Abstract
We provide a guide for identification of Comatulidae, a family of crinoid echinoderms, incorporating morphological and molecular evidence. A non-dichotomous key for all genera is included, as well as photographs of species most likely to be encountered in the tropical western Pacific Ocean. Based on sequencing of cytochrome oxidase subunit I (COI), and other genes when necessary, we identified four cases where taxonomic revision was needed. We synonymized Comaster nobilis under Comaster schlegelii, and Clarkcomanthus exilis under Clarkcomanthus comanthipinnus, and re-described Clarkcomanthus albinotus. We also showed the variation of COI sequences within and among species, which varies from lineage to lineage. In some cases, specimens with obvious morphological disparity possessed very little intraspecific molecular diversity (<1%). In others, specimens with nearly identical external appearances exhibited quite divergent COI sequences (up to 6%). These results, combined with the non-dichotomous key herein, offer guidelines for identification and discussion of existing and new species of Comatulidae.
References
Agassiz, J.L.R. (1836) Prodrome d’une Monographie des Radiaires ou Echinodermes. Mémoires de la Société des Sciences Naturelles de Neuchatel, 1835, 168–199.
Agassiz, A. (1865) Crinoids. In: Agassiz, E.C. & Agassiz, A. (Eds.), Sea-side studies in natural history. Marine Animals of Massachusetts Bay. Radiates. Ticknor & Fields, Boston, pp. 120–122.
Bell, F.J. (1882) An attempt to apply a method of formulation to the species of the Comatulidae; with a description of a new species. Proceedings of the Zoological Society of London, 1882, 530–536.
https://doi.org/10.1111/j.1096-3642.1882.tb02757.xBell, F.J. (1884) Echinodermata. In: Coppinger, R.W. (Ed.), Report on the zoological collections made in the Indo-Pacific Ocean during the voyage of H.M.S. “Alert”, 1881–2. Taylor & Francis, London, pp. 117–177.
Britayev, T.A. & Mekhova, E.S. (2010) Assessment of hidden diversity of crinoids and their symbionts in the Bay of Nhatrang, Vietnam. Marine Biodiversity, 41, 261–285.
Carpenter, P.H. (1879) On the genus Actinometra, Müller, with a morphological account of a new species, A. polymorpha, from the Philippine Islands. Part I. Transactions of the Linnean Society of London, 2, 1–122.
https://doi.org/10.1111/j.1096-3642.1879.tb00318.xCarpenter, P.H. (1881) The Comatulae of the Leyden Museum. Notes from the Leyden Museum, 3, 173–217.
Carpenter, P.H. (1882) Description of new or little known Comatulae. I. On the species of Atelecrinus and Eudiocrinus. II. The Comatulae of the Hamburg Museum. Journal of the Linnean Society of London (Zoology), 16, 487–526.
https://doi.org/10.1111/j.1096-3642.1882.tb02394.xCarpenter, P.H. (1884) Report on the Crinoidea collected during the Voyage of H.M.S. Challenger, during the years 1873–1876.—The Stalked Crinoids. Part XXXII. In: Thomson, C.W. & Murray, J. (Eds.), Report of the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–1876. Zoology—Vol. XI. Neill & Company, Edinburgh, 442 pp.
Carpenter, P.H. (1888) Report on the Crinoidea collected during the Voyage of H.M.S. Challenger, during the years 1873–1876. Part II—The Comatulae. In: Thomson, C.W. & Murray, J. (Eds.), Report of the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–1876, Zoology—Vol. XXVI. Neill & Company, Edinburgh, pp. 1–400.
Chen, J.C., Chang, K.H. & Chen, C.P. (1988) Shallow-water crinoids of Kenting National Park, Taiwan. Bulletin of the Institute of Zoology Academia Sinica, 27, 73–90.
Clark, A.H. (1907a) New genera of Recent free crinoids. Smithsonian Miscellaneous Collection, Quarterly Issue, 50, 343–364.
Clark, A.H. (1907b) Description of new species of recent unstalked crinoids from the coast of northeastern Asia. Proceedings of the United States National Museum, 33, 127–156.
https://doi.org/10.5479/si.00963801.33-1561.127Clark, A.H. (1908a) The genus Comatula Lamarck; with a note on the Encrinus parrae of Guerin. Proceedings of the United States National Museum, 33, 683–688.
https://doi.org/10.5479/si.00963801.33-1585.683Clark, A.H. (1908b) New genera of unstalked crinoids. Proceedings of the Biological Society of Washington, 21, 125–136.
