Cladobranchia

Cladobranchia és un dels dos grans llinatges evolutius de Nudibranchia, juntament amb Doridina. Comprèn un grup extraordinàriament divers de llimacs marins, caracteritzat per una gran varietat de formes corporals i d’estructures dorsals i laterals especialitzades.

Candiella striata per Pontes, Miquel

Cladobranchia presenta una notable diversitat morfològica. Moltes espècies posseeixen apèndixs dorsals especialitzats relacionats amb la respiració, la defensa, l’alimentació o les funcions sensorials. Els aeòlids es caracteritzen típicament per nombrosos cerata distribuïts per la superfície dorsal. Aquestes estructures contenen prolongacions de la glàndula digestiva i, en moltes espècies, incorporen cnidosacs, estructures especialitzades relacionades amb la defensa. Altres cladobranquis presenten processos dorsals ramificats o dendrítics, estructures orals eixamplades, beines rinofòriques o altres apèndixs especialitzats.

El nom Cladobranchia va ser introduït per Willan i Morton (1984) per a un grup de nudibranquis que incloïa els principals llinatges tradicionalment reconeguts com a aeòlids, arminacis i dendronotacis. Posteriorment, el concepte del grup va ser revisat mitjançant estudis anatòmics comparatius i filogenètics. L’arribada de les dades moleculars va permetre revisar les relacions entre diversos llinatges i va posar de manifest que algunes agrupacions tradicionals basades en la morfologia no corresponien necessàriament a grups evolutius naturals.

La filogenètica molecular ha comportat nombrosos canvis en la classificació superior de Cladobranchia. Diverses famílies i superfamílies han estat creades, redefinides o reorganitzades a mesura que s’han incorporat noves dades. Les relacions entre alguns dels principals llinatges del grup continuen sent objecte d’estudi, i la classificació ha anat evolucionant amb l’acumulació de noves evidències.

La classificació proposada per Korshunova et al. (2025) va introduir una important reorganització de la classificació superior dels nudibranquis i va adoptar un concepte restringit de Nudibranchia. Posteriorment, Moles et al. (2026) van reconsiderar aquesta proposta en el context de la seva anàlisi de l’estabilitat taxonòmica dels heterobranquis marins. En el tractament adoptat aquí, es manté el concepte ampli de Nudibranchia i Cladobranchia es considera un dels seus dos grans llinatges principals.

Cladobranchia comprèn una gran diversitat de grups, entre els quals es troben nombrosos llinatges d’aeòlids i altres formes tradicionalment agrupades entre els arminacis i dendronotacis. La classificació superior ha experimentat diverses modificacions i, per aquest motiu, la delimitació i l’assignació d’alguns grups poden variar segons la font taxonòmica utilitzada.

Els aeòlids són especialment coneguts pels seus cerata, que poden exercir funcions respiratòries, digestives i defensives. Algunes espècies són capaces de retenir nematocists funcionals obtinguts de les seves preses cnidàries i incorporar-los als cnidosacs situats als extrems dels cerata. Altres cladobranquis presenten mecanismes defensius diferents, entre els quals hi ha defenses químiques, coloracions críptiques i estructures corporals especialitzades.

Cladobranchia ocupa una àmplia varietat d’ambients marins, des d’aigües costaneres poc profundes i zones intermareals fins a ambients més profunds de la plataforma i el talús continental. Moltes espècies mantenen relacions tròfiques especialitzades amb determinats organismes, entre els quals hi ha hidrozoos, octocoralls i altres cnidaris, esponges i briozous. Aquestes especialitzacions alimentàries poden estar associades amb adaptacions morfològiques, comportamentals i químiques.

La diversitat de Cladobranchia es manifesta tant en la seva morfologia com en les seves estratègies ecològiques. La gran varietat d’estructures corporals, formes d’alimentació i mecanismes defensius que presenta el grup reflecteix una història evolutiva complexa. Els estudis anatòmics, del desenvolupament, moleculars i ecològics continuen aportant informació per comprendre millor les relacions entre els seus diferents llinatges i la seva diversificació.

La classificació que es presenta a continuació reflecteix el marc taxonòmic actual utilitzat al World Register of Marine Species (WoRMS), tenint en compte el concepte més ampli de Nudibranchia avalat per 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

Grups principals de Cladobranchia:

Bibliografia

    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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    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.