Knowledge (XXG)

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).
1602: 1637: 1846:, 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. 1272:, 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. 1995:. 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. 490: 751: 1885:, 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. 834: 477: 1655: 144: 912: 2240:, 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. 1929:, 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 1676: 1014: 1969:(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 967: 814: 891: 1619: 544:
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.
709: 120: 994: 1257: 2117:. 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 1797: 1382: 1084: 1313:. 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 1854: 854: 2175: 1372: 1849:
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
1818:. 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 1744:
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.
1480:. 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. 1460:
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
1908:. 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. 1317:
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
1297:, 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 4597:
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".
1904:. 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 1346:
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
2224:. 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. 1586:. 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 671:
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
456:. 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 2368:
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
1329:, 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 2683: 1762:, trapping tiny crustacea, nematodes, and other small invertebrates with the muscular lobes which surround their buccal siphons. Certain tropical species in the family 5209: 2779: 4362: 4197: 3474: 1636: 7380: 1511:. 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 1318:
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
1472:. 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 7452: 4844: 5394: 2269:
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.
7457: 1268:, integrate with one another. In the simplest systems, the individual animals are widely separated, but linked together by horizontal connections called 667:
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
1716:. 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 4998: 2793:
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
1358:. The 70 species of larvaceans superficially resemble the tadpole larvae of amphibians, although the tail is at right angles to the body. The 4075: 4048: 2886: 3571: 4471: 4065: 1986:(class Ascidiacea) is a colonial tunicate, a member of the only group of chordates that are able to reproduce both sexually and asexually. 1354:(or Appendicularia), is the only group of tunicates to retain their chordate characteristics in the adult state, a product of extensive 2145:, a didemnin effective against various types of cancer; as of late January 2021 undergoing Phase III trials as a treatment for COVID-19 5985: 5424: 4307: 4871: 4845:"International team of scientists identifies new treatment for COVID-19 that appears to be far more effective than drugs in use now" 4653: 3851: 3612: 2693: 2317: 1322:
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
464:. Their name derives from their unique outer covering or "tunic", which is formed from proteins and carbohydrates, and acts as an 7419: 6022: 2482: 506:; fewer than 100 species of these are found at depths greater than 200 m (660 ft). Some are solitary animals leading a 6017: 3698:
Syvanen, M.; Ducore, J. (2010). "Whole genome comparisons reveals a possible chimeric origin for a major metazoan assemblage".
1567: 1476:. The blood may appear pale green, but this is not due to any respiratory pigments, and oxygen is transported dissolved in the 4254:"Is there intracellular cellulose in the appendicularian tail epidermis? A tale of the adult tail of an invertebrate chordate" 3174:
Jefferies, R. P. S. (1991) in Biological Asymmetry and Handedness (eds Bock, G. R.; Marsh, J.) pp. 94–127 (Wiley, Chichester).
3831: 7372: 2924: 1523:, although it is unclear whether this is due to the presence of the metal or low pH. Other species of tunicates concentrate 502:
About 3,000 species of tunicate exist in the world's oceans, living mostly in shallow water. The most numerous group is the
6101: 6091: 6074: 1519:. The vanadocytes are later deposited just below the outer surface of the tunic, where their presence is thought to deter 1618: 5317:
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).
1881:. Some of these are known as trophozooids, have a nutritional function, and are arranged in lateral rows. Others are 1562:
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.
1654: 4765: 4726:"The Oikopleura coenocyst, a unique chordate germ cell permitting rapid, extensive modulation of oocyte production" 3641:
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.
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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
1712:. The net is rolled up on the dorsal side of the pharynx, and it and the trapped particles are drawn into the 4091:
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
468:. In some species, it is thin, translucent, and gelatinous, while in others it is thick, tough, and stiff. 7255: 5978: 5417: 5165: 4487:
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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Ostrovsky, A. N.; Lidgard, S.; Gordon, D. P.; Schwaha, T.; Genikhovich, G.; Ereskovsky, A. V. (2015).
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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
1145:, shows that the tunicate basic body design had already been established 500 million years ago. 608:
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).
7398: 6011: 5330: 5291: 5250: 5240: 5175: 5105: 5052: 5044: 4930: 4737: 4690: 4641: 4607: 4558: 4496: 4389: 4379: 4324: 4316: 4273: 4265: 4224: 4214: 4173: 4112: 3996: 3988: 3947: 3937: 3896: 3888: 3877:"Tunicates: Exploring the sea shores and roaming the open ocean. A tribute to Thomas Huxley" 3839: 3804: 3763: 3747: 3707: 3672: 3501: 3491: 3440: 3430: 3389: 3379: 3340: 3296: 3286: 3242: 3232: 3213:"An updated 18S rRNA phylogeny of tunicates based on mixture and secondary structure models" 3193: 3147: 3137: 3084: 3037: 2810: 2730: 2648: 2613: 2573: 2555: 2514: 2434: 2387: 2334: 2209: 2065: 1953: 1644: 1544: 1540: 911: 511: 403: 387:
is still sometimes used for these animals. They are the only chordates that have lost their
309: 129: 29: 7237: 7143: 6994: 6897: 6754: 6287: 5826: 5580: 5546: 5473: 5084:(Urochordata, Ascidiacea, Enterogona): Comparison of gene arrangement and tRNA genes with 4636:
Bernstein, H; Hopf, FA; Michod, RE (1987). "The Molecular Basis of the Evolution of Sex".
4303:"Transcriptional regulation of a horizontally transferred gene from bacterium to chordate" 2711:"On the Structure and Innervation of the Muscle Bands of Doliolum (Tunicata: Cyclomyaria)" 2204: 1759: 1334: 949: 906: 674:
The following cladogram is based on the 2018 phylogenomic study of Delsuc and colleagues.
624: 616: 453: 3652: 3616: 1575:, and do not have any obvious means of disposing of the material during their lifetimes. 1448:
developing into a barrel-like and usually sedentary adult form. The species in the class
5388: 5161: 5101: 4554: 4375: 4210: 4159: 4108: 3952: 3925: 3743: 3668: 3487: 3228: 3033: 2984: 2806: 2726: 2430: 2383: 2330: 422:, immobile and permanently attached to rocks or other hard surfaces on the ocean floor. 7293: 6902: 6698: 6610: 6588: 6551: 6042: 5695: 5667: 5574: 5255: 5224: 4896:"Sea Squirt, Heal Thyself: Scientists Make Major Breakthrough in Regenerative Medicine" 4394: 4353: 4329: 4302: 4278: 4253: 4040: 4001: 3976: 3901: 3876: 3768: 3727: 3445: 3418: 3394: 3367: 3301: 3274: 3247: 3212: 3152: 3121: 2937: 2578: 2541: 2255: 2244: 2090: 2033: 1785: 1781: 1661: 1285: 1227:
in 2010 proposed that sea squirts are descended from a hybrid between a chordate and a
1209: 1154: 886: 746: 731: 620: 391:
segmentation, with the possible exception of the seriation of the gill slits. However,
4645: 4229: 4192: 4177: 3506: 3469: 3419:"Phylogenomics resolves the evolutionary chronicle of our squirting closest relatives" 2542:"Phylogenomics resolves the evolutionary chronicle of our squirting closest relatives" 1675: 1013: 7441: 7047: 6951: 6450: 5938: 5843: 5706: 5650: 4710: 4417:
Response of Marine Ecosystems to Global Change: Ecological Impact of Appendicularians
3861: 3684: 3106: 2902: 2830: 2617: 2518: 2456: 2399: 2354: 2213: 2114: 2025: 1843: 1807: 1771: 1516: 704: 664: 411: 72: 5195: 5125: 4958: 4917:
Kürn, Ulrich; Rendulic, Snjezana; Tiozzo, Stefano; Lauzon, Robert J. (August 2011).
3352: 3015:"Tunicates and not cephalochordates are the closest living relatives of vertebrates" 2959: 2758: 2625: 2526: 2415:"New Ediacaran fossils from the Ukraine, some with a putative tunicate relationship" 452:
The earliest probable species of tunicate appears in the fossil record in the early
7320: 7153: 7148: 7036: 6892: 6778: 6664: 6567: 6546: 6416: 6367: 6323: 6304: 5818: 5773: 5678: 5622: 5459: 4580: 3653:"The first tunicate with a calcareous exoskeleton (Upper Triassic, northern Italy)" 3072: 3057: 2668: 2282: 2236:
is being researched. The cellulose body wall can be broken down and converted into
2084: 2075: 2037: 1874: 1555: 1515:
of less than 2 for stability, and this is achieved by the vacuoles also containing
1492: 1477: 1453: 1306: 1232: 1141:
skeletons of the upper midwestern United States. A well-preserved Cambrian fossil,
1107: 966: 813: 668: 591: 586: 581: 431: 296: 263: 5295: 4132: 890: 4999:"Biofuel made from marine filter feeders? Tunicates usable as source of biofuels" 2308:
Yang, Chuan; Li, Xian-Hua; Zhu, Maoyan; Condon, Daniel J.; Chen, Junyuan (2018).
