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Type: Article
Published: 2021-06-01
Page range: 256–268
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An updated species list for “Smoky Bears”: Tardigrades of the Great Smoky Mountains National Park, USA

Department of Biology, Warren Wilson College, Asheville, NC 28815, USA.
Department of Biological Sciences, East Tennessee State University, Johnson City, TN 37614, USA.
Department of Animal Taxonomy and Ecology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61–614 Poznań, Poland.
Tardigrada All Taxa Biodiversity Inventory biodiversity Nearctic North America water bears

Abstract

One of the largest inventories of tardigrades ever conducted occurred from 2000–2010 in the Great Smoky Mountains National Park, USA. Over 16,000 specimens were catalogued, 85 species were identified, 11 species new to science were described, and 16 other possible new species await further study. More than 20 papers have resulted from the GSMNP tardigrade inventory, making the Smokies the most thoroughly studied area in North America for tardigrades. Several species lists have been published over this 20-year period, but many taxonomic revisions and new identifications have led to significant changes to the list. Biogeographical studies citing species records from earlier studies could yield serious errors. Here we update the species list from the Smokies to accommodate the many recent changes in tardigrade taxonomy, we re-analyze some species in light of delineations of cryptic species groups that have occurred recently via integrative taxonomy, and we provide a table of all synonyms that have been used in previous publications. We also make available, for the first time, the Smokies tardigrade database, complete with all locations, elevations, and substrates.

 

References

  1. Arcidiacono, R. (1962) Contributo alla conoscenza dei Tardigradi dei Monti Nebrodi e descrizione di una nuova specie di Itaquascon. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 3, 123–134.

    Bartels, P.J. (2005) Little known water bears. ATBI Quarterly, 6 (2), 4.

    Bartels, P.J. & Nelson, D.R. (2006) A large-scale, multihabitat inventory of the Phylum Tardigrada in the Great Smoky Mountains National Park, USA: A preliminary report. Hydrobiologia, 558 (1), 111–118.

    https://doi.org/10.1007/s10750-005-1405-9

    Bartels, P.J. & Nelson, D.R. (2007) An evaluation of species richness estimators for tardigrades of the Great Smoky Mountains National Park, Tennessee and North Carolina, USA. Journal of Limnology, 66 (Supplement 1), 104–110.

    https://doi.org/10.4081/jlimnol.2007.s1.104

    Bartels, P.J. & Nelson, D.R. (2012) An online key and field guide to the tardigrades of the Great Smoky Mountains National Park (North Carolina & Tennessee, USA, North America) with taxonomic revisions of two species. Zootaxa, 3249 (1), 67–68. [https://keys.lucidcentral.org/keys/v3/smokey_bears/Tardigrades_of_the_Great_Smoky_Mountains_National_Park.html]

    https://doi.org/10.11646/zootaxa.3249.1.7

    Bartels, P.J., Nelson, D.R. & Exline, R.P. (2011a) Allometry and the removal of body size effects in the morphometric analysis of tardigrades. Journal of Zoological Systematics and Evolutionary Research, 49, 17–25.

    https://doi.org/10.1111/j.1439-0469.2010.00593.x

    Bartels, P.J., Nelson, D.R., Kaczmarek, Ł. & Michalczyk, Ł. (2007) Doryphoribius smokiensis, a new species of Eutardigrada (Hypsibiidae) from the Great Smoky Mountains National Park, TN, USA (North America). Zootaxa, 1646 (1), 56–65.

    https://doi.org/10.11646/zootaxa.1646.1.6

    Bartels, P.J., Nelson, D.R., Kaczmarek, Ł. & Michalczyk, Ł. (2008) Three new species and one new record of the genus Doryphoribius Pilato, 1969 (Tardigrada: Eutardigrada: Hypsibiidae) from the Great Smoky Mountains National Park (Tennessee, USA). Journal of Natural History, 42 (41–42), 2595–2608.

    https://doi.org/10.1080/00222930802357343

    Bartels, P.J. Nelson, D.R., Kaczmarek, Ł. & Michalczyk, Ł. (2011b) Ramazzottius belubellus, a new species of Tardigrada (Eutardigrada: Parachela: Hypsibiidae) from the Great Smoky Mountains National Park (North Carolina, USA). Proceedings of the Biological Society of Washington, 124 (1), 23–27.

