Abstract
Helicometra Odhner, 1902 (Digenea: Opecoelidae) is one of the largest genera within Plagioporinae Manter, 1947. All 34 nominal species possess a single polar filament on the egg, a unique, conspicuous helical uterus, and they parasitize the intestinal tract of 75 families, 172 genera, and 296 species of mostly marine piscine hosts from shallow to deep waters and from equatorial to polar latitudes. This study describes a new species of Helicometra from the cusk-eel Luciobrotula corethromycter Cohen, 1964 (Ophidiiformes: Ophidiidae) collected from 1,280 m depth in the northwestern Gulf of Mexico, and it represents the first report of a parasite from the genus Luciobrotula Smith and Radcliffe in Radcliffe, 1913. A key is presented for Helicometra, all species of which are placed into one of four groups based on oral sucker shape, cirrus-sac length, and vitelline follicle distribution. Analyses and conclusions are given for trends observed in zoogeography, life history, and host families (definitive and intermediate) of Helicometra as well as the presence of this parasite in the deep sea and in Ophidiidae Rafinesque, 1810. A prospective life cycle for the new species includes a gastropod as first intermediate host, a caridean shrimp as second intermediate host, and an ophidiid cusk-eel as a definitive host. We provide a comprehensive historical faunal review of all species of Helicometra with information presented on taxonomic status (synonymies, validity of species), definitive and intermediate hosts, infection sites, geographic localities, and museum accession numbers for deposited specimens.
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References
Abdalah LGB, Elbohli S, Maamouri F (2010) Digenean diversity in labrid fish from the Bay of Bizerte in Tunisia. J Helminthol 84:27–33
Abollo E, Estévez J, Soto M, Gestal C, Rodríguez H, Pascual S, Arias-Fernández MC (1996) Diversity of parasite communities of Gobius paganellus L. in the Ria of Vigo (N.W. Spain). Parassitologia 38:11 [Abstract]
Agrawal V (1964) On a trematode, Helicometra indica n. sp. from the intestine of the gurnard fish, Trigla gurnardus. Proc Helminthol Soc Wash 31:216–218
Aken’Ova TOL, Cribb TH, Bray RA (2006) Helicometra Odhner, 1902 (Digenea: Opecoelidae) in Australian waters: problems of species identification and a description of H. sprenti n. sp. Syst Parasitol 63:17–27
Akmirza A (2000) Seasonal distribution of parasites detected in fish belonging to the Sparidae family found near Gökçeada. Türk Parazitol Derg 24:435–441
Akmirza A (2001) The samples from metazoan parasites detected in fish around Gökçeada. In: The national culture of the Aegean Islands 2001 proceedings book. Turkey, No. 7, pp 85–96
Akmirza A (2004) Parasite fauna of greater weever (Trachinus draco Linnaeus, 1758). Acta Adriat (Split) 45:35–41
Akmirza A (2012) Metazoan parasite fauna of conger eel (Conger conger L.) near Gökçeada, Northeastern Aegean Sea, Turkey. Kafkas Univ Vet Fak Derg 18:845–848
Aleshkina LD, Gaevskaia AV (1985) Trematodes of fish from the Atlantic coast of Africa. Nauchn Dokl Vyss Shk Biol Nauk 3:35–40
Appeltans W, Bouchet P, Boxshall GA, De Broyer C, de Voogd NJ, Gordon DP, Hoeksema BW, Horton T, Kennedy M, Mees J, Poore GCB, Read G, Stöhr S, Walter TC, Costello MJ (eds) (2012) World register of marine species. http://www.marinespecies.org. Accessed 19 July 2013
Arai HP (1963) Tremátodos digeneos de peces marinos de Baja California, México. Ann Inst Biol Univ Nac Méx 33:113–130
Arai HP (1969) Preliminary report on the parasites of certain marine fishes of British Columbia. J Fish Res Board Can 26:2319–2337
Arculeo M, Hristovski N, Riggio S (1997) Helminth infestation of three fishes (Serranus scriba, Mullus surmuletus, Scorpaena porcus) from a coastal seaground in the Gulf of Palermo (Tyrrhenian Sea). Ital J Zool 64:283–286
Arthur JR, Lumanlan-Mayo S (1997) Checklist of the parasites of fishes of the Philippines. FAO Fish Tech Pap Rome 369:1–102
Baeva OM (1968) Helminth fauna of Pleurogrammus azonus Jordan et Metz in the Sea of Japan. In: Skrjabin KI, Mamaev YL (eds) Helminths of animals of Pacific Ocean. Nauka, Moscow, pp 80–88
Barbagallo P, Drago U (1903) Primo contributo allo studio della fauna elmintologica dei pesci dell Sisalia orientale. Arch Parasitol Paris 7:408–427
Bartoli P, Boudouresque CF (1997) Transmission failure of parasites (Digenea) in sites colonized by the recently introduced invasive alga Caulerpa taxifolia. Mar Ecol Prog Ser 154:253–260
Bartoli P, Gibson DI, Bray RA (2005) Digenean species diversity in teleost fish from a nature reserve of Corsica, France (Western Mediterranean), and a comparison with other Mediterranean regions. J Nat Hist 39:47–70
Baylis HA (1939) Further records of parasitic worms from British vertebrates. Ann Mag Nat Hist 11:473–498
Baylis HA, Jones EI (1933) Some records of parasitic worms from marine fishes at Plymouth. J Mar Biol Assoc U K 18:627–634
Bona F, Buriola E, Cerioni S, Orecchia P, Paggi L (2000) Fascicolo 5. Digenea. In: Alessandro M, Sandro R, Sandro LP (eds) Checklist delle specie della fauna Italiana. Italian Ministry of Environment, Rome, pp 1–31
Braun MGCC (1893) II. Bericht über Thierische Parasiten. Zentralbl Bakteriol Parasitenkd 13:176–190
Bravo-Hollis M, Manter HW (1957) Trematodes of marine fishes of Mexican waters. X. Thirteen Digenea, including nine new species and two new genera, from the Pacific coast. Proc Helminthol Soc Wash 24:35–48
Bray RA (1973) Some digenetic trematodes in fishes from the Bay of Biscay and nearby waters. Bull Br Mus Nat Hist 26:151–183
Bray RA (1979) Dignenea in marine fishes from the eastern seaboard of Canada. J Nat Hist 13:399–431
Bray RA (1987a) Some helminth parasites of marine fishes of South Africa: Family Opecoelidae (Digenea). J Nat Hist 21:1049–1075
Bray RA (1987b) A study of the helminth parasites of Anarhichas lupus (Perciformes: Anarhichadidae) in the North Atlantic. J Fish Biol 31:237–264
Bray RA (2005) Family Acanthocolpidae Lühe, 1906. In: Jones A, Bray RA, Gibson DI (eds) Keys to the Trematoda, vol 2. CABI and Natural History Museum, Wallingford, pp 603–619
Bray RA, Cribb TH (1989) Digeneans of the Family Opecoelidae Ozaki, 1925 from the Southern Great Barrier Reef Australia including a new genus and three new species. J Nat Hist 23:429–474
Bray RA, Justine JL (2007) Pseudopycnadena tendu sp. nov. (Digenea, Opecoelidae) in the yellow-spotted triggerfish Pseudobalistes fuscus (Perciformes, Balistidae) and additional opecoelids parasitizing fishes from the waters off New Caledonia. Acta Parasitol 52:13–17
Brickle P, MacKenzie K, Pike A (2005) Parasites of the Patagonian toothfish, Dissostichus eleginoides Smitt 1898, in different parts of the Subantarctic. Polar Biol 28:663–671
Bush AO, Lafferty KD, Lotz JM, Shostak AW (1997) Parasitology meets ecology on its own terms: Margolis et al. revisited. J Parasitol 83:575–583
Campos A, Carbonell E (1994) Parasite community diversity in two Mediterranean labrid fishes Symphodus tinca and Labrus merula. J Fish Biol 44:409–413
Campos A, Carbonell E, Pellicer M (1990) Helminth fauna of Symphodus tinca L. and Labrus merula L. (Pisces: Labridae) from the Valencia coast. I. Trematoda. Rev Iber Parasitol 50:37–42
Carus JV (1884) Prodomus faunae mediterraneae. Vol. 1. E. Schweizerbart’sche Verlagshandlung, Stuttgart
Chaplina OM, Antsishkina LM (1961) Data on the fish parasites in small rivers of the northern Azov area. Dopov Akad Nauk Ukr RSR 2:247–250
Chatterji RC (1936) The helminths parasitic in the fresh-water turtles of Rangoon. Rec Indian Mus 38:81–94
Chertkova AN, Kosupko GA, Khrustalev AV (1994) Catalogue of type specimens of species of helminths from the collection of the Central Helminthological Museum. VIGIS, Moscow
Cobbold TS (1860) Synopsis of the Distomidae. J Proc Linn Soc Lond Zool 5:1–56
Cordero del Campillo M, Castañón Ordóñez L, Reguera Feo A (1994) Índice-catálogo de zooparásitos ibéricos. Universidad de León, León
Costello MJ, Deady S, Pike A, Fives JM (1996) Parasites and diseases of wrasse being used as cleaner-fish on salmon farms in Ireland and Scotland. In: Sayer MDJ, Treasurer JW, Costello MJ (eds) Wrasse: biology and use in aquaculture. Fishing News Books, Oxford, pp 211–227
Cribb TH (2005) Family Opecoelidae Ozaki, 1925. In: Jones A, Bray RA, Gibson DI (eds) Keys to the Trematoda, vol. 2. CABI and Natural History Museum, Wallingford, pp 443–531
Cribb TH, Bray RA, Wright T, Pichelin S (2002) The trematodes of groupers (Serranidae: Epinephelinae): knowledge, nature and evolution. Parasitology 124:23–42
Crofton HD (1941) A record of trematode parasites from Mola mola and Raniceps raninus (Linn.). Parasitology 33:209–210
Crofton HD (1947) The parasites of some littoral fishes of Northumberland. Rep Dove Mar Lab 9:59–64
Crowcroft PW (1947a) Some digenetic trematodes from fishes of shallow Tasmanian waters. Pap Proc R Soc Tasman 1946:5–25
Crowcroft PW (1947b) The anatomy of two new digenetic trematodes from Tasmanian food fishes. Proc Linnean Soc NSW 71:108–118
Culurgioni J, D’Amico V, Coluccia E, Mulas A, Figus V (2006) Metazoan parasite fauna of conger eel Conger conger L. from Sardinian waters (Italy). Ittiopatologia 3:253–261
Dawes B (1956) The Trematoda with special reference to British and other European forms. Cambridge Univ. Press, London
De Buron I, Maillard C (1987) Experimental transfer by ichthyophagy and cannibalism of adult helminths, parasite of fishes. Ann Parasitol Hum Comp 62:188–191
De Giusti DL, Nasir P (1960) Trematode parasites of the fish family Gobiidae from Naples, Italy and Bimini, B.W.I. J Parasitol 46:13 [Abstract]
Dezfuli BS, Manera M, Onestini S, Rossi R (1997) Histopathology of the alimentary canal of Anguilla anguilla L. associated with digenetic trematodes: a light and electron microscopic study. J Fish Dis 20:317–322
Dezfuli BS, Lui A, Boldrini P, Pironi F, Giari L (2008) The inflammatory response of fish to helminth parasites. Parasite 15:426–433
Dezfuli BS, Giari L, Castaldelli G, Lanzoni M, Rossi R, Lorenzoni M, Kennedy CR (2014) Temporal and spatial changes in the composition and structure of helminth component communities in European eels Anguilla anguilla in an Adriatic coastal lagoon and some freshwaters in Italy. Parasitol Res 113:113–120
Di Cave D, Berrilli F, De Liberato C, Orecchia P, Kennedy CR (2001) Helminth communities in eels Anguilla anguilla from Adriatic coastal lagoons in Italy. J Helminthol 75:7–13
Diesing KM (1850) Systema helminthum. Vindobonae 1:1–680
Dolgikh AV, Naĭdenova NN (1968) On the fauna of helminths of Gaidropsarus mediterraneus (L.) from the Black Sea. Parazitologia (Leningrad) 2:448–453
Dujardin F (1845) Histoire naturelle des helminthes ou vers intestinaux. Paris
Durio WO, Manter HW (1968) Some digenetic trematodes of marine fishes of New Caledonia. Part II. Opecoelidae and Lepocreadiidae. J Parasitol 54:747–756
Dyer WG, Williams EH Jr, Bunkley-Williams L (1985) Digenetic trematodes of marine fishes of the western and southwestern coasts of Puerto Rico. Proc Helminthol Soc Wash 52:85–94
Dyer WG, Williams EH Jr, Bunkley-Williams L (1988) Digenetic trematodes of marine fishes of Okinawa, Japan. J Parasitol 74:638–645
Dyer WG, Williams EH Jr, Bunkley-Williams L (1992) Homalometron dowgialloi sp. n. (Homalometridae) from Haemulon flavolineatum and additional records of digenetic trematodes of marine fishes in the West Indies. J Helminthol Soc Wash 59:182–189
El-Labadi SN, Ismail NS, Khalaf M (2006) Intestinal digenetic trematodes of some fishes from the Gulf of Aqaba, Red Sea. Pak J Zool 38:43–48
Ergens R (1960) Helminth fauna of fish from Alabania. Česk Parasitol 7:49–90
Fischthal JH (1972) Zoogeography of digenetic trematodes from West African marine fishes. Proc Helminthol Soc Wash 39:192–203
Fischthal JH (1977) Some digenetic trematodes of marine fishes from the barrier reef and reef lagoon of Belize. Zool Scr 6:81–88
Fischthal JH (1978) Allometric growth in four species of digenetic trematodes of marine fishes from Belize. Zool Scr 7:13–18
Fischthal JH, Kuntz RE (1964) Digenetic trematodes of fishes from Palawan Island, Philippines. Part II. Five Opecoelidae, including three new species. Proc Helminthol Soc Wash 31:40–46
Fischthal JH, Kuntz RE (1965) Digenetic trematodes of fishes from North Borneo (Malaysia). Proc Helminthol Soc Wash 32:63–71
Fischthal JH, Thomas JD (1970) Digenetic trematodes of marine fishes from Ghana: Family Opecoelidae. Proc Helminthol Soc Wash 37:129–141
Fischthal JH, Thomas JD (1972) Digenetic trematodes of marine fishes from Senegal. Bull Inst Fond Afr Noire 34:292–322
Froese R, Pauly D (eds) (2013) FishBase (version 04/2013). http://www.fishbase.org. Accessed 12 Feb 2014
Fuhrmann O (1928) Zweite Klasse des Cladus Plathelminthes: Trematoda. Handb Zool 2:1–128
Gaevskaia AV (1981) Helicometra indica (Trematoda: Opecoelidae), a parasite of fish of the West European shelf. Parazitologia 15:178–181
Gaevskaia AV (1995) New data on fish trematodes from the Black Sea. In: Program and book of abstracts, IVth International Symposium on Fish Parasitology, 3–6 October 1995. Munich, p 116
Gaevskaia AV (2002) New data on trematodes of the families Opecoelidae and Accacoelidae from fishes in Atlantic Ocean and its seas. Parazitologia 36:219–223
Gaevskaia AV, Umnova BA (1977) Parasitic fauna of the principal commercial fishes of the Northwest Atlantic. Biol Moria (Vladivost) 4:40–48
Galtsoff PS (1954) Gulf of Mexico, it’s origin, waters, and marine life. Fish Bull Fish Wildl Serv (No 89) 55:1–604
Gibson DI (1996) Part IV. Trematoda. In: Margolis L, Kabata Z (eds) Guide to the parasites of fishes of Canada. NRC Research Press, Ottawa, pp 1–373
Gibson DI, Bray RA (1982) A study and reorganization of Plagioporus Stafford, 1904 (Digenea: Opecoelidae) and related genera, with special reference to forms from European Atlantic waters. J Nat Hist 16:529–559
Gupta NK (1956) Studies on the digenetic trematodes of marine fishes from the Gulf of Manaar (India). Res Bull Panj Univ Zool 89:61–83
Hafeezullah M (1971) A review on the validity of Helicometrina Linton, 1910 and Stenopera Manter, 1933 (Trematoda). Acta Parasitol Pol 19:133–139
Hanumantha Rao K, Kishore S, Shyamasundari K (1993) Observations on reproductive effort in trematode parasites. Riv Parassitol (Roma) 10:487–501
Hassanine RME (2001) Digenean trematodes of marine teleost fishes: the genus Helicometra Odhner, 1902 (Opecoelidae): on the validity of two known species from the Red Sea. Egypt J Aquat Biol Fish 5:83–96
Hassanine RME (2005) Trematodes of Red Sea fishes: Gibsonius aegyptensis gen. nov., sp. nov. (Lepocreadiidae Odhner, 1905) and Helicometra interrupta sp. nov. (Opecoelidae Ozaki, 1925). Acta Parasitol 50:276–280
Hassanine RME (2007) Trematodes from Red Sea fishes: on the validity of Helicometra marmoratae Nagaty et Abdel-Aal, 1962 and the description of H. aegyptense sp. nov. (Opecoelidae Ozaki, 1925). Acta Parasitol 52:18–23
Hermida M, Saraiva A, Cruz C (2008) Metazoan parasite community of a European eel (Anguilla anguilla) population from an estuary in Portugal. Bull Eur Assoc Fish Pathol 28:35–40
Hewitt CC, Hine PM (1972) Checklist of parasites of New Zealand fishes and of their hosts. N Z J Mar Freshw Res 6:69–114
Hofer B (1904) Handbuch der Fischkrankheiten. Munich
Holloway HL Jr, Bier JW (1968) Helicometra antarcticae sp. nov. from Antarctic coastal fishes. Proc Helminthol Soc Wash 35:30–34
Holmes JC (1990) Helminth communities in marine fishes. In: Esch G, Bush A, Aho J (eds) Parasite communities: patterns and processes. Chapman Hall, London, pp 101–130
Holmes JC (1991) Spatial scale and important species in the analysis of communities of parasites. In: Second International School: parasite–host environment, Sofia, 28 May–1 June 1991, Bulgarian Academy of Sciences, Sofia
Hopkins SH (1941) The excretory systems of Helicometra and Cymbephallus (Trematoda), with remarks on their relationships. Trans Am Microsc Soc 60:41–44
Hossam N, Haridy M, El-Ezz AA, El-Sayed A, Ibrahim A (2012) Histopathological studies on the internal parasites of Arabian Mallas (Thalassoma klunzengri) from Red Sea. In: Proceedings of the Fifth Global Fisheries and Aquaculture Research Conference, Giza, 1–3 October 2012. Faculty of Agriculture, Cairo University, Giza, pp 33–41
Hristovski N, Jardas I (1991) Endohelminths of fishes from the mid-Dalmatian region of the Adriatic Sea (a review). Acta Adriat (Split) 32:671–681
Iannacone J (2004) Metazoan parasites of the minor stardrum, Stellifer minor (Tschudi) (Osteichthyes, Sciaenidae), caught by artisanal fishery on Chorrillos, Lima, Peru. Rev Bras Zool 21:815–820
Iannacone J, Moron L, Guizado S (2010) Between-year variation of metazoan parasite fauna on Sciaena deliciosa (Tschudi, 1846) (Perciformes: Sciaenidae) in Lima, Peru. Lat Am J Aquat Res 38:218–226
Iannacone J, Cerapio JP, Cardenas-Callirgos J, Sanchez K, Briceno F, Duenas A (2011) Parasite communities of the trambollo, Labrisomus philippii (Steindachner, 1866) (Perciformes: Labrisomidae) off coastal zone of Chorrillos, Lima, Peru. Neotrop Helminthol 5:73–84
Isaĭchikov IM (1928) Kpoznani paraziticheskikh chervei nekotorykh grupp pozvonochnykh Russkoi Arktiki. A. Trematodes. [Parasitic worms of some vertebrate groups of the Russian Arctic. A. Trematodes]. Tr Mor Nauch Inst Mosk 3:5–79
Janiszewska J (1953) Some Adriatic Sea fish trematodes. Zool Pol 6:20–48
Jousson O (2001) Les trématodes digénétiques parasites de poissons en Méditerranée occidentale: cycles biologiques, diversité cryptique et évolution moléculaire. Dissertation, Université de Genève
Jousson O, Bartoli P, Pawlowski J (1999) Molecular identification of developmental stages in Opecoelidae (Digenea). Int J Parasitol 29:1853–1858
Kalantan AMN, Arfin M, Alyousif MS (1992) Chemical nature and mode of stabilization of egg-shell in Helicometra pulchella (Rudolphi, 1819) (Digenea: Opecoelidae). Acta Zool 73:153–156
Klimpel S, Seehagen A, Palm HW, Rosenthal H (2001) Deep-water metazoan fish parasites of the world. Logos Verlag, Berlin
Klimpel S, Busch MW, Kellermanns E, Kleinertz S, Palm HW (2009) Metazoan deep-sea fish parasites. Verlag Natur and Wissenschaft, Solingen
Ko RC, Chan SW (2002) A preliminary study on the parasite fauna of three common marine fish of Hong Kong (Siganus fuscescens, Sebasticus marmoratus, Epinephelus akaara). University of Hong Kong Occasional Publication, Hong Kong
Køie M (2000) Metazoan parasites of teleost fishes from Atlantic waters off the Faroe Islands. Ophelia 52:25–44
Korniychuk JM (1997) Trematode fauna of four common blenniid species in the Black Sea. Ekol Morya (Kiev) 46:43–46
Korniychuk JM (1999) Phenotypic hostal differentiation of Helicometra fasciata (Rud., 1891) maritae. Ekol Morya (Kiev) 49:44–48
Korniychuk JM (2000a) Morphological variability of genus Helicometra (Trematoda: Opecoelidae) maritae from the Black Sea. Ekol Morya (Kiev) 51:40–44
Korniychuk JM (2000b) Interspecific interactions of trematodes in the gut of Symphodus tinca. Ekol Morya (Kiev) 52:34–38
Korniychuk JM (2001a) Structure of the trematode fauna of the Black Sea labrid fishes (Pisces: Labridae). Ekol Morya (Kiev) 58:32–36
Korniychuk JM (2001b) New data on trematode fauna of the Black Sea labrid fishes. In: Sixth National Conference of Parasitology, Sofia, 5–7 October 2001, programs and abstracts. Pensoft, Sofia, pp 24–25
Korniychuk JM (2008) Parthenogenetic generations of Helicometra fasciata Rud., 1819 (Trematoda: Opecoelidae) in the Black Sea molluscs Gibbula adriatica. Parazitologia 42:41–52
Korniychuk JM, Gaevskaia AV (1999) Reproductive strategy of trematode, Helicometra fasciata (Trematoda: Opecoelidae) as an index of favourability of its environment. Ekol Morya (Kiev) 48:42–47
Koubkova B, Barus V (2000) Metazoan parasites of the recently established tubenose goby (Proterorhinus marmoratus: Gobiidae) population from the South Moravian reservoir, Czech Republic. Helminthol 37:89–95
Kovaleva AA (1970) Some features of the helminth fauna of horse mackerel of the genus Trachurus (Carangidae, Perciformes). Biol Morya (Parazit Mor Mollyuskov Ryb I Mleko) 20:66–74
Kuramochi T (2006) Digenean trematodes of fishes caught in the Sagami Sea, central Japan. Mem Natl Sci Mus (Tokyo) 40:175–186
Lester RJG, Sewell KB (1989) Checklist of parasites from Heron Island, Great Barrier Reef. Aust J Zool 37:101–128
Levron C, Ternengo S, Marchand B (2003) Ultrastructure of spermiogenesis and the spermatozoon of Helicometra fasciata (Digenea, Opecoelidae), a parasite of Labrus merula (Pisces, Teleostei). Acta Parasitol 48:255–264
Li QK, Qui ZZ, Zhang RS (1989) Digenetic trematodes of fishes from the Bo-Hai Sea, China VI. (Trematoda: Opecoelidae). Acta Zootaxon Sin 14:12–16
Linstow OFB von (1889) Compendium der Helminthologie. Nachtrag. Die Litteratur der Jahre 1878–1889. Hannover
Linton E (1910) Helminth fauna of the Dry Tortugas. 2. Trematodes. Pap Tortugas Lab Carneg Inst Wash Publ 133:11–98
Little PA (1929) The trematode parasites of Irish marine fishes. Parasitol 21:22–30
Liu S-F, Peng W-F, Gao P, Fu M, H-Z W, Lu M-K, Gao J-Q, Xiao J (2010) Digenean parasites of Chinese marine fishes: a list of species, hosts and geographical distribution. Syst Parasitol 75:1–52
Looss A (1899) Weitere Beiträge zur Kenntnis der Trematoden-Fauna Aegyptens, zugleich Versuch einter natürlichen Gliederung des Genus Distomum Retzius. Zool Jahrb Jena Abt Syst 12:521–784
Lopez-Roman R, Guevara Pozo D (1974) Incidence of digenean parasitism in various marine teleosts from the sea around Alboran. Rev Iber Parasitol 34:147
Lopez-Roman R, Guevara Pozo D (1977) Algunos Opecoelidae Ozaki, 1925 de teleosteos del Mar de Alboran. In: Caballero y Caballero E (eds) Excerta parasitológica en memoria del doctor Eduardo Caballero y Caballero, Universidad Nacional Autónoma de México, México, pp 223–231
Love MS, Moser M (1976) Parasites of California marine and estuarine fish. University of California, Santa Barbara
Love MS, Moser M (1983) A checklist of parasites of California, Oregon, and Washington marine and estuarine fishes. NOAA Tech Rep, No. NMFS SSRF-777:1–576
Love MS, Yoklavich M, Thorsteinson L (2002) The rockfishes of the northeast Pacific. Univ. of California Press, Berkeley
Lozano C, Ubeda JM, Rojas M, de Ariza C, Guevara DC (2002) Opecoelidae Ozaki, 1925 (Platyhelminthes, Trematoda, Digenea) parasites of marine fishes from Southern Iberian Peninsula coasts. Bol Real Soc Esp Hist Nat Secc Biol 97:27–32
Luque JL (1994) Population dynamics and community structure of metazoan parasites of Menticirrhus ophicephalus (Pisces: Sciaenidae) from the Peruvian coast. Rev Biol Trop (Costa Rica) 42:21–29
Luque JL, Oliva M (1993a) Structural and quantitative analysis of the parasitic community of Paralonchurus peruanus (Pisces: Sciaenidae) from the coast of Peru. Parasitol Dia 17:107–111
Luque JL, Oliva M (1993b) Trematodes of marine fishes from the Peruvian Faunistic Province (Peru and Chile), with description of Lecithochirium callaoensis n. sp. and new new records. Rev Biol Mar 28:271–286
Luque JL, Iannacone J, Farfan C (1991) Parasitos de peces oseos marinos en el Peru: lista de especies conocidas. Bol Lima 74:17–28
Machida M (1984) Trematodes of marine fishes from depth of 200–400 m off Yamagata, the Japan Sea. Mem Natl Sci Mus (Tokyo) 17:101–110
