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Tunicate

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Rash, S.; Saiga, H.; Satake, M.; Terry, A.; Yamada L.; Wang, H. G.; Awazu, S.; Azumi, K.; Boore, J.; Branno, M.; Chin-Bow, S.; DeSantis, R.; Doyle, S., Francino, P.; Keys, D. N.; Haga, S.; Hayashi, H.; Hino, K.; Imai, K. S.; Inaba, K.; Kano, S.; Kobayashi, K.; Kobayashi, M.; Lee, B. I.; Makabe, K. W.; Manohar, C.; Matassi, G.; Medina, M.; Mochizuki, Y.; Mount, S.; Morishita, T.; Miura, S.; Nakayama, A.; Nishizaka, S.; Nomoto, H.; Ohta, F.; Oishi, K.; Rigoutsos, I.; Sano, M.; Sasaki, A.; Sasakura, Y.; Shoguchi, E.; Shin-i, T.; Spagnuolo, A.; Stainier, D.; Suzuki, M. M.; Tassy, O.; Takatori, N.; Tokuoka, M.; Yagi, K.; Yoshizaki, F.; Wada, S.; Zhang C.; Hyatt, P. D.; Larimer, F.; Detter, C.; Doggett, N.; Glavina, T.; Hawkins, T.; Richardson, P.; Lucas, S.; Kohara, Y.; Levine, M.; Satoh, N.; Rokhsar, D. S. (2002).
1613: 1648: 501: 1857:, one of the most significant being the reduction of the cerebral ganglion, which controls movement and is the equivalent of the vertebrate brain. From this comes the common saying that the sea squirt "eats its own brain". However, the adult does possess a cerebral ganglion adapted to lack of self-locomotion. In the Thaliacea, the larval stage is rudimentary or suppressed, and the adults are pelagic (swimming or drifting in the open sea). Colonial forms also increase the size of the colony by budding off new individuals to share the same tunic. 762: 845: 1283:, which grow along the seabed. Other species have the zooids growing closer together in a tuft or clustered together and sharing a common base. The most advanced colonies involve the integration of the zooids into a common structure surrounded by the tunic. These may have separate buccal siphons and a single central atrial siphon and may be organized into larger systems, with hundreds of star-shaped units. Often, the zooids in a colony are tiny but very numerous, and the colonies can form large encrusting or mat-like patches. 488: 2006:. Self-fertilized eggs develop with a substantially higher frequency of anomalies during cleavage than cross-fertilized eggs (23% vs. 1.6%). Also a significantly lower percentage of larvae derived from self-fertilized eggs metamorphose, and the growth of the colonies derived from their metamorphosis is significantly lower. These findings suggest that self-fertilization gives rise to inbreeding depression associated with developmental deficits that are likely caused by expression of deleterious recessive mutations. 1666: 155: 1896:, have a transport function, and are arranged in a single central row. Other zooids link to the phorozooids, which then detach themselves from the nurse. These zooids develop into gonozooids, and when these are mature, they separate from the phorozooids to live independently and start the cycle over again. Meanwhile, the phorozooids have served their purpose and disintegrate. The asexual phase in the lifecycle allows the doliolid to multiply very rapidly when conditions are favourable. 923: 1687: 1025: 825: 978: 902: 1630: 720: 131: 2251:, and other parts of the animal are protein-rich and can be converted into fish feed. Culturing tunicates on a large scale may be possible and the economics of doing so are attractive. As tunicates have few predators, their removal from the sea may not have profound ecological impacts. Being sea-based, their production does not compete with food production as does the cultivation of land-based crops for biofuel projects. 1005: 1268: 1940:, and a larva resembles the tadpole larva of ascidians. Once the trunk is fully developed, the larva undergoes "tail shift", in which the tail moves from a rearward position to a ventral orientation and twists through 90° relative to the trunk. The larva consists of a small, fixed number of cells, and grows by enlargement of these rather than cell division. Development is very rapid and only takes seven hours for a 1980:(class Ascidiacea) is a hermaphrodite that releases sperm and eggs into the surrounding seawater almost simultaneously. It is self-sterile, and thus has been used for studies on the mechanism of self-incompatibility. Self/non-self-recognition molecules play a key role in the process of interaction between sperm and the vitelline coat of the egg. It appears that self/non-self recognition in ascidians such as 1808: 1095: 1393: 1865: 865: 555:
precedence, and grounds for superseding the name never existed. Accordingly, the current (formally correct) trend is to abandon the name Urochorda or Urochordata in favour of the original Tunicata, and the name Tunicata is almost invariably used in modern scientific works. It is accepted as valid by the World Register of Marine Species but not by the Integrated Taxonomic Information System.
2186: 1383: 2128:. Another possible means of introduction is on the shells of molluscs brought in for marine cultivation. Current research indicates many tunicates previously thought to be indigenous to Europe and the Americas are, in fact, invaders. Some of these invasions may have occurred centuries or even millennia ago. In some areas, tunicates are proving to be a major threat to 1324:. Two openings are found in the body wall: the buccal siphon at the top through which water flows into the interior, and the atrial siphon on the ventral side through which it is expelled. A large pharynx occupies most of the interior of the body. It is a muscular tube linking the buccal opening with the rest of the gut. It has a ciliated groove known as an 1860:
Pyrosome colonies grow by budding off new zooids near the posterior end of the colony. Sexual reproduction starts within a zooid with an internally fertilized egg. This develops directly into an oozooid without any intervening larval form. This buds precociously to form four blastozooids which become
1829:. In others, especially colonial species, sperm is released into the water and drawn into the atria of other individuals with the incoming water current. Fertilization takes place here and the eggs are brooded through their early developmental stages. Some larval forms appear very much like primitive 1755:
part of the peribranchial cavity near the atrial siphon. Here, the faeces are caught up by the constant stream of water which carries the waste to the exterior. The animal orientates itself to the current in such a way that the buccal siphon is always upstream and does not draw in contaminated water.
1491:. Exact details of the circulatory system are unclear, but the gut, pharynx, gills, gonads, and nervous system seem to be arranged in series rather than in parallel, as happens in most other animals. Every few minutes, the heart stops beating and then restarts, pumping fluid in the reverse direction. 1471:
is pelagic throughout their lives and may have complex lifecycles. In this class a free living larval stage is absent: Doliolids and pyrosomatids are viviparous–lecithotrophic, and salpids are viviparous–matrotrophic. Only some species of doliolids still have a rudimentary tailed tadpole stage, which
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Dehal, P.; Satou, Y.; Campbell, R. K.; Chapman, J., Degnan, B., De Tomaso, A.; Davidson, B.; Di Gregorio, A.; Gelpke, M.; Goodstein, D. M.; Harafuji, N.; Hastings, K. E.; Ho, I.; Hotta, K.; Huang, W.; Kawashima, T.; Lemaire, P.; Martinez, D.; Meinertzhagen, I. A.; Necula, S.; Nonaka, M.; Putnam, N.;
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Various common names are used for different species. Sea tulips are tunicates with colourful bodies supported on slender stalks. Sea squirts are so named because of their habit of contracting their bodies sharply and squirting out water when disturbed. Sea liver and sea pork get their names from the
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When sufficiently developed, the larva of the sessile species finds a suitable rock and cements itself in place. The larval form is not capable of feeding, though it may have a rudimentary digestive system, and is only a dispersal mechanism. Many physical changes occur to the tunicate's body during
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Fossils of tunicates are rare because their bodies decay soon after death, but in some tunicate families, microscopic spicules are present, which may be preserved as microfossils. These spicules have occasionally been found in Jurassic and later rocks, but, as few palaeontologists are familiar with
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introduced another name for the same group, "Urochorda", to emphasize the affinity of the group to other chordates. No doubt largely because of his influence, various authors supported the term, either as such, or as the slightly older "Urochordata", but this usage is invalid because "Tunicata" has
1919:. An egg in each is fertilized internally by a sperm from another colony. The egg develops in a brood sac inside the blastozooid and has a placental connection to the circulating blood of its "nurse". When it fills the blastozooid's body, it is released to start the independent life of an oozooid. 1328:
on its ventral surface, and this secretes a mucous net which collects food particles and is wound up on the dorsal side of the pharynx. The gullet, at the lower end of the pharynx, links it to a loop of gut which terminates near the atrial siphon. The walls of the pharynx are perforated by several
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Adult tunicates have a hollow cerebral ganglion, equivalent to a brain, and a hollow structure known as a neural gland. Both originate from the embryonic neural tube and are located between the two siphons. Nerves arise from the two ends of the ganglion; those from the anterior end innervate the
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Fedonkin, M. A.; Vickers-Rich, P.; Swalla, B.; Trusler, P.; Hall, M. (2008). "A Neoproterozoic chordate with possible affinity to the ascidians: New fossil evidence from the Vendian of the White Sea, Russia and its evolutionary and ecological implications". HPF-07 Rise and fall of the Ediacaran
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lining the gill slits. To obtain enough food, an average ascidian needs to process one body-volume of water per second. This is drawn through a net lining the pharynx which is being continuously secreted by the endostyle. The net is made of sticky mucus threads with holes about 0.5 μm in
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that resembles a tadpole, with the exception of some members of the families Styelidae and Molgulidae which has direct development. The latter also have several species with tail-less larval forms. The ascidian larvae very rapidly settle down and attach themselves to a suitable surface, later
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is mechanistically similar to self-incompatibility systems in flowering plants. Self-incompatibility promotes out-crossing, and thus provides the adaptive advantage at each generation of the masking of deleterious recessive mutations (i.e. genetic complementation) and the avoidance of
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have a very complex life cycle that includes various zooids with different functions. The sexually reproducing members of the colony are known as gonozooids. Each one is a hermaphrodite with the eggs being fertilised by sperm from another individual. The gonozooid is
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Vickers-Rich P. (2007). "Chapter 4. The Nama Fauna of Southern Africa". In: Fedonkin, M. A.; Gehling, J. G.; Grey, K.; Narbonne, G. M.; Vickers-Rich, P. "The Rise of Animals: Evolution and Diversification of the Kingdom Animalia", Johns Hopkins University Press. pp.
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derives its name. This varies in thickness between species but may be tough, resembling cartilage, thin and delicate, or transparent and gelatinous. The tunic is composed of proteins, crosslinked by phenoloxidase reaction, and complex carbohydrates, and includes
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is a sequential (protogynous) hermaphrodite, and in a colony, eggs are ovulated about two days before the peak of sperm emission. Thus self-fertilization is avoided, and cross-fertilization is favored. Although avoided, self-fertilization is still possible in
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buccal siphon and those from the posterior end supply the rest of the body, the atrial siphon, organs, gut and the musculature of the body wall. There are no sense organs but there are sensory cells on the siphons, the buccal tentacles and in the atrium.
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is retained, and the animals, mostly under 1 cm long, are propelled by undulations of the tail. They secrete an external mucous net known as a house, which may completely surround them and is very efficient at trapping planktonic particles.
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https://www.cambridge.org/core/services/aop-cambridge-core/content/view/0FE5DCCCDFDD464B92DCA4AF68F36F2B/S0022336019001094a.pdf/rare_case_of_an_evolutionary_late_and_ephemeral_biomineralization_tunicates_with_composite_calcareous_skeletons.pdf
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that occupy the rectal loop, and the structure has no duct. Each vesicle is a remnant of a part of the primitive coelom, and its cells extract nitrogenous waste matter from circulating blood. They accumulate the wastes inside the vesicles as
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Gasparini, F; Manni, L; Cima, F; Zaniolo, G; Burighel, P; Caicci, F; Franchi, N; Schiavon, F; Rigon, F; Campagna, D; Ballarin, L (July 2014). "Sexual and asexual reproduction in the colonial ascidian Botryllus schlosseri".
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particles by filtering sea water through their bodies. Ascidians are typical in their digestive processes, but other tunicates have similar systems. Water is drawn into the body through the buccal siphon by the action of
1308:, a variety of cellulose. The tunic is unique among invertebrate exoskeletons in that it can grow as the animal enlarges and does not need to be periodically shed. Inside the tunic is the body wall or mantle composed of 4608:
Sawada H, Morita M, Iwano M (August 2014). "Self/non-self recognition mechanisms in sexual reproduction: new insight into the self-incompatibility system shared by flowering plants and hermaphroditic animals".
1915:. The chain of salps is the 'aggregate' portion of the lifecycle. The aggregate individuals, known as blastozooids, remain attached together while swimming and feeding and growing larger. The blastozooids are 1357:
are also small, under 4 cm (1.6 in) long, and found in the surface waters of both warm and cold seas. They also move by jet propulsion, and often form long chains by budding off new individuals.
