Inicio Guía de Opistobranquios Cephalaspidea

Cephalaspidea

Los Cefalaspídeos (CEPHALASPIDEA) son los opistobranquios más primitivos y tradicionalmente se han considerado como un grupo de transición entre los prosobranquios y los demás opistobranquios. Se conocen algo más de 800 especies en todo el mundo. Hace un tiempo se realizaron estudios filogenéticos que demostraron que los cefalaspídeos clásicos (sensu lato) no eran un grupo monofilético, por lo que lo escindieron en tres, los Cephalaspidea sensu stricto, los Runcinida (antiguos Runcinacea) y los Heterobranquios [no asignados] (antiguos Architectibranchia). Dentro de este último grupo, según estos estudios, familias como Acteonidae, Aplustridae y Ringiculidae ya no se consideran realmente opistobranquios, grupo dentro del cual se habían considerado hasta hace muy poco, aunque sí se clasifican como Heterobranquios.

Melanochlamys miqueli por Enric Madrenas

Los cefalaspídeos en sentido amplio (sensu lato) son los opistobranquios típicamente testáceos ya que muchas de sus especies poseen concha. Las especies de Scaphander y Haminoea, tienen una concha externa, bien calcificada, pero que está poco espiralizada y el animal no puede retraerse completamente dentro de ella. La concha puede estar muy poco calcificada y ser interna al haber quedado completamente cubierta por el manto, como ocurre en la familia Philinidae. La cabeza de los cefalaspídeos tiene un disco o escudo cefálico (Cephalaspidea= Cabeza con escudo) que facilita el enterramiento del animal en el substrato blando, hábitat habitual de los representantes de este grupo. Precisamente por ésto la cabeza no suele disponer de tentáculos u otras expansiones sensoriales. La cavidad paleal suele estar bien desarrollada y puede estar situada anteriormente o bien en posición posterior. Dentro de ella se encuentra una branquia de tipo plegado. En el tubo digestivo hay una molleja que posee placas endurecidas para triturar el alimento y la rádula puede faltar en algunas especies. En el aparato reproductor, el orificio genital está situado a cierta distancia del órgano copulador masculino por lo que existe entre ellos un surco seminal ciliado por donde discurre el esperma que se emite en la cópula, como ocurre también en los Aplysiida. El pie es de tipo reptante y diferencia, en algunas especies, un par de lóbulos laterales o parapodiales, como ocurre en Gastropteron rubrum, que se aplican sobre la concha en estado de reposo y pueden ser utilizados también para la natación.

Biología
Los cefalaspídeos generalmente son habitantes de substratos blandos, donde viven enterrados a varios centímetros de profundidad en busca de alimento. Algunas especies habitan en fondos de fango o cascajo que son caladeros de arrastre de barcos de pesca y por ello suelen aparecer entre los restos de los descartes no comerciales de la pesca, como ocurre con Scaphander lignarius, que puede alcanzar 80 mm de longitud y que se alimenta de foraminíferos, poliquetos, gasterópodos y pequeños bivalvos. En fondos similares viven  Philine quadripartita y Ph. catena, que se alimentan de poliquetos, foraminíferos y bivalvos. Las especies de Retusa y Cylichna viven en fondos de arena y fango o praderas de Posidonia oceanica alimentándose de foraminíferos y pequeños gasterópodos. Las especies del género Haminoea y Bulla son herbívoras y viven en fondos fangosos del interior de puertos o fondos de bahías someras con fanerógamas marinas, alimentándose de algas verdes, pardas y rojas y fanerógamas como Cymodocea nodosa, además de cianobacterias y diatomeas. Las especies de los géneros Navanax, Philinopsis, Aglaja y otros (familia  Aglajidae) son depredadores muy activos de turbelarios, nemertinos, poliquetos e incluso de nudibranquios y otros cefalaspídeos. En diferentes especies de cefalaspídeos se han encontrado metabolitos secundarios implicados en la defensa química del animal y también otros que funcionan como feromonas de alarma para la comunicación entre los diferentes ejemplares, como ocurre en las especies de Haminoea. Los cefalaspídeos son hermafroditas simultáneos y después de la cópula efectúan puestas en forma de cinta o cordón con numerosos huevos que depositan sobre la arena.