Machida M, Araki J (1994) Some trematodes and cestodes in fishes from off eastern Hokkaido, northern Japan. Mem Natl Sci Mus (Tokyo) 27:87–92
Machida M, Ichihara A, Kamegai S (1970) Digenetic trematodes collected from the fishes in the sea north of the Tsushima Islands. Mem Natl Sci Mus (Tokyo) 3:101–112
Machida M, Araki J, Kamiya H, Ohbayashi M (1972) Trematodes collected from sea fishes of the Hidaka district, Hokkaido. Mem Natl Sci Mus (Tokyo) 5:1–9
Machkevsky VK (1990) Helminthofauna of labrids in the places of cultivation of Black-Sea mussel, Mytilus galloprovincialis. Ekol Morya (Kiev) 36:75–82
Machkevsky VK, Gaevskaia AV (1997) Response to some parasite systems from the Black Sea coastal zone to aquatic pollution. Ekol Morya (Kiev) 46:51–57
Machkevsky V, Gaevskaia A, Korniychuk J (1996) Population dynamic of the trematode, Helicometra fasciata from the Black Sea. Parassitologia (Roma) 38:40
Machkevsky VK, Pronkina NV, Gaevskaia AV, Korniychuk JM (1997) Life cycle of Helicometra fasciata (Rud., 1819) (Trematoda:Opecoelidae) in the rock biocenosis from the Black Sea. Ekol Morya (Kiev) 46:58–62
Madhavi R (1975) Digenetic trematodes from marine fishes of Waltair Coast, Bay of Bengal. Family Opecoelidae. Riv Parassitol (Roma) 36:153–164
Madhavi R (2011) Checklist of digenean trematodes reported from Indian marine fishes. Syst Parasitol 78:163–232
Maillard C, Aussel JP (1988) Host specificity of fish trematodes investigated by experimental ichthyophagy. Int J Parasitol 18:493–498
Maillard C, Lambert M (1978) Plagioporus novellus n. sp. (Trematoda, Opecoelidae), parasite de Conger conger (Teleostei). Ann Parasitol 53:367–371
Mamaev YL, Oshmarin PG (1966) Trematodes of the family Acanthocolpidae Luhe, 1901 in herrings of the North-Vietnam Bay. Helminthol 7:155–164
Manter HW (1933) The genus Helicometra and related trematodes from Tortugas, Florida. Pap Tortugas Lab 28:167–182
Manter HW (1934) Some digenetic trematodes from deep-water fish of Tortugas, Florida. Pap Tortugas Lab 28:257–345
Manter HW (1940) Digenetic trematodes of fishes from the Galapagos Islands and the neighboring Pacific. Rep Allan Hancock Pac Exped (1932–1938) 2:325–497
Manter HW (1947) The digenetic trematodes of marine fishes of Tortugas, Florida. Am Midl Nat 38:257–416
Manter HW (1954) Some digenetic trematodes from fishes of New Zealand. Trans R Soc N Z 82:475–568
Manter HW, Pritchard MH (1960) Some digenetic trematodes of eels of Hawaii. J Parasitol 46:651–658
Margolis L, Arthur JR (1979) Synopsis of the parasites of fishes of Canada. Bull Fish Res Board Can 199:1–269
Markevich AP (1952) Parasite fauna of freshwater fish of Ukrainian SSR. Academy of Sciences of the Ukrainian SSR Zoological Institute, Kiev
Mateo C (1972) La Ictiopatología y su importante rol en la productividad pesquera. Ed Univ Fed Vill Perú, pp 122
Mathias P (1934) Sur quelques trématodes de poissons marins de la region de Banyuls. Arch Zool Exp Gén 75:567–581
McEachran JD, Fechhelm JD (1998) Fishes of the Gulf of Mexico. Vol 1: Myxiniformes to Gasterosteiformes. Univ. of Texas Press, Austin
Meenakshi M, Madhavi R, Swarnakumari VGM (1993) The life-cycle of Helicometra gibsoni n. sp. (Digenea: Opecoelidae). Syst Parasitol 25:63–72
Monticelli FS (1893) Studii sui trematodi endoparassiti: primo contributo di osservazioni sui distomidi. Zool Jahrb Abt Syst Ökol Geogr Tiere 7:1–230
Moravec F, Vidal-Martinez VM, Vargas-Vázquez J, Vivas-Rodríguez C, González-Solis D, Mendoza-Franco E, Simá-Alvarez R, Güemez-Ricalde J (1997) Helminth parasites of Epinephelus morio (Pisces: Serranidae) of the Yucatan Peninsula, southeastern Mexico. Folia Parasitol 44:255–266
Mordvinova TN (1979) Higher Crustacea as intermediate hosts of fish helminths. In: VII Vsesoyuznoe soveshchanie po parazitam i boleznyam ryb, Leningrad, Sentyabr’, (Tezisy dokladov), Nauka, Leningrad, pp 74–75 [Abstract]
Mordvinova TN (2000) Helminthofauna of myctophid fishes from the World Ocean. Ekol Morya (Kiev) 53:59–63
Mordvinova TN (2001) The place and role of Myctophidae in the parasitic systems of the World Ocean. In: Sixth National Conference of Parasitology, Sofia, 5–7 October 2001, programs and abstracts. Pensoft, Sofia, p 25
Muñoz G, Cribb TH (2006) Parasite communities and diet of Coris batuensis (Pisces: Labridae) from Lizard Island, Great Barrier Reef. Mem Qld Mus 52:191–198
Nagasawa K (1989) Literature on fish parasites of Japan, published in 1988–89. Parasite–host list. Newsl Ichthyo Parasitol Group Jpn (Yoichi) 4:5–17
Nagaty HF (1956a) Trematodes of fishes from the Red Sea. Part 6. On five distomes including one new genus and four new species. J Parasitol 42:151–155
Nagaty HF (1956b) Trematodes of fishes from the Red Sea. Part 7. On two gyliauchenids and three allocreadoids, including four new species. J Parasitol 42:523–527
Nagaty HF, Abdel Aal TM (1962) Trematodes of fishes of the Red Sea. Part 17. On three allocreadiid sp. and one schistorchiid sp. J Arab Vet Med Assoc 22:307–314
Nahhas FM (1993) Some Acanthocephala and Digenea of marine fish from Grand Cayman, Cayman Islands, British West Indies. J Helminthol Soc Wash 60:270–272
Nahhas FM, Cable RM (1964) Digenetic and aspidogastrid trematodes from marine fishes of Curaçao and Jamaica. Tulane Stud Zool 11:169–228
Nahhas FM, Carlson K (1994) Digenetic trematodes of marine fishes of Jamaica, West Indies. Publi Hofstra Univ Mar Lab Ecol Surv Jam 2:1–60
Naĭdenova NN (1966) Two cases of teratology in helminths. In: Voprosy morskoi biologii Tezisy simpoziuma molodykh uchenykh Instituta Biologii Yuzhnykh Morei, Sevastopol’, 13–16 Aprelya, 1966, pp 88–89 [Abstract]
Naĭdenova NN (1974) Parasite fauna of gobiid fish in the Black and Azov Seas. Naukova Dumka, Kiev
Naĭdenova NN, Dolgikh AV (1969) Revision of some Helicometra species (Trematoda: Opecoelidae). Nauch Dokl Vyss Shk Biol Nauk 12:7–12
Naĭdenova NN, Mordvinova TN (1997) Helminth fauna of Mediterranean Sea fish upon the data of the IBSS’s expeditions (1959–1973). Ekol Morya (Kiev) 46:69–74
Nguyễn VH (2012) The description of two new species Helicometra pisodonophi sp. n. and Opecoelus haduyngoi sp. n. (Trematoda: Opecoelidae) from marine fishes in Ha Long Bay, Vietnam. Tạp Chí Sinh Học 34:133–138
Nicoll W (1909) Studies on the structure and classification of the digenetic trematodes. Q J Microsc Sci 53:391–487
Nicoll W (1910) On the entozoa of fishes from the Firth of Clyde. Parasitol 3:322–359
Nicoll W (1914) The trematode parasites of fishes from the English Channel. J Mar Biol Assoc U K 10:466–505
Nicoll W (1915) A list of the trematode parasites of British marine fishes. Parasitol 7:339–378
Nielsen JG (2009) A revision of the bathyal genus Luciobrotula (Teleostei, Ophidiidae) with two new species. Galathea Rep 22:141–156
Niewiadomska K (2002) Family Liolopidae Odhner, 1912. In: Gibson DI, Jones A, Bray RA (eds) Keys to the Trematoda, vol 1. CABI and Natural History Museum, Wallingford, pp 121–125
Nikolaeva VM, Solonchenko AI (1970) On the helminth fauna of some benthopelagic fishes of the Black Sea. Biol Morya (Parazit Mor Mollyuskov Ryb I Mleko) No. 20, pp 129–166
Odhner T (1901) Revision einiger Arten der Distomengattung Allocreadium Lss. Zool Jahrb Jena Abt Syst 14:483–520
Odhner T (1902a) Revision einiger Arten der Distomengattung Allocreadium Lss. Zool Cent 9:404 [Abstract]
Odhner T (1902b) Mittielungen zur Kenntnis der Distomen. 1. Cent Bakt 1. Abt Orig 31:58–69
Odhner T (1902c) Mittielungen zur Kenntnis der Distomen. 2. Cent Bakt 1. Abt Orig 31:152–162
Oguz MC (1995) Mudanya kiyikarindaki bazi teleost baliklarda rastlanilan helmintler. Dissertation, Uludag University
Oguz MC, Bray RA (2006) Digenetic trematodes of some teleost fish off the Mudanya Coast (Sea of Marmara, Turkey). Helminthologia 43:161–167
Oliva ME, Luque JL (1998) Metazoan parasite infracommunities in five sciaenids from the central Peruvian coast. Mem Inst Oswaldo Cruz 93:175–180
Oliva ME, Luque JL (2002) Endohelminth parasites of the trambollo Labrisomus philippii (Steindachner) (Osteichthyes: Labrisomidae) from the central Peruvian coast. Comp Parasitol 69:100–104
Oliva M, Muñoz M (1985) Helicometra fasciata (Rudolphi, 1819) y Helicometrina nimia Linton, 1910 (Opecoelidae) en peces marinos de la II Región-Chile. Parasitol Día 9:107–111
Oliva ME, Luque JL, Iannacone JA (1990) The metazoan parasites of Stellifer minor (Tschudi, 1844): an ecological approach. Mem Inst Oswaldo Cruz 85:271–274
Olsson P (1868) Entozoa, iakttagua hos Skandineviska hafsfiskar. I. Platyelminthes. Lund’s Universitets Arsskrift. Math Naturvidensk Vetensk 4:1–64
Orecchia P, Paggi L (1978) Aspetti di systematica e di ecologia degli elminti parassiti di pesci marini studiati presso l’Istituto di Parassitologia dell’Universita di Roma. Parassitologia 20:73–89
Orecchia P, Paggi L, Radujkovic BM (1988) Digeneans of fishes from the Adriatic Sea with a description of Lecithaster atherinae n. sp. from Atherina (Hepsetia) boyeri. Parassitologia 30:225–229
Ortis M, Amelio D, Mariniello L, Sequi R, Ippoliti P, Paggi L (2002) Metazoan parasite component communities in fish from the Natural Marine Reserve “Island of Ustica”, South Tyrrhenian Sea (Mediterranean). Parassitologia 44(Suppl):1–120
Osmanov SO (1940) Data on the parasitic fauna of fishes from the Black Sea. Uch Zap Leningr Gos Pedagog Inst Im NK Krupskoi 30:187–267
Outeiral S, Alvarez F, Paniagua E, Leiro L, Sanmartin ML (2001) Digenean parasites of the European eel from estuaries in North-West Spain. Helminthologia 38:85–92
Overstreet RM (1969) Digenetic trematodes of marine teleost fishes from Biscayne Bay, Florida. Tulane Stud Zool Bot 15:119–176
Overstreet RM, Martin DM (1974) Some digenetic trematodes from synaphobranchid eels. J Parasitol 60:80–84
Overstreet RM, Cook JO, Heard RW (2009) Trematoda (Platyhelminthes) of the Gulf of Mexico. In: Felder DL, Camp CK (eds) Gulf of Mexico origin, waters and biota. Volume 1. Biodiversity. Texas A&M University Press, College Station, pp 419–486
Palombi A (1929a) Ricerche sul ciclo evolutivo di Helicometra fasciata (Rud.). Revisione delle specie del genere Helicometra Odhner. Pubblici Stn Zool (Napoli) 9:237–292
Palombi A (1929b) Le specie del genere Helicometra essistenti nella collezione elmintologica central Italiana. Ann Mus Zool R Univ Napoli 5:1–19
Palombi A (1931) Il polimorfismo nei Trematodi Ricerche sperimentali su Helicometra fasciata (Rud.). Helicometra fasciata (Rud.) = H. pulchella (Rud.) = H. sinuata (Rud.). Ann Mus Zool R Univ Napoli 6:1–8
Palombi A (1934a) Bacciger bacciger (Rud.). Trematode dignetico: fam. Steringophoridae Odhner. Anatomia, sistematica e biologia. Pubblici Stn Zool (Napoli) 13:438–478
Palombi A (1934b) Gli stadi larvali dei trematodi gel Golfo di Napoli. 1°. Contributo allo studio della morfologia, biologia e sistematica delle cercarie marine [Larval stages of the trematodes of the Gulf of Naples. 1. Contribution to the study of the morphology, biology, and taxonomy of the marine cercariae]. Pubblici Stn Zool (Napoli) 14:51–94
Palombi A (1937) La cercaria de Mesometra orbicularis (Rud.) e la sua trasformazione in metacercaria. Appunit sul ciclo evolutivo. Riv Parassitol (Roma) 1:13–17
Palombi A (1938) Metodi impiegati per lo studio dei cicli evolutivi dei trematodi digenetici. Materiale per la conoscenza della biologia di Podocotyle atomon (Rud.). In: Neiva A (ed) Livro Jubilar do Professor Lauro Travassos. Editado para commemorar o 25° anniversario de suas actividades scientificas (1913–1938). Typographia do Instituto do Oswaldo Cruz, Rio de Janeiro, pp 371–379
Pan JH, Zhang JY, Li ZC (1990) Parasitology of fishes: a survey of study of parasites appearing in Chinese freshwater fishes. Science Press, Beijing
Paniagua E, Vilas R, Sanmartin ML, Santamarina MT, Leiro JM, Ubeira FM (1999) Characterization by enzyme electrophoresis of specimens of the genus Helicometra (Trematoda, Opecoelidae) from fish caught off the coast of northwest Spain. Parasite 6:89–91
Papoutsoglou SE (1976) Metazoan parasites of fishes from Saronicos Gulf Athens-Greece. Thalassographica 1:69–102
Paradižnik V, Radujković B (2007) Digenea trematodes in fish of the North Adriatic Sea. Acta Adriat 48:115–129
Parona C (1887) Vermi parassiti in animali della Liguria. Nota preventiva a contributo di una elmintologia ligure. (Res Ligusticae. 2). Ann Mus Civ Storia Nat Genova 4:483–501
Parona C (1902) Catalogo di elminti raccolti in vertebrati dell’ Isola d’Elba (Seconda nota). Boll Mus Zool (Genova) 113:1–20
Parona C (1912) L’elmintologia italiana da’suoi primi tempi all’anno 1910. Bibliografia, sistematica, corologia, storia. V. 2: Sistematic, corologia, storia. Novara, Italy
Parukhin AM (1966) On the species composition of the helminth fauna of fishes in the South Atlantic. Mater Nauch Konf Vses Obs Gel’mintol 3:219–222
Parukhin AM (1968) Helminth fauna of fishes of the South Atlantic. Biol Morya (Parazit Mor Zhivot) No 14:96–113
Parukhin AM (1970) On a study of the trematode fauna of fishes of the Red Sea and the Gulf of Aden. Biol Morya (Parazit Mor Mollyuskov Ryb I Mleko) No 20:187–213
Parukhin AM (1989) Parasitic worms of bottom fishes of the southern seas. Naukova Dumka, Kiev
Pérez-Ponce de León G, García-Prieto L, Mendoza-Garfias B, León-Règagnon V, Pulido-Flores G, Aranda-Cruz C, García-Vargas F (1999) Listados faunísticos de México IX. Biodiversidad de helmintos parásitos de peces marinos y estuarinos de la Bahía de Chamela, Jalisco. UNAM, Mexico City
Pogorel’tseva TP (1954) New species of digenetic trematodes from fishes in the Black Sea. Nauk Zap Kiev Derjavnogo Pedagog Inst 15:133–137
Polianskii II (1955) Materials on the parasitology of fish of the northern seas of SSSR. Parasites of fish of the Barents Sea. Tr Zool Inst Akad Nauk SSSR 19:5–170
Pozdnyakov SE (1990) Helminths of scombrid-like fishes of the world’s oceans. DVO AN SSSR, Vladivostok
Pritchard MH (1966) Studies on digenetic trematodes of Hawaiian fishes: Family Opecoelidae Ozaki, 1925. Zool Jahrb Jena Abt Syst 93:173–202
Prudhoe S, Bray RA (1973) Digenetic trematodes from fishes. Rep Ser BANZ Antarct Res Exped Ser B 8:199–225
Quilichini Y, Foata J, Justine JL, Bray RA, Marchand B (2011) Sperm ultrastructure of Helicometra epinepheli (Platyhelminthes, Digenea, Opecoelidae), parasite of Epinephelus fasciatus (Pisces, Teleostei). Histol Histopathol 26:1019–1028
Radujkovic BM, Raibaut A (1989) Parasites des poissons marins du Montenegro: liste des especes de poissons avec leurs parasites. Acta Adriat (Split) 30:307–320
Radujkovic BM, Orecchia P, Paggi L (1989) Parasites des poissons marins du Montenegro: Digènes. Acta Adriat (Split) 30:137–187
Ramadan MM (1986) Trematodes of the genera Helicometra Odhner, 1902 (Opecoelidae) and Apharyngogyliauchen Yamaguti, 1942 (Gyliauchenidae) from the Red Sea fishes. Jpn J Parasitol 35:483–490
Rees FG (1945) A record of parasitic worms from fishes in rock pools at Aberystwyth. Parasitology 36:165–167
Reimer LW (1974) Pecularities of parasitocoenoses of deep-sea fishes. In: Proceedings of the Third International Congress of Parasitology, Munich, 25–31 August 1974, proceedings (vol. III). Facta Publication, Munich, pp 1648–1649
Reimer LW (1975) Helminthen von fischen des mesopelagials von Nordwestafrika. Wiss Z Pädagog Hochsch Güstrow 2:151–172
Reversat J (1990) Spécificité parasitaire des Trématodes de poissons: le genre Helicometra (Opecoelidae) dans l’étang de Thau (France). Dissertation, Université de Montpellier
Reversat J, Silan P (1991) Comparative population biology of Digenea and their first intermediate host mollusk: the case of three Helicometra (Trematoda: Opecoelidae) endoparasites of marine prosobranchs (Gastropoda). Ann Parasitol Hum Comp 66:219–225
Reversat J, Silan P (1993) Comparative population biology of fish digeneans: the case of three Helicometra (Trematoda: Opecoelidae) mesoparasites of marine teleosts in a Mediterranean lagoon. Ann Parasitol Hum Comp 68:128–135
Reversat J, Renaud F, Maillard C (1989) Biology of parasite populations: the differential specificity of the genus Helicometra Odhner, 1902 (Trematoda: Opecoelidae) in the Mediterranean Sea demonstrated by enzyme electrophoresis. Int J Parasitol 19:885–890
Reversat J, Leducq R, Marin R, Renaud F (1991a) A new methodology for studying parasite specificity and life cycles of trematodes. Int J Parasitol 21:467–469
Reversat J, Maillard C, Silan P (1991b) Phenotypic and enzymatic polymorphism: use and limitations in the descriptions of species of Helicometra, mesoparasites of marine teleosts. Syst Parasitol 19:147–158
Rigby M, Holmes JC, Cribb TH, Morand S (1997) Pattern of species diversity in the gastrointestinal helminths of a coral reef fish, Epinephalus merra (Serranidae), from French Polynesia and the South Pacific Ocean. Can J Zool 75:1818–1827
Rigby MC, Lo CM, Cribb TH, Euzet L, Faliex E, Galzin R, Holmes JC, Morand S (1999) Checklist of the parasites of coral reef fishes from French Polynesia, with considerations on their potential role in these fish communities. Cybium 23:273–284
Ronald K (1959) A checklist of the metazoan parasites of the Heterosomata. Dept Fish Que Contrib 67:1–152
Rudolphi CA (1819) Entozoorum synopsis cui accedunt mantissa duplex et indices locupletissimi. Sumtibus A. Rücker, Berolini
Sadovy Y, Eklund AM (1999) Synopsis of biological data on the Nassau grouper, Epinephelus striatus (Bloch, 1792), and the jewfish, E. itajara (Lichtenstein, 1822). NOAA Tech Rep NMFS-Seattle 146:1–65
Santos MJ, Eiras JC (1995) A seasonal study on the parasitization of Lipophrys pholis (Pisces: Blenniidae) by Helicometra fasciata (Digenea: Opecoelidae) and Lecithochirium furcolabiatum (Digenea: Hemiuridae) in Portugal. Aquac 132:175–181
Sasal P, Faliex E, Morand S (1996) Parasitism of Gobius bucchichii Steindachner, 1870 (Teleostei, Gobiidae) in protected and unprotected marine environments. J Wildl Dis 32:607–613
Sasal P, Niquil N, Bartoli P (1999) Community structure of digenean parasites of sparid and labrid fishes of the Mediterranean Sea: a new approach. Parasitol 119:635–648
Satya Gopal K, Vijayalakshmi C, Hanumantha Rao K (1985) Male reproductive system of some digenetic trematodes. Proc Indian Acad Sci (Anim Sci) 94:145–151
Schell SC (1970) How to know the trematodes. Wm. C. Brown, Dubuque
Schell SC (1973) Three new species of digenetic trematodes from Puget Sound fishes. Proc Helminthol Soc Wash 40:227–230
Schell SC (1985) Handbook of trematodes of North America north of Mexico. Univ. Press of Idaho, Moscow
Schroeder RE (1965) Ecological studies of the intestinal trematodes of the gray snapper, Lutjanus griseus (Linnaeus), in the vicinity of lower Matecumbe Key, Florida. Diss Abstr 25:5460
Schroeder RE (1971) Ecology of the intestinal trematodes of the gray snapper, Lutjanus griseus, near lower Matecumbe Key, Florida, with a description of a new species. In: Starck WA, Schroeder RE (eds) Studies in tropical oceanography No. 10: investigations on the gray snapper, Lutjanus griseus. University of Miami Press, Coral Gables, pp 151–218
Sekerak AD (1975) Parasites as indicators of populations and species of rockfishes (Sebastes: Scorpaenidae) of the Northeastern Pacific Ocean. Dissertation, University of Calgary
Sekerak AD, Arai HP (1973) Helminths of Sebastes alutus (Pisces: Teleostei) from the northeastern Pacific. Can J Zool 51:475–477
Sekerak AD, Arai HP (1974) A revision of Helicometra Odhner, 1902 and related genera (Trematoda: Opecoelidae), including a description of Neohelicometra sebastis n. sp. Can J Zool 52:707–738
Sekerak AD, Arai HP (1977) Some metazoan parasites of rockfishes of the genus Sebastes from the northeastern Pacific Ocean. Syesis 10:139–144
Sey O (1970) Parasitic helminths occurring in Adriatic fishes. Part II (Flukes and tapeworms). Acta Adriat (Split) 13:3–16
Shchepkina AM, Tkachuk LP (2001) Parasite influence on egg lipid content of the Black Sea shrimp Palaemon elegans. In: Sixth National Conference of Parasitology, Sofia, 5–7 October 2001, programs and abstracts. Pensoft, Sofia, p 49
Shen JW (1986) Opecoelidae trematodes of marine fishes from Hainan Island. Oceanol Limnol Sin 17:379–385
Shen JW (1990) Digenetic trematodes of marine fishes from Hainan Island. Science Press, Beijing
Shen JW, Qui ZZ (1995) Studies on the trematodes of fishes from the Yellow Sea and the Bo Hai Sea. Science Press, Beijing
Shul’man SS, Shul’man-Al’bova RE (1953) Parasites of fish of the White Sea. Moskva, Leningrad
Siddiqi AH, Cable RM (1960) Digenetic trematodes of marine fishes of Puerto Rico. Sci Surv P R Virg-Isl 17:257–369
Skrjabin KI (1964) Keys to the trematodes of animals and man. Univ. Illinois Press, Urbana
Skrjabin KI, Koval VP (1958) Subfamily Coitocaecinae Poche, 1926; subfamily Plagioporinae Manter, 1947. In: Skrjabin KI (ed) Trematodes of animals and man, vol 15. Akademiya Nauk, Moscow-Leningrad, pp 329–818
Sogandares-Bernal F (1959) Studies on marine fish trematodes from the gulf of Panama and Bimini, British West Indies. Diss Abstr 20:1–429
Solonchenko AI (1969) A study of the helminth fauna of Symphodus seina from the Crimean shore of the Black Sea. In: Voprosy morskoi biologii Tezisy II vsesoyuznogo symposiuma molodykh uchenykh, Sevastopol’, 1969, pp 118–119 [Abstract]
Sonsino P (1890) Notizie di trematodi della collezione del Museo di Pisa. Proc Verb Soc Tosc Sci Nat 7:137–143
Sonsino P (1891) Parassiti animali del Mugil cephalus e di altri pesci della collezione del Museo di Pisa. Atti Soc Tosc Sci Nat Proc Verb 7:253–265
Sparks AK (1957) Some digenetic trematodes of marine fishes of the Bahama Islands. Bull Mar Sci Gulf Caribb 7:255–265
Sparks AK (1972) Invertebrate pathology. Noncommunicable diseases. Academic, New York
Sproston NG (1938) Notes sur la faune parasitaire des poissons à Roscoff. Trav Stn Biol Roscoff 16:33–58
Stanley RD, Lee DL, Whitaker DJ (1992) Parasites of yellowtail rockfish, Sebastes flavidus (Ayres, 1862) (Pisces: Teleostei), from the Pacific coast of North America as potential biological tags for stock indentification. Can J Zool 70:1086–1096
Stiles CW, Hassall A (1908) Index-catalogue of medical and veterinary zoology. Subjects: Trematoda and trematode diseases. Hygienic Laboratory, Washington
Stone MAB, MacDiarmid SC, Pharo HJ (1997) Import health risk analysis: salmonids for human consumption. Ministry of Agriculture Regulatory Authority, Wellington
Stossich M (1883) Brani di elmintologia tergestina. Serie prima. Boll Soc Adriat Sci Nat Trieste 8:111–121
Stossich M (1885) Brani di elmintologia tergestina. Serie seconda. Boll Soc Adriat Sci Nat Trieste 9:156–164
Stossich M (1886) I distomi dei pesci marini e d’acqua dolce. Tipografia del Lloyd Austro-Ungarico, Trieste, p 66
Stossich M (1887a) Brani di elmintologia tergestina. Serie quarta. Boll Soc Adriat Sci Nat Trieste 10:90–96
Stossich M (1887b) Brani di elmintologia tergestina. Serie quinta. Boll Soc Adriat Sci Nat Trieste 10:184–192
Stossich M (1888) Appendice al mio lavaro. I distomi dei pesci marini e d’acqua dolce. Programma Ginn-Comun Sup, Trieste, p 14
Stossich M (1889) I distomi degli anfibi Lavoro monografico. Boll Soc Adriat Sci Nat 11:60–74
Stossich M (1890) Brani di elmintologia tergestina. Serie settina. Boll Soc Adriat Sci Nat Trieste 12:39–46
