Abstract
Between 2015 and 2018, the NOAA ship Okeanos Explorer embarked on the CAPSTONE expedition, that collected video and specimen data from the Hawaiian Islands region and other American territories in the Central and South Pacific Ocean. Ultimately 20 species, 12 of which are newly described, from 14 genera in 3 families are described herein with one further undescribed species of Pythonaster currently in preparation. Okeanos Explorer collected a total of 17 sea star specimens, of which 9 represent new species and 4 new genera. Additionally, one species imaged by Okeanos Explorer during the campaign was discovered in a museum collection is described herein. Supplementary collections from the tropical Pacific by the R/V Nautilus resulted in a further 11th newly described species which had been originally observed by Okeanos Explorer. A total of 21 specimens were involved as part of this effort. Heligmaster n. gen. is the first new genus of the rarely encountered deep-sea Myxasteridae since Fisher (1906). The Goniasteridae, particularly the corallivorous subfamily Hippasterinae were most heavily represented in the collected material. Video imagery was extensive and provided new occurrence data for rarely seen species as well as several new records that likely represent undescribed species. Many of the new species collected were observed preying on of deep-sea octocorals and sponges.
References
Acloque, A. (1900) Faune de France contenant la description de toutes les espèces indigènes-avec un preface par E. Perrier. Bailliere, Paris, 500 pp. [1899]
Alcock, A. (1893) Natural History notes from the HM Indian Marine Survey Steamer Investigator. 7. An account of the collection of deep-sea Asteroidea. Annals of the Magazine of Natural History, Series 6, 11 (6), 73–121. https://doi.org/10.1080/00222939308677478
Alton, M.S. (1966) A new sea-star from the northeastern Pacific Ocean, Asthenactis fisheri n. sp., with a review of the family Myxasteridae. Deep-Sea Research, 13 (4), 687–697. https://doi.org/10.1016/0011-7471(66)90600-0
Asbjornsen, P.C. (1856) Brisinga endecacnemos. In: Sars, G.O., Koren, J. & Danielssen, D.C., Fauna Littoralis Norvegiae, 2, pp. 101.
Aziz, A. & Jangoux, M. (1985) On the status and affinities of the goniasterid genus Calliaster Gray, 1840. In: Keegan, B.F. & O’Connor, B.D.S. (Eds.), Proceedings of the 5th International Echinoderm Conference, Galway, 1984. Balkema, Rotterdam, pp 585–586.
Bernasconi, I. (1964) Asteroideos argentinos. Claves para los ordenes, familias, subfamilies y géneros. Physis Buenos Aires, 24, 241–277.
Bernasconi, I. (1970) Equinodermos antárticos. 2. Asteroideos. 3. asteroides de la extremidad norte de la Peninsula Antartica. Revista del Museo Argentino de Ciencias Naturales e Instituto Nacional de Investigation de las Ciencias naturales, 9, 211–281, 20 pls., 3 figs.
Birkeland, C. & Lucas, J.S. (1990) Acanthaster planci: Major Management Problem of Coral Reefs. CRC Press, Boca Raton, Florida, 257 pp.
Blake D.B. (1978) The taxonomic position of the modern sea-star Cistina Gray, 1840. Proceedings of the Biological Society of Washington, 91 (1), 234–241.
Bo, M., Canese, S. & Bavestrello, G. (2018) On the coral-feeding habit of the sea star Peltaster placenta. Marine Biodiversity, 49, 2009–2012. https://doi.org/10.1007/s12526-018-0931-4
Breton, G. (1992) Les goniasteridae (Asteroidea, Echinodermata) Jurassiques et Cretaces de France: taphonomie, systematique, biostratigraphie, paleobiogeographie, evolution. Thése de Doctorat d’ Etat Sciences, Editions du Museum du Havre, Université de Caen, Caen, 588 pp.
Cairns, S.D. & Bayer, F.M. (2008) A Review of the Octocorallia (Cnidaria: Anthozoa) from Hawai‘i and Adjacent Seamounts: The Genus Narella. Pacific Science, 62 (1), 83–115. https://doi.org/10.2984/1534-6188(2008)62[83:AROTOC]2.0.CO;2
Chave, E.H. & Malahoff, A. (1998) In Deeper Waters: photographic studies of Hawaiian deep-sea habitats and life forms. University of Hawaii Press, Honolulu, 127 pp.
