Ukrainian Antarctic Journal

Vol 24 No 1(32) (2026): Ukrainian Antarctic Journal
Articles

Diversity and host specificity of monogeneans in Antarctic fishes: a review

Ivanna Dudliv
Institute of Marine Biology, National Academy of Science of Ukraine, Odesa, 65048, Ukraine; Institute of Vertebrate Biology, Czech Academy of Sciences, Brno, 603 00, Czech Republic
Yuriy Kvach
Institute of Marine Biology, National Academy of Science of Ukraine, Odesa, 65048, Ukraine; Institute of Vertebrate Biology, Czech Academy of Sciences, Brno, 603 00, Czech Republic
Published July 2, 2026
Keywords
  • Antarctica,
  • marine parasitology,
  • Monogenea,
  • Nototheniidae,
  • fish parasite,
  • diversity baseline
  • ...More
    Less
How to Cite
Dudliv, I., & Kvach, Y. (2026). Diversity and host specificity of monogeneans in Antarctic fishes: a review. Ukrainian Antarctic Journal, 24(1(32), 60-76. https://doi.org/10.33275/1727-7485.1.2026.758

Abstract

We compiled and harmonised published records of monogeneans from Antarctic and sub-Antarctic fishes, standardising host names, georeferencing localities, deduplicating records, and excluding ambiguous entries (genuslevel hosts or “not specified” hosts) from specificity metrics. We aggregated occurrences from 49 unique localities spanning the Scotia Arc, Antarctic Peninsula, East Antarctica, Ross Sea, and the Kerguelen–Crozet–Heard–Macquarie region. The dataset comprises 31 species (35 including genus-level records), representing 11 genera in 7 families and two subclasses, drawn from 46 publications; species richness within the Subclass Monopisthocotylea was concentrated in Capsalidae (10 species), Gyrodactylidae (9), and Tetraonchoididae (8), with one species each in Mazocraeidae, Hexabothriidae, and Diclidophoridae within Polyopisthocotylea. We identified 29 species-level taxa with unambiguous hosts; of these, 15 (51.7%) were recorded from a single fish host, 6 (20.7%) from two hosts, and 8 (27.6%) from three or more (median = 1; maximum = 12 for Pseudobenedenia nototheniae). The spatial distribution of records was uneven, with the Kerguelen Plateau and the Ross Sea containing the densest clusters. In contrast, many other records occurred as isolated points, limiting the reliability of inferring species ranges. Overall, our analysis provides a curated baseline of diversity, host associations, and regional coverage for Antarctic monogeneans, highlights gaps caused by underspecified hosts and localities, and supplies standardised counts for future taxonomic and ecological work.

References

  1. Bargelloni, L., Ritchie, P. A., Patarnello, T., Battaglia, B., Lambert, D. M., & Meyer, A. (1994). Molecular evolution at subzero temperatures: mitochondrial and nuclear phylogenies of fishes from Antarctica (suborder Notothenioidei), and the evolution of antifreeze glycopeptides. Molecular Biology and Evolution, 11(6), 854–863. https://doi.org/10.1093/oxfordjournals.molbev.a040168
  2. Boulenger, G. A. (1902). XXXIII. – Notes on the classification of Teleostean fishes – II. On the Berycidæ. Annals and Magazine of Natural History, Ser. 7, 9(51), 197–204. https://doi.org/10.1080/00222930208678568
  3. Brickle, P., MacKenzie, K., & Pike, A. (2005). Parasites of the Patagonian toothfish, Dissostichus eleginoides Smitt 1898, in different parts of the Subantarctic. Polar Biology, 28(8), 663–671. https://doi.org/10.1007/s00300-005-0737-2