Stossich M (1892) Osservazioni elmintologiche. Glas Hrvat Naravosl Druš 7:64–73
Stossich M (1898) Saggio di una fauna elmintologica di Trieste e province contermini. Programma Civ Scuola R Sup, Trieste, p 162
Stossich M (1902) Sopra una nuova specie delle Allocreadiinae. Osservazioni. Arch Parasitol (Paris) 5:578–582
Stossich M (1903a) Una nuova specie de Helicometra Odhner. Arch Parasitol (Paris) 7:373–376
Stossich M (1903b) Note distomologiche. Boll Soc Adriat Sci Nat Trieste 21:193–201
Stossich M (1904a) Sopra alcuni nematodi. Zool Cent 11:246
Stossich M (1904b) Alcuni distomi della collezione elmintologica del museo zoologico di Napoli. Ann Mus Zool R Univ Napoli (NS) 1:1–14
Tantalean VM, Carvajal G, Martinez R, Huiza A (1982) Helmintos parasitos de peces marinos de la costa Peruana. NCTL, Lima
Tantalean VM, Sarmiento BL, Huiza PA (1992) Digeneos (Trematoda) del Peru. Bol Lima 80:47–84
Teresa-Gonzalez M, Henriquez V, Lopez Z (2013) Variations in the fecundity and body size of digenean (Opecoelidae) species parasitizing fishes from Northern Chile. Rev Biol Mar Oceanogr 48:421–429
Ternengo S, Levron C, Mouillot D, Marchand B (2009) Site influence in parasite distribution from fishes of the Bonifacio Strait Marine Reserve (Corsica Island, Mediterranean Sea). Parasitol Res 104:1279–1287
Thapar GS, Dayal J (1934) A new species of the genus Helicometra from the intestine of Trigla gurnardus. Proc 21 Indian Sci Congr, Bombay, January 2–8, p 261
Timon-David J (1937) Etude sur les trématodes parasites des poisons du golfe de Marseille (Première liste). Bull Inst Océanogr, No. 717, 24 pp
Tkachuk L (2000) The influence of environmental factor (pollution) on abundance of the dominant parasite species of shrimp Palaemon elegans Rathke in the Black Sea. Bull Scand Soc Parasitol 10:134
Tkachuk LP, Mordvinova TN (1999) On parasitic infection of the shrimp, Palaemon elegans from two coastal regions of the Black Sea. Ekol Morya (Kiev) 49:21–23
Travassos LP (1922) Informações sôbre a fauna helmintológica de Mato Grosso. Fôlha Méd 3:187–190
Travassos LP (1928) Fauna helmintológica de Matto Grosso (Trematódeos – I.a parte). Mem Inst Oswaldo Cruz 21:309–372
Travassos L, Teixeira de Freitas JF, Kohn A (1969) Trematódeos do Brasil. Mem Inst Oswaldo Cruz 67:1–886
Treasurer JW (1997) Parasites of wrasse (Labridae) in inshore waters of west Scotland stocked as cleaner fish of sea lice (Caligidae) on farmed Atlantic salmon. J Fish Biol 50:895–899
UNESCO (1973) Check-list of fishes of the north-eastern Atlantic and of the Mediterranean (C.L.O.F.N.A.M.). UNESCO, Paris
van Beneden PJ (1871) Les poisons des côtes de Belgiques, leurs parasites et leurs commensaux. Mém Acad Roy Sci Belg 38:1–100
Vassiliades G (1985) Helminthes parasites des poissons de mer des cotes du Senegal. Bull Inst Fondam Afr Noire 44:78–99
Vidal-Martínez VM, Aguirre-Macedo ML, Vivas-Rodriguez CM, Moravec F (1998) Las comunidades de macroparásitos del mero Epinephelus morio (Pisces: Serranidae) en la península de Yucatán, México. Proc Gulf Caribb Fish Inst 50:764–779
Vlasenko PV (1931) Zur Helminthofauna de Schwarzmeerfische [On the parasitic worm fauna of fishes of the Black Sea]. Tr Karadag Biol Stantsii 4:88–136
Wallin IE (1909) A new species of the trematode genus Allocreadium. With a revision of the genus and a key to the sub-family Allocreadiinae. Trans Am Microsc Soc 29:50–66
Wang P (1981) Notes on some trematodes from freshwater fishes in Fujian Province. J Fujian Norm Univ 2:81–90
Wang PQ (1982) Some digenetic trematodes of marine fishes from Fujian Province, China. Oceanol Limnol Sin 13:179–194
Wang YY, Wang PQ, Zhang WH (1992) Opecoelid trematodes of marine fishes from Fujian Province. Wuyi Sci J 9:67–89
Willemoes-Suhm R (1871) Ueber einige Trematoden und Nemathelminthen. Z Wiss Zool 21:175–203
Woolcock V (1935) Digenetic trematodes from some Australian fishes. Parasitology 27:309–331
Yamaguti S (1934) Studies on the helminth fauna of Japan. Part 2. Trematodes of fishes, I. Jpn J Zool 5:249–541
Yamaguti S (1939) Studies on the helminth fauna of Japan. Part 26. Trematodes of fishes, VI. Jpn J Zool 8:211–230
Yamaguti S (1940) Studies on the helminth fauna of Japan. Part 31. Trematodes of fishes, VII. Jpn J Zool 9:35–108
Yamaguti S (1942) Studies on the helminth fauna of Japan. Part 39. Trematodes of fishes mainly from Naha. Trans Biogeogr Soc Jpn 3:329–398
Yamaguti S (1953) Parasitic worms mainly from Celebes. Part 3. Digenetic trematodes of fishes, II. Acta Med Okayama 8:257–295
Yamaguti S (1958a) Systema helminthum. Vol 1. The digenetic trematodes of vertebrates—parts I & II. Interscience, New York
Yamaguti S (1958b) Studies on the helminth fauna of Japan. Part 52. Trematodes of fishes, XI. Publ Seto Mar Biol Lab 6:369–384
Yamaguti S (1970) Digenetic trematodes of Hawaiian fishes. Keigaku Publishing, Tokyo
Yamaguti S (1971) Synopsis of digenetic trematodes of vertebrates. Vols I & II. Keigaku Publishing, Tokyo
Young MR (1939) Helminth parasites of Australia: a bibliography with alphabetical lists of authors, hosts and parasites. Imperial Bureau of Agricultural Parasitology (Helminthology), Winches Farm
Yu SZ, Liu SF (2007) Population ecology of Helicometra execta parasiting in Sebastiscus marmoratus in Taiwan Strait, China. J Oceanogr Taiwan Strait 26:256–260
Zdzitowiecki K (1993) A re-examination of some Antarctic and subantarctic fish digeneans from the collection of the British Museum (Natural History). Acta Parasitol 38:157–160
Zdzitowiecki K (1997a) Antarctic Digenea, parasites of fishes. In: Wägele JW, Sieg J (eds) Synopses of the Antarctic Benthos, vol 8. Koeltz Scientific Books, Koenigstein
Zdzitowiecki K (1997b) Digenea of fishes of the Weddell Sea. IV. Three opecoelid species of the genera Neolebouria, Helicometra and Stenakron. Acta Parasitol 42:138–143
Zdzitowiecki K (1998) Helicometra pisanoae sp. n. (Digenea, Opecoelidae), a parasite of a fish, Trematomus hansoni, in the Eastern Antarctic. Acta Parasitol 43:26–29
Zdzitowiecki K (2001) New data on the occurrence of fish endoparasitic worms off Adelie Land, Antarctica. Pol Polar Res 22:159–165
Zdzitowiecki K (2002) Occurrence of Digenea in fishes of the family Nototheniidae in the Weddell Sea. Acta Parasitol 47:154–158
Zdzitowiecki K, Rocka A, Pisano E, Ozouf-Costaz C (1998) A list of fish parasitic worms collected off Adelie Land (Antarctic). Acta Parasitol 43:71–74
Zdzitowiecki K, Palladino S, Vacchi M (1999) Preliminary results on Digenea found in fishes in the coastal waters of Terra Nova Bay, Antarctica. Parassitologia 41:575–578
Zhang JY, Qiu ZZ, Ding XQ (1999) Parasites and parasitic diseases of fishes. Science Press, Beijing
Zhukov EV (1960) Endoparasitic worms of the fishes in the Sea of Japan and South Kuril shallow-waters. Trudy Zool Inst Leningr 28:3–146
Zongguo H (ed) (2001) Marine species and their distribution in China’s seas. Krieger, Malabar
Zschokke F (1887) Helminthologische Bemerkungen. Mitth Zool Stat Neapel 7:264–271
Acknowledgments
The authors wish to thank Heather L. Prestridge, BRTC/TCWC, Texas A&M University, College Station, TX, USA, for providing NOD access to fish and host collection data. We are grateful to Dr. Mark Pokras and Robert Ertsgard, Cummings School of Veterinary Medicine, and the Tisch Library, Tufts University, Medford, MA, USA, for access to and use of their on-line literature databases. We thank Mrs. Patricia Pilitt, USNPC, for help in depositing specimens. Special appreciation to Lorraine Portch, Library & Archives Division, BM(NH), for help in obtaining literature for H. dalianensis. Final thanks are given to the reviewers who provided helpful comments to improve the manuscript and to Gordon College, Wenham, MA, USA for the loan of microscopes and other equipment to CKB.
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FAUNAL REVIEW OF SPECIES OF THE GENUS HELICOMETRA ODHNER, 1902
FAUNAL REVIEW OF SPECIES OF THE GENUS HELICOMETRA ODHNER, 1902
Helicometra Odhner, 1902 (Digenea: Opecoelidae)
(Syns. Allostenopera Baeva, 1958; Loborchis Lühe in Stossich, 1902; Metahelicometra Yamaguti, 1971; Stenopera Manter, 1933).
Reports: Keys to the genus and species of Helicometra: Wallin (1909), Linton (1910), Isaĭchikov (1928), Palombi (1929a), Manter (1933), Chatterji (1936), Skrjabin and Koval (1958), Yamaguti (1958a, 1970, 1971), Skrjabin (1964), Pritchard (1966), Schell (1970, 1985), Sekerak and Arai (1974), Gibson and Bray (1982), Ramadan (1986), Bray and Cribb (1989), Gibson (1996), Cribb (2005), Aken’Ova et al. (2006) and Hassanine (2007). Synonyms of the genus Helicometra: Skrjabin and Koval (1958), Yamaguti (1958a, 1971), Siddiqi and Cable (1960), Hafeezullah (1971), Sekerak and Arai (1974), Gibson and Bray (1982) and Cribb (2005).
Remarks: Odhner (1902c) created this genus with Helicometra pulchella (Rudolphi, 1819) as type species. Stossich (1903b) considered it to belong within Allocreadiidae Looss, 1902, and Woolcock (1935) included it within Allocreadiinae Looss, 1902 as did Chatterji (1936). Manter (1947, p. 286) created Plagioporinae Manter, 1947 within Opecoelidae Ozaki, 1925 and placed Helicometra and Stenopera in this subfamily where the former genus is considered to belong today (Cribb 2005). Gupta (1956) modified the generic diagnosis of Stenopera which is now considered a junior syn. of Helicometra (see Siddiqi and Cable 1960; Hafeezullah 1971; Sekerak and Arai 1974; Gibson and Bray 1982). The first listings we could locate of Loborchis as a junior syn. of Helicometra were in Skrjabin and Koval (1958) and in Yamaguti (1958a). Sekerak and Arai (1974) supported this same synonymous position for Loborchis and also considered Allostenopera a junior syn. of Neohelicometra Siddiqi and Cable, 1960; furthermore, those species of Helicometra possessing a terminal, funnel-shaped oral sucker were moved into Neohelicometra by the same authors—a move later refuted by Bray (1979). Gibson and Bray (1982, p. 554) later considered Allostenopera a junior synonym of Helicometra. Yamaguti (1971) erected the subgenera Helicometra (Odhner, 1902) and Metahelicometra Yamaguti, 1971; however, neither subgenera were adopted by later authors and the latter subgenus is now considered a junior synonym of Helicometra (see Cribb 2005).
Helicometra sp. of Vlasenko (1931)
Hosts: Labridae: Symphodus tinca (Linnaeus, 1758) (=Crenilabrus tinca [Linnaeus, 1758])—East Atlantic peacock wrasse; Lotidae: Gaidropsarus vulgaris (Cloquet, 1824) (=Onos tricirrata [non Brünnich, 1768])—Three-bearded rockling; Scorpaenidae: Scorpaena porcus Linnaeus 1758—Black scorpionfish.
Site of infection: Intestine.
Distribution: Black Sea (Vlasenko 1931).
Specimens deposited: See Remarks below. We could find no information for deposition of specimens; however, based on Vlasenko (1931, p. 127), the materials for this study were collected at the Karadag Biological Station, located near Feodosiya, Crimea, Ukraine along the Black Sea. Additional work was performed at the Kharkiv branch of the All-Ukrainian Biol.-Zool. Institute for Science and within the Helminthology Dept. of the State/Civil (=“Stattsinst.”) Center for Experimental Veterinary Science, Moscow, Russia.
Remarks: Sekerak and Arai (1974) commented that Vlasenko (1931, p. 115) thought his material resembled H. pulchella of Nicoll (1910), but perhaps Nicoll was incorrect in his identification. In addition, “Vlasenko’s specimens were probably within the concept of H. pulchella, but since they are not available for examination, and no description accompanied the report, their specific identity cannot be determined.” (see Sekerak and Arai 1974, p. 710).
Helicometra sp. of Arai (1969)
(Syns. Helicometra sebastis [Sekerak and Arai, 1974]; Neohelicometra sebastis Sekerak and Arai, 1974).
Host: Sebastidae: Sebastes maliger (Jordan & Gilbert, 1880)—Quillback rockfish.
Site of infection: Intestine.
Distribution: British Columbia, Canada (Arai 1969).
Reports: Love and Moser (1976, 1983).
Specimens deposited: No information from Arai (1969) was provided for deposition of specimens; however, the study was carried out at the Fisheries Research Board of Canada Biological Station, Nanaimo, British Columbia, Canada, and the address for Hisao P. Arai at the time of the study was the Dept. of Biology, University of Calgary, Calgary, Alberta, Canada.
Remarks: Sekerak and Arai (1974, p. 733) reexamined the two specimens of this species reported by Arai (1969) and considered them to be synonymous with Neohelicometra sebastis Sekerak and Arai, 1974. Gibson (1996, p. 170) considered Helicometra sp. of Arai (1969) to be a junior syn. of Helicometra sebastis (Sekerak and Arai, 1974).
Helicometra sp. of Machida et al. (1970)
Host: Sebastidae: Sebastiscus marmoratus (Cuvier, 1829)—False kelpfish.
Sites of infection: Intestine; pyloric caecum.
Distribution: Sea north of Tsushima Islands, Japan (Machida et al. 1970).
Specimens deposited: Materials from this study expected to have been deposited in Dept. of Zoology, National Science Museum, Tokyo, Japan and/or Meguro Parasitological Museum, Tokyo, Japan.
Helicometra sp. of Prudhoe and Bray (1973)
(Syns. Helicometra antarcticae Holloway and Bier, 1968; Helicometra fasciata [Rudolphi, 1819]; Helicometra rakusai Zdzitowiecki, 1997).
Hosts: Nototheniidae: Trematomus hansoni Boulenger, 1902—Striped rockcod; Trematomus loennbergii Regan, 1913 (=Trematomus loennbergi [sic] Regan, 1913)—Scaly rockcod; Trematomus scotti (Boulenger, 1907)—Crowned rockcod.
Site of infection: Intestine.
Distribution: Off Antarctica in far southern Indian Ocean (Prudhoe and Bray 1973).
Specimens deposited: BM(NH) 1975.1.30.29–32.
Remarks: Prudhoe and Bray (1973) note that due to the contracted nature of these worms, no species name was given to these specimens. Later, Bray (1987a) considered this species to be a junior syn. of H. fasciata. Zdzitowiecki (1993) identified these specimens as Helicometra antarcticae Holloway and Bier, 1968 but later (Zdzitowiecki, 1998) suggested they might be H. rakusai Zdzitowiecki, 1997.
Helicometra sp. of Sekerak and Arai (1973)
(Syns. Helicometra sebastis [Sekerak and Arai, 1974]; Neohelicometra sebastis Sekerak and Arai, 1974).
Host: Sebastidae: Sebastes alutus (Gilbert, 1890)—Pacific Ocean perch.
Sites of infection: Intestine; pyloric caecum.
Distribution: Northeastern Pacific Ocean (Sekerak and Arai 1973).
Specimens deposited: No information was given as to deposition of specimens; however, this work was performed at the Pacific Biological Station, Nanaimo, British Columbia, Canada, and the address given for the first author was Dept. of Biology, University of Calgary, Calgary, Alberta, Canada.
Remarks: Sekerak and Arai (1974, p. 733) reexamined their specimens of this species and considered them to be synonymous with Neohelicometra sebastis Sekerak and Arai, 1974. Gibson (1996, p. 170) considered Helicometra sp. of Sekerak and Arai (1973) to be a junior syn. of Helicometra sebastis (Sekerak and Arai, 1974).
Helicometra sp. of Reimer (1974)
Hosts: Myctophidae: Ceratoscopelus maderensis (Lowe, 1839)—Madeira lantern fish; Nemichthyidae: Nemichthys scolopaceus Richardson, 1848—Slender snipe eel.
Site of infection: Stomach.
Distribution: Cape Blanc and English Channel in Atlantic Ocean (Reimer 1974).
Reports: (Klimpel et al. 2001, 2009).
Specimens deposited: As this reference is an abstract from proceedings of the Third International Congress of Parasitology, no information was given on deposition of specimens.
Remarks: Specimens collected were juveniles.
Helicometra sp. of Reimer (1975)
Host: Nemichthyidae: Nemichthys scolopaceus Richardson, 1848—Slender snipe eel.
Site of infection: Digestive tract.
Distribution: Cape Blanc of northwestern Africa in Atlantic Ocean (Reimer 1975).
Reports: (Klimpel et al. 2001, 2009).
Specimens deposited: We were unable to confirm the presence or location of any deposited specimens.
Remarks: Helicometra species collected in this study was preadult.
Helicometra sp. of Gaevskaia and Umnova (1977)
Host: Sebastidae: Sebastes norvegicus (Ascanius, 1772) (=Sebastes marinus [not of Linnaeus, 1758])—Golden redfish.
Site of infection: Intestine.
Distribution: Grand Newfoundland Bank, NW Atlantic Ocean (Gaevskaia and Umnova 1977).
Report: Margolis and Arthur (1979).
Specimens deposited: We could find no information in Gaevskaia and Umnova (1977) on deposition of specimens; however, the address of both authors given for inquiries was Laboratory of Parastology, Atlantic Research Institute of Marine Fisheries and Oceanography, Kaliningrad, Russia.
Helicometra sp. of De Buron and Maillard (1987)
Hosts: Gobiidae: Gobius niger Linnaeus, 1758—Black goby; Zosterisessor ophiocephalus (Pallas, 1814)—Grass goby; Labridae: Symphodus cinereus (Bonnaterre, 1788)—Grey wrasse.
Site of infection: “Digestive tube” = intestine.
Distribution: “Etang de Thau” lagoon basin and Mauguio pond in the Mediterranean Sea in southern France (De Buron and Maillard 1987).
Specimens deposited: We could find no evidence in this publication of where specimens, if any, were deposited; however, we note that the address for both authors at that time was the Laboratoire de Parasitologie Comparée, Université des Sciences et Techniques du Languedoc, place E.-Bataillon, F 34060, Montpellier Cedex, France.
Remarks: The authors comment that several species of Helicometra (H. fasciata [Rudolphi, 1819], H. pulchella [Rudolphi, 1819], and H. gobii [Stossich, 1883]) occur in gobies, wrasse and anguillids within the location of their study; however, they assigned all individuals within this study to “Helicometra sp.” This study also allowed the authors to suggest strongly that the species of Helicometra parasitizing the two goby species (G. niger and Z. ophiocephalus) was not the same species of Helicometra present in the wrasse species (Symphodus cinereus).
Helicometra sp. of Maillard and Aussel (1988)
Intermediate host: Palaemonidae: Palaemon serratus (Pennant, 1777)—Common prawn.
Site of infection: Expected to be thoracic muscles (see Meenakshi et al. 1993).
Distribution: Lagoon of Languedoc (“Etang de Thau”) in southern France (Maillard and Aussel 1988).
Specimens deposited: We could find no evidence in this publication of where specimens, if any, were deposited; however, we note that the address for both authors for inquiries was the Laboratoire de Parasitologie Comparée, Université des Sciences et Techniques du Languedoc, place E.-Bataillon, 34060, Montpellier Cedex, France.
Helicometra spp. of Reversat et al. (1989)
Hosts: Anguillidae: Anguilla anguilla (Linnaeus, 1758)—European eel; Blenniidae: Salaria pavo (Risso, 1810) (=Blennius pavo Risso, 1810)—Peacock blenny; Gobiidae: Gobius niger Linnaeus, 1758—Black goby; Zosterisessor ophiocephalus (Pallas, 1814)—Grass goby; Labridae: Symphodus cinereus (Bonnaterre, 1788)—Grey wrasse.
Site of infection: Intestine.
Distribution: “Etang de Thau” in Hérault, France (Reversat et al. 1989).
Specimens deposited: We could find no evidence in this publication of where specimens, if any, were deposited; however, we note that the address for all authors for inquiries was the Laboratoire de Parasitologie Comparée (CNRS-URA 698), Université des Sciences et Techniques du Languedoc, place Eugène Bataillon, 34060 Monpellier Cedex, France.
Remarks: Reversat et al. (1989) discovered at least 3 “species” of Helicometra; however, they were identified using alleles (not Linnaean taxonomic designations) corresponding to locus Mdh-1 coding for malate dehydrogenase, obtained through enzyme electrophoresis, and are as follows: (1) Mdh-1 a found only in S. cinereus; (2) Mdh-1 c and Mdh-1 d parasitizing G. niger, Z. ophiocephalus, A. anguilla and S. pavo; and (3) Mdh-1 b found in all five host fish studied (S. cinereus, G. niger, Z. ophiocephalus, A. anguilla and S. pavo). Reversat et al. (1991b) later found that allele Mdh-1 a corresponded with H. pulchella, allele Mdh-1 b corresponded with H. gobii and alleles Mdh-1 c and Mdh-1 d corresponded with H. fasciata (see below).
Helicometra spp. of Reversat (1990)
Intermediate hosts: Crangonidae: Crangon crangon (Linnaeus, 1758)—Brown shrimp; Gammaridae: Gammarus insensibilis Stock, 1966—Lagoon sand shrimp; Palaemonidae: Palaemon serratus (Pennant, 1777)—Common prawn.
Site of infection: Expected to be thoracic muscles (see Meenakshi et al. 1993).
Distribution: Lagoon of Languedoc (“Etang de Thau”) in southern France (Reversat 1990).
Specimens deposited: See Remarks below.
Remarks: This reference is the dissertation of Dr. Jerome Reversat of the Laboratoire de Parasitologie Comparée, Université de Montpellier, France.
Helicometra sp. of Shen (1990)
Host: Serranidae: Plectropomus leopardus (Lacepède, 1802)—Leopard coralgrouper.
Site of infection: Expected to be intestine.
Distribution: Off Hainan Island, South China Sea (Shen 1990).
Specimens deposited: WorldCat listed only one location (Natural History Museum, London) that possessed this reference; however, an earlier publication by the same author (Shen 1986) focusing on digenetic trematodes from marine fishes of the same locality (Hainan Island) listed that type specimens were deposited in the Institute of Oceanology, Academia Sinica, China.
Remarks: This is a report of a “larval form” of an unidentified species of Helicometra (Shen 1990).
Helicometra sp. of Dyer et al. (1992)
Host: Mullidae: Pseudupeneus maculatus (Bloch, 1793)—Spotted goatfish.
Site of infection: Intestine.
Distribution: La Parguera, Puerto Rico (Dyer et al. 1992).
Specimen deposited: Voucher: USNPC #81628.
Helicometra sp. of Abollo et al. (1996)
Host: Gobiidae: Gobius paganellus Linnaeus, 1758—Rock goby.
Site of infection: Intestine.
Distribution: Ria of Vigo, Spain, along Western Iberian coast (Abollo et al. 1996).
Specimens deposited: We could find no evidence of where specimens, if any, were deposited; however, we note that the address for all authors for inquiries was the Laboratorio de Parasitología, Facultad de Ciencias del Mar, Universidad de Vigo, 36200 Vigo, Spain.
Remarks: This report is from an abstract included at a conference entitled the VII European Multicolloquium of Parasitology (EMOP VII) held in Parma, Italy (Abollo et al. 1996).
Helicometra sp. of Sasal et al. (1996)
Host: Gobiidae: Gobius bucchichi Steindachner, 1870 (=Gobius bucchichii [sic] Steindachner, 1870)—Bucchich’s goby.
Site of infection: Intestine.
Distribution: In and around the Cerbère-Banyuls Marine Reserve in southeastern France (Sasal et al. 1996).
Specimens deposited: Vouchers: Muséum National d’Histoires Naturelles of Paris, France #494 HF-Tg 112, 495 HF-Tg 113; Symbiotype: Muséum National d’Histoires Naturelles of Paris, France #MNHN 1996-127.
Helicometra spp. of Rigby et al. (1999)
Hosts: Balistidae: Balistapus undulatus (Park, 1797)—Orange-lined triggerfish; Holocentridae: Sargocentron diadema (Lacepède, 1802)—Crown squirrelfish; Sargocentron microstoma (Günther, 1859)—Smallmouth squirrelfish; Sargocentron spiniferum (Forsskål, 1775)—Sabre squirrelfish; Lutjanidae: Lutjanus gibbus (Forsskål, 1775)—Humpback red snapper; Muraenidae: Gymnothorax javanicus (Bleeker, 1859)—Giant moray.
Site of infection: Intestine.
Distribution: Moorea, Society Islands and Rangiroa, Tuamotu Islands, French Polynesia in South Pacific Ocean (Rigby et al. 1999).
Specimens deposited: We could find no evidence in this publication of where specimens, if any, were deposited; however, we noted the following addresses for the authors where inquiries can be made: ETH Zürich, Experimentelle Ökologie, ETH-Zentrum NW, CH-8092 Zürich, Switzerland; CRIOBE, B.P. 1013, Moorea, French Polynesia; Univ. of Queensland, Dept. of Parasitology, Brisbane Qld 4072, Australia; Université des Sciences et Techniques du Languedoc Monpellier 2, Station Méditerranéenne de l’Environnement Littoral, CNRS UMR 5555, 1 Quai de la Daurade, 34200 Sète, France; Université de Perpignan, Centre de Biologie et d’Éecologie tropicale et méditerranéenne, CNRS UMR 5555, Av. de Villeneuve, 66860 Perpignan Cedex, France; École Pratique des Hautes Etudes, CNRS URA 1453, Université de Perpignan, 66860 Perpignan Cedex, France.
Helicometra sp. of Mordvinova (2000)
Host: Myctophidae: Ceratoscopelus maderensis (Lowe, 1839)—Madeira lantern fish.
Site of infection: Expected to be in musculature.
Distribution: World Ocean (Mordvinova 2000)
Report: Klimpel et al. (2009).
Specimens deposited: We were unable to confirm the presence or location of any deposited specimens.
Remarks: This is a report of a juvenile(s) of Helicometra.
Helicometra spp. of Jousson (2001)
Hosts: Labridae: Labrus viridis Linnaeus, 1758—Wrasse sp.; Symphodus rostratus (Bloch, 1791)—Wrasse sp.; Scorpaenidae: Scorpaena scrofa Linnaeus, 1758—Red scorpionfish.
Intermediate hosts: Hippolytidae: Hippolyte inermis Leach, 1816—Seaweed shrimp; Trochidae: Gibbula umbilicalis (da Costa, 1778)—Flat top shell; Jujubinus striatus (Linnaeus, 1758) (=Calliostoma striatum [Linnaeus, 1758])—Grooved top shell.
Site of infection: Expected to be intestine (definitive hosts); expected to be thoracic muscles (shrimp intermediate host—see Meenakshi et al. 1993).
Distribution: Western Mediterranean Sea (Jousson 2001).
Specimens deposited: See Remarks below.
Remarks: This reference is the dissertation of Dr. Olivier Jousson of the Université de Genève, Switzerland.
Helicometra sp. of Mordvinova (2001)
Host: Myctophidae: Myctophidae sp.—Lanternfish sp.
Site of infection: Expected to be in musculature (metacercariae).
Distribution: World Ocean (Mordvinova 2001).
Specimens deposited: As this reference is an abstract from proceedings of the Sixth National Conference of Parasitology, no information was given on deposition of specimens.
Remarks: This report, which is from an abstract in the proceedings of a conference, simply mentions that myctophids, in general, are paratenic hosts for metacercariae of species of Helicometra (Mordvinova 2001).
Helicometra sp. of Ko and Chan (2002)
Host: Sebastidae: Sebasticus marmoratus (Cuvier, 1829)—False kelpfish.
Site of infection: Expected to be intestine.
Distribution: Hong Kong, China (Ko and Chan 2002).
Specimens deposited: We were unable to confirm the presence or location of any deposited specimens.
Helicometra spp.
Hosts: Blenniidae: Lipophrys pholis (Linnaeus, 1758) (=Blennius pholis Linnaeus, 1758)—Shanny; Bothidae: Arnoglossus laterna (Walbaum, 1792)—Mediterranean scaldfish; Carapidae: Echiodon drummondii Thompson, 1837—Pearlfish sp.; Clinidae: Clinus rotundifrons Barnard, 1937 (=Gynutoclinus rotundifrons [Barnard, 1937])—Kelp klipfish; Congridae: Conger conger (Linnaeus, 1758)—European conger; Gobiidae: Gobius niger Linnaeus, 1758—Black goby; Gobius paganellus Linnaeus, 1758—Rock goby; Labridae: Labridae sp.—Wrasse sp.; Labrus bergylta Ascanius, 1767—Ballan wrasse; Thalassoma lunare (Linnaeus, 1758)—Moon wrasse; “Swallow-tailed wrasse”; Lutjanidae: Lutjanus synagris (Linnaeus, 1758)—Lane snapper; Pinguipedidae: Parapercis hexophtalma (Cuvier, 1829) (=Parapercis hexophthalmus [sic] [Cuvier, 1829])—Speckled sandperch; Platycephalidae: Platycephalus bassensis Cuvier, 1829—Sand flathead; Pomacentridae: Hypsypops rubicundus (Girard, 1854) (=Hypsypos [sic] rubicundus [Girard, 1854])—Garibaldi damselfish; Psychrolutidae: Cottunculus microps Collett, 1875—Polar sculpin; Sebastidae: Helicolenus dactylopterus (Delaroche, 1809)—Blackbelly rosefish; Serranidae: Epinephelus guttatus (Linnaeus, 1758)—Red hind; Epinephelus striatus (Bloch, 1792)—Nassau grouper; Sparidae: Pachymetopon blochii (Valenciennes, 1830)—Hottentot seabream; Triglidae: Chelidonichthys cuculus (Linnaeus, 1758) (=Aspitrigla cuculus [Linnaeus, 1758])—Red gurnard; Eutrigla gurnardus (Linnaeus, 1758) (=Trigla gurnardus Linnaeus, 1758)—Grey gurnard; Zoarcidae: Lycodes esmarkii Collett, 1875—Greater eelpout; Lycodes sp.—Eelpout sp.
Sites of infection: Intestine; “mid gut”; pyloric caecum.
Distribution: New York Aquarium (Coll: MacCallum); La Jolla, California (Coll: Young); Queensland, Australia (Coll: Manter); Bahamas (Sogandares-Bernal); Additional localities obtained from BM(NH) collections: Bay of Biscay; East Greenland; Essex and Plymouth, England; Northeast Atlantic Ocean; South Africa; South Australia; St. Kilda, Scotland.
Specimens deposited: Vouchers: USNPC #36411, 48970; HWML #2306–2308, 2381, 44284, 44296, 44322, 44438; BM(NH) #1963.6.20.4; 1982.6.9.80–101; 1982.11.5.3–4,23; 1983.7.28.74–75; 1987.9.18.55; 1988.10.24.2–7; 1989.1.5.15; 1989.8.31.12.
Remarks: Our searchings of the USNPC and HWML databases revealed two and eight deposited specimens, respectively, from unidentified species of Helicometra; all were collected by individuals with no indication of subsequent publication(s). Our searchings of the BM(NH) holdings revealed 38 slides of unidentified species of Helicometra collected from 17 host spp; we were unable to determine if these materials were subsequently published.
Helicometra aegyptense Hassanine, 2007
Host: Holocentridae: Sargocentron spiniferum (Forsskål, 1775)—Sabre squirrelfish [type host].
Site of infection: Upper intestine.
Distribution: Red Sea off coast of Sharm El-Sheikh, South Sinai, Egypt (Hassanine 2007).
Specimens deposited: Holotype and paratypes: Helminthological Coll. of Red Sea Fishes, Marine Science Dept., Faculty of Science, Suez Canal University, Ismailia, Egypt #2006.2.25.1–4 and 2006.2.25.5–10.
Remark: Hassanine (2007) placed this species in Group III.
Helicometra antarcticae Holloway and Bier, 1968
(Syns. Helicometra sp. Prudhoe and Bray, 1973; Neohelicometra antarcticae [Holloway and Bier, 1968] [see Hassanine 2007, p. 22]).
Hosts: Nototheniidae: Dissostichus eleginoides Smitt, 1898—Patagonian toothfish; Dissostichus mawsoni Norman, 1937—Antarctic toothfish [type host]; Trematomus hansoni Boulenger, 1902 (=Pagothenia hansoni [Boulenger, 1902])—Striped rockcod; Trematomus loennbergii Regan, 1913 (=Trematomus loennbergi [sic] Regan, 1913)—Scaly rockcod; Trematomus scotti (Boulenger, 1907)—Crowned rockcod; Zoarcidae: Lycodichthys dearborni (DeWitt, 1962) (=Rhigophila dearborni DeWitt, 1962)—Eelpout sp.
Sites of infection: Intestine; pyloric caecum.
Distribution: McMurdo Sound, Antarctica (Holloway and Bier 1968); Antarctic Ocean (Zdzitowiecki 1993, 1997a); Ross Sea, Antarctica (Brickle et al. 2005).
Report: Klimpel et al. (2009).
Specimens deposited: Holotype and paratype: USNPC #63113, 63114; Vouchers: BM(NH) #1975.1.30.29–32.
Remarks: This species is included in key of Sekerak and Arai (1974), who transferred it to Neohelicometra Siddiqi and Cable, 1960. Bray (1979), noting that fixation method produced variability in the terminal vs subterminal appearance of the oral sucker (the main feature used in the key by Sekerak and Arai [1974] to distinguish Neohelicometra and Helicometra), returned N. antarcticae to Helicometra. Zdzitowiecki (1993) identified specimens of Helicometra sp. of Prudhoe and Bray (1973) as this species but later (Zdzitowiecki, 1998) believed this same H. sp. to be also similar to H. rakusai Zdzitowiecki, 1997. Hassanine (2007) placed this species in Group IV.
Helicometra aposinuata Pritchard, 1966
(Syns. Helicometra [Helicometra] aposinuata Pritchard, 1966; Helicometra pulchella [Rudolphi, 1819]).
Hosts: Holocentridae: Neoniphon aurolineatus (Liénard, 1839) (=Flammeo scythrops Jordan and Evermann, 1903; Holocentrus scythrops [Jordan and Evermann, 1903])—Yellowstriped squirrelfish [type host]; Sargocentron spiniferum (Forsskål, 1775) (=Holocentrus spinifer [Forsskål, 1775])—Sabre squirrelfish; “Holocentrid”; Labridae: Thalassoma duperrey (Quoy & Gaimard, 1824) (=Thalassoma duperreyi [Quoy & Gaimard, 1824])—Saddle wrasse; Serranidae: Epinephelus awoara (Temminck and Schlegel, 1842)—Yellow grouper.
Site of infection: Intestine.
Distribution: Hawaii (Pritchard 1966; Yamaguti 1970); South China Sea (Wang et al. 1992).
Reports: Yamaguti (1971), Zhang et al. (1999), Cribb et al. (2002).
Specimens deposited: Holotype: USNPC #60354; Paratypes: HWML #1639–1641.
Remarks: Included in key to Helicometra species in Hawaiian fishes by Yamaguti (1970). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella.
Helicometra bassensis Woolcock, 1935
(Syn. Helicometra [Helicometra] bassensis Woolcock, 1935).
Host: Platycephalidae: Platycephalus bassensis Cuvier, 1829—Sand flathead [type host].
Site of infection: Intestine.
Distribution: Port Philip Bay, Victoria, Australia (Woolcock 1935); Darwent Estuary, Tasmania (Crowcroft 1947a).
Reports: Young (1939), Yamaguti (1958a, 1971).
Specimens deposited: Whole-mount specimens from P.W. Crowcroft’s Collection #TMAG No. Rg15–Rg20.
Remarks: This species is included in keys of Skrjabin and Koval (1958), Skrjabin (1964) and Sekerak and Arai (1974). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group II.
Helicometra borneoensis Fischthal and Kuntz, 1965
(Syn. Helicometra [Helicometra] borneoensis Fischthal and Kuntz, 1965).
Host: Serranidae: Epinephelus fasciatus (Forsskål, 1775)—Blacktip grouper [type host].
Site of infection: Small intestine.
Distribution: Jesselton (=Kota Kinabalu), North Borneo (Fischthal and Kuntz 1965).
Reports: Yamaguti (1971) and Cribb et al. (2002).
Specimen deposited: Holotype: USNPC #60079.
Remarks: Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. This species is included in key of Sekerak and Arai (1974). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra boseli Nagaty, 1956
(Syn. Helicometra equilata [Manter, 1933]; Helicometra nasae Nagaty and Abdel Aal, 1962; Stenopera boseli [Nagaty, 1956]).
Hosts: Acanthuridae: Naso unicornis (Forsskål, 1775)—Bluespine unicornfish; Holocentridae: Neoniphon sammara (Forsskål, 1775) (=Holocentrus sammara [Forsskål, 1775]; Holocentrus samara [sic] [Forsskål, 1775]; Egyptian name for species is “boseli”)—Sammara squirrelfish [type host]; Sargocentron punctatissimum (Cuvier, 1829) (=Holocentrus lacteoguttatus Cuvier, 1829)—Speckled squirrelfish; Sargocentron spiniferum (Forsskål, 1775) (=Holocentrus spinifer [Forsskål, 1775])—Sabre squirrelfish; Sargocentron xantherythrum (Jordan and Evermann, 1903) (=Holocentrus xantherythrus Jordan and Evermann, 1903)—Hawaiian squirrelfish.
Site of infection: Intestine.
Distribution: Hurghada, Egypt, Red Sea (Nagaty 1956b); Hawaii (Pritchard 1966; Yamaguti 1970).
Report: Yamaguti (1971).
Specimens deposited: Paratype and voucher: USNPC #59554, 60355; Paratype: HWML #1584; Vouchers: HWML #1646–1648; Type materials are in the Parasitology Dept., Faculty of Medicine, Abbasia, Cairo, Egypt.
Remarks: Pritchard (1966, p. 197–198) disagreed with Siddiqi and Cable (1960), who synonymized Stenopera with Helicometra, and re-established Stenopera as a valid genus moving H. boseli in to Stenopera (validity of this species in Stenopera supported by Yamaguti [1971]); however, later authors still consider this genus as synonymous with Helicometra. This species is included in key of Pritchard (1966) and a key to Helicometra species in Hawaiian fishes by Yamaguti (1970). Sekerak and Arai (1974) believed this species to be a junior syn. of H. equilata. Ramadan (1986) considered H. nasae to be a junior syn. of H. boseli and included the latter species in a key to the species of Helicometra from Red Sea fishes. Aken’Ova et al. (2006) placed this species in Group III.
Helicometra colligata (Wallin, 1909)
(Syn: Allocreadium colligatum Wallin, 1909; Allocreadium (Allocreadium) colligatum Wallin, 1909; Distomum fasciatum Rudolphi of Stossich, 1885, 1886, 1898 [see Palombi 1929a; Yamaguti 1971, p. 134]; Peracreadium fasciatum [Stossich, 1885] [see Palombi 1929a]).
Host: Labridae: Labrus mixtus Linnaeus, 1758—Cuckoo wrasse [type host].
Site of infection: Intestine.
Distribution: Unknown. Host species distributed though out NE Atlantic Ocean and Mediterranean Sea (Froese and Pauly 2013).
Reports: Wallin (1909), Nicoll (1915) and Isaĭchikov (1928).
Specimens deposited: We could find no evidence as to the exact location(s) of deposited specimens. However, Wallin (1909, p. 63) stated that his “study was carried out in the Zoological Laboratory of the University of Nebraska” and he wanted “to express [his] sincere thanks to Prof. Henry B. Ward for … the preserved materials …”
Remarks: Palombi (1929a) transferred this species to Peracreadium Nicoll, 1909. Yamaguti (1958a, p. 102) listed this species as Allocreadium colligatum Wallin, 1909 from the stomach of the fallfish, Semotilus corporalis (Mitchill, 1817) (=Semotilus buliaria [sic] [Rafinesque, 1817]; Cyprinidae) collected from Sebago Lake, Maine (freshwater). Yamaguti (1971, p. 134) listed this species as Allocreadium (Allocreadium) colligatum Wallin, 1909 also from the stomach of S. corporalis (=S. bullaris) from Sebago Lake, Maine; however, he noted that the “generic identity” of this species was “not verified yet.”
Helicometra dalianensis (Li, Qui and Zhang, 1989)
(Syn. Neohelicometra dalianensis Li, Qui and Zhang, 1989 [see Hassanine 2007, p. 22]).
Host: Hexagrammidae: Hexagrammos otakii Jordan and Starks, 1895—Greenling sp. [type host].
Site of infection: Intestine.
Distribution: Yellow Sea and Bo-Hai Sea (Li et al. 1989).
Reports: Shen and Qui (1995), Liu et al. (2010).
Specimens deposited: Dept. of Biology, Nankai Univ., Tianjin, China Holotype 1, Paratype 2.
Remarks: This species was transferred by Cribb (2005, p. 471) from Neohelicometra Siddiqi and Cable, 1960 to Helicometra as it possesses caeca that end blindly rather than open via separate ani. Hassanine (2007) placed this species in Group IV.
Helicometra dochmosorchis Manter and Pritchard, 1960
(Syns. Helicometra [Helicometra] dochmosorchis Manter and Pritchard, 1960; Helicometra pulchella [Rudolphi, 1819]).
Hosts: Cirrhitidae: Paracirrhites arcatus (Cuvier, 1829)—Arc-eye hawkfish; Paracirrhites forsteri (Schneider, 1801)—Blackside hawkfish; Muraenidae: Gymnothorax buroensis (Bleeker, 1857)—Vagrant moray [type host]; Scorpaenidae: Sebastapistes ballieui (Sauvage, 1875) (=Scorpaena ballieui Sauvage, 1875)—Spotfin scorpionfish.
Site of infection: Intestine.
Distribution: Hawaii (Manter and Pritchard 1960; Pritchard 1966).
Report: Yamaguti (1971).
Specimens deposited: Holotype: USNPC #39157; Vouchers: HWML #1642–1644.
Remarks: Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella.
Helicometra dorabi (Mamaev and Oshmarin, 1966)
(Syn. Stenopera dorabi Mamaev and Oshmarin, 1966).
Host: Chirocentridae: Chirocentrus dorab (Forsskål, 1775)—Dorab wolf-herring [type host].
Intermediate host: Clupeidae: Sardinella sp.—Sardinella sp.
Site of infection: Stomach (definitive host).
Distribution: North Vietnam Bay (Mamaev and Oshmarin 1966).
Specimens deposited: Authors stated that this reference provided descriptions of new or little known Acanthocolpidae Lühe, 1909 from the collection of helminths obtained in 1960–1961 from an expedition to Vietnam by the Pacific Research Institute of Fisheries and Oceanography (TINRO).
Remarks: We became aware of this taxon via Zoological Records, which listed Stenopera dorabi Mamaev and Oshmarin, 1966 parasitizing C. dorab and Sardinella sp. (encysted metacercariae infecting the gills of latter host) from Vietnam with the source reference as Mamaev and Oshmarin (1966, p. 160–163). We obtained this reference and found no mention of Stenopera (=Helicometra) dorabi; furthermore, p. 160–163 described Stephanostomoides dorabi Mamaev and Oshmarin, 1966 from the same hosts, locality and infection sites (see Satya Gopal et al. [1985, p. 149] who described the male reproductive system of Stephanostomoides dorabi citing Mamaev and Oshmarin [1966]). As Stephanostomoides Mamaev and Oshmarin, 1966 is within Acanthocolpidae (see Yamaguti 1971, p. 127; Bray 2005, p. 608) and not Opecoelidae and because we are neither aware of another species with this taxon name nor found any other reference(s) for Stenopera dorabi, we believe this name cannot be accepted and declare it a nomen nudum.
Helicometra epinepheli Yamaguti, 1934
(Syns. Helicometra [Helicometra] epinepheli Yamaguti, 1934; Helicometra pulchella [Rudolphi, 1819]).
Hosts: Balistidae: Rhinecanthus aculeatus (Linnaeus, 1758) (=Balistes aculeatus Linnaeus, 1758)—White-banded triggerfish; Labridae: Anampses caeruleopunctatus Rüppell, 1829—Bluespotted wrasse; Thalassoma purpureum (Forsskål, 1775)—Surge wrasse; Sebastidae: Sebastes inermis Cuvier, 1829 (=Sebastodes inermis Cuvier, 1829)—Dark-banded rockfish; Serranidae: Epinephelus akaara (Temminck and Schlegel, 1842)—Hong Kong grouper [type host]; Epinephelus fasciatus (Forsskål, 1775) (=Epinephelus tsirimenara [Temminck and Schlegel, 1842])—Blacktip grouper; Epinephelus merra Bloch, 1793 (=Serranus merra [Bloch, 1793])—Honeycomb grouper; Family?: “Zeus sp. with longitudinal stripes”.
Site of infection: Intestine.
Distribution: Inland Sea of Japan (Yamaguti 1934); Hamazima, Japan (Yamaguti 1940); Naha and Tokushima, Japan (Yamaguti 1942); Makassar and Celebes, South Sulawesi, Indonesia (Yamaguti 1953); Japan (Yamaguti 1958b); Fujian Province, China (Wang 1982); Al-Ghardaga, Red Sea (Ramadan 1986); South China Sea (Wang et al. 1992); New Caledonia (Bray and Justine 2007).
Reports: Yamaguti (1958a, 1971), Pan et al. (1990), Zhang et al. (1999), Zongguo (2001), Cribb et al. (2002) and Quilichini et al. (2011).
Specimens deposited: Holotype and paratypes: Yamaguti (1934, p. 301) stated “type and paratypes in my collection” and now at Meguro Parasitological Museum, Tokyo, Japan, Holotype: MPM Coll. #22253; Paratypes: MPM Coll. #22253, 22310; HWML #1830; Vouchers: Helminthology Collection, Zoology Dept., Faculty of Science, Ain Shams University (#348, 349).
Remarks: This species is included in keys of Skrjabin and Koval (1958) and Skrjabin (1964). Manter (1954), Dawes (1956), Durio and Manter (1968) and Bray (1979; see Bray and Cribb 1989, p. 448) considered this species a junior syn. of H. fasciata, while Sekerak and Arai (1974) considered this species a junior syn. of H. pulchella. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors, and noted that while Durio and Manter (1968) synonymized H. epinepheli with H. fasciata, there is a “definite difference in egg length” between the two species. Ramadan (1986) included this species in a key to the species of Helicometra from Red Sea fishes.
Helicometra equilata (Manter, 1933)
(Syn: Helicometra boseli Nagaty, 1956 [see Sekerak and Arai 1974; p. 727; Hassanine 2007, p. 22]; Helicometra nasae Nagaty and Abdel Aal, 1962 [see Hassanine 2001]; Stenopera equilata Manter, 1933 [see Hassanine 2007, p. 22]).
Hosts: Holocentridae: Holocentrus adscensionis (Osbeck, 1765) (=Holocentrus ascensionis [Osbeck, 1765])—Squirrelfish [type host]; Holocentrus rufus (Walbaum, 1792)—Longspine squirrelfish; Neoniphon marianus (Cuvier, 1829) (=Holocentrus marianus Cuvier, 1829)—Longjaw squirrelfish.
Sites of infection: Intestine; pyloric caecum.
Distribution: Tortugas, Florida (Manter 1933, 1947); Bahamas (Sparks 1957); Near Cat Cay & N. Bimini, Bahamas, British West Indies (Sogandares-Bernal 1959); Mona Island, Puerto Rico (Siddiqi and Cable 1960); Jamaica (Nahhas and Cable 1964); Caribbean Sea (Drowned Cays) off Belize (Fischthal 1977, 1978); Mario Island (Dyer et al. 1985); Grand Cayman, Cayman Islands, British West Indies (Nahhas 1993); Caribbean Sea (Nahhas and Carlson 1994).
Reports: Galtsoff (1954), Skrjabin and Koval (1958), Yamaguti (1958a [Pl. 35, Fig. 455], 1971 [Fig. 421]) and Overstreet et al. (2009).
Specimens deposited: “Type” and Vouchers: USNPC #29956, 39358, 74239, 77783, 82475; Syntype: HWML# 267; Vouchers: HWML #266, 22538, 22539, 35210, 124177.
Remarks: Manter (1933) erected Stenopera with Stenopera equilata Manter, 1933 as type species; however, this genus is now considered a junior synonym of Helicometra (see Siddiqi and Cable 1960, p. 299). Pritchard (1966, p. 198) disagreed and re-established Stenopera as a valid genus returning H. equilata to Stenopera (validity of Stenopera supported by Yamaguti [1958a, 1971]); however, later authors still consider this genus as synonymous with Helicometra. This species is included in keys of Pritchard (1966) and Sekerak and Arai (1974). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group III.
Helicometra execta Linton, 1910
(Syns. Helicometra [Helicometra] execta Linton, 1910; Helicometrina execta [Linton, 1910]; Helicometrina parva Manter, 1933; Helicometrina trachinoti Siddiqi and Cable, 1960).
Hosts: Blenniidae: Scartella cristata (Linnaeus, 1758) (=Blennius cristatus Linnaeus, 1758)—Molly miller; Carangidae: Trachinotus falcatus (Linnaeus, 1758)—Permit; Gobiidae: Bathygobius soporator (Valenciennes, 1837)—Frillfin goby; Haemulidae: Anisotremus virginicus (Linnaeus, 1758)—Porkfish; Haemulon plumierii (Lacepède, 1801) (=Haemulon plumieri [Lacepède, 1801])—White grunt [type host]; Haemulon sciurus (Shaw, 1803)—Bluestriped grunt; Labridae: Doratonotus megalepis Günther, 1862—Dwarf wrasse; Halichoeres bivittatus (Bloch, 1791) (=Iridio bivittatus [Bloch ?])—Slippery dick; Halichoeres pictus (Poey, 1860)—Rainbow wrasse; Halichoeres poeyi (Steindachner, 1867) (=Halichoeres kirschii [Evermann and Jordan, 1898]; Iridio kirschii Evermann and Jordan, 1898)—Blackear wrasse; Halichoeres radiatus (Linnaeus, 1758) (=Iridio radiatus Jordan and Evermann, 1896)—Puddingwife wrasse; Lachnolaimus maximus (Walbaum, 1792)—Hogfish; Thalassoma bifasciatum (Bloch, 1791) (=Chlorichthys bifasciatus ?)—Bluehead; Labrisomidae: Labrisomus bucciferus Poey, 1868—Puffcheek blenny; Labrisomus haitiensis Beebe and Tee-Van, 1928 (=Labrisomis [sic] haitensis Beebe and Tee-Van, 1928)—Longfin blenny; Labrisomus kalisherae (Jordan, 1904)—Downy blenny; Lutjanidae: Lutjanus griseus (Linnaeus, 1758)—Grey snapper; Malacanthidae: Malacanthus plumieri (Bloch, 1786)—Sand tilefish; Muraenidae: Gymnothorax vicinus (Castelnau, 1855)—Purplemouth moray; Sciaenidae: Pareques umbrosus (Jordan and Eigenmann, 1889) (=Eques acuminatus [non Bloch and Schneider, 1801]; Pareques acuminatus [non Bloch and Schneider, 1801])—Cubbyu; Sebastidae: Sebastiscus marmoratus (Cuvier, 1829)—False kelpfish; Serranidae: Mycteroperca bonaci (Poey, 1860)—Black grouper; Mycteroperca venenosa (Linnaeus, 1758)—Yellowfin grouper; Synodontidae: Saurida elongata (Temminck and Schlegel, 1846)—Slender lizardfish.
Sites of infection: Intestine; pyloric caecum; rectum.
Distribution: Tortugas, Florida (Linton 1910; Manter 1933, 1947); North Bimini, Bahamas, British West Indies (Sogandares-Bernal 1959); Jamaica (Nahhas and Cable 1964); Lower Matecumbe Key, Florida (Schroeder 1965); Biscayne Bay, Florida (Overstreet 1969); Near lower Matecumbe Key, Florida (Schroeder 1971); Bahamas (Yamaguti 1971); Taiwan Strait (Wang et al. 1992; Yu and Liu 2007).
Reports: Galtsoff (1954), Yamaguti (1958a, 1971), Zhang et al. (1999); Overstreet et al. (2009—as Helicometrina execta [Linton, 1910]); Liu et al. (2010).
Specimens deposited: Syntypes: USNPC #08446; Vouchers: USNPC #08447–08449; HWML #254–256, 258, 259, 262, 263, 1827, 22532, 22533.
Remarks: This species is included in keys of Linton (1910), Palombi (1929a), Manter (1933), Skrjabin and Koval (1958), and Skrjabin (1964). Overstreet (1969, p. 144) transferred this species to Helicometrina Linton, 1910. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors, and he commented on (1) the moving of this species to Helicometrina by Overstreet (1969, p. 144) based on the “great variability in the number of testes (one to nine) and low host-specificity” of this species and (2) the synonymization of H. parva and H. trachinoti with H. execta by Overstreet (1969) “in spite of the fact that the egg length is 42–54 μm in execta but 54–59 μm in parva and 56–70 μm in trachinoti.” As can be understood in part from above, Yamaguti (1971) did not support the taxonomic changes of Overstreet (1969), choosing to keep this species in Helicometra and recognizing neither the synonymies of Helicometrina parva Manter, 1933 nor Helicometrina trachinoti Siddiqi and Cable, 1960 with H. execta. It is worth noting that Cribb (2005) distinguishes species of Helicometra from Helicometrina based, in part, on the number of testes (consistently two vs. three to nine, rarely two or none) and that this species was not recognized by Aken’Ova et al. (2006) and Hassanine (2007).
Helicometra fasciata (Rudolphi, 1819)
(Syns: Allocreadium fasciatum [Rudolphi, 1819] Odhner, 1902 [see Odhner 1901; Palombi 1929a]; Bacciger bacciger [Rudolphi, 1819] of Palombi 1928 [see Palombi 1934a]; Distoma fasciatum Rudolphi, 1819 [see Rudolphi 1819; Dujardin 1845; Olsson 1868; van Beneden 1871; Willemoes-Suhm 1871; Palombi 1929a]; Distomum [Dicrocoelium] fasciatum [Rudolphi, 1819] Barbagallo and Drago, 1903 [see Barbagallo and Drago 1903; Palombi 1929a]; Distomum fasciatum Rudolphi, 1819 nec Stossich, 1885 [see Diesing 1850; Carus 1884; Parona 1887; Sonsino 1890; Monticelli 1893; Palombi 1929a]; Distomum gobii Stossich, 1883 [see Stossich 1883, 1886, 1890; Carus 1884; Sonsino 1891; Odhner 1901; Palombi 1929a]; Distomum palaemonis Linstow, 1877 [cercaria—see Palombi 1929a, p. 259, 1934b, p. 86; Yamaguti 1958a, p. 109]; Helicometra sp. Prudhoe and Bray, 1973; Helicometra aposinuata Pritchard, 1966 [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra dochmosorchis Manter and Pritchard, 1960 [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra epinepheli Yamaguti, 1934 [see Durio and Manter 1968, p. 748; Fischthal and Thomas 1972, p. 315; Madhavi 1975, p. 159; Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra filamentosa Madhavi, 1975; Helicometra [Helicometra] fasciata [Rudolphi, 1819]; Helicometra flava Stossich, 1903 [see Stossich 1903a, 1904b; Wallin 1909; Palombi 1929a, b; Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra gobii [Stossich, 1883] [see Stossich 1904b; Wallin 1909; Palombi 1929a, b; Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra hypodytis Yamaguti, 1934 [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra indica Agrawal, 1964; Helicometra labri [Stossich, 1886] [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra markewitschi Pogorel’tseva, 1954 [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra marmoratae Nagaty and Abdel Aal, 1962 [see Hassanine 2007, p. 20, 22]; Helicometra mutabilis [Stossich, 1902] [see Stossich 1904b; Wallin 1909; Palombi 1929a, b; Akmirza 2004, p. 37; Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra pulchella [Rudolphi, 1819] [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra pulchella of Nicoll 1910 [see Nicoll 1910; Palombi 1929a; Yamaguti 1971, p. 190]; Helicometra scorpaenae Prudhoe and Bray, 1973 nec Wang, 1982; Helicometra scorpaenae Wang, 1982 nec Prudhoe & Bray, 1973; Helicometra sinuata [Rudolphi, 1819] [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Helicometra upapula Yamaguti, 1970 [see Aken’Ova et al. 2006, p. 21; Hassanine 2007, p. 22]; Loborchis fasciatum [Rudolphi, 1819]; Loborchis mutabilis Stossich, 1902 [see Stossich 1902; Palombi 1929a]; “MDH- 1c” and “MDH- 1d” [see Reversat et al. 1989, 1991b]; From Palombi 1929a, p. 260—see Odhner 1902c; Stossich 1904b; Stiles and Hassall 1908; Wallin 1909; Palombi 1929b).
Hosts: Anguillidae: Anguilla anguilla (Linnaeus, 1758) (=Anguilla vulgaris Rafinesque, 1810; Muraena anguilla Linnaeus, 1758)—European eel; Anarhichadidae: Myoxocephalus scorpius (Linnaeus, 1758)—Shorthorn sculpin; Antennariidae: Antennarius striatus (Shaw, 1794)—Striated frogfish; Apogonidae: Apogon limenus Randall and Hoese, 1988—Sydney cardinalfish; Ariidae: Galeichthys feliceps Valenciennes, 1840 (=Arius feliceps [Valenciennes, 1840])—White barbel; Balistidae: Balistes capriscus Gmelin, 1789 (=Balistes carolinensis Gmelin, 1789)—Grey triggerfish; Blenniidae: Aidablennius sphynx (Valenciennes, 1836) (=Blennius sphynx Valenciennes, 1836)—Combtooth blenny sp.; Lipophrys pholis (Linnaeus, 1758) (=Blennius pholis Linnaeus, 1758)—Shanny; Parablennius gattorugine (Linnaeus, 1758) (=Blennius gattorugine Linnaeus, 1758)—Tompot blenny; Parablennius sanguinolentus (Pallas, 1814) (=Blennius sanguinolentus Pallas, 1814)—Rusty blenny; Parablennius tentacularis (Brünnich, 1768) (=Blennius tentacularis Brünnich, 1768)—Tentacled blenny; Salaria pavo (Risso, 1810) (=Blennius pavo Risso, 1810)—Peacock blenny; Bothidae: Bothus podas (Delaroche, 1809)—Wide-eyed flounder; Carangidae: Carangoides plagiotaenia Bleeker, 1857 (=Caranx compressus Day, 1871)—Barcheek trevally; Cheilodactylidae: Chirodactylus brachydactylus (Cuvier, 1830)—Two-tone fingerfin; Cichlidae: Cichlasoma bimaculatum (Linnaeus, 1758) (=Labrus bimaculatus Linnaeus, 1758)—Black acara; Clinidae: Cirrhibarbis capensis Valenciennes 1836 (=Clinus [Cirrhibarbis] capensis Valenciennes, 1836)—Barbelled klipfish; Clinus cottoides Valenciennes, 1836 (=Clinus [Clinus] cottoides Valenciennes, 1836)—Bluntnose klipfish; Clinus superciliosus (Linnaeus, 1758) (=Clinus [Clinus] superciliosus [Linnaeus, 1758])—Clinid sp.; Clinus rotundifrons Barnard, 1937 (=Gynutoclinus rotundifrons [Barnard, 1937])—Kelp klipfish; Xenopoclinus kochi Smith, 1948—Platanna klipfish; Xenopoclinus leprosus Smith, 1961—Leprous platanna-klipfish; Congridae: Conger conger (Linnaeus, 1758)—European conger; Cottidae: Taurulus bubalis (Euphrasen, 1786) (=Cottus bubalis Euphrasen, 1786)—Longspined bullhead; Gadidae: Microgadus tomcod (Walbaum, 1792)—Atlantic tomcod; Raniceps raninus (Linnaeus, 1758)—Tadpole fish; Gobiesocidae: Lepadogaster candolii Risso, 1810 (=Lepadogaster decandollei Risso, 1827)—Connemarra clingfish; Lepadogaster lepadogaster (Bonnaterre, 1788) (=Lepadogaster gouanii Risso, 1810)—Shore clingfish; Gobiidae: Bathygobius soporator (Valenciennes, 1837)—Frillfin goby; Gobius cobitis Pallas, 1814—Giant goby; Gobius niger Linnaeus, 1758 (=Gobius jozo Linnaeus, 1758; Gobius niger jozo Linnaeus, 1758)—Black goby; Gobius paganellus Linnaeus, 1758—Rock goby; Mesogobius batrachocephalus (Pallas, 1814) (=Gobius batrachocephalus Pallas, 1814)—Knout goby; Neogobius melanostomus (Pallas, 1814) (=Gobius cephalarges Pallas, 1814; Gobius exanthematicus [sic] [=exanthematosus] Pallas, 1814; Gobius melanostomus Pallas, 1814)—Round goby; Proterorhinus marmoratus (Pallas, 1814)—Tubenose goby; Zosterisessor ophiocephalus (Pallas, 1814) (=Gobius ophiocephalus Pallas, 1814)—Grass goby; Haemulidae: Pomadasys olivaceus (Day, 1875) (=Pomadasys olivaceum Day, 1875)—Olive grunt; Holocentridae: Myripristis murdjan (Forsskål, 1775)—Pinecone soldierfish; Kyphosidae: Microcanthus strigatus (Cuvier, 1831)—Stripey; Labridae: Cheilinus abudjubbe Rüppell, 1835 (=Cheilinius [sic] abudjubbe Rüppell, 1835)—Wrasse sp.; Cheilinus lunulatus (Forsskål, 1775)—Broomtail wrasse; Coris batuensis (Bleeker, 1856)—Batu coris; Coris julis (Linnaeus, 1758) (=Julis giofredi Risso, 1826)—Mediterranean rainbow wrasse; Ctenolabrus rupestris (Linnaeus, 1758)—Goldsinny wrasse; Labridae sp.—Wrasse sp.; Labrus bergylta Ascanius, 1767 (=Labrus maculatus Bloch, 1792)—Ballan wrasse; Labrus merula Linnaeus 1758—Brown wrasse; Labrus mixtus Linnaeus, 1758—Cuckoo wrasse; Labrus viridis Linnaeus, 1758 (=Labrus turdus Linnaeus, 1758)—Wrasse sp.; Oxycheilinus digramma (Lacepède, 1801) (=Cheilinus diagrammus [Lacepède, 1801])—Cheeklined wrasse; Pseudolabrus guentheri Bleeker, 1862—Günther’s wrasse; Symphodus cinereus (Bonnaterre, 1788) (=Crenilabrus cinereus [Bonnaterre, 1788]; Crenilabrus griseus [Gmelin, 1789]; Symphodus griseus Gratzianov, 1907?)—Grey wrasse; Symphodus doderleini Jordan, 1890 (=Crenilabrus doderleini [Jordan, 1890])—Wrasse sp.; Symphodus melanocercus (Risso, 1810) (=Crenilabrus coeruleus [non Ascanius, 1772])—Wrasse sp.; Symphodus melops (Linnaeus, 1758) (=Crenilabrus melops [Linnaeus, 1758]; Labrus melops Linnaeus, 1758)—Corkwing wrasse; Symphodus ocellatus (Linnaeus, 1758) (=Crenilabrus ocellatus [Forsskål, 1775])—Wrasse sp.; Symphodus roissali (Risso, 1810) (=Crenilabrus quinquemaculatus Risso, 1827)—Five-spotted wrasse; Symphodus rostratus (Bloch, 1791) (=Crenilabrus scina [Forsskål, 1775]; Symphodus scina [Forsskål, 1775])—Wrasse sp.; Symphodus tinca (Linnaeus, 1758) (=Crenilabrus tinca [Linnaeus, 1758]; Labrus tinca Linnaeus, 1758)—East Atlantic peacock wrasse [type host]; Thalassoma lunare (Linnaeus, 1758)—Moon wrasse; Thalassoma pavo (Linnaeus, 1758) (=Crenilabrus pavo [Linnaeus, 1758])—Ornate wrasse; Thalassoma rueppellii (Klunzinger, 1871) (=Thalassoma klunzingeri Fowler and Steinitz, 1956)—Klunzinger’s wrasse; Labrisomidae: Labrisomus nuchipinnis (Quoy & Gaimard, 1824)—Hairy blenny; Labrisomus philippii (Steindachner, 1866) (=Labrisomus philippi [sic] [Steindachner, 1866])—Chalapo clinid or “trambollo”; Lotidae: Ciliata mustela (Linnaeus, 1758)—Fivebeard rockling; Gaidropsarus mediterraneus (Linnaeus, 1758)—Shore rockling; Gaidropsarus vulgaris (Cloquet, 1824) (=Onos tricciratus [sic] [non Brünnich, 1768])—Three-bearded rockling; Lutjanidae: Lutjanus agennes Bleeker, 1863 (=Lutjanus modestus Bleeker, 1863)—African red snapper; Mugilidae: Mugil cephalus Linnaeus, 1758—Flathead grey mullet; Mullidae: Mullus barbatus barbatus Linnaeus, 1758—Red mullet; Muraenidae: Gymnothorax porphyreus (Guichenot, 1848) (=Gymnothorax wieneri Sauvage, 1883)—Lowfin moray; Neosebastidae: Neosebastes thetidis (Waite, 1899)—Thetis fish; Ophidiidae: Ophidion barbatum Linnaeus, 1758—Snake blenny; Paralichthyidae: Pseudorhombus arsius (Hamilton, 1822)—Largetooth flounder; Pholidae: Pholis gunnellus (Linnaeus, 1758)—Rock gunnel; Pinguipedidae: Parapercis nebulosa (Quoy and Gaimard, 1825)—Barred sandperch; Pleuronectidae: Platichthys flesus (Linnaeus, 1758) (=Platichthys flesus luscus [Pallas, 1814])—European flounder; Pleuronectes platessa Linnaeus, 1758—European plaice; Pomacentridae: Abudefduf saxatilis (Linnaeus, 1758) (=Glyphidodon saxatilis [Linnaeus, 1758])—Sergeant-major; Pomatomidae: Pomatomus saltatrix (Linnaeus, 1766) (=Pomatomus saltator [sic] [Linnaeus, 1766])—Bluefish; Pristigasteridae: Ilisha africana (Bloch, 1795) (=Ilisha melanota Derscheid, 1924)—West African ilisha; Sciaenidae: Cheilotrema fasciatum Tschudi, 1846 (=Sciaena fasciata [Tschudi, 1846])—Arnillo drum; Menticirrhus ophicephalus (Jenyns, 1840)—Snakehead kingcroaker; Paralonchurus peruanus (Steindachner, 1875)—Peruvian banded croaker; Pteroscion peli (Bleeker, 1863) (=Larimus peli Bleeker, 1863)—Boe drum; Sciaena deliciosa (Tschudi, 1846)—Lorna drum; Sciaena umbra Linnaeus, 1758—Brown meager; Stellifer minor (Tschudi, 1846)—Minor stardrum; Scombridae: Scomber scombrus Linnaeus, 1758—Atlantic mackerel; Scophthalmidae: Zeugopterus punctatus (Bloch, 1787)—Topknot; Scorpaenidae: Scorpaena notata Rafinesque, 1810 (=Scorpaena ustulata Lowe, 1841)—Small red scorpionfish; Scorpaena porcus Linnaeus, 1758—Black scorpionfish; Scorpaena scrofa Linnaeus, 1758—Red scorpionfish; Scorpaenopsis cirrosa (Thunberg, 1793) (=Scorpaenopsis cirrhosus [sic] [Thunberg, 1793])—Weedy stingfish; Scorpaenopsis gibbosa (Bloch and Schneider, 1801) (=Scorpaenopsis gibbosus [sic] [Bloch and Schneider, 1801])—Humpbacked scorpionfish; Sebastapistes sp. (=Sebastispistes [sic] sp.)—Scorpionfish sp.; Sebastidae: Helicolenus dactylopterus (Delaroche, 1809)—Blackbelly rosefish; Serranidae: Epinephelus akaara (Temminck and Schlegel, 1842)—Hong Kong grouper; Epinephelus awoara (Temminck and Schlegel, 1842)—Yellow grouper; Epinephelus fasciatus (Forsskål, 1775)—Blacktip grouper; Epinephelus merra Bloch, 1793—Honeycomb grouper; Epinephelus quoyanus (Valenciennes, 1830)—Longfin grouper; Epinephelus summana (Forsskål, 1775)—Summan grouper; Epinephelus sp. (=Epinephelis [sic] sp.)—Grouper sp.; Paralabrax auroguttatus Walford, 1936—Goldspotted sand bass; Paralabrax humeralis (Valenciennes, 1828) (=Palabrax [sic] humeralis [Valenciennes, 1828])—Peruvian rock seabass; Paralabrax maculatofasciatus (Steindachner, 1868)—Spotted sand bass; Plectropomus maculatus (Bloch, 1790) (=Plectropomus maculata [sic] [Bloch, 1790])—Spotted coralgrouper; Serranus atricauda Günther, 1874—Blacktail comber; Serranus cabrilla (Linnaeus, 1758) (=Serranellus cabrilla Linnaeus, 1758)—Comber; Serranus hepatus (Linnaeus, 1758)—Brown comber; Serranus scriba (Linnaeus, 1758) (=Perca marina Linnaeus, 1758; Serranus seriba Linnaeus, 1758)—Painted comber; Soleidae: Microchirus ocellatus (Linnaeus, 1758)—Foureyed sole; Microchirus variegatus (Donovan, 1808) (=Solea variegata [Donovan, 1808])—Thickback sole; Monochirus hispidus Rafinesque, 1814—Whiskered sole; Pegusa lascaris (Risso, 1810) (=Solea lascaris [Risso, 1810])—Sand sole; Solea solea (Linnaeus, 1758) (=Solea vulgaris Quensel, 1806)—Common sole; Sparidae: Acanthopagrus australis (Günther, 1859)—Surf bream; Dentex dentex (Linnaeus, 1758)—Common dentex; Pachymetopon blochii (Valenciennes, 1830)—Hottentot seabream; Rhabdosargus sarba (Forsskål, 1775)—Goldlined seabream; Tetrarogidae: Centropogon australis (White, 1790)—Fortescue; Trachichthyidae: Hoplostethus mediterraneus mediterraneus Cuvier, 1829 (=Hoplostethus mediterraneus Cuvier, 1829)—Mediterranean slimehead; Trachinidae: Trachinus draco Linnaeus, 1758 (=Trachinus lineatus Bloch and Schneider, 1801)—Greater weever; Triglidae: Bellator militaris (Goode and Bean, 1896)—Horned searobin; Chelidonichthys capensis (Cuvier, 1829) (=Trigla capensis Cuvier, 1829)—Cape gurnard; Chelidonichthys cuculus (Linnaeus, 1758) (=Aspitrigla cuculus [Linnaeus, 1758]; Trigla pini Bloch, 1793)—Red gurnard; Chelidonichthys lucerna (Linnaeus, 1758) (=Trigla corax Bonaparte, 1834; Trigla lucerna Linnaeus, 1758)—Tub gurnard; Chelidonichthys obscurus (Walbaum, 1792) (=Aspitrigla obscura [Bloch and Schneider, 1801]; Trigla obscura Walbaum, 1792)—Longfin gurnard; Eutrigla gurnardus (Linnaeus, 1758)—Grey gurnard; Lepidotrigla cavillone (Lacepède, 1801) (=Trigla aspera Cuvier, 1829)—Large-scaled gurnard; Prionotus alatus Goode and Bean, 1883—Spiny searobin; Prionotus carolinus (Linnaeus, 1771)—Northern searobin; “Prionodes sp.”—Searobin sp.; Trigla lyra Linnaeus, 1758—Piper gurnard; Trigloporus lastoviza (Bonnaterre, 1788) (=Trigla lineata Gmelin, 1789)—Streaked gurnard; Tripterygiidae: Tripterygion tripteronotum (Risso, 1810) (=Tripterygion tripteronotus [sic] [Risso, 1810])—Threefin blenny sp.; Uranoscopidae: “Unidentified stargazer”; Zeidae: Zeus faber Linnaeus, 1758—John dory; Miscellaneous: “Flounder”; “Unknown host”.
Intermediate hosts: Hippolytidae: Eualus cranchii (Leach, 1817 [in Leach, 1815–1875]) (=Hippolyte cranchii Leach, 1817; Spirontocaris [=Hyppolyte] cranchii [Leach, 1817])—Humpback prawn sp.; Palaemonidae: Palaemon adspersus (Rathke, 1837) (=Leander squilla Linnaeus, 1758; Palaemon squilla [Linnaeus, 1758])—Baltic prawn; Palaemon elegans Rathke, 1837—Grass prawn; Palaemon serratus (Pennant, 1777) (=Leander serratus Pennant, 1777)—Common prawn; Palaemon sp.—Prawn sp.; Palaemon xiphias Risso, 1816 (=Leander xiphias [Scheer, 1960])—Posidonia prawn; Trochidae: Gibbula adansonii (Payreadeau, 1826) (=Gibbula adansoni [sic] [Payreadeau, 1826])—Marine gastropod sp.; Gibbula adriatica (Philippi, 1844)—Marine gastropod sp.
Sites of infection: Intestine; pyloric caecum; rectum; stomach (definitive hosts); expected to be thoracic muscles (decapod intermediate hosts—see Meenakshi et al. 1993).
Distribution: Gulf of Naples off SW coast of Italy (Rudolphi 1819; Palombi 1929a, b, 1931, 1934b); Gulf of Genoa (Willemoes-Suhm 1871); Trieste, Italy (Stossich 1885, 1892; Palombi 1929b); Italy (Collection of the Museum of Pisa) (Sonsino 1890); Unknown locality (Braun 1893; Monticelli 1893); Catania, Sicily (Barbagallo and Drago 1903); Plymouth (Baylis and Jones 1933); Tortugas, Florida (Manter 1933, 1934, 1947); Banyula in “Pyrénées orientales” (southern France) (Mathias 1934); Gulf of Marseille, France (Timon-David 1937); Roscoff in NW France (Sproston 1938); S. Devon, British Isles (Baylis 1939); Santa Maria and Tenacatita Bay, Mexico (Manter 1940); Sevastopol’, Ukraine, Black Sea (Osmanov 1940; Naĭdenova 1966); Beaufort, North Carolina (Hopkins 1941); Aberystwyth, Wales in U.K. (Rees 1945); Northumberland, Cullercoats, England (Crofton 1947); Tasmanian waters (Crowcroft 1947a); Black Sea (Dolgikh and Naĭdenova 1968; Naĭdenova and Dolgikh 1969; Machkevsky 1990; Gaevskaia 1995, 2002; Machkevsky et al. 1996, 1997; Korniychuk 1997, 1999, 2000a, b, 2001a, b, 2008; Machkevsky and Gaevskaia 1997; Korniychuk and Gaevskaia 1999; Tkachuk and Mordvinova 1999; Tkachuk 2000; Shchepkina and Tkachuk 2001); Split, Yugoslavia (=Croatia), Adriatic Sea (Janiszewska 1953; Sey 1970); Stazione Zoologica, Naples, Italy and Lerner Marine Lab, Bimini, Bahamas, British West Indies (De Giusti and Nasir 1960); Vlorë, Albania (Ergens 1960); Baja California, México (Arai 1963); South Atlantic (Parukhin 1966, 1968, 1989); New Caledonia (Durio and Manter 1968); Crimean shore of Black Sea, U.S.S.R. (Solonchenko 1969; Gaevskaia 2002); Cape Coast, Elmina, Tema of Ghana (Fischthal and Thomas 1970); Sevastopol’, Ukraine and Karadag, Turkey, Black Sea (Nikolaeva and Solonchenko 1970); Red Sea (Parukhin 1970); Ghana and Senegal (Fischthal 1972); Gorée, Senegal (Fischthal and Thomas 1972); Peruvian coast (Mateo 1972); Bay of Biscay (Bray 1973); Alboran Sea in western Meditteranean (Lopez-Roman and Guevara Pozo 1974, 1977); Waltair Coast, Bay of Bengal, India (Madhavi 1975); Saronicos Gulf, Athens, Greece (Papoutsoglou 1976); Golfe du Lion in Méditerranée Occidentale (Western Mediterranean Sea) (Maillard and Lambert 1978); Taiwan Strait (Wang 1982; Wang et al. 1992); Atlantic coast of Africa off Morocco (Aleshkina and Gaevskaia 1985); II Región, Chile (Oliva and Muñoz 1985); Northeast Atlantic Ocean (Senegal) (Vassiliades 1985); Blue Hole, Port Elizabeth, eastern Cape Province, Oudekrall, western Cape Province, South Pier, Durban, Natal (=Kwazulu-Natal) Province, South Africa (Bray 1987a); Mediterranean Sea (Orecchia et al. 1988; Parukhin 1989; Naĭdenova and Mordvinova 1997); Heron and Fairfax Islands off Queensland, Australia (Bray and Cribb 1989); Western Pacific Ocean (Heron Island) (Lester and Sewell 1989); Adriatic Sea (Radujkovic and Raibaut 1989; Hristovski and Jardas 1991; Di Cave et al. 2001; Paradižnik and Radujković 2007); Kotor, Yugoslavia (=Montenegro) (Radujkovic et al. 1989); Central American and Mexican Pacific coasts (Campos et al. 1990); Coast of Valencia, Spain (Campos et al. 1990; Campos and Carbonell 1994); Off coast of Chorrillos, Peru (Oliva et al. 1990; Iannacone et al. 2010, 2011); Etang de Thau, France in Mediterranean Sea (Reversat 1990; Reversat and Silan 1991, 1993; Reversat et al. 1991a, b); Bahia de Chorrillos, Peru and central Peruvian coast (Luque et al. 1991; Luque and Oliva 1993a, b; Luque 1994; Oliva and Luque 1998, 2002); Off Mudanya Coast, Sea of Marmara, Turkey (Oguz 1995; Oguz and Bray 2006); Foz do Douro on west coast of Portugal (Santos and Eiras 1995); Western coast of British Isles (Ireland, Scotland) (Costello et al. 1996); Gulf of Palermo, Tyrrhenian Sea (Arculeo et al. 1997); Villefranche Bay and Saint-Jean-Cap-Ferrat, southeast France (Bartoli and Boudouresque 1997); Comacchio lagoons, northern Adriatic Sea, Italy (Dezfuli et al. 1997, 2014); Off coast of Queensland, Australia and South Pacific Isles (Rigby et al. 1997); Western Mediterranean (Jousson et al. 1999; Sasal et al. 1999; Culurgioni et al. 2006); Off coast of northwest Spain (Paniagua et al. 1999); Gökçeada, Turkey (Akmirza 2000, 2001, 2004, 2012); Arousa and Ferrol estuaries in northwest Spain (Outeiral et al. 2001); Southern Iberian Peninsula coasts (Lozano et al. 2002); Natural Marine Reserve ‘Island of Ustica’, South Tyrrhenian Sea (Mediterranean) (Ortis et al. 2002); Chorrillos Fish Market, Lima, Peru (Iannacone 2004); Scandola Nature Reserve of Corsica, France (Bartoli et al. 2005); Off Tangalooma and Stradbroke Island, Central Moreton Bay, Queensland and off New Brighton, New South Wales, Australia (Aken’Ova et al. 2006); Gulf of Aqaba, Red Sea (El-Labadi et al. 2006); Lizard Island, Great Barrier Reef, Australia (Muñoz and Cribb 2006); Ria de Aveiro, Portugal (Hermida et al. 2008); Cerbicales Islands, Bonifacio Strait Marine Reserve, Corsica Island (France) in Mediterranean Sea (Ternengo et al. 2009); Bay of Bizerte, Tunisia (Abdalah et al. 2010); Northern coast of Chile (Teresa-Gonzalez et al. 2013); Additional localities obtained from BM(NH) collections: Cardigan Bay off Wales; County Wexford and Galway, Ireland; Greenland; London Zoo; St. Kilda, Scotland; Off coast proximate to York, England; Sweden.
Reports: Odhner (1902c), Stossich (1902), Nicoll (1915), Palombi (1929b, 1937, 1938), Galtsoff (1954), Yamaguti (1958a, 1971), Love and Moser (1976, 1983—erroneously as Helicometrina fasciata), Pan et al. (1990), Pozdnyakov (1990), Cordero del Campillo et al. (1994), Zhang et al. (1999), Bona et al. (2000), Koubkova and Barus (2000), Klimpel et al. (2001, 2009), Cribb et al. (2002), Levron et al. (2003), Dezfuli et al. (2008), Overstreet et al. (2009) and Liu et al. (2010).
Specimens deposited: Vouchers: BM(NH) #1932.11.22.44; 1952.12.5.176–179; 1963.6.20.1; 1972.3.27.17; 1973.5.17.72; 1976.4.8.176–178; 1978.7.18.4–6; 1979.1.12.16–18; 1980.7.24.11–12; 1982.4.6.73; 1982.6.9.102–114; 1982.11.5.3–4,23; 1982.12.14.51–54; 1983.1.13.16–17; 1986.1.13.6,16–22; 1986.4.3.39; 1986.8.7.10 (“Scotia” 1985 Cruise); 1987.11.11.47–50; 1988.3.14.39 (“Scotia” 1987 Cruise); 1988.9.9.21–22 (C.A. Wright Coll.); 1989.4.13.16; 1990.10.19.40–41; 1992.7.1.284–285; 2005.3.11.1–5; USNPC #51659, 67915, 67945, 68044, 68050, 70681–70686, 71896; HWML #260, 261, 1831, 1832, 1932, 2304, 2305, 2309–2312; 216129–216133; Australian Helm. Coll., South Australian Museum, Adelaide, Australia #10065; Heron Island Workshop #A14, D77; Queensland Museum, Brisbane, Australia #GL 11760; QM G222965–222980; Thomas Cribb Coll. #596; Colección Helmintológica de la Universidad Ricardo Palma, Lima, Perú #CHURP N:587 (see Luque and Oliva 1993b, p. 276); Gaevskaia (2002, p. 221) stated that specimens of this species were kept in the Institute of Medicine, National Academy of Sciences, Sevastopol, Ukraine.
Remarks: This cosmopolitan and widely-reported species was first described by Rudolphi (1819) as Distoma fasciatum Rudolphi, 1819 from the intestine of the East Atlantic peacock wrasse (type host), Symphodus tinca (Linnaeus, 1758) (=Labrus tinca Linnaeus, 1758), the corkwing wrasse, Symphodus melops (Linnaeus, 1758) (=Labrus melops Linnaeus, 1758), and the painted comber, Serranus scriba (Linnaeus, 1758) (=Perca marina Linnaeus, 1758) from Naples, Italy (see Palombi 1929a, p. 244; Bray and Cribb 1989, p. 448). This species was in Loborchis; however, this genus is now considered a junior synonym of Helicometra (see Skrjabin and Koval 1958; Yamaguti 1958a). This species is included in keys of Wallin (1909), Isaĭchikov (1928), Palombi (1929a); Manter (1933), Skrjabin and Koval (1958) and Skrjabin (1964). Palombi (1929a) concluded that H. gobii, H. mutabilis, H. flava and H. pulchella of Nicoll (1910—specimens collected from Millport, a town on the island of Great Cumbrae in the Firth of Clyde off coast of North Ayrshire, Scotland) were junior synonyms of H. fasciata; furthermore, Palombi (1929b) reported H. fasciata in the helminthological collection of Stossich, who previously had identified his material as H. sinuata (see Sekerak and Arai 1974, p. 715). Based on variation in testes texture (lobed to entire), Palombi (1931) regarded H. pulchella and H. sinuata as junior synonyms of H. fasciata. Palombi (1934a) commented that he erroneously reported in 1928 this species as Bacciger bacciger (Rudolphi, 1819). Manter (1954) and Dawes (1956) considered H. hypodytis to be a junior syn. of H. fasciata, while Dawes (1956), Durio and Manter (1968), Fischthal and Thomas (1972) and Madhavi (1975) considered H. epinepheli to be a junior syn. also of H. fasciata. Naĭdenova and Dolgikh (1969) stated that H. markewitschi was a junior syn. of H. fasciata, while Sekerak and Arai (1974) believed H. fasciata to be a junior syn. of H. pulchella—this latter synonymy was refuted by Bray (1979) (see H. pulchella below). Naĭdenova (1974) considered H. pulchella to be junior syn. of H. fasciata. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Bray (1987a, p. 1068–1069) considered H. indica, H. scorpaenae Prudhoe & Bray, 1973 nec Wang, 1982, Helicometra sp. Prudhoe & Bray, 1973, H. filamentosa and H. scorpaenae Wang, 1982 nec Prudhoe & Bray, 1973 all to be junior syns. of H. fasciata; furthermore, he hypothesized that H. fasciata may represent a species complex whose members are indistinguishable morphologically and/or that “clines” may occur in this form. This species is included in a key to opecoelids from fishes of the southern Great Barrier Reef (Bray and Cribb 1989). Meenakshi et al. (1993) considered H. fasciata of Madhavi (1975) to be a junior syn. of H. gibsoni Meenakshi, Madhavi and Swarnakumari, 1993. Aken’Ova et al. (2006) placed H. fasciata in Group I and did not see evidence for a species-complex in their specimens from Australian waters; they suspected a single widely-distributed species with very low host specificity rather than a species-complex. Furthermore, while H. fasciata is a “morphologically variable form, not all the forms that fall within its range of variation are necessarily of the same species” and “it seems likely that ultimately they [species of H. fasciata from various locations worldwide] will prove to represent a different taxon” (Aken’Ova et al. 2006). Hassanine (2007) recognized H. fasciata as type species and also in Group I.
Helicometra filamentosa Madhavi, 1975
(Syn. Helicometra fasciata [Rudolphi, 1819]).
Host: Lutjanidae: Lutjanus sp. (=Lutianus [sic] sp.)—Snapper sp. [type host].
Site of infection: Intestine.
Distribution: Waltair Coast, Bay of Bengal, India (Madhavi 1975).
Report: Madhavi (2011).
Specimen deposited: Holotype: USNPC #73301.
Remarks: Bray (1987a) considered this species to be a junior syn. of H. fasciata. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra flava Stossich, 1903
(Syns. Helicometra fasciata [Rudolphi, 1819] [see Yamaguti 1971, p. 190]; Helicometra [Helicometra] flava Stossich, 1903; Helicometra pulchella [Rudolphi, 1819]).
Host: Serranidae: Serranus hepatus (Linnaeus, 1758) (=Centropristis hepatus [Linnaeus, 1758])—Brown comber [type host].
Site of infection: Intestine.
Distribution: Trieste in NE Italy (Stossich 1903a); Mediterranean Sea (Reversat et al. 1991b).
Reports: Palombi (1929a, b), Yamaguti (1958a, 1971) and Bona et al. (2000).
Specimens deposited: We could find no information regarding deposition of specimens from the original description of H. flava by Professor Michele Stossich (Stossich 1903a). Reversat et al. (1991b, p. 154, Table VII) recently noted this species, and based on this reference, inquiries can be made at the Université Montpellier II, Sciences et Techniques de Languedoc, Laboratoire de Parasitologie Comparée, URA CNRS 698, place Eugène Battaillon, 34095 Montpellier Cedex 5, France.
Remarks: This species is included in keys of Wallin (1909) and Isaĭchikov (1928). Palombi (1929a) and Skrjabin and Koval (1958) considered this species a junior syn. of H. fasciata while Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors.
Helicometra fusiformis Wang, Wang and Zhang, 1992
Host: Hapalogenyidae: Hapalogenys mucronatus (Eydoux and Souleyet, 1850)—Barbeled grunter sp. [type host].
Site of infection: Intestine.
Distribution: Off Fujian Province, China in Taiwan Strait (Wang et al. 1992).
Reports: Zhang et al. (1999); Liu et al. (2010).
Specimens deposited: Type specimens deposited in the Parasitological Research Laboratory, Fujian Teachers University, China.
Remarks: Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra gibsoni Meenakshi, Madhavi and Swarnakumari, 1993
(Syn: Helicometra fasciata of Madhavi, 1975 [see Madhavi 2011, p. 195]).
Hosts: Scorpaenidae: Scorpaenopsis cirrosa (Thunberg, 1793) (=Scorpaenopsis cirrhosis [sic] [Thunberg, 1793]; Scorpaenopsis cirrhosus [sic] [Thunberg, 1793])—Weedy stingfish [type host]; Terapontidae: Therapon jarbua (Forsskål, 1775)—Jarbua terapon (experimental infection).
Intermediate hosts: Alpheidae: Alpheus malabaricus (Fabricius, 1775) (=Alphaeus [sic] malabaricus [Fabricius, 1775])—Shooting shrimp; Columbellidae: Anachis terpsichore (Sowerby, 1822)—Intertidal snail sp.
Sites of infection: Intestine (definitive hosts); thoracic muscles (shrimp intermediate host).
Distribution: Lawson’s Bay Coast, Bay of Bengal, India (Meenakshi et al. 1993).
Report: Madhavi (2011).
Specimen deposited: Holotype: BM(NH) #1992.4.30.1.
Remarks: Meenakshi et al. (1993) stated that the adult of H. gibsoni was morphologically identical with H. fasciata; however, both species were differentiated on the basis of first intermediate host specificity and cercarial morphology (first intermediate host of H. gibsoni = A. terpsichore and cercaria with long extensile tail and four pairs of penetration glands; first intermediate host of H. fasciata = Gibbula adansonii [Payreadeau, 1826] [=Gibbula adamsoni Payr, 1899] and cercaria with short stumpy tail and cluster of penetration glands). Meenakshi et al. (1993) also considered H. fasciata of Madhavi (1975; host = S. cirrosa; locality = Bay of Bengal, India) to be junior syn. of H. gibsoni. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra gobii (Stossich, 1883)
(Syns: Distoma gobii Stossich, 1883; Distomum gobii Stossich, 1883; Helicometra fasciata [Rudolphi, 1819] [see Yamaguti 1971, p. 190]; Helicometra [Helicometra] gobii (Stossich, 1883); Helicometra pulchella [Rudolphi, 1819]; Loborchis gobii [Stossich, 1883]; “MDH- 1b” [see Reversat et al. 1989, 1991b]).
Hosts: Anguillidae: Anguilla anguilla (Linnaeus, 1758)—European eel; Gobiidae: Gobius niger Linnaeus, 1758 (=Gobius jozo Linnaeus, 1758)—Black goby [type host]; Zosterisessor ophiocephalus (Pallas, 1814)—Grass goby; Labridae: Symphodus cinereus (Bonnaterre, 1788)—Grey wrasse; Triglidae: Chelidonichthys lucerna (Linnaeus, 1758) (=Trigla corax Bonaparte, 1834)—Tub gurnard.
Intermediate host: Trochidae: Jujubinus striatus (Linnaeus, 1758) (=Calliostoma striatum [Linnaeus, 1758])—Grooved top shell.
Site of infection: Intestine (definitive hosts).
Distribution: Trieste in NE Italy (Stossich 1883, 1904a); Lagoon of Languedoc (“Etang de Thau”) in southern France (Maillard and Aussel 1988); Etang de Thau, France in Western Mediterranean Sea (Reversat 1990; Reversat and Silan 1991, 1993; Reversat et al. 1991a, b).
Reports: Stossich (1902), Palombi (1929a), Yamaguti (1958a, 1971) and Bona et al. (2000).
Specimens deposited: We could find no information regarding deposition of specimens from the description of H. gobii (as Distomum gobii Stossich, 1883) provided by Professor Michele Stossich (Stossich 1883). Reversat et al. (1991b, p. 154, Table IV) performed isoenzyme electrophoresis on this species (i.e. allele MDH-1b) collected from four species of fish from Etang de Thau, France, and based on this reference, inquiries for deposited materials can be made at the Université Montpellier II, Sciences et Techniques de Languedoc, Laboratoire de Parasitologie Comparée, URA CNRS 698, place Eugène Battaillon, 34095 Montpellier Cedex 5, France.
Remarks: This species was in Loborchis; however this genus is now considered a junior synonym of Helicometra (see Skrjabin and Koval 1958; Yamaguti 1958a). This species is included in keys of Wallin (1909) and Isaĭchikov (1928). Palombi (1929a) considered this species a junior syn. of H. fasciata. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella.
Helicometra gomphosi Yamaguti, 1970
(Syn. Helicometra [Helicometra] gomphosi Yamaguti, 1970).
Host: Labridae: Gomphosus varius Lacepède, 1801—Bird wrasse [type host].
Site of infection: Intestine.
Distribution: Hanauma Bay, Oahu, Hawaii (Yamaguti 1970).
Report: Yamaguti (1971).
Specimen deposited: Holotype: USNPC #63579; Paratype: Meguro Parasitological Museum, Tokyo, Japan, MPM Coll. #15023.
Remarks: Included in key to Helicometra species in Hawaiian fishes by Yamaguti (1970). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. This species is included in key of Sekerak and Arai (1974). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra grandora Manter, 1954
(Syns: Helicometra [Helicometra] grandora Manter, 1954; Helicometra magnora Manter, 1954).
Hosts: Scorpaenidae: Scorpaena papillosa (Schneider and Forster, 1801) (=Helicolenus papillosus [Schneider and Forster, 1801])—Red rock cod; Sebastidae: Helicolenus percoides (Richardson and Solander, 1842)—Red gurnard perch; Triglidae: Chelidonichthys kumu (Cuvier, 1829)—Bluefin gurnard [type host].
Site of infection: Intestine.
Distribution: Portobello and Wellington in New Zealand (Manter 1954).
Reports: Yamaguti (1958a, 1971), Hewitt and Hine (1972) and Stone et al. (1997).
Specimens deposited: Holotype: USNPC #49135; Paratypes: Meguro Parasitological Museum, Tokyo, Japan, MPM Coll. #22250; HWML #1217, 1218, 124227.
Remarks: This species is included in keys of Skrjabin and Koval (1958), Skrjabin (1964), and Sekerak and Arai (1974). Holloway and Bier (1968, p. 31) considered H. magnora to be a junior syn. of H. grandora stating that “Manter (1954) employed two names for a single taxon in the genus, Helicometra grandora and Helicometra magnora. The former is more prominent [used in description on p. 518–519], whereas the latter was employed in the explanation of the figures [Figs. 41, 42]. After consultation (H. W. Manter, personal communication) and for reasons previously considered, Helicometra magnora is synonymized with Helicometra grandora.” Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra gurnardus Thapar and Dayal, 1934
(Syn. Helicometra [Helicometra] gurnardus Thapar and Dayal, 1934).
Host: Triglidae: Eutrigla gurnardus (Linnaeus, 1758) (=Trigla gurnardus Linnaeus, 1758)—Grey gurnard [type host].
Site of infection: Intestine.
Distribution: Type locality unknown. Host species distributed though out coastal areas of NE Atlantic Ocean (Norway to Morocco, Madeira, and Iceland), Mediterranean and Black Seas (Froese and Pauly 2013).
Reports: Thapar and Dayal (1934), Yamaguti (1958a, 1971).
Specimens deposited: See Remarks below.
Remarks: This species is a nomen nudum in that it was first erected by Thapar and Dayal (1934) in an abstract published in the conference proceedings for the 21st Indian Science Congress. This view was supported by Manter (1954, p. 519) and by Agrawal (1964, p. 218), both of which noted that Thapar and Dayal (1934) failed to provide a description of their new species. Sekerak and Arai (1974, p. 710) remarked that this species “was reported from Trigla gurnardus from the aquarium of the Zoological Society, London…no description was published” and “this species must be considered as a nomen nudum.” Skrjabin and Koval (1958) and Skrjabin (1964, p. 135) mentioned this species but made note that H. gurnardus was omitted from their keys. Yamaguti (1958a, 1971) listed this species from India, and Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors; he also listed with this species the note “No measurements.”
Helicometra hapalogenyos Wang, Wang and Zhang, 1992
(Syn. Helicometra hapogenyos Wang, Wang and Zhang, 1992; Helicometra hapolegenyos Wang, Wang and Zhang, 1992).
Host: Hapalogenyidae: Hapalogenys mucronatus (Eydoux and Souleyet, 1850)—Barbeled grunter sp. [type host].
Site of infection: Intestine.
Distribution: Off Fujian Province, China in Taiwan Strait (Wang et al. 1992).
Reports: Zhang et al. (1999), Liu et al. (2010).
Specimens deposited: Type specimens deposited in the Parasitological Research Laboratory, Fujian Teachers University, China.
Remarks: Aken’Ova et al. (2006, p. 18) commented that for the species designation “hapalogenyos”, there have been several spellings used (e.g. “hapogenyos”), but the spelling used above is “the most satisfactory if, as seems certain, the name is based on the host genus Hapalogenys Richardson, 1844.” Aken’Ova et al. (2006) and Hassanine (2007) also placed this species in Group I.
Helicometra hypodytis Yamaguti, 1934
(Syns. Helicometra [Helicometra] hypodytis Yamaguti, 1934; Helicometra pulchella [Rudolphi, 1819]).
Hosts: Apogonidae: Apogon doederleini Jordan and Snyder, 1901—Doederlein’s cardinalfish; Cheilodipterus quinquelineatus Cuvier, 1828—Five-lined cardinalfish; Labridae: Thalassoma rueppellii (Klunzinger, 1871) (=Thalassoma klunzengri [sic] Fowler and Steinitz, 1956)—Klunzinger’s wrasse; Lethrinidae: Lethrinus harak (Forsskål, 1775)—Thumbprint emperor; Pomacentridae: Chromis notata (Temminck and Schlegel, 1843) (=Chromis notatus [Temminck and Schlegel, 1843])—Pearl-spot chromis; Serranidae: Serranus (=Epinephelus) sp.—Grouper sp. and locally called “Koshar”; Tetrarogidae: Paracentropogon rubripinnis (Temminck and Schlegel, 1843) (=Hypodytes rubripinnis [Temminck and Schlegel, 1843])—wasp fish sp [type host].
Site of infection: Small intestine.
Distribution: Inland Sea of Japan (Yamaguti 1934); Tarumi, Japan (Yamaguti 1939); Hurghada, Egypt, Red Sea (Nagaty 1956a; Hossam et al. 2012); Japan (Dyer et al. 1988; Nagasawa 1989); Taiwan Strait (Wang et al. 1992); Bay of Bengal (Hanumantha Rao et al. 1993).
Reports: Yamaguti (1958a [Pl. 10, Fig. 122], 1971 [Fig. 407]), Pan et al. (1990), Zhang et al. (1999), Liu et al. (2010).
Specimens deposited: Vouchers: USNPC #59548, 79879–79881, 79961; Yamaguti (1934, p. 301–302) described this species from a single specimen, which we expect was deposited in his collection.
Remarks: This species is included in keys of Skrjabin and Koval (1958) and Skrjabin (1964). Manter (1954) and Dawes (1956) considered this species (and H. epinepheli) a junior syn. of H. fasciata; however, Yamaguti (1958a, 1971) refuted this by stating that “the Laurer’s canal forms a very peculiar convolution in H. hypodytis, but nothing like that in H. epinepheli.” Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Ramadan (1986) included this species in a key to the species of Helicometra from Red Sea fishes.
Helicometra indica Agrawal, 1964
(Syns. Helicometra fasciata [Rudolphi, 1819]; Helicometra [Helicometra] indica Agrawal, 1964).
Host: Triglidae: Eutrigla gurnardus (Linnaeus, 1758) (=Trigla gurnardus Linnaeus, 1758)—Grey gurnard [type host].
Sites of infection: Intestine; pyloric caecum.
Distribution: Type locality unknown. West European shelf (Gaevskaia 1981); Bay of Biscay, Plymouth, England and St. Kilda, Scotland (Bray 1987a). Host species distributed thoughout coastal areas of the Eastern Atlantic (Norway to Morocco, Madeira, and Iceland) as well as from the Mediterranean and Black Seas (Froese and Pauly 2013).
Reports: Agrawal (1964), Yamaguti (1971).
Specimens deposited: BM(NH) 1963.6.20.1,4; 1982.6.9.95,100; see Remarks below.
Remarks: As stated by Agrawal (1964, p. 216) regarding H. indica, “Only one stained specimen of the genus Helicometra Odhner, 1902 was made available to me through the courtesy of Dr. S. P. Gupta, from the Helminthological collection of Professor G. S. Thapar, Lucknow University, Lucknow.” Sekerak and Arai (1974) included this species in their key but noted that the location of where the host was collected was not included in the original description. Both authors also noted that “[i]t is possible that H. indica is based on, or is one, of the specimens reported, but not described, by Thapar and Dayal (1934) as H. gurnardus” (see H. gurnardus above which was collected from the same host species also from an unknown locality). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors, and stated its locality as India. Bray (1987a) considered this species to be a junior syn. of H. fasciata. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I. Agrawal (1964, p. 216) described the oral sucker of H. indica as terminal; however, it is clearly subterminal in the whole mount illustrated in Fig. 1 (p. 217).
Helicometra insolita Polianskii, 1955
(Syns. Helicometra [Helicometra] insolita Polianskii, 1955; Helicometra pleurogrammi [Baeva, 1968]; Neohelicometra insolita [Polianskii, 1955] [see Hassanine 2007, p. 22]).
Hosts: Agonidae: Percis japonica (Pallas, 1769) (=Percis japonicus [Pallas, 1769])—Dragon poacher; Cottidae: Icelus bicornis (Reinhardt, 1840)—Twohorn sculpin; Labridae: Symphodus tinca (Linnaeus, 1758)—East Atlantic peacock wrasse; Stichaeidae: Lumpenus fabricii Reinhardt, 1836—Slender eelblenny [type host]; Lumpenus lampretaeformis (Walbaum, 1792)—Snakeblenny; Stichaeus grigorjewi Herzenstein, 1890—Prickleback sp.
Site of infection: Intestine.
Distribution: Barents Sea (Polianskii 1955); Putiatin Island (an island in Peter the Great Gulf in the Sea of Japan belonging to the USSR) (Skrjabin and Koval 1958; Zhukov 1960); Hidaka District, Hokkaido, Japan (Machida et al. 1972); Grand Bank off Newfoundland and Hamilton Inlet Bank off Labrador (Bray 1979); Sea of Marmara, Turkey (Oguz 1995).
Reports: Yamaguti (1971), Gibson (1996).
Specimens deposited: Vouchers: BM(NH) #1977.1.17.1–7.
Remarks: This species is included in keys of Skrjabin and Koval (1958), Skrjabin (1964), and Sekerak and Arai (1974), who transferred it to Neohelicometra Siddiqi and Cable, 1960. Bray (1979), noting that fixation method produced variability in the terminal vs subterminal appearance of the oral sucker (the main feature used in the key by Sekerak and Arai [1974] to distinguish Neohelicometra and Helicometra), returned N. insolita to Helicometra. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Machida (1984) considered H. pleurogrammi as probably conspecific with H. insolita. Gibson (1996, p. 170) included this species in a key to three species of Helicometra. Hassanine (2007) placed this species in Group IV.
Helicometra interrupta Hassanine, 2005
Host: Platycephalidae: Cociella crocodilus (Cuvier, 1829) (=Cociella crocodila [sic] [Tilesius, 1814])—Crocodile flathead [type host].
Site of infection: Intestine.
Distribution: Red Sea off coast of Sharm El-Sheikh, South Sinai, Egypt (Hassanine 2005).
Specimens deposited: Holotype and paratypes: Helminthological Collection of the Red Sea Fishes, Marine Science Dept., Faculty of Science, Suez Canal University, Ismailia, Egypt.
Remark: Hassanine (2007) placed this species in Group III.
Helicometra kyliotrema Pritchard, 1966
(Syn. Helicometra [Helicometra] kyliotrema Pritchard, 1966).
Host: Muraenidae: Moray eel sp. [type host].
Site of infection: Intestine.
Distribution: Hawaii (Pritchard 1966).
Report: Yamaguti (1971).
Specimens deposited: Holotype: USNPC #60356; Paratype: HWML #1645.
Remarks: Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. This species is included in key of Sekerak and Arai (1974). While acknowledging H. kyliotrema from Hawaiian fishes (see Pritchard 1966), it was not included in a key to Helicometra species in Hawaiian fishes by Yamaguti (1970) because he personally did not collect any specimens of this species. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra labri (Stossich, 1886)
(Syn: Allocreadium labri [Stossich, 1886] of Odhner, 1901; Distomum labri Stossich, 1886; Helicometra pulchella [Rudolphi, 1819]; Loborchis labri [Stossich, 1886]).
Hosts: Labridae: Labrus mixtus Linnaeus, 1758—Cuckoo wrasse [type host]; Trachinidae: Trachinus draco Linnaeus, 1758—Greater weever (see Remarks below).
Site of infection: Digestive tract.
Distribution: Triest, Italy (Stossich 1886, 1887a, b); Russian Arctic (Isaĭchikov 1928); Mediterranean Sea (Reversat et al. 1991b).
Reports: Stossich (1902), Bona et al. (2000).
