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Nudibranchia

Nudibranchia és un grup divers de llimacs marins que tradicionalment comprèn dos grans llinatges evolutius, Doridina i Cladobranchia. Els nudibranquis es caracteritzen per l’absència de closca en l’estat adult i per una gran diversitat d’estructures externes especialitzades relacionades amb la respiració, l’alimentació, la defensa i les funcions sensorials.

Felimare tricolor per Enric Madrenas

La història taxonòmica dels nudibranquis (Nudibranchia Cuvier, 1817) il·lustra la transició des de classificacions basades principalment en la morfologia i l’anatomia cap a enfocaments filogenètics que incorporen evidències anatòmiques, moleculars i del desenvolupament. Tradicionalment, els nudibranquis s’incloïen entre els «opistobranquis», una agrupació informal de gasteròpodes marins caracteritzada per la freqüent reducció o pèrdua de la closca i per diverses estructures respiratòries especialitzades. Aquest concepte tradicional va persistir durant bona part del segle XX, quan els estudis anatòmics comparatius van demostrar progressivament la complexitat de les relacions entre els grups inclosos dins d’Opisthobranchia i van qüestionar-ne la consideració com a grup evolutiu natural.

Wägele i Willan (2000), a partir d’una àmplia anàlisi de caràcters morfològics i anatòmics, van proporcionar un marc filogenètic influent per a Nudibranchia. La seva classificació reconeixia dues grans divisions, Anthobranchia, corresponent en termes generals als nudibranquis doridacis, i Cladobranchia, que comprenia els grups tradicionalment reconeguts com a aeòlids, arminacis i dendronotacis. Aquest marc va influir posteriorment en diverses classificacions dels nudibranquis i dels gasteròpodes heterobranquis.

El desenvolupament dels mètodes moleculars i transcriptòmics durant el segle XXI va proporcionar noves evidències sobre les relacions entre els diferents llinatges de nudibranquis i va conduir a nombrosos canvis en la seva classificació. Els estudis moleculars van demostrar que algunes agrupacions tradicionals no es corresponien amb llinatges evolutius i van aportar evidències de relacions que no sempre podien inferir-se únicament a partir de la morfologia externa. En conseqüència, la classificació de Nudibranchia ha continuat revisant-se a mesura que s’han incorporat noves evidències filogenètiques.

Korshunova et al. (2025) van proposar una important reorganització de la classificació superior dels nudibranquis. En el seu tractament, el nom Nudibranchia es restringia al llinatge corresponent essencialment a Cladobranchia, mentre que els nudibranquis doridacis eren separats i reconeguts com l’ordre Doridida. Els autors també van reorganitzar substancialment la classificació superior dels llinatges cladobranquis i van recuperar diversos subordres tradicionals.

Aquesta interpretació va ser posteriorment reconsiderada per Moles et al. (2026) en la seva anàlisi més àmplia de l’estabilitat taxonòmica dels heterobranquis marins. Els autors van argumentar que les decisions taxonòmiques han de tenir en compte no només les relacions filogenètiques, sinó també la capacitat diagnòstica i l’estabilitat dels tàxons, la continuïtat de la nomenclatura establerta i les conseqüències dels canvis nomenclaturals de gran abast. En el cas dels nudibranquis, van donar suport al manteniment del nom establert Nudibranchia en el seu sentit ampli, en lloc de restringir-lo exclusivament a Cladobranchia.

El marc adoptat aquí segueix, per tant, el concepte ampli de Nudibranchia avalat per Moles et al. (2026). Segons aquesta interpretació, l’ordre manté el seu àmbit tradicional i inclou Doridina i Cladobranchia com els seus dos principals llinatges evolutius.

Els nudibranquis presenten una extraordinària diversitat de formes corporals i estructures especialitzades. Els llinatges d’aeòlids i altres cladobranquis posseeixen habitualment cerata o altres apèndixs dorsals, mentre que les formes dendronotàcies i tritoníides poden desenvolupar elaborats processos ramificats o dendrítics. Els nudibranquis doridacis presenten generalment un cos més compacte i una disposició posterior de les brànquies secundàries. Aquests diferents plans corporals estan associats amb diverses estratègies alimentàries, mecanismes defensius, adaptacions sensorials i relacions ecològiques.

Nudibranchia ocupa una àmplia varietat d’ambients marins, des d’hàbitats costaners soms i intermareals fins a regions més profundes de la plataforma i el talús continental. Moltes espècies estan estretament associades amb determinats organismes presa, entre els quals hi ha cnidaris, esponges, briozous i altres invertebrats sèssils. Aquestes associacions tròfiques es reflecteixen sovint en la seva morfologia, comportament i estratègies defensives.

La diversitat dels nudibranquis també es reflecteix en la seva classificació. Les relacions entre diversos dels seus principals llinatges han estat revisades repetidament a mesura que s’han acumulat noves evidències anatòmiques, del desenvolupament i moleculars. Per això, la classificació superior del grup s’ha de considerar un marc que pot continuar modificant-se a mesura que es disposi de noves evidències filogenètiques.

La sinopsi sistemàtica de Nudibranchia adoptada aquí segueix el concepte ampli avalat per Moles et al. (2026), segons el qual l’ordre inclou els dos grans llinatges Doridina i Cladobranchia.

Grups principals de nudibranquis:

Bibliografia

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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.