  4. Brickle, P., MacKenzie, K., & Pike, A. (2006). Variations in the parasite fauna of the Patagonian toothfish (Dissostichus eleginoides Smitt, 1898), with length, season, and depth of habitat around the Falkland Islands. Journal of Parasitology, 92(2), 282–291. https://doi.org/10.1645/GE-539R.1
  5. Buchmann, K., & Bresciani, J. (2006). Monogenea (phylum Platyhelminthes). In P. T. K. Woo (Ed.), Fish Diseases and Disorders. Vol. 1,Protozoan and Metazoan Infections. (pp. 297–344). Wallingford: CABI Publishing. https://doi.org/10.1079/9780851990156.0297
  6. Bychowskii, B. E., Guzev, A. V., & Nagibina, L. F. (1967). Monogenetic trematodes of the family Tetraonchoididae Bychowsky, 1951. (Translation series, No. 17). Virginia Institute of Marine Science, William & Mary. https://scholarworks.wm.edu/handle/internal/16293
  7. Clarke, A., Griffiths, H. J., Linse, K., Barnes, D. K. A., & Crame, J. A. (2007). How well do we know the Antarctic marine fauna? A preliminary study of macroecological and biogeographical patterns in Southern Ocean gastropod and bivalve molluscs. Diversity and Distributions, 13(5), 620–632. https://doi.org/10.1111/j.1472-4642.2007.00380.x
  8. Convey, P., & Peck, L. S. (2019). Antarctic environmental change and biological responses. Science Advances, 5(11), eaaz0888. https://doi.org/10.1126/sciadv.aaz0888
  9. De Broyer, C., Koubbi, P., Griffiths, H. J., Raymond, B., d’Udekem d’Acoz, C., Van de Putte, A. P., Danis, B., David, B., Grant, S., Gutt, J., Held, C., Hosie, G., Huettmann, F., Post, A., & Ropert-Coudert, Y. (Eds). (2014). Biogeographic Atlas of the Southern Ocean. Cambridge, Scientific Committee on Antarctic Research. https://epic.awi.de/43647/1/Wiencke.pdf
  10. Desvignes, T., Lauridsen, H., Valdivieso, A., Fontenele, R. S., Kraberger, S., Murray, K. N., Le François, N. R., Detrich, H. W. III, Kent, M. L., Varsani, A., & Postlethwait, J. H. (2022). A parasite outbreak in notothenioid fish in an Antarctic fjord. iScience, 25(7), 104588. https://doi.org/10.1016/j.isci.2022.104588
  11. Dillon, W. A., & Hargis Jr., W. J. (1967). Helminth parasites of Antarctic vertebrates: Part III. Monogenetic trematodes from Antarctic fishes: The superfamily Tetraonchoidea Yamaguti, 1963. In G. A. Llano, & W. L. Schmitt (Eds.), Biology of the Antarctic Seas III (Vol. 11, pp. 101–124). American Geophysical Union. https://doi.org/10.1029/AR011p0101
  12. Dillon, W. A., & Hargis Jr., W. J. (1968). Monogenetic trematodes from the southern Pacific Ocean. Part IV. Polyopisthocotyleids from New Zealand fishes: The families Mazocraeidae, Diclidophoridae and Hexabothriidae. Proceedings of the Biological Society of Washington, 81, 351–366. https://biostor.org/reference/74078
  13. Evdokimova, E. B. (1969). New species of monogeneans from bony fishes of the Patagonian Shelf. Parazitologiya, 3, 415–419. (In Russian)
  14. Froese, R., & Pauly, D. (Eds). (2025). FishBase (Version 08/2025). World Wide Web electronic publication. Retrieved August 25, 2025, from https://www.fishbase.org
  15. Gaevskaja, A., & Rodjuk, G.N. (1988). New trematode genera from the South Atlantic fishes. Parazitologiya, 22(6), 509–513. (In Russian)
  16. Gaevskaya, A. V., Rodyuk, G. N., & Parukhin, A. M. (1990). Peculiarities and formation of parasitofauna of the Patagonian toothfish Dissostichus eleginoides. Biologiya Morya, 4, 23–28. (In Russian).
