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

• Autorzy: Zawierucha K. , Stec D. , Lachowska-Cierlik D. , Takeuchi N. , Li Z. , Michalczyk L.
• Języki publikacji: EN

Abstrakty

EN

Glaciers and ice sheets are considered a biome with unique organism assemblages. Tardigrada (water bears) are micrometazoans that play the function of apex consumers on glaciers. Cryoconite samples with the dark-pigmented tardigrade Cryoconicus gen. nov. kaczmareki sp. nov. were collected from four locations on glaciers in China and Kyrgyzstan. The erection of the new genus is based on a unique combination of morphological traits as well as on phylogenetic analyses. The analysis of COI sequences in the new species revealed high genetic differentiation with 9 haplotypes shared among 13 sequenced individuals from three sequenced populations. There was no apparent geographic structure in COI haplotype diversity, which might indicate effective dispersal abilities of the new species. A recovery of numerous live individuals from a sample that was frozen for 11 years suggests high survival rates in the natural environment. The ability to withstand low temperatures, combined with dark pigmentation that is hypothesised to protect from intense UV radiation, could explain how the new taxon is able to dwell in an extreme glacial habitat. We also found that a rare mountain tardigrade Ramazzottius cataphractus (Maucci, 1974) is morphologically similar to the new species, therefore we propose to transfer it to the new genus. Our study indicates that glacier invertebrate fauna is still poorly known and requires intense research.

• Wydawca: -
• Czasopismo: Annales Zoologici
• Rocznik: 2018
• Tom: 68
• Numer: 1
• Opis fizyczny: p.179-201,fig.,ref.

Twórcy

autor

Zawierucha K.

• Department of Animal Taxonomy and Ecology, Adam Mickiewicz University, Umultowska 89, 61-614, Poznań, PolandLaboratory of Fish Genetics, Institute of Animal Physiology and Genetics, Academy of Sciences of the Czech Republic, Rumburska 89, 277-21 Libechov, Czech Republic

autor

Stec D.

• Institute of Zoology and Biomedical Research, Jagiellonian University, Gronostajowa 9, 30-387 Krakow, Poland

autor

Lachowska-Cierlik D.

• Institute of Zoology and Biomedical Research, Jagiellonian University, Gronostajowa 9, 30-387 Kraków, Poland

autor

Takeuchi N.

• Department of Earth Sciences, Graduate School of Science, Chiba University, Chiba, Japan

autor

Li Z.

• State Key Laboratory of Cryospheric Sciences and Tien Shan Glaciological Station, Chinese Academy of Sciences, 320 Donggang West Road, Lanzhou 730000, China

autor

Michalczyk L.

• Institute of Zoology and Biomedical Research, Jagiellonian University, Gronostajowa 9, 30-387 Kraków, Poland

