Professional Documents
Culture Documents
Introduction
More than one thousand Tardigrada species were included in the published checklist
(Guidetti, R. & Bertolani, R. 2005. Tardigrade taxonomy: an updated check list of the taxa
and a list of characters for their identification. Zootaxa, 845, 146.) plus the additions and
corrections to this checklist (Degma, P. & Guidetti, R. 2007. Notes to the current checklist of
Tardigrada. Zootaxa, 1579, 4153.). For practical reasons, we have joined these two papers
(without comments added to particular taxa as well as without references published in these
papers) into an accurate combined version of the checklists.
This checklist is free for all users, but utilization of it requires the citation of the two
original papers. This checklist is the platform for occasional upgrades (date of latest upgrade is in
the title). All changes, additions and corrections in comparison with both papers are written in red
font, the changings respect to the previous version of the checklist have yellow background.
If you also use these changes, please also cite this checklist (Degma, P., Bertolani, R. &
Guidetti, R. Actual checklist of Tardigrada species. http://www.tardigrada.modena.unimo.it
/miscellanea/Actual%20checklist%20of%20Tardigrada.pdf, pp. 40. Accessed date).
Please write us if you find any mistake or missing data in this checklist. Your help in its
improvement will be acknowledged.
Acknowledgements
We are very grateful to Professor Clark Beasley (McMurry University, Abilene, Texas, U.S.A.)
for his suggestions as well as for help with improving the English in the last version of the
manuscript. We also thank to Nigel Marley (University of Plymouth, U.K.), Gary Grothman (St.
Marys University, Calgary, Canada), Paulo Fontoura (University of Porto, Portugal), Harry
Meyer (McNeese State University, Lake Charles, Louisiana, U.S.A.) and Sandra McInnes
(British Antarctic Survey, U.K.) for their valuable comments to this checklist.
This study was partly supported by the Slovak Scientific Grant Agency (VEGA) (grant No.
1/3265/06 and 1/0294/09 to PD), by University of Modena and Reggio Emilia (50% FAR) and by
Fondazione Cassa di Risparmio di Modena (MoDNA project: Morphology and DNA) to RB and
RG.
Checklist of Tardigrada
(The assignment of synonyms are reported within square parenthesis [ ]. Amendments are
reported within angle brackets < > . Comments are reported within brace parenthesis { } .
The species written in bold characters are the genus type species)
TARDIGRADA Doyre, 1840
HETEROTARDIGRADA Marcus, 1927
ARTHROTARDIGRADA Marcus, 1927
Archechiniscidae Binda, 1978
<Jrgensen et al. 2010 reconsider valid the family>
[Archechiniscinae Grimaldi de Zio & DAddabbo Gallo, 1987]
Archechiniscus Schulz, 1953
Archechiniscus biscaynei Miller, Clark & C. Miller, 2012
Archechiniscus marci Schulz, 1953
Archechiniscus minutus Grimaldi de Zio & D'Addabbo Gallo, 1987
Archechiniscus symbalanus Chang & Rho, 1998
Batillipedidae Ramazzotti, 1962
[Discopodidae Marcus, 1934]
Batillipes Richters, 1909
Batillipes acaudatus Pollock, 1971
