扁形动物门

Features common to all subgroups

The lack of circulatory and respiratory organs limits platyhelminths to sizes and shapes that enable oxygen to reach and carbon dioxide to leave all parts of their bodies by simple diffusion. Hence, many are microscopic and the large species have flat ribbon-like or leaf-like shapes. The guts of large species have many branches, allowing nutrients to diffuse to all parts of the body. Respiration through the whole surface of the body makes them vulnerable to fluid loss, and restricts them to environments where dehydration is unlikely: sea and freshwater, moist terrestrial environments such as leaf litter or between grains of soil, and as parasites within other animals.
The space between the skin and gut is filled with mesenchyme, a connective tissue made of cells and reinforced by collagen fibers that act as a type of skeleton, providing attachment points for muscles. The mesenchyme contains all the internal organs and allows the passage of oxygen, nutrients and waste products. It consists of two main types of cell: fixed cells, some of which have fluid-filled vacuoles; and stem cells, which can transform into any other type of cell, and are used in regenerating tissues after injury or asexual reproduction.
Most platyhelminths have no anus and regurgitate undigested material through the mouth. However, some long species have an anus and some with complex, branched guts have more than one anus, since excretion only through the mouth would be difficult for them. The gut is lined with a single layer of endodermal cells that absorb and digest food. Some species break up and soften food first by secreting enzymes in the gut or pharynx (throat).
All animals need to keep the concentration of dissolved substances in their body fluids at a fairly constant level. Internal parasites and free-living marine animals live in environments with high concentrations of dissolved material, and generally let their tissues have the same level of concentration as the environment, while freshwater animals need to prevent their body fluids from becoming too dilute. Despite this difference in environments, most platyhelminths use the same system to control the concentration of their body fluids. Flame cells, so called because the beating of their flagella looks like a flickering candle flame, extract from the mesenchyme 
water that contains wastes and some reusable material, and drive it into networks of tube cells which are lined with flagella and microvilli. The tube cells' flagella drive the water towards exits called nephridiopores, while their microvilli reabsorb reusable materials and as much water as is needed to keep the body fluids at the right concentration. These combinations of flame cells and tube cells are called protonephridia.
In all platyhelminths, the nervous system is concentrated at the head end. This is least marked in the acoels, which have nerve nets rather like those of cnidarians and ctenophores, but densest around the head. Other platyhelminths have rings of ganglia in the head and main nerve trunks running along their bodies.

主要分组

早期分类将扁形动物分为四组，即：涡虫纲、吸虫纲、单殖纲及绦虫纲。这个分类长久以来都被认为是一种人工强行分类，所以Ehlers (1985)提出一个比较符合支序亲缘学观点的分类，将涡虫纲分为12个目，然后再将吸虫类、单殖类和绦虫类等寄生虫组合成一个新的Neodermata目。可是这个分类仍然主要只为科学文献采用，所以以下我们仍然以传统分类去介绍扁形动物的分类。

涡虫纲

涡虫纲包括有物种4500种均能自由迁徙，长度从1 mm（0.039英寸）到600 mm（24英寸）不等。大多数为掠食者或者是食腐动物。多居住在阴暗潮湿的地方，如落叶底下。然而，部分物种会与其他动物（例如：甲壳类动物）共生、甚或是寄生。Free-living turbellarians are mostly black, brown or gray, but some larger ones are brightly colored. The Acoela and Nemertodermatida were traditionally regarded as turbellarians, but are now regarded as members of a separate phylum, the Acoelomorpha, or as two separate phyla. Xenoturbella, a genus of very simple animals, has also been reclassified as a separate phylum.
Some turbellarians have a simple pharynx lined with cilia and generally feed by using cilia to sweep food particles and small prey into their mouths, which are usually in the middle of their undersides. Most other turbellarians have a pharynx that is eversible (can be extended by being turned inside-out), and the mouths of different species can be anywhere along the underside. The freshwater species Microstomum caudatum can open its mouth almost as wide as its body is long, to swallow prey about as large as itself.
Most turbellarians have pigment-cup ocelli ("little eyes"), one pair in most species, but two or even three pairs in some. A few large species have many eyes in clusters over the brain, mounted on tentacles, or spaced uniformly around the edge of the body. The ocelli can only distinguish the direction from which light is coming and enable the animals to avoid it. A few groups have statocysts, fluid-filled chambers containing a small, solid particle or, in a few groups, two. These statocysts are thought to be balance and acceleration sensors, as that is the function they perform in cnidarian medusae and in ctenophores. However, turbellarian statocysts have no sensory cilia, and how they sense the movements and positions of the solid particles is unknown. On the other hand, most have ciliated touch-sensor cells scattered over their bodies, especially on tentacles and around the edges. Specialized cells in pits or grooves on the head are probably smell sensors.
Planarians, a subgroup of seriates, are famous for their ability to regenerate if divided by cuts across their bodies. Experiments show that, in fragments that do not already have a head, a new head grows most quickly on those closest to the original head. This suggests the growth of a head is controlled by a chemical whose concentration diminishes from head to tail. Many turbellarians clone themselves by transverse or longitudinal division, and others, especially acoels, reproduce by budding.
The vast majority of turbellarians are hermaphrodites (have both female and male reproductive cells), and fertilize eggs internally by copulation. Some of the larger aquatic species mate by penis fencing, a duel in which each tries to impregnate the other, and the loser adopts the female role of developing the eggs. In most species, "miniature adults" emerge when the eggs hatch, but a few large species produce plankton-like larvae.