  17. Gibson, D. I. (1976). Monogenea and Digenea from fishes. Discovery Reports, 36, 179–266.
  18. Gusev, A. V. (1967). Two new species of genus Gyrodactylus Nordmann (Monogenoidea) near off the coast of the Antarctic continent. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR. Issledovaniya Fauny Morey, IV(XII): Biologicheskie Otchety Sovetskoy Antarkticheskoy Ekspeditsii (1955–1958), 3, 187–189. (In Russian)
  19. Hargis Jr., W. J., & Dillon, W. A. (1967). Helminth parasites of Antarctic vertebrates: Part II. Monogenetic trematodes from Antarctic fishes: The superfamily Gyrodactyloidea Johnston and Tiegs, 1922. In G. A. Llano & W. L. Schmitt (Eds.), Biology of the Antarctic Seas, III (Vol. 11, pp. 91–99). American Geophysical Union. https://doi.org/10.1029/AR011p0091
  20. Hargis, W. J., & Dillon, W. A. (1968). Helminth parasites of Antarctic vertebrates: Part IV. Monogenetic trematodes from Antarctic fishes: The superfamily Capsaloidea Price, 1936. Proceedings of the Biological Society of Washington, 81, 403–412. https://doi.org/10.5281/zenodo.13669323
  21. Heglasová, I., Nezhybová, V., & Přikrylová, I. (2020). An amended description of two Gyrodactylus species (Platyhelminthes: Monogenea) parasitizing Antarctic Notothenioid fish. Journal of Helminthology, 94, e20. https://doi.org/10.1017/S0022149X18001098
  22. Hernández-Orts, J. S., Alama-Bermejo, G., Crespo, E. A., Raga, J. A., & Montero, F. E. (2014). A revision of the genus Neogrubea Dillon & Hargis, 1968 (Monogenea: Mazocraeidae): New morphological and molecular data from off the Patagonian coast of Argentina. Systematic Parasitology, 89(1), 59–72. https://doi.org/10.1007/s11230-014-9511-6
  23. Hoogester, J. N., & White, M. G. (1981). Notes on parasite infestation of inshore fish at Signy Island, South Orkney Islands. British Antarctic Survey Bulletin, 54, 23–31. https://nora.nerc.ac.uk/id/eprint/524681
  24. Hudson, P. J., Dobson, A. P., & Lafferty, K. D. (2006). Is a healthy ecosystem one that is rich in parasites? Trends in Ecology & Evolution, 21(7), 381–385. https://doi.org/10.1016/j.tree.2006.04.007
  25. Johnston, T. H. (1931). New trematodes from the Sub-antarctic and Antarctic. Australian Journal of Experimental Biology and Medical Science, 8(2), 91–98. https://doi.org/10.1038/icb.1931.7
  26. Kovaljova, A. A., & Gaevskaya, A. V. (1977). Two new species of monogeneans, parasites of Antarctic fishes. Zoological Journal, 56(5), 783–786. (In Russian).
  27. Kuzmina, T., Dykyy, I. V., Salganskij, O. O., Lisitsyna, O. I., Korol, E. M., & Kuzmin, Y. I. (2021). Helminth diversity in teleost fishes from the area of the Ukrainian Antarctic station “Akademik Vernadsky”, Argentine Islands, West Antarctica. Zoodiversity, 55(3), 251—264. https://doi.org/10.15407/zoo2021.03.251
  28. Kuzmina, T., Salganskiy, O., Vishnyakova, K., Ivanchikova, J., Lisitsyna, O., Korol, E., & Kuzmin, Y. (2022). Helminth diversity in teleost fishes from the South Orkney Islands region, West Antarctica. Zoodiversity, 56(2), 135–152. https://doi.org/10.15407/zoo2022.02.135
  29. Kvach, Y., & Kuzmina, T. (2020). Parasitological research in Antarctica: review of the issues and future prospects. Ukrainian Antarctic Journal, (1), 102—110. (In Ukrainian). https://doi.org/10.33275/1727-7485.1.2020.383
  30. Kvach, Y., Ondračková, M., Janáč, M., & Jurajda, P. (2016). Methodological issues affecting the study of fish parasites. I. Duration of live fish storage prior to dissection. Diseases of Aquatic Organisms,119, 107–115. https://doi.org/10.3354/dao02990
  31. Lutnicka, H., & Zdzitowiecki, K. (1984). On some problems related to the occurrence of Pseudobenedenia nototheniae Johnston, 1931 (Monogenea, Capsalidae, Trochopodinae) off the South Shetlands. Acta Ichthyologica et Piscatoria, 14(1), 141–147. https://doi.org/10.3750/AIP1984.14.1-2.09
  32. Lyadov, V. N. (1985). Zoogeographical characteristics of the helminths of fishes from the Antarctic zone of the World Ocean. In W. J. Hardis Jr. (Ed.), NOAA Technical Report NMFS 25: Parasitology and pathology of marine organisms of the World Ocean (pp. 41–43). https://repository.library.noaa.gov/view/noaa/5661
  33. MacKenzie, K. (2017). The history of Antarctic parasitological research. In S. Klimpel, T. Kuhn, & H. Mehlhorn (Eds), Biodiversity and evolution of parasitic life in the Southern Ocean. Parasitology Research Monographs (Vol. 9, pp. 13–31). Springer, Cham. https://doi.org/10.1007/978-3-319-46343-8_3
  34. Mamaev, Y. L. (1976). The system and phylogeny of monogeneans of the fam. Diclidophoridae. Trudy Biologo-Pochvennogo Instituta, 35(138), 57–80. (In Russian).