Batillipes adriaticus Grimaldi de Zio, Morone De Lucia, D'Addabbo Gallo & Grimaldi, 1979
Batillipes africanus Morone De Lucia, DAddabbo Gallo & Grimaldi de Zio, 1988
Batillipes annulatus de Zio, 1962
Batillipes bullacaudatus McGinty & Higgins, 1968
Batillipes carnonensis Fize, 1957
Batillipes crassipes Tchesunov & Mokievsky, 1995
Batillipes dicrocercus Pollock, 1970
Batillipes friaufi Riggin, 1962
Batillipes gilmartini McGinty, 1969
Batillipes lesteri Kristensen & Mackness, 2000
Batillipes littoralis Renaud-Debyser, 1959
Batillipes longispinosus Chang & Rho, 1997
Batillipes marcelli Morone De Lucia, D'Addabbo Gallo & Grimaldi de Zio, 1988
Batillipes mirus Richters, 1909 [Batillipes caudatus Hay, 1917]
Batillipes noerrevangi Kristensen, 1978
Batillipes orientalis Chang & Rho, 1997
Batillipes pennaki Marcus, 1946
Batillipes philippinensis Chang & Rho, 1997
Batillipes phreaticus Renaud Debyser, 1959 [Batillipes littoralis submersus D'Hondt, 1970]
Batillipes roscoffensis Kristensen, 1978
2
Paratanarctus D'Addabbo Gallo, Grimaldi de Zio, Morone De Lucia & Troccoli, 1992
Paratanarctus kristenseni D'Addabbo Gallo, Grimaldi de Zio, Morone De Lucia & Troccoli,
1992
Pleocola Cantacuzne, 1951
Pleocola limnoriae Cantacuzne, 1951
Raiarctus Renaud-Mornant, 1981
<Amended by Jrgensen et al. 2014>
Raiarctus aureolatus Renaud-Mornant, 1981
Raiarctus colurus Renaud-Mornant, 1981
Raiarctus jesperi Jrgensen, Boesgaard, Mbjerg & Kristensen, 2014
Raiarctus katrinae Jrgensen, Boesgaard, Mbjerg & Kristensen, 2014
Raiarctus variabilis DAddabbo Gallo, Grimaldi de Zio & Morone De Lucia, 1986
Rhomboarctus Renaud-Mornant, 1984
Rhomboarctus aslaki Hansen, Gallo DAddabbo & de Zio Grimaldi, 2003
Rhomboarctus duplicicaudatus Hansen, Gallo DAddabbo & de Zio Grimaldi, 2003
Rhomboarctus thomassini Renaud-Mornant, 1984
Styraconyx Thulin, 1942
Styraconyx craticuliformis Chang & Rho, 1998
Styraconyx craticulus (Pollock, 1983)
Styraconyx hallasi Kristensen, 1977
Styraconyx haploceros Thulin, 1942
Styraconyx kristenseni kristenseni Renaud-Mornant, 1981
Styraconyx kristenseni neocaledonensis Renaud-Mornant, 1981
Styraconyx nanoqsunguak Kristensen & Higgins, 1984
Styraconyx paulae Robotti, 1971
Styraconyx qivitoq Kristensen & Higgins, 1984
Styraconyx sardiniae D'Addabbo Gallo, Morone De Lucia & Grimaldi de Zio, 1989
Styraconyx sargassi Thulin, 1942
Styraconyx testudo D'Addabbo Gallo, Grimaldi de Zio & Morone De Lucia, 1984
Styraconyx tyrrhenus D'Addabbo Gallo, Morone De Lucia & Grimaldi de Zio, 1989
Tetrakentron Cunot, 1892
Tetrakentron synaptae Cunot, 1892
Tholoarctus Kristensen & Renaud-Mornant, 1983
<Amended by Jrgensen et al. 2014>
Tholoarctus natans natans Kristensen & Renaud-Mornant, 1983
Tholoarctus natans pedunculatus D'Addabbo Gallo, de Zio Grimaldi, Morone De Lucia &
Troccoli, 1992
Tholoarctus oleseni Jrgensen, Boesgaard, Mbjerg & Kristensen, 2014
Tanarctinae Renaud-Mornant, 1980
6
16
Pilatobius rugosus (Barto, 1935) <Transferred from Diphascon by Bertolani et al. 2014>
Pilatobius secchii (Bertolani & Rebecchi, 1996) <Transferred from Diphascon by Bertolani et al.
