Inicio Guía de Opistobranquios Nudibranchia Doridina

Doridina

Doridina es uno de los dos grandes linajes evolutivos de los Nudibranchia, junto con Cladobranchia. Comprende los nudibranquios doridáceos, un grupo muy diverso de babosas marinas caracterizado por un plano corporal compacto y por la presencia de un penacho de branquias secundarias dispuesto alrededor del ano. Los miembros del grupo son conocidos habitualmente como dóridos o doridáceos.

Peltodoris atromaculata @ Costa Brava, Spain by Enric Madrenas por Enric Madrenas

Los dóridos presentan generalmente un cuerpo ovalado, alargado o aplanado dorsoventralmente, con el notum dorsal bien diferenciado de los lados del cuerpo. Los rinóforos pueden ser contráctiles y, en muchas especies, retraerse completamente dentro de cavidades rinofóricas especializadas. La superficie dorsal puede ser lisa o presentar tubérculos de formas diversas, incluyendo estructuras redondeadas, cónicas o digitiformes. En algunos géneros, como Jorunna, Taringa y Rostanga, los tubérculos contienen espículas conspicuas que sobresalen por sus márgenes y forman las estructuras conocidas como cariophyllidia.

El penacho branquial se sitúa en la región posterior de la superficie dorsal y rodea la abertura anal. Las branquias pueden estar formadas por hojas simples o ramificadas y pueden permanecer permanentemente expuestas o retraerse dentro de una cavidad branquial especializada. Tradicionalmente, los dóridos con las branquias expuestas se denominaban fanerobranquios (phanerobranchs), mientras que aquellos capaces de retraerlas dentro de una cavidad branquial se denominaban criptobranquios (cryptobranchs). Sin embargo, los estudios anatómicos y del desarrollo han demostrado que esta distinción no representa una secuencia evolutiva simple y que la evolución de la cavidad branquial es más compleja de lo que se consideraba anteriormente.

La glándula digestiva de los dóridos es compacta y está formada por un número variable de lóbulos. El grupo presenta una considerable diversidad en la ornamentación corporal, la coloración, las estructuras alimentarias, la anatomía reproductora y los mecanismos de defensa química.

Los dóridos son predominantemente carnívoros y muchas especies presentan un elevado grado de especialización alimentaria. Las esponjas constituyen el alimento principal de numerosos linajes, mientras que otras especies se alimentan de briozoos, ascidias, cnidarios u otros invertebrados marinos sésiles. Estas relaciones alimentarias especializadas pueden estar asociadas con la adquisición, modificación o síntesis de compuestos químicos defensivos.

La clasificación tradicional de Doridina reconoce dos grandes infraórdenes: Bathydoridoidei y Doridoidei. Bathydoridoidei comprende un pequeño linaje de dóridos predominantemente asociado a ambientes profundos, mientras que Doridoidei concentra la mayor parte de la diversidad del grupo e incluye numerosos linajes de aguas someras. Dentro de Doridoidei, la clasificación de Bouchet et al. (2017) reconoce las superfamilias Chromodoridoidea, Doridoidea, Onchidoridoidea, Phyllidioidea y Polyceroidea, junto con algunos taxones sin asignación a una superfamilia.

La delimitación taxonómica de Doridina ha sido objeto de debate reciente. Korshunova et al. (2025) propusieron separar este linaje y reconocerlo como el orden Doridida. Posteriormente, Moles et al. (2026) defendieron la conservación de la clasificación tradicional y el mantenimiento de Doridina dentro de Nudibranchia. En este tratamiento se sigue esta última interpretación.

Doridina presenta una notable diversidad morfológica y ecológica. Las especies van desde formas relativamente lisas y de coloración críptica hasta animales muy ornamentados, con tubérculos, crestas, apéndices y patrones de coloración conspicuos. El grupo ocupa principalmente ambientes bentónicos e incluye tanto especies de aguas someras asociadas a fondos rocosos y comunidades bentónicas como linajes especializados de aguas profundas.

La especialización alimentaria, la diversidad morfológica y los diversos mecanismos de defensa química de los dóridos han desempeñado un papel importante en su diversificación y constituyen ámbitos destacados de investigación en anatomía comparada, ecología química, desarrollo y evolución.