  35. Marcogliese, D. J. (2002). Food webs and the transmission of parasites to marine fish. Parasitology, 124(7), S83–S99. https://doi.org/10.1017/S003118200200149X
  36. Mintenbeck, K., Barrera-Oro, E. R., Brey, T., Jacob, U., Knust, R., Mark, F. C., Moreira, E., Strobel, A., & Arntz, W. E. (2012). 5 – Impact of climate change on fishes in complex Antarctic ecosystems. Advances in Ecological Research, 46, 351–426. https://doi.org/10.1016/B978-0-12-396992-7.00006-X
  37. Moore, J. K., Abbott, M. R., & Richman, J. G. (1999). Location and dynamics of the Antarctic Polar Front from satellite sea surface temperature data. Journal of Geophysical Research: Oceans, 104(C2), 3059–3073. https://doi.org/10.1029/1998JC900032
  38. Morley, S. A., Abele, D., Barnes, D. K. A., Cárdenas, C. A., Cotté, C., Gutt, J., Henley, S. F., Höfer, J., Hughes, K. A., Martin, S. M., Moffat, C., Raphael, M., Stammerjohn, S. E., Suckling, C. C., Tulloch, V. J. D., Waller, C. L., & Constable, A. J. (2020). Global Drivers on Southern Ocean Ecosystems: Changing Physical Environments and Anthropogenic Pressures in an Earth System. Frontiers in Marine Science, 7, 547188. https://doi.org/10.3389/fmars.2020.547188
  39. Moser, M., & Cowen, R. K. (1991). The Effects of Periodic Eutrophication on Parasitism and Stock Identification of Trematomus bernacchii (Pisces: Nototheniidae) in McMurdo Sound, Antarctica. Journal of Parasitology, 77(4), 551–556. https://doi.org/10.2307/3283158
  40. Murphy, E. J., Johnston, N. M., Hofmann, E. E., Phillips, R. A., Jackson, J. A., Constable, A. J., Henley, S. F., Melbourne-Thomas, J., Trebilco, R., Cavanagh, R. D., Tarling, G. A., Saunders, R. A., Barnes, D. K. A., Costa, D. P., Corney, S. P., Fraser, C. I., Höfer, J., Hughes, K. A., Sands, C. J., ..., & Xavier, J. C. (2021). Global connectivity of Southern Ocean ecosystems. Frontiers in Ecology and Evolution, 9, 624451. https://doi.org/10.3389/fevo.2021.624451
  41. Near, T. J. (2009). Notothenioid fishes (Notothenioidei). In S. B. Hedges, & S. Kumar (Eds), The Time-tree of Life (pp. 339–343). Oxford University Press.
  42. Oguz, M. C., Heckmann, R. A., Cheng, C. H. C., El-Naggar, A., & Tepe, Y. (2012). Ecto and endoparasites of some fishes from the Antarctic Region. Scientia Parasitologica,13(3), 119–128. https://scientia.zooparaz.net/2012_13_03/sp2012-119-128-Oguz.pdf
  43. Oğuz, M. C., Tepe, Y., Belk, M. C., Heckmann, R. A., Aslan, B., Gürgen, M., Bray, R. A., & Akgül, Ü. (2015). Metazoan Parasites of Antarctic Fishes. Turkiye Parazitoloji Dergisi, 39(2), 174–178. https://doi.org/10.5152/tpd.2015.3661
  44. Palm, H. W., Reimann, N., Spindler, M., & Plötz, J. (1998). The role of the rock cod Notothenia coriiceps Richardson, 1844 in the lifecycle of Antarctic parasites. Polar Biology, 19, 399–406. https://doi.org/10.1007/s003000050265
  45. Parukhin, A. M. (1986). Peculiarities of nototheniid fish helminth fauna in Subantarctic sector of Indian Ocean. Vestnik Zoologii, 3, 6–9. (In Russian).
  46. Parukhin, A. M., & Lyadov, V. N. (1981). Parasitofauna of Notothenioidei from waters of the Atlantic and Indian Oceans. Vestnik Zoologii, 3, 90–94. (In Russian).
  47. Parukhin, A. M., & Lyadov, V. N. (1982). Helminth fauna of food Nototheniidae fishes from Kerguelen subregion. Ekologiya Morya, 10, 49–57. (In Russian).
