【扁形动物门】
扁形动物门（学名：Platyhelminthes；语源：πλατύ platy 扁平 + ἑλμινθ- helminth- 虫）是动物界的一个门，是一类简单的无环节两侧对称动物，属于无脊椎动物，有三胚层，无体腔，无呼吸及循环系统，有口，但无肛门，所以必须保持身体扁平，以使氧气及养料能够透过渗透来吸收。消化腔只有一个开口，同时用于进食及排泄；所以食物在其体内无法有效处理。
已记录的扁形动物约有29500种。生活于淡水、海水等潮湿处，体前端有两个可感光的色素点。体表部分或全部分布有纤毛。
传统的医学文献会将扁形动物划分为非寄生的涡虫纲（例如：真涡虫科的物种）和三个会寄生的物种的纲：绦虫纲、吸虫纲及单殖纲。然而，由于涡虫纲已证实并非单系群，这种划分方式在动物学来看已经过时。
非寄生的扁虫都是捕食者，栖息于水中或遮荫的陆上潮湿环境，例如落叶堆等。寄生种如绦虫和吸虫的生命周期比较复杂：牠们的成熟阶段会以寄生虫的形式居住在鱼类或陆上脊椎动物的消化系统里；而中间宿主阶段会寻找可被感染的中间宿主。吸虫的卵从最终宿主体内排出，而成年绦虫会产生大量雌雄同体的节片，在成熟后会分离，排出宿主，再释放卵。与其他寄生的类群不同，单殖纲是水生生物的体外寄生虫，其幼虫在附着于合适宿主之后，再变态为成虫。
因为扁形动物没有体腔，它们曾被认为是最原始的两侧对称动物。但是，在1980年代中期以来的研究发现原来被分类为扁形动物的一个群体──无腔动物门，离最初的两侧对称动物较任何其他现代类群更近。除去无腔动物后的扁形动物门是一个单系群，即是有一个共同祖先及其所有后裔组成的。扁形动物门属于冠轮动物，是较复杂的两侧对称动物的三个进化支之一。近年来的分析显示出已重新定义、不再包含无腔动物门的扁形动物门，是由两个单系群，即小锁状目和被杆体纲所组成的；而绦虫纲、吸虫纲和单殖纲则组成了被杆体纲下的一个分支──新皮类底下的单系群，至于传统的涡虫纲物种范围则涵盖了除新皮类外的所有扁形动物。因此，传统上认定的涡虫纲现在被认为是并系群，因为其不包含从传统涡虫纲分支出去的寄生性物种之类群。
超过一半已知的扁形动物属于寄生虫, 而且某些物种会对人类以及牲畜造成很大的伤害。血吸虫病是由吸虫纲裂体属的物种所造成的，是所有由寄生虫造成的人类疾病中第二致命的，仅次于疟疾。囊虫脑病变是当猪肉绦虫的幼体（囊尾幼虫）侵入中枢神经系统时产生, 是后天获得之癫痫的主因之一。这些寄生性物种在开发中国家的威胁来自于生食与未完全煮熟之食物受到欢迎，以及从高风险地区的进口。在低度开发的国家中，人们可能无法负担用以煮熟食物之燃料的费用，再加上设计不良之供水系统以及灌溉工程，使得不良的公共卫生及农业卫生提高被感染之风险。
两种扁形动物在菲律宾、印尼、夏威夷、新几内亚和关岛被用来控制威胁原生种蜗牛生存的外来种非洲大蜗牛之族群大小，并收到良好成效。但是，最近发现这些扁形动物可能自身就会威胁原生种蜗牛的生存。在西北欧地区，以蚯蚓为食的纽西兰扁虫（Arthurdendyus triangulatus）的散播引起各方担忧。
==描述==
Varied flatworm species from Kunstformen der Natur (1904), plate 75
===分辨性特征===
扁形动物皆为
动物：牠们的左半侧与右半侧互为镜像；这亦表示了这些动物可以分辨出其顶部与底部 、以及其头部及尾部。就像其他的两侧对称动物，牠们的细胞可分成三个胚层，即：内胚层、中胚层及外胚层，而刺胞动物及栉水母等辐射对称动物则只有两个胚层。Beyond that, they are "defined more by what they do not have than by any particular series of specializations."。
不同于其他两侧对称动物，扁形动物并没有体腔，
所以被称为无体腔动物。牠们也没有特化的循环系统及呼吸系统器官，而这些都是为扁形动物分类时的解剖特征。牠们的身体是柔软且没有分节的。
===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.
