Enterogona

Aplidium solidum, copyright James Watanabe.

Belongs within: Urochordata.
Contains: Didemnidae.

The Enterogona are a group of tunicates united by features of the larva, in which the atrial cavity develops from paired dorsal invaginations.

<==Enterogona
    |--Phlebobranchiata S79
    |    |--Rhopalaea MG-H11 [Diazonidae S79]
    |    |    `--R. crassa (Herdman 1880) MG-H11
    |    |--Perophora [Perophoridae] S79
    |    |    `--P. listeri M62
    |    |--‘Agnesia’ Michaelsen 1898 non de Koninck 1883 [Agnesiidae] M86
    |    |    `--A. tenue Monniot & Monniot 1983 M86
    |    |--Ciona [Cionidae] HJ08
    |    |    |--C. intestinalis (Linnaeus 1767) HJ08
    |    |    `--C. savignyi J-GP07
    |    |--Corellidae M86
    |    |    |--Dextrogaster Monniot 1962 M86
    |    |    |    `--D. suecica Monniot 1962 M86
    |    |    `--Corella Alder & Hancock 1870 M86
    |    |         |--C. halli Kott 1951 M86
    |    |         `--C. parallellogramma BK77
    |    `--Ascidiidae HJ08
    |         |--Ascidiella aspersa (Mueller 1776) HJ08
    |         |--Psammascidia Monniot 1962 M86
    |         |    `--P. teissieri Monniot 1962 M86
    |         `--Ascidia CGS00
    |              |--A. callosa K92
    |              |--A. canina BK77
    |              |--A. ceratodes CGS00
    |              |--A. mentula Müller 1776 W01
    |              `--A. nigra S79
    `--Aplousobranchiata S79
         |--Didemnidae S79
         |--Clavelina MG-H11 [Clavelinidae S79]
         |    |--C. gemmae Turón 2005 F05
         |    |--C. lepadiformis CH97
         |    `--C. meridionalis (Herdman 1891) MG-H11
         `--Polyclinidae S79
              |--Parascidia turbinata (see below for synonymy) M62
              |--Polyclinum constellatum H04
              |--Amaroucium M62
              |    |--A. constellatum B79
              |    `--A. pellucidum B79
              |--Synoicum MG-H11
              |    |--S. chrysanthemum Kott 1992 MG-H11
              |    |--S. implicatum Kott 2008 MG-H11
              |    |--S. laboutei Monniot & Monniot 2006 MG-H11
              |    |--S. macroglossum (Hartmeyer 1919) MG-H11
              |    |--S. obscurum Kott 1992 MG-H11
              |    |--S. pseudogrisiatum Kott 2008 MG-H11
              |    |--S. rapum Kott 2008 MG-H11
              |    |--S. sphinctorum Kott 2006 MG-H11
              |    `--S. vesica Kott 2008 MG-H11
              `--Aplidium MG-H11
                   |--A. caelestis Monniot 1987 MG-H11
                   |--A. clivosum Kott 1992 MG-H11
                   |--A. crateriferum (Sluiter 1909) MG-H11
                   |--A. crustum Kott 2008 MG-H11
                   |--A. eudistomum Kott 2008 MG-H11
                   |--A. hyacinthum Kott 2008 MG-H11
                   |--A. magnilarvum Kott 1992 MG-H11
                   |--A. panis Kott 2008 MG-H11
                   |--A. solidum (Herdman 1891) MG-H11
                   `--A. tuberosum Kott 2008 MG-H11

Enterogona incertae sedis:
  Sycozoa [Holozoidae] MG-H11
    `--S. seiziwadai Tokioka 1952 MG-H11
  Polycitoridae MG-H11
    |--Cystodytes dellachiajei (Della Valle 1877) MG-H11
    |--Eucoelium coronarium (Monniot 1988) MG-H11
    `--Eudistoma MG-H11
         |--E. convocatum Kott 2008 MG-H11
         `--E. eboreum Kott 1990 MG-H11
  Protopolyclinidae MG-H11
    |--Pseudodiazona longigona (Tokioka 1959) MG-H11
    `--Condominium MG-H11
         |--C. areolatum (Kott 1963) MG-H11
         `--C. floreum Kott 2008 MG-H11
  Pseudodistomidae MG-H11
    |--Pseudodistoma australe Kott 1957 MG-H11
    `--Sigillina MG-H11
         |--S. australis Savigny 1816 MG-H11
         |--S. cyanea (Herdman 1899) MG-H11
         `--S. grandissima Kott 1990 MG-H11

