Insectivorous bats are the best pollinators of cactus flowers. Pollination of flowers by bats Some flowers are pollinated by bats

29.10.2019

The flowers pollinated by bats are usually large, strong, produce a lot of nectar, are not brightly colored, or often open only after sunset, since bats feed only at night. Many of the flowers are tubular or have other structures to conserve nectar. In many plants that attract bats for pollination or seed dispersal, flowers or fruits either hang on long stalks below the foliage, where it is easier for bats to fly, or form on trunks. Bats search for flowers using their sense of smell, so the flowers have a very strong smell of fermentation or fruit. These animals, flying from tree to tree, lick nectar, eat parts of the flower and pollen, while transferring it on their fur from one plant to another. They pollinate and distribute the seeds of at least 130 angiosperm genera. In North America, long-nosed bats pollinate over 60 species of agave, including those used in Mexican tequila. Flower bats pollinate mainly cacti (Pachycereen) and agaves. The sausage tree, or Ethiopian Kigelia, growing in tropical Africa and Madagascar, is pollinated by bats. Bats pollinate plants such as:
Couroupita guianensis, Cephalocereus (Cephalocereus senilis), African Baobab (Adansonia digitata), Sausage Tree (Kigelia pinnata), Trianea (Trianaea), Breadfruit (Artocarpus altilis), Liana Mucuna holtonii., Blue Agave (Аgave tequilana weber azul), Cocoa (Theobroma cacao), Dracula orchids, Chorisia speciosa, Durian zibethinus.

Pachycereus Pringle pollinated by bats of the Sonoran Desert (Central America)

Selenicereus is another cactus pollinated by bats at night and by bees during the day.

Bats that pollinate flowers feed on nectar. As an adaptation, they developed an elongated muzzle. In North America, there is a genus of bats, which are called so - long-nosed.

Pollination

What is pollination? Bloom- this is the state of plants from the beginning of the opening of flowers to the drying of their stamens and petals . During flowering, pollination of plants occurs.

Pollinationcalled the transfer of pollen from the stamens to the stigma of the pistil. If pollen is transferred from the stamens of one flower to the stigma of the pistil of another flower, then cross pollination . If pollen falls on the stigma of the pistil of the same flower, this is self-pollination .

Cross pollination. With cross-pollination, two options are possible: pollen is transferred to flowers located on the same plant, pollen is transferred to flowers of another plant. In the latter case, it must be borne in mind that pollination occurs only between individuals of the same species!

Cross-pollination can be carried out by wind, water (these plants grow in water or near water: hornwort, naiad, vallisneria, elodea ), insects, and in tropical countries also birds and bats.

Cross-pollination is biologically more appropriate, because the offspring, combining the characteristics of both parents, can better adapt to the environment. Self-pollination has its advantages: it does not depend on external conditions, and the offspring stably retains parental characteristics. For example, if yellow tomatoes are grown, then next year, using their seeds, you can again get the same yellow tomatoes ( tomatoes are usually self-pollinators). Most plants cross-pollinate, although there are few strictly cross-pollinated plants (e.g., rye), more often cross-pollination is combined with self-pollination, which further increases the fitness of plants for survival.

Flower pollination types: self-pollination, cross-pollination

Wind pollinated plants. Plants whose flowers are pollinated by the wind are called wind pollinated . Usually their inconspicuous flowers are collected in compact inflorescences, for example, in a complex spike, or in panicles. They produce a huge amount of small, light pollen. Wind pollinated plants often grow in large groups. Among them are herbs. (timothy grass, bluegrass, sedge) and shrubs and trees (hazel, alder, oak, poplar, birch) . Moreover, these trees and shrubs bloom at the same time as the leaves bloom (or even earlier).

In wind-pollinated plants, the stamens usually have a long filament and carry the anther outside the flower. The stigmas of the pistils are also long, "shaggy" - to catch dust particles flying in the air. These plants also have certain adaptations to ensure that pollen is not wasted, but rather falls on the stigmas of flowers of its own species. Many of them bloom by the hour: some bloom early in the morning, others in the afternoon.

Insect pollinated plants. Insects (bees, bumblebees, flies, butterflies, beetles) are attracted by sweet juice - nectar, which is secreted by special glands - nectaries. Moreover, they are located in such a way that the insect, getting to the nectaries, must touch the anthers and stigma of the pistil. Insects feed on nectar and pollen. And some (bees) even store them for the winter.

Therefore, the presence of nectaries is an important feature of an insect pollinated plant. In addition, their flowers are usually bisexual, their pollen is sticky with outgrowths on the shell to cling to the insect's body. Insects find flowers by a strong smell, by bright colors, by large flowers or inflorescences.

In a number of plants, nectar, which attracts insects, is available to many of them. So on blooming poppies, jasmine, buzulnik, nivyanika you can see bees, and bumblebees, and butterflies, and beetles.

But there are plants that have adapted to a particular pollinator. However, they may have a special structure of the flower. Carnation, with its long corolla, is pollinated only by butterflies, whose long proboscis can reach the nectar. Only bumblebees can pollinate flaxseed, snapdragon : under their weight, the lower petals of the flowers are bent and the insect, reaching the nectar, collects pollen with its shaggy body. The stigma of the pistil is located so that the pollen brought by the bumblebee from another flower must remain on it.

Flowers can smell attractive to different insects or smell particularly strong at different times of the day. Many white or light flowers smell especially strongly in the evening and at night - they are pollinated by moths. Bees are attracted to sweet, “honey” smells, and flies are often not very pleasant smells for us: many umbrella plants smell like this. (snyt, cow parsnip, kupyr) .

Scientists have conducted studies that have shown that insects see colors in a special way and each species has its own preferences. It is not for nothing that in nature all shades of red reign among daytime flowers (but in the dark red is almost indistinguishable), and blue and white are much less.

Why so many devices? In order to have a better chance that pollen will not be wasted, but will fall on the pistil of a flower of a plant of the same species.

Having studied the structure and features of the flower, we can assume which animals will pollinate it. So, fragrant tobacco flowers have a very long tube of fused petals. Therefore, only insects with a long proboscis can reach the nectar. The flowers are white, well visible in the dark. They smell especially strong in the evening and at night. Pollinators - hawk moths, night butterflies, which have a proboscis up to 25 cm long.

The largest flower in the world - rafflesia - painted red with dark spots. It smells like rotten meat. But for flies there is no smell more pleasant. They pollinate this wonderful, rare flower.

Self-pollination. Majority self-pollinating plants are crops (peas, flax, oats, wheat, tomato) , although there are self-pollinating plants among the wild ones.

Some of the flowers are already pollinated in bud. If you open a pea bud, you can see that the pistil is covered with orange pollen. In flax, pollination takes place in an open flower. The flower blooms early in the morning and after a few hours the petals fall off. During the day, the air temperature rises and the filaments twist, the anthers touch the stigma, burst, and the pollen spills out on the stigma. Self-pollinating plants, including linen, can be pollinated and cross-pollinated. Conversely, under unfavorable conditions, self-pollination can occur in cross-pollinated plants.

Cross-pollinated plants in the flower have devices that prevent self-pollination: the anthers mature and shed pollen before the pistil develops; the stigma is located above the anthers; pistils and stamens can develop in different flowers and even on different plants (dioecious).

artificial pollination. In certain cases, a person carries out artificial pollination, that is, he himself transfers pollen from the stamens to the stigma of the pistils. Artificial pollination is carried out for different purposes: to breed new varieties, to increase the yield of some plants. In calm weather, a person pollinates wind-pollinated crops. (corn), and in cold or wet weather - insect pollinated plants (sunflower) . Both wind- and insect-pollinated plants are artificially pollinated; both cross- and self-pollinated.

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Birds, elephants and turtles

The relationship between trees and animals is most often expressed in the fact that birds, monkeys, deer, sheep, cattle, pigs, etc. contribute to the dispersal of seeds, but we will only consider the effect of animal digestive juices on ingested seeds.

Homeowners in Florida have a strong dislike for the Brazilian pepper tree (Schinus terebinthifolius), a beautiful evergreen that turns red berries in December, peeping from dark green scented leaves in such numbers that it resembles a holly. In this magnificent dress, the trees stand for several weeks. Seeds ripen, fall to the ground, but young shoots never appear under the tree.

Arriving in large flocks, the red-throated thrushes descend on pepper trees and fill full crops with tiny berries. Then they flit to the lawns and walk among the sprinklers there. In the spring, they fly north, leaving numerous business cards on Florida lawns, and a few weeks later, pepper trees begin to grow everywhere - and especially in flowerbeds where thrushes searched for worms. A weary gardener has to pull out thousands of sprouts so that the pepper trees do not take over the whole garden. The gastric juice of the red-throated thrush somehow affected the seeds.

Formerly in the United States, all pencils were made from juniper wood (Juniperus silicicola), which grew abundantly on the plains of the Atlantic coast from Virginia to Georgia. Soon, the insatiable demands of industry led to the extermination of all large trees and it was necessary to look for another source of wood. True, a few surviving young junipers reached maturity and began to bear seeds, but under these trees, which in America to this day are called "pencil cedars", not a single sprout appeared.

But driving along rural roads in South and North Carolina, you can see millions of "pencil cedars" growing in straight rows along wire fences, where their seeds have fallen in the excrement of tens of thousands of sparrows and meadow trupials. Without the help of feathered intermediaries, juniper forests would forever remain only a fragrant memory.

This service that birds have rendered to the juniper makes us wonder: to what extent do the digestive processes of animals act on plant seeds? A. Kerner found that most of the seeds, passing through the digestive tract of animals, lose their germination. In Rossler, out of 40,025 seeds of various plants fed to California oatmeal, only 7 germinated.

In the Galapagos Islands off the west coast of South America, a large, long-lived perennial tomato (Lycopersicum esculentum var. minor) grows, which is of particular interest because careful scientific experiments have shown that less than one percent of its seeds naturally germinate. But in the event that the ripe fruits were eaten by giant tortoises, which are found on the island, and remained in their digestive organs for two to three weeks or longer, 80% of the seeds germinated. Experiments have suggested that the giant tortoise is a very important natural mediator, not only because it stimulates the germination of seeds, but also because it ensures their efficient dispersal. The scientists also concluded that seed germination was due not to mechanical, but to enzymatic action on the seeds during their passage through the turtle's digestive tract.

In Ghana Baker ( Herbert J. Baker - Director of the Botanical Gardens of the University of California (Berkeley).) experimented with the germination of baobab and sausage tree seeds. He found that these seeds practically did not germinate without special treatment, while their numerous young shoots were found on stony slopes at a considerable distance from adult trees. These places served as a favorite habitat for baboons, and fruit cores indicated that they were included in the diet of monkeys. The strong jaws of baboons allow them to easily gnaw through the very hard fruits of these trees; since the fruits themselves do not open, without such assistance the seeds would not have the opportunity to disperse. The percentage of germination in seeds extracted from baboon dung was noticeably higher.

In Southern Rhodesia, there is a large, beautiful ricinodendron tree (Ricinodendron rautanenii), which is also called "Zambezian almond" and "Manketti's nut". It bears fruits the size of plums, with a thin layer of pulp surrounding very hard nuts - "edible if you can crack them open," as one forest ranger wrote. The wood of this tree is only slightly heavier than balsa (see ch. 15). The package of seeds that was sent to me said: "Collected from elephant droppings." Naturally, these seeds rarely germinate, but there are a lot of young shoots, since elephants are addicted to these fruits. Passing through the digestive tract of an elephant does not seem to have any mechanical effect on the nuts, although the surface of the samples sent to me was covered with grooves, as if made with the tip of a sharpened pencil. Perhaps these are traces of the action of the gastric juice of an elephant?

C. Taylor wrote to me that the ricinodendron growing in Ghana produces seeds that germinate very easily. However, he adds that musanga seeds may “need to pass through the digestive tract of some animal, as it is extremely difficult to germinate them in nurseries, and in natural conditions the tree reproduces very well.”

Although elephants in Southern Rhodesia cause great damage to the forests of the savannahs, they at the same time ensure the distribution of certain plants. Elephants love camelthorn beans and eat them in large quantities. The seeds come out undigested. During the rainy season, dung beetles bury elephant droppings. Thus, most of the seeds end up in an excellent bed. This is how thick-skinned giants at least partly compensate for the damage they cause to trees, tearing off the bark from them and causing all sorts of other damage to them.

