Abstract
In two moss samples collected in El Cajas National Park (Azuay Province, Ecuador), 33 specimens and 19 eggs of a new species of Macrobiotus and 56 heterotardigrades belonging to a new Echiniscus species were extracted. Macrobiotus dulciporus sp. nov. belongs to the hufelandi group and is most similar to Macrobiotus maculatus Iharos, 1973. It differs from M. maculatus mainly based on the presence of an oral cavity armature of the patagonicus type, smooth egg chorion surface, lower number of processes on the egg circumference and some morphometric characters of the egg processes. Echiniscus hannae sp. nov. belongs to the bigranulatus group, but differs from all species in this group due to the presence of large, smooth areas on its dorsal plate.
References
Bąkowski, M., Roszkowska, M., Gawlak, M. & Kaczmarek, Ł. (2016) Macrobiotus naskreckii sp. nov. (Eutardigrada: Macrobiotidae: hufelandi group) from Mozambique. Annales Zoologici, 66 (2), 155–164.
https://doi.org/10.3161/00034541ANZ2016.66.2.001
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.
https://doi.org/10.1111/j.1463-6409.1993.tb00347.x
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. & Pilato, G. (1972) Tardigradi muscicoli di Sicilia (IV Nota). Bollettino delle sedute della Accademia Gioenia di Scienze Naturali in Catania, Series 4, 11, 47–60.
Biserov, V.I. (1990) On the revision of the genus Macrobiotus. The subgenus Macrobiotus s. str.: a new systematic status of the group hufelandi (Tardigrada, Macrobiotidae). Communication 1. Zoologicheskii Zhurnal, 69 (12), 5–17.
Biserov, V., Pilato, G. & Lisi, O. (2011) Macrobiotus trunovae sp.n., a new species of tardigrade from Russia. Invertebrate Zoology, 8 (1), 57–62.
https://doi.org/10.15298/invertzool.08.1.07
Cuénot, L. (1932) Tardigrades. In: Lechevalier, P. (Ed.), Faune de France, 24, pp. 1–96.
Dastych, H. (1984). The Tardigrada from Antarctica with description of several new species. Acta Zoologica Cracoviensia, 27, 377–436.
Degma, P. & Guidetti, R. (2007) Notes to the current checklist of Tardigrada. Zootaxa, 1579, 41–53.
Degma, P., Bertolani, R. & Guidetti, R. (2009–2017) Actual checklist of Tardigrada species. Version 33. 15 October 2017. www.tardigrada.modena.unimo.it/miscellanea/Actual%20checklist%20of%20Tardigrada.pdf (accessed 10 December 2018)
Doyère, L.M.F. (1840) Memoire sur les Tardigrades. I. Annales des Sciences Naturelles, Paris, Series 2, 14, 269–362.
Durante Pasa, M.V. & Maucci, W. (1979) Moss Tardigrada from the Scandinavian Peninsula. Zeszyty Naukowe Uniwersytetu Jagielloskiego, Prace Zoologiczme, 79 (25), 47–85.
Fontoura, P. & Morais, P. (2011) Assessment of traditional and geometric morphometrics for discriminating cryptic species of the Pseudechiniscus suillus complex (Tardigrada, Echiniscidae). Journal of Zoological Systematics and Evolutionary Research, 49 (Supplement 1), 26–33.
https://doi.org/10.1111/j.1439-0469.2010.00594.x
Fontoura, P., Rubal, M. & Veiga, P. (2017) Two new species of Tardigrada (Eutardigrada: Ramazzottiidae, Macrobiotidae) from the supralittoral zone of the Atlantic Iberian Peninsula rocky shores. Zootaxa, 4263 (3), 450–466.
https://doi.org/10.11646/zootaxa.4263.3.2Guidetti, 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. & Moly de 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
Iharos, G. (1973) Neuere Daten zur Kenntnis der Tardigraden-Fauna von Neuguinea. Opuscula Zoologica, Budapest, 11, 65–73.
Kaczmarek, Ł. & Michalczyk, Ł. (2004) New records of Tardigrada from Cyprus with a description of the new species Macrobiotus marlenae (hufelandi group) (Eutardigrada: Macrobiotidae). Genus, 15 (1), 141–152.
Kaczmarek, Ł. & Michalczyk, Ł. (2009) Two new species of Macrobiotidae, Macrobiotus szeptyckii (harmsworthi group) and Macrobiotus kazmierskii (hufelandi group) from Argentina. Acta Zoologica Cracoviensia, 52B, 87–99.
https://doi.org/10.3409/azc.52b_1-2.87-99
Kaczmarek, Ł. & Michalczyk, Ł. (2017) The Macrobiotus hufelandi group (Tardigrada) revisited. Zootaxa, 4363 (1), 101–123.
https://doi.org/10.11646/zootaxa.4363.1.4
Kaczmarek, Ł., Michalczyk, Ł. & McInnes, S.J. (2014a). 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., Michalczyk, Ł. & McInnes, S.J. (2015) 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.
Kaczmarek, Ł., Kosicki, J.Z. & Roszkowska, M. (2018b) Tardigrada of Bory Tucholskie National Park, Zaborski Landscape Park and their surroundings (Pomerania Province, Poland). Turkish Journal of Zoology, 42, 6–17.
https://doi.org/10.3906/zoo-1705-44
Kaczmarek, Ł., Cytan, J., Zawierucha, K., Diduszko, D. & Michalczyk, Ł. (2014b) Tardigrades from Peru (South America), with descriptions of three new species of Parachela. Zootaxa, 3790 (2), 357–379.
https://doi.org/10.11646/zootaxa.3790.2.5
Kaczmarek, Ł., Gawlak, M., Bartels, P.J., Nelson, D.R. & Roszkowska, M. (2017) Revision of the genus Paramacrobiotus Guidetti et al., 2009 with the description of a new species, re-descriptions and a key. Annales Zoologici, 67, 627–656.
https://doi.org/10.3161/00034541ANZ2017.67.4.001
Kaczmarek, Ł., Parnikoza, I., Gawlak, M., Esefeld, J., Peter, H., Kozeretska, I. & Roszkowska, M. (2018a) Tardigrades from Larus dominicanus Lichtenstein, 1823 nests on the Argentine Islands (maritime Antarctic). Polar Biology, 41, 283–301.
https://doi.org/10.1007/s00300-017-2190-4
Kristensen, R.M. (1987) Generic revision of the Echiniscidae (Heterotardigrada), with a discussion of the origin of the family. In: Bertolani, R. (Ed.), Biology of Tardigrada. Selected Symposia and Monographs. U.Z.I., Mucchi Modena, pp. 261–335.
Lisi, O. (2011) Remarks on Doryphoribius flavus (Iharos, 1966), and description of three new species (Tardigrada, Hypsibiidae). Zootaxa, 2834, 17–32.
Marcus, E. (1927) Zur Anatomie und Ökologie mariner Tardigraden. Zoologische Jahrbücher. Abteilungfür Systematik, 53, 487–558.
Marley, N.J. (2006) A new species of Tardigrada, Platicrista ramsayi sp. nov. from the páramo of Volcán Chiles, Ecuador. Zootaxa, 1166, 35–48.
Marley, N.J., McInnes, S.J. & Sands, C.J. (2011) Phylum Tardigrada: a re-evaluation of the Parachela. Zootaxa, 2819, 51–64.
Maucci, W. (1988) Tardigrada from Patagonia (Southern South America) with description of three new species. Revista Chilena de Entomologia, 16, 5–13.