==主要亚群==
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)提出一个比较符合系统发生学观点的分类，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===
The turbellarian Pseudoceros dimidiatus
涡虫纲包括有物种4500种均能自由迁徙，长度从abbr=on到abbr=on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.
Two turbellarians are mating by penis fencing. Each has two penises, the white spikes on the undersides of their heads.
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===
Life cycle of the diagenean Metagonimus
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 abbr=on and abbr=on 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====
Silhouettes of bodies of various polyopisthocotylean ns
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著器官，s (Greek ἅπτειν, haptein, means "catch"), which have , , 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）。
====绦虫纲====
带绦虫属物种的生命周期。图5显示的是4条猪带绦虫，在其末端有盘状倒钩。图6显示的是一整条绦虫。
绦虫有扁瘦但非常长的身体。其学名
“cestode”的字根源于拉丁语的 cestus，意思就是「带子」。这3400个绦虫的物种的成虫都是内部寄生虫。牠们没有口部和肠脏，
and the syncitial skin absorbs nutrients – mainly carbohydrates and amino acids – from the host, and also disguises it chemically to avoid attacks by the host's immune system. Shortage of carbohydrates in the host's diet stunts the growth of the parasites and kills some. Their metabolisms generally use simple but inefficient chemical processes, and they compensate by consuming large amounts of food relative to their size.
In the majority of species, known as eucestodes ("true tapeworms"), the neck produces a chain of segments called proglottids by a process known as strobilation. Hence, the most mature proglottids are furthest from the scolex. Adults of Taenia saginata, which infests humans, can form proglottid chains over ft long, although ft is more typical. Each proglottid has both male and female reproductive organs. If the host's gut contains two or more adults of the same cestode species, they generally fertilize each other, but proglottids of the same worm can fertilize each other and even fertilize themselves. When the eggs are fully developed, the proglottids separate and are excreted by the host. The eucestode life cycle is less complex than that of digeneans, but varies depending on the species. For example:
*Adults of Diphyllobothrium infest fish, and the juveniles use copepod crustaceans as intermediate hosts. Excreted proglottids release their eggs into the water, and the eggs hatch into ciliated, swimming larvae. If a larva is swallowed by a copepod, it sheds the cilia and the skin becomes a syncitium and the larva makes its way into the copepod's hemocoel (internal cavity that is the main part of the circulatory system) and attached itself with three small hooks. If the copepod is eaten by a fish, the larva metamorphoses into a small, unsegmented tapeworm, drills through to the gut and becomes an adult.
*Various species of Taenia infest the guts of humans, cats and dogs. The juveniles use herbivores – for example pigs, cattle and rabbits – as intermediate hosts. Excreted proglottids release eggs that stick to grass leaves and hatch after being swallowed by a herbivore. The larva makes its way to the herbivore's muscles and metamorphoses into an oval worm about in long, with a scolex that is kept inside. When the definitive host eats infested and raw or undercooked meat from an intermediate host, the worm's scolex pops out and attaches itself to the gut, and the adult tapeworm develops.
A members of the smaller group known as Cestodaria have no scolex, do not produce proglottids, and have body shapes like those of diageneans. Cestodarians parasitize fish and turtles.
==Classification and evolutionary relationships==
Relationships of Platyhelminthes to other Bilateria:Note: Bold indicates members of traditional "Platyhelminthes".
Relationships of Platyhelminthes (excluding Acoelomorpha) to each other
The oldest confidently identified parasitic flatworm fossils are cestode eggs found in a Permian shark coprolite, but helminth hooks still attached to Devonian acanthodians and placoderms might also represent parasitic flatworms with simple life cycles. The oldest known free-living platyhelminth specimen is a fossil preserved in Eocene age Baltic amber and placed in the monotypic species Micropalaeosoma balticus, while the oldest subfossil specimens are schistosome eggs discovered in ancient Egyptian mummies. The Platyhelminthes have very few synapomorphies, distinguishing features that all Platyhelminthes and no other animals have. This makes it difficult to work out both their relationships with other groups of animals and the relationships between different groups that are described as members of the Platyhelminthes.
