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Original Articles

A trans-isthmus survey of marine tardigrades from Costa Rica (Central America) with descriptions of seven new species

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Pages 120-166 | Received 13 Oct 2020, Accepted 05 Mar 2021, Published online: 04 May 2021

References

  • Accogli G, Gallo M, D’Addabbo R, Hansen JG. 2011. Diversity and ecology of the marine tardigrades along the Apulian Coast. Journal of Zoological Systematics and Evolutionary Research. 49(Suppl. 1):53–57. doi:10.1111/j.1439-0469.2010.00598.x.
  • Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ. 1990. Basic local alignment search tool. Journal of Molecular Biology. 215:403–410. doi:10.1016/S0022-2836(05)80360-2.
  • Ataide MB, Venekey V, Filho JSR, dos Santos PJP. 2014. Sandy reefs of Sabellaria wilsoni (Polychaeta: Sabellariidae) as ecosystem engineers for meiofauna in the Amazon coastal region, Brazil. Marine Biodiversity. 44(3):403–413. doi:10.1007/s12526-014-0248-x.
  • Bartels PJ, Apodaca JJ, Mora C, Nelson DR. 2016. A global biodiversity estimate of a little known taxon: Phylum Tardigrada. Zoological Journal of the Linnean Society. 178:730–736. doi:10.1111/zoj.12441.
  • Bartels PJ, Fontaneto D, Roszkowska M, Nelson DR, Kaczmarek Ł. 2020. Latitudinal gradients in body size in marine tardigrades. Zoological Journal of the Linnean Society. 188(3):820–838. doi:10.1093/zoolinnean/zlz080.
  • Bartels PJ, Fontoura P, Nelson DR. 2015b. New records of marine tardigrades from Moorea, French Polynesia, with the description of Styraconyx turbinarium sp. nov. (Arthrotardigrada, Halechniscidae). Zootaxa. 3955(3):389–402. doi:10.11646/zootaxa.3955.3.6.
  • Bartels PJ, Fontoura P, Nelson DR. 2018. Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus. Zootaxa. 4420:43–70. doi:10.11646/zootaxa.4420.1.3.
  • Bartels PJ, Kaczmarek Ł, Roszkowska M, Nelson DR. 2015a. Interactive map of marine tardigrades of the world. https://paul-bartels.shinyapps.io/marine-tardigrades/. Accessed 3-Oct-20.
  • Bartylak T, Kulpa AD, Grobys D, Kepel MA, Kepel A, Kmita HA, Gawlak M, Grabiński WO, Roszkowska MI, Kaczmarek ŁU. 2019. Variability of Echiniscus tristis Gąsiorek & Kristensen. 2018. Is morphology sufficient for taxonomic differentiation of Echiniscidae? Zootaxa. 4701(1):1–24. doi:10.11646/zootaxa.4701.1.1.
  • Binda MG. 1978. Risistemazione di alcuni tardigradi con l’istituzione di un nuovo genere di Oreellidae e della nuova famiglia Archechiniscidae. Animalia. 5:307–314. doi:10.5962/bhl.title.63039.
  • Biserov VI. 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. [Google Scholar].
  • Casquet J, Thebaud C, Gillespie RG. 2012. Chelex without boiling, a rapid and easy technique to obtain stable amplifiable DNA from small amounts of ethanol-stored spiders. Molecular Ecology Resources. 12:136–141. doi:10.1111/j.1755-0998.2011.03073.x.
  • Chang CY, Rho HS. 1998a. Three new tardigrade species associated with barnacles from the Thai coast of Andaman Sea. Korean Journal of Biological Sciences. 2:323–331. doi:10.1080/12265071.1998.9647426.
  • Cornils A, Held C. 2014. Evidence of cryptic and pseudocryptic speciation in the Paracalanus parvus species complex (Crustacea, Copepoda, Calanoida). Frontiers in Zoology. 11:19. doi:10.1186/1742-9994-11-19.
  • Cortés J. 2016a. The Caribbean coastal and marine ecosystems. In: Kappelle M, editor. Costa Rican ecosystems. Chicago: University of Chicago Press; p. 591–617. doi:10.7208/chicago/9780226121642.003.0017.
  • Cortés J. 2016b. The Pacific coastal and marine ecosystems. In: Kappelle M, editor. Costa Rican ecosystems. Chicago: University of Chicago Press; p. 97–138. doi:10.7208/chicago/9780226121642.003.0005.
  • Curini-Galletti M, Stocchino GA, Norenburg JL. 2019. New species of Duploinona Karling, 1966 and Pseudominona Karling, 1978 (Platyhelminthes: Proseriata) from the Caribbean. Zootaxa. 4657(1):127–147. doi:10.11646/zootaxa.4657.1.5.
  • D'Addabbo Gallo M, Grimaldi de Zio S, Morone De Lucia MR. 1986. Description of two species of Halechiniscidae belonging to the sub-families Halechiniscinae and Styraconyxinae (Tardigrada: Heterotardigrada). Oebalia. 13:69–75. doi:10.1016/0198-0254(87)90958-7.
  • D'Addabbo Gallo M, Grimaldi de Zio S, Morone De Lucia MR, Troccoli A. 1992. Halechiniscidae and Echiniscoididae from Western Mediterranean Sea (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine. 33:299–318. [Google Scholar].
  • D'Addabbo Gallo M, Morone De Lucia MR, de Zio Grimaldi S. 1989. Two new species of the genus Styraconyx (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine. 30:17–33. [Google Scholar].
  • D'Addabbo Gallo M, Morone De Lucia MR, Grimaldi de Zio S. 1987. Heterotardigrada of the Amendolara Shoal, High Ionian Sea. In: Bertolani R, editor. Biology of Tardigrades. Vol. 1. Modena, Italy: Mucchi. Selected Symposia and Monographs U.Z. I.; p. 93–101. [Google Scholar].
  • D’Addabbo Gallo M, Grimaldi de Zio S, Morone De Lucia MR. 1984. Styraconyx testudo n. sp., a new Styraconyxinae of the Mediterranean Sea (Arthrotardigrada: Halechiniscidae). Oebalia. 10:95–103. [Google Scholar].
  • Danovaro R, Scopa M, Gambi C, Frascheti S. 2007. Trophic importance of subtidal metazoan meiofauna: evidence from in situ exclusion experiments on soft and rocky substrates. Marine Biology. 152:339–35. doi:10.1007/s00227-007-0696-y.
  • Darriba D, Taboada GL, Doallo R, Posada D. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods. 9:772. doi:10.1038/nmeth.2109.
  • de Zio Grimaldi S, D`Addabbo Gallo M, Pietanza R. 2000. Two new sub-Antarctic Echiniscoididae from Marion Island (Heterotardigrada, Echiniscoidea). Italian Journal of Zoology. 67:221–228. doi:10.1080/11250000009356315.
  • Degma P, Bertolani R, Guidetti R. 2009–2020. Actual checklist of Tardigrada species (2009–2020, 38th Edition:18-08-2020). http://dx.doi.org/10.25431/11380_1178608.
  • Degma P, Guidetti R. 2007. Notes to the current checklist of Tardigrada. Zootaxa. 1579:41–53. doi:10.11646/zootaxa.1579.1.2.
  • Derycke S, De Ley P, Tandingan De Ley I, Holovachov O, Rigaux A, Moens T. 2010. Linking DNA sequences to morphology: cryptic diversity and population genetic structure in the marine nematode Thoracostoma trachygaster (Nematoda, Leptosomatidae). Zoologica Scripta. 39(3):276–289. doi:10.1111/j.1463-6409.2009.00420.x.
  • Doyère LM. 1840. Mémoire sur les Tardigrades. Annales des Sciences Naturelles (Paris), Sér. 2. 14:269–361. [Google Scholar].
  • du Bois-Reymond Marcus E. 1952. On South American Malacopoda. Boletins da Faculdade de Philosophia, Sciencias e Letras, Universidade de São Paulo. Zoologia. 17:189–209. doi:10.11606/issn.2526-4877.bsffclzoologia.1952.125190.
  • Egger C, Neusser TP, Norenburg J, Leasi F, Buge B, Vannozzi A, Cunha RL, Cox CJ, Jörger KM. 2020. Uncovering the shell game with barcodes: diversity of meiofaunal Caecidae snails (Truncatelloidea, Caenogastropoda) from Central America. ZooKeys. 968:1–42. doi:10.3897/zookeys.968.52986.
  • Faurby S, Barber PH. 2015. Extreme population subdivision despite high colonization ability: Contrasting regional patterns in intertidal tardigrades from the west coast of North America. Journal of Biogeography. 42:1006–1017. doi:10.1111/jbi.12500.
  • Faurby S, Jørgensen A, Kristensen RM, Funch P. 2011. Phylogeography of North Atlantic intertidal tardigrades: refugia, cryptic speciation and the history of the Mid-Atlantic Islands. Journal of Biogeography. 38:1613–1624. doi:10.1111/j.1365-2699.2011.02533.x.
  • Faurby S, Jørgensen A, Kristensen RM, Funch P. 2012. Distribution and speciation in marine intertidal tardigrades: testing the roles of climatic and geographical isolation. Journal of Biogeography. 39(9):1596–1607. doi:10.1111/j.1365-2699.2012.02720.x.
  • 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. [Google Scholar].
  • Fontoura P, Bartels PJ, Jørgensen A, Kristensen RM, Hansen JG. 2017. An illustrated dichotomous key to the genera of marine heterotardigrades (Tardigrada). Zootaxa. 4294:1–45. doi:10.11646/zootaxa.4294.1.1.
  • Fujimoto S, Jørgensen A, Hansen JG. 2016. A molecular approach to arthrotardigrade phylogeny (Heterotardigrada, Tardigrada). ZoolScr. 46(4):496–505. doi:10.1111/zsc.12221.
  • Fujimoto S, Miyazaki K, Suzuki AC. 2013. A new marine tardigrade, Tanarctus diplocerus (Arthrotardigrada: Halechiniscidae) from Japan. Journal of the Marine Biological Association UK. 93(4):955–961. doi:10.1017/S0025315412000689.
  • Fujimoto S, Suzuki AC, Ito M, Tamura T, Tsujimoto M. 2020. Marine tardigrades from Lutzow-Holm Bay, East Antarctica with the description of a new species. Polar Biology. 43:679–693. doi:10.1007/s00300-020-02671-w.
