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Cladobranchia

Cladobranchia

Cladobranchia es uno de los dos grandes linajes evolutivos de Nudibranchia, siendo el otro Doridina. Comprende un conjunto muy diverso de babosas marinas caracterizado por una extraordinaria variedad de formas corporales y de estructuras dorsales y laterales especializadas. A diferencia de los nudibranquios doridáceos, que poseen una característica corona o círculo de branquias secundarias alrededor del ano, los cladobranquios presentan una gran diversidad de estructuras respiratorias y sensoriales, entre ellas cerata, procesos dorsales ramificados y otros apéndices especializados.

Candiella striata por Pontes, Miquel

El nombre Cladobranchia fue introducido por Willan y Morton (1984) para un grupo importante de nudibranquios que incluía los tradicionales linajes de aeólidos, armináceos y dendronotáceos. El nombre hace referencia a las estructuras respiratorias ramificadas o de otro modo diferenciadas que caracterizan a muchos miembros del grupo. Posteriormente, Cladobranchia se consolidó como uno de los dos principales linajes evolutivos de Nudibranchia, junto con el linaje doridáceo, tradicionalmente denominado Anthobranchia y posteriormente Doridina.

Cladobranchia presenta una enorme diversidad morfológica. Muchas especies poseen apéndices dorsales especializados que intervienen en la respiración, la defensa, la alimentación o las funciones sensoriales. Los nudibranquios 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 estructuras defensivas especializadas denominadas cnidosacos. Otros cladobranquios poseen procesos dorsales ramificados o dendríticos, estructuras orales expandidas, vainas rinofóricas u otros apéndices especializados. Esta extraordinaria diversidad morfológica refleja la extensa radiación evolutiva que ha experimentado el grupo.

El concepto tradicional de Cladobranchia fue progresivamente refinado mediante estudios anatómicos comparativos y filogenéticos. Wägele y Willan (2000) reconocieron Cladobranchia como una de las dos divisiones principales de Nudibranchia, junto con Anthobranchia. Los posteriores estudios moleculares y transcriptómicos ampliaron considerablemente nuestro conocimiento de las relaciones dentro de Cladobranchia y demostraron que varios grupos tradicionales eran más complejos de lo que se había supuesto. Los datos moleculares también revelaron relaciones entre linajes que no siempre podían inferirse únicamente a partir de la morfología externa.

El desarrollo de la filogenética molecular durante el siglo XXI condujo, por tanto, a importantes cambios en la clasificación superior de Cladobranchia. Los agrupamientos tradicionales basados en la forma corporal general, la disposición de los cerata o determinadas estructuras relacionadas con la alimentación demostraron, en algunos casos, no corresponderse con linajes evolutivos monofiléticos. Como consecuencia, se establecieron varias familias y superfamilias nuevas, mientras que otras fueron redefinidas o reorganizadas a medida que se acumulaban nuevas evidencias filogenéticas.

La clasificación propuesta por Korshunova et al. (2025) representó una de las reorganizaciones más extensas de Cladobranchia realizadas en los últimos años. En su tratamiento, el orden Nudibranchia quedó restringido esencialmente a Cladobranchia, mientras que los nudibranquios doridáceos fueron excluidos del orden y reconocidos como un orden separado, Doridida. Dentro de este concepto restringido de Nudibranchia, los autores reinstauraron varios subórdenes tradicionales, entre ellos Arminina, Tritonina, Dendronotina, Janolina y Aeolidina, y reorganizaron sustancialmente la taxonomía superior del grupo.

Esta interpretación fue posteriormente reconsiderada por Moles et al. (2026) en su análisis de la estabilidad taxonómica de los heterobranquios marinos. Los autores argumentaron que las decisiones taxonómicas deben tener en cuenta no solo las relaciones filogenéticas, sino también la capacidad de diagnóstico, la estabilidad nomenclatural, la compatibilidad con las clasificaciones existentes y los posibles efectos colaterales de los cambios taxonómicos. En el caso de Nudibranchia, concluyeron que restringir el nombre establecido Nudibranchia a Cladobranchia no proporciona una ventaja filogenética o práctica suficiente que justifique sustituir su uso tradicional más amplio.

