The Flabellinidae, a heterogeneous assembly of supposedly plesiomorphic to very derived sea slug groups, have not yet been addressed by integrative studies. Here novel material of rarely seen Arctic taxa as well as North Atlantic, North and South Pacific, and tropical Indo-West Pacific flabellinid species is investigated morpho-anatomically and with multi-locus markers (partial COI, 16S rDNA, 28S rDNA and H3) which were generated and analysed in a comprehensive aeolid taxon sampling. It was found that the current fam- RESEARCH ARTICLE Launched to accelerate biodiversity research A peer-reviewed open-access journalTatiana Korshunova et al. / ZooKeys 717: 1-139 (2017) 2 ily Flabellinidae is polyphyletic and its phylogeny and taxonomic patterns cannot be understood without considering members from all the Aeolidacean families and, based on a robust phylogenetic hypothesis, morpho-anatomical evolution of aeolids is more complex than suspected in earlier works and requires reclassification of the taxon. Morphological diversity of Flabellinidae is corroborated by molecular divergence rates and supports establishing three new families (Apataidae fam. n., Flabellinopsidae fam. n., Samlidae fam. n.), 16 new genera, 13 new species, and two new subspecies among the former Flabellinidae. Two families, namely Coryphellidae and Paracoryphellidae, are restored and traditional Flabellinidae is considerably restricted. The distinctness of the recently described family Unidentiidae is confirmed by both morphological and molecular data. Several species complexes among all ex-"Flabellinidae" lineages are recognised using both morphological and molecular data. The present study shows that Facelinidae and Aeolidiidae, together with traditional "Tergipedidae", deeply divide traditional "Flabellinidae." Diagnoses for all aeolidacean families are therefore provided and additionally two new non-flabellinid families (Abronicidae fam. n. and Murmaniidae fam. n.) within traditional tergipedids are established to accommodate molecular and morphological disparity. To address relationships and disparity, we propose a new family system for aeolids. Here the aeolidacean species are classified into at least 102 genera and 24 families. Operational rules for integration of morphological and molecular data for taxonomy are suggested.

‘Cryptic’ species are an emerging biological problem that is broadly discussed in the present study. Recently, a cryptic species definition was suggested for those species which manifest low morphological, but considerable genetic, disparity. As a case study we present unique material from a charismatic group of nudibranch molluscs of the genus Trinchesia from European waters to reveal three new species and demonstrate that they show a dual nature: on one hand, they can be considered a ‘cryptic’ species complex due to their overall similarity, but on the other hand, stable morphological differences as well as molecular differences are demonstrated for every species in that complex. Thus, this species complex can equally be named ‘cryptic’, ‘pseudocryptic’ or ‘non-cryptic’. We also present evidence for an extremely rapid speciation rate in this species complex and link the species problem with epigenetics. Available metazoan-wide data, which are broadly discussed in the present study, show the unsuitability of a ‘cryptic’ species concept because the degree of crypticity represents a continuum when a finer multilevel morphological and molecular scale is applied to uncover more narrowly defined species making the ‘cryptic’ addition to ‘species’ redundant. Morphological and molecular methods should be applied in concordance to form a fine-scale multilevel taxonomic framework, and not necessarily implying only an a posteriori transformation of exclusively molecular-based ‘cryptic’ species into morphologically-defined ‘pseudocryptic’ ones. Implications of the present study have importance for many fields, including conservation biology and fine-scale biodiversity assessments.

Dorids are one of the largest nudibranch groups, encompassing more than 2000 species. One of the crucial problems with tracing the evolution of dorids is the relationship between cryptobranch dorids (gill cavity present) and phanerobranch dorids (gill cavity absent). Integrative morphological and molecular studies of the enigmatic Japanese dorid species of the cryptobranch genus Cadlina, C. japonica and ‘C.’ sagamiensis, which were collected by the Emperor of Japan (Shōwa era), are presented here for the first time. It is shown that while C. japonica does belong to the Cadlina clade, another Japanese species ‘C.’ sagamiensis is not part of the cryptobranch dorids of the family Cadlinidae, but is related to both the phanerobranch dorid family, Hexabranchidae, and to the cryptobranch family Cadlinellidae stat. nov. A new genus, Showajidaia gen. nov., and new family, Showajidaiidae fam. nov., are proposed for ‘C.’ sagamiensis, and four new species of the genus Cadlina are described based on a dorid-wide molecular phylogenetic analysis, which is the first substantial update of the dorid family system since 2010. Integration of phylogenetic data with an ontogenetic model of dorid evolution suggests that cryptobranch organization can be most reliably assessed as the ancestral state for the majority of dorids.

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