C. White reports that the seeds of the Australian quondong (Elaeocarpus grandis) germinate only after being in the stomach of emus, which love to feast on fleshy, plum-like pericarp.

wasp trees

One of the most misunderstood groups of tropical trees is the fig tree. Most of them come from Malaysia and Polynesia. Corner writes:

“All members of this family (Moraceae) have small flowers. In some, such as breadfruit, mulberries, and fig trees, the flowers are united in dense inflorescences that develop into fleshy buds. In breadfruit and mulberries, the flowers are placed outside the fleshy stem that supports them; the fig trees have them within it. The fig is formed as a result of the growth of the stem of the inflorescence, the edge of which then bends and contracts until a calyx or a jug with a narrow mouth is formed - something like a hollow pear, and the flowers are inside ... The pharynx of the fig is closed by many scales superimposed on each other ...

The flowers of these fig trees are of three types: male with stamens, female, which produce seeds, and gall flowers, so called because they develop larvae of small wasps that pollinate the fig tree. Gallic flowers are sterile female flowers; breaking a ripe fig, they are easy to recognize, as they look like tiny balloons on pedicels, and on the side you can see the hole through which the wasp got out. The female flowers are recognized by the small, flat, hard, yellowish seed they contain, and the male flowers by the stamens...

Pollination of fig blossoms is perhaps the most interesting form of interrelationship between plants and animals known so far. Only tiny insects called fig wasps (Blastophaga) can pollinate the flowers of the fig tree, so the reproduction of fig trees depends entirely on them ... If such a fig tree grows in a place where these wasps are not found, the tree will not be able to reproduce with the help of seeds ... ( Recent studies have established that some fig trees, such as figs, are characterized by the phenomenon of apomixis (fetal development without fertilization). - Approx. ed. But fig wasps, in turn, are completely dependent on the fig tree, since their larvae develop inside gall flowers and the entire life of adults passes inside the fruit - excluding the flight of females from a ripening fig on one plant to a young fig on another. Males, almost or completely blind and wingless, live in the adult stage for only a few hours. If the female fails to find a suitable fig tree, she cannot lay her eggs and dies. There are many varieties of these wasps, each of which appears to serve one or more related species of the fig tree. These insects are called wasps because they are distantly related to true wasps, but they do not sting and their tiny black bodies are no more than a millimeter long...

When the figs on the gall plant ripen, adult wasps hatch from the ovaries of the gall flowers, gnawing through the wall of the ovary. The males fertilize the females inside the fetus and die soon after. The females get out between the scales covering the mouth of the fig. Male flowers are usually located near the throat and open by the time the fig ripens, so that their pollen falls on the female wasps. The wasps, showered with pollen, fly to the same tree, on which young figs begin to develop, and which they probably find with the help of smell. They penetrate into young figs, squeezing between the scales that cover the throat. This is a difficult process ... If a wasp climbs into a fig-gall, its ovipositor easily penetrates through a short style into the ovule, in which one egg is laid ... The wasp moves from flower to flower until its supply of eggs runs out; then she dies of exhaustion, because, having hatched, she does not eat anything ... "

Trees pollinated by bats

In the temperate zones, the pollination of flowers is in most cases done by insects, and it is believed that the lion's share of this work falls on the bee. However, in the tropics, many species of trees, especially those that bloom at night, rely on bats for pollination. Scientists have proven that "bats that feed on flowers at night ... apparently play the same ecological role that hummingbirds play during the day."

This phenomenon has been studied in detail in Trinidad, Java, India, Costa Rica, and many other places; observations revealed the following facts:

1. The smell of most flowers pollinated by bats is very unpleasant for humans. This applies primarily to the flowers of Oroxylon indicum, baobab, as well as some types of kigelia, parkia, durian, etc.

2. Bats come in different sizes - from animals smaller than a human palm to giants with a wingspan of more than a meter. The little ones, launching long red tongues into the nectar, either soar above the flower, or wrap their wings around it. Big bats stick their muzzles into the flower and begin to quickly lick the juice, but the branch sinks under their weight, and they fly up into the air.

3. Bat-attracting flowers belong almost exclusively to three families: Bignonia (Bignoniacea), Mulberry Cotton (Bombacaceae) and Mimosa (Leguminoseae). The exception is Phagrea from the Loganiaceae family and the giant cereus.

Rat "tree"

The climbing pandanus (Freycinetia arborea), found in the Pacific Islands, is not a tree, but a liana, although if its many trailing roots can find suitable support, it stands so straight that it looks like a tree. Otto Degener wrote about him:

“Freycinetia is quite widespread in the forests of the Hawaiian Islands, especially in the foothills. It is not found anywhere else, although more than thirty related species have been found on the islands located to the southwest and east.

The road from Hilo to Kilauea Crater is teeming with yeye ( Hawaiian name for climbing pandanus. - Approx. transl.), which are especially conspicuous in summer when they bloom. Some of these plants climb the trees, reaching the very tops - the main stem wraps around the trunk with thin aerial roots, and the branches, bending, get out into the sun. Other individuals crawl along the ground, forming impenetrable plexuses.

The woody yellow stems of the yeye are 2-3 cm in diameter and are surrounded by scars left from fallen leaves. They produce many long adventitious aerial roots of almost the same thickness along the entire length, which not only supply the plant with nutrients, but also enable it to cling to a support. The stems branch every meter and a half, ending in bunches of thin glossy green leaves. The leaves are pointed and covered with spines along the edges and along the underside of the main vein ...

The method developed by the yeye to ensure cross-pollination is so unusual that it is worth talking about in more detail.

During the flowering period, bracts consisting of a dozen orange-red leaves develop at the ends of some yeye branches. They are fleshy and sweet at the base. Three bright plumes stick out inside the bract. Each sultan consists of hundreds of small inflorescences, which are six combined flowers, of which only tightly fused pistils have survived. On other individuals, the same bright stipules develop, also with sultans. But these sultans do not carry pistils, but stamens in which pollen develops. Thus, the yeye, dividing into male and female individuals, completely secured themselves from the possibility of self-pollination ...

Inspection of the flowering branches of these individuals shows that they are most often damaged - most of the fragrant, brightly colored fleshy leaves of the bract disappear without a trace. They are eaten by rats, which, in search of food, move from one flowering branch to another. Eating fleshy bracts, rodents stain their whiskers and hair with pollen, which then falls on the stigmas of females in the same way. Yeye is the only plant in the Hawaiian Islands (and one of the few in the world) that is pollinated by mammals. Some of its relatives are pollinated by flying foxes - fruit-eating bats that find these fleshy bracts tasty enough.

Ant trees

Some tropical trees are attacked by ants. This phenomenon is completely unknown in the temperate zone, where the ants are just harmless bugs that climb into the sugar bowl.

Everywhere in the rain forests there are countless ants of the most varied sizes and with the most varied habits - ferocious and gluttonous, ready to bite, sting, or in some other way destroy their enemies. They prefer to settle in trees and for this purpose they choose certain species in the diverse plant world. Almost all of their chosen ones are united by the common name "ant trees". A study of the relationship between tropical ants and trees has shown that their union is beneficial for both parties ( For lack of space, we will not touch here on the part played by ants in the pollination of some flowers or in the dispersal of seeds, nor on the ways in which some flowers protect their pollen from ants.).

Trees shelter and often feed ants. In some cases, trees secrete lumps of nutrients, and ants eat them; in others, the ants feed on tiny insects, such as aphids, that live off the tree. In forests that are subject to periodic flooding, trees are especially important for ants, as they save their homes from flooding.

Trees undoubtedly extract some nutrients from the debris that accumulates in ant nests - very often an aerial root grows into such a nest. In addition, ants protect the tree from all kinds of enemies - caterpillars, larvae, grinder bugs, other ants (leaf cutters) and even from people.

Regarding the latter, Darwin wrote:

“The protection of the foliage is provided ... by the presence of entire armies of painfully stinging ants, whose tiny size only makes them more formidable.

Belt, in his book The Naturalist in Nicaragua, gives a description and drawings of the leaves of one of the plants of the Melastomae family with swollen petioles and indicates that, in addition to small ants living on these plants in large numbers, he noticed dark-colored Aphides several times. In his opinion, these small, painfully stinging ants bring great benefits to plants, as they protect them from enemies that eat leaves - from caterpillars, slugs and even herbivorous mammals, and most importantly, from the ubiquitous sauba, that is, leaf-cutting ants, which, according to he said, they are very afraid of their small relatives.

This union of trees and ants is carried out in three ways:

1. In some ant trees, the twigs are hollow, or their core is so soft that the ants, arranging a nest, easily remove it. Ants look for a hole or a soft spot at the base of such a branch, if necessary, gnaw their way and settle inside the branch, often expanding both the inlet and the branch itself. Some trees even seem to prepare entrances for ants in advance. On thorny trees, ants sometimes settle inside the thorns.

2. Other ant trees place their tenants inside the leaves. This is done in two ways. Usually ants find or gnaw the entrance at the base of the leaf blade, where it connects to the petiole; they climb inside, pushing the top and bottom covers of the sheet apart, like two pages glued together - here's your nest. Botanists say that the leaf "invaginates", that is, it simply expands, like a paper bag, if you blow into it.

The second way of using leaves, which is observed much less often, is that ants bend the edges of the leaf, glue them together and settle inside.

3. And finally, there are ant trees that do not themselves provide dwellings for ants, but instead ants settle in those epiphytes and vines that they support. When you stumble upon an ant tree in the jungle, you usually don't waste time checking whether the ant streams are coming from the leaves of the tree itself or from its epiphyte.

Ants in the branches

Spruce detailed his introduction to ant trees in the Amazon:

“Ant nests in the thickening of the branches are in most cases on low trees with soft wood, especially at the base of the branches. In these cases, you will almost certainly find ant nests either at each node or on the tops of the shoots. These anthills are an expanded cavity inside the branch, and communication between them is sometimes carried out along the passages laid inside the branch, but in the overwhelming majority of cases - through covered passages built outside.

Cordia gerascantha almost always has pouches at the point of branching, in which very vicious ants live - the Brazilians call them "takhi". C. nodosa is usually inhabited by small fire ants, but sometimes takhi. Perhaps the fire ants were the first inhabitants in all cases, and the takhs are pushing them out.

All tree-like plants of the buckwheat family (Polygonaceae), Spruce continues, are affected by ants:

“The entire core of each plant, from the roots to the apical shoot, is almost completely scraped out by these insects. Ants settle in a young stem of a tree or shrub, and as it grows, releasing branch after branch, they make their moves through all its branches. These ants all seem to belong to the same genus, and their bite is extremely painful. In Brazil they are called "tahi" or "tasiba" and in Peru "tangarana", and in both these countries the same name is commonly used for both the ants and the tree in which they live.

In Triplaris surinamensis, a fast-growing tree throughout the Amazon, and in T. schomburgkiana, a small tree in the upper Orinoco and Ca-siquiare, the thin, long tube-like branches are almost always perforated with many tiny holes that can be found in the stipule of almost every leaf. This is the gate, from which, at a signal from the sentinels constantly walking along the trunk, a formidable garrison is ready to appear at any second - as a carefree traveler can easily see from his own experience, if, seduced by the smooth bark of a takhi tree, he decides to lean against it.

Almost all tree ants, even those that sometimes descend to the ground during the dry season and build summer anthills there, always keep the above-mentioned passages and bags as their permanent homes, and some species of ants do not leave trees at all all year round. Perhaps the same applies to ants who build anthills on a branch of foreign materials. Apparently, some ants always live in their aerial dwellings, and the inhabitants of the tokoki (see p. 211) do not leave their tree even where they are not threatened by any floods.

