Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2021-02-12
Page range: 234–256
Abstract views: 280
PDF downloaded: 10

Morphological and molecular characterization of adult hairworms (Phylum Nematomorpha) from Iceland and the Faroe Islands, and documentation of their non-adult stages and hosts

School of Biological Sciences, University of Nebraska Lincoln, Lincoln, NE 68588, U.S.A.
Biology Department, Northern Michigan University, 1401 Presque Isle Avenue, Marquette, MI 49855, U.S.A.
Department of Integrative Biology, 501 Life Sciences West, Oklahoma State University, Stillwater, Oklahoma 74078, U.S.A.
Nematoda Gordiida hairworm Gordian worm scanning electron microscopy oviposition behavior non-adult life stages Gordionus Amara Radix peregra phylogeny

Abstract

No species of freshwater Nematomorpha have been described from Iceland, but they have been identified anecdotally. Recent surveys in Iceland using freshwater gastropods as biodiversity indicators resulted in the collection of adult free-living hairworms and their non-adult stages including cysts in snail paratenic hosts and juvenile worms in ground beetle definitive hosts. Additionally, specimens acquired from the Icelandic Institute of Natural History indicate nematomorphs are common in Iceland. A single specimen from the Faroe Islands National Museum represents a new species record. Our morphological and molecular characterization indicated all the samples belong to the species Gordionus wolterstorffii, a common nematomorph found throughout Europe. Also, we provide the first descriptions of the cyst stage for the genus Gordionus. Molecular phylogenetic analysis based on 10 species of Gordionus and one species of the closely related genus Parachordodes indicates that Gordionus is not monophyletic. Combining our morphological and phylogenetic investigations, we discuss the lack of clarity in diagnostic morphological characters and the need for additional global collections to clarify the taxonomy of Gordionus.

 

References

  1. Anaya, C., Schmidt-Rhaesa, A., Hanelt, B. & Bolek, M.G. (2019) A new species of Gordius (Phylum Nematomorpha) from terrestrial habitats in North America. ZooKeys, 892, 59–75.

    https://doi.org/10.3897/zookeys.892.38868

    Anaya, C., Hanelt, B. & Bolek, M.G. (2021) Field and laboratory observations on the life history of Gordius terrestris (Phylum Nematomorpha), a terrestrial nematomorph. Journal of Parasitology. [in press]

    Annandale, N. (1905) Note on the hair-worms in oriental and European folklore. Journal of the Asiatic Society of Bengal, 73, 75–77.

    Baird, W. (1853) Description of some new species of Entozoa from the collection of the British Museum. Proceedings of the Zoological Society of London, 21 (1), 18–25.

    https://doi.org/10.1111/j.1469-7998.1853.tb07169.x

    Begay, A., Schmidt-Rhaesa, A., Bolek, M.G. & Hanelt, B. (2012) Two new Gordionus species (Nematomorpha: Gordiidae) from the southern Rocky Mountains, USA. Zootaxa, 3406 (1), 30–38.

    https://doi.org/10.11646/zootaxa.3406.1.2

    Bleidorn, C., Schmidt-Rhaesa, A. & Garey, J.R. (2002) Systematic relationships of Nematomorpha based on molecular and morphological data. Invertebrate Biology, 121 (4), 357–364.

    https://doi.org/10.1111/j.1744-7410.2002.tb00136.x

    Bolek, M.G. & Coggins, J.R. (2002) Seasonal occurrence, morphology, and observations on the life history of Gordius difficilis (Nematomorpha: Gordioidea) from southeastern Wisconsin, United States. Journal of Parasitology, 88 (2), 287–294.

    https://doi.org/10.1645/0022-3395(2002)088[0287:SOMAOO]2.0.CO;2

    Bolek, M.G., Szmygiel C., Kubat, A., Schmidt-Rhaesa, A. & Hanelt, B. (2013) Novel techniques for biodiversity studies of gordiids and description of a new species of Chordodes (Gordiida, Nematomorpha) from Kenya, Africa. Zootaxa, 3717 (1), 23–38.

    https://doi.org/10.11646/zootaxa.3717.1.2

    Bolek, M.G., Schmidt-Rhaesa, A., De Villalobos, L.C. & Hanelt, B. (2015) Phylum Nematomorpha. In: Thorp, J., and Rogers, D.C. (Eds.), Ecology and General Biology. Vol. I. Thorp and Covich’s Freshwater Invertebrates. 4th Edition. Academic Press, New York, pp. 303–326.

    https://doi.org/10.1016/B978-0-12-385026-3.00015-2

    Bolotov, I.N., Aksenova, O.V., Bespalaya, Y.V., Gofarov, M.Y., Kondakov, A.V., Paltser, I.S., Stefansson, A., Travina, O.V. & Vinarski, M.V. (2017) Origin of a divergent mtDNA lineage of a freshwater snail species, Radix balthica, in Iceland: cryptic glacial refugia or a postglacial founder event? Hydrobiologia, 787, 73–98.

    https://doi.org/10.1007/s10750-016-2946-9

    Brooks, D. R., Hoberg, E.P. & Boeger, W.A. (2019) The Stockholm Paradigm: Climate Change and Emerging Disease. University of Chicago Press, Chicago, Illinois, 400pp.

    Camerano, L. (1888) Ricerche sopra i Gordii d´Europa e descrizione di due nuove specie. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Universitá di Torino 3, Nr. 42, 1–10.

    Camerano, L. (1897) Monografia dei Gordii. Memorie della Reale Accademia delle Scienze de Torino, 47, 339–415.

    Carvalho, J.C.M. (1942) Studies on some Gordiacea of North and South America. Journal of Parasitology, 28 (3) 213–222.

    https://doi.org/10.2307/3272775

    Cham, S.A., Seymour, M.K. & D.J. Hooper (1983) Observations on a British hairworm, Parachordodes wolterstorffi (Nematomoprha: Gordiidae) Journal of Zoology, 199 (3), 275–285.

    https://doi.org/10.1111/j.1469-7998.1983.tb02096.x

    Chiu, M.C., Huang, C.G., Wu, W.J. & Shiao, S.F. (2011) A new horsehair worm, Chordodes formosanus sp. n. (Nematomorpha, Gordiida) from Hierodula mantids of Taiwan and Japan with redescription of a closely related species, Chordodes japonensis. ZooKeys, 160, 1–22.

    https://doi.org/10.3897/zookeys.160.2290

    Chiu, M.C., Huang, C.G., Wu, W.J. & Shiao, S.F. (2016) Annual survey of horsehair worm cysts in northern Taiwan, with notes on a single seasonal infection peak in Chironomid larvae (Diptera: Chironomidae). Journal of Parasitology, 102 (3) 319–326. https://doi.org/10.1645/15-907

    Chiu, M.C., Huang, C.G., Wu, W.J. & Shiao, S.F. (2017) A new orthopteran-parasitizing horsehair worm, Acutogordius taiwanensis sp. n., with a redescription of Chordodes formosanus and novel host records from Taiwan (Nematomorpha, Gordiida). ZooKeys, 683, 1–23.

    https://doi.org/10.3897/zookeys.683.12673

    Chiu, M.C., Huang, C.G., Wu, W.J., Lin, Z.H., Chen, H.W. & Shiao, S.F. (2020) A new millipede-parasitizing horsehair worm, Gordius chiashanus sp. nov., at medium altitudes in Taiwan (Nematomorpha, Gordiida). ZooKeys, 941, 25–48.

    https://doi.org/10.3897/zookeys.941.49100

    Creplin, C.H. (1847) Chordodes parasitus, ein Schmarotzerwurm aus einer Heuschrecke. Notizen aus dem Gebiete der Natur- und Heilkunde, 3, 161–166.

    Dorier, A. (1929) Sur les Gordiacés des Myriapods. Comptes Rendus de l’Académie des Sciences Paris, 188, 743–745.

    Dorier, A. (1930) Recherches biologiques et systématiques sur les Gordiacés. Trav. Lab. Hydrobiol. Piscicult. Université Grenoble Alpes, 22, 1–183.

    Dorier, A. (1935) Sur le passage à la vie latente des larves de Gordiacés. Comptes Renus de l’Academie des Sciences Paris, 200, 492–494.

    Dujardin, F. (1842) Mémoire sur la structure anatomique des Gordius et d´un autre Helminthe, le Mermis, qu´on a confondu avec eux. Annales de Sciences Naturelles, 18, 129–151.

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

    https://doi.org/10.1093/nar/gkh340

    Efeykin, B.D., Schmatko, V.Y. & Spiridonov, S.E. (2016) Comparative Phylogenetic informativity of single ribosomal cluster regions in freshwater horsehair worms (Gordiacea, Nematomorpha). Biology Bulletin, Zoology, 43 (1), 41–49.

    https://doi.org/10.1134/S1062359016010040

    Ernst, C.M., Hanelt, B. & Buddle, C.M. (2016) Parasitism of ground beetles (Coleoptera: Carabidae) by a new species of hairworm (Nematomorpha: Gordiida) in Arctic Canada. Journal of Parasitology, 10 (3), 327–335.

    https://doi.org/10.1645/15-863.

    Eversham B (2013) Identifying freshwater molluscs. Version 2.2. 8 pp. Available from : https://www.naturespot.org.uk/sites/default/files/downloads/Pondsnails%20Key%20version%20%202.2%20iv2013%20illustrated%20PDF.pdf (accessed 29 December 2020)

    Folmer, O., Black, M.., Hoeh, W, Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology Biotechnology, 3 (5) 294–299.

    https://doi.org/10.1371/journal.pone.0013102

    Galbreath, K.E., Hafner, D.J. & Zamudio, K.R. (2009) When cold is better: climate-driven elevation shifts yield complex patterns of diversification and demography in an alpine specialist (American pika, Ochotona princeps). Evolution, 63 (11) 2848–2863.

    https://doi.org/10.1111/j.1558-5646.2009.00803.x

    Glöer, P. (2019) The freshwater gastropods of the West-Palaearctis. Volume I. Fresh- and brackish waters except spring and subterranean snails. Vol. I. Peter Glöer, Hetlingen, 216 pp.

    Hanelt, B. &Janovy, J. (1999) The life cycle of a horsehair worm, Gordius robustus (Nematomorpha: Gordioidea). Journal of Parasitology, 85 (1), 139–141.

    https://doi.org/10.2307/3285720

    Hanelt, B., Grother, L.E. & Janovy, J. Jr. (2001) Physid snails as sentinels of freshwater Nematomorphs. Journal of Parasitology, 87 (5), 1049–1053.

    https://doi.org/10.1645/0022-3395(2001)087[1049:PSASOF]2.0.CO;2

    Hanelt, B. & Janovy, J. Jr. (2002) Morphometric analysis of nonadult characters of common species of American gordiids (Nematomorpha: Gordioidea). Journal of Parasitology, 88 (3), 557–562.

    https://doi.org/10.1645/0022-3395(2002)088[0557:MAONCO]2.0.CO;2

    Hanelt, B. & Janovy, J. (2003) Spanning the gap: experimental determination of paratenic host specificity of horsehair worms (Nematomorpha: Gordiida). Invertebrate Biology, 122 (1), 12–18.

    https://doi.org/10.1111/j.1744-7410.2003.tb00068.x

    Hanelt, B. & Janovy, J. (2004a) Life cycle and paratenesis of American gordiids (Nematomorpha: Gordiida). Journal of Parasitology, 90 (2), 240–244.

    https://doi.org/10.1645/GE-78R

    Hanelt, B. & Janovy, J. (2004b) Untying a Gordian knot: the domestication and laboratory maintenance of a Gordian worm, Paragordius varius (Nematomorpha: Gordiida). Journal of Natural History, 38 (8), 939–950.

    https://doi.org/10.1080/0022293021000058718

    Hanelt, B., Thomas, F. & Schmidt-Rhaesa, A. (2005) Biology of the phylum Nematomorpha. Advances in Parasitology, 59, 243–305.

    https://doi.org/10.1016/S0065-308X(05)59004-3

    Hanelt, B., Bolek, M.G. & Schmidt-Rhaesa, A. (2012) Going solo: discovery of the first parthenogenetic gordiid (Nematomorpha: Gordiida). PlosONE, 7, (4) e34472

    https://doi.org/10.1371/journal.pone.0034472

    Hanelt, B., Schmidt-Rhaesa, A. & Bolek, M.G. (2015) Cryptic species of hairworm parasites revealed by molecular data and crowdsourcing of specimen collections. Molecular Phylogenetics and Evolution, 82, 211–218.

    https://doi.org/10.1016/j.ympev.2014.09.010

    Harkins, C., Shannon, R., Papeş, M., Schmidt-Rhaesa, A., Hanelt, B. & Bolek, M.G. (2016) Using Gordiid cysts to discover the hidden diversity, potential distribution, and new species of Gordiids (Phylum Nematomorpha). Zootaxa, 4088 (4), 515–530.

    https://doi.org/10.11646/zootaxa.4088.4.3

    Heinze, K. (1935) Über Gordiiden (Species inquirendae und Neubeschreibungen). Zoologischer Anzeiger, 111, 23–32.

