Doridina

Doridina és un dels dos grans llinatges evolutius dels Nudibranchia, essent l’altre Cladobranchia. Comprèn els nudibranquis doridacis, un grup molt divers de llimacs marins caracteritzat per un pla corporal compacte i per la presència d’un conjunt de brànquies secundàries disposades al voltant de l’anus. Els nudibranquis doridacis també són coneguts habitualment com a doridacis o doridis.

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

Els nudibranquis doridacis presenten generalment un cos allargat, ovalat o aplanat dorsoventralment, en el qual el notum dorsal està clarament diferenciat dels costats del cos. Els rinòfors poden ser contràctils i, en moltes espècies, poden retraure’s completament dins de cavitats rinofòriques especialitzades. La superfície dorsal pot ser llisa o estar coberta de tubercles de formes diverses, incloent-hi tubercles arrodonits, cònics o digitiformes. En alguns gèneres, com Jorunna, Taringa i Rostanga, els tubercles contenen espícules conspícues que sobresurten al voltant dels seus marges i que reben habitualment el nom de cariòfil·lidis (caryophyllidia).

El plomall branquial es troba a la regió posterior de la superfície dorsal i envolta l’obertura anal. Les brànquies secundàries poden estar formades per fulles simples o ramificades i poden estar permanentment exposades o ser capaces de retraure’s dins d’una cavitat branquial especialitzada. Tradicionalment, els nudibranquis doridacis 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 moderns han demostrat que aquesta distinció no representa una seqüència evolutiva simple i que la relació entre la morfologia branquial i el desenvolupament de la cavitat branquial és més complexa del que es suposava anteriorment.

La glàndula digestiva dels nudibranquis doridacis és compacta, en lloc d’estar estesa pels cerata característics dels nudibranquis cladobranquis. Aquesta diferència en l’anatomia digestiva és una de les principals característiques utilitzades històricament per distingir els dos grans llinatges de nudibranquis. Els doridacis també presenten una considerable diversitat en l’ornamentació corporal, la coloració, la química defensiva, les estructures alimentàries i l’anatomia reproductora.

Els nudibranquis doridacis són predominantment carnívors i moltes espècies presenten un elevat grau d’especialització en la seva dieta. Les esponges constitueixen l’aliment principal de nombrosos llinatges de doridacis, mentre que altres espècies s’alimenten de briozous, ascídies, cnidaris o altres invertebrats marins sèssils. Aquestes relacions alimentàries especialitzades estan sovint associades amb l’adquisició, modificació o síntesi de compostos químics defensius i han tingut un paper important en la diversificació ecològica del grup.

La classificació tradicional dels nudibranquis doridacis reconeix dos grans infraordres: Bathydoridoidei i Doridoidei. Bathydoridoidei comprèn un petit llinatge de doridacis predominantment d’aigües profundes, mentre que Doridoidei conté la gran majoria de la diversitat dels doridacis i inclou els principals 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 no assignats a cap superfamília.

La posició taxonòmica dels nudibranquis doridacis ha experimentat un canvi important en els darrers anys. En la classificació proposada per Korshunova et al. (2025), el llinatge tradicional dels doridacis va ser eliminat de Nudibranchia i elevat a la categoria d’un ordre independent, Doridida. Segons aquesta interpretació, l’ordre Nudibranchia restringit estava constituït únicament per Cladobranchia, mentre que Doridida representava el llinatge dels doridacis com un ordre independent.

Aquesta interpretació va ser posteriorment reconsiderada per Moles et al. (2026) en el marc d’una anàlisi més àmplia sobre l’estabilitat taxonòmica dels llimacs marins. Aquests autors van defensar que el nom establert Nudibranchia s’havia de conservar en el seu sentit ampli i històricament reconegut quan correspon a un grup evolutiu coherent, en lloc de restringir-lo innecessàriament a un dels seus principals llinatges. En conseqüència, van donar suport al manteniment de Doridina i Cladobranchia dins de Nudibranchia.

En el marc adoptat aquí, els nudibranquis doridacis es consideren, per tant, Doridina, un dels dos llinatges principals inclosos dins de l’ordre Nudibranchia. Aquesta disposició conserva el concepte tradicional i ampli de Nudibranchia, alhora que manté Doridina com un llinatge diferenciat i filogenèticament significatiu.

Aquest marc és coherent amb la interpretació filogenètica més àmplia segons la qual Nudibranchia comprèn dos llinatges principals, Doridina i Cladobranchia. Els estudis moleculars recents també han recuperat aquests dos grups com a branques principals fortament recolzades dins de Nudibranchia, reforçant la distinció entre els llinatges doridaci i cladobranqui i donant suport alhora a la seva inclusió dins d’un únic ordre.

Doridina presenta una notable diversitat morfològica i ecològica malgrat la similitud fonamental de la seva organització respiratòria. Les espècies comprenen des de formes relativament llises i de coloració críptica fins a animals molt ornamentats, amb elaborats tubercles, crestes, apèndixs i patrons de coloració aposemàtica molt vistosos. El grup inclou tant espècies d’aigües somes associades a fons rocosos, esponges, briozous i altres comunitats bentòniques com llinatges especialitzats d’aigües profundes representats per Bathydoridoidei.

L’evolució de Doridina ha estat especialment important en els estudis sobre morfologia, desenvolupament i ecologia química dels nudibranquis. Les seves relacions alimentàries especialitzades i els seus diversos mecanismes defensius han contribuït a l’extraordinària radiació del grup, mentre que els estudis anatòmics i moleculars comparatius continuen perfeccionant la nostra comprensió de les relacions entre els seus principals llinatges.

La classificació sistemàtica de Doridina presentada a continuació segueix el concepte ampli de Nudibranchia recolzat per Moles et al. (2026), en el qual Doridina i Cladobranchia són reconeguts com els dos llinatges principals de l’ordre. La classificació es basa en Bouchet et al. (2017) i en el seu tractament posterior 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:

Doridoidei

Bathydoridoidei

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.