Clark, A.H. (1908c) The nomenclature of the recent crinoids. Proceedings of the United States National Museum, 34, 435–542.
https://doi.org/10.5479/si.00963801.34-1623.435Clark, A.H. (1908d) Preliminary notice of a collection of recent crinoids from the Philippine Islands. Smithsonian Miscellaneous Collections Quarterly, 52, 199–234.
Clark, A.H. (1908e) On a collection of feather stars, or comatulids, from Japan. Proceedings of the United States National Museum, 34, 305–319.
https://doi.org/10.5479/si.00963801.34-1615.305Clark, A.H. (1908f) New genera and species of crinoids. Proceedings of the Biological Society of Washington, 21, 219–232.
Clark, A.H. (1909a) Revision of the crinoid family Comasteridae, with description of new genera and species. Proceedings of the United States National Museum, 36, 493–507.
https://doi.org/10.5479/si.00963801.36-1685.493Clark, A.H. (1911a) A new unstalked crinoid from Christmas Island. Annals and Magazine of Natural History, 8, 644–645.
https://doi.org/10.1080/00222931108692988Clark, A.H. (1909b) New recent Indian crinoids. Proceedings of the Biological Society of Washington, 22, 143–152.
Clark, A.H. (1909c) On a collection of crinoids from the Zoological Museum of Copenhagen. Videnskabelige Meddelelser Fra Dansk Naturhistorisk Forening, 61, 115–195.
Clark, A.H. (1909d) New genera and higher groups of unstalked crinoids. Proceedings of the Biological Society of Washington, 22, 173–178.
Clark, A.H. (1909e) The type of the genus Comaster. Proceedings of the Biological Society of Washington, 22, 87–90.
Clark, A.H. (1909f) On a collection of recent crinoids from the Philippine Islands. Proceedings of the United States National Museum, 36, 391–410.
https://doi.org/10.5479/si.00963801.36-1673.391Clark, A.H. (1909g) Comatilia, a remarkable new genus of unstalked crinoids. Proceedings of the United States National Museum, 36, 361–367.
https://doi.org/10.5479/si.00963801.36-1668.361Clark, A.H. (1911a) Crinoidea. In: Michaelsen, W. & Hartmeyer, R. (Eds.), Die fauna südwest-Australischen forschungsreise 1905—3(13). Gustav Fischer, Jena, pp. 435–467.
Clark, A.H. (1911b) On a collection of unstalked crinoids made by the United States Fisheries Steamer “Albatross” in the vicinity of the Philippine Islands. Proceedings of the United States National Museum, 36, 529–563.
https://doi.org/10.5479/si.00963801.39-1798.529Clark, A.H. (1911c) The recent crinoids of Australia. Australia Museum Memoir, 4, 705–804.
https://doi.org/10.3853/j.0067-1967.4.1911.1511Clark, A.H. (1911d) The recent crinoids of the Leyden Museum. Notes from the Leyden Museum, 33, 175–192.
Clark, A.H. (1911e) A new unstalked crinoid from the Philippine Islands. Proceedings of the United States National Museum, 41, 171–173.
https://doi.org/10.5479/si.00963801.41-1849.171Clark, A.H. (1912a) Seventeen new East Indian crinoids belonging to the families Comasteridae and Zygometridae. Proceedings of the Biological Society of Washington, 25, 17–28.
Clark, A.H. (1912b) The crinoids of the Indian Ocean. Echinoderma of the Indian Museum, Part VII. Trustees of the Indian Museum, Calcutta, 325 pp.
Clark, A.H. (1913a) On a collection of recent crinoids from the waters around Ireland. Department of Agriculture & Technical Instruction for Ireland, 1912, 1–5.
Clark, A.H. (1913b) Notes on the Recent crinoids in the British Museum. Smithsonian Miscellaneous Collections, 61, 1–89.
Clark, A.H. (1915) A monograph of the existing crinoids 1(1). Bulletin of the United States National Museum, 82, 1–406.
https://doi.org/10.5962/bhl.title.1326Clark, A.H. (1916a) Three interesting additions to the crinoid fauna of Sagami Bay and Suruga Gulf, Japan. Proceedings of the Biological Society of Washington, 29, 105–108.
