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Doridina

Doridina és un dels dos grans llinatges evolutius dels Nudibranchia, juntament amb Cladobranchia. Comprèn els nudibranquis doridacis, un grup molt divers de llimacs marins caracteritzat per un pla corporal compacte i per la presència d’un plomall de brànquies secundàries disposat al voltant de l’anus. Els membres del grup són coneguts habitualment com a dòrids o doridacis.

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

Els dòrids presenten generalment un cos ovalat, allargat o aplanat dorsoventralment, amb el notum dorsal ben diferenciat dels costats del cos. Els rinòfors poden ser contràctils i, en moltes espècies, retraure’s completament dins de cavitats rinofòriques especialitzades. La superfície dorsal pot ser llisa o presentar tubercles de formes diverses, incloent-hi estructures arrodonides, còniques o digitiformes. En alguns gèneres, com Jorunna, Taringa i Rostanga, els tubercles contenen espícules conspícues que sobresurten pels seus marges i formen les estructures conegudes com a cariòfil·lidis (caryophyllidia).

El plomall branquial se situa a la regió posterior de la superfície dorsal i envolta l’obertura anal. Les brànquies poden estar formades per fulles simples o ramificades i poden romandre permanentment exposades o retraure’s dins d’una cavitat branquial especialitzada. Tradicionalment, els dòrids amb les brànquies exposades es denominaven fanerobranquis (phanerobranchs), mentre que aquells capaços de retraure-les dins d’una cavitat branquial es denominaven criptobranquis (cryptobranchs). Tanmateix, els estudis anatòmics i del desenvolupament han demostrat que aquesta distinció no representa una seqüència evolutiva simple i que l’evolució de la cavitat branquial és més complexa del que es considerava anteriorment.

La glàndula digestiva dels dòrids és compacta i està formada per un nombre variable de lòbuls. El grup presenta una considerable diversitat en l’ornamentació corporal, la coloració, les estructures alimentàries, l’anatomia reproductora i els mecanismes de defensa química.

Els dòrids són predominantment carnívors i moltes espècies presenten un elevat grau d’especialització alimentària. Les esponges constitueixen l’aliment principal de nombrosos llinatges, mentre que altres espècies s’alimenten de briozous, ascídies, cnidaris o altres invertebrats marins sèssils. Aquestes relacions alimentàries especialitzades poden estar associades amb l’adquisició, modificació o síntesi de compostos químics defensius.

La classificació tradicional de Doridina reconeix dos grans infraordres: Bathydoridoidei i Doridoidei. Bathydoridoidei comprèn un petit llinatge de dòrids predominantment associat a ambients profunds, mentre que Doridoidei concentra la major part de la diversitat del grup i inclou nombrosos llinatges d’aigües somes. Dins de Doridoidei, la classificació de Bouchet et al. (2017) reconeix les superfamílies Chromodoridoidea, Doridoidea, Onchidoridoidea, Phyllidioidea i Polyceroidea, juntament amb alguns tàxons sense assignació a una superfamília.

La delimitació taxonòmica de Doridina ha estat objecte de debat recent. Korshunova et al. (2025) van proposar separar aquest llinatge i reconèixer-lo com l’ordre Doridida. Posteriorment, Moles et al. (2026) van defensar la conservació de la classificació tradicional i el manteniment de Doridina dins de Nudibranchia. En aquest tractament se segueix aquesta darrera interpretació.

Doridina presenta una notable diversitat morfològica i ecològica. Les espècies van des de formes relativament llises i de coloració críptica fins a animals molt ornamentats, amb tubercles, crestes, apèndixs i patrons de coloració conspicus. El grup ocupa principalment ambients bentònics i inclou tant espècies d’aigües somes associades a fons rocosos i comunitats bentòniques com llinatges especialitzats d’aigües profundes.

L’especialització alimentària, la diversitat morfològica i els diversos mecanismes de defensa química dels dòrids han tingut un paper important en la seva diversificació i constitueixen àmbits destacats de recerca en anatomia comparada, ecologia química, desenvolupament i evolució.

La classificació sistemàtica de Doridina presentada a continuació es basa en Bouchet et al. (2017) i en el seu tractament al World Register of Marine Species (WoRMS), tenint en compte les revisions sistemàtiques posteriors:

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

Grups principals de Doridina:

Bibliografia

    1246567 Doridina 1 eunomia-revista-en-cultura-de-la-legalidad 50 creator asc 1 7700 https://opistobranquis.info/wp-content/plugins/zotpress/
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    Bibliografia basada en els treballs de Steve Long, 2006. Bibliography of Opisthobranchia 1554-2000 i Gary McDonald, 2009. Bibliographia Nudibranchia, amb actualitzacions posteriors procedents d´altres fonts.