  • Gallo D’Addabbo M, D’Addabbo R, de Zio Grimaldi S. 2000. Redescription of Batillipes dicrocercus Pollock 1970 and revision of the genus Batillipes (Tardigrada, Heterotardigrada). Zoologischer Anzeiger. 239:329–330. [Google Scholar].
  • Gallo D’Addabbo M, Sandulli R, de Zio Grimaldi S. 2005. New Batillipedidae (Tardigrada, Heterotardigrada) from the Orosei Gulf, Sardinia, Tyrrhenian Sea. Zoologischer Anzeiger. 243:219–225. doi:10.1016/j.jcz.2004.07.004.
  • Gallo M, D’Addabbo R, De Leonardis C, Sandulli R, de Zio Grimaldi S. 2007. The diversity of Indian Ocean Heterotardigrada. Journal of Limnology. 66(1s):60–64. doi:10.4081/jlimnol.2007.s1.60.
  • Garraffoni AR, Balsamo M. 2017. Is the ubiquitous distribution real for marine gastrotrichs? Detection of areas of endemism using parsimony analysis of endemicity (PAE). Proceedings of the Biological Society of Washington. 130:197–210.
  • Gąsiorek P, Kristensen RM. 2018. Echiniscidae (Heterotardigrada) of Tanzania and Uganda. Tropical Zoology. 31(3):131–160. doi:10.1080/03946975.2018.1477350.
  • Giere O. 2009. Meiobenthology, 2nd ed. Berlin: Springer-Verlag. doi:10.1007/978-3-662-02912-1.
  • Goujon M, McWilliam H, Li W, Valentin F, Squizzato S, Paern J, Lopez R. 2010. A new bioinformatics analysis tools framework at EMBL–EBI. Nucleic Acids Research. 38:W695–W699. doi:10.1093/nar/gkq313.
  • Grimaldi de Zio S, D`Addabbo Gallo M. 1975. Post-embryonic development and moults in Batillipes pennaki Marcus (Heterotardigrada). Rivista di Biologia. 68:243–274. [PubMed].
  • Grimaldi de Zio S, D'Addabbo Gallo M. 1987. Archechiniscus minutus n. sp. and its systematic position within Arthrotardigrada (Tardigrada: Heterotardigrada). In: Bertolani R, editor. Biology of Tardigrades. Modena, Italy: Mucchi. Selected Symposia and Monographs U.Z.I.; p. 253–260. [Google Scholar].
  • Grimaldi de Zio S, Gallo D’Addabbo M. 2001. Further data on the Mediterranean Sea tardigrade fauna. Zoologischer Anzeiger. 240(3):345–360. doi:10.1078/0044-5231-00042.
  • Grimaldi de Zio S, Gallo D’Addabbo M, Sandulli R, D’Addabbo R. 2003. Checklist of the Italian marine Tardigrada. Meiofauna Marina. 12:97–135. [Google Scholar].
  • Grimaldi de Zio S, Morone De Lucia MR, D'Addabbo Gallo M. 1983. Marine tardigrades of the ‘Secca dell’Ar meleia’ (Ionian Sea), and redescription of Raiarctus colurus Renaud-Mornant 1981 (Heterotardigrada). Oebalia. 9:33–42. [Google Scholar].
  • Grimaldi de Zio S, D’Addabbo Gallo M, Morone DeLucia RM. 1990. Revision of the genus Halechiniscus (Halechiniscidae, Arthrotardigrada). Cahiers de Biologie Marine. 31:271–279. [Google Scholar].
  • 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. doi:10.1093/zoolinnean/zlz171.
  • Gross V, Miller WR, Hochberg R. 2014. A new tardigrade, Mutaparadoxipus duodigifinis gen. nov., sp. nov. (Heterotardigrada: Arthrotardigrada), from the Southeastern United States. Zootaxa. 3835:263–272. doi:10.11646/zootaxa.3835.2.6.
  • Guidetti R, Bertolani R. 2005. Tardigrade taxonomy: an updated check list of the taxa and a list of characters for their identification. Zootaxa. 845:1–46. doi:10.11646/zootaxa.845.1.1.
  • 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. doi:10.1186/s40851-018-0113-z.
  • Guil N, Giribet G. 2009. Fine scale population structure in the Echiniscus blumi-canadensis series (Heterotardigrada, Tardigrada) in an Iberian mountain range—When morphology fails to explain genetic structure. Molecular Phylogenetics and Evolution. 51(3):606–613. doi:10.1016/j.ympev.2009.02.019.
  • Guzmán HM, Obando V, Cortés J. 1987. Meiofauna associated with a Pacific coral reef in Costa Rica. Coral Reefs. 6:107–112. doi:10.1007/BF00301379.
  • Hall TA. 1999. BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series. 41:95–98. [Google Scholar].
  • Hallas T, Kristensen RM. 1982. Two new species of the tidal genus Echiniscoides from Rhode Island, U.S.A. (Echiniscoididae, Heterotardigrada). In: Nelson D, editor. Proceedings of the Third International Symposium on Tardigrada. Johnson City, TN: East Tennessee State University Press; p. 179–192. [Google Scholar].
