Halanonchus scintillatulus sp. nov. from New Zealand and a review of the suborder Trefusiina (Nematoda: Enoplida)

Daniel Leduc, Zeng Qi Zhao, Frederic Sinniger


We provide a review of the enoplid suborder Trefusiina Siddiqi, 1983, based on morphological considerations and analyses of new and published 18S rDNA sequences. We also describe Halanonchus scintillatulus Leduc sp. nov. from the Hauraki Gulf, northern New Zealand, as well as females of Trefusialaimus idrisi Leduc, 2013 from the continental slope of New Zealand. We show for the first time that the structure of the female reproductive system of Trefusialaimus Riemann, 1974 consists of two opposed and outstretched ovaries, an unusual feature for the Enoplida. The Trefusiina did not form a monophyletic group in the 18S rDNA phylogeny due to the placement of Lauratonema Gerlach, 1953 and Trefusialaimus sequences well away from the main Trefusiina clade. However, due to generally weak Maximum Likelihood support values, we refrain from changing the classification of these taxa until more comprehensive analyses can be conducted. Our phylogenetic analysis supports the inclusion of the Trischistomatidae Andrássy, 2007 within the Trefusiina, meaning that all of the enoplid suborders now include at least some terrestrial/freshwater representatives. The Trefusiina currently comprises five families, 14 genera and 92 valid species.


Trefusialaimus; Halanonchinae; Trefusiidae; Triplonchida; continental slope

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DOI: https://doi.org/10.5852/ejt.2020.661


Allgén C.A. 1932. Über einige freilebende marine Nematoden aus der Umgebung der Biologischen Station auf der Insel Herdla (Norweg.). Mit Anhang: Zur Richtigstellung älterer und neuerer mariner Nematodengenera I. Archiv fur Naturgeschichte 1: 399–434.

Allgén C.A. 1933. Freilebende Nematoden aus dem Trondhjemsfjord. Capita Zoologica 4: 1–162.

Allgén C. A. 1935. Die freilebenden Nematoden des Öresunds. Capita Zoologica 6: 1–192.

Allgén C.A. 1947. Zur Kenntnis norwegischer Nematoden XII. Weitere freilebende Nematoden aus der Strandzone bei Rörvik. Det Kongelige Norske Videnskabers Selskabs Forhandlinger 19: 60–63.

Allgén C.A. 1953. Über eine ganz bemerkenswerte neue Art der Oxystomatidengattung Trefusia de Man, T. axonolaimoides n. sp. von Süd-Georgien. Det Kongelige Norske Videnskabers Selskabs Forhandlinger 26: 48–50.

Allgén C.A. 1954. Über eine neue Art der Gattung Xennella Cobb 1920, Xennella filicaudata n. sp. aus Südgeorgien. Zoologischer Anzeiger 152: 93–94.

Altherr E. 1963. Contribution à la connaissance de la faune des sables submergés en Lorraine. Nématodes. Annales de Spéléologie 18: 53–98.

Andrássy I. 1964. Onchulidae n. fam., eine neue Familie der Ordnung Enoplida (Nematoda). Opuscula Zoologica Budapestinensis 5: 25–41.

Andrássy I. 1977. Die Gattungen Amphidelus Thorne, 1939, Paramphidelus n. gen. und Etamphidelus n. gen. (Nematoda: Alaimidae). Opuscula Zoologica Budapestinensis 14: 3–43.

Andrássy I. 1985. A dozen new nematode species from Hungary. Opuscula Zoologica Budapestinensis 19: 3–39.

Andrássy I. 2006. Three new species of the family Tripylidae (Penetrantia: Enoplida) from South America. International Journal of Nematology 16: 208–216.

Andrássy I. 2007. Free-living Nematodes of Hungary II (Nematoda Errantia). Pedozoologica Hungarica. Hungarian Natural History Museum, Systematic Zoology Research Group of the Hungarian Academy of Sciences, Budapest.

Andrássy I. 2011 Two unusually slender nematode species of Trischistoma Cobb, 1913 (Enoplida: Tripylidae). Nematology 13: 561–567.

Argo A.D. & Heyns J. 1973. New and little known species of the nematode families Monhysteridae and Tripylidae from South Africa. Phytophylactica 5: 149–154.