1539:, which may serve a similar function. Other tunicate species produce distasteful 7354: 7307: 7287: 7114: 7085: 7071: 7061: 6646: 6462: 6375: 6157: 5735: 5635: 4742: 4725: 3808: 2546: 2278: 2188: 2179: 2148: 2142: 2118: 2095: 2041: 2013: 1749: 1401: 708: 465: 376: 47: 5334: 4611: 3751: 3572:"A 500 million-year-old fossil reveals the amazing secrets of tunicate origins" 3325: 2439: 2414: 7211: 7196: 7191: 7172: 7167: 7095: 7042: 7012: 7004: 6946: 6820: 6812: 6795: 6575: 6487: 6440: 6380: 6342: 6316: 6162: 6147: 5931: 5870: 5723: 5608: 5562: 5496: 5109: 5048: 4116: 4033: 3711: 3435: 3291: 2652: 2560: 2391: 2195: 2169: 2099: 2074:) has taken over a 6.5 sq mi (17 km) area of the seabed on the 2060: 1926: 1866: 1776: 1763: 1590:, encoded by a gene horizontally transferred from a bacterium. When, in 1845, 1551: 1508: 1488: 1473: 1437: 1433: 1281: 1239:). This study was based on a quartet partitioning approach designed to reveal 1228: 1135: 1047: 993: 962: 809: 660: 612: 602: 577: 569: 442: 438: 253: 222: 188: 119: 92: 57: 7278: 4942: 4881: 3843: 3759: 3384: 3275:"A phylogenomic framework and timescale for comparative studies of tunicates" 3142: 2795:
Journal of Experimental Zoology Part B: Molecular and Developmental Evolution
2569: 2448: 2346: 7341: 7177: 7124: 7119: 7090: 7079: 7028: 7023: 6975: 6887: 6880: 6674: 6669: 6631: 6480: 6472: 6401: 6353: 6274: 6261: 6242: 6237: 6196: 5862: 5552: 5526: 5453: 5245: 5179: 4384: 4219: 3496: 3237: 2221: 2021: 2009: 1945: 1940:, where the fate of the cells is set early on with reduced cell numbers and 1893: 1861: 1835: 1823: 1767: 1725: 1713: 1626: 1583: 1572: 1520: 1457: 1397: 1359: 1339: 1326: 1314: 1289: 1256: 1149: 989: 849: 796: 691: 648: 638: 630: 503: 457: 446: 427: 423: 392: 372: 300: 233: 228: 155: 109: 97: 41: 5342: 5303: 5264: 5187: 5117: 5066: 4950: 4751: 4702: 4619: 4572: 4403: 4338: 4320: 4287: 4238: 4124: 4010: 3961: 3910: 3816: 3777: 3515: 3454: 3403: 3310: 3256: 3161: 3098: 3049: 2822: 2734: 2660: 2587: 2310:"Geochronological constraint on the Cambrian Chengjiang biota, South China" 5402: 4663: 3942: 2750: 1796: 1381: 1093:) in this Upper Ordovician bryozoan may represent a tunicate preserved by 1083: 7272: 7201: 7185: 7108: 7100: 7055: 7018: 6988: 6970: 6964: 6825: 6766: 6742: 6639: 6520: 6409: 6293: 6184: 6175: 6081: 5832: 5760: 5748: 5481: 5434: 5380: 4934: 2715:
Proceedings of the Royal Society of London. Series B, Biological Sciences
2054: 2032:
can be maintained in laboratory culture, and is of growing interest as a
2016:
organism, reproducing only once in its lifetime. It employs an original
1870: 1819: 1815: 1709: 1699: 1587: 1536: 1500: 1496: 1449: 1413: 1409: 1405: 1393: 1351: 1330: 1236: 1192: 1178: 1164: 1112: 1009: 652: 634: 573: 557: 523: 360: 242: 175: 87: 82: 67: 62: 52: 33: 5080:
Yokobori, S.; Watanabe, Y.; Oshima, T. (2003). "Mitochondrial genome of
4769: 3892: 3041: 2814: 2338: 2220:) is the main species eaten. It is cultivated on dangling cords made of 1853: 853: 7161: 7137: 7129: 7066: 6980: 6800: 6787: 6720: 6711: 6656: 6581: 5945: 5784: 5630: 3676: 2237: 2233: 1952:
relatively late in development and cell cleavage is indeterminate. The
1831: 1827: 1721: 1559: 1532: 1524: 1504: 1443:
Ascidian tunicates begin life as a lecithotrophic (non-feeding) mobile
1429: 1355: 1343: 1294: 1159: 1138: 1119: 1098: 829: 565: 561: 515: 461: 399: 388: 102: 77: 7359: 5963: 5383:(1901), British Museum (Natural History). Department of Zoology et al. 5225:"A haplome alignment and reference sequence of the highly polymorphic 5057: 4694: 4269: 3992: 3089: 2174: 1830:. These swim by undulations of the tail and may have a simple eye, an 1456:, and the general larval form is kept throughout life. Also the class 7206: 6940: 6230: 6140: 6127: 5998: 5995: 5851: 5808: 5797: 5447: 4799: 3793:"Delineating metamorphic pathways in the ascidian Ciona intestinalis" 2742: 1941: 1930: 1901: 1741: 1729: 1717: 1704: 1563: 1417: 1302: 1269: 1265: 1173: 1127: 1094: 518:. Some are supported by a stalk, but most are attached directly to a 398:
Some tunicates live as solitary individuals, but others replicate by
364: 165: 7249: 4563: 4538: 4500: 3531:"Parasitism of Ordovician bryozoans and the origin of pseudoborings" 3197: 2710: 1432:(or branchial sac), the organs are enclosed in a membrane called an 1371: 7346: 5364:
Dutch Ascidians: Extensive database of images from around the world
3344: 3071:
Delsuc, F.; Tsagkogeorga, G.; Lartillot, N.; Philippe, H. (2008).
2184: 2173: 1852: 1811: 1795: 1733: 1674: 1558:. Most have no excretory structures, but rely on the diffusion of 1484: 1465: 1444: 1425: 1370: 1310: 1255: 1082: 598: 488: 475: 407: 213: 7333: 3615:. University of California Museum of Paleontology. Archived from 5876: 5774:
Lissamphibia (modern amphibians: frogs, salamanders, caecilians)
3126:
mitochondrial genome and support for the new chordate phylogeny"
1737: 1528: 642: 441:, the most well-known class of tunicates, are commonly known as 7253: 6259: 5967: 5406: 5387: 3977:"Matrotrophy and placentation in invertebrates: A new paradigm" 3602:(Vendian) biota. International Geological Congress - Oslo 2008. 3122:"Tunicate mitogenomics and phylogenetics: peculiarities of the 3013:
Delsuc, F.; Brinkmann, H.; Chourrout, D.; Philippe, H. (2006).
2113:
on the hulls of ships, but may also be introduced as larvae in
5368: 4724:
Ganot P, Bouquet JM, Kallesøe T, Thompson EM (February 2007).
2927:
Integrated Taxonomic Information System. Retrieved 2017-03-30.
2503:
Nielsen, C. (2012). "The authorship of higher chordate taxa".
3073:"Additional molecular support for the new chordate phylogeny" 318: 5668:
Chondrichthyes (cartilaginous fish: sharks, rays, chimaeras)
4919:"Asexual Propagation and Regeneration in Colonial Ascidians" 2052:
Over the past few decades, tunicates (notably of the genera
1933:
to develop into a house-building juvenile starting to feed.
1495:, it contains high concentrations of the transitional metal 1424:, renal, and gonadal cavities of the adults. Except for the 5369:
Aniseed: A model organism database for ascidians including
3470:"The first tunicate from the Early Cambrian of South China" 2685:
Before the Backbone: Views on the origin of the vertebrates
1512: 1260:
Colonial tunicate with multiple openings in a single tunic.
339: 333: 315: 4768:. NOAA Fisheries Service. 19 November 2004. Archived from 5223:
Small, K. S.; Brudno, M.; Hill, M. M.; Sidow, A. (2007).
5024:
Iannelli, F.; Pesole, G.; Sordino, P.; Gissi, C. (2007).
4193:"A functional cellulose synthase from ascidian epidermis" 3981:
Biological Reviews of the Cambridge Philosophical Society
2212:, both raw and in seafood stews. In Japan and Korea, the 324: 5363: 4301:
Sasakura, Y.; Ogura, Y.; Treen, N.; et al. (2016).
532:, which can grow to be over 1 metre (3.3 ft) tall. 4640:. Advances in Genetics. Vol. 24. pp. 323–70. 4426: 4424: 2915:
World Register of Marine Species. Retrieved 2011-11-12.
2285:
which are possibly the sister group of modern tunicates
2068:
coastal waters in many countries. The carpet tunicate (
1857:
A 1901 comparison of frog tadpole and a tunicate larva.
1708:
diameter which can trap planktonic particles including
3651:
Wendt, Jobst (25 July 2018). Hautmann, Michael (ed.).
3368:"Immunity in Protochordates: The Tunicate Perspective" 1956:
of amphioxus and vertebrates is also relatively slow.
548:
resemblance of their dead colonies to pieces of meat.
4539:"Developmental biology: A chordate with a difference" 4031:
Dorit, R.L.; Walker, W.F. & Barnes, R.D. (1991).
3728:"New putative phenol oxidase in ascidian blood cells" 3333:
Publications of the Seto Marine Biological Laboratory
2413:
Martyshyn, Andrej; Uchman, Alfred (1 December 2021).
348: 330: 321: 312: 4252:
Hirose, E.; Nakashima, K. & Nishino, A. (2011).
2881:(7th ed.). Cengage Learning. pp. 940–956. 2877:
Ruppert, E.E.; Fox, R.S. & Barnes, R.D. (2004).
