{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T01:58:19Z","timestamp":1787018299517,"version":"build-2736575974"},"reference-count":57,"publisher":"Magnolia Press","issue":"3","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zootaxa"],"abstract":"<jats:p>The Rhoptrobothriidae are one of the more enigmatic families of cestodes of elasmobranchs. Opinions on the taxonomic status of the family\u2019s three original genera (i.e., Myzophyllobothrium, Rhoptrobothrium, and Myzocephalus) have varied over the 115 years since they were erected. Some authors have considered all three valid, others have considered Rhoptrobothrium to be a synonym of Myzopyllobothrium or a genus inquirendum, yet others have considered Myzocephalus to be a synonym of the phyllobothriid genus Thysanocephalum. All three genera were established for specimens collected from eagle rays off Sri Lanka. The erection of Mixophyllobothrium for two specimens from a cowtail stingray off India three decades ago added additional confusion to the situation, with some authors considering it valid and others a synonym of Myzocephalus. These disagreements stem largely from differences in interpretation of the complex morphology of the scolex of members of these genera. Furthermore, with the exception of Rhoptrobothrium comprising four species, each genus is monotypic. All but Rhoptrobothrium has not been considered in detail for nearly a century, largely because of a lack of available material. The taxonomic status of these genera is assessed here based on light and scanning electron microscopy, and molecular data generated from new material collected from eagle rays off Indonesian and Malaysian Borneo, Japan, Sri Lanka, and Viet Nam. Morphological work indicates that the genera differ largely only in the degree of folding of the four remi that extend from the cephalic peduncle. A molecular phylogeny based on sequence data for the D1\u2013D3 region of the 28S rRNA gene, which include new data for eight specimens of four species, indicates that Myzophyllobothrium, Myzocephalus, and Rhoptrobothrium are not mutually monophyletic. The latter two genera and Mixophyllobothrium are considered synonyms of Myzophyllobothrium and five species are transferred to that genus. Myzophyllobothrium okamuri n. comb. is considered a species inquirendum. Myzophyllobothrium nagasawai n. sp. is described from Aetobatus narutobiei off Japan. Myzophyllobothrium narinari n. comb. is re-described based on newly collected cestodes from the type host and locality (i.e., Aetobatus ocellatus off Sri Lanka). Despite consisting of only a single genus, the family status of the group is retained in recognition of the unusual configuration of the scolex, which bears four biloculate bothridia and four remi extending from the cephalic peduncle. The ordinal placement of the family remains uncertain, but affinities with the Phyllobothriidea, rather than \u201cTetraphyllidea\u201d are considered.