Cladobranchia

The Cladobranchia are a group of nudibranchs that includes traditional cladohepatic nudibranchs (eolidaceans, arminaceans and dendronotaceans) plus the enigmatic group of Doridoxidae, all characterized by having the anus in the right lateral position and a more or less branched digestive gland.

Janolus cristatus by Enric Madrenas

Cladobranchia is diverse group of marine molluscs that includes approximately 1000 species, but the lack of studies results in that this group is poorly understood.

According to Goodheart et al. (2015), to date, there have been only two large-scale phylogenies published specifically on Cladobranchia: (Pola & Gosliner, 2010) which used the three genes most commonly used in nudibranch phylogenetics and (Goodheart et al, 2015), that used all data publicly available through GenBank to understand the evolutionary relationships of members of this group, thus adding two additional genes and increasing taxon sampling by approximately 200 taxa. In both of these phylogenetic inferences of Cladobranchia, the majority of relationships remained unclear, both between and within the three traditional taxonomic divisions of Cladobranchia (Arminaceans, with about 100 species; Dendronotaceans, with about 250 species; and Aeolidaceans, with about 600 species).

In addition to the phylogenies specifically aimed at understanding relationships within Cladobranchia, phylogenetic inferences have been attempted to address the evolutionary history of Nudibranchia or the larger clade Euthyneura (Wollscheid E & Wägele H. 1999; Wollscheid et al. 2001; Thollesson, 1999; Wägele & Willan, 2000; Wägele et al, 2013; Mahguib & Valdés, 2015 and Schrödl et al, 2001). The results regarding the major groupings within Cladobranchia were inconsistent in all of these studies, with all three major divisions considered both paraphyletic and monophyletic in different publications.

Though the more basal evolutionary history of Cladobranchia has been problematic for phylogenetic inference, a number of studies (both morphological and molecular) have provided evidence to support relationships at both the family and generic levels. Recently several phylogenies of individual and at the genus level have been published, focused on particular aspects of the Cladobranchia tree, and although they have supported some relationships, it is clear that the overall relationships of higher level groups in Cladobranchia are not well understood. In addition, individually these studies cover very little of the overall diversity of Cladobranchia.

The latest work by Bouchet et al. (2017) tries to put some order based on a series of works published in 2017 that sought to clarify the phylogenetic relationships of relatively large groups, such as the Flabellinoidea or the Fionoidea, and eliminating the “catchall group” of unassigned Cladobranchia by adscribing the families that it contained to the other groups. Thus, we get a greater granularity with the aim to improve our understanding of these evolutionary lines.

The Cladobranchia, as a group, lack an homogeneous body organization: Aeolidioidea, Fionoidea, Flabellinoidea and Proctonotoidea have an elongated body and groups of papillae called “cerata” on both sides of the body. The Arminoidea have a roughly triangular shaped body and live in soft bottoms. Dendronotoidea i Tritonioidea have an elongated body and dendritic extensions in both sides of the body. The Doridoxoidea have the body shape of doridacean nudibranchs, that is, dorsoventrally flattened.

Current taxonomy of Cladobranchia proposed by Bouchet et al.(2017) and accepted at WoRMS is:

