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Type: Article
Published: 2020-09-11
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Tardigrades of Finland: new records and an annotated checklist

Department of Biological and Environmental Science, University of Jyvaskyla, PO Box 35, FI-40014, Finland
Department of Life Science, University of Modena and Reggio Emilia, via G. Campi 213/D, 41125 Modena, Italy
Department of Biological and Environmental Science, University of Jyvaskyla, PO Box 35, FI-40014, Finland
Department of Biological and Environmental Science, University of Jyvaskyla, PO Box 35, FI-40014, Finland
Tardigrada biodiversity Fennoscandia Finnish fauna microinvertebrates sampling effort

Abstract

Species checklists are powerful and important tools of communication between taxonomists and applied environmental biologists, which in turn lead to well-planned and successful conservation strategies and ecological studies. Despite this, only recently the interest on compiling systematic checklists is growing among taxonomists who study tardigrades—micrometazoans that inhabit almost every habitat worldwide. As the Finnish records of tardigrades (a.k.a. water bears) species are incomplete, outdated and no checklist has ever been compiled for this country, an easy-to-consult checklist is here reported. This checklist covers all Finnish tardigrade taxa identified in the past and in the 13 samples collected for this study. A total of 68 tardigrade species are recorded from Finland, with 6 of them being new records presented in this contribution. Of these species, four have their loci tipici in Finland and we provide an English translation of their original German descriptions. A Generalised Linear Model was used to test the effect of sampling effort and area size on the number of species recorded in each biogeographical province of Finland. The results showed that geographical differences in species richness can be explained solely by sampling effort. The number of tardigrade species recorded in Finland corresponds to about 5% of all described species in the phylum, thus indicating a potential high richness for this country. However, the results of the Generalised Linear Model highlight that a reliable knowledge of the tardigrade diversity in Finland will be reached only with a more uniform and intensive sampling effort.

 

References

  1. Arakawa, K. (2020) Simultaneous metabarcoding of Eukaryotes and Prokaryotes to elucidate the community structures within tardigrade microhabitats. Diversity, 12 (3), 110.

    https://doi.org/10.3390/d12030110

    Arcidiacono, R. (1962) Contributo alla conoscenza dei Tardigradi dei Monti Nebrodi e desrizione di una nuova specie di Itaquascon. Bollettino delle Sedute dell’Accedemia Gioenia di Scienze Naturali, Catania, 4 (7), 123–134

    Arrhenius, O. (1921) Species and area. Journal of Ecology, 9 (1), 95–99.

    https://doi.org/10.2307/2255763

    Berdi, D. & Altýndað, A. (2020) Contribution to the knowledge on distribution of Tardigrada in Turkey. Diversity, 12 (3), 95.

    https://doi.org/10.3390/d12030095

    Bertolani, R. (1981) A new genus and five new species of Italian fresh-water tardigrades. Bolletino del Museo Civico di Storia Naturale di Verona, 8, 249–254.

    Bertolani, R. & Rebecchi, L. (1993) A revision of the Macrobiotus hufelandi group (Tardigrada, Macrobiotidae), with some observations on the taxonomic characters of eutardigrades. Zoologica Scripta, 22 (2), 127–152.

    https://doi.org/10.1111/j.1463-6409.1993.tb00347.x

    Bertolani, R., Biserov, V., Rebecchi, L. & Cesari, M. (2011a) Taxonomy and biogeography of tardigrades using an integrated approach: new results on species of the Macrobiotus hufelandi group. Invertebrate Zoology, 8 (1), 23–36.

    https://doi.org/10.15298/invertzool.08.1.05

    Bertolani, R., Rebecchi, L., Giovannini, I. & Cesari, M. (2011b) DNA barcoding and integrative taxonomy of Macrobiotus hufelandi C.A.S. Schultze 1834, the first tardigrade species to be described, and some related species. Zootaxa, 2997 (1), 19–36.

    https://doi.org/10.11646/zootaxa.2997.1.2

    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. Molecular Phylogenetics and Evolution, 76, 110–126.

