Tardigrade Taxa

  • Peter DegmaEmail author
  • Roberto Guidetti
Part of the Zoological Monographs book series (ZM, volume 2)


The list of tardigrade taxa up to a sub-generic level, with the type species of each genus and the numbers of lower taxa in each taxon (known in the end of June 2018) is here presented together with the main environments in which taxa were found.

Each listed taxon is characterized by characters common to all lower taxa using predominantly most recent taxa definitions. Taxa definitions were eventually adjusted using original descriptions.

This review of tardigrade taxa underlines as taxonomy and systematics of the phylum are continuously updated by researchers and new species are constantly discovered. Currently, there are 1265 species (2 of them fossil) of Tardigrada belonging to 129 genera and 28 families including one fossil.



We thank anonymous reviewer for the valuable suggestions which significantly helped us to improve the chapter.


  1. Bartels PJ, Fontoura P, Nelson DR (2018) Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus. Zootaxa 4420:43–70PubMedCrossRefPubMedCentralGoogle Scholar
  2. Bartels PJ, Nelson DR, Kaczmarek Ł, Michalczyk Ł (2011) Ramazzottius belubellus, a new species of Tardigrada (Eutardigrada: Parachela: Hypsibiidae) from the Great Smoky Mountains National Park (North Carolina, USA). Proc Biol Soc Wash 124:23–27CrossRefGoogle Scholar
  3. Bello G, de Zio Grimaldi S (1998) Phylogeny of the genera of the Stygarctidae and related families (Tardigrada: Heterotardigrada). Zool Anz 237:171–183Google Scholar
  4. Bertolani R, Guidetti R, Marchioro T, Altiero T, Rebecchi L, Cesari M (2014) Phylogeny of Eutardigrada: New molecular data and their morphological support lead to the identification of new evolutionary lineages. Mol Phylogenet Evol 76:110–126CrossRefPubMedPubMedCentralGoogle Scholar
  5. Binda MG (1978) Risistemazione di alcuni tardigradi con l'istituzione di un nuovo genere di Oreellidae e della nuova famiglia Archechiniscidae. Animalia 5:307–314Google Scholar
  6. Binda MG, Kristensen RM (1986) Notes on the genus Oreella (Oreellidae) and the systematic position of Carphania fluviatilis Binda, 1978 (Carphanidae fam. nov., Heterotardigrada). Animalia 13:9–20Google Scholar
  7. Biserov VI (1992) A new genus and three new species of tardigrades (Tardigrada: Eutardigrada) from the USSR. Boll Zool 59:95–103CrossRefGoogle Scholar
  8. Boesgaard TM, Kristensen RM (2001) Tardigrades from Australian marine caves. With a redescription of Actinarctus neretinus (Arthrotardigrada). Zool Anz 240:253–264CrossRefGoogle Scholar
  9. Bussau C (1992) New deep sea Tardigrada (Arthrotardigrada, Halechiniscidae) from a manganese nodule area of the eastern South Pacific. Zool Scr 21:79–92CrossRefGoogle Scholar
  10. Cantacuzène MA (1951) Tardigrade marin noveau, commensal de Limnoria lignorum (Rathke). C R Hebd Seances Acad Sci 232:1699–1700PubMedPubMedCentralGoogle Scholar
  11. Cesari M, Vecchi M, Palmer A, Bertolani R, Pilato G, Rebecchi L, Guidetti R (2016) What if the claws are reduced? Morphological and molecular phylogenetic relationships of the genus Haplomacrobiotus May, 1948 (Eutardigrada, Parachela). Zool J Linn Soc Lond 178:819–827CrossRefGoogle Scholar