McInnes, S.J. (1994) Zoogeographic distribution of terrestrial/freshwater tardigrades from current literature. Journal of the Natural History, 28, 257–352.
https://doi.org/10.1080/00222939400770131
Meyer, H.A., Domingue, M.N. & Hinton, J.G. (2014) Tardigrada of the West Gulf Coastal Plain, with descriptions of two new species from Louisiana. Southeastern Naturalist, 13 (5), 117–130.
https://doi.org/10.1656/058.013.s511
Michalczyk, Ł. & Kaczmarek, Ł. (2003) A description of the new tardigrade Macrobiotus reinhardti (Eutardigrada: Macrobiotidae, harmsworthi group) with some remarks on the oral cavity armature within the genus Macrobiotus Schultze. Zootaxa, 331 (1), 1–24.
https://doi.org/10.11646/zootaxa.331.1.1
Michalczyk, Ł. & Kaczmarek, Ł. (2006) Revision of the Echiniscus bigranulatus group with a description of a new species Echiniscus madonnae (Tardigrada: Heterotardigrada: Echiniscidae) from South America. Zootaxa, 1154, 1–26.
Michalczyk, Ł. & Kaczmarek, Ł. (2007) Echiniscus ganczareki, a new species of Tardigrada (Heterotardigrada: Echiniscidae, bigranulatus group) from Costa Rica. Zootaxa, 1471, 15–25.
Michalczyk, Ł. & Kaczmarek, Ł. (2013) The Tardigrada Register: a comprehensive online data repository for tardigrade taxonomy. Journal of Limnology, 72 (Supplement 1), 175–181.
https://doi.org/10.4081/jlimnol.2013.s1.e22
Nelson, D.R., Guidetti, R. & Rebecchi, L. (2015) Phylum Tardigrada. In: Thorp, J. & Rogers, D.C. (Eds.), Ecology and General Biology: Thorp and Covich's Freshwater Invertebrates. Academic Press, London, pp. 347–380.
https://doi.org/10.1016/B978-0-12-385026-3.00017-6
Nickel, K., Miller, W.R. & Marley, N.J. (2001) Tardigrades of South America: Machu Picchu and Ollantaytambo, Peru. Zoologischer Anzeiger, 240 (3–4), 505–509.
https://doi.org/10.1078/0044-5231-00060
Pilato, G. (1981) Analisi di nuovi caratteri nello studio degli Eutardigradi. Animalia, 8, 51–57.
Pilato, G. (2000) Macrobiotus centesimus, new species of eutardigrade from the South America. Bollettino delle sedute della Accademia Gioenia di Scienze Naturali in Catania, 33, 97–101.
Pilato, G. (2007) Echiniscus quitensis, a new species of tardigrade from Ecuador (Heterotardigrada: Echiniscidae). Zootaxa, 1389 (4), 55–60.
https://doi.org/10.11646/zootaxa.1389.4
Pilato, G. & Bertolani, R. (2004) Macrobiotus dariae sp. n., a new species of eutardigrade (Eutardigrada, Macrobiotidae) from Cyprus. Zootaxa, 638 (1), 1–7.
https://doi.org/10.11646/zootaxa.638.1.1
Pilato, G. & Binda, M.G. (2010) Definition of families, subfamilies, genera and subgenera of the Eutardigrada, and keys to their identification. Zootaxa, 2404 (1), 1–52.
https://doi.org/10.11646/zootaxa.2404.1.1
Pilato, G., Binda, M.G. & Lisi, O. (2003b) Remarks on some species of tardigrades from South America with description of Minibiotus sidereus n. sp. Zootaxa, 195 (1), 1–8.
https://doi.org/10.11646/zootaxa.195.1.1
Pilato, G., Kaczmarek, Ł., Michalczyk, Ł. & Lisi, O. (2003a) Macrobiotus polonicus, a new species of Tardigrada from Poland (Eutardigrada, Macrobiotidae, 'hufelandi group'). Zootaxa, 258 (1), 1–8.
https://doi.org/10.11646/zootaxa.258.1.1
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). Bollettine Delle Sedute Dell’ Accademia Gioenia di Scienze Naturali in Catania, 33, 103–120.
