Abstract
Marine tardigrades were sampled at three sites on Mexico’s Caribbean coast. Eleven taxa were collected, one of which is described as a new species. Styraconyx robertoi sp. nov. is characterized by: asymmetric primary clavae; dorsal cuticle with a grid-like sculpture; claws with reduced accessory hooks; females with peduncles on only two digits (the external) of legs I–IV; males with peduncles only on the external digits of legs I–III and peduncles on all four digits of leg IV. Styraconyx robertoi sp. nov. is most similar to S. craticuliformis Chang & Rho, 1998 and S. kristenseni Renaud-Mornant, 1981 by having asymmetric primary clavae but differs from S. craticuliformis mainly by the number of peduncles and from S. kristenseni mainly by the presence of a grid-like dorsal sculpture. Additionally, a comparison of material collected from the same region, but reported previously only as genus level records, was carried out in order to produce a refined list of the known Mexican marine tardigrade species.
References
Álvarez-Hernández, J.H. (2003) Trophic model of a fringing coral reef in the Southern Mexican Caribbean. Fisheries Centre Research Reports, 11, 227–235.
Ardisson, P.L., May-Kú, M.A., Herrera-Dorantes, M.T. & Arellano-Guillermo, A. (2011) El Sistema Arrecifal Mesoamericano-México: consideraciones para su designación como Zona Marítima Especialmente Sensible. Hidrobiológica, 21, 261–280.
Bartels, P.J., Fontoura, P. & Nelson, D.R. (2015) New records of marine tardigrades from Moorea, French Polynesia, with the description of Styraconyx turbinarium sp. nov. (Arthrotardigrada, Halechiniscidae). Zootaxa, 3955 (3), 389–402.
https://doi.org/10.11646/zootaxa.3955.3.6
Bartels, P.J., Fontoura, P. & Nelson, D.R. (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 (1), 43–70.
https://doi.org/10.11646/zootaxa.4420.1.3
Bello, G. & de Zio Grimaldi, S. (1998) Phylogeny of the genera of the Stygarctidae and related families (Tardigrada: Heterotardigrada). Zoologischer Anzeiger, 237, 171–183.
Binda, M.G. (1978) Risistemazione di alcuni tardigradi con l’istituzione di un nuovo genere di Oreellidae e della nuova famiglia Archechiniscidae. Animalia, 5, 307–314.
Chang, C.Y. & Rho, H.S. (1998) Three new tardigrade species associated with barnacles from the Thai coast of Andaman Sea. Korean Journal of Biological Sciences, 2, 323–331.
https://doi.org/10.1080/12265071.1998.9647426
D’Addabbo Gallo, M., de Zio Grimaldi, S., Morone De Lucia, R.M. & Troccoli, A. (1992) Halechiniscidae and Echiniscoididae from the Western Mediterranean Sea (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine, 33, 299–318.
D’Addabbo Gallo, M., Morone De Lucia, M.R. & de Zio Grimaldi, S. (1989) Two new species of the genus Styraconyx (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine, 30, 17–33.
de Jesús-Navarrete, A. (2003) Diversity of Nematoda in a Caribbean atoll: Banco Chinchorro, Mexico. Bulletin of Marine Science, 73, 47–56.
de Jesús-Navarrete, A. (2007) Nematodos de los arrecifes de Isla Mujeres y Banco Chinchorro, Quintana Roo, México. Revista de Biología Marina y Oceanografía, 42, 193–200.
https://doi.org/10.4067/s0718-19572007000200006
Doyère, L.M. (1840) Mémoire sur les Tardigrades. I. Annales des Sciences Naturelles, Paris, Série 2, 14, 269–361.
Grimaldi de Zio, S., D’Addabbo Gallo, M., Morone de Lucia, R. & D’Addabo, L. (1987) Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean. Bollettino di Zoologia, 54 (4), 347–357.
https://doi.org/10.1080/11250008709355608
Jordán-Dahlgren, E. (1993) El ecosistema arrecifal coralino del Atlántico Mexicano. Revista de la Sociedad Mexicana de Historia Natural, 45, 157–175.