  12. Claxton SK (1999) Milnesioides exsertum gen. n. sp. n., a new tardigrade from Australia (Tardigrada: Milnesiidae). Zool Anz 238:183–190Google Scholar
  13. Cooper KW (1964) The first fossil tardigrade: Beorn leggi Cooper, from cretaceous amber. Psyche 71:41–48CrossRefGoogle Scholar
  14. Dastych H (1992) Paradiphascon manningi gen. n. sp. n., a new water-bear from South Africa, with the erecting of a new subfamily Diphasconinae (Tardigrada). Mitt Hamb Zool Mus Inst 89:125–139Google Scholar
  15. Dastych H (2009) Thalerius konradi gen. nov., sp. nov., a new tardigrade from the periglacial area of the Ötztal Alps, Austria (Tardigrada). Contrib Nat Hist 12:391–402Google Scholar
  16. Dastych H (2011) Bergtrollus dzimbowski gen. n., sp. n., a remarkable new tardigrade genus and species from the nival zone of the Lyngen Alps, Norway (Tardigrada: Milnesiidae). Entomol Mitt Zool Mus Hamburg 15(186):335–359Google Scholar
  17. Dastych H, McInnes SJ, Claxton SK (1998) Oreella mollis Murray, 1910 (Tardigrada): a redescription and revision of Oreella. Mitt Hamb Zool Mus Inst 95:89–113Google Scholar
  18. D’Addabbo Gallo M, Grimaldi de Zio S, Morone de Lucia MR, Troccoli A (1992) Halechiniscidae and Echiniscoididae from the Western Mediterranean Sea (Tardigrada: Heterotardigrada). Cah Biol Mar 33:299–318Google Scholar
  19. Degma P, Guidetti R (2007) Notes to the current checklist of Tardigrada. Zootaxa 1579:41–53CrossRefGoogle Scholar
  20. Degma P, Bertolani R, Guidetti R (2018) Actual checklist of Tardigrada species, p 48. Accessed 20 Aug 2018
  21. de Zio Grimaldi S, Lamarca A, D'addabbo Gallo M, Pietanza R (1999) Florarctinae of Asdhu Island, Maldives, Indian Ocean (Tardigrada, Heterotardigrada). Ital J Zool 66:383–391CrossRefGoogle Scholar
  22. du Bois-Reymond Marcus E (1952) On South American Malacopoda. Bol Fac Fil Ciên Letr Univ S Paulo Zool 17:189–209Google Scholar
  23. Fontoura P, Bartels PJ, Jørgensen A, Kristensen RM, Hansen JG (2017) A dichotomous key to the genera of the marine Heterotardigrades (Tardigrada). Zootaxa 4294(1):1–45CrossRefGoogle Scholar
  24. Fujimoto S (2015) Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan. Zootaxa 3948:145–150PubMedCrossRefPubMedCentralGoogle Scholar
  25. Fujimoto S, Yamasaki H (2017) A new genus and species of Renaudarctidae (Heterotardigrada: Arthrotardigrada) from Ryukyu Archipelago, Japan. Mar Biol Res 13(3):288–299CrossRefGoogle Scholar
  26. Fujimoto S, Jørgensen A, Hansen JG (2016) A molecular approach to arthrotardigrade phylogeny (Heterotardigrada, Tardigrada). Zool Scr 46:496–505CrossRefGoogle Scholar
  27. Gallo D’Addabbo M, Sandulli R, de Zio Grimaldi S (2005) A new Batillipedidae (Tardigrada, Heterotardigrada) from the Orosei Gulf, Sardinia, Tyrrhenian Sea. Zool Anz 243:219–225CrossRefGoogle Scholar
  28. Gąsiorek P, Stec D, Morek W, Michalczyk Ł (2017) An integrative redescription of Echiniscus testudo (Doyère, 1840), the nominal taxon for the class Heterotardigrada (Ecdysozoa: Panarthropoda: Tardigrada). Zool Anz 270:107–122CrossRefGoogle Scholar
  29. Gąsiorek P, Stec D, Morek W, Michalczyk Ł (2018) An integrative redescription of Hypsibius dujardini (Doyère, 1840), the nominal taxon for Hypsibioidea (Tardigrada: Eutardigrada). Zootaxa 4415:45–75PubMedCrossRefGoogle Scholar