Clark, A.M. (1993) An index of names of recent Asteroidea—Part 2: Valvatida. Echinoderm Studies, 4,187–366.
Clark, A.M. (1996) An index of names of recent Asteroidea—Part 3: Velatida and Spinulosida. Echinoderm Studies, 5, 183–250.
Clark, A.M. & Courtman-Stock, J. (1976) The Echinoderms of Southern Africa. British Museum of Natural History, London, 277 pp.
Clark, A.M. & Downey, M.E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp.
Danielssen, D.C. & Koren, J. (1882) Fra den norske Nordhavsexpedition, Echinodermer. Nyt Magazin for Naturvidenskaberne, 27, 267–299.
Djakonov, A.M. (1950) s.n. Morskie Zvezdy Morei SSSR Tableauxanalytiques de la faune de U.R.S.S., 34, 1–203. [translated as Dyakonov. A.M. 1968. Sea stars (Asteroids) of the USSR Seas. Keys to the Fauna of the USSR 34. Zoological Institute of the Academy of Sciences of the USSR (Ed.) By Strelkov, A.A., Israel Program for scientific translations Ltd., Jerusalem, 183 pp.]
Downey, M.E. (1986) Revision of the Atlantic Brisingida (Echinodermata:Asteroidea), with description of a new genus and family. Smithsonian Contributions to Zoology, 435, 1–57.
Emson, R.H. & Young, C.M. (1994) Feeding mechanism of the brisingid starfish Novodinia antillensis. Marine Biology, 118, 433–442. https://doi.org/10.1007/BF00350300
Fisher, W.K. (1906) The starfishes of the Hawaiian islands. Bulletin of the United States Fish Commission, 23, 987–1130.
Fisher, W.K. (1911a) Hyalinothrix, a new genus of starfishes from the Hawaiian Islands. Proceedings of the US National Museum, 39, 659-664.
Fisher, W.K. (1911b) Asteroidea of the North Pacific and adjacent waters. 1. Phanerozonia and Spinulosida. Bulletin of the US National Museum, 76, xiii + 1–420.
Fisher, W.K. (1913) New starfishes from the Philippine Islands, Celebes, and the Moluccas. Proceedings of the Untied States National Museum, 46, 201–224.
Fisher, W.K. (1919) Starfishes of the Philippine seas and adjacent waters. Bulletin of the United States National Museum, 3 (100), 1–547, 156 pls.
Fisher, W.K. (1925) Sea stars of the tropical Central Pacific. Bulletin of the Bernice P. Bishop Museum, 10 (1), 1–9.
Fisher, W.K. (1928) Asteroidea of the North Pacific and adjacent waters, Part 2: Forcipulata (Part). Bulletin of the U. S. National Museum, 76, 1–245.
Forbes, E. (1839) On the Asteriadae of the Irish Sea. Memoirs of the Wernerian Natural History Society of Edinburgh, 8, 114–129.
Gray, J.E. (1840) XXII. A synopsis of the genera and species of the class Hypostoma (Asterias, Linnaeus). Annals and Magazine of Natural History, Series 1, 6 (36), 175–184. https://doi.org/10.1080/03745484009443282
Halpern, J.A. (1969) Biological Investigations of the deep sea. 46. The genus Litonotaster (Echinodermata, Asteroidea). Proceedings of the Biological Society of Washington, 82, 129–142.
Halpern, J.A. (1970) Goniasteridae (Echinodermata: Asteroidea) of the Straits of Florida. Bulletin of Marine Science, 20 (1), 193–286.
Hayashi, R. (1939) Solasterids in Japanese waters. Journal of the Faculty of Science of Hokkaido University, 6 (3), 297–311.
Hayashi, R. (1940) Contributions to the Classification of the sea-stars of Japan. I. Spinulosa. Journal of the Faculty of Imperial Science of Hokkaido University, 6 (7), 107–204.
