Cladobranchia es uno de los dos grandes linajes evolutivos de Nudibranchia, junto con Doridina. Comprende un grupo extraordinariamente diverso de babosas marinas, caracterizado por una gran variedad de formas corporales y de estructuras dorsales y laterales especializadas.
Candiella striata por Pontes, Miquel
Cladobranchia presenta una notable diversidad morfológica. Muchas especies poseen apéndices dorsales especializados relacionados con la respiración, la defensa, la alimentación o las funciones sensoriales. Los aeólidos se caracterizan típicamente por numerosos cerata distribuidos por la superficie dorsal. Estas estructuras contienen prolongaciones de la glándula digestiva y, en muchas especies, incorporan cnidosacos, estructuras especializadas relacionadas con la defensa. Otros cladobranquios presentan procesos dorsales ramificados o dendríticos, estructuras orales ensanchadas, vainas rinofóricas u otros apéndices especializados.
El nombre Cladobranchia fue introducido por Willan y Morton (1984) para un grupo de nudibranquios que incluía los principales linajes tradicionalmente reconocidos como aeólidos, armináceos y dendronotáceos. Posteriormente, el concepto del grupo fue revisado mediante estudios anatómicos comparativos y filogenéticos. La incorporación de datos moleculares permitió revisar las relaciones entre diversos linajes y puso de manifiesto que algunas agrupaciones tradicionales basadas en la morfología no se correspondían necesariamente con grupos evolutivos naturales.
La filogenética molecular ha comportado numerosos cambios en la clasificación superior de Cladobranchia. Diversas familias y superfamilias han sido creadas, redefinidas o reorganizadas a medida que se han incorporado nuevos datos. Las relaciones entre algunos de los principales linajes del grupo continúan siendo objeto de estudio, y la clasificación ha ido evolucionando con la acumulación de nuevas evidencias.
La clasificación propuesta por Korshunova et al. (2025) introdujo una importante reorganización de la clasificación superior de los nudibranquios y adoptó un concepto restringido de Nudibranchia. Posteriormente, Moles et al. (2026) reconsideraron esta propuesta en el contexto de su análisis de la estabilidad taxonómica de los heterobranquios marinos. En el tratamiento adoptado aquí, se mantiene el concepto amplio de Nudibranchia y Cladobranchia se considera uno de sus dos grandes linajes principales.
Cladobranchia comprende una gran diversidad de grupos, entre los que se encuentran numerosos linajes de aeólidos y otras formas tradicionalmente agrupadas entre los armináceos y dendronotáceos. La clasificación superior ha experimentado diversas modificaciones y, por este motivo, la delimitación y asignación de algunos grupos pueden variar según la fuente taxonómica utilizada.
Los aeólidos son especialmente conocidos por sus cerata, que pueden desempeñar funciones respiratorias, digestivas y defensivas. Algunas especies son capaces de retener nematocistos funcionales obtenidos de sus presas cnidarias e incorporarlos a los cnidosacos situados en los extremos de los cerata. Otros cladobranquios presentan mecanismos defensivos diferentes, entre los que se encuentran las defensas químicas, las coloraciones crípticas y las estructuras corporales especializadas.
Cladobranchia ocupa una amplia variedad de ambientes marinos, desde aguas costeras poco profundas y zonas intermareales hasta ambientes más profundos de la plataforma y el talud continental. Muchas especies mantienen relaciones tróficas especializadas con determinados organismos, entre los que se encuentran hidrozoos, octocorales y otros cnidarios, esponjas y briozoos. Estas especializaciones alimentarias pueden estar asociadas con adaptaciones morfológicas, comportamentales y químicas.
La diversidad de Cladobranchia se manifiesta tanto en su morfología como en sus estrategias ecológicas. La gran variedad de estructuras corporales, formas de alimentación y mecanismos defensivos que presenta el grupo refleja una historia evolutiva compleja. Los estudios anatómicos, del desarrollo, moleculares y ecológicos continúan aportando información para comprender mejor las relaciones entre sus diferentes linajes y su diversificación.