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colonial tunicates with a hollow cylindrical structure. The buccal siphons are on the outside and the atrial siphons inside. About 10 species are known, and all are found in the tropics. The 23 species of
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Daugavet, M. A.; Dobrynina, M. I.; Shaposhnikova, T. G.; Solovyeva, A. I.; Mittenberg, A. G.; Shabelnikov, S. V.; Babkina, I. Yu; Grinchenko, A. V.; Ilyaskina, D. V.; Podgornaya, O. I. (22 August 2022).
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first announced the presence in the test of some ascidians of a substance very similar to cellulose, he called it "tunicine", but it is now recognized as cellulose rather than any alternative substance.
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process may be possible for humans. The mechanisms underlying the phenomenon may lead to insights about the potential of cells and tissues to be reprogrammed and to regenerate compromised human organs.
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Tatián, Marcos; Lagger, Cristian; Demarchi, Milagros; Mattoni, Camilo (2011). "Molecular phylogeny endorses the relationship between carnivorous and filter-feeding tunicates (Tunicata, Ascidiacea)".
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root, dock, piling, or ship's hull. They are found in a range of solid or translucent colours and may resemble seeds, grapes, peaches, barrels, or bottles. One of the largest is a stalked sea tulip,
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lived in shallow coastal waters slightly more than 555 to 548 million years ago, and are believed to be the oldest evidence of the chordate lineage of metazoans. The Russian Precambrian fossil
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Seo, H. C.; Kube, M.; Edvardsen, R. B.; Jensen, M. F.; Beck, A.; Spriet, E.; Gorsky, G.; Thompson. E. M.; Lehrach, H.; Reinhardt, R.; Chourrout, D. (2001). "Miniature genome in the marine chordate
2235:. In 1994, over 42,000 tons were produced, but since then, mass mortality events have occurred among the farmed sea squirts (the tunics becoming soft), and only 4,500 tons were produced in 2004. 1597:. The production in animals of cellulose is so unusual that at first some researchers denied its presence outside of plants, but the tunicates were later found to possess a functional cellulose 682:
status. A close relationship between Thaliacea and Ascidiacea, with the former possibly emerging from the latter, had already been proposed since the early 20th century under the name of Acopa.
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are small, mostly under 2 cm (0.79 in) long. They are solitary, have the two siphons at opposite ends of their barrel-shaped bodies, and swim by jet propulsion. The 40 species of
467:. Despite their simple appearance and very different adult form, their close relationship to the vertebrates is evidenced by the fact that during their mobile larval stage, they possess a 2379:
Fedonkin, M. A.; Vickers-Rich, P.; Swalla, B. J.; Trusler, P.; Hall, M. (2012). "A new metazoan from the Vendian of the White Sea, Russia, with possible affinities to the ascidians".
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and feeding, they take in water through the incurrent (or inhalant) siphon and expel the filtered water through the excurrent (or exhalant) siphon. Adult ascidian tunicates are
1340:, the other main class of tunicates, is characterised by free-swimming, pelagic individuals. They are all filter feeders using a pharyngeal mucous net to catch their prey. The 2694: 1773:, trapping tiny crustacea, nematodes, and other small invertebrates with the muscular lobes which surround their buccal siphons. Certain tropical species in the family 1522:. Some tunicates can concentrate vanadium up to a level ten million times that of the surrounding seawater. It is stored in a +3 oxidation form that requires a 1329:
bands of slits, known as stigmata, through which water escapes into the surrounding water-filled cavity, the atrium. This is criss-crossed by various rope-like
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Foster, M. (ed.); Sedgwick, Adam (ed.); The Works of Francis Maitland Balfour. Vol. III. Memorial edition. Pub: Macmillan and co. 1885. May be downloaded from
1100: 1483:. The heart is a double U-shaped tube situated just below the gut. The blood vessels are simple connective tissue tubes, and their blood has several types of 4855: 2280:
appears to be one of the smallest among metazoans and this species has been used to study gene regulation and the evolution and development of chordates.
1279:, integrate with one another. In the simplest systems, the individual animals are widely separated, but linked together by horizontal connections called 678:
sequencing studies. Although the traditional classification is provisionally accepted, newer evidence suggests the Ascidiacea are an artificial group of
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and each has a single ovary and testis, either near the gut or on the body wall. In some solitary species, sperm and eggs are shed into the sea and the
1727:. The gut is U-shaped and also ciliated to move the contents along. The stomach is an enlarged region at the lowest part of the U-bend. Here, digestive 5009: 2804:
Alié, Alexandre; Hiebert, Laurel S.; Scelzo, Marta; Tiozzo, Stefano (19 March 2020). "The eventful history of nonembryonic development in tunicates".
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detached in a single unit when the oozoid disintegrates. The atrial siphon of the oozoid becomes the exhalent siphon for the new, four-zooid colony.
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Section through the wall of an ascidian pyrosoma showing several zooids; (br) buccal siphon; (at) atrial siphon; (tp) test process;
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with a water-filled, sac-like body structure and two tubular openings, known as siphons, through which they draw in and expel water. During their
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Tsagkogeorga, G.; Turon, X.; Hopcroft, R. R.; Tilak, M. K.; Feldstein, T.; Shenkar, N.; Loya, Y.; Huchon, D.; Douzery, E. J.; Delsuc, F. (2009).
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during their early development, but it is lost by the time they have completed their metamorphosis. As members of the Chordata, they are true
1369:. The 70 species of larvaceans superficially resemble the tadpole larvae of amphibians, although the tail is at right angles to the body. The 7391: 3582: 7463: 4076: 1997:(class Ascidiacea) is a colonial tunicate, a member of the only group of chordates that are able to reproduce both sexually and asexually. 5405: 1365:(or Appendicularia), is the only group of tunicates to retain their chordate characteristics in the adult state, a product of extensive 7468: 2156:, a didemnin effective against various types of cancer; as of late January 2021 undergoing Phase III trials as a treatment for COVID-19 4856:"International team of scientists identifies new treatment for COVID-19 that appears to be far more effective than drugs in use now" 3623: 1333:
which extend from the mantle and provide support for the pharynx, preventing it from collapsing, and also hold up the other organs.
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Pisut, D.P. & Pawlik, J.R. (2002). "Anti-predatory chemical defenses of ascidians: Secondary metabolites or inorganic acids?".
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Delsuc F, Philippe H, Tsagkogeorga G, Simion P, Tilak MK, Turon X, López-Legentil S, Piette J, Lemaire P, Douzery EJ (April 2018).
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Fodor ACA; Powers, M. M.; Andrykovich, K.; Liu, J.; Lowe, E. K.; Brown, C. T.; Di Gregorio, A.; Stolfi, A.; Swalla, B. J. (2021).
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Tunicates are unusual among animals in that they produce a large fraction of their tunic and some other structures in the form of
475:. Their name derives from their unique outer covering or "tunic", which is formed from proteins and carbohydrates, and acts as an 7378: 517:; fewer than 100 species of these are found at depths greater than 200 m (660 ft). Some are solitary animals leading a 3709:
Syvanen, M.; Ducore, J. (2010). "Whole genome comparisons reveals a possible chimeric origin for a major metazoan assemblage".
1487:. The blood may appear pale green, but this is not due to any respiratory pigments, and oxygen is transported dissolved in the 4265:"Is there intracellular cellulose in the appendicularian tail epidermis? A tale of the adult tail of an invertebrate chordate" 3185:
Jefferies, R. P. S. (1991) in Biological Asymmetry and Handedness (eds Bock, G. R.; Marsh, J.) pp. 94–127 (Wiley, Chichester).
4086: 4059: 3842: 2897: 7383: 2935: 1534:, although it is unclear whether this is due to the presence of the metal or low pH. Other species of tunicates concentrate 513:
About 3,000 species of tunicate exist in the world's oceans, living mostly in shallow water. The most numerous group is the
4482: 1530:. The vanadocytes are later deposited just below the outer surface of the tunic, where their presence is thought to deter 5328:
Clarke, T.; Bouquet, JM; Fu, X; Kallesøe, T.; Schmid, M; Thompson, E.M. (2007). "Rapidly evolving lamins in a chordate,
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body cavities, if any at all. Whether they do or not, by the end of their larval development, all that remain are the
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Singh, T. R.; Tsagkogeorga, G.; Delsuc, F.; Blanquart, S.; Shenkar, N.; Loya, Y.; Douzery, E. J.; Huchon, D. (2009).
2704: 2328: 1892:. Some of these are known as trophozooids, have a nutritional function, and are arranged in lateral rows. Others are 1573:
across their tissues to rid themselves of nitrogenous waste, though some have a simple excretory system. The typical
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In Permian and Triassic there were also forms with a calcareous exoskeleton. At first they were mistaken for corals.
358: 7430: 6033: 4776: 4737:"The Oikopleura coenocyst, a unique chordate germ cell permitting rapid, extensive modulation of oocyte production" 3652:
A rare case of an evolutionary late and ephemeral biomineralization: tunicates with composite calcareous skeletons]
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off the northeast coast of North America, covering stones, molluscs, and other stationary objects in a dense mat.
7396: 6028: 4365:"Transposon-mediated insertional mutagenesis revealed the functions of animal cellulose synthase in the ascidian 2274:. Both species' mitochondrial and nuclear genomes have been sequenced. The nuclear genome of the appendicularian 1578: 1207:
is identified as a tunicate only tentatively, because its fossils are nowhere near as well-preserved as those of
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Colonies of tunicates occur in a range of forms, and vary in the degree to which individual organisms, known as
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A Guide to the Shell and Starfish Galleries: (Mollusca, Polyzoa, Brachiopoda, Tunicata, Echinoderma, and Worms)
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before being released into the sea as a free-swimming, tadpole-like larva. This undergoes metamorphosis in the
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gland (absent in appendicularians) adds further secretions. After digestion, the food is moved on through the
1723:. The net is rolled up on the dorsal side of the pharynx, and it and the trapped particles are drawn into the 6112: 6102: 6085: 4102:
Odate, S. & Pawlik, J.R. (2007). "The role of vanadium in the chemical defense of the solitary tunicate,
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Tunicates are able to correct their own cellular abnormalities over a series of generations, and a similar
479:. In some species, it is thin, translucent, and gelatinous, while in others it is thick, tough, and stiff. 4498:
Mackie, G. O.; Burighel, P. (2005). "The nervous system in adult tunicates: current research directions".
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Undisputed fossils of tunicates are rare. The best known and earliest unequivocally identified species is
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Nakayama-Ishimura, Akie; Chambon, Jean-Phillippe; Horie, Takeo; Satoh, Nori; Sasakura, Yasunori (2009).
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Sasakura, Y.; Nakashima, K.; Awazu, S.; Matsuoka, T.; Nakayama, A.; Azuma, J. & Satoh, N. (2005).
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Ostrovsky, A. N.; Lidgard, S.; Gordon, D. P.; Schwaha, T.; Genikhovich, G.; Ereskovsky, A. V. (2015).
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Onai T (2018). "The evolutionary origin of chordate segmentation: revisiting the enterocoel theory".
2485: 1927: 1665: 1193:. Results of a new study have shown possible affinity of these Ediacaran organisms to the ascidians. 154: 5181: 4179: 2923: 4832: 4078:
Evolutionary Origin of Sensory and Neurosecretory Cell Types: Vertebrate Cranial Placodes, volume 2
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in its "house". Arrows indicate water movement and (x) the lateral reticulated parts of the house.
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Chen, Jun-Yuan; Huang, Di-Ying; Peng, Qing-Qing; Chi, Hui-Mei; Wang,Xiu-Qiang; Feng, Man (2003).
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Zeng, L.; Swalla, B. J. (2005). "Molecular phylogeny of the protochordates: chordate evolution".
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into an oozooid. This is known as a "nurse" as it develops a tail of zooids produced by budding
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The Tunicata contain roughly 3,051 described species, traditionally divided into these classes:
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Matthysse, A.G.; Deschet, K.; Williams, M.; Marry, M.; White, A.R. & Smith, W.C. (2004).
2784: 2271: 2028: 1986: 1619: 547: 492: 214: 7409: 3888:"Tunicates: Exploring the sea shores and roaming the open ocean. A tribute to Thomas Huxley" 3224:"An updated 18S rRNA phylogeny of tunicates based on mixture and secondary structure models" 398:
is still sometimes used for these animals. They are the only chordates that have lost their
7339: 5168: 5108: 4561: 4382: 4217: 4166: 4115: 3750: 3675: 3494: 3235: 3040: 2813: 2733: 2437: 2390: 2337: 2299: 1993: 1948: 1908: 1889: 539: 5095:(Urochordata, Ascidiacea, Enterogona): Comparison of gene arrangement and tRNA genes with 4647:
Bernstein, H; Hopf, FA; Michod, RE (1987). "The Molecular Basis of the Evolution of Sex".