Distribución
Los Cefalaspídeos viven en todo tipo de substratos blandos, desde la zona intermareal hasta fondos circalitorales. Numerosas especies habitan aguas cálidas y templadas de los dos hemisferios e incluso se conocen algunas que prefieren las aguas frías polares, como por ejemplo Philine alata, de aguas antárticas. Scaphander tiene una veintena de especies que se distribuyen por los fondos blandos de cierta profundidad de todos los mares del mundo, desde el Ártico hasta las aguas antárticas. En la Península Ibérica, Baleares y Canarias se han citado algo más de un centenar de especies de Cefalaspídeos, que corresponden al 90% de todas las especies presentes en aguas europeas según el ERMS (European Register of Marine Species). Una treintena de estas especies se han observado en aguas catalanas.

Clasificación
La taxonomia actual de los Cephalaspidea, propuesta por Bouchet et al. (2017) y publicada en WoRMS, es:

Cephalaspidea P. Fischer, 1883

  • Bulloidea Gray, 1827
    • Bullidae J. E. Gray, 1827
      • Bulla Linnaeus, 1758
    • Retusidae Thiele, 1925
      • Pyrunculus Pilsbry, 1895
      • Relichna Rudman, 1971
      • Retusa T. Brown, 1827
      • Spissitydeus Iredale, 1936
    • Rhizoridae Dell, 1952
      • Rhizorus Montfort, 1810
      • Volvulella R. B. Newton, 1891
    • Tornatinidae P. Fischer, 1883
      • Acteocina J. E. Gray, 1847
  • Cylichnoidea H. Adams & A. Adams, 1854
    • Colinatydidae Oskars, Bouchet & Malaquias, 2015
      • Colinatys Ortea, Moro & Espinosa, 2013
    • Cylichnidae H. Adams & A. Adams, 1854
      • Bogasonia Warén, 1989
      • Cylichna Lovén, 1846
      • Cylichnella Gabb, 1873
      • Decorifer Iredale, 1937
      • Mamillocylichna F. Nordsieck, 1972
      • Paracteocina Minichev, 1966
      • Semiretusa Thiele, 1925
      • Sphaerocylichna Thiele, 1925
      • Toledonia Dall, 1902
      • Truncacteocina Kuroda & T. Habe, 1955
    • Diaphanidae Odhner, 1914 (1857)
      • Diaphaninae Odhner, 1914 (1857)
        • Diaphana T. Brown, 1827
    • Eoscaphandridae Chaban & Kijashko, 2016
      • Cylichnoides Minichev, 1977
      • Eoscaphander T. Habe, 1952
      • Hamineobulla Habe, 1950
      • Pseudocylichna Chaban & Kijashko, 2016
    • Mnestiidae Oskars, Bouchet & Malaquias, 2015
      • Mnestia H. Adams & A. Adams, 1854
  • Haminoeoidea Pilsbry, 1895
    • Haminoeidae Pilsbry, 1895
      • Aliculastrum Pilsbry, 1896
      • Atys Montfort, 1810
      • Bakawan Oskars & Malaquias, 2019
      • Bullacta Bergh, 1901
      • Confettiella Oskars, 2024
      • Cylichnatys Habe, 1952
      • Diniatys Iredale, 1936
      • Haloa Pilsbry, 1921
      • Haminella Thiele, 1925
      • Haminoea W. Turton & Kingston, 1830
      • Lamprohaminoea T. Habe, 1952
      • Liloa Pilsbry, 1921
      • Papawera Oskars & Malaquias, 2019
      • Phanerophthalmus A. Adams, 1850
      • Roxaniella Monterosato, 1884
      • Smaragdinella A. Adams, 1848
