Dead man’s fingers point to new taxa: Two new genera of New Zealand soft corals (Anthozoa, Octocorallia) and a revision of Alcyonium aurantiacum Quoy & Gaimard, 1833

Keywords: Alcyonacea, integrative taxonomy, phylogenetics, species complex, species delimitation


The taxonomic status of Alcyonium aurantiacum Quoy & Gaimard, 1833, an octocoral endemic to New Zealand, was reviewed through morpho-molecular data comparisons in an integrative approach. Molecular phylogenetic analyses (nuclear 28S and mitochondrial mtMutS) resolved New Zealand taxa as more closely related to other genera and nominal Alcyonium Linnaeus, 1758 from South America than to the genus’ North Atlantic type species. Due to low genetic variation, species delimitation relied predominantly on identifying consistent differences in sclerite and colony morphology. The former A. aurantiacum is reassigned to Kotatea gen. nov. as K. aurantiaca gen. et comb. nov. and seven new species are described in this genus (K. amicispongia gen. et sp. nov., K. lobata gen. et sp. nov., K. kapotaiora gen. et sp. nov., K. kurakootingotingo gen. et sp. nov., K. niwa gen. et sp. nov., K. raekura gen. et sp. nov., and K. teorowai gen. et sp. nov.). Three new species in Ushanaia gen. nov. are also described (U. ferruginea gen. et sp. nov., U. fervens gen. et sp. nov. and U. solida gen. et sp. nov. ). These descriptions increase our understanding of New Zealand’s endemic octocoral diversity and contribute to ongoing systematic revisions of Alcyonium.


Alderslade P. 2000. Four new genera of soft corals (Coelenterata: Octocorallia), with notes on the classification of some established taxa. Zoologische Mededelingen (Leiden) 74 (16): 237–249.

Baco A.R. & Cairns S.D. 2012. Comparing molecular variation to morphological species designations in the deep-sea coral Narella reveals new insights into seamount coral ranges. PLoS ONE 7 (9): e45555.

Bayer F.M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa), with diagnoses of new taxa. Proceedings of the Biological Society of Washington 94 (3): 902-–948.

Benayahu Y. & Schleyer M.H. 1995. Corals of the south-west Indian Ocean. 2. Eleutherobia aurea spec. nov. (Cnidaria, Alcyonacea) from deep reefs on the KwaZulu-Natal coast, South Africa. Oceanographic Research Institute (Durban) Investigational Report 68: 1–12.

Benham W.B. 1928. On some alcyonarians from New Zealand waters. Transactions and Proceedings of the New Zealand Institute 59 (1): 67–84.

Bilewitch J.P. & Degnan S.M. 2011. A unique horizontal gene transfer event has provided the octocoral mitochondrial genome with an active mismatch repair gene that has potential for an unusual self-contained function. BMC Evolutionary Biology 11: e228.

Brazeau D.A. & Harvell C.D. 1994. Genetic structure of local populations and divergence between growth forms in a clonal invertebrate, the Caribbean octocoral Briareum asbestinum. Marine Biology 119: 53–60.

Cairns S.D. 2012. The Marine Fauna of New Zealand: New Zealand Primnoidae (Anthozoa: Alcyonacea). Part 1. Genera Narella, Narelloides, Metanarella, Calyptrophora, and Helicoprimnoa. NIWA Biodiversity Memoir 126, National Institute of Water and Atmospheric Research, Wellington.

Cairns S.D. 2016. The Marine Fauna of New Zealand: Primnoid Octocorals (Anthozoa: Alcyonacea). Part 2. Primnoella, Callozostrom, Metafannyella, Callogorgia, Fanellia and other Genera. NIWA Biodiversity Memoir 129, National Institute of Water and Atmospheric Research, Wellingotn.

Cairns S.D. & Baco A. 2007. Review and five new Alaskan species of the deep-water octocoral Narella (Octocorallia: Primnoidae). Systematics and Biodiversity 5 (4): 391–407.

Cairns S.D. & Bayer F.M. 2005. A review of the genus Primnoa (Octocorallia: Gorgonacea: Primnoidae), with the description of two new species. Bulletin of Marine Science 77 (2): 225–256.

Cairns S., Gershwin L., Brook F., Pugh P., Dawson E., Ocaña O., Vervoort W., Williams G., Watson J., Opresko D., Schuchert P., Hine P., Gordon D., Campbell H., Wright C., Sánchez J. & Fautin D. 2009. Phylum Cnidaria: Corals, medusae, hydroids, myxozoans. In: Gordon D.P. (ed.) New Zealand Inventory of Biodiversity, Volume 1: Kingdom Animalia: 59–101. Canterbury University Press, Christchurch.