2014>
Pilatobius sexbullatus (Ito, 1995) <Transferred from Diphascon by Bertolani et al. 2014>
Pilatobius trachydorsatus (Barto, 1937) <Transferred from Diphascon by Bertolani et al. 2014>
Incerta subfamilia {according to Bertolani et al. 2014}
Acutuncus Pilato & Binda, 1997
<Transferred from Hypsibiinae by Bertolani et al. 2014>
Acutuncus antarcticus (Richters, 1904) [Hypsibius simoizumii Sudzuki, 1964]
Mixibius Pilato, 1992
{to be confirmed in the family Isohypsibiidae, see Marley et al. 2011}
<Transferred from Isohypsibiidae by Bertolani et al. 2014>
Mixibius fueginus Pilato & Binda, 1996
Mixibius ninguidus Biserov, 1999
Mixibius ornatus Pilato, Binda, Napolitano & Moncada, 2002
Mixibius parvus Lisi, Sabella & Pilato, 2014
Mixibius pilatoi L. Wang, 2009
Mixibius saracenus (Pilato, 1973)
Mixibius schnurae Pilato, Lisi & Binda, 2010
Mixibius sutirae Pilato, Binda & Lisi 2004
Mixibius tibetanus H. Li & X. Li, 2008
Microhypsibiidae Pilato, 1998
Fractonotus Pilato, 1998
Fractonotus caelatus (Marcus, 1928)
Microhypsibius Thulin, 1928
Microhypsibius bertolanii Kristensen, 1982
Microhypsibius japonicus Ito, 1991
Microhypsibius minimus Kristensen, 1982
Microhypsibius truncatus Thulin, 1928
Ramazzottiidae Sands, McInnes, Marley, Goodall-Copestake, Convey & Linse, 2008
{Sands et al. (2008) reported as taxon name Ramazzottidae but the correct name is
Ramazzottiidae in compliance with the International Code of Zoological Nomenclature}
Hebesuncus Pilato, 1987
Hebesuncus conjungens (Thulin, 1911)
Hebesuncus mollispinus Pilato, McInnes & Lisi, 2012
Hebesuncus ryani Dastych & Harris, 1994
Hebesuncus schusteri (Dastych, 1984)
Ramajendas Pilato & Binda, 1990
23
2010>
Eremobiotus Biserov, 1992
Eremobiotus alicatai (Binda, 1969)
Eremobiotus ovezovae Biserov, 1992
Halobiotus Kristensen, 1982
Halobiotus arcturulius Crisp & Kristensen, 1983
Halobiotus crispae Kristensen, 1982
Halobiotus stenostomus (Richters, 1908) [Macrobiotus appelloefi Richters, 1908; Hypsibius
(Isohypsibius) geddesi Hallas, 1971]
Haplohexapodibius Pilato & Beasley, 1987
<Transferred from Calohypsibiidae by Bertolani et al. 2014>
Haplohexapodibius seductor Pilato & Beasley, 1987 [Hexapodibius beasleyi Maucci, 1988]
Haplomacrobiotus May, 1948
<Transferred from Calohypsibiidae by Bertolani et al. 2014>
Haplomacrobiotus hermosillensis May, 1948
Haplomacrobiotus utahensis Pilato & Beasley, 2005
Hexapodibius Pilato, 1969
<Transferred from Calohypsibiidae by Bertolani et al. 2014>
Hexapodibius bindae Pilato, 1982
Hexapodibius boothi Dastych & McInnes, 1994
Hexapodibius christenberryae Pilato & Binda, 2003
Hexapodibius micronyx Pilato, 1969
Hexapodibius pseudomicronyx Robotti, 1972
Hexapodibius reginae Vargha, 1995
Isohypsibius Thulin, 1928
Isohypsibius altai Kaczmarek & Michalczyk, 2006
Isohypsibius annulatus annulatus (Murray, 1905)
Isohypsibius annulatus minor (Ramazzotti, 1945)
Isohypsibius arbiter Binda, 1980
Isohypsibius archangajensis Kaczmarek & Michalczyk, 2004
Isohypsibius arcuatus (Barto, 1934)
Isohypsibius asper (Murray, 1906)
Isohypsibius austriacus (Iharos, 1966)
Isohypsibius baicalensis (Ramazzotti, 1966)
Isohypsibius baldii (Ramazzotti, 1945)
Isohypsibius baldiioides Tumanov, 2003
Isohypsibius barbarae Pilato & Binda, 2002
Isohypsibius bartosi (Iharos, 1966)
Isohypsibius basalovoi (Durante & Maucci, 1973)
Isohypsibius belliformis (Miheli, 1971)
Isohypsibius bellus (Miheli, 1971)
26
28
Isohypsibius sattleri (Richters, 1902) [Hypsibius bakonyiensis Iharos, 1964] {the correct
reference of Dastych 1990 cited in the previous check lists is: Dastych, H. (1991)
Isohypsibius sattleri (Richters 1902), a valid species (Tardigrada). Senckenbergiana
biologica, 71, 181189.}
Isohypsibius schaudinni (Richters, 1909)
