Tatiana Korshunova

Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda)

Abstract 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 family 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.

External diversity is restrained by internal conservatism: New nudibranch mollusc contributes to the cryptic species problem

Cryptic species are an important concept of modern biodiversity studies and a potential basis for immensely increasing the amount of biological diversity worldwide; however, the conceptual grounds of cryptic species still need to be refined. Here, using a remarkable example from marine invertebrates, the case of a species complex of common large‐sized nudibranchs of the genus Dendronotus in European waters is presented, which is relevant for the ongoing cryptic species discussion. Based on extensive sampling in the seas of Northern Hemisphere from United Kingdom to the Sea of Japan, a morphological and molecular framework is constructed for understanding the diversity of European Dendronotus nudibranchs. The present study discovered notable multilevel character diversity within the common Dendronotus “frondosus” species complex. This implies the highest degree of variation of external characters with parallel colour patterns among different species and significantly constrained conservatism of internal characters. The molecular data are congruent with the stable differences in the internal features and confirm the presence of three distinct species in this complex. While two species were already known, the third species is evidently new, based on morphological and molecular evidence and is described here as Dendronotus europaeus sp. n. In the light of multilevel character diversity, the general problem of plausibility of the distinction and practical usage of the terms “cryptic” and “pseudocryptic” species is discussed.

Two new species and a remarkable record of the genus Dendronotus from the North Pacific and Arctic oceans (Nudibranchia)

Abstract Two new species of the nudibranch genus Dendronotus, Dendronotus arcticus sp. n. and Dendronotus robilliardi sp. n., are described from the Arctic and North Pacific oceans respectively, based on morphological and molecular data, and the North Pacific Dendronotus albus is revealed to be a species complex. The species Dendronotus robilliardi sp. n. is described from the northwestern Pacific (Kamchatka) differing from the northeastern Pacific Dendronotus albus by molecular and morphological data. The synonymy of Dendronotus diversicolor with Dendronotus albus is confirmed by analysis of their original descriptions. An endemic Arctic species Dendronotus arcticus sp. n. is also described here, differing substantially from all species of the genus Dendronotus using morphological and molecular data. An unusual record of the recently described Dendronotus kamchaticus Ekimova, Korshunova, Schepetov, Neretina, Sanamyan, Martynov, 2015 is also presented, the first from the northeastern Pacific, geographically separated from the type locality of this species in the northwestern Pacific by a distance ca. 6000 km; molecular data show them to belong to the same species.

Zachsia zenkewitschi (Teredinidae), a Rare and Unusual Seagrass Boring Bivalve Revisited and Redescribed

The sea-grass borer Zachsia zenkewitschi belongs to a group of economically and ecologically important bivalves, commonly referred to as shipworms. The sole recognized representative of the genus Zachsia, this species displays an unusual life history and reproductive strategy that is now understood to include: environmental sex determination of free swimming larvae, extreme sexual and size dimorphism between males and females, internal fertilization, maintenance of often large harems of male dwarfs within a specialized cavity of the female mantle, and complex maternal care of larvae in specialized brood pouches within the gill. It is also the only shipworm species known to burrow in sea grass rhizomes rather than terrestrial wood. Although Z. zenkewitschi is rare and little studied, understanding of its biology and anatomy has evolved substantially, rendering some aspects of its original description inaccurate. Moreover, no existing type specimens are known for this species. In light of these facts, we designate a neotype from among specimens recently collected at the type location, and undertake a re-description of this species, accounting for recent reinterpretation of its life history and functional anatomy.

Unique pleuroproctic taxa of the nudibranch family Aeolidiidae from the Atlantic and Pacific Oceans, with description of a new genus and species

Morphological and molecular data on a deep-sea pleuroproctic taxon of the family Aeolidiidae from the Barents Sea (subarctic North Atlantic) are presented for the first time. Zeusia n. gen. is established for the North Atlantic and North Pacific pleuroproctic species of the family Aeolidiidae with two species: Z. hyperborea n. sp. and Z. herculea (Bergh, 1894) n. comb. Zeusia was recovered as sister to the genus Aeolidia in molecular analyses. The two included species of Zeusia share the potential synapomorphies of very large seminal receptacle and long vas deferens. A pleuroproctic anal opening is an important basal character of the group Nudibranchia and the family Aeolidiidae is discussed in the light of the present discovery.

Morphological and molecular evidence indicate Dendronotus primorjensis is a valid species that has priority over D. dudkai (Nudibranchia)

Abstract Morphological and molecular data of type material of the nudibranch mollusc Dendronotus primorjensis Martynov, Sanamyan, Korshunova, 2015 from the Sea of Japan are summarised and compared with those of Dendronotus dudkai Ekimova, Schepetov, Chichvarkhina, Chichvarkhin, 2016. The clear conclusion is that the latter is a junior synonym of Dendronotus primorjensis.

First true brackish-water nudibranch mollusc provides new insights for phylogeny and biogeography and reveals paedomorphosis-driven evolution

A unique example of brackish water fjord-related diversification of a new nudibranch genus and species Bohuslania matsmichaeli gen. n., sp. n. is presented. There are only few previously known brackish-water opisthobranchs and B. matsmichaeli gen. n., sp. n. is the first ever described brackish-water nudibranch with such an extremely limited known geographical range and apparently strict adherence to salinity levels lower than 20 per mille. Up to date the new taxon has been found only in a very restricted area in the Idefjord, bordering Sweden and Norway, but not in any other apparently suitable localities along the Swedish and Norwegian coasts. We also show in this study for the first time the molecular phylogenetic sister relationship between the newly discovered genus Bohuslania and the genus Cuthona. This supports the validity of the family Cuthonidae, which was re-established recently. Furthermore, it contributes to the understanding of the evolutionary patterns and classification of the whole group Nudibranchia. Molecular and morphological data indicate that brackish water speciation was triggered by paedomorphic evolution among aeolidacean nudibranchs at least two times independently. Thus, the present discovery of this new nudibranch genus contributes to several biological fields, including integration of molecular and morphological data as well as phylogenetic and biogeographical patterns.

Corrigenda: Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda). https://doi.org/10.3897/zookeys.717.21885

[This corrects the article DOI: 10.3897/zookeys.717.21885.].