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)
Pablo J. López-González
A * and Jim Drewery B
+ Author Affiliations
- Author Affiliations
A Biodiversidad y Ecología Acuática, Departamento de Zoología, Facultad de Biología, Universidad de Sevilla, Reina Mercedes 6, E-41012 Sevilla, Spain.
B Marine Scotland Science, Marine Laboratory, 375 Victoria Road, Aberdeen, AB11 9DB, Scotland, UK.
Invertebrate Systematics 36(3) 199-225 https://doi.org/10.1071/IS21040
Submitted: 23 May 2021 Accepted: 5 November 2021 Published: 23 February 2022
© 2022 The Author(s) (or their employer(s)). Published by CSIRO Publishing.
Abstract
Among octocorals, colonies of the deep-sea pennatulacean genus Umbellula Gray, 1870 are some of the most instantly recognisable forms. Historically however, species identification in this genus has been usually based on few morphological characters with very little knowledge of associated intraspecific variability. This fact, combined with the very limited access to these deep-sea organisms, has resulted in numerous uncertainties about the true characters that should be used in species determination and recognition of synonyms and questionable species. Recent phylogenetic analyses based on mitochondrial and nuclear DNA markers has shown to be an excellent complementary source of information to morphological examination, being able to detect incongruent taxonomic assignments in classifications based only on morphological characters. Molecular analyses can reveal the presence of paraphyletic or polyphyletic groupings of taxa that may then be the subject of further research integrating morphological and molecular techniques. This paper addresses the existence of a set of specimens initially assigned to the genus Umbellula Gray, 1870 but that have been shown to be distantly related to the type species Umbellula encrinus (Linnaeus, 1758) based on molecular phylogenetic hypotheses. Phylogenetic analyses based on four genetic markers, three mitochondrial (mtMutS, ND2, Cox1) and one nuclear (28S), validate the definition of a new family (Pseudumbellulidae fam. nov.) and two new genera (Pseudumbellula gen. nov. and Solumbellula gen. nov). These analyses also justify the segregation of some of the morphological characters previously included in the diagnosis of the genus Umbellula and the monotypic family Umbellulidae Kölliker, 1880. Moreover, a new species, Pseudumbellula scotiae sp. nov. is described and illustrated with material from the North Eastern Atlantic and compared with congeners. Additionally, the well-known but atypical species Umbellula monocephalus Pasternak, 1964 is transferred and described here as Solumbellula monocephalus (Pasternak, 1964), comb. nov., based on both molecular data and morphology.
Keywords: deep sea, Integrated approach, new taxa, Octocorallia, Pennatulacea, Pseudumbellula gen. nov., Pseudumbellulidae fam. nov., sea pen, Solumbellula gen. nov., taxonomy, Umbellula.
References
Baker, KD, Wareham, VE, Snelgrove, PVR, Haedrich, RL, Fifield, DA, Edinger, EN, and Gilkinson, KD (2012). Distributional patterns of deep-sea coral assemblages in three submarine canyons off Newfoundland, Canada. Marine Ecology Progress Series 445, 235–249.| Distributional patterns of deep-sea coral assemblages in three submarine canyons off Newfoundland, Canada.Crossref | GoogleScholarGoogle Scholar |
Bayer, FM (1993). Taxonomic status of the octocoral genus Bathyalcyon (Alcyoniidae: Anthomastinae), with descriptions of a new subspecies from the Gulf of Mexico and a new species of Anthomastus from Antarctic waters. Precious Corals & Octocoral Research 1, 3–13.
Bayer, FM, and Grasshoff, M (1997). Umbellula Cuvier, [1797] (Cnidaria, Anthozoa): proposed conservation as the correct original spelling, and corrections to the entries relating to Umbellularia Lamarck, 1801 on the Official Lists and Indexes of Names in Zoology. Bulletin of Zoological Nomenclature 54, 14–18.