4314:"Transcriptional regulation of a horizontally transferred gene from bacterium to chordate" 2722:"On the Structure and Innervation of the Muscle Bands of Doliolum (Tunicata: Cyclomyaria)" 685:
The following cladogram is based on the 2018 phylogenomic study of Delsuc and colleagues.
8: 3663: 3627: 1923: 1765: 1697: 1586:, and do not have any obvious means of disposing of the material during their lifetimes. 1459:
developing into a barrel-like and usually sedentary adult form. The species in the class
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in 2010 proposed that sea squirts are descended from a hybrid between a chordate and a
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segmentation, with the possible exception of the seriation of the gill slits. However,
284: 149: 4656: 4240: 4203: 4188: 3517: 3480: 3430:"Phylogenomics resolves the evolutionary chronicle of our squirting closest relatives" 2553:"Phylogenomics resolves the evolutionary chronicle of our squirting closest relatives" 7404: 7326: 6967: 6887: 6237: 6107: 6070: 6058: 6048: 5982: 5421: 5349: 5310: 5271: 5194: 5124: 5073: 4957: 4949: 4888: 4878: 4758: 4721: 4709: 4670: 4660: 4626: 4579: 4478: 4428:
Response of Marine Ecosystems to Global Change: Ecological Impact of Appendicularians
4410: 4345: 4294: 4245: 4131: 4082: 4055: 4017: 3968: 3917: 3872: 3858: 3823: 3784: 3766: 3695: 3522: 3461: 3410: 3317: 3263: 3168: 3117: 3105: 3056: 2913: 2893: 2841: 2829: 2757: 2700: 2667: 2628: 2594: 2576: 2529: 2516: 2467: 2455: 2410: 2365: 2353: 2276: 2146: 2121: 2114: 2081: 2015: 1932: 1706: 1691: 1309: 1126: 379: 5206: 5136: 4969: 4928:
Kürn, Ulrich; Rendulic, Snjezana; Tiozzo, Stefano; Lauzon, Robert J. (August 2011).
3363: 3221: 3026:"Tunicates and not cephalochordates are the closest living relatives of vertebrates" 2970: 2769: 2636: 2537: 2426:"New Ediacaran fossils from the Ukraine, some with a putative tunicate relationship" 463:
The earliest probable species of tunicate appears in the fossil record in the early
7331: 6022: 5341: 5302: 5261: 5251: 5186: 5116: 5063: 5055: 4941: 4748: 4701: 4652: 4618: 4591: 4569: 4507: 4400: 4390: 4335: 4327: 4284: 4276: 4235: 4225: 4184: 4123: 4007: 3999: 3958: 3948: 3907: 3899: 3850: 3815: 3774: 3758: 3718: 3683: 3664:"The first tunicate with a calcareous exoskeleton (Upper Triassic, northern Italy)" 3512: 3502: 3451: 3441: 3400: 3390: 3351: 3307: 3297: 3253: 3243: 3204: 3158: 3148: 3130: 3095: 3083: 3081: 3068: 3048: 2821: 2741: 2679: 2659: 2624: 2584: 2566: 2525: 2445: 2398: 2345: 2247:
is being researched. The cellulose body wall can be broken down and converted into
2220: 2076: 1964: 1655: 1555: 1551: 1526:
of less than 2 for stability, and this is achieved by the vacuoles also containing
1152:
skeletons of the upper midwestern United States. A well-preserved Cambrian fossil,
522: 500: 414: 320: 140: 40: 6241: 5306: 4143: 844: 761: 487: 7248: 7154: 7005: 6908: 6765: 6298: 5837: 5591: 5557: 5484: 5010:"Biofuel made from marine filter feeders? Tunicates usable as source of biofuels" 2319:
Yang, Chuan; Li, Xian-Hua; Zhu, Maoyan; Condon, Daniel J.; Chen, Junyuan (2018).
2215: 1770: 1550:, which may serve a similar function. Other tunicate species produce distasteful 1345: 960: 917: 635: 627: 464: 7318: 4753: 4736: 3819: 3023: 2180: 6913: 6891: 6709: 6621: 6599: 6562: 6053: 5706: 5678: 5585: 5345: 5288: 4622: 4201: 3762: 3583:"A 500 million-year-old fossil reveals the amazing secrets of tunicate origins" 3336: 2450: 2425: 2266: 2255: 2101: 2044: 1796: 1792: 1672: 1296: 1220: 1165: 897: 757: 742: 631: 5120: 5059: 4362: 4127: 4044: 3722: 3446: 3302: 2663: 2571: 2402: 2085:) has taken over a 6.5 sq mi (17 km) area of the seabed on the 1601:, encoded by a gene horizontally transferred from a bacterium. When, in 1845, 1250:). This study was based on a quartet partitioning approach designed to reveal 7452: 7289: 7058: 6962: 6461: 5949: 5854: 5717: 5661: 4953: 4892: 3854: 3770: 3395: 3286:"A phylogenomic framework and timescale for comparative studies of tunicates" 3153: 2806:
Journal of Experimental Zoology Part B: Molecular and Developmental Evolution
2580: 2459: 2357: 2224: 2125: 2036: 1854: 1818: 1782: 1527: 1138: 1105: 922: 715: 675: 422: 83: 5256: 5190: 4395: 4230: 3507: 3248: 1951:, where the fate of the cells is set early on with reduced cell numbers and 1904: 7164: 7159: 7047: 6903: 6789: 6675: 6578: 6557: 6427: 6378: 6334: 6315: 5829: 5784: 5689: 5633: 5470: 5353: 5314: 5275: 5198: 5128: 5077: 4961: 4762: 4713: 4630: 4583: 4414: 4349: 4331: 4298: 4249: 4135: 4021: 3972: 3921: 3827: 3788: 3526: 3465: 3414: 3321: 3267: 3172: 3109: 3060: 2833: 2745: 2671: 2598: 2321:"Geochronological constraint on the Cambrian Chengjiang biota, South China" 2293: 2095: 2086: 2048: 1885: 1566: 1503: 1488: 1464: 1317: 1243: 1118: 679: 602: 597: 592: 442: 307: 274: 5413: 5233: 5034: 4674: 3953: 2761: 1104:) in this Upper Ordovician bryozoan may represent a tunicate preserved by 7365: 7298: 7125: 7096: 7082: 7072: 6657: 6473: 6386: 6168: 5746: 5646: 5391: 4945: 2726:
Proceedings of the Royal Society of London. Series B, Biological Sciences
2557: 2289: 2199: 2190: 2159: 2153: 2129: 2106: 2052: 2043:
can be maintained in laboratory culture, and is of growing interest as a
2027:
organism, reproducing only once in its lifetime. It employs an original
2024: 1760: 1412: 476: 387: 58: 5091:
Yokobori, S.; Watanabe, Y.; Oshima, T. (2003). "Mitochondrial genome of
4780: 3903: 3052: 2825: 2349: 2231:) is the main species eaten. It is cultivated on dangling cords made of 1686: 1024: 7222: 7207: 7202: 7183: 7178: 7106: 7053: 7023: 7015: 6957: 6831: 6823: 6806: 6586: 6498: 6451: 6391: 6353: 6327: 6173: 6158: 5942: 5881: 5734: 5619: 5573: 5507: 3735: 3687: 2206: 2110: 2071: 1963:
relatively late in development and cell cleavage is indeterminate. The
1937: 1877: 1787: 1774: 1562: 1519: 1499: 1484: 1454:
Ascidian tunicates begin life as a lecithotrophic (non-feeding) mobile
1448: 1444: 1292: 1239: 1146: 1058: 973: 820: 671: 623: 613: 588: 580: 453: 449: 264: 233: 199: 103: 68: 7370: 5974: 5394:(1901), British Museum (Natural History). Department of Zoology et al. 5236:"A haplome alignment and reference sequence of the highly polymorphic 5068: 4705: 4280: 4003: 3100: 1841:. These swim by undulations of the tail and may have a simple eye, an 1467:, and the general larval form is kept throughout life. Also the class 977: 824: 7352: 7188: 7135: 7130: 7101: 7090: 7039: 7034: 6986: 6898: 6685: 6680: 6642: 6491: 6483: 6412: 6364: 6285: 6272: 6253: 6248: 6207: 5873: 5563: 5537: 5464: 4810: 4690: 3804:"Delineating metamorphic pathways in the ascidian Ciona intestinalis" 2753: 2232: 2032: 2020: 1956: 1872: 1846: 1834: 1778: 1736: 1724: 1637: 1594: 1583: 1531: 1468: 1408: 1370: 1350: 1337: 1325: 1300: 1160: 1000: 901: 860: 807: 702: 659: 649: 641: 529:. Some are supported by a stalk, but most are attached directly to a 514: 468: 457: 438: 434: 409:
Some tunicates live as solitary individuals, but others replicate by
403: 383: 311: 244: 239: 166: 120: 108: 52: 7260: 4574: 4549: 4511: 3542:"Parasitism of Ordovician bryozoans and the origin of pseudoborings" 3208: 2721: 1443:(or branchial sac), the organs are enclosed in a membrane called an 719: 7357: 7283: 7212: 7196: 7119: 7111: 7066: 7029: 6999: 6981: 6975: 6836: 6777: 6753: 6650: 6531: 6420: 6304: 6195: 6186: 6092: 5843: 5771: 5759: 5492: 5445: 5375:
Dutch Ascidians: Extensive database of images from around the world
3355: 2065: 1881: 1830: 1826: 1720: 1710: 1598: 1547: 1511: 1507: 1460: 1424: 1420: 1416: 1404: 1362: 1341: 1247: 1203: 1189: 1175: 1123: 1020: 663: 645: 584: 568: 534: 371: 253: 186: 98: 93: 78: 73: 63: 44: 5090: 4262: 3801: 3082:
Delsuc, F.; Tsagkogeorga, G.; Lartillot, N.; Philippe, H. (2008).
1569:. Most have no excretory structures, but rely on the diffusion of 1004: 130: 7344: 7172: 7148: 7140: 7077: 6991: 6811: 6798: 6731: 6722: 6667: 6592: 5956: 5795: 5641: 3626:. University of California Museum of Paleontology. Archived from 2378: 2248: 2244: 1842: 1838: 1732: 1570: 1543: 1535: 1515: 1440: 1366: 1354: 1305: 1267: 1170: 1149: 1130: 1109: 840: 576: 572: 526: 472: 410: 399: 113: 88: 5785:
Lissamphibia (modern amphibians: frogs, salamanders, caecilians)
3283: 3137:
mitochondrial genome and support for the new chordate phylogeny"
452:, the most well-known class of tunicates, are commonly known as 7217: 6951: 6151: 6138: 6009: 6006: 5862: 5819: 5808: 5458: 5398: 3988:"Matrotrophy and placentation in invertebrates: A new paradigm" 3613:(Vendian) biota. International Geological Congress - Oslo 2008. 3133:"Tunicate mitogenomics and phylogenetics: peculiarities of the 3024:
Delsuc, F.; Brinkmann, H.; Chourrout, D.; Philippe, H. (2006).
2124:
on the hulls of ships, but may also be introduced as larvae in
1952: 1941: 1912: 1807: 1752: 1740: 1728: 1715: 1574: 1428: 1392: 1313: 1280: 1276: 1184: 1094: 375: 176: 5379: 4735:
Ganot P, Bouquet JM, Kallesøe T, Thompson EM (February 2007).
4311: 2938:
Integrated Taxonomic Information System. Retrieved 2017-03-30.
2514:
Nielsen, C. (2012). "The authorship of higher chordate taxa".
1864: 864: 3985: 3934: 3084:"Additional molecular support for the new chordate phylogeny" 2195: 1822: 1744: 1495: 1476: 1455: 1436: 1321: 609: 418: 329: 224: 5679:
Chondrichthyes (cartilaginous fish: sharks, rays, chimaeras)
4930:"Asexual Propagation and Regeneration in Colonial Ascidians" 4734: 2185: 2063:
Over the past few decades, tunicates (notably of the genera
1944:
to develop into a house-building juvenile starting to feed.