Isohypsibius sculptus (Ramazzotti, 1962)
Isohypsibius sellnicki (Miheli, 1962)
Isohypsibius septentrionalis Thulin, 1928
Isohypsibius silvicola (Iharos, 1966)
Isohypsibius sismicus (Maucci, 1978)
Isohypsibius solidus (Miheli, 1971)
Isohypsibius taibaiensis X. Li & L. Wang, 2005
Isohypsibius tetradactyloides (Richters, 1907)
Isohypsibius theresiae (Iharos, 1964)
Isohypsibius torulosus (Miheli, 1959)
Isohypsibius truncorum (Iharos, 1964)
Isohypsibius tuberculatus (Plate, 1888)
Isohypsibius tuberculoides (Miheli, 1951)
Isohypsibius tubereticulatus Pilato & Catanzaro, 1989
Isohypsibius tucumanensis Claps & Rossi, 1984
Isohypsibius undulatus Thulin, 1928
Isohypsibius vejdovskyi (Barto, 1939)
Isohypsibius verae Pilato & Catanzaro, 1989
Isohypsibius verrucosus (Della Valle, 1915) [Isohypsibius gibbus (Marcus, 1928)]
Isohypsibius wilsoni (Horning, Schuster & Grigarick, 1978)
Isohypsibius woodsae Kathman, 1990
Isohypsibius yunnanensis Yang, 2002
Isohypsibius zappalai Pilato, Sabella & Lisi, 2015
Paradiphascon Dastych, 1992
<Transferred from Diphasconinae by Bertolani et al. 2014>
Paradiphascon manningi Dastych, 1992
Parhexapodibius Pilato, 1969
<Transferred from Calohypsibiidae by Bertolani et al. 2014>
Parhexapodibius bactrianus Biserov, 1999
Parhexapodibius castrii (Ramazzotti, 1964)
Parhexapodibius lagrecai (Binda & Pilato, 1969)
Parhexapodibius pilatoi (Bernard, 1977)
Parhexapodibius ramazzottii Manicardi & Bertolani, 1987
Pseudobiotus Nelson, 1980
Pseudobiotus hirsutellus Pilato, Lisi & Binda, 2010
Pseudobiotus kathmanae Nelson, Marley & Bertolani, 1999
Pseudobiotus longiunguis (Iharos, 1968)
Pseudobiotus matici (Pilato, 1971)
Pseudobiotus megalonyx (Thulin, 1928)
29
30
36
629.
Christenberry, D. & Mason W.H. (1979) Redescription of Echiniscus virginicus Riggin
(Tardigrada) with notes on life history, range, and geographic variation. Journal of the
Alabama Academy of Science, 50, 4761.
Claxton, S.K., Beasley, C.W., Kaczmarek, L. & Michalczyk, L. (2010) Redescription of
Doryphoribius zyxiglobus (Horning, Schuster & grigarick 1978) (Eutardigrada:
Hypsibiidae) and extension of its geographic range. New Zealand Journal of Zoology, 37,
243253.
Dabert, M., Dastych, H., Hohberg, K., Dabert, J. (2013) Phylogenetic position of the enigmatic
clawless eutardigrade genus Apodibius Dastych, 1983 (Tardigrada), based on 18S and 28S
rRNA sequence data from its type species A. confusus. Molecular Phylogenetics and
Evolution, 70, 7075.
Dastych, H. (2009) Notes on the African limno-terrestrial tardigrade Ramazzottius szeptycki
(Dastych, 1980) (Tardigrada). Entomologische Mitteilungen aus dem Zoologischen
Museum Hamburg, 15, 8791.
Degma, P. (2013) Notes to the species composition of the genus Paramacrobiotus Guidetti et al.,
2009 (Tardigrada, Eutardigrada, Macrobiotidae). Zootaxa, 3681(1), 9698.
Fujimoto, S. & Miyazaki, K. (2013) Neostygarctus lovedeluxe n. sp. from the Miyako Islands,
Japan: The First Record of Neostygarctidae (Heterotardigrada: Arthrotardigrada) from the
Pacific. Zoological Science, 30(5), 414419.
Gross, V., Miller, W.R. & Hochberg, R. (2014) A new tardigrade, Mutaparadoxipus duodigifinis
gen. nov., sp. nov. (Heterotardigrada: Arthrotardigrada), from the Southeastern United
States. Zootaxa, 3835(2), 263272.
Guidetti, R., Schill, R.O., Bertolani, R., Dandekar, T. & Wolf, M. (2009) New molecular data for
tardigrade phylogeny, with the erection of Paramacrobiotus gen. n. Journal of Zoological
Systematics and Evolutionary Research (doi: 10.1111/j.1439-0469.2009.00526.x).
Hansen, J.G., Kristensen, R.M. & Jrgensen, A. (2012) The armoured marine tardigrades
(Arthrotardigrada, Tardigrada). Scientia Danica, series B Biologica 2, The Royal Danish
Academy of Sciences and Letters, pp. 91.