| Umbellula Cuvier, [1797] (Cnidaria, Anthozoa): proposed conservation as the correct original spelling, and corrections to the entries relating to Umbellularia Lamarck, 1801 on the Official Lists and Indexes of Names in Zoology.Crossref | GoogleScholarGoogle Scholar |
Bayer FM, Grasshoff M, Verseveldt J (1983) ‘Illustrated trilingual glossary of morphological and anatomical terms applied to Octocorallia.’ (Brill/Backhuys: Leiden, Netherlands)
Berntson, EA, France, SC, and Mullineaux, LS (1999). Phylogenetic relationships within the class Anthozoa (phylum Cnidaria) based on nuclear 18S rDNA sequences. Molecular Phylogenetics and Evolution 13, 417–433.
| Phylogenetic relationships within the class Anthozoa (phylum Cnidaria) based on nuclear 18S rDNA sequences.Crossref | GoogleScholarGoogle Scholar | 10603268PubMed |
Berntson, EA, Bayer, FM, McArthur, AG, and France, SC (2001). Phylogenetic relationships within the Octocorallia (Cnidaria: Anthozoa) based on nuclear 18S rRNA sequences. Marine Biology 138, 235–246.
| Phylogenetic relationships within the Octocorallia (Cnidaria: Anthozoa) based on nuclear 18S rRNA sequences.Crossref | GoogleScholarGoogle Scholar |
Bessho-Uehara, M, Francis, WR, and Haddock, SH (2020). Biochemical characterization of diverse deep-sea anthozoan bioluminescence systems. Marine Biology 167, 114.
| Biochemical characterization of diverse deep-sea anthozoan bioluminescence systems.Crossref | GoogleScholarGoogle Scholar |
Billett, DSM, and Rice, AL (2001). The BENGAL programme: introduction and overview. Progress in Oceanography 50, 13–25.
| The BENGAL programme: introduction and overview.Crossref | GoogleScholarGoogle Scholar |
Broch, H (1957). Pennatularians (Umbellula). Reports of the Swedish Deep Sea Expedition 1947–48 2, 348–364.
Broch, H (1958). The pennatularian genus Umbellula. Nature 182, 267–268.
| The pennatularian genus Umbellula.Crossref | GoogleScholarGoogle Scholar |
Brockman, SA, and McFadden, CS (2012). The mitochondrial genome of Paraminabea aldersladei (Cnidaria: Anthozoa: Octocorallia) supports intramolecular recombination as the primary mechanism of gene rearrangement in octocoral mitochondrial genomes. Genome Biology and Evolution 4, 994–1006.
| The mitochondrial genome of Paraminabea aldersladei (Cnidaria: Anthozoa: Octocorallia) supports intramolecular recombination as the primary mechanism of gene rearrangement in octocoral mitochondrial genomes.Crossref | GoogleScholarGoogle Scholar | 22975720PubMed |
Brugler, MR, and France, SC (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.
| 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.Crossref | GoogleScholarGoogle Scholar | 18506498PubMed |
Cordeiro R, McFadden C, van Ofwegen L, Williams GC (2021) World List of Octocorallia. Umbellula Gray, 1870. In ‘World Register of Marine Species’. Available at http://www.marinespecies.org/aphia.php?p=taxdetails&id=128499 [Verified 19 January 2021]
Cuvier G (1798) ‘Tableau élémentaire de l’histoire nauturelle des animaux.’ (Baudouin: Paris, France)
Dolan E (2008) Phylogenetic, systematics and biogeography of deepsea Pennatulacea (Anthozoa: Octocorallia). Evidences from molecules and morphology. PhD thesis, University of Southampton, Southampton, UK. Available at https://eprints.soton.ac.uk/65669/1/Dolan_2008_PhD.pdf
Dolan, JR (2020). The Origins of Oceanography in France: the scientific expeditions of Travailleur and Talisman (1880–1883). Oceanography 33, 126–133.
| The Origins of Oceanography in France: the scientific expeditions of Travailleur and Talisman (1880–1883).Crossref | GoogleScholarGoogle Scholar |
Dolan, E, Tyler, PA, Yesson, C, and Rogers, AD (2013). Phylogeny and systematics of deep-sea sea pens (Anthozoa: Octocorallia: Pennatulacea). Molecular Phylogenetics and Evolution 69, 610–618.
| Phylogeny and systematics of deep-sea sea pens (Anthozoa: Octocorallia: Pennatulacea).Crossref | GoogleScholarGoogle Scholar | 23906600PubMed |
Felsenstein, J (1985). Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39, 783–791.