Ant trees exist throughout the tropics. Among the most famous is the cecropia (Cecropia peltata) of tropical America, which is called the "trumpet tree" because the Waupa Indians make their wind pipes from its hollow stems. Ferocious Azteca ants often live inside its stems, which, as soon as the tree is swayed, run out and. pounce on the daredevil who disturbed their peace. These ants protect cecropia from leaf cutters. The internodes of the stem are hollow, but they do not communicate directly with the outside air. However, near the apex of the internode, the wall becomes thinner. A fertilized female gnaws through it and hatches her offspring inside the stem. The base of the petiole is swollen, outgrowths are formed on its inner side, which the ants feed on. As the outgrowths are eaten, new ones appear. A similar phenomenon is observed in several related species. Undoubtedly, this is a form of mutual accommodation, as evidenced by the following interesting fact: the stem of one species, which is never "ant-like", is covered with a wax coating that prevents leaf cutters from climbing it. In these plants, the walls of the internodes do not become thinner and edible outgrowths do not appear.

In some acacias, the stipules are replaced by large spines swollen at the base. In Acacia sphaerocephala in Central America, ants enter these spines, clean them of internal tissues and settle there. According to J. Willis, the tree provides them with food: "Additional nectaries are found on the petioles, and edible outgrowths are found on the tips of the leaves." Willis adds that any attempt to damage the tree in any way causes the ants to pour out in masses.

The old riddle of which came first, the chicken or the egg, is repeated in the example of the Kenyan black gall locust (A. propanolobium), also known as the whistling thorn. The branches of this small shrub-like tree are covered with straight white thorns up to 8 cm long. Large galls form on these thorns. At first, they are soft and greenish-purple, and then harden, blacken, and ants settle in them. Dale and Greenway report: “The galls at the base of the thorns... are said to be due to ants that gnaw them from the inside. When the wind hits the holes of the Gauls, a whistle is heard, which is why the name "whistling thorn" arose. J. Salt, who examined the galls on many acacias, found no evidence that their formation was stimulated by ants; the plant forms swollen bases, and the ants use them.

Ant tree in Ceylon and southern India is Humboldtia laurifolia from the legume family. In him, cavities appear only in flowering shoots, and ants settle in them; the structure of non-flowering shoots is normal.

Considering the South American species of Duroia from the madder family, Willis notes that two of them - D. petiolaris and D. hlrsuta - have swollen stems right under the inflorescence, and ants can enter the cavity through the cracks that appear. A third species, D. saccifera, has anthills on leaves. The entrance, located on the upper side, is protected from rain by a small valve.

Corner describes the different types of macaranga (locally called mahang), the main ant tree of Malaya:

“Their leaves are hollow, and ants live inside. They gnaw their way out in the shoot between the leaves, and in their dark galleries they keep a mass of aphids, like herds of blind cows. The aphids suck the sugary sap of the shoot, and their bodies secrete a sweetish liquid that the ants eat. In addition, the plant produces so-called "edible outgrowths", which are tiny white balls (1 mm in diameter), which consist of oily tissue - it also serves as food for ants ... In any case, ants are protected from rain ... If you cut escape, they run out and bite ... Ants penetrate young plants - winged females gnaw their way inside the shoot. They settle in plants that have not reached even half a meter in height, while the internodes are swollen and look like sausages. The voids in the shoots arise as a result of the drying of the wide core between the nodes, like in bamboos, and the ants turn individual voids into galleries, gnawing through the partitions in the nodes.

J. Baker, who studied ants on macaranga trees, discovered that it was possible to cause a war by bringing two trees inhabited by ants into contact. Apparently, the ants of each tree recognize each other by the specific smell of the nest.

Ants inside leaves

Richard Spruce points out that spreading tissues and integuments, which form suitable sites for the emergence of ant colonies, are found mainly in some South American melastomas. The most interesting of these is the tokoka, whose numerous species and varieties grow in abundance along the banks of the Amazon. They are found mainly in those parts of the forest that are flooded during floods of rivers and lakes or during rains. Describing bags formed on leaves, he says:

“The leaves of most species have only three veins; some have five or even seven; however, the first pair of veins always departs from the main one about 2.5 cm from the base of the leaf, and the bag occupies precisely this part of it - from the first pair of lateral veins down.

This is where the ants settle in. Spruce reported that he found only one species - Tososa planifolia - without such swellings on the leaves, and trees of this species, as he noticed, grow so close to rivers that they are undoubtedly under water for several months of the year. These trees, in his opinion, “cannot serve as a permanent residence for ants, and therefore the temporary appearance of the latter would not leave any imprint on them, even if instinct did not force the ants to avoid these trees altogether. Trees of other species of Tosos, growing so far from the shore that their tops remain above the water even at the moment of its highest rise, and therefore suitable for the constant habitation of ants, always have leaves with bags and are not free from them in any of the seasons. . I know this from bitter experience, for I have had many skirmishes with these belligerent bugs when I damaged their dwellings while collecting specimens.

Bag-like dwellings of ants also exist in the leaves of plants of other families.

Ant nests on epiphytes and vines

The most notable of the epiphytes that harbor ants high among the branches of tropical trees are the eighteen species of Myrmecodia, which are found everywhere from New Guinea to Malaya and the far north of Australia. They often coexist with another epiphyte, Hydnophytum, a genus of forty species. Both of these genera are included in the madder family. Merril reports that some of them are found in lowlands and even in mangroves, while others grow in primary forests at high altitudes. He continues:

“The bases of these trees, sometimes armed with short thorns, are very enlarged, and this enlarged part is penetrated by wide tunnels into which small holes lead; inside the strongly swollen bases of these plants myriads of small black ants find shelter. From the top of the tuberous, tunnelled base rises stems, sometimes thick and unbranched, sometimes thin and very branched; small white flowers and small fleshy fruits develop in the axils of the leaves.

“Perhaps the most peculiar adaptation of the leaves is noted in groups such as Hoya, Dlschidia and Conchophyllum. These are all creepers with abundant milky juice belonging to the family Asclepmdaceae. Some of them hang on trees as epiphytes or semi-epiphytes, but in Conchophyllum and some species of Noua, the thin stems lie close to the trunk or branches of the depewa, and the round leaves, arranged in two rows along the stem, are arched and their edges are closely pressed to the bark. Roots grow from their sinuses, often completely covering a piece of bark under the leaf - these roots hold the plant in place and, in addition, absorb the moisture and nutrients it needs; under each such leaf in a finished dwelling, colonies of small ants live.

Dischidia rafflesiana, a peculiar pitcher plant of Southeast Asia, provides shelter to ants. Some of its leaves are iloski, others are swollen and reminiscent of jugs. Willis describes them as follows:

“Each leaf is a jug with an edge turned inside, about 10 cm deep. An adventitious root grows into it, developing near on the stem or on the petiole. The jug ... usually contains various debris caused by ants nesting there. Rainwater accumulates in most pitchers ... The inner surface is covered with a wax coating, so that the pitcher itself cannot absorb water and it is sucked up by the roots.

The study of the development of the pitcher shows that it is a leaf, the lower part of which is invaginated.

Introduction

Each organism, including plants, has the ability to reproduce its own kind, which ensures the existence of a species in space and time, sometimes for a very long time. With the loss of the ability to reproduce, species die out, which has repeatedly occurred in the course of the evolution of the plant world.

Plants reproduce both sexually and asexually. Sexual reproduction consists in the fact that two cells, called gametes, merge, and, in addition to the fusion of protoplasms, the fusion of nuclei is necessary for sexual reproduction. Thus, the fusion of nuclei is the most important stage of the sexual process, otherwise called fertilization.

Pollination plays a major role in plant reproduction. Pollination is the process of transferring pollen grains from the stamens to the stigma of the pistil. This process can occur with the help of various factors, both biotic and abiotic.

In this paper, we will consider the definition of pollination, its types. Cross-pollination and morphological adaptations of plants to it will be considered and studied in more detail.

The purpose of the course work is to consider and study the morphological adaptations of angiosperms to cross-pollination.

1. Review the definition of pollination.

2. Study the types of pollination.

3. Consider cross-pollination in more detail.

4. Consider the morphological adaptations of plants to cross-pollination.

Chapter 1. Pollination as a way of reproduction of angiosperms

1.1 Pollination as a mode of reproduction

Pollination is the process of transferring pollen grains from the stamen to the stigma of the pistil. This process can occur with the help of various factors, both biotic and abiotic.

In classical works on the ecology of pollination, two concepts are distinguished: autogamy, or self-pollination, in which pollen from the same flower falls on the stigma. If the flowers are on the same plant, pollination is called heitenogamy, if on different plants - xenogamy.

There are no sharp differences between these variants of pollination. Geitenogamy is genetically equivalent to autogamy, but requires the participation of certain pollinators, depending on the structure of the flower. In this respect, it is similar to xenogamy. In turn, xenogamy can be identical to autogamy if the pollinated plants belong to the same clone, i.e. arose as a result of vegetative reproduction of one maternal individual.

In this regard, pollination is reduced to two types: autogamy, or self-pollination, and cross-pollination.

1.2 Autogamy, or self-pollination

This type of pollination is characteristic only of bisexual flowers. Autogamy can be random or regular.

Random autogamy is not uncommon. It is difficult to enumerate all the factors contributing to its implementation. It is only important that there is a physiological compatibility of pollen grains and the stigma of the pistil.

Regular autogamy can be gravitational if the pollen grain, due to its gravity, falls on the stigma from the anther located above it. The carriers of pollen grains inside the flower can be raindrops, small insects - thrips, which settle in the flower. The most common is contact autogamy, in which the opening anther comes into contact with the stigma of the pistil (hoof). Autogamy is closely related to the time factor and environmental conditions. In Dortmann's lobelia (Lobelia dortmanna) (see Fig. 1), it occurs before flowering, although it develops chasmogamous flowers with external attributes to attract pollinators.

Figure 1 - Lobelia Dortmann (Lobelia dortmanna)

In the small mousetail (Myosurus minimus L.) (see Fig. 2), self-pollination occurs in the first half of flowering, later it is impossible. In flowers in which self-pollination occurs before flowering, certain elements are often reduced. The extreme degree of such reduction is represented by cleistogamous flowers.

Figure 2 - Small mousetail (Myosurus minimus L.)

In oxalis (Oxalis) (see Fig. 3), about a month after flowering, when seeds are already developing in their ovaries, small (up to 3 mm) cleistogamous flowers appear with perianth in the form of small scales. An important feature of the cleistogamous flower is that anthers never open in it, but pollen tubes grow from the pollen grains in them, piercing the anther wall and growing towards the stigma, often bending at the same time. The stigma is often located at the top of the ovary, there is no style.

Figure 3 - Common Oxalis (Oxalisacetosella)

Often, cleistogamy is optional and appears in plants only under certain weather conditions. This is found in plantain chastukha (Alismaplantago-aguatica), sundew, feather grass, in which cleistogamous flowers develop during soil drought and low temperatures. In wheat, chasmogamous flowers are formed in warm, humid weather, and cleistogamous in dry and hot weather.

In most cases, cleistogamy occurs in unstable habitat conditions unfavorable for cross-pollination.

1.3 Cross-pollination

Cross-pollination, or allogamy, is a type of pollination in angiosperms in which pollen from the androecium of one flower is transferred to the stigma of the pistil of another flower.

There are two forms of cross-pollination:

1. Geitonogamy (adjacent pollination) - pollination in which pollen from a flower of one plant is transferred to the stigma of the pistil of another flower on the same plant;

2. Xenogamy - cross-pollination, in which pollen from the flower of one plant is transferred to the stigma of the pistil in the flower of another plant.

With the help of cross-pollination, genes are exchanged, which maintains a high level of heterozygosity in the population, determines the unity and integrity of the species. With cross-pollination, the possibilities of recombination of genetic material increase, more diverse genotypes of offspring are formed as a result of the combination of hereditarily diverse gametes, therefore, more viable than with self-pollination, offspring with a greater amplitude of variability and adaptability to various conditions of existence. Thus, cross-pollination is biologically more beneficial than self-pollination, therefore it was fixed by natural selection and became dominant in the plant world. Cross-pollination exists in over 90% of plant species.

Cross-pollination can be carried out both biotically (with the help of living organisms) and abiotic (through air or water currents).

Cross-pollination is carried out in the following ways:

a) Anemophily (pollination by wind)

b) Hydrophilia (pollination with water)

c) Ornithophilia (pollination by birds)

d) Chiropterophilia (pollination by bats)

e) Entomophily (pollination by insects)

Chapter 2. Morphological adaptations of plants to cross-pollination.

2.1 Anemophily or wind pollination

Wind-pollinated plants often grow in large clusters, for example, hazel thickets, birch groves. A person sows rye and corn on hundreds of hectares, and sometimes thousands of hectares of land.