Howell, K.L., Rogers, A.D., Tyler, P.A. & Billett, D.S.M. (2004) Reproductive isolation among morphotypes of the Atlantic sea star species Zoroaster fulgens (Asteroidea: Echinodermata). Marine Biology, 144, 977–984. https://doi.org/10.1007/s00227-003-1248-8
Jangoux, M. (1982) Food and feeding mechanisms: Asteroidea. In: Jangoux, M. & Lawrence, J. (Eds.), Echinoderm Nutrition Balkema, Rotterdam. pp. 117–159.
Kelley, C., Mah, C. Malik, M. & Elliott, K. (2018) Laulima O Ka Moana: Exploring deep
monuments around Johnston Atoll. New frontiers in ocean exploration: the E/V Nautilus, NOAA ship Okeanos Explorer and R/V Falkor 2017 field season. Raineault et al. eds. Oceanography, 31 (1), 80–81. https://doi.org/10.5670/oceanog.2018.supplement.01
Kennedy, B.R., Cantrell, K., Malik, M., Kelly, C., Potter, J., Elliott, K., Lobecker, E., McKenna, Gray, L., Sowers, D., White, M., France, S., Auscavitch, S., Mah, C., Moriwake, V., Bingo, S., Putts, M. & Rotjan, R.D. (2019) The unknown and the unexplored: insights into the Pacific deep-sea following NOAA CAPSTONE expeditions. Frontiers in Marine Science. [published online] https://doi.org/10.3389/fmars.2019.00480
Koehler, R. (1909) An account of the deep-sea Asteroidea collected by the Royal Indian Marine Survey Ship Investigator. Echinoderma of the Indian Museum, Part 5, 115–131.
Krieger, K.J. & Wing, B.L., (2002) Megafauna associations with deepwater corals (Primnoa spp.) in the Gulf of Alaska. Hydrobiologia, 471, 83–90.
Linchangco, G.V., Foltz, D.W., Reid, R., Williams, J., Nodzak, C., Kerr, A.M., Miller, A.K., Hunter, R., Wilson, N.G., Nielsen, W.J., Mah, C.L., Rouse, G.W., Wray, G.A. & Janies, D.A. (2017) The phylogeny of extant starfish (Asteroidea: Echinodermta) including Xyloplax based on comparative transcriptomics. Molecular Phylogenetics and Evolution, 115, 161–170. https://doi.org/10.1016/j.ympev.2017.07.022
de Loriol, P. (1883) Catalogue Raisonne des Echinodermes recueillis par M.V. de Robillard a l’iIle Maurice. Memoires de la Société de physique et d’histoire naturelle de Genève, 28 (8), 1–64.
Ludwig, H. (1905) Asteroidea. Memoirs of the Museum of Comparative Zoology at Harvard, 32, vii–xii + 1–292, pls.
Madsen, F.J. (1956) Echinoidea, Asteroidea, and Ophiuroidea from depths exceeding 6000 m. Galathea Reports, 2, 23–32.
Mah, C. (1998) Preliminary phylogeny and taxonomic revision of the Brisingida (Asteroidea: Forcipulatacea). In: Mooi, R. & Telford, M. (Eds.), Proceedings of the 9th International Echinoderm Conference. Ballkema, San Francisco, California, pp. 273–277.
Mah, C. (1999) Re-description and taxonomic notes on the South Pacific brisingidan Brisingaster robillardi (Asteroidea) with new ontogenetic and phylogenetic information. Zoosystema, 21 (3), 535–546.
Mah, C.L. (2021) World Asteroidea Database. Brisingidae G.O. Sars, 1875. Accessed from: http://www.marinespecies.org/asteroidea/aphia.php?p=taxdetails&id=123119 (accessed 10 March 2022)
Mah, C.L. (2006) Phylogeny and biogeography of the deep-sea goniasterid, Circeaster (Echinodermata: Asteroidea) including descriptions of six new species. Zoosystema, 28 (4), 917–954.
