Glaciambulata neumayeri gen. et sp. nov., a new Antarctic trachymedusa (Cnidaria: Hydrozoa), with a revision of the family Ptychogastriidae

  • Horia R. Galea Hydrozoan Research Laboratory, 405 Chemin des Gatiers, 83170 Tourves
  • Cornelia Roder Alfred-Wegener-Institut, Helmholtz Center for Polar and Marine Research Centre for Scientific Diving, Am Binnenhafen 1117, 27498 Helgoland
  • Christoph Walcher Alfred-Wegener-Institut, Helmholtz Center for Polar and Marine Research Centre for Scientific Diving, Am Binnenhafen 1117, 27498 Helgoland
  • Marco Warmuth Alfred-Wegener-Institut, Helmholtz Center for Polar and Marine Research Centre for Scientific Diving, Am Binnenhafen 1117, 27498 Helgoland
  • Eberhard Kohlberg Alfred-Wegener-Institut, Helmholtz Center for Polar and Marine Research, Am Handelshafen 12, 27570 Bremerhaven
  • Philipp F. Fischer Alfred-Wegener-Institut, Helmholtz Center for Polar and Marine Research Centre for Scientific Diving, Am Binnenhafen 1117, 27498 Helgoland
Keywords: trachymedusae, Rhopalonematidae, hydromedusa, sympagic, Atka Bay


A new genus and species of sympagic trachymedusa, Glaciambulata neumayeri gen. et sp. nov., are described based on material from Antarctica. Its generic features are compared to those of its relatives, Ptychogastria Allman, 1878 and Tesserogastria Beyer, 1959, and a review of the family Ptychogastriidae Mayer, 1910, based on literature data, is provided. From this, it results that the poorly-known Pectis antarctica Haeckel, 1879, formerly assigned to Ptychogastria by a number of authors, displays all characters of the contemporary rhopalonematid genus Voragonema Naumov, 1971. It is additionally demonstrated that V. laciniata Bouillon et al., 2001 is coterminous with P. antarctica, a finding that clarifies, 137 years later, the morphology and the taxonomic status of Haeckel’s medusa. In accordance with the Principle of Priority, Pectis is recognized as the valid name of the genus, and Voragonema is assigned to its synonymy. Similarly so, the specific name introduced by Haeckel has priority over laciniata.


Allman G.J. 1878. Appendix No. XI. Hydrozoa. In: Feilden H.W. (ed.) Narrative of a voyage to the Polar Sea during 1875-6 in H.M. ships “Alert” and “Discovery” by Capt. G.S. Nares 2: 290–292. Sampson Low, Marston, Searle & Rivington, London.

Antsulevich A.E. 1991. Ob endemizme Belomorskoĭ fauny gidroidov i svyazi mezhdu faunami Belogo i Barentseva moreĭ. On the White Sea hydrozoan fauna endemismus: White and Barentz Seas faunas connection. In: Bentos Belogo Morya. Populyatsii, biotsenozy, fauna. Trudy Zoologicheskogo Instituta, Akademiya Nauk SSSR 233: 35–43. Akademiya Nauk, Moscow.

Antsulevich A.E. 2015. Hydrozoa (Hydroids and Hydromedusae) of Russian seas. St. Petersburg University Press, St. Petersburg.

Aurivillius C.W.S. 1896. Das Plankton der Baffins Bay und Davis’ Strait. In: Almqvist & Wiksellz Boktryckeri-Aktiebolag (eds) Zoologiska Studier. Festskrift Wilhelm Lilljeborg tillegnad på hans åttionde födelsedag af Svenska Zoologer: 179–212, Almqvist & Wiksells, Uppsala.

Aurivillius C.W.S. 1899. Animalisches Plankton aus dem Meere zwischen Jan Mayen - Spitzbergen - K. Karls Land und der Nordküste Norwegens. Kongliga Svenska Vetenskaps-akademiens Handlingar 32 (6): 1–71.

Beliaev G.M. 1989. Deep-sea ocean trenches and their fauna. Nauka Publishing House, Moscow.

Beyer F. 1959. A new, bottom-living trachymedusa from the Oslofjord. Description of the species, and a general discussion of the life conditions and fauna of the fjord deeps. Nytt Magasin for Zoologi 6: 121–143.

Bernstein T. 1934. Zooplankton des nördlichen Teiles des Karischen Meeres. Trudy Arkticheskogo nauchno-issledovaniya Instituta (= Transactions of the Arctic Institute) 9: 3–58.

Bigelow H.B. 1909. Coelenterates from Labrador and Newfoundland, collected by Mr. Owen Bryant from July to October, 1908. Proceedings of the United States National Museum 37 (1706): 301–320.

Bigelow H.B. 1913. Medusae and Siphonophorae collected by the U.S. Fisheries steamer “Albatross” in the northwestern Pacific, 1906. Proceedings of the Unites States National Museum 44 (1946): 1–119.

Boero F. & Bouillon J. 1993. Zoogeography and life cycle patterns of Mediterranean hydromedusae (Cnidaria). Biological Journal of the Linnean Society 48: 239–266.

Bouillon J. 1985. Essai de classification des Hydropolypes-Hydroméduses (Hydrozoa-Cnidaria). Indo-Malayan Zoology 1: 29–243.

Bouillon J. 1993. Classe des Hydrozoaires. In: Doumenc D. (ed) Traité de Zoologie. Anatomie, Systématique, biologie. Tome III. Cnidaires. Cténaires. Fascicule 2. Masson, Paris.

Bouillon J. & Boero F. 2000. Phylogeny and classification of Hydroidomedusae. Synopsis of the families and genera of the Hydroidomedusae of the world, with a list of the worldwide species. Thalassia Salentina 24: 47–296.

Bouillon J., Pagès F. & Gili J.M. 2001. New species of benthopelagic hydromedusae from the Weddel Sea. Polar Biology 24: 839–845.

Bouillon J., Gravili C., Pagès F., Gili J.M. & Boero F. 2006. An introduction to Hydrozoa. Mémoires du Muséum national d’Histoire naturelle 194, Muséum national d’Histoire naturelle, Paris.

Bouillon J., Medel M.D., Pagès F., Gili J.M., Boero F. & Gravili C. 2004. Fauna of the Mediterranean Hydrozoa. Scientia Marina 68 (Suppl. 2): 5–438.

Bouillon J., Pagès F., Gili J.M., Palanques A., Puig P. & Heussner S. 2000. Deep-water Hydromedusae from the Lacaze-Duthiers submarine canyon (Banyuls, northwestern Mediterranean) and description of two new genera, Guillea and Parateclaia. Scientia Marina 64 (Suppl. 1): 87–95.

Broch H. 1907. Hydroiden und Medusen. In: Brøgger T.O. (ed.) Report of the Second Norwegian Arctic Expedition in the “Fram” 1898-1902. 2 (12): 1–12. Videnskabs-Selskabet i Kristiania, Oslo.

Broch H. 1929. Craspedote Medusen. Teil II: Trachylinen (Trachymedusen und Narcomedusen). Nordisches Plankton 12: 481–539.

Browne E.T. 1903. Report on some medusae from Norway and Spitzbergen. Bergens Museum Årbog 1903: 1–36.

Chen Y., Lin Y., Zheng L. & Cao W. 2015. The assemblage and abundance distribution of jellyfish in northern Beibu Gulf. Acta Ecologica Sinica 35 (10): 3382–3393.

Chiu S.Y. 1954. Studies on medusae fauna of south‑eastern China coast, with notes on their geographical distribution. Acta Zoologica Sinica 6 (1): 49–57.

Collins A.G., Bentlage B., Lindner A., Lindsay D., Haddock S.H.D., Jarms G., Norenburg J.L., Jankowski T. & Cartwright P. 2008. Phylogenetics of Trachylina (Cnidaria: Hydrozoa) with new insights on the evolution of some problematical taxa. Journal of the Marine Biological Association of the United Kingdom 88 (8): 1673–1685.

Dunbar J.W. 1942. Marine macroplankton from the Canadian Eastern Arctic. II. Medusae, Siphonophorae, Ctenophora, Pteropoda and Chaetognatha. Canadian Journal of Research (D) 20 (3): 71–77.

Fewkes J.W. 1888. Medusae. Appendix No. 132. Report on the Proceedings of the United States Expedition to Lady Franklin Bay, Grinell Land 2: 39–45.

Foerster R.E. 1923. The Hydromedusae of the West coast of North America, with special reference to those of the Vancouver Islands Region. Contributions to Canadian Biology, n. ser. 1 (12): 219–277.

Galea H.R. 2007. Hydroids and hydromedusae (Cnidaria: Hydrozoa) from the fjords region of southern Chile. Zootaxa 1597: 1–116.

Gili J.M., Bouillon J., Pagès F., Palanques A. & Puig P. 1999. Submarine canyons as habitats of prolific plankton: three new deep-sea Hydroidomedusae in the western Mediterranean. Zoological Journal of the Linnean Society 125: 313–329.