La clasificación que se presenta a continuación refleja el marco taxonómico actual utilizado en el World Register of Marine Species (WoRMS), teniendo en cuenta el concepto más amplio de Nudibranchia avalado por Moles et al. (2026):
Cladobranchia
- Aeolidioidea J. E. Gray, 1827
- Aeolidiidae J. E. Gray, 1827
- Aeolidia Cuvier, 1798
- Aeolidiella Bergh, 1867
- Aeolidiopsis Pruvot-Fol, 1956
- Anteaeolidiella M. C. Miller, 2001
- Baeolidia Bergh, 1888
- Berghia Trinchese, 1877
- Bulbaeolidia Carmona, Pola, Gosliner & Cervera, 2013
- Cerberilla Bergh, 1873
- Limenandra Haefelfinger & Stamm, 1958
- Spurilla Bergh, 1864
- Zeusia Korshunova, Zimina & Martynov, 2017
- Babakinidae Roller, 1973
- Babakina Roller, 1973
- Facelinidae Bergh, 1889
- Adfacelina Millen & Hermosillo, 2012
- Algarvia García-Gómez & Cervera, 1990
- Amanda Macnae, 1954
- Anetarca Gosliner, 1991
- Antonietta Schmekel, 1966
- Austraeolis Burn, 1962
- Bajaeolis Gosliner & Behrens, 1986
- Burnaia M. C. Miller, 2001
- Caloria Trinchese, 1888
- Cratena Bergh, 1864
- Crateninae Bergh, 1889
- Dicata Schmekel, 1967
- Echinopsole Macnae, 1954
- Emarcusia Roller, 1972
- Facelina Alder & Hancock, 1855
- Facelininae Bergh, 1889
- Facelinopsis Pruvot-Fol, 1954
- Favorininae Bergh, 1889
- Favorinus J. E. Gray, 1850
- Hermosita Gosliner & Behrens, 1986
- Herviella Baba, 1949
- Herviellinae Burn, 1967
- Janssonius Ortea & Moro, 2022
- Jason M. C. Miller, 1974
- Learchis Bergh, 1896
- Moridilla Bergh, 1888
- Myja Bergh, 1896
- Noumeaella Risbec, 1937
- Palisa Edmunds, 1964
- Pauleo Millen & Hamann, 1992
- Phidiana Gray, 1850
- Pruvotfolia J. Tardy, 1970
- Pteraeolidiinae Risbec, 1953
- Pteraeolidia Bergh, 1875
- Sakuraeolis Baba, 1965
- Setoeolis Baba, 1965
- Flabellinidae Bergh, 1889
- Calmella Eliot, 1906
- Carronella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Coryphellina O’Donoghue, 1929
- Edmundsella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Flabellina McMurtrie, 1831
- Paraflabellina Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Flabellinopsidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Baenopsis Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Flabellinopsis MacFarland, 1966
- Glaucidae Gray, 1827
- Glaucilla Bergh, 1861
- Glaucus Forster, 1777
- Hantazuidae Korshunova, Fletcher & Martynov, 2025
- Hantazuia Korshunova, Fletcher & Martynov, 2025
- Myrrhinidae Bergh, 1905
- Dondice Er. Marcus, 1958
- Godiva Macnae, 1954
- Hermissenda Bergh, 1879
- Nanuca Er. Marcus, 1957
- Nemesignis Furfaro & Mariottini, 2021
- Phyllodesmium Ehrenberg, 1831
- Piseinotecidae Edmunds, 1970
- Piseinotecus Er. Marcus, 1955
- Pleurolidiidae Burn, 1966
- Pleurolidia Burn, 1966
- Protaeolidiella Baba, 1955
- Aeolidiidae J. E. Gray, 1827
- Apataoidea Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Apataidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Apata Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Tularia Burn, 1966
- Apataidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Chudoidea Korshunova, Fletcher & Martynov, 2025
- Chudidae Korshunova, Fletcher & Martynov, 2025
- Chudo Korshunova, Fletcher & Martynov, 2025