  • Hansen JG, Jørgensen A, Kristensen RM. 2001. Preliminary studies of the tardigrade fauna of the Faroe Bank. Zoologischer Anzeiger. 240:385–393. doi:10.1078/0044-5231-00046.
  • Hansen JG, Kristensen RM, Jørgensen A. 2012. The armoured marine tardigrades (Arthrotardigrada, Tardigrada). The Royal Danish Academy of Sciences and Letters. Scientia Danica, Series B, Biologica. 2:1–91. [Google Scholar].
  • Harris RP. 1972. Seasonal changes in the meiofauna population of an intertidal sand beach. Journal of the Marine Biological Association UK. 52:389–403. doi:10.1017/S0025315400018750.
  • Hartman O. 1944. Polychaetous annelids. Part VI. Paraonidae, Magelonidae, Longosomidae, Ctenodrilidae, and Sabellariidae. Allan Hancock Pacific Expeditions. 10(3):311–389. [Google Scholar].
  • Hygum TL, Clausen L, Halberg KA, Jørgensen A, Møbjerg N. 2016. Tun formation is not a prerequisite for desiccation tolerance in the marine tidal tardigrade Echiniscoides. Zoological Journal of the Linnean Society. 178:907–911. doi:10.1111/zoj.12444.
  • Jørgensen A, Boesgaard TM, Møbjerg N, Kristensen RM. 2014. The tardigrade fauna of Australian marine caves: With descriptions of nine new species of Arthrotardigrada. Zootaxa. 3802(4):401–443. doi:10.11646/zootaxa.3802.4.1.
  • Jørgensen A, Faurby S, Hansen JG, Møbjerg N, Kristensen RM. 2010. Molecular phylogeny of Arthrotardigrada (Tardigrada). Molecular Phylogenetics and Evolution. 54:1006–1015. doi:10.1016/j.ympev.2009.10.006.
  • Jørgensen A, Møbjerg N. 2015. Notes on the cryptobiotic capability of the marine arthrotardigrades Styraconyx haploceros (Halechiniscidae) and Batillipes pennaki (Batillipedidae) from the tidal zone in Roscoff, France. Marine Biology Research. 11(2):214–217. doi:10.1080/17451000.2014.904883.
  • Jørgensen A, Møbjerg N, Kristensen RM. 2007. A molecular study of the tardigrade Echiniscus testudo (Echiniscidae) reveals low DNA sequence diversity over a large geographical area. The Journal of Limnology. 66(Suppl. 1):77–83. doi:10.4081/jlimnol.2007.s1.77.
  • Kaczmarek Ł, Roszkowska M 2016. A new eutardigrade from Costa Rica with taxonomical and zoogeographical remarks on Costa Rican tardigrades. New Zealand Journal of Zoology. 43(3):234–245. doi:10.1080/03014223.2016.1156553.
  • Kaczmarek Ł, Bartels P, Roszkowska M. 2018a. First records of marine tardigrades (Arthrotardigrada) from Fuerteventura (Canary Islands, Spain). African Journal of Zoology. 53(2):75–81. doi:10.1080/15627020.2018.1493705.
  • Kaczmarek Ł, Bartels PJ, Roszkowska M, Nelson DR. 2015. The zoogeography of marine Tardigrada. Zootaxa. 4037:1–189. doi:10.11646/zootaxa.4037.1.1.
  • Kaczmarek Ł, Gołdyn B, McInnes SJ, Michalczyk Ł. 2016. Diversity of limno-terrestrial tardigrades of the Americas in relation to the Great American Biotic Interchange hypothesis (GABI). Zoological Journal of the Linnean Society. 178(4):737–746. doi:10.1111/zoj.12422.
  • Kaczmarek Ł, Grobys D, Kulpa A, Bartylak T, Kmita H, Kepel M, Kepel A, Roszkowska M. 2019. Two new species of the genus Milnesium Doyère, 1840 (Tardigrada, Apochela, Milnesiidae) from Madagascar. ZooKeys. 884:1–22. doi:10.3897/zookeys.884.29469.
  • Kaczmarek Ł, Parnikoza I, Gawlak M, Esefeld J, Peter H-U, Kozeretska I, Roszkowska M. 2018b. Tardigrades from Larus dominicanus Lichtenstein, 1823 nests on the Argentine Islands (maritime Antarctic). Polar Biology. 41(2):283–301. doi:10.1007/s00300-017-2190-4.
  • Katoh K, Misawa K, Kuma KI, Miyata T. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research. 30:3059–3066. doi:10.1093/nar/gkf436.
  • Katoh K, Toh H. 2008. Recent developments in the MAFFT multiple sequence alignment program. Brief Bioinform. 9:286–298. doi:10.1126/science.8503007.
  • Knowlton N, Weight LA, Solórzano LA, Mills DK, Bermingham E. 1993. Divergence in proteins, mitochondrial DNA, and reproductive compatibility across the isthmus of Panama. Science. 260:1629–1772.
  • Kristensen RM. 1978. Notes on marine heterotardigrades. 1. Description of two new Batillipes species, using the electron microscope. Zoologischer Anzeiger. 200:1–17. [Google Scholar].
  • Kristensen RM. 1984. On the biology of Wingstrandarctus corallinus nov. gen. et spec., with notes on the symbiotic bacteria in the subfamily Florarctinae (Arthrotardigrada). Videnskabelige Meddelelser Naturhistorisk Forening. 145:201–218. [Google scholar].