Casas C., Ramil F. & van Ofwegen L.P. 1997. Octocorallia (Cnidaria: Anthozoa) from the Scotia Arc, South Atlantic Ocean. I. The genus Alcyonium Linnaeus, 1758. Zoologische Mededelingen 71: 299–311.

Cryer M., O’Shea S., Gordon D., Kelly M., Drury J., Morrison M., Hill A., Saunders H., Shankar U., Wilkinson M. & Foster G. 2000. Distribution and Structure of Benthic Invertebrate Communities Between North Cape and Cape Reinga. Final Research Report for Ministry of Fisheries Research Project ENV9805 Objectives 1–4, National Institute of Water and Atmospheric Research, Wellington.

Daly M., Brugler M.R., Cartwright P., Collins A.G., Dawson M.N., Fautin D.G., France S.C., McFadden C.S., Opresko D.M., Rodriguez E., Romano S.L. & Stake J.L. 2007. The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus. Zootaxa 1668 (1): 127–182.

D’Archino R., Ngauma B., Norman J.H. & Zuccarello G.C. 2020. A new species of Polycoelia (Kallymeniaceae, Rhodophyta), P. kurariirapa, from the Three Kings Islands/Manawatāwhi, New Zealand. Phycologia 59 (2): 127–132.

Darriba D., Taboada G.L., Doallo R. & Posada D. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9 (8): 772.

Doak W. 1971. Beneath New Zealand Seas. Reed, Auckland.

Dueñas L.F., Alderslade P. & Sánchez J.A. 2014. Molecular systematics of the deep-sea bamboo corals (Octocorallia: Isididae: Keratoisidinae) from New Zealand with descriptions of two new species of Keratoisis. Molecular Phylogenetics and Evolution 74: 15–28.

Dueñas L.F., Tracey D.M., Crawford A.J., Wilke T., Alderslade P. & Sánchez J.A. 2016. The Antarctic Circumpolar Current as a diversification trigger for deep-sea octocorals. BMC Evolutionary Biology 16: e2.

Erickson K.L., Pentico A., Quattrini A.M. & McFadden C.S. 2020. New approaches to species delimitation and population structure of anthozoans: Two case studies of octocorals using ultraconserved elements and exons. Molecular Ecology Resources 21 (4): 1404–1404.

Fabricius K.E. & Alderslade P. 2001. Soft Corals and Sea Fans: a Comprehensive Guide to the Tropical Shallow-water Genera of the Central West-Pacific, the Indian Ocean and the Red Sea. Australian Institute of Marine Science, Townsville.

France S.C. & Hoover L.L. 2002. DNA sequences of the mitochondrial COI gene have low levels of divergence among deep-sea octocorals (Cnidaria: Anthozoa). Hydrobiologia 471: 149–155.

Galbreath R. 2020. Why have so few Māori or Moriori names been used in taxonomic description? New Zealand Journal of Ecology 45 (1): 3429.

Geneious Development Team 2020. Geneious Prime. Ver. 2020.1.1. Available from [accessed Jul. 2020].

Gillman L.N. & Wright S.D. 2020. Restoring indigenous names in taxonomy. Communications Biology 3 (1): e609.

GIMP Development Team 2020. GIMP GNU Image Manipulation Program. Ver. 2.8.22. Available from [accessed Jul. 2020].

Goldberg W.M., Grange K.R., Taylor G.T. & Zuniga A.L. 1990. The structure of sweeper tentacles in the black coral Antipathes fiordensis. The Biological Bulletin 179 (1): 96–104.

Grange K.R., Singleton R.I., Richardson J.R., Hill P.J. & Main W.D.L. 1981. Shallow rock-wall biological associations of some southern fiords of New Zealand. New Zealand Journal of Zoology 8 (2): 209–227.

Grange K.R., Watson J., Cook S.D.C., Barnett T.J., Brook F.J. & Cairns S.D. 2010. Phylum Cnidaria. In: Cook S.D.C. (ed.) New Zealand Coastal Marine Invertebrates 1: 137---–-----247. Canterbury University Press, Christchurch.

Grehan J. 2020. Lucy Cranwell, Leon Croizat, and the biogeography of Manawatāwhi (Three Kings Islands). Botanical Society of Otago Newsletter 91: 14–19.