<\/jats:p>","DOI":"10.11646\/zootaxa.4999.3.1","type":"journal-article","created":{"date-parts":[[2021,7,11]],"date-time":"2021-07-11T20:46:33Z","timestamp":1626036393000},"page":"201-218","source":"Crossref","is-referenced-by-count":5,"title":["A revision of the Rhoptrobothriidae (Cestoda: \u201cTetraphyllidea\u201d)"],"prefix":"10.11646","volume":"4999","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0294-8471","authenticated-orcid":false,"given":"KIRSTEN","family":"JENSEN","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4409-8156","authenticated-orcid":false,"given":"ISABEL A. M.","family":"PEN","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9597-6978","authenticated-orcid":false,"given":"JANINE N.","family":"CAIRA","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4913","published-online":{"date-parts":[[2021,7,12]]},"reference":[{"key":"64599","unstructured":"<p>Baer, J.G. &amp; Euzet, L. (1962) Revision critique des cestodes t\u00e9traphyllides d\u00e9crits par T. Southwell (1<sup>re<\/sup> partie). <em>Bulletin de la Societe Neuchateloise des Sciences Naturelles<\/em>, 85, 143\u2013172.<\/p>"},{"key":"64600","doi-asserted-by":"crossref","unstructured":"<p>Beveridge, I. 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(2021) Emerging global novelty in phyllobothriidean tapeworms (Cestoda: Phyllobothriidea) from sharks and skates (Elasmobranchii). <em>Zoological Journal of the Linnean Society<\/em>, zlaa185.&nbsp; https:\/\/doi.org\/10.1093\/zoolinnean\/zlaa185<\/p>","DOI":"10.1093\/zoolinnean\/zlaa185"},{"key":"64603","unstructured":"<p>Caira, J.N., Jensen, K. &amp; Barbeau, E. (2020a) Global Cestode Database. Available from: http:\/\/www.elasmobranchs.tapewormdb.uconn.edu (accessed 10 January 2021)<\/p>"},{"key":"64604","doi-asserted-by":"crossref","unstructured":"<p>Caira, J.N., Jensen, K., Hayes, C. &amp; Ruhnke, T.R. (2020c) Insights from new cestodes of the crocodile shark, <em>Pseudocarcharias kamoharai <\/em>(Lamniformes: Pseudocarchariidae), prompt expansion of <em>Scyphyophyllidium<\/em> and formal synonymization of seven phyllobothriidean genera\u2014at last! <em>Journal of Helminthology<\/em>, 94, e132. https:\/\/doi.org\/10.1017\/S0022149X20000036<\/p>","DOI":"10.1017\/S0022149X20000036"},{"key":"64605","doi-asserted-by":"crossref","unstructured":"<p>Caira, J.N., Jensen, K. &amp; Healy, C.J. (1999) On the phylogenetic relationships among tetraphyllidean, lecanicephalidean and diphyllidean tapeworm genera. <em>Systematic Parasitology<\/em>, 42, 77\u2013151. https:\/\/doi.org\/10.1023\/a:1006192603349<\/p>","DOI":"10.1023\/A:1006192603349"},{"key":"64606","unstructured":"<p>Caira, J.N., Jensen, K. &amp; Healy, C.J. (2001) Interrelationships among Tetraphyllidean and Lecanicephalidean cestodes. <em>In:<\/em> Littlewood, D.T.J. &amp; Bray, R.A. (Eds.), <em>Interrelationships of the Platyhelminthes<\/em>. Taylor &amp; Francis, London, pp. 135\u2013158.<\/p>"},{"key":"64607","doi-asserted-by":"crossref","unstructured":"<p>Caira, J.N., Jensen, K., Pickering, M., Ruhnke, T.R. &amp; Gallagher, K.A. (2020b) Intrigue surrounding the life-cycles of species of <em>Clistobothrium<\/em> (Cestoda: Phyllobothriidea) parasitising large pelagic sharks. <em>International Journal for Parasitology<\/em>, 50, 1043\u20131055. https:\/\/doi.org\/10.1016\/j.ijpara.2020.08.002<\/p>","DOI":"10.1016\/j.ijpara.2020.08.002"},{"key":"64608","unstructured":"<p>Caira, J.N., Jensen, K. &amp; Ruhnke, T.R. (2017) \u201cTetraphyllidea\u201d van Beneden, 1849 relics. <em>In:<\/em> Caira, J.N. &amp; Jensen, K. (Eds.), <em>Planetary Biodiversity Inventory (2008\u20132017): Tapeworms from Vertebrate Bowels of the Earth.