Jørgensen, A., Boesgaard, T.M., Møbjerg, N. & Kristensen, R.M. (2014) The tardigrade fauna of Australian marine caves: With descriptions of nine new species of Arthrotardigrada. Zootaxa, 3802 (4), 401–443.
https://doi.org/10.11646/zootaxa.3802.4.1
Kristensen, R.M. (1983) Loricifera, a new phylum with Aschelminthes characters from the meiobenthos. Journal of Zoological Systematics and Evolutionary Research, 21 (3), 163–180.
https://doi.org/10.1111/j.1439-0469.1983.tb00285.x
Kristensen, R.M. (1984) On the biology of Wingstrandarctus corallinus nov. gen. et spec., with notes on the symbiontic bacteria in the subfamily Florarctinae (Arthrotardigrada). Videnskabelige Meddelelser Dansk naturhistorisk Forening, 145, 201–218.
Kristensen, R.M. & Hallas, T.E. (1980) The tidal genus Echiniscoides and its variability, with erection of Echiniscoididae fam. n. (Tardigrada). Zoologica Scripta, 9, 113–127.
https://doi.org/10.1111/j.1463-6409.1980.tb00657.x
Kristensen, R.M. & Higgins, R.P. (1984) Revision of Styraconyx (Tardigrada: Halechiniscidae) with description of two new species from Disko Bay, West Greenland. Smithsonian Contributions to Zoology, 391, 1–40.
https://doi.org/10.5479/si.00810282.391
Kristensen, R.M. & Higgins, R.P. (1989) Marine Tardigrada from the Southeastern United States coastal waters. I. Paradoxipus orzeliscoides n. gen., n. sp. (Arthrotardigrada: Halechiniscidae). Transactions of the American Microscopical Society, 108, 262–282.
https://doi.org/10.2307/3226344
Kristensen, R.M. & Mackness, B.S. (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 (2), 73–87.
Kristensen, R.M. & 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.
Marcus, E. (1927) Zur Anatomie und Ökologie mariner Tardigraden. Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere, 53, 487–588.
McKirdy, D.J., Schmidt, P. & McGinty-Bayly, M. (1976) Interstitielle Fauna von Galapagos. XVI. Tardigrada. Mikrofauna des Meeresbodens, Mainz, 58, 409–449.
Miller, W.R. & Perry, E.S. (2016) The coastal marine Tardigrada of the Americas. Zootaxa, 4126 (3), 375–396.
https://doi.org/10.11646/zootaxa.4126.3.4
Møbjerg, N., Jørgensen, A. & Kristensen, R.M. (2019). Ongoing revision of Echiniscoididae (Heterotardigrada: Echiniscoidea), with the description of a new interstitial species and genus with unique anal structures. Zoological Journal of Linnean Society, 181, 1–18.
https://doi.org/10.1093/zoolinnean/zlz122
Pérez-Pech, W.A., Anguas-Escalante A., de Jesús-Navarrete A. & Hansen J.G. (2018) Primer registro genérico de tardígrados marinos en costas de Quintana Roo, México. Academia Journals, 4 (10), 1909–1912.
Pérez-Pech, W.A., Hansen, J.G., DeMilio, E., de Jesús-Navarrete, A., Martínez Mendoza, I., Rocha Olivares, A. & Vargas-Espositos, A. (2020) First records of marine tardigrades of the genus Coronarctus (Tardigrada, Heterotardigrada, Arthrotardigrada) from Mexico. Check List. [in press]
Plate, L.H. (1888) Beiträge zur Naturgeschichte der Tardigraden. Zoologische Jahrbücher, 3, 487–550.
https://doi.org/10.5962/bhl.part.1265
Pollock, L.W. (1970) Batillipes dicrocercus n. sp., Stygarctus granulatus n. sp. and other Tardigrada from Woods Hole, Massachusetts, U.S.A. Transactions of the American Microscopical Society, 89 (1), 38–52.
https://doi.org/10.2307/3224613
Pollock, L.W. (1975) Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean. Cahiers de Biologie Marine, 16, 121–132.