1506:, it contains high concentrations of the transitional metal 1435:, renal, and gonadal cavities of the adults. Except for the 5887: 5380:
Aniseed: A model organism database for ascidians including
3481:"The first tunicate from the Early Cambrian of South China" 2696:
Before the Backbone: Views on the origin of the vertebrates
1748: 1539: 1382: 1271:
Colonial tunicate with multiple openings in a single tunic.
653: 350: 344: 326: 4779:. NOAA Fisheries Service. 19 November 2004. Archived from 3539: 5234:
Small, K. S.; Brudno, M.; Hill, M. M.; Sidow, A. (2007).
5035:
Iannelli, F.; Pesole, G.; Sordino, P.; Gissi, C. (2007).
4204:"A functional cellulose synthase from ascidian epidermis" 3992:
Biological Reviews of the Cambridge Philosophical Society
2223:, both raw and in seafood stews. In Japan and Korea, the 335: 5374: 4927: 4312:
Sasakura, Y.; Ogura, Y.; Treen, N.; et al. (2016).
2719: 543:, which can grow to be over 1 metre (3.3 ft) tall. 6941: 5327: 4651:. Advances in Genetics. Vol. 24. pp. 323–70. 4437: 4435: 2926:
World Register of Marine Species. Retrieved 2011-11-12.
2803: 2296:
which are possibly the sister group of modern tunicates
2079:
coastal waters in many countries. The carpet tunicate (
1868:
A 1901 comparison of frog tadpole and a tunicate larva.
1719:
diameter which can trap planktonic particles including
1523: 3662:
Wendt, Jobst (25 July 2018). Hautmann, Michael (ed.).
3379:"Immunity in Protochordates: The Tunicate Perspective" 1967:
of amphioxus and vertebrates is also relatively slow.
559:
resemblance of their dead colonies to pieces of meat.
4550:"Developmental biology: A chordate with a difference" 4441: 4042:
Dorit, R.L.; Walker, W.F. & Barnes, R.D. (1991).
3739:"New putative phenol oxidase in ascidian blood cells" 3708: 3344:
Publications of the Seto Marine Biological Laboratory
3194: 2424:
Martyshyn, Andrej; Uchman, Alfred (1 December 2021).
359: 341: 332: 323: 4432: 4263:
Hirose, E.; Nakashima, K. & Nishino, A. (2011).
2892:(7th ed.). Cengage Learning. pp. 940–956. 2888:
Ruppert, E.E.; Fox, R.S. & Barnes, R.D. (2004).
2035:
is contained within an ovary that is a single giant
509:
showing oral tentacles at openings of buccal siphons
347: 270: 260: 249: 4646: 2209:species are consumed as food around the world. The 1880:, and at first, the developing embryo feeds on its 1709:(the larval form usually does not feed), capturing 674:were included in Ascidiacea in 2011 as a result of 338: 4466: 4464: 4159:Journal of Experimental Marine Biology and Ecology 4043: 1837:(stiffening rod) and superficially resemble small 1377: 612:consisting of tunicates and vertebrates is called 4769: 4607: 4547: 4041: 1183:-like genus from the Onega Peninsula of northern 1159:Three enigmatic species were also found from the 521:existence attached to the seabed, but others are 7450: 5157:: insights into chordate and vertebrate origins" 2887: 1947:During embryonic development, tunicates exhibit 1295:. The body of an ascidiacean is surrounded by a 406:still display segmentation of the muscle bands. 4686: 4684: 4603: 4601: 4524: 4374:Proceedings of the National Academy of Sciences 4209:Proceedings of the National Academy of Sciences 3486:Proceedings of the National Academy of Sciences 3376: 3019: 3017: 2483: 2423: 2318: 2145:Tunicates contain a host of potentially useful 1759:Some ascidians that live on soft sediments are 5483: 4642: 4640: 4497: 1498:has some unusual features. In some species of 1215:, so this identification has been questioned. 533:, which may be a rock, shell, coral, seaweed, 5990: 5429: 5037:"Mitogenomics reveals two cryptic species in 4804: 3885: 3279: 3277: 1785:in their tunics, and one of these symbionts, 1291:By far the largest class of tunicates is the 5282: 5227: 5219:: CS1 maint: multiple names: authors list ( 5143: 5084: 5028: 4728: 4681: 4598: 3608: 3606: 3593: 3472: 3215: 3124: 3075: 3014: 2789:: CS1 maint: multiple names: authors list ( 2051:position within the closest sister group to 1970: 1955:that are rapidly evolving. In contrast, the 1899:Salps also have a complex lifecycle with an 1795:products are assumed to be available to the 6377: 5632: 5443: 4800: 4798: 4637: 4156: 4101: 3840: 3377:Franchi, Nicola; Ballarin, Loriano (2017). 2883: 2881: 2879: 2877: 2875: 2873: 2871: 2513: 1743:, where undigested remains are formed into 6351: 5997: 5983: 5872: 5436: 5422: 3886:Lemaire, Patrick; Piette, Jacques (2015). 3337:"Phylogenetic Speculation of the Tunicata" 3274: 2869: 2867: 2865: 2863: 2861: 2859: 2857: 2855: 2853: 2851: 2507: 1387:Internal anatomy of a generalised tunicate 129: 6544: 5265: 5255: 5180: 5067: 4752: 4573: 4404: 4394: 4339: 4288: 4239: 4229: 4178: 4074: 4011: 3962: 3952: 3911: 3778: 3603: 3580: 3516: 3506: 3455: 3445: 3404: 3394: 3311: 3301: 3257: 3247: 3162: 3152: 3099: 2699:. Springer Science & Business Media. 2588: 2570: 2449: 1145:), of a possible tunicate found in Upper 6411: 6335:Hemichordata (acorn worms and relatives) 6283: 4807:"Invasive Tunicates of Washington State" 4795: 4050:. Saunders College Publishing. pp.  4037: 4035: 4033: 4031: 3188: 2963: 2610: 2608: 2184: 1863: 1806: 1739:, where absorption takes place, and the 1685: 1472:is never free-living and lacks a brain. 1447:, which is surrounded by the jelly-like 1391: 1381: 1266: 1229: 1093: 499: 486: 390:). The subphylum was at one time called 6136: 6004: 5564:Thaliacea (pyrosomes, salps, doliolids) 4269:Communicative & Integrative Biology 3937:"The Degenerate Tale of Ascidian Tails" 3427: 3334: 2848: 2550: 2479: 2477: 14: 7451: 6329:Echinodermata (starfish and relatives) 6299:Xenacoelomorpha (acoels and relatives) 5332:, with unusual nuclear architecture". 4986:. Korea-US Aquaculture. Archived from 2649: 1577:organ is a mass of large clear-walled 1219:them, they may have been mistaken for 567:Tunicates are more closely related to 437:(pyrosomes, doliolids, and salps) and 27:Marine animals, subphylum of chordates 7265: 7264: 6270: 5978: 5417: 4872: 4028: 3847:Fouling Organisms of the Indian Ocean 3661: 3540:Palmer, T. J.; Wilson, M. A. (1988). 2605: 2120:Invasive tunicates usually arrive as 1427:, but they do not develop very clear 6600:Acanthocephala (thorny-headed worms) 2474: 2243:The use of tunicates as a source of 7464:Cambrian Series 2 first appearances 4875:Stem cells : from hydra to man 4853: 4075:Schlosser, Gerhard (17 June 2021). 3941:Integrative and Comparative Biology 2692: 2058: 2009: 1763:. A few deepwater species, such as 1242:ancestor (before the divergence of 24: 6634: 6440: 6197:Cnidaria (jellyfish and relatives) 4319:Proceedings of the Royal Society B 2551:Giribet, Gonzalo (27 April 2018). 2162:, an FDA approved anticancer drug. 2105:have appeared and are thriving in 25: 7480: 7469:Extant Cambrian first appearances 5363: 4909:. Sciencedaily.com. 24 April 2007 4649:Molecular Genetics of Development 4473:. Little Brown & Co. p.  3624:"Introduction to the Urochordata" 2720:Bone, Q., and K. P. Ryan (1974). 2329:Journal of the Geological Society 1791:, is unique to tunicates. Excess 1653:Fluorescent-colored sea squirts, 1286: 1129:at Shankou village, Anning, near 562: 546:The Tunicata were established by 471:or stiffening rod and resemble a 7241:Phyla with ≥1000 extant species 5707:Actinopterygii (ray-finned fish) 4527:"Botryllus: A colonial ascidian" 3581:Nagalwade, Vidya (7 July 2023). 3335:Tokioka, Takasi (30 June 1971). 2629:10.1111/j.1463-6409.2011.00493.x 2530:10.1111/j.1463-6409.2012.00536.x 2494:World Register of Marine Species 2194:tunicates for sale at a market, 2135: 1664: 1646: 1635:Blue sea squirts from the genus 1628: 1611: 1475:Tunicates have a well-developed 1089: 1023: 1003: 976: 921: 900: 863: 843: 823: 760: 718: 378:which includes all animals with 370:). This grouping is part of the 319: 153: 56: 5321: 5002: 4976: 4921: 4899: 4866: 4847: 4825: 4541: 4518: 4491: 4458: 4421: 4356: 4305: 4256: 4195: 4150: 4095: 4068: 3979: 3928: 3879: 3834: 3795: 3729: 3702: 3655: 3642: 3616: 3574: 3533: 3421: 3370: 3328: 3179: 2988: 2941: 2929: 2917: 2906: 2797: 2713: 2140: 1705:Nearly all adult tunicates are 1561:Tunicates lack the kidney-like 1378:Physiology and internal anatomy 441:on the other hand, swim in the 6428:Nematomorpha (horsehair worms) 4777:"Have You Seen This Tunicate?" 4442:Cavanihac, Jean-Marie (2000). 2686: 2643: 2544: 2417: 2372: 2312: 1907:life history phase, an oozoid 13: 1: 6271: 5307:10.1126/science.294.5551.2506 4657:10.1016/S0065-2660(08)60012-7 4611:Biochem. Biophys. Res. Commun 4189:10.1016/S0022-0981(02)00023-0 3711:Journal of Biological Systems 2306: 2238: 2039:cell termed the "coenocyst". 1802: 1254:events among metazoan phyla. 