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Morphological and Molecular Description of a New Genus And Zootaxa 4821 (3): 553–569 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4821.3.7 http://zoobank.org/urn:lsid:zoobank.org:pub:DB0D720E-F076-456E-8FCF-03B62ACF0CE4 Morphological and molecular description of a new genus and species of black coral (Cnidaria: Anthozoa: Hexacorallia: Antipatharia: Antipathidae: Blastopathes) from Papua New Guinea JEREMY HOROWITZ1,5, MERCER R. BRUGLER2,3, TOM C.L. BRIDGE1,4 & PETER F. COWMAN1,6 1Australian Research Council Centre of Excellence for Coral Reef Studies, James Cook University, 101 Angus Smith Drive, Townsville, QLD, 4811, Australia 2Department of Natural Sciences, University of South Carolina Beaufort, 801 Carteret Street, Beaufort, SC, 29902, USA �[email protected]; https://orcid.org/0000-0003-3676-1226 3Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, USA 4Biodiversity and Geosciences Program, Museum of Tropical Queensland, Queensland Museum, 70-102 Flinders St, Townsville, QLD, 4810, Australia. �[email protected]; https://orcid.org/0000-0003-3951-284X 5 �[email protected]; https://orcid.org/0000-0002-2643-5200 6 �[email protected]; https://orcid.org/0000-0001-5977-5327 Abstract Blastopathes medusa gen. nov., sp. nov., is described from Kimbe Bay, Papua New Guinea, based on morphological and molecular data. Blastopathes, assigned to the Antipathidae, is a large, mythology-inspiring black coral characterized by clusters of elongate stem-like branches that extend out at their base and then curve upward. Colonies are not pinnulate and contain single branches, which could represent new branch cluster formations. Morphological and molecular (mitochondrial DNA and targeted capture of nuclear loci) evidence supporting the establishment of a new genus is discussed. This is the first study to utilize the target capture of ultraconserved elements (UCEs) and exonic loci to elucidate phylogenetic relationships among black corals and to identify and place a new genus and species. Key words: taxonomy, systematics, ultraconserved elements, targeted capture, igrN, phylogeny, Kimbe Bay Introduction Black corals (Anthozoa: Antipatharia) are colonial hexacorallians that live in all oceans and marine habitats, from shallow water to at least 8,600 m depth (Molodtsova 2006; Wagner et al. 2012a). Black corals are characterized by polyps with six tentacles and microscopic spines along a skeleton composed of chitin and scleroproteins (Bo et al. 2012). A majority of the 45 genera (containing around 273 species) occur beyond recreational SCUBA diving depths (> 50 m) (Cairns 2007; Opresko 2019; Wagner et al. 2012b), which makes collecting and conducting in-situ experiments on black corals difficult. Black corals also have few and variable morphological features, leading to uncertainty regarding the number of extant species, their distributions, and phylogenetic relationships within the Order. In addition, slow evolution of mitochondrial genes in black corals (2.3 times slower than octocoral mtDNA) results in low genus and species-level resolution in corresponding phylogenetic trees, making it difficult to properly resolve the taxonomy and systematics of the Antipatharia (Bo et al. 2018; Brugler et al. 2013). Due to the many challenges associated with sampling and identifying black corals, the group is often ignored during exploratory expeditions. For example, taxonomists have been exploring Kimbe Bay, Papua New Guinea for over 50 years which has led to the description of many new fish and hard coral species (Allen & Munday 1995; Allen & Randall 1996; Hemond & Vollmer 2010; Wallace 1999); however, not a single black coral taxonomic de- scription has resulted from these efforts. Our first visit to Kimbe Bay in 2018 led to the discovery of a large, mythol- ogy-inspiring black coral that has to date gone unreported by researchers. In this study, we integrate morphological and molecular data to compare Blastopathes gen. nov., sp. nov. with other black coral genera to formally describe the new genus based on the new species Blastopathes medusa. This Accepted by M. Daly: 16 Jul. 2020; published: 3 Aug. 2020 553 new genus is characterized by clusters of elongate stem-like branches that extend out at their base and then curve upward. Based on mitochondrial DNA (mtDNA) nad5-IGR-nad1 (IGR = intergenic region) and the targeted capture of ultraconserved elements (UCEs) and exonic loci, B. medusa was found to group within the Antipathidae. The targeted capture of UCE/exon loci was recently used to successfully reconstruct the evolutionary origins of Antho- zoa (Quattrini et al. 2018), with the inclusion of a number of black coral specimens. Subsequently, an enhanced