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Tunicate
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=== Classification === Tunicates are more closely related to [[craniate]]s (including [[hagfish]], [[lamprey]]s, and jawed [[vertebrate]]s) than to [[lancelet]]s, [[echinoderm]]s, [[hemichordate]]s, ''[[Xenoturbella]]'' or other [[invertebrate]]s.<ref name="pmid16495997">{{cite journal |author1=Delsuc, F. |author2=Brinkmann, H. |author3=Chourrout, D. |author4=Philippe, H. |title=Tunicates and not cephalochordates are the closest living relatives of vertebrates |journal=Nature |volume=439 |issue=7079 |pages=965–968 |year=2006 |pmid=16495997 |doi=10.1038/nature04336 |bibcode=2006Natur.439..965D |s2cid=4382758 |url=http://hal-sde.archives-ouvertes.fr/docs/00/31/54/36/PDF/Delsuc-Nature06_HAL.pdf }}</ref><ref name="pmid19003928">{{cite journal |author1=Delsuc, F. |author2=Tsagkogeorga, G. |author3=Lartillot, N. |author4=Philippe, H. |title=Additional molecular support for the new chordate phylogeny |journal=Genesis |volume=46 |issue=11 |pages=592–604 | year=2008 |pmid=19003928 |doi=10.1002/dvg.20450 |s2cid=205771088 |url=https://hal.archives-ouvertes.fr/halsde-00338411 |doi-access=free }}</ref><ref name="pmid19922605">{{cite journal |author1=Singh, T. R. |author2=Tsagkogeorga, G. |author3=Delsuc, F. |author4=Blanquart, S. |author5=Shenkar, N. |author6=Loya, Y. |author7=Douzery, E. J. |author8=Huchon, D. |title=Tunicate mitogenomics and phylogenetics: peculiarities of the ''Herdmania momus'' mitochondrial genome and support for the new chordate phylogeny |journal=BMC Genomics |volume=10 |page=534 |year=2009 |pmid=19922605 |doi=10.1186/1471-2164-10-534 |pmc=2785839 |doi-access=free }}</ref> The [[clade]] consisting of tunicates and vertebrates is called [[Olfactores]].<ref>Jefferies, R. P. S. (1991) in Biological Asymmetry and Handedness (eds Bock, G. R.; Marsh, J.) pp. 94–127 (Wiley, Chichester).</ref> The Tunicata contain roughly 3,051 described species,<ref name=Ruppert/> traditionally divided into these classes: * [[Ascidiacea]] ([[Aplousobranchia]], [[Phlebobranchia]], and [[Stolidobranchia]]) * [[Thaliacea]] ([[Pyrosome|Pyrosomida]], [[Doliolida]], and [[Salp]]ida) * [[Larvacean|Appendicularia]] ([[Copelata]]) Members of the [[Sorberacea]] were included in Ascidiacea in 2011 as a result of [[ribosomal DNA|rDNA]] sequencing studies.<ref name="Tatian">{{cite journal |author1=Tatián, Marcos |author2=Lagger, Cristian |author3=Demarchi, Milagros |author4=Mattoni, Camilo |year=2011 |title=Molecular phylogeny endorses the relationship between carnivorous and filter-feeding tunicates (Tunicata, Ascidiacea) |journal=Zoologica Scripta |volume=40 |issue=6 |pages=603–612 |doi=10.1111/j.1463-6409.2011.00493.x |s2cid=86421513 }}</ref> Although the traditional classification is provisionally accepted, newer evidence suggests the Ascidiacea are an artificial group of [[paraphyletic]] status.