2024:
is contained within an ovary that is a single giant
498:
showing oral tentacles at openings of buccal siphons
336: 259: 249: 238: 7262: 6930: 6811: 6786: 6777: 6753: 6731: 6696: 6655: 6630: 6623: 6608: 6560: 6533: 6518: 6471: 6449: 6429: 6400: 6393: 6366: 6351: 6340: 6303: 6272: 6195: 6156: 6125: 5861: 5842: 5817: 5806: 5782: 5758: 5733: 5704: 5676: 5648: 5621: 5606: 5561: 5535: 5509: 5494: 5472: 2198:species are consumed as food around the world. The 1869:, and at first, the developing embryo feeds on its 1698:(the larval form usually does not feed), capturing 663:were included in Ascidiacea in 2011 as a result of 327: 4455: 4148:Journal of Experimental Marine Biology and Ecology 4032: 1826:(stiffening rod) and superficially resemble small 601:consisting of tunicates and vertebrates is called 1172:-like genus from the Onega Peninsula of northern 1148:Three enigmatic species were also found from the 510:existence attached to the seabed, but others are 5146:: insights into chordate and vertebrate origins" 1936:During embryonic development, tunicates exhibit 1284:. The body of an ascidiacean is surrounded by a 395:still display segmentation of the muscle bands. 4675: 4673: 4592: 4590: 4363:Proceedings of the National Academy of Sciences 4198:Proceedings of the National Academy of Sciences 3475:Proceedings of the National Academy of Sciences 3008: 3006: 2134:Tunicates contain a host of potentially useful 1748:Some ascidians that live on soft sediments are 4631: 4629: 1487:has some unusual features. In some species of 1204:, so this identification has been questioned. 522:, which may be a rock, shell, coral, seaweed, 5979: 5418: 5026:"Mitogenomics reveals two cryptic species in 3268: 3266: 1774:in their tunics, and one of these symbionts, 1280:By far the largest class of tunicates is the 8: 5208:: CS1 maint: multiple names: authors list ( 3597: 3595: 2778:: CS1 maint: multiple names: authors list ( 2040:position within the closest sister group to 1944:that are rapidly evolving. In contrast, the 1888:Salps also have a complex lifecycle with an 1784:products are assumed to be available to the 4789: 4787: 3366:Franchi, Nicola; Ballarin, Loriano (2017). 2872: 2870: 2868: 2866: 2864: 2862: 2860: 1732:, where undigested remains are formed into 7250: 6927: 6783: 6759: 6735: 6704: 6627: 6620: 6616: 6539: 6530: 6526: 6455: 6433: 6426: 6397: 6363: 6359: 6348: 6337: 6309: 6280: 6269: 6256: 6168: 6133: 6122: 5986: 5972: 5964: 5858: 5814: 5790: 5766: 5741: 5716: 5688: 5660: 5618: 5614: 5567: 5539: 5519: 5506: 5502: 5469: 5425: 5411: 5403: 3875:Lemaire, Patrick; Piette, Jacques (2015). 3326:"Phylogenetic Speculation of the Tunicata" 2858: 2856: 2854: 2852: 2850: 2848: 2846: 2844: 2842: 2840: 1376:Internal anatomy of a generalised tunicate 118: 20: 5254: 5244: 5169: 5056: 4741: 4562: 4393: 4383: 4328: 4277: 4228: 4218: 4167: 4000: 3951: 3941: 3900: 3767: 3505: 3495: 3444: 3434: 3393: 3383: 3300: 3290: 3246: 3236: 3151: 3141: 3088: 2688:. Springer Science & Business Media. 2577: 2559: 2438: 1134:), of a possible tunicate found in Upper 6324:Hemichordata (acorn worms and relatives) 4796:"Invasive Tunicates of Washington State" 4039:. Saunders College Publishing. pp.  4026: 4024: 4022: 4020: 2599: 2597: 1728:, where absorption takes place, and the 1461:is never free-living and lacks a brain. 1436:, which is surrounded by the jelly-like 1380: 379:). The subphylum was at one time called 5553:Thaliacea (pyrosomes, salps, doliolids) 4258:Communicative & Integrative Biology 3926:"The Degenerate Tale of Ascidian Tails" 2468: 2466: 2300: 1597: 6318:Echinodermata (starfish and relatives) 6288:Xenacoelomorpha (acoels and relatives) 5321:, with unusual nuclear architecture". 5201: 4975:. Korea-US Aquaculture. Archived from 2771: 1566:organ is a mass of large clear-walled 1208:them, they may have been mistaken for 556:Tunicates are more closely related to 426:(pyrosomes, doliolids, and salps) and 16:Marine animals, subphylum of chordates 3836:Fouling Organisms of the Indian Ocean 3529:Palmer, T. J.; Wilson, M. A. (1988). 2109:Invasive tunicates usually arrive as 1416:, but they do not develop very clear 7: 6589:Acanthocephala (thorny-headed worms) 2232:The use of tunicates as a source of 7453:Cambrian Series 2 first appearances 4864:Stem cells : from hydra to man 4064:Schlosser, Gerhard (17 June 2021). 3930:Integrative and Comparative Biology 1752:. A few deepwater species, such as 1231:ancestor (before the divergence of 982: 955: 945: 879: 802: 792: 785: 724: 697: 687: 677: 6186:Cnidaria (jellyfish and relatives) 4308:Proceedings of the Royal Society B 2540:Giribet, Gonzalo (27 April 2018). 2151:, an FDA approved anticancer drug. 2094:have appeared and are thriving in 14: 7458:Extant Cambrian first appearances 4898:. Sciencedaily.com. 24 April 2007 4638:Molecular Genetics of Development 4462:. Little Brown & Co. p.  3613:"Introduction to the Urochordata" 2709:Bone, Q., and K. P. Ryan (1974). 2318:Journal of the Geological Society 1780:, is unique to tunicates. Excess 1642:Fluorescent-colored sea squirts, 1118:at Shankou village, Anning, near 535:The Tunicata were established by 460:or stiffening rod and resemble a 7230:Phyla with ≥1000 extant species 5696:Actinopterygii (ray-finned fish) 4516:"Botryllus: A colonial ascidian" 3570:Nagalwade, Vidya (7 July 2023). 3324:Tokioka, Takasi (30 June 1971). 2618:10.1111/j.1463-6409.2011.00493.x 2519:10.1111/j.1463-6409.2012.00536.x 2483:World Register of Marine Species 2183:tunicates for sale at a market, 1653: 1635: 1624:Blue sea squirts from the genus 1617: 1600: 1464:Tunicates have a well-developed 1012: 992: 965: 910: 889: 852: 832: 812: 749: 707: 367:which includes all animals with 359:). This grouping is part of the 308: 142: 1694:Nearly all adult tunicates are 1550:Tunicates lack the kidney-like 1367:Physiology and internal anatomy 430:on the other hand, swim in the 6417:Nematomorpha (horsehair worms) 4766:"Have You Seen This Tunicate?" 4431:Cavanihac, Jean-Marie (2000). 1896:life history phase, an oozoid 1: 5296:10.1126/science.294.5551.2506 4646:10.1016/S0065-2660(08)60012-7 4600:Biochem. Biophys. Res. Commun 4178:10.1016/S0022-0981(02)00023-0 3700:Journal of Biological Systems 2028:cell termed the "coenocyst". 1243:events among metazoan phyla. 406:, each unit being known as a 7029:Echinozoans inc. sea urchins 4862:Bosch, Thomas C. G. (2008). 4794:Dornfeld, Ann (1 May 2008). 2991:. Smithsonian at Fort Pierce 2938:"Sea squirts and sea tulips" 1801:Anatomy of a larval tunicate 1501:vanadium-associated proteins 1168:, and one from a second new 445:, sea pork, sea livers, or 7086:Chelicerates inc. arachnids 6826:Phoronida (horseshoe worms) 6641:Platyhelminthes (flatworms) 6552:Gnathostomulida (jaw worms) 6441:Loricifera (corset animals) 5527:Appendicularia (larvaceans) 4824:. Woods Hole Science Center 4743:10.1016/j.ydbio.2006.10.021 4489:Canadian Journal of Zoology 4454:Dennett, Daniel C. (1991). 