Bibliografia

• Altschul, S. F., Gish, W., Miller, W., Myers, Е. W. and D. J. Lipman. 1990. Basic local alignment search tool. Journal of Molecular Biology, 215: 403-410.
• Anesio, A. M. and J. Laybourn-Parry. 2012. Glaciers and ice sheets as a biome. Trends in Ecology and Evolution, 4: 219-225.
• Bandelt, H., Forster, P. and A. Röhl. 1999. Median-joining networks for inferring intraspecific phytogenies. Molecular Biology and Evolution, 16: 37-48.
• Beasley, C. W., Kaczmarek, Ł. and Ł. Michalczyk. 2008. Doryphoribius mexicanus, a new species of Tardigrada (Eutardigrada, Hypsibiidae) from Mexico (North America). Proceedings of the Biological Society of Washington, 121: 34-40.
• Bertolani, R., Guidetti, R., Marchioro, T., Altiero, T., Rebecchi, L. and M. Cesari. 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.
• Bertolani, R. and I. M. Kinchin. 1993. A new species of Ramazzottius (Tardigrada, Hypsibiidae) in a rain gutter sediment from England. Zoological Journal of the Linnean Society, 109: 327-333.
• Biserov, V. I.1985. Hypsibius subanomalus sp. n. (Eutardigrada, Hypsibiidae) from the Astrakhan District. Zoologicheskii Zhurnal, 64: 131-135.
• Biserov, V. I. 1999. A review of the Tardigrada from Novaya Zemlya; with description of three new species and an evaluation of the state of the environment in the region. Zoologischer Anzeiger, 238: 169-182.
• Blumthaler, M., Ambach, W. and R. Ellinger. 1997. Increase in solar UV radiation with altitude. Journal of Photochemistry and Photobiology B: Biology, 39:130-134.
• Boetius, A., Anesio, A. M., Deming, J. W., Mikucki, J. A. and J. Z. Rapp. 2015. Microbial ecology of the cryosphere: sea ice and glacial habitats. Nature Reviews Microbiology, 13: 677-690.
• Bolch, T. 2007. Climate change and glacier retreat in northern Tien Shan (Kazakhstan/Kyrgyzstan) using remote sensing data. Global and Planetary Change, 56:1-12.
• Cesari, M., McInnes, S. J., Bertolani, R., Rebecchi, L. and R. Guidetti. 2016. Genetic diversity and biogeography of the south polar water bear Acutuncus antaroticus (Eutardigrada : Hypsibiidae) - evidence that it is a truly pan-Antarctic species. Invertebrate Systematics, 30(6): 635-649.
• Cook, J. M., Edwards, A., Bulling, M., Mur, L. A. J., Cook S., Gokul, J. K., Cameron, K. A., Sweet, M., and T. D. L. Irvine-Fynn. 2016. Metabolome-mediated biocryomorphic evolution promotes carbon fixation in Greenlandic cryoconite holes. Environmental Microbiology, 18: 4674-4686.
• Cook, J., Edwards, A., Takeuchi, N. and T. Irvine-Fynn. 2015. Cryoconite. The dark biological secret of the cryosphere. Progress in Physical Geography, 40: 66-111.
• D'Haese, J., Traore-Freitag, A., Kiehl, E., Prasath, T. and H. Greven. 2011. The actin gene of Hypsibius klebelsbergi (Eutardigrada) - complete sequence and comparison with actin from related and non-related taxa. Journal of Zoological Systematics and Evolutionary Research, 49: 84-89.
• Dabert, M., Dastych, H. and J. Dabert. 2015. Molecular data support the dispersal ability of the glacier tardigrade Hypsibius klebelsbergi Mihelčič, 1959 across the environmental barrier (Tardigrada). Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 17: 233-240.
• Dastych, H. 1985. West Spitsbergen Tardigrada. Acta Zoologica Cracoviensia, 28: 169-214.
• Dastych, H. 2004. Hypsibius thaleri sp. nov., a new species of a glacier-dwelling tardigrade from the Himalayas, Nepal (Tardigrada). Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 101: 169-183.
• Dastych, H. 2009. Thalerius konradi gen. nov., sp. nov., a new tardigrade from the periglacial area of the Otztal Alps, Austria (Tardigrada). Contributions to Natural History, 12: 391-402.
• Dastych, H., Kraus, H. J. and K. Thaler. 2003. Redescription and notes on the biology of the glacier tardigrade Hypsibius klebelsbergi Mihelčič, 1959 (Tardigrada), based on material from Ötztal Alps, Austria. Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 100: 73-100.
• Di Mauro, В., Baccolo, G., Garzonio, R., Giardino, C., Massabo, D., Piazzalunga, A., Rossini, M. and R. Colombo. Impact of impurities and cryoconite on the optical properties of the Morteratsch Glacier (Swiss Alps). The Cryosphere, 11: 2393-2409.
• Degma, P., Bertolani, R. and R. Guidetti. 2015. Actual checklist of Tardigrada species. http://www.tardigrada.modena.unimo.it/miscellanea /Actual%20checklist%20of%20Tardi grada.pdf
• Degma, P. and R. Guidetti. 2007. Notes to the current checklist of Tardigrada. Zootaxa, 1579: 41-53.
• Dial, C. R., Dial, R. J., Saunders, R., Lang, S. A., Lee, B., Wimberger, P., Dinapoli, M. S., Egiazarov, A. S., Gipple, S. L., Maghirang, M. R., Swartley-McArdle, D. J., Yudkovitz, S. R., and D. H. Shain. 2012. Historical biogeography of the North American glacier ice worm, Mesenchytraeus solifugus (Annelida: Oligochaeta: Enchytraeidae). Molecular Phylogenetics and Evolution, 63: 577-584.
• Drygalski, Е. V. 1897. Die Kryoconitlcher. In: Grönland - Expedition der Gesellschaft für Erdkunde zu Berlin 1891-1893: 93-103. Kuhl, W.H. (Ed.). Berlin: Heeausgegebex von dek.
• Doyère, P. L. N. 1840. Memoire sur les Tardigrades. Annales des Sciences Naturelles, Zoologie, Paris, Series 2, 14: 269-362.
• Eibye-Jacobsen, J. 2001. A new method for making SEM preparations of the tardigrade buccopharyngeal apparatus. Zoologischer Anzeiger, 240: 309-319.
• Farinotti, D., Longuevergne, L., Moholdt, G., Duethmann, D., Mölg, T., Bolch, T., Vorogushyn, S. and A. Güntner. 2015. Substantial glacier mass loss in the Tien Shan over the past 50 years. Nature Geoscience, 8: 716-722.
• Faurby, S., Jonsson, K. I., Rebecchi, L. and P. Funch. 2008. Variation in anhydrobiotic survival of two eutardigrade morphospecies: a story of cryptic species and their dispersal. Journal of Zoology, 275:139-145.
• Ferrario, C., Pittino, F., Tagliaferri, I., Gandolfi, I., Bestetti, G., Azzoni, R. S., Diolaiuti, G., Franzetti, A., Ambrosini, R. and S. Villa. 2017. Bacteria contribute to pesticide degradation in cryoconite holes in an Alpine glacier. Environmental Pollution, 230: 919-926.
• Folmer, O., Black, M., Hoeh, W., Lutz, R. and R. Vrijenhoek. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3: 294-299.
• Franzetti, F., Tagliaferri, I., Gandolfi, I., Bestetti, G., Minora, G., Mayer, C., Azzoni, R. S., Diolaiuti, G., Smiraglia, C., and R. Ambrosini. 2016. Light-dependent microbial metabolisms drive carbon fluxes on glacier surfaces. The ISME Journal 10: 1-5.
• Gawor, J., Grzesiak, J., Sasin-Kurowska, J., Borsuk, P., Gromadka, R., Górniak, D., Świątecki, A., Aleksandrzak- Piekarczyk, T. and M. K. Zdanowski. 2016. Evidence of adaptation, niche separation and microevolution within the genus Polaromonas on Arctic and Antarctic glacial surfaces. Extremophiles, 20: 403-413.
• Gąsiorek, P., Stec, D., Morek, W. and Ł. Michalczyk. 2017. An integrative redescription of Echiniscus testudo (Doyére, 1840), the nominal taxon for the class Heterotardigrada (Ecdysozoa: Panarthropoda: Tardigrada). Zoologischer Anzeiger, 270: 107-122.
• Gąsiorek, P., Stec, D., Morek, W. and L. Michalczyk. In press. An integrative redescription of Hypsibius dujardini Doyère, 1840, the nominal taxon for Hypsibioidea (Tardigrada: Eutardigrada). Zootaxa.
• Gąsiorek, P., Stec, D., P., Morek, W., Zawierucha, K, Kaczmarek, Ł., Lachowska-Cierlik, D. and Ł. Michalczyk. 2016. An integrative revision of Mesocrista Pilato, 1987 (Tardigrada: Eutardigrada: Hypsibiidae). Journal of Natural History, 50: 2803-2828.
• Gąsiorek, P., Zawierucha, K., Stec, D. and L. Michalczyk. 2017. Integrative redescription of a common Arctic water bear Pilatobius recamieri (Richters, 1911). Polar Biology, 40: 2239-2252.
• Greven, H., Dastych, H. and H. J. Kraus. 2005. Notes on the integument of the glacier-dwelling tardigrade Hypsibius klebelsbergi Mihelčič, 1959 (Tardigrada). Mitteilungen aus den Hamburgischen Zoologischen Museum und Institut, 102: 11-20.
• Grzesiak, J., Górniak, D., Świątecki, A., Aleksandrzak-Piekarczyk, T., Szatraj, K. and M. K. Zdanowski. 2015. Microbial community development on the surface of Hans and Werenskiold Glaciers (Svalbard, Arctic): a comparison. Extremophiles, 19: 885-897.
• Guidetti, R., Altiero, T., Marchioro, T., Sarzi Amade, L., Avdonina, A. M., Bertolani, R. and L. Rebecchi. 2012. Form and function of the feeding apparatus in Eutardigrada (Tardigrada). Zoomorphology, 131:127-148.
• Guidetti, R., Rizzo, A. M., Altiero, T. and L. Rebecchi. 2012. What can we learn from the toughest animals of the Earth? Water bears (tardigrades) as multicellular model organisms in order to perform scientific preparations for lunar exploration. Planetary and Space Science, 74: 97-102.
• Guidetti, R. and R. Bertolani. 2005. Tardigrade taxonomy: an updated check list of the taxa and a list of characters for their identification. Zootaxa, 845:1-46.
• Guil, N. and S. Sanchez-Moreno. 2013. Fine-scale patterns in micrometazoans: tardigrade diversity, community composition and trophic dynamics in leaf litter. Systematics and Biodiversity, 11: 181-193.
• Guil, N. and G. Giribet. 2012. A comprehensive molecular phylogeny of tardigrades-adding genes and taxa to a poorly resolved phylum-level phytogeny. Cladistics, 27: 1-29.
• Guil, N., Machordom, A. and R. Guidetti. 2013. High level of phenotypic homoplasy amongst eutardigrades (Tardigrada) based on morphological and total evidence phylogenetic analyses. Zoological Journal of the Linnean Society, 169: 1-26.
• Hall, T. A 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41: 95-98.
• Hartzell, P. L., Nghiem, J. V., Richio, K. J., and D. H. Shain. 2005. Distribution and phytogeny of glacier ice worms (Mesenchytraeus solifugus and Mesenchytraeus solifugus rainierensis). Canadian Journal of Zoology, 83: 1206-1213.
• Hashimoto, T., Horikawa, D. D., Saito, Y., Kuwahara, H., Kozuka-Hata, H., et al. 2016. Extremotolerant tardigrade genome and improved radiotolerance of human cultured cells by tardigrade-unique protein. Nature Communications, 7: 12808.
• Hodson, A., Anesio, A. M., Tranter, M., Fountain, A., Osborn, M., Priscu, J., Laybourn-Parry, J. and B. Sattler. 2008. Glacial ecosystems. Ecological Monographs, 78: 41-67.
• Hodson, A., Cameron, K., Bøggild, C., Irvine-Fynn, T., Langford, H., Pearce, D. and S. Banwart. 2010. The structure, biological activity and biogeochemistry of cryoconite aggregates upon an Arctic valley glacier: Longyearbreen, Svalbard. Journal of Glaciology, 56: 349-362.
• Hotaling, S., Hood, E. and T. E. Hamilton. 2017. Microbial ecology of mountain glacier ecosystems: biodiversity, ecological connections, and implications of a warming climate. Environmental Microbiology, doi:10.1111/l462-2920.13766.
