A new family for the enigmatic sea pen genus Gyrophyllum Studer, 1891 (Octocorallia, Pennatulacea), a molecular and morphological approach

Keywords: biodiversity, coral, morphology, molecular analyses, integrated approach


The description in 1891 of the sea pen genus Gyrophyllum Studer, 1891 and also the type species G. hirondellei Studer, 1891 was based on a single colony collected in the Azores Archipelago. During the 19th and 20th centuries, the family placement of this genus became controversial as the set of morphological features present in Gyrophyllum could justify its assignation to both the families Pennatulidae Ehrenberg, 1834 and Pteroeididae Kölliker, 1880. Deliberations over this intermediate set of characters finally ended in the reunification of the genera and species of both families under Pennatulidae by principle of priority. The use of molecular sources of information based on a series of sequencing techniques presents a different but promising phylogenetic scenario in order to go further in the understanding of pennatulacean systematics. In this paper, a complementary morphological and molecular study (multiloci sequences with three mitochondrial and one nuclear markers) based mainly on newly collected material is carried out. This study re-confirms from a molecular point of view previously published results that indicate the position of Gyrophyllum as being distant from Pennatula Linnaeus, 1758 and Pteroeides Herklots, 1858 (type genera of the families Pennatulidae and Pteroeididae, respectively). This fact together with the results of a detailed morphological examination strongly supports the placement of the enigmatic genus Gyrophyllum in a separate family: Gyrophyllidae fam. nov. and resolves the nomenclatural uncertainty at family level for this genus. Moreover, the characters previously considered useful in the distinction of the two currently recognised species G. hirondellei in the Atlantic and G. sibogae Hickson, 1916 in the Indo-western Pacific are revisited.


Bayer F.M. & Stefani J. 1988. Primnoidae (Gorgonacea) de Nouvelle-Caledonie. Bulletin du Muséum national d’histoire naturelle, Paris 10 (A) 3: 449–476.

Bayer F.M., Grasshoff M. & Verseveldt J. 1983. Illustrated Trilingual Glossary of Morphological and Anatomical Terms Applied to Octocorallia. Brill / Backhuys, Leiden.

Cuvier C. 1817. Le Règne Animal. Tome IV. Les Zoophytes. A. Belin, Paris.

Dana D.J. 1846. Zoophytes. United States Exploring Expedition during the years 1838–1842, under the command of Charles Wilkes, U.S.N., Vol. 7. C. Sherman, Philadelphia.

Deichmann E. 1936. The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of the Museum of Comparative Zoology, Harvard 53: 1–317. https://doi.org/10.5962/bhl.title.49348

Dolan E., Tyler P.A., Yesson C. & Rogers A.D. 2013. Phylogeny and systematics of deep-sea sea pens (Anthozoa: Octocorallia: Pennatulacea). Molecular Phylogenetic and Evolution 69: 610–618. https://doi.org/10.1016/j.ympev.2013.07.018

Ehrenberg C.G. 1834. Beitrage 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 zu Berlin 1: 225–380.

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. http://doi.org/10.1023/A:1016517724749

García-Cárdenas F.J., Drewery J. & López-González P.J. 2019. Resurrection of the sea pen genus Ptilella Gray, 1870 and description of Ptilella grayi n. sp. from the NE Atlantic (Octocorallia, Pennatulacea). Scientia Marina 83 (3): 261–276. https://doi.org/10.3989/scimar.04845.26A

García-Cárdenas F.J., Núñez-Flores M., López-González P.J. 2020. Molecular phylogeny and divergence time estimates in pennatulaceans (Cnidaria: Octocorallia: Pennatulacea). Scientia Marina 84 (4): 317–330. https://doi.org/10.3989/scimar.05067.28A

Gray J.E. 1848. Description of Sarcoptilus, a new genus of Pennatulidae. Proceedings of the Zoological Society of London 1848 (16): 45. https://doi.org/10.1111/j.1469-7998.1848.tb00171.x

Gray J.E 1870. Catalogue of Sea-pens or Pennatulariidae in the Collection of the British Museum, London. British Museum. London. https://doi.org/10.5962/bhl.title.11307

Haeckel E. 1866. Generelle Morphologie der organismen. Georg Reimer, Berlin. https://doi.org/10.1515/9783110848281

Herklots J.A. 1858. Notices pour servir à l’étude des polypiers nageurs ou pennatulidés. Bijdragen tot de Dierkunde 7 (1): 1–31. https://doi.org/10.1163/26660644-00701001

Hickson S.J. 1916. The Pennatulacea of the Siboga expedition, with a general survey of the order. Siboga Expeditie Monographs (14) 77: 1–265.

