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Korshunova, T.; Fletcher, K.; Martynov, A. (2025). The endless forms are the most differentiated — how taxonomic pseudo-optimization masked natural diversity and evolution: the nudibranch case. Zoological Journal of the Linnean Society. 204(4): zlaf057: 1-93.
514054
10.1093/zoolinnean/zlaf057 [view]
Korshunova, T.; Fletcher, K.; Martynov, A.
2025
The endless forms are the most differentiated � how taxonomic pseudo-optimization masked natural diversity and evolution: the nudibranch case
Zoological Journal of the Linnean Society
204(4): zlaf057: 1-93
Publication
Stated date: 18 August 2025
Available for editors  PDF available
In this study, a global-encompassing, fine-scale differentiated taxonomy is consistently presented from the high level of a major nudibranch suborder, Aeolidacea, and then consequentially to lower levels of superfamilies, families, genera, and species to conjoin the underlying evolutionary pathways of aeolidacean nudibranchs and systematic representation to a maximal degree. The suborder Aeolidacea is reinstated and the superfamily system of Aeolidacea is reformed; 10 aeolidacean superfamilies are recognized encompassing 29 families in an all-family Synopsis, and the superfamily Embletonioidea is included in addition. Two new families, Chudidae fam. nov. and Hantazuidae fam. nov., are described, which show unique morphological and molecular patterns. Seven new species are described within various superfamilies, and five species are reinstated. The genus-level taxonomy of the family Coryphellidae is revisited, 11 genera are recognized, including nine reinstated and two new genera. We show that recent synonymization of all valid Coryphellidae genera into just a single genus was based on an erroneous phylogenetic analysis, the careless treatment of morphological characters, and established preconceptions. Uniting fine-scale morphological and molecular data, order level nudibranch taxonomy is refined, order Nudibranchia is restricted, and order Doridida is reinstated. Within the order Nudibranchia, five suborders are recognized: Arminacea, restricted, reinstated; Tritoniacea, reinstated; Dendronotacea, restricted, reinstated; Janolacea, reinstated; and Aeolidacea, revised, reinstated.
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Date
action
by
2025-08-20 17:20:27Z
created
Picton, Bernard
2025-08-21 11:30:48Z
changed
Bouchet, Philippe
2025-09-03 08:51:45Z
changed
Picton, Bernard
2025-11-13 14:52:55Z
changed
Gofas, Serge

Aeolidina accepted as Aeolidioidea J. E. Gray, 1827 (status source)
Amphorina Quatrefages, 1844 (status source)
Amphorina amazighi (Tamsouri, Carmona, Moukrim & Cervera, 2015) (new combination reference)
Amphorina andra Korshunova, Malmberg, Prkic, Petani, Fletcher, Lundin & Martynov, 2020 (status source)
Amphorina farrani (Alder & Hancock, 1844) (additional source)
Amphorina linensis (García-Gómez, Cervera & F. J. Garcia, 1990) (additional source)
Amphorina pallida (Alder & Hancock, 1842) (additional source)
Amphorina viriola Korshunova, Malmberg, Prkic, Petani, Fletcher, Lundin & Martynov, 2020 (status source)
Arminina accepted as Arminoidea Iredale & O'Donoghue, 1923 (1841) (status source)
Borealea Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Coryphella J. E. Gray, 1850 (status source)
Borealea nobilis (A. E. Verrill, 1880) accepted as Coryphella nobilis A. E. Verrill, 1880 (status source)
Calmidae Iredale & O'Donoghue, 1923 accepted as Flabellinidae Bergh, 1889 (additional source)
Capellinia Trinchese, 1873 (status source)
Capellinia vittata (Alder & Hancock, 1842) (new combination reference)
Catriona aurantia (Alder & Hancock, 1842) (status source)
Coryphella borealis Odhner, 1922 (status source)
Coryphella browni Picton, 1980 (status source)