Asghari R., Pourjam E., Heydari R., Zhao Z.Q. & Ramaji A. 2012. Tripylina gorganensis n. sp. (Triplonchida: Tripylidae) from northern Iran. Nematology 14: 613–621.

Asghari R., Eskandari A., Maafi Z.T., Zhao Z.Q., Alvarez-Ortega S. & Nadirkhanloo M. 2015. Description of Trischistoma abharensis n. sp. (Nematoda: Trischistomatidae) and first record of Tripylella intermedia (Bütschli, 1873) Brzeski & Winiszewska-Ślipinska, 1993 (Nematoda: Tripylidae) from Iran. Zootaxa 3986 (3): 357–372. https://doi.org/10.11646/zootaxa.3986.3.5

Bastian H.C. 1865. Monograph on the Anguillutidae, or free nematoids, marine, land, and freshwater, with descriptions of 100 new species. Transactions of the Linnean Society 25: 73–180.

Bezerra T.N., Decraemer W., Eisendle-Flöckner U., Hodda M., Holovachov O., Leduc D., Miljutin D., Mokievsky V., Peña Santiago R., Sharma J., Smol N., Tchesunov A., Venekey V., Zhao Z. & Vanreusel A. 2020. Nemys: World Database of Nematodes. Availale from http://nemys.ugent.be [accessed 15 Jan. 2020]. https://doi.org/10.14284/366

Bik H.M., Lambshead P.J.D., Thomas W.K. & Hunt D.H. 2010. Moving towards a complete molecular framework on the Nematoda: a focus on the Enoplida and early-branching clades. BMC Evolutionary Biology 10: e353. https://doi.org/10.1186/1471-2148-10-353

Blome D. & Schrage M. 1985. Freilebende Nematoden aus der Antarktis: Mit einer Beschreibung der Simpliconematidae nov. fam. (Trefusiida) und einer Revision von Filipjeva Ditlevsen, 1928 (Monhysterida, Xyalidae). Veröffenlichungen des Instituts für Meeresforschung in Bremerhaven 21: 71–96.

Bouwman L.A. 1981. A survey of nematodes from the Ems Estuary Part I: Systematics. Zoologische Jahrbücher, Systematik, Ökologie und Geographie der Tiere 108: 335–385.

Bresslau E. & Schuurmans Stekhoven J.H. 1940. Marine Freilebende Nematoden aus der Nordsee. Musée d’Histoire Naturelle de Bruxelles.

Brzeski M.W. 1963. Nematode genera of the family Tripylidae (Nematoda, Enoplida). Acta Zoologica Cracoviensia 8: 295–308.

Brzeski M.W. & Winiszewska-Ślipińska G. 1993. Taxonomy of Tripylidae (Nematoda: Enoplia). Nematologica 39: 12–52.

Bussau C. 1990. Freilebende Nematoden aus Küstendünen und angrenzenden Biotopen der deutschen und dänischen Küsten. II. Monhysterida, Enoplida und Trefusiida (Nematoda). Zoologischer Anzeiger 225: 189–209.

Bussau C. 1993. Taxonomische und ökologische Untersuchungen an Nematoden des Peru-Beckens. PhD Thesis. Christian-Albrechts-Universitat, Kiel.

Bütschli O. 1874. Zur Kenntniss der freilebenden Nematoden, insbesondere der des Kieler Hafens. Abhandlungen, Herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft 9: 236–292.

Castresana J. 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17: 540–552.

Chen Y. & Guo Y. 2015. Two new species of Lauratonema (Nematoda: Lauratonematidae) from the intertidal zone of the East China Sea. Journal of Natural History 49: 1777–1788. https://doi.org/10.1080/00222933.2015.1006285

Chitwood B.G. 1935. Nomenclatorial notes, I. Proceedings of the Helminthological Society of Washington 2: 51–54.

Chitwood B.G. 1936. Some marine nematodes from North Carolina. Proceedings of the Helminthological Society of Washington 3: 1–16.

Chitwood B.G. 1951. North American marine nematodes. Texas Journal of Science 3: 617–672.

Cid del Prado-Vera I., Ferris H. & Nadler S.A. 2010. Soil inhabiting nematodes of the genera Trischistoma, Tripylina and Tripyla from México and the USA with descriptions of new species. Journal of Nematode Morphology and Systematics 13: 29–49.

Cid del Prado-Vera I., Ferris H., Nadler S.A. & Lamothe-Argumedo R. 2012. Four new species of Tripylina Brzeski, 1963 (Enoplida: Tripylidae) from Mexico, with an emended diagnosis of the genus. Journal of Nematode Morphology and Systematics 15: 71–86.

Cid del Prado-Vera I., Ferris H, & Nadler S.A. 2016. Five new species of the family Trischistomatidae (Nematoda: Enoplida) from North and Central America, with keys to the species of Trischistoma and Tripylina. Zootaxa 4109 (2): 173–197. https://doi.org/10.11646/zootaxa.4109.2.4

Clark W.C. 1961. A revised classification of the order Enoplida (Nematoda). New Zealand Journal of Science 4: 123–150.

Cobb N.A. 1894. Tricoma and other nematode genera. Proceedings of the Linnean Society of New South Wales 8: 389–421.

Cobb N.A. 1913. New nematode genera found inhabiting fresh water and nonbrackish soils. Journal of the Washington Academy of Sciences 3: 432–444.

Cobb N.A. 1920. One hundred new nemas (type species of 100 new genera). Contributions to a Science of Nematology 9: 217–343.

Cobb N.A. 1933. New nemic genera and species, with taxonomic notes. Journal of Parasitology 20: 81–94.

Coomans A. 1979. A proposal for a more precise terminology of the body regions in the nematode. Annales de la Société Royale Zoologique de Belgique 108: 115–117.

Darriba D., Taboada G.L., Doallo R. & Posada D. 2012. jModelTest2: more models, new heuristics and parallel computing. Nature Methods 9: 772. https://doi.org/10.1038/nmeth.2109

De Coninck L.A. 1937. Sur le male de Xennella suecica Allgén 1935 (Nematodes), ainsi que sur la position systématique du genre Xennella Cobb, 1920. Biologisch Jaarboek (Dodonaea) 4: 378–387.

De Coninck L.A. 1965. Systématique des nématodes. In: Grassé P.P. (ed.) Traité de Zoologie: Anatomie, Systématique, Biologie. Nemathelminthes (Nematodes): 586–731. Masson et Cie, Paris.

De Coninck L.A. & Schuurmans Stekhoven J.H. 1933. The freeliving marine nemas of the Belgian Coast. II. With general remarks on the structure and the system of nemas. Mémoires du Musée Royal d’Histoire Naturelle de Belgique 58: 3–163.

De Ley P. & Blaxter M.L. 2002. Systematic position and phylogeny. In: Lee D.L. (ed.) The Biology of Nematodes: 1–30. Taylor and Francis, London.

De Ley P. & Blaxter M.L. 2004. A new system for Nematoda: combining morphological characters with molecular trees, and translating clades into ranks and taxa. Nematology Monographs & Perspectives 2: 633–653.

De Ley P., De Ley I.T., Morris K., Abebe E., Mundo-Ocampo M., Yoder M., Heras J., Waumann D., Rocha-Olivares A., Jay Burr A.H., Baldwin J.G. & Thomas W.K. 2005. An integrated approach to fast and informative morphological vouchering of nematodes for applications in molecular barcoding. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 360: 1945–1958.

de Man J.G. 1876. Onderzoekingen over vrij in de aarde levende nematoden. Tijdschrift der Nederlandsche Dierkundige Vereeniging 2: 78–196. Available from https://www.biodiversitylibrary.org/page/9771869 [accessed 18 May 2020].

de Man J.G. 1880. Die einheimischen, frei in der reinen Erde und im süßen Wasser lebende Nematoden monographisch bearbeitet. Vorläufiger Bericht und descriptiv-systematischer Theil. Tijdschrift der Nederlandsche Dierkundige Vereeniging 5: 1–104.

Available from https://www.biodiversitylibrary.org/page/9964525 [accessed 18 May 2020].

de Man J.G. 1888. Sur quelques nématodes libres de la Mer du Nord, nouveaux ou peu connus. Mémoires de la Société zoologique de France 1: 1–51.

de Man J.G. 1893. Cinquième note sur les nématodes libres de la Mer du Nord et de la Manche. Mémoires de la Société zoologique de France 6: 81–125.

Edgar R.C. 2004a. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32: 1792–1797.

Edgar R.C. 2004b. MUSCLE: a multiple sequence alignment method with reduced time and space complexity. BMC Bioinformatics 5: e113. https://doi.org/10.1186/1471-2105-5-113

Fadeeva N. 1989. Free-living nematode family Lauratonematidae (Nematoda Enoplida) from the Sea of Japan. Biologicheskie Nauki 33–39.

Filipjev I.N. 1918. Free-living marine nematodes of the Sevastopol area. Transactions of the Zoological Laboratory and the Sevastopol Biological Station of the Russian Academy of Sciences 4: 1–350.

Filipjev I.N. 1922. Encore sur les nématodes libres de la Mer Noire. Acta Instituti Agronomomici Stavropol 1: 83–184.

Filipjev I.N. 1925. Les nématodes libres des mers septentrionales appartenant a la famille des Enoplidae. Archiv für Naturgeschichte 91: 1–216.

Filipjev I.N. 1928. Free-living nematodes from the Oka River. Okskaia Biologicheskaia Stantsiia (Oka Biological Station) 5: 81–112.

Filipjev I.N. 1929. Classification of free-living Nematoda and relations to parasitic forms. Journal of Parasitology (Urbana) 15: 281–282.

Filipjev I.N. 1934. The classification of the free-living nematodes and their relation to the parasitic nematodes. Smithsonian Miscellaneous Collections 89 (6): 1–63.

Filipjev I.N. 1946. Free-living nematodes from the Northern Arctic Ocean. Trudy, Dreifuiushchaia Ekspeditsiia Glavsevmorputi na Ledokol’nom Parokhode “G. Sedov” 1937–1940 3: 158–184.

Gerlach S.A. 1953. Lauratonema nov. gen., Vertreter einer neuen Familie mariner Nematoden aus dem Küstengrundwasser. Zoologischer Anzeiger 151: 43–52.

Gerlach S.A. 1955. Zur Kenntnis der freilebenden marinen Nematoden von San Salvador. Zeitschrift für wissenschaftliche Zoologie 158: 249–303.

Gerlach S.A. 1956. Brasilianische Meeres-Nematoden I. Boletim do Instituto Oceanográfico 5: 3–69.

Gerlach S.A. 1957. Die Nematodenfauna des Sandstrandes an der Küste von Mittelbrasilien (Brasilianische Meeres-Nematoden IV). Mitteilungen aus dem Zoologischen Museum Berlin 33: 411–459.

Gerlach S.A. 1958. Deuxième contribution à la faune des nématodes des eaux interstitielles littorales de Madagascar. Mémoires de l’Institut Scientifique de Madagascar 2: 343–365.

Gerlach S.A. 1959. Drei neue Nematoden aus dem Küstengrundwasser der Insel Abd el-Kuri (Golf von Aden). Zoologischer Anzeiger 163: 360–364.

Gerlach S.A. 1962. Freilebende Meeresnematoden von den Malediven. Kieler Meeresforschungen 18: 81–108.

Gerlach S.A. 1964. Freilebende Nematoden aus dem Roten Meer. Kieler Meeresforschungen 20: 18–34.

Gerlach S.A. 1966. Bemerkungen zur Phylogenie der Nematoden. Mitteilungen aus der Biologischen Bundesanstalt für Land- und Forstwirtschaft 118: 25–39.

Gerlach S.A. & Riemann F. 1973/1974. The Bremerhaven Checklist of Aquatic Nematodes. A Catalogue of Nematoda Adenophorea excluding the Dorylaimida. Veröffentlichungen des Instituts für Meeresforschung in Bremerhaven, Supplement 4 (1–2): 1–736.

Gourbault N. & Vincx M. 1985. Deux espèces nouvelles d`Ironidae marins; observations sur les spermatozoïdes flagellés des nématodes. Bulletin du Musée national d’Histoire naturelle 4: 109–118.

Groza-Rojancovski E. 1972. Free-living marine nematodes from the Black Sea: Description of three new species. Revue Roumaine de Biologie, Série de Zoologie 17: 79–85.