417:, each unit being known as a 7040:Echinozoans inc. sea urchins 4873:Bosch, Thomas C. G. (2008). 4805:Dornfeld, Ann (1 May 2008). 3002:. Smithsonian at Fort Pierce 2949:"Sea squirts and sea tulips" 1812:Anatomy of a larval tunicate 1512:vanadium-associated proteins 1262: 1179:, and one from a second new 456:, sea pork, sea livers, or 7: 7097:Chelicerates inc. arachnids 6837:Phoronida (horseshoe worms) 6652:Platyhelminthes (flatworms) 6563:Gnathostomulida (jaw worms) 6452:Loricifera (corset animals) 5538:Appendicularia (larvaceans) 4835:. Woods Hole Science Center 4754:10.1016/j.ydbio.2006.10.021 4500:Canadian Journal of Zoology 4465:Dennett, Daniel C. (1991). 4108:Journal of Chemical Ecology 3820:10.1016/j.ydbio.2008.11.026 2283: 2254:Some tunicates are used as 2031:in which the entire female 1845:, and a balancing organ, a 482: 10: 7485: 6987:Medusozoans inc. jellyfish 6733:Annelida (segmented worms) 6558:Chaetognatha (arrow worms) 6474:Onychophora (velvet worms) 5965:denote paraphyletic groups 5844:Squamata (scaled reptiles) 5346:10.1016/j.gene.2007.03.006 4623:10.1016/j.bbrc.2014.05.099 4548:Holland, Linda Z. (2007). 4444:"Tunicates extraordinaire" 3841:Renganathan, T.K. (2020). 3763:10.1038/s41598-022-18283-9 2451:10.1007/s12542-021-00596-1 2181:Ascidiacea § Culinary 2178: 2174: 1928:protandrous hermaphrodites 1922:Larvaceans only reproduce 1901:alternation of generations 1681: 1257: 1137:). There is also a common 7273: 7237: 7035:Asterozoans inc. starfish 6937: 6930: 6881: 6832:Brachiopoda (lamp shells) 6822: 6813:Ectoprocta (moss animals) 6797: 6788: 6773: 6764: 6749: 6742: 6718: 6707: 6666: 6658:Gastrotricha (hairybacks) 6641: 6630: 6619: 6571: 6553: 6540: 6529: 6482: 6469: 6460: 6447: 6436: 6404: 6387:Kinorhyncha (mud dragons) 6373: 6362: 6347: 6323: 6314: 6294: 6279: 6266: 6231: 6206: 6182: 6167: 6159:Ctenophora (comb jellies) 6147: 6132: 6016: 5936: 5882:Crocodilia (crocodilians) 5853: 5838:Rhynchocephalia (tuatara) 5828: 5817: 5804: 5793: 5780: 5769: 5755: 5744: 5730: 5715: 5702: 5687: 5674: 5659: 5628: 5617: 5581: 5572: 5553: 5546: 5533: 5520: 5516: 5505: 5479: 5453: 5121:10.1007/s00239-003-2511-9 5060:10.1016/j.tig.2007.07.001 4833:"Marine Nuisance Species" 4128:10.1007/s10886-007-9251-z 3723:10.1142/S0218339010003408 3555:: 939–949. Archived from 3447:10.1186/s12915-018-0517-4 3428:Giribet, Gonzalo (2018). 3303:10.1186/s12915-018-0499-2 2709:– via Google Books. 2664:10.1007/s12064-018-0260-y 2572:10.1186/s12915-018-0517-4 2403:10.1134/S0031030112010042 1971:Promotion of out-crossing 1917:sequential hermaphrodites 1817:Ascidians are almost all 1751:opens into the dorsal or 1701:(orange) showing siphons. 1299:or tunic, from which the 1057: 1018: 998: 991: 971: 964: 954: 915: 895: 888: 858: 838: 818: 811: 801: 794: 755: 740: 733: 713: 706: 696: 306:is an exclusively marine 290: 283: 230: 222: 150:Scientific classification 148: 137: 128: 34: 6593:Rotifera (wheel animals) 6392:Priapulida (penis worms) 5943:Ascidiacea (sea squirts) 5735:Actinistia (coelacanths) 4525:Parmentier, Jan (1998). 4064:– via archive.org. 3855:10.1201/9781003077992-16 3396:10.3389/fimmu.2017.00674 3228:BMC Evolutionary Biology 3154:10.1186/1471-2164-10-534 2693:Gee, H. (27 July 2007). 2484:Sanamyan, Karen (2013). 1747:pellets or strings. The 1518:in blood cells known as 1252:horizontal gene transfer 1119:Shankouclava shankouense 552:Francis Maitland Balfour 310:animal, a member of the 6807:Entoprocta or Kamptozoa 6779:Nemertea (ribbon worms) 6500:Tardigrada (waterbears) 6493:Arthropoda (arthropods) 5493:Leptocardii (lancelets) 5406:Encyclopædia Britannica 5370:The Tunicate Web Portal 5257:10.1186/gb-2007-8-3-r41 5191:10.1126/science.1080049 5099:mitochondrial genome". 4934:The Biological Bulletin 4877:. Dordrecht: Springer. 4469:Consciousness Explained 4396:10.1073/pnas.0503640102 4231:10.1073/pnas.0303623101 3508:10.1073/pnas.1431177100 3383:Frontiers in Immunology 3249:10.1186/1471-2148-9-187 2382:Paleontological Journal 2302:– claimed hybridization 1169:from the Nama Group of 1098:The star-shaped holes ( 496:, the bluebell tunicate 138:Gold-mouth sea squirt ( 6982:Anthozoans inc. corals 6681:Dicyemida or Rhombozoa 5647:Hyperoartia (lampreys) 5409:(11th ed.). 1911. 4332:10.1098/rspb.2016.1712 2936:Tunicata Lamarck, 1816 2746:10.1098/rspb.1974.0077 2202: 1869: 1814: 1771:sit-and-wait predators 1702: 1695:(black and white) and 1400: 1389: 1272: 1113: 510: 506:Botrylloides violaceus 497: 445:of the sea as adults. 7418:Paleobiology Database 7112:Hexapods inc. insects 6422:Nematoda (roundworms) 5153:"The draft genome of 4809:. NPR. Archived from 3808:Developmental Biology 3698:– via CrossRef. 2272:developmental studies 2188: 2029:reproductive strategy 1987:inbreeding depression 1959:and vertebrates show 1867: 1810: 1689: 1620:Oikopleura cophocerca 1395: 1385: 1270: 1230:Hybridization studies 1176:Yarnemia ascidiformis 1097: 548:Jean-Baptiste Lamarck 503: 493:Clavelina moluccensis 490: 223:Classes and unplaced 6305:Chordata (chordates) 5863:Testudines (turtles) 4946:10.1086/BBLv221n1p43 3849:. pp. 507–534. 3562:on 27 September 2013 2890:Invertebrate Zoology 2300:Donald I. Williamson 1994:Botryllus schlosseri 1949:determinate cleavage 1909:reproduces asexually 1143:Catellocaula vallata 1101:Catellocaula vallata 540:Pyura pachydermatina 6755:Mollusca (molluscs) 6240:of the animal root 5957:Reptilia (reptiles) 5665:(jawed vertebrates) 5386:Halocynthia roretzi 5173:2002Sci...298.2157D 5167:(5601): 2157–2167. 5113:2003JMolE..57..574Y 5097:Halocynthia roretzi 4566:2007Natur.447..153H 4387:2005PNAS..10215134S 4381:(42): 15134–15139. 4222:2004PNAS..101..986M 4171:2002JEMBE.270..203P 4120:2007JCEco..33..643O 3954:10.1093/icb/icab022 3904:10.1098/rsob.150053 3755:2022NatSR..1214326D 3680:2018Palgy..61..575W 3499:2003PNAS..100.8314C 3240:2009BMCEE...9..187T 3053:10.1038/nature04336 3045:2006Natur.439..965D 2951:. Australian Museum 2826:10.1002/jez.b.22940 2818:2021JEZB..336..250A 2738:1974RSPSB.187..315B 2442:2021PalZ...95..623M 2395:2012PalJ...46....1F 2350:10.1144/jgs2017-103 2342:2018JGSoc.175..659Y 2270:have been used for 2229:Halocynthia roretzi 1766:Megalodicopia hians 1731:are secreted and a 1698:Pycnoclavella flava 1599:synthesizing enzyme 1558:against predators. 1407:, tunicates have a 1361:A third class, the 1154:Megasiphon thylakos 448:Various species of 6968:Calcareous sponges 6153:Porifera (sponges) 5809:Mammalia (mammals) 5721:(lobe-finned fish) 5693:(bony vertebrates) 5382:Ciona intestinalis 5155:Ciona intestinalis 5039:Ciona intestinalis 4367:Ciona intestinalis 4326:(1845): 20161712. 3743:Scientific Reports 3688:10.1111/pala.12356 2998:Aplidium stellatum 2261:Ciona intestinalis 2203: 2147:chemical compounds 1977:Ciona intestinalis 1961:cell determination 1870: 1815: 1707:suspension feeders 1703: 1565:organs typical of 1481:circulatory system 1401: 1390: 1273: 1114: 550:in 1816. In 1881, 511: 498: 421:. They are marine 380:dorsal nerve cords 7446: 7445: 7405:Open Tree of Life 7267:Taxon identifiers 7258: 7257: 7233: 7232: 6926: 6925: 6922: 6921: 6877: 6876: 6873: 6872: 6869: 6868: 6865: 6864: 6861: 6860: 6857: 6856: 6853: 6852: 6849: 6848: 6845: 6844: 6703: 6702: 6699: 6698: 6615: 6614: 6611: 6610: 6525: 6524: 6521: 6520: 6517: 6516: 6513: 6512: 6509: 6508: 6400: 6399: 6343: 6342: 6262: 6261: 6227: 6226: 6223: 6222: 5972: 5971: 5954:³orders of class 5940:¹orders of class 5932: 5931: 5928: 5927: 5924: 5923: 5920: 5919: 5916: 5915: 5912: 5911: 5908: 5907: 5904: 5903: 5900: 5899: 5896: 5895: 5760:Dipnoi (lungfish) 5655: 5654: 5613: 5612: 5609: 5608: 5605: 5604: 5601: 5600: 5501: 5500: 5330:Oikopleura dioica 5291:Oikopleura dioica 4783:on 9 January 2009 4706:10.1002/dvg.22802 4281:10.4161/cib.17757 4088:978-1-000-36913-7 4061:978-0-03-030504-7 4004:10.1111/brv.12189 3493:(14): 8314–8318. 3101:10.1002/dvg.20450 3039:(7079): 965–968. 2899:978-81-315-0104-7 2732:(1088): 315–327. 