In summer, flower pollen rises above the rye field in a cloud. Wind pollinated plants produce a lot of pollen. Part of the dry and light pollen necessarily falls on the stigmas. But most of the pollen is wasted without pollinating the flowers. The same can be seen in spring when hazel, birch and other wind-pollinated trees and shrubs bloom. Poplar, alder, rye, corn and other plants with inconspicuous flowers are pollinated by the wind.

Most wind-pollinated trees bloom in early spring, before the leaves appear. This ensures that the pollen gets to the stigma better.

Plants pollinated by the wind do not have bright and fragrant flowers. Inconspicuous, usually small flowers, anthers on long hanging threads, very small, light, dry pollen - all these are adaptations for wind pollination.

2.2 Hydrophilia or water pollination

Hydrophilia is of more ancient origin, since it is believed that the first higher plants appeared in water. However, most aquatic plants are air-pollinated, just like their terrestrial relatives. Plants such as Nymphaea, Alisma, and Hottonia are entomophilous, Potamogeton or Myriophyllum anemophilous, and Lobelia dortman self-pollinating. But for pollination of some aquatic plants, an aquatic environment is necessary.

Hydrophilia can occur both on the surface of the water (ephidrophilia) and in the water (hyphydrophilia). These two types of pollination represent a further development of anemophily or entomophily. Many small, self-pollinating land plants can flower while submerged in water; at the same time, the self-pollination mechanism functions, usually enclosed in an air sac inside the flower. Cleistogamous flowers represent the highest stage of such development.

Ephydrophily is a unique type of abiotic pollination, since in this case pollination occurs in a two-dimensional environment. Compared to the three-dimensional environment in which anemophily or hyphydrophilia occurs, this type of pollination provides a greater economy of pollen. In epihydrophilia, pollen is released from the anthers in the water and floats to the surface where the stigmas (Ruppia, Callitriche autumnalis) are found. Pollen grains quickly spread over the surface film of water. This is easy to see when watching Ruppia bloom: small yellow drops appear on the surface of the water and spread quickly, like drops of fat; this is facilitated by an oily layer covering the shell of the pollen grain.

An interesting case of pollination in Vallisneria is widely known, in which, instead of individual pollen grains, the entire male flower comes to the surface of the water; therefore, the pollen does not even touch the surface of the water. Small funnels form around the emerging female flowers; male flowers floating nearby slide from the edge of such a funnel to its center; while the anthers touch the stigmas. Due to this efficient method of pollination, the number of pollen grains in male flowers is greatly reduced. Vallisneria-type mechanisms are also found in various representatives of the Hydrocharitaceae, sometimes, as in Hydrilla, along with exploding anthers. A similar mechanism of pollination is also observed in Lemna trisulca, only the entire plant rises to the surface of the water; and in Elodea, with a similar mechanism of pollination, staminate flowers are brought to the surface of the water, which are partly attached and partly free-floating.

Hyphydrophilia has been described in a very few plants, such as Najas, Halophila, Callitriche hamulata and Ceratophyllum. So far, they are treated simply as separate cases, since there is probably little in common between them, except for the extreme reduction of the exine. In Najas, the slowly descending pollen grains are "caught" by the stigma.

The dispersing pollen unit in Zostera is 2500 µm long and much more like a pollen tube than a pollen grain. Being very mobile, it quickly wraps itself around any object encountered on the way, for example, around a stigma. However, this reaction is completely passive. The pollen grain morphology of Zostera can be seen as an extreme case of a trend that seems to be shared by other hyphydrophilic plants: a rapidly growing pollen tube ensures that the pollen grains spread rapidly. In Cymodoceae, even more elongated pollen grains (5000-6000 µm) have been described.

2.3 Ornithophily or bird pollination

Since the birds fly well and the surface of their body is not smooth, they have good external conditions for pollination. No one is surprised that insects get food from flowers, but the corresponding actions of birds cause great surprise and reflection on how they got the “idea” to use the nectar of flowers. One of the ideas put forward was the idea that pollination arose as a result of the eating of flowers by birds, and that food may have been primarily fruits. It has also been suggested that woodpeckers or sap-eating woodpeckers (Sphyrapicus) sometimes change their diet and switch to juices flowing from hollows (some of them also peck fruits; Dendrocopus analis - fruits of Cassia grandis). A third group of "explanations" suggests that the birds pursued insects in flowers and happened to find nectar or pierce succulent tissues; or at first they drank water collected in flowers to quench their thirst, since in tropical forests water is difficult to access for animals living in the crowns of trees. The fact that hummingbirds originally pursued insects in flowers can be seen even today. The rapid absorption of nectar makes it difficult to identify it in the stomach of birds, while indigestible remains of insects are easily recognized. However, in the ornithological literature there is a large amount of data indicating that the digestive systems of birds are filled with nectar. Extraction of nectar by piercing the base of the corolla is further evidence that all this is done for the sake of extracting nectar. Insects cannot obtain nectar in this way. Some hummingbirds are addicted to piercing flowers, like some hymenoptera. None of the insects get nectar from the closed flowers of the Loranthaceae from Java, which open only when struck by nectar-seeking birds. The fact that birds visit flowers can be confirmed even on very old museum preparations by the presence of pollen grains in feathers or on the beak.

Hummingbirds need a large amount of energy, especially when hovering. It is precisely such a large expenditure of energy for soaring and flying that can explain the small size of these birds. After a period of fasting, nutrient stores can be severely reduced despite low metabolic rates during sleep.

In pollinators with different energy budgets, the efficiency of nectar uptake and its metabolism are different. The presence of flowers with a large amount of nectar is a signal forcing hummingbirds to seize and defend territories. One could refer to the migration of hummingbirds to those places where these flowers are numerous, especially during the breeding season.

From a pollination point of view, it didn't really matter whether the birds visited the flowers for nectar or to catch insects, until these visits became regular. Whether nectar or insect is the reason for the visit is a problem of adaptation, not function. In Java, Zosterops visits the non-ornithophilous Elaeocarpus ganitrus to collect mites, which are abundant in flowers.

There is no doubt that birds perched on flowers for all the reasons mentioned above. Even if, from the gardener's point of view, the flowers were damaged, they were successfully pollinated. Damage to the flower itself is of little consequence as long as the pistil is not damaged. After all, explosive flowers are also destroyed themselves.

Other similar occasional flower visits by dystrophic birds have recently been recorded in birds migrating to England from more southerly areas. Campbell observed various birds in England chasing insects in flowers while landing very small amounts of pollen.

From these examples of dystropic visits to flowers, it appears that there is a gradual transition through certain allotropic birds with a mixed diet, in which nectar is one of the ingredients, to eutropic ones, as a result of which true ornithophily is established.

For a long time, observations were made of visits to hummingbird flowers. Ornithophilia as a scientifically recognized phenomenon was established by Treleese at the end of the last century, and Johaw, Freese, and chiefly Werth studied it in more detail. However, it was only when Porsche in the 1920s collected a huge amount of data and made convincing conclusions about the now well-known phenomena that ornithophilia was unanimously recognized, even if its origin is still a matter of controversy.

The habit of collecting nectar is obviously polyphyletic, having arisen in different groups of birds in different regions. The best-known example of high adaptation are the hummingbirds (Trochilidae) of North and South America. Hummingbirds were probably originally insectivorous, but later switched to nectar; their chicks still eat insects in addition to nectar. The same is observed in insects.

Another American group of more or less eutropic flower-eating birds are the much less important sugar-eating birds (Coerebidae). In the Old World, other families have developed the same characteristics as hummingbirds, even if their adaptations are usually less significant. In Africa and Asia live nectaries (Nectarinidae), in Hawaii - Hawaiian flower girls (Drepanididae), closely related to local lobelia, in the Indo-Australian region - honey badgers (Meliphagidae) and brush-tongued honey parrots or small loris parrots (Trichoglossidaei).

Less specialized pollinators of flowers with a mixed diet (allotropic pollinators) are also active, but as pollinators to a much lesser extent, especially in simpler bird-pollinated flowers (Bombax, Spathodea); this shows that flowers and their birds may have evolved in parallel, influencing each other. Pollinators are found in many other families, such as some tropical nightingales (Pycnonotidae), starlings (Sturnidae), orioles (Oriolidae), and even among tropical woodpeckers (Picidae), where the fringe at the tip of the tongue is the first sign of morphological adaptation.

The flower-suckers (Dicaeidae) visit a variety of flowers, while showing a curious "specialization" to one group of plants, namely the tropical Loranthoideae, in which they not only visit ornithophilous flowers, but also adapt to the digestion of fruits and the dispersal of seeds. The oldest observations of bird pollination in the New World were made by Catesby and Ramphius in the Old World.

The areas in which any type of ornithophilia is found practically cover the American continent and Australia and further tropical Asia and the deserts of South Africa. According to Werth, Israel is the northern limit of this area, with Cinnyris visiting the flowers of the red Loranthus as well. recently Galil reported the abundance of these birds on plants growing in gardens.

In the mountains of Central and South America, the number of ornithophilous species is unusually high. If bees are present in the high elevations of Mexico, they are just as effective as pollinators as birds, except that birds are more efficient under adverse conditions. However, Bombus species are not very sensitive to climate. Their presence can completely change the picture, as shown by van Leeuwen. Stevens points out similar results of Rhododendron pollination in the mountains of Papua.

Obviously, in Australia and New Zealand, the number of eutropic pollinating insects is also low, and the function of higher bees performed by them on other continents is taken over by birds.

Individual cases of feeding on flowers in various groups of birds, their geographical distribution and single cases of ornithophilous types of flowers in many groups of plants - all this indicates that ornithophily arose relatively recently.

The ability to soar, well developed in hummingbirds, is rare in other groups of birds; it is observed, for example, in the honey-eating Acanthorhynchus, and is poorly developed in the Asiatic Arachnothera. Some birds can soar in strong headwinds.

The brightness of the plumage, leading to a significant similarity in the color of birds and flowers, may seem rather strange. We have reason to consider this fact from the point of view of protective coloration. Van der Piel observed that a highly conspicuous flock of red-green Loriculus (brightly colored hanging parrots) becomes invisible when landing on a flowering Erythrina. Obviously, these animals are largely vulnerable when they are immobile while eating.

Grant argued that "persistence" to flowers is underdeveloped in birds and that their feeding habits are too complex. Information about the evolution of constancy to flowers is different for different authors. Snow and Snow suggest a very close relationship - monotropic, in our current terminology - between Passijloramixta and Ensiferaensifera. Obviously, the relationship between different species of hummingbirds and the plants that provide them with food varies greatly, ranging from strict territoriality to a highly inefficient strategy of successive visits, when birds use any available source of nectar. It is also necessary to take into account the possibility of learning in birds. If diversity is allowed, then impermanence may be due to the lack of a proper distinction between deceit and preferable constancy. Birds feed on any kind of food, so it is natural that if there is a profuse bloom and a large amount of nectar is available, the apparent preference of the birds in this case will simply be a matter of statistics and will not depend on the food itself. If there is no such flowering, then they can fly from one species to another or even use other food. Any observed consistency will be impressive even though flower tube length, beak length, nectar composition, etc. may play a role in flower selection. In emergencies, birds eat flowers. Johow noticed in Chile that hummingbirds can even switch to European fruit trees or Citrus species. Hemitropic birds switch to fruits more frequently. In the tropics, birds especially prefer fresh flowering trees. The ecological significance of this, of course, is not absolute, but relative and can be of selective significance.

The phylogenetic development of tropical plant species and the most highly developed groups of pollinators has led to a distinct and easily recognizable bird pollination syndrome that excludes other pollinators. Any random combinations in this case are impossible. The mutual dependence is well seen in the example of the Hawaiian flower girls Drepanididae and the flowers pollinated by them, which, when the birds were exterminated, became autogamous.

For the differential diagnosis of classes of ornithophilous flowers and flowers pollinated by diurnal Lepidoptera. The differences are rather indistinct, especially in American plants.

Some bird-pollinated flowers are brush-like (Eucalyptus, heads of Proteaceae and Compositae), others are slanted-mouthed (Epiphyllum) or tubular (Fuchsiafulgens). Some moths are typically ornithophilous.