Clark, A.H. (1916b) The first New Zealand crinoid. Proceedings of the Biological Society of Washington, 29, 48.
Clark, A.H. (1918a) The unstalked crinoids of the Siboga Expedition. Siboga Expedition, 42b, 1–300.
Clark, A.H. (1918b) A new crinoid from New Zealand, and another from Tasmania. Proceedings of the Biological Society of Washington, 31, 41–44.
Clark, A.H. (1921a) A monograph of the existing crinoids 1(2). Bulletin of the United States National Museum, 82, 1–795.
Clark, A.H. (1931) A monograph of the existing crinoids 1(3). Bulletin of the United States National Museum, 82, 1–916.
Clark, A.M. (1972) Some crinoids from the Indian Ocean. Bulletin of the British Museum of Natural History, Zoology, 24 (2), 75–156.
Clark, A.M. (1975) The Swain Reefs expedition: Crinoidea. Records of the Australian Museum, 29, 391–406.
https://doi.org/10.3853/j.0067-1975.29.1975.189Clark, A.M. (1980) Crinoidea collected by the Meteor and Discovery in the NE Atlantic. Bulletin of the British Museum of National History (Zoology), 38, 187–210.
Clark, A.M. & Rowe, F.W.E. (1971) Shallow-water Indo-West Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 238 pp.
Clark, H.L. (1915) The comatulids of Torres Strait: with special reference to their habits and reactions. Papers from the Department of Marine Biology Carnegie Institution, 8, 67–125.
Clark, H.L. (1916) Report on the sea-lilies, starfishes, brittle-stars and sea urchins obtained by the F.I.S. “Endeavour” on the coasts of Queensland, New South Wales, Tasmania, Victoria, South Australia, and Western Australia. Biological results of the fishing experiments carried on by the F.I.S. “Endeavour,” 1909–14, 4, 1–123.
Clark, H.L. (1921) The echinoderm fauna of Torres Strait: Its composition and origin. Papers from the Department of Marine Biology of the Carnegie Institution of Washington, 10, 1–223.
Clark, H.L. (1932) Echinodermata (other than Asteroidea) of the Great Barrier Reef Expedition, 1928–1929. Scientific Reports of the Great Barrier Reef Expedition, 4, 197–239.
Clark, H.L. (1946) The echinoderm fauna of Australia. Its composition and its origin. Carnegie Institution of Washington, 566, 1–568.
Deheyn, D.D., Lyskin, S. & Eeckhaut, I. (2006) Assemblages of symbionts in tropical shallow-water crinoids and assessment of symbionts' host-specificity. Symbiosis, 42, 161–168.
Fleming, J. (1828) A history of British animals, exhibiting the descriptive characters and systematical arrangement of the genera and species of quadrupeds, birds, reptiles, fishes, mollusca, and radiata of the United Kingdom; including the indigenous, extirpated, and extinct kinds, together with periodical and occasional visitants. Bell & Bradfute, Edinburgh, 565 pp.
Freycinet, L. (1815) Voyage de découvertes aux Terres Australes, exécuté sure les corvettes le Géographe, le Naturaliste, et la goëlette le Casuarina, pendant les années 1800, 1801, 1802, 1803 et 1804; sous le commandement du Capitaine de vaisseau N. Baudin. Navigation et Géographie. L’Imprimerie Royale, Paris, 578 pp.
Fujita, Y. & Obuchi, M. (2012) Comanthus kumi, a new shallow-water comatulid (Echinodermata: Crinoidea: Comatulida: Comasteridae) from the Ryukyu Islands, Japan. Zootaxa, 3367, 261–268.
Gislén, T. (1922) The crinoids from Dr. S. Bock’s expedition to Japan 1914. Nova Acta Societatis Scientiarum Upsaliensis, 5, 1–183.
https://doi.org/10.5962/bhl.title.11735Hartlaub, C. (1890) Beitrag zur Kenntniss der Comatuliden-Fauna des indischen Archipels. Nachrichten von der Königlichen Gesellschaft der Wissenschaften und der Georgs-August-Universität zu Göttingen, 5, 168–187.
Hartlaub, C. (1891) Beitrag zur Kenntniss der Comatulidenfauna des Indischen Archipels. Nova Acta der Kaiserlische Leopoldinisch-Carolinisch Deutschen Akademie der Naturforscher, 58, 5–120.
https://doi.org/10.5962/bhl.title.11733Hartlaub, C. (1893) Beitrag zur Kenntnis der Comatulidenfauna des Indischen Archipels. Nova Acta Kaiserlichen Leopold-Carol Deutschen Akademie der Naturforschungen, 58, 1–120.
Hemery, L.G., Roux, M., Améziane, N. & Eléaume, M. (2013) High-resolution crinoid phyletic inter-relationships derived from molecular data. Cahiers de Biologie Marine, 54, 511–523.
Hoggett, A.K. & Rowe, F.W.E. (1986) A reappraisal of the family Comasteridae A.H. Clark, 1908 (Echinodermata: Crinoidea), with description of a new subfamily and a new genus. Zoological Journal of the Linnean Society, 88, 103–142.
https://doi.org/10.1111/j.1096-3642.1986.tb01183.xKirkendale, L. & Messing, C.G. (2003) An annotated checklist and key to the Crinoidea of Guam and Commonwealth of the Northern Marianas Islands. Micronesica, 35–36, 523–546.
Kogo, I. (1998) Crinoids from Japan and its adjacent waters. Special Publication of the Osaka Museum of National History, 30, 1–148.
Kogo, I. (2002) Report on the crinoids collected from the Nansei Islands, southern Japan, during a cruise of the training vessel Toyoshio Maru in 1999 (Crinoidea). Bulletin of the Osaka Museum of Natural History, 56, 1–44.
Kogo, I. & Fujita, T. (2014) The Feather Stars of Sagami Bay. Tokai University Press, Toyko, 162 pp.
Lamarck, J.B.P. (1816) Histoire naturelle des animaux sans vertèbres. Vol. 2. Verdière, Paris, 568 pp.
Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Laurentius Salvius, Holmiae, 824 pp.
Lovén, S. (1866) Phanogenia, ett hittills okändt slägte af fria Crinoideer. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 23, 223–233.
Macurda, Jr., D.B. (1975) Bathymetry and zoogeography of shallow-water crinoids in the Bahama Islands. Hydro-lab Journal, 3, 5–24.
Macurda, Jr., D.B. & Meyer, D.L. (1977) Crinoids of West Indian coral reefs. American Association of Petroleum Geologists Studies in Geology, 4, 231–237.
McKnight, D.G. (1977a) Additions to the New Zealand crinoid fauna. New Zealand Oceanographic Institute Records, 3, 93–112.
McKnight, D.G. (1977b) Crinoids from Norfolk Island and Wanganella Bank. New Zealand Oceanographic Institute Records, 3, 129–137.
Messing, C.G. (1978) A revision of the comatulid genus Comactinia A.H. Clark (Crinoidea: Echinodermata). Bulletin of Marine Science, 28, 49–80.
Messing, C.G. (1984) Brooding and paedomorphosis in the deep-water feather star Comatilia iridometriformis (Echinodermata: Crinoidea). Marine Biology, 80, 83–91.
https://doi.org/10.1007/BF00393131Messing, C.G. (1995a) Alloeocomatella, a new genus of reef-dwelling feather star from the tropical Indo-west Pacific (Echinodermata, Crinoidea, Comasteridae). Proceedings of the Biological Society of Washington, 108, 436–450.
Messing, C.G. (1995b) Redescription of a unique feather star (Echinodermata: Crinoidea: Comatulida: Comasteridae) with the diagnosis of a new genus. Proceedings of the Biological Society of Washington, 108, 656–661.
Messing, C.G. (1998a) Revision of the recent Indo-West Pacific comatulid genus Comaster Agassiz - Part 1 - the type species of Comaster and Phanogenia Løven (Echinodermata, Crinoidea, Comasteridae). Invertebrate Taxonomy, 12, 191–209.
https://doi.org/10.1071/IT97004Messing, C.G. (1998b) An initial re-assessment of the distribution and diversity of the East Indian shallow-water crinoid fauna. Pp. 187-192. In: Mooi, R. & Telford, M. (eds.) Echinoderms: San Francisco. Balkema, Rotterdam.