2617:Zoologica Scripta 2517:Zoologica Scripta 2277:Oikopleura dioica 2219:) is used in the 2122:fouling organisms 2115:Pacific Northwest 2082:Didemnum vexillum 2016:Oikopleura dioica 1933:Oikopleura dioica 1692:Clavelina robusta 1556:chemical defenses 1552:organic compounds 1310:connective tissue 1127:Maotianshan Shale 1086: 1085: 1077: 1076: 1068: 1067: 1054: 1053: 1045: 1044: 1036: 1035: 943: 942: 934: 933: 876: 875: 782: 781: 773: 772: 300: 299: 295: 218: 16:(Redirected from 7476: 7439: 7438: 7426: 7425: 7413: 7412: 7400: 7399: 7387: 7386: 7374: 7373: 7361: 7360: 7348: 7347: 7335: 7334: 7322: 7321: 7309: 7308: 7307: 7294: 7293: 7292: 7262: 7261: 6939: 6938: 6895: 6795: 6794: 6771: 6770: 6747: 6746: 6716: 6715: 6712: 6639: 6638: 6632: 6631: 6628: 6627: 6624: 6551: 6550: 6542: 6541: 6538: 6537: 6534: 6467: 6466: 6445: 6444: 6438: 6437: 6409: 6408: 6375: 6374: 6371: 6370: 6367: 6360: 6359: 6356: 6349: 6348: 6321: 6320: 6292: 6291: 6288: 6281: 6280: 6268: 6267: 6245: 6180: 6179: 6145: 6144: 6141: 6134: 6133: 6045: 6025: 5999: 5992: 5985: 5976: 5975: 5870: 5869: 5826: 5825: 5822: 5802: 5801: 5798: 5778: 5777: 5774: 5753: 5752: 5749: 5728: 5727: 5724: 5722: 5700: 5699: 5696: 5694: 5672: 5671: 5668: 5666: 5642:Myxini (hagfish) 5630: 5629: 5626: 5625: 5622: 5579: 5578: 5551: 5550: 5531: 5530: 5526: 5518: 5517: 5514: 5513: 5510: 5481: 5480: 5438: 5431: 5424: 5415: 5414: 5410: 5402: 5400:"Tunicata"  5358: 5357: 5325: 5319: 5318: 5286: 5280: 5279: 5269: 5259: 5231: 5225: 5224: 5218: 5210: 5184: 5147: 5141: 5140: 5088: 5082: 5081: 5071: 5045: 5032: 5026: 5025: 5023: 5021: 5006: 5000: 4999: 4997: 4995: 4980: 4974: 4973: 4925: 4919: 4918: 4916: 4914: 4903: 4897: 4896: 4870: 4864: 4863: 4860:Journal Sentinel 4851: 4845: 4844: 4842: 4840: 4829: 4823: 4822: 4820: 4818: 4802: 4793: 4792: 4790: 4788: 4773: 4767: 4766: 4756: 4732: 4726: 4725: 4688: 4679: 4678: 4644: 4635: 4634: 4605: 4596: 4595: 4577: 4545: 4539: 4538: 4536: 4534: 4522: 4516: 4515: 4495: 4489: 4488: 4484:978-0316-18065-8 4472: 4462: 4456: 4455: 4453: 4451: 4439: 4430: 4425: 4419: 4418: 4408: 4398: 4360: 4354: 4353: 4343: 4309: 4303: 4302: 4292: 4260: 4254: 4253: 4243: 4233: 4199: 4193: 4192: 4182: 4154: 4148: 4147: 4099: 4093: 4092: 4072: 4066: 4065: 4049: 4039: 4026: 4025: 4015: 3983: 3977: 3976: 3966: 3956: 3932: 3926: 3925: 3915: 3883: 3877: 3876: 3838: 3832: 3831: 3799: 3793: 3792: 3782: 3733: 3727: 3726: 3706: 3700: 3699: 3659: 3653: 3646: 3640: 3639: 3637: 3635: 3630:on 21 April 2009 3620: 3614: 3610: 3601: 3597: 3591: 3590: 3578: 3572: 3571: 3569: 3567: 3561: 3546: 3537: 3531: 3530: 3520: 3510: 3476: 3470: 3469: 3459: 3449: 3425: 3419: 3418: 3408: 3398: 3374: 3368: 3367: 3341: 3332: 3326: 3325: 3315: 3305: 3281: 3272: 3271: 3261: 3251: 3219: 3213: 3212: 3192: 3186: 3183: 3177: 3176: 3166: 3156: 3128: 3122: 3121: 3103: 3079: 3073: 3072: 3030: 3021: 3012: 3011: 3009: 3007: 2992: 2986: 2985: 2983: 2981: 2967: 2961: 2960: 2958: 2956: 2945: 2939: 2933: 2927: 2921: 2915: 2910: 2904: 2903: 2885: 2846: 2845: 2801: 2795: 2794: 2788: 2780: 2778: 2776: 2717: 2711: 2710: 2690: 2684: 2683: 2647: 2641: 2640: 2612: 2603: 2602: 2592: 2574: 2548: 2542: 2541: 2511: 2505: 2504: 2502: 2500: 2481: 2472: 2471: 2453: 2421: 2415: 2414: 2376: 2370: 2369: 2325: 2316: 2221:cuisine of Chile 2059:Invasive species 2010:A model tunicate 1965:genome evolution 1668: 1656:Rhopalaea crassa 1650: 1632: 1615: 1399: 1166:Ausia fenestrata 1027: 1007: 994: 993: 980: 967: 966: 957: 956: 925: 904: 891: 890: 867: 847: 827: 814: 813: 804: 803: 797: 796: 764: 736: 735: 722: 709: 708: 699: 698: 689: 688: 366: 362: 357: 356: 353: 352: 349: 346: 343: 340: 337: 334: 331: 328: 325: 293: 272: 262: 251: 213: 158: 157: 141:Polycarpa aurata 133: 118: 55: 41:Cambrian Stage 3 38:Temporal range: 32: 31: 21: 7484: 7483: 7479: 7478: 7477: 7475: 7474: 7473: 7449: 7448: 7447: 7442: 7434: 7429: 7421: 7416: 7408: 7403: 7395: 7390: 7382: 7377: 7369: 7364: 7356: 7351: 7343: 7338: 7330: 7325: 7317: 7312: 7303: 7302: 7297: 7288: 7287: 7282: 7269: 7259: 7254: 7229: 7155:Phylactolaemata 6943: 6933: 6932: 6918: 6909:Xenambulacraria 6885: 6841: 6818: 6784: 6766:Kryptotrochozoa 6760: 6738: 6708: 6695: 6686:Monoblastozoa ( 6662: 6620: 6607: 6579:Micrognathozoa 6567: 6530: 6505: 6478: 6456: 6432: 6396: 6363: 6352: 6339: 6310: 6284: 6275: 6258: 6235: 6219: 6202: 6171: 6163: 6137: 6128: 6127: 6043: 6021: 6012: 6003: 5973: 5968: 5948:²subclasses of 5892: 5868: 5849: 5818: 5813: 5794: 5789: 5770: 5765: 5745: 5740: 5720: 5719: 5716: 5711: 5692: 5691: 5688: 5683: 5664: 5663: 5660: 5651: 5618: 5597: 5592:Aplousobranchia 5568: 5558:Stolidobranchia 5542: 5524: 5523: 5506: 5497: 5485:Cephalochordata 5475: 5449: 5442: 5397: 5366: 5361: 5326: 5322: 5287: 5283: 5232: 5228: 5212: 5211: 5182:10.1.1.319.2643 5148: 5144: 5089: 5085: 5043: 5033: 5029: 5019: 5017: 5016:. 26 March 2013 5008: 5007: 5003: 4993: 4991: 4990:on 2 March 2013 4982: 4981: 4977: 4926: 4922: 4912: 4910: 4905: 4904: 4900: 4885: 4871: 4867: 4854:Johnson, Mark. 4852: 4848: 4838: 4836: 4831: 4830: 4826: 4816: 4814: 4813:on 14 July 2014 4803: 4796: 4786: 4784: 4775: 4774: 4770: 4733: 4729: 4689: 4682: 4667: 4645: 4638: 4606: 4599: 4575:10.1038/447153a 4546: 4542: 4532: 4530: 4529:. Microscope UK 4523: 4519: 4512:10.1139/z04-177 4496: 4492: 4485: 4463: 4459: 4449: 4447: 4446:. Microscope UK 4440: 4433: 4426: 4422: 4361: 4357: 4310: 4306: 4261: 4257: 4200: 4196: 4180:10.1.1.558.3639 4155: 4151: 4104:Phallusia nigra 4100: 4096: 4089: 4073: 4069: 4062: 4040: 4029: 3984: 3980: 3933: 3929: 3884: 3880: 3865: 3839: 3835: 3800: 3796: 3734: 3730: 3707: 3703: 3660: 3656: 3647: 3643: 3633: 3631: 3622: 3621: 3617: 3611: 3604: 3598: 3594: 3579: 3575: 3565: 3563: 3559: 3544: 3538: 3534: 3477: 3473: 3426: 3422: 3375: 3371: 3339: 3333: 3329: 3282: 3275: 3220: 3216: 3209:10.1139/z05-010 3193: 3189: 3184: 3180: 3135:Herdmania momus 3129: 3125: 3094:(11): 592–604. 3080: 3076: 3028: 3022: 3015: 3005: 3003: 2994: 2993: 2989: 2979: 2977: 2969: 2968: 2964: 2954: 2952: 2947: 2946: 2942: 2934: 2930: 2922: 2918: 2911: 2907: 2900: 2886: 2849: 2802: 2798: 2782: 2781: 2774: 2772: 2718: 2714: 2707: 2691: 2687: 2648: 2644: 2613: 2606: 2549: 2545: 2512: 2508: 2498: 2496: 2482: 2475: 2422: 2418: 2377: 2373: 2323: 2317: 2313: 2309: 2286: 2256:model organisms 2241: 2216:Pyura chilensis 2183: 2177: 2143: 2138: 2061: 2047:because of its 2012: 1982:C. intestinalis 1973: 1813: 1805: 1781:green algae or 1684: 1677: 1669: 1660: 1651: 1642: 1633: 1624: 1616: 1403:Like all other 1398:(br s) pharynx. 1397: 1388: 1380: 1289: 1265: 1260: 1236:molecular study 1232: 1221:sponge spicules 1173:, the sac-like 1122:from the Lower 1092: 1087: 1078: 1069: 1055: 1046: 1037: 961:Stolidobranchia 944: 935: 918:Aplousobranchia 877: 783: 774: 670:Members of the 636:Stolidobranchia 628:Aplousobranchia 565: 485: 465:Cambrian period 394:, and the term 364: 360: 322: 318: 294:Lankester, 1877 212: 152: 124: 123:record, 557 Ma) 117: 116: 111: 106: 101: 96: 91: 86: 81: 76: 71: 66: 61: 50: 49: 48: 39: 36: 28: 23: 22: 15: 12: 11: 5: 7482: 7472: 7471: 7466: 7461: 7444: 7443: 7441: 7440: 7427: 7414: 7401: 7388: 7375: 7362: 7349: 7336: 7323: 7310: 7295: 7279: 7277: 7271: 7270: 7256: 7255: 7253: 7252: 7245: 7238: 7235: 7234: 7231: 7230: 7228: 7227: 7226: 7225: 7220: 7215: 7210: 7205: 7193: 7192: 7191: 7186: 7181: 7169: 7168: 7167: 7162: 7157: 7145: 7144: 7143: 7138: 7133: 7128: 7120:Platyhelminths 7116: 7115: 7114: 7109: 7104: 7099: 7087: 7086: 7085: 7080: 7075: 7063: 7062: 7061: 7056: 7044: 7043: 7042: 7037: 7032: 7020: 7019: 7018: 7013: 7008: 6996: 6995: 6994: 6989: 6984: 6972: 6971: 6970: 6965: 6960: 6947: 6945: 6935: 6934: 6931: 6928: 6927: 6924: 6923: 6920: 6919: 6917: 6916: 6914:Centroneuralia 6911: 6906: 6901: 6896: 6882: 6879: 6878: 6875: 6874: 6871: 6870: 6867: 6866: 6863: 6862: 6859: 6858: 6855: 6854: 6851: 6850: 6847: 6846: 6843: 6842: 6840: 6839: 6834: 6828: 6826: 6820: 6819: 6817: 6816: 6809: 6803: 6801: 6792: 6786: 6785: 6783: 6782: 6774: 6768: 6762: 6761: 6759: 6758: 6750: 6744: 6740: 6739: 6737: 6736: 6729: 6719: 6713: 6710:Lophotrochozoa 6705: 6704: 6701: 6700: 6697: 6696: 6694: 6693: 6683: 6678: 6672: 6670: 6664: 6663: 6661: 6660: 6655: 6647: 6645: 6636: 6625: 6622:Platytrochozoa 6617: 6616: 6613: 6612: 6609: 6608: 6606: 6605: 6604: 6603: 6596: 6584: 6581:(Limnognathia) 6575: 6573: 6569: 6568: 6566: 6565: 6560: 6554: 6548: 6535: 6527: 6526: 6523: 6522: 6519: 6518: 6515: 6514: 6511: 6510: 6507: 6506: 6504: 6503: 6496: 6488: 6486: 6480: 6479: 6477: 6476: 6470: 6464: 6458: 6457: 6455: 6454: 6448: 6442: 6434: 6433: 6431: 6430: 6425: 6417: 6415: 6406: 6402: 6401: 6398: 6397: 6395: 6394: 6389: 6383: 6381: 6368: 6357: 6345: 6344: 6341: 6340: 6338: 6337: 6332: 6324: 6318: 6312: 6311: 6309: 6308: 6301: 6295: 6289: 6277: 6276: 6264: 6263: 6260: 6259: 6257: 6256: 6251: 6246: 6232: 6229: 6228: 6225: 6224: 6221: 6220: 6218: 6217: 6213: 6211: 6210:(Triploblasts) 6204: 6203: 6201: 6200: 6193: 6191:and relatives) 6183: 6177: 6165: 6164: 6162: 6161: 6156: 6148: 6142: 6130: 6129: 6126: 6125: 6124: 6123: 6122: 6121: 6118:Mesomycetozoea 6115: 6110: 6100: 6090: 6089: 6088: 6083: 6078: 6073: 6068: 6067: 6066: 6054:Diaphoretickes 6051: 6046: 6041: 6036: 6031: 6026: 6018: 6017: 6014: 6013: 6002: 6001: 5994: 5987: 5979: 5970: 5969: 5967: 5966: 5960: 5952: 5946: 5937: 5934: 5933: 5930: 5929: 5926: 5925: 5922: 5921: 5918: 5917: 5914: 5913: 5910: 5909: 5906: 5905: 5902: 5901: 5898: 5897: 5894: 5893: 5891: 5890: 5885: 5878: 5876: 5867: 5866: 5859: 5857: 5851: 5850: 5848: 5847: 5841: 5834: 5832: 5823: 5815: 5814: 5812: 5811: 5805: 5799: 5791: 5790: 5788: 5787: 5781: 5775: 5767: 5766: 5764: 5763: 5756: 5750: 5742: 5741: 5739: 5738: 5731: 5725: 5713: 5712: 5710: 5709: 5703: 5697: 5685: 5684: 5682: 5681: 5675: 5669: 5657: 5656: 5653: 5652: 5650: 5649: 5644: 5638: 5636: 5623: 5615: 5614: 5611: 5610: 5607: 5606: 5603: 5602: 5599: 5598: 5596: 5595: 5589: 5586:Phlebobranchia 5582: 5576: 5570: 5569: 5567: 5566: 5561: 5554: 5548: 5544: 5543: 5541: 5540: 5534: 5528: 5511: 5503: 5502: 5499: 5498: 5496: 5495: 5489: 5487: 5477: 5476: 5474: 5473: 5467: 5461: 5454: 5451: 5450: 5441: 5440: 5433: 5426: 5418: 5412: 5411: 5395: 5389: 5377: 5372: 5365: 5364:External links 5362: 5360: 5359: 5340:(1): 159–169. 