Huelsenbeck J.P. & Ronquist F. 2001. MrBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754–755. https://doi.org/10.1163/26660644-00701001

ICZN 1944. International Commission on Zoological Nomenclature. Opinion 185. Suppression of Bohadsch (J. B.) De quibusdam animalibus marinis, 1761 and of the German translation there of published by Leske (N. G.) in 1776. Opinions and Declarations Rendered by the International Commission on Zoological Nomenclature 3 (4): 37–52. Available from https://www.biodiversitylibrary.org/page/34653700#page/59/mode/1up [accessed 11 Nov. 2022].

Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide-sequences. Journal of Molecular Evolution 16 (2): 111–120.

Kölliker R.A. 1870. Anatomisch-Systematische Beschreibung der Alcyonararien. I. Die Pennatuliden. Abhandlungen von der Senckenbergischen naturforschenden Gesellschaft 7: 109–255.

Kölliker von R.A. 1880. Report on the Pennatulida dredged by H.M.S. Challenger during the years 1873–1876. Report on the Scientific Results of the Voyage of H.M.S. Challenger 1873–76. Zoology 1 (2): 1–41.

Kükenthal W. 1915. Pennatularia. In: Das Tierreich Lfg. 43. Verlag von R. Friedländer und Sohn, Berlin. https://doi.org/10.5962/bhl.title.1092

Kükenthal W. & Broch H. 1911. Pennatulacea. Wissenschaftiliche Ergebnisse Deutsche Tiefsee-Expedition „Valdivia“ 13 (2): 113–576.

Kushida Y. & Reimer J.D. 2019. Molecular phylogeny and diversity of sea pens (Cnidaria: Octocorallia: Pennatulacea) with a focus on shallow water species of the northwestern Pacific Ocean. Molecular Phylogenetic and Evolution 131: 233–244. https://doi.org/10.1016/j.ympev.2018.10.032

Kushida Y. & Reimer J.D. 2020. Description of the sea pen Calibelemnon hinoenma sp. nov. from shallow waters in southern Japan. Marine Biodiversity 50: 107. https://doi.org/10.1007/s12526-020-01132-1

Lamarck J.B.P.A. 1801. Systême des Animaux sans Vertèbres, ou Tableau Général des Classes, des Orders et des Genres de ces Animaux. Muséum d’histoire naturelle, Paris.

Lamarck J.B.P.A. 1816. Histoire Naturelle des Animaux sans Vertèbres. J.B. Ballière, Libraire, Paris.

Li Y., Zhan Z. & Xu K. 2020. Morphology and molecular phylogenetic analysis of deep-sea purple gorgonians (Octocorallia: Victorgorgiidae) from seamounts in the tropical western pacific, with description of three new species. Frontiers in Marine Science 7: 701. https://doi.org/10.3389/fmars.2020.00701

Li Y., Zhan Z. & Xu K. 2021. Establishment of Alloptilella splendida gen. et sp. nov. and resurrection of Scytalium veneris (Thomson & Henderson, 1906), two sea pens (Cnidaria: Pennatulacea) from seamounts in the tropical Western Pacific. Journal of Oceanology and Limnology 39 (5): 1790–1804. https://doi.org/10.1007/s00343-021-1083-0

Linnaeus C. 1758. Systema Naturae. Editio Decima, Reformata. Salvus, Holmiae [Stockholm].

López-González P.J. 2020. A new calcaxonian genus and family for Trichogorgia utinomii Cordeiro, 2019 (Octocorallia, Alcyonacea): new recordsof a scleriteless gorgonian species from Antarctica. Marine Biodiversity 50: 96. https://doi.org/10.1007/s12526-020-01109-0

López-González P.J. 2021. Scytalium herklotsi sp. nov. (Anthozoa, Octocorallia, Pennatulacea), the first Atlantic species in the genus Scytalium Herklots, 1858. Marine Biodiversity 51:62. https://doi.org/10.1007/s12526-021-01200-0

López-González P.J. 2022. Molecular phylogeny and morphological comparison of the deep-sea genus Alloptilella Li, Zhan & Xu, 2021 (Octocorallia, Pennatulacea). Marine Biodiversity 52: 41. https://doi.org/10.1007/s12526-022-01260-w

López-González P.J. & Drewery J. 2022. When distant relatives look too alike: a new family, two new genera and a new species of deep-sea Umbellula-like sea pens (Anthozoa, Octocorallia, Pennatulacea). Invertebrate Systematics 36 (3): 199–225. http://doi.org/10.1071/IS21040

Marion A.F. 1906. Étude des Coelentérés atlantiques recueillis par la Commission des dragages de l’aviso Le Travailleur durant les campagnes 1880–1881. Expédition scientifique du Travailleur et du Talisman. Paris, Marson: 103–151.