| Confidence limits on phylogenies: an approach using the bootstrap.Crossref | GoogleScholarGoogle Scholar | 28561359PubMed |
Filhol H (1885) ‘La vie au fond des mers: Les explorations sous-marines et les voyages du Travailleur et du Talisman.’ (Masson: Paris, France)
France, SC, and Hoover, LL (2002). DNA sequences of the mitochondrial COI gene have low levels of divergence among deep-sea octocorals (Cnidaria: Anthozoa). Hydrobiologia 471, 149–155.
| DNA sequences of the mitochondrial COI gene have low levels of divergence among deep-sea octocorals (Cnidaria: Anthozoa).Crossref | GoogleScholarGoogle Scholar |
García-Cárdenas, FJ, Drewery, J, and López-González, PJ (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, 261–276.
| Resurrection of the sea pen genus Ptilella Gray, 1870 and description of Ptilella grayi n. sp. from the NE Atlantic (Octocorallia, Pennatulacea).Crossref | GoogleScholarGoogle Scholar |
García-Cárdenas, FJ, Núñez-Flores, M, and López-González, PJ (2020). Molecular phylogeny and divergence time estimates in pennatulaceans (Cnidaria: Octocorallia: Pennatulacea). Scientia Marina 84, 317–330.
| Molecular phylogeny and divergence time estimates in pennatulaceans (Cnidaria: Octocorallia: Pennatulacea).Crossref | GoogleScholarGoogle Scholar |
Grasshoff, M (1972). Eine Seefeder mit einem einzigen Polypen: Umbellula thieli n. sp. Die von F.S. ‘Meteor’ 1967–1970 im östlichen Nordatlantik gedredschten Pennatularia (Cnidaria: Anthozoa). Meteor Forschungsergebnisse – Reihe D 12, 1–11.
Grasshoff, M (1982a). Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der Biskaya (Cnidaria, Anthozoa). Ergebnisse der französischen Expeditionen Biogas, Polygas, Geomanche, Incal, Noratlante und Fahrten der ‘Thalassa’. I. Allgemeiner Teil. Bulletin du Muséum National d’Histoire Naturelle, Section A 3, 731–766.
Grasshoff, M (1982b). Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der Biskaya (Cnidaria, Anthozoa). Ergebnisse der französischen Expeditionen Biogas, Polygas, Geomanche, Incal, Nortatlante und Fahrten der ‘Thalassa’. II. Taxonomischer Teil. Bulletin du Muséum National d’Histoire Naturelle, Section A 3, 941–978.
Gray JE (1870) ‘Catalogue of sea-pens or Pennatulariidae in the collection of the British Museum.’ (British Museum: London, UK)
Haeckel E (1866) ‘Generelle Morphologie der organismen.’ (Georg Reimer: Berlin, German Empire)
Hickson, SJ (1916). The Pennatulacea of the Siboga expedition, with a general survey of the order. Siboga Expeditie Monographs 14, 1–265.
Hogan, RI, Hopkins, K, Wheeler, AJ, Allcock, AL, and Yesson, C (2019). Novel diversity in mitochondrial genomes of deep-sea Pennatulacea (Cnidaria: Anthozoa: Octocorallia). Mitochondrial DNA – A 30, 764–777.
| Novel diversity in mitochondrial genomes of deep-sea Pennatulacea (Cnidaria: Anthozoa: Octocorallia).Crossref | GoogleScholarGoogle Scholar |
Huelsenbeck, JP, and Ronquist, F (2001). MrBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17, 754–755.
| MrBAYES: Bayesian inference of phylogenetic trees.Crossref | GoogleScholarGoogle Scholar | 11524383PubMed |
(1944). Opinion 185. Suppression of Bohadsch (J. B.) De quibusdam animalibus marinis, 1761 and of the German translation thereof published by Leske (N. G.) in 1776. Opinions and Declarations Rendered by the International Commission on Zoological Nomenclature 3, 37–52.
