Wind: Many flowers are wind pollinated.

It is not an efficient method of pollination because much pollen must be produced with the hope that some of it may land on a receptive stigma of the right species. Characteristic features of wind pollinated flowering plants include: They produce huge amounts of non sticky pollen They often lack a large and showy calyx or corolla They have many flowers packed into a inflorescence They have large stigmas They have large, well exposed anthers Water: Pollination by water is not common but a few seed grasses and water weeds release their pollen into the water which is passively carried to other flowers by water currents. Insects: A great many flowers are pollinated by insects. These flowers do not tend to have any common characteristics because many different types of insects have very different ways of pollinating flowers e.g. bees (the most common insect pollinators), butterflies, moths, beetles and wasps. Many insect pollinated flowers not only have bright colours, but their petals have nectar guides which contain UV absorbing pigments. The nectar guides lead the insect towards the nectar - the reward the insect receives for visiting the flower. Pollen is deposited on the insect from the stamens when it visits the flower to collect or drink the nectar, and is deposited on the stigma of the next flower it visits. Flowers which are visited by nocturnal insects have less showy corollas but are often strongly scented. Other flowers are brown in colour and smell like carrion and attract flies, which pollinate them. Some flowers may get robbed of their nectar by insects which do not pollinate them. Some plants have therefore developed complex structures which prevent all but specific insects species from reaching the nectar and getting pollen deposited on them, which will transferred to the stigma of the next flower they visit. Mammals: Some flowers are pollinated by small mammals such as bats and rodents. Mammal pollinated flowers have the following characteristics: They often have a strong scent e.g. those which attach mice have a yeasty odour. They are often brown or white in colour. They are quite sturdy in structure in order to bear the vigorous activity of the small mammals while they are feeding on the nectar they provide. They offer their mammal pollinators a reward of large amounts of nectar

Birds: Bird pollinated flowers are much more common than mammal pollinated flowers. Two large groups of birds which pollinate flowers are the sunbirds of Africa and Asia and the Hummingbirds of the Americas. Both groups of birds have long beaks which allow them to reach inside the corolla tubes of flowers. Hummingbirds are well known for their ability to hover in

front of the flowers while drinking the nectar. Sunbirds however sit on the flower stalk and collect the nectar. Bird pollinated flowers have the following characteristics: They often have a red, orange or yellow corolla, calyx, bracts or stamens which are attractive to birds The are not usually scented because most birds do not have a well developed sense of smell They provide a large amount of nectar as a reward for the bird pollinators

A pollinator is the biotic agent (vector) that moves pollen from the male anthers of a flower to the female stigma of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain. Though the terms are sometimes confused, a pollinator is different from a pollenizer, which is a plant that is a source of pollen for the pollination process. Plants fall into pollination syndromes 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 that are visible only in ultraviolet light), the scent, amount of nectar, composition of nectar, etc.[1] For example, birds visit red flowers with long narrow tubes and lots of 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.[2][3] Types of pollinators [edit] Bees

Lipotriches sp. bee pollinating flowers The most recognized pollinators are the various species of bees, 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, which is on the hind legs of most bees, and/or the lower abdomen (e.g., of megachilid bees), made up of thick, plumose setae. Honey bees, bumblebees, 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 inadvertently 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 bees such as honey bees need an abundant and steady source of pollen to multiply. [edit] Honey bees

Honey bee with pollen adhering. 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 (beekeepers say hives are "building"), bees deliberately gather pollen to meet the nutritional needs of the brood. A honey bee that is deliberately gathering pollen is up to ten times more efficient as a pollinator than one that is primarily gathering nectar and only unintentionally transferring pollen. 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 hives of honey bees are contracted out as pollinators by beekeepers, and honey bees are by far the most important commercial pollinating agents, but many other kinds of pollinators, from bluebottle 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 it should be remembered that honey bees are not native to the Western Hemisphere; all pollination of native plants in the Americas has been historically performed by various native bees.

An Australian painted lady with its proboscis extended during feeding. Butterflies are recognised pollinators though not as effective as bees. [edit] Other insects Lepidoptera (butterflies and moths) also pollinate plants to various degrees.[4] They are not major pollinators of food crops, but various moths are important pollinators of other commercial crops such as tobacco. Pollination by certain moths may be important however, even crucial, for some wildflowers mutually adapted to specialist pollinators. Spectacular examples include orchids such as Angraecum sesquipedale, dependant on a particular hawk moth, Morgan's Sphinx. Yucca species provide other examples, being fertilised in elaborate ecological interactions with particular species of yucca moths.