The fact that various types of flowers are ornithophilous indicates a recent development of ornithophily, which is on top of the previous ecomorphological organizations that determine the types of structure, etc., but leading to a secondary convergence of the style. Isolated instances of resemblance between unrelated flowers, regarded by some morphologists as a mysterious "repeated pair" and by others as orthogenetic, probably represent a parallel adaptation in the field of pollination.

The effectiveness of this syndrome is shown by the fact that typical bird-pollinated flowers growing in European gardens attract the attention of short-beaked, unadapted dystrophic birds, and also by the fact that flower-pollinating birds immediately recognize and then try to use the flowers of introduced bird-pollinated plants. Flower size is not included in the syndrome. Many flowers pollinated by birds are relatively small. The flowers pollinated by birds are usually deep, and do not belong to any one particular class, but the most characteristic of them are brush-like and tubular.

Sensitivity to different regions of the spectrum in different species of birds varies. In one species of hummingbird (Huth), a shift to the short-wavelength region of the spectrum was found compared to the human visible spectrum.

In Columneaflorida birds are attracted by red spots on the leaves, while the flowers themselves are hidden. Since this spot does not reproduce the shape of the flower, a high degree of mental integration can be assumed in birds pollinating Columneaflorida.

Flowers with a bright, contrasting color should include flowers in the species Aloe, Strelitzia and many bromeliads.

The transition to ornithophily is mostly recent, but in some groups the ornithophily appears to be older. Porsche identified a suprageneric group in Cactaceae (Andine Loxantocerei), in which, apparently, ornithophily in the tribe was fixed. Snow and Snow give other examples of the coevolution of ornithophilous flowers and their pollinators.

Among Euphorbiaceae with dense cyathium, Poinsettia has large glands and red bracts that attract hummingbirds. The genus Pedilanthus is characterized by an even higher specialization, which appeared from the beginning of the Tertiary period, and in this genus the glands are in spurs, the flowers are erect and zygomorphic.

Even among orchids, which have excellent pollinators - bees, some species have switched to ornithophily in an endless search for new pollinators typical of this family. In the South African genus Disa, some species have probably become ornithophilous. Therefore, the flowers of this genus pollinated by butterflies are already red, with a spur and with a reduced upper lip. The same occurs in Cattleyaaurantiaca and in some species of Dendrobium in the mountains of New Guinea. Birds visiting the flowers of Elleanthuscapitatus and Masdevalliarosea were observed by Dodson.

2.4 Chiropterophilia or bat pollination

Like birds, bats' body surfaces are not smooth, so they have a great ability to retain pollen. They also fly fast and can travel long distances. Pollen from plants located at a distance of 30 km was found in the faeces of bats. Therefore, it is not surprising that bats are good pollinators.

The first conscious observations of bats visiting flowers were made by Bürk in the Biitenzorg (now Bogor) Botanical Garden. He observed that frugivorous bats (probably Cynopterus) visited the inflorescences of Freycinetia insignis, a plant now known to be entirely chiropterophilic, in contrast to its closely related ornithophilous species.

Later, some authors described other cases, and the example of Kigelia (Kigelia) has become a classic. As early as 1922, Porsche was expressing certain considerations regarding chiropterophilia, noting its characteristic features and predicting many possible examples.

Thanks to the work of van der Pijl in Java, Vogel in South America, Jaeger, and Baker and Harris in Africa, bat pollination has now been identified in many plant families. It turned out that some plants, previously considered ornithophilous, are pollinated by bats (for example, species of Marcgravia).

Bats are generally insectivorous, but herbivorous bats independently appeared in both the Old and New Worlds. Perhaps the evolution went through frugivorousness to the use of flowers for food. Fruit-eating bats are known in two suborders inhabiting different continents, while African Pteropinae are characterized by a mixed diet. Like hummingbirds, nectar feeding is thought to have evolved from hunting insects in flowers.

Hart's observations in Trinidad in 1897 on Bauhiniamegalandra and Eperuafalcata are often mentioned in the literature, confusingly with incorrect conclusions.

Relationships between fruit and flower feeding Megalochiroptera are still partly dystropic. In Java, Cynopterus has been found to eat Durio flowers and parts of Parkia inflorescences.

In eastern Indonesia and Australia, Cynopterus and Pteropus destroy many Eucalyptus flowers, indicating hitherto unbalanced pollination conditions.

Macroglossinae are more adapted to the flower than even hummingbirds. In the stomachs of these animals caught in Java, only nectar and pollen were found, the latter in such large quantities that its accidental use is completely excluded. Obviously, pollen is in this case a source of protein, which their ancestors received from fruit juice. In the Glossophaginae, the use of pollen, although found, seems to be less significant.

Howell is of the opinion that Leptonycteris satisfies its protein requirements from pollen, and the protein in the pollen is not only of high quality, but also in sufficient quantity. She also states that the chemical composition of the pollen of flowers pollinated by bats is adapted to the use of it by these animals and differs from the composition of the pollen of related species that are pollinated by other animals. This can be seen as a floral part of the co-evolution of the chiropterophilia syndrome. Until now, the issue of African fruit-eating bats that swallow pollen has not been clarified.

The class of flowers pollinated by bats has been found to have an early side branch of evolution, forming its own subclass, for which the only pollinator is Pteropineae. In these flowers, solid food (with a characteristic odor) is represented only by specialized structures. There is neither nectar nor large masses of pollen. Freycinetiainsignis has a sweet bract, the Bassia species is a very sweet and easily separated corolla. Perhaps another species of Sapotaceae, namely the African Dumoriaheckelii, also belongs to this subclass.

Possibility of bat pollination of white-flowered tree strelitzia (Strelitzianicolai) in the eastern region of Cape Cod needs to be investigated.

Pteropinae extends beyond the latitude of Canton. The Eastern Pacific border runs in a sharp ridge through the Caroline Islands to Fiji. Macroglossinae are known to have visited flowers in Northern Australia (introduced by Agave), but the native Adansoniagregorii has all the characteristics of chiropterophilia; therefore, chiropterophilia must also exist on this continent.

Knowing the characteristics of pollination by bats can help in solving the mysteries of the origin of plants. The chiropterophilic flower of Musafehi is evidence that the species was introduced to Hawaii, where there are no bats. Chiropterophilia could have taken place in his homeland, in New Caledonia, from where, as several botanists have established, he comes from.

Similarly, some species of the New World Glossophaginae have a longer snout and tongue than their insectivorous relatives. Musonycterisharrisonii has a tongue length of 76 mm and a body length of 80 mm. Vogel believes that the hairs of the Glossophaga's coat are particularly well adapted to carrying pollen, since they are equipped with scales similar in size to those on the hairs that cover the belly of a bumblebee.

The physiology of Megachiroptera's sense organs deviates from what we normally see in bats. The eyes are large, sometimes with a folded retina (allowing rapid accommodation), with many rods but no cones (causing color blindness). In night photographs, fruit-eating Epomopsfranqueti show huge eyes, almost the same as those of a lemur. Smell perception probably plays a more important role than usual (large nasal cavities separated by septa), and the sonar (hearing) apparatus is less developed. According to Novik, sonar locating organs are present in Leptonycteris and other pollinating Microchiroptera. In American bats with a mixed diet - nectar, fruits and insects - the sonar apparatus is intact. They make long flights with very short visits to sometimes rather poor flowers with a less rigid corolla (in this case, soaring visits are more often observed).

Macroglossinae have a powerful flight, which at first glance resembles the flight of swallows. Some species can hover in much the same way as hummingbirds. Similar data have been obtained for the Glossophaginae.

The presence of a certain harmony between the flower and animals in structure and physiology allows you to create the concept of the existence of a special type of flower pollinated by bats. Secondary self-pollination in Ceiba, or even parthenocarpy, as in cultivated Musa, can only cause harm.

It is noteworthy that although the development of chiropterophilia in America occurred independently and probably much later than elsewhere, and although the bats in question developed as an independent lineage rather late, the basic features that make up the syndrome of chiropterophilia are the same throughout the world. In all regions, bat-pollinated flowers and flower-pollinating bats are mutually adapted. This indicates common features in the physiology of all the bats under consideration. Sometimes, the development of chiropterophilia in different lines may also be based on common features of plant families.

Many flowers open shortly before dark and fall off in the early morning. Since the times of activity of diurnal birds and dusky bats, as well as the opening times of flowers pollinated by birds and bats, overlap, it is not surprising that some chiropterophilous plants are visited by birds. Werth apparently never made nocturnal observations and therefore lists Ceiba and Kigelia in the list of ornithophilous plants, although birds only plunder these flowers.

Flowers pollinated by bats are similar in appearance to flowers pollinated by hummingbirds, but only more pronounced. Flagellifloria (pendulifloria) is often observed, with flowers hanging freely on long hanging stems (Adansonia, Parkia, Marcgravia, Kigelia, Musa, Eperua). This is most evident in some species of Misipa, in which shoots up to 10 m long or more bring attraction elements out of the foliage.

In Markhamia, Oroxylum there is also a pincushion type with tight stems that lift the flowers up. The giant agave blossom speaks for itself. Favorable is also the pagoda-like structure of some Bombacaceae.

The phenomenon of chiropterophilia also explains why caulifloria, best adapted to visiting bats, is practically limited to the tropics, with only 1,000 cases found. Good examples are Cres "centia, Parmentiera, Durio and Amphitecna. In many genera (Kigelia, Misipa), flagellifloria and caulifloria are observed simultaneously in the same species; in other cases, these signs occur in different species.

Caulifloria is a secondary phenomenon. Its ecological nature is consistent with the results of studies of its morphological basis. Numerous cases had no taxonomic morphological, anatomical and physiological commonality.

In most examples of cauliflory where the flower was not chiropterophilous, another connection with bats was found, namely chiropterochory, the dispersal of seeds by fruit-eating bats. In this case, bats had an earlier and more widespread effect on tropical fruit, including color, position, and smell. This older syndrome corresponds exactly to the newer chiropterophilia syndrome. Basicaulicarpy may also be related to saurochory syndrome (seed dispersal by reptiles), a phenomenon older than angiosperms.

The sequence of flowering periods is necessary for both the plant and the bats. In Java, on large plantations of Ceiba, which has a certain flowering period, bats visited the flowers only in places close to gardens with Musa, Parkia, etc., where they could feed when Ceiba was not in bloom.

In general, the relatively young nature of chiropterophily is reflected in the distribution of bat-pollinated flowers among plant families. So, in Ranales, bats eat fruits, but do not visit flowers. Pollination of flowers by bats occurs in highly evolutionarily advanced families ranging from the Capparidaceae and Cactaceae, and is concentrated mainly in the Bignoniaceae, Bombacaceae and Sapotaceae. Many cases are completely isolated.

Some families (Bombacaceae and Bignoniaceae), characterized by chiropterophilia, apparently developed independently of each other in the Old and New Worlds, probably on the basis of some kind of preadaptations. It may also have happened in some genera, such as Misipa and especially Parkia, which Baker and Harris considered in terms of the representations noted.

Similarly, Bignoniacae and Bombacaceae, like Misipa and Musa, are characterized by some intermediate types which are pollinated by both birds and bats. Bombaxmalabaricum (Gossampinusheptaphylla) is ornithophilous, but not completely so it has open red cup-shaped daytime flowers. The flowers of this plant, however, have a bat-smell, which is characteristic of the chiropterophilic related species valetonii. In Java, malabaricum flowers are neglected by bats, but in the tropical regions of southern China they are eaten by Pteropinae. Chiropterophilia appears to have evolved from ornithophilia in the Bignoniaceae; Bombacaceae and Musa have probably reverted and subtropical species are being pollinated by birds. The transition from hawk-pollinated flowers in Cactaceae has already been considered.

It is still too early to try to quantify the links and their genetic implications. Sometimes bats (especially the slow Pteropinae) confine themselves to a single tree, resulting in self-pollination. Macroglossinae, characterized by rapid flight, make circles around trees, and apparently remember spatial relationships very well. However, in the study of pollen on wool and especially large accumulations of pollen in the stomachs, it was found that they are not characterized by constancy to flowers. It is also not clear how genetic purity is maintained in related chiropterophilic species, such as the wild species Musa, or whether it is maintained at all.