  • Kristensen RM, Hallas TE. 1980. The tidal genus Echiniscoides and its variability, with erection of Echiniscoididae fam. n. (Tardigrada). Zoologica Scripta. 9:113–127. doi:10.1111/j.1463-6409.1980.tb00657.x.
  • Kristensen RM, Higgins RP. 1984a. A new family of Arthrotardigrada (Tardigrada: Heterotardigrada) from the Atlantic Coast of Florida, U.S.A. Transactions of the American Microscopical Society . 103:295–311. doi:10.2307/3226191.
  • Kristensen RM, Higgins RP. 1984b. Revision of Styraconyx (Tardigrada: Halechiniscidae) with description of two new species from Disko Bay, West Greenland. Smithsonian Contributions to Zoology. 391:1–40. doi:10.5479/si.00810282.391.
  • Kristensen RM, Mackness BS. 2000. First record of the marine tardigrade genus Batillipes (Arthrotardigrada: Batillipedidae) from South Australia with a description of a new species. Records of the South Australian Museum. 33:73–87. [Google Scholar].
  • Kristensen RM, Persson DK, Møbjerg N, Jørgensen A. 2012. The marine tardigrades of Al Zubarah. In: Bangsgaard P, Kristensen RM, Møller PR, Nymann H, editor. Qatar Islamic archaeology and heritage project: end of season report environmental studies 2011–2012. Copenhagen: University of Copenhagen; p. 13–24.
  • Kristensen RM, Renaud-Mornant J. 1983. Existence d'Arthrotardigrades semi-benthiques de genres nouveaux de la sous-famille des Styraconyxinae subfam. nov.. Cahiers de Biologie Marine. 24:337–353. [Google Scholar].
  • Kristensen RM, Sørensen MV, Hansen JG, Zeppilli D. 2015. A new species of Neostygarctus (Arthrotardigrada) from the Condor Seamount in the Azores, Northeast Atlantic. Marine Biodiversity. 45(3):453–467. doi:10.1007/s12526-015-0323-y.
  • Kumar S, Stecher G, Li M, Knyaz C, Tamura K. 2018. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution. 35(6):1547–1549. doi:10.1093/molbev/msy096.
  • Leasi F, Andrade SdS, Norenburg J. 2016. At least some meiofaunal species are not everywhere. Indication of geographic, ecological and geological barriers affecting the dispersion of species of Ototyphlonemertes (Nemertea, Hoplonemertea). Molecular Ecology. 25(6):1381–1397. doi:10.1111/mec.13568.
  • Leasi F, Norenburg JL. 2014. The necessity of DNA taxonomy to reveal cryptic diversity and spatial distribution of meiofauna, with a focus on Nemertea. PLoS ONE. 9(8):e104385. doi:10.1371/journal.pone.0104385.
  • Lee J, Hyun Soo Rho HS, Cheon Young Chang CY. 2017. Taxonomic study of marine tardigrades from Korea III. A new species of the genus Orzeliscus (Heterotardigrada, Halechiniscidae). Animal Systematics, Evolution and Diversity. 33(1):26–32. doi:10.5635/ASED.2017.33.1.056.
  • Lessios H. 2008. The great American schism: divergence of marine organisms after the rise of the Central American Isthmus. Annual Review of Ecology, Evolution, and Systematics. 39:63–91. doi:10.1146/annurev.ecolsys.38.091206.095815.
  • Librado P, Rozas J. 2009. DnaSP v5: A software for comprehensive analysis of DNA polymorphism data. Bioinformatics. 25:1451–1452. doi:10.1093/bioinformatics/btp187.
  • Marcus E. 1927. Zur Anatomie und Ökologie mariner Tardigraden. Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere. 53:487–588. [Google Scholar].
  • McGinty M, Higgins R. 1968. Ontogenetic variation of taxonomic characters of two marine tardigrades with the description of Batillipes bullacaudatus n. sp. Transactions of the American Microscopical Society. 87:252–262. doi:10.2307/3224449.
  • McKirdy DJ. 1975. Batillipes (Heterotardigrada): Comparison of six species from Florida (USA) and a discussion of taxonomic characters within the genus. Memorie dell'Istituto Italiano di Idrobiologia. 32(Supp l):177–223. [Google Scholar].
  • Meyer HA. 2015. Water bears (Phylum Tardigrada) of Oceania, with the description of a new species of Milnesium. New Zealand Journal of Zoology. 42(3):173–186. doi:10.1080/03014223.2015.1062402.
  • Meyer HA, Hinton JG. 2012. Terrestrial Tardigrada of the Island of Barbados in the West Indies, with the description of Milnesium barbadosense sp. n. (Eutardigrada: Apochela: Milnesiidae). Caribbean Journal of Science. 46(2-3):194–202. doi:10.18475/cjos.v46i2.a8.
  • Miller WR, Clark T, Miller C. 2012. Tardigrades of North America: Archechiniscus biscaynei, sp. nov. (Arthrotardigrada: Archechiniscidae), a marine tardigrade from Biscayne National Park, Florida. Southeastern Naturalist. 11:279–276. doi:10.1656/058.011.0209.
