, Volume 558, Issue 1, pp 53–60 | Cite as

A Family Level Analysis of Tardigrade Phylogeny

  • P. Brent Nichols
  • Diane R. Nelson
  • James R. Garey


In the present study a character data set suitable for cladistic analysis at the family level was developed. A data matrix consisting of 50 morphological characters from 15 families of tardigrades was analyzed by maximum parsimony. Kinorhynchs, loriciferans, and gastrotrichs were used as outgroups. The results agree with the currently accepted hypothesis that Eutardigrada and Heterotardigrada are distinct monophyletic groups. Among the eutardigrades, Eohypsibiidae was found to be a sister group to Macrobiotidae+Hypsibiidae, while Milnesiidae was the basal eutardigrade family. The basal heterotardigrade family was found to be Oreellidae. Echiniscoideans grouped with some traditional Arthrotardigrada (Renaudarctidae, Coronarctidae+Batillipedidae) suggesting that the arthrotardigrades are not monophyletic. The 18S rRNA gene sequence of Batillipes mirus Richters, 1909 and Calohypsibius schusteri Nelson & McGlothlin, 1996 were obtained and their addition to a previously published dataset supports the monophyly of Heterotardigrada and of Parachela versus Apochela within the Eutardigrada.


tardigrada systematics cladistics evolution phylogeny 18S rRNA 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Aguinaldo, A. M., Turbeville, J. M., Linford, L. S., Rivera, M. C., Garey, J. R., Raff, R. A., Lake, J. A. 1997Evidence for a clade of nematodes, arthropods, and other moulting animalsNature387489493PubMedCrossRefGoogle Scholar
  2. Bertolani, R. 2003Thulinius, new generic name substituting for Thulinia Bertolani, 1981 (Tardigrada, Eutardigrada)Zootaxa31414Google Scholar
  3. Bertolani, R., Biserov, V. I. 1996Leg and claw adaptations in soil tardigrades, with erection of two new genera of Eutardigrada, Macrobiotidae: Pseudohexapodibius and XerobiotusInvertebrate Biology115299304Google Scholar
  4. Bertolani, R., Rebecchi, L. 1993A revision of the Macrobiotus hufelandi group (Tardigrada, Macrobiotidae), with some observations on the taxonomic character of eutardigradesZoologica Scripta22127152CrossRefGoogle Scholar
  5. Binda, M. G., Kristensen, R. M. 1987Notes on the genus Carphania fluviatilis Binda, 1978 (Carphanidae fam. nov.), HeterotardigradaAnimalia13920Google Scholar
  6. Blaxter, M. L., Ley, P., Garey, J. R., Liu, L. X., Scheldeman, P., Vierstraete, A., Vanfleteren, J. R., Mackey, L. Y., Dorris, M., Frisse, L. M., Vida, J. T., Thomas, W. K. 1998A molecular evolutionary framework for the phylum NematodaNature3927175PubMedCrossRefGoogle Scholar
  7. Rosa, R., Grenier, J. K., Andreeva, T., Cook, C., Adoutte, A., Akam, M., Carroll, S. B., Balavoine, G. 1999Hox genes in brachiopods and priapulids and protostome evolutionNature399772776PubMedGoogle Scholar
  8. Dewel, R. A., Clark, W. H. 1973aStudies on the tardigrades. II. Fine structure of the pharynx of Milnesium tardigradum DoyèreTissue and Cell5147159Google Scholar
  9. Dewel, R. A., Clark, W. H. 1973bStudies on the tardigrades. III. Fine structure of the sophagus of Milnesium tardigradum DoyèreTissue and Cell5161169Google Scholar
  10. Eibye-Jacobsen, J. 2001Are the supportive structures of the tardigrade pharynx homologous throughout the entire group?Journal of Zoological Systematics and Evolutionary Research39111CrossRefGoogle Scholar
  11. Garey, J. R., Nelson, D. R., Mackey, L. Y., Li, J. 1999Tardigrade phylogeny: congruency of morphological and molecular evidenceZoologischer Anzeiger238205210Google Scholar
  12. Garey, J. R., Krotec, M., Nelson, D. R., Brooks, J. 1996Molecular analysis supports a tardigrade arthropod associationInvertebrate Biology1157988Google Scholar
  13. Giribet, G., Carranza, S., Baguna, J., Riutort, M., Ribera, C. 1996First molecular evidence for the existence of a Tardigrada+Arthropoda cladeMolecular Biology and Evolution137684PubMedGoogle Scholar
  14. Grenier, J. K., Carroll, S. B. 2000Functional evolution of the Ultrabithorax proteinProceedings of the National Academy of Sciences of the United States of America97704709PubMedCrossRefGoogle Scholar
