What is really out there? Review of the genus Okenia Menke, 1830 (Nudibranchia: Goniodorididae) in the Mediterranean Sea with description of two new species

The precise number of Okenia taxa inhabiting the Mediterranean Sea, as well as their general taxonomy, varies according to different specialists. So far, eight valid species have been reported from the area: Okenia aspersa (Alder & Hancock, 1845), Okenia cupella (Vogel & Schultz, 1970), Okenia elegans (Leuckart, 1828), Okenia hispanica Valdes & Ortea, 1995, Okenia impexa Er. Marcus, 1957, Okenia leachii (Alder & Hancock, 1854), Okenia mediterranea (Ihering, 1886), and Okenia zoobotryon (Smallwood, 1910). Of these, only three (O. elegans, O. hispanica, and O. mediterranea) have their type localities in the Mediterranean Sea, whereas the others were described from different biogeographic areas and later included in the Mediterranean biota. We carried out a review on Mediterranean Okenia species through an integrative approach, based on a wide literature search and a morphological and molecular analysis of available type material and samples collected recently. The present study confirmed the presence of O. aspersa, O. elegans, O. hispanica, and O. mediterranea in the Mediterranean Sea, although leaving remaining questions about some of those taxa. The distribution of O. cupella, O. impexa, and O. zoobotryon is limited to the western Atlantic, and of O. leachii to the eastern Atlantic. All specimens previously identified as O. cupella, O. impexa, and O. zoobotryon by different authors in the Mediterranean Sea were repeatedly misidentified. Thus, we describe Okenia problematica sp. nov. and Okenia longiductis sp. nov., from the Mediterranean Okenia cupella/impexa and O. zoobotryon. We also consider here Okenia pusilla Sordi, 1974 a nomen dubium and include a redescription of the holotype of O. cupella. A molecular phylogeny, including all the sequenced Okenia species, was performed in order to evaluate the evolutionary relationships of the newly described species with the other congeneric taxa.