Pollock, L.W. (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. The Bulletin of Marine Science, 33, 109–117.
Pollock, L.W. (1995) New marine tardigrades from Hawaiian beach sand and phylogeny of the family Halechiniscidae. Invertebrate Biology, 114 (3), 220–235.
https://doi.org/10.2307/3226877
Ramazzotti, G. (1962) Il Phylum Tardigrada. Memorie dell’Istituto Italiano di Idrobiologia, 14, 1–595.
Renaud-Debyser, J. (1959) Études sur la faune interstitielle des Iles Bahamas. III. Tardigrades. Vie et Milieu, 10, 296–302.
Renaud-Mornant, J. (1974) Une nouvelle famille de Tardigrades marins abyssaux: les Coronarctidae fam nov (Heterotardigrada). Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences, Paris, 278, 3087–3090.
Renaud-Mornant, J. (1981) Tardigrades marins (Arthrotardigrada) du Pacifique Sud. Bulletin du Muséum National d’Histoire Naturelle, Paris, Série 4e, Section A, 3, 799–813.
Renaud-Mornant, J. (1982) Sous-famille et genre nouveaux de Tardigrades marins (Arthrotardigrada). Bulletin du Muséum National d’Histoire Naturelle, Paris, Série 4e, Section A, 1–2, 89–94.
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. (Eds.), Handbuch der Zoologie. Vol. 3. Walter de Gruyter & Co., Berlin and Leipzig, pp. 58–61.
Roberts, C.M., McClean, C.J., Vernon, J.E.N., Hawkins, J.P., Allen, G.R., McAllister, D.E., Mittermeier, C.G., Schueler, F.W., Spalding, M., Wells, F., Vynne, C. & Werner, T.B. (2002) Marine Biodiversity Hotspots and Conservation Priorities for Tropical Reefs. Science, 295 (5558), 1280–1284.
https://doi.org/10.1126/science.1067728
Romano III, F.A., Gallo, M., D’Addabbo, R., Accogli, G., Baguley, J. & Montagna, P. (2011) Deep-sea tardigrades in the northern Gulf of Mexico with a description of a new species of Coronarctidae (Tardigrada : Arthrotardigrada), Coronarctus mexicus. Journal of Zoological Systematics and Evolutionary Research, 49, 48–52.
https://doi.org/10.1111/j.1439-0469.2010.00597.x
Santos, E., Rubal, M., Veiga, P., da Rocha, C.M.C. & Fontoura, P. (2018) Batillipes (Tardigrada, Arthrotardigrada) from the Portuguese coast with the description of two new species and a new dichotomous key for all species. European Journal of Taxonomy, 425, 1–32.
https://doi.org/10.5852/ejt.2018.425
Schulz, E. (1951) Über Stygarctus bradypus n. g. n. sp., einen Tardigraden aus dem Kustengrundwasser, und seine phylogenetische Bedeutung. Kieler Meeresforschungen, 8, 86–97.
Schulz, E. (1953) Eine neue Tardigraden-Gattung von der pazifischen Küste. Zoologischer Anzeiger, 151, 306–310.
Schulz, E. (1955) Studien an marinen Tardigraden. Kieler Meeresforschungen, 11, 73–79.
Schulz, E. (1963) Über die Tardigraden. Zoologischer Anzeiger, 171, 3–12.
Suárez-Morales, E. & Gasca, R. (1998) Updated checklist of the free-living marine Copepoda (Crustacea) of Mexico. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, Serie Zoología, 69 (1), 105–109.
Thulin, G. (1928) Über die Phylogenie und das System der Tardigraden. Hereditas Genetiskt Arkiv, 11, 207–266.
https://doi.org/10.1111/j.1601-5223.1928.tb02488.x
Thulin, G. (1942) Ein neuer mariner Tardigrad. Meddelanden från Göteborgs Musei Zoologiska Avdeling, 99, 1–10.