Guindon S., Dufayard J.F., Lefort V., Anisimova M., Hordijk W. & Gascuel O. 2010. New algorithms and methods to estimate Maximum-Likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology 59: 307–321. https://doi.org/10.1093/sysbio/syq010

Hoeppli R. & Chu H.J. 1934. Greenia orientalis (Corrigendum). Hong Kong Naturalist 5: 161.

Holovachov O. 2019. Campydoroides manautei gen. et sp. nov. from New Caledonia and a reappraisal of the suborder Campydorina (Nematoda). European Journal of Taxonomy 518: 1–23. https://doi.org/10.5852/ejt.2019.518

Holovachov O. & Shoshin A. 2014. Order Triplonchida. In: Schmidt-Rhaesa A. (ed.) Handbook of Zoology Volume 2: Nematoda: 251–276. De Gruyter, Hamburg.

Holterman M., van der Wurff A., van den Elsen S., van Megen H., Bongers T., Holovachov O., Bakker J, & Helder J. 2006. Phylum-wide analysis of SSU rDNA reveals deep phylogenetic relationships among nematodes and accelerated evolution towards crown clades. Molecular Biology and Evolution 23: 1792–1800. https://doi.org/10.1093/molbev/msl044

Holterman M., Schratzberger M. & Helder J. 2019. Nematodes as evolutionary commuters between marine, freshwater and terrestrial habitats. Biological Journal of the Linnaean Society 128: 756-767. https://doi.org/10.1093/biolinnean/blz107

Hope W.D. & Murphy D.G. 1972. A taxonomic hierarchy and checklist of the genera and higher taxa of marine nematodes. Smithsonian Contributions to Zoology 137: 1–101.

Hopper B.E. 1961. Marine nematodes from the coast line of the Gulf of Mexico. Canadian Journal of Zoology 39 (2): 183–199. https://doi.org/10.1139/z61-023

Huelsenbeck J.P. & Ronquist F. 2001. MR BAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17: 1754–755.

Jairajpuri M.S. 1983. Observations on Campydora (Nematoda: Dorylaimida). Nematologia Mediterranea 11: 33–42.

Jayasree K. & Warwick R.M. 1977. Free-living marine nematodes of a polluted sandy beach in the Firth of Clyde, Scotland. Description of seven new species. Journal of Natural History 11 (3): 289–302. https://doi.org/10.1080/00222937700770211

Juario J.V. 1974. Neue freilebende Nematoden aus dem Sublitoral der Deutschen Bucht. Veröffentlichungen des Instituts für Meeresforschung in Bremerhaven 14: 275–303.

Kearse M., Moir R., Wilson A., Stones-Havas S., Cheung M., Sturrock S., Buxton S., Cooper A., Markowitz S., Duran C., Thierer T., Ashton B., Mentjies P. & Drummond A. 2012. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1647–1649. https://doi.org/10.1093/bioinformatics/bts199

Keppner E.J. 1986. New species of free-living marine nematodes (Nematoda: Enoplida) from Bay County, Florida, U.S.A. Transactions of the American Microscopical Society 105: 319–337.

Keppner E.J. 1992. Some free-living marine nematodes from Northwest Florida, U.S.A. with descriptions of three new species (Nematoda: Chromadorida, Trefusiida). Transactions of the American Microscopical Society 111: 199–210.

Khera S. 1970. Studies on the genus Tripyla Bastian (Nematoda: Tripylidae). Proceedings of the Zoological Society 23: 195–200.

Khera S. 1971. Nematodes from the banks of still and running water VII. Family Monhysteridae. Nematologica 16: 495–502.

Leduc D. 2013. Two new free-living nematode species (Trefusiina: Trefusiidae) from the Chatham Rise crest, Southwest Pacific Ocean. European Journal of Taxonomy 55: 1–13. https://doi.org/10.5852/ejt.2013.55

Leduc D. & Zhao Z. 2018. Phylogenetic relationships within the Cyatholaimidae (Nematoda: Chromadorida), the taxonomic significance of pore and pore-like structures, and a description of two new species. Marine Biodiversity 48: 217–230. https://doi.org/10.1007/s12526-016-0605-z

Lorenzen S. 1981. Entwurf eines phylogenetischen Systems der freilebenden Nematoden. Veröffentlichungen des Instituts fur Meeresforschung in Bremerhaven 7: 1–472.