Specimens deposited: We could find no information regarding deposition of specimens from the descriptions of Helicometra labri (as Distomum labri Stossich, 1886) by Professor Michele Stossich (Stossich 1886, 1887a, b). Reversat et al. (1991b, p. 154, Table VI) recently noted this species, and based on this reference, inquiries can be made at the Université Montpellier II, Sciences et Techniques de Languedoc, Laboratoire de Parasitologie Comparée, URA CNRS 698, place Eugène Battaillon, 34095 Montpellier Cedex 5, France.
Remarks: This species was in Loborchis, a genus now considered a junior synonym of Helicometra (see Skrjabin and Koval 1958; Yamaguti 1958a). Yamaguti (1958a, p. 1,502), Sekerak and Arai (1974), Sonsino (1891) and Odhner (1902c) considered this species (the latter as Distomum labri Stossich, 1886) a junior syn. of H. pulchella. Yamaguti (1971, p. 189) recognized this species as Allocreadium labri (Stossich, 1886) of Odhner, 1901 and considered it a junior syn. of H. pulchella. Reversat et al. (1991b, p. 148) reported that Stossich (1886, p. 30) originally described D. labri from the cuckoo wrasse, Labrus mixtus Linnaeus, 1758 (Labridae), from off Trieste, Italy. These same authors noted that in Stossich’s time L. mixtus was also referred to as Labrus bimaculatus Linnaeus, 1758 (see Reversat et al. 1991b, p. 148, 154, Table VI). With this in mind, it is important to state that today the taxon L. bimaculatus is considered a junior syn. of the taxon Cichlasoma bimaculatum (Linnaeus, 1758) which is the black acara within Cichlidae. This host is not distributed in the Old World but in various locations throughout South America including the Orinoco River basin (in the Caroni River in Venezuela), Guianas (from the Essequibo River to the Sinnamary River), and the Amazon River basin (in the upper Branco River basin) (see Froese and Pauly 2013).
Helicometra markewitschi Pogorel’tseva, 1954
(Syns. Helicometra [Helicometra] markewitschi Pogorel’tseva, 1954; Helicometra pulchella [Rudolphi, 1819]).
Hosts: Scorpaenidae: Scorpaena porcus Linnaeus, 1758—Black scorpionfish [type host]; Trachinidae: Trachinus draco Linnaeus, 1758—Greater weever.
Site of infection: Intestine.
Distribution: Black Sea (Pogorel’tseva 1954; Skrjabin and Koval 1958); Mediterranean Sea (Reversat et al. 1991b).
Report: Yamaguti (1971).
Specimens deposited: We were unable to confirm the presence or location of any deposited specimens. However, Reversat et al. (1991b, p. 154, Table V) recently noted this species, and based on this reference, inquiries can be made at the Université Montpellier II, Sciences et Techniques de Languedoc, Laboratoire de Parasitologie Comparée, URA CNRS 698, place Eugène Battaillon, 34095 Montpellier Cedex 5, France.
Remarks: Index Catalogue of Medical and Veterinary Zoology (Suppl. 19, Part 3, p. 102) lists this species as a nomen nudum. This species is included in keys of Skrjabin and Koval (1958) and Skrjabin (1964). Naĭdenova and Dolgikh (1969) stated that H. markewitschi was a junior syn. of H. fasciata, while Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors.
Helicometra marmoratae Nagaty and Abdel Aal, 1962
(Syns. Helicometra fasciata [Rudolphi, 1819]; Helicometra [Helicometra] marmoratae Nagaty and Abdel Aal, 1962; Helicometra pulchella [Rudolphi, 1819] [see Hassanine 2001]).
Host: Siganidae: Siganus spinus (Linnaeus, 1758) (=Teuthis marmorata [Quoy and Gaimard, 1825])—Little spinefoot [type host].
Site of infection: Middle intestine.
Distribution: Hurghada, Egypt in Red Sea (Nagaty and Abdel Aal 1962; Hassanine 2001); Red Sea off coast of Sharm El-Sheikh, South Sinai, Egypt (Hassanine 2007).
Report: Yamaguti (1971).
Specimens deposited: Holotype: Dept. of Parasitology, Faculty of Medicine, Cairo, Egypt #TRSF.1962.12; Vouchers: Helminthological Coll. of Red Sea Fishes, Marine Science Dept., Faculty of Science, Suez Canal University, Ismailia, Egypt #2006.2.25.1–30.
Remarks: Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. This species is included in keys of Sekerak and Arai (1974), who also mentioned that H. marmoratae was the largest species of Helicometra (total length = 6,320 μm). Ramadan (1986) included this species in a key to the species of Helicometra from Red Sea fishes. Hassanine (2001, p. 87) considered this species to be junior syn. of H. pulchella (Rudolphi, 1819). Aken’Ova et al. (2006) placed this species in Group I. Hassanine (2007) examined the type specimen of this species and noted that the genital pore is not postbifurcal, as in the original description, but situated at the mid-esophageal level. Furthermore, Hassanine (2007) considered H. marmoratae to be a junior syn. of H. fasciata.
Helicometra mutabilis (Stossich, 1902)
(Syns: Helicometra fasciata [Rudolphi, 1819] [see Yamaguti 1971, p. 190]; Helicometra [Helicometra] mutabilis [Stossich, 1902]; Helicometra pulchella [Rudolphi, 1819]; Loborchis mutabilis Stossich, 1902).
Host: Anguillidae: Anguilla anguilla (Linnaeus, 1758) (=Anguilla vulgaris Rafinesque, 1810)—European eel [type host].
Site of infection: Intestine.
Distribution: Trieste, Italy (Stossich 1902); Lagoon of Languedoc (“Etang de Thau”) in southern France (Maillard and Aussel 1988); Mediterranean Sea (Reversat et al. 1991b).
Reports: Stossich (1903a), Nicoll (1910), Palombi (1929a), Yamaguti (1958a, 1971) and Bona et al. (2000).
Specimens deposited: We could find no information regarding deposition of specimens from the original description of H. mutabilis (as Loborchis mutabilis Stossich, 1902) by Professor Michele Stossich (Stossich 1902). Reversat et al. (1991b, p. 155, Table VIII) recently noted this species, and based on this reference, inquiries can be made at the Université Montpellier II, Sciences et Techniques de Languedoc, Laboratoire de Parasitologie Comparée, URA CNRS 698, place Eugène Battaillon, 34095 Montpellier Cedex 5, France.
Remarks: Lühe in Stossich (1902) erected Loborchis with Loborchis mutabilis Stossich, 1902 as type species; however, this genus is now considered a junior synonym of Helicometra (see Skrjabin and Koval 1958; Yamaguti 1958a, 1971). This species is included in keys of Wallin (1909) and Isaĭchikov (1928). Palombi (1929a) considered this species a junior syn. of H. fasciata. Markevich (1952) and Sekerak and Arai (1974) considered this species a junior syn. of H. pulchella. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors.
Helicometra nasae Nagaty and Abdel Aal, 1962
(Syns. Helicometra boseli Nagaty, 1956; Helicometra equilata (Manter, 1933) [see Hassanine 2001]; Helicometra [Helicometra] nasae Nagaty and Abdel Aal, 1962; Helicometra rectisaccus [Fischthal and Kuntz, 1964] [see Sekerak and Arai 1974, p. 729; Hassanine 2005, p. 279]; Helicometra [=Stenopera] rectisaccus [Fischthal and Kuntz, 1964] [see Pritchard 1966, p. 198; Hassanine 2007, p. 22]; Stenopera nasae [Nagaty and Abdel Aal, 1962]; Stenopera rectisaccus Fischthal and Kuntz, 1964 [see Hassanine 2005, p. 279]).
Hosts: Acanthuridae: Naso sp.? (=Nasa [sic] sp.; locally called “dakkara”)—Unicornfish sp.; Holocentridae: Sargocentron violaceum (Bleeker, 1853)—Violet squirrelfish; Serranidae: Epinephelus fasciatus (Forsskål, 1775) (=Epenepfelus [sic] fasciatus [Forsskål, 1775]; locally called “koshari abu luli”)—Blacktip grouper [type host].
Site of infection: Digestive tract.
Distribution: Hurghada, Egypt in Red Sea (Nagaty and Abdel Aal 1962; Hassanine 2001).
Reports: Yamaguti (1971), Arthur and Lumanlan-Mayo (1997) and Cribb et al. (2002).
Specimens deposited: Type specimens in Dept. of Parasitology, Faculty of Medicine, Abbassia, Cairo, U.A.R.; Vouchers: Helminthological Collection of the Red Sea Fishes, Marine Science Dept., Faculty of Science, Suez Canal University, Ismailia, Egypt.
Remarks: Pritchard (1966, p. 198) disagreed with Siddiqi and Cable (1960), who synonymized Stenopera with Helicometra, and re-established Stenopera as a valid genus moving H. nasae in to Stenopera; however, later authors still consider this genus as synonymous with Helicometra. This species is included in keys of Pritchard (1966) and Sekerak and Arai (1974). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Ramadan (1986) considered H. nasae to be a junior syn. of H. boseli while Hassanine (2001, p. 89) considered H. nasae a junior syn. of H. equilata (Manter, 1933). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group III.
Helicometra neoscorpanae Wang, Wang and Zhang, 1992
(Syns. Helicometra fasciata [Rudolphi, 1819] [see Liu et al. 2010, p. 40]; Helicometra scorpaenae Wang, 1982, preoccupied).
Host: Scorpaenidae: Scorpaena neglecta Temminck & Schlegel, 1843—Scorpaenid sp. [type host].
Site of infection: Intestine.
Distribution: Fujian Province, China in Taiwan Strait (Wang 1982; Wang et al. 1992).
Reports: Zhang et al. (1999), Zongguo (2001) and Liu et al. (2010).
Specimens deposited: All type specimens were deposited in the Parasitological Research Laboratory, Fujian Normal University, China.
Remarks: Bray (1987a) considered this species to be a junior syn. of H. fasciata. Wang et al. (1992, p. 72) proposed H. neoscorpanae nom. nov. to replace H. scorpaenae Wang, 1982 that is itself an illegitimate name being a younger homonym of H. scorpaenae Prudhoe and Bray, 1973. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra neosebastodis Crowcroft, 1947
(Syns. Helicometra [Metahelicometra] neosebastodis Crowcroft, 1947; Helicometra tenuifolia Woolcock, 1935).
Host: Neosebastidae: Neosebastes thetidis (Waite, 1899)—Thetis fish [type host].
Site of infection: Gut immediately behind stomach.
Distribution: Hobart fish market, Tasmania (off southern coast of Australia) (Crowcroft 1947b).
Reports: Yamaguti (1958a, 1971 [Fig. 392]).
Specimens deposited: Syntypes from P.W. Crowcroft’s Collection #TMAG Nos. Rg9, Rg14, T/Tr9.
Remarks: This species is included in keys of Skrjabin and Koval (1958) and Skrjabin (1964). Yamaguti (1971) considered this species to be the type species within the subgenus Metahelicometra Yamaguti, 1971, a subgenus now considered a junior syn. of Helicometra (Cribb 2005). Sekerak and Arai (1974) believed this species to be a junior syn. of H. tenuifolia. Aken’Ova et al. (2006) placed this species in Group II.
Helicometra paracirrhiti Pritchard, 1966
(Syn. Helicometra [Helicometra] paracirrhiti Pritchard, 1966).
Host: Cirrhitidae: Paracirrhites forsteri (Schneider, 1801)—Blackside hawkfish [type host].
Site of infection: Intestine.
Distribution: Hawaii (Pritchard 1966).
Report: Yamaguti (1971).
Specimens deposited: Holotype: USNPC #60357; Paratype: HWML #1638.
Remarks: Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. This species is included in keys of Sekerak and Arai (1974). While acknowledging H. paracirrhiti from Hawaiian fishes (see Pritchard 1966), it was not included in a key to Helicometra species in Hawaiian fishes by Yamaguti (1970) because he personally did not collect any specimens of this species. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra pisanoae Zdzitowiecki, 1998
Host: Nototheniidae: Trematomus hansoni Boulenger, 1902—Striped rockcod [type host].
Sites of infection: Anterior half of small intestine and large intestine; pyloric caecum.
Distribution: Off Adelie Land, Terre Adelie, East Antarctica (Zdzitowiecki 1998, 2001; Zdzitowiecki et al. 1998); Coastal waters of Terra Nova Bay, Ross Sea, Antarctica (Zdzitowiecki et al. 1999).
Specimens deposited: Holotype: Museum of Zoology, Warszawa #MZPW 1796; Paratypes: BM(NH) #1997.11.3.5–14; Museum of Zoology, Warszawa MZPW #1797–1804.
Remarks: Aken’Ova et al. (2006) placed this species in Group II, while Hassanine (2007) placed it in Group IV.
Helicometra pisodonophi Nguyễn Văn Hà, 2012
Host: Ophichthidae: Pisodonophis cancrivorus (Richardson, 1848)—Longfin snake-eel [type host].
Site of infection: Stomach.
Distribution: Ha Long Bay, Vietnam (Nguyễn Văn Hà, 2012).
Specimens deposited: Type specimens in the Institute of Ecology and Biological Resources, VAST.
Remarks: This species was published after the revision by Hassanine (2007) and we have placed it in Group I.
Helicometra pleurogrammi (Baeva, 1968)
(Syns. Allostenopera pleurogrammi Baeva, 1968 [see Hassanine 2007, p. 22]; Helicometra insolita Polianskii, 1955; Helicometra pugetensis Schell, 1973 [see Hassanine 2007, p. 22]; Neohelicometra pleurogrammi [Baeva, 1968] [see Hassanine 2007, p. 22]).
Hosts: Hexagrammidae: Hexagrammos octogrammus (Pallas, 1814)—Masked greenling; Hexagrammos otakii Jordan and Starks, 1895—Greenling sp.; Pleurogrammus azonus Jordan and Metz, 1913—Okhotsk atka mackerel [type host]; Psychrolutidae: Ebinania vermiculata Sakamoto, 1932—Fathead sp.; Stichaeidae: Acantholumpenus mackayi (Gilbert, 1896)—Pighead prickleback; Stichaeus nozawae Jordan and Snyder, 1902 (=Stichaeus nozawai [sic] Jordan and Snyder, 1902)—Prickleback sp.
Sites of infection: Intestine; pyloric caecum.
Distribution: Peter the Great Bay, Sea of Japan (Baeva 1968); Hidaka District, Hokkaido, Japan (Machida et al. 1972); Off Kesen-numa of Miyagi Prefecture in northern North Pacific, off Yamagata Prefecture in Sea of Japan, and off Kushiro, Hokkaido in Sea of Okhotsk (Machida 1984; Machida and Araki 1994); Uraga Channel, Tokyo Bay (Kuramochi 2006).
Reports: Yamaguti (1971 [Fig. 1784]), Love and Moser (1976, 1983).
Specimens deposited: Vouchers: National Science Museum, Tokyo, Japan #NSMT-P1 2662, 2699, 2824, 4442.
Remarks: Baeva (1968) erected Allostenopera Baeva, 1968 within Allocreadiidae with Allostenopera pleurogrammi Baeva, 1968 as type species (validity of genus supported by Yamaguti [1971, p. 185–186] but he placed it in Opecoelidae); however, this genus is now considered a junior synonym of Helicometra (see Cribb 2005). Sekerak and Arai (1974) considered Allostenopera to be synonymous with Neohelicometra Siddiqi and Cable, 1960 and moved A. pleurogrammi in to Neohelicometra as Neohelicometra pleurogrammi (Baeva, 1968) while also establishing H. pugetensis Schell, 1973 as a junior syn. of this species. N. pleurogrammi included in key of Sekerak and Arai (1974). Bray (1979), noting that fixation method produced variability in the terminal vs subterminal appearance of the oral sucker (the main feature used in the key by Sekerak and Arai [1974] to distinguish Neohelicometra and Helicometra), moved N. pleurogrammi in to Helicometra. Machida (1984, p. 109) considered H. pleurogrammi as probably conspecific with H. insolita. Hassanine (2007) placed this species in Group IV.
Helicometra plovmornini Isaĭchikov, 1928
(Syns: Helicometra [Helicometra] pulvomornini [sic] Isaĭchikov, 1928; Helicometra pulvomornini [sic] Isaĭchikov, 1928).
Hosts: Agonidae: Ulcina olrikii (Lütken, 1877) (=Aspidophoroides olriki Lütkin, 1877)—Arctic alligatorfish; Anarhichadidae: Anarhichas lupus Linnaeus, 1758—Atlantic wolfish; Cottidae: Artediellus atlanticus Jordan & Evermann, 1898 (=Artediellus europeus [sic] Knipowitsch, 1907)—Atlantic hookear sculpin; Myoxocephalus scorpius (Linnaeus, 1758)—Shorthorn sculpin; Triglops murrayi Günther, 1888—Moustache sculpin; Triglops pingelii Reinhardt, 1837—Ribbed sculpin; Psychrolutidae: Cottunculus microps Collett, 1875—Polar sculpin; Zoarcidae: Lycodes esmarkii Collett, 1875—Greater eelpout; Lycodes gracilis Sars, 1867 (=Lycodes vahli gracilis Sars, 1867)—Eelpout sp.; Lycodes polaris (Sabine, 1824) (=Lycodes agnostus Jensen, 1902)—Canadian eelpout [type host]; Lycodes reticulatus Reinhardt, 1835—Arctic eelpout; Lycodes vahlii Reinhardt, 1831 (=Lycodes vahli [sic] Reinhardt, 1831)—Vahl’s eelpout.
Sites of infection: Intestine; stomach.
Distribution: Eastern Barents Sea, Kara Sea & Gulf of Dvinsk, White Sea (Isaĭchikov 1928); White Sea (Shul’man and Shul’man-Al’bova 1953); Barents Sea (Polianskii 1955); Grand and Green Banks off Newfoundland and Banquereau Bank off Nova Scotia (Bray 1979); North Atlantic Ocean (Bray 1987b); Faroese coast (Faroe Islands) (Køie 2000).
Reports: Yamaguti (1958a, 1971), Chertkova et al. (1994), Gibson (1996) and Klimpel et al. (2001, 2009).
Specimens deposited: Vouchers: BM(NH) #1976.12.10.1–67.
Remarks: This species is included in keys of Isaĭchikov (1928), Manter (1933), Skrjabin and Koval (1958), Skrjabin (1964), and Sekerak and Arai (1974). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Gibson (1996, p. 170) included this species in a key to three species of Helicometra. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra pretiosa Bravo-Hollis and Manter, 1957
(Syns. Helicometra [Metahelicometra] pretiosa Bravo-Hollis and Manter, 1957; Helicometra torta Linton, 1910).
Host: Serranidae: Paralabrax maculatofasciatus (Steindachner, 1868) (=Paralabrax maculofasciatus [sic] [Steindachner, 1868])—Spotted sand bass [type host].
Site of infection: Intestine.
Distribution: Puerto Vallarta, Jaslico, Mexico (Bravo-Hollis and Manter 1957).
Reports: Yamaguti (1971), Love and Moser (1976, 1983).
Specimens deposited: Holotype: USNPC #38181; Paratype: HWML #1833.
Remarks: This species is included in keys of Skrjabin and Koval (1958) and Skrjabin (1964). Overstreet (1969, p. 144) and Sekerak and Arai (1974) believed this species to be a junior syn. of H. torta. Yamaguti (1971) considered this species to be within the subgenus Metahelicometra Yamaguti, 1971, a subgenus now considered a junior syn. of Helicometra (Cribb 2005), and he stated that by “[j]udging from the measurements, especially of eggs, this species cannot be identical with H. torta Linton, though synonymized with the latter by Overstreet (1969).” Aken’Ova et al. (2006) placed this species in Group II.
Helicometra pteroisi (Gupta, 1956)
(Syn. Stenopera pteroisi Gupta, 1956 [see Hassanine 2007, p. 22; Madhavi 2011, p. 195])
Host: Scorpaenidae: Pterois russelii Bennett, 1831—Plaintail turkeyfish [type host].
Site of infection: Intestine.
Distribution: Gulf of Manaar (between India and Sri Lanka) (Gupta 1956).
Reports: Fischthal and Kuntz (1965), Yamaguti (1971) and Madhavi (2011).
Specimens deposited: In the type description of this species (Gupta 1956), there was no indication of where materials were deposited; however, the author, from Panjab University College, Hoshiarpur, Panjab, India, carried out the study in the Dept. of Parasitology, London School of Hygiene and Tropical Medicine, London, England.
Remarks: This species was in Stenopera; however, this genus is now considered a junior synonym of Helicometra (see Siddiqi and Cable 1960). Pritchard (1966, p. 198) disagreed and re-established Stenopera as a valid genus returning H. pteroisi to Stenopera (validity of this species in Stenopera supported by Yamaguti [1971]); however, later authors still consider this genus as synonymous with Helicometra. This species is included in keys of Pritchard (1966) and Sekerak and Arai (1974). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group III.
Helicometra pugetensis Schell, 1973
(Syns. Allostenopera pleurogrammi Baeva, 1968; Helicometra pleurogrammi [Baeva, 1968]; Neohelicometra pleurogrammi [Baeva, 1968]).
Host: Hexagrammidae: Hexagrammos decagrammus (Pallas, 1810)—Kelp greenling [type host].
Site of infection: Intestine.
Distribution: Puget Sound, vicinity of San Juan Islands, Washington (Schell 1973).
Reports: Love and Moser (1976, 1983).
Specimens deposited: Holotype and paratypes: USNPC #72447, 72448; HWML #23782.
Remarks: This species was transferred by Sekerak and Arai (1974) to Neohelicometra Siddiqi and Cable, 1960 as a junior syn. of N. pleurogrammi; however, Bray (1979, p. 402) returned this latter species to Helicometra.
Helicometra pulchella (Rudolphi, 1819)
(Syns: Allocreadium alacre Stossich, 1905 [in schedis] non Looss, 1901 [see Palombi 1929a, b]; Allocreadium fasciatum [Rudolphi, 1819] Odhner, 1901; Allocreadium labri [Stossich, 1886] Odhner, 1901 [see Odhner 1901; Palombi 1929a]; Allocreadium sinuatum [Rudolphi, 1819] Odhner, 1901; Distoma [Dicrocoelium] fasciatum [Rudolphi, 1819] Barbagallo and Drago, 1903; Distoma [Dicrocoelium] pulchellum [Rudolphi, 1819] Barbagallo and Drago, 1903 [see Barbagallo and Drago 1903; Palombi 1929a]; Distoma fasciatum Rudolphi, 1819; Distoma labri Stossich, 1886 nec van Beneden, 1890 (see Sonsino 1891; Reversat et al. 1991b, p. 148); Distoma pulchellum Rudolphi, 1819 [see Rudolphi 1819; Dujardin 1845; Palombi 1929a]; Distoma sinuatum Rudolphi, 1819; Distomum fasciatum Rudolphi, 1819 nec Stossich, 1885; Distomum gobii Stossich, 1883; Distomum labri Rudolphi, 1819; Distomum labri Stossich, 1886 nec Rudolphi, 1819 [see Stossich 1886, 1887a, 1888; Sonsino 1891; Palombi 1929a]; Distomum pulchellum Diesing, 1850 [see Diesing 1850; Carus 1884; Stossich 1886, 1898; Looss 1899; Zschokke 1887; Palombi 1929a]; Helicometra aposinuata Pritchard, 1966 [see Sekerak and Arai 1974, p. 713]; Helicometra dochmosorchis Manter and Pritchard, 1960 [see Sekerak and Arai 1974, p. 713]; Helicometra epinepheli Yamaguti, 1934 [see Sekerak and Arai 1974, p. 710]; Helicometra fasciata [Rudolphi, 1819] Odhner, 1902 [see Sekerak and Arai 1974, p. 710]; Helicometra fasciata of Nicoll 1914 [see Nicoll 1914; Manter 1933]; Helicometra flava Stossich, 1903 [see Sekerak and Arai 1974, p. 710]; Helicometra gobii [Stossich, 1883] Stossich, 1904 [see Sekerak and Arai 1974, p. 710]; Helicometra [Helicometra] pulchella [Rudolphi, 1819]; Helicometra hypodytis Yamaguti, 1934 [see Sekerak and Arai 1974, p. 713]; Helicometra labri [Stossich, 1886] Isaĭchikov, 1928 [see Sekerak and Arai 1974, p. 710]; Helicometra markewitschi Pogorel’tseva, 1954 [see Sekerak and Arai 1974, p. 710]; Helicometra marmoratae Nagaty and Abdel-Aal, 1962 [see Hassanine 2001]; Helicometra mutabilis [Stossich, 1902] Stossich, 1903 [see Sekerak and Arai 1974, p. 710]; Helicometra sinuata [Rudolphi, 1819] Odhner, 1902 [see Sekerak and Arai 1974, p. 710]); Helicometra upapalu Yamaguti, 1970 [see Sekerak and Arai 1974, p. 713]; Loborchis fasciatum [Rudolphi, 1819] Stossich, 1902 [see Sekerak and Arai 1974, p. 710]; Loborchis gobii [Stossich, 1883] Stossich, 1902 [see Sekerak and Arai 1974, p. 710]; Loborchis labri [Stossich, 1886] Stossich, 1902 [see Sekerak and Arai 1974, p. 710]; Loborchis mutabilis Stossich, 1902 [see Nicoll 1915, p. 344; Sekerak and Arai 1974, p. 710]; “MDH- 1a” [see Reversat et al. 1989, 1991b]; From Palombi 1929a, p. 277—see Odhner 1902c; Stossich 1904b; Wallin 1909; Palombi 1929b).