Kimura, M (1980). A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide-sequences. Journal of Molecular Evolution 16, 111–120.
| A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide-sequences.Crossref | GoogleScholarGoogle Scholar | 7463489PubMed |
Kükenthal, W (1902). Diagnosen neuer Umbelluliden aus der Ausbeuter der deutschen Tiefsee-Expedition. Zoologischer Anzeiger 25, 593–597.
Kükenthal W (1915) ‘Pennatularia. Das Tierreich.’ (Verlag von R. Friedländer und Sohn: Berlin, German Empire)
Kushida, Y, and Reimer, JD (2019). Molecular phylogeny and diversity of sea pens (Cnidaria: Octocorallia: Pennatulacea) with a focus on shallow water species of the northwestern Pacific Ocean. Molecular Phylogenetics and Evolution 131, 233–244.
| Molecular phylogeny and diversity of sea pens (Cnidaria: Octocorallia: Pennatulacea) with a focus on shallow water species of the northwestern Pacific Ocean.Crossref | GoogleScholarGoogle Scholar | 30471843PubMed |
Kushida, Y, and Reimer, JD (2020). Description of the sea pen Calibelemnon hinoenma sp. nov. from shallow waters in southern Japan. Marine Biodiversity 50, 107.
| Description of the sea pen Calibelemnon hinoenma sp. nov. from shallow waters in southern Japan.Crossref | GoogleScholarGoogle Scholar |
Linnaeus C (1758) Systema naturae. Editio decima, reformata, Holmiae.
López-González, PJ (2020). A new calcaxonian genus and family for Trichogorgia utinomii Cordeiro, 2019 (Octocorallia, Alcyonacea): new records of a scleriteless gorgonian species from Antarctica. Marine Biodiversity 50, 96.
| A new calcaxonian genus and family for Trichogorgia utinomii Cordeiro, 2019 (Octocorallia, Alcyonacea): new records of a scleriteless gorgonian species from Antarctica.Crossref | GoogleScholarGoogle Scholar |
López-González, PJ, and Williams, GC (2011). A new deep-sea pennatulacean (Anthozoa: Octocorallia: Chunellidae) from the Porcupine Abyssal Plain (NE Atlantic). Helgoland Marine Research 65, 309–318.
| A new deep-sea pennatulacean (Anthozoa: Octocorallia: Chunellidae) from the Porcupine Abyssal Plain (NE Atlantic).Crossref | GoogleScholarGoogle Scholar |
Manuel RL (1981) ‘British Anthozoa. Synopses of the British Fauna (New Series). Number 18.’ (Linnean Society of London: London, UK)
McFadden, CS, and van Ofwegen, LP (2013). Molecular phylogenetic evidence supports a new family of octocorals and a new genus of Alcyoniidae (Octocorallia, Alcyonacea). ZooKeys 346, 59–83.
| Molecular phylogenetic evidence supports a new family of octocorals and a new genus of Alcyoniidae (Octocorallia, Alcyonacea).Crossref | GoogleScholarGoogle Scholar |
McFadden, CS, Tullis, ID, Hutchinson, MB, Winner, K, and Sohm, JA (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.
| Variation in coding (NADH dehydrogenase subunits 2, 3, and 6) and noncoding intergenic spacer regions of the mitochondrial genome in Octocorallia (Cnidaria: Anthozoa).Crossref | GoogleScholarGoogle Scholar | 15723254PubMed |
McFadden, CS, France, SC, Sánchez, JA, and Alderslade, PA (2006). A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa) based on mitochondrial protein-coding sequences. Molecular Phylogenetics and Evolution 41, 513–527.
| A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa) based on mitochondrial protein-coding sequences.Crossref | GoogleScholarGoogle Scholar | 16876445PubMed |
McFadden, CS, Brown, AS, Brayton, C, and van Ofwegen, LP (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, 275–286.
| Application of DNA barcoding in biodiversity studies of shallow-water octocorals: molecular proxies agree with morphological estimates of species richness in Palau.Crossref | GoogleScholarGoogle Scholar |
Nutting, CC (1912). Descriptions of the Alcyonaria collected by the U.S. Bureau of Fisheries steamer ‘Albatross’, mainly in Japanese waters, during 1906. Proceedings of the United States National Museum 43, 1–104.