    Heinze, K. (1937) Die Saitenwürmer (Gordioidea) Deutschlands. Eine systematisch-faunistische Studie über Insektenparasiten aus der Gruppe der Nematomorpha. Zeitschrift für Parasitenkunde, 9, 263–344.

    https://doi.org/10.1007/BF02119887

    Heinze, K. (1952) Über Gordioidea, eine systematische Studie über Insektenparasiten aus der Gruppe der Nematomorpha. Zeitschrift für Parasitenkunde, 15, 183–202.

    https://doi.org/10.1007/BF00260452

    Hoberg, E. P. & D. R. Brooks. (2010) Beyond vicariance: integrating taxon pulses, ecological fitting, and oscillation in evolution and historical biogeography. In: Morand, S. & Krasnov, B.R. (Eds.), The Biogeography of Host-Parasite Interactions. Oxford University Press, Oxford, pp. 7–20.

    Hoberg, E.P. & Brooks, D.R. (2013) Episodic processes, invasion, and faunal mosaics in evolutionary and ecological time. In: Rohde, K. (Ed.), The Balance of Nature and Human Impact. Cambridge University Press, Cambridge, pp. 199–213.

    https://doi.org/10.1017/CBO9781139095075.021

    Hoberg, E.P., Agosta, S.J., Boeger, W.A. & Brooks, D.R. (2015) An integrated parasitology: revealing the elephant through tradition and invention. Trends in Parasitology, 31 (4), 128–133.

    https://doi.org/10.1016/j.pt.2014.11.005

    Inoue, I. (1962) Nematomorpha (or Gordiacea). In: Uchida, T. (Ed.), Systematic Zoology. Vol. 4. Nakayama Shoten, Tokyo, pp 192–220.

    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., Meintjes, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28 (12), 1647–1649.

    https://doi.org/10.1093/bioinformatics/bts199

    Kumar, S., Stecher, G. & Tamura, K. 2016. MEGA7: Molecular evolutionary genetics analysis Version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874.

    https://doi.org/10.1093/molbev/msw054

    Leidy, J. (1851) On the Gordiaceae. Proceedings of the Academy of Natural Sciences of Philadelphia, 5, 262–263 + 266 + 275.

    Linné, C. (1758) Systema Naturae. Vol. 1. 10th Edition. Trustees of the British Museum, Natural History, London, 824 pp. [from Facsimile 1956]

    Looney, C., Hanelt, B. & Zack, R.S. (2012) New Records of Nematomorph parasites (Nematomorpha: Grodiida) of ground beetles (Coleoptera: Carabidae) and camel crickets (Orthoptera: Rhaphidophoridae) in Washington State. Journal of Parasitology, 98 (3), 554–559.

    https://doi.org/10.1645/GE-2929.1

    May, H.G. (1919) Contributions to the life histories of Gordius robustus Leidy and Paragordius varius (Leidy). Illinois Biological Monographs, 5, 1–119.

    Meguro, N., O Kishida, O., Utsumi, S., Niwa, S., Igarashi, S., Kozuka, C., Naniwa, A. & Sato, T. (2020) Host phenologies and the life history of horsehair worms (Nematomorpha, Gordiida) in a mountain stream in northern Japan. Ecological Research 35 (3), 482–493.

    https://doi.org/10.1111/1440-1703.12122

    Mikhailov, K.V., Efeykin, B.D., Panchin, A.Y., Knorre, D.A., Logacheva, M.D., Penin, A.A., Muntyan, M.S., Nikitin, M.A., Popova, O.V., Zanegina, O.N., Vyssokikh, M.Y., Spiridonov, S.E., Aleoshin, V.V. & Panchin, Y.V. (2019) Coding palindfromes in mitochondrial genes of Nematomorpha. Nucleic Acids Research, 47 (13), 6858–6870.

    https://doi.org/10.1093/nar/gkz517

    Montén, E. (1951) Nematomorpha. Acta University Lund, 46, 162–170.

    Müller, G.W. (1926) Über Gordiaceen. Zeitschrift für. Morpholgie und Ökologie der Tiere, 7, 134–220.

    https://doi.org/10.1007/BF00540721

    Muus, B.J. (1956) A note on the Danish hairworms, Nematomorpha: Gordioidea. Videnskabelige Meddelelser fra den Naturhistoriske Forening Kjöbenhavn, 118, 33–39.