Lorenzen S. 1994. The Phylogenetic Systematics of Freeliving Nematodes. The Ray Society, London.

Meldal B.H.M., Debenham N.J., De Ley P., De Ley I.T., Vanfleteren J.R., Vierstraete A.R., Bert W., Borgonie G., Moens T., Tyler P.A., Austen M.C., Blaxter M.L., Rogers A.D. & Lambshead P.J.D. 2007. An improved molecular phylogeny of the Nematoda with special emphasis on marine taxa. Molecular Phylogenetics and Evolution 42: 622–636. https://doi.org/10.1016/j.ympev.2006.08.025

Micoletzky H. 1922. Die freilebenden Erd-Nematoden mit besonderer Berücksichtigung der Steiermark und der Bukowina, zugleich mit einer Revision sämtlicher nicht mariner, freilebender Nematoden in Form von Genus-Beschreibungen und Bestimmungsschlüsseln. Archiv für Naturgeschichte 87: 1–650.

Miljutin D., Gunnar G., Miljutina M.M., Mokievsky V., Fonseca-Genevois V. & Esteves A.M. 2010. The state of knowledge on deep-sea nematode taxonomy: how many valid species are known down there? Marine Biodiversity 40: 143–159. https://doi.org/10.1007/s12526-010-0041-4

Mullin P., Harris T. & Powers T. 2003. Systematic status of Campydora Cobb, 1920 (Nematoda: Campydorina). Nematology 5: 699–711.

Murphy D.G. & Jensen H.J. 1961. Lauratonema obtusicaudatum, n. sp. (Nemata: Enoploidea), a marine nematode from the coast of Oregon. Proceedings of the Helminthological Society of Washington 28: 167–169.

Nylander J.A.A. 2004. MrModeltest Version 2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University, Uppsala.

Ott J.A. 1972. Determination of Fauna boundaries of nematodes in an intertidal sand flat. Internationale Revue der Gesamten Hydrobiologie 57: 645–663.

Ott J.A. 1977. New freeliving marine nematodes from the West Atlantic I. Four new species from Bermuda with a discussion of the genera Cytolaimium and Rhabdocoma Cobb, 1920. Zoologischer Anzeiger 198: 120–138.

Ott J.A. & Sciemer F. 1973. Respiration and anaerobiosis of free-living nematodes from marine and limnic sediments. Netherlands Journal of Sea Research 7: 233–243.

Pavlyuk O.N. 1984. New species of marine free-living nematodes in the Sea of Japan and comments to the genus Halanonchus. Zoologicheskii Zhurnal 63: 1144–1149.

Pearse A.S. 1942. Introduction to Parasitology. Bailliere, Tindall & Cox, London.

Pham T.H., Wang H.T., Zhao Z.Q. & Zheng J.W. 2013. A new species of the genus Tripylina Brzeski, 1963 (Nematoda: Enoplida: Trischistomatidae) from Zhejiang Province, eastern China. Zootaxa 3717 (2): 158–168. https://doi.org/10.11646/zootaxa.3717.2.2

Platonova T.A. 1971. Exploration of the fauna of the seas VIII (XVI). Fauna and flora of the Possjet Bay of the Sea of Japan. Zoological Institute Academy of Science of the USSR 8: 72–108.

Rambaut A. & Drummond A.J. 2007. Tracer v 1.4. Available from http://beast.bio.ed.ac.uk/Tracer [accessed 28 Apr. 2020].

Riemann F. 1966. Die Gattung Trefusia De Man, 1893 (Enoplida: Oxystominidae). Beitrag zum natürlichen System freilebender Nematoden. Veröffentlichungen des Instituts für Meeresforschung in Bremerhaven 10: 1–30.

Riemann F. 1974. Trefusialaimus nov. gen. (Nematoda) aus der iberischen Tiefsee mit Diskussion des männlichen Genitalapparates von Enoplida Tripyloidea. Meteor Forschungsergebnisse 18: 39–43.