  48. Peck, L. S. (2018). Antarctic marine biodiversity: adaptations, environments and responses to change. In S. J. Hawkins, A. J. Evans, A. C. Dale, L. B. Firth, & I. P. Smith (Eds.), Oceanography and Marine Biology, 56. CRC Press. https://doi.org/10.1201/9780429454455
  49. Perkins, E. M., Donnellan, S. C., Bertozzi, T., Chisholm, L. A., & Whittington, I. D. (2009). Looks candeceive: Molecular phylogeny of a family of flatworm ectoparasites (Monogenea: Capsalidae) does not reflect current morphological classification. Molecular Phylogenetics and Evolution, 52(3), 705–714. https://doi.org/10.1016/j.ympev.2009.05.008
  50. Poulin, R. (1999). The functional importance of parasites in animal communities: Many roles at many levels? International Journal for Parasitology, 29(6), 903–914. https://doi.org/10.1016/S0020-7519(99)00045-4
  51. Poulin, R., & Mouritsen, K. N. (2006). Climate change, parasitism and the structure of intertidal ecosystems. Journal of Helminthology, 80(2), 183–191. https://doi.org/10.1079/JOH2006341
  52. Raffel, T. R., Martin, L. B., & Rohr, J. R. (2008). Parasites as predators: unifying natural enemy ecology. Trends in Ecology & Evolution, 23(11), 610–618. https://doi.org/10.1016/j.tree.2008.06.015
  53. Regan, C. T. (1914). II – Diagnoses of new marine fishes collected by the British Antarctic (‘Terra Nova’) expedition. The Annals and Magazine of Natural History: zoology, botany, and geology, 13(73), 11–17. https://www.biodiversitylibrary.org/page/18640770#page/23/mode/1up
  54. Rocka, A. (2006). Helminths of Antarctic fishes: Life cycle biology, specificity and geographical distribution. Acta Parasitologica, 51(1), 26–35. https://doi.org/10.2478/s11686-006-0003-y
  55. Rodjuk, G. N. (1985). Parasitic fauna of the fishes of the Atlantic part of the Antarctic (South Georgia Island and South Shetland Isles). In W. J. Hargis Jr. (Ed.), NOAA Technical Report NMFS 25: Parasitology and pathology of marine organisms of the World ocean (pp. 31–32). U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service. https://spo.nmfs.noaa.gov/Technical%20Report/tr25opt.pdf
  56. Rohde, K. (Ed.). (2005). Marine Parasitology. CSIRO Publishing. http://www.publish.csiro.au/pid/5045.htm
  57. Rohde, K., Hayward, C., & Heap, M. (1995). Aspects of the ecology of metazoan ectoparasites of marine fishes. International Journal for Parasitology, 25(8), 945–970. https://doi.org/10.1016/0020-7519(95)00015-T
  58. Rohde, K., Ho, J.-S., Smales, L., & Williams, R. (1998). Parasites of Antarctic fishes: Monogenea, Copepoda and Acanthocephala. Marine and Freshwater Research, 49(2), 121–125. https://doi.org/10.1071/MF97133
  59. Rokicka, M., Lumme, J., & Ziętara, M. S. (2009). Two new Antarctic Gyrodactylus species (Monogenoidea): description and phylogenetic characterization. Journal of Parasitology, 95(5), 1112–1119. https://doi.org/10.1645/GE-2002.1
  60. Rubtsova, N. Y., & Marcotegui, P. (2024). Redescription of Pseudobenedenia lauriei Szidat 1965 (Monogenea: Capsalidae) from Notothenia neglecta in Laurie, Peterman, and South Orkney Islands, South Antarctica. International Journal of Zoology and Animal Biology, 7(3), 000590. http://dx.doi.org/10.23880/izab-16000590
  61. Rubtsova, N. Y., Chaudhary, A., Glotov, S., & Kuzmina, T. A. (2024). Description of Pseudobenedeniella johnstoni sp. n. (Monogenea: Capsalidae) from the gills of Antarctic black rockcod, Notothenia coriiceps Richardson in coastal waters of West Antarctica. Helminthologia, 61(4), 327–344. https://doi.org/10.2478/helm-2024-0037
  62. Rubtsova, N. Y., Chaudhary, A., Glotov, S., & Kuzmina, T. A. (2026). Description of Pseudobenedenia galindezensis sp. n. (Monopisthocotyla: Capsalidae) from the skin of Notothenia coriiceps off the coastal waters of Galindez Island, West Antarctica. Systematic Parasitology, 103, 1. https://doi.org/10.1007/s11230-025-10261-9
