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Mesoplodon eueu sp. nov. Holotype NMNZ MM003000, a pregnant, 5.06 m long adult female named Nihongore by Te Rūnanga o Makaawhio. Collected by Ramari Stewart, Nathaniel Scott and Don Neale after beachcast on 27 November 2011. The complete skeletons of the female and fetus are held by Museum of New Zealand Te Papa Tongarewa (NMNZ, Wellington, Aotearoa New Zealand; specimen MM003000), and a tissue sample is held in the New Zealand Cetacean Tissue Archive (NZCeTA, Universityof Auckland, Auckland, Aotearoa New Zealand; all institutional abbreviations in electronic supplementary material, S1). Type locality Waiatoto Spit, South Westland, Aotearoa New Zealand. Paratypes Adult females (PEM N0136 and PEM N3438) and adult male (PEM N1114) held at Port Elizabeth Museum (Gqeberha, South Africa) and adult males (SAM-ZM-041596 and SAM- ZM-039840) held at Iziko South African Museum (Cape Town, South Africa). Full description of paratypes is found in electronic supplementary material S2. Etymology The scientific and common names acknowledge links with Indigenous communities in South Africa and Aotearoa New Zealand, respectively, and were chosen in consultation with these peoples. Most of the South African strandings come from territory inhabited by the Khoisan peoples. Guided by the Khoisan Council, we chose the name //eu//’eu (simplified to eueu to fit nomenclature standards; correct pronunciation available in associated audio clip a 1 in the electronic supplementary material), which means ‘big fish’ in Khwedam (from the Khoe language family). In Aotearoa New Zealand, Māori cultural expert Brad Haami developed a shortlist of potential names meaningful in the Māori language, whichwasthen sent for comment to Te Rūnanga o Ngāi Tahu. The selected common name, Ramari’ s beaked whale, pays homage to Māoritohunga (expert) Ramari Stewart, who has kept traditional knowledge alive, contributed extensively to scientific research on marine mammals, and helped prepare the skeleton of the holotype. The word ‘Ramari’ means a rare event in the Māori language, reflecting the elusive nature of most beaked whales. Diagnosis Molecular characteristics M. eueu differs from M. mirus based on nuclear DNA markers, and from M. mirus and its closest relatives M. europaeus, M. ginkgodens and M. bidens using mtDNA markers (figure 2). M. mirus is distinct from all other mesoplodont species based on previous mitochondrial and nuclear DNA trees [7,20,24]. Mitochondrial DNA: analysis of mitochondrial data includes sequences from the holotypes of both M. mirus and M. eueu at all sequence lengths. Over the 304 bp mitochondrial control region segment, M. eueu is distinguished by seven fixed differences from M. mirus, with F ST = 0.85 (p dA = 0.04 between the two species. At the full mitochondrial genome lengths, M. eueu is distinguished by 579 fixed differences from M. mirus with F ST = 0.96 (p dA = 0.04 (electronic supplementary material, table S2). Nuclear DNA: reduced representation sequencing with ddRAD showed M. eueu had a distinct admixture pattern to M. mirus (figure 2), and an F ST = 0.64 (p M. mirus and M. eueu weredistinguished by 1909 fixed differences (12%, per SNP allele error rate = 0.002), across a dataset of 15671 SNPs found between or within both species. Comparison of one whole nuclear genome each from M. mirus and M. eueu showed a level of nucleotide divergence of 0.28%. Morphological characters M. eueu is a larger (5.3 m) species of Mesoplodon differing from all other members of the genus except M. mirus, M. hectori and M. perrini in having tusks positioned at the tip of the mandible. It also differs from M. hectori and M. perrini in having smaller, less triangular tusks and from M. mirus in having a relatively shorter rostrum with a wider base, a shorter mandibular symphysis, wider premaxillary sac fossae and crests, and a taller cranium. External appearance The external appearances of M. eueu and M. mirus are not known to be consistently distinguishable. Both species are rotund mesoplodonts with bodies that taper towards the tail and rostrum, with