<\/em> <em>University of Kansas, Natural History Museum, Special Publication No. 25<\/em>. University of Kansas, Natural History Museum, Lawrence, Kansas, pp. 371\u2013400.<\/p>"},{"key":"64609","doi-asserted-by":"crossref","unstructured":"<p>Caira, J.N., Jensen, K., Waeschenbach, A., Olson, P.D. &amp; Littlewood, D.T.J. (2014) Orders out of chaos\u2014molecular phylogenetics reveals the complexity of shark and stingray tapeworm relationships. <em>International Journal for Parasitology<\/em>, 44, 55\u201373. https:\/\/doi.org\/10.1016\/j.ijpara.2013.10.004<\/p>","DOI":"10.1016\/j.ijpara.2013.10.004"},{"key":"64610","doi-asserted-by":"crossref","unstructured":"<p>Campbell, R.A. &amp; Beveridge, I. (2002) The genus <em>Acanthobothrium<\/em> (Cestoda: Tetraphyllidea: Onchobothriidae) parasitic in Australian elasmobranch fishes. <em>Invertebrate Systematics<\/em>, 16, 237\u2013344. https:\/\/doi.org\/10.1071\/it01004<\/p>","DOI":"10.1071\/IT01004"},{"key":"64611","doi-asserted-by":"crossref","unstructured":"<p>Chervy, L. (2009) Unified terminology for cestode microtriches: A proposal from the International Workshops on Cestode Systematics in 2002\u20132008. <em>Folia Parasitologica<\/em>, 56, 199\u2013230. https:\/\/doi.org\/10.14411\/fp.2009.025<\/p>","DOI":"10.14411\/fp.2009.025"},{"key":"64612","doi-asserted-by":"crossref","unstructured":"<p>Coleman, G.M., Beveridge, I. &amp; Campbell, R.A. (2019) <em>Caulobothrium pedunculatum<\/em> sp. nov., a new species of cestode (Platyhelminthes) parasitic in Australian stingrays (Elasmobranchii: Batoidea). <em>Transactions of the Royal Society of South Australia<\/em>, 143, 167\u2013174. https:\/\/doi.org\/10.1080\/03721426.2019.1624932<\/p>","DOI":"10.1080\/03721426.2019.1624932"},{"key":"64613","doi-asserted-by":"crossref","unstructured":"<p>Darriba, D., Taboada, G.L., Doallo, R. &amp; Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. <em>Nature Methods<\/em>, 9, 772. https:\/\/doi.org\/10.1038\/nmeth.2109<\/p>","DOI":"10.1038\/nmeth.2109"},{"key":"64614","unstructured":"<p>Deshmukh, R.A. (1979) On three new species of <em>Uncibilocularis<\/em> Southwell, 1925 (Cestoda: Onchobothriidae) from marine fishes with a key to the species of the genus. <em>Proceedings of the National Academy of Sciences, India<\/em>, 49 (B), 227\u2013236.<\/p>"},{"key":"64615","unstructured":"<p>Deshmukh, R.A. &amp; Shinde, G.B. (1979) Three new species of <em>Tetragonocephalum<\/em> Shipley and Hornell, 1905 (Cestoda: Tetragonocephalidae) from marine fishes of west coast of India. <em>Bioresearch<\/em>, 3, 19\u201323.<\/p>"},{"key":"64616","unstructured":"<p>Euzet, L. (1994) Order Tetraphyllidea Carus, 1863. <em>In:<\/em> Khalil, L.F., Jones, A. &amp; Bray, R.A. (Eds.), <em>Keys to the Cestode Parasites of Vertebrates<\/em>. CAB International, Wallingford, pp. 149\u2013194.<\/p>"},{"key":"64617","doi-asserted-by":"crossref","unstructured":"<p>Fernando, D., Bown, R.M.K., Tanna, A., Gobiraj, R., Ralicki, H., Jockusch, E.L., Ebert, D.A., Jensen, K. &amp; Caira, J.N. (2019) New insights into the identities of the elasmobranch fauna of Sri Lanka. <em>Zootaxa<\/em>, 4585 (2), 201\u2013238. https:\/\/doi.org\/10.11646\/zootaxa.4585.2.1<\/p>","DOI":"10.11646\/zootaxa.4585.2.1"},{"key":"64618","doi-asserted-by":"crossref","unstructured":"<p>Guindon, S. &amp; Gascuel, O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. <em>Systematic Biology<\/em>, 52, 