Pollen wasp foraging; note hairy exterior, useful in pollen gathering Many other insects 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. For example, males of many species of Hymenoptera, including many hunting wasps and solitary bees, rely on freely flowering plants as sources of energy (in the form of nectar) and also as territories for meeting nubile females that visit the flowers. Prominent examples are Wasps (especially Sphecidae and Vespidae). Pollen wasps in particular are widely regarded as comprising the Masarinae, a subfamily of the Vespidae, and they are remarkable in that they specialise in gathering pollen for feeding their larvae. Many of them are atypically hairy for wasps, an adaptation for collecting large quantities of pollen. [[Syrphidae|syrphid flies, such as hoverflies and drone flies, are important non-selective pollinators. Many bee flies, some Tabanidae and Nemestrinidae are particularly adapted to pollinating fynbos and Karoo plants with narrow, deep corolla tubes, such as Lapeirousia species. Part of the adaptation takes the form of remarkably long proboscises.

Tabanid on thistle flower Beetles of species that specialise in eating pollen, nectar, or flowers themselves, are important crosspollinators of some plants such as members of the Araceae and Zamiaceae, that produce prodigious amounts of pollen. Others, for example the Hopliini, specialise in free-flowering species of the Asteraceae and Aizoaceae. Various midges, and thrips are comparatively minor opportunist pollinators. Ants also pollinate some kinds of flowers, but for the most part they are parasites, robbing nectar without conveying useful amounts of pollen to a stigma. Whole groups of plants, such as certain fynbos Moraeas and Ericas produce flowers on sticky [[Peduncle (botany)|peduncles] or with sticky corolla tubes that only permit access to flying pollinators, whether bird, bat or insect. Carrion flies and flesh flies in families such as Calliphoridae and Sarcophagidae are important for some species of plants whose flowers 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.

Some male Bactrocera fruit flies are exclusive pollinators of some wild Bulbophyllum orchids that have a specific chemical attractant present in their floral fragrance.[5][6] A class of strategy of great biological interest is that of sexual deception, where plants, generally orchids, produce remarkably complex combinations of pheromonal attractants and physical mimicry that induce male bees or wasps to attempt to mate with them, conveying pollinia in the process. Examples are known from all continents apart from Antarctica, though Australia appears to be exceptionally rich in examples.[7] Some Diptera (flies) may be the main pollinators in higher elevations of mountains whereas Bombus species are the only pollinators among Apoidea in alpine regions at timberline and beyond. Other insect orders are rarely pollinators, and then typically only incidentally (e.g., Hemiptera such as Anthocoridae, Miridae). [edit] Vertebrates Bats are important pollinators of some tropical flowers. Birds, particularly hummingbirds, honeyeaters and sunbirds also accomplish much pollination, especially of deep-throated flowers. Other vertebrates, such as monkeys, lemurs, possums, rodents and lizards[8] have been recorded pollinating some plants. Humans can be pollinators, as many gardeners have discovered that they must hand pollinate garden vegetables, whether because of pollinator decline (as has been occurring in parts of the U.S. since the mid-20th century) 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 pores. This can be done by wind, by humans, or by a 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. [edit] Pollinator population declines and conservation In 1999 the Convention on Biological Diversity issued the So Paulo Declaration on Pollinators, recognizing the critical role that these species play in supporting and maintaining terrestrial productivity as well as the survival challenges they face due to anthropogenic change. Today pollinators are considered to be in a state of decline; some species, such as Franklins bumble bee (Bombus franklini) have been red-listed and are in danger of extinction. Declines in the health and population of pollinators pose what could be a significant threat to the integrity of biodiversity, to global food webs, and to human health. At least 80% of our world's crop species require pollination to set seed. An estimated one out of every three bites of food comes to us through the work of animal pollinators.

are insects.[3] Entomophily, pollination by insects, often occurs on plants that have developed colored petals and a strong scent to attract insects such as, bees, wasps and occasionally ants (Hymenoptera), beetles (Coleoptera), moths and butterflies (Lepidoptera), and flies (Diptera). In zoophily, pollination is performed by vertebrates such as birds and bats, particularly, hummingbirds, sunbirds, spiderhunters, honeyeaters, and fruit bats. Plants adapted to using bats or moths as pollinators typically have white petals and a strong scent, while plants that use birds as pollinators tend to develop red petals and rarely develop a scent (few birds rely on a sense of smell to find plantbased food). Insect pollinators such as honeybees (Apis mellifera),[4] bumblebees (Bombus terrestris),[5][6] and butterflies (Thymelicus flavus) [7] have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants.[8] This can be beneficial for the pollenisers, as flower constancy prevents the loss of pollen during interspecific flights and pollinators from clogging stigmas with pollen of other flower species.[9]

