Inicio Guía de Opistobranquios Nudibranchia Doridina

Doridina

Doridina es uno de los dos grandes linajes evolutivos de los Nudibranchia, siendo el otro Cladobranchia. Comprende los nudibranquios doridáceos, un grupo muy diverso de babosas marinas caracterizado por un plan corporal compacto y por la presencia de un conjunto de branquias secundarias dispuestas alrededor del ano. Los nudibranquios doridáceos también son conocidos comúnmente como doridáceos o doridos.

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

Los nudibranquios doridáceos presentan generalmente un cuerpo alargado, ovalado o aplanado dorsoventralmente, en el que el notum dorsal está claramente diferenciado de los lados del cuerpo. Los rinóforos pueden ser contráctiles y, en muchas especies, pueden retraerse completamente en cavidades rinofóricas especializadas. La superficie dorsal puede ser lisa o estar cubierta de tubérculos de diferentes formas, incluidos tubérculos redondeados, cónicos o digitiformes. En algunos géneros, como Jorunna, Taringa y Rostanga, los tubérculos contienen espículas conspicuas que sobresalen alrededor de sus márgenes y que reciben comúnmente el nombre de caryophyllidia.

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

La glándula digestiva de los nudibranquios doridáceos es compacta, en lugar de estar extendida por los cerata característicos de los nudibranquios cladobranquios. Esta diferencia en la anatomía digestiva es una de las principales características utilizadas históricamente para distinguir los dos grandes linajes de nudibranquios. Los doridáceos también presentan una considerable diversidad en la ornamentación corporal, la coloración, la química defensiva, las estructuras alimentarias y la anatomía reproductora.

Los nudibranquios doridáceos son predominantemente carnívoros y muchas especies presentan un elevado grado de especialización en su dieta. Las esponjas constituyen el principal alimento de numerosos linajes de doridáceos, mientras que otras especies se alimentan de briozoos, ascidias, cnidarios u otros invertebrados marinos sésiles. Estas relaciones alimentarias especializadas están frecuentemente asociadas con la adquisición, modificación o síntesis de compuestos químicos defensivos y han desempeñado un papel importante en la diversificación ecológica del grupo.

La clasificación tradicional de los nudibranquios doridáceos reconoce dos grandes infraórdenes: Bathydoridoidei y Doridoidei. Bathydoridoidei comprende un pequeño linaje de doridáceos predominantemente de aguas profundas, mientras que Doridoidei contiene la gran mayoría de la diversidad de doridáceos e incluye los principales 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 no asignados a ninguna superfamilia.

La posición taxonómica de los nudibranquios doridáceos ha experimentado un cambio importante en los últimos años. En la clasificación propuesta por Korshunova et al. (2025), el linaje tradicional de los doridáceos fue eliminado de Nudibranchia y elevado a la categoría de un orden independiente, Doridida. Según esta interpretación, el orden Nudibranchia restringido estaba constituido únicamente por Cladobranchia, mientras que Doridida representaba el linaje de los doridáceos como un orden independiente.

Esta interpretación fue posteriormente reconsiderada por Moles et al. (2026) en el marco de un análisis más amplio sobre la estabilidad taxonómica de las babosas marinas. Estos autores defendieron que el nombre establecido Nudibranchia debía conservarse en su sentido amplio e históricamente reconocido cuando corresponde a un grupo evolutivo coherente, en lugar de restringirlo innecesariamente a uno de sus principales linajes. En consecuencia, apoyaron el mantenimiento de Doridina y Cladobranchia dentro de Nudibranchia.

En el marco adoptado aquí, los nudibranquios doridáceos se consideran, por tanto, Doridina, uno de los dos linajes principales incluidos dentro del orden Nudibranchia. Esta disposición conserva el concepto tradicional y amplio de Nudibranchia, al tiempo que mantiene Doridina como un linaje distinto y filogenéticamente significativo.

Este marco es coherente con la interpretación filogenética más amplia según la cual Nudibranchia comprende dos linajes principales, Doridina y Cladobranchia. Los estudios moleculares recientes también han recuperado estos dos grupos como ramas principales fuertemente respaldadas dentro de Nudibranchia, reforzando la distinción entre los linajes doridáceo y cladobranquio y apoyando al mismo tiempo su inclusión dentro de un único orden.

Doridina presenta una notable diversidad morfológica y ecológica a pesar de la similitud fundamental de su organización respiratoria. Las especies abarcan desde formas relativamente lisas y de coloración críptica hasta animales muy ornamentados, con elaborados tubérculos, crestas, apéndices y llamativos patrones de coloración aposemática. El grupo incluye tanto especies de aguas someras asociadas a fondos rocosos, esponjas, briozoos y otras comunidades bentónicas como linajes especializados de aguas profundas representados por Bathydoridoidei.

La evolución de Doridina ha sido especialmente importante en los estudios sobre morfología, desarrollo y ecología química de los nudibranquios. Sus relaciones alimentarias especializadas y sus diversos mecanismos defensivos han contribuido a la extraordinaria radiación del grupo, mientras que los estudios anatómicos y moleculares comparativos continúan perfeccionando nuestra comprensión de las relaciones entre sus principales linajes.

La clasificación sistemática de Doridina presentada a continuación sigue el concepto amplio de Nudibranchia respaldado por Moles et al. (2026), en el que Doridina y Cladobranchia son reconocidos como los dos linajes principales del orden. La clasificación se basa en Bouchet et al. (2017) y en su tratamiento posterior 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.