Abstract
Members of Chrysogorgia Duchassaing & Michelotti, 1864, known as golden corals, are distributed in the deep sea worldwide. Based on specimens obtained from the tropical western Pacific and an integrated morphological-molecular approach, we established a new genus Parachrysogorgia, which includes 13 species separated from Chrysogorgia, and described nine species of Chrysogorgia including three new species C. arboriformis sp. nov., C. cylindrata sp. nov. and C. tenuis sp. nov., and two new combinations of Parachrysogorgia. The genus differs from Chrysogorgia by possessing a bare, sclerite-free area at the base of each tentacle and eight distinct projections beneath tentacles. Chrysogorgia cylindrata sp. nov. is distinguished by a slender bottlebrush-shaped colony in adults, elongate and lobed scales in coenenchyme, and warty and thick sclerites near the polyp mouth area. Chrysogorgia arboriformis sp. nov. is characterized by a tree-shaped colony with a long unbranched stem, slender and thin scales usually with large warts in coenenchyme. Chrysogorgia tenuis sp. nov. can be separated by a broad bottlebrush-shaped colony, long interbranch distance, elongate scales with broad lobes in polyp body wall. In contrast to the intraspecific variation of colony form, branch internodes, branching sequence and polyps, the sclerite forms and arrangement in different parts can be viewed as the most important character to separate Chrysogorgia species. For convenient identification, we divided the Chrysogorgia species into 12 morphological groups and divided Parachrysogorgia into three groups based on their sclerite forms and arrangement.
References
- Bayer, F.M., Grasshoff, M. & Verseveldt, J. (1983) Illustrated Trilingual Glossary of Morphological and Anatomical Terms Applied to Octocorallia. E.J. Brill/Dr. W. Backhuys, Leiden, 75 pp.
- Bayer, F.M. & Stefani, J. (1988) A new species of Chrysogorgia (Octocorallia: Gorgonacea) from New Caledonia, with descriptions of some other species from the western Pacific. Proceedings of the Biological Society of Washington, 101, 257–279.
- Berntson, E.A., Bayer, F.M., McArthur, A.G. & France, S.C. (2001) Phylogenetic relationships within the Octocorallia (Cnidaria: Anthozoa) based on nuclear 18S rDNA sequences. Marine Biology, 138, 235–246. https://doi.org/10.1007/s002270000457
- Brugler, M.R. & France, S.C. (2008) The mitochondrial genome of a deep-sea bamboo coral (Cnidaria, Anthozoa, Octocorallia, Isididae): Genome structure and putative origins of replication are not conserved among octocorals. Journal of Molecular Evolution, 67, 125–136. https://doi.org/10.1007/s00239-008-9116-2
- Cairns, S.D. (2001) Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 1: the genus Chrysogorgia Duchassaing & Michelloti, 1864. Proceedings of the Biological Society of Washington, 114, 746–787. [https://repository.si.edu/handle/10088/2462]
- Cairns, S.D. (2002) A new species of Chrysogorgia (Anthozoa: Octocorallia) from the Antarctic. Proceedings of the Biological Society of Washington, 115, 217–222.
- Cairns, S.D. & Wirshing, H.H. (2018) A phylogenetic analysis of the Primnoidae (Anthozoa: Octocorallia: Calcaxonia), with a key to the genera and subgenera. BMC Evolutionary Biology, 18 (66), 1–20. https://doi.org/10.1186/s12862-018-1182-5
- Cairns, S.D., Cordeiro, R.T.S., Xu, Y., Zhan, Z. & Alderslade, P. (2021) A new family and two new genera of calcaxonian octocoral, including a redescription of Pleurogorgia militaris (Cnidaria: Octocorallia: Chrysogorgiidae) and its placement in a new genus. Invertebrate Systematics, 35, 282–297. https://doi.org/10.1071/IS20066
- Cordeiro, R.T.S., Castro, C.B. & Pérez, C.D. (2015) Deep-water octocorals (Cnidaria: Octocorallia) from Brazil: family Chrysogorgiidae Verrill, 1883. Zootaxa, 4058 (1), 81–100. https://doi.org/10.11646/zootaxa.4058.1.4
- Duchassaing, P. & Michelotti, J. (1864) Supplément au mémoire sur les coralliaires des Antilles. Mémoires de l’ Académie des Sciences de Turin, 2, 97–206. https://doi.org/10.5962/bhl.title.105196
- Ehrenberg, C.G. (1834) Beiträge zur physiologischen Kenntniss der Corallenthiere im allgemeinen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben. Abhandlungen der Königlichen Akademie der Wissenschaften, Berlin, 1, 225–380.