somewhat bulbous and well-defined melons, a mostly straight beak and short, straight gape. Colouration is generally grey with a dark eye patch in both species, but there may be specific colouration patterns linked to M. eueu; for example, a female that stranded in South Africa showed a whitish dorsal colouration from the fin to caudal peduncle [14]. However, Aguilar et al. [6] reported M. mirus with varying dorsal and ventral white colouration in the Canary and Azores Islands, so colouration patterns linked to species cannot yet be conclusively defined. Distribution M. eueu probably occurs throughout temperate SH waters, with reports from several locations. Genetic methods have confirmed at least some of these records in South Africa ([20], this study), Australia [20] and Aotearoa New Zealand [11]. Nomenclatural acts This published work and the nomenclatural acts it contains are registered in ZooBank (http://zoobank.org/), the online registration system for the International Commission on Zoological Nomenclature (ICZN). The ZooBank Life Science Identifier is urn:lsid:zoobank.org:pub: C61A1A33-B234-476E- AA24-86C6ED130C10, and for the new species is urn:lsid:zoo bank.org:act: C56121C4-1E15-4A07-A270-D92AF43AE74A., Published as part of Carroll, Emma L., McGowen, Michael R., McCarthy, Morgan L., Marx, Felix G., Aguilar, Natacha, Dalebout, Merel L., Dreyer, Sascha, Gaggiotti, Oscar E., Hansen, Sabine S., Helden, Anton van, Onoufriou, Aubrie B., Baird, Robin W., Baker, C. Scott, Berrow, Simon, Cholewiak, Danielle, Claridge, Diane, Constantine, Rochelle, Davison, Nicholas J., Eira, Catarina, Fordyce, R. Ewan, Gatesy, John, Greg Hofmeyr, G. J., Martín, Vidal, Mead, James G., Mignucci-Giannoni, Antonio A., Morin, Phillip A., Reyes, Cristel, Rogan, Emer, Rosso, Massimiliano, Silva, Mónica A., Springer, Mark S., Steel, Debbie & Olsen, Morten Tange, 2021, Speciation in the deep: genomics and morphology reveal a new species of beaked whale Mesoplodon eueu., pp. 1-10 in Proceedings of the Royal Society B 288 (1961) on pages 5-6, DOI: 10.1098/rspb.2021.1213

This work was supported by ONR grants N000141613017 to E.L.C. and N.A. and N00014-18-1-2808 to C.S.B.; funds from the NMNH Rebecca G. Mead and James G. Mead Marine Mammal Endowment, NSF (USA) grant no. DEB-1457735 to M.S.S., P.A.M. and J.G.; Brothers Hartmann Foundation grant no. AB28148 to M.T.O.; NMFS, BOEM, and USA Navy funding to D.Ch. under the Atlantic Marine Assessment Program for Protected Species. M.L.M. was funded under the Marie Skłodowska-Curie grant agreement no 801199; E.L.C. by a Rutherford Discovery Fellowship from the Royal Society of New Zealand Te Apārangi. Irish Whale and Dolphin Group Cetacean Stranding scheme is part-funded by the National Parks and Wildlife Service. The deep sea has been described as the last major ecological frontier, as much of its biodiversity is yet to be discovered and described. Beaked whales (ziphiids) are among the most visible inhabitants of the deep sea, due to their large size and worldwide distribution, and their taxonomic diversity and much about their natural history remain poorly understood. We combine genomic and morphometric analyses to reveal a new Southern Hemisphere ziphiid species, Ramari's beaked whale, Mesoplodon eueu, whose name is linked to the Indigenous peoples of the lands from which the species holotype and paratypes were recovered. Mitogenome and ddRAD-derived phylogenies demonstrate reciprocally monophyletic divergence between M. eueu and True's beaked whale (M. mirus) from the North Atlantic, with which it was previously subsumed. Morphometric analyses of skulls also distinguish the two species. A time-calibrated mitogenome phylogeny and analysis of two nuclear genomes indicate divergence began circa 2 million years ago (Ma), with geneflow ceasing 0.35–0.55 Ma. This is an example of how deep sea biodiversity can be unravelled through increasing international collaboration and genome sequencing of archival specimens. Our consultation and involvement with Indigenous peoples offers a model for broadening the cultural scope of the scientific naming process. Publisher PDF