2.5 Entomophily or insect pollination

Insects in the flowers are attracted to pollen and the sweet juice of nectar. It is secreted by special glands - nectaries. They are located inside the flower, often at the base of the petals. Pollen and sweet nectar are the food of many insects.

Here a bee sat on the inflorescence. She quickly makes her way to the nectar stores hidden in the depths of the flower. Squeezing among the anthers and touching the stigma, the bee sucks nectar with its proboscis. Her furry body was covered with yellow pollen. In addition, the bee collected pollen in special baskets on its hind legs. A few seconds pass, and the bee leaves one flower, flies to another, third, etc.

Large single flowers, small flowers collected in inflorescences, bright color of petals or tepals, nectar and aroma are signs of insect pollinated plants. Fragrant tobacco flowers open only at dusk. They smell a lot. By night, the aroma intensifies, and white large flowers still attract night butterflies from afar.

Large, brightly colored poppy petals and an abundance of pollen in the flower are a good bait for beautiful golden-green bronze beetles. They feed on pollen. Smeared in pollen, bronzes fly from one plant to another and transfer the dust particles adhering to the body to the stigmas of the pistils of neighboring flowers.

There are plants whose flowers are pollinated only by certain insects. For example, snapdragons are pollinated by bumblebees. During flowering, hives with bees are brought to the gardens. Bees in search of food pollinate the flowers of fruit trees, and the yield of fruits increases.

The flowers, relying on insects for such an important matter, amaze with a variety of shapes and shades, and almost all of them are brightly colored. However, in all this diversity, one can trace the structure common to all. A typical flower is a receptacle surrounded by leaves that have taken the form of petals and stamens.

Some resemblance to the leaves was retained only by the calyx, formed from green sepals and forming the outer circle of the perianth. The sepals hiding the bud in poppies fall off when the flower blooms, while in tomatoes or strawberries they remain until the fruit is fully ripe.

Above the calyx are larger and brightly colored petals, although wind-pollinated flowers such as the single-flowered coastal (Littorella unijlora) do not have them at all. Hidden within some of the modified petals are nectaries, groups of cells that produce sweet nectar to attract insects. Nectaries may be pouches at the base of the petals, like buttercups, or long spurs, like violets. Spurs usually attract pollinators with long proboscises - hawks and butterflies.

The sepals and petals together form a perianth, although gardeners more often use this term to designate fused perianths, as in daffodils. The totality of all the petals is called the corolla. The reproductive organs of the flower are also located here. The female organ - pistil - consists of an ovary, a style and a stigma, on which pollen settles. The column is surrounded by male organs (stamens), each of which is a thin stalked filament with an anther at the top.

Depending on the position of the ovary, the upper one is distinguished when the petals and sepals are located below it, and the lower one, when parts of the flower are above the ovary. In some flowers - for example, in buttercups - several pistils are collected in one corolla, containing all the female organs; others may have fused pistils, sometimes with one style for all, sometimes with several.

Most flowering plants are bisexual, but some of them have chosen a different path of development. Almost all species of sedge (all wind-pollinated) have male and female flowers on the same plant, while the insect-pollinated holly has same-sex flowers on separate male and female plants.

If a tulip throws out only one flower, then, for example, lily of the valley flowers are collected in an inflorescence on one pedicel, attracting insects with their appearance and delicate fragrance. Some inconspicuous flowering plants lure pollinators by surrounding the flowers with brightly colored leaves. The fiery red "petals" of the poinsettia (Euphorbia pulcherti) are actually modified leaves, or bracts. No one, except for insects, usually notices real flowers.

Conclusion

Having done this work, we found out that pollination is the main method of reproduction of angiosperms, there are 2 types of pollination: autogamy (self-pollination) and cross-pollination.

In the work, the morphological adaptations of flowering plants to cross-pollination, such as wind, water, bird, insect and bat pollination, were considered and studied.

In this work, the goal was achieved and all tasks were disclosed.

pollination angiosperm plant morphological

Bibliography

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3. A. E. Vasil’ev, N. S. Voronin, A. G. Elenevsky, and M. I. Serebryakova, Russ. Botany. Morphology and anatomy of plants. - M. Education, 1988, 528 p.

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11.2.1. Pollination by birds. Ornithophilia

Since the birds fly well and the surface of their body is not smooth, they have good external conditions for pollination. No one is surprised that insects get their food from flowers, but the corresponding actions of birds cause great surprise and reflection on how they got the "idea" to use the nectar of flowers (probably the absence of pollinating birds in Europe led to this attitude). One of the ideas put forward was the idea that pollination arose as a result of the eating of flowers by birds, and may have been primarily fruit feeding *. It has also been suggested that woodpeckers or sap-eating woodpeckers (Sphyrapicus) sometimes change their diet and switch to juices flowing from hollows (some of them also peck fruits; Dendrocopus analis - fruits of Cassia grandis). A third group of "explanations" suggests that the birds pursued insects in the flowers and happened to find nectar or pierce the succulent tissues; or at first they drank water collected in flowers to quench their thirst, since in tropical forests water is difficult to access for animals living in the crowns of trees. The fact that hummingbirds originally pursued insects in flowers can be seen even today. The rapid absorption of nectar makes it difficult to identify it in the stomach of birds, while indigestible remains of insects are easily recognized. However, in the ornithological literature there is a large amount of data indicating that the digestive systems of birds are filled with nectar. Extraction of nectar by piercing the base of the corolla is further evidence that all this is done for the sake of extracting nectar. Insects cannot obtain nectar in this way. Some hummingbirds have become addicted to flower piercing, similar to some hymenoptera (Snow and Snow, 1980). None of the insects obtain nectar from the closed flowers of the Loranthaceae from Java, which open only when attacked by nectar-seeking birds (Docters van Leeuwen, 1954). The fact that birds visit flowers can be confirmed even on very old museum preparations by the presence of pollen grains in feathers or on the beak (Iwarsson, 1979).

* (Some examples of this we find 1) in nightingales (Pychonotus), which eat the fleshy bracts of Freycinetia funicular is and act as legitimate pollinators. Characteristically, this species has fiery red, odorless daytime flowers; 2) in semi-dystropic birds that drink from little specialized flowers such as Bombax (Gossampinus) or pluck petals in Dillenia species; 3) cases of Boerlagiodendron (Beccari, 1877), which is said to attract pollinating birds (pigeons) by mimicking fruits (sterile flowers); 4) in birds that pollinate nectarless flowers of Calceolaria uniflora, in which they bite off food bodies (Vogel, 1974).)

Hummingbirds need a large amount of energy, especially when hovering (215 cal / h per 1 g of body weight). It is precisely such a large expenditure of energy for soaring and flying (plus rest) that can explain the small size of these birds. After a period of fasting, nutrient stores can be severely reduced despite low metabolic rates during sleep.

In pollinators with different energy budgets (Schlising et al., 1972), the efficiency of nectar uptake and its metabolism are different. The presence of flowers with a large amount of nectar is a signal forcing hummingbirds to seize and defend territories (Grant and Grant, 1968; Stiles, 1971). One could refer to the migration of hummingbirds to those places where these flowers are numerous, especially during the breeding season.

Anyone who has witnessed how sparrows completely destroy a crocus bed in the spring knows that these birds eat any food; therefore it is natural that birds "loving" sugar will sooner or later discover its sources in flowers, just as sparrows do. The way in which plants and birds themselves adapted to each other is remarkable, but again this is no more, but no less remarkable, than the mutual adaptation of plants and insects.

* (Sparrows and finches in New Zealand have been noted (McCann, 1952) to teach birds to rob flowers by piercing them at the base (see also Swynnerton, 1915; Iyengar, 1923). In the gardens of southern Europe, local poorly adapted birds have often been observed plundering introduced ornithophilous plants (Abutilon, Erythrina), mainly damaging the flowers, but sometimes also pollinating them. Compare with the remarkable way in which blackbirds on the islands have adapted to take nectar from the Chilean Riua cultivated on these islands (Ebbels, 1969).)

From the point of view of pollination, it was absolutely irrelevant whether the birds visited the flowers for nectar or for the sake of capturing an insect, as long as these visits did not become regular. Whether the nectar or the insect is the reason for the visit is a problem of adaptation, not function. In Java, Zosterops visits the non-ornithophilous Elaeocarpus ganitrus to collect mites, which are in abundance in flowers (van der Pijl).

There is no doubt that birds perched on flowers for all the reasons mentioned above. The sparrow example shows that this is still the case today. Even if, from the gardener's point of view, the flowers were damaged, they were successfully pollinated. Damage to the flower itself is of little consequence as long as the pistil is not damaged. After all, explosive flowers are also destroyed themselves. There are observations that sparrows pollinated pear trees (K. Fægri).

Other similar occasional flower visits by dystrophic birds have recently been recorded in birds migrating to England from more southerly areas (Ash et al., 1961). Campbell (1963) observed various birds in England chasing insects in flowers with very little pollen.

From these examples of dystropic visits to flowers, it can be seen that there is a bed transition through certain allotropic birds with a mixed diet, in which nectar is one of the ingredients (Porsch, 1924), to eutropic ones, as a result of which true ornithophily is established.

For a long time, observations were made of visits to hummingbird flowers. Ornithophily as a scientifically recognized phenomenon was established by Trelease (Trelease, 1881) at the end of the last century, and Johow (Johow, 1900), Fries (Fries, 1903) and mainly Werth (Werth, 1915) studied it in more detail. . However, it was only when Porsche in the 1920s (see references) collected a huge amount of data and drew convincing conclusions about now well-known phenomena that ornithophilia was unanimously recognized, even if its origin is still a matter of controversy.

* (Marden (1963) relates a wonderful story of flies attracted by the smell of Stanhopea graveolens flowers, hunted by a hidden spider, which in turn was hunted by a hummingbird (Glaucis hirsuta) that pollinated the flower.)

The flowering plants (Dicaeidae) visit a variety of flowers, while showing a curious "specialization" to one group of plants, namely the tropical Loranthoideae, in which they not only visit ornithophilous flowers, but also adapt to fruit digestion and seed dispersal (Docters van Leeuwen, 1954 ). The oldest observations of bird pollination in the New World were made by Catesby (1731-1743) and Rumphius (1747) in the Old World.

The areas in which any type of ornithophilia is found practically cover the American continent and Australia and further tropical Asia and the deserts of South Africa. According to Werth (1956b), Israel is the northern limit of this area, with Cinnyris visiting the flowers of the red Loranthus, and recently Galil (Galil, in press) reported to us the abundance of these birds on plants growing in gardens.

In the mountains of Central and South America, the number of ornithophilous species is unusually high. If bees are present in the highlands of Mexico, they are just as efficient as birds as pollinators, except that birds are more efficient under unfavorable conditions (Cruden, 1972b). However, Bombus species are not very sensitive to climate. Their presence can completely change the picture, as shown by Docters van Leeuwen (1933). Stevens (Stevens, 1976) points out similar results of Rhododendron pollination in the mountains of Papua.

It is obvious that in Australia and New Zealand the number of eutropic pollinating insects is also low, and the function of higher bees performed by them on other continents is taken over by birds (cf. the predominant role of the ornithophilous genus Eucalyptus). We have fairly accurate data on the confinement of ornithophilous plant families only to certain areas (percentage).

The ability to soar, well developed in hummingbirds (Greenewait, 1963), is rare in other groups of birds; it is observed, for example, in the honey-eating Acanthorhynchus, and is poorly developed in the Asiatic Arachnothera. Some birds can soar in strong headwinds.

Grant (1949b) argued that "persistence" to flowers is poorly developed in birds and that their feeding habits are too complex. Information about the evolution of constancy to flowers is different for different authors. Snow and Snow (1980) suggest a very close relationship - monotropic, in our current terminology - between Passiflora mixta and Ensifera ensifera. Obviously, the relationship between different species of hummingbirds and the plants that provide them with food varies greatly, ranging from strict territoriality to a very inefficient strategy of successive visits, when birds use any available source of nectar (Snow and Snow, 1980). It is also necessary to take into account the possibility of learning in birds. If diversity is allowed, then impermanence may be due to the lack of a proper distinction between deceit and preferable constancy. Birds feed on any kind of food, so it is natural that if there is a profuse bloom and a large amount of nectar is available, the apparent preference of the birds in this case will simply be a matter of statistics and will not depend on the food itself. If there is no such flowering, then they can fly from one species to another or even use other food. Any observed consistency will be impressive even though flower tube length, beak length, nectar composition, etc. may play a role in flower selection. Under extreme circumstances (migration and nesting), birds eat (various?) flowers. Johow (1900) noted in Chile that hummingbirds may even switch to European fruit trees or Citrus species. Hemitropic birds switch to fruits more often (causing some damage). In the tropics, birds especially prefer fresh flowering trees. The ecological significance of this, of course, is not absolute, but relative and can be of selective significance.