Parascidia turbinata [=Sydnium turbinatum; incl. Circinalium concrescens, Parascidia flemingi, Pa. forbesi, Amaroucium rubicondum, Polyclinum succineum] M62

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[B79] Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: why colonial animals have more hiding places. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 459–497. Academic Press: London.

[CGS00] Cameron, C. B., J. R. Garey & B. J. Swalla. 2000. Evolution of the chordate body plan: new insights from phylogenetic analyses of deuterostome phyla. Proceedings of the National Academy of Sciences of the USA 97 (9): 4469–4474.

[CH97] Castro, P., & M. E. Huber. 1997. Marine Biology 2^nd ed. WCB McGraw-Hill: Boston.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[HJ08] Huisman, J. M., D. S. Jones, F. E. Wells & T. Burton. 2008. Introduced marine biota in Western Australian waters. Records of the Western Australian Museum 25 (1): 1–44.

[J-GP07] Jiménez-Guri, E., H. Philippe, B. Okamura & P. W. H. Holland. 2007. Buddenbrockia is a cnidarian worm. Science 317: 116–118.

[K92] Kozloff, E. N. 1992. The genera of the phylum Orthonectida. Cahiers de Biologie Marine 33: 377–406.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[M62] Monniot, F. 1962. Recherches sur les graviers a Amphioxus de la région de Banyuls-sur-Mer. Vie et Milieu 13: 231–322.

[M86] Monniot, F. 1986. Urochordata=Tunicata: les ascidies interstitielles. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 705–708. E. J. Brill/Dr. W. Backhuys: Leiden.

[S79] Sabbadin, A. 1979. Colonial structure and genetic patterns in ascidians. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 433–444. Academic Press: London.

[W01] Williamson, D. I. 2001. Larval transfer and the origins of larvae. Zoological Journal of the Linnean Society 131: 111–122.

Urochordata

Colony of Salpa fusiformis, copyright Mark Rosenstein.

Belongs within: Chordata.
Contains: Enterogona, Pleurogona.

The Urochordata, tunicates, are spineless chordates that possess a notochord and postanal tail in the tadpole-like larval stage but usually loose them when they metamorphose into adults.

Characters (from Miller & Harley 1996): Notochord, nerve cord and postanal tail present in free-swimming larvae; adults sessile or planktonic, usually enclosed in a tunic containing some cellulose.

<==Urochordata
    |  i. s.: Octacnemus [Sorberacea] MH96
    |--Shankouclava Chen, Huang et al. 2003 EL11, CH03
    |    `--*S. anningense Chen, Huang et al. 2003 [=S. shankouense Chen, Huang et al. 2003 (n. n.)] CH03
    `--+--Oikopleura GD00 [Appendicularia C-S98, Larvacea]
       |    |--O. dioica LG03
       |    |--O. labradoriensis B26
       |    `--O. vanhoeffeni B26
       `--Tunicata C-S98
            |--Thaliacea [Thaliae] C-S98
            |    |  i. s.: Thalia democratica BN03
            |    |--Pyrosoma [Pyrosomatidae, Pyrosomida] MG-H11
            |    |    |--P. atlanticum Péron 1804 MG-H11
            |    |    `--P. spinosum MH96
            |    |--Doliolidae [Dolioloda] MG-H11
            |    |    |--Dolioletta gegenbauri (Uljanin 1884) MG-H11
            |    |    `--Doliolum W01
            |    |         |--D. mulleri Krohn 1852 W01
            |    |         `--D. nationalis CGS00
            |    `--Salpa [Salpida, Salpidae] MG-H11
            |         |--S. fusiformis B26
            |         |--S. maxima Forskål 1775 MG-H11
            |         |--S. tilesii B26
            |         `--S. zonaria B26
            `--Ascidiacea [Ascidiae] C-S98
                 |  i. s.: Clarelina molucensis RZ03
                 |         Phallusia CGS00
                 |           |--P. mamillata CGS00
                 |           `--P. nigra B79
                 |         Polyandrocarpa B79
                 |           |--P. gravei B79
                 |           |--P. tincta B79
                 |           `--P. zorritensis GO78
                 |         Symplegma viride B79
                 |         Diplosoma listerianum B79
                 |         Ctenicella appendiculata M62
                 |         Distomus variolosus M62
                 |         Asterocarpa coerulea HS01
                 |         Permosoma EL11
                 |         Cynthia melocactus H04
                 |         Fragarium elegans H04
                 |         Synoecum turgens H04
                 |         Polycyclus cyaneus H04
                 |--Enterogona S79
                 `--Pleurogona S79