• Huelsenbeck, J. P. and F. Ronquist. 2001. MrBAYES: Bayesian inference of phytogeny. Bioinformatics, 17: 754-755.
• Huelsenbeck, J. P., Ronquist, F., Nielsen, R. and J. P. Bolback. 2001. Bayesian inference of phytogeny and its impact on evolutionary biology. Science, 294: 2310-2314.
• Jørgensen, A., Faurby, S., Persson, D. K, Halberg, K. A., Kristensen, R. M. and N. Møbjerg. 2013. Genetic diversity in the parthenogenetic reproducing tardigrade Echiniscus testudo (Heterotardigrada: Echiniscoidea). Journal of Limnology, 72: 136-143.
• Jørgensen, M. and R. M. Kristensen. 1991. Meiofauna investigations from Igloolik, N.W.T., Arctic Canada. In: Arctic Biology Course 1989, Igloolik N.W.T. Canada. M. Jørgensen (ed.): 61-80 pp.
• Jørgensen, A. and R. M. Kristensen. 2004 Molecular phytogeny of Tardigrada - investigation of the monophyly of Heterotardigrada. Molecular Phylogenetics and Evolution, 32: 666-670.
• Jørgensen, A., Møbjerg, N. and R. M. Kristensen. 2007. A molecular study of the tardigrade Echiniscus testudo (Echiniscidae) reveals low DNA sequence diversity over a large geographical area. Journal of Limnology, 66: 77-83.
• Jönsson, K. I. and R. Bertolani. 2001. Facts and fiction about long-term survival in tardigrades. Journal of Zoology, 255: 121-123.
• Kaczmarek, Ł., Cytan, J., Zawierucha, K., Diduszko, D. and Ł. Michalczyk. 2014a. Tardigrades from Peru (South America), with descriptions of three new species of Parachela. Zootaxa, 3790: 357-379, doi:10.11646/zootaxa.3790.2.5.
• Kaczmarek, Ł., Janko, K., Smykla, J. and Ł. Michalczyk. 2014b. Soil tardigrades from the Antarctic Peninsula with a description of a new species and some remarks on the genus Ramajendas (Eutardigrada: Isohypsibiidae). Polar Record, 50: 176-182.
• Kaczmarek, Ł., Michalczyk, Ł. and D. Diduszko. 2006. Ramazzottius bunikowskae, a new species of Tardigrada (Eutardigrada, Hypsibiidae) from Russia. Zootaxa, 1229: 49-57.
• Kibbe, W. A. 2007. OligoCalc: an online oligonucleotide properties calculator. Nucleid Acids Research, 35: 43-46.
• Kiehl, E., Dastych, H., D'Haese, J. and H. Greven. 2007. The 18S rDNA sequences support polyphyly of the Hypsibiidae (Eutardigrada). Journal of Limnology, 66: 21-25.
• Li, Z. and Li, H. and Y. J. Chen. 2011. Mechanisms and simulation of accelerated shrinkage of continental glaciers: A case study of Urumqi Glacier No. 1 in eastern Tianshan, Central Asia. Journal of Earth Science, 22: 423. doi: 10. 1007/S12583-011-0194-5
• Makowska, N., Zawierucha, K., Mokracka, J. and R. Koczura. 2016. First report of microorganisms of Caucasus glaciers (Georgia). Biologia, 71: 620-625.
• Marley, N. J., McInnes, S. J. and C. J. Sands. 2011. Phylum Tardigrada: A re-evaluation of the Parachela. Zootaxa, 2819: 51-64.
• Maucci, W. 1974. Hypsibius (H.) cataphractus (Tardigrada: Macrobiotidae) und weitere Nachrichten uber Tardigraden aus Österreich. Bericht des Naturwissenschaftlisch- Medizinischen Vereins in Innsbruck, 61: 83-86.
• Maucci, W. (1991). Tre nuove specie di Eutardigradi delia Groenlandia meridionale. Bollettino del Museo Civico di Storia Naturale di Verona, 15: 279-289.
• Maucci, W. 1996. Tardigrada of the Arctic tundra with description of two new species. Zoological Journal of the Linnean Society, 116:185-204.
• McIntyre, N. F. 1984. Cryoconite hole thermodynamics. Canadian Journal of Earth Science, 21:152-156.
• Michalczyk, L. and L. Kaczmarek. 2013. The Tardigrada Register: a comprehensive online data repository for tardigrade taxonomy. Journal of Limnology, 72: 175-181.
