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Research Article

Diverse eggs, diverse species? Production of two egg morphotypes in Paramacrobiotus bifrons, a new eutardigrade species within the areolatus group

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Pages 274-297 | Received 07 Aug 2023, Accepted 19 Jan 2024, Published online: 01 Mar 2024

References

  • Altiero T, Rebecchi L. 2001. Rearing tardigrades: results and problems. Zoologischer Anzeiger 240(3–4):217–221. DOI: 10.1078/0044-5231-00028.
  • Baumann H. 1966. Lebenslauf und Lebensweise von Hypsibius (H.) oberhaeuseri Doyère (Tardigrada). Veröffentlichen Überseemuseum Bremen 3:245–258.
  • Bertolani R, Cesari M, Giovannini I, Rebecchi L, Guidetti R, Kaczmarek Ł, Pilato G. 2023. The Macrobiotus persimilis-polonicus complex (Eutardigrada, Macrobiotidae), another example of problematic species identification, with the description of four new species. Organisms Diversity & Evolution 23:329–368. DOI: 10.1007/s13127-022-00599-z.
  • Bertolani R, Rebecchi L. 1988. The tardigrades of Emilia (Italy). I. Rossena. Bolletino di zoologia 55:367–371. DOI: 10.1080/11250008809386634.
  • 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 22:127–152. DOI: 10.1111/j.1463-6409.1993.tb00347.x.
  • Bertolani R, Rebecchi L, Claxton SK. 1996. Phylogenetic significance of eggshell variation in tardigrades. Zoological Journal of the Linnean Society 116(1–2):139–148. DOI: 10.1111/j.1096-3642.1996.tb02339.x.
  • Bertolani R, Rebecchi L, Giovannini I, Cesari M. 2011. DNA barcoding and integrative taxonomy of Macrobiotus hufelandi CAS Schultze 1834, the first tardigrade species to be described, and some related species. Zootaxa 2997:19–36. DOI: 10.11646/zootaxa.2997.1.2.
  • Binda MG. 1980. Tardigradi di Lucania. Animalia 7:79–91.
  • Binda MG, Guglielmino A. 1982. Tardigradi muscicoli e dulcacquicoli di Sardegna. Animalia 9:199–221.
  • Binda MG, Pilato G. 1971. Nuovo contributo alla conoscenza dei Tardigradi di Sicilia. Bollettino delle sedute dell’Accademia Gioenia di Scienze Naturali, Catania 10:896–909.
  • Biserov VI. 1985. Hypsibius subanomalus n. sp. (Eutardigrada, Hypsibiidae) du district d’Astrakhan. Zoologičeskij žurnal 64:131–135.
  • Biserov.1997/98. Tardigrades of the Caucasus with a taxonomic analysis of the genus Ramazzottius (Parachela: Hypsibiidae). Zoologischer Anzeiger 236:139–159.
  • Biserov VI. 1991. An annotated list of Tardigrada from European Russia. Zoologische Jahrbücher System 118:193–216.
  • Bussers JC, Jeuniaux C. 1973. Chitinous cuticle and systematic position of Tardigrada. Biochemical Systematics and Ecology 1:77–78. DOI: 10.1016/0305-1978(73)90040-9.
  • Camarda D, Lisi O, Mifsud D. 2022. First faunistic report of limno-terrestrial tardigrades (Tardigrada Doyère, 1840) from the Maltese Islands. Check List 18(4):779–792. DOI: 10.15560/18.4.779.
  • Camarda D, Massa E, Guidetti R, Lisi O. 2023. A new, simplified, drying protocol to prepare tardigrades for scanning electron microscopy. Microscopy Research and Technique 1–11. DOI: 10.1002/jemt.24460.
  • Cesari M, Bertolani R, Rebecchi L, Guidetti R. 2009. DNA barcoding in Tardigrada: the first case study on Macrobiotus macrocalix Bertolani & Rebecchi 1993 (Eutardigrada, Macrobiotidae). Molecular Ecology Resources 9(3):699–706. DOI: 10.1111/j.1755-0998.2009.02538.x.
  • Cesari M, Giovannini I, Altiero T, Guidetti R, Cornette R, Kikawada T, Rebecchi L. 2022. Resistance to extreme stresses by a newly discovered Japanese tardigrade species, Macrobiotus kyoukenus (Eutardigrada, Macrobiotidae). Insects 13:1–21. DOI: 10.3390/insects13070634.
  • Cesari M, Guidetti R, Rebecchi L, Giovannini I, Bertolani R. 2013. A DNA barcoding approach in the study of tardigrades. Journal of Limnology 72:182–198. DOI: 10.4081/jlimnol.2013.s1.e23.
  • Claxton SK. 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. DOI: 10.3853/j.0067-1975.50.1998.1276.
  • Clement M, Posada D, Crandall K. 2000. TCS: A computer program to estimate gene genealogies. Molecular Ecology 9:1657–1660. DOI: 10.1046/j.1365-294x.2000.01020.x.
  • Clopper CJ, Pearson ES. 1934. The use of confidence or fiducial limits illustrated in the case of the binomial. Biometrika 26(4):404–413. DOI: 10.1093/biomet/26.4.404.
  • Dastych H. 1983. Two new Eutardigrada species from west Spitsbergen and the Tatra MTS. Bulletin de la Société des Amis des Sciences et des Lettres de Poznan 23:195–200.
  • Dastych H. 2011. Ramazzottius agannae sp. nov., a new tardigrade species from the nival zone of the Austrian Central Alps (Tardigrada). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 15:237–253.
  • Degma P, Guidetti R. 2023. Actual checklist of Tardigrada species. DOI: 10.25431/11380_1178608.
  • Devleesschauwer B, Torgerson P, Charlier J, Levecke B, Praet N, Roelandt S, Smit S, Dorny P, Berkvens D, Speybroeck N (2022). prevalence: Tools for prevalence assessment studies. R Package Version 0.4.1. https://cran.r-project.org/package=prevalence
  • Edgar RC. 2004. MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5):1792–1797. DOI: 10.1093/nar/gkh340.
  • Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3:294–299.
  • Gąsiorek P, Stec D, Morek W, Michalczyk Ł. 2017. An integrative redescription of Echiniscus testudo (Doyère, 1840), the nominal taxon for the class Heterotardigrada (Ecdysozoa: Panarthropoda: Tardigrada). Zoologischer Anzeiger 270:107–122. DOI: 10.1016/j.jcz.2017.09.006.
  • Greven H, Kaya M, Baran T. 2016. The presence of α-chitin in Tardigrada with comments on chitin in the Ecdysozoa. Zoologischer Anzeiger 264:11–16. DOI: 10.1016/j.jcz.2016.06.003.
  • Greven H, Peters W. 1986. Localization of chitin in the cuticle of Tardigrada using wheat germ agglutinin-gold conjugate as a specific electron-dense marker. Tissue & Cell 18:297–304. DOI: 10.1016/0040-8166(86)90037-6.
  • Guidetti R, Altiero T, Hansen JG. 2006. A new species of freshwater tardigrades from Disko Island (Greenland) increases an unsolved paradox in tardigrade systematics. Hydrobiologia 558(1):69–79. DOI: 10.1007/s10750-005-1408-6.
  • Guidetti R, Altiero T, Rebecchi L. 2011. On dormancy strategies in tardigrades. Journal of Insect Physiology 57(5):567–576. DOI: 10.1016/j.jinsphys.2011.03.003.
  • Guidetti R, Cesari M, Bertolani R, Altiero T, Rebecchi L. 2019. High diversity in species, reproductive modes and distribution within the Paramacrobiotus richtersi complex (Eutardigrada, Macrobiotidae). Zoological Letters 5:1–28. DOI: 10.1186/s40851-018-0113-z.
  • Guidetti R, Massa E, Bertolani R, Rebecchi L, Cesari M. 2019. Increasing knowledge of Antarctic biodiversity: New endemic taxa of tardigrades (Eutardigrada; Ramazzottiidae) and their evolutionary relationships. Systematics & Biodiversity 17(6):573–593. DOI: 10.1080/14772000.2019.1649737.
  • Guidetti R, Schill RO, Bertolani R, Dandekar T, Wolf M. 2009. New molecular data for tardigrade phylogeny, with the erection of Paramacrobiotus gen. nov. Journal of Zoological Systematics & Evolutionary Research 47(4):315–321. DOI: 10.1111/j.1439-0469.2009.00526.x.
  • Guidetti R, Schill RO, Giovannini I, Massa E, Goldoni SE, Ebel C, Förschler MI, Rebecchi L, Cesari M. 2021. When DNA sequence data and morphological results fit together: Phylogenetic position of Crenubiotus within Macrobiotoidea (Eutardigrada) with description of Crenubiotus ruhesteini sp. nov. Journal of Zoological Systematics & Evolutionary Research 59(3):576–587. DOI: 10.1111/jzs.12449.
  • Hansen J, Katholm AK. 2002. A study of the genus Amphibolus from Disko Island with special attention on the life cycle of Amphibolus nebulosus (Eutardigrada, Eohypsibiidae). In: Hansen JG, editor Arctic biology field course, Qeqertarsuaq, 2002. Copenhagen: Zoological Museum of Denmark, University of Copenhagen. pp. 129–163.
  • Hansen JG, Kristensen RM, Bertolani R, Guidetti R. 2017. Comparative analyses of bertolanius species (Eohypsibiidae; eutardigrada) with the description of Bertolanius birnae sp. nov. From northern polar regions. Polar biology 40:123–140.
  • Hart MW, Sunday J. 2007. Things fall apart: Biological species form unconnected parsimony networks. Biology Letters 3(5):509–512. DOI: 10.1098/rsbl.2007.0307.
  • Iharos G. 1966a. Neue Tardigraden-arten aus Ungarn. Acta Zoologica Academiae Scientiarum Hungaricae 12:111–122.
  • Iharos G. 1966b. A Bakony-hegyseg Tardigrada-faunaja. III. Kulonlenyomat az Allattani Kozlemenyek 53:69–78.
  • Kaczmarek Ł, Bartylak T, Stec D, Kulpa A, Kepel M, Kepel A, Roszkowska M. 2020. Revisiting the genus Mesobiotus Vecchi et al., 2016 (Eutardigrada, Macrobiotidae)–remarks, updated dichotomous key and an integrative description of new species from Madagascar. Zoologischer Anzeiger 287:121–146.
  • Kaczmarek Ł, Cytan J, Zawierucha K, Diduszko D, Michalczyk Ł. 2014. Tardigrades from Peru (South America), with descriptions of three new species of Parachela. Zootaxa 3790(2):357–379. DOI: 10.11646/zootaxa.3790.2.5.
  • Kaczmarek Ł, Gawlak M, Bartels PJ, Nelson DR, 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 67:627–656. DOI: 10.3161/00034541ANZ2017.67.4.001.
  • Kaczmarek Ł, Michalczyk Ł. 2017. The Macrobiotus hufelandi group (Tardigrada) revisited. Zootaxa 4363(1):101–123. DOI: 10.11646/zootaxa.4363.1.4.
  • Kaczmarek Ł, Zawierucha K, Buda J, Stec D, Gawlak M, Michalczyk Ł, Roszkowska M, Rubal 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. Public Library of Science ONE 13(10):e0204756. DOI: 10.1371/journal.pone.0204756.
  • Kayastha P, Mioduchowska M, Warguła J, Kaczmarek Ł. 2023a. A review on the genus Paramacrobiotus (Tardigrada) with a New Diagnostic Key. Diversity 15(9):977. DOI: 10.3390/d15090977.
  • Kayastha P, Stec D, Sługocki Ł, Gawlak M, Mioduchowska M, Kaczmarek Ł. 2023b. Integrative taxonomy reveals new, widely distributed tardigrade species of the genus Paramacrobiotus (Eutardigrada: Macrobiotidae). Scientific Reports 13(1):2196. DOI: 10.1038/s41598-023-28714-w.
  • Kihm JH, Kim S, McInnes SJ, Zawierucha K, Rho HS, Kang P, Park TYS. 2020. Integrative description of a new Dactylobiotus (Eutardigrada: Parachela) from Antarctica that reveals an intraspecific variation in tardigrade egg morphology. Scientific Reports 10:1–11. DOI: 10.1038/s41598-020-65573-1.
  • Kristensen RM, Neuhaus B. 1999. Special issue on tardigrada - the ultrastructure of the tardigrade cuticle with special attention to marine species. Zoologischer Anzeiger 238:261–282.
  • Mapalo MA, Stec D, Mirano-Bascos D, Michalczik Ł. 2017. An integrative description of a limnoterrestrial tardigrade from the Philippines, Mesobiotus insanis, new species (Eutardigrada: Macrobiotidae: Harmsworthi group). The Raffles Bulletin of Zoology 65:440–454.
  • Mapalo MA, Stec D, Mirano-Bascos D, Michalczyk Ł. 2016. Mesobiotus philippinicus sp. nov., the first limnoterrestrial tardigrade from the Philippines. Zootaxa 4126(3):411–426. DOI: 10.11646/zootaxa.4126.3.6.
  • Massa E, Guidetti R, Cesari M, Rebecchi L, Jönsson KI. 2021. Tardigrades of Kristianstads Vattenrike biosphere reserve with description of four new species from Sweden. Scientific Reports 11(1):4861. DOI: 10.1038/s41598-021-83627-w.
  • Massa E, Rebecchi L, Guidetti R. 2023. Effects of synthetic acid rain and organic and inorganic acids on survival and CaCO3 piercing stylets in tardigrades. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology 339:578–589. DOI: 10.1002/jez.2701.
  • Massa E, Rebecchi L, Guidetti R. 2024. Composition and structural organization of tardigrades feeding apparatus focusing on chitin and other autofluorescent molecules. Zoological Journal of the Linnean Society 200(1):186–199. DOI: 10.1093/zoolinnean/zlad028.
  • Maucci W, Durante Pasa MV. 1984. Macrobiotus lusitanicus sp. nov. nuova specie di Eutardigrado del Portogallo Nord-Occidentale (Tardigrada, Macrobiotidae). Bollettino del Museo Civico di Storia Naturale di Verona 11:319–326.
  • McGreevy KM, Heikes KL, Kult S, Tharp ME, Goldstein B. 2018. Fluorescent cell staining methods for living Hypsibius exemplaris embryos. Cold Spring Harbor Protocols 11:db–prot106021. DOI: 10.1101/pdb.prot106021.
  • Michalczyk Ł, Kaczmarek Ł. 2013. The Tardigrada Register: a comprehensive online data repository for tardigrade taxonomy. Journal of Limnology 72:175–181. DOI: 10.4081/jlimnol.2013.s1.e22.
  • 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 3154(1):1–20. DOI: 10.11646/zootaxa.3154.1.1.
  • Murray J. 1907. XXV.—Arctic Tardigrada, collected by wm. S. Bruce. Transactions of the Royal Society of Edinburgh 45:669–681. DOI: 10.1017/S0080456800011789.
  • Murray J. 1910. Canadian Tardigrada. In: Shackleton, C.V.O. Report for the scientific investigations. Murray J, editors. British Antarctic Expedition 1907–1909 under the command sir, Vol. I. London: William Heinemann, pp. 158–178. C.V.O. Report for the scientific investigations.
  • Nelson DR, Bartels PJ, Guil N. 2019. Tardigrade ecology. In: Schill R, editor Water bears: The biology of Tardigrades. zoological monographs. Vol. 2. Cham: Springer. pp. 163–210.
  • Pennak RW. 1953. Fresh-water invertebrates of the United States. New York: Ronald Press.
  • Pilato G. 1972. Structure, intraspecific variability and systematic value of the buccal armature of eutardigrades. Journal of Zoological Systematics & Evolutionary Research 10(1):65–78. DOI: 10.1111/j.1439-0469.1972.tb00785.x.
  • Pilato G, Kiosya Y, Lisi O, Sabella G. 2012. New records of Eutardigrada from Belarus with the description of three new species. Zootaxa 3179(1):39–60. DOI: 10.11646/zootaxa.3179.1.2.
  • Pleijel F, Jondelius U, Norlinder E, Nygren A, Oxelman B, Schander C, Sundberg P, Thollesson M. 2008. Phylogenies without roots? A plea for the use of vouchers in molecular phylogenetic studies. Molecular Phylogenetics and Evolution 48(1):369–371. DOI: 10.1016/j.ympev.2008.03.024.
  • Prendini L, Weygoldt P, Wheeler WC. 2005. Systematics of the Damon variegatus group of African whip spiders (Chelicerata: Amblypygi): Evidence from behaviour, morphology and DNA. Organisms Diversity & Evolution 5:203–236. DOI: 10.1016/j.ode.2004.12.004.
  • Puillandre N, Brouillet S, Achaz G. 2021. ASAP: Assemble species by automatic partitioning. Molecular Ecology Resources 21(2):609–620. DOI: 10.1111/1755-0998.13281.
  • R Core Team. 2023. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. Available: https://www.R-project.org/.
  • Ramazzotti G. 1956. Tre nuove specie di Tardigradi ed altre specie poco comuni. Atti della Società Italiana di Scienze Naturali 95:284–291.
  • Rebecchi L, Bertolani R. 1994. Maturative pattern of ovary and testis in eutardigrades of freshwater and terrestrial habitats. Invertebrate Reproduction & Development 26:107–117. DOI: 10.1080/07924259.1994.9672407.
  • Rebecchi L, Guidi A. 1991. First SEM studies on tardigrade spermatozoa. Invertebrate Reproduction & Development 19:151–156. DOI: 10.1080/07924259.1991.9672169.
  • Santos AM, Cabezas MP, Tavares AI, Xavier R, Branco M. 2016. tcsBU: A tool to extend TCS network layout and visualization. Bioinformatics 32(4):627–628. DOI: 10.1093/bioinformatics/btv636.
  • Schindelin J, Arganda-Carreras I, Frise E, Kaynig V, Longair M, Pietzsch T, Preibisch S, Rueden C, Saalfeld S, Schmid B, Tinevez JY, White DJ, Hartenstein V, Eliceiri K, Tomancak P, Cardona A. 2012. Fiji: An open-source platform for biological-image analysis. Nature Methods 9(7):676–682. DOI: 10.1038/nmeth.2019.
  • Schuetz G. 1987. A one year study on the population dynamics of milnesium tardigradum doyère in the lichen Xanthoria parietina (L.). In: Bertolani R, editor Biology of tardigrades: Selected symposia and monographs, UZI. Modena: Mucchi. pp. 217–228.
  • Schultze CAS. 1834. Macrobiotus hufelandii animal e Crustaceorum Classe Novum, Reviviscendi Post diuturnam asphixiam et Aridiatem Potens. Berlin: Apud Carolum Curths.
  • Schuster RO, Nelson DR, Grigarick AA, Christenberry D. 1980. Systematic criteria of the Eutardigrada. Transactions of the American Microscopical Society 1980:284–303. DOI: 10.2307/3226004.
  • Stamatakis A. 2014. RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30(9):1312–1313. DOI: 10.1093/bioinformatics/btu033.
  • Stamatakis A, Hoover P, Rougemont J, Renner S. 2008. A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57(5):758–771. DOI: 10.1080/10635150802429642.
  • Stec D, Kristensen RM, Michalczyk Ł. 2020a. An integrative description of Minibiotus ioculator sp. nov. From the Republic of South Africa with notes on Minibiotus pentannulatus Londoño et al., 2017 (Tardigrada: Macrobiotidae). Zoologischer Anzeiger 286:117–134. DOI: 10.1016/j.jcz.2020.03.007.
  • Stec D, Krzywański Ł, Zawierucha K, Michalczyk Ł. 2020b. 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. DOI: 10.1093/zoolinnean/zlz163.
  • Stec D, Morek W, Gąsiorek P, Kaczmarek Ł, Michalczyk Ł. 2016. Determinants and taxonomic consequences of extreme egg shell variability in Ramazzottius subanomalus (Biserov, 1985) (Tardigrada). Zootaxa 4208:zootaxa–4208. DOI: 10.11646/zootaxa.4208.2.5.
  • 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 & Biodiversity 16(4):357–376. DOI: 10.1080/14772000.2018.1424267.
  • Stec D, Roszkowska M, Kaczmarek Ł, Michalczyk Ł. 2018. Paramacrobiotus lachowskae, a new species of Tardigrada from Colombia (Eutardigrada: Parachela: Macrobiotidae). New Zealand Journal of Zoology 45(1):43–60. DOI: 10.1080/03014223.2017.1354896.
  • Tamura K, Stecher G, Kumar S, Battistuzzi FU. 2021. MEGA11: Molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution 38(7):3022–3027. DOI: 10.1093/molbev/msab120.
  • Templeton AR, Crandall KA, Sing CF. 1992. A cladistic analysis of phenotypic association with haplotypes inferred from restriction endonuclease mapping and DNA sequence data. III. Cladogram estimation. Genetics 132:619–633. DOI: 10.1093/genetics/132.2.619.
  • Thorne G. 1938. Notes on free-living and plant-parasitic nematodes. Proceedings of the Helminthological Society of Washington 5:64–65.
  • Trygvadottir BV, Kristensen RM. 2011. Eohypsibiidae (Eutardigrada, Tardigrada) from the Faroe Islands with the description of a new genus containing three new species. Zootaxa 2886(1):39–62.
  • Vecchi M, Cesari M, Bertolani R, Jönsson KI, Rebecchi L, Guidetti R. 2016. Corrigendum to: Integrative systematic studies on tardigrades from Antarctica identify new genera and new species within Macrobiotoidea and Echiniscoidea. Invertebrate Systematics 30:521–521. DOI: 10.1071/IS15033_CO.
  • Vecchi M, Tsvetkova A, Stec D, Ferrari C, Calhim S, Tumanov D. 2023. Expanding Acutuncus: Phylogenetics and morphological analyses reveal a considerably wider distribution for this tardigrade genus. Molecular Phylogenetics and Evolution 180:107707. DOI: 10.1016/j.ympev.2023.107707.
  • Wełnicz W, Grohme MA, Kaczmarek Ł, Schill RO, Frohme M. 2011. ITS–2 and 18S rRNA data from Macrobiotus polonicus and Milnesium tardigradum (Eutardigrada, Tardigrada). Journal of Zoological Systematics & Evolutionary Research 49:34–39. DOI: 10.1111/j.1439-0469.2010.00595.x.
  • White TJ, Bruns T, Lee S, Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis M, Gelfand D, Sninsky J, White T, editors PCR protocols: A guide to methods and applications. Orlando: Academic. pp. 315–322.
  • Zawierucha K, Kolicka M, Kaczmarek Ł. 2016. Re-description of the Arctic tardigrade Tenuibiotus voronkovi (Tumanov, 2007) (Eutardigrada; Macrobiotidea), with the first molecular data for the genus. Zootaxa 4196:zootaxa–4196. DOI: 10.11646/zootaxa.4196.4.2.