Tylodina corticalis, from Sea Slug Forum.

Belongs within: Heterobranchia.
Contains: Acteonidae, Cephalaspidea, Acochlidiacea, Thecosomata, Gymnosomata, Aplysiomorpha, Oxynooidea, Placobranchacea, Euctenidiacea, Dexiarchia.

The Opisthobranchia are a group of gastropods exhibiting a tendency towards reduction of the shell and detorsion of the body, with the sea slugs completely missing the shell and appearing bilateral externally. Within the Opisthobranchia, the Nudipleura are distinguished as a clade including the shell-less nudibranchs and the shelled Pleurobranchidae by the possession of a blood gland and the absence of the osphradium. The remaining, often shelled, opisthobranchs are commonly classified as the Tectibranchia but this group is of uncertain monophyly.

Characters (from Cox 1960): Shell usually reduced or lost; anatomy euthyneurous, more or less detorted, externally bilaterally symmetrical, nerve ganglia usually concentrated around esophagus. Mantle cavity migrating to right and posteriorly before opening out; loss of mantle cavity associated with loss of pallial organs; secondary gills may be present. Gut more or less straight; stomach reduced, gizzard or crop may be present. Single auricle present; renal and reproductive systems asymmetrical. Radula multiseriate, teeth reduced in carnivores; esophageal glands and style-sac absent. Reproductive system invariably hermaphroditic; retractile penis usually present; pallial region of hermaphrodite duct typically divided longitudinally into male and female passages; albumen gland an appendage to female tract; fertilized eggs are laid in jelly-like masses.

Opisthobranchia (see below for synonymy)
    |--Tectibranchia [Anaspidea] APS-P86
    |    |--Cephalaspidea APS-P86
    |    |--Acochlidiacea APS-P86
    |    |--Thecosomata APS-P86
    |    |--Gymnosomata APS-P86
    |    |--Aplysiomorpha APS-P86
    |    |--Sacoglossa APS-P86
    |    |    |--Oxynooidea BR05
    |    |    `--Placobranchacea BR05
    |    |--Cylindrobulla Fischer 1857 (see below for synonymy) BR05
    |    |    |--C. fischeri Adams & Angas 1865 H09
    |    |    |--C. gigas Mikkelsen 1998 BC01
    |    |    `--C. pusilla Nevill 1869 H09
    |    `--Umbraculoidea [Umbraculacea, Umbraculida] BR05
    |         |--Tylodina Rafinesque 1814 [Tylodinacea, Tylodinana, Tylodinidae, Tylodininae] BR05
    |         |    |--*T. punctulata [incl. T. citrina] O27
    |         |    |--T. corticalis (Tate 1889) MG-H11
    |         |    `--T. fungina Gabb 1865 O27
    |         `--Umbraculidae BR05
    |              |--Operculatum Mörch 1852 [Operculatinae] BR05
    |              `--Umbraculum Schumacher 1817 [=Umbrella Lamarck 1819; Umbraculinae, Umbrellidae, Umbrellinae] BR05
    |                   |--‘Patella’ umbraculum Solander 1786 F27 (see below for synonymy)
    |                   `--U. botanicum Hedley 1923 F27
    `--Nudipleura [Eleutherobranchia, Eleutherobranchiatae] BR05
         |--Nudibranchia [Bifaribranchiata, Gnathodoridacea, Gnathodoridoidea, Hypobranchia, Hypobranchiata] BR05
         |    |  i. s.: Trapania rudmani HS01
         |    |         Rhodope Koelliker 1847 [Apneusta, Rhodopacea, Rhodopemorpha, Rhodopidae, Rhodopida] BR05
         |    |           `--*R. veranyi Kölliker 1847 APS-P86
         |    |         Polycerca quadrilineata D56
         |    |         Idalia elegans H99
         |    |--Euctenidiacea BR05
         |    `--Dexiarchia BR05
         `--Pleurobranchidae [Pleurobranchoidea, Pleurobranchomorpha, Subaplysiacea] BR05
              |  i. s.: Berthellina citrina (Rüppell & Leuckart 1828) BW09
              |         Pleurobranchidium BR05
              |         Bouvieria Vayssiere 1896 P61
              |           |--*B. aurantiaca (Risso 1818) [=Pleurobranchus aurantiacus] P61
              |           `--B. ornata (Cheeseman 1878) P61
              |--Pleurobranchaea Meckel 1813 [Pleurobranchaeidae, Pleurobranchaeinae] BR05
              |    |--*P. meckelii P61
              |    |--P. californica (MacFarland 1966) RD77
              |    |--P. maculata (Quoy & Gaimard 1832) P61
              |    |--P. novaezelandiae Cheeseman 1878 F27
              |    |    |--P. n. novaezelandiae P61
              |    |    `--P. n. granulosa Berg 1900 P61
              |    `--P. tarda B26
              `--Pleurobranchinae BR05
                   |--Bathyberthella Willan 1983 [Bathyberthellini] BR05
                   |--Berthella Blainville 1824 [Berthellinae, Berthellini] BR05
                   `--Pleurobranchus Cuvier 1804 [Pleurobrancheae, Pleurobranchini] BR05
                        |--*P. peronii O27
                        |--P. areolatus (Mörch 1863) PP78
                        |--P. californicus Dall 1900 O27
                        `--P. forskalii (Ruppell & Leuckart 1828) BW09

