called the Eastern Hemisphere. Geopolitically, ...
; 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.


Butterflies and moths

Lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...
(
butterflies Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
and
moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...
s) may also pollinate to various degrees. They are not major pollinators of food crops, but various moths are important pollinators of other commercial crops such as
tobacco Tobacco is the common name of several plants in the genus '' Nicotiana'' of the family Solanaceae, and the general term for any product prepared from the cured leaves of these plants. More than 70 species of tobacco are known, but the ...
. 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 hawk moth, Morgan's sphinx. ''
Yucca ''Yucca'' ( , YUCK-uh) is both the scientific name and common name for a genus native to North America from Panama to southern Canada. It contains 50 accepted species. In addition to yucca, they are also known as Adam's needle or Spanish-bayon ...
'' species provide other examples, being fertilised in elaborate ecological interactions with particular species of yucca moths.


Flies

Many bee flies, and some Tabanidae and Nemestrinidae are particularly adapted to pollinating
fynbos Fynbos (; , ) is a small belt of natural shrubland or heathland vegetation located in the Western Cape and Eastern Cape provinces of South Africa. The area is predominantly coastal and mountainous, with a Mediterranean climate. The fynbos ...
and
Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe Khoemana (also known as !Orakobab or Korana) word is a semidesert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its extent is ...
plants with narrow, deep 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 woodland geranium ( ''Geranium sylvaticum'' L.) as effectively as bees. Carrion flies and flesh flies in families such as
Calliphoridae The Calliphoridae (commonly known as blowflies, blow flies, blow-flies, carrion flies, bluebottles, or greenbottles) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing bait, ...
and
Sarcophagidae Sarcophagidae () are a family (biology), family of fly, flies commonly known as flesh flies. They differ from most flies in that they are Ovoviviparity, ovoviviparous, opportunistically depositing hatched or hatching maggots instead of eggs on c ...
are important for some species of plants whose flowers exude a fetid odor. The plants' ecological strategy varies; several species of ''
Stapelia ''Stapelia'' is a genus of low-growing, spineless, stem succulent plants, predominantly from South Africa with a few from other parts of Africa. Several Asian and Latin American species were formerly included but they have all now been transfer ...
'', for example, attract carrion flies that futilely lay their eggs on the flower, where their larvae promptly starve for lack of
carrion Carrion (), also known as a carcass, is the decaying flesh of dead animals. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...
. 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.
Hoverflies Hoverflies, also called flower flies or syrphids, make up the insect family (biology), family Syrphidae. As their common name suggests, they are often seen Hover (behaviour), hovering or nectaring at flowers; the adults of many species feed main ...
are important pollinators of flowering plants worldwide. Often hoverflies are considered to be the second most important pollinators after wild bees. 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. Another plant, the slipper orchid in southwest China, also achieves pollination by deceit by exploiting the
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