*Type species of generic name indicated

REFERENCES

[B26] Bigelow, H. B. 1926. Plankton of the offshore waters of the Gulf of Maine. Bulletin of the Bureau of Fisheries 40 (2): 1–509.

[BN03] Bourlat, S. J., C. Nielsen, A. E. Lockyer, D. T. J. Littlewood & M. J. Telford. 2003. Xenoturbella is a deuterostome that eats molluscs. Nature 424: 925–928.

[B79] Buss, L. W. 1979. Habitat selection, directional growth and spatial refuges: why colonial animals have more hiding places. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 459–497. Academic Press: London.

[CGS00] Cameron, C. B., J. R. Garey & B. J. Swalla. 2000. Evolution of the chordate body plan: new insights from phylogenetic analyses of deuterostome phyla. Proceedings of the National Academy of Sciences of the USA 97 (9): 4469–4474.

[C-S98] Cavalier-Smith, T. 1998. A revised six-kingdom system of life. Biological Reviews 73: 203–266.

[CH03] Chen, J.-Y., D.-Y. Huang, Q.-Q. Peng, H.-M. Chi, Z.-Q. Wang & M. Feng. 2003. The first tunicate from the Early Cambrian of South China. Proceedings of the National Academy of Sciences of the USA 100 (14): 8314–8318.

[EL11] Erwin, D. H., M. Laflamme, S. M. Tweedt, E. A. Sperling, D. Pisani & K. J. Peterson. 2011. The Cambrian conundrum: early divergence and later ecological success in the early history of animals. Science 334: 1091–1097.

[GO78] Gallardo S., C., & C. Osorio R. 1978. Hiatella solida (Sowerby, 1834) (Mollusca: Hiatellidae) on Concholepas concholepas (Bruguière, 1789) and other substrates. Veliger 20 (3): 274–278.

[GD00] Giribet, G., D. L. Distel, M. Polz, W. Sterrer & W. C. Wheeler. 2000. Triploblastic relationships with emphasis on the acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: A combined approach of 18S rDNA sequences and morphology. Systematic Biology 49: 539–562.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[LG03] Lancelot, C., & R. Groben. 2003. Meeting report: Regional conference on the ecology and impact assessment of Phaeocystis blooms in the Eastern Channel and Southern Bight of the North Sea. Protist 154 (2): 173–176.

]MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[MH96] Miller, S. A., & J. P. Harley. 1996. Zoology 3^rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[M62] Monniot, F. 1962. Recherches sur les graviers a Amphioxus de la région de Banyuls-sur-Mer. Vie et Milieu 13: 231–322.

[RZ03] Romanenko, L. A., N. V. Zhukova, M. Rhode, A. M. Lysenko, V. M. Mikhailov & E. Stackebrandt. 2003. Pseudoalteromonas agarivorans sp. nov., a novel marine agarolytic bacterium. International Journal of Systematic and Evolutionary Microbiology 53: 125–131.

[S79] Sabbadin, A. 1979. Colonial structure and genetic patterns in ascidians. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 433–444. Academic Press: London.

[W01] Williamson, D. I. 2001. Larval transfer and the origins of larvae. Zoological Journal of the Linnean Society 131: 111–122.