- Chudidae Korshunova, Fletcher & Martynov, 2025
- Embletonioidea Pruvot-Fol, 1954
- Embletoniidae Pruvot-Fol, 1954
- Embletonia Alder & Hancock, 1851
- Embletoniidae Pruvot-Fol, 1954
- Fionoidea J. E. Gray, 1857
- Abronicidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Abronica Cella, Carmona, Ekimova, Chichvarkhin, Schepetov & Gosliner, 2016
- Calmidae Iredale & O’Donoghue, 1923
- Calma Alder & Hancock, 1855
- Coryphellidae Bergh, 1889
- Borealea Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Corrupta Korshunova, Fletcher & Martynov, 2025
- Coryphella J. E. Gray, 1850
- Fjordia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Gulenia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Himatina Thiele, 1931
- Itaxia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Microchlamylla Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Occidenthella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Orienthella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Portorchardia Korshunova, Fletcher & Martynov, 2025
- Cumanotidae Odhner, 1907
- Cumanotus Odhner, 1907
- Cuthonellidae M. C. Miller, 1977
- Cratenopsis Lemche, 1935
- Cuthonella Bergh, 1884
- Fiocuthona Martynov, 1992
- Cuthonidae Odhner, 1934
- Bohuslania Korshunova, Lundin, Malmberg, Picton & Martynov, 2018
- Cuthona Alder & Hancock, 1855
- Eubranchidae Odhner, 1934
- Aenigmastyletus Martynov, 1998
- Amphorina Quatrefages, 1844
- Annulorhina K. P. Rao, 1968
- Capellinia Trinchese, 1873
- Corruptobranchus Martynov & Korshunova, 2025
- Dunga Eliot, 1902
- Eubranchopsis Baba, 1949
- Eubranchulus Martynov & Korshunova, 2025
- Eubranchus Forbes, 1838
- Galvinella Eliot, 1907
- Karavellia Martynov & Korshunova, 2025
- Leostyletus Martynov, 1998
- Nudibranchus Martynov, 1998
- Produnga Martynov, 1998
- Fionidae J. E. Gray, 1857
- Fiona Alder & Hancock, 1853
- Tergiposacca Cella, Carmona, Ekimova, Chichvarkhin, Schepetov & Gosliner, 2016
- Zatteria Eliot, 1902
- Magallanidae Ortea & Moro, 2020
- Magallanes Ortea & Moro, 2020
- Murmaniidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Guyvalvoria Vayssière, 1906
- Murmania Martynov, 2006
- Paracoryphellidae M. C. Miller, 1971
- Chlamylla Bergh, 1886
- Paracoryphella M. C. Miller, 1971
- Polaria Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Ziminella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Pinufiidae Er. Marcus & Ev. Marcus, 1960
- Pinufius Er. Marcus & Ev. Marcus, 1960
- Pseudovermidae Thiele, 1931
- Pseudovermis Perejaslavtzeva, 1891
- Tergipedidae Bergh, 1889
- Tergipes Cuvier, 1805
- Trinchesiidae F. Nordsieck, 1972
- Catriona Winckworth, 1941
- Diaphoreolis Iredale & O’Donoghue, 1923
- Phestilla Bergh, 1874
- Rubramoena Cella, Carmona, Ekimova, Chichvarkhin, Schepetov & Gosliner, 2016
- Selva Edmunds, 1964
- Tenellia A. Costa, 1866
- Trinchesia Ihering, 1879
- Xenocratenidae Martynov, Lundin, Picton, Fletcher, Malmberg & Korshunova, 2020
- Xenocratena Odhner, 1940
- Abronicidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Notaeolidioidea Eliot, 1910
- Notaeolidiidae Eliot, 1910
- Notaeolidia Eliot, 1905
- Notaeolidiidae Eliot, 1910