Rusin L.Y., Aleshin V.V., Vladychenskaya N.S., Milyutina I.A., Kedrova O.S. & Petrov N.B. 2001. Trefusiidae are a subtaxon of marine Enoplida (Nematoda): evidence from primary structure of hairpin 35 and 48 loops of SSU rRNA gene. Molecular Biology 35: 778–784.

Schneider W. 1939. Würmer oder Vermes. II. Fadenwürmer oder Nematoden. Freilebende und pflanzenparasitische Nematoden. In: Dahl F. (ed.) Die Tierwelt Deutschlands 36. Gustav Fischer, Jena.

Schuurmans Stekhoven J.H. Jr. 1942. The free living nematodes of the Mediterranean. III. The Balearic Islands. Zoölogische Mededeelingen 23: 229–262.

Schuurmans Stekhoven J.H. Jr. 1950. The freeliving marine nemas of the Mediterranean: I. The Bay of Villefranche. Mémoires de l’Institut royal des Sciences naturelles de Belgique 2: 1–220.

Schuurmans Stekhoven J.H. Jr. 1951. Nématodes saprozoaires et libres du Congo Belge. Mémoires de l’Institut royal des Sciences naturelle de Belgique 39: 1–79.

Shi B. & Xu K. 2017. Morphological and molecular characterizations of Africanema multipapillatum sp. nov. (Nematoda, Enoplida) in intertidal sediment from the East China Sea. Marine Biodiversity 48: 281–288. https://doi.org/10.1007/s12526-017-0690-7

Siddiqi M.R. 1983. Phylogenetic relationships of the soil nematode orders Dorylaimida, Mononchida, Triplonchida and Alaimida, with a revised classification of the subclass Enoplia. Pakistan Journal of Nematology 1 (2): 79–110.

Sinniger F., Reimer J.D. & Pawlowski J. 2009. The Parazoanthidae (Hexacorallia: Zoantharia) DNA taxonomy: description of two new genera. Marine Biodiversity 40: 57–70. https://doi.org/10.1007/s12526-009-0034-3

Somerfield P.J. & Warwick R.M. 1996. Meiofauna in Marine Pollution Monitoring Programmes: a Laboratory Manual. Ministry of Agriculture, Fisheries and Food, Lowestoft UK.

Smol N. & Coomans A. 2006. Order Enoplida. In: Eyualem-Abebe, Traunspurger W. & Andrássy I. (eds) Freshwater Nematodes: Ecology and Taxonomy: 225–292. CABI Publishing, Oxfordshire.

Steiner G. 1916. Freilebende Nematoden aus der Barentssee. Zoologische Jahrbücher 39: 511–664.

Swofford D.L. 2002. PAUP*: Phylogenetic Analysis Using Parsimony (*and other methods), Ver. 4. Sinauer Associates, Sunderland MA.

Tahseen Q. & Nusrat T. 2010. Some new and known species of genera Tripylina Brzeski and Trischistoma Cobb, 1913 (Nematoda) with a discussion on their relationships. Journal of Nematology 42: 128–138.

Tahseen Q., Sultana R., Khan R. & Hussain A. 2012. A new genus and species of the family Rhabdolaimidae (Nematoda), with descriptions of two known species and taxonomic discussion. Journal of Nematology 44: 302–312.

Talavera G. & Castresana J. 2007. Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Systematic Biology 56: 564–577.

Tchesunov A.V. 1984. Data for a revision of marine free-living nematodes of the family Lauratonematidae. Proceedings of the Zoological Institute of the USSR Academy of Science (Leningrad) 126: 79–96.

Tchesunov A.V. 1988. New species of nematodes from the White Sea. Proceedings of the Zoological Institute of the USSR Academy of Science (Leningrad) 180: 68–76.

Tchesunov A.V. 1990. New taxa of marine free-living nematodes of the Family Xyalidae Chitwood, 1951 (Nematoda, Chromadorida, Monhysterida) from the White Sea. In: Gagarin V.G. (ed.) Fauna, Biology and Systematics of Free-living Lower Worms. Institute of Inland Water Biology, Proceedings 64: 101–117. Academy of Sciences of the USSR, St Petersburg.

Timm R.W. 1961. The marine nematodes of the Bay of Bengal. Proceedings of the Pakistan Academy of Science 1: 25–88.