    https://doi.org/10.2988/10-13.1

    Bartels, P.J., Nelson, D.R., Kaczmarek, Ł. & Michalczyk, Ł. (2014) The genus Milnesium (Tardigrada: Eutardigrada: Milnesi-idae) in the Great Smoky Mountains National Park (North Carolina and Tennessee, USA), with the description of Milnesium bohleberi sp. nov. Zootaxa, 3826 (2), 356–368.

    https://doi.org/10.11646/zootaxa.3826.2.5

    Bartels, P.J., Pilato, G., Lisi, O. & Nelson, D.R. (2009) Macrobiotus (Eutardigrada, Macrobiotidae) from the Great Smoky Mountains National Park, Tennessee/North Carolina, USA (North America): two new species and six new records. Zootaxa, 2022 (1), 45–57.

    https://doi.org/10.11646/zootaxa.2022.1.4

    Bertolani, R. (1982) A new genus and five new species of Italian fresh-water tardigrades. Bollettino del Museo Civico di Storia Naturale, Verona, 114, 249–254.

    Bertolani, R., Bartels, P.J., Guidetti, R., Cesari, M. & Nelson, D.R. (2014a) Aquatic tardigrades in the Great Smoky Mountains National Park, North Carolina and Tennessee, U.S.A., with the description of a new species of Thulinius (Tardigrada, Isohypsibiidae). Zootaxa, 3764 (5), 524–536.

    https://doi.org/10.11646/zootaxa.3764.5.2

    Bertolani, R., Guidetti, R., Marchioro, T., Altiero, T., Rebecchi, L. & Cesari, M. (2014b) Phylogeny of Eutardigrada: New molecular data and their morphological support lead to the identification of new evolutionary lineages. Molecular Phylogenetics and Evolution, 76, 110–126.

    https://doi.org/10.1016/j.ympev.2014.03.006

    Binda, M.G. (1969) Nuovi dati su tardigradi di Sicilia con descrizione di due nuove specie. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 9, 623–633.

    Binda, M.G. (1971) Su alcuni Tardigradi muscicoli del Nord-Africa. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 10, 759–765.

    Binda, M.G. & Pilato, G. (1969a) Ulteriore contributo alla conoscenza dei Tardigradi di Sicilia con descrizione di due nuove specie. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 10, 205–214.

    Binda, M.G. & Pilato, G. (1969b) Tardigradi muscicoli dell’Isola di Ustica (Sicilia), con descrizione di due specie nuove. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 10, 171–180.

    Binda, M.G. & Pilato, G. (1971) Nuove osservazioni sui Tardigradi delle Isole Eolie. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania, 10, 766–774.

    Dastych, H. (1972) Isohypsibius weglarskae sp. n. a new species of Tardigrada from Poland. Bulletin de l’Académie polonaise des sciences. Série des Sciences Biologiques, 20, 271–274.

    Degma, P., Bertolani, R., Guidetti, R. (2020) Actual checklist of Tardigrada species. 39th Edition. Available from: http://www.tardigrada.modena.unimo.it/miscellanea/Actual%20checklist%20of%20Tardigrada.pdf (accessed 16 February 2021)

    Doyère, P.L.N. (1840) Memoire sur les Tardigrades. Annales des Sciences Naturelles, Zoologie, Serie 2, 14, 269–362.

    Durante Pasa, M.V. & Maucci, W. (1972) Descrizione di Hypsibius (Isohypsibius) basalovoi sp. nov. e altre notizie su tardigradi del Veronese. Memorie del Museo Civico di Storia Naturale, Verona, 20, 275–281.

    Ehrenberg, C.G. (1853) Diagnoses novarum formarum. Verhandlungen der Königlich Preussische Akademie der Wissenschaften zu Berlin, 8, 526–533.

    Gąsiorek, P., Oczkowski, A., Blagde,B., Kristensen, R.M., Bartels, P.J., Nelson, D.R., Suzuki, A.C & Michalczyk, Ł. (2020) New Asian and Nearctic Hypechiniscus species (Heterotardigrada: Echiniscidae) indicate a pseudocryptic horn of plenty. Zoological Journal of the Linnean Society, zlaa110, 59.

    https://doi.org/10.1093/zoolinnean/zlaa110

    Gąsiorek, P., Stec, D., Morek, W. & Michalczyk, Ł. (2019) Deceptive conservatism of claws: distinct phyletic lineages concealed within Isohypsibioidea (Eutardigrada) revealed by molecular and morphological evidence. Contributions to Zoology, 88, 78–132.

    https://doi.org/10.1163/18759866-20191350

    Greven, H. (1972) Tardigraden des nördlichen Sauerlandes. Zoologischer Anzeiger, 189, 368–381.

    Grigarick, A.A., Schuster, R.O. & Nelson, D.R. (1983) Heterotardigrada of Venezuela (Tardigrada). Pan-Pacific Entomologist, 59, 64–77.

    Grobys, D., Roszkowska, M., Gawlak, M., Kmita, H., Kepel, A., Kepel, M., Parnikoza, I., Bartylak, T. & Kaczmarek, Ł. (2020) High diversity in the Pseudechiniscus suillus—facettalis complex (Heterotardigrada: Echiniscidae) with remarks on the morphology of the genus Pseudechiniscus. Zoological Journal of the Linnean Society, 188 (3), 733–752.

    https://doi.org/10.1093/zoolinnean/zlz171

    Guidetti, R. (1998) Two new species of Macrobiotidae (Tardigrada: Eutardigrada) from the United States of America, and some taxonomic considerations of the genus Murrayon. Proceedings of the Biological Society of Washington, 111, 663–673.

    Guidetti, R., Rebecchi, L., Bertolani, R., Jönsson, K.I., Kristensen, R.M. & 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. Zoological Journal of the Linnean Society, 178 (4), 834–845.

    https://doi.org/10.1111/zoj.12428

    Iharos, G. (1966) Neue Tardigraden-Arten aus Ungarn. Acta Zoologica Academiae Scientiarum Hungaricae, 12, 111–122.

    Kaczmarek, Ł. & Michalczyk, Ł. (2003) Platicrista horribilis, a new species of Tardigrada (Eutardigrada: Hypsibiidae) from Mongolia. Zootaxa, 132 (1), 1–8.

    https://doi.org/10.11646/zootaxa.132.1.1

    Kaczmarek, Ł. & Michalczyk, Ł. (2017) The Macrobiotus hufelandi group (Tardigrada) revisited. Zootaxa, 4363 (1), 101–123.

    https://doi.org/10.11646/zootaxa.4363.1.4

    Kaczmarek, Ł., Michalczyk, Ł. & Degma, P. (2005) A new species of Tardigrada Bryodelphax brevidentatus sp. nov. (Heterotardigrada: Echiniscidae) from China (Asia). Zootaxa, 1080 (1), 33–38.

    https://doi.org/10.11646/zootaxa.1080.1.3

    Kaczmarek, Ł., Michalczyk, Ł. & McInnes, S.J. (2016) Annotated zoogeography of non-marine Tardigrada. Part III: North America and Greenland. Zootaxa, 4203 (1), 1–249.

    https://doi.org/10.11646/zootaxa.4203.1.1

    Kaczmarek, Ł., Gawlak, M., Bartels, P.J., Nelson, D.R., & Roszkowska, M. (2017) Revision of the genus Paramacrobiotus Guidetti et al., 2009 with the description of a new species, re-descriptions and a key. Annales Zoologici, 67 (4), 627–656. https://doi.org/10.3161/00034541ANZ2017.67.4.001

    Kaczmarek, Ł., Kayashta, P., Gawlak, M., Mioduchowska, M., & Roszkowska, M. (2021) An integrative description of a Diploechiniscus oihonnae (Richters, 1903) population from near the original type locality in Merok (Norway). Zootaxa, 4964 (1), 83–102.

    https://doi.org/10.11646/zootaxa.4964.1.4

    Kaczmarek, Ł., Zawierucha, K., Buda, J., Stec, D., Gawlak, M., Michalczyk, Ł. & Roszkowska, M. (2018). An integrative redescription of the nominal taxon for the Mesobiotus harmsworthi group (Tardigrada: Macrobiotidae) leads to descriptions of two new Mesobiotus species from Arctic. PLoS One, 13 (10), e0204756.

    https://doi.org/10.1371/journal.pone.0204756

    Kendall-Fite, K. & Nelson, D.R. (1996) Two new species of tardigrades from Short Mountain, Tennessee, USA. Zoological Journal of the Linnean Society, 116 (1–2), 205–214.

    https://doi.org/10.1111/j.1096-3642.1996.tb02344.x

    Kristensen, R.M. (1982) New aberrant eutardigrades from homothermic springs on Disko Island, West Greenland. In: Nelson, D.R. (Ed.), Proceedings of the Third International Symposium on Tardigrada. East Tennessee State University Press, Johnson City, Tennessee, pp. 203–220.

    Lisi, O., Londoño, R. & Quiroga, S. (2020) Description of a new genus and species (Eutardigrada: Richtersiidae) from Colombia, with comments on the family Richtersiidae. Zootaxa, 4822 (4), 531–550.

    https://doi.org/10.11646/zootaxa.4822.4.4

    Marcus, E. (1936) Tardigrada. Das Tierreich, 66, 1–340.

    Maucci, W. (1954) Tardigradi nuovi della fauna Italiana. Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 93, 576–585.

    Maucci, W. (1986) Tardigrada. In: Fauna d`Italia. Vol. 24. Bologna, Calderini, pp. 1–388.

    McFatter, M.M., Meyer, H.A. & Hinton, J.G. (2007) Nearctic freshwater tardigrades: A review. Journal of Limnology, 66 (Supplement 1), 84–89.

    https://doi.org/10.4081/jlimnol.2007.s1.84

    Meyer, H.A. (2013) Terrestrial and freshwater Tardigrada of the Americas. Zootaxa, 3747 (1), 1–71.

    https://doi.org/10.11646/zootaxa.3747.1.1

    Murray, J. (1905a) The Tardigrada of the Forth Valley. Annales of Scottish Natural History, 55, 160–164.

    Murray, J. (1905b) The Tardigrada of the Scottish Lochs. Transactions of the Royal Society of Edinburgh, 41, 677–698.

    https://doi.org/10.1017/S0080456800035547

    Murray, J. (1907a) Scottish Tardigrada collected by the Lake Survey. Transactions of the Royal Society of Edinburgh, 45, 641–668.

    https://doi.org/10.1017/S0080456800011777

    Murray, J. (1907b) Arctic Tardigrada, collected by Wm. S. Bruce. Transactions of the Royal Society of Edinburgh, 45, 669–681.

    https://doi.org/10.1017/S0080456800011789

    Murray, J. (1910) Tardigrada. British Antarctic Expedition 1907–1909. Reports on the Scientific Investigations, 1, Biology (Part V), 83–187.

    Murray, J. (1911) Arctiscoida. Proceedings of the Royal Irish Academy, 31, 1–16.

    Nelson, D.R. & Bartels, P.J. (2007) “Smoky Bears—Tardigrades of Great Smoky Mountains National Park. Southeastern Natur-alist, 6 (Supplement 2), 229–238.

    https://doi.org/10.1656/1528-7092(2007)6[229:SBOGSM]2.0.CO;2

    Nelson, D.R. & Bartels, P.J. (2013) Species richness of soil and leaf litter tardigrades in the Great Smoky Mountains National Park (North Carolina/Tennessee, USA). Journal of Limnology, 72 (Supplement 1), e18.

    https://doi.org/10.4081/jlimnol.2013.s1.e18

    Nelson, D.R. & McGlothlin, K.L. (1993) A new species of Hypsibius (phylum Tardigrada) from Roan Mountain, Tennessee, USA. Transactions of the American Microscopical Society, 112, 140–144.

    https://doi.org/10.2307/3226827

    Nelson, D.R. & McGlothlin, K.L. (1996) A new species of Calohypsibius (Phylum Tardigrada, Eutardigrada) from Roan Mountain, Tennessee—North Carolina, USA. Zoological Journal of the Linnean Society, 116, 167–174.

    https://doi.org/10.1111/j.1096-3642.1996.tb02341.x

    Nelson, D.R., Bartels, P.J. & Fegley, S.R. (2020) Environmental correlates of tardigrade community structure in mosses and lichens in the Great Smoky Mountains National Park (Tennessee and North Carolina, USA). Zoological Journal of the Linnean Society, 188 (3), 913–924.

    https://doi.org/10.1093/zoolinnean/zlz043

    Perry, E., Miller, W.R. & Kaczmarek, Ł. (2019) Recommended abbreviations for the names of genera of the phylum Tardigrada. Zootaxa, 4608 (1), 145–154.

    https://doi.org/10.11646/zootaxa.4608.1.8

    Péterfi, F. (1956) Contributioni la cunoasterea Tardigradelor din R.P.R. Studii si Cercetari de Biologie, 7, 149–155.

    Pilato, G. (1971) Tardigradi delle acque dolci siciliane. Nota prima. Bollettino delle Sedute dell′Accademia Gioenia di Scienze Naturale Catania, 9, 126–134.

    Pilato, G. (1973) Tardigradi delle acque dolci siciliane—Nota Seconda. Bollettino delle Sedute dell′Accademia Gioenia di Scienze Naturale Catania, 12, 177–186.

    Pilato, G. & Binda, M.G. (1997/98) A comparison of Diphascon (D.) alpinum Murray, 1906, D. (D.) chilenense Plate, 1889 and D. (D.) pingue Marcus, 1936 (Tardigrada), and description of a new species. Zoologischer Anzeiger, 236, 181–185.

    Pilato, G., Fontoura, P. & Lisi, O. (2007) Remarks on the Echiniscus viridis group, with the description of a new species (Tardigrada, Echiniscidae). Journal of Limnology, 66 (Supplement 1), 33–39.

    https://doi.org/10.4081/jlimnol.2007.s1.33

    Plate, L. (1888) Beiträge zur Naturgeschichte der Tardigraden. Zoologische Jahrbücher, 3, 487–550.

    https://doi.org/10.5962/bhl.part.1265

    Ramazzotti, G. (1956) Tre nouve specie di Tardigradi ed altre specie poco comuni. Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 95, 284–291.

    Ramazzotti, G. (1957) Due nuove specie di Tardigradi extra-europei. Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 96, 188–191.

    Ramazzotti, G. (1959) Il gruppo dell’ Echiniscus viridis con la nuova species E. perviridis e Macrobiotus pustulatus altra nuova specie (Tardigrada). Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 98, 303–309.

    Ramazzotti, G. (1962) Tardigradi del Cile, con descrizione di quattro nuove specie e di una variet. Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 101, 275–287.

    Richters, F. (1900) Beiträge zur Kenntnis der Fauna der Umgebung von Frankfurt a. M. Bericht der Senckenbergischen Naturforschenden gesellschaft in Frankfurt am Main, 1900, 21–44.

    Richters, F. (1902) Beiträge zur Kenntnis der Fauua der Umgebung von Frankfurt a. M. Bericht der Senckenbergischen Naturforschenden gesellschaft in Frankfurt am Main, 1902, 3–21.

    Richters, F. (1903) Nordische Tardigraden. Zoologisher Anzeiger, 27, 168–172.

    Richters, F. (1904a) Arktische Tardigraden. Fauna Arctica, 3, 495–508.

    Richters, F. (1904b) Isländische Tardigraden. Zoologischer Anzeiger, 28, 373–377.

    Richters, F. (1911) Faune des mousses. Tardigrades. Duc d’Orleans. Campagne arctique de 1907. Impr. Sci. C. Buelens, Bruxelles, 20 pp.

    Riggin, G.T. (1962) Tardigrada of Southwest Virginia: with the addition of a new marine tardigrade species from Florida. Virginia Agricultural Experiment Station Technical Bulletin, 152, 1–145.

    Robotti, C. (1970) Hypsibius (D.) ramazzotti sp. nov. e Macrobiotus aviglianae sp. nov. Primo contributo alla conoscenza dei Tardigradi del Piemonte. Atti della Societ Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 110, 251–255.

    Roszkowska, M., Grobys, D., Bartylak, T., Gawlak, M., Kmita, H., Kepel, A., Kepel, M., Parnikoza, I. & Kaczmarek, Ł. (2020) Integrative description of five Pseudechiniscus species (Heterotardigrada: Echiniscidae: the suillus-facettalis complex). Zootaxa, 4763 (4), 451–484.

    https://doi.org/10.11646/zootaxa.4763.4.1

    Roszkowska, M., Bartels, P.J., Gołdyn, B., Ciobanu, D.A., Fontoura, P., Michalczyk, Ł., Nelson, D.R., Ostrowska, M., Moreno-Talamantes, A. & Kaczmarek, Ł. (2016) Is the gut content of Milnesium (Eutardigrada) related to buccal tube size? Zoological Journal of the Linnean Society, 178 (4), 794–803.

    https://doi.org/10.1111/zoj.12459

    Schultze, C.A.S. (1834) Macrobiotus Hufelandii animal e crustaceorum classe novum, reviviscendi post diuturnam asphixiam et aridiatem potens, etc. C. Curths, Berlin, 8 pp., 1 tab.

    Schuster, R.O. & Grigarick, A.A. (1971) Two new species of Echiniscus from the Pacific Northwest (Tardigrada: Echiniscidae). Proceedings of the Entomological Society of Washington, 73, 105–110.

    Schuster R.O., Toftner, E.C. & Grigarick, A.A. (1978) Tardigrada of Pope Beach, Lake Tahoe, California. The Wasmann Journal of Biology, 35, 115–136.

    Sigovini, M., Keppel, E. & Tagliapietra, D. (2016) Open Nomenclature in the biodiversity era. Methods in Ecology and Evolution, 7 (10), 1217–1225.

    https://doi.org/10.1111/2041-210X.12594

    Stec, D., Morek, W., Gąsiorek, P. & Michalczyk, Ł. (2018) Unmasking hidden species diversity within the Ramazzottius oberhaeuseri complex, with an integrative redescription of the nominal species for the family Ramazzottiidae (Tardigrada: Eutardigrada: Parachela). Systematics and Biodiversity, 16 (4), 357–376.

    https://doi.org/10.1080/14772000.2018.1424267

    Stec, D., Vecchi, M., Maciejowski, W. & Michalczyk, Ł. (2020) Resolving the systematics of Richtersiidae by multilocus phylogeny and an integrative redescription of the nominal species for the genus Crenubiotus (Tardigrada). Scientific Reports, 10, 19418.

    https://doi.org/10.1038/s41598-020-75962-1

    Stec, D., Vecchi, M., Dudziak, M., Bartels, P.J., Calhim, S. & Michalczyk, Ł. (2021) Integrative taxonomy resolves species identities within the Macrobiotus pallarii complex (Eutardigrada: Macrobiotidae). Zoological Letters.

    https://doi.org/10.1186/s40851-021-00176-w

    Thulin, G. (1911) Beitrag zur Kenntnis der Tardigradenfauna Schwedens. Arkiv för zoologi, 7, 1–60.

    https://doi.org/10.5962/bhl.part.1270

    Thulin, G. (1928) Über die Phylogenie und das System der Tardigraden. Hereditas, 11, 207–266.

    https://doi.org/10.1111/j.1601-5223.1928.tb02488.x

    Urbanowicz, C. (1925) Sur la variabilit de Macrobiotus oberhaeuseri. Bulletin Biologique de la France et de la Belgique, Paris, 59, 124–142.

    Węglarska, B. (1970) Hypsibius (Isohypsibius) smreczynskii spec. nov., a new species of freshwater tardigrade. Zeszyty Uniwersytetu Jagiellońskiego, Prace Zoologiczne, 16, 107–114.

    Węglarska, B. (1973) Tardigrada in high Tatra localities bare of snow with a description of Itaquascon pawlowskii sp. n. Vĕstnk Československ Společnosti Zoologick, 37, 150–154.