  30. Gomes-Júnior E, Santos É, da Rocha CMC, Santos PJP, Fontoura P (2018) A new species of Ligiarctus (Tardigrada, Arthrotardigrada) from the Brazilian continental shelf, Southwestern Atlantic Ocean. Mar Biodivers 48:5–12CrossRefGoogle Scholar
  31. Grimaldi de Zio S, D’Addabbo Gallo M, Morone De Lucia MR (1990) Revision of the genus Halechiniscus (Halechiniscidae, Arthrotardigrada). Cah Biol Mar 31:271–279Google Scholar
  32. Gross V, Miller WR, Hochberg R (2014) A new tardigrade, Mutaparadoxipus duodigifinis gen. nov., sp. nov. (Heterotardigrada: Arthrotardigrada), from the Southeastern United States. Zootaxa 3835:263–272PubMedCrossRefPubMedCentralGoogle Scholar
  33. Grothman GT, Johansson C, Chilton G, Kagoshima H, Tsujimoto M, Suzuki AC (2017) Gilbert Rahm and the status of Mesotardigrada Rahm, 1937. Zool Sci 34:5–10CrossRefGoogle Scholar
  34. Guidetti R, Bertolani R (2001) An evolutionary line of the Macrobiotinae (Tardigrada, Macrobiotidae): Calcarobiotus and related species. Ital J Zool 68(3):229–233CrossRefGoogle Scholar
  35. Guidetti R, Bertolani R (2005) Tardigrade taxonomy: an updated check list of the taxa and a list of characters for their identification. Zootaxa 845:1–46CrossRefGoogle Scholar
  36. Guidetti R, Pilato G (2003) Revision of the genus Pseudodiphascon (Tardigrada, Macrobiotidae), with the erection of three new genera. J Nat Hist 37:1679–1690CrossRefGoogle Scholar
  37. Guidetti R, Altiero T, Marchioro T, Amadè LS, Avdonina AM, Bertolani R, Rebecchi L (2012) Form and function of the feeding apparatus in Eutardigrada (Tardigrada). Zoomorphology 131(2):127–148CrossRefGoogle Scholar
  38. Guidetti R, Rebecchi L, Bertolani R, Jönsson KI, Kristensen RM, Cesari M (2016) Morphological and molecular analyses on Richtersius (Eutardigrada) diversity reveal its new systematic position and lead to the establishment of a new genus and a new family within Macrobiotoidea. Zool J Linn Soc Lond 178:834–845CrossRefGoogle Scholar
  39. Guil N, Rodrigo E, Machordom A (2015) Soil tardigrade biodiversity with the description of a new eutardigrade genus and its phylogenetic position. Syst Biodivers 13:234–256CrossRefGoogle Scholar
  40. Hansen JG (2007) The deep sea elements of the Faroe Bank tardigrade fauna with a description of two new species. J Limnol 66:12–20CrossRefGoogle Scholar
  41. Hansen JG, Kristensen RM (2006) The ‘hyena female’of tardigrades and descriptions of two new species of Megastygarctides (Arthrotardigrada: Stygarctidae) from Saudi Arabia. Hydrobiologia 558(1):81–101CrossRefGoogle Scholar
  42. Hansen JG, D’Addabbo Gallo M, de Zio Grimaldi S (2003) A comparison of morphological characters whitin the genus Rhomboarctus (Tardigrada: Heterotardigrada) with the description of two new species. Zool Anz 242:83–96CrossRefGoogle Scholar
  43. Hansen JG, Kristensen RM, Jørgensen A (2012) The armoured marine tardigrades (Arthrotardigrada, Tardigrada). Scientia Danica, Series B Biologica, vol 2. The Royal Danish Academy of Sciences and Letters, p 91Google Scholar
  44. Jørgensen A, Kristensen RM (2001) A new tanarctid arthrotardigrade with buoyant bodies. Zool Anz 240:425–439CrossRefGoogle Scholar
  45. Jørgensen A, Boesgaard TM, Møbjerg N, Kristensen RM (2014) The tardigrade fauna of Australian marine caves: With descriptions of nine new species of Arthrotardigrada. Zootaxa 3802:401–443CrossRefGoogle Scholar
  46. Kaczmarek Ł, Michalczyk Ł (2006) Redescription of Macrobiotus tridigitus Schuster, 1983 and the erection of a new genus of Tardigrada (Eutardigrada, Macrobiotidae). J Nat Hist 40:1223–1229CrossRefGoogle Scholar
  47. Kaczmarek Ł, Gawlak M, Bartels PJ, Nelson DR, Roszkowska M (2017) Revision of the genus Paramacrobiotus Guidetti et al., 2009 with the description of a new species, re-descriptions and a key. Ann Zool 67:627–656CrossRefGoogle Scholar
  48. Kristensen RM (1982a) New aberrant eutardigrades from homothermic springs on Disko Island, West Greenland. In: Proceedings of the 3rd international symposium on the Tardigrada, Johnson City, TN, USA, pp 203–220Google Scholar
  49. Kristensen RM (1982b) The first record of cyclomorphosis in Tardigrada based on a new genus and species from Arctic meiobenthos. J Zool Syst Evol Res 20:249–270CrossRefGoogle Scholar
  50. Kristensen RM (1984) On the biology of Wingstrandarctus corallinus nov. gen. et spec., with notes on the symbiontic bacteria in the subfamily Florarctinae (Arthrotardigrada). Vidensk Meddr Dansk Naturh Foren 145:201–218Google Scholar
  51. Kristensen RM (1987) Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. In: Bertolani R (ed) Biology of Tardigrades. Selected Symposia and Monographs U.Z.I. 1. Mucchi, Modena, pp 261–335Google Scholar
  52. Kristensen RM, Hallas TE (1980) The tidal genus Echiniscoides and its variability, with erection of Echiniscoididae fam. n. (Tardigrada). Zool Scr 9:113–127CrossRefGoogle Scholar
  53. Kristensen RM, Hansen JG (2005) Tardigrada (water bears). Chapter 5 Minor groups and fossils. In: Rohde K (ed) Marine parasitology. CSIRO, Melbourne, pp 230–235Google Scholar
  54. Kristensen RM, Higgins RP (1984) Revision of Styraconyx (Tardigrada: Halechiniscidae), with descriptions of two new species from Disko Bay, West Greenland. Smithson Contr Zool 391:1–40CrossRefGoogle Scholar
  55. Kristensen RM, Higgins RP (1989) Marine Tardigrada from the Southeastern United States coastal waters. I. Paradoxipus orzeliscoides n. gen., n. sp. (Arthrotardigrada: Halechiniscidae). Trans Am Microsc Soc 108:262–282CrossRefGoogle Scholar
  56. Kristensen RM, Renaud-Mornant J (1983) Existence d’Arthrotardigrades semi-benthiques de genres nouveaux de la sous-famille des Styraconyxinae, subfam. nov. Cah Biol Mar 24:337–353Google Scholar
  57. Kristensen RM, Sørensen MV, Hansen JG, Zeppilli D (2015) A new species of Neostygarctus (Arthrotardigrada) from the Condor Seamount in the Azores, Northeast Atlantic. Mar Biodivers 45:453–467CrossRefGoogle Scholar
  58. Lisi O (2011) Remarks on Doryphoribius flavus (Iharos, 1966), and description of three new species (Tardigrada, Hypsibiidae). Zootaxa 2834:17–32CrossRefGoogle Scholar
  59. Lisi O, Sabella G, Pilato G (2014) Mixibius parvus sp. nov. and Diphascon (Diphascon) ziliense sp. nov., two new species of Eutardigrada from Sicily. Zootaxa 3802:459–468CrossRefGoogle Scholar
