Zoophily

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A rufous hummingbird (Selasphorus rufus) is attracted to brightly colored flowers and assists the pollination of the plant. Selasphorus rufus1.jpg
A rufous hummingbird ( Selasphorus rufus ) is attracted to brightly colored flowers and assists the pollination of the plant.

Zoophily, or zoogamy, is a form of pollination whereby pollen is transferred by animals, usually by invertebrates but in some cases vertebrates, [1] particularly birds and bats, but also by other animals. Zoophilous species frequently have evolved mechanisms to make themselves more appealing to the particular type of pollinator, e.g. brightly colored or scented flowers, nectar, and appealing shapes and patterns. These plant-animal relationships are often mutually beneficial because of the food source provided in exchange for pollination.

Contents

Pollination is defined as the transfer of pollen from the anther to the stigma. [2] There are many vectors for pollination, including abiotic (wind and water) and biotic (animal). There are benefits and costs associated with any vector. For instance, using animal pollination is beneficial because the process is more directed and often results in pollination. At the same time it is costly for the plant to produce rewards, such as nectar, to attract animal pollinators. Not producing such rewards is one benefit of using abiotic pollinators, but a cost associated with this approach is that the pollen may be distributed more randomly. In general, pollination by animals occurs after they reach inside the flowers for nectar. While feeding on the nectar, the animal rubs or touches the stamens and is covered in pollen. Some of this pollen will be deposited on the stigma of the next flower it visits, pollinating the flower. [3]

Insect pollination

This is known as entomophily. There are many different subtypes.

Bee pollination (melittophily)

There are diverse types of bees (such as honeybees, bumblebees, and orchid bees), forming large groups that are quite distinctive in size, tongue length and behaviour (some solitary, some colonial); thus generalization about bee pollination is difficult. [4] Some plants can only be pollinated by bees because their anthers release pollen internally, and it must be shaken out by buzz pollination (also known as "sonication"). Bees are the only animals that perform this behaviour. Bumblebees and solitary bees sonicate, but honeybees do not. [5] About 9% of the flowers of the world are primarily pollinated using buzz pollination. [6]

Wasp pollination

Wasps are also responsible for the pollination of several plants species, being important pollen vectors, and in some cases, even more efficient pollinators than bees. [7]

Butterfly pollination (psychophily)

Despite their complete dependence on flowers for sustenance as imagoes, butterflies are generally poor pollinators, lacking specific structures to carry pollen. [8] Nonetheless, some plants appear to have specialised on attracting butterflies. Buddleja is a well-known example. [9] The species in the orchid genus Bonatea are all pollinated by moths, except for Bonatea cassidea which has evolved into a psychophile. This orchid affixes its pollinaria firmly between the palpi of visiting butterflies. Unlike its relatives, this orchid species exhibits diurnal anthesis, a weak scent which is virtually absent at night, and has short spurs containing small amounts of relatively dilute sucrose-rich nectar -these are all considered psychophilous traits. B. cassidea has white flowers, [10] but butterfly-attracting flowers are often coloured. Unlike bees and wasps, some butterflies such as swallowtails are able to see the colour red. Butterflies also require a platform on which to land. [8] [9]

Moth pollination (phalaenophily)

Among the more important moth pollinators are the hawk moths (Sphingidae). Their behaviour is similar to hummingbirds: they hover in front of flowers with rapid wingbeats. Most are nocturnal or crepuscular.

Other moths (Noctuids, Geometrids, Pyralids, for example) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads). [11]

Fly pollination (myophily and sapromyophily)

Flies tend to be important pollinators in high-altitude and high-latitude systems, where they are numerous and other insect groups may be lacking. [12] There are two main types of fly pollination: myophily and sapromyophily.