    Örley, L. (1881) On hair-worms in the collection of the British Museum. Annals and Magazine of Natural History, 8, 325–332.

    https://doi.org/10.1080/00222938109487466

    Poinar, G. (2008) Global diversity of hairworms (Nematomorpha: Gordiaceae) in freshwater. Hydrobiologia, 595, 79–83.

    https://doi.org/10.1007/s10750-007-9112-3

    Poinar, G.O. Jr. (1991) Hairworm (Nematomorpha: Gordioidea) parasites of New Zealand wetas (Orthoptera: Stenopelmatidae). Canadian Journal of Zoology, 69 (6), 1592–1599.

    https://doi.org/10.1139/z91-223

    Poinar, G., Rykken, J. & LaBonte, J. (2004) Parachordodes tegonotus n. sp. (Gordioidea: Nematomorpha), a hairworm parasite of ground beetles (Carabidae: Coleoptera), with a summary of gordiid parasites of carabids. Systematic Parasitology, 58, 139–148.

    https://doi.org/10.1023/B:SYPA.0000029420.56409.a5

    Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67 (5), 901–904.

    https://doi.org/10.1093/sysbio/syy032

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.4: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542.

    https://doi.org/10.1093/sysbio/sys029

    Sato, T., Watanabe, K., Tamotsu, S., Ichikawa, A. & Schmidt-Rhaesa, A. (2012) Diversity of nematomorph and cohabiting nematode parasites in riparian ecosystems around the Kii Peninsula, Japan. Canadian Journal of Zoology, 90 (7), 829–838. https://doi.org/10.1139/z2012-048

    Schmidt-Rhaesa, A. (1997) Nematomorpha. In: Schwoerbel, J. & Zwick, P. (Eds.), Süßwasserfauna Mitteleuropas. Gustav Fischer Verlag, Stuttgart, pp. 1–124.

    Schmidt-Rhaesa, A. (2001) Variation of cuticular characters in the nematomorpha: studies on Gordionus violaceus (Baird, 1853) and G. wolterstorffii (Camerano, 1888) from Britain and Ireland. Systematic Parasitology, 49, 41–57.

    https://doi.org/10.1023/A:1010600200665

    Schmidt-Rhaesa, A. (2013) Nematomorpha. In: Schmidt-Rhaesa, A. (Ed.), Handbook of Zoology. Gastrotricha, Cycloneuralia and Gnathifera. Nematomorpha, Priapulida, Kinorhyncha and Loricifera. Vol 1. De Gruyter, Berlin, pp. 29–145.

    https://doi.org/10.1515/9783110272536.1

    Schmidt-Rhaesa, A. (2016) The collection of Nematomorpha in the Zoological Museum Hamburg, including description of a new species, Chordodes jelkae sp. n. Zoosystematics and Evolution, 92 (2), 211–250.

    https://doi.org/10.3897/zse.92.10351

    Schmidt-Rhaesa, A., Hanelt, B. & Reeves, W.K. (2003) Redescription and compilation of Nearctic freshwater Nematomorpha (Gordiida), with the description of two new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 153 (1), 77–117.

    https://doi.org/10.1635/0097-3157(2003)153[0077:RACONF]2.0.CO;2

    Schmidt-Rhaesa, A. & Bryant, M.S. (2004) Filling the Gondwana gaps: new species and new reports of the genus Beatogordius (Nematomorpha) from Australia and Madagascar. Systematic Parasitology, 57, 173–181.

    https://doi.org/10.1023/B:SYPA.0000019084.58675.e0

    Schmidt-Rhaesa, A. & Sato, T. (2009) Gordionus chinensis (Villot, 1874) and Gordionus kii sp.n. (Nematomorpha, Gordiida), new reports of the genus Gordionus in Japan. Species Diversity, 14 (1), 61–67.

    https://doi.org/10.12782/specdiv.14.61

    Schmidt-Rhaesa, A., Nishi, H., Tanabe, A.S. & Urabe, M. (2009) New records of hairworms (Nematomorpha: Gordiida) from Japan. Species Diversity, 14, 131–135.

    https://doi.org/10.12782/specdiv.14.131

    Schmidt-Rhaesa, A. & Gusich, V.V. (2010) A new Gordionus Müller, 1927 from Switzerland (Nematomorpha, Gordiida). Revue Suisse Zoologie, 117, 77–82.

    https://doi.org/10.5962/bhl.part.117589

    Schmidt-Rhaesa, A. & Yadav, A.K. (2013) One new species and a new record of the genus Chordodes (Nematomorpha; Gordiida) from North-East India. Zootaxa, 3693 (2), 293–300.