Mah, C.L. (2015a) A new Atlantic species of Evoplosoma with taxonomic summary and in situ observations of Atlantic deep-sea corallivorous Goniasteridae (Valvatida; Asteroidea) Marine Biodiversity Records, 8 (e5), 1–8. https://doi.org/10.1017/S1755267214001407
Mah, C.L. (2015b) New species, corallivory, in situ video observations and overview of the Goniasteridae (Valvatida, Asteroidea) in the Hawaiian Region. Zootaxa, 3926 (2), 211–228. https://doi.org/10.11646/zootaxa.3926.2.3
Mah, C.L. (2016a) Deep-sea (>1000 m) Goniasteridae (Valvatida; Asteroidea) from the North Pacific, including an overview of Sibogaster, Bathyceramaster n. gen. and three new species. Zootaxa, 4175 (2), 101–141. https://doi.org/10.11646/zootaxa.4175.2.1
Mah, C.L. (2016b) A new species of Brisingenes from the Hawaii Undersea Military Munitions Assessment Area with an overview of Hawaiian brisingid in situ video observations and functional morphology of subambulacral spines (Forcipulatacea; Asteroidea). Deep-Sea Research, Part II, 128, 43–52. https://doi.org/10.1016/j.dsr2.2014.06.003
Mah, C.L. (2018) New genera, species and occurrence records of Goniasteridae (Asteroidea; Echinodermata) from the Indian Ocean. Zootaxa, 4539 (1), 1–116. https://doi.org/10.11646/zootaxa.4539.1.1
Mah, C.L. (2020) New Species, Occurrence Records and Observations of Predation by Deep-sea Asteroidea (Echinodermata) from the North Atlantic by NOAA Ship Okeanos Explorer. Zootaxa, 4766 (2), 201–260. https://doi.org/10.11646/zootaxa.4766.2.1
Mah, C. & Blake, D.B. (2012) Global Diversity and Phylogeny of the Asteroidea (Echinodermata). PLoS ONE, 7 (4), e35644. https://doi.org/10.1371/journal.pone.0035644
Mah, C.L. & Foltz, D.W. (2011) Molecular Phylogeny of the Valvatacea (Asteroidea, Echinodermata). Zoological Journal of the Linnean Society, 161, 769–788. https://doi.org/10.1111/j.1096-3642.2010.00659.x
Mah, C. & Fujita, T. (2020) New species and occurrence records of Japanese Solasteridae and Ganeriidae including a new species of Paralophaster from the North Pacific with an overview of Hyalinothrix. Zootaxa, 4750 (1), 67–100. https://doi.org/10.11646/zootaxa.4750.1.4
Mah, C.L., Mercier, A., Hamel, J.-F. & Nizinski, M. (2012) Range Extensions and Taxonomic Note on Atlantic Myxasteridae) Velatida; Asteroidea, Zootaxa, 3572 (1), 55–62. https://doi.org/10.11646/zootaxa.3572.1.7
Mah, C.L., Neill, K, Eleaume, M. & Foltz, D. (2014) New Species and global revision of Hippasteria (Hippasterinae: Goniasteridae; Asteroidea; Echinodermata). Zoological Journal of the Linnean Society, 171, 422–456. https://doi.org/10.1111/zoj.12131
Mah, C.L., Nizinski, M. & Lundsten, L. (2010) Phylogenetic Revision of the Hippasterinae (Goniasteridae; Asteroidea): Systematics of Deep Sea Corallivores, including one new genus and three new species. Zoological Journal of the Linnean Society, 160, 266–301. https://doi.org/10.1111/j.1096-3642.2010.00638.x
Mauzey, K.P., Birkeland, C. & Dayton, P.K. (1968) Feeding behavior of asteroids and escape responses of their prey in the Puget Sound region. Ecology, 49, 603–619. https://doi.org/10.2307/1935526
McKnight, D.G. (1973) Additions to the asteroid fauna of New Zealand: Family Goniasteridae. NZOI Records, 1 (13), 171–195.
