{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T03:08:30Z","timestamp":1787022510074,"version":"build-2736575974"},"reference-count":49,"publisher":"Springer Science and Business Media LLC","issue":"4","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2015,1,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>There are three major groups of parasitic platyhelminths (flatworms). The digeneans and cestodes are endoparasites, while the monogeneans are ectoparasites mostly on the gills or skin of fishes. Monogeneans are hermaphrodite and, with the exception of the gyrodactylids, mostly protandrous, the male reproductive system maturing before the female system. Their ectoparasitic life-style provides unique opportunities to observe the reproductive biology of living platyhelminths, opportunities restricted in digeneans and cestodes by their endoparasitic habits. Moreover, the male copulatory organs (MCOs) of monogeneans are of special interest because of their perplexing diversity, ranging from sclerotised penis tubes, many with accessory sclerites, to cirruses and genital atrium armature (hooks and spines). The relatively few accounts in the literature of mating in monogeneans are reproduced in this review, together with consideration of the following aspects of sperm transfer: structure and function of MCOs; self-insemination; spermatophores and pseudospermatophores; \u201chypodermic\u201d and transtegumental insemination; tissue fusion; glands associated with MCOs and vaginae; finding a mating partner.<\/jats:p>","DOI":"10.1515\/ap-2015-0082","type":"journal-article","created":{"date-parts":[[2015,6,16]],"date-time":"2015-06-16T13:39:53Z","timestamp":1434461993000},"source":"Crossref","is-referenced-by-count":11,"title":["Sperm transfer in monogenean (platyhelminth) parasites"],"prefix":"10.1007","volume":"60","author":[{"given":"Graham","family":"Kearn","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ian","family":"Whittington","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","reference":[{"key":"ref141","first-page":"102","article-title":"Swimming in Gyrodactylus rysavyi from the Nile catfish Clarias gariepinus","volume":"49","author":"El","year":"2004","journal-title":"Acta Parasitologica"},{"key":"ref711","doi-asserted-by":"publisher","first-page":"915","DOI":"10.1017\/S0025315400071320","article-title":"The biology of Atrispinum labracis n comb on the gills of the bass Dicentrachus labrax of the Marine Biological Association of the United Kingdom","volume":"63","author":"Winch","year":"1983","journal-title":"Journal"},{"key":"ref291","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/0020-7519(84)90013-4","article-title":"The migration of the monogenean Entobdella soleae on the surface of its host Solea solea for","volume":"14","author":"Kearn","year":"1984","journal-title":"International Journal Parasitology"},{"key":"ref551","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1016\/j.parint.2013.10.009","article-title":"Insemination of the monogenean Neobenedenia girellae","volume":"63","author":"Ogawa","year":"2014","journal-title":"Parasitology International"},{"key":"ref31","doi-asserted-by":"publisher","first-page":"590","DOI":"10.2307\/2420295","article-title":"Ectoparasitic trematodes of Puget Sound fishes Acanthocotyle","volume":"20","author":"Bonham","year":"1938","journal-title":"American Midland Naturalist"},{"key":"ref461","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1098\/rspb.1983.0083","article-title":"Sperm transfer in the monogenean gill parasite Gastrocotyle trachuri of the of","author":"Llewellyn","year":"1983","journal-title":"Proceedings Royal Society London"},{"key":"ref651","doi-asserted-by":"publisher","first-page":"1113","DOI":"10.1080\/00222939600770611","article-title":"A revision of Neobenedenia Yamaguti including a redescription of N melleni Yamaguti Natural","volume":"30","author":"Whittington","year":"1996","journal-title":"Journal of History"},{"key":"ref521","doi-asserted-by":"publisher","first-page":"613","DOI":"10.2307\/3279944","article-title":"Studies on the reproductive system of Philophthalmus gralli using techniques of transplantation and autoradiography of","volume":"64","author":"Nollen","year":"1978","journal-title":"Journal 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Australia","volume":"53","author":"Whittington","year":"2008","journal-title":"Acta Parasitologica"},{"key":"ref51","article-title":"Monogenetic Trematodes their Classification and Phylogeny of Leningrad","author":"Bychowsky","year":"1957","journal-title":"Academy Sciences"},{"key":"ref331","doi-asserted-by":"publisher","first-page":"1227","DOI":"10.1016\/0020-7519(94)90193-7","article-title":"Evolutionary expansion of the Monogenea for","volume":"24","author":"Kearn","year":"1994","journal-title":"International Journal Parasitology"},{"key":"ref271","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1017\/S0031182000069183","article-title":"The biology of Leptocotyle minor a skin parasite of the dogfish Scyliorhinus canicula","volume":"55","author":"Kearn","year":"1965","journal-title":"Parasitology"},{"key":"ref481","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1017\/S0031182000067962","article-title":"Spermatophores in the monogenean Entobdella diadema Monticelli from the skin of sting - rays with a note on the taxonomy of the parasite","volume":"54","author":"Llewellyn","year":"1964","journal-title":"Parasitology"},{"key":"ref511","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1017\/S0025315400024139","article-title":"Reproduction