Myophily includes flies that feed on nectar and pollen as adults - particularly bee flies (Bombyliidae), hoverflies (Syrphidae), and others - and these regularly visit flowers. In contrast, male fruit flies (Tephritidae) are enticed by specific floral attractants emitted by some wild orchids which do not produce nectar. Chemicals emitted by the orchid act as the fly's sex pheromone precursor or booster. [13] [14]

Sapromyophiles, on the other hand, normally visit dead animals or dung. They are attracted to flowers which mimic the odor of such objects. The plant provides them with no reward and they leave quickly unless it has traps to slow them down. Such plants are far less common than myophilous ones. [15]

Beetle pollination (cantharophily)

Beetles are particularly important in some parts of the world such as semi-arid areas of Southern Africa and southern California [15] and the montane grasslands of KwaZulu-Natal in South Africa. [16]

Cantharophily is often the main pollination system in the Araceae family. It occurs in genera such as Amorphophallus , [17] Dieffenbachia , [18] Monstera , [19] Philodendron [20] and Theriophonum . [21] A well-known example is the gigantic inflorescence of Amorphophallus titanum . This bloom appears like column sticking out of a vast sheet of rotting flesh. It is able to generate heat, which it uses to exude a powerful foetid and revolting odour at night. This attracts necrophagous beetles, and also specialised beetle predators of these beetles -the plant is essentially tricking the beetles into believing that there is food or a place to lay their eggs. Araceae flowers often trap beetles in a compartment with the pollen: the beetles must pass through a constriction of the spathe to get inside, but the plant can tighten the spathe against the spadix and thus close the constriction for a time. [22] There is also some evidence that the giant inflorescence, which heats itself to 36°C, thus shines like an invisible infrared beacon in the dark of night on the jungle floor, unseen by humans but detectable by insects. [23] The blooms of Philodendron adamantinum are able to stick a glob of resin on the otherwise smooth back of the beetles it attracts, modifying them so they are better equipped to carry pollen to the next inflorescence. [20]

Other

In Dioscorea chouardii , pollination is performed by ants. [24]

Lizard pollination

According to one 2003 paper, it is possible that lizard pollination is underestimated. Lizards have been known to feed on nectar since 1977. However, only two species of lizard from New Zealand has been shown to carry pollen as of 2003, although it is unknown if they actually pollinate flowers. There is also circumstantial evidence Podarcis lilfordi may help pollinate Euphorbia dendroides on the Balearic Islands; one study found seed set higher in areas with a higher lizard density. Both these species are not dependant on nectar as a food source, and do not appear to have evolved specific adaptations to exploit it. Despite the lack of evidence, the authors nonetheless theorise that some plants on small islands may have mutualistically evolved to accommodate lizard pollination. [25]

The lizards known to carry pollen are the geckoes Hoplodactylus duvauceli and Dactylocnemis pacificus . Between them, they visit the flowers of at least four species of plant: Metrosideros excelsa , Phormium tenax , Myoporum laetum and in one case Hebe bollonsii , although the structure of the flowers of the last three species do not allow pollen transfer to occur during feeding by lizards, which are better seen as robbers of nectar, these plants are adapted for bird or insect pollination. The only pollen ever found on these lizards is that of Metrosideros excelsa, which has been found on 3% to 47% of the lizards caught on these trees, depending on the night. These lizards are only attracted to the nectar on flowers, not the pollen. If it is proved that lizards are the main pollinators of these plants, the authors of the study theorise that the possible pollination syndrome associated with lizard pollination could be copious nectar and possibly scented flowers in the case of nocturnal lizards. Flowers or inflorescences must also be robust enough to support the weight of the pollinator while feeding. They also theorise that the colour red that the flowers of most species Metrosideros have, which generally attracts birds, might somehow also serve to repel insects and thus leave more nectar for the geckoes. Metrosideros excelsa is a generalist, which is pollinated by both birds and insects. There does not appear to be any mutualistic relationship between the geckoes and Metrosideros excelsa, neither species requires the presence of the other to thrive. [26]

Bird pollination

The pollen-laden anthers of a Tapinanthus mistletoe rub against the forecrown of a nectar-feeding amethyst sunbird. Amethyst sunbird, Chalcomitra amethystina, female at Kloofendal Nature Reserve, Johannesburg, South Africa (21354148246), crop.jpg
The pollen-laden anthers of a Tapinanthus mistletoe rub against the forecrown of a nectar-feeding amethyst sunbird.

The term ornithophily is used to describe pollination specifically by birds. Bird pollination is done primarily by bird species that specialize on eating nectar, which is known as nectarivory. Hummingbirds, found only in the Americas, and many other bird species throughout the world are obligate nectarivores and important pollinators. These include sunbirds, sugarbirds, honeyeaters, flowerpeckers and honeycreepers, these have long narrow bills suited for probing flowers. However, many shorter-billed birds can also pollinate, including white-eyes, bananaquits, flowerpiercers, lories and lorikeets, many of which have more generalist diets and also feed on insects, fruits, and seeds (short-billed birds can also steal nectar from long flowers, as suggested by the name 'flowerpiercer'). Hummingbirds are the oldest group of nectar-specialist birds, with the greatest degree of specialization on nectar. [27] The trumpet creeper ( Campsis radicans ) is a plant species adapted specifically for hummingbirds. [28]

Plants pollinated by birds often have elongated or tube-shaped, brightly colored diurnal flowers that are red or orange, but no odor because birds have a poor sense of smell.