La clasificación sistemática de Doridina presentada a continuación se basa en Bouchet et al. (2017) y en su tratamiento en el World Register of Marine Species (WoRMS), teniendo en cuenta las revisiones sistemáticas posteriores:

Doridina

  • Bathydoridoidei
    • Bathydoridoidea Bergh, 1891
      • Bathydorididae Bergh, 1891
        • Bathydoris Bergh, 1884
        • Prodoris Baranetz & Minichev, 1995
  • Doridoidei
    • Chromodoridoidea Bergh, 1891
      • Actinocyclidae O’Donoghue, 1929
        • Actinocyclus Ehrenberg, 1831
        • Hallaxa Eliot, 1909
      • Cadlinellidae Odhner, 1934
        • Cadlinella Thiele, 1931
      • Cadlinidae Bergh, 1891
        • Aldisa Bergh, 1878
        • Cadlina Bergh, 1879
        • Inuda Ev. Marcus & Er. Marcus, 1967
      • Chromodorididae Bergh, 1891
        • Chromodoridinae Bergh, 1891
          • Ardeadoris Rudman, 1984
          • Chromodoris Alder & Hancock, 1855
          • Chromolaichma Bertsch, 1977
          • Diversidoris Rudman, 1987
          • Doriprismatica A. d’Orbigny, 1839
          • Felimida Ev. Marcus, 1971
          • Glossodoris Ehrenberg, 1831
          • Goniobranchus Pease, 1866
          • Tyrinna Bergh, 1898
          • Verconia Pruvot-Fol, 1931
        • Miamirinae Bergh, 1891
          • Ceratosoma Gray, 1850
          • Felimare Ev. Marcus & Er. Marcus, 1967
          • Hypselodoris W. Stimpson, 1855
          • Mexichromis Bertsch, 1977
          • Miamira Bergh, 1874
          • Thorunna Bergh, 1878
      • Hexabranchidae Bergh, 1891
        • Hexabranchus Ehrenberg, 1828
      • Showajidaiidae Korshunova, Fletcher, Picton, Lundin, Kashio, N. Sanamyan, K. Sanamyan, Padula, Schrödl & Martynov, 2020
        • Showajidaia Korshunova, Fletcher, Picton, Lundin, Kashio, N. Sanamyan, K. Sanamyan, Padula, Schrödl & Martynov, 2020
    • Doridoidea Rafinesque, 1815
      • Discodorididae Bergh, 1891
        • Alloiodoris Bergh, 1904
        • Aphelodoris Bergh, 1879
        • Aporodoris Ihering, 1886
        • Asteronotus Ehrenberg, 1831
        • Atagema Gray, 1850
        • Avaldesia Donohoo & Gosliner, 2024
        • Baptodoris Bergh, 1884
        • Carminodoris Bergh, 1889
        • Diaulula Bergh, 1878
        • Dictyodoris Bergh, 1880
        • Discodoris Bergh, 1877
        • Gargamella Bergh, 1894
        • Geitodoris Bergh, 1891
        • Halgerda Bergh, 1880
        • Hiatodoris Dayrat, 2010
        • Hoplodoris Bergh, 1880
        • Jorunna Bergh, 1876
        • Nirva Bergh, 1905
        • Nophodoris Á. Valdés & Gosliner, 2001
        • Otinodoris K. M. White, 1948
        • Paradoris Bergh, 1884
        • Peltodoris Bergh, 1880
        • Peronodoris Bergh, 1904
        • Platydoris Bergh, 1877
        • Rostanga Bergh, 1879
        • Sclerodoris Eliot, 1904
        • Sebadoris Er. Marcus & Ev. Marcus, 1960
        • Taringa Er. Marcus, 1955
        • Tayuva Ev. Marcus & Er. Marcus, 1967
        • Thordisa Bergh, 1877
        • Thorybopus Bouchet, 1977
      • Dorididae Rafinesque, 1815
        • Archidoris Bergh, 1878
        • Artachaea Bergh, 1881
        • Austrodoris Odhner, 1926
        • Conualevia Collier & Farmer, 1964
        • Doriopsis Pease, 1860
        • Doris Linnaeus, 1758
        • Goslineria Á. Valdés, 2001
        • Pharodoris Á. Valdés, 2001
    • Onchidoridoidea Gray, 1827
      • Acanthodorididae P. Fischer, 1883
        • Acanthodoris J. E. Gray, 1850
      • Acanthomiridae Martynov & Korshunova, 2025
      • Aegiridae P. Fischer, 1883