Groot S. & Weinberg S. 1982. Biogeography, taxonomical status and ecology of Alcyonium (Parerythropodium) coralloides (Pallas, 1766). Marine Ecology 3 (4): 293–312.

Hadley A. 2020. CombineZP. Available from [accessed Jul. 2020].

Halász A., Reynolds A.M., McFadden C.S., Toonen R.J. & Benayahu Y. 2014. Could polyp pulsation be the key to species boundaries in the genus Ovabunda (Octocorallia: Alcyonacea: Xeniidae)? Hydrobiologia 759: 95–107.

Herrera S. & Shank T.M. 2016. RAD sequencing enables unprecedented phylogenetic resolution and objective species delimitation in recalcitrant divergent taxa. Molecular Phylogenetics and Evolution 100: 70–79.

Herrera S., Shank T.M. & Sánchez J.A. 2012. Spatial and temporal patterns of genetic variation in the widespread antitropical deep-sea coral Paragorgia arborea. Molecular Ecology 21 (24): 6053–6067.

Hickson S.J. 1895. The anatomy of Alcyonium digitatum. Quarterly Journal of Microscopical Science 37: 343–388.

Huelsenbeck J.P. & Ronquist F. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17 (8): 754–755.

Jenkins T.L., Ellis C.D. & Stevens J.R. 2019. SNP discovery in European lobster (Homarus gammarus) using RAD sequencing. Conservation Genetics Resources 11: 253–257.

Katoh K. & Standley D.M. 2013. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30 (4): 772–780.

Katoh K., Misawa K., Kuma K. & Miyata T. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30 (14): 3059– 3066.

Kim E., Lasker H.R., Coffroth M.A. & Kim K. 2004. Morphological and genetic variation across reef habitats in a broadcast-spawning octocoral. Hydrobiologia 530: 423–432.

Kumar S., Stecher G. & Tamura K. 2016. MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33 (7): 1870–1874.

Li Y., Wang W., Liu X., Luo W., Zhang J. & Gul Y. 2011. DNA extraction from crayfish exoskeleton. Indian Journal of Experimental Biology 49 (12): 953–957.

Maffi L. 2005. Linguistic, cultural and biological diversity. Annual Review of Anthropology 34: 599–617.

McAllister T.G., Beggs J.R., Ogilvie S., Kirikiri R., Black A. & Wehi P.M. 2019. Kua takoto te mānuka: mātauranga Māori in New Zealand ecology. New Zealand Journal of Ecology 43 (3): e3393.

McFadden C.S. 1999. Genetic and taxonomic relationships among Northeastern Atlantic and Mediterranean populations of the soft coral Alcyonium coralloides. Marine Biology 133 (2): 171–184.

McFadden C.S. & Hutchinson M.B. 2004. Molecular evidence for the hybrid origin of species in the soft coral genus Alcyonium (Cnidaria: Anthozoa: Octocorallia). Molecular Ecology 13 (6): 1495–1505.

McFadden C.S. & van Ofwegen L.P. 2012. A revision of the soft coral genus, Eunephthya Verrill, 1869 (Anthozoa: Octocorallia: Nephtheidae), with a description of four new species from South Africa. Zootaxa 3485 (1): 1---–-25.

McFadden C.S. & van Ofwegen L.P. 2013a. Molecular phylogenetic evidence supports a new family of octocorals and a new genus of Alcyoniidae (Octocorallia, Alcyonacea). ZooKeys 346: 59–83.

McFadden C.S. & van Ofwegen L.P. 2013b. A second, cryptic species of the soft coral genus Incrustatus (Anthozoa: Octocorallia: Clavulariidae) from Tierra del Fuego, Argentina, revealed by DNA barcoding. Helgoland Marine Research 67: 137–147.

McFadden C.S. & van Ofwegen L.P. 2017. Revisionary systematics of the endemic soft coral fauna (Octocorallia: Alcyonacea: Alcyoniina) of the Agulhas Bioregion, South Africa. Zootaxa 4363 (4): 451–488.

McFadden C.S., Donahue R., Hadland B.K. & Weston R. 2001. A molecular phylogenetic analysis of reproductive trait evolution in the soft coral genus Alcyonium. Evolution 55 (1): 54–67. Available from [accessed 27 Jul. 2022].