in Acanthocotyle greeni n sp from the skin of Raia spp at Plymouth of the Marine Biological Association of the United Kingdom","volume":"60","author":"Macdonald","year":"1980","journal-title":"Journal"},{"key":"ref81","doi-asserted-by":"publisher","first-page":"1169","DOI":"10.1016\/S0020-7519(96)00113-0","article-title":"A revision of Heterocotyle with a description of Heterocotyle capricornensis n sp from Himantura fai from Heron Island Great Barrier Reef Australia for","volume":"26","author":"Chisholm","year":"1996","journal-title":"International Journal Parasitology"},{"key":"ref411","doi-asserted-by":"publisher","first-page":"115","DOI":"10.2478\/s11686-010-0019-1","article-title":"A new genus and new species of microbothriid monogenean with a functionally enigmatic reproductive system parasitic on the skin and mouth lining of the largetooth sawfish Pristis microdon in Australia","volume":"55","author":"Kearn","year":"2010","journal-title":"Acta Parasitologica"},{"key":"ref111","first-page":"2348","article-title":"Three new species of Benedenia Diesing from the Great Barrier Reef Australia with a key to species of the genus","volume":"1858","author":"Deveney","year":"2010","journal-title":"Zootaxa"},{"key":"ref281","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1017\/S0031182000078136","article-title":"The production transfer and assimilation of spermatophores by Entobdella soleae a monogenean skin parasite of the common sole","volume":"60","author":"Kearn","year":"1970","journal-title":"Parasitology"},{"key":"ref701","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1007\/BF00006905","article-title":"Benedenia rohdei n sp from the gills of Lutjanus 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Parasitology"},{"key":"ref221","doi-asserted-by":"crossref","first-page":"71","DOI":"10.5962\/bhl.part.91371","article-title":"Bicotylophora baeri n sp parasite branchial du teleosteen Trachinotus falcatus en Cote - d Ivoire de","volume":"84","author":"Euzet","year":"1977","journal-title":"Revue Suisse Zoologie"},{"key":"ref421","first-page":"371","article-title":"Spermatophores in Dermopristis cairae Whittington et Kearn","volume":"56","author":"Kearn","year":"2011","journal-title":"Acta Parasitologica"},{"key":"ref241","doi-asserted-by":"publisher","first-page":"998","DOI":"10.2307\/3283449","article-title":"A new species of Metabenedeniella from the dorsal fin of Diagramma pictum from the Great Barrier Reef Australia with a revision of the genus of","volume":"80","author":"Horton","year":"1994","journal-title":"Journal Parasitology"},{"key":"ref151","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/S0020-7519(83)80067-8","article-title":"Glands associated with the anterior adhesive areas and body margins in the skin - parasitic monogenean Entobdella soleae for","volume":"13","author":"El","year":"1983","journal-title":"International Journal Parasitology"},{"key":"ref591","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1017\/S0031182000070979","article-title":"The functional morphology of the genitalia of the spermatophore - producing monogenean parasite Diplectanum aequans Diesing with a note on the copulation of the parasite","volume":"56","author":"Paling","year":"1966","journal-title":"Parasitology"},{"key":"ref261","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/0889-1605(85)90126-0","article-title":"The aflagellate spermatozoon of Diplozoon : a demonstrative case of relationship between sperm ultrastructure and biology of reproduction of","volume":"92","author":"Justine","year":"1985","journal-title":"Journal Ultrastructure 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Parasitology"},{"key":"ref691","doi-asserted-by":"publisher","first-page":"29","DOI":"10.14411\/fp.2009.005","article-title":"Two new species of Neoentobdella from the skin of Australian stingrays","volume":"56","author":"Whittington","year":"2009","journal-title":"Folia Parasitologica"},{"key":"ref21","doi-asserted-by":"publisher","first-page":"616","DOI":"10.2307\/3283862","article-title":"Clemacotyle australis from the white - spotted eagle ray Aetobatus narinari on the Great Barrier Reef : redescription emended generic diagnosis and oncomiracidium of","volume":"81","author":"Beverley","year":"1995","journal-title":"Journal Parasitology"},{"key":"ref91","doi-asserted-by":"publisher","first-page":"1041","DOI":"10.1016\/S0020-7519(97)00072-6","article-title":"A revision of Neoheterocotyle with descriptions of the larvae of N rhinobatis and N rhynchobatis from Heron Island Great Barrier Reef Australia for","volume":"27","author":"Chisholm","year":"1997","journal-title":"International 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labrosus","volume":"88","author":"Llewellyn","year":"1984","journal-title":"Parasitology"},{"key":"ref11","doi-asserted-by":"publisher","first-page":"965","DOI":"10.1016\/0020-7519(87)90016-6","article-title":"Generic diagnosis of Asthenocotyle Robinson and description of Asthenocotyle taranakiensis new species from Oxynotus bruniensis taken in New Zealand waters for","volume":"17","author":"Beverley","year":"1987","journal-title":"International Journal Parasitology"},{"key":"ref631","first-page":"2519","article-title":"Revision of Benedeniella Johnston its assignment to Entobdellinae Bychowsky and comments on subfamilial composition","author":"Whittington","year":"2010","journal-title":"Zootaxa"}],"container-title":["Acta 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