Gili J.M., Bouillon J., Pagès F., Palanques A., Puig P. & Heussner S. 1998. Origin and biogeography of the deep-water Mediterranean hydromedusae including the description of two new species collected in submarine canyons of Northwestern Mediterranean. Scientia Marina 62 (1–2): 113–134.

Gili J.M., Pagès F., Bouillon J., Palanques A., Puig P., Heussner S., Calafat A., Canals M. & Monaco A. 2000. A multidisciplinary approach to the understanding of hydromedusan populations inhabiting Mediterranean submarine canyons. Deep-Sea Res I 47: 1513–1533.

Grönberg G. 1898. Die Hydroid-Medusen des arktischen Gebiets. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 11 (5): 451–468.

Haeckel E. 1879. Das System der Medusen. Erster Theil einer Monographie der Medusen. Mit einem Atlas von vierzig Tafeln. Gustav Fisher, Jena.

Haeckel E. 1881a. Monographie der Medusen. Zweiter Theil. Erste Hälfte: Die Tiefsee-Medusen der Challenger-Reise. Zweite Hälfte: Der Organismus der Medusen. Gustav Fischer, Jena.

Haeckel E. 1881b. Report on the deep-sea medusae dredged by H.M.S. Challenger, during the years 1873-1876. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76, Zoology 4 (2): 1–154.

Hesthagen I.H. 1971. On the biology of the bottom-dwelling Trachymedusa Tesserogastria musculosa Beyer. Norwegian Journal of Zoology 19 (1): 1–19.

Hosia A., Stemmann L. & Youngbluth M. 2008. Distribution of net-collected planktonic cnidarians along the northern Mid-Atlantic Ridge and their associations with the main water masses. Deep Sea Research Part II: Tropical Studies in Oceanography 55 (1–2): 106–118.

Jamieson A. 2015. The hadal zone: life in the deepest oceans. Cambridge University Press, Cambridge.

Kramp P.L. 1914. Meduser og Siphonophorer. In: Conspectus faunae Groenlandicae. Meddelelser om Grønland 23: 381–456.

Kramp P.L. 1942. Medusae. In: The Godthaab Expedition, 1928. Meddelelser om Grønland 81 (1): 1–168.

Kramp P.L. 1943. Medusae, Siphonophora, and Ctenophora. In: The Zoology of East Greenland. Meddelelser om Grønland 121 (12): 1–20.

Kramp P.L. 1947. Medusae. III. Trachylina and Scyphozoa. Danish Ingolf Expedition 5D (14): 1–66.

Kramp P.L. 1955. A revision of Ernst Haeckel’s determinations of a collection of Medusae belonging to the Zoological Museum of Copenhagen. In: Papers in marine Biology and Oceanography. Deep‑Sea Research 3 (Suppl.): 149–168.

Kramp P.L. 1957. Hydromedusae from the Discovery collections. “Discovery” Reports 29: 1–128.

Kramp P.L. 1959. The hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report 46: 1–283.

Kramp P.L. 1961. Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom 40: 1–469.

Kramp P.L. 1968. The hydromedusae of the Pacific and Indian Oceans. Sections II and III. Dana Report 72: 1–200.

Kramp P.L. & Damas D. 1925. Les Méduses de la Norvège. Introduction et partie spéciale. Videnskabelige Meddelelser fra dansk Naturhistorisk Forening i Kjøbenhavn 80: 217–323.

Langhorne P.J., Hughes K.G., Gough A.J., Smith I.J., Williams M.J.M., Robinson N.J., Stevens C.L., Rack W., Price D., Leonard G.H., Mahoney A.R., Haas C. & Haskell T.G. 2015. Observed platelet ice distribution in Antarctic sea ice: An index for ocean-ice shelf heat flux. Geophysical Research Letters 42: 5442–5451.

Larson R.J. & Harbison G.R. 1990. Medusae from McMurdo Sound, Ross Sea including the descriptions of two new species, Leuckartiara brownei and Benthocodon hyalinus. Polar Biology 11: 19–25.

Larson R.J., Mills C.E. & Harbison G.R. 1991. Western Atlantic midwater hydrozoan and scyphozoan medusae: in situ studies using manned submersibles. Hydrobiologia 216/217: 311–317.

Larson R.J., Matsumoto G.I., Madin L.P. & Lewis L.M. 1992. Deep-sea benthic and benthopelagic medusae: recent observations from submersibles and a remotely operated vehicle. Bulletin of Marine Science 51 (3): 277–286.