Según el marco respaldado por Moles et al. (2026), Nudibranchia se mantiene, por tanto, en su sentido amplio e históricamente establecido, incluyendo dos grandes linajes principales: Doridina y Cladobranchia. En consecuencia, Cladobranchia conserva su condición de gran linaje monofilético dentro de Nudibranchia, en lugar de considerarse equivalente a todo el orden. Esta disposición preserva tanto el nombre históricamente establecido Nudibranchia como la identidad evolutiva bien sustentada de Cladobranchia.

Las evidencias moleculares recientes son coherentes con este marco amplio. Los análisis mitogenómicos han recuperado Doridina y Cladobranchia como dos grandes ramas fuertemente sustentadas de Nudibranchia. Al mismo tiempo, las relaciones entre los principales linajes cladobranquios siguen siendo objeto de investigación activa, y los estudios moleculares continúan revelando relaciones evolutivas anteriormente desconocidas y casos en los que los grupos tradicionales no son monofiléticos.

La diversidad de Cladobranchia se refleja en su clasificación superior. El grupo incluye las superfamilias de aeólidos Aeolidioidea, Apataoidea, Chudoidea, Embletonioidea, Fionoidea, Notaeolidioidea, Samloidea y Unidentioidea, junto con Arminoidea, Heterodoridoidea, Dendronotoidea, Proctonotoidea y Tritonioidea. Estos grupos abarcan una extraordinaria variedad de formas corporales y estrategias ecológicas, desde los aeólidos provistos de cerata altamente especializados hasta las formas dendríticas de Dendronotoidea y Tritonioidea y las formas a menudo alargadas o aplanadas que se encuentran en Arminoidea y Proctonotoidea.

Los nudibranquios aeólidos son especialmente conocidos por sus cerata, que pueden actuar simultáneamente como estructuras respiratorias, prolongaciones de la glándula digestiva y órganos defensivos. Algunas especies son capaces de retener nematocistos funcionales obtenidos de sus presas cnidarias e incorporarlos a cnidosacos situados en los extremos de los cerata. Otros cladobranquios han desarrollado mecanismos defensivos y respiratorios diferentes, entre ellos elaborados procesos ramificados, formas corporales crípticas, defensas químicas y asociaciones especializadas con determinadas presas.

Cladobranchia presenta también una extraordinaria diversidad ecológica. Sus miembros habitan desde aguas costeras poco profundas y ambientes intermareales hasta comunidades más profundas de la plataforma y el talud continental. Muchas especies están estrechamente asociadas a determinados organismos presa, entre ellos hidrozoos, octocorales, otros cnidarios, esponjas y briozoos. Estas relaciones tróficas especializadas han desempeñado un papel importante en la diversificación de varios linajes cladobranquios y se reflejan con frecuencia en su morfología, comportamiento y ecología química.

La historia evolutiva de Cladobranchia constituye, por tanto, un excelente ejemplo de la complejidad de la diversificación de los nudibranquios. La amplia variedad de arquitecturas corporales, estructuras respiratorias, estrategias alimentarias y mecanismos defensivos que se encuentran dentro del grupo no puede explicarse adecuadamente mediante una clasificación basada únicamente en la morfología. La integración de la anatomía comparada, la biología del desarrollo, la filogenética molecular y la información ecológica continúa revelando nuevas relaciones evolutivas y permite perfeccionar la clasificación del grupo.

Según la clasificación adoptada aquí, Cladobranchia se considera uno de los dos grandes linajes de Nudibranchia, junto con Doridina. 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 respaldado 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
  • 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
      • 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
  • Notaeolidioidea Eliot, 1910
    • Notaeolidiidae Eliot, 1910
      • Notaeolidia Eliot, 1905
  • 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
  • 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
  • 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
  • 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
  • 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 Cladobranchia:

Bibliografía

    1246567 Cladobranchia 1 eunomia-revista-en-cultura-de-la-legalidad 50 creator asc 1 48603 https://opistobranquis.info/wp-content/plugins/zotpress/
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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.