The phylogenetic development of tropical plant species and the most highly developed groups of pollinators has led to a distinct and easily recognizable bird pollination syndrome that excludes other pollinators*. Any random combinations in this case are impossible. The mutual dependence is well seen in the example of the Hawaiian flower girls Drepanididae and the flowers pollinated by them, which, when the birds were exterminated, became autogamous (Porsch, 1930; Amadon, 1947).

* (For the differential diagnosis of classes of ornithophilous flowers and flowers pollinated by diurnal Lepidoptera. The differences are rather indistinct, especially in American plants.)

Some bird-pollinated flowers are brush-like (Eucalyptus, heads of Proteaceae and Compositae; Mutisia), others are slant-mouthed (Epiphyllum) or tubular (Fuchsia fulgens). Some moths (Mucuna spp., Erythrina) are typically ornithophilous.

The fact that different types of flowers are ornithophilous indicates a recent development of ornithophily, which is on top of the previous ecomorphological organizations that determine the types of structure, etc., but leading to a secondary convergence of the style. Isolated instances of resemblance between unrelated flowers, regarded by some morphologists as a mysterious "repeated pair" and by others as orthogenetic, probably represent parallel adaptations in the realm of pollination. Considering the phylogeny of these convergent changes, we can say that in some phylogenetic lineages they often occur independently of each other.

Ornithophilia syndrome is described in Table. 7, which illustrates the correspondence of the flowers to the ethology of the birds in question (cf. also the discussion of the genus Salvia).

Table 7. Ornithophilia syndrome

Flowers pollinated by birds	Birds pollinating flowers
1. Daytime bloom	Daily
2. Bright colors, often scarlet or contrasting	Visual with red sensitivity, not ultraviolet
3. Lip or margin absent or recurved, flowers tubular and/or pendulous, necessarily zygomorphic	Too big to sit on a flower
4. Solid walls of the flower, filaments rigid or fused, protected ovary, nectar hidden	strong beak
5. No smell	Probably no sensitivity to smell
6. Abundance of nectar	Large, consume a lot of nectar
7. The capillary system lifts the nectar up or prevents it from leaking	*
8. Possibly the deep tube or spur is wider than that of a butterfly-pollinated flower.	Long beak and tongue
9. Remoteness of nectar - the genital area can be large	Big long beak, big body
10. Nectar indicator is very simple or absent	Show "intelligence" when finding the entrance to the flower

Some comments on the table will not be superfluous. Relationships are partly positive (attractive), partly negative (excluding competing visitors). Hymenoptera's neglect of bird-pollinated flowers is an exception, seen in Mimulus cardinalis, Monarda species, and Salvia splendens in any botanical garden. Darwin had already noted that the bees neglected Lobelia fulgens, which grows in the garden among the melittophilic species.

The effectiveness of this syndrome is shown by the fact that typical bird-pollinated flowers growing in European gardens attract the attention of short-billed, unadapted dystrophic birds, and also by the fact that flower-pollinating birds immediately recognize and then try to use the flowers of introduced bird-pollinated plants (Porsch, 1924). Flower size is not included in the syndrome. Many flowers pollinated by birds are relatively small. The flowers pollinated by birds are usually deep, and do not belong to any one particular class, but the most characteristic of them are brush-like and tubular.

Birds that pollinate flowers are not always limited to those types of flowers that have this syndrome. As already mentioned, if there is no nectar, they will also eat "non-adapted" flowers.

This table needs one clarification. There are regionally differentiated flower traits for hummingbirds and other birds. In the former (American), the flowers are erect or drooping, with open organs ready for pollination by soaring pollinators (cf. Pedilanthus, Quassia). It is believed that hummingbirds are reluctant to sit on erect flowers (Frankie, 1975). In the latter (Asian and African), planting is carried out near the flower, and the flower itself indicates the landing site (Spathodea campanulata, Protea, Aloe). From this point of view, we could analyze the species Fuchsia and Erythrina (Toledo, 1974) to confirm their "American" or "Old World" appearance, as Linnaeus put it: Hie flos facien americanam habet (or something like that). spirit). There are "American" flowers with landing pads, such as Heliconia rostrata.

In the Chilean Puya (subgenus Puya) the outer part of each incomplete inflorescence is sterile and forms a peculiar perching site * used by legitimate pollinators, members of the Icteridae (Gourlay, 1950) and thrushes in England (Ebbels, 1969). An excellent example of a specially formed structure of a similar type is found in Antholyza ringens, a member of the African flora. Due to the lack of places to plant, the flowers of some American plants grown in Java are inaccessible to nectar-eating animals, so they pierce them (van der Pijl, 1937a). The African Aloë ferox in Chile is not pollinated by hummingbirds, but by tyrants (Elaeina) (Johow, 1901). Cruden (1976) provides other examples (Eucalyptus and Leonotis species) where adaptations for planting birds adversely affect pollination by hummingbirds of plants introduced to the Americas. However, the American continents have many plants that are pollinated by perching birds (Toledo, 1975). At the same time, many ornithophilous flowers of the Old World that do not have landing sites should be considered, at least in this respect, as flowers exhibiting the hummingbird pollination syndrome. One of the characteristic features of flowers is that their reproductive organs are hidden (Gill and Conway, 1979).

* (There is a beautiful illustration (Fig. 13) in J. Roy. Horticult. Soc. 87 (1962).)

Regarding paragraph 2 of Table. 7 we can say that many flowers pollinated by birds are white. The relationship between bird and color is not absolute. In some geographical areas, the flowers pollinated by birds are mostly not red (eg Hawaii). However, the general significance of red is evidenced by statistical data showing its relative predominance in the tropics, especially in the Andes (see Porsch, 1931a; data for South Africa - Vogel, 1954). Let us also mention the color preference of Trochilidae, known to every observer, and in addition to general sensory-physiological studies, which indicate a high sensitivity of birds to red and a much lower sensitivity to blue. Since true red is invisible to most or even all pollinating insects, the red flowers seen by birds (and humans) represent a free ecological niche open to exploitation (K. Grant, 1966). For migratory American bird pollinators—both seasonal and occasional—red is usually a common cue indicating the availability of a suitable nectar source (like a hotel roadside sign), usually increasing visit efficiency. These issues are discussed in more detail in the work on pollinating birds with illustrations and extensive information by K. and V. Grants (Grant, Grant, 1968); see also (Raven, 1972).

Sensitivity to different regions of the spectrum in different species of birds varies. In one species of hummingbird (Huth and Burkhardt, 1972), a shift to the short wavelength region of the spectrum compared to the human visible spectrum was found (from 363 to about 740 nm compared to 390 and 750 nm).

In Columnea florida birds are attracted by the red spots on the leaves, while the flowers themselves are hidden. Since this spot does not reproduce the shape of the flower, a high degree of mental integration can be assumed in birds pollinating Columnea florida (Jones and Rich, 1972).

Flowers with bright, contrasting coloration include flowers in the species Aloë, Strelitzia and many bromeliads.

On point 3, it should be noted that the zygomorphy (the usual sign of entomophily) in ornithophilous flowers is formed in two aspects when the sometimes dangerous lower edge is removed. This typical form is characteristic even of the ornithophilous Cactaceae, the rest of which usually have actinomorphic flowers (see below). Flowers that lack elements, usually insect landing sites and bird obstacles, can be admired in flower shops, such as the red flowers of Corytholoma.

Item 4 is questioned by Snow and Snow (1980), although it is partially recognized as possible (Datura). However, their point of view that the rigid basal parts of the floral tubes protect the nectar from "theft" is quite acceptable.

Regarding point 5, we may add that the smell itself is not an obstacle, but its absence is characteristic of ornithophilia. It is still found in transitional flowers such as Bombax and Spathodea. According to some data (von Aufsess, 1960), the pollen and nectar of ornithophilous flowers have such a weak smell that it is impossible to teach bees to distinguish it.

In order to get an idea of the amount of nectar in ornithophilous flowers (point 6), one should (in temperate botanical gardens) recall Phormium or Aloë, from which the nectar literally drips, or Protea from Cape Cod. The nectar of ornithophilous plants cannot be too viscous, even if it is more concentrated than the nectar of flowers pollinated by butterflies. Otherwise, the capillary vascular system of the plant could not function, delivering nutrients to various organs (Baker, 1975).

Tubular flowers (item 8) often occur in the interpetalous, but they are "improvised" in many Choripetalae, such as Cuphea, species of Cadaba, Tropeolum, Fuchsia and Malvaviscus. In contrast to the short tubes of the melittophilic species, the ornithophilous Iris fulva found by Vogel (1967) has a long tube with strong walls. Birds can stick out their tongues and use flower tubes that are longer than their beaks. Short-billed hummingbirds usually pierce flowers and steal nectar.

In Pitcairnia, which exhibits an extreme degree of ornithophily, the common, unspecialized, rather short-tubular bromeliad flower forms a long tube with a yawn by twisting the inner petal, which joins with the two upper petals to form a vault of the flower with anthers and stigma located at the top, as in a flower with a yawn , not in the center, as is typical for this family. The similarity between the fiery red flowers of P. nabilis and the flowers of Salvia splendens or Anapaline (Iridaceae) is striking (botanical garden, Berlin, K. Fægri).

As regards point 10 - the absence of a nectar indicator, it should be noted that the strong reduction and flexion of the corolla tip in one way or another makes it difficult to locate the nectar indicator.

We have already mentioned that the transition to ornithophily is mostly recent, but in some groups ornithophily seems to be older. Porsche (Porsch, 1937a), alas, without any evidence obtained under natural conditions, identified a suprageneric group in Cactaceae (Andine Loxantocerei), in which, apparently, ornithophily in the tribe was fixed. Snow and Snow (1980) provide other examples of the coevolution of ornithophilous flowers and their pollinators.

Among Euphorbiaceae with dense cyathium, Poinsettia has large glands and red bracts that attract hummingbirds. The genus Pedilanthus (Dressier, 1957) is characterized by an even higher specialization, which appeared from the beginning of the Tertiary period, and in this genus the glands are in spurs, the flowers are erect and zygomorphic.

Even among orchids, which have excellent pollinators - bees, some species have switched to ornithophily in an endless search for new pollinators typical of this family. In the South African genus Disa, some species have probably become ornithophilous (Vogel, 1954). Therefore, the flowers of this genus pollinated by butterflies are already red, with a spur and with a reduced upper lip. We believe that the same occurs in Cattleya aurantiaca and in some species of Dendrobium in the mountains of New Guinea (van der Pijl and Dodson, 1966). Birds visiting the flowers of Elleanthus capitatus and Masdevallia rosea were observed by Dodson (Dodson, 1966).

Dressier (1971) gives a list of orchids that are pollinated by birds and suggests that the dark color of their pollinia (as opposed to the usual yellow) does not contrast with the color of the hummingbird's beak, and therefore the birds have no desire to brush them off.

11.2.2. Pollination by bats. Chiropterophilia

The first conscious observations of bats visiting flowers were made by Burck (Burck, 1892) in the Biitenzorg (now Bogorsk) botanical garden. He observed that frugivorous bats (probably Cynopterus) visited the inflorescences of Freycinetia insignis, a plant now known to be entirely chiropterophilic, in contrast to its closely related ornithophilous species (section 11.2.1)*.

* (Hart's observations in Trinidad in 1897, carried out by him on Bauhinia megalandra and Eperua falcata, are often mentioned in the literature, confusingly with incorrect conclusions.)

Later, some authors (Cleghorn, McCann - India; Bartels, Heide, Danser, Boedijn - Java) described other cases, and the Kigelia example became a classic. As early as 1922, Porsche expressed certain considerations regarding chiropterophilia, noting its characteristic features and predicting many possible examples. After visiting South America, he published in his homeland (Porsch, 1931b) the first well-studied case (Crescentia cujete in Costa Rica, see also Porsch, 1934-1936).