696\u2013704. https:\/\/doi.org\/10.1080\/10635150390235520<\/p>","DOI":"10.1080\/10635150390235520"},{"key":"64619","doi-asserted-by":"crossref","unstructured":"<p>Hahn, C., Bachmann, L. &amp; Chevreux, B. (2013) Reconstructing mitochondrial genomes directly from genomic next-generation sequencing reads\u2014a baiting and iterative mapping approach. <em>Nucleic Acids Research<\/em>, 41, e129. https:\/\/doi.org\/10.1093\/nar\/gkt371<\/p>","DOI":"10.1093\/nar\/gkt371"},{"key":"64620","doi-asserted-by":"crossref","unstructured":"<p>Haseli, M. (2013) Trypanorhynch cestodes from elasmobranchs from the Gulf of Oman, with the description of <em>Prochristianella garshaspi <\/em>n. sp. (Eutetrarhynchidae). <em>Systematic Parasitology<\/em>, 85, 271\u2013279. https:\/\/doi.org\/10.1007\/s11230-013-9425-8<\/p>","DOI":"10.1007\/s11230-013-9425-8"},{"key":"64621","doi-asserted-by":"crossref","unstructured":"<p>Haseli, M. &amp; Palm, H.W. (2015) <em>Dollfusiella<\/em> <em>qeshmiensis<\/em> n. sp (Cestoda: Trypanorhyncha) from the cowtail stingray <em>Pastinachus sephen<\/em> (Forssk\u00e5l) in the Persian Gulf, with a key to the species of <em>Dollfusiella<\/em> Campbell &amp; Beveridge, 1994. <em>Systematic Parasitology<\/em>, 92, 161\u2013169. https:\/\/doi.org\/10.1007\/s11230-015-9592-x<\/p>","DOI":"10.1007\/s11230-015-9592-x"},{"key":"64622","unstructured":"<p>Hassan, S.H. (1983) <em>Acanthobothrium manteri <\/em>sp. n. a tetraphyllidean cestode (Oncobothriidae [sic]) from <em>Dasyatis sephen<\/em>. <em>Journal of the Egyptian Society of Parasitology<\/em>, 13, 75\u201380.<\/p>"},{"key":"64623","doi-asserted-by":"crossref","unstructured":"<p>Hornell, J. (1912) New cestodes from Indian fishes. <em>Records of the Indian Museum<\/em>, 7, 197\u2013204.<\/p>","DOI":"10.5962\/bhl.part.28231"},{"key":"64624","unstructured":"<p>ICZN. (1999) <em>International Code of Zoological Nomenclature, 4th edition<\/em>. The International Trust for Zoological Nomenclature, London, 306 pp.<\/p>"},{"key":"64625","doi-asserted-by":"crossref","unstructured":"<p>Jensen, K. &amp; Caira, J.N. (2006) The status of <em>Rhoptrobothrium<\/em> Shipley et Hornell, 1906 (Cestoda: Tetraphyllidea), with redescription of the type species, <em>R. myliobatidis<\/em>, and description of three new species from two species of <em>Aetomylaeus<\/em> (Myliobatiformes: Myliobatidae) from Malaysian Borneo. <em>Folia Parasitologica<\/em>, 53, 189\u2013207. https:\/\/doi.org\/10.14411\/fp.2006.025<\/p>","DOI":"10.14411\/fp.2006.025"},{"key":"64626","doi-asserted-by":"crossref","unstructured":"<p>Jensen, K. &amp; Caira, J.N. (2008) A revision of <em>Uncibilocularis<\/em> Southwell, 1925 (Tetraphyllidea: Onchobothriidae) with the description of four new species. <em>Comparative Parasitology<\/em>, 75, 157\u2013173. https:\/\/doi.org\/10.1654\/4347.1<\/p>","DOI":"10.1654\/4347.1"},{"key":"64627","unstructured":"<p>Joyeux, C. &amp; Baer, J.G. (1961) Classe de Cestodes. Cestoidea Rudolphi. <em>In:<\/em> Baer, J.-G., de Beauchamp, P., Caullery, M., Euzet, L., Gontcharoff, M., Grass, P.-P. &amp; Joyeux, C. (Eds.), <em>Trait de Zoologie. Anatomie, syst\u00e9matique, biologie. Tome IV. Plathelminthes, M\u00e9sozoaires, Acanthoc\u00e9phales, N\u00e9mertiens<\/em>. Masson et C<sup>ie<\/sup>, Paris, pp. 347\u2013560.