Levinsen G.M.R. 1893. Meduser, Ctenophorer og Hydroider fra Grønlands Vestkyst, tilligemed Bemaerkninger om Hydroidernes Systematik. Videnskabelige Meddelelser fra dansk Naturhistorisk Forening i Kjøbenhavn (5) 4: 143–212.

Lindsay D.J. & Miyake H. 2009. A checklist of midwater cnidarians and ctenophores from Japanese waters ‒ species sampled during submersible surveys from 1993-2008 with notes on their taxonomy. Kaiyo Monthly 41 (8): 417–438.

Lindsay D. & Pagès F. 2010. Voragonema tatsunoko (Trachymedusae: Rhopalonematidae), a new species of benthopelagic medusa, host to the hyperiid amphipod Mimonectes spandli (Physosomata: Mimonectidae). Zootaxa 2671: 31–39.

Linko A.K. 1904. Zoologische Studien im Barents-Meere. Auf Grund der Untersuchungen der wissenschaftlichen Murman-Expedition. Zoologischer Anzeiger 28 (6): 210–220.

Linko A.K. 1913. Zooplankton’ Sibirskogo ledovitago okeana po sboram’ russkoĭ polyarnoĭ ekspeditsii 1900‑1903 g. Zooplancton de la Mer Glaciale de Sibérie après les récoltes de l’Expédition Polaire Russe en 1900‑1903. Zapiski Imperatorskoĭ akademīi nauk (= Mémoires de l’Académie des Sciences de Saint-Petersbourg) 29 (4): 1–53.

Maas O. 1893. Die craspedoten Medusen der Plankton-Expedition. Ergebnisse der in dem Atlantischen Ocean von Mitte Juli bis Anfang November 1889 ausgeführten Plankton-Expedition der Humboldt-Stiftung 2 (K): 1–17.

Maas O. 1906a. Die arktischen Medusen (ausschlieβlich der Polypenmedusen). Fauna Arctica 4 (3): 479–526.

Maas O. 1906b. Medusen. In: Résultats du voyage du S.Y. Belgica en 1897-1898-1899 sous le commandement de A. de Gerlache de Gomery. Rapports Scientifiques, Zoologie 9 (21): 1–32.

Mackie G.O. 1985. Midwater macroplankton of British Columbia studied by submersible PISCES IV. Journal of Plankton Research 7 (6): 753–777.

Mahuzier S. & Sylvestre J.P. 2016. Cap sur le grand continent blanc. Quae, Versailles.

Mayer A.G .1910. Medusae of the world. Vol. II. The hydromedusae. Carnegie Institution of Washington Publication 109: 231–498.

Minemizu R., Kubota S., Hirano Y. & Lindsay D.J. 2015. A photographic guide to the jellyfishes of Japan. Heibonsha, Tokyo.

Miyake H., Lindsay D.J. & Kubota S. 2004. Midwater and bentho-pelagic animals on the south slope of Shiribeshi Seamount off the west coast of Hokkaido. JAMSTEC Journal of Deep Sea Research 24: 37–42.

Naumov D.V. 1956. [Medusae in the far eastern waters of the Soviet Union]. Trudy Problemnykh i tematicheskikh soveshchanii (=Transactions of Special Problems Conferences) 6: 36–41.

Naumov D.V. 1969. Hydroids and Hydromedusae of the USSR. Israel Program for Scientific Translation, Jerusalem.

Naumov D.V. 1971. Gydroidnye i stsifoidnye medusy iz Kurilo‑Kamchatskogo zhelova (Hydromedusae and Scyphomedusae from the Kurile‑Kamchatka Trench). Trudy Instituta Okeanologii 92: 9–17.

Pagès F., Corbera J. & Lindsay D. 2007. Piggybacking pycnogonids and parasitic narcomedusae on Pandea rubra (Anthomedusae, Pandeidae). Plankton and Benthos Research 2 (2): 83–90.

Panteleeva N.N., Frolova E.A. & Sheiko O.V. 1999. New records of the benthic medusa Ptychogastria polaris Allman, 1878 (Trachylida, Hydroidea) in the Barents Sea and off the Kurile Islands (Pacific Ocean). Polar Biology 22: 372–378.

Picard J. 1955. Nouvelles recherches sur les hydroméduses des herbiers méditerranéens de posidonies. Recueil des Travaux de la Station Marine d’Endoume 15: 59–71.

Rees W.J. 1953. Records of hydroids and medusae taken at Herdla, Bergen in 1937. Universitetet i Bergen. Arbok, 1952, Naturvitenskapelig Rekke 16: 1–8.

