Los nudibranquios son un grupo de babosas marinas que incluye a los tradicionales nudibranquios cladohepáticos (eolidáceos, armináceos y dendronotáceos), además del enigmático grupo de los Doridoxidae, todos ellos caracterizados por presentar el ano en posición lateral derecha y una glándula digestiva más o menos ramificada.
Felimare tricolor por Enric Madrenas
La historia taxonómica de los nudibranquios (Nudibranchia Cuvier, 1817) es un ejemplo paradigmático de cómo la biología marina ha evolucionado desde un enfoque esencialmente morfológico hacia una síntesis filogenética que integra datos anatómicos, moleculares y ontogenéticos. Los nudibranquios, tradicionalmente incluidos dentro de los “opistobranquios”, fueron considerados durante décadas un orden de este gran grupo de moluscos gasterópodos marinos caracterizados por la pérdida de la concha en el estado adulto y por la presencia de branquias externas. Esta concepción clásica —heredada de la zoología del siglo XIX— se mantuvo vigente hasta bien entrado el siglo XX, cuando los primeros análisis morfológicos sistemáticos comenzaron a revelar la complejidad interna del grupo y la falta de monofilia de los “Opisthobranchia”.
Los estudios de Heike Wägele (2000), basados en más de un centenar de caracteres anatómicos e histológicos, fueron decisivos para consolidar la monofilia de Nudibranchia y establecer una división primaria en dos grandes linajes: Anthobranchia y Cladobranchia, diferenciados por la morfología corporal y la disposición de las branquias. Esta estructura básica dominó la sistemática del grupo durante casi dos décadas, hasta que la llegada de las técnicas moleculares transformó radicalmente la manera de entender las relaciones internas de los heterobranquios.
A partir de los primeros trabajos moleculares y transcriptómicos del siglo XXI (como los de Karmeinski et al., 2021), se puso de manifiesto que muchas de las clasificaciones tradicionales eran artificiales o parafiléticas. Grupos que habían sido definidos por caracteres superficiales -como la forma del cuerpo o la estructura de los cerata- resultaron ser heterogéneos, mientras que otros linajes aparentemente dispersos mostraban una afinidad filogenética inesperada. En este contexto, la noción clásica de Nudibranchia como un conjunto de “opistobranquios sin concha” perdió sentido, y el foco se desplazó hacia una delimitación filogenéticamente coherente dentro del gran clado Heterobranchia.
El trabajo de Korshunova et al. (2025) representa, en este sentido, un punto de inflexión fundamental. Los autores proponen una revisión profunda de la taxonomía de los nudibranquios basada en una combinación de datos morfológicos detallados, observaciones ontogenéticas y secuencias moleculares. Su estudio denuncia lo que denominan “pseudo-optimización taxonómica”: una tendencia histórica a simplificar en exceso la clasificación para ajustarla a modelos jerárquicos rígidos, suprimiendo así buena parte de la diversidad real del grupo.
Korshunova et al. (2025) restringen de manera explícita el alcance del orden Nudibranchia y reinstauran cinco subórdenes clásicos que habían sido progresivamente fusionados o abandonados: Arminina, Tritonina, Dendronotina, Janolina y Aeolidina. Esta reestructuración devuelve coherencia filogenética a la clasificación y reconoce la independencia evolutiva de diversos linajes que antes se habían agrupado bajo denominaciones artificiales como “Cladobranchia”. Especialmente significativa es la revisión del suborden Aeolidina, dentro del cual los autores identifican diez superfamilias claramente definidas y veintinueve familias, incluyendo dos de nueva descripción (Chudidae fam. nov. y Hantazuidae fam. nov.). También se restablece la superfamilia Embletonioidea, cuya posición había sido objeto de debate persistente en la literatura molecular.
Esta nueva clasificación, aunque compleja, ofrece un marco evolutivo más realista y pone de manifiesto que la diversidad de los nudibranquios es mucho mayor de lo que se había supuesto. El enfoque integrador de Korshunova et al. (2025) sostiene que solo el análisis conjunto de morfología, genética y desarrollo puede evitar distorsiones derivadas de la “optimización” taxonómica y restituir así la auténtica riqueza evolutiva del grupo. En definitiva, su trabajo redefine Nudibranchia no como un residuo de los antiguos “opistobranquios”, sino como un clado bien delimitado dentro de los Heterobranchia, en el que la evolución de las formas corporales y de los sistemas defensivos químicos ha generado una diversidad que, como sugiere el título del artículo, es realmente “inacabable y extraordinariamente diferenciada”.
Los Nudibranchia, como grupo, no presentan una organización corporal homogénea: los Aeolidioidea, Fionoidea, Flabellinoidea y Proctonotoidea tienen el cuerpo alargado y grupos de papilas llamados cerata a ambos lados del cuerpo. Los Arminoidea presentan un cuerpo de forma más o menos triangular y viven en fondos blandos. Los Dendronotoidea y Tritonioidea tienen el cuerpo alargado y prolongaciones dendríticas a los lados. Los Doridoxoidea muestran una morfología corporal semejante a la de los nudibranquios dorídidos, es decir, aplanada dorsoventralmente.
La sinopsis sistemática de los Nudibranchia según Bouchet et al. (2017) y reflejada en WoRMS es como sigue:
Nudibranchia Cuvier, 1817
- Aeolidina Eliot, 1910
- 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
- 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
- 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
- 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
- Chudoidea Korshunova, Fletcher & Martynov, 2025
- Chudidae Korshunova, Fletcher & Martynov, 2025
- Chudo Korshunova, Fletcher & Martynov, 2025
- Embletonioidea Pruvot-Fol, 1954
- Embletoniidae Pruvot-Fol, 1954
- Embletonia Alder & Hancock, 1851
- 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
- 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
- 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
- Notaeolidioidea Eliot, 1910
- 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
- 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
- Arminina
- 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
- Heterodoridoidea A. E. Verrill & Emerton, 1882
- Doridomorphidae Er. Marcus & Ev. Marcus, 1960 (1908)
- Doridoxidae Bergh, 1899
- Heterodorididae A. E. Verrill & Emerton, 1882
- Heterodoris A. E. Verrill & Emerton, 1882
- Dendronotina Odhner, 1934
- Dendronotoidea Allman, 1845
- Bornellidae Bergh, 1874
- 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
- Lomanotidae Bergh, 1890
- 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
- Janolina
- Goniaeolididae Odhner, 1907
- Proctonotoidea J. E. Gray, 1853
- Curnonidae d’Udekem d’Acoz, 2017
- Curnon d’Udekem d’Acoz, 2017
- Pseudotritonia Thiele, 1912
- Dironidae Eliot, 1910
- 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
- Tritonina
- 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
Grupos principales de nudibranquios:
Bibliografía
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Nudibranchia
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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.