Messing, C.G. (2001) A key to the genera of Comasteridae with a description of a new genus. Bulletin of the Biological Society of Washington, 10, 277–300.
Messing, C.G. (2003) Three new species of Comasteridae (Echinodermata, Crinoidea) from the tropical western Pacific. Zoosystema, 25, 149–162.
Messing, C.G. (2007) The crinoid fauna (Echinodermata: Crinoidea) of Palau. Pacific Science, 61, 91–111.
https://doi.org/10.1353/psc.2007.0010Messing, C.G., Améziane, N. & Eleaume, M. (2000) Echinodermata Crinoidea: Comatulid Crinoids of the KARUBAR Expedition to Indonesia. The families Comasteridae, Asterometridae, Calometridae and Thalassometridae. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 21. Memoires du Museum National d’Histoire Naturelle, 184, 627–702.
Messing C.G. & Dearborn, J. (1990) Marine Flora and Fauna of the northeastern United States. Echinodermata: Crinoidea. NOAA Technical Report, National Marine Fisheries Service No. 91. US Department of Commerce, Springfield, 33 pp.
Messing, C.G., Hoggett, A.K., Vail, L.L., Rouse, G.W. & Rowe, F.W.E. (2017) Class Crinoidea. In: Byrne, M. & O’Hara, T.D. (Eds.), Australian Echinoderms: Biology, Ecology & Evolution. CISIRO Publishing, Canberra, 624 pp.
Messing, C.G., Meyer, D.L., Siebeck, U.E., Jermiin, L.S., Vaney, D.I. & Rouse, G.W. (2006) A modern soft-bottom, shallow-water crinoid fauna (Echinodermata) from the Great Barrier Reef, Australia. Coral Reefs, 25, 164–168.
https://doi.org/10.1007/s00338-005-0076-3Messing C.G., Neumann A.C. & Lang J.C. (1990) Biozonation of deep-water lithoherms and associated hardgrounds in the northeastern Straits of Florida. Palaios, 5, 15–33.
https://doi.org/10.2307/3514994Messing, C.G. & Tay, T.S. (2016) Extant Crinoidea (Echinodermata) of Singapore. Raffles Bulletin of Zoology Supplement, 34, 270–301.
Meyer, D.L. (1973a) Feeding behavior and ecology of shallow-water unstalked crinoids (Echinodermata) in the Caribbean Sea. Marine Biology, 22, 105–129.
https://doi.org/10.1007/BF00391776Meyer, D.L. (1973b) Distribution and living habits of comatulid crinoids near Discovery Bay, Jamaica. Bulletin of Marine Science, 23, 244–259.
Meyer, D.L. (1986) Les crinoïdes. In: Guille, A., Laboute, P. & Menou J.-L. (Eds.), Guide des étoiles de mer, oursins, et autres échinodermes du lagon de Nouvelle-Calédonie, Faune Tropicale. ORSTOM, Paris, pp. 199–225.
Meyer, D.L. & Macurda, Jr., D.B. (1980) Ecology and distribution of the shallow-water crinoids of Palau and Guam. Micronesica, 16, 59–99.
Meyer, D.L., Messing, C.G. & Macurda, Jr., D.B. (1978) Zoogeography of tropical western Atlantic Crinoidea (Echinodermata). Bulletin of Marine Science, 28, 412–441.
Mladenov, P.V. & Brady, K. (1987) Reproductive cycle of the Caribbean feather star Nemaster rubiginosa (Echinodermata: Crinoidea). Marine Ecology, 8, 313–325.
https://doi.org/10.1111/j.1439-0485.1987.tb00191.xMüller, J. (1840) Über den Bau des Pentacrinus Caput Medusae. Archiv für Naturgeschichte, 1, 307–318.
Müller, J. (1841) Über die Gattungen und Arten der Comatulen. Monatsberichte Königlich Preussische Akademie der Wissenschaften zu Berlin, 1841, 179–189.
Müller, J. (1843) Neue Beiträge zur Kenntnis der Arten der Comatulen. Archiv für Naturgeschichte, 9, 131–136.
Müller, J. (1846) Nachtrag zu der Abhandlungen uber die Comatulen. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglichen. Preussische Akademie der Wissenschaften zu Berlin, 1846, 177–179.