The "traditional" view before the 1990s was that Platyhelminthes formed the sister group to all the other bilaterians, which include, for example, arthropods, molluscs, annelids and chordates. Since then molecular phylogenetics, which aims to work out evolutionary "family trees" by comparing different organisms' biochemicals such as DNA, RNA and proteins, has radically changed scientists' view of evolutionary relationships between animals. Detailed morphological analyses of anatomical features in the mid-1980s and molecular phylogenetics analyses since 2000 using different sections of DNA agree that Acoelomorpha, consisting of Acoela (traditionally regarded as very simple "turbellarians") and Nemertodermatida (another small group previously classified as "turbellarians") are the sister group to all other bilaterians, including the rest of the Platyhelminthes. However, a 2007 study concluded that Acoela and Nemertodermatida were two distinct groups of bilaterians, although it agreed that both are more closely related to cnidarians (jellyfish, etc.) than other bilaterians are.
Xenoturbella, a bilaterian whose only well-defined organ is a statocyst, was originally classified as a "primitive turbellarian". However, it has recently been reclassified as a deuterostome.
The Platyhelminthes excluding Acoelomorpha contain two main groups, Catenulida and Rhabditophora, both of which are generally agreed to be monophyletic (each contains all and only the descendants of an ancestor that is a member of the same group). Early molecular phylogenetics analyses of the Catenulida and Rhabditophora left uncertainties about whether these could be combined in a single monophyletic group, but a study in 2008 concluded they could, therefore Platyhelminthes could be redefined as Catenulida plus Rhabditophora, excluding the Acoelomorpha.
Other molecular phylogenetics analyses agree the redefined Platyhelminthes are most closely related to Gastrotricha, and both are part of a grouping known as Platyzoa. Platyzoa are generally agreed to be at least closely related to the Lophotrochozoa, a superphylum that includes molluscs and annelid worms. The majority view is that Platyzoa are part of Lophotrochozoa, but a significant minority of researchers regard Platyzoa as a sister group of Lophotrochozoa.
It has been agreed since 1985 that each of the wholly parasitic platyhelminth groups (Cestoda, Monogenea and Trematoda) is monophyletic, and that together these form a larger monophyletic grouping, the Neodermata, in which the adults of all members have syncitial skins. However, there is debate about whether the Cestoda and Monogenea can be combined as an intermediate monophyletic group, the Cercomeromorpha, within the Neodermata. It is generally agreed that the Neodermata are a sub-group a few levels down in the "family tree" of the Rhabditophora. Hence the traditional sub-phylum "Turbellaria" is paraphyletic, since it does not include the Neodermata although these are descendants of a sub-group of "turbellarians".
==演化==
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. They were found in coprolites (fossilised faeces) from an elasmobranch.
== 与人的联系 ==
=== 寄生 ===
Magnetic resonance image of a patient with neurocysticercosis demonstrating multiple cysticerci within the brain
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.
==特点==
*体形小至1毫米，大可至25公分。
*可在水中，地面生活，或行寄生生活
*初生表皮层全部为外胚层细胞，后来又中胚层细胞移入。
*躯干表面有纤毛表皮，可用于运动。
*皮层底下有长轴，环形和对角走向的肌肉，腹背部薄壁组织中的肌肉可拮抗静水压的作用，成为静水骨骼
*大部分为雌雄同体，体内受精。
*行断裂生殖。
*不完整的消化道
==分类==
===传统分类===
以下为按照传统分类方法最新近的分类：
*涡虫纲（Turbellaria）
*吸虫纲（Trematoda）
**复殖亚纲（Digenea）
**楯盘纲（Aspidocotylea）
*Cercomeromorpha
**绦虫纲（Cestoda）
**单殖纲（Monogenea）
===现时分类===
根据分子亲缘支序学，本门的物种都被重新分类过，与传统分类的差异很大。现时本门物种被分为下列两大支：
*小锁状目 Catenulida
*被杆体纲 Rhabditophora
=== 演化树 ===
==参看==
*Miracidium
* 血吸虫
* 再生医学（Regenerative medicine）
==参考文献==
==延伸阅读==
*Campbell, Neil A., Biology: Fourth Edition (Benjamin/Cummings Publishing, New York; 1996; page 599) 0-8053-1957-3
*Crawley, John L., and Kent M. Van De Graff. (editors); A Photographic Atlas for the Zoology Laboratory: Fourth Edition) (Morton Publishing Company; Colorado; 2002) 0-89582-613-5
*The Columbia Electronic Encyclopedia, 6th ed. (Columbia University Press; 2004) Retrieved 8 February 2005 date=20121013054049
*Evers, Christine A., Lisa Starr. Biology: Concepts and Applications. 6th ed. United States:Thomson, 2006. 0-534-46224-3.
==外部连结==
* Marine flatworms of the world date=20171117162959