Coryphella chriskaugei (Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017) (status source)
Coryphella insolita (García-Gómez & Cervera, 1989) (additional source)
Coryphella lineata (Lovén, 1846) (status source)
Coryphella monicae (Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017) (additional source)
Coryphella nobilis A. E. Verrill, 1880 (additional source)
Coryphella orjani (Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017) (additional source)
Cuthona Alder & Hancock, 1855 (additional source)
Cuthonellidae M. C. Miller, 1977 (additional source)
Cuthonidae Odhner, 1934 (additional source)
Dendronotina accepted as Dendronotoidea Allman, 1845 (status source)
Diaphoreolis stipata (Alder & Hancock, 1843) (status source)
Doridida accepted as Doridina (status source)
Embletonioidea Pruvot-Fol, 1954 (basis of record)
Eubranchidae Odhner, 1934 (additional source)
Eubranchulus olivaceus (O'Donoghue, 1922) (status source)
Eubranchulus rupium (Møller, 1842) (status source)
Eubranchus Forbes, 1838 (additional source)
Favorinidae Bergh, 1889 accepted as Facelinidae Bergh, 1889 (additional source)
Fionoidea J. E. Gray, 1857 (basis of record)
Fjordia Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Coryphella J. E. Gray, 1850 (status source)
Fjordia browni (Picton, 1980) accepted as Coryphella browni Picton, 1980 (status source)
Fjordia chriskaugei Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Coryphella chriskaugei (Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017) (status source)
Fjordia insolita (García-Gómez & Cervera, 1989) accepted as Coryphella insolita (García-Gómez & Cervera, 1989) (status source)
Fjordia lineata (Lovén, 1846) accepted as Coryphella lineata (Lovén, 1846) (additional source)
Goniaeolididae Odhner, 1907 (status source)
Gulenia borealis (Odhner, 1922) accepted as Coryphella borealis Odhner, 1922 (status source)
Gulenia monicae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Coryphella monicae (Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017) (additional source)
Gulenia orjani Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Coryphella orjani (Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017) (additional source)
Janolina accepted as Proctonotoidea J. E. Gray, 1853 (status source)
Microchlamylla Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Coryphella J. E. Gray, 1850 (status source)
Microchlamylla gracilis (Alder & Hancock, 1844) accepted as Coryphella gracilis (Alder & Hancock, 1844) (status source)
Murmaniidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 (additional source)
Nudibranchus Martynov, 1998 (status source)
Nudibranchus exiguus (Alder & Hancock, 1848) (basis of record)
Phetia Korshunova, Fletcher & Martynov, 2025 accepted as Piseinotecus Er. Marcus, 1955 (original description)
Phetia soussi (Tamsouri, Carmona, Moukrim & Cervera, 2014) accepted as Piseinotecus soussi Tamsouri, Carmona, Moukrim & Cervera, 2014 (new combination reference)
Samlidae Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 (additional source)
Samloidea Korshunova, Martynov, Bakken, Evertsen, Fletcher, Mudianta, H. Saito, Lundin, Schrödl & Picton, 2017 accepted as Fionoidea J. E. Gray, 1857 (basis of record)
Trinchesia Ihering, 1879 (status source)
Trinchesia caerulea (Montagu, 1804) (status source)
Trinchesia morrowae Korshunova, Picton, Furfaro, Mariottini, Pontes, Prkic, Fletcher, Malmberg, Lundin & Martynov, 2019 (status source)
Tritonina accepted as Tritonioidea Lamarck, 1809 (status source)
Unidentiidae Millen & Hermosillo, 2012 accepted as Piseinotecidae Edmunds, 1970 (additional source)
Unidentioidea Millen & Hermosillo, 2012 accepted as Fionoidea J. E. Gray, 1857 (basis of record)
Xenocratenidae Martynov, Lundin, Picton, Fletcher, Malmberg & Korshunova, 2020 (additional source)