Winfield, I. & Escobar-Briones, E. (2007) Anfípodos (Crustacea: Gammaridea) del sector norte del Mar Caribe: listado faunístico, registros nuevos y distribución espacial. Revista Mexicana de Biodiversidad, 78, 51–61.
https://doi.org/10.22201/ib.20078706e.2007.001.411
Álvarez-Hernández, J.H. (2003) Trophic model of a fringing coral reef in the Southern Mexican Caribbean. Fisheries Centre Research Reports, 11, 227–235.
Ardisson, P.L., May-Kú, M.A., Herrera-Dorantes, M.T. & Arellano-Guillermo, A. (2011) El Sistema Arrecifal Mesoamericano-México: consideraciones para su designación como Zona Marítima Especialmente Sensible. Hidrobiológica, 21, 261–280.
Bartels, P.J., Fontoura, P. & Nelson, D.R. (2015) New records of marine tardigrades from Moorea, French Polynesia, with the description of Styraconyx turbinarium sp. nov. (Arthrotardigrada, Halechiniscidae). Zootaxa, 3955 (3), 389–402.
https://doi.org/10.11646/zootaxa.3955.3.6
Bartels, P.J., Fontoura, P. & Nelson, D.R. (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 (1), 43–70.
https://doi.org/10.11646/zootaxa.4420.1.3
Bello, G. & de Zio Grimaldi, S. (1998) Phylogeny of the genera of the Stygarctidae and related families (Tardigrada: Heterotardigrada). Zoologischer Anzeiger, 237, 171–183.
Binda, M.G. (1978) Risistemazione di alcuni tardigradi con l’istituzione di un nuovo genere di Oreellidae e della nuova famiglia Archechiniscidae. Animalia, 5, 307–314.
Chang, C.Y. & Rho, H.S. (1998) Three new tardigrade species associated with barnacles from the Thai coast of Andaman Sea. Korean Journal of Biological Sciences, 2, 323–331.
https://doi.org/10.1080/12265071.1998.9647426
D’Addabbo Gallo, M., de Zio Grimaldi, S., Morone De Lucia, R.M. & Troccoli, A. (1992) Halechiniscidae and Echiniscoididae from the Western Mediterranean Sea (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine, 33, 299–318.
D’Addabbo Gallo, M., Morone De Lucia, M.R. & de Zio Grimaldi, S. (1989) Two new species of the genus Styraconyx (Tardigrada: Heterotardigrada). Cahiers de Biologie Marine, 30, 17–33.
de Jesús-Navarrete, A. (2003) Diversity of Nematoda in a Caribbean atoll: Banco Chinchorro, Mexico. Bulletin of Marine Science, 73, 47–56.
de Jesús-Navarrete, A. (2007) Nematodos de los arrecifes de Isla Mujeres y Banco Chinchorro, Quintana Roo, México. Revista de Biología Marina y Oceanografía, 42, 193–200.
https://doi.org/10.4067/s0718-19572007000200006
Doyère, L.M. (1840) Mémoire sur les Tardigrades. I. Annales des Sciences Naturelles, Paris, Série 2, 14, 269–361.
Grimaldi de Zio, S., D’Addabbo Gallo, M., Morone de Lucia, R. & D’Addabo, L. (1987) Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean. Bollettino di Zoologia, 54 (4), 347–357.
https://doi.org/10.1080/11250008709355608
Jordán-Dahlgren, E. (1993) El ecosistema arrecifal coralino del Atlántico Mexicano. Revista de la Sociedad Mexicana de Historia Natural, 45, 157–175.
Jørgensen, A., Boesgaard, T.M., Møbjerg, N. & Kristensen, R.M. (2014) The tardigrade fauna of Australian marine caves: With descriptions of nine new species of Arthrotardigrada. Zootaxa, 3802 (4), 401–443.
https://doi.org/10.11646/zootaxa.3802.4.1
Kristensen, R.M. (1983) Loricifera, a new phylum with Aschelminthes characters from the meiobenthos. Journal of Zoological Systematics and Evolutionary Research, 21 (3), 163–180.
https://doi.org/10.1111/j.1439-0469.1983.tb00285.x
Kristensen, R.M. (1984) On the biology of Wingstrandarctus corallinus nov. gen. et spec., with notes on the symbiontic bacteria in the subfamily Florarctinae (Arthrotardigrada). Videnskabelige Meddelelser Dansk naturhistorisk Forening, 145, 201–218.
Kristensen, R.M. & Hallas, T.E. (1980) The tidal genus Echiniscoides and its variability, with erection of Echiniscoididae fam. n. (Tardigrada). Zoologica Scripta, 9, 113–127.
https://doi.org/10.1111/j.1463-6409.1980.tb00657.x
Kristensen, R.M. & Higgins, R.P. (1984) Revision of Styraconyx (Tardigrada: Halechiniscidae) with description of two new species from Disko Bay, West Greenland. Smithsonian Contributions to Zoology, 391, 1–40.
https://doi.org/10.5479/si.00810282.391
Kristensen, R.M. & Higgins, R.P. (1989) Marine Tardigrada from the Southeastern United States coastal waters. I. Paradoxipus orzeliscoides n. gen., n. sp. (Arthrotardigrada: Halechiniscidae). Transactions of the American Microscopical Society, 108, 262–282.
https://doi.org/10.2307/3226344
Kristensen, R.M. & Mackness, B.S. (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 (2), 73–87.
Kristensen, R.M. & 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.
Marcus, E. (1927) Zur Anatomie und Ökologie mariner Tardigraden. Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere, 53, 487–588.
McKirdy, D.J., Schmidt, P. & McGinty-Bayly, M. (1976) Interstitielle Fauna von Galapagos. XVI. Tardigrada. Mikrofauna des Meeresbodens, Mainz, 58, 409–449.
Miller, W.R. & Perry, E.S. (2016) The coastal marine Tardigrada of the Americas. Zootaxa, 4126 (3), 375–396.
https://doi.org/10.11646/zootaxa.4126.3.4
Møbjerg, N., Jørgensen, A. & Kristensen, R.M. (2019). Ongoing revision of Echiniscoididae (Heterotardigrada: Echiniscoidea), with the description of a new interstitial species and genus with unique anal structures. Zoological Journal of Linnean Society, 181, 1–18.
https://doi.org/10.1093/zoolinnean/zlz122
Pérez-Pech, W.A., Anguas-Escalante A., de Jesús-Navarrete A. & Hansen J.G. (2018) Primer registro genérico de tardígrados marinos en costas de Quintana Roo, México. Academia Journals, 4 (10), 1909–1912.
Pérez-Pech, W.A., Hansen, J.G., DeMilio, E., de Jesús-Navarrete, A., Martínez Mendoza, I., Rocha Olivares, A. & Vargas-Espositos, A. (2020) First records of marine tardigrades of the genus Coronarctus (Tardigrada, Heterotardigrada, Arthrotardigrada) from Mexico. Check List. [in press]
Plate, L.H. (1888) Beiträge zur Naturgeschichte der Tardigraden. Zoologische Jahrbücher, 3, 487–550.
https://doi.org/10.5962/bhl.part.1265
Pollock, L.W. (1970) Batillipes dicrocercus n. sp., Stygarctus granulatus n. sp. and other Tardigrada from Woods Hole, Massachusetts, U.S.A. Transactions of the American Microscopical Society, 89 (1), 38–52.
https://doi.org/10.2307/3224613
Pollock, L.W. (1975) Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean. Cahiers de Biologie Marine, 16, 121–132.
Pollock, L.W. (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. The Bulletin of Marine Science, 33, 109–117.