- Everett, M.V., Park, L.K., Berntson, E.A., Elz, A.E. & Clarke, M.E. (2016) Large-scale genotyping-by-sequencing indicates high levels of gene flow in the deep-sea octocoral Swiftia simplex (Nutting, 1909) on the west coast of the united states. PLoS One, 11 (10), e0165279. https://doi.org/10.1371/journal.pone.0165279
- Folmer, O., Black, M., Hoeh, W., Lutz, R., Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome coxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
- Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321. https://doi.org/10.1093/sysbio/syq010
- Haeckel, E. (1866) Generelle morphologie der Organismen (Vol. 2). Verlag von Georg Reimer, Berlin, 652 pp. https://doi.org/10.1515/9783110848281
- Herrera, S., Baco, A. & Sánchez, J.A. (2010) Molecular systematics of the bubblegum coral genera (Paragorgiidae, Octocorallia) and description of a new deep-sea species. Molecular Phylogenetics and Evolution, 55 (1), 123–135. https://doi.org/10.1016/j.ympev.2009.12.007
- Kinoshita, K. (1913) Studien über einige Chrysogorgiiden Japans. Journal of the College of Science, Tokyo Imperial University, 33, 1–47.
- Kükenthal, W. (1908) Diagnosen neuer Gorgoniden aus der Gattung Chrysogorgia (6. Mitteilung). Zoologischer Anzeiger, 33, 704–708.
- Kükenthal, W. (1919) Gorgonaria. Wissenschaftliche Ergennisse der Deutschen Tiefsee - Expedition auf dem Dampfer “Valdivia” 1898-1899, 13 (2), 1–946.
- Lamouroux, J.V.F. (1812) Extrait d’un mémoire sur la classification des polypiers coralligènes non entièrement piérreux. Nouveau Bulletin des Sciences, Société Philomathique de Paris, 3 (63), 181–188.
- Madsen, F.J. (1944) Octocorallia. Danish Ingolf-Exped, 5 (13), 1–65.
- McFadden, C.S., Benayahu, Y., Pante, E., Thoma, J.N., Nevarez, P.A. & France, S.C. (2011) Limitations of mitochondrial gene barcoding in Octocorallia. Molecular Ecology Resources, 11 (1), 19–31. https://doi.org/10.1111/j.1755-0998.2010.02875.x
- McFadden, C.S. & van Ofwegen, L.P. (2012) Stoloniferous octocorals (Anthozoa, Octocorallia) from South Africa, with descriptions of a new family of Alcyonacea, a new genus of Clavulariidae, and a new species of Cornularia (Cornulariidae). Invertebrate Systematics, 26 (4), 331–356. https://doi.org/10.1071/IS12035
- McFadden, C.S., Brown, A.S., Brayton, C., Hunt, C.B. & van Ofwegen, L.P. (2014) Application of DNA barcoding to biodiversity studies of shallow-water octocorals: molecular proxies agree with morphological estimates of species richness in Palau. Coral Reefs, 33, 275–286. https://doi.org/10.1007/s00338-013-1123-0
- McFadden, C.S., van Ofwegen, L.P. & Quattrini, A.M. (2022) Revisionary systematics of Octocorallia (Cnidaria: Anthozoa) guided by phylogenomics. Bulletin of the Society of Systematic Biologists, 1 (3), 8735. https://doi.org/10.18061/bssb.v1i3.8735
- Mosher, C.V., Watling, L. (2009) Partners for life: a brittle star and its octocoral host. Marine Ecology Progress Series, 397, 81–88. https://doi.org/10.3354/meps08113
- Nutting, C.C. (1908) Descriptions of the Alcyonaria collected by the U.S. Bureau of Fisheries steamer Albatross in the vicinity of the Hawaiian Islands in 1902. Proceedings of the United States National Museum, 34, 543–601. https://doi.org/10.5479/si.00963801.34-1624.543
- Pante, E., Watling, L. (2012) Chrysogorgia from the New England and Corner Seamounts: Atlantic–Pacific connections. Journal of the Marine Biological Association of the United Kingdom, 92 (5), 911–927. https://doi.org/10.1017/S0025315411001354
- Pante, E., France, S.C., Couloux, A., Cruaud, C., McFadden, C.S., Samadi, S. & Watling, L. (2012) Deep-sea origin and in-situ diversification of chrysogorgiid octocorals. PLoS One, 7 (6), e38357. https://doi.org/10.1371/journal.pone.0038357
- Pante, E., France, S.C., Gey, D., Cruaud, C. & Samadi, S. (2015) An inter-ocean comparison of coral endemism on seamounts: the case of Chrysogorgia. Journal of Biogeography, 42, 1907–1918. https://doi.org/10.1111/jbi.12564
- Pasternak, F.A. (1981) Alcyonacea and Gorgonacea. In: Benthos of the Submarine Mountains of the Marcus-Necker and adjacent Pacific Regions. Academy of Sciences of the USSR, P.P. Shirshov Institute of Oceanology, Moscow, 1981, pp. 40–56.
- Quattrini, A.M., Georgian, S.E., Byrnes, L., Stevens, A. & Cordes, E.E. (2013) Niche divergence by deep-sea octocorals in the genus Callogorgia across the continental slope of the Gulf of Mexico. Molecular Ecology, 22 (15), 4123–4140. https://doi.org/10.1111/mec.12370
- Quattrini, A.M., Gómez, C.E. & Cordes, E.E. (2017) Environmental filtering and neutral processes shape octocoral community assembly in the deep sea. Oecologia, 183, 221–236. https://doi.org/10.1007/s00442-016-3765-4
- Rambaut, A. & Drummond, A.J. (2022) Tracer. Version 1.4. Available from: https://beast.bio.ed.ac.uk/Tracer (accessed 1 September 2022)
- Ronquist, F.R. & Huelsenbeck, J.P. (2003) Mrbayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
- Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 30 (12), 2725–2729. https://doi.org/10.1093/molbev/mst197
- Thoma, J.N., Pante, E., Brugler, M.R. & France, S.C. (2009) Deep-sea octocorals and antipatharians show no evidence of seamount-scale endemism in the NW Atlantic. Marine Ecology Progress Series, 397, 25–35. https://doi.org/10.3354/meps08318
- Untiedt, C.B., Quattrini, A.M., McFadden, C.S., Alderslade, P.A., Pante, E. & Burridge, C.P. (2021) Phylogenetic relationships within Chrysogorgia (Alcyonacea: Octocorallia), a morphologically diverse genus of octocoral, revealed using a target enrichment approach. Frontiers in Marine Science, 7, 599984. https://doi.org/10.3389/fmars.2020.599984
- Verrill, A.E. (1883) Report on the Anthozoa, and on some additional species dredged by the “Blake” in 1877–1879, and by the U.S. Fish Commission steamer “Fish Hawk” in 1880–1882. Bulletin of the Museum of Comparative Zoology at Harvard College, 11, 1–72.
- Versluys, J. (1902) Die Gorgoniden der Siboga-Expedition. 1. Die Chrysogorgiiden. Siboga-Expeditie, 13, 1–120.
- Watling, L., France, S.C., Pante, E. & Simpson, A. (2011) Chapter Two-Biology of Deep-Water Octocorals. Advances in Marine Biology, 60 (3), 41–122. https://doi.org/10.1016/B978-0-12-385529-9.00002-0
- Wright, E.P. & Studer, T. (1889) Report on the Alcyonaria collected by H.M.S. Challenger during the years 1873–76. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76, Zoology, 31 (64), 1–314.
- Xu, Y., Li, Y., Zhan, Z. & Xu, K. (2019) Morphology and phylogenetic analysis of two new deep-sea species of Chrysogorgia (Cnidaria, Octocorallia, Chrysogorgiidae) from Kocebu Guyot (Magellan seamounts) in the Pacific Ocean. Zookeys, 881, 91–107. https://doi.org/10.3897/zookeys.881.34759
- Xu, Y., Zhan, Z., Li, Y. & Xu, K. (2020) Morphology and phylogenetic analysis of two new species of deep-sea golden gorgonians (Cnidaria: Octocorallia: Chrysogorgiidae) from seamounts in the Western Pacific Ocean. Zootaxa, 4731 (2), 249–262. https://doi.org/10.11646/zootaxa.4731.2.4
- Xu, Y., Zhan, Z. & Xu, K. (2020a) Morphology and molecular phylogeny of three new deep-sea species of Chrysogorgia (Cnidaria, Octocorallia) from seamounts in the tropical Western Pacific Ocean. PeerJ, 8, e8832. https://doi.org/10.7717/peerj.8832
- Xu, Y., Zhan, Z. & Xu, K. (2020b) Morphology and phylogenetic analysis of five deep-sea golden gorgonians (Cnidaria, Octocorallia, Chrysogorgiidae) in the Western Pacific Ocean, with the description of a new species. ZooKeys, 989, 1–37. https://doi.org/10.3897/zookeys.989.53104
- Xu, Y., Zhan, Z. & Xu, K. (2021) Morphology and phylogeny of Chrysogorgia pinniformis sp. nov. and C. varians sp. nov., two golden corals from the Caroline seamounts in the tropical Western Pacific Ocean. Journal of Oceanology and Limnology, 39, 1767–1789. https://doi.org/10.1007/s00343-021-0386-5