Owen, C.L., Messing, C.G., Rouse, G.W. & Shivji, M.S. (2009) Using a combined approach to explain the morphological and ecological diversity in Phanogenia gracilis Hartlaub, 1893 (Echinodermata: Crinoidea) sensu lato: two species or intraspecific variation? Marine Biology, 156, 1517–1529.
https://doi.org/10.1007/s00227-009-1173-6Pilcher, N.J. & Messing, C.G. (2001) A preliminary checklist of the shallow-water crinoids of the Kerama Islands, Okinawa, Japan. Report of Akajima Marine Science Laboratory, 12, 15–17.
Pourtalès, L.F. de (1869) List of the crinoids obtained on the coasts of Florida and Cuba, by the United States Coast Survey Gulf Stream Expeditions, in 1867, 1868, 1869. Bulletin of the Museum of Comparative Zoology at Harvard College, 1, 355–358.
Pourtalès, L.F. de (1878) Reports on the dredging operations of the U.S. Coast Survey Steamer "Blake." Corals and crinoids. Bulletin of the Museum of Comparative Zoology at Harvard College, 5, 213–216.
Rowe, F.W.E. (1974) Catalogue of the Sladen Collection in the Royal Albert Memorial Museum, Exeter, Devon. Biological Journal of the Linnean Society, 6, 179–243.
https://doi.org/10.1111/j.1095-8312.1974.tb00722.xRowe, F.W.E. (1989) Nine new deep-water species of Echinodermata from Norfolk Island and Wanganella Bank, northeastern Tasman Sea, with a checklist of the echinoderm fauna. Proceedings of the Linnean Society of New South Wales, 111, 257–291.
Rowe, F.W.E. & Gates, J. (1995) Echinodermata. Zoological Catalogue of Australia. CSIRO, Melbourne, 510 pp.
Rowe, F.W.E., Hoggett, A.K., Birtles, R.A. & Vail, L.L. (1986) Revision of some comasterid genera from Australia (Echinodermata: Crinoidea), with descriptions of two new genera and nine new species. Zoological Journal of the Linnean Society, 86, 197–277.
https://doi.org/10.1111/j.1096-3642.1986.tb01812.xSchmeltz, J.D.E. (1874) Museum Godeffroy. Catalog V. Nebst einer Beiläge enthaltend topographische und zoologische Notizen. L. Friedrichsen & Co., Hamburg, 215 pp.
Summers, M.M., Messing C.G. & Rouse G.W. (2014a). Phylogeny of Comatulidae (Echinodermata: Crinoidea: Comatulida). Molecular Phylogenetics & Evolution, 80, 319–339.
https://doi.org/10.1016/j.ympev.2014.06.030Summers, M.M., Al-Hakim, I., & Rouse, G.W. (2014b). Turbo-taxonomy: 21 new species of Myzostomida (Annelida). Zootaxa, 3873 (4), 301–344.
https://doi.org/10.11646/zootaxa.3873.4.1Summers, M.M., & Rouse, G.W. (2014). Phylogeny of Myzostomida (Annelida) and their relationships with echinoderm hosts. BMC Evolutionary Biology, 14, 170.
https://doi.org/10.1186/s12862-014-0170-7Tommasi, L.R. (1965) Lista dos crinóides recentes do Brasil. Contribuições avulsas do Instituto Oceanográfico: Oceanografia biológica, 9, 1–33.
Utinomi, H. & Kogo, I. (1965) On some comatulids from the coastal sea of Kii Peninsula. Publications of the Seto Marine Biological Laboratory, 134, 263–286.
White, C.M., Messing, C.G., Shivji, M. & Forstner, M.R.J. (2001) Initial phylogeny of the Comasteridae (Crinoidea) from mtDNA sequences. In: Barker, M. (Ed.) Echinoderms 2000; Proceedings of the tenth International Echinoderm Conference, Dunedin, N.Z., 2000. Balkema, Rotterdam, pp. 135–138.
Zmarzly, D.L. (1984) Distribution and ecology of shallow-water crinoids at Enewetak Atoll, Marshall Islands, with an annotated checklist of their symbionts. Pacific Science, 38, 105–122.
Zmarzly, D.L. (1985) The shallow-water crinoid fauna of Kwajalein Atoll, Marshall Islands: ecological observations, interatoll comparisons, and zoogeographic affinities. Pacific Science, 39, 340–358.