bait set was designed to increase capture efficiency when targeting members of the Subclass Hexcorallia (Cowman et al. 2020). Using this enhanced target enrichment bait set for Hexacorallia (Cowman et al. 2020), we captured both UCE and exonic loci for phylogenomic analysis within the Order Antipatharia. The resulting phylogeny, in combination with morphological evidence, is used to describe and systematically place the newly discovered genus and species. Our results demonstrate that much about black coral biodiversity remains to be discovered, even in a relatively shallow and well-explored habitat like Kimbe Bay, Papua New Guinea. Materials and methods Specimens. The new species was collected from Kimbe Bay, West New Britain Province, Papua New Guinea. Kimbe Bay is located on the north coast of the volcanic island of New Britain in the Bismarck Sea, Papua New Guinea. Kimbe Bay supports shallow reefs along the coast and underwater mountains (seamounts) whose bases can reach depths > 500 m. Five specimens belonging to the new species were collected in March 2019 by SCUBA diving at shallow depths (<40 m) in Kimbe Bay (Figure 1). Small (~5 cm) fragments were subsampled from distal sections of terminal branches of each colony for Scanning Electron Microscope (SEM) imaging of the spines. Small tissue samples were also preserved in 100% ethanol for molecular analyses. Morphological descriptions were based on colony branching, spine morphology, and polyp characteristics. Polyp size was measured as the distance between the distal edge of distal lateral tentacles and the proximal edge of proximal lateral tentacles of one polyp. Spine height was measured as the distance between the apex and the center of the base of one spine. The distance between spines was measured from the center of the base of one spine to the center of the base of an adjoining spine in the same axial row. Accessioning of type and molecular sequences. Type material is accessioned at the Museum of Tropical Queensland, Townsville, Australia (registration numbers: MTQ G74904–MTQ G74915) and the Papua New Guin- ea National Museum and Art Gallery (registration numbers: NMAG 1892, NMAG 1893, NMAG 1895–NMAG 1898. All molecular data were deposited in GenBank under accession numbers MN974021–MN974022 (mtDNA), and SRA Genbank BioProject submission PRJNA644402, BioSamples #SAMN15459017 and SAMN15461031– 15461062 (UCE/exon). Molecular analyses. Three specimens representing the new genus collected from Kimbe Bay in March 2019 were sampled for genomic DNA, in addition to non-Blastopathes museum specimens collected from all three Oceans (46 and 31 specimens for mitochondrial and UCE/exon analyses, respectively) to place the new genus in the Antipatharia. Antipathes grandis Verrill, 1928, (sample ANT14) was also included in the UCE/exon analysis from Quattrini et al. (2018) to have an additional member of Antipathes compared with the new genus. Occurrence data for Blastopathes medusa specimens and Antipatharian specimens included in UCE/exon analysis are detailed in Table 1 and specimens included in nad5-IGR-nad1 (hereafter, “igrN”) analysis can be found in Brugler, Opresko and France (2013). The specimens underwent amplification and Sanger sequencing of the mitochondrial nad5-IGR- nad1 region; and high throughput next generation sequencing to target UCEs and exons. The mitochondrial region chosen was based on the results of Brugler, Opresko and France (2013) showing that igrN was the most variable region within the black coral mitogenome. The targeted capture approach for UCE/exon loci was chosen due to their ability to resolve phylogenetic relationships across shallow and deep time scales in corals (Cowman et al. 2020; Quattrini et al. 2018). DNA extraction for all specimens followed the protocol detailed in MacIsaac et al. (2013). With regard to the igrN analysis, DNA quantification, PCR primers and reagents, PCR thermocycling profiles, PCR cleanup, cycle sequencing, cycle sequencing cleanup, traditional Sanger sequencing on an ABI‐3730xL, and multiple sequence alignment (exception: Gap opening penalty: 1.0; Offset value: 0.0) followed the protocol detailed in MacIsaac et al. (2013). While faint PCR products for igrN were obtained for the holotype (MTQ G74904), the resulting DNA chromatograms were unreadable; therefore, we were only able to obtain mtDNA sequence data for two specimens (NMAG 1893 and NMAG 1895). The newly-obtained sequence data were added to the igrN mul- 554 · Zootaxa 4821 (3) © 2020 Magnolia Press HOROWITZ ET AL. tiple sequence alignment from Brugler, Opresko and France (2013) in the form of a single representative because the two sequences shared identical haplotypes
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