吸虫纲

吸虫纲的学名Trematoda源自希腊语的τρῆμα，意思就是一个孔洞，refers to the cavities in their holdfasts which resemble suckers and anchor them within their hosts. 。
The skin of all species is a syncitium, a layer of cells that shares a single external membrane. Trematodes are divided into two groups, Digenea and Aspidogastrea (also known as Aspodibothrea).

复殖亚纲

These are often called flukes, as most have flat rhomboid shapes like that of a flounder (Old English flóc). There are about 11,000 species, more than all other platyhelminthes combined, and second only to roundworms among parasites on metazoans. Adults usually have two holdfasts, a ring around the mouth and a larger sucker midway along what would be the underside in a free-living flatworm.
Although the name "Digeneans" means "two generations", most have very complex life cycles with up to seven stages, depending on what combinations of environments the early stages encounter – most importantly whether the eggs are deposited on land or in water. The intermediate stages transfer the parasites from one host to another. The definitive host in which adults develop is a land vertebrate, the earliest host of juvenile stages is usually a snail that may live on land or in water, and in many cases a fish or arthropod is the second host. For example, the adjoining illustration shows the life cycle of the intestinal fluke metagonimus, which hatches in the intestine of a snail; moves to a fish, where it penetrates the body and encysts in the flesh; then moves to the small intestine of a land animal that eats the fish raw; and then generates eggs that are excreted and ingested by snails, thereby completing the cycle. Schistosomes, which cause the devastating tropical disease bilharzia, belong to this group.
Adults range between 0.2 mm（0.0079英寸） and 6 mm（0.24英寸） in length. Individual adult digeneans are of a single sex, and in some species, slender females live in enclosed grooves that run along the bodies of the males, and partially emerge to lay eggs. In all species, the adults have complex reproductive systems and can produce between 10,000 and 100,000 times as many eggs as a free-living flatworm. In addition, the intermediate stages that live in snails reproduce asexually.
Adults of different species infest different parts of the definitive host, for example the intestine, lungs, large blood vessels, and liver. The adults use a relatively large, muscular pharynx to ingest cells, cell fragments, mucus, body fluids or blood. In both the adults and the stages that live in snails, the external syncytium absorbs dissolved nutrients from the host. Adult digeneans can live without oxygen for long periods.

Aspidogastrea

Members of this small group have either a single divided sucker or a row of suckers that cover the underside. They infest the guts of bony or cartilaginous fish and of turtles, and the body cavities of marine and freshwater bivalves and gastropods. Their eggs produce ciliated swimming larvae, and the life cycle has one or two hosts.

Cercomeromorpha

These parasites attach themselves to their hosts by means of disks that bear crescent-shaped hooks. They are divided into Monogenea and Cestoda.

单殖纲

单殖纲，旧作单殖目，目前包括约1100个物种，绝大多数都是外部寄生虫，需要特定物种作为中间宿主。体型细小，约2～3 cm长。这些中间宿主以鱼类最为常见，但有时也可以是两栖类或水生爬虫类动物。少数属于内部寄生虫。单殖纲的成虫在其后端有大型的附着器官，haptors (Greek ἅπτειν, haptein, means "catch"), which have Sucker (parasitic worm anatomy), clamp (zoology), and hooks. They often have flattened bodies. In some species, the 咽 secretes enzymes to digest the host's skin, allowing the parasite to feed on blood and cellular debris. Others graze externally on mucus and flakes of the hosts' skins. 
“单殖纲”这名称源由于这些寄生虫的生命周期只有一个非蚴阶段（Nonlarval stage）。

Evolution

An outline of the origins of the parasitic life style has been proposed. Epithelial feeding monopisthocotyleans on fish hosts are basal in the Neodermata and were the first shift to parasitism from free living ancestors. The next evolutionary step was a dietary change from epithelium to blood. The last common ancestor of Digenea + Cestoda was monogenean and most likely sanguinivorous.
The earliest known fossils of tapeworms have been dated to 270百万年前. They were found in coprolites (fossilised faeces) from an elasmobranch.