Some 500 genera of plants are pollinated by birds. [29]

Bat pollination

Bat pollination is called chiropterophily. Hundreds of tropical plant species completely, or partially, dependent on bats for pollination in tropical regions. [30] As of 2009, 28 orders, 67 families and about 528 species of angiosperms in some 250 genera are known to be pollinated by nectar-feeding bats. [29] In some cases nectivorous bats are do not pollinate certain species, even while they do pollinate others, but instead act as 'nectar robbers' and exploit other pollination systems. [31] Only two families of bats (not including the somewhat bizarre Mystacinidae) contain nectivores, and morphologically specialized nectivores are in the minority in both of these families, Pteropodidae (15 species) and Phyllostomidae (perhaps up to 38 species in the subfamily called Glossophaginae). Pteropodidae are large fruit bats from the Old World which must perch on the plant to access the nectar and do not have the ability to echolocate, whereas the much smaller Phyllostomidae only occur in the New World and have the ability to hover and echolocate. [29]

Plants pollinated by bats often have white or pale nocturnal flowers that are large and bell shaped. Many of these flowers have large amounts of nectar, and emit a smell that attracts bats, such as a strong fruity or musky odor. Bats use certain chemical cues to locate food sources. They are attracted to odors that contain esters, alcohols, aldehydes, and aliphatic acids. [32] Bats often have excellent spatial memory and will visit specific flowering plants repeatedly. [33]

Pollination by other mammals

Non-flying mammals (i.e. all mammals except bats) have been found to feed on the nectar of several species of plant. [34] This is known as therophily. [35] Though some of these mammals are pollinators, others do not carry or transfer enough pollen to be considered pollinators. The group of non-flying pollinators is composed of marsupials, lemurs, rodents, shrews, and elephant shrews. [31] [34] [36] [37] As of 1997 studies have documented non-flying mammal pollination involving at least 59 species of mammal distributed among 19 families and six orders. As of 1997, there were 85 species of plants from 43 genera and 19 families which were visited by these mammals. In many cases, a plant species is visited by a range of mammals. Two examples of multiple mammal pollination are the genus Quararibea which is visited by twelve species and Combretum which is visited by eight (although not all these animals actually pollinate the flowers). [38]

The honey possum from southwestern Australia is the only entirely nectarivorous mammal which is not a bat. [34]

One example of a plant using animal pollinators is the bulb Massonia depressa . At least four rodent species were found to be visiting M. depressa during the night. Traits of the M. depressa flowers support non-flying mammal pollination: it has a dull-colored and very sturdy inflorescence at ground level, has a strong yeasty odor, and secretes copious amounts of sucrose-dominant nectar during the night. The nectar of M. depressa was also found to be 400 times as viscous, i.e. sticky, as an equivalent sugar solution. This jelly-like consistency of the nectar may discourage insect consumption while also facilitating lapping by rodents. [37]

See also

Related Research Articles

<span class="mw-page-title-main">Hypanthium</span> Structure in angiosperms where basal portions form a cup-shaped tube

In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. It often contains the nectaries of the plant. It is present in many plant families, although varies in structural dimensions and appearance. This differentiation between the hypanthium in particular species is useful for identification. Some geometric forms are obconic shapes as in toyon, whereas some are saucer-shaped as in Mitella caulescens.

<span class="mw-page-title-main">Pollinator</span> Animal that moves pollen from the male anther of a flower to the female stigma

A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains.

<span class="mw-page-title-main">Petal</span> Part of most types of flower

Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the corolla. Petals are usually accompanied by another set of modified leaves called sepals, that collectively form the calyx and lie just beneath the corolla. The calyx and the corolla together make up the perianth, the non-reproductive portion of a flower. When the petals and sepals of a flower are difficult to distinguish, they are collectively called tepals. Examples of plants in which the term tepal is appropriate include genera such as Aloe and Tulipa. Conversely, genera such as Rosa and Phaseolus have well-distinguished sepals and petals. When the undifferentiated tepals resemble petals, they are referred to as "petaloid", as in petaloid monocots, orders of monocots with brightly colored tepals. Since they include Liliales, an alternative name is lilioid monocots.

