Home Guide to Opisthobranchs Nudibranchia Doridina

Doridina

Doridina is one of the two major evolutionary lineages of Nudibranchia, together with Cladobranchia. It comprises the doridacean nudibranchs, a highly diverse group of sea slugs characterized by a compact body plan and a secondary gill plume arranged around the anus. Members of the group are commonly known as dorids or doridaceans.

Peltodoris atromaculata @ Costa Brava, Spain by Enric Madrenas by Enric Madrenas

Dorids generally have an oval, elongated, or dorsoventrally flattened body, with the dorsal notum clearly differentiated from the sides of the body. The rhinophores may be contractile and, in many species, can retract completely into specialized rhinophoral cavities. The dorsal surface may be smooth or bear tubercles of various shapes, including rounded, conical, or digitiform structures. In some genera, such as Jorunna, Taringa, and Rostanga, the tubercles contain conspicuous spicules that project from their margins and form the structures known as caryophyllidia.

The gill plume is located in the posterior region of the dorsal surface and surrounds the anal opening. The gills may consist of simple or branched leaves and may remain permanently exposed or retract into a specialized branchial cavity. Traditionally, dorids with exposed gills were referred to as phanerobranchs, whereas those capable of retracting their gills into a branchial cavity were referred to as cryptobranchs. However, anatomical and developmental studies have shown that this distinction does not represent a simple evolutionary sequence and that the evolution of the branchial cavity is more complex than previously considered.

The digestive gland of dorids is compact and consists of a variable number of lobes. The group exhibits considerable diversity in body ornamentation, coloration, feeding structures, reproductive anatomy, and chemical defence mechanisms.

Dorids are predominantly carnivorous, and many species exhibit a high degree of dietary specialization. Sponges constitute the main food source for numerous lineages, while other species feed on bryozoans, ascidians, cnidarians, or other sessile marine invertebrates. These specialized feeding relationships may be associated with the acquisition, modification, or synthesis of defensive chemical compounds.

The traditional classification of Doridina recognizes two major infraorders: Bathydoridoidei and Doridoidei. Bathydoridoidei comprises a small lineage of dorids predominantly associated with deep-water environments, whereas Doridoidei contains most of the group’s diversity and includes numerous shallow-water lineages. Within Doridoidei, the classification of Bouchet et al. (2017) recognizes the superfamilies Chromodoridoidea, Doridoidea, Onchidoridoidea, Phyllidioidea, and Polyceroidea, together with some taxa not assigned to a superfamily.

The taxonomic delimitation of Doridina has been the subject of recent debate. Korshunova et al. (2025) proposed separating this lineage and recognizing it as the order Doridida. Subsequently, Moles et al. (2026) advocated retaining the traditional classification and maintaining Doridina within Nudibranchia. The latter interpretation is followed here.

Doridina exhibits remarkable morphological and ecological diversity. Species range from relatively smooth forms with cryptic coloration to highly ornamented animals bearing tubercles, ridges, appendages, and conspicuous colour patterns. The group occupies primarily benthic environments and includes both shallow-water species associated with rocky substrates and benthic communities and specialized deep-water lineages.

Dietary specialization, morphological diversity, and the various chemical defence mechanisms of dorids have played an important role in their diversification and remain prominent areas of research in comparative anatomy, chemical ecology, development, and evolution.

The systematic classification of Doridina presented below is based on Bouchet et al. (2017) and its treatment in the World Register of Marine Species (WoRMS), with consideration of subsequent systematic revisions:

Doridina

  • Bathydoridoidei
    • Bathydoridoidea Bergh, 1891
      • Bathydorididae Bergh, 1891
        • Bathydoris Bergh, 1884
        • Prodoris Baranetz & Minichev, 1995
  • Doridoidei
    • Chromodoridoidea Bergh, 1891
      • Actinocyclidae O’Donoghue, 1929
        • Actinocyclus Ehrenberg, 1831
        • Hallaxa Eliot, 1909
      • Cadlinellidae Odhner, 1934
        • Cadlinella Thiele, 1931
      • Cadlinidae Bergh, 1891
        • Aldisa Bergh, 1878
        • Cadlina Bergh, 1879
        • Inuda Ev. Marcus & Er. Marcus, 1967
      • Chromodorididae Bergh, 1891
        • Chromodoridinae Bergh, 1891
          • Ardeadoris Rudman, 1984
          • Chromodoris Alder & Hancock, 1855
          • Chromolaichma Bertsch, 1977
          • Diversidoris Rudman, 1987
          • Doriprismatica A. d’Orbigny, 1839
          • Felimida Ev. Marcus, 1971
          • Glossodoris Ehrenberg, 1831
          • Goniobranchus Pease, 1866
          • Tyrinna Bergh, 1898
          • Verconia Pruvot-Fol, 1931
        • Miamirinae Bergh, 1891
          • Ceratosoma Gray, 1850
          • Felimare Ev. Marcus & Er. Marcus, 1967
          • Hypselodoris W. Stimpson, 1855
          • Mexichromis Bertsch, 1977
          • Miamira Bergh, 1874
          • Thorunna Bergh, 1878
      • Hexabranchidae Bergh, 1891
        • Hexabranchus Ehrenberg, 1828
      • Showajidaiidae Korshunova, Fletcher, Picton, Lundin, Kashio, N. Sanamyan, K. Sanamyan, Padula, Schrödl & Martynov, 2020
        • Showajidaia Korshunova, Fletcher, Picton, Lundin, Kashio, N. Sanamyan, K. Sanamyan, Padula, Schrödl & Martynov, 2020
    • Doridoidea Rafinesque, 1815
      • Discodorididae Bergh, 1891
        • Alloiodoris Bergh, 1904
        • Aphelodoris Bergh, 1879
        • Aporodoris Ihering, 1886
        • Asteronotus Ehrenberg, 1831
        • Atagema Gray, 1850
        • Avaldesia Donohoo & Gosliner, 2024
        • Baptodoris Bergh, 1884
        • Carminodoris Bergh, 1889
        • Diaulula Bergh, 1878
        • Dictyodoris Bergh, 1880
        • Discodoris Bergh, 1877
        • Gargamella Bergh, 1894
        • Geitodoris Bergh, 1891
        • Halgerda Bergh, 1880
        • Hiatodoris Dayrat, 2010
        • Hoplodoris Bergh, 1880
        • Jorunna Bergh, 1876
        • Nirva Bergh, 1905
        • Nophodoris Á. Valdés & Gosliner, 2001
        • Otinodoris K. M. White, 1948
        • Paradoris Bergh, 1884
        • Peltodoris Bergh, 1880
        • Peronodoris Bergh, 1904
        • Platydoris Bergh, 1877
        • Rostanga Bergh, 1879
        • Sclerodoris Eliot, 1904
        • Sebadoris Er. Marcus & Ev. Marcus, 1960
        • Taringa Er. Marcus, 1955
        • Tayuva Ev. Marcus & Er. Marcus, 1967
        • Thordisa Bergh, 1877
        • Thorybopus Bouchet, 1977
      • Dorididae Rafinesque, 1815
        • Archidoris Bergh, 1878
        • Artachaea Bergh, 1881
        • Austrodoris Odhner, 1926
        • Conualevia Collier & Farmer, 1964
        • Doriopsis Pease, 1860
        • Doris Linnaeus, 1758
        • Goslineria Á. Valdés, 2001
        • Pharodoris Á. Valdés, 2001
    • Onchidoridoidea Gray, 1827
      • Acanthodorididae P. Fischer, 1883
        • Acanthodoris J. E. Gray, 1850
      • Acanthomiridae Martynov & Korshunova, 2025
      • Aegiridae P. Fischer, 1883
        • Aegires Lovén, 1844
        • Notodoris Bergh, 1875
      • Akiodorididae Millen & Martynov, 2005
        • Akiodoris Bergh, 1879