Opisthobranchia incertae sedis:
  Pneumodermopsis (Pneumodermopsis) paucidens (Boas 1866) H01
  Oscanius tuberculatus M62
  Acteonoidea BR05 [Acteonacea C60, Acteonidea, Pleurocoela]
    |--Acteonidae BR05
    |--Zardinella Bandel 1994 [Zardinellidae] BR05
    |--Aplustridae BR05
    |    |--Aplustrum Schumacher 1817 [=Amplustrum (l. c.); Amplustrina, Aplustrinae] BR05
    |    |    `--A. aplustre (Linnaeus 1758) [=Bulla aplustris] H09
    |    `--Hydatina Schumacher 1817 BR05
    |         `--H. physis (Linnaeus 1758) F27 [=Bulla physis H09]
    |--Bullinidae BR05
    |    |--Nonacteonina Stephenson 1941 [Nonacteoninidae] BR05
    |    |--Bullina Férussac 1822 BR05
    |    |    `--B. lineata (Gray 1825) HS01
    |    `--Sulcoactaeon Cossmann 1895 [Sulcoactaeonidae] BR05
    |         `--S. pulloides (Hudleston 1896) GK02
    `--Acteonellidae [Actaeonellidae] BR05
         |--Itieria Matheron 1842 [Itieriacea, Itieriidae, Itieriinae] BR05
         |--Cylindrobullina Ammon 1878 (see below for synonymy) BR05
         |    `--C. convexa Batten & Stokes 1986 NE02
         `--Acteonellinae BR05
              |--‘Orthostoma’ Deshayes 1850 (see below for synonymy) BR05
              |--Trochactaeon Meek 1863 [Trochactaeonidae, Trochactaeoninae] BR05
              `--Acteonella d’Orbigny 1843 [=Actaeonella (l. c.)] BR05
                   `--A. gigantea C60

Cylindrobulla Fischer 1857 [Cylindrobullacea, Cylindrobullida, Cylindrobullidae, Cylindrobullinae, Cylindrobulloidea] BR05

Cylindrobullina Ammon 1878 [Cylindrobullinidae, Cylindrobullininae, Cylindrobullinoidea] BR05

Opisthobranchia [Acera, Aceridae, Aiolobranchiata, Angiophora, Apneumonophora, Aporobranchiata, Aptera, Enterobranchiata, Notaspidea, Opisthobranchiata, Pomatobranchiata, Tetracerata]

‘Orthostoma’ Deshayes 1850 nec Ehrenberg 1831 nec Audinet-Serville 1834 nec Conrad 1838 [Orthostomatidae, Orthostomatoidea] BR05

‘Patella’ umbraculum Solander 1786 F27 [incl. P. umbrella Martyn 1786 H09, *Umbraculum umbrella P61]

*Type species of generic name indicated


[APS-P86] Arnaud, P. M., Cl. Poizat & L. v. Salvini-Plawen. 1986. Marine-interstitial Gastropoda (including one freshwater interstitial species).  In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 153–176. E. J. Brill/Dr. W. Backhuys: Leiden.

[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.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1-2): 1–397.

[BC01] Boyko, C. B., & J. R. Cordeiro. 2001. Catalog of Recent type specimens in the Division of Invertebrate Zoology, American Museum of Natural History. V. Mollusca, part 2 (class Gastropoda [exclusive of Opisthobranchia and Pulmonata], with supplements to Gastropoda [Opisthobranchia], and Bivalvia). Bulletin of the American Museum of Natural History 262: 1–170.

[BW09] Bryce, C., & C. Whisson. 2009. The macromolluscs of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 177–208.

[C60] Cox, L. R. 1960. Gastropoda: general characteristics of Gastropoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I84–I169. Geological Society of America, and University of Kansas Press.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[F27] Finlay, H. J. 1927. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[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.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[H01] Huys, R. 2001. Splanchnotrophid systematics: a case of polyphyly and taxonomic myopia. Journal of Crustacean Biology 21 (1): 106–156.

[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.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. II pt I. Stanford University Press: Stanford University (California).

[PP78] Poorman, F. L., & L. H. Poorman. 1978. Additional molluscan records from Bahía de Los Angeles, Baja California Norte. Veliger 20 (4): 369–374.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[RD77] Ram, J. L., & W. J. Davis. 1977. Mechanisms underlying "singleness of action" in the feeding behavior of Pleurobranchaea californica (MacFarland, 1966). Veliger 20 (1): 55–56.

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