Animal vectors From this need widespread and complex kinds of mutualism (mutually beneficial interactions) have evolved between plants and animals. The pollen-transporting agent is frequently an insect or other flying animal. (Flying animals are more mobile than grounded species, and thus more likely to visit widely-separated plants.) In order to get pollinated, a flower must both make its presence known (advertise), and provide an incentive (a reward) for an animal to make repeated visits to flowers of the same species. The advertisements are fragrance and/or conspicuous color. Two kinds of food are the usual reward. Nectar is a sugar solution that provides energy for flight. Flying requires much more energy than terrestrial locomotion. Pollen, besides being the male gene-bearer of a flower, is also rich in proteins essential for maintaining animal tissues and for raising young. In place of nectar some flowers offer oil (fat), another energy food. Others provide fragrances that the pollinator gathers to use for its own reproductive advertisement, and a few fascinating species employ deceit and provide no reward (see the species account on pipevine for an example).

White lined sphinx mothan important pollinator in the Sonoran Desert. (Source: Arizona-Sonora Desert Museum) The sugar in nectar and the protein in pollen are expensive to produce, so there is selective pressure to use these resources efficiently. It is important that animals other than the pollinators do not eat (steal) the nectar and pollen, and that the pollinators transport pollen to other flowers of the same species and deposit it in the right place. Natural selection has produced specialization: most plants with animal-pollinated flowers attract only a few species of animals which have the right size and behavior to reach the reward and pick up pollen. The more than 100 million years of coevolution

between flowering plants and their pollinators has greatly contributed to the huge number of species in both kingdoms (300,000 flowering plants, 350 hummingbirds, and 15,000 known bees in the world). It also explains why there are so many different shapes and colors of flowers. Flowers can be classified into several pollination syndromes according to their pollinators. (A syndrome is a set of characteristics associated with a specific phenomenon.) This is not the same classification as systematic taxonomy and does not reflect the evolutionary relationships among plants. Species in the same family or even the same genus may attract different pollinators. The hummingbird pollination syndrome is one of the most easily recognized. Hummingbirds are large compared to most insects, almost unique in their ability to feed while hovering, and daytime-active; they have no sense of smell, but have long narrow beaks and tongues that can probe deep narrow tubes, and excellent color vision. Hummingbird flowers tend to be long-tubular, non-fragrant, sideways- or downward-facing, day-blooming, and brightly colored. Bees and most other animals cannot easily land on a hanging flower, and even if they succeed they cannot reach the nectar at the base of the narrow tube. There are common misconceptions that all hummingbird flowers are red and that hummingbirds can see only the warm colors of the spectrum. It is true that most hummingbird flowers in the temperate biomes are red, but in the tropics they come in many colors. The predominance of red in temperate hummingbird flowers may be a disincentive to bees. Bees are aggressive pollen collectors in temperate climates. But they cannot see red, so red flowers do not appear conspicuous to them.

Pollination is simply the transfer of pollen from anthers to stigmas of flowers. It is the next step after flower production in the sexual reproduction of plants, both angiosperms (flowering plants) and gymnosperms (conifers and their relatives). Pollination is effected by many means. Plants may be pollinated by wind, water, or gravity. Pollination by insects is thought to be basic to the evolution of flowering plants, although even primitive Gymnospermae may have been pollinated by insects prior to the time of the evolution of flowering plants (see Kevan, 2001 for review). In short, pollination is an important mutualism and ecosystem service that links the dominant flora with the dominant fauna, the insects of the world, from the tropics to the arctic. This ancient and co-evolved process arose at about, or before, the age of the dinosaurs and has allowed the marvelous radiation of flowering plants and pollinating insects that now comprise part of the green biofilm of the terrestrial surface, robed in the worlds largest membrane, the atmosphere (Thomas, 1974).

Wind pollination and pollination by vertebrates are considered to be further refinements of the basic pattern. Recently, it has become apparent that pollination mutualisms involving various plants and pollinating bees, beetles, flies, butterflies, other insects, birds, bats, and other mammals are becoming jeopardized as the worlds biological diversity is being so rapidly eroded by human activities (see Buchmann and Nabhan, 1996; Kearns et al., 1998; Kevan, 1999, 2001; Kevan et al. [eds], 2002; Kevan and Wojcik, in press; for reviews).