McFadden C.S., Tullis L.D., Hutchinson M.B., Winner K. & Sohm J.A. 2004. Variation in coding (NADH dehydrogenase subunits 2, 3, and 6) and noncoding intergenic spacer regions of the mitochondrial genome in Octocorallia (Cnidaria: Anthozoa). Marine Biotechnology 6 (6): 516–-526.

McFadden C.S., Rettig P.M. & Beckman E.J. 2005. Molecular evidence for hybridization between two alcyoniid soft coral species with contrasting life histories. Integrative and Comparative Biology 45 (6): 1042. [Abstract.]

McFadden C.S., Alderslade P., van Ofwegen L.P., Johnsen H. & Rusmevichientong A. 2006a. Phylogenetic relationships within the tropical soft coral genera Sarcophyton and Lobophytum (Anthozoa, Octocorallia). Invertebrate Biology 125 (4): 288–305.

McFadden C.S., France S.C., Sánchez J.A. & Alderslade P. 2006b. A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa) based on mitochondrial protein-coding sequences. Molecular Phylogenetics and Evolution 41 (3): 513–527.

McFadden C.S., van Ofwegen L.P., Beckman E.J., Benayahu Y. & Alderslade P. 2009. Molecular systematics of the speciose Indo-Pacific soft coral genus, Sinularia (Anthozoa: Octocorallia). Invertebrate Biology 128 (4): 303–323.

McFadden C.S., Sánchez J.A. & France S.C. 2010. Molecular phylogenetic insights into the evolution of Octocorallia: a review. Integrative and Comparative Biology 50 (3): 389–410.

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: 19–31.

McFadden C.S., Brown A.S., Brayton C., Hunt C.B. & van Ofwegen L.P. 2014. Application of DNA barcoding in biodiversity studies of shallow-water octocorals: molecular proxies agree with morphological estimates of species richness in Palau. Coral Reefs 33 (2): 275–286.

McFadden C.S., Haverkort-Yeh R., Reynolds A.M., Halász A., Quattrini A.M., Zac H., Forsman Z.H., Benayahu Y. & Toonen R.J. 2017. Species boundaries in the absence of morphological, ecological or geographical differentiation in the Red Sea octocoral genus Ovabunda (Alcyonacea: Xeniidae). Molecular Phylogenetics and Evolution 112: 174–184.

Mills S., Tracey D., Bilewitch J., Sinniger F., Cairns S., Opresko D. & Kitahara M. 2019. Chapter 1: Taxonomic coral classification. In: Tracey D.M. & Hjorvarsdottir F. (eds) The State of Knowledge of Deep-Sea Corals in the New Zealand Region: 16–19. NIWA Science and Technology Series Number 84, Auckland.

Moore K.M., Alderslade P. & Miller K.J. 2016. A taxonomic revision of the genus Primnoisis Studer

[& Wright], 1887 (Coelenterata: Octocorallia: Isididae) using morphological and molecular data. Zootaxa 4075 (1): 1–141.

Moore K.M., Alderslade P. & Miller K.J. 2017. A taxonomic revision of Anthothela (Octocorallia: Scleraxonia: Anthothelidae) and related genera, with the addition of new taxa, using morphological and molecular data. Zootaxa 4304 (1): 1–212.

Morton J. 2004. Seashore Ecology of New Zealand and the Pacific. David Bateman Ltd, Auckland.

Morton J. & Miller M. 1973. The New Zealand Sea Shore: Second Edition. William Collins Sons and Co. Ltd, Glasgow.

Nelson W.A., Sutherland J.E., Ringham S. & Murupaenga H. 2019. Dictyota korowai sp. nov. (Dictyotales, Phaeophyceae) from Manawatāwhi/Three Kings Islands, northern New Zealand, previously confused with Dictyota intermedia. Phycologia 58 (4): 433–442.

Ngāti Kurī Trust Board 2013. Treaty Settlement Offer: Ratification Guide. Ngāti Kurī Trust Board inc., Ngataki. Available from [accessed 15 Jul. 2022].

Núñez-Flores M., Gomez-Uchida D. & López-González P.J. 2020. Molecular and morphological data reveal three new species of Thouarella Gray, 1870 (Anthozoa: Octocorallia: Primnoidae) from the Southern Ocean. Marine Biodiversity 50: e30.

Pante E., Abdelkrim J., Viricel A., Gey D., France S.C., Boisselier M.C. & Samadi S. 2015. Use of RAD sequencing for delimiting species. Heredity 114: 450–459.

Powell A.W.B. 1947. Native Animals of New Zealand, Auckland Museum Handbook of Zoology. The Unity Press Ltd, Auckland.