        • Armodoris Minichev, 1972
        • Doridunculus G. O. Sars, 1878
        • Echinocorambe Á. Valdés & Bouchet, 1998
        • Prodoridunculus Thiele, 1912
      • Calycidorididae Roginskaya, 1972
        • Calycidoris Abraham, 1876
        • Diaphorodoris Iredale & O’Donoghue, 1923
      • Corambidae Bergh, 1871
        • Corambe Bergh, 1869
        • Loy Martynov, 1994
      • Goniodorididae H. Adams & A. Adams, 1854
        • Anculinae Pruvot-Fol, 1954
          • Ancula Lovén, 1846
          • Trapania Pruvot-Fol, 1931
        • Goniodoridinae H. Adams & A. Adams, 1854
          • Bermudella Odhner, 1941
          • Cargoa Vogel & L. P. Schultz, 1970
          • Ceratodoris J. E. Gray, 1850
          • Goniodoridella Pruvot-Fol, 1933
          • Lophodoris G. O. Sars, 1878
          • Murphydoris Sigurdsson, 1991
          • Naisdoris Paz-Sedano, Moles, Smirnoff, Gosliner & Pola, 2024
          • Okenia Menke, 1830
          • Pelagella J. E. Gray, 1850
        • Spahria Risbec, 1928
      • Onchidorididae J. E. Gray, 1827
        • Acanthomira Ekimova, Nikitenko, Stanovova, Schepetov, Antokhina & Á. Valdés, 2024
        • Adalaria Bergh, 1878
        • Arctadalaria Roginskaya, 1971
        • Atalodoris Iredale & O’Donoghue, 1923
        • Idaliadoris Furfaro & Trainito, 2022
        • Onchidoris Blainville, 1816
        • Onchimira Martynov, Korshunova, N. Sanamyan & K. Sanamyan, 2009
    • Phyllidioidea Rafinesque, 1814
      • Dendrodorididae O’Donoghue, 1924 (1864)
        • Dendrodoris Ehrenberg, 1831
        • Doriopsilla Bergh, 1880
      • Forjacellidae Ortea & Moro, 2023
        • Forjacella Ortea & Moro, 2023
      • Mandeliidae Á. Valdés & Gosliner, 1999
        • Mandelia Á. Valdés & Gosliner, 1999
      • Phyllidiidae Rafinesque, 1814
        • Ceratophyllidia Eliot, 1903
        • Phyllidia Cuvier, 1797
        • Phyllidiella Bergh, 1869
        • Phyllidiopsis Bergh, 1876
        • Reticulidia Brunckhorst, 1990
    • Polyceroidea Alder & Hancock, 1845
      • Polyceridae Alder & Hancock, 1845
        • Kalinginae Pruvot-Fol, 1956
          • Kalinga Alder & Hancock, 1864
        • Kankelibranchinae Ortea, Espinosa & Caballer, 2005
          • Kankelibranchus Ortea, Espinosa & Caballer, 2005
        • Nembrothinae Burn, 1967
          • Martadoris Willan & Y.-W. Chang, 2017
          • Nembrotha Bergh, 1877
          • Roboastra Bergh, 1877
          • Tambja Burn, 1962
          • Tyrannodoris Willan & Y.-W. Chang, 2017
        • Okadaiinae Baba, 1930
          • Vayssierea Risbec, 1928
        • Polycerinae Alder & Hancock, 1845
          • Gymnodoris W. Stimpson, 1855
          • Lamellana G.-Y. Lin, 1992
          • Lecithophorus Macnae, 1958
          • Palio Gray, 1857
          • Paliolla Burn, 1958
          • Paliota Pola, Miguel-González & Paz-Sedano, 2023
          • Polycera Cuvier, 1816
          • Polycerella A. E. Verrill, 1880
          • Thecacera J. Fleming, 1828
        • Triophinae Odhner, 1941
          • Colga Bergh, 1880
          • Crimora Alder & Hancock, 1862
          • Holoplocamus Odhner, 1926
          • Joubiniopsis Risbec, 1928
          • Kaloplocamus Bergh, 1880
          • Limacia O. F. Müller, 1781
          • Plocamopherus Rüppell & Leuckart, 1828
          • Triopha Bergh, 1880

Main Doridina groups:

Bibliography

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