  60. Lisi O, Daza A, Londoño R, Quiroga S (2017) Echiniscidae from the Sierra Nevada de Santa Marta, Colombia, new records and a new species of Bryodelphax Thulin, 1928 (Tardigrada). ZooKeys 703:1–14CrossRefGoogle Scholar
  61. Marley NJ, McInnes SJ, Sands CJ (2011) Phylum Tardigrada: A re-evaluation of the Parachela. Zootaxa 2819:51–64CrossRefGoogle Scholar
  62. Marley NJ, Kaczmarek Ł, Gawlak M, Bartels PJ, Nelson DR, Roszkowska M, Stec D, Degma P (2018) A clarification for the subgenera of Paramacrobiotus Guidetti, Schill, Bertolani, Dandekar and Wolf, 2009, with respect to the International Code of Zoological Nomenclature. Zootaxa 4407:130–134PubMedCrossRefPubMedCentralGoogle Scholar
  63. Miller WR, Schulte R, Johansson C (2012) Tardigrades of North America: further description of the genus Multipseudechiniscus Schulte and Miller, 2011 (Heterotardigrada: Echiniscoidea: Echiniscidae) from California. Proc Biol Soc Wash 125:153–164CrossRefGoogle Scholar
  64. Møbjerg N, Kristensen RM, Jørgensen A (2016) Data from new taxa infer Isoechiniscoides gen. nov. and increase the phylogenetic and evolutionary understanding of echiniscoidid tardigrades (Echiniscoidea: Tardigrada). Zool J Linn Soc Lond 178:804–818CrossRefGoogle Scholar
  65. Nelson DR, Marley NJ, Bertolani R (1999) Re-description of the genus Pseudobiotus (Eutardigrada, Hypsibiidae) and of the new type species Pseudobiotus kathmanae sp. n. Zool Anz 238:311–317Google Scholar
  66. Pilato G (1972) Structure, intraspecific variability and systematic value of the buccal armature of eutardigrades. J Zool Syst Evol 10:65–78CrossRefGoogle Scholar
  67. Pilato G (2013) The taxonomic value of the structures for the insertion of the stylet muscles in the Eutardigrada, and description of a new genus. Zootaxa 3721:365–378PubMedCrossRefPubMedCentralGoogle Scholar
  68. Pilato G, Binda MG (2010) Definition of families, subfamilies, genera and subgenera of the Eutardigrada, and keys to their identification. Zootaxa 2404:1–54CrossRefGoogle Scholar
  69. Pilato G, Lisi O (2011) Tenuibiotus, a new genus of Macrobiotidae (Tardigrada). Zootaxa 2761:34–40CrossRefGoogle Scholar
  70. Pilato G, Lisi O, Binda MG (2010) Tardigrades of Israel with description of four new species. Zootaxa 2665:1–28CrossRefGoogle Scholar
  71. Pilato G, Kiosya Y, Lisi O, Inshina V, Biserov V (2011) Annotated list of Tardigrada records from Ukraine with the description of three new species. Zootaxa 3123:1–31Google Scholar
  72. Pilato G, Kiosya Y, Lisi O, Sabella G (2012) New records of Eutardigrada from Belarus with the description of three new species. Zootaxa 3179:39–60Google Scholar
  73. Pollock LW (1983) A closer look at some marine Heterotardigrada II. The morphology and taxonomy of Bathyechiniscus, with a description of B. craticulus n. sp. from the Caribbean. Bull Mar Sci 33:109–117Google Scholar
  74. Pollock LW (1989) Marine interstitial Heterotadrigrada from the Pacific coast of the United States, including a description of Batillipes tridentatus n. sp. Trans Am Microsc Soc 108:169–189CrossRefGoogle Scholar
  75. Ramazzotti G, Maucci W (1983) Il phylum Tardigrada (III edizione riveduta e aggiornata). Mem Ist Ital Idrobiol 41:1–1016Google Scholar