4097:Journal of Chemical Ecology 3809:10.1016/j.ydbio.2008.11.026 2243:Some tunicates are used as 2020:in which the entire female 1834:, and a balancing organ, a 7474: 6976:Medusozoans inc. jellyfish 6722:Annelida (segmented worms) 6547:Chaetognatha (arrow worms) 6463:Onychophora (velvet worms) 5954:denote paraphyletic groups 5833:Squamata (scaled reptiles) 5335:10.1016/j.gene.2007.03.006 4612:10.1016/j.bbrc.2014.05.099 4537:Holland, Linda Z. (2007). 4433:"Tunicates extraordinaire" 3830:Renganathan, T.K. (2020). 3752:10.1038/s41598-022-18283-9 2440:10.1007/s12542-021-00596-1 2170:Ascidiacea § Culinary 2167: 1917:protandrous hermaphrodites 1911:Larvaceans only reproduce 1890:alternation of generations 1126:). There is also a common 7226: 7024:Asterozoans inc. starfish 6926: 6919: 6870: 6821:Brachiopoda (lamp shells) 6802:Ectoprocta (moss animals) 6762: 6738: 6707: 6647:Gastrotricha (hairybacks) 6619: 6542: 6529: 6458: 6436: 6425: 6376:Kinorhyncha (mud dragons) 6362: 6336: 6312: 6283: 6268: 6255: 6220: 6171: 6148:Ctenophora (comb jellies) 6136: 6121: 6005: 5925: 5871:Crocodilia (crocodilians) 5827:Rhynchocephalia (tuatara) 5793: 5769: 5744: 5719: 5691: 5663: 5617: 5570: 5542: 5522: 5505: 5468: 5442: 5110:10.1007/s00239-003-2511-9 5049:10.1016/j.tig.2007.07.001 4822:"Marine Nuisance Species" 4117:10.1007/s10886-007-9251-z 3712:10.1142/S0218339010003408 3544:: 939–949. Archived from 3436:10.1186/s12915-018-0517-4 3417:Giribet, Gonzalo (2018). 3292:10.1186/s12915-018-0499-2 2698:– via Google Books. 2653:10.1007/s12064-018-0260-y 2561:10.1186/s12915-018-0517-4 2392:10.1134/S0031030112010042 1960:Promotion of out-crossing 1906:sequential hermaphrodites 1806:Ascidians are almost all 1740:opens into the dorsal or 1690:(orange) showing siphons. 1288:or tunic, from which the 1046: 1007: 987: 980: 960: 953: 943: 904: 884: 877: 847: 827: 807: 800: 790: 783: 744: 729: 722: 702: 695: 685: 295:is an exclusively marine 279: 272: 219: 211: 139:Scientific classification 137: 126: 117: 23: 6582:Rotifera (wheel animals) 6381:Priapulida (penis worms) 5932:Ascidiacea (sea squirts) 5724:Actinistia (coelacanths) 4514:Parmentier, Jan (1998). 4053:– via archive.org. 3844:10.1201/9781003077992-16 3385:10.3389/fimmu.2017.00674 3217:BMC Evolutionary Biology 3143:10.1186/1471-2164-10-534 2682:Gee, H. (27 July 2007). 2473:Sanamyan, Karen (2013). 1736:pellets or strings. The 1507:in blood cells known as 1241:horizontal gene transfer 1108:Shankouclava shankouense 541:Francis Maitland Balfour 299:animal, a member of the 6796:Entoprocta or Kamptozoa 6768:Nemertea (ribbon worms) 6489:Tardigrada (waterbears) 6482:Arthropoda (arthropods) 5482:Leptocardii (lancelets) 5395:Encyclopædia Britannica 5359:The Tunicate Web Portal 5246:10.1186/gb-2007-8-3-r41 5180:10.1126/science.1080049 5088:mitochondrial genome". 4923:The Biological Bulletin 4866:. Dordrecht: Springer. 4458:Consciousness Explained 4385:10.1073/pnas.0503640102 4220:10.1073/pnas.0303623101 3497:10.1073/pnas.1431177100 3372:Frontiers in Immunology 3238:10.1186/1471-2148-9-187 2371:Paleontological Journal 2291:– claimed hybridization 1158:from the Nama Group of 1087:The star-shaped holes ( 485:, the bluebell tunicate 127:Gold-mouth sea squirt ( 6971:Anthozoans inc. corals 6670:Dicyemida or Rhombozoa 5636:Hyperoartia (lampreys) 5398:(11th ed.). 1911. 4321:10.1098/rspb.2016.1712 2925:Tunicata Lamarck, 1816 2735:10.1098/rspb.1974.0077 2191: 1858: 1803: 1760:sit-and-wait predators 1691: 1684:(black and white) and 1389: 1378: 1261: 1102: 499: 495:Botrylloides violaceus 486: 434:of the sea as adults. 7407:Paleobiology Database 7101:Hexapods inc. insects 6411:Nematoda (roundworms) 5142:"The draft genome of 4798:. NPR. Archived from 3797:Developmental Biology 3687:– via CrossRef. 2261:developmental studies 2177: 2018:reproductive strategy 1976:inbreeding depression 1948:and vertebrates show 1856: 1799: 1678: 1609:Oikopleura cophocerca 1384: 1374: 1259: 1219:Hybridization studies 1165:Yarnemia ascidiformis 1086: 537:Jean-Baptiste Lamarck 492: 482:Clavelina moluccensis 479: 212:Classes and unplaced 6294:Chordata (chordates) 5852:Testudines (turtles) 4935:10.1086/BBLv221n1p43 3838:. pp. 507–534. 3551:on 27 September 2013 2879:Invertebrate Zoology 2289:Donald I. Williamson 1983:Botryllus schlosseri 1938:determinate cleavage 1898:reproduces asexually 1132:Catellocaula vallata 1090:Catellocaula vallata 529:Pyura pachydermatina 6744:Mollusca (molluscs) 6229:of the animal root 5946:Reptilia (reptiles) 5654:(jawed vertebrates) 5375:Halocynthia roretzi 5162:2002Sci...298.2157D 5156:(5601): 2157–2167. 5102:2003JMolE..57..574Y 5086:Halocynthia roretzi 4555:2007Natur.447..153H 4376:2005PNAS..10215134S 4370:(42): 15134–15139. 4211:2004PNAS..101..986M 4160:2002JEMBE.270..203P 4109:2007JCEco..33..643O 3943:10.1093/icb/icab022 3893:10.1098/rsob.150053 3744:2022NatSR..1214326D 3669:2018Palgy..61..575W 3488:2003PNAS..100.8314C 3229:2009BMCEE...9..187T 3042:10.1038/nature04336 3034:2006Natur.439..965D 2940:. Australian Museum 2815:10.1002/jez.b.22940 2807:2021JEZB..336..250A 2727:1974RSPSB.187..315B 2431:2021PalZ...95..623M 2384:2012PalJ...46....1F 2339:10.1144/jgs2017-103 2331:2018JGSoc.175..659Y 2259:have been used for 2218:Halocynthia roretzi 1755:Megalodicopia hians 1720:are secreted and a 1687:Pycnoclavella flava 1588:synthesizing enzyme 1547:against predators. 1396:, tunicates have a 1350:A third class, the 1143:Megasiphon thylakos 437:Various species of 6957:Calcareous sponges 6142:Porifera (sponges) 5798:Mammalia (mammals) 5710:(lobe-finned fish) 5682:(bony vertebrates) 5371:Ciona intestinalis 5144:Ciona intestinalis 5028:Ciona intestinalis 4356:Ciona intestinalis 4315:(1845): 20161712. 3732:Scientific Reports 3677:10.1111/pala.12356 2987:Aplidium stellatum 2250:Ciona intestinalis 2192: 2136:chemical compounds 1966:Ciona intestinalis 1950:cell determination 1859: 1804: 1696:suspension feeders 1692: 1554:organs typical of 1470:circulatory system 1390: 1379: 1262: 1103: 539:in 1816. In 1881, 500: 487: 410:. They are marine 369:dorsal nerve cords 7435: 7434: 7394:Open Tree of Life 7256:Taxon identifiers 7247: 7246: 7222: 7221: 6915: 6914: 6911: 6910: 6866: 6865: 6862: 6861: 6858: 6857: 6854: 6853: 6850: 6849: 6846: 6845: 6842: 6841: 6838: 6837: 6834: 6833: 6692: 6691: 6688: 6687: 6604: 6603: 6600: 6599: 6514: 6513: 6510: 6509: 6506: 6505: 6502: 6501: 6498: 6497: 6389: 6388: 6332: 6331: 6251: 6250: 6216: 6215: 6212: 6211: 5961: 5960: 5943:³orders of class 5929:¹orders of class 5921: 5920: 5917: 5916: 5913: 5912: 5909: 5908: 5905: 5904: 5901: 5900: 5897: 5896: 5893: 5892: 5889: 5888: 5885: 5884: 5749:Dipnoi (lungfish) 5644: 5643: 5602: 5601: 5598: 5597: 5594: 5593: 5590: 5589: 5490: 5489: 5319:Oikopleura dioica 5280:Oikopleura dioica 4772:on 9 January 2009 4695:10.1002/dvg.22802 4270:10.4161/cib.17757 4077:978-1-000-36913-7 4050:978-0-03-030504-7 3993:10.1111/brv.12189 3482:(14): 8314–8318. 