- Samloidea Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Samlidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Luisella Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Samla Bergh, 1900
- Samlidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Unidentioidea Millen & Hermosillo, 2012
- Unidentiidae Millen & Hermosillo, 2012
- Pacifia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017
- Phetia Korshunova, Fletcher & Martynov, 2025
- Unidentia Millen & Hermosillo, 2012
- Unidentiidae Millen & Hermosillo, 2012
- Arminoidea Iredale & O’Donoghue, 1923 (1841)
- Arminidae Iredale & O’Donoghue, 1923 (1841)
- Armina Rafinesque, 1814
- Atthila Bergh, 1899
- Dermatobranchus van Hasselt, 1824
- Histiomena Mörch, 1859
- Pleurophyllidiella Eliot, 1903
- Pleurophyllidiopsis S. Tchang, 1934
- Arminidae Iredale & O’Donoghue, 1923 (1841)
- Dendronotoidea Allman, 1845
- Bornellidae Bergh, 1874
- Bornella Gray, 1850
- Dendronotidae Allman, 1845
- Cabangus Korshunova, Bakken, Grøtan, K. B. Johnson, Lundin & Martynov, 2020
- Dendronotus Alder & Hancock, 1845
- Pseudobornella Baba, 1932
- Dotidae Gray, 1853
- Caecinella Bergh, 1870
- Doto Oken, 1815
- Kabeiro Shipman & Gosliner, 2015
- Miesea Er. Marcus, 1961
- Hancockiidae MacFarland, 1923
- Hancockia Gosse, 1877
- Lomanotidae Bergh, 1890
- Lomanotus Vérany, 1844
- Phylliroidae Menke, 1830
- Cephalopyge Hanel, 1905
- Phylliroe F. Péron & Lesueur, 1810
- Scyllaeidae Alder & Hancock, 1855
- Crosslandia Eliot, 1902
- Notobryon Odhner, 1936
- Scyllaea Linnaeus, 1758
- Tethydidae Rafinesque, 1815
- Melibe Rang, 1829
- Tethys Linnaeus, 1767
- Bornellidae Bergh, 1874
- Proctonotoidea J. E. Gray, 1853
- Curnonidae d’Udekem d’Acoz, 2017
- Curnon d’Udekem d’Acoz, 2017
- Pseudotritonia Thiele, 1912
- Dironidae Eliot, 1910
- Dirona MacFarland, 1905
- Goniaeolididae Odhner, 1907
- Goniaeolis M. Sars, 1861
- Janolidae Pruvot-Fol, 1933
- Antiopella Hoyle, 1902
- Bonisa Gosliner, 1981
- Galeojanolus M. C. Miller, 1971
- Janolus Bergh, 1884
- Lemindidae R. J. Griffiths, 1985
- Leminda R. J. Griffiths, 1985
- Madrellidae Preston, 1911
- Eliotia Vayssière, 1909
- Madrella Alder & Hancock, 1864
- Proctonotidae J. E. Gray, 1853
- Caldukia Burn & M. C. Miller, 1969
- Proctonotus Alder & Hancock, 1844
- Curnonidae d’Udekem d’Acoz, 2017
- Tritonioidea Lamarck, 1809
- Tritoniidae Lamarck, 1809
- Marioniinae F. V. Silva, Pola & Cervera, 2023
- Marionia Vayssière, 1877
- Marioniopsis Odhner, 1934
- Paratritonia Baba, 1949
- Tritoniinae Lamarck, 1809
- Candiella J. E. Gray, 1850
- Marianina Pruvot-Fol, 1931
- Myrella Odhner, 1963
- Tochuina Odhner, 1963
- Tritonia Cuvier, 1797
- Tritonicula Korshunova & Martynov, 2020
- Tritonidoxa Bergh, 1907
- Tritoniella Eliot, 1907
- Tritoniopsis Eliot, 1905
- Marioniinae F. V. Silva, Pola & Cervera, 2023
- Tritoniidae Lamarck, 1809
Grupos principales de Cladobranchia:
Bibliografía
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Bibliografía basada en los trabajos de Steve Long, 2006. Bibliography of Opisthobranchia 1554-2000 y Gary McDonald, 2009. Bibliographia Nudibranchia, con actualizaciones posteriores procedentes de otras fuentes.
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