<span class="mw-page-title-main">Hoverfly</span> Family of insects

Hoverflies, also called flower flies or syrphids, make up the insect family Syrphidae. As their common name suggests, they are often seen hovering or nectaring at flowers; the adults of many species feed mainly on nectar and pollen, while the larvae (maggots) eat a wide range of foods. In some species, the larvae are saprotrophs, eating decaying plant and animal matter in the soil or in ponds and streams. In other species, the larvae are insectivores and prey on aphids, thrips, and other plant-sucking insects.

<span class="mw-page-title-main">Pollination</span> Biological process occurring in plants

Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, for example beetles; birds, butterflies, and bats; water; wind; and even plants themselves. Pollinating animals travel from plant to plant carrying pollen on their bodies in a vital interaction that allows the transfer of genetic material critical to the reproductive system of most flowering plants. When self-pollination occurs within a closed flower. Pollination often occurs within a species. When pollination occurs between species, it can produce hybrid offspring in nature and in plant breeding work.

<i>Philodendron</i> Genus of flowering plants

Philodendron is a large genus of flowering plants in the family Araceae. As of September 2015, the World Checklist of Selected Plant Families accepted 489 species; other sources accept different numbers. Regardless of number of species, the genus is the second-largest member of the family Araceae, after genus Anthurium. Taxonomically, the genus Philodendron is still poorly known, with many undescribed species. Many are grown as ornamental and indoor plants. The name derives from the Greek words philo- 'love, affection' and dendron 'tree'. The generic name, Philodendron, is often used as the English name.

<span class="mw-page-title-main">Entomophily</span> Form of pollination by insects

Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, sometimes with conspicuous patterns leading to rewards of pollen and nectar; they may also have an attractive scent which in some cases mimics insect pheromones. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, and in some species also pollen baskets on their hind legs. This required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups. Both the size and the density of a population are known to affect pollination and subsequent reproductive performance.

<span class="mw-page-title-main">Carrion flower</span> Flowers that smell like rotting flesh

Carrion flowers, also known as corpse flowers or stinking flowers, are mimetic flowers that emit an odor that smells like rotting flesh. Apart from the scent, carrion flowers often display additional characteristics that contribute to the mimesis of a decaying corpse. These include their specific coloration, the presence of setae and orifice-like flower architecture. Carrion flowers attract mostly scavenging flies and beetles as pollinators. Some species may trap the insects temporarily to ensure the gathering and transfer of pollen.

<span class="mw-page-title-main">Nectar</span> Sugar-rich liquid produced by many flowering plants, that attracts pollinators and insects

Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, which in turn provide herbivore protection. Common nectar-consuming pollinators include mosquitoes, hoverflies, wasps, bees, butterflies and moths, hummingbirds, honeyeaters and bats. Nectar plays a crucial role in the foraging economics and evolution of nectar-eating species; for example, nectar foraging behavior is largely responsible for the divergent evolution of the African honey bee, A. m. scutellata and the western honey bee.

<span class="mw-page-title-main">Flower</span> Reproductive structure in flowering plants

A flower, also known as a bloom or blossom, is the reproductive structure found in flowering plants. Flowers consist of a combination of vegetative organs – sepals that enclose and protect the developing flower, petals that attract pollinators, and reproductive organs that produce gametophytes, which in flowering plants produce gametes. The male gametophytes, which produce sperm, are enclosed within pollen grains produced in the anthers. The female gametophytes are contained within the ovules produced in the carpels.

<span class="mw-page-title-main">Nectarivore</span> Animal in which nectar is a main source of nutrition in their diet

In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants.

<span class="mw-page-title-main">Ornithophily</span> Pollination by birds

Ornithophily or bird pollination is the pollination of flowering plants by birds. This sometimes 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. 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.