5320: 5301:(5551): 2506. 5281: 5238:Ciona savignyi 5226: 5142: 5107:(5): 574–587. 5093:Ciona savignyi 5083: 5054:(9): 419–422. 5027: 5001: 4975: 4920: 4898: 4883: 4865: 4846: 4824: 4794: 4768: 4747:(2): 591–600. 4727: 4680: 4665: 4636: 4597: 4560:(1): 153–155. 4540: 4517: 4506:(1): 151–183. 4490: 4483: 4457: 4431: 4420: 4355: 4304: 4275:(6): 768–771. 4255: 4216:(4): 986–991. 4194: 4165:(2): 203–214. 4149: 4114:(3): 643–654. 4094: 4087: 4067: 4060: 4027: 3998:(3): 673–711. 3978: 3947:(2): 358–369. 3927: 3878: 3863: 3833: 3814:(2): 357–367. 3794: 3728: 3717:(2): 261–275. 3701: 3674:(4): 575–595. 3654: 3641: 3615: 3602: 3592: 3587:Tech Explorist 3573: 3532: 3471: 3420: 3369: 3356:10.5134/175655 3327: 3273: 3214: 3187: 3178: 3123: 3074: 3013: 2987: 2975:Dictionary.com 2962: 2940: 2928: 2916: 2905: 2898: 2847: 2812:(3): 250–266. 2796: 2712: 2705: 2685: 2642: 2623:(6): 603–612. 2604: 2543: 2524:(4): 435–436. 2506: 2473: 2436:(4): 623–639. 2416: 2371: 2336:(4): 659–666. 2310: 2308: 2305: 2304: 2303: 2297: 2285: 2282: 2267:Ciona savignyi 2240: 2237: 2179:Main article: 2176: 2173: 2164: 2163: 2157: 2142: 2139: 2137: 2134: 2102:Ciona savignyi 2060: 2057: 2045:model organism 2021:Appendicularia 2011: 2008: 1972: 1969: 1819:hermaphrodites 1811: 1804: 1801: 1793:photosynthetic 1683: 1680: 1679: 1678: 1673:Didemnum molle 1670: 1663: 1661: 1652: 1645: 1643: 1634: 1627: 1625: 1617: 1610: 1584:urate crystals 1563:metanephridial 1461:Appendicularia 1439:, gonads, and 1386: 1379: 1376: 1288: 1287:Body structure 1285: 1264: 1261: 1259: 1256: 1234:A multi-taxon 1231: 1228: 1190:Burykhia hunti 1091: 1088: 1084: 1083: 1080: 1079: 1075: 1074: 1071: 1070: 1066: 1065: 1062: 1061: 1056: 1052: 1051: 1048: 1047: 1043: 1042: 1039: 1038: 1034: 1033: 1030: 1029: 1017: 1014: 1013: 1010: 1009: 997: 992: 990: 987: 986: 983: 982: 970: 965: 963: 955: 953: 950: 949: 946: 945: 941: 940: 937: 936: 932: 931: 928: 927: 914: 911: 910: 907: 906: 898:Phlebobranchia 894: 889: 887: 883: 882: 879: 878: 874: 873: 870: 869: 857: 854: 853: 850: 849: 837: 834: 833: 830: 829: 817: 812: 810: 802: 800: 795: 793: 789: 788: 785: 784: 780: 779: 776: 775: 771: 770: 767: 766: 758:Fritillariidae 754: 751: 750: 747: 746: 743:Kowalevskiidae 739: 734: 732: 729: 728: 725: 724: 712: 707: 705: 703:Appendicularia 697: 695: 687: 668: 667: 660:Appendicularia 657: 639: 632:Phlebobranchia 564: 563:Classification 561: 525:and a few are 484: 481: 423:filter feeders 298: 297: 288: 287: 281: 280: 279: 278: 268: 258: 247: 245:Appendicularia 242: 237: 236:(polyphyletic) 228: 227: 220: 219: 207: 203: 202: 197: 190: 189: 184: 180: 179: 174: 170: 169: 164: 160: 159: 146: 145: 135: 134: 126: 125: 112: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 37: 26: 9: 6: 4: 3: 2: 7481: 7470: 7467: 7465: 7462: 7460: 7457: 7456: 7454: 7437: 7432: 7428: 7424: 7419: 7415: 7411: 7406: 7402: 7398: 7393: 7389: 7385: 7380: 7376: 7372: 7367: 7363: 7359: 7354: 7350: 7346: 7341: 7337: 7333: 7328: 7324: 7320: 7315: 7311: 7306: 7300: 7296: 7291: 7285: 7281: 7280: 7278: 7276: 7272: 7268: 7263: 7250: 7249:dubious phyla 7246: 7244: 7240: 7239: 7236: 7224: 7221: 7219: 7216: 7214: 7211: 7209: 7206: 7204: 7201: 7200: 7199: 7198: 7194: 7190: 7187: 7185: 7182: 7180: 7177: 7176: 7175: 7174: 7170: 7166: 7163: 7161: 7158: 7156: 7153: 7152: 7151: 7150: 7146: 7142: 7139: 7137: 7134: 7132: 7129: 7127: 7124: 7123: 7122: 7121: 7117: 7113: 7110: 7108: 7105: 7103: 7100: 7098: 7095: 7094: 7093: 7092: 7088: 7084: 7081: 7079: 7076: 7074: 7071: 7070: 7069: 7068: 7064: 7060: 7057: 7055: 7052: 7051: 7050: 7049: 7048:Hemichordates 7045: 7041: 7038: 7036: 7033: 7031: 7028: 7027: 7026: 7025: 7021: 7017: 7014: 7012: 7009: 7007: 7004: 7003: 7002: 7001: 6997: 6993: 6990: 6988: 6985: 6983: 6980: 6979: 6978: 6977: 6973: 6969: 6966: 6964: 6963:Glass sponges 6961: 6959: 6956: 6955: 6954: 6953: 6949: 6948: 6946: 6940: 6936: 6929: 6915: 6912: 6910: 6907: 6905: 6904:Deuterostomia 6902: 6900: 6897: 6893: 6890:of Bilateria 6889: 6884: 6883: 6880: 6838: 6835: 6833: 6830: 6829: 6827: 6825: 6821: 6815: 6814: 6810: 6808: 6805: 6804: 6802: 6800: 6796: 6793: 6791: 6787: 6781: 6780: 6776: 6775: 6772: 6769: 6767: 6763: 6757: 6756: 6752: 6751: 6748: 6745: 6741: 6735: 6734: 6730: 6728: 6726: 6723:Cycliophora ( 6721: 6720: 6717: 6714: 6711: 6706: 6691: 6689: 6684: 6682: 6679: 6677: 6674: 6673: 6671: 6669: 6665: 6659: 6656: 6654: 6653: 6649: 6648: 6646: 6644: 6640: 6637: 6633: 6629: 6626: 6623: 6618: 6602: 6601: 6597: 6595: 6594: 6590: 6589: 6588: 6585: 6583: 6582: 6577: 6576: 6574: 6570: 6564: 6561: 6559: 6556: 6555: 6552: 6549: 6547: 6543: 6539: 6536: 6533: 6528: 6502: 6501: 6497: 6495: 6494: 6490: 6489: 6487: 6485: 6481: 6475: 6472: 6471: 6468: 6465: 6463: 6462:Panarthropoda 6459: 6453: 6450: 6449: 6446: 6443: 6439: 6435: 6429: 6426: 6424: 6423: 6419: 6418: 6416: 6414: 6410: 6407: 6403: 6393: 6390: 6388: 6385: 6384: 6382: 6380: 6376: 6372: 6369: 6366: 6361: 6358: 6355: 6350: 6346: 6336: 6333: 6331: 6330: 6326: 6325: 6322: 6319: 6317: 6313: 6307: 6306: 6302: 6300: 6297: 6296: 6293: 6290: 6287: 6282: 6278: 6274: 6269: 6265: 6255: 6252: 6250: 6247: 6243: 6239: 6234: 6233: 6230: 6215: 6214: 6212: 6209: 6205: 6199: 6198: 6194: 6192: 6190: 6185: 6184: 6181: 6178: 6175: 6170: 6166: 6160: 6157: 6155: 6154: 6150: 6149: 6146: 6143: 6140: 6135: 6131: 6119: 6116: 6114: 6111: 6109: 6106: 6105: 6104: 6101: 6099: 6096: 6095: 6094: 6091: 6087: 6084: 6082: 6081:Stramenopiles 6079: 6077: 6074: 6072: 6069: 6065: 6062: 6061: 6060: 6057: 6056: 6055: 6052: 6050: 6047: 6044:(major groups 6042: 6040: 6037: 6035: 6032: 6030: 6027: 6024: 6020: 6019: 6015: 6011: 6008: 6000: 5995: 5993: 5988: 5986: 5981: 5980: 5977: 5964: 5961: 5959: 5958: 5953: 5951: 5950:Sarcopterygii 5947: 5945: 5944: 5939: 5938: 5935: 5889: 5886: 5883: 5880: 5879: 5877: 5875: 5871: 5864: 5861: 5860: 5858: 5856: 5855:Archelosauria 5852: 5845: 5842: 5839: 5836: 5835: 5833: 5831: 5827: 5824: 5821: 5816: 5810: 5807: 5806: 5803: 5800: 5797: 5792: 5786: 5783: 5782: 5779: 5776: 5773: 5768: 5761: 5758: 5757: 5754: 5751: 5748: 5743: 5736: 5733: 5732: 5729: 5726: 5723: 5718:Sarcopterygii 5714: 5708: 5705: 5704: 5701: 5698: 5695: 5686: 5680: 5677: 5676: 5673: 5670: 5667: 5662:Gnathostomata 5658: 5648: 5645: 5643: 5640: 5639: 5637: 5635: 5631: 5627: 5624: 5621: 5616: 5593: 5590: 5587: 5584: 5583: 5580: 5577: 5575: 5571: 5565: 5562: 5559: 5556: 5555: 5552: 5549: 5545: 5539: 5536: 5535: 5532: 5529: 5527: 5525:(Urochordata) 5519: 5515: 5512: 5509: 5504: 5494: 5491: 5490: 5488: 5486: 5482: 5478: 5472: 5471:Deuterostomia 5468: 5466: 5462: 5460: 5456: 5455: 5452: 5447: 5439: 5434: 5432: 5427: 5425: 5420: 5419: 5416: 5408: 5407: 5401: 5396: 5393: 5390: 5388: 5387: 5383: 5378: 5376: 5373: 5371: 5368: 5367: 5355: 5351: 5347: 5343: 5339: 5335: 5331: 5324: 5316: 5312: 5308: 5304: 5300: 5296: 5292: 5285: 5277: 5273: 5268: 5263: 5258: 5253: 5249: 5245: 5241: 5239: 5230: 5222: 5216: 5208: 5204: 5200: 5196: 5192: 5188: 5183: 5178: 5174: 5170: 5166: 5162: 5158: 5156: 5146: 5138: 5134: 5130: 5126: 5122: 5118: 5114: 5110: 5106: 5102: 5098: 5094: 5087: 5079: 5075: 5070: 5065: 5061: 5057: 5053: 5049: 5042: 5040: 5031: 5015: 5014:Cleantechnica 5011: 5005: 4989: 4985: 4979: 4971: 4967: 4963: 4959: 4955: 4951: 4947: 4943: 4939: 4935: 4931: 4924: 4908: 4902: 4894: 4890: 4886: 4884:9781402082740 4880: 4876: 4869: 4861: 4857: 4850: 4834: 4828: 4812: 4808: 4801: 4799: 4782: 4778: 4772: 4764: 4760: 4755: 4750: 4746: 4742: 4738: 4731: 4723: 4719: 4715: 4711: 4707: 4703: 4700:(1): 105–20. 4699: 4695: 4687: 4685: 4676: 4672: 4668: 4666:9780120176243 4662: 4658: 4654: 4650: 4643: 4641: 4632: 4628: 4624: 4620: 4617:(3): 1142–8. 