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1 Biosphere Sci (2016) 総説日本産コイ科魚類に寄生する単生類フタゴムシ Eudiplozoon nipponicum と近縁未同定種に関する解説 [ 付録 : 亀谷了博士の研究業績目録 ] * 要旨 Eudiplozoon nipponicum (Goto, 1891) Diplozoon sp. Diplozoon sp. Paradilpozoon skrjabini Akhmerov, 1974 キーワード Diplozoon sp. Eudiplozoon nipponicum Paradilpozoon skrjabini 緒言 Nagasawa et al., 2007, 2013, 2014 Maneepitaksanti and Nagasawa, 2012 Nagasawa and Obe, Eudiplozoon nipponicum (Goto, 1891) Paradilpozoon skrjabini Akhmerov, 1974 Shimazu et al., Diplozoon sp * ornatus@hiroshima-u.ac.jp

2 40 Tribolodon spp. Diplozoon sp Nagasawa et al., 1987 Ogawa, 1994 Diplozoon sp Diplozoon sp Diplozoon sp Diplozoon sp Diplozoon sp フタゴムシ類の分類学的位置 2 2 * 1 Khotenovsky 1985 Shimazu et al Class Monogenea van Beneden, 1858 Subclass Polyopisthocotylea Odhner, 1912 Order Mazocraeidea Bychowsky, 1937 Family Diplozoidae Palombi, 1949 Subfamily Diplozoinae Palombi, 1949 Genus Eudiplozoon Khotenovsky, 1985 Eudiplozoon nipponicum (Goto, 1891) Khotenovsky, 1985 Diplozoon nipponicum Goto, 1891 Genus Paradiplozoon Akhmerov, 1974 鯎 Paradiplozoon skrjabini Akhmerov, Diplozoon Diplozoon nipponicum Goto Diplozoon nipponicum X 2 Diplozoidae Diplozoinae Diplozoidae Goto 1891 Diplozoon Khotenovsky 1985 Eudiplozoon *

3 Paradiplozoon Sicard et al Eudiplozoon sp. Shimazu et al Eudiplozoon nipponicum フタゴムシの生物学五島淸太郎による記載 24 Diplozoon nipponicum Goto, Carassius vulgaris 2013 Carassius sp. Carassius buergeri subsp. 2 Shimazu et al most likely referring to Gin-buna Goto, Kamegai a pond in the Shimura district, a suburb of Tokyo City 19 形態 Goto Fig. 1A 1B 2 X 6.7 mm mouth 1 sticky gland 1 oral sucker pharynx intestine vitellaria 2 1 opisthaptor 3 1 ovary testis 2 3 clamp

4 42 m A B os sg ph i v i ov t op c Fig. 1. Eudiplozoon nipponicum. A, entire body; B, flattened entire body. (Illustrated from Goto [1891, figs. 1 2]). c, clamp; i, intestine; m, mouth; op, opisthaptor; os, oral sucker; ov, ovary; ph, pharynx; sg, sticky gland; t, testis; v, vitellaria. 宿主 Carassius spp. = Carassius vulgaris Goto, 1891 Carassius carassius Kamegai, 1968 Carassius spp Carassius carassius 1976 crucian carp Kawatsu, 1978 Carassius sp. = Carassius auratus langsdorfii 1986, 1987 Carassius auratus subsp. Nagasawa et al., 1989, Ogawa, Carassius auratus Sicard et al., 2003 Shimazu et al., Carassius buergeri subsp. 2 = Carassius auratus subsp Cyprinus carpio 1966 Kamegai, 1968, 1970a, 1970b, * Carassius auratus * Diplozoon sp. Dilpozoon sp. * 2 Carassius cuvieri = 1977a Carassius buergeri grandoculis = Carassius auratus grandoculus 1975b * a Diplozoon sp. 3.3 mm * c, 1982b, 1983a, 1983b, 1985b, 1987a 1981a, 1981b 1984

5 43 Sea of Okhotsk 2 1 Sea of Japan North Pacific Ocean Fig. 2. Map of the Japanese Archipelago, showing the collection sites of Eudiplozoon nipponicum from crucian carps (Carassius spp.) and common carp (Cyprinus carpio). 1, Lake Tôro (Hokkaido); 2, Ishikari- Furukawa (Hokkaido); 3, Yabuki Town (Fukushima); 4, Lake Yunoko (Tochigi); 5, Lake Kitaura and Itako (Ibaraki); 6, Lake Kasumigaura (Tochigi); 7, Isezaki City (Gunma); 8, Ogawa Town (Saitama); 9, Tama River system (Tokyo); 10, Tsurumi River (Kanagawa); 11, Sakai River (Kanagawa); 12, Chikuma River (Nagano); 13, Lake Suwa (Nagano); 14, Lake Biwa (Shiga); 15, Kurose River (Hiroshima). Dotted lines show prefectural boundaries. 国内分布 Nagasawa et al., 1989, Ogawa, 1994 Nagasawa et al., 1989, Ogawa, 1994 Kamegai, 1968 Kamegai, , Kamegai, , Kamegai, 1968, 1970b Kamegai, 1968 Kamegai, 1968, 1986, Kawatsu, 1978

6 Sicard et al., 2003 Shimazu et al., Kamegai, 1968 Fig. 2 * 5 Goto Kamegai, 1968 Kamegai, 1968 生活史 oncomiracidium diporpa oncomiracidium Kamegai, 1968 Kamegai 1968 fig m filament ºC 8 Kamegai, ºC ºC m Kamegai, larval hook Kamegai, * ºC mm 2 X Goto, 成熟産卵の季節変化 Kamegai, * b 1974b * 6 * 4

7 45 Kawatsu, ºC ºC , 1986 Kamegai, 1970b 宿主サイズと寄生状況との関係 * 7 10 g % 10 g 100% Kawatsu, 1978 病害性と対策 Kawatsu, Kawatsu, ppm 48 87% 水産業との関係 1966 Kamegai, % 30.3% Kamegai, cm フタゴムシに関する解説 * b 9 cm 15 cm

8 46 コイ科魚類から報告された近縁未同定種 Diplozoon sp. Shimazu et al Tribolodon hakonensis Tribolodon sachalinensis Tribolodon brandtii Phoxinus lagowskii steindachneri = Phoxinus steindachneri Phoxinus oxycephalus jouyi = Phoxinus oxycephalus Diplozoon sp. Diplozoon sp. Diplozoon sp. Diplozoon sp. 同定と宿主 Tribolodon spp. Diplozoon sp Nagasawa et al., 1989 Ogawa, 1994 Nagasawa et al Ogawa 1994 Ogawa 1994 Shimazu et al Diplozoon sp. * 8-9 Diplozoon sp Diplozoon sp. Diplozoon sp. Gnathopogon elongatus elongatus Squalidus chankaensis biwae = Squalidus biwae Opsariichthys platypus = Zacco platypus Candidia temminckii = Zacco temminckii Pseudogobio esocinus esocinus = Pseudogobio esocinus Hemibarbus barbus Carassius sp. = Carassius carassius Diplozoon sp % % % Diplozoon sp Diplozoon sp. 3 0% 1 7.7% 100% * mm 1977 * 9 Diplozoon sp. 1994c

9 47 Sea of Okhotsk 1 Sea of Japan 2 North Pacific Ocean Fig. 3. Map of the Japanese Archipelago, showing the collection sites of Diplozoon sp. from wild (closed circles) and aquarium-held (open circle) cyprinids. 1, Lake Barato (Hokkaido); 2, Niigata City Aquarium (Niigata); 3, Tone River system (Saitama); 4, Ara River system (Saitama); 5, Oh-oka River (Kanagawa); 6, Sakai River (Kanagawa); 7, Kiso River system (Gifu); 8, Takami River (Nara). Dotted lines show prefectural boundaries. * 10 1 Diplozoon sp Tribolodon nakamurai 1 Diplozoon sp. X mm mm Shimazu et al Diplozoon sp Diplozoon nipponicum Diplozoon sp. Diplozoon * 10 * a Diplozoon sp. 3.3 mm

10 48 sp. Diplozoon sp Shimazu et al Diplozoon sp. Diplozoon sp., Diplozoon sp Diplozoon sp. Shimazu et al., Shimazu et al., 2015 Diplozoon sp. 8 国内分布 Diplozoon sp. Nagasawa et al., Fig 寄生状況の季節変化と経年変化 Diplozoon sp % Diplozoon sp , % 70.0% 0% Diplozoon sp. Diplozoon sp. 病害性と対策 Diplozoon sp.

11 Diplozoon sp Diplozoon sp Diplozoon sp. Diplozoon sp. 0.5 ppm % Diplozoon sp. 100 ppm 2.5% 5.0% 1988 Diplozoon sp. 250 ppm 45 B 今後の課題 Diplozoon sp. 5 8 Diplozoon sp. 7 mm 3 mm 2

12 50 謝辞引用文献 Goto, S., On Diplozoon nipponicum, n. sp. Journal of the College of Science, Imperial University of Tokyo. 4: , 3 pls Diplozoon nipponicum 53: pp. Kamegai, S., On Diplozoon nipponicum Goto, Part II. The distribution in Japan and the developmental observation. Research Bulletin of the Meguro Parasitological Museum. (2): 1-8. Kamegai, S., 1970a. On Diplozoon nipponicum Goto, Part III. The seasonal development of the reproductive organs of Diplozoon nipponicum parasitic on Cyprinus carpio. Research Bulletin of the Meguro Parasitological Museum. (3): Kamegai, S., 1970b. An abnormal shape of reproductive organs of Diplozoon nipponicum Goto, 1891 from an old carp at Lake Kasumigaura, Japan. Research Bulletin of the Meguro Parasitological Museum. (4): Diplozoon (49): pp Diplozoon nipponicum Goto, Cyprinus carpio (83/84) pp. Kawatsu, H., Studies on the anemia of fish-ix. Hypochromic microcytic anemia of crucian carp caused by infestation with a trematode, Diplozoon nipponicum. Bulletin of the Japanese Society of Scientific Fisheries. 44: Diplozoon nipponicum Goto, 1891 (128): Diplozoon nipponicum Khotenovsky, I. A., Suborder Octomacrinea Khotenovsky. Fauna of the USSR. Monogenea. Izdatel stvo, Nauka, Leningrad. New Series, No. 132, 262 pp. (In Russian) Maneepitaksanti, W., Nagasawa, K., Actinocleidus fergusoni (Monogenea: Ancyrocephalidae), a gill parasite of bluegill (Lepomis macrochirus), new to Japan. Biogeography. 14: (23):

13 pp. Nagasawa, K., Obe, M., Spatial distribution of Neoergasilus japonicus (Copepoda: Ergasilidae) on the fins of the bluegill (Lepomis macrochirus). Journal of Natural History. 47: Nagasawa, K., Awakura, T., Urawa, S., A checklist and bibliography of parasites of freshwater fishes of Hokkaido. Scientific Reports of the Hokkaido Fish Hatchery. (44): Nagasawa, K., Inoue, A., Myat, S., Umino, T., New host records for Lernaea cyprinacea (Copepoda), a parasite of freshwater fishes, with a checklist of the Lernaeidae in Japan ( ). Journal of the Graduate School of Biosphere Science, Hiroshima University. 46: Nagasawa, K., Katahira, H., Nitta, M., Isoparorchis hypselobagri (Trematoda: Isoparorchiidae) from freshwater fishes in western Japan, with a review of its host-parasite relationships in Japan ( ). Biogeography. 15: Nagasawa, K., Sato, H., Nitta, M., The parasitic copepod Neoergasilus japonicus (Ergasilidae) from a new host Candidia sieboldii (Cyprinidae). Biogeography. 16: Ogawa, K., Monogenean parasites of freshwater fishes of Hokkaido, Japan. Scientific Reports of the Hokkaido Fish Hatchery. 48: Diplozoon sp. (44): Diplozoon sp. (44): Shimazu, T., Kobayashi, K., Tojo, K., Besprozvannkh, V. V., Ogawa, K., Paradiplozoon skrjabini (Monogenea, Diplozoidae), an ectoparasite on the gills of freshwater fishes (Cyprinidae, Leuciscinae) of Japan and Primorsky Region, Russia: a morphological and molecular study. Bulletin of the National Museum of Nature and Science, Series A. 41: : Sicard, M., Desmarais, E., Vigneux, F., Shimazu, T., Lambert, A., Molecular phylogeny of the Diplozoidae (Monogenea, Polyopisthocotylea) parasitizing 12 species of Cyprinidae (Teleostei): new data about speciation. In: Taxonomy, Ecology and Evolution of Metazoan Parasites, Eds. Combes, C. and Jourdane. 2: Presses Unversitaires de Perpignan, Perpignan Diplozoon (46): Diplozoon (47): pp pp : Diplozoon nipponicum Goto Diplozoon nipponicum Goto Diplozoon nipponicum Goto 124.

14 52 付録亀谷了博士によるフタゴムシ類に関する研究業績目録 ( 年 ) 学術論文著書 1966 Diplozoon nipponicum Goto, Cyprinus carpio (83/84): 2-9 Kamegai, S., On Diplozoon nipponicum Goto, Part II. The distribution in Japan and the developmental observation. Research Bulletin of the Meguro Parasitological Museum. (2): 1-8. Kamegai, S., 1970a. On Diplozoon nipponicum Goto, Part III. The seasonal development of the reproductive organs of Diplozoon nipponicum parasitic on Cyprinus carpio. Research Bulletin of the Meguro Parasitological Museum. (3): Kamegai, S., 1970b. An abnormal shape of reproductive organs of Diplozoon nipponicum Goto, 1891 from an old carp at Lake Kasumigaura, Japan. Research Bulletin of the Meguro Parasitological Museum. (4): Diplozoon (49): pp. 学会講演 Diplozoon nipponicum Goto, Diplozoon nipponicum Digramma alterans 14(7): alternans 1966 Diplozoon nipponicum Goto (4): Diplozoon nipponicum Goto (4): a Diplozoon nipponicum Goto larval hook biporpa 17(4): biporpa diporpa 1968b Diplozoon nipponicum Goto (6): Diplozoon nipponicum Goto (4): Diplozoon nipponicum Goto (4): Tribolodon hakonensis Diplozoon 20(4): a Diplozoon

15 b Diplozoon nipponicum Oncomiracidium a Diplozoon (46): b Diplozoon nipponicum clamp a Diplozoon nipponicum Goto filament b Diplozoon nipponicum Goto Kamegai, S., 1974 Studies on Diplozoon nipponicum Goto, 1891 Abstracts of ICOPA III, Munich, Germany pp a Diplozoon b Diplozoon 16 Diplozoon nipponicum Goto Diplozoon Diplozoon a Diplozoon b Diplozoon Diplozoon 21 Oncomiracidium Diplozoon nipponicum Goto Diplozoon 21 Oncomiracidium Diplozoon a Diplozoon b Diplozