A Checklist of the Parasites of Ayu (Plecoglossus Altivelis Altivelis) (Salmoniformes: Plecoglossidae) in Japan (1912-2007)
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J. Grad. Sch. Biosp. Sci. Hiroshima Univ. (2007), 46:59~89 REVIEW A Checklist of the Parasites of Ayu (Plecoglossus altivelis altivelis) (Salmoniformes: Plecoglossidae) in Japan (1912-2007) 1) 1) 2) Kazuya Nagasawa , Tetsuya Umino and Mark J. Grygier 1)Graduate School of Biosphere Science, Hiroshima University 1-4-4 Kagamiyama, Higashi-Hiroshima, Hiroshima 739-8528, Japan 2)Lake Biwa Museum, 1091 Oroshimo, Kusatsu, Shiga 525-001, Japan Abstract The ayu or sweetfish Plecoglossus altivelis altivelis is distributed in many rivers and some lakes in Japan and also occurs in rivers of the Korean Peninsula and along the east coast of China and northern Vietnam. In Japan, this species is one of the most important freshwater fishes for commercial fisheries, aquaculture, and recreational fishing. In the present paper, information on the protistan and metazoan parasites of ayu in Japan is compiled based on the literature published for 96 years between 1912 and 2007, and the parasites, including 29 named species and those not identified to species level, are listed by higher taxon as follows: Ciliophora (no named species), Microspora (1), Myxozoa (1), Trematoda (13), Monogenea (3), Cestoda (1), Nematoda (1), Acantho- cephala (3), Copepoda (4), and Branchiura (2). For each taxon of parasite, the following information is given: its currently recognized scientific name, any original combination, synonym(s), or other previous identification used for the parasite occurring in ayu; habitat (freshwater, brackish, or marine); site(s) of infection within or on the host; known geographical distribution in Japanese waters; and the published source of each locality record. There has been no record of parasites from a subspecies of ayu, the ryukyu-ayu Plecoglossus altivelis ryukyuensis. The myxozoan Mitraspora plecoglossi Fujita is transferred to the genus Hoferellus (as H. plecoglossi n. comb.), Acanthocephalus opsariichthydis Yamaguti, 1935 is determined to be the nomenclaturally correct spelling and date of that acanthocephalan species, and synonymy of the acanthocephalan Echinorhynchus oblitus Golvan with E. cotti Yamaguti is formally accepted. Key words: ayu; checklist; Echinorhynchus oblitus; Hoferellus plecoglossi n. com.; parasites; Plecoglossus altivelis altivelis INTRODUCTION The ayu Plecoglossus altivelis altivelis Temminck and Schlegel, also called sweetfish, is a fish of the order Salmoniformes that belongs to the family Plecoglossidae (Hosoya, 2002). In Japan, ayu are distributed in many rivers and some lakes from western Hokkaido to southern Kyushu. A land- locked population of ayu inhabits Lake Biwa, central Japan. Ayu also occur in rivers of the Korean Peninsula and along the east coast of China and northern Vietnam. A population of ayu that previously inhabited Taiwan has gone extinct. The ryukyu-ayu Plecoglossus altivelis ryukyuensis Nishida is a subspecies of ayu that currently occurs only in streams on the island of Amami-Oshima Accepted : September 4, 2007 07Kazuya Nagasawa他2名.indd 59 2007/12/08 11:17:19 60 Kazuya Nagasawa, Tetsuya Umino and Mark J. Grygier in southern Japan (Hosoya, 2002). This subspecies currently has been successfully reintroduced into streams and lakes on the island of Okinawa (Ikehara, 2004), where it had gone extinct around 1980. Ayu are one of the most important freshwater fishes for commercial fisheries, aquaculture, and recreational fishing in Japan. Much work has been conducted on the parasites of ayu in Japan because of their importance in fisheries and human parasitology, but the only review of these parasites (Kobayashi, 1938) is long out of date. To provide a comprehensive update, information on the protistan and metazoan parasites of ayu in Japan is compiled anew here based on the literature published for 96 years between 1912 and 2007. In total, 29 named species of parasites are listed herein along with those parasites not identified to species level. As far as we know, there has been no record of parasites from ryukyu-ayu P. altivelis ryukyuensis. In the present checklist, parasites are arranged by higher taxon in the following order: Ciliophora, Microspora, Myxozoa, Trematoda, Monogenea, Cestoda, Nematoda, Acanthocephala, Copepoda, and Branchiura. The format is almost the same as in the checklists of Margolis and Arthur (1979) and McDonald and Margolis (1995). Within each higher taxon, genera and species are listed alphabetically. For each taxon of parasite, the following information is provided: 1) The current scientific name, including author(s) and date(s), followed by any original combination, recognized synonym(s), or other identifications(s) that have been used in establishing records from ayu in Japan. No attempt has been made to evaluate the taxonomic validity of the published reports; however, a generic reassignment is formally affirmed for the myxozoan Mitraspora plecoglossi as an absolute necessity under the current classification of this group. In addition, Harada’s (1935) view that the acanthocephalan Acanthocephalus aculeatus is a junior synonym of A. echigoensis in not accepted herein because his proposal seems to require reassessment based on abundant material. On the other hand, the synonymy suggested by Amin et al. (2007) of two nominal species of the acanthocephalan genus Echinorhynchus is formally adopted herein. 