Abstract
The Capsalinae Baird, 1853 (Monogenea: Capsalidae) is revised based on a thorough review of original descriptions and examination of type museum material, where available, to validate species. A total of 262 type and voucher specimens was studied representing apparently 42 of the 60 currently described capsaline species. A combination of characters that should be independent of variation due to specimen preparation techniques was chosen to discriminate species. These characters include the presence/absence of papillae on the ventral surface of the haptor, the presence/absence and the morphology of haptoral accessory sclerites and the presence/absence of dorsomarginal body sclerites and their morphology and distribution. We consider that only 36 of the 60 nominal capsaline species are valid. We could find no support for Caballerocotyla Price, 1960 and therefore we synonymise it with Capsala Bosc, 1811. Under the current concept we recognise 22 species of Capsala, 7 species of Capsaloides Price, 1938, 3 species of Nasicola Yamaguti, 1968 and 4 species of Tristoma Cuvier, 1817. The following Capsala species are considered valid: C. albsmithi (Dollfus, 1962) n. comb.; C. biparasitica (Goto, 1894) Price, 1938; C. caballeroi Winter, 1955; C. foliacea (Goto, 1894) Price, 1938; C. gouri Chauhan, 1951; C. gregalis (Wagner & Carter, 1967) n. comb.; C. interrupta (Monticelli, 1891) Johnston, 1929; C. katsuwoni (Ishii, 1936) Price, 1938; C. laevis (Verrill, 1875) Johnston, 1929; C. maccallumi Price, 1939; C. magronum (Ishii, 1936) Price, 1938; C. manteri Price, 1951; C. manteriaffinis (Mamaev, 1968) n. comb.; C. martinierei Bosc, 1811; C. notosinense (Mamaev, 1968) n. comb.; C. nozawae (Goto, 1894) Price, 1938; C. onchidiocotyle (Setti, 1899) Johnston, 1929; C. ovalis (Goto, 1894) Price, 1938; C. paucispinosa (Mamaev, 1968) n. comb.; C. pelamydis (Taschenberg, 1878) Price, 1938; C. poeyi (Pérez-Vigueras, 1935) Price, 1938; C. pricei Hildago-Escalente, 1950. We consider the following Capsaloides species valid: C. cornutus (Verrill, 1875) Price, 1938; C. cristatus Yamaguti, 1968; C. hoffmannae Lamothe-Argumedo, 1996; C. magnaspinosus Price, 1939; C. nairagi Yamaguti, 1968; C. perugiai (Setti, 1898) Price, 1938; C. sinuatus (Goto, 1894) Price, 1938. The following Nasicola species are deemed valid: N. brasiliensis Kohn, Baptista-Farias, Santos & Gibson, 2004; N. hogansi Wheeler & Beverley-Burton, 1987; N. klawei Stunkard, 1962. Presently, we consider the following Tristoma species valid: T. adcoccineum Yamaguti, 1968; T. adintegrum Yamaguti, 1968; T. coccineum Cuvier, 1817; T. integrum Diesing, 1850. A list of new and re-established synonyms is provided. The status of each species is discussed in detail and a key to all capsaline species that we consider valid is presented. The following 5 capsaline species are considered to be species inquirendae: Caballerocotyla phillippina Velasquez, 1982; Capsala megacotyle (Linstow, 1906) Johnston, 1929; Tristoma fuhrmanni Guiart, 1938; T. levinsenii Monticelli, 1891; T. uncinatum Monticelli, 1889. The importance of careful character selection to discriminate between capsaline species and the need for studies of live parasites to obtain additional characters based on reproductive structures is addressed. Hostspecificity in the Capsalinae is also discussed.References
Brinkmann, A. Jr. (1952) Fish trematodes from Norwegian waters. Universitetet Bergen Årbok, Naturvitenskapelig rekke Nr, 1, 1–134.
Bussiéras, J. (1972) Les Monogènes Capsalinae parasites des thons de l’Atlantique tropical oriental. Annales de Parasitologie (Paris), 47, 29–49.
Bussiéras, J. & Aldrin, J.-F. (1967) Caballerocotyla klawei Stunkard 1962, Monogène parasite des sac nasaux du thon albacore de l’Océan Atlantique. Revue d’Élevage et de Médicine Vétérinaire des Pays Tropicaux, 20, 105–108.
Bussiéras, J. & Baudin-Laurencin, F. (1970) Caballerocotyla abidjani n. sp. (Monogenea: Capsalidae), parasite des opercules du thon albacore, Thunnus albacares. Cahiers Office de la Recherche Scientifique et Technique Outre, Série Océanographie, 3, 47–51.
Bussiéras, J. & Baudin-Laurencin, F. (1973) Les helminthes parasites des thons tropicaux. Revue d’Élevage et de Médicine Vétérinaire des Pays Tropicaux, 26, 13a–19a.