| Descriptions of the Alcyonaria collected by the U.S. Bureau of Fisheries steamer ‘Albatross’, mainly in Japanese waters, during 1906.Crossref | GoogleScholarGoogle Scholar |
Pante, E, and France, SC (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.
| Pseudochrysogorgia bellona n. gen. n. sp.: a new genus and species of chrysogorgiid octocoral (Coelenterata: Anthozoa) from the Coral Sea.Crossref | GoogleScholarGoogle Scholar |
Pante, E, France, SC, Couloux, A, Cruaud, C, McFadden, CS, Samadi, S, and Watling, L (2012). Deep-sea origin and in-situ diversification of chrysogorgiid octocorals. PLoS One 7, e38357.
| Deep-sea origin and in-situ diversification of chrysogorgiid octocorals.Crossref | GoogleScholarGoogle Scholar | 22723855PubMed |
Pante, E, Saucier, EH, and France, SC (2013). Molecular and morphological data support reclassification of the octocoral genus Isidoides. Invertebrate Systematics 27, 365–378.
| Molecular and morphological data support reclassification of the octocoral genus Isidoides.Crossref | GoogleScholarGoogle Scholar |
Pasternak, FA (1962). Pennatularia of the genus Umbellula Cuvier (Coelenterata, Octocorallia) from Antarctic and Subantarctic Waters. Biol Rep Soviet Antarctic Expedition (1955–1958) 1, 107–130.
Pasternak, FA (1964). The deep-sea pennatularians and antipatharians obtained by R/S ‘Vitjaz’ in the Indian Ocean and the resemblance between the faunas of the pennatularians of the Indian Ocean and the Pacific. Trudy Instituta okeanologii, Akademiya nauk SSSR 69, 183–215.
Pasternak, FA (1970). Sea pens (Octocorallia, Pennatularia) of the hadal zone of the Kurile-Kamtaschatka Trench. Trudy Instituta okeanologii, Akademiya nauk SSSR 86, 236–248.
Pasternak, FA (1975). Deep-sea pennatularian genus Umbellula from the Caribbean Sea and Puerto-Rican Trench. In: Scientific studies Caribbean Sea, Gulf of Mexico and adjacent waters. Transactions of the P.P. Shirsov Institute of Oceanology 100, 160–173.
Pasternak, FA (1993). Pennatularians, gorgonians and antipatharians collected in 43 cruises of R/V “Dimitri Mendelev” in Argentine Basin, Orkney Trench and in the western part of African-Antarctic Basin. Trudy Instituta okeanologii, Akademii nauk SSSR 127, 82–88.
Pérez, CD, Cordeiro, RTS, Williams, GC, and Gomes, PB (2021). Revised nomenclature of the sea pen genus Balticina Gray, 1870 (= Halipteris Kölliker, 1870) (Anthozoa: Octocorallia). Zootaxa 4966, 237–244.
| Revised nomenclature of the sea pen genus Balticina Gray, 1870 (= Halipteris Kölliker, 1870) (Anthozoa: Octocorallia).Crossref | GoogleScholarGoogle Scholar |
Rambaut, A, Drummond, AJ, Xie, D, Baele, G, and Suchard, MA (2018). Posterior summarization in bayesian phylogenetics using Tracer 1.7. Systematic Biology 67, 901–904.
| Posterior summarization in bayesian phylogenetics using Tracer 1.7.Crossref | GoogleScholarGoogle Scholar | 29718447PubMed |
Risaro, J, Williams, GC, Pereyra, D, and Lauretta, D (2020). Umbellula pomona sp. nov., a new sea pen from Mar del Plata Submarine Canyon (Cnidaria: Octocorallia: Pennatulacea). European Journal of Taxonomy 720, 121–143.
| Umbellula pomona sp. nov., a new sea pen from Mar del Plata Submarine Canyon (Cnidaria: Octocorallia: Pennatulacea).Crossref | GoogleScholarGoogle Scholar |
Ronquist, F, and Huelsenbeck, JP (2003). MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574.
| MrBayes 3: Bayesian phylogenetic inference under mixed models.Crossref | GoogleScholarGoogle Scholar | 12912839PubMed |
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 11, 454–458.