  • Suborder Cladobranchia
    • Superfamily Aeolidioidea Gray, 1827
      • Family Aeolidiidae Gray, 1827
        • Genus Aeolidia Cuvier, 1798
        • Genus Aeolidiella Bergh, 1867
        • Genus Anteaeolidiella M. C. Miller, 2001
        • Genus Baeolidia Bergh, 1888
        • Genus Berghia Trinchese, 1877
        • Genus Bulbaeolidia Carmona, Pola, Gosliner & Cervera, 2013
        • Genus Cerberilla Bergh, 1873
        • Genus Limenandra Haefelfinger & Stamm, 1958
        • Genus Spurilla Bergh, 1864
        • Genus Zeusia Korshunova, Zimina & Martynov, 2017
      • Family Babakinidae Roller, 1973
        • Genus Babakina Roller, 1973
      • Family Facelinidae Bergh, 1889
        • Genus Adfacelina Millen & Hermosillo, 2012
        • Genus Algarvia Garcia-Gomez & Cervera, 1990
        • Genus Amanda Macnae, 1954
        • Genus Anetarca Gosliner, 1991
        • Genus Antonietta Schmekel, 1966
        • Genus Austraeolis Burn, 1962
        • Genus Bajaeolis Gosliner & Behrens, 1986
        • Genus Burnaia M. C. Miller, 2001
        • Genus Caloria Trinchese, 1888
        • Genus Cratena Bergh, 1864
        • Subfamily Crateninae Bergh, 1889
          • Genus Echinopsole Macnae, 1954
          • Genus Emarcusia Roller, 1972
          • Genus Facelina Alder & Hancock, 1855
        • Subfamily Facelininae Bergh, 1889
          • Genus Facelinopsis Pruvot-Fol, 1954
        • Subfamily Favorininae Bergh, 1889
          • Genus Dicata Schmekel, 1967
          • Genus Dondice Marcus, 1958
          • Genus Godiva Macnae, 1954
          • Genus Favorinus Gray, 1850
          • Genus Hermissenda Bergh, 1879
          • Genus Hermosita Gosliner & Behrens, 1986
          • Genus Herviella Baba, 1949
        • Subfamily Herviellinae Burn, 1967
          • Genus Jason M. C. Miller, 1974
          • Genus Learchis Bergh, 1896
          • Genus Moridilla Bergh, 1888
          • Genus Nanuca Marcus, 1957
          • Genus Noumeaella Risbec, 1937
          • Genus Palisa Edmunds, 1964
          • Genus Pauleo Millen & Hamann, 1992
          • Genus Phidiana Gray, 1850
          • Genus Phyllodesmium Ehrenberg, 1831
          • Genus Pruvotfolia Tardy, 1969
          • Genus Pteraeolidia Bergh, 1875
        • Subfamily Pteraeolidiinae Risbec, 1953
          • Genus Sakuraeolis Baba, 1965
          • Genus Setoeolis Baba & Hamatani, 1965
      • Family Glaucidae Gray, 1827
        • Genus Glaucus Forster, 1777
      • Family Piseinotecidae Edmunds, 1970
        • Genus Piseinotecus Er. Marcus, 1955
      • Family Pleurolidiidae Burn, 1966
        • Genus Pleurolidia Burn, 1966
        • Genus Protaeolidiella Baba, 1955
      • Family Unidentiidae Millen & Hermosillo, 2012
        • Genus Pacifia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Unidentia Millen & Hermosillo, 2012
    • Superfamily Arminoidea Iredale & O’Donoghue, 1923 (1841)
      • Family Arminidae Iredale & O’Donoghue, 1923 (1841)
        • Genus Armina Rafinesque, 1814
        • Genus Atthila Bergh, 1899
        • Genus Dermatobranchus van Hasselt, 1824
        • Genus Heterodoris Verrill & Emerton, 1882
        • Genus Histiomena Mörch, 1860
        • Genus Pleurophyllidiella Eliot, 1903
        • Genus Pleurophyllidiopsis Tchang, 1934
      • Family Doridomorphidae Er. Marcus & Ev. Marcus, 1960 (1908)
        • Genus Doridomorpha Eliot, 1903
    • Superfamily Dendronotoidea Allman, 1845
      • 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
    • Superfamily Doridoxoidea Bergh, 1899
      • Family Doridoxidae Bergh, 1899
        • Genus Doridoxa Bergh, 1899
    • Superfamily Fionoidea Gray, 1857
      • Family Abronicidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Abronica Cella, Carmona, Ekimova, Chichvarkhin, Schepetov & Gosliner, 2016
      • Family Calmidae Iredale & O’Donoghue, 1923
        • Genus Calma Alder & Hancock, 1855
      • Family Cuthonellidae Miller, 1971
        • Genus Cuthonella Bergh, 1884
      • Family Cuthonidae Odhner, 1934
        • Genus Cuthona Alder & Hancock, 1855
      • Family Eubranchidae Odhner, 1934
        • Genus Eubranchus Forbes, 1838
        • Genus Leostyletus Martynov, 1998
      • Family Fionidae Gray, 1857
        • Genus Fiona Alder & Hancock [in Forbes & Hanley], 1853
        • Genus Tergiposacca Cella, Carmona, Ekimova, Chichvarkhin, Schepetov & Gosliner, 2016
        • Genus Zatteria Eliot, 1902
      • Family Murmaniidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Murmania Martynov, 2006
      • Family Tergipedidae Bergh, 1889
        • Genus Tergipes Cuvier, 1805
      • Family Trinchesiidae F. Nordsieck, 1972
        • Genus Catriona Winckworth, 1941
        • Genus Diaphoreolis Iredale & O’Donoghue, 1923
        • Genus Phestilla Bergh, 1874
