Dendronotaceans have a long and narrow body and have a row of branched or tuberculate processes on each side of the body, in dorsolateral position. The rhinophores can be smooth or have lamellae but their base is always surrounded by a high sheath, into which they can be retracted. Head may differentiate either rounded lateral expansions or a face veil with several digitations. The anus and the genital openings are located to the right of the body.

Doto rosea 3mm by Enric Madrenas

They feed primarily on cnidarians, such as hydroid colonies, soft corals and gorgonians. Species of the family Tethydidae, however, tend to eat small crustaceans such as amphipods, copepods, isopods and ostracods. Within this family, a big sizes species, Tethys fimbria, has a wider food spectrum as it has been cited feeding on small echinoderms, gastropods, polychaetes and small fish. Most dendronotacean are benthic but some species can swim by undulating movements of the body; species of the family Phylliroidae are pelagic and prey on jellyfish and siphonophores.


Wägele & Willan (2000) concluded that the Dendronotida (including Tritonioidea and Dendronotoidea) are monophyletic, but Martin et al. (2009, 2010) and Goodheart (2017) obtained different results. Already Healy & Willan (1991) identified wide variations in sperm morphology that they questioned its monophyly.

Although the family Dotidae is consistently excluded from the Dendronotida in morphocladistic analyses (Wägele & Willan, 2000), Doto and Hancockia cluster with Dendronotus and Scyllaea in the molecular phylogeny of Mahguib & Valdés (2015), which supports the recovery of Doto as sister to a clade of Melibe and Dendronotus (Goodheart, 2017).

The relationship of Hancockiidae to other Cladobranchia was unresolved in morphocladistic (Martin et al., 2009) and multilocus approaches (Pola & Gosliner, 2010). They were included in Dendronotoidea by Bouchet et al. (2017) based on the tree of Mahguib & Valdés (2015). Similarly, the relationship of Tethys to other Cladobranchia was unresolved in multilocus phylogeny of Pola & Gosliner (2010), but resulted sister to Dendronotus in the phylogenomic study by Goodheart et al. (2015) and Goodheart (2017).

Actual classification of the Dendronotaceans as proposed by Bouchet et al.(2017) and accepted at WoRMS is:

  • Superfamily Dendronotoidea Allman, 1845
    • Family Bornellidae Bergh, 1874
      • Genus Bornella Gray, 1850
      • Genus Bornellopsis O’Donoghue, 1929
    • Family Dendronotidae Allman, 1845
      • Genus Dendronotus Alder & Hancock, 1845
    • Family Dotidae Gray, 1853
      • Genus Caecinella Bergh, 1870
      • Genus Doto Oken, 1815
      • Genus Kabeiro Shipman & Gosliner, 2015
      • Genus Miesea Er. Marcus, 1961
    • Family Hancockiidae MacFarland, 1923
      • Genus Hancockia Gosse, 1877
    • Family Lomanotidae Bergh, 1890
      • Genus Lomanotus Vérany, 1844
    • Family Scyllaeidae Alder & Hancock, 1855
      • Genus Crosslandia Eliot, 1902
      • Genus Notobryon Odhner, 1936
      • Genus Scyllaea Linnaeus, 1758
    • Family Tethydidae Rafinesque, 1815
      • Genus Melibe Rang, 1829
      • Genus Tethys Linnaeus, 1767

Dendronotacean species recorded in the Mediterranean Sea or around the Iberian Peninsula:

Dendronotus frondosus by Klas Malmberg (Sweden)

Dendronotus frondosus

Doto alidrisi @ Ceuta, Spain 24-04-2010 by Leopoldo Moro Abad

Doto alidrisi

Doto caballa @ Ceuta, Spain 25-04-2010 by Leopoldo Moro Abad

Doto caballa

Doto cervicenigra @ Sta. Maria al Bagno, Lecce, Italy by Cataldo Licchelli

Doto cervicenigra

Doto coronata @ L'Escala by Enric Madrenas

Doto coronata

Doto cuspidata by Klas Malmberg (Sweden)

Doto cuspidata

Doto dunnei @ “Depuradora” (L’Escala, Girona), 21/04/2012 by Enric Madrenas

Doto dunnei

Doto eireana - Photograph ©Bernard Picton

Doto eireana

Doto floridicola by Enric Madrenas

Doto floridicola

Doto fragaria @ Santa Maria al Bagno (Lecce), Jonian Sea, Italy 7m on hydroids and porifera, 2013 by Cataldo Licchelli

Doto fragaria

Doto fragilis @ L'Escala, Girona, Spain 6.03.2021 by Paqui Serra i Oriol Planes

Doto fragilis

Doto furva @ Tarifa (Spain) by Manuel Martínez Chacón

Doto furva

Doto koenneckeri by Bernard Picton

Doto koenneckeri

Doto lemchei

Doto lemchei

Doto maculata by Bernard Picton

Doto maculata

Doto millbayana by Klas Malmberg (Sweden)

Doto millbayana

Doto paulinae by Enric Madrenas

Doto paulinae

Doto pinnatifida by João Pedro Silva

Doto pinnatifida

Doto pita on hydroid @ Puerto del Carmen, Lanzarote, Spain by Dennis Rabeling

Doto pita

Doto pygmaea @ Lanzarote, Canary Islands by Dennis Rabeling

Doto pygmaea

Doto rosea by Enric Madrenas

Doto rosea

Doto tuberculata by Klas Malmberg (Sweden)

Doto tuberculata

Doto verdicioi @ Lapamán (Ría de Pontevedra, Spain) 28.03.1979 by Victoriano Urgorri

Doto verdicioi

Hancockia uncinata @ Costa Brava, Spain by Enric Madrenas

Hancockia uncinata

Lomanotus barlettai by José Carlos García Gómez

Lomanotus barlettai

Lomanotus genei Adult with parasitic copepod eggs protruding from back. Mulroy Bay, Co Donegal, Ireland. - Photograph ©Bernard Picton

Lomanotus genei

Lomanotus marmoratus by Klas Malmberg (Sweden)

Lomanotus marmoratus

Melibe viridis @ Punta Japlenicka-island Ugljan, Croatia, length 45cm, depth 6m by Đani Iglic

Melibe viridis

Scyllaea pelagica by Enrico Ricchitelli

Scyllaea pelagica

Tethys fimbria @ Badalona (Spain) by Enric Madrenas

Tethys fimbria


    Bouchet P, Rocroi JP, Hausdorf B, et al. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia. 61(1–2):1–526.

    Bibliography based on the works by Steve Long, 2006. Bibliography of Opisthobranchia 1554-2000 and Gary McDonald, 2009. Bibliographia Nudibranchia, with later updates from other resources.