<ref name="Zeng2005">{{cite journal |author1=Zeng, L. |author2=Swalla, B. J. |title=Molecular phylogeny of the protochordates: chordate evolution |journal=Can. J. Zool. |volume=83 |pages=24–33 |year=2005 |doi=10.1139/z05-010 }}</ref><ref name="pmid19656395">{{cite journal |author1=Tsagkogeorga, G. |author2=Turon, X. |author3=Hopcroft, R. R. |author4=Tilak, M. K. |author5=Feldstein, T. |author6=Shenkar, N. |author7=Loya, Y. |author8=Huchon, D. |author9=Douzery, E. J. |author10=Delsuc, F. |title=An updated 18S rRNA phylogeny of tunicates based on mixture and secondary structure models|journal=BMC Evolutionary Biology |volume=9 |page=187 |year=2009 |issue=1 |pmid=19656395 |doi=10.1186/1471-2148-9-187 |pmc=2739199 |bibcode=2009BMCEE...9..187T |doi-access=free }}</ref><ref name=Delsuc2018>{{cite journal |vauthors=Delsuc F, Philippe H, Tsagkogeorga G, Simion P, Tilak MK, Turon X, López-Legentil S, Piette J, Lemaire P, Douzery EJ |date=April 2018 |title=A phylogenomic framework and timescale for comparative studies of tunicates |journal=BMC Biology |volume=16 |issue=1 |page=39 |doi=10.1186/s12915-018-0499-2 |pmc=5899321 |pmid=29653534 |doi-access=free }}</ref> 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.<ref>{{cite journal |url=https://core.ac.uk/download/pdf/39301635.pdf |title=Phylogenetic Speculation of the Tunicata |last1=Tokioka |first1=Takasi |date=1971-06-30 |journal=Publications of the Seto Marine Biological Laboratory |volume=19 |issue=1 |page=47 |doi=10.5134/175655 |s2cid=55491438 }}</ref> The following cladogram is based on the 2018 phylogenomic study of Delsuc and colleagues.<ref>{{Cite journal |last1=Franchi |first1=Nicola |last2=Ballarin |first2=Loriano |year=2017 |title=Immunity in Protochordates: The Tunicate Perspective |journal=Frontiers in Immunology |volume=8 |page=674 |doi=10.3389/fimmu.2017.00674 |pmc=5465252 |pmid=28649250 |doi-access=free}}</ref><ref name=Delsuc2018/><ref>{{Cite journal |last=Giribet |first=Gonzalo |year=2018 |title=Phylogenomics resolves the evolutionary chronicle of our squirting closest relatives |journal=BMC Biology |volume=16 |issue=1 |page=49 |doi=10.1186/s12915-018-0517-4 |pmc=5924484 |pmid=29703197 |doi-access=free}}</ref> {{clade |label1='''Tunicata''' |1={{clade |label1=[[Larvacean|Appendicularia]] |1={{clade |1=[[Oikopleuridae]] [[File:Oikopleura_dioica.gif|40 px]] |2={{clade |1=[[Kowalevskiidae]] |2=[[Fritillariidae]] [[File:Appendicularia_%28YPM_IZ_096169%29.jpeg|60 px]] }} }} |label2=Acopa |2={{clade |1={{clade |label1=[[Thaliacea]] |1={{clade |1=[[Pyrosomida]] [[File:Pyrosoma_atlanticum.JPG|60 px]] |2=[[Salpida]] [[File:Salp_colony,_Aorangaia_PA171899.JPG|60 px]] |3=[[Doliolida]] [[File: Cyclomyaria.jpg|60 px]] }} |label2=Enterogona |2={{clade |1=[[Phlebobranchia]] [[File: CionaintestinalisR.jpg|60 px]] |bar1=green |2=[[Aplousobranchia]] [[File: Sea_Squirts_Didemnum_molle.jpg|60 px]] |bar2=green }} }} |grouplabel2=[[Ascidiacea]] |grouplabelstyle2=vertical-align:top;|bar2=green |2={{clade |label1=[[Stolidobranchia]] |1={{clade |1=[[Molgulidae]] [[File: Molgula_oculata_001.png|60 px]] |2={{clade |1=[[Styelidae]] [[File:Tunicate_komodo.jpg|60 px]] |2=[[Pyuridae]] [[File:Microcosmus_sabatieri.jpg|60 px]] }} }} }} }} }} |style=font-size:100%;line-height:100%}}
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