  76. Rebecchi L, Altiero T, Eibye-Jacobsen J, Bertolani R, Kristensen RM (2008) A new discovery of Novechiniscus armadilloides (Schuster, 1975) (Tardigrada, Echiniscidae) from Utah, USA with considerations on non-marine Heterotardigrada phylogeny and biogeography. Org Divers Evol 8:58–65CrossRefGoogle Scholar
  77. Renaud-Mornant J (1983) Tardigrades abyssaux nouveaux de la sous-famille des Euclavarctinae n. subfam. (Arthrotardigrada, Halechiniscidae). Bull Mus Natn Hist Nat Paris 5:201–219Google Scholar
  78. Renaud-Mornant J (1984) Halechiniscidae (Heterotardigrada) de la campagne Benthedi, canal du Mozambique. Bull Mus Natn Hist Nat Paris 6:67–88Google Scholar
  79. Renaud-Mornant J (1987) Halechiniscidae nouveaux de sables coralliens tropicaux (Tardigrada, Arthrotardigrada). Bull Mus Natn Hist Nat Paris, 4e Serie Miscellanea 9:353–373Google Scholar
  80. Renaud-Mornant J (1989) Opydorscus, un nouveau genre d’Orzeliscinae et sa signification phylogénique (Tardigrada, Arthrotardigrada). Bull Mus Natn Hist Nat Paris, 4e Serie Miscellanea 11:763–771Google Scholar
  81. Richters F (1926) Tardigrada. Handbuch der Zoologie, III, pp 1–68Google Scholar
  82. Rubal M, Veiga P, Fontoura P, Sousa-Pinto I (2013) A new intertidal arthrotardigrade, Prostygarctus aculeatus gen. nov., sp. nov. (Tardigrada: Heterotardigrada) from the North of Portugal (Atlantic ocean). Proceedings of the 12th international symposium on Tardigrada. J Limnol 72(Suppl. 1):8–14Google Scholar
  83. Trygvadóttir BV, Kristensen RM (2011) Eohypsibiidae (Eutardigrada, Tardigrada) from the Faroe Islands with the description of a new genus containing three new species. Zootaxa 2886:39–62CrossRefGoogle Scholar
  84. Van der Land J (1975) The parasitic marine tardigrade Tetrakentron synaptae. Mem Ist Ital Idrobiol 32 Suppl.:413–423Google Scholar
  85. Vecchi M, Cesari M, Bertolani R, Jönsson KI, Rebecchi L, Guidetti R (2016) Integrative systematic studies on tardigrades from Antarctica identify new genera and new species within Macrobiotoidea and Echiniscoidea. Invertebr Syst 30:303–322CrossRefGoogle Scholar
  86. Vicente F, Fontoura P, Cesari M, Rebecchi L, Guidetti R, Serrano A, Bertolani R (2013) Integrative taxonomy allows the identification of synonymous species and the erection of a new genus of Echiniscidae (Tardigrada, Heterotardigrada). Zootaxa 3613:557–572PubMedPubMedCentralCrossRefGoogle Scholar
  87. Villora-Moreno S (1996) A new genus and species of the deep-sea family Coronarctidae (Tardigrada) from a submarine cave with a deep-sea like condition. Sarsia 81:275–283CrossRefGoogle Scholar
  88. Zawierucha K, Stec D, Lachowska-Cierlik D, Takeuchi N, Li Z, Michalczyk Ł (2018) High mitochondrial diversity in a new water bear species (Tardigrada: Eutardigrada) from mountain glaciers in central Asia, with the erection of a new genus Cryoconicus. Ann Zool 68:179–201CrossRefGoogle Scholar

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© Springer Nature Switzerland AG 2018

Authors and Affiliations

  1. 1.Department of ZoologyComenius University in BratislavaBratislavaSlovakia
  2. 2.Department of Life SciencesUniversity of Modena and Reggio EmiliaModenaItaly

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