3090:10.1002/dvg.20450 3028:(7079): 965–968. 2888:978-81-315-0104-7 2721:(1088): 315–327. 2606:Zoologica Scripta 2506:Zoologica Scripta 2266:Oikopleura dioica 2208:) is used in the 2111:fouling organisms 2104:Pacific Northwest 2071:Didemnum vexillum 2005:Oikopleura dioica 1922:Oikopleura dioica 1681:Clavelina robusta 1545:chemical defenses 1541:organic compounds 1299:connective tissue 1116:Maotianshan Shale 1075: 1074: 1066: 1065: 1057: 1056: 1043: 1042: 1034: 1033: 1025: 1024: 932: 931: 923: 922: 865: 864: 771: 770: 762: 761: 289: 288: 284: 207: 7465: 7428: 7427: 7415: 7414: 7402: 7401: 7389: 7388: 7376: 7375: 7363: 7362: 7350: 7349: 7337: 7336: 7324: 7323: 7311: 7310: 7298: 7297: 7296: 7283: 7282: 7281: 7251: 6928: 6884: 6784: 6760: 6736: 6705: 6701: 6628: 6621: 6617: 6613: 6540: 6531: 6527: 6523: 6456: 6434: 6427: 6398: 6364: 6360: 6356: 6349: 6345: 6338: 6310: 6281: 6277: 6270: 6257: 6234: 6169: 6134: 6130: 6123: 6034: 6014: 5988: 5981: 5974: 5965: 5859: 5815: 5811: 5791: 5787: 5767: 5763: 5742: 5738: 5717: 5713: 5711: 5689: 5685: 5683: 5661: 5657: 5655: 5631:Myxini (hagfish) 5619: 5615: 5611: 5568: 5540: 5520: 5515: 5507: 5503: 5499: 5470: 5427: 5420: 5413: 5404: 5399: 5391: 5389:"Tunicata"  5347: 5346: 5314: 5308: 5307: 5275: 5269: 5268: 5258: 5248: 5220: 5214: 5213: 5207: 5199: 5173: 5136: 5130: 5129: 5077: 5071: 5070: 5060: 5034: 5021: 5015: 5014: 5012: 5010: 4995: 4989: 4988: 4986: 4984: 4969: 4963: 4962: 4914: 4908: 4907: 4905: 4903: 4892: 4886: 4885: 4859: 4853: 4852: 4849:Journal Sentinel 4840: 4834: 4833: 4831: 4829: 4818: 4812: 4811: 4809: 4807: 4791: 4782: 4781: 4779: 4777: 4762: 4756: 4755: 4745: 4721: 4715: 4714: 4677: 4668: 4667: 4633: 4624: 4623: 4594: 4585: 4584: 4566: 4534: 4528: 4527: 4525: 4523: 4511: 4505: 4504: 4484: 4478: 4477: 4473:978-0316-18065-8 4461: 4451: 4445: 4444: 4442: 4440: 4428: 4419: 4414: 4408: 4407: 4397: 4387: 4349: 4343: 4342: 4332: 4298: 4292: 4291: 4281: 4249: 4243: 4242: 4232: 4222: 4188: 4182: 4181: 4171: 4143: 4137: 4136: 4088: 4082: 4081: 4061: 4055: 4054: 4038: 4028: 4015: 4014: 4004: 3972: 3966: 3965: 3955: 3945: 3921: 3915: 3914: 3904: 3872: 3866: 3865: 3827: 3821: 3820: 3788: 3782: 3781: 3771: 3722: 3716: 3715: 3695: 3689: 3688: 3648: 3642: 3635: 3629: 3628: 3626: 3624: 3619:on 21 April 2009 3609: 3603: 3599: 3590: 3586: 3580: 3579: 3567: 3561: 3560: 3558: 3556: 3550: 3535: 3526: 3520: 3519: 3509: 3499: 3465: 3459: 3458: 3448: 3438: 3414: 3408: 3407: 3397: 3387: 3363: 3357: 3356: 3330: 3321: 3315: 3314: 3304: 3294: 3270: 3261: 3260: 3250: 3240: 3208: 3202: 3201: 3181: 3175: 3172: 3166: 3165: 3155: 3145: 3117: 3111: 3110: 3092: 3068: 3062: 3061: 3019: 3010: 3001: 3000: 2998: 2996: 2981: 2975: 2974: 2972: 2970: 2956: 2950: 2949: 2947: 2945: 2934: 2928: 2922: 2916: 2910: 2904: 2899: 2893: 2892: 2874: 2835: 2834: 2790: 2784: 2783: 2777: 2769: 2767: 2765: 2706: 2700: 2699: 2679: 2673: 2672: 2636: 2630: 2629: 2601: 2592: 2591: 2581: 2563: 2537: 2531: 2530: 2500: 2494: 2493: 2491: 2489: 2470: 2461: 2460: 2442: 2410: 2404: 2403: 2365: 2359: 2358: 2314: 2305: 2210:cuisine of Chile 2048:Invasive species 1999:A model tunicate 1954:genome evolution 1657: 1645:Rhopalaea crassa 1639: 1621: 1604: 1388: 1155:Ausia fenestrata 1016: 996: 983: 969: 956: 946: 914: 893: 880: 856: 836: 816: 803: 793: 786: 753: 725: 711: 698: 688: 678: 355: 351: 346: 345: 342: 341: 338: 335: 332: 329: 326: 323: 320: 317: 314: 282: 261: 251: 240: 202: 147: 146: 130:Polycarpa aurata 122: 107: 44: 30:Cambrian Stage 3 27:Temporal range: 21: 7473: 7472: 7468: 7467: 7466: 7464: 7463: 7462: 7438: 7437: 7436: 7431: 7423: 7418: 7410: 7405: 7397: 7392: 7384: 7379: 7371: 7366: 7358: 7353: 7345: 7340: 7332: 7327: 7319: 7314: 7306: 7301: 7292: 7291: 7286: 7277: 7276: 7271: 7258: 7248: 7243: 7218: 7144:Phylactolaemata 6932: 6922: 6921: 6907: 6898:Xenambulacraria 6874: 6830: 6807: 6773: 6755:Kryptotrochozoa 6749: 6727: 6697: 6684: 6675:Monoblastozoa ( 6651: 6609: 6596: 6568:Micrognathozoa 6556: 6519: 6494: 6467: 6445: 6421: 6385: 6352: 6341: 6328: 6299: 6273: 6264: 6247: 6224: 6208: 6191: 6160: 6152: 6126: 6117: 6116: 6032: 6010: 6001: 5992: 5962: 5957: 5937:²subclasses of 5881: 5857: 5838: 5807: 5802: 5783: 5778: 5759: 5754: 5734: 5729: 5709: 5708: 5705: 5700: 5681: 5680: 5677: 5672: 5653: 5652: 5649: 5640: 5607: 5586: 5581:Aplousobranchia 5557: 5547:Stolidobranchia 5531: 5513: 5512: 5495: 5486: 5474:Cephalochordata 5464: 5438: 5431: 5386: 5355: 5350: 5316: 5315: 5311: 5277: 5276: 5272: 5222: 5221: 5217: 5200: 5171:10.1.1.319.2643 5138: 5137: 5133: 5079: 5078: 5074: 5032: 5023: 5022: 5018: 5008: 5006: 5005:. 26 March 2013 4997: 4996: 4992: 4982: 4980: 4979:on 2 March 2013 4971: 4970: 4966: 4916: 4915: 4911: 4901: 4899: 4894: 4893: 4889: 4874: 4861: 4860: 4856: 4843:Johnson, Mark. 4842: 4841: 4837: 4827: 4825: 4820: 4819: 4815: 4805: 4803: 4802:on 14 July 2014 4793: 4792: 4785: 4775: 4773: 4764: 4763: 4759: 4723: 4722: 4718: 4679: 4678: 4671: 4656: 4635: 4634: 4627: 4596: 4595: 4588: 4564:10.1038/447153a 4536: 4535: 4531: 4521: 4519: 4518:. Microscope UK 4513: 4512: 4508: 4501:10.1139/z04-177 4486: 4485: 4481: 4474: 4453: 4452: 4448: 4438: 4436: 4435:. Microscope UK 4430: 4429: 4422: 4415: 4411: 4351: 4350: 4346: 4300: 4299: 4295: 4251: 4250: 4246: 4190: 4189: 4185: 4169:10.1.1.558.3639 4145: 4144: 4140: 4093:Phallusia nigra 4090: 4089: 4085: 4078: 4063: 4062: 4058: 4051: 4030: 4029: 4018: 3974: 3973: 3969: 3923: 3922: 3918: 3874: 3873: 3869: 3854: 3829: 3828: 3824: 3790: 3789: 3785: 3724: 3723: 3719: 3697: 3696: 3692: 3650: 3649: 3645: 3636: 3632: 3622: 3620: 3611: 3610: 3606: 3600: 3593: 3587: 3583: 3569: 3568: 3564: 3554: 3552: 3548: 3533: 3528: 3527: 3523: 3467: 3466: 3462: 3416: 3415: 3411: 3365: 3364: 3360: 3328: 3323: 3322: 3318: 3272: 3271: 3264: 3210: 3209: 3205: 3198:10.1139/z05-010 3183: 3182: 3178: 3173: 3169: 3124:Herdmania momus 3119: 3118: 3114: 3083:(11): 592–604. 3070: 3069: 3065: 3017: 3012: 3011: 3004: 2994: 2992: 2983: 2982: 2978: 2968: 2966: 2958: 2957: 2953: 2943: 2941: 2936: 2935: 2931: 2923: 2919: 2911: 2907: 2900: 2896: 2889: 2876: 2875: 2838: 2792: 2791: 2787: 2770: 2763: 2761: 2708: 2707: 2703: 2696: 2681: 2680: 2676: 2638: 2637: 2633: 2603: 2602: 2595: 2539: 2538: 2534: 2502: 2501: 2497: 2487: 2485: 2472: 2471: 2464: 2412: 2411: 2407: 2367: 2366: 2362: 2312: 2307: 2306: 2302: 2298: 2275: 2245:model organisms 2230: 2205:Pyura chilensis 2172: 2166: 2132: 2127: 2050: 2036:because of its 2001: 1971:C. intestinalis 1962: 1802: 1794: 1770:green algae or 1673: 1666: 1658: 1649: 1640: 1631: 1622: 1613: 1605: 1392:Like all other 1387:(br s) pharynx. 