Thanks to the work of van der Pijl (1936, 1956) in Java, Vogel (Vogel, 1958, 1968, 1969) in South America, Jaeger (Jaeger, 1954), and Baker and Harris (Baker and Harris, 1959) in Africa, bat pollination has now been identified in many plant families. It turned out that some plants, previously considered ornithophilous, are pollinated by bats (for example, species of Marcgravia).

Relationships between fruit and flower feeding Megalochiroptera are still partly dystropic. In Java, Cynopterus has been found to eat Durio flowers and parts of Parkia inflorescences. In eastern Indonesia and Australia, Cynopterus and Pteropus destroy many Eucalyptus flowers, indicating hitherto unbalanced pollination conditions.

Howell (Howell, 1974) is of the opinion that Leptonycteris satisfies its protein requirements from pollen, and the protein in the pollen is not only of high quality, but also in sufficient quantity. She also states that the chemical composition of the pollen of flowers pollinated by bats is adapted to the use of it by these animals and differs from the composition of the pollen of related species that are pollinated by other animals. This can be seen as a floral part of the co-evolution of the chiropterophilia syndrome. Until now, the issue of African fruit-eating bats that swallow pollen has not been clarified.

In the class of flowers pollinated by bats, an early side branch of evolution was found to form its own subclass, for which the only pollinator is Pteropineae. In these flowers, solid food (with a characteristic odor) is represented only by specialized structures. We find neither nectar nor large masses of pollen here. Freycinetia insignis has a sweet bract, Bassia and Madhuca species have a very sweet and easily detachable corolla. It is possible that another species of Sapotaceae, namely the African Dumoria heckelii, also belongs to this subclass.

Nectar-eating New World bats are typically found in the tropics, but some migrate to the southern US during the summer, visiting cacti and agaves in Arizona. There is no record of bat pollination in Africa from the north of the Sahara, while Ipomoea albivena at Southpansbergen in South Africa just grows in the tropics*. In Asia, the northern limit of bat pollination is in the north of the Philippines and the island of Hainan, with a small Pteropinae going beyond the latitude of Canton. The Eastern Pacific border runs in a sharp ridge through the Caroline Islands to Fiji. Macroglossinae are known to have visited flowers in Northern Australia (introduced by Agave), but the native Adansonia gregorii has all the characteristics of chiropterophilia; therefore, chiropterophilia must also exist on this continent.

* (The possibility of bat pollination of the white-flowered tree strelitzia (Strelitzia nicolai) in the eastern region of Cape Cod needs to be investigated.)

Knowing the characteristics of pollination by bats can help in solving the mysteries of the origin of plants. The chiropterophilic flower of Musa fehi is evidence that the species was introduced to Hawaii, where there are no bats. Chiropterophilia could have taken place in his homeland, in New Caledonia, from where, as several botanists have established, he comes from.

Nectar-eating bats are characterized by a variety of adaptations. Thus, the Macroglossinae of the Old World have adapted to life on flowers, namely, they have decreased in size (the mass of Macroglossus minimus is 20-25 g), they have reduced molars, a long muzzle, a very elongated tongue with long soft papillae at the end (and not hard bristles, as noted in older publications). Our description is based on observations of the life of bats, while the denial of chiropterophilia is based on studies of animals preserved in alcohol.

Similarly, some species of the New World Glossophaginae have a longer snout and tongue than their insectivorous relatives. Musonycteris harrisonii has a tongue length of 76 mm and a body length of 80 mm (Vogel, 1969a). Vogel (1958, 1968, 1969) considers that the coat hairs of Glossophaga are particularly well adapted to carry pollen, since they are provided with scales similar in size to those on the hairs covering the belly of a bumblebee.

The physiology of Megachiroptera's sense organs deviates from what we normally see in bats. The eyes are large, sometimes with a folded retina (allowing rapid accommodation), with many rods but no cones (causing color blindness). Fruit-eating Epomops franqueti (Ayensu, 1974) at night shows huge eyes, almost the same as those of a lemur. Smell perception probably plays a more important role than usual (large nasal cavities separated by septa), and the sonar (hearing) apparatus is less developed. According to Novick (cited in Vogel, 1969a), sonar locating organs are present in Leptonycteris and other pollinating Microchiroptera. In American bats with a mixed diet - nectar, fruits and insects - the sonar apparatus is intact. They make long flights with very short visits to sometimes rather poor flowers with a less rigid corolla (in this case, soaring visits are more often observed).

Macroglossinae have a powerful flight, which at first glance resembles the flight of swallows. Some species can hover in much the same way as hummingbirds. Similar data were obtained for Glossophaginae (Heithaus et al., 1974).

In the comparative table. 8 again list the adaptive syndrome, partly positive, partly negative.

Table 8. Chiropterophilia syndrome

Flowers pollinated by bats	Bats pollinating flowers
1. Night flowering, mostly only one night	nocturnal lifestyle
2. Sometimes whitish or creamy	Good vision, probably for close orientation
3. Often dull brown, greenish or purple, rarely pink	color blindness
4. Strong smell at night	Good sense of smell for long distance orientation
5. Musty smell, reminiscent of the smell of fermentation	Glands with a stale (heavy) odor as attractants
6. Large mouth and strong single flowers, often hard (brush-shaped) inflorescences of small flowers	Large animals clinging with thumb claws
7. A very large amount of nectar	Large with a high metabolic rate
8. Large amount of pollen, large or many anthers	Pollen as the only source of protein
9. Peculiar arrangement on top of foliage	Hearing organs are poorly developed, flights inside the foliage are difficult

Some remarks should be made about the table. eight.

To point 1. Night flowering is easy to observe in bananas, where large bracts that cover the flowers open every night.

Many flowers open shortly before dark and fall off in the early morning. Since the times of activity of diurnal birds and dusky bats, as well as the opening times of flowers pollinated by birds and bats, overlap, it is not surprising that some chiropterophilous plants are visited by birds. Werth (1956a) apparently never made nocturnal observations and therefore lists Musa paradisiaca, Ceiba and Kigelia as ornithophilous plants, although birds only plunder these flowers.

To points 4 and 5. A researcher with some experience can easily determine the smell of flowers pollinated by bats. It has much in common with the smell of the animals themselves, which probably has some kind of social function in the formation of accumulations of animals and also has some kind of stimulating effect. This odor has been found to have a strong effect on captive-bred Pteropus.

The same smell, reminiscent of the smell of butyric acid, was found in fruits distributed by bats (for example, guava). This circumstance, as well as the manner in which fruits are represented, has served as a starting point for the development of chiropterophily, primarily in those taxa in which fruits are dispersed by bats, a condition often found in the tropics (van der Pijl, 1957). In many Sapotaceae, Sonneratiaceae, and Bignoniaceae, this odorous substance possibly aids in bonding. Vogel (Vogel, 1958) found the presence of a pronounced bat smell in the fruits of the species Drymonia, while other Gesneriaceae (Satrapea) have flowers pollinated by bats.

Bat-smell, still or already characteristic of some ornithophilous species of Gossampinus, Mucuna and Spathodea, is associated with species pollinated by bats.

The transition from nocturnal sphingophilic odors seems relatively easy. Porsche (1939) suggested this chemical change in some Cactaceae, where night flowering, successful cauliflory and a large number of anthers were already characteristic as organizational characters. This assumption was confirmed by Alcorn (Alcorn et al., 1961) on the example of a giant cactus, Carnegiea, in Arizona. Pollen had previously been found in the bat Leptonycteris nivalis, and the authors confirmed its presence, albeit under artificial conditions.

An odor, sometimes reminiscent of mold, is found in Musa, and cabbage in Agave. A chemical study is needed.

Go to point 6. Typical claw prints usually betray nocturnal visits to flowers that have been shed. In the inflorescences of bananas, the number of impressions on the bracts makes it possible to count the number of visits. Random hovering may explain the lack of claw marks (Carnegiea).

To point 7. Nectar is even more abundant than in flowers pollinated by birds. In Ochroma lagopus, 7 ml was found, in O. grandiflora - up to 15 ml. We do not have data on its possible composition. In the cold morning time, banana nectar forms a colloidal structure. Heithaus et al. (1974) describe two strategies for nectar feeding in Bauhinia pauletti. Large bats gather in groups, land and spend quite a lot of time to collect nectar from flowers. Small bats hover in front of the flowers and consume nectar during repeated, very short visits. Obviously, in this case, no traces remain on the flower, indicating a visit. Sazima and Sazima (1975) describe a strategy that is more like a successive visit strategy.

To point 8. Elongation of anthers is evident in Ceiba, Bauhinia, Agave, Eugenia cauliflora and Cactaceae, and an increase in their number - in Adansonia, which has up to 1500-2000 anthers.

To point 9. The need for open space for landing and takeoff and the relative inability to echolocate in Megachiroptera have been proven in experiments with placing obstacles in front of flowers; at the same time, collisions of mice with an obstacle were observed; in addition, hunters catch Megachiroptera more easily than Microchiroptera.

Flowers pollinated by bats are similar in appearance to flowers pollinated by hummingbirds, but only more pronounced. Flagellifloria (pendulifloria) is often observed, with flowers hanging freely on long hanging stems (Adansonia, Parkia, Marcgravia, Kigelia, Musa, Eperua). This is most evident in some species of Mucuna, in which shoots up to 10 m long or more bring attraction elements out of the foliage.

The phenomenon of chiropterophilia also explains why caulifloria, best adapted to visiting bats, is practically limited to the tropics, with only 1,000 cases found. Good examples are Cressentia, Parmentiera, Durio and Amphitecna. In many genera (Kigelia, Mucuna), flagellifloria and caulifloria are observed simultaneously in the same species; in other cases, these characters are found in different species.

Our previous articles have discussed all the known theories of cauliflory in the tropics and have spoken of its extremely wide distribution (van der Pijl, 1936, 1956). Caulifloria is a secondary phenomenon. Its ecological nature is consistent with the results of studies of its morphological basis. Numerous cases had no taxonomic morphological, anatomical and physiological commonality.

In most examples of cauliflory where the flower was not chiropterophilous, another association with bats was found, namely chiropterochory, seed dispersal by fruit bats (van der Pijl, 1957). In this case, the bats had an earlier and more widespread influence on tropical fruits (and thus flower position), including color, position, and scent. This older syndrome corresponds exactly to the newer chiropterophilia syndrome. Basicaulicarpy may also be related to saurochory syndrome (seed dispersal by reptiles), a phenomenon older than angiosperms.

Some families (Bombacaceae and Bignoniaceae), characterized by chiropterophilia, apparently developed independently of each other in the Old and New Worlds, probably on the basis of some kind of preadaptations, as already mentioned in the previous sections. It may also have happened in some genera, such as Mucuna and especially Parkia, which Baker and Harris (1957) reviewed in terms of noted representations.

Similarly, Bignoniacae and Bombacaceae, like Mucuna and Musa, are characterized by some intermediate types which are pollinated by both birds and bats. Bombax malabaricum (Gossampinus heptaphylla) is ornithophilous but not completely so it has open red cup-shaped daytime flowers. The flowers of this plant, however, have the smell of a bat, which is characteristic of the chiropterophilic related species B. valetonii. In Java, the flowers of B. malabaricum are neglected by bats, but in the tropical regions of southern China they are eaten by Pteropinae (Mell, 1922). Chiropterophilia appears to have evolved from ornithophilia in the Bignoniaceae; Bombaceae and Musa have probably reverted and the subtropical species are being pollinated by birds. The transition from hawk-pollinated flowers in Cactaceae has already been considered.

It is still too early to try to quantify the links and their genetic implications. Sometimes bats (especially the slow Pteropinae observed by Baker and Harris) confine themselves to a single tree, resulting in self-pollination. Macroglossinae, characterized by rapid flight, make circles around trees, and apparently remember spatial relationships very well. However, in the study of pollen on wool and especially large accumulations of pollen in the stomachs, it was found that they are not characterized by constancy to flowers. It is also not clear how genetic purity is maintained in related chiropterophilic species, such as the wild species Musa, or whether it is maintained at all.

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