Pollock, L.W. (1995) New marine tardigrades from Hawaiian beach sand and phylogeny of the family Halechiniscidae. Invertebrate Biology, 114 (3), 220–235.
https://doi.org/10.2307/3226877
Ramazzotti, G. (1962) Il Phylum Tardigrada. Memorie dell’Istituto Italiano di Idrobiologia, 14, 1–595.
Renaud-Debyser, J. (1959) Études sur la faune interstitielle des Iles Bahamas. III. Tardigrades. Vie et Milieu, 10, 296–302.
Renaud-Mornant, J. (1974) Une nouvelle famille de Tardigrades marins abyssaux: les Coronarctidae fam nov (Heterotardigrada). Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences, Paris, 278, 3087–3090.
Renaud-Mornant, J. (1981) Tardigrades marins (Arthrotardigrada) du Pacifique Sud. Bulletin du Muséum National d’Histoire Naturelle, Paris, Série 4e, Section A, 3, 799–813.
Renaud-Mornant, J. (1982) Sous-famille et genre nouveaux de Tardigrades marins (Arthrotardigrada). Bulletin du Muséum National d’Histoire Naturelle, Paris, Série 4e, Section A, 1–2, 89–94.
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. (Eds.), Handbuch der Zoologie. Vol. 3. Walter de Gruyter & Co., Berlin and Leipzig, pp. 58–61.
Roberts, C.M., McClean, C.J., Vernon, J.E.N., Hawkins, J.P., Allen, G.R., McAllister, D.E., Mittermeier, C.G., Schueler, F.W., Spalding, M., Wells, F., Vynne, C. & Werner, T.B. (2002) Marine Biodiversity Hotspots and Conservation Priorities for Tropical Reefs. Science, 295 (5558), 1280–1284.
https://doi.org/10.1126/science.1067728
Romano III, F.A., Gallo, M., D’Addabbo, R., Accogli, G., Baguley, J. & Montagna, P. (2011) Deep-sea tardigrades in the northern Gulf of Mexico with a description of a new species of Coronarctidae (Tardigrada : Arthrotardigrada), Coronarctus mexicus. Journal of Zoological Systematics and Evolutionary Research, 49, 48–52.
https://doi.org/10.1111/j.1439-0469.2010.00597.x
Santos, E., Rubal, M., Veiga, P., da Rocha, C.M.C. & Fontoura, P. (2018) Batillipes (Tardigrada, Arthrotardigrada) from the Portuguese coast with the description of two new species and a new dichotomous key for all species. European Journal of Taxonomy, 425, 1–32.
https://doi.org/10.5852/ejt.2018.425
Schulz, E. (1951) Über Stygarctus bradypus n. g. n. sp., einen Tardigraden aus dem Kustengrundwasser, und seine phylogenetische Bedeutung. Kieler Meeresforschungen, 8, 86–97.
Schulz, E. (1953) Eine neue Tardigraden-Gattung von der pazifischen Küste. Zoologischer Anzeiger, 151, 306–310.
Schulz, E. (1955) Studien an marinen Tardigraden. Kieler Meeresforschungen, 11, 73–79.
Schulz, E. (1963) Über die Tardigraden. Zoologischer Anzeiger, 171, 3–12.
Suárez-Morales, E. & Gasca, R. (1998) Updated checklist of the free-living marine Copepoda (Crustacea) of Mexico. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, Serie Zoología, 69 (1), 105–109.
Thulin, G. (1928) Über die Phylogenie und das System der Tardigraden. Hereditas Genetiskt Arkiv, 11, 207–266.
https://doi.org/10.1111/j.1601-5223.1928.tb02488.x
Thulin, G. (1942) Ein neuer mariner Tardigrad. Meddelanden från Göteborgs Musei Zoologiska Avdeling, 99, 1–10.
Winfield, I. & Escobar-Briones, E. (2007) Anfípodos (Crustacea: Gammaridea) del sector norte del Mar Caribe: listado faunístico, registros nuevos y distribución espacial. Revista Mexicana de Biodiversidad, 78, 51–61.
https://doi.org/10.22201/ib.20078706e.2007.001.411