1386: 1377: 1369: 1278: 1254: 1249: 1225:molecular study 1221: 1210:sponge spicules 1162:, the sac-like 1111:from the Lower 1081: 1076: 1067: 1058: 1044: 1035: 1026: 950:Stolidobranchia 933: 924: 907:Aplousobranchia 866: 772: 763: 659:Members of the 625:Stolidobranchia 617:Aplousobranchia 554: 474: 454:Cambrian period 383:, and the term 353: 349: 311: 307: 283:Lankester, 1877 201: 141: 113: 112:record, 557 Ma) 106: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 55: 50: 39: 38: 37: 28: 25: 17: 12: 11: 5: 7471: 7469: 7461: 7460: 7455: 7450: 7440: 7439: 7433: 7432: 7430: 7429: 7416: 7403: 7390: 7377: 7364: 7351: 7338: 7325: 7312: 7299: 7284: 7268: 7266: 7260: 7259: 7254: 7245: 7244: 7242: 7241: 7234: 7227: 7224: 7223: 7220: 7219: 7217: 7216: 7215: 7214: 7209: 7204: 7199: 7194: 7182: 7181: 7180: 7175: 7170: 7158: 7157: 7156: 7151: 7146: 7134: 7133: 7132: 7127: 7122: 7117: 7109:Platyhelminths 7105: 7104: 7103: 7098: 7093: 7088: 7076: 7075: 7074: 7069: 7064: 7052: 7051: 7050: 7045: 7033: 7032: 7031: 7026: 7021: 7009: 7008: 7007: 7002: 6997: 6985: 6984: 6983: 6978: 6973: 6961: 6960: 6959: 6954: 6949: 6936: 6934: 6924: 6923: 6920: 6917: 6916: 6913: 6912: 6909: 6908: 6906: 6905: 6903:Centroneuralia 6900: 6895: 6890: 6885: 6871: 6868: 6867: 6864: 6863: 6860: 6859: 6856: 6855: 6852: 6851: 6848: 6847: 6844: 6843: 6840: 6839: 6836: 6835: 6832: 6831: 6829: 6828: 6823: 6817: 6815: 6809: 6808: 6806: 6805: 6798: 6792: 6790: 6781: 6775: 6774: 6772: 6771: 6763: 6757: 6751: 6750: 6748: 6747: 6739: 6733: 6729: 6728: 6726: 6725: 6718: 6708: 6702: 6699:Lophotrochozoa 6694: 6693: 6690: 6689: 6686: 6685: 6683: 6682: 6672: 6667: 6661: 6659: 6653: 6652: 6650: 6649: 6644: 6636: 6634: 6625: 6614: 6611:Platytrochozoa 6606: 6605: 6602: 6601: 6598: 6597: 6595: 6594: 6593: 6592: 6585: 6573: 6570:(Limnognathia) 6564: 6562: 6558: 6557: 6555: 6554: 6549: 6543: 6537: 6524: 6516: 6515: 6512: 6511: 6508: 6507: 6504: 6503: 6500: 6499: 6496: 6495: 6493: 6492: 6485: 6477: 6475: 6469: 6468: 6466: 6465: 6459: 6453: 6447: 6446: 6444: 6443: 6437: 6431: 6423: 6422: 6420: 6419: 6414: 6406: 6404: 6395: 6391: 6390: 6387: 6386: 6384: 6383: 6378: 6372: 6370: 6357: 6346: 6334: 6333: 6330: 6329: 6327: 6326: 6321: 6313: 6307: 6301: 6300: 6298: 6297: 6290: 6284: 6278: 6266: 6265: 6260: 6253: 6252: 6249: 6248: 6246: 6245: 6240: 6235: 6221: 6218: 6217: 6214: 6213: 6210: 6209: 6207: 6206: 6202: 6200: 6199:(Triploblasts) 6193: 6192: 6190: 6189: 6182: 6180:and relatives) 6172: 6166: 6154: 6153: 6151: 6150: 6145: 6137: 6131: 6119: 6118: 6115: 6114: 6113: 6112: 6111: 6110: 6107:Mesomycetozoea 6104: 6099: 6089: 6079: 6078: 6077: 6072: 6067: 6062: 6057: 6056: 6055: 6043:Diaphoretickes 6040: 6035: 6030: 6025: 6020: 6015: 6007: 6006: 6003: 6002: 5993: 5991: 5990: 5983: 5976: 5968: 5959: 5958: 5956: 5955: 5949: 5941: 5935: 5926: 5923: 5922: 5919: 5918: 5915: 5914: 5911: 5910: 5907: 5906: 5903: 5902: 5899: 5898: 5895: 5894: 5891: 5890: 5887: 5886: 5883: 5882: 5880: 5879: 5874: 5867: 5865: 5856: 5855: 5848: 5846: 5840: 5839: 5837: 5836: 5830: 5823: 5821: 5812: 5804: 5803: 5801: 5800: 5794: 5788: 5780: 5779: 5777: 5776: 5770: 5764: 5756: 5755: 5753: 5752: 5745: 5739: 5731: 5730: 5728: 5727: 5720: 5714: 5702: 5701: 5699: 5698: 5692: 5686: 5674: 5673: 5671: 5670: 5664: 5658: 5646: 5645: 5642: 5641: 5639: 5638: 5633: 5627: 5625: 5612: 5604: 5603: 5600: 5599: 5596: 5595: 5592: 5591: 5588: 5587: 5585: 5584: 5578: 5575:Phlebobranchia 5571: 5565: 5559: 5558: 5556: 5555: 5550: 5543: 5537: 5533: 5532: 5530: 5529: 5523: 5517: 5500: 5492: 5491: 5488: 5487: 5485: 5484: 5478: 5476: 5466: 5465: 5463: 5462: 5456: 5450: 5443: 5440: 5439: 5432: 5430: 5429: 5422: 5415: 5407: 5401: 5400: 5384: 5378: 5366: 5361: 5354: 5353:External links 5351: 5349: 5348: 5329:(1): 159–169. 5309: 5290:(5551): 2506. 5270: 5227:Ciona savignyi 5215: 5131: 5096:(5): 574–587. 5082:Ciona savignyi 5072: 5043:(9): 419–422. 5016: 4990: 4964: 4909: 4887: 4872: 4854: 4835: 4813: 4783: 4757: 4736:(2): 591–600. 4716: 4669: 4654: 4625: 4586: 4549:(1): 153–155. 4529: 4506: 4495:(1): 151–183. 4479: 4472: 4446: 4420: 4409: 4344: 4293: 4264:(6): 768–771. 4244: 4205:(4): 986–991. 4183: 4154:(2): 203–214. 4138: 4103:(3): 643–654. 4083: 4076: 4056: 4049: 4016: 3987:(3): 673–711. 3967: 3936:(2): 358–369. 3916: 3867: 3852: 3822: 3803:(2): 357–367. 3783: 3717: 3706:(2): 261–275. 3690: 3663:(4): 575–595. 3643: 3630: 3604: 3591: 3581: 3576:Tech Explorist 3562: 3521: 3460: 3409: 3358: 3345:10.5134/175655 3316: 3262: 3203: 3176: 3167: 3112: 3063: 3002: 2976: 2964:Dictionary.com 2951: 2929: 2917: 2905: 2894: 2887: 2836: 2801:(3): 250–266. 2785: 2701: 2694: 2674: 2631: 2612:(6): 603–612. 2593: 2532: 2513:(4): 435–436. 2495: 2462: 2425:(4): 623–639. 2405: 2360: 2325:(4): 659–666. 2299: 2297: 2294: 2293: 2292: 2286: 2274: 2271: 2256:Ciona savignyi 2229: 2226: 2168:Main article: 2165: 2162: 2153: 2152: 2146: 2131: 2128: 2126: 2123: 2091:Ciona savignyi 2049: 2046: 2034:model organism 2010:Appendicularia 2000: 1997: 1961: 1958: 1808:hermaphrodites 1800: 1793: 1790: 1782:photosynthetic 1672: 1669: 1668: 1667: 1662:Didemnum molle 1659: 1652: 1650: 1641: 1634: 1632: 1623: 1616: 1614: 1606: 1599: 1573:urate crystals 1552:metanephridial 1450:Appendicularia 1428:, gonads, and 1375: 1368: 1365: 1277: 1276:Body structure 1274: 1253: 1250: 1248: 1245: 1223:A multi-taxon 1220: 1217: 1179:Burykhia hunti 1080: 1077: 1073: 1072: 1069: 1068: 1064: 1063: 1060: 1059: 1055: 1054: 1051: 1050: 1045: 1041: 1040: 1037: 1036: 1032: 1031: 1028: 1027: 1023: 1022: 1019: 1018: 1006: 1003: 1002: 999: 998: 986: 981: 979: 976: 975: 972: 971: 959: 954: 952: 944: 942: 939: 938: 935: 934: 930: 929: 926: 925: 921: 920: 917: 916: 903: 900: 899: 896: 895: 887:Phlebobranchia 883: 878: 876: 872: 871: 868: 867: 863: 862: 859: 858: 846: 843: 842: 839: 838: 826: 823: 822: 819: 818: 806: 801: 799: 791: 789: 784: 782: 778: 777: 774: 773: 769: 768: 765: 764: 760: 759: 756: 755: 747:Fritillariidae 743: 740: 739: 736: 735: 732:Kowalevskiidae 728: 723: 721: 718: 717: 714: 713: 701: 696: 694: 692:Appendicularia 686: 684: 676: 657: 656: 649:Appendicularia 646: 628: 621:Phlebobranchia 553: 552:Classification 550: 514:and a few are 473: 470: 412:filter feeders 287: 286: 277: 276: 270: 269: 268: 267: 257: 247: 236: 234:Appendicularia 231: 226: 225:(polyphyletic) 217: 216: 209: 208: 196: 192: 191: 186: 179: 178: 173: 169: 168: 163: 159: 158: 153: 149: 148: 135: 134: 124: 123: 115: 114: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 51: 46: 45: 26: 15: 13: 10: 9: 6: 4: 3: 2: 7470: 7459: 7456: 7454: 7451: 7449: 7446: 7445: 7443: 7426: 7421: 7417: 7413: 7408: 7404: 7400: 7395: 7391: 7387: 7382: 7378: 7374: 7369: 7365: 7361: 7356: 7352: 7348: 7343: 7339: 7335: 7330: 7326: 7322: 7317: 7313: 7309: 7304: 7300: 7295: 7289: 7285: 7280: 7274: 7270: 7269: 7267: 7265: 7261: 7257: 7252: 7239: 7238:dubious phyla 7235: 7233: 7229: 7228: 7225: 7213: 7210: 7208: 7205: 7203: 7200: 7198: 7195: 7193: 7190: 7189: 7188: 7187: 7183: 7179: 7176: 7174: 7171: 7169: 7166: 7165: 7164: 7163: 7159: 