Prada C., Schizas N.V. & Yoshioka P.M. 2008. Phenotypic plasticity or speciation? A case from a clonal marine organism. BMC Evolutionary Biology 8: e47.

Quattrini A.M., Faircloth B.C., Dueñas L.F., Bridge T.C.L., Brugler M.R., Calixto-Botía I.F., DeLeo D.M., Forêt S., Herrera S., Lee S.M.Y., Miller D.J., Prada C., Rádis-Baptista G., Ramírez-Portilla C., Sánchez J.A., Rodríguez E. & McFadden C.S. 2018. Universal target-enrichment baits for anthozoan (Cnidaria) phylogenomics: New approaches to long-standing problems. Molecular Ecology Resources 18 (2): 281–295.

Quattrini A.M., Wu T., Soong K., Jeng M.S., Benayahu Y. & McFadden C.S. 2019. A next generation approach to species delimitation reveals the role of hybridization in a cryptic species complex of corals. BMC Evolutionary Biology 19: e116.

Quattrini A.M., Rodriguez E., Faircloth B.C., Cowman P.F., Brugler M.R., Farfan G.A., Hellberg M.E., Kitahara M.V., Morrison C.L., Paz-García D.A., Reimer J.D. & McFadden C.S. 2020. Palaeoclimate ocean conditions shaped the evolution of corals and their skeletons through deep time. Nature Ecology and Evolution 4 (11): 1531–1538.

Quoy J.R.C. & Gaimard J.P. 1833. Voyage de découvertes de l’Astrolabe: exécuté par ordre du roi, pendant les années 1826-1827-1828-1829, sous le commandement de M.J. Dumont d’Urville, zoologie, tome quatrième. J. Tastu, Paris.

Rowden A.A., Warwick R.M. & Gordon D.P. 2004. Bryozoan biodiversity in the New Zealand region and implications for marine conservation. Biodiversity and Conservation 13 (14): 2695–2721.

Sánchez J.A. 2005. Systematics of the bubblegum corals (Cnidaria : Octocorallia : Paragorgiidae) with description of new species from New Zealand and the Eastern Pacific. Zootaxa 1014 (1): 1–72.

Sánchez J.A. & Rowden A.A. 2006. Octocoral diversity on New Zealand seamounts. In: Gakkai N.S. (ed.) Proceedings of the 10th International Coral Reef Symposium, Okinawa, Japan, June 28 – July 2 2004: 1812–1820. International Society for Reef Studies.

Sánchez J.A., McFadden C.S., France S.C. & Lasker H.R. 2003. Molecular phylogenetic analyses of shallow-water Caribbean octocorals. Marine Biology 142: 975–987.

Sánchez J.A., Aguilar C., Dorado D. & Manrique N. 2007. Phenotypic plasticity and morphological integration in a marine modular invertebrate. BMC Evolutionary Biology 7: e122.

Stokvis F.R. & van Ofwegen L.P. 2006. New and redescribed encrusting species of Alcyonium from the Atlantic Ocean (Octocorallia: Alcyonacea: Alcyoniidae). Zoologische Mededelingen 80 (4): 165–183.

Sutherland W.J. 2003. Parallel extinction risk and global distribution of languages and species. Nature 423 (6937): 276–279.

Tracey D., Stewart R., Bilewitch J., Macpherson D., Chin C., Mills S., Kessel G., Aguirre D. & Liggins L. 2019. Chapter 12: Key coral groups. In: Tracey D.M. & Hjorvarsdottir F. (eds) The State of Knowledge of Deep-Sea Corals in the New Zealand Region: 62–77. NIWA Science and Technology Series Number 84, Auckland.

UNGA 2007. United Nations Declaration on the Rights of Indigenous Peoples: A/RES/61/295. Available from [accessed 18 Jul. 2022].

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: e599984.

van Ofwegen L.P., Häussermann V. & Försterra G. 2006. A new genus of soft coral (Octocorallia: Alcyonacea: Clavulariidae) from Chile. Zootaxa 1219 (1): 47–57.

van Ofwegen L.P., Häussermann V. & Försterra G. 2007. The genus Alcyonium (Octocorallia: Alcyonacea: Alcyoniidae) in Chile. Zootaxa 1607 (1): 1–19.

Veale A.J., de Lange P., Buckley T.R., Cracknell M., Hohaia H., Parry K., Raharaha-Nehemia K., Reihana K., Seldon D., Tawiri K. & Walker L. 2019. Using te reo Māori and ta re Moriori in taxonomy. New Zealand Journal of Ecology 43 (3): e3388.