<span class="mw-page-title-main">Palynivore</span> Group of herbivorous animals

In zoology, a palynivore /pəˈlɪnəvɔːɹ/, meaning "pollen eater" is an herbivorous animal which selectively eats the nutrient-rich pollen produced by angiosperms and gymnosperms. Most true palynivores are insects or mites. The category in its strictest application includes most bees, and a few kinds of wasps, as pollen is often the only solid food consumed by all life stages in these insects. However, the category can be extended to include more diverse species. For example, palynivorous mites and thrips typically feed on the liquid content of the pollen grains without actually consuming the exine, or the solid portion of the grain. Additionally, the list is expanded greatly if one takes into consideration species where either the larval or adult stage feeds on pollen, but not both. There are other wasps which are in this category, as well as many beetles, flies, butterflies, and moths. One such example of a bee species that only consumes pollen in its larval stage is the Apis mellifera carnica. There is a vast array of insects that will feed opportunistically on pollen, as will various birds, orb-weaving spiders and other nectarivores.

<span class="mw-page-title-main">Pollination syndrome</span> Flower traits that attract pollinators

Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different pollen vectors, which can be abiotic or biotic, such as birds, bees, flies, and so forth through a process called pollinator-mediated selection. These traits include flower shape, size, colour, odour, reward type and amount, nectar composition, timing of flowering, etc. For example, tubular red flowers with copious nectar often attract birds; foul smelling flowers attract carrion flies or beetles, etc.

<span class="mw-page-title-main">Nectar robbing</span> Foraging behavior

Nectar robbing is a foraging behavior utilized by some organisms that feed on floral nectar, carried out by feeding from holes bitten in flowers, rather than by entering through the flowers' natural openings. "Nectar robbers" usually feed in this way, avoiding contact with the floral reproductive structures, and therefore do not facilitate plant reproduction via pollination. Because many species that act as pollinators also act as nectar robbers, nectar robbing is considered to be a form of exploitation of plant-pollinator mutualism. While there is variation in the dependency on nectar for robber species, most species rob facultatively.

<span class="mw-page-title-main">Pollination trap</span>

Pollination traps or trap-flowers are plant flower structures that aid the trapping of insects, mainly flies, so as to enhance their effectiveness in pollination. The structures of pollination traps can include deep tubular corollas with downward pointing hairs, slippery surfaces, adhesive liquid, attractants, flower closing and other mechanisms.

<span class="mw-page-title-main">Monocotyledon reproduction</span> Flowering plant reproduction system

The monocots are one of the two major groups of flowering plants, the other being the dicots. In order to reproduce they utilize various strategies such as employing forms of asexual reproduction, restricting which individuals they are sexually compatible with, or influencing how they are pollinated. Nearly all reproductive strategies that evolved in the dicots have independently evolved in monocots as well. Despite these similarities and their close relatedness, monocots and dicots have distinct traits in their reproductive biologies.

<span class="mw-page-title-main">Pollination of orchids</span>

The pollination of orchids is a complex chapter in the biology of this family of plants that are distinguished by the complexity of their flowers and by intricate ecological interactions with their pollinator agents. It has captured the attention of numerous scientists over time, including Charles Darwin, father of the theory of evolution by natural selection. Darwin published in 1862 the first observations of the fundamental role of insects in orchid pollination, in his book The Fertilization of Orchids. Darwin stated that the varied stratagems orchids use to attract their pollinators transcend the imagination of any human being.

<span class="mw-page-title-main">Pollinator garden</span> Type of garden

A pollinator garden is a type of garden designed with the intent of growing specific nectar and pollen-producing plants, in a way that attracts pollinating insects known as pollinators. Pollinators aid in the production of one out of every three bites of food consumed by humans, and pollinator gardens are a way to offer support for these species. In order for a garden to be considered a pollinator garden, it should provide various nectar producing flowers, shelter or shelter-providing plants for pollinators, and avoid the use of pesticides.

<span class="mw-page-title-main">UV coloration in flowers</span> Natural phenomenon

UV coloration is a natural phenomenon that leads to unique interactions between organisms that have evolved the ability to perceive these wavelengths of light. It serves as one method to attract pollinators to the flower along with scent, shape, and nectar quality. Flowers are known for their range of visible colors that humans can see with their eyes and observe an array of different shades and patterns. The naked eye cannot see the ultraviolet coloration many flowers employ to bring attention to themselves. By either reflecting or absorbing UV light waves, flowers are able to communicate with pollinators. This allows plants that may require an animal pollinator to stand out from other flowers or distinguish where their flowers are in a muddied background of other plant parts. For the plant, it is important to share and receive pollen so they can reproduce, maintain their ecological role, and guide the evolutionary history of the population.

References

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