4616: 4612: 4604: 4602: 4593: 4589: 4585: 4581: 4576: 4571: 4567: 4563: 4559: 4555: 4551: 4544: 4528: 4521: 4513: 4509: 4505: 4501: 4494: 4486: 4480: 4476: 4471: 4470: 4461: 4445: 4438: 4436: 4429: 4424: 4416: 4412: 4407: 4402: 4397: 4392: 4388: 4384: 4380: 4376: 4375: 4370: 4368: 4359: 4351: 4347: 4342: 4337: 4333: 4329: 4325: 4321: 4320: 4315: 4308: 4300: 4296: 4291: 4286: 4282: 4278: 4274: 4270: 4266: 4259: 4251: 4247: 4242: 4237: 4232: 4227: 4223: 4219: 4215: 4211: 4210: 4205: 4198: 4190: 4186: 4181: 4176: 4172: 4168: 4164: 4160: 4153: 4145: 4141: 4137: 4133: 4129: 4125: 4121: 4117: 4113: 4109: 4105: 4098: 4090: 4084: 4081:. CRC Press. 4080: 4079: 4071: 4063: 4057: 4053: 4048: 4047: 4038: 4036: 4034: 4032: 4023: 4019: 4014: 4009: 4005: 4001: 3997: 3993: 3989: 3982: 3974: 3970: 3965: 3960: 3955: 3950: 3946: 3942: 3938: 3931: 3923: 3919: 3914: 3909: 3905: 3901: 3898:(6): 150053. 3897: 3893: 3889: 3882: 3874: 3870: 3866: 3864:9781003077992 3860: 3856: 3852: 3848: 3844: 3837: 3829: 3825: 3821: 3817: 3813: 3809: 3805: 3798: 3790: 3786: 3781: 3776: 3772: 3768: 3764: 3760: 3756: 3752: 3748: 3744: 3740: 3732: 3724: 3720: 3716: 3712: 3705: 3697: 3693: 3689: 3685: 3681: 3677: 3673: 3669: 3668:Palaeontology 3665: 3658: 3651: 3645: 3629: 3625: 3619: 3609: 3607: 3596: 3588: 3584: 3577: 3558: 3554: 3550: 3549:Palaeontology 3543: 3536: 3528: 3524: 3519: 3514: 3509: 3504: 3500: 3496: 3492: 3488: 3487: 3482: 3475: 3467: 3463: 3458: 3453: 3448: 3443: 3439: 3435: 3431: 3424: 3416: 3412: 3407: 3402: 3397: 3392: 3388: 3384: 3380: 3373: 3365: 3361: 3357: 3353: 3349: 3345: 3338: 3331: 3323: 3319: 3314: 3309: 3304: 3299: 3295: 3291: 3287: 3280: 3278: 3269: 3265: 3260: 3255: 3250: 3245: 3241: 3237: 3233: 3229: 3225: 3218: 3210: 3206: 3202: 3198: 3191: 3182: 3174: 3170: 3165: 3160: 3155: 3150: 3146: 3142: 3138: 3136: 3127: 3119: 3115: 3111: 3107: 3102: 3097: 3093: 3089: 3085: 3078: 3070: 3066: 3062: 3058: 3054: 3050: 3046: 3042: 3038: 3034: 3027: 3020: 3018: 3001: 2999: 2991: 2976: 2972: 2966: 2950: 2944: 2937: 2932: 2925: 2920: 2914: 2909: 2901: 2895: 2891: 2884: 2882: 2880: 2878: 2876: 2874: 2872: 2870: 2868: 2866: 2864: 2862: 2860: 2858: 2856: 2854: 2852: 2843: 2839: 2835: 2831: 2827: 2823: 2819: 2815: 2811: 2807: 2800: 2792: 2786: 2771: 2767: 2763: 2759: 2755: 2751: 2747: 2743: 2739: 2735: 2731: 2727: 2723: 2716: 2708: 2706:9780585252728 2702: 2698: 2697: 2689: 2681: 2677: 2673: 2669: 2665: 2661: 2657: 2653: 2652:Theory Biosci 2646: 2638: 2634: 2630: 2626: 2622: 2618: 2611: 2609: 2600: 2596: 2591: 2586: 2582: 2578: 2573: 2568: 2564: 2560: 2559: 2554: 2547: 2539: 2535: 2531: 2527: 2523: 2519: 2518: 2510: 2495: 2491: 2487: 2480: 2478: 2469: 2465: 2461: 2457: 2452: 2447: 2443: 2439: 2435: 2431: 2427: 2420: 2412: 2408: 2404: 2400: 2396: 2392: 2388: 2384: 2383: 2375: 2367: 2363: 2359: 2355: 2351: 2347: 2343: 2339: 2335: 2331: 2330: 2322: 2315: 2311: 2301: 2298: 2295: 2294:deuterostomes 2291: 2288: 2287: 2281: 2279: 2278: 2273: 2269: 2268: 2263: 2262: 2257: 2252: 2250: 2246: 2236: 2234: 2230: 2226: 2225:sea pineapple 2222: 2218: 2217: 2212: 2208: 2201: 2197: 2193: 2192: 2187: 2182: 2172: 2169: 2161: 2158: 2155: 2152: 2151: 2150: 2149:, including: 2148: 2136:Use by humans 2133: 2131: 2127: 2126:ballast water 2123: 2118: 2116: 2112: 2108: 2104: 2103: 2098: 2097: 2092: 2088: 2084: 2083: 2078: 2074: 2073: 2068: 2067: 2056: 2054: 2050: 2046: 2042: 2038: 2037:multinucleate 2034: 2030: 2026: 2022: 2018: 2017: 2007: 2005: 2004:B. schlosseri 2000: 1999:B. schlosseri 1996: 1995: 1990: 1988: 1983: 1979: 1978: 1968: 1966: 1962: 1958: 1954: 1950: 1945: 1943: 1939: 1935: 1934: 1930:, except for 1929: 1925: 1920: 1918: 1914: 1910: 1906: 1902: 1897: 1895: 1891: 1887: 1883: 1879: 1874: 1866: 1862: 1858: 1856: 1855:metamorphosis 1850: 1848: 1844: 1840: 1836: 1832: 1828: 1824: 1820: 1809: 1800: 1798: 1794: 1790: 1789: 1784: 1783:cyanobacteria 1780: 1776: 1772: 1768: 1767: 1762: 1757: 1754: 1750: 1746: 1742: 1738: 1734: 1730: 1726: 1722: 1717: 1712: 1708: 1700: 1699: 1694: 1693: 1688: 1675: 1674: 1667: 1662: 1658: 1657: 1649: 1644: 1640: 1639: 1631: 1626: 1622: 1621: 1614: 1609: 1608: 1607: 1604: 1600: 1596: 1591: 1587: 1585: 1580: 1576: 1572: 1568: 1567:deuterostomes 1564: 1559: 1557: 1553: 1549: 1545: 1541: 1537: 1533: 1529: 1528:sulfuric acid 1525: 1521: 1517: 1513: 1509: 1505: 1501: 1497: 1492: 1490: 1486: 1482: 1478: 1473: 1470: 1466: 1462: 1457: 1452: 1450: 1446: 1442: 1438: 1434: 1430: 1426: 1422: 1418: 1414: 1410: 1406: 1394: 1384: 1375: 1372: 1368: 1364: 1359: 1356: 1352: 1347: 1343: 1339: 1334: 1332: 1327: 1323: 1319: 1318:blood vessels 1315: 1311: 1307: 1302: 1298: 1294: 1284: 1282: 1278: 1269: 1255: 1253: 1249: 1245: 1244:panarthropods 1241: 1237: 1227: 1224: 1222: 1216: 1214: 1210: 1206: 1205: 1200: 1196: 1192: 1191: 1186: 1182: 1178: 1177: 1172: 1168: 1167: 1162: 1157: 1155: 1151: 1148: 1144: 1140: 1139:bioimmuration 1136: 1132: 1128: 1125: 1121: 1120: 1111: 1107: 1106:bioimmuration 1103: 1102: 1096: 1090:Fossil record 1082: 1081: 1073: 1072: 1064: 1063: 1060: 1050: 1049: 1041: 1040: 1032: 1031: 1028: 1026: 1022: 1016: 1015: 1012: 1011: 1008: 1006: 1002: 996: 995: 989: 988: 985: 984: 981: 979: 975: 969: 968: 962: 959: 958: 952: 951: 948: 947: 939: 938: 930: 929: 926: 924: 919: 913: 912: 909: 908: 905: 903: 899: 893: 892: 885: 884: 881: 880: 872: 871: 868: 866: 862: 856: 855: 852: 851: 848: 846: 842: 836: 835: 832: 831: 828: 826: 822: 816: 815: 809: 806: 805: 799: 798: 791: 790: 787: 786: 778: 777: 769: 768: 765: 763: 759: 753: 752: 749: 748: 745: 744: 738: 737: 731: 730: 727: 726: 723: 721: 717: 716:Oikopleuridae 711: 710: 704: 701: 700: 694: 691: 690: 686: 683: 681: 677: 673: 665: 661: 658: 655: 651: 647: 643: 640: 637: 633: 629: 625: 622: 621: 620: 617: 615: 611: 606: 604: 603:invertebrates 600: 599: 594: 593:hemichordates 590: 586: 582: 578: 574: 570: 560: 556: 553: 549: 544: 542: 541: 536: 532: 528: 524: 520: 516: 508: 507: 502: 495: 494: 489: 480: 478: 474: 470: 466: 461: 459: 455: 451: 446: 444: 440: 436: 432: 428: 424: 420: 416: 412: 407: 405: 401: 397: 393: 389: 385: 381: 377: 373: 369: 368: 355: 316: 313: 309: 305: 296: 289: 286: 282: 277: 276: 269: 267: 266: 259: 256: 255: 248: 246: 243: 241: 238: 235: 232: 231: 229: 226: 221: 216: 211: 208: 205: 204: 201: 198: 195: 192: 191: 188: 185: 182: 181: 178: 175: 172: 171: 168: 165: 162: 161: 156: 151: 147: 143: 142: 136: 132: 127: 122: 115: 110: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 54: 46: 42: 33: 30: 19: 7274: 7247:Potentially 7242: 7195: 7171: 7165:Gymnolaemata 7160:Stenolaemata 7149:Ectoproctans 7147: 7118: 7089: 7065: 7059:Pterobranchs 7046: 7022: 7010: 6998: 6974: 6950: 6944:within phyla 6942:Major groups 6812: 6799:Bryozoa s.l. 6790:Lophophorata 6778: 6754: 6732: 6724: 6687: 6676:Orthonectida 6651: 6598: 6591: 6580: 6499: 6492: 6421: 6379:Scalidophora 6328: 6316:Ambulacraria 6303: 6216:(see below↓) 6196: 6188: 6152: 6103:Opisthokonta 5962: 5955: 5941: 5888:Aves (birds) 5830:Lepidosauria 5690:Euteleostomi 5634:Cyclostomata 5521: 5469:Superphylum 5404: 5385: 5381: 5337: 5333: 5329: 5323: 5298: 5294: 5290: 5284: 5247: 5243: 5237: 5229: 5215:cite journal 5164: 5160: 5154: 5145: 5104: 5101:J. Mol. Evol 5100: 5096: 5092: 5086: 5051: 5048:Trends Genet 5047: 5038: 5030: 5018:. Retrieved 5013: 5004: 4992:. Retrieved 4988:the original 4984:"Sea squirt" 4978: 4940:(1): 43–61. 4937: 4933: 4923: 4911:. Retrieved 4901: 4874: 4868: 4859: 4849: 4837:. Retrieved 4827: 4815:. Retrieved 4811:the original 4785:. Retrieved 4781:the original 4771: 4744: 4740: 4730: 4697: 4693: 4648: 4614: 4610: 4557: 4553: 4543: 4531:. Retrieved 4520: 4503: 4499: 4493: 4468: 4460: 4448:. Retrieved 4423: 4378: 4372: 4366: 4358: 4323: 4317: 4307: 4272: 4268: 4258: 4213: 4207: 4197: 4162: 4158: 4152: 4111: 4107: 4103: 4097: 4077: 4070: 4045: 3995: 3991: 3981: 3944: 3940: 3930: 3895: 3892:Open Biology 3891: 3881: 3846: 3836: 3811: 3807: 3797: 3749:(1): 14326. 3746: 3742: 3731: 3714: 3710: 3704: 3671: 3667: 3657: 3644: 3632:. Retrieved 3628:the original 3618: 3595: 3586: 3576: 3564:. Retrieved 3557:the original 3552: 3548: 3535: 3490: 3484: 3474: 3437: 3433: 3423: 3386: 3382: 3372: 3347: 3343: 3330: 3293: 3289: 3231: 3227: 3217: 3200: 3197:Can. J. Zool 3196: 3190: 3181: 3144: 3141:BMC Genomics 3140: 3134: 3126: 3091: 3087: 3077: 3036: 3032: 3006:25 September 3004:. Retrieved 2997: 2990: 2980:25 September 2978:. Retrieved 2974: 2971:"Sea squirt" 2965: 2955:25 September 2953:. Retrieved 2943: 2931: 2919: 2908: 2889: 2809: 2805: 2799: 2785:cite journal 2773:. Retrieved 2729: 2725: 2715: 2695: 2688: 2655: 2651: 2645: 2620: 2616: 2562: 2556: 2546: 2521: 2515: 2509: 2497:. 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Index

Tunicata
Cambrian Stage 3
Present
Ma
PreꞒ

O
S
D
C
P
T
J
K
Pg
N
Ediacaran

Polycarpa aurata
Scientific classification
Edit this classification
Eukaryota
Animalia
Chordata
Olfactores
Tunicata
Lamarck
genera
Ascidiacea
Thaliacea

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