Pleurogona

Sea grape Molgula manhattensis, copyright Wilfried Bay-Nouailhat.

Belongs within: Urochordata.

The Pleurogona are a group of tunicates characterised by a folded pharyngeal basket, and the lack of an abdomen.

<==Pleurogona [Stolidobranchiata]
    |--Molgulidae MG-H11
    |    |--Eugyra arenosa M62
    |    |--Bostrichobranchus SM10
    |    `--Molgula Forbes & Hanley 1848 M86
    |         |--M. citrina SM10
    |         |--M. elva Kott 2008 MG-H11
    |         |--M. hirta Monniot 1965 M86
    |         |--M. manhattensis (De Kay 1843) L70
    |         |--M. occidentalis SM10
    |         |--M. oculata CGS00
    |         |--M. retortiformis SM10
    |         |--M. tectiformis J-GP07
    |         `--M. tubulosa H04
    |--Pyuridae MG-H11
    |    |--Cratostigma gravellophila M62
    |    |--Halocynthia SM10
    |    |    |--H. aurantiaca RZ03
    |    |    `--H. roretzi SM10
    |    |--Pyura MG-H11
    |    |    |--P. chilensis GO78
    |    |    |--P. pachydermatina KG03
    |    |    |--P. scortea Kott 1985 MG-H11
    |    |    `--P. stolonifera G75
    |    |--Herdmania MG-H11
    |    |    |--H. curvata CGS00
    |    |    |--H. grandis (Heller 1878) MG-H11
    |    |    `--H. momus (Savigny 1816) MG-H11
    |    `--Heterostigma Ärnbäck-Christie-Linde 1924 M86
    |         |--H. fagei Monniot & Monniot 1961 M86
    |         |--H. gonochorica Monniot 1965 M86
    |         |--H. reptans Monniot & Monniot 1963 M86
    |         `--H. separ Ärnbäck-Christie-Linde 1924 M86
    `--Styelidae MG-H11
         |  i. s.: Polycarpa Heller 1877 M86
         |           |--P. argentata (Sluiter 1890) MG-H11
         |           |--P. arnbackae Monniot 1964 M86
         |           |--P. aurita (Sluiter 1890) MG-H11
         |           |--P. chinensis (Tokioka 1967) MG-H11
         |           |--P. decipiens Herdman 1906 MG-H11
         |           |--P. pentarhiza Monniot 1965 M86
         |           `--P. viridis Herdman 1881 MG-H11
         |         Cnemidocarpa MG-H11
         |           |--C. bicornuata HS01
         |           |--C. completa Kott 1985 MG-H11
         |           `--C. radicosa (Herdman 1882) MG-H11
         |         Stolonica aluta Kott 1985 MG-H11
         |--Botryllinae S79
         |    |--Botrylloides S79
         |    |    |--B. leachi (Savigny 1816) HJ08
         |    |    `--B. purpureus H04
         |    `--Botryllus HJ08
         |         |--B. helleborus H04
         |         |--B. marionis H04
         |         |--B. polycyclus H04
         |         |--B. primigenius S79
         |         |--B. rubigo H04
         |         `--B. schlosseri (Pallas 1766) HJ08
         `--Styelinae M62
              |--Psammostyela Weinstein 1961 M62, M86
              |    `--P. delamarei Weinstein 1961 M86
              `--Styela MG-H11
                   |--S. clava Herdman 1881 HJ08 MG-H11
                   `--S. plicata (Lesueur 1823) MG-H11

*Type species of generic name indicated

REFERENCES

[CGS00] Cameron, C. B., J. R. Garey & B. J. Swalla. 2000. Evolution of the chordate body plan: new insights from phylogenetic analyses of deuterostome phyla. Proceedings of the National Academy of Sciences of the USA 97 (9): 4469–4474.

[GO78] Gallardo S., C., & C. Osorio R. 1978. Hiatella solida (Sowerby, 1834) (Mollusca: Hiatellidae) on Concholepas concholepas (Bruguière, 1789) and other substrates. Veliger 20 (3): 274–278.