        • Aegires Lovén, 1844
        • Notodoris Bergh, 1875
      • Akiodorididae Millen & Martynov, 2005
        • Akiodoris Bergh, 1879
        • Armodoris Minichev, 1972
        • Doridunculus G. O. Sars, 1878
        • Echinocorambe Á. Valdés & Bouchet, 1998
        • Prodoridunculus Thiele, 1912
      • Calycidorididae Roginskaya, 1972
        • Calycidoris Abraham, 1876
        • Diaphorodoris Iredale & O’Donoghue, 1923
      • Corambidae Bergh, 1871
        • Corambe Bergh, 1869
        • Loy Martynov, 1994
      • Goniodorididae H. Adams & A. Adams, 1854
        • Anculinae Pruvot-Fol, 1954
          • Ancula Lovén, 1846
          • Trapania Pruvot-Fol, 1931
        • Goniodoridinae H. Adams & A. Adams, 1854
          • Bermudella Odhner, 1941
          • Cargoa Vogel & L. P. Schultz, 1970
          • Ceratodoris J. E. Gray, 1850
          • Goniodoridella Pruvot-Fol, 1933
          • Lophodoris G. O. Sars, 1878
          • Murphydoris Sigurdsson, 1991
          • Naisdoris Paz-Sedano, Moles, Smirnoff, Gosliner & Pola, 2024
          • Okenia Menke, 1830
          • Pelagella J. E. Gray, 1850
        • Spahria Risbec, 1928
      • Onchidorididae J. E. Gray, 1827
        • Acanthomira Ekimova, Nikitenko, Stanovova, Schepetov, Antokhina & Á. Valdés, 2024
        • Adalaria Bergh, 1878
        • Arctadalaria Roginskaya, 1971
        • Atalodoris Iredale & O’Donoghue, 1923
        • Idaliadoris Furfaro & Trainito, 2022
        • Onchidoris Blainville, 1816
        • Onchimira Martynov, Korshunova, N. Sanamyan & K. Sanamyan, 2009
    • Phyllidioidea Rafinesque, 1814
      • Dendrodorididae O’Donoghue, 1924 (1864)
        • Dendrodoris Ehrenberg, 1831
        • Doriopsilla Bergh, 1880
      • Forjacellidae Ortea & Moro, 2023
        • Forjacella Ortea & Moro, 2023
      • Mandeliidae Á. Valdés & Gosliner, 1999
        • Mandelia Á. Valdés & Gosliner, 1999
      • Phyllidiidae Rafinesque, 1814
        • Ceratophyllidia Eliot, 1903
        • Phyllidia Cuvier, 1797
        • Phyllidiella Bergh, 1869
        • Phyllidiopsis Bergh, 1876
        • Reticulidia Brunckhorst, 1990
    • Polyceroidea Alder & Hancock, 1845
      • Polyceridae Alder & Hancock, 1845
        • Kalinginae Pruvot-Fol, 1956
          • Kalinga Alder & Hancock, 1864
        • Kankelibranchinae Ortea, Espinosa & Caballer, 2005
          • Kankelibranchus Ortea, Espinosa & Caballer, 2005
        • Nembrothinae Burn, 1967
          • Martadoris Willan & Y.-W. Chang, 2017
          • Nembrotha Bergh, 1877
          • Roboastra Bergh, 1877
          • Tambja Burn, 1962
          • Tyrannodoris Willan & Y.-W. Chang, 2017
        • Okadaiinae Baba, 1930
          • Vayssierea Risbec, 1928
        • Polycerinae Alder & Hancock, 1845
          • Gymnodoris W. Stimpson, 1855
          • Lamellana G.-Y. Lin, 1992
          • Lecithophorus Macnae, 1958
          • Palio Gray, 1857
          • Paliolla Burn, 1958
          • Paliota Pola, Miguel-González & Paz-Sedano, 2023
          • Polycera Cuvier, 1816
          • Polycerella A. E. Verrill, 1880
          • Thecacera J. Fleming, 1828
        • Triophinae Odhner, 1941
          • Colga Bergh, 1880
          • Crimora Alder & Hancock, 1862
          • Holoplocamus Odhner, 1926
          • Joubiniopsis Risbec, 1928
          • Kaloplocamus Bergh, 1880
          • Limacia O. F. Müller, 1781
          • Plocamopherus Rüppell & Leuckart, 1828
          • Triopha Bergh, 1880

Grupos principales de Doridina:

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.