  63. Rubtsova, N. Y., Chaudhary, A., Salganskiy, O. O., & Kuzmina, T. A. (2023). Description of Pseudobenedenia coriicepsi sp. n. (Monogenea: Capsalidae) from the Antarctic black rockcod, Notothenia coriiceps Richardson in coastal waters of West Antarctica using novel SEM images, energy dispersive X-Ray analysis and molecular analysis. International Journal of Zoology and Animal Biology, 6(5), 000512. https://doi.org/10.23880/izab-16000512
  64. Rye, C. D., Marshall, J., Kelley, M., Russell, G., Nazarenko, L. S., Kostov, Y., Schmidt, G. A., & Hansen, J. (2020). Antarctic glacial melt as a driver of recent Southern Ocean climate trends. Geophysical Research Letters, 47(11), e2019GL086892. https://doi.org/10.1029/2019GL086892
  65. Sures, B., & Nachev, M. (2022). Effects of multiple stressors in fish: how parasites and contaminants interact. Parasitology, 149(14), 1822–1828. https://doi.org/10.1017/S0031182022001172
  66. Sures, B., Nachev, M., Selbach, C., & Marcogliese, D. J. (2017). Parasite responses to pollution: what we know and where we go in ‘Environmental Parasitology’. Parasites & Vectors, 10, 65. https://doi.org/10.1186/s13071-017-2001-3
  67. Suydam, L. S. (1972). Ectoparasites of fishes of the Antarctic Peninsular Area [Degree of Masters of Arts, The College of William and Mary in Virginia]. William & Mary Institutional Repository. https://doi.org/10.25773/v5-rk58-5s65
  68. Swart, N. C., Gille, S. T., Fyfe, J. C., & Gillett, N. P. (2018). Recent Southern Ocean warming and freshening driven by greenhouse gas emissions and ozone depletion. Nature Geoscience, 11(11), 836–841. https://doi.org/10.1038/s41561-018-0226-1
  69. Szidat, L. (1965). Estudios sobre la fauna de parásitos de peces antarticos. I. Los parásitos de Notothenia neglecta Nybellin. Buenos Aires: Secretaría de Marina. Servicio de Hidrografía Naval, 910, 1–84. https://www.sidalc.net/search/Record/cat-unco-ar-55199/Description
  70. Szidat, L. (1967). Echinococcus pampeanus una nueva especie de la Argentina, parasita de Felis colocolo pajeros Desmarest, 1916 (Cestoda). Neotropica, 13(11), 90–96.
  71. Szidat, L. (1969). Structure, development, and behaviour of new strigeatoid metacercariae from subtropical fishes of South America. Journal of the Fisheries Research Board of Canada, 26(4), 753–786. https://doi.org/10.1139/f69-074
  72. Timofeeva, T. A., Gaevskaya, A. V., & Kovaleva, A. A. (1987). Capsalids of the notothenioid fishes from the Atlantic region of Antarctica and Subantarctica (Monogenea, Capsalidae). Transactions of the Zoological Institute, 161, 78–93. (In Russian).
  73. van de Lagemaat, S. H. A., Swart, M. L. A., Vaes, B., Kosters, M. E., Boschman, L. M., Burton-Johnson, A., Bijl, P. K., Spakman, W., & van Hinsbergen, D. J. J. (2021). Subduction initiation in the Scotia Sea region and opening of the Drake Passage: When and why? Earth-Science Reviews, 221, 103551. https://doi.org/10.1016/j.earsci-rev.2021.103551
  74. von Graff, L. (1884). Report of the scientific results of the voyage of H.M.S. Challenger during the years 1873–76. Zoology Part XXVII. Retrieved August 12, 2025, from https://www.19thcenturyscience.org/HMSC/HMSC-Reports/Zool-27/htm/doc.htmlvon
  75. Linstow, O. A. H. (1891). Helminthen von Süd-Georgien: Nach der Ausbeute der Deutschen Expedition von 1882–1883. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 9(2), 59–75. Retrieved August 9, 2025, from https://www.zobodat.at
  76. WoRMS Editorial Board. (2025). World Register of Marine Species. Accessed August 25, 2025, at https://www.marinespecies.org, https://doi.org/10.14284/170
  77. Zdzitowiecki, K., & Laskowski, Z. (2004). Helminths of an Antarctic fish, Notothenia coriiceps, from the Vernadsky station (Western Antarctica) in comparison with Admiralty Bay (South Shetland Islands). Helminthologia, 41(4), 201–207.