  • Miller WR, Johansson C, Adams B. 2014. Tardigrades of North America: records for five marine species from the Pacific Coast. Transactions of the Kansas Academy of Science. 117(3):281–286. doi:10.1660/062.117.0315.
  • Møbjerg N, Halberg KA, Jørgensen A, Persson D, Bjørn M, Ramløv H, Kristensen RM. 2011. Survival in extreme environments – on the current knowledge of adaptations in tardigrades: Adaptation to extreme environments in tardigrades. Acta Physiologica. 202(3):409–420. doi:10.1111/j.1748-1716.2011.02252.x.
  • Møbjerg N, Jørgensen A, Kristensen RM. 2019. Ongoing revision of Echiniscoididae (Heterotardigrada: Echiniscoidea), with the description of a new interstitial species and genus with unique anal structures. Zoological Journal of the Linnean Society. 188(2):663–680. doi:10.1093/zoolinnean/zlz122.
  • Møbjerg N, Kristensen RM, Jørgensen A. 2016. Data from new taxa infer Isoechiniscoides gen. nov. and increase the phylogenetic and evolutionary understanding of echiniscoidid tardigrades (Echiniscoidea: Tardigrada). Zoological Journal of the Linnean Society. 178:804–818. doi:10.1111/zoj.12500.
  • Mogle MJ, Kimball SA, Miller WR, McKown RD. 2018. Evidence of avian-mediated long distance dispersal in American tardigrades. PeerJ. 6:e5035. doi:10.7717/peerj.5035.
  • Montes C, Cardona A, Jaramillo C, Pardo A, Silva JC, Valencia V. 2015. Middle Miocene closure of the Central American Seaway. Science. 348(6231):226–229. doi:10.1126/science.aaa2815.
  • Morek W, Ciosek JA, 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. 288:107–117. doi: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. doi:10.1163/18759866-08502003.
  • Moreno-Talamantes A, Roszkowska M, García-Aranda MA, Flores-Maldonado JJ, Kaczmarek Ł. 2019. Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840. Zootaxa. 4691(5):501–524. doi:10.11646/zootaxa.4691.5.5.
  • Morone de Lucia RM, D'Addabbo Gallo M, Grimaldi De Zio S. 1988. Descrizione di due nuove specie di Batillipedidae (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine. 29(3):361–373.
  • Nelson DR, Bartels PJ, Guil N. 2018. Tardigrade ecology. In: Schill RO, editor. Water Bears: The Biology of Tardigrades. Ecol Monogr. 2:163–210. Switzerland: Springer Nature; doi:10.1007/978-3-319-95702-9_7.
  • Pardos F, Herranz M, Sánchez N. 2016. Two sides of a coin: the phylum Kinorhyncha in Panama. II) Pacific Panama. Zoologischer Anzeiger. 265:26–47.
  • Perry E, Miller WR, Kaczmarek Ł. 2019. Recommended abbreviations for the names of genera of the phylum Tardigrada. Zootaxa. 4608(1):145. doi:10.11646/zootaxa.4608.1.8.
  • Perry ES, Miller WR. 2015. Echiniscoides wyethi, a new marine tardigrade from Maine, U.S.A. (Heterotardigrada: Echiniscoidea: Echiniscoididae). Proceedings of the Biological Society of Washington. 128(1):103–110. doi:10.2988/0006-324X-128.1.103.
  • Perry ES, Rawson P, Ameral NJ, Miller WR, Miller JD. 2018. Echiniscoides rugostellatus, a new marine tardigrade from Washington, U.S.A. (Heterotardigrada: Echiniscoidea: Echiniscoididae: Echiniscoidinae). Proceedings of the Biological Society of Washington. 131:182–193. doi:10.2988/18-00004.
  • Pérez-Pech WA, De Jesús-Navarrate A, Demilio E, Anguas-Escalante A, Hansen JG. 2020. Marine Tardigrada from the Mexican Caribbean with the description of Styraconyx robertoi sp. nov. (Arthrotardigrada: Styraconyxidae). Zootaxa. 4731(4):492–508. doi:10.11646/zootaxa.4731.4.3.
  • Plate L. 1888. Beiträge zur Naturgeschichte der Tardigraden. Zoologische Jahrbücher. Abteilung für Anatomie und Ontogenie der Tiere. 3:487–550. doi:10.5962/bhl.part.1265.
  • Pollock LW. 1970. Batillipes dicrocercus n. sp., Stygarctus granulatus n. sp. and other Tardigrada from Woods Hole, Massachusetts, USA. Transactions of the American Microscopical Society. 89:38–52. doi:10.2307/3224613.
  • Pollock LW. 1971. On some British marine Tardigrada including two new species of Batillipes. Journal of the Marine Biological Association UK. 51:93–103. doi:10.1017/S0025315400006482.
  • Pollock LW. 1975. Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean. Cahiers de Biologie Marine. 16:121–132. [Google Scholar].
  • Pollock LW. 1983. A closer look at some marine Heterotardigrada. II: The morphology and taxonomy of Bathyechiniscus, with a description of B. craticulus n. sp. from the Caribbean. Bulletin of Marine Science. 33:109–117. [Google Scholar].