Hosts: Anguillidae: Anguilla anguilla (Linnaeus, 1758) (=Anguilla vulgaris Rafinesque, 1810)—European eel; Blenniidae: Blennius ocellaris Linnaeus, 1758—Butterfly blenny; Lipophrys pholis (Linnaeus, 1758) (=Blennius pholis Linnaeus, 1758)—Shanny; Parablennius gattorugine (Linnaeus, 1758) (=Blennius gattorugine Linnaeus, 1758)—Tompot blenny; Carangidae: Trachurus trachurus (Linnaeus, 1758)—Atlantic horse mackerel; Trachurus sp.—Mackerel sp.; Cichlidae: Cichlasoma bimaculatum (Linnaeus, 1758) (=Labrus bimaculatus Linnaeus, 1758)—Black acara; Congridae: Conger conger (Linnaeus 1758)—European conger; Cyprinidae: Leucaspius delineatus (Heckel, 1843)—Belica; Tinca tinca (Linnaeus, 1758) (=Tinca vulgaris Fleming, 1828)—Tench; Gadidae: Raniceps raninus (Linnaeus, 1758)—Tadpole fish; Gobiesocidae: Lepadogaster lepadogaster (Bonnaterre, 1788) (=Lepidogaster gouanii Risso, 1810)—Shore clingfish; Gobiidae: Gobius cobitis Pallas, 1814—Giant goby; Gobius niger Linnaeus, 1758 (=Gobius jozo Linnaeus, 1758)—Black goby; Gobius paganellus Linnaeus, 1758—Rock goby; Zosterisessor ophiocephalus (Pallas, 1814) (=Gobius ophiocephalus Pallas, 1814)—Grass goby; Haemulidae: Plectorhinchus pictus (Tortonese, 1936) (=Plectorhynchus [sic] pictus [Tortonese, 1936])—Trout sweetlips; Labridae: Ctenolabrus rupestris (Linnaeus, 1758) (=Labrus cynaedus? Linnaeus, 1758; Labrus rupestris Linnaeus, 1758)—Goldsinny wrasse [type host]; Labridae sp.—Wrasse sp.; Labrus bergylta Ascanius, 1767 (=Labrus berggylta Ascanius, 1767)—Ballan wrasse; Labrus merula Linnaeus, 1758—Brown wrasse; Labrus mixtus Linnaeus, 1758—Cuckoo wrasse; Symphodus cinereus (Bonnaterre, 1788)—Grey wrasse; Symphodus mediterraneus (Linnaeus, 1758) (=Crenilabrus mediterraneus [Linnaeus, 1758])—Axillary wrasse; Symphodus melops (Linnaeus, 1758) (=Crenilabrus melops [Linnaeus, 1758])—Corkwing wrasse; Symphodus ocellatus (Linnaeus, 1758) (=Crenilabrus ocellatus [Forsskål, 1775])—Wrasse sp.; Symphodus tinca (Linnaeus, 1758)—East Atlantic peacock wrasse; Thalassoma pavo (Linnaeus, 1758) (=Crenilabrus pavo [Linnaeus, 1758])—Ornate wrasse; Lotidae: Gaidropsarus vulgaris (Cloquet, 1824) (=Onos tricirrata [sic] [non Brünnich, 1768])—Three-bearded rockling; Lutjanidae: Lutjanus lutjanus Bloch, 1790 (=Lutjanus lineolatus [Rüppell, 1829])—Bigeye snapper; Muraenidae: Gymnothorax porphyreus (Guichenot, 1848) (=Gymnothorax wieneri Sauvage, 1883)—Lowfin moray; Ophidiidae: Lepophidium sp.—Cusk-eel sp.; Ophidion barbatum Linnaeus, 1758 (=Ophidium [sic] barbatum Linnaeus, 1758)—Snake blenny; Oplegnathidae: Oplegnathus fasciatus (Temminck and Schlegel, 1844)—Barred knifejaw; Sciaenidae: Cilus gilberti (Abbott, 1899) (=Sciaena gilberti Abbott, 1899)—Corvina drum; Sciaena umbra Linnaeus, 1758 (=Corvina nigra non Cuvier, 1829)—Brown meager; Stellifer minor (Tschudi, 1846)—Minor stardrum; Scophthalmidae: Scophthalmus maeoticus (Pallas, 1814) (=Bothus maeoticus Pallas, 1814)—Turbot sp.; Zeugopterus punctatus (Bloch, 1787)—Topknot; Scorpaenidae: Scorpaena porcus Linnaeus 1758—Black scorpionfish; Scorpaena scrofa Linnaeus, 1758 (=Scorpaena natalensis [Regan, 1906])—Red scorpionfish; Serranidae: Paralabrax humeralis (Valenciennes, 1828)—Peruvian rock seabass; Serranus cabrilla (Linnaeus, 1758)—Comber; Serranus hepatus (Linnaeus, 1758)—Brown comber; Syngnathidae: Syngnathus abaster Risso, 1827 (=Syngnathus nigrolineatus Eichwald, 1831)—Black-striped pipefish; Trachinidae: Trachinus draco Linnaeus, 1758—Greater weever; Triglidae: Chelidonichthys cuculus (Linnaeus, 1758) (=Trigla cuculus Linnaeus, 1758; Trigla pini Bloch, 1793)—Red gurnard; Chelidonichthys lucerna (Linnaeus, 1758) (=Trigla hirundo Linnaeus, 1758; Trigla lucerna Linnaeus, 1758)—Tub gurnard; Chelidonichthys obscurus (Walbaum, 1792) (=Trigla obscura Walbaum, 1792)—Longfin gurnard; Trigloporus lastoviza (Bonnaterre, 1788) (=Trigla lineata Gmelin, 1789)—Streaked gurnard.
Intermediate hosts: Palaemonidae: Palaemon adspersus Rathke, 1837—Baltic prawn; Palaemon elegans Rathke, 1837—Grass prawn; Phasianellidae: Tricolia tenuis (Michaud, 1829)—Marine gastropod sp.
Sites of infection: Intestine; pylorica caeca; stomach (definitive hosts); expected to be thoracic muscles (prawn intermediate hosts—see Meenakshi et al. 1993).
Distribution: Gulf of Naples (Rudolphi 1819); France? (Diesing 1850); Bergen, Norway (Olsson 1868); Gulf of Genoa (Willemoes-Suhm 1871); Trieste (Stossich 1883, 1889); Germany? (Linstow 1889); Italy (Collection of the Museum of Pisa) (Sonsino 1891); Elba, Italy (Parona 1902); Catania, Sicily (Barbagallo and Drago 1903); Unknown locality (Hofer 1904; Fuhrmann 1928); Firth of Clyde, Scotland (Nicoll 1910); Plymouth, England (Nicoll 1914; Baylis and Jones 1933); Ireland (Little 1929); South Devon, British Isles (Baylis 1939); Sevastopol’, Ukraine, Black Sea (Osmanov 1940; Markevich 1952); Black Sea (Osmanov 1940; Naĭdenova and Dolgikh 1969; Korniychuk 2000a, 2001a, b); Cullercoats, England (Crofton 1941); Aberystwyth, Wales in U.K. (Rees 1945); Northern Azov (in small rivers) (Chaplina and Antsishkina 1961); Atlantic Ocean (Kovaleva 1970); Adriatic Sea near Split, Yugoslavia (=Croatia) (Sey 1970); Saronicos Gulf, Athens, Greece (Papoutsoglou 1976); Italian seas (Orecchia and Paggi 1978); Azov and Black Seas, U.S.S.R. (Mordvinova 1979); Lagoon of Languedoc (“Etang de Thau”) in southern France (Maillard and Aussel 1988; Reversat 1990; Reversat and Silan 1991, 1993; Reversat et al. 1991a, b); Southwest Indian Ocean & Mediterranean Sea (Parukhin 1989); Persian Gulf (Kalantan et al. 1992); Peruvian coast (Tantalean et al. 1992); Taiwan Strait (Wang et al. 1992); Mediterranean Sea (Naĭdenova and Mordvinova 1997); Western coast of British Isles (Treasurer 1997); Scandola Nature Reserve of Corsica (Bartoli et al. 2005); Bay of Bizerte, Tunisia (Abdalah et al. 2010).
Reports: Cobbold (1860; see Isaĭchikov 1928, p. 46), Odhner (1902b), Linton (1910), Nicoll (1915), Crofton (1941), Skrjabin and Koval (1958), Yamaguti (1958a [Pl. 12, Fig. 151], 1971 [Fig. 405]), Ronald (1959), Tantalean et al. (1982), Zhang et al. (1999), Bona et al. (2000) and Liu et al. (2010).
Specimens deposited: Vouchers: BM(NH): #1932.11.22.45,46; 1938.11.4.206; HWML #16366.
Remarks: This species is considered the type species for the genus Helicometra (see Cribb 2005, p. 501). Odhner (1902c) regarded Helicometra labri (Stossich, 1886) (=Distomum labri Stossich, 1886) as a junior syn. of H. pulchella. Palombi (1929a, b, 1931) considered this species to be synonymous with H. fasciata and Manter (1933) considered H. pulchella of Nicoll (1914) to be a synonym of H. fasciata. Markevich (1952) listed Distoma pulchellum Rudolphi, 1819, Distoma labri Stossich, 1886 nec van Beneden, 1890, Allocreadium labri (Stossich, 1886) Odhner, 1901 and Helicometra mutabilis (Stossich, 1902) as junior synonyms of H. pulchella. Yamaguti (1971) considered this species to be the type species within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors; furthermore, he noted that the specimens of H. pulchella of Nicoll (1910) may be referable to H. fasciata. Naĭdenova (1974) considered H. pulchella to be junior syn. of H. fasciata. By “page priority”, Sekerak and Arai (1974) considered H. sinuata and H. fasciata junior syns. of H. pulchella as well as H. epinepheli, H. hypodytis, H. markewitschi, H. aposinuata, H. dochmosorchis and H. upapalu. Bray (1979, p. 401) restated that Sekerak and Arai (1974) used the name H. pulchella, rejecting the name H. fasciata used by Palombi (1931) because of “page priority”; however, he quoted ICZN Art 24(a) which states “that the priority in cases of simultaneous publication in the same work is determined by the actions of the first reviser, and, in the case of Helicometra, Palombi (1931) qualifies on all counts. As Palombi considers H. pulchella a junior synonym of H. fasciata, and as the name H. fasciata has been used more than twice as often as H. pulchella (see Sekerak and Arai’s list), it seems more appropriate that H. fasciata should continue to be used.” This species is included in keys of Wallin (1909), Isaĭchikov (1928), Palombi (1929a), Manter (1933), Skrjabin and Koval (1958), Skrjabin (1964), and Sekerak and Arai (1974). It is important to note that identifying the type host for H. pulchella, reported by Rudolphi (1819) as “Labrus cynaedus”, is problematic. Maillard and Aussel (1988, p. 497) stated that the species Labrus cynaedus Linnaeus, 1758 was not indexed in UNESCO (1973), but this reference stated that this taxon designation was synonymous with Labrus rupestris Linnaeus, 1758, which Froese and Pauly (2013) identify as the goldskinny wrasse, Ctenolabrus rupestris (Linnaeus, 1758).
Helicometra rakusai Zdzitowiecki, 1997
(Syn. Helicometra sp. Prudhoe and Bray, 1973).
Host: Nototheniidae: Trematomus loennbergii Regan, 1913 (=Trematomus loennbergi [sic] Regan, 1913)—Scaly rockcod [type host].
Sites of infection: Pyloric caecum; small intestine.
Distribution: Halley Bay and Kap Norvegia, Weddell Sea off Antarctica (Zdzitowiecki 1997b, 2002).
Reports: Klimpel et al. (2001, 2009).
Specimens deposited: Holotype: Museum of Zoology, Warszawa #MZPW 1790; Paratypes: BM(NH) #1996.12.13.1–4; Museum of Zoology, Warszawa #MZPW 1791–1793.
Remarks: Zdzitowiecki (1998, p. 28) commented that Helicometra sp. of Prudhoe and Bray (1973) appears similar to H. rakusai; however, the “specimens deposited in the Natural History Museum in London are strongly contracted and their identification should be accepted with caution.” Aken’Ova et al. (2006) placed this species in Group I, while Hassanine (2007) placed it in Group IV.
Helicometra rectisaccus (Fischthal and Kuntz, 1964)
(Syns. Helicometra [Helicometra] rectisaccus Fischthal and Kuntz, 1964; Helicometra nasae Nagaty and Abdel Aal, 1962; Stenopera rectisaccus Fischthal and Kuntz, 1964).
Host: Holocentridae: Sargocentron violaceum (Bleeker, 1853) (=Holocentrus violaceus Bleeker, 1853)—Violet squirrelfish [type host].
Site of infection: Small intestine.
Distribution: Puerto Princesa, Palawan Island, Philippines (Fischthal and Kuntz 1964).
Reports: Fischthal and Kuntz (1965), Yamaguti (1971).
Specimens deposited: “Type” and paratype: USNPC #37896, 37897.
Remarks: This species was in Stenopera; however, this genus is now considered a junior synonym of Helicometra (see Siddiqi and Cable 1960). Pritchard (1966, p. 198) disagreed and re-established Stenopera as a valid genus returning H. rectisaccus to Stenopera; however, later authors still consider this genus as synonymous with Helicometra. Pritchard (1966) also stated that “Stenopera rectisaccus is probably a synonym of S. nasae” (a view supported by Sekerak and Arai [1974, p. 729]) and included this species in her key. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Aken’Ova et al. (2006) placed this species in Group III.
Helicometra robinsorum Overstreet and Martin, 1974
Hosts: Synaphobranchidae: Ilyophis brunneus Gilbert, 1891—Muddy arrowtooth eel; Synaphobranchus oregoni Castle, 1960—Cutthroat eel sp. [type host].
Sites of infection: Intestine; stomach.
Distribution: Gulf of Mexico, Straits of Florida (Overstreet and Martin 1974).
Reports: Klimpel et al. (2001, 2009), Overstreet et al. (2009).
Specimens deposited: Holotype and paratypes: USNPC #72847, 72848; HWML #20030.
Remarks: Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra scorpaenae Prudhoe and Bray, 1973
(Syn. Helicometra fasciata [Rudolphi, 1819]).
Host: Scorpaenidae: Scorpaena papillosa (Schneider and Forster, 1801) (=Scorpaena cruenta Solander, 1842)—Red rock cod [type host].
Site of infection: Expected to be intestine.
Distribution: Maria Island, Tasmania (Prudhoe and Bray 1973).
Specimens deposited: Syntype: BM(NH) 1975.1.30.28. Prudhoe and Bray (1973) stated that material described in their report was divided between the BM(NH) and the South Australian Museum with no holotypes selected; descriptions of new species (e.g. H. scorpaenae) was based on designated syntypes.
Remarks: Bray (1987a) considered this species to be a junior syn. of H. fasciata. Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I. This species is not to be confused with Neohelicometra scorpaenae Siddiqi and Cable, 1960, which is currently the only species within Neohelicometra (Siddiqi and Cable, 1960), a genus which differs from Helicometra in having caeca that open through separate ani as opposed to ending blindly (see Cribb 2005; Hassanine 2007, p. 19).
Helicometra sebastis (Sekerak and Arai, 1974)
(Syns. Helicometra sp. Arai, 1969; Helicometra sp. Sekerak and Arai, 1973; Neohelicometra sebastis Sekerak and Arai, 1974 [see Hassanine 2007, p. 22]).
Hosts: Sebastidae: Sebastes aleutianus (Jordan and Evermann, 1898)—Rougheye rockfish; Sebastes alutus (Gilbert, 1890)—Pacific ocean perch [type host]; Sebastes babcocki (Thompson, 1915)—Redbanded rockfish; Sebastes caurinus Richardson, 1844—Copper rockfish; Sebastes diploproa (Gilbert, 1890)—Splitnose rockfish; Sebastes flavidus (Ayers, 1862)—Yellowtail rockfish; Sebastes helvomaculatus Ayers, 1859—Rosethorn rockfish; Sebastes maliger (Jordan & Gilbert, 1880)—Quillback rockfish; Sebastes nebulosus Ayres, 1854—China rockfish; Sebastes nigrocinctus Ayres, 1859 (=Sebastes nigroncintus [sic] Ayers, 1859)—Tiger rockfish; Sebastes reedi (Westrheim & Tsuyuki, 1967)—Yellowmouth rockfish; Sebastes ruberrimus (Cramer, 1895)—Yelloweye rockfish; Sebastes variegatus Quast, 1971—Harlequin rockfish; Sebastes zacentrus (Gilbert, 1890)—Sharpchin rockfish.
Sites of infection: Intestine; pyloric caecum.
Distribution: Northeastern Pacific Ocean, from southwestern Oregon to Cape Ommaney, Alaska (Sekerak and Arai 1974); Northeastern Pacific Ocean (Sekerak 1975); La Pérouse Bank and Strait of Georgia, British Columbia, Cape Ommaney and Icy Bay, Alaska in northeastern Pacific Ocean (Sekerak and Arai 1977); Northern Pacific Ocean (Holmes 1990, 1991); Pacific coast of North America (Stanley et al. 1992).
Reports: Love and Moser (1976, 1983), Margolis and Arthur (1979), Holmes (1990), Gibson (1996), Klimpel et al. (2001, 2009) and Love et al. (2002).
Specimens deposited: Holotype and paratypes: National Museums of Canada, Ottawa #65, 66.
Remarks: Bray (1979), noting that fixation method produced variability in the terminal vs subterminal appearance of the oral sucker (the main feature used in the key by Sekerak and Arai [1974] to distinguish Neohelicometra and Helicometra), moved N. sebastis in to Helicometra. Gibson (1996, p. 170) considered Helicometra sp. of Arai (1969) and of Sekerak and Arai (1973) as well as Neohelicometra sebastis Sekerak and Arai, 1974 all as junior syns. of this species, which he included in a key to three species of Helicometra. Hassanine (2007) placed this species in Group IV.
Helicometra selaroidis Shen, 1986
Host: Carangidae: Selaroides leptolepis (Cuvier, 1833)—Yellowstripe scad [type host].
Site of infection: Intestine.
Distribution: Off Hainan Island, South China Sea (Shen 1986, 1990).
Reports: Zhang et al. (1999), Zongguo (2001) and Liu et al. (2010).
Specimens deposited: Type specimens in the Institute of Oceanology, Academia Sinica.
Remarks: Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra sinipercae Wang, 1981
Host: Percichthyidae: Siniperca chuatsi (Basilewsky, 1855)—Mandarin fish [type host].
Site of infection: Intestine.
Distribution: Fujian Province, China (Wang 1981).
Reports: Pan et al. (1990), Zhang et al. (1999).
Specimens deposited: Type specimens in the Parasitological Research Laboratory, Fujian Teachers University.
Remarks: Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group I.
Helicometra sinuata (Rudolphi, 1819)
(Syns. Allocreadium sinuatum [Rudolphi, 1819] [see Odhner 1901; Palombi 1929a; Yamaguti 1971, p. 190]; Distoma sinuatum Rudolphi, 1819 [see Rudolphi 1819; Dujardin 1845; Willemoes-Suhm 1871; Palombi 1929a]; Distomum sinuatum Rudolphi, 1819 [see Diesing 1850; Carus 1884; Stossich 1886; Parona 1887, 1902; Palombi 1929a]; Helicometra fasciata Stossich, 1885 [in schedis] non Rudolphi, 1819 [see Palombi 1929a, b]; Helicometra [Helicometra] sinuata [Rudolphi, 1819]; Helicometra pulchella [Rudolphi, 1819]; Helicometra sinuatum [sic] [Rudolphi, 1819]; From Palombi 1929a, p. 281—see Odhner 1902c; Stossich 1904b; Wallin 1909; Palombi 1929b).
Hosts: Carapidae: Carapus acus (Brünnich, 1768) (=Fierasfer acus [Brünnich, 1768]; Fierasfer imberbis [Cuvier, 1815]; Ophidium imberba [sic] Cuvier, 1815; Ophidium imberbe [sic] Cuvier, 1815)—Pearl fish [type host]; Muraenidae: Moray eel sp.; Ophidiidae: Ophidion barbatum Linnaeus, 1758 (=Ophidium [sic] barbatum Linnaeus, 1758)—Snake blenny; Syngnathidae: Hippocampus hippocampus (Linnaeus, 1758) (=Hippocampus antiquorum Leach, 1814)—Short snouted seahorse; Trachinidae: Trachinus draco Linnaeus, 1758—Greater weever.
Site of infection: Intestine.
Distribution: Gulf of Naples (Rudolphi 1819; Parona 1912); Gènes (=Genoa), Italy (Willemoes-Suhm 1871); Island of Elba, Italy (Parona 1902); Roscoff in NW France (Sproston 1938); Europe and Port Culebra, Costa Rica (Manter 1940); Mediterranean Sea (Reversat et al. 1991b).
Reports: Odhner (1902a), Palombi (1929a, b), Yamaguti (1958a, 1971) and Bona et al. (2000).
Specimen deposited: Voucher: HWML #1829.
Remarks: This species is included in keys of Wallin (1909), Isaĭchikov (1928), Palombi (1929a), Manter (1933), Skrjabin and Koval (1958), and Skrjabin (1964). Palombi (1929a) redescribed this species from Rudolphi’s types, discovered four specimens of this species in the 1901 collection of Stossich (see Palombi 1929b) and later, based on variation in testes texture, Palombi (1931) regarded H. sinuata as a junior syn. of H. fasciata. Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella.
Helicometra sprenti Aken’Ova, Cribb and Bray, 2006
Host: Platycephalidae: Platycephalus speculator Klunzinger, 1872—Yank flathead [type host].
Site of infection: Intestine.
Distribution: Off Point Peron, Fremantle, Western Australia (Aken’Ova et al. 2006).
Specimens deposited: Holotype: Queensland Museum, Reg. No. QM G222961; Paratypes: Queensland Museum, Reg. Nos. QM G222962–G222964, BM(NH) #2005.3.11.1–2.
Remarks: Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group II.
Helicometra tenuifolia Woolcock, 1935
(Syns. Helicometra [Metahelicometra] tenuifolia Woolcock, 1935; Helicometra neosebastodis Crowcroft, 1947 [see Sekerak and Arai 1974, p. 726; Hassanine 2007, p. 22]).
Host: Sebastidae: Helicolenus percoides (Richardson & Solander, 1842)—Red gurnard perch [type host].
Site of infection: Intestine.
Distribution: Port Philip Bay, Victoria, Australia (Woolcock 1935).
Reports: Young (1939), Yamaguti (1958a, 1971).
Specimens deposited: Sekerak and Arai (1974, p. 726) stated that “the entire collection of Woolcock (1935) seems to be missing; hence, type specimens of H. tenuifolia are unavailable for examination.” Indeed, we could find no information regarding deposition of specimens for this species; however, Violet Woolcock, at the time of publishing the type description of H. tenuifolia, was in the Dept. of Zoology, University of Melbourne, Australia.
Remarks: This species is included in keys of Skrjabin and Koval (1958), Skrjabin (1964), and Sekerak and Arai (1974). Yamaguti (1971) considered this species to be within the subgenus Metahelicometra Yamaguti, 1971, a subgenus now considered a junior syn. of Helicometra (Cribb 2005). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group II.
Helicometra torta Linton, 1910
(Syns. Helicometra [Metahelicometra] torta Linton, 1910; Helicometra pretiosa Bravo-Hollis and Manter, 1957 [see Overstreet 1969, p. 144; Yamaguti 1971, p. 190; Sekerak and Arai 1974, p. 724; Hassanine 2007, p. 22]).
Hosts: Lutjanidae: Lutjanus griseus (Linnaeus, 1758)—Grey snapper; Mullidae: Pseudupeneus maculatus (Bloch, 1793)—Spotted goatfish; Serranidae: Cephalopholis panamensis (Steindachner, 1877) (=Epinephelus panamensis [Steindachner, 1877])—Pacific graysby; Epinephelus adscensionis (Osbeck, 1765)—Rock hind; Epinephelus labriformis (Jenyns, 1840)—Starry grouper; Epinephelus morio (Valenciennes, 1828)—Red grouper; Epinephelus striatus (Bloch, 1792) (=Epinepheles [sic] striatus [Bloch, 1792])—Nassau grouper [type host]; Paralabrax maculatofasciatus (Steindachner, 1868) (=Paralabrax maculofasciatus [sic] [Steindachner, 1868])—Spotted sand bass; Sparidae: Calamus calamus (Valenciennes, 1830)—Saucereye porgy.
Sites of infection: Intestine; pyloric caecum; stomach.
Distribution: Dry Tortugas, Florida (Linton 1910; Manter 1933, 1947); Cape Elena, Ecuador (Manter 1940); N. Bimini, Bahamas, British West Indies (Sogandares-Bernal 1959); Cabo Rojo, Puerto Rico (Siddiqi and Cable 1960); Biscayne Bay, Florida (Overstreet 1969); Caribbean Sea off Belize (Fischthal 1977); Playa Sardinero, Mona Island, and Media Luna and Enrique Reefs, La Parguera in Puerto Rico (Dyer et al. 1985, 1992); Gulf of Mexico (Yucatan Peninsula) (Moravec et al. 1997); Off Yucatan Peninsula, Mexico (Vidal-Martínez et al. 1998); Central American and Mexican Pacific coasts (Pérez-Ponce de León et al. 1999).
Reports: Galtsoff (1954), Yamaguti (1958a, 1971), Sparks (1972), Sadovy and Eklund (1999), Cribb et al. (2002) and Overstreet et al. (2009).
Specimens deposited: Syntypes: USNPC #08438; Vouchers: USNPC #08439–08440, 36211, 39359, 74233, 77784, 77957, 81621, 81629; HWML #67, 257, 264, 265, 1214, 22534, 124756; CNHE #HE3160, 3219–3220.
Remarks: This species is included in keys of Linton (1910), Palombi (1929a), Manter (1933), Skrjabin and Koval (1958), Skrjabin (1964), and Sekerak and Arai (1974). Yamaguti (1971) considered this species to be within the subgenus Metahelicometra Yamaguti, 1971, a subgenus now considered a junior syn. of Helicometra (Cribb 2005). Aken’Ova et al. (2006) and Hassanine (2007) placed this species in Group II.
Helicometra upapalu Yamaguti, 1970
(Syns. Helicometra [Helicometra] upapalu Yamaguti, 1970; Helicometra pulchella [Rudolphi, 1819]).
Host: Apogonidae: Apogon taeniopterus Bennett, 1836 (=Apogon menesemus Jenkins, 1903)—Bandfin cardinalfish [type host].
Site of infection: Intestine.
Distribution: Hawaii (Yamaguti 1970).
Report: Yamaguti (1971).
Specimen deposited: Holotype: USNPC #63580; Paratype: Meguro Parasitological Museum, Tokyo, Japan, MPM Coll. #15024.
Remarks: Included in key to Helicometra species in Hawaiian fishes by Yamaguti (1970). Yamaguti (1971) considered this species to be within the subgenus Helicometra (Odhner, 1902), a subgenus not recognized by later authors. Sekerak and Arai (1974) believed this species to be a junior syn. of H. pulchella.
Additional Comments:
Travassos et al. (1969, p. 391–393) referred to “Helicometra asymmetrica Travassos, 1922” and “Helicometra asymetrica Travassos, 1928” as junior syns. of Helicotrema asymmetricum (Travassos, 1922). We believe the authors here mistakenly used the taxon Helicometra Odhner, 1902 instead of the correct genus designation Helicotrema Odhner, 1912 in these descriptions (see Travassos 1922, 1928) as the former genus is within Opecoelidae Ozaki, 1925 and infects many varieties of fish while the latter genus is within Liolopidae Odhner, 1912 and parasitizes lizards (e.g. the green iguana, Iguana iguana [Linnaeus, 1758]) (see Yamaguti 1971; Niewiadomska 2002).
On that same note, Woolcock (1935, p. 313) pointed out that Travassos (1928) used the name “Helicometra spirale” when undoubtedly Helicotrema spirale (Diesing, 1850) was meant.
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Blend, C.K., Dronen, N.O. A review of the genus Helicometra Odhner, 1902 (Digenea: Opecoelidae: Plagioporinae) with a key to species including Helicometra overstreeti n. sp. from the cusk-eel Luciobrotula corethromycter Cohen, 1964 (Ophidiiformes: Ophidiidae) from the Gulf of Mexico. Mar Biodiv 45, 183–270 (2015). https://doi.org/10.1007/s12526-014-0250-3
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DOI: https://doi.org/10.1007/s12526-014-0250-3