<\/p>"},{"key":"64628","unstructured":"<p>Khambata, F.S. &amp; Bal, D.V. (1954) Studies on the cestodes of marine fishes from Bombay. 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(1892) The anatomy of <em>Thysanocephalum crispum<\/em>, Linton, a parasite of the tiger shark. <em>Report of the United States Commissioner of Fish and Fisheries for 1888<\/em>, 1892, 543\u2013556.<\/p>"},{"key":"64632","doi-asserted-by":"crossref","unstructured":"<p>L\u00f6ytynoja, A. &amp; Goldman, N. (2010) webPRANK: a phylogeny-aware multiple sequence aligner with interactive alignment browser. <em>BMC Bioinformatics<\/em>, 11, 579. https:\/\/doi.org\/10.1186\/1471-2105-11-579<\/p>","DOI":"10.1186\/1471-2105-11-579"},{"key":"64633","doi-asserted-by":"crossref","unstructured":"<p>Maleki, L., Malek, M. &amp; Palm, H.W. (2013) Two new species of <em>Acanthobothrium<\/em> (Tetraphyllidea: Onchobothriidae) from <em>Pastinachus<\/em> cf. <em>sephen<\/em> (Myliobatiformes: Dasyatidae) from the Persian Gulf and Gulf of Oman. <em>Folia Parasitologica<\/em>, 60, 448\u2013456. https:\/\/doi.org\/10.14411\/fp.2013.048<\/p>","DOI":"10.14411\/fp.2013.048"},{"key":"64634","unstructured":"<p>Maleki, L., Malek, M. &amp; Rastgoo, A. (2018) <em>Acanthobothrium chabahariensis<\/em> n. sp. (Cestoda: Onchoproteocephalidea) in the cowtail stingray <em>Pastinachus <\/em>cf. <em>sephen<\/em> (Myliobatiformes: Dasyatidae) from the Gulf of Oman, Iran. <em>Journal of Genetic Resources<\/em>, 4, 114\u2013121. https:\/\/doi.org\/10.22080\/jgr.2019.15565.1120<\/p>"},{"key":"64635","doi-asserted-by":"crossref","unstructured":"<p>Naylor, G.J.P., Caira, J.N., Jensen, K., Rosana, K.A.M., White, W.T. &amp; Last, P.R. 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A plea for the use of vouchers in molecular phylogenetic studies. <em>Molecular Phylogenetics and Evolution<\/em>, 48, 369\u2013371. https:\/\/doi.org\/10.1016\/j.ympev.2008.03.024<\/p>","DOI":"10.1016\/j.ympev.2008.03.024"},{"key":"64638","doi-asserted-by":"crossref","unstructured":"<p>Reyda, F., Healy, C., Haslach, A., Ruhnke, T., Aprill, T., Bergman, M., Daigler, A., Dedrick, E., Delgado, I., Forti, K., Herzog, K., Russell, R. &amp; Willsey, D. (2016) A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations. <em>Folia Parasitologica<\/em>, 63, 038. https:\/\/doi.org\/10.14411\/fp.2016.038<\/p>","DOI":"10.14411\/fp.2016.038"},{"key":"64639","doi-asserted-by":"crossref","unstructured":"<p>Roohi Aminjan, A. &amp; Malek, M. (2017) Two new species of <em>Tetragonocephalum<\/em> (Cestoda: Lecanicephalidea) from <em>Pastinachus sephen<\/em> (Myliobatiformes: Dasyatidae) from the Gulf of Oman. <em>Folia Parasitologica<\/em>, 64, 014. https:\/\/doi.org\/10.14411\/fp.2017.014<\/p>","DOI":"10.14411\/fp.2017.014"},{"key":"64640","unstructured":"<p>Ruhnke, T.R., Caira, J.N. &amp; Pickering, M. (2017) Phyllobothriidea Caira, Jensen, Waeschenbach, Olson &amp; Littlewood, 2014. <em>In:<\/em> Caira, J.N. &amp; Jensen, K. (Eds.), <em>Planetary Biodiversity Inventory (2008\u20132017): Tapeworms from Vertebrate Bowels of the Earth.<\/em> <em>University of Kansas, Natural History Museum, Special Publication No. 25<\/em>. University of Kansas, Natural History Museum, Lawrence, Kansas, pp. 305\u2013326.<\/p>"},{"key":"64641","doi-asserted-by":"crossref","unstructured":"<p>Schaeffner, B.C. &amp; Beveridge, I. 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