  • Posada D, Buckley TR. 2004. Model selection and model averaging in phylogenetics: advantages of the AIC and Bayesian approaches over likelihood ratio tests. Systematic Biology. 535:793–808. doi:10.1080/10635150490522304.
  • Ramazzotti G. 1962. Il Phylum Tardigrada. Memorie dell'Istituto Italiano di Idrobiologia. 16 : 1–595.
  • Renaud-Debyser J. 1959a. Etudes sur la faune interstitielle des Iles Bahamas. III. Tardigrades. Vie Milieu. 10:297–302. [Google Scholar].
  • Renaud-Debyser J. 1959b. Sur quelques Tardigrades du Bassin d'Arcachon. Vie Milieu. 10:135–146. [Google Scholar].
  • Renaud-Debyser J. 1965. Parastygarctus higginsi n. g., n. sp., Tardigrade marin Interstitiel de Madagascar. Comptes Rendus de l'Académie des Sciences. 260:955–957. [Google Scholar].
  • Renaud-Mornant J. 1970. Parastygarctus sterreri n. sp., Tardigrade marin nouveau de l'Adriatique. Cahiers de Biologie Marine. 11:355–360.
  • Renaud-Mornant J. 1975. Occurrence of the genus Tanarctus Renaud-Debyser, 1959 in Northeastern Atlantic waters with a description of T. ramazzotti n. sp. (Arthrotardigrada). Memorie dell'Istituto Italiano di Idrobiologia. 32(Suppl.):325–332. [Google Scholar].
  • Renaud-Mornant J. 1976. Tardigrades marins de Polynesie. Cahiers du Pacifique sud contemporain. 19:289–297. [Google Scholar].
  • Renaud-Mornant J. 1979. Tardigrades marins de Madagascar. II. Stygarctidae et Orellidae, III. Considérations écologiques générales. Bulletin du Muséum National d'Histoire Naturelle Paris, Sér 4e. 1(2):339–351. [Google Scholar].
  • Renaud-Mornant J. 1980. Description de trois espécies nouvelles du genre Tanarctus Renaud-Debyser, 1959, et création de la sous-famille des Tanarctinae subfam. nov. (Tardigrada, Heterotardigrada). Bulletin du Muséum National d'Histoire Naturelle Paris, Sér 4e. 2:129–141. [Google Scholar].
  • Renaud-Mornant J. 1981a. Raiarctus colurus n. g., n. sp., et R. aureolatus n. sp., Tardigrades (Arthrotardigrada) marins nouveaux de sédiments calcaires. Bulletin du Muséum National d'Histoire Naturelle Paris, Sér 4e. 3:512–522. [Google Scholar].
  • Renaud-Mornant J. 1981b. Stygarctus gourbaultae n. sp. un nouveau Tardigrade marin (Arthrotardigrada) de la Guadeloupe. Bulletin du Muséum National d'Histoire Naturelle Paris, Sér 4e. 3(1):175–180. [Google Scholar].
  • Renaud-Mornant J. 1982. Sous-famille et genre nouveaux de Tardigrades marins (Arthrotardigrada). Bulletin du Muséum National d'Histoire Naturelle Paris, Sér 4e. 4(1–2):89–94. [Google Scholar].
  • Renaud-Mornant J. 1984. Nouveaux Arthrotardigrades des Antilles. Bulletin du Muséum National d'Histoire Naturelle Paris, Sér 4e. 6(4):975–988. [Google Scholar].
  • Renaud-Mornant J, Gourbault N. 1981. Premières prospections méiofaunistiques en Guadeloupe I: Les biotopes et leurs peuplements. Bulletin du Muséum National d'Histoire Naturelle, Paris. Sér 3. 4:1011–1034. [Google Schoolar].
  • Renaud-Mornant J, Gourbault N. 1984. Premières prospections méiofaunistiques en Guadeloupe II: Communautés des sables littoraux. Hydrobiologia. 18:113–118. doi:10.1007/BF00031793. [Google Scholar].
  • Renaud-Mornant J, Gourbault N, Helléouet MN. 1983. Premieres prospections faunistiques en Martinique. I. Les biotopes et leurs peuplements. Bulletin du Muséum National d'Histoire Naturelle, 4, Serie 5A. 1:221–234. [Google Scholar].
  • Rice P, Longden I, Bleasby A. 2000. EMBOSS: The European molecular biology open software suite. Trends in Genetics. 16:276–277. doi:10.1016/S0168-9525(00)02024-2. [Google Scholar].
  • Richters F. 1903. Nordische Tardigraden. Zoologischer Anzeiger. 27:168–172. [Google Scholar].
  • Richters F. 1908. Marine Tardigraden. Zoologischer Anzeiger. 33:77–85.
  • Richters F. 1909. Tardigraden-Studien. Bericht der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main. 40:28–48.
  • Richters F. 1926. Tardigrada. In: Kükenthal W, Krumbach T, editor. Handbuch der Zoologie, Vol. 3. Berlin: Walter de Gruyter & Co; p. 58–61. [Google Scholar].
  • Riggin GT. 1962. Tardigrada of Southwest Virginia: with the addition of a new marine tardigrade species from Florida. Virginia Agricultural Experiment Station. 152:1–145. [Google Scholar].