Russell F.S. 1980. Trachymedusae. Families: Geryonidae, Ptychogastriidae, Halicreatidae. Fiches d’Identification du Zooplancton 164: 1–4.

Runnström S. 1932. Eine Uebersicht über das Zooplankton des Herdla- und Hjeltefjordes. Bergens Museum Årbog 1931 (7): 1–67.

Sirenko B.I., Markhaseva E.L., Buzhinskaya G.N., Golikov A.A., Menshutkina T.V., Petryashov V.V., Semenova T.N., Stepanjants S.D. & Vassilenko S.V. 1996. Preliminary data on suprabenthic invertebrates collected during the RV Polarstern cruise in the Laptev Sea. Polar Biology 16: 345–352.

Stepanjants S.D. 1989. Hydrozoa of the Eurasian Arctic Seas. In: Herman Y. (ed) The Arctic Seas: 397–430. Van Nostrand Reinhold, New York.

Stepanjants S.D. 2007. Conception of the bipolar distribution of the organisms: origin and modern views, on the Medusozoa example. Problemy Arktiki i Antraktiki 77: 89–96.

Stepanjants S., Coretese G., Kruglikova SB. & Bjørklund K.R. 2006. A review of bipolarity concepts: history and examples from Radiolaria and Medusozoa (Cnidaria). Marine Biology Research 2 (3): 200–241.

Stübing D. & Piepenburg D. 1998. Occurrence of the benthic trachymedusa Ptychogastria polaris Allman, 1878 (Cnidaria: Hydrozoa) off Northeast Greenland and in the northern Barents Sea. Polar Biology 19: 193–197.

Tanasijčuk N. 1927. Zoologische Ergebnisse der Terminalfahrt bis zum 75° n Br. Längs des Kola-Meridians (Barents-Meer) in August, 1925. Dokladi Akademii Nauk USSR 1927: 359–363.

Thiel M.E. 1932a. Uebersicht über die Hydromedusen der Arktis. Fauna Arctica 6 (2): 121–158.

Thiel M.E. 1932b. Die Hydromedusen‑Fauna des Nördlichen Eismeeres in tiergeographischer Betrachtung. Archiv für Naturgeschichte, n. ser. 1 (3): 435–514.

Thuesen E.V. 1993. Vampyrocrossota childressi, a new genus and species of black medusa from the bathypelagic zone off California (Cnidaria: Trachymedusae: Rhopalonematidae). Proceedings of the Biological Society of Washington 106 (1): 190–194.

Thuesen E.V. 2003. Crossota millsae (Cnidaria: Trachymedusae: Rhopalonematidae), a new species of viviparous hydromedusa from the deep sea off California and Hawaii. Zootaxa 309: 1–12.

Vanhöffen E. 1902. Die craspedoten Medusen der deutschen Tiefsee-Expedition 1898–1899. I. Trachymedusen. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition 1898–1899 3: 55–86.

Vanhöffen E. 1912. Die craspedoten Medusen der deutschen Südpolar-Expedition. Deutsche Südpolar-Expedition 1901–1903, Zoologie 5 (3): 351–396.

Vibe C. 1950. The marine mammals and the marine fauna in the Thule district (Northwest Greenland) with observations on ice conditions in 1933‑41. Meddelelser om Grønland 150 (6): 1–116.

Yashnov V.A. 1940. Hydromedusy Sibirskogo poberezh’ya Ledovitogo Okeana (Hydroméduses de l‘Océan glacial de Sibérie). Biulleten‘ Moskovskogo Obshchestva Ispytatelei Prirody (= Bulletin de la Société Impériale des Naturalistes de Moscou, section Biologie), n. ser. 48 (2–3): 107–114.

Yashnov V.A. 1948. Hydromedusae ‑ Gidroidnye medusy. In: N.S. Gaevskoi M.S. (ed) Opredelitel’ fauny i flory severnykh morei SSSR: 65–74, State Publishing House for Soviet Science, Moscow.

Zeidler W. 2012. A review of the hyperiidean families Mimonectidae and Proscinidae (Crustacea: Amphipoda: Hyperiidea: Scinoidea). Zootaxa 3533: 1–74.

How to Cite
Galea, H. R., Roder, C., Walcher, C., Warmuth, M., Kohlberg, E., & Fischer, P. F. (2016). Glaciambulata neumayeri gen. et sp. nov., a new Antarctic trachymedusa (Cnidaria: Hydrozoa), with a revision of the family Ptychogastriidae. European Journal of Taxonomy, (252).