Chauhan, B.S. (1951) Trematodes from Indian marine fishes. Part VII. On monogenetic parasites of the family Capsalidae Baird, 1853 (Capsaloidea) from Indian region, with description of a new species of the genus Capsala Bosc, 1811. Records of the Indian Museum, 49, 45–54.
Chisholm, L.A. & Whittington, I.D. (2006) Revision of Capsaloides (Monogenea: Capsalidae) with a redescription of C. magnaspinosus Price, 1939 from the nasal tissue of Tetrapterus audax (Istiophoridae) collected off Nelson Bay, New South Wales, Australia. Zootaxa, 1160, 1–20.
Chisholm, L.A. & Whittington, I.D. (2007) The oncomiracidium of Nasicola klawei (Monogenea: Capsalidae: Capsalinae). Parasitology Research, 100, 467–471.
Egorova, T.P. (1989) A taxonomic analysis of the subfamily Capsalinae (Monogenoidea, Capsalidae). In: Lebedev, B.I. (Ed.), Parazitologicheskie issledovaniya: sbornik nauchnykh trudov. Dal’nevostochneo otdelenie. Akademiya Nauk SSSR, Vladivostok, pp. 46–54 (In Russian).
Froese, R. & Pauly, D. (2007) FishBase. Version 02/2007. Available from: http://www.fishbase.org (accessed April 5, 2007).
Fuhrmann, O. (1928) Zweite klasse des Cladus Platyhelminthes: Trematoda. In: Kűkenthal’s Handbuch der Zoologie. Berlin und Leipzig, pp. 1–140.
Goto, S. (1894) Studies on the ectoparasitic trematodes of Japan. Journal of the College of Science, Imperial University of Tokyo, 8, 1–273.
Goto, S. (1899) Notes on some exotic species of ectoparasitic trematodes. Journal of the College of Science, Imperial University of Tokyo, 12, 263–295.
Guiart, J. (1938) Trematodes parasites provenant des Campagnes scientifiques du Prince Albert ler de Monaco (1886-1912). Resultats des Campagnes Scientifiques du Prince de Monaco, 100, 1–84.
Ishii, N. (1936) Some new ectoparasitic trematodes of marine fishes. Zoological Magazine, 48, 781–790.
Kardousha, M.M. (2002) Monogenea of Arabian Gulf fishes 1. Descriptions of three Capsala spp. (Capsalidae) including Capsala naffari n. sp. infecting mackerel tuna Euthynnus affinis from coasts of Emirates. Parasitology International, 51, 327–335.
Kohn, A., Baptista-Farias, M.F., dos Santos, A.L. & Gibson, D.I. (2004) A new species of Nasicola Yamaguti, 1968 (Monogenea: Capsalidae) from the nasal cavities of Thunnus obesus and a redescription of N. klawei (Stunkard, 1962) from T. albacares off Brazil. Systematic Parasitology, 57, 51–58.
Kohn, A. & Justo, M.C.N. (2006) Caballerocotyla llewelyni n. sp. and Caballerocotyla neothunni (Yamaguti, 1968) (Monogenea: Capsalidae) parasites of Brazilian tunas (Scombridae). Zootaxa, 1139, 19–26.
Lamothe-Argumedo, R. (1997) Nuevo arreglo taxonómico de la subfamilia Capsalinae (Monogenea: Capsalinae), clave para los géneros y dos combinaciones nuevas. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, Serie Zoologia, 68, 207–223.
Lawler, A.R. (1981) Zoogeography and host-specificity of the superfamily Capsaloidea Price, 1936 (Monogenea: Monopisthocotylea). An evaluation of the host-parasite records of the superfamily Capsaloidea Price, 1936, and their utility in determinations of host-specificity and zoogeography. Special Publications in Marine Science 6, Virginia Institute of Marine Science, Virginia, U.S.A, 650 pp.
Linstow, von O. (1906) Helminths from the collection of the Colombo museum. Spoilia Seylanica, 3, 163–188.
Mamaev, Yu L. (1968) Helminths of tuna fish in the South China Sea. In: Skryabin, K. I. & Mamaev, Yu L. (Eds), Helminths of Animals of the Pacific Ocean. Moscow: Izdat. Nauka, pp. 5–27. (In Russian).
Monticelli, F.S. (1889) Tristomum uncinatum n. sp. Bolletino della Società dei Naturalisti in Napoli 1, Ser 3(2), 117–119.
Monticelli, F.S. (1891) Di alcuni organi di tattoo nei tristomidi. Contributo allo studio dei trematodi monogenetici Parte I. Bolletino della Società dei Naturalisti in Napoli (1891) 1, 5, 99–134.
Mogrovejo, C.D. & Santos, C.P. (2002) Caballerocotyla lenti n. sp., a capsalid monogenean from Auxis thazard (Scombridae) from off the southeastern coast of Brazil. Memorias do Instituto Oswaldo Cruz, 97, 1067–1074.