Jrgensen, A., Boesgaard, T.M., Mbjerg, N. & Kristensen, R.M. (2014) The tardigrade fauna of
Australian marine caves: With descriptions of nine new species of Arthrotardigrada.
Zootaxa, 3802(4), 401443.
Jrgensen A, Faurby, S., Hansen, J.G., Mbjerg, N. & Kristensen R.M. (2010) Molecular
phylogeny of Arthrotardigrada (Tardigrada). Molecular Phylogenetics and Evolution, 54,
10061015.
Kaczmarek, . & Michalczyk, . (2009) Redescription of Hypsibius microps Thulin, 1928 and H.
pallidus Thulin, 1911 (Eutardigrada: Hypsibiidae) based on the type material from the
Thulin collection. Zootaxa, 2275, 6068.
Kaczmarek, . & Michalczyk, . (2010) Redescription of Dactylobiotus grandipes (Schuster,
Toftner & Grigarick, 1978), the type species for the genus Dactylobiotus (Tardigrada:
Macrobiotidae). Genus, 21, 155162.
Kaczmarek, ., Bertolani, R., Ndzyska-Stygar, M. (2010) Thulinius saltursus comb. nov.: a
new systematic position for Isohypsibius saltursus Schuster, Toftner & Grigarick, 1978
(Eutardigrada: Hypsibiidae) and a key for the genus Thulinius. Zootaxa, 2483, 5864.
Kristensen, R.M. (1984) On the biology of Wingstrandarctus corallinus nov. gen. et spec., with
notes on the symbiontic bacteria in the subfamily Florarctinae (Arthrotardigrada).
Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 145, 201218.
39
Li, X. (2009) Revision of Biserovus xiae Li, Su and Yu, 2004 (Tardigrada: Macrobiotidae).
Zootaxa, 2076, 6368.
Lisi, O. (2011) Remarks on Doryphoribius flavus (Iharos, 1966), and description of three new
species (Tardigrada, Hypsibiidae). Zootaxa, 2834, 1732.
Marley, N.J., McInnes, S.J. & Sands, C.J. (2011) Phylum Tardigrada: A re-evaluation of the
Parachela. Zootaxa, 2819, 5164.
Michalczyk, ., Wenicz, W., Frohme M. & Kaczmarek, . (2012) Redescriptions of three
Milnesium Doyre, 1840 taxa (Tardigrada: Eutardigrada: Milnesiidae), including the
nominal species for the genus. Zootaxa 3154, 120.
Miller, W.R. & Mehlen, R.H. (2007) Tardigrades of North America: re-description of Echiniscus
tamus Mehlen, 1969 (Tardigrada: Heterotardigrada: Echiniscoidae: Echiniscidae).
Proceedings of the Biological Society of Washington, 120(2), 184188.
Miller, W.R., Schulte, R., Johansson, C. (2012) Tardigrades of North America: further
description of the genus Multipseudechiniscus Schulte and Miller, 2011 (Heterotardigrada:
Echiniscoidea: Echiniscidae) from California. Proceedings of the Biological Society of
Washington, 125(2), 153164.
zdikmen, H. (2008) Bertolanius nom. nov., a replacement name for the genus Amphibolus
Bertolani, 1981 (Tardigrada: Parachela) with type species designation. Munis Entomology
& Zoology, 3(1), 330332.
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(4), 365378.
Pilato, G., Fontoura, P. & Lisi, O. (2008a) New description of Echiniscus viridis Murray, 1910
and remarks on the viridis group. New Zealand Journal of Zoology, 35, 8592.
Pilato, G., Fontoura, P., Lisi, O. & Beasley, C. (2008b) New description of Echiniscus
scabrospinosus Fontoura, 1982, and description of a new species of Echiniscus
(Heterotardigrada) from China. Zootaxa, 1856, 4154 and Zootaxa, 1931, 68.
Pilato, G. & Lisi, O. (2011) Tenuibiotus, a new genus of Macrobiotidae (Tardigrada). Zootaxa,
2761, 3440.
Renaud-Mornant, J. (1989) Espces nouvelles de Florarctinae de l`Atlantique Nord-Est et du
Pacifique Sud (Tardigrada, Arthrotardigrada). Bulletin du Muse d'Histoire Naturelle de
Paris, 11, 571592.
Trygvadttir, B.V. & Kristensen, R.M. (2011) Eohypsibiidae (Eutardigrada, Tardigrada) from the
Faroe Islands with the description of a new genus containing three new species. Zootaxa,
2886, 3962.
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(6),
557572.
40