Parasitism

Cestodes (tapeworms) and digeneans (flukes) cause important diseases in humans and their livestock, and monogeneans can cause serious losses of stocks in fish farms. Schistosomiasis, also known as bilharzia or snail fever, is the second-most devastating parasitic disease in tropical countries, behind malaria. The Carter Center estimated 200 million people in 74 countries are infected with the disease, and half the victims live in Africa. The condition has a low mortality rate, but often is a chronic illness that can damage internal organs. It can impair the growth and cognitive development of children, and increase the risk of bladder cancer in adults. The disease is caused by several flukes of the genus Schistosoma, which can bore through human skin. The people most at risk are those who use infected bodies of water for recreation or laundry.
In 2000, an estimated 45 million people were infected with the beef tapeworm Taenia saginata and 3 million with the pork tapeworm Taenia solium. Infection of the digestive system by adult tapeworms causes abdominal symptoms that are unpleasant but not disabling or life-threatening. However, neurocysticercosis resulting from penetration of T. solium larvae into the central nervous system is the major cause of acquired epilepsy worldwide. In 2000, about 39 million people were infected with trematodes (flukes) that naturally parasitize fish and crustaceans, but can pass to humans who eat raw or lightly cooked seafood. Infection of humans by the broad fish tapeworm Diphyllobothrium latum occasionally causes vitamin B₁₂ deficiency and, in severe cases, megaloblastic anemia.
The threat to humans in developed countries is rising as a result of social trends: the increase in organic farming, which uses manure and sewage sludge rather than artificial fertilizers, and spreads parasites both directly and via the droppings of seagulls which feed on manure and sludge; the increasing popularity of raw or lightly cooked foods; imports of meat, seafood and salad vegetables from high-risk areas; and, as an underlying cause, reduced awareness of parasites compared with other public health issues such as pollution. In less-developed countries, inadequate sanitation and the use of human feces (night soil) as fertilizer and to enrich fish farm ponds continues to spread parasitic platyhelminths, and poorly designed water-supply and irrigation projects have provided additional channels for their spread. People in these countries often cannot afford the cost of fuel required to cook food thoroughly enough to kill parasites. Controlling parasites that infect humans and livestock has become more difficult, as many species have become resistant to drugs that used to be effective, mainly for killing juveniles in meat. While poorer countries still struggle with unintentional infection, cases have been reported of intentional infection in the US by dieters desperate for rapid weight-loss.

Pests

There is concern about the proliferation in northwest Europe, including the British Isles, of the New Zealand planarian Arthurdendyus triangulatus and the Australian flatworm Australoplana sanguinea, both of which prey on earthworms. A. triangulatus is thought to have reached Europe in containers of plants imported by botanical gardens.

Benefits

In Hawaii, the planarian Endeavouria septemlineata has been used to control the imported giant African snail Achatina fulica, which was displacing native snails, and Platydemus manokwari, another planarian, has been used for the same purpose in Philippines, Indonesia, New Guinea and Guam. Although A. fulica has declined sharply in Hawaii, there are doubts about how much E. septemlineata contributed to this. However, P. manokwari is given credit for severely reducing, and in places exterminating, A. fulica – achieving much greater success than most biological pest control programs, which generally aim for a low, stable population of the pest species. The ability of planarians to take different kinds of prey and to resist starvation may account for their ability to decimate A. fulica. However, these abilities have raised concerns that planarians may themselves become a serious threat to native snails.
A study in La Plata, Argentina shows the potential for planarians such as Girardia anceps, Mesostoma ehrenbergii, and Bothromesostoma evelinae to reduce populations of the mosquito species Aedes aegypti and Culex pipiens. The experiment showed that G. anceps in particular can prey on all instars of both mosquito species and maintain a steady predation rate over time. The ability for these flatworms to live in artificial containers showed the potential of placing these species in popular mosquito breeding sites, which would ideally reduce the amount of mosquito-borne diseases.