  15. Guidetti, R., Bertolani, R. 2001Phylogenetic relationships in the Macrobiotidae (Tardigrada: Eutardigrada: Parachela)Zoologischer Anzeiger240371376Google Scholar
  16. Jorgensen, A. 2000Cladistic analysis of the Echiniscidae Thulin, 1928 (Tardigrada: Heterotardigrada: Echiniscoidea)Streenstrupia251123Google Scholar
  17. Kinchin, I. M. 1994The Biology of TardigradesPortland PressLondonGoogle Scholar
  18. Kristensen, R. M. 1987Generic revision of the Echiniscidae (Herterotardigrada), with a discussion of the origin of the familyBertolani, R. eds. Selected Symposia and Monographs U.Z.I., 1MucchiModena261335Google Scholar
  19. Kristensen, R. M. 1991LoriciferaBarnes, R. D.Harrison, F. W. eds. Microscopic Anatomy of Invertebrates, Volume 4: AschelminthesWiley-LissNew York351375Google Scholar
  20. Kristensen, R. M., Higgins, R. P. 1984A new family of Arthrotardigrada (Tardigrada: Heterotardigrada) from the Atlantic Coast of Florida, U.S.ATransactions of American Microscopical Society103295311Google Scholar
  21. Kumar, S., Tamura, K. I., Jakobsen, I. B., Nei, M. 2001MEGA2: Molecular Evolutionary Genetics Analysis softwareBioinformatics1212441245Google Scholar
  22. Mallat, J. M., Winchell, C. J. 2002Testing the new animal phylogeny: first use of combined large-subunit and small-subunit rRNA gene sequences to classify the protostomesMolecular Biology and Evolution19289301Google Scholar
  23. Mallat, J. M., Garey, J. R., Shultz, J. W. 2004Ecdysozoan phylogeny and Bayesian inference: first use of nearly complete 28S and 18S rRNA gene sequences to classify the arthropods and their kinMolecular Phylogenetics and Evolution31178191Google Scholar
  24. Marcus, E., 1929. Tardigrada. In Bronn, H.G. (ed), Klassen und Ordnungen des tierreichs. Vol 5, Section 4, Part 3: 1–608Google Scholar
  25. Neefs, J. M., Peer, Y., Ruijk, P., Chapelle, S., Wachter, R. 1993Compilation of small Ribosomal subunit RNA sequencesNucleic Acids Research2130253049PubMedGoogle Scholar
  26. Nelson, D. 2002Current status of the Tardigrada: evolution and ecologyIntegrative and Comparative Biology42652659Google Scholar
  27. Panganiban, G., Irvine, S. M., Lowe, C., Roehl, H., Corley, L. S., Sherbon, B., Grenier, J. K., Fallon, J. F., Kimble, J., Walker, M., Wray, G. A., Swalla, B. J., Martindale, M. Q., Carroll, S. B. 1997The origin and evolution of animal appendagesProceedings of the National Academy of Sciences of the United States of America9451625166PubMedCrossRefGoogle Scholar
  28. Pollock, L. W. 1995New marine tardigrades from Hawaiian beach sand and phylogeny of the family HalechiniscidaeInvertebrate Biology114220235Google Scholar
  29. Ramazzotti, G., W. Maucci, 1983. Il Phylum Tardigrada, 3rd ed (C. W. Beasley, English Translation). Memorie dell’Istitato idrobiologia 41: 1–985Google Scholar
  30. Renaud-Mornant, J. 1982Species diversity in marine TardigradaNelson, D. R. eds. Proceedings of the Third International Symposium on TardigradaEast Tennessee State University PressJohnson City, Tennessee149178Google Scholar
  31. Schmidt-Rhaesa, A., Bartolomaeus, T., Lemburg, C., Ehlers, U., Garey, J. R. 1998The position of the arthropoda in the phylogenetic systemJournal of Morphology238263285CrossRefGoogle Scholar
  32. Schuster, R. O., Nelson, D. R., Grigarick, A. A., Christenberry, D. 1980Systematic criteria of the EutardigradaTransactions of the American Microscopical Society99284303Google Scholar
  33. Swofford, D. L., 1998. PAUP. Phylogenetic Analysis Using Parsimony (and Other Methods). Version 4. Sinauer Associates, Sunderland, MAGoogle Scholar
  34. Thulin, G. 1928Ûber die phylogenie und das system der tardigradenHereditas11207Google Scholar

Copyright information

© Springer 2006

Authors and Affiliations

  • P. Brent Nichols
    • 1
  • Diane R. Nelson
    • 2
  • James R. Garey
    • 1
  1. 1.Department of BiologyUniversity of South FloridaTampaUSA
  2. 2.Department of Biological SciencesEast Tennessee State UniversityJohnson CityUSA

Personalised recommendations