2) The habitat in which the parasite was acquired and normally completes its life cycle is given as FW for fresh waters, B for brackish waters, and M for marine waters. 3) The Site(s) of infection of the parasite in or on its host. If the site was not given in the original record, the likely site was determined from other records and is enclosed in square brackets. 4) The Distribution of the parasite is indicated by prefecture (boundaries shown in Fig. 1), in geographical order from northeast to southwest. For marine or brackish-water species, the name of the prefecture nearest the collection site is given. 5) The Record(s). The authors responsible for the records are listed in chronological order. If a parasite has been reported more than once, the references are numbered, but not when there has been only one record of the parasite. Each reference is followed by the locality or localities given in two parts, first the prefecture(s) and then the detailed collection locality or localities from which the parasite was reported. If no locality record was given, the geographical locality is shown by a dash (-). When all records are from the same prefecture, only the detailed collection locality or localities are listed. 6) Under Remarks, explanatory comments are given on systematics, nomenclature, useful references, and notes on specific items such as tentative parasite identifications in the original reports. 7) The References section includes works directly cited in the Parasite List; the Supplementary References are publications dealing with various aspects of parasites of Japanese ayu but not 07Kazuya Nagasawa他2名.indd 60 2007/12/08 11:17:19 Checklist of the parasites of ayu in Japan 61 Fig. 1. Map of Japan showing the prefectural boundaries. The following prefectural names are arranged in alphabetical order: Aichi-24; Akita-7; Aomori-5; Chiba-15; Ehime-41; Etorofu Island-1; Fukui-26; Fukuoka-43; Fukushima-10; Gifu-23; Gunma-14; Hiroshima-37; Hokkaido-4; Hyogo-33; Ibaraki-12; Ishikawa-25; Iwate-6; Kagawa-39; Kagoshima-49; Kanagawa-18; Kochi-42; Kumamoto-47; Kunashiri Island-3; Kyoto-29; Mie-28; Miyagi-8; Miyazaki-48; Nagano-20; Nagasaki-45; Nara-30; Niigata-11; Oita-46; Okayama-35; Okinawa-50; Osaka-31; Saga-44; Saitama-16; Shiga-27; Shikotan Island-2; Shimane-36; Shizuoka-21; Tochigi-13; Tokushima-40; Tokyo-17; Tottori-34; Toyama-22; Wakayama-32; Yamagata-9; Yamaguchi-38; and Yamanashi-19. containing original parasite records. If only a Japanese title was given by the original author(s), our translation of the title into English is provided in square brackets. This checklist is the fifth in the following series of published synopses of the parasites of commercially important fishes and shellfishes in Japan: Nagasawa et al. (1987) for the parasites of salmonids; Nagasawa et al. (1989) for the parasites of freshwater fishes in Hokkaido; Nagasawa (1993a) for the parasites of squids and cuttlefishes; and Nagasawa (1993b) for the parasites of gadids. PARASITE LIST Ciliophora Trichodina sp. (FW) Site of infection: gills Distribution: Aichi 07Kazuya Nagasawa他2名.indd 61 2007/12/08 11:17:19 62 Kazuya Nagasawa, Tetsuya Umino and Mark J. Grygier Record: Nakajima et al. 1974 (Mito) Remarks: There has been no study of the morphology and taxonomy of the species. The infected fish found in Aichi were transported as juveniles from Lake Biwa (Shiga) (Nakajima et al., 1974). The parasite was often found together with other gill parasites such as monogeneans Dactylogyrus sp. and copepods Ergasilus (as “Pseudergasilus”) zacconis (Nakajima et al., 1974). Although there is a record of a similar parasite (as “Cyclochaeta sp.”) from ayu in Taiwan (Abe, 1933; Kobayashi, 1938), Basson and Van As (1994) did not find any trichodinid infection of ayu from the Feitsui Reservoir in Taiwan. Microspora Glugea plecoglossi Takahashi and Egusa, 1977 (FW) Includes: Glugea sp. Awakura, 1974; Suzuki et al., 1974; Takahashi et al., 1975, 1979; Nakajima and Egusa, 1975a, 1975b, 1975c, 1979; Shiose and Tominaga, 1976; Inada and Hamaguchi, 1987 Sites of infection: visceral organs (pyloric caeca, abdominal cavity, abdominal membrane, ovary, testis, fat, spleen, liver, heart), musculature, gills, fins,