McFadden C.S. & Ofwegen van L.P. 2013. Molecular phylogenetic evidence supports a new family of octocorals and a new genus of Alcyoniidae (Octocorallia, Alcyonacea). ZooKeys 346: 59–83. https://doi.org/10.3897/zookeys.346.6270

McFadden C.S., France S.C., Sánchez J.A. & Alderslade P.A. 2006. A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa) based on mitochondrial protein-coding sequences. Molecular Phylogenetic and Evolution 41: 513–527. https://doi.org/10.1016/j.ympev.2006.06.010

McFadden C.S., Tullis I.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: 516–526. https://doi.org/10.1007/s10126-002-0102-1

Pante E. & France S.C. 2010. Pseudochrysogorgia bellona n. gen., n. sp.: a new genus and species of chrysogorgiid octocoral (Coelenterata: Anthozoa) from the Coral Sea. Zoosystema 32: 595–612. https://doi.org/10.5252/z2010n4a4

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

Rambaut A., Drummond A.J., Xie D., Baele G. & Suchard M.A. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematics Biology 67: 901–904. https://doi.org/10.1093/sysbio/syy032

Ronquist F. & Huelsenbeck J.P. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574. https://doi.org/10.1093/bioinformatics/btg180

Roule L. 1905. Notice préliminaire sur les pennatulides recueillis par le “Travailleur” et le “Talisman”, dans l’Océan Atlantique, au large du Maroc. Bulletin du Muséum national d’histoire naturelle, Paris 11: 454–458.

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. https://doi.org/10.1007/s00227-003-1018-7

Studer T. 1891. Note préliminaire sur les Alcyonaires provenant des campagnes du yacht l’Hirondelle 1886, 1887, 1888. Part 2. Alcyonacea and Pennatulacea. Mémoires de la Société Zoologique de France 4 (2): 86–95.

Studer T. 1901. Alcyonaires provenant des campagnes de l’Hirondelle (1886–88). Résultats des Campagnes Scientifiques du Prince Albert Ier de Monaco 20: 1–64.

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

Thomson J.A. 1927. Alcyonaires provenant des campagnes scientifiques du Prince Albert 1er de Monaco. Résultats des Campagnes Scientifiques Prince Albert Ier de Monaco 73: 1–77.

Tixier-Durivault A. 1961. Crassophyllum cristatum n. gen. et n. sp., type d’un genre de Pteroeididae (Pennatulacea). Bulletin du Muséum national d’histoire naturelle, Paris (2) 33 (4): 428–433.

Tixier-Durivault A. & d’Hondt M.-J. 1974a. Nouvelles récoltes d‘octocoralliaires a Madagascar. Tethys 5 (2–3) 1973: 251–266.

Tixier-Durivault A. & d‘Hondt M.-J. 1974b. Les octocoralliaires de la campagne Biaçores. Bulletin du Muséum national d‘histoire naturelle, Paris 174: 1361–1433.

Verrill A.E. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expedition, under Commodore C. Ringgold and Captain John Rogers, U.S.N., from 1853 to 1856. Collected by Dr. W. Stimpson, naturalist of the Expedition. With descriptions of some additional species from the west coast of North America. Proceedings of the Essex Institute 4: 181–196.

Williams G.C. 1995a. Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key and synopses. Zoological Journal of the Linnean Society 114: 93–140. https://doi.org/10.1111/j.1096-3642.1995.tb00929.x

Williams G.C. 1995b. The enigmatic sea pen genus Gyrophyllum – a phylogenetic reassessment and description of G. sibogae from Tasmanian waters (Coelenterata: Octocorallia). Proceedings of the California Academy of Sciences 48 (15): 315–128.

Williams G.C. 2011. The Global Diversity of Sea Pens (Cnidaria: Octocorallia: Pennatulacea). PLoS ONE 6: e22747. https://doi.org/10.1371/journal.pone.0022747

Williams G.C, Tracey D. & Mackay E. 2014. Pennatulacea (Sea Pens) Descriptions for the New Zealand Region. New Zealand Aquatic Environment and Biodiversity Report 132: 1–22.

WoRMS. 2022. Pennatulidae Ehrenberg, 1834. Available from: https://www.marinespecies.org/aphia.php?p=taxdetails&id=128484 [accessed 25 Oct. 2022].

How to Cite
López-Gózalez, P. J., Drewery, J., & Williams, G. C. (2022). A new family for the enigmatic sea pen genus Gyrophyllum Studer, 1891 (Octocorallia, Pennatulacea), a molecular and morphological approach. European Journal of Taxonomy, 847(1), 73–103. https://doi.org/10.5852/ejt.2022.847.1983
Research article