    https://doi.org/10.11646/zootaxa.3693.2.10

    Schneider, C.A., Rasband, W.S. & Eliceiri, K.W. (2012) NIH Image to ImageJ: 25 years of Image Analysis. Nature Methods, 9 (7), 671–675.

    https://doi.org/10.1038/nmeth.2089

    Smith, D.G. (1991) Observations on the morphology and taxonomy of two Parachordodes species (Nematomorpha, Gordioida, Chordodidae) in southern New England (USA). Journal of Zoology, 225 (3), 469–480.

    https://doi.org/10.1111/j.1469-7998.1991.tb03829.x

    Snyder, S.D. & Tkach, V.V. (2001) Phylogenetic and biogeographical relationships among some Holarctic frog lung flukes (Digenea: Haematoloechidae). Journal of Parasitology, 87 (6), 1433–1440.

    https://doi.org/10.1645/0022-3395(2001)087[1433:PABRAS]2.0.CO;2

    Spiridonov, S. & Schmatko, V. (2013) Morphological and molecular characterization of Gordionus alpestris (Nematomorpha) from the North-West Caucasus, Russia. Helminthologia, 50 (1), 67–72.

    https://doi.org/10.2478/s11687-013-0109-3

    Swanteson-Franz, R.J., Marquez, D.A., Goldstein, C.I., Schmidt-Rhaesa, A., Bolek, M.G. & Hanelt, B. (2018) New Hairworm (Nematomorpha, Gordiida) species described from the Arizona Madrean Sky Islands. ZooKeys, 733, 131–145.

    https://doi.org/10.3897/zookeys.733.22798

    Swanteson-Franz, R.J., Schmidt-Rhaesa, A., Bolek, M.G. & Hanelt, B. (2020) A new species of Gordionus (Nematomorpha: Gordiida) from the Rocky Mountains of New Mexico. Journal of Parasitology, 106 (4), 471–477.

    https://doi.org/10.1645/18-36

    Swofford, D. L. (2003) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program]

    Szmygiel, C., Schmidt-Rhaesa, A., Hanelt, B. & Bolek, M.G. (2014) Comparative descriptions of non-adult stages of four genera of Gordiids (Phylum: Nematomorpha). Zootaxa, 3768 (2), 101–118.

    https://doi.org/10.11646/zootaxa.3768.2.1

    Tobias, Z.J.C., Yadav, A.K., Schmidt-Rhaesa, A. & Poulin, R. (2017) Intra- and interspecific genetic diversity of New Zealand hairworms (Nematomorpha). Parasitology, 144 (8), 1026–1040.

    https://doi.org/10.1017/S0031182017000233

    Triplehorn, C.A. & Johnson, N.F. (2005) Borror and DeLong’s introduction to the study of insects. Thomson Brooks/Cole, Belmont, California, x + 864 pp.

    de Villalobos, L.C., Ribera, I. & Bilton, D.T. (2001) First data of Iberian Nematomorpha, with Redescription of Gordius aquaticus Linnaeus, G. plicatulus Heinze, Gordionus wolterstorffii (Camerano) and Paragordius tricuspidatus (Dufour). Contributions to Zoology, 70 (2), 73–84.

    https://doi.org/10.1163/18759866-07002002

    Villot, A. (1874) Monographie des Dragonneaux (Genre Gordius, Dujardin), introduction + première partie. Taxonomie et histoire naturelle. Archives Zoologie Expérimentale et généale, 3, 39–72.

    Villot, A. (1885) Nouvelles espéces de gordiens. Bulletin de la Société des Sciences Naturelles de l’Ouest de la France, 3, 43–48.

    White, D.A. (1969) The infection of immature aquatic insects by larval Paragordius (Nematomorpha). Western North American Naturalist, 29 (1), 44.

    Yadav, A.K., Tobias, Z.J.C. & Schmidt-Rhaesa, A. (2018) Gordionus maori (Nematomorpha: Gordiida), a new species of horsehair worm from New Zealand. New Zealand Journal of Zoology, 45 (1), 29–42.

    https://doi.org/10.1080/03014223.2017.1329155

    Yamashita, J., Sato, T. & Watanabe, K. (2017) Hairworm infection and seasonal changes in paratenic hosts in a mountain stream in Japan. Journal of Parasitology, 103 (1), 32–37.

    https://doi.org/10.1645/15-887