Verseveldt J. 1973. On the validity of Alcyonium siderium Verrill (Coelenterata: Octocorallia). Zoologische Mededelingen 46 (16): 209–216.

Verseveldt J. & Alderslade P. 1982. Descriptions of types and other alcyonacean material (Coelenterata: Octocorallia) in the Australian Museum, Sydney. Records of the Australian Museum 34 (15): 619–647.

Verseveldt J. & van Ofwegen L.P. 1992. New and redescribed species of Alcyonium Linnaeus, 1758 (Anthozoa: Alcyonacea). Zoologische Mededelingen 66 (7): 155–181.

Waitangi Tribunal Report Wai 262. 2011. A Report in Claims Concerning New Zealand Law and Policy Affecting Maori Culture and Identity. Legislation Direct, Wellington.

West J.M. 1997. Plasticity in the sclerites of a gorgonian coral: Tests of water motion, light level, and damage cues. The Biological Bulletin 192 (2): 279–289.

West J.M., Harvell C.D. & Walls A.M. 1993. Morphological plasticity in a gorgonian coral (Briareum asbestinum) over a depth cline. Marine Ecology Progress Series 94: 61-–69.

Westerskov K. & Probert K. 1981. The Seas around New Zealand. Reed, Wellington.

Whyte K.P., Brewer J.P. & Johnson J.T. 2016. Weaving indigenous science, protocols and sustainability science. Sustainability Science 11: 25–32.

Wiens J.J. & Morrill M.C. 2011. Missing data in phylogenetic analysis: reconciling results from simulations and empirical data. Systematic Biology 60 (5): 719–731.

Williams G.C. 1986a. A new species of the octocorallian genus Alcyonium (Anthozoa: Alcyonacea) from southern Africa, with a revised diagnosis of the genus. Journal of Natural History 20: 53–63.

Williams G.C. 1986b. Morphology, systematics and variability of the southern African soft coral Alcyonium variabile (J. Stuart Thomson, 1921) (Octocorallia, Alcyoniidae). Annals of the South African Museum 96 (6): 241–270.

Williams G.C. 1988. Four new species of southern African octocorals (Cnidaria: Alcyonacea), with a further diagnostic revision of the genus Alcyonium Linnaeus, 1758. Zoological Journal of the Linnean Society 92 (1): 1–26.

Williams G.C. 1992. The Alcyonacea of southern Africa: Gorgonian octocorals (Coelenterata, Anthozoa). Annals of the South African Museum 101 (8): 181–296.

Williams G.C. 2000. Two new genera of soft corals (Anthozoa: Alcyoniidae) from South Africa, with a discussion of diversity and endemism in the Southern African Octocorallian fauna. Proceedings of the California Academy of Sciences 52 (6): 65–75.

Williams G.C. & Lundsten L. 2009. The nephtheid soft coral genus Gersemia Marenzeller, 1878, with the description of a new species from the northeast Pacific and a review of two additional species (Octocorallia: Alcyonacea). Zoologische Mededelingen (Leiden) 83 (4): 1067–1081.

Wirshing H.H., Messing C.G., Douady C.J., Reed J., Stanhope M.J. & Shivji M.S. 2005. Molecular evidence for multiple lineages in the gorgonian family Plexauridae (Anthozoa: Octocorallia). Marine Biology 47 (2): 497–508.

Xu Y., Zhan Z. & Xu K. 2020. 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.

Zachos F.E. 2016. Chapter 4: An annotated list of species concepts. In: Zachos F.E. (ed.) Species Concepts in Biology: Historical Development, Theoretical Foundations and Practical Relevance: 77–96. Springer, Cham, Switzerland.

Zwickl D.J. 2006. Genetic Algorithm Approaches for the Phylogenetic Analysis of Large Biological Sequence Datasets under the Maximum Likelihood Criterion. PhD Thesis, University of Texas, USA.

How to Cite
Kessel, G. M., Alderslade, P., Bilewitch, J. P., Schnabel, K. E., Norman, J., Tekaharoa Potts, R., & Gardner, J. P. (2022). Dead man’s fingers point to new taxa: Two new genera of New Zealand soft corals (Anthozoa, Octocorallia) and a revision of Alcyonium aurantiacum Quoy & Gaimard, 1833. European Journal of Taxonomy, 837(1), 1-85.