7155: 7152: 7150: 7147: 7145: 7142: 7141: 7140: 7139: 7135: 7131: 7128: 7126: 7123: 7121: 7118: 7116: 7113: 7112: 7111: 7110: 7106: 7102: 7099: 7097: 7094: 7092: 7089: 7087: 7084: 7083: 7082: 7081: 7077: 7073: 7070: 7068: 7065: 7063: 7060: 7059: 7058: 7057: 7053: 7049: 7046: 7044: 7041: 7040: 7039: 7038: 7037:Hemichordates 7034: 7030: 7027: 7025: 7022: 7020: 7017: 7016: 7015: 7014: 7010: 7006: 7003: 7001: 6998: 6996: 6993: 6992: 6991: 6990: 6986: 6982: 6979: 6977: 6974: 6972: 6969: 6968: 6967: 6966: 6962: 6958: 6955: 6953: 6952:Glass sponges 6950: 6948: 6945: 6944: 6943: 6942: 6938: 6937: 6935: 6929: 6925: 6918: 6904: 6901: 6899: 6896: 6894: 6893:Deuterostomia 6891: 6889: 6886: 6882: 6879:of Bilateria 6878: 6873: 6872: 6869: 6827: 6824: 6822: 6819: 6818: 6816: 6814: 6810: 6804: 6803: 6799: 6797: 6794: 6793: 6791: 6789: 6785: 6782: 6780: 6776: 6770: 6769: 6765: 6764: 6761: 6758: 6756: 6752: 6746: 6745: 6741: 6740: 6737: 6734: 6730: 6724: 6723: 6719: 6717: 6715: 6712:Cycliophora ( 6710: 6709: 6706: 6703: 6700: 6695: 6680: 6678: 6673: 6671: 6668: 6666: 6663: 6662: 6660: 6658: 6654: 6648: 6645: 6643: 6642: 6638: 6637: 6635: 6633: 6629: 6626: 6622: 6618: 6615: 6612: 6607: 6591: 6590: 6586: 6584: 6583: 6579: 6578: 6577: 6574: 6572: 6571: 6566: 6565: 6563: 6559: 6553: 6550: 6548: 6545: 6544: 6541: 6538: 6536: 6532: 6528: 6525: 6522: 6517: 6491: 6490: 6486: 6484: 6483: 6479: 6478: 6476: 6474: 6470: 6464: 6461: 6460: 6457: 6454: 6452: 6451:Panarthropoda 6448: 6442: 6439: 6438: 6435: 6432: 6428: 6424: 6418: 6415: 6413: 6412: 6408: 6407: 6405: 6403: 6399: 6396: 6392: 6382: 6379: 6377: 6374: 6373: 6371: 6369: 6365: 6361: 6358: 6355: 6350: 6347: 6344: 6339: 6335: 6325: 6322: 6320: 6319: 6315: 6314: 6311: 6308: 6306: 6302: 6296: 6295: 6291: 6289: 6286: 6285: 6282: 6279: 6276: 6271: 6267: 6263: 6258: 6254: 6244: 6241: 6239: 6236: 6232: 6228: 6223: 6222: 6219: 6204: 6203: 6201: 6198: 6194: 6188: 6187: 6183: 6181: 6179: 6174: 6173: 6170: 6167: 6164: 6159: 6155: 6149: 6146: 6144: 6143: 6139: 6138: 6135: 6132: 6129: 6124: 6120: 6108: 6105: 6103: 6100: 6098: 6095: 6094: 6093: 6090: 6088: 6085: 6084: 6083: 6080: 6076: 6073: 6071: 6070:Stramenopiles 6068: 6066: 6063: 6061: 6058: 6054: 6051: 6050: 6049: 6046: 6045: 6044: 6041: 6039: 6036: 6033:(major groups 6031: 6029: 6026: 6024: 6021: 6019: 6016: 6013: 6009: 6008: 6004: 6000: 5997: 5989: 5984: 5982: 5977: 5975: 5970: 5969: 5966: 5953: 5950: 5948: 5947: 5942: 5940: 5939:Sarcopterygii 5936: 5934: 5933: 5928: 5927: 5924: 5878: 5875: 5872: 5869: 5868: 5866: 5864: 5860: 5853: 5850: 5849: 5847: 5845: 5844:Archelosauria 5841: 5834: 5831: 5828: 5825: 5824: 5822: 5820: 5816: 5813: 5810: 5805: 5799: 5796: 5795: 5792: 5789: 5786: 5781: 5775: 5772: 5771: 5768: 5765: 5762: 5757: 5750: 5747: 5746: 5743: 5740: 5737: 5732: 5725: 5722: 5721: 5718: 5715: 5712: 5707:Sarcopterygii 5703: 5697: 5694: 5693: 5690: 5687: 5684: 5675: 5669: 5666: 5665: 5662: 5659: 5656: 5651:Gnathostomata 5647: 5637: 5634: 5632: 5629: 5628: 5626: 5624: 5620: 5616: 5613: 5610: 5605: 5582: 5579: 5576: 5573: 5572: 5569: 5566: 5564: 5560: 5554: 5551: 5548: 5545: 5544: 5541: 5538: 5534: 5528: 5525: 5524: 5521: 5518: 5516: 5514:(Urochordata) 5508: 5504: 5501: 5498: 5493: 5483: 5480: 5479: 5477: 5475: 5471: 5467: 5461: 5460:Deuterostomia 5457: 5455: 5451: 5449: 5445: 5444: 5441: 5436: 5428: 5423: 5421: 5416: 5414: 5409: 5408: 5405: 5397: 5396: 5390: 5385: 5382: 5379: 5377: 5376: 5372: 5367: 5365: 5362: 5360: 5357: 5356: 5352: 5344: 5340: 5336: 5332: 5328: 5324: 5320: 5313: 5310: 5305: 5301: 5297: 5293: 5289: 5285: 5281: 5274: 5271: 5266: 5262: 5257: 5252: 5247: 5242: 5238: 5234: 5230: 5228: 5219: 5216: 5211: 5205: 5197: 5193: 5189: 5185: 5181: 5177: 5172: 5167: 5163: 5159: 5155: 5151: 5147: 5145: 5135: 5132: 5127: 5123: 5119: 5115: 5111: 5107: 5103: 5099: 5095: 5091: 5087: 5083: 5076: 5073: 5068: 5064: 5059: 5054: 5050: 5046: 5042: 5038: 5031: 5029: 5020: 5017: 5004: 5003:Cleantechnica 5000: 4994: 4991: 4978: 4974: 4968: 4965: 4960: 4956: 4952: 4948: 4944: 4940: 4936: 4932: 4928: 4924: 4920: 4913: 4910: 4897: 4891: 4888: 4883: 4879: 4875: 4873:9781402082740 4869: 4865: 4858: 4855: 4850: 4846: 4839: 4836: 4823: 4817: 4814: 4801: 4797: 4790: 4788: 4784: 4771: 4767: 4761: 4758: 4753: 4749: 4744: 4739: 4735: 4731: 4727: 4720: 4717: 4712: 4708: 4704: 4700: 4696: 4692: 4689:(1): 105–20. 4688: 4684: 4676: 4674: 4670: 4665: 4661: 4657: 4655:9780120176243 4651: 4647: 4643: 4639: 4632: 4630: 4626: 4621: 4617: 4613: 4609: 4606:(3): 1142–8. 4605: 4601: 4593: 4591: 4587: 4582: 4578: 4574: 4570: 4565: 4560: 4556: 4552: 4548: 4544: 4540: 4533: 4530: 4517: 4510: 4507: 4502: 4498: 4494: 4490: 4483: 4480: 4475: 4469: 4465: 4460: 4459: 4450: 4447: 4434: 4427: 4425: 4421: 4418: 4413: 4410: 4405: 4401: 4396: 4391: 4386: 4381: 4377: 4373: 4369: 4365: 4364: 4359: 4357: 4348: 4345: 4340: 4336: 4331: 4326: 4322: 4318: 4314: 4310: 4309: 4304: 4297: 4294: 4289: 4285: 4280: 4275: 4271: 4267: 4263: 4259: 4255: 4248: 4245: 4240: 4236: 4231: 4226: 4221: 4216: 4212: 4208: 4204: 4200: 4199: 4194: 4187: 4184: 4179: 4175: 4170: 4165: 4161: 4157: 4153: 4149: 4142: 4139: 4134: 4130: 4126: 4122: 4118: 4114: 4110: 4106: 4102: 4098: 4094: 4087: 4084: 4079: 4073: 4070:. CRC Press. 4069: 4068: 4060: 4057: 4052: 4046: 4042: 4037: 4036: 4027: 4025: 4023: 4021: 4017: 4012: 4008: 4003: 3998: 3994: 3990: 3986: 3982: 3978: 3971: 3968: 3963: 3959: 3954: 3949: 3944: 3939: 3935: 3931: 3927: 3920: 3917: 3912: 3908: 3903: 3898: 3894: 3890: 3887:(6): 150053. 3886: 3882: 3878: 3871: 3868: 3863: 3859: 3855: 3853:9781003077992 3849: 3845: 3841: 3837: 3833: 3826: 3823: 3818: 3814: 3810: 3806: 3802: 3798: 3794: 3787: 3784: 3779: 3775: 3770: 3765: 3761: 3757: 3753: 3749: 3745: 3741: 3737: 3733: 3729: 3721: 3718: 3713: 3709: 3705: 3701: 3694: 3691: 3686: 3682: 3678: 3674: 3670: 3666: 3662: 3658: 3657:Palaeontology 3654: 3647: 3644: 3640: 3634: 3631: 3618: 3614: 3608: 3605: 3598: 3596: 3592: 3585: 3582: 3577: 3573: 3566: 3563: 3547: 3543: 3539: 3538:Palaeontology 3532: 3525: 3522: 3517: 3513: 3508: 3503: 3498: 3493: 3489: 3485: 3481: 3477: 3476: 3471: 3464: 3461: 3456: 3452: 3447: 3442: 3437: 3432: 3428: 3424: 3420: 3413: 3410: 3405: 3401: 3396: 3391: 3386: 3381: 3377: 3373: 3369: 3362: 3359: 3354: 3350: 3346: 3342: 3338: 3334: 3327: 3320: 3317: 3312: 3308: 3303: 3298: 3293: 3288: 3284: 3280: 3276: 3269: 3267: 3263: 3258: 3254: 3249: 3244: 3239: 3234: 3230: 3226: 3222: 3218: 3214: 3207: 3204: 3199: 3195: 3191: 3187: 3180: 3177: 3171: 3168: 3163: 3159: 3154: 3149: 3144: 3139: 3135: 3131: 3127: 3125: 3116: 3113: 3108: 3104: 3100: 3096: 3091: 3086: 3082: 3078: 3074: 3067: 3064: 3059: 3055: 3051: 3047: 3043: 3039: 3035: 3031: 3027: 3023: 3016: 3009: 3007: 3003: 2990: 2988: 2980: 2977: 2965: 2961: 2955: 2952: 2939: 2933: 2930: 2926: 2921: 2918: 2914: 2909: 2906: 2903: 2898: 2895: 2890: 2884: 2880: 2873: 2871: 2869: 2867: 