      • Vellicolla Oskars, Too, D. Rees, P. M. Mikkelsen, Willassen & Malaquias, 2019
      • Weinkauffia Weinkauff, 1873
  • Newnesioidea Moles, Wägele, Schrödl & C. Ávila, 2017
    • Newnesiidae Moles, Wägele, Schrödl & C. Ávila, 2017
      • Hocius Moles, C. Ávila & Malaquias, 2017
      • Newnesia E. A. Smith, 1902
  • Philinoidea J. E. Gray, 1850 (1815)
    • Aglajidae Pilsbry, 1895 (1847)
      • Aglaja Renier, 1807
      • Aglaona Chaban, Ekimova, Schepetov & Chernyshev, 2022
      • Biuve Zamora-Silva & Malaquias, 2017
      • Camachoaglaja Zamora-Silva & Malaquias, 2017
      • Chelidonura A. Adams, 1850
      • Mannesia Zamora-Silva & Malaquias, 2017
      • Mariaglaja Zamora-Silva & Malaquias, 2017
      • Melanochlamys Cheeseman, 1881
      • Nakamigawaia Kuroda & T. Habe, 1961
      • Navanax Pilsbry, 1895
      • Niparaya Zamora-Silva & Malaquias, 2017
      • Odontoglaja Rudman, 1978
      • Philinissima Chaban, Ekimova & Schepetov, 2023
      • Philinopsis Pease, 1860
      • Spinoaglaja Ortea, Moro & Espinosa, 2007
      • Tubulophilinopsis Zamora-Silva & Malaquias, 2017
    • Alacuppidae Oskars, Bouchet & Malaquias, 2015
      • Alacuppa Oskars, Bouchet & Malaquias, 2015
      • Mimatys Habe, 1952
      • Roxania Leach, 1847
    • Antarctophilinidae Moles, C. Ávila & Malaquias, 2019
      • Antarctophiline Chaban, 2016
      • Waegelea Moles, C. Ávila & Malaquias, 2019
    • Colpodaspididae Oskars, Bouchet & Malaquias, 2015
      • Colobocephalus M. Sars, 1870
      • Colpodaspis M. Sars, 1870
    • Gastropteridae Swainson, 1840
      • Africanopteron Ortea & Moro, 2020
      • Enotepteron Minichev, 1967
      • Gastropteron Kosse, 1813
      • Sagaminopteron Tokioka & Baba, 1964
      • Siphopteron Gosliner, 1989
    • Laonidae Pruvot-Fol, 1954
      • Laona A. Adams, 1865
      • Retusophiline F. Nordsieck, 1972
    • Philinidae J. E. Gray, 1850 (1815)
      • Hermaniinae Chaban, Ekimova, Schepetov, Kohnert, Schrödl & Chernyshev, 2019
        • Hermania Monterosato, 1884
        • Spiniphiline Gosliner, 1988
      • Philininae J. E. Gray, 1850 (1815)
        • Globophiline Habe, 1958
        • Philine Ascanius, 1772
        • Praephiline Chaban & Soldatenko, 2009
        • Spiraphiline Moles, C. Ávila & Malaquias, 2019
        • Yokoyamaia T. Habe, 1950
    • Philinoglossidae Hertling, 1932
      • Abavopsis Salvini-Plawen, 1973
      • Philinoglossa Hertling, 1932
      • Pluscula Er. Marcus, 1953
      • Sapha Er. Marcus, 1959
    • Philinorbidae Oskars, Bouchet & Malaquias, 2015
      • Philinorbis Habe, 1950
      • Pseudophiline T. Habe, 1976
    • Scaphandridae G. O. Sars, 1878
      • Kaitoa Marwick, 1931
      • Nipponoscaphander Kuroda & Habe, 1971
      • Sabatia Bellardi, 1877
      • Scaphander Montfort, 1810

Superfamilias de cefalaspideos citadas en el Mediterráneo o en la Península Ibérica:

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.