Pilato, G., Binda, M.G., Napolitano, A. & Moncada. E. (2001) Notes on South American tardigrades with the description of two new species: Pseudechiniscus spinerectus and Macrobiotus danielae. Tropical Zoology, 14, 223–231.
https://doi.org/10.1080/03946975.2001.10531154
Pilato, G., Binda, M.G., Napolitano, A. & Moncada, E. (2002) Tardigrades from Ecuador, with the description of two new species: Mixibius ornatus n. sp. and Diphascon (Adropion) onorei n. sp. (Eutardigrada, Hypsibiidae). Studies on Neotropical Fauna and Environment, 37, 175–179.
https://doi.org/10.1076/snfe.37.2.175.8584
Pilato, G., Binda, M.G., Napolitano, A. & Moncada, E. (2004) Remarks on some species of tardigrades from South America with description of two new species. Journal of Natural History, 38, 1081–1086.
https://doi.org/10.1080/0022293031000071541
Ramazzotti, G. (1964) Tardigradi del Cile II, con descrizione di due nuove specie e note sulla scultura degli Echiniscidae. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 103, 89–100.
Ramazzotti, G. & Maucci, W. (1983) Il Phylum Tardigrada. Memorie dell’Istituto Italiano di Idrobiologia, Pallanza, 41, 1–1012.
Ramazzotti, G. & Maucci, W. (1995) The phylum Tardigrada. 3rd Edition. Abilene: University of Texas, pp. [translated by Beasley, C.W.]
Richters, F. (1908) Moosbewohner. Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition, 6, 1–16.
Richters, F. (1926) Tardigrada. In: Kükenthalund, W. & Krumbach, T. (Eds.), Handbuch der Zoologie. Walter de Gruyter & Co., Berlin, pp. 1–68.
Roszkowska, M., Stec, D., Ciobanu, D.A. & Kaczmarek, Ł. (2016) Tardigrades from Nahuel Huapi National Park (Argentina, South America) with descriptions of two new Macrobiotidae species. Zootaxa, 4105 (3), 243–260.
https://doi.org/10.11646/zootaxa.4105.3.2
Roszkowska, M., Ostrowska, M., Stec, D., Janko, K. & Kaczmarek, Ł. (2017) Macrobiotus polypiformis, a new tardigrade (Macrobiotidae; hufelandi group) from the Ecuadorian Pacific coast, with remarks on the claw abnormalities in eutardigrades. Journal of European Taxonomy, 327, 1–19.
https://doi.org/10.5852/ejt.2017.327
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.
Schultze, C.A.S. (1840) Echiniscus bellermanni, animal crustaceum. Scribebam Berolini, 1–8.
Schuster, R.O. & Grigarick, A.A. (1966) Tardigrada from the Galápagos and Cocos Islands. Proceedings of the California Academy of Sciences, 34, 315–328.
Schuster, R.O., Nelson, D.R., Grigarick, A.A. & Christenberry, D. (1980) Systematic criteria of Eutardigrada. Transactions of the American Microscopical Society, 99, 284–303.
https://doi.org/10.2307/3226004
Stec, D., Smolak, R., Kaczmarek, Ł. & Michalczyk, Ł. (2015) An integrative description of Macrobiotus paulinae sp. nov. (Tardigrada: Eutardigrada: Macrobiotidae: hufelandi group) from Kenya. Zootaxa, 4052 (5), 501–526.
https://doi.org/10.11646/zootaxa.4052.5.1
St Louis, R., Burningham, L., Dowl, A. & Grosberg, M. (2009) Lonely Planet Guide: Ecuador & the Galápagos Islands. Lonely Planet, 2009, 1–432.
Thulin, G. (1928) Über die Phylogenie und das System der Tardigraden. Hereditas, Lund, 11, 207–266.
https://doi.org/10.1111/j.1601-5223.1928.tb02488.x