Murugesh, M. (1995) Monogenetic trematodes from scombrid fishes of the Visakhapatnam coast, Bay of Bengal. Journal of Natural History, 29, 1–26.
Oliva, M. (1986) Monogenea in marine fishes from Antofagasta, Chile, with description of Caballerocotyla australis n. sp. (Capsalidae). Revista Chilena de Historia Natural, 59, 87–94.
Palombi, A. (1949) I trematodi d'Italia. Parte I. Trematodi monogenetici. Archivio Zoologico Italiano, 34, 203–408.
Pérez-Vigueras, I. (1935) Tristomum poeyi n. sp. (Trematoda), parasito de Makaira ampla Poey (Pisces). Memorias de la Sociedad Cubana de Historia Natural " Felipe Poey", 9, 43–44.
Perugia, A. & Parona, C. (1890) Di alcuni Trematodi ectoparassiti di pesci Adriatici. Annali di Museo Civico di Storia Naturale Giacomo Doria, 29, 16–32.
Price, E.W. (1938) The monogenetic trematodes of Latin America. In: Livro Jubilar Prof. Travassos. Rio de Janerio, Brazil, pp. 407–413.
Price, E.W. (1939) North American monogenetic trematodes III. The family Capsalidae (Capsaloidea). Journal of the Washington Academy of Sciences, 29, 63–92.
Price, E.W. (1960) The giant marlin, Makiara marlina Jordan and Evermann, a new host for Capsala pricei Hidalgo, 1959, with a review of the subfamily Capsalinae. In: Libro Homenaje al Doctor Eduardo Caballero y Caballero, Jubileo 1930-1960. Instituto Politécnico Nacional Escuela Nacional de Ciencias Biológicas México, pp. 237–244.
Price, E.W. (1962a) What is Capsala martinieri Bosc 1811? Proceedings of the 1st All-India Congress of Zoology (1959) Part 2, Section 9, 440–445.
Price, E.W. (1962b) A description of Tricotyla molae (Blanchard), with a discussion of the monogenetic trematodes of the sunfish (Mola mola L.). Journal of Parasitology, 48, 748–751.
Price, E.W. (1963) Redescriptions of some species of Caballerocotyla Price (Monogenea; Capsalidae). Proceedings of the Helminthological Society of Washington, 30, 149–154.
Setti, E. (1899) Secondo contributo per una revisione dei Tristomi e descrizione di una nuova specie. Atti della Società Ligustica di Scienze Naturali e Geografiche, 10, 71–84; 117–125.
Sproston, N. (1946) A synopsis of the monogenetic trematodes. Transactions of the Zoological Society of London, 25, 185–600.
Sproston, N. (1969) A contribution to the ecology of the Capsala on the southern bluefin tuna (Thunnus maccoyi). Parazitologicheskii sbornik, 24, 242–243.
Stunkard, H.W. (1962) Caballerocotyla klawei sp. n., a monogenetic trematode from the nasal capsule of Neothunnus macropterus. Journal of Parasitology, 48, 883–890.
Taschenberg, E.O. (1878) Ueber die Geschlechtorgane ectoparasitische mariner Trematoden. Vorlaugfige Mittheilung. Zoologische Anzeiger, 1, 1–176.
Velasquez, C.C. (1982) Monogenea (Capsaloidea) from Philippine marine fishes. Proceedings of the Helminthogical Society of Washington, 49, 176–184.
Verrill, A.E. (1875) Brief contributions to zoology from the museum of Yale College. 33. Results of dredging expeditions off the New England coast in 1874. American Journal of Science and Arts, 110, 36–43.
Verrill, A.E. (1885) Results of explorations made by the steamer “Albatross” of the northern coast of the United States, in 1883. Report of the United States Fish Commission, Washington, 11, 503–699.
Wagner, E.D. & Carter, C.E. (1967) Caballerocotyla gregalis sp. n. (Trematoda: Monogenea) from the gills of Sarda lineolata (Girard). Journal of Parasitology, 53, 277–279.
Whittington, I.D. (2004) The Capsalidae (Monogenea:Monopisthocotylea): a review of diversity, classification and phylogeny with a note about species complexes. Folia Parasitologica, 51, 109–122.
Winter, H.A. (1955) Capsala caballeroi sp. n., parásito de Sarda orientalis, con un catálogo de los tremátodos monogéneos de los peces del Océano Pacífico de las Américas. Revista Brasileira de Biologia, 15, 9–32.
Yamaguti, S. (1963) Systema Helminthum. Volume IV. Monogenea and Aspidocotylea. Interscience Publishers, New York, 699 pp.
Yamaguti, S. (1968) Monogenetic Trematodes of Hawaiian Fishes. University of Hawaii Press, Hawaii, 287 pp.