2865: 2863: 2861: 2859: 2857: 2855: 2853: 2851: 2849: 2847: 2845: 2843: 2841: 2837: 2832: 2828: 2824: 2820: 2816: 2812: 2808: 2804: 2800: 2796: 2789: 2786: 2781: 2775: 2760: 2756: 2752: 2748: 2744: 2740: 2736: 2732: 2728: 2724: 2720: 2716: 2712: 2705: 2702: 2697: 2695:9780585252728 2691: 2687: 2686: 2678: 2675: 2670: 2666: 2662: 2658: 2654: 2650: 2646: 2642: 2641:Theory Biosci 2635: 2632: 2627: 2623: 2619: 2615: 2611: 2607: 2600: 2598: 2594: 2589: 2585: 2580: 2575: 2571: 2567: 2562: 2557: 2553: 2549: 2548: 2543: 2536: 2533: 2528: 2524: 2520: 2516: 2512: 2508: 2507: 2499: 2496: 2484: 2480: 2476: 2469: 2467: 2463: 2458: 2454: 2450: 2446: 2441: 2436: 2432: 2428: 2424: 2420: 2416: 2409: 2406: 2401: 2397: 2393: 2389: 2385: 2381: 2377: 2373: 2372: 2364: 2361: 2356: 2352: 2348: 2344: 2340: 2336: 2332: 2328: 2324: 2320: 2319: 2311: 2304: 2301: 2295: 2290: 2287: 2284: 2283:deuterostomes 2280: 2277: 2276: 2272: 2270: 2268: 2267: 2262: 2258: 2257: 2252: 2251: 2246: 2241: 2239: 2235: 2227: 2225: 2223: 2219: 2215: 2214:sea pineapple 2211: 2207: 2206: 2201: 2197: 2190: 2186: 2182: 2181: 2176: 2171: 2163: 2161: 2158: 2150: 2147: 2144: 2141: 2140: 2139: 2138:, including: 2137: 2129: 2125:Use by humans 2124: 2122: 2120: 2116: 2115:ballast water 2112: 2107: 2105: 2101: 2097: 2093: 2092: 2087: 2086: 2081: 2077: 2073: 2072: 2067: 2063: 2062: 2057: 2056: 2047: 2045: 2043: 2039: 2035: 2031: 2027: 2026:multinucleate 2023: 2019: 2015: 2011: 2007: 2006: 1998: 1996: 1994: 1993:B. schlosseri 1989: 1988:B. schlosseri 1985: 1984: 1979: 1977: 1972: 1968: 1967: 1959: 1957: 1955: 1951: 1947: 1943: 1939: 1934: 1932: 1928: 1924: 1923: 1919:, except for 1918: 1914: 1909: 1907: 1903: 1899: 1895: 1891: 1886: 1884: 1880: 1876: 1872: 1868: 1863: 1855: 1851: 1847: 1845: 1844:metamorphosis 1839: 1837: 1833: 1829: 1825: 1821: 1817: 1813: 1809: 1798: 1791: 1789: 1787: 1783: 1779: 1778: 1773: 1772:cyanobacteria 1769: 1765: 1761: 1757: 1756: 1751: 1746: 1743: 1739: 1735: 1731: 1727: 1723: 1719: 1715: 1711: 1706: 1701: 1697: 1689: 1688: 1683: 1682: 1677: 1670: 1664: 1663: 1656: 1651: 1647: 1646: 1638: 1633: 1629: 1628: 1620: 1615: 1611: 1610: 1603: 1598: 1596: 1593: 1589: 1585: 1580: 1576: 1574: 1569: 1565: 1561: 1557: 1556:deuterostomes 1553: 1548: 1546: 1542: 1538: 1534: 1530: 1526: 1522: 1518: 1517:sulfuric acid 1514: 1510: 1506: 1502: 1498: 1494: 1490: 1486: 1481: 1479: 1475: 1471: 1467: 1462: 1459: 1455: 1451: 1446: 1441: 1439: 1435: 1431: 1427: 1423: 1419: 1415: 1411: 1407: 1403: 1399: 1395: 1383: 1373: 1366: 1364: 1361: 1357: 1353: 1348: 1345: 1341: 1336: 1332: 1328: 1323: 1321: 1316: 1312: 1308: 1307:blood vessels 1304: 1300: 1296: 1291: 1287: 1283: 1275: 1273: 1271: 1267: 1258: 1251: 1246: 1244: 1242: 1238: 1234: 1233:panarthropods 1230: 1226: 1218: 1216: 1213: 1211: 1205: 1203: 1199: 1195: 1194: 1189: 1185: 1181: 1180: 1175: 1171: 1167: 1166: 1161: 1157: 1156: 1151: 1146: 1144: 1140: 1137: 1133: 1129: 1128:bioimmuration 1125: 1121: 1117: 1114: 1110: 1109: 1100: 1096: 1095:bioimmuration 1092: 1091: 1085: 1079:Fossil record 1078: 1071: 1070: 1062: 1061: 1053: 1052: 1049: 1039: 1038: 1030: 1029: 1021: 1020: 1017: 1015: 1011: 1005: 1004: 1001: 1000: 997: 995: 991: 985: 984: 978: 977: 974: 973: 970: 968: 964: 958: 957: 951: 948: 947: 941: 940: 937: 936: 928: 927: 919: 918: 915: 913: 908: 902: 901: 898: 897: 894: 892: 888: 882: 881: 874: 873: 870: 869: 861: 860: 857: 855: 851: 845: 844: 841: 840: 837: 835: 831: 825: 824: 821: 820: 817: 815: 811: 805: 804: 798: 795: 794: 788: 787: 780: 779: 776: 775: 767: 766: 758: 757: 754: 752: 748: 742: 741: 738: 737: 734: 733: 727: 726: 720: 719: 716: 715: 712: 710: 706: 705:Oikopleuridae 700: 699: 693: 690: 689: 683: 680: 679: 675: 672: 670: 666: 662: 654: 650: 647: 644: 640: 636: 632: 629: 626: 622: 618: 614: 611: 610: 609: 606: 604: 600: 595: 593: 592:invertebrates 589: 588: 583: 582:hemichordates 579: 575: 571: 567: 563: 559: 551: 549: 545: 542: 538: 533: 531: 530: 525: 521: 517: 513: 509: 505: 497: 496: 491: 484: 483: 478: 471: 469: 467: 463: 459: 455: 450: 448: 444: 440: 435: 433: 429: 425: 421: 417: 413: 409: 405: 401: 396: 394: 390: 386: 382: 378: 374: 370: 366: 362: 358: 357: 344: 305: 302: 298: 294: 285: 278: 275: 271: 266: 265: 258: 256: 255: 248: 245: 244: 237: 235: 232: 230: 227: 224: 221: 220: 218: 215: 210: 205: 200: 197: 194: 193: 190: 187: 184: 181: 180: 177: 174: 171: 170: 167: 164: 161: 160: 157: 154: 151: 150: 145: 140: 136: 132: 131: 125: 121: 116: 111: 104: 99: 94: 89: 84: 79: 74: 69: 64: 59: 54: 49: 43: 35: 31: 22: 19: 7263: 7236:Potentially 7231: 7184: 7160: 7154:Gymnolaemata 7149:Stenolaemata 7138:Ectoproctans 7136: 7107: 7078: 7054: 7048:Pterobranchs 7035: 7011: 6999: 6987: 6963: 6939: 6933:within phyla 6931:Major groups 6801: 6788:Bryozoa s.l. 6779:Lophophorata 6767: 6743: 6721: 6713: 6676: 6665:Orthonectida 6640: 6587: 6580: 6569: 6488: 6481: 6410: 6368:Scalidophora 6317: 6305:Ambulacraria 6292: 6205:(see below↓) 6185: 6177: 6141: 6092:Opisthokonta 5951: 5944: 5930: 5877:Aves (birds) 5819:Lepidosauria 5679:Euteleostomi 5623:Cyclostomata 5510: 5458:Superphylum 5393: 5374: 5370: 5326: 5322: 5318: 5312: 5287: 5283: 5279: 5273: 5236: 5232: 5226: 5218: 5204:cite journal 5153: 5149: 5143: 5134: 5093: 5090:J. Mol. Evol 5089: 5085: 5081: 5075: 5040: 5037:Trends Genet 5036: 5027: 5019: 5007:. Retrieved 5002: 4993: 4981:. Retrieved 4977:the original 4973:"Sea squirt" 4967: 4929:(1): 43–61. 4926: 4922: 4912: 4900:. Retrieved 4890: 4863: 4857: 4848: 4838: 4826:. Retrieved 4816: 4804:. Retrieved 4800:the original 4774:. Retrieved 4770:the original 4760: 4733: 4729: 4719: 4686: 4682: 4637: 4603: 4599: 4546: 4542: 4532: 4520:. Retrieved 4509: 4492: 4488: 4482: 4457: 4449: 4437:. Retrieved 4412: 4367: 4361: 4355: 4347: 4312: 4306: 4296: 4261: 4257: 4247: 4202: 4196: 4186: 4151: 4147: 4141: 4100: 4096: 4092: 4086: 4066: 4059: 4034: 3984: 3980: 3970: 3933: 3929: 3919: 3884: 3881:Open Biology 3880: 3870: 3835: 3825: 3800: 3796: 3786: 3738:(1): 14326. 3735: 3731: 3720: 3703: 3699: 3693: 3660: 3656: 3646: 3633: 3621:. Retrieved 3617:the original 3607: 3584: 3575: 3565: 3553:. Retrieved 3546:the original 3541: 3537: 3524: 3479: 3473: 3463: 3426: 3422: 3412: 3375: 3371: 3361: 3336: 3332: 3319: 3282: 3278: 3220: 3216: 3206: 3189: 3186:Can. J. Zool 3185: 3179: 3170: 3133: 3130:BMC Genomics 3129: 3123: 3115: 3080: 3076: 3066: 3025: 3021: 2995:25 September 2993:. Retrieved 2986: 2979: 2969:25 September 2967:. Retrieved 2963: 2960:"Sea squirt" 2954: 2944:25 September 2942:. Retrieved 2932: 2920: 2908: 2897: 2878: 2798: 2794: 2788: 2774:cite journal 2762:. Retrieved 2718: 2714: 2704: 2684: 2677: 2644: 2640: 2634: 2609: 2605: 2551: 2545: 2535: 2510: 2504: 2498: 2486:. 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Index

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
Appendicularia

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