• Michalczyk, L., Wełnicz, W., Frohme, M. and Ł. Kaczmarek. 2012. Redescriptions of three Milnesium Doyère, 1840 taxa (Tardigrada: Eutardigrada: Milnesiidae), including the nominal species for the genus. Zootaxa, 3154: 1-20.
• Mihelčič, F. 1959. Zwei neue Tardigraden aus der Gattung Hypsibius Thulin aus Osttirol (Österreich). Systematisches zur Gattung Hypsibius Thulin. Zoologischer Anzeiger, 163: 254-261.
• Mironov, S. V., Dabert, J. and M. Dabert. 2012. A new feather mite species of the genus Proctophyllodes Robin, 1877 (Astigmata: Proctophyllodidae) from the Long-tailed Tit Aegithalos caudatus (Passeriformes: Aegithalidae): morphological description with DNA barcode data. Zootaxa, 3253: 54-61.
• Morek, W., Gąsiorek, P., Stec, D., Blagden, В. and L. Michalczyk. 2016. Experimental taxonomy exposes ontogenetic variability and elucidates the taxonomie value of claw configuration in Milnesium Doyère, 1840 (Tardigrada: Eutardigrada: Apochela). Contributions to Zoology, 85(2): 173-200.
• Morek, W., Stec, D., Gąsiorek, P., Schill, R. O., Kaczmarek, L. and L. Michalczyk. 2016. An experimental test of eutardigrade preparation methods for light microscopy. Zoological Journal of the Linnean Society, 178(4): 785-793.
• Mueller, D. R., Vincent, W. F., Pollard, W. H. and C. H. Fristen. 2001. Glacial cryoconite ecosystems: a bipolar comparison of algal communities and habitats. Nova Hedwigia. Beiheft, 123: 173-197.
• Narama, Ch., Kääb, A., Duishonakunov, M. and K. Abdrakhmatov. 2010. Spatial variability of recent glacier area changes in the Tien Shan Mountains, Central Asia, using Corona (~ 1970), Landsat (~ 2000), and ALOS (~ 2007) satellite data. Global and Planetary Change, 71: 42-54.
• Nelson, D. R., Guidetti, R. and L. Rebecchi. 2015. Phylum Tardigrada. In: J. Thorp and D.C. Rogers (eds) Ecology and General Biology. Thorp and Covich's Freshwater Invertebrates, Academic Press, Massachusetts, 347-380 pp.
• Nylander, J. A. A., Ronquist, F., Huelsenbeck, J. P. and J. L. Nieves. 2004. Bayesian phylogenetic analysis of comined data. Systematic Biology, 53: 47-67.
• Page, R. D. 1996. TreeView: an application to display phylogenetic trees on personal computers. Computer Applications in the Biosciences, 12: 357-358.
• Pilato, G. 1981. Analisi di nuovi caratteri nello studio degli Eutardigradi. Animalia, 8: 51-57.
• Pilato, G. 1987. Revision of the genus Diphascon Plate, 1889, with remarks on the subfamily Itaquasconinae (Eutardigrada, Hypsibiidae). Biology of Tardigrades. Selected Symposia and Monographs U.Z.I., 1: 337-357
• Pilato, G. 2013. The taxonomie value of the structures for the insertion of the stylet muscles in the Eutardigrada, and description of a new genus. Zootaxa, 3721(4): 365-378.
• Pilato, G. and M. G. Binda. 2010. Definition of families, subfamilies, genera and subgenera of the Eutardigrada, and keys to their identification. Zootaxa 2404: 1-54.
• Pilato, G. and M. G. Binda. 1990. Tardigradi dell Antartide. I. Ramajendas, nuovo genere di Eutardigrado. Nuova posizione sistematica di Hypsibius renaudi Ramazzotti, 1972 е descrizione di Ramajendas frigidus n. sp. Animalia, 17: 61-71.
• PopART ver.1.7. 2016. Available via http://popart.otago.ac.nz
• Potapow, M. 2001. Isotomidae. In: Dunger, W. (ed.). Pp. synopses on Palearctic collembolan 3: 1-603. Görlitz: Abhandlungen und Berichte des Naturkundemuseums Görlitz.
• Prendini, L., Weygoldt, P. and W. C. Wheeler. 2005. Systematics of the Damon variegatus group of African whip spiders (Chelicerata: Amblypygi): evidence from behaviour, morphology and DNA. Organisms, Diversity and Evolution, 5: 203-236.
• Ramazzotti, G. 1968. Tardigradi dei pozzetti glaciali di fusione (Kryokonitlöcher) dell'Himalaya. Khumbu Himal: Universitätsverlag, Wagner, Innsbrück, München, 3: 1-3.
• Ramazzotti, G. and W. Maucci. 1983. II Phylum Tardigrada. Memorie dell'Istituto Italiano di Idrobiologia, Pallanza, 41: 1-1012.
• Rambaut, A. and A. J. Drummond. 2009. Tracer ver. 1.5. Available at: http://beast.bio.ed.ac.uk/Tracer.
• Richters, F. 1904. Vorläufiger Bericht über die antarktische Moosfauna. Verhandlungen der deutschen Zoologisehen Gesellschaft EV, 236-239.
• Richters, F. 1926. Tardigrada. In: Handbuch der Zoologie Vol. 3. Ed. By Kükenthal, W. & Krumbach, T. Walter de Gruyter & Co. Berlin and Leipzig, pp. 58-61.
• Sands, C. J., McInnes, S. J., Marley, N. J., Goodall-Copestake, W., Convey, P. and K. Linse. 2008. Phylum Tardigarda: an "individual" approach. Cladistics, 24: 1-18.
• Schuster, R. O., Nelson, D. R., Grigarick, A. A. and D. Christenberry. 1980. Systematic criteria of the Eutardigrada. Transactions of the American Microscopical Society, 99: 284-303.
• Shain, D. H., Carter, M. R., Murray, K. P., Maleski, K. A, Smith, N. R., McBride, T. R., Michalewicz, L. A. and W. M. Saidel. 2000. Morphologic characterization of the ice worm Mesenchytraeus solifugus. Journal of Morphology, 246: 192-197.
• Shain, D. H., Halldórsdóttir, K, Pálsson, F., Aðalgeirsdóttir, G., Gunnarsson, A., Jónsson, P., et al. 2016. Colonization of maritime glacier ice by bdelloid Rotifera. Molecular Phylogenetics and Evolution, 98: 280-287.
• Sommaruga, R. 2001. The role of solar UV radiation in the ecology of alpine lakes. Journal of Photochemistry and Photobiology B: Biology, 62: 35-42.
• Stamatakis, A. 2014. RAxML version 8: a tool for phylogenetic analysis and postanalysis of large phytogenies. Bioinformatics, 30: 1312-1313.
• Stec, D., Gąsiorek, P., Morek, W., Kosztyla, P., Zawierucha, K., Michno, K., Kaczmarek, Ł., Prokop, Z. M. and M. Michalczyk. 2016. Estimating optimal sample size for tardigrade morphometry. Zoological Journal of the Linnean Society, 178(4): 776-784.
• Stec, D., Morek, W., Gąsiorek, P., Kaczmarek, Ł. and Ł. Michalczyk. 2016. Determinants and taxonomie consequences of extreme egg shell variability in Ramazzottins subanomalus (Biserov, 1985) (Tardigrada). Zootaxa, 4208(2): 176-188.
• Stec, D., Smolak, R., Kaczmarek, Ł. and Ł. Michalczyk. 2015. An integrative description of Macrobiotus paulinae sp. nov. (Tardigrada: Eutardigrada: Macrobiotidae: hufelandi group) from Kenya. Zootaxa, 4052(2): 501-526.
• Stec, D., Zawierucha, K. and L. Michalczyk. 2017. An integrative description of Ramazzottins subanomalus (Biserov, 1985) (Tardigrada) from Poland. Zootaxa, 4300(3): 403-420.
• Stibal, M., Lawson, E. C., Lis, G. P., Mak, K M., Wadham, J. L., A. M. Anesio. 2010. Organic matter content and quality in supraglacial debris across the ablation zone of the Greenland ice sheet. The Annals of Glaciology, 51:1-8.
• Stibal, M., Telling, J., Cook, J., Mak, K. M., Hodson, A. and A. M. Anesio. 2012. Environmental Controls on Microbial Abundance and Activity on the Greenland Ice Sheet: A Multivariate Analysis Approach. Microbial Ecology, 63: 74-84.
• Stibal, M., Schostag, M., Cameron, K. A., Hansen, L. H., Chandler, D. M., Wadham, J. L. and C. S. Jacobsen. 2015. Different bulk and active bacterial communities in cryoconite from the margin and interior of the Greenland ice sheet. Environmental Microbiology Reports, 7: 293-300.
• Swofford, D. L. 2002. PAUP: Phylogenetic Analysis Using Parsimony and Other Methods, Version 4.0. Sinauer Associates, Sunderland, Massachusetts.
• Takeuchi, N., Kohshima, S. and K. Seko. 2001. Structure, formation, and darkening process of albedo-reducing material (cryoconite) on a Himalayan glacier: a granular algal mat growing on the glacier. Arctic Antarctic and Alpine Research, 33: 115-122.
• Takeuchi, N., and Z. Li. 2008. Characteristics of surface dust on Ürümqi Glacier No. 1 in the Tien Shan Mountains, China. Arctic, Antarctic, and Alpine Research, 40(4), 744-750.
• Takeuchi, N., Matsuda, Y., Sakai, A. and K. Fujita. 2005. A large amount of biogenic surface dust (cryoconite) on a glacier in the Qilian Mountains, China. Bulletin of Glaciological Research, 22, 1-8.
• Takeuchi, N., Nakawo, M., Narita, H. and J. Han. 2008. Miaoergou Glaciers in the Kalik Mountains, western China: Report of a reconnaissance for future ice core drilling and biological study. Bulletin of Glaciological Research, 26: 33-40.
• Tamura, K., Stecher, G., Peterson, D., Filipski, A. and S. Kumar. 2013. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30: 2725-2729.
• Thompson, J. D., Higgins, D. G. and T. J. Gibson. 1994. CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research, 22: 4673-4680.
• Tsujimoto, M., Imura, S. and H. Kanda. 2016. Recovery and reproduction of an Antarctic tardigrade retrieved from a moss sample frozen for over 30 years. Cryobiology, 72(1): 78-81.
• Vaughan, D. G., Comiso, J. C., Allison, I., Carrasco, J., Kaser, G., Mote, P., et al. 2013. Observations: Cryosphere. In: Stocker, T. F., Qin, D., Plattner, G. K., Tignor, M., Allen, S. K., Boschung, J. (ed). Contribution of working group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press 2013, Cambridge, United Kingdom and New York, USA.
• Wełnicz, W., Grohme, M. A., Kaczmarek, Ł., Schill, R. O. and M. Frohme. 2011. ITS-2 and 18S rRNA data from Macrobiotus polonicus and Milnesium tardigradum (Eutardigrada, Tardigrada). Journal of Zoological Systematics and Evolutionary Research, 49: 34-39.
• White, T. J., Bruns, T., Lee, S. and J. Taylor. 1990. PCR protocols: a guide to methods and application. Academic Press, San Diego, California, 315-322 pp.
• World Glacier Monitoring Service 2008. Global glacier changes: facts and Fig. URL: www.grid.unep.ch/glaciers.
• Wharton, R. A., McKay, C. P., Simmons, G. M. and B. C. Parker. 1985. Cryoconite holes on glaciers. Bioscience, 35: 499-503.
• Wright, J. C. 2001. Cryptobiosis 300 years on from van Leeuwenhoek: what have we learned about tardigrades? Zoologischer Anzeiger, 240: 563-582.
• Yallop, M. L., Anesio, A. M., Perkins, R. G., Cook, J., Telling, J., Fagan, D., MacFarlane, J., Stibal, M., Baker, G., Bellas, C., Hodson, A., Tranter, M., Wadham, J. and N. W. Roberts. 2012. Photophysiology and albedo-changing potential of the ice-algal community on the surface of the Greenland ice sheet. ISME Journal, 6: 2302-2313.
• Zawierucha, K., Buda, J., Pietryka, M., Richter, D., Łokas, E., Lehmann-Konera, S., Makowska, N. and M. Bogdziewicz. 2018. Snapshot of micro-animals and associated biotic and abiotic environmental variables on the edge of south-west Greenland ice sheet. Limnology, 19: 141-150.
• Zawierucha, K., Kolicka, M., Takeuchi, N. and Ł. Kaczmarek. 2015. What animals can live in cryoconite holes? A faunal review. Journal of Zoology, 295:159-169.
• Zawierucha, K., Ostrowska, M., Vonnahme, T. R., Devetter, M., Nawrot, A. P., Lehmann, S. and M. Kolicka. 2016. Diversity and distribution of Tardigrada in Arctic cryoconite holes. Journal of Limnology, 75(3): 545-559.

Identyfikatory

DOI

10.3161/00034541ANZ2018.68.1.007