Tsalolikhin S.J. 1983. Nematodes of the Families Tobrilidae and Tripylidae of the World Fauna. Nauka, Leningrad.

Van Megen H., van den Elsen S., Holterman M., Karssen G., Mooyman P., Bongers T., Holovachov O., Bakker J. & Helder J. 2009. A phylogenetic tree of nematodes based on about 1200 full-length small subunit ribosomal DNA sequences. Nematology 11: 927–950. https://doi.org/10.1163/156854109X456862

Vinciguerra M.T. & La Fauci G. 1978. Nematodi muscicoli dell’isola di Lampedusa. Animalia 5: 13–37.

Vincx M. & Furstenberg J.P. 1988. Africanema interstitialis gen. nov., sp. nov., a species which indicates the relationship between the Trefusiidae (Halanonchinae) and the Tripyloididae (Nematoda). Stygologia 4: 10–16.

Vincx M. & Vanreusel A. 1989. Free-living marine nematodes from the Southern Bight of the North Sea II. Notes on species of the Trefusiidae Gerlach, 1966. Hydrobiologia 175: 213–221.

Vitiello P. 1970. Nématodes libres marins des vases profondes du Golfe du Lion. I. Enoplida. Téthys 2: 139–210.

Vitiello P. 1971. Nématodes nouveaux des vases terrigènes côtières des côtes provenҫales. Thetys 2: 859–875.

Wieser W. 1956. Free-living marine nematodes III. Axonolaimoidea and Monhysteroidea. Acta Universitatis Lundensis 52: 1–115.

Wieser W. 1959. Free-living nematodes and other small invertebrates of Puget Sound beaches. University of Washington Publications in Biology 19: 1–179.

Wieser W. & Hopper B. 1967. Marine nematodes of the east coast of North America. I. Florida. Bulletin of the Museum of Comparative Zoology 135: 239–344.

Xu Y.M., Zhao Z.Q., Wang J.M. & Zheng J.W. 2013. A new species of the genus Tripylina Brzeski, 1963 (Nematoda: Enoplida: Trischistomatidae) from Shanxi province, China. Zootaxa 3630 (3): 561–570. https://doi.org/10.11646/zootaxa.3630.3.10

Xu Y.M., Zhao Z.Q. & Wang J.M. 2015. A new species and a new record of Trischistoma Cobb, 1913 (Nematoda: Enoplida: Trischistomatidae) from Shanxi Province, China. Zootaxa 3937 (3): 564–576. https://doi.org/10.11646/zootaxa.3937.3.8

Yeates G.W. 1972. Trischistoma stramenti n. sp. (Nematoda: Enoplida) from leaf litter. New Zealand Journal of Science 14: 578–900.

Zhao Z.Q. 2009. A review of the genus Tripylina Brzeski, 1963 (Nematoda: Triplonchida), with descriptions of five new species from New Zealand. Zootaxa 2238 (1): 1–24. https://doi.org/10.11646/zootaxa.2238.1.1

Zhao Z.Q. 2011. A review of the genus Trischistoma Cobb, 1913 (Nematoda: Enoplida), with descriptions of four new species from New Zealand. Zootaxa 3045 (1): 1–25. https://doi.org/10.11646/zootaxa.3045.1.1

Zhao Z.Q. & Buckley T.R. 2009. Phylogenetic analyses of nematode nuclear 18S rDNA sequences indicates the genus Tripylina Brzeski, 1963 (Nematoda: Tripylidae de Man, 1876) should be placed in Enoplida. Zootaxa 2238 (1): 25–32. https://doi.org/10.11646/zootaxa.2238.1.2

Zhao Z., Li D. & Buckley T.R. 2012. Analysis of primary structure loops from hairpins 35 and 48 of the Nematoda SSU rRNA gene provides further evidence that the genera Tripylina Brzeski, 1963, Trischistoma Cobb, 1913 and Rhabdolaimus de Man, 1880 are members of Enoplida. Zootaxa 3208: 41–57 (1). https://doi.org/10.11646/zootaxa.3208.1.3

Zullini A. 2006. Order Triplonchida. In: Eyualem-Abebe, Traunspurger W. & Andrássy I. (eds) Freshwater Nematodes: Ecology and Taxonomy: 293–325. CABI Publishing, Wallingford UK.


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