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

    Binda, M.G. (1971) Su alcuni tardigradi muscicoli del Nord-Africa. Bollettino delle sedute dell’Accademia Gioenia di Scienze Naturali, Catania, 10, 759–765.

    Blagden, B., DeMilio, E., Hansen, J.G. & Kristensen, R.M. (2020) First records of tardigrades (Tardigrada) from Irish and Scottish leaf litter. Glasgow Naturalist, 27 (2), 1–16.

    https://doi.org/10.37208/tgn27202

    Cesari, M., Montanari, M., Kristensen, R.M., Bertolani, R., Guidetti, R. & Rebecchi, L. (2020) An integrated study of the biodiversity within the Pseudechiniscus suillus–facettalis group (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society, 188 (3), 717–732.

    https://doi.org/10.1093/zoolinnean/zlz045

    Claxton, S.K. (1998) A revision of the genus Minibiotus (Tardigrada: Macrobiotidae) with descriptions of eleven new species from Australia. Records of the Australian Museum, 50 (2), 125–160.

    https://doi.org/10.3853/j.0067-1975.50.1998.1276

    Dastych, H. (1997) Redescription of Isohypsibius sculptus Ramazzotti, 1962 (Tardigrada). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg, 12 (155), 163–167.

    Degma, P. & Pečalková, M. (2003) First records of Tardigrada in Slovakia from Stužica National Nature Reserve (Bukovské vrchy Mts, NE Slovakia). Biologia—Section Zoology, 58 (2), 274–286.

    Degma, P. & Guidetti, R. (2007) Notes to the current checklist of Tardigrada. Zootaxa, 53 (1579), 41–53.

    https://doi.org/10.11646/zootaxa.1579.1.2

    Degma, P. (2018) Field and Laboratory Methods. In: Schill, R.O. (Ed.), Zoological Monographs. Vol. 2. Water Bears: The Biology of Tardigrades. Springer International Publishing, Cham, pp. 349–369.

    https://doi.org/10.1007/978-3-319-95702-9_14

    Degma, P., Bertolani, R. & Guidetti, R. (2019) Actual checklist of Tardigrada species (2009–2019, 36th Edition. 1 September 2019). Available from: https://iris.unimore.it/handle/11380/1178608 (accessed 1 March 2020)

    https://doi.org/10.25431/11380_1178608

    DeMilio, E., Lawton, C. & Marley, N.J. (2016) Tardigrada of Ireland: A review of records and an updated checklist of species including a new addition to the Irish fauna. ZooKeys, 616, 77–101.

    https://doi.org/10.3897/zookeys.616.8222

    Doyère, L.M.F. (1840) Mémoire sur les Tardigrades. Annales des Sciences Naturelles, Zoologie, Series 2, 14, 269–362.

    Dujardin, F. (1851) Sur les Tardigrades et sur une espèce a longs pieds dans l’eau de mer. Annales Des Sciences Naturelles, 15, 160–166.

    Durante Pasa, M.V. & Maucci, W. (1979) Moss Tardigrada from the Scandinavian peninsula. Zeszyty Naukwe Uniwersytetu Jagiellońskiego, Prace Zoologiczne Kraków, 79 (25), 47–85.

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

    Faurby, S., Jönsson, K.I., Rebecchi, L. & Funch, P. (2008) Variation in anhydrobiotic survival of two eutardigrade morphospecies: a story of cryptic species and their dispersal. Journal of Zoology, 275 (2), 139–145.

    https://doi.org/10.1111/j.1469-7998.2008.00420.x

    Gąsiorek, P., Stec, D., Morek, W., Zawierucha, K., Kaczmarek, Ł., Lachowska-Cierlik, D. & Michalczyk, Ł. (2016) An integrative revision of Mesocrista Pilato, 1987 (Tardigrada: Eutardigrada: Hypsibiidae). Journal of Natural History, 50 (45–46), 2803–2828.

    https://doi.org/10.1080/00222933.2016.1234654

    Gąsiorek, P., Stec, D., Morek, W., Marnissi, J. & Michalczyk, Ł. (2017) The tardigrade fauna of Tunisia, with an integrative description of Bryodelphax maculatus sp. nov. (Heterotardigrada: Echiniscidae). African Zoology, 52 (2), 77–89.

    https://doi.org/10.1080/15627020.2017.1297688

    Gąsiorek, P., Stec, D., Morek, W. & Michalczyk, Ł. (2018a) An integrative redescription of Hypsibius dujardini (Doyère, 1840), the nominal taxon for Hypsibioidea (Tardigrada: Eutardigrada). Zootaxa, 4415 (1), 45–75.

    https://doi.org/10.11646/zootaxa.4415.1.2

    Gąsiorek, P., Stec, D., Zawierucha, K., Kristensen, R.M. & Michalczyk, Ł. (2018b) Revision of Testechiniscus Kristensen, 1987 (Heterotardigrada: Echiniscidae) refutes the polar-temperate distribution of the genus. Zootaxa, 4472 (2), 261–297.

    https://doi.org/10.11646/zootaxa.4472.2.3

    Gąsiorek, P., Morek, W., Stec, D. & Michalczyk, Ł. (2019a) Untangling the Echiniscus Gordian knot: paraphyly of the “arctomys group” (Heterotardigrada: Echiniscidae). Cladistics, 35 (6), 633–653.

    https://doi.org/10.1111/cla.12377

    Gąsiorek, P., Stec, D., Morek, W. & Michalczyk, Ł. (2019b) Deceptive conservatism of claws: distinct phyletic lineages concealed within Isohypsibioidea (Eutardigrada) revealed by molecular and morphological evidence. Contributions to Zoology, 88 (1), 78–132.

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

    Gąsiorek, P. & Michalczyk, Ł. (2020) Revised Cornechiniscus (Heterotardigrada) and new phylogenetic analyses negate echiniscid subfamilies and tribes. Royal Society Open Science, 7 (6), 1–37.

    https://doi.org/10.1098/rsos.200581

    Gąsiorek, P. & Michalczyk, Ł. (2020) Phylogeny of Itaquasconinae in the light of the evolution of the flexible pharyngeal tube in Tardigrada. Zoologica Scripta, 49 (4), 499–515. [in press]

    https://doi.org/10.1111/zsc.12424

    Grigarick, A.A., Schuster, R. O. & Toftner, E. C. (1973) Descriptive morphology of eggs of some species in the Macrobiotus hufelandii group (Tardigrada: Macrobiotidae). Pan-Pacific Entomologist, 49 (1–4), 258–263

    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

    Grothman, G.T., Johansson, C., Chilton, G., Kagoshima, H., Tsujimoto, M. & Suzuki, A.C. (2017) Gilbert Rahm and the status of Mesotardigrada Rahm, 1937. Zoological science, 34 (1), 5–10.

    https://doi.org/10.2108/zs160109

    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), 1–46.

    https://doi.org/10.11646/zootaxa.845.1.1

    Guidetti, R., Peluffo, J.R., Rocha, A.M., Cesari, M. & De Moly Peluffo, M.C. (2013) The morphological and molecular analyses of a new South American urban tardigrade offer new insights on the biological meaning of the Macrobiotus hufelandi group of species (Tardigrada: Macrobiotidae). Journal of Natural History, 47 (37–38), 2409–2426.

    https://doi.org/10.1080/00222933.2013.800610

    Guidetti, R., Jönsson, K.I. & Kristensen, R.M. (2015) Tardigrades of Sweden; an updated check-list. Zootaxa, 3981 (4), 491–507.

    https://doi.org/10.11646/zootaxa.3981.4.2

    Guidetti, R., Rebecchi, L., Bertolani, R., Jönsson, K.I., Møbjerg Kristensen, R. & 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

    Guidetti, R., Cesari, M., Bertolani, R., Altiero, T. & Rebecchi, L. (2019a) High diversity in species, reproductive modes and distribution within the Paramacrobiotus richtersi complex (Eutardigrada, Macrobiotidae). Zoological letters, 5 (1), 1–28.

    https://doi.org/10.1186/s40851-018-0113-z

    Guidetti, R., Massa, E., Bertolani, R., Rebecchi, L. & Cesari, M. (2019b) Increasing knowledge of Antarctic biodiversity: new endemic taxa of tardigrades (Eutardigrada; Ramazzottiidae) and their evolutionary relationships. Systematics and Biodiversity, 17 (6), 573-593.

    https://doi.org/10.1080/14772000.2019.1649737

    Hallas, T. E. (1975) Interstitial water and Tardigrada in a moss cushion. Annales Zoologici Fennici, 12 (4), 255–259.

    Hallas, T. E. (1977) Survey of the Tardigrada of Finland. Annales Zoologici Fennici, 14 (3), 173–183

    Hallas, T. E. (1978) Habitat preference in terrestrial tardigrades. Annales Zoologici Fennici, 15 (1), 66–68.

    Heinis, F. (1910) Systematik und Biologie der moosbewohnenden Rhizopoden, Rotatorien und Tardigraden der Umgebung von Basel mit Berücksichtigung der übrigen Schweiz. Archiv für Hydrobiologie und Planktonkunde, 5, 127–256

    Hohberg, K., Russell, D.J. & Elmer, M. (2011) Mass occurrence of algal‐feeding tardigrade Apodibius confusus, in the young soils of a post‐mining site. Journal of Zoological Systematics and Evolutionary Research, 49, 62–65.

    https://doi.org/10.1111/j.1439-0469.2010.00600.x

    Holopainen, I.J. & Paasivirta, L. (1977) Abundance and biomass of the meiozoobenthos in the oligotrophic and mesohumic lake Pääjärvi, southern Finland. Annales Zoologici Fennici, 14 (3), 124–134.

    Huhta, V. (1976) Effects of clear-cutting on numbers, biomass and community respiration of soil invertebrates. Annales Zoologici Fennici, 13 (1), 63–80

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

    Inkscape Project (2020) Inkscape. Available from: https://inkscape.org (accessed 4 August 2020)

    Kaczmarek, Ł., Michalczyk, Ł. & McInnes, S.J. (2014) Annotated zoogeography of non-marine Tardigrada. Part I: Central America. Zootaxa, 3763 (1), 1–62.

    https://doi.org/10.11646/zootaxa.3763.1.1

    Kaczmarek, Ł., Bartels, P.J., Roszkowska, M. & Nelson, D.R. (2015a) The zoogeography of marine Tardigrada. Zootaxa, 4037 (1), 1–189.

    https://doi.org/10.11646/zootaxa.4037.1.1

    Kaczmarek, Ł., Michalczyk, Ł. & McInnes, S.J. (2015b) Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa, 3923 (1), 1–107.

    https://doi.org/10.11646/zootaxa.3923.1.1

    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. & 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., 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

    Karling, T.G. (1955) Uber einige Kleintiere des Meeressandes des Nordsee-und Ostsee-Gebietes. Arkiv för Zoologi, 14, 241–249

    Lowry, P.P. & Smith, P.P. (2003) Closing the Gulf between Botanists and Conservationists. Conservation Biology, 17 (4), 1175–1176.

    https://doi.org/10.1046/j.1523-1739.2003.02378.x

    Marcus, E. (1928) Spinnentiere oder Arachoides. IV Bärtierchen (Tardigrada). Tierwelt Deutschlands und der angrenzenden Meeresteile Jena, 12, 1–230.

    Marcus, E. (1936) Das Tierreich. Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen. Artropoda. Tardigrada. Vol. 66. Preußischen Akademie der Wissenschaften zu Berlin, Berlin and Walter de Gruiter & Co., Leipzig, 341 pp.

    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.

    McInnes, S.J. (1994) Zoogeographic distribution of terrestrial/freshwater tardigrades from current literature. Journal of Natural History, 28 (2), 257–352.

    https://doi.org/10.1080/00222939400770131

    McInnes, S.J., Michalczyk, Ł. & Kaczmarek, Ł. (2017) Annotated zoogeography of non-marine Tardigrada. Part IV: Africa. Zootaxa, 4284 (1), 1–74.

    https://doi.org/10.11646/zootaxa.4284.1.1

    Meier, T. (2017) A checklist of norwegian tardigrada. Fauna Norvegica, 37, 25–42.

    https://doi.org/10.5324/fn.v37i0.2269

    Michalczyk, Ł., Wełnicz, W., Frohme, M. & Kaczmarek, Ł. (2012) Redescriptions of three Milnesium Doyère, 1840 taxa (Tardigrada: Eutardigrada: Milnesiidae), including the nominal species for the genus. Zootaxa, 3393 (1), 66–68.

    https://doi.org/10.11646/zootaxa.3154.1.1

    Mihelčič, F. (1938) Beitrage zur Kenntnis der Tardigraden Jugoslawiens, V. Die Tardigraden der Hohlen und Grotten des Karstes. Zoologischer Anzeiger, 123 (10/11), 318–319.

    Mihelčič, F. (1951) Beitrag zur Systematik der Tardigraden. Archivio Zoologico Italiano, 36, 57–103

    Mihelcic, F. (1971) Süsswassertardigraden aus Nordeuropa. Insect Systematics & Evolution, 2 (3), 205–214.

    https://doi.org/10.1163/187631271X00211

    Mihelčič, F. (1971/72) Ein beitrag zur kenntnis der Susswassertardigraden Nordeuropas. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 110 (111), 37–45.

    Morek, W., Ciosek, J. & Michalczyk, Ł. (2020) Description of Milnesium pentapapillatum sp. nov., with an amendment of the diagnosis of the order Apochela and abolition of the class Apotardigrada (Tardigrada). Zoologischer Anzeiger. [in press]

    https://doi.org/10.1016/j.jcz.2020.07.002

    Morek, W., Gąsiorek, P., Stec, D., Blagden, B. & Michalczyk, Ł. (2016) Experimental taxonomy exposes ontogenetic variability and elucidates the taxonomic value of claw configuration in Milnesium Doyère, 1840 (Tardigrada: Eutardigrada: Apochela). Contributions to Zoology, 85 (2), 173–200.

    https://doi.org/10.1163/18759866-08502003

    Morek, W., Stec, D., Gąsiorek, P., Surmacz, B. & Michalczyk, Ł. (2019a) Milnesium tardigradum Doyère, 1840: The first integrative study of interpopulation variability in a tardigrade species. Journal of Zoological Systematics and Evolutionary Research, 57 (1), 1–23.

    https://doi.org/10.1111/jzs.12233

    Morek, W., Suzuki, A. C., Schill, R. O., Georgiev, D., Yankova, M., Marley, N. J. & Michalczyk, Ł. (2019b) Redescription of Milnesium alpigenum Ehrenberg, 1853 (Tardigrada: Apochela) and a description of Milnesium inceptum sp. nov., a tardigrade laboratory model. Zootaxa, 4586 (1), 57–62.

    https://doi.org/10.11646/zootaxa.4586.1.2

    Morek, W. & Michalczyk, Ł. (2020) First extensive multilocus phylogeny of the genus Milnesium (Tardigrada) reveals no congruence between genetic markers and morphological traits. Zoological Journal of the Linnean Society, 188 (3), 681–693.

    https://doi.org/10.1093/zoolinnean/zlz040

    Morek, W., Ciosek, J.A. & Michalczyk, Ł. (2020) Description of Milnesium pentapapillatum sp. nov., with an amendment of the diagnosis of the order Apochela and abolition of the class Apotardigrada (Tardigrada). Zoologischer Anzeiger. [in press]

    https://doi.org/10.1016/j.jcz.2020.07.002

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

    Murray, J. (1905b) XXVII. The Tardigrada of the Scottish Lochs. Transactions of The Royal Society of Edinburgh, 41 (3), 677–698.

    https://doi.org/10.1017/s0080456800035547

    Murray J. (1906a) The Tardigrada of the Forth Valley (Second Paper). The Annals of Scottish Natural History, 15, 214–217.

    Murray, J. (1906b) Scottish Alpine Tardigrada. The Annals of Scottish Natural History, 57, 25–30.

    Murray, J. (1907a) XXIV. Scottish Tardigrada, collected by the Lake Survey. Transactions of The Royal Society of Edinburgh, 45 (3), 641–668.

    https://doi.org/10.1017/S0080456800011777

    Murray, J. (1907b) The encystment of Macrobiotus. Zoologist, 11 (4), 4–11.

    Murray, J. (1907c) XXV. Arctic Tardigrada, collected by Wm. S. Bruce. Transactions of The Royal Society of Edinburgh, 45 (3), 669–681.

    https://doi.org/10.1017/s0080456800011789

    Murray, J. (1910) Tardigrada of New Zeland. In: Murray, J. (Ed.), British Antarctic Expedition 1907–1909. Uder the command of Sir E. H. Shackleton. Reports on the Scientific Investigations. Biology. Vol. 1. V. Heinemann, London, pp. 107–122.

    https://doi.org/10.5962/bhl.title.22427

    Murray, J. (1911) Arctiscoida. Proceedings of the Royal Irish Academy, 31 (37), 1–16.

    Nederström, P. (1919) Die bis jetzt aus Finland bekannten Tardigraden. Acta Societatis pro Fauna et Flora Fennica, 46 (8), 1–25.

    Nelson, D.R., Bartels, P.J. & Guil, N. (2018) Tardigrade ecology. In: Schill, R.O. (Ed.), Zoological Monographs. Vol. 2. Water Bears: The Biology of Tardigrades. Springer International Publishing, Cham, pp. 163–210.

    https://doi.org/10.1007/978-3-319-95702-9_7

    Nowak, B. & Stec D. (2018) An integrative description of Macrobiotus hannae sp. nov.(Tardigrada: Eutardigrada: Macrobiotidae: hufelandi group) from Poland. Turkish Journal of Zoology, 42 (3), 269–286.

    https://doi.org/10.3906/zoo-1712-31

    Paasivirta, L. & Särkkä, J. (1978) Effects of pulp mill and municipial effluents and humus load on the macro-and meiozoobenthos of some Finnish lakes: Internationale Vereinigung für theoretische und angewandte Limnologie: Verhandlungen, 20 (3), 1779–1788.

    https://doi.org/10.1080/03680770.1977.11896767

    Pilato, G. & Binda, M.G. (1999) Three new species of Diphascon of the pingue group (Eutardigrada, Hypsibiidae) from Antarctica. Polar Biology, 21 (5), 335–342.

    https://doi.org/10.1007/s003000050370

    Pilato, G., Binda, M. G., Napolitano, A. & Moncada, E. (2000) The specific value of Macrobiotus coronatus DeBarros, 1942, and description of two new species of the harmsworthi group (Eutardigrada). Bollettino delle sedute della Accademia Gioenia di Scienze Naturali, Catania, 33, 103–120.

    Pilato, G., Guidetti, R., Rebecchi, L., Lisi, O., Hansen, J.G. & Bertolani, R. (2006) Geonemy, ecology, reproductive biology and morphology of the tardigrade Hypsibius zetlandicus (Eutardigrada: Hypsibiidae) with erection of Borealibius gen. n. Polar Biology, 29 (7), 595–603.

    https://doi.org/10.1007/s00300-005-0094-1

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

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

    Pogwizd, J. & Stec, D. (2020) New Records of Dactylobiotus parthenogeneticus Bertolani, 1982 Provide Insight into Its Genetic Variability and Geographic Distribution. Folia Biologica, Krakòw, 68 (2), 57–72.

    https://doi.org/10.3409/fb_68-2.08

    Pollock, L.W. (1971) On some British marine Tardigrada, including two new species of Batillipes. Journal of the Marine Biological Association of the United Kingdom, 51 (1), 93–103.

    https://doi.org/10.1017/S0025315400006482

    Purasjoki, K. (1953) Zwei verbesserte Apparate zum Auffangen von Kleintieren des See-und Meeresbodens. Suomal. eläin-ja kasvit. Seur. van. Tiedon, Archivum Societatis Zoologicae Botanicae Fennicae ”Vanamo”, 8, 110–114.

    Rahm, G. (1931). Tardigrada of the South of America (esp. Chile). Revista Chilena de Historia Natural, 35, 118–141.

    Ramazzotti, G. (1962) Tardigradi del Cile, con descrizione di quattro nuove specie e di una nuova varietà. Atti della Società italiana di Scienze naturali e Museo civico di Storia naturale di Milano, 101, 275–287.

    Ramazzotti, G. (1972) Il Phylum Tardigrada. Memorie dell’Isituto Italiano di Idrobiologia Dott, Marco De Marchi, 28, 1–732.

    Ramazzotti, G. & Maucci, W. (1983) Il Phylum Tardigrada. Terza edizione riveduta e corretta. Memorie dell’Isituto Italiano di Idrobiologia Dott. Marco De Marchi, 41, 1012 pp.

    R Core Team (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Availeble from: https://www.R-project.org (accessed 29 March 2020)

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

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

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

    Richters, F. (1904) Arktische Tardigraden. Fauna Arctica, 3, 494–508.

    Richters, F. (1908) Moosbewohner. Wissenschaftliche Ergebnisse der Schwedischen Südpolar-expedition 1901–1903, 6 (2), 1–16.

    Richters, F. (1909) Marine Tardigraden. Bericht der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main, 1909, 28–48.

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

    Rodríguez-Roda, J. (1952) Tardígrados de la fauna Española. Vol. 1. Museo de Ciencias Naturales, Barcelona, 108 pp.

    Roszkowska, M., Ostrowska, M., Grobys, D. Kmita, H. & Kaczmarek, Ł. (2019) Some Tardigrades from Italy, with an Updated Checklist of Limno-terrestrial Species from the Country. Acta Zoologica Bulgarica, 71 (2), 167–174.

    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

    Särkkä, J. & Paasivirta, L. (1972) Vertical distribution and abundance of the macro- and meiofauna in the profundal sediments of Lake Päijänne, Finland. Annales Zoologici Fennici, 9 (1), 1–9.

    Särkkä, J., Levonen, L. & Mäkelä, J. (1997) Meiofauna of springs in Finland in relation to environmental factors. Hydrobiologia, 347 (1–3), 139–150.

    https://doi.org/10.1023/A:1003031621659

    Särkkä, J. & Mäkelä, J. (1998) Troglochaetus beranecki Delachaux (Polychaeta, Archiannelida) in esker groundwaters of Finland: a new class of limnic animals for northern Europe. Hydrobiologia, 379 (1–3), 17–21.

    https://doi.org/10.1023/A:1003292202048

    Särkkä, J. & Mäkelä, J. (1999) Meiofauna of esker groundwaters in Finland. Hydrobiologia, 405, 25–37.

    https://doi.org/10.1023/A:1003710829888

    Schill, R.O. (2018) Water Bears: The Biology of Tardigrades. Springer International Publishing, Cham, 419 pp.

    https://doi.org/10.1007/978-3-319-95702-9

    Schill, R.O., Förster, F., Dandekar, T. & Wolf, M. (2010) Using compensatory base change analysis of internal transcribed spacer 2 secondary structures to identify three new species in Paramacrobiotus (Tardigrada). Organisms Diversity & Evolution, 10 (4), 287–296.

    https://doi.org/10.1007/s13127-010-0025-z

    Schultze, C.A.S. (1834) Macrobiotus Hufelandii animal e crustaceorum classe novum, reviviscendi post diuturnam asphixiam et aridiatem potens. Berolini Apud Carolum Curths, 1834, 1–8.

    Scourfield, D.J. (1897) Contributions to the non-marine fauna of Spitsbergen. Part I. Preliminary notes, and report on the Rhizopoda, Tardigrada, Entomostraca. Proceedings of the Zoological Society of London, 1897, 784–792.

    https://doi.org/10.1111/j.1096-3642.1897.tb03120.x

    Stark, S. & Kristensen, R.M. (1999) Tardigrades in the soil of Greenland. Berichte zur Polar- und Meeresforschung, 330, 44–63.

    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). Systematic and Biodiversity, 16 (4), 357–376.

    https://doi.org/10.1080/14772000.2018.1424267

    Stec, D., Krzywański, Ł., Zawierucha, K. & Michalczyk, Ł. (2020a)a Untangling systematics of the Paramacrobiotus areolatus species complex by an integrative redescription of the nominal species for the group, with multilocus phylogeny and species delineation in the genus Paramacrobiotus. Zoological Journal of the Linnean Society, 188 (3), 694–716.

    https://doi.org/10.1093/zoolinnean/zlz163

    Stec, D., Tumanov, D.T. & Kristensen, R.M. (2020b) Integrative taxonomy identifies two new tardigrade species (Eutardigrada: Macrobiotidae) from Greenland. European Journal of Taxonomy, 614, 1–40.

    https://doi.org/10.5852/ejt.2020.614

    Suzuki, A.C. (2016) Specimens with an artifact appearing as ‘three spines’ in Milnesium tardigradum var. trispinosa Rahm, 1931 (Tardigrada). Zoological science, 33 (4), 431–433.

    https://doi.org/10.2108/zs150184

    Suzuki, A.C., Tsujimoto, M., Kagoshima, H. & Imura, S. (2016) An undescribed tardigrade of genus Milnesium from Inhovde, East Antarctica. The 7th Symposium on Polar Science, National Institute of Polar Research, Tachikawa, Japan, 2016, 1. Available from: https://www.nipr.ac.jp/symposium2016/abstract/abstract.zip (accessed 3 August 2020)

    Thulin, G. (1911) Beiträge zur Kenntnis der Tardigradenfauna Schwedens. Arkiv för Zoologi, 7 (16), 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

    Toftner, E.C., Grigarick, A.A. & Schuster, R.O. (1975) Analysis of scanning electron microscope images of Macrobiotus eggs. Memorie dell’Istituto Italiano di Idrobiologia, 32, 393–411.

    Vecchi, M., Cesari, M., Bertolani, R., Jönsson, K.I., Rebecchi, L. & Guidetti, R. (2016a) Corrigendum to: Integrative systematic studies on tardigrades from Antarctica identify new genera and new species within Macrobiotoidea and Echiniscoidea. Invertebrate Systematics, 30 (5), 521–521.

    https://doi.org/10.1071/IS15033_CO

    Vecchi, M., Vicente, F., Guidetti, R., Bertolani, R., Rebecchi, L. & Cesari, M. (2016b) Interspecific relationships of tardigrades with bacteria, fungi and protozoans, with a focus on the phylogenetic position of Pyxidium tardigradum (Ciliophora). Zoological Journal of the Linnean Society, 178 (4), 846–855.

    https://doi.org/10.1111/zoj.12446

    Venables, W.N. & Ripley, B.D. (2002) Modern Applied Statistics with S. Springer, New York, 498 pp.

    https://doi.org/10.1007/978-0-387-21706-2

    Vicente, F., Fontoura, P., Cesari, M., Rebecchi, L., Guidetti, R., Serrano, A. & Bertolani, R. (2012) Integrative taxonomy allows the identification of synonymous species and a new genus of Tardigrada Echiniscidae (Heterotardigrada). Zootaxa, 3613 (6), 557–572.

    https://doi.org/10.11646/zootaxa.3613.6.3