Sánchez, JA, McFadden, CS, France, SC, and Lasker, HR (2003). Molecular phylogenetic analyses of shallow-water Caribbean octocorals. Marine Biology 142, 975–987.
| Molecular phylogenetic analyses of shallow-water Caribbean octocorals.Crossref | GoogleScholarGoogle Scholar |
Tamura, K (1992). Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G+C-content biases. Molecular Biology and Evolution 9, 678–687.
| 1630306PubMed |
Tamura, K, Stecher, G, Peterson, D, Filipski, A, and Kumar, S (2013). MEGA6: molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution 30, 2725–2729.
| MEGA6: molecular evolutionary genetics analysis version 6.0.Crossref | GoogleScholarGoogle Scholar | 24132122PubMed |
Thomson, JS (1915). The Pennatulaceae of the Cape of Good Hope and Natal. Memoirs and Proceedings of the Manchester Literary & Philosophical Society. Manchester Literary and Philosophical Society 59, 1–26.
Thomson JA, Henderson WD (1906) ‘An account of the alcyonarians collected by the Royal Indian Marine Survey Ship Investigator in the Indian Ocean. Part I. The alcyonarians of the deep sea.’ (Trustees of the Indian Museum: Calcutta, India)
Tiefenbacher, L (2001). Umbellula monocephalus Pasternak, 1964, eine seltene Pennatularia aus dem südlichen Westeuropäischen Becken (Anthozoa, Octocorallia, Pennatularia). Spixiana 24, 1–4.
Tyler, PA (2002). Deep-sea eukaryote ecology of the semi-isolated basins off Japan. Journal of Oceanography 58, 333–341.
| Deep-sea eukaryote ecology of the semi-isolated basins off Japan.Crossref | GoogleScholarGoogle Scholar |
Tyler, PA, Bronsdon, SK, Young, CM, and Rice, AL (1995). Ecology and gametogenic biology of the genus Umbellula (Pennatulacea) in the North Atlantic Ocean. International Review of Hydrobiology 80, 187–199.
| Ecology and gametogenic biology of the genus Umbellula (Pennatulacea) in the North Atlantic Ocean.Crossref | GoogleScholarGoogle Scholar |
Uda, K, Komeda, Y, Koyama, H, Koga, K, Fujita, T, Iwasaki, N, and Suzuki, T (2011). Complete mitochondrial genomes of two Japanese precious corals, Paracorallium japonicum and Corallium konojoi (Cnidaria, Octocorallia, Coralliidae): notable differences in gene arrangement. Gene 476, 27–37.
| Complete mitochondrial genomes of two Japanese precious corals, Paracorallium japonicum and Corallium konojoi (Cnidaria, Octocorallia, Coralliidae): notable differences in gene arrangement.Crossref | GoogleScholarGoogle Scholar | 21310221PubMed |
Verrill, AE (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.
Versluys, J (1906). Bathyalcyon robustum nov. gen. nov. spec. Ein neuer Alcyonarier der Siboga-Sammlung. Zoologischer Anzeiger 30, 549–553.
von Kölliker, RA (1869). Anatomisch-Systematische Beschreibung der Alcyonararien. I. Die Pennatuliden. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 7–8, 1–458.
von Kölliker, RA (1874). Über den Bau und die systematische Stellung der Gattung Umbellularia; eine vorläufige Mittheilung. Verhandlungen der Physikalisch-medizinischen Gesellschaft zu Würzburg 8, 13–18.
von Kölliker, RA (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, 1–41.
Williams, GC (1990). The Pennatulacea of southern Africa (Coelenterata, Anthozoa). Annals of the South African Museum 99, 31–119.
Williams, GC (1995). Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key and synopses. Zoological Journal of the Linnean Society 113, 93–140.
| Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key and synopses.Crossref | GoogleScholarGoogle Scholar |
Williams, GC, and Alderslade, P (2011). Three new species of pennatulacean octocorals with the ability to attach to rocky substrata (Cnidaria: Anthozoa: Pennatulacea). Zootaxa 3001, 33–48.
| Three new species of pennatulacean octocorals with the ability to attach to rocky substrata (Cnidaria: Anthozoa: Pennatulacea).Crossref | GoogleScholarGoogle Scholar |