Meadows, P.S., Meadows, A. & Murray, J.M.H. (2012) Biological modifiers of marine benthic seascapes: Their role as ecosystem engineers. Geomorphology, 157–158, 31–48. https://doi.org/10.1016/j.geomorph.2011.07.007
Menge, B.A. (1972) Competition for food between two intertidal starfish species and its effect on body size and feeding. Ecology, 53, 635–644. https://doi.org/10.2307/1934777
Mooi, R. & David, B. (2000) What a new model of skeletal homologies tells us about asteroid evolution. American Zoologist, 40 (3), 326–339.
https://doi.org/10.1093/icb/40.3.326
Paine, R.T. (1966) Food web complexity and species diversity. American Naturalist, 100, 65–75. https://doi.org/10.1086/282400
Paine, R.T. (1969) The Pisaster-Tegula interaction: prey patches, predator food preference and intertidal community structure. Ecology, 50, 950–961. https://doi.org/10.2307/1936888
Perrier, E. (1894). Stellérides. Expéditions Scientifique Travailleur et du Talisman, 3, 1–431, 26 pls.
Quattrini, A.M. & Demopoulos, A.W.J. (2016) Ectoparasitism on deep-sea fishes in the western North Atlantic: In situ observations from ROV surveys. International Journal for Parasitology: Parasites and Wildlife, 5 (3), 217–228. https://doi.org/10.1016/j.ijppaw.2016.07.004
Roux, M. (1994) The CALSUB cruise on the bathyal slopes off New Caledonia. In: Crosnier, A. (Ed.), Résultats des Campagnes MUSORSTOM 12. Mémoires du Muséum National d’Histoire Naturelle, 161, 9–47.
Rowe, 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(4), 257–291.
Rowe, F.W.E. & Gates, J. (1995) Echinodermata in Wells, A. (Ed.), Zoological Catalogue of Australia. Vol. 33. CSIRO, Melbourne, 510 pp.
Sars, G.O. (1875) On Some Remarkable Forms of Animal Life From the Great Deeps off the Norwegian Coast. II. Researches on the Structure and Affinity of the genus Brisinga Based on the study of a New Species, Brisinga coronata. Christiana University, Christiania, 112 pp.
Sheild, C.J. & Witman, J.D. (1993) The impact of Henricia sanguinolenta (O.F. Müller) (Echinodermata:Asteroidea) predation on the finger sponges, Isodictya spp. Journal of Experimental Marine Biology and Ecology, 166 (1), 107–133. https://doi.org/10.1016/0022-0981(93)90081-X
Sladen, W.P. (1889) Asteroidea. Report of the Scientific Results of H.M.S. Challenger, 30, 1–893.
Spencer, W.K. & Wright, C.W. (1966) Asterozoans, Part U: Echinodermata. In: Moore, R.C. (Ed.), Treatise on Invertebrate Paleontology. 3 (1). University of Kansas Press, Lawrence, Kansas, pp. U4–U107.
Sumida, P.Y.G., Tyler, P.A. & Billett, D.S.M. (2001) Early juvenile development of deep-sea asteroids of the NE Atlantic Ocean, with notes on juvenile bathymetric distributions. Acta Zoologica, 82, 11–40. https://doi.org/10.1046/j.1463-6395.2001.00058.x
Tortonese, E. & Clark, A.M. (1956) On the generic position of the asteroid Goniodiscus placenta Müller and Troschel. Annals of the Magazine of Natural History, Series 12, 9 (101), 347–352. https://doi.org/10.1080/00222935608655821
Turner, R.L. & Dearborn, J.H. (1972) Skeletal morphology of the mud star, Ctenodiscus crispatus (Echinodermata: Asteroidea). Journal of Morphology, 138, 239–262. https://doi.org/10.1002/jmor.1051380207F
VanVeldhuizen, H.D. & Oakes, V.J. (1981) Behavioral responses of seven species of asteroids to the asteroid predator, Solaster dawsoni (responses of asteroids to the predator Solaster dawsoni). Oecologia, 48 (2), 214–220. https://doi.org/10.1007/BF00347967
Verrill, A.E. (1899) Revision of certain genera and species of starfishes, with descriptions of new forms. Transactions of the Connecticut Academy of Arts and Sciences, 10 (1), 145–234.
Zhang, R., Wang, C., Zhou, Y. & Zhang, H. (2019) Morphology and molecular phylogeny of two new species in genus Freyastera (Asteroidea: Brisingida: Freyellidae), with a revised key to close species and ecological remarks. Deep-Sea Research, Part I, 154. https://doi.org/10.1016/j.dsr.2019.103163