        • Genus Rubramoena Cella, Carmona, Ekimova, Chichvarkhin, Schepetov & Gosliner, 2016
        • Genus Selva Edmunds, 1964
        • Genus Tenellia A. Costa, 1866
        • Genus Trinchesia Ihering, 1879
        • Genus Zelentia Korshunova, Martynov & Picton, 2017
    • Superfamily Flabellinoidea Bergh, 1889
      • Family Apataidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Apata Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Tularia Burn, 1966
      • Family Coryphellidae Bergh, 1889
        • Genus Borealea Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Coryphella M. E. Gray, 1850
        • Genus Fjordia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Gulenia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Himatina Thiele, 1931
        • Genus Itaxia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Microchlamylla Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Occidenthella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Orienthella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
      • Family Cumanotidae Odhner, 1907
        • Genus Cumanotus Odhner, 1907
      • Family Flabellinidae Bergh, 1889
        • Genus Calmella Eliot, 1910
        • Genus Carronella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Coryphellina O’Donoghue, 1929
        • Genus Edmundsella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Flabellina Gray, 1833
        • Genus Paraflabellina Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
      • Family Flabellinopsidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Baenopsis Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Flabellinopsis MacFarland, 1966
      • Family Notaeolidiidae Eliot, 1910
        • Genus Notaeolidia Eliot, 1905
      • Family Paracoryphellidae M. C. Miller, 1971
        • Genus Chlamylla Bergh, 1886
        • Genus Paracoryphella M. C. Miller, 1971
        • Genus Polaria Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Ziminella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
      • Family Samlidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Luisella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, Saito, Lundin, Schrödl & Picton, 2017
        • Genus Samla Bergh, 1900
    • Superfamily Proctonotoidea Gray, 1853
      • Family Curnonidae d’Udekem d’Acoz, 2017
        • Genus Curnon d’Udekem d’Acoz, 2017
        • Genus Pseudotritonia Thiele, 1912
      • Family Dironidae Eliot, 1910
        • Genus Dirona MacFarland, 1905
      • Family Lemindidae Griffiths, 1985
        • Genus Leminda Griffiths, 1985
      • Family Proctonotidae Gray, 1853
        • Genus Bonisa Gosliner, 1981
        • Genus Caldukia Burn & M. C. Miller, 1969
        • Genus Galeojanolus M. C. Miller, 1971
        • Genus Janolus Bergh, 1884
        • Genus Proctonotus Alder & Hancock, 1844
    • Superfamily Tritonioidea Lamarck, 1809
      • Family Tritoniidae Lamarck, 1809
        • Genus Marianina Pruvot-Fol, 1931
        • Genus Marionia Vayssière, 1877
        • Genus Marioniopsis Odhner, 1934
        • Genus Paratritonia Baba, 1949
        • Genus Tochuina Odhner, 1963
        • Genus Tritonia Cuvier, 1798
        • Genus Tritonidoxa Bergh, 1907
        • Genus Tritoniella Eliot, 1907
        • Genus Tritoniopsis Eliot, 1905
    • Superfamily Unassigned Cladobranchia
      • Family Bornellidae Bergh, 1874
        • Genus Bornella Gray, 1850
        • Genus Bornellopsis O’Donoghue, 1929
      • Family Embletoniidae Pruvot-Fol, 1954
        • Genus Embletonia Alder & Hancock, 1851
      • Family Goniaeolididae Odhner, 1907
        • Genus Goniaeolis M. Sars, 1861
      • Family Heroidae Gray, 1857
        • Genus Hero Alder & Hancock, 1855
      • Family Madrellidae Preston, 1911
        • Genus Eliotia Vayssière, 1909
        • Genus Madrella Alder & Hancock, 1864
      • Family Phylliroidae Menke, 1830
        • Genus Cephalopyge Hanel, 1905
        • Genus Phylliroe Péron & Lesueur, 1810
      • Family Pinufiidae Er. Marcus & Ev. Marcus, 1960
        • Genus Pinufius Er. Marcus & Ev. Marcus, 1960
      • Family Pseudovermidae Thiele, 1931
        • Genus Pseudovermis Perejaslavtzeva, 1891
      • Genus Trivettea Bertsch, 2014

Main Cladobranchia groups:

Dondice banyulensis

Aeolidioidea

Armina neapolitana

Arminoidea

Doto rosea 3mm by Enric Madrenas

Dendronotoidea

Eubranchus farrani (forma A) @ Galicia by Carlos Fernández-Cid

Fionoidea

Doridoxa ingolfiana

Doridoxoidea

Fjordia lineata by Enric Madrenas

Flabellinoidea

Janolus cristatus @ Salpatxot, Illes Medes, Spain 31-03-2012 by Miquel Pontes

Proctonotoidea

Tritonia striata

Tritonioidea

Phylliroe lichtensteinii 3-01-2017 by Linda Ianello

Unassigned Cladobranchia

Bibliography