[G75] Grant, E. M. 1975. Guide to Fishes. The Co-ordinator-General’s Department: Brisbane (Australia).

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

[HJ08] Huisman, J. M., D. S. Jones, F. E. Wells & T. Burton. 2008. Introduced marine biota in Western Australian waters. Records of the Western Australian Museum 25 (1): 1–44.

[J-GP07] Jiménez-Guri, E., H. Philippe, B. Okamura & P. W. H. Holland. 2007. Buddenbrockia is a cnidarian worm. Science 317: 116–118.

[KG03] Kingsley, R. J., R. van Gilder, R. Z. LeGros & N. Watabe. 2003. Multimineral calcareous deposits in the marine alga Acetabularia acetabulum (Chlorophyta; Dasycladaceae). Journal of Phycology 39: 937–947.

[L70] Leloup, E. 1970. Recherches sur l’ostreiculture dans le bassin de chasse d’Ostende en 1968. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 46 (6): 1–24.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[M62] Monniot, F. 1962. Recherches sur les graviers a Amphioxus de la région de Banyuls-sur-Mer. Vie et Milieu 13: 231–322.

[M86] Monniot, F. 1986. Urochordata=Tunicata: les ascidies interstitielles. In: Botosaneanu, L.(ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 705–708. E. J. Brill/Dr. W. Backhuys: Leiden.

[RZ03] Romanenko, L. A., N. V. Zhukova, M. Rhode, A. M. Lysenko, V. M. Mikhailov & E. Stackebrandt. 2003. Pseudoalteromonas agarivorans sp. nov., a novel marine agarolytic bacterium. International Journal of Systematic and Evolutionary Microbiology 53: 125–131.

[S79] Sabbadin, A. 1979. Colonial structure and genetic patterns in ascidians. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 433–444. Academic Press: London.

[SM10] Saffo, M. B., A. M. McCoy, C. Rieken & C. H. Slamovits. 2010. Nephromyces, a beneficial apicomplexan symbiont in marine animals. Proceedings of the National Academy of Sciences of the USA 107 (37): 16190–16195.

Didemnidae

Didemnum candidum, copyright marine_p.

Belongs within: Enterogona.

The Didemnidae are a group of tunicates whose small zooids form encrusting colonies.

<==Didemnidae
    |--Lissoclinum maculatum Kott 2001 MG-H11
    |--Atriolum MG-H11
    |    |--A. bucinum Kott 2001 MG-H11
    |    `--A. tubiporum Kott 2001 MG-H11
    |--Trididemnum MG-H11
    |    |--T. grandistellatum Kott 2008 MG-H11
    |    `--T. sibogae (Hartmeyer 1910) MG-H11
    |--Leptoclinides MG-H11
    |    |--L. fungiformis Kott 1972 MG-H11
    |    |--L. multilobatus Kott 1954 MG-H11
    |    `--L. vesica Kott 2008 MG-H11
    |--Polysyncraton MG-H11
    |    |--P. calculum Kott 2008 MG-H11
    |    |--P. dealbatum Kott 2008 MG-H11
    |    |--P. fistulum Kott 2008 MG-H11
    |    |--P. millepore Vaseur 1969 MG-H11
    |    |--P. palliolum Kott 2001 MG-H11
    |    |--P. pavimentum Monniot 1993 MG-H11
    |    `--P. solidum MG-H11
    `--Didemnum MG-H11
         |--D. candidum Savigny 1816 MG-H11
         |--D. cygnus Kott 2001 MG-H11
         |--D. dolium Kott 2008 MG-H11
         |--D. jedanense Sluiter 1909 MG-H11
         |--D. moseleyi (Herdman 1886) MG-H11
         |--D. multiampullae Kott 2008 MG-H11
         |--D. ossium Kott 2001 MG-H11
         |--D. perplexum Kott 2001 MG-H11
         |--D. plebeium Kott 2005 MG-H11
         |--D. tapetum Kott 2008 MG-H11
         `--D. velum Kott 2008 MG-H11

*Type species of generic name indicated

REFERENCES

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1-191.