  • Robertson MW, Russo NJ, McInnes SJ, Goffinet B, Jiménez JE. 2020. Potential dispersal of tardigrades by birds through endozoochory: evidence from Sub-Antarctic White-bellied Seedsnipe (Attagis malouinus). Polar Biology. 43:899–902. doi:10.1007/s00300-020-02680-9.
  • Ronquist F, Huelsenbeck JP 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics. 19(12):1572–1574. doi:10.1093/bioinformatics/btg180.
  • 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. doi:10.11646/zootaxa.4763.4.1.
  • Roszkowska M, Stec D, Gawlak M, Kaczmarek Ł. 2018. An integrative description of a new tardigrade species Mesobiotus romani new species (Macrobiotidae: harmsworthi group) from the Ecuadorian Pacific coast. Zootaxa. 4450(5):550–564. doi:10.11646/zootaxa.4450.5.2.
  • Santos É, Rocha CMC, Gomes Jr., E, Fontoura P. 2017. Three new Batillipes species (Arthrotardigrada: Batillipedidae) from the Brazilian coast. Zootaxa. 4243(3):483. doi:10.11646/zootaxa.4243.3.4.
  • Santos É, Rubal M, Veiga P, Bartels PJ, da Rocha CMC, Fontoura P. 2018b. On the distribution of Batillipes tubernatis Pollock, 1971 (Arthrotardigrada: Batillipedidae) in the Atlantic Basin. Marine Biodiversity. 49(2):621–631. doi:10.1007/s12526-017-0834-9.
  • Santos É, Rubal M, Veiga P, da Rocha CMC, Fontoura P. 2018a. Batillipes (Tardigrada, Arthrotardigrada) from the Portuguese coast with the description of two new species and a new dichotomous key for all species. The European Journal of Taxonomy. 425:1–32. doi:10.5852/ejt.2018.425.
  • Santos É, Veiga P, Rubal M, Bartels PJ, da Rocha CMC, Fontoura P. 2019. Batillipes pennaki Marcus, 1946 (Arthrotardigrada: Batillipedidae): deciphering a species complex. Zootaxa. 4648(3):549–567. doi:10.11646/zootaxa.4648.3.9.
  • Scarpa F, Cossu P, Sanna D, Lai T, Norenburg JL, Curini-Galletti M, Casu M. 2015. An 18S and 28S-based clock calibration for marine Proseriata (Platyhelminthes). Journal of Experimental Marine Biology and Ecology. 463:22–31. doi:10.1016/j.jembe.2014.10.020.
  • Schratzberger M, Ingels J. 2017. Meiofauna matters: the roles of meiofauna in benthic ecosystems. Journal of Experimental Marine Biology and Ecology. 502:12–25. doi:10.1016/j.jembe.2017.01.007.
  • Schultze M. 1865. Echiniscus sigismundi, ein Arctiscoide der Nordsee. Archiv für Mikroskopische Anatomie. 1:428–436.
  • Schulz E. 1951. Über Stygarctus bradypus n. g. n. sp., einen Tardigraden aus Küsten grundwasser und seine phylogenetische Bedeutung. Kieler Meeresforschung. 8:86–97. [Google Scholar].
  • Schulz E. 1953. Eine neue Tardigraden-Gattung von der pazifischen Küste. Zoologischer Anzeiger. 151:306–310. [Google Scholar].
  • Schulz E. 1955. Studien an marinen Tardigraden. Kieler Meeresforschung. 11:73–79. [Google Scholar].
  • Schulz E. 1963. Über die Tardigraden. Zoologischer Anzeiger. 171:3–12.
  • Schuster RO, Nelson DR, Grigarick AA, Christenberry D. 1980. Systematic criteria of the Eutardigrada. Transactions of the American Microscopical Society. 99(3):284–303. doi:10.2307/3226004.
  • Stec D, Krzywański Ł, Arakawa K, Michalczyk Ł. 2020. A new redescription of Richtersius coronifer, supported by transcriptome, provides resources for describing concealed species diversity within the monotypic genus Richtersius (Eutardigrada). Zoological Letters. 6:2. doi:10.1186/s40851-020-0154-y.
  • Stehli FG, Webb SD. 1985. The Great American Interchange. New York: Plenum.
  • Surmacz B, Morek W, Michalczyk Ł. 2019. What if multiple claw configurations are present in a sample? A case study with the description of Milnesium pseudotardigradum sp. nov. (Tardigrada) with unique developmental variability. Zoological Studies. 58:32. doi:10.6620/ZS.2019.58-32.
  • Surmacz B, Morek W, Michalczyk Ł. 2020. What to do when ontogenetic tracking is unavailable: a morphometric method to classify instars in Milnesium (Tardigrada). Zoological Journal of the Linnean Society. 188:797–808. doi:10.1093/zoolinnean/zlz099.
  • Thulin G. 1928. Über die Phylogenie und das System der Tardigraden. Hereditas. 11:207–266. doi:10.1111/j.1601-5223.1928.tb02488.x.
  • Thulin G. 1942. Ein neuer mariner Tardigrad. Meddelanden fran Göteborgs Musei Zoologiska Avdelning. 99:1–10. [Google Scholar].
  • Vargas JA. 1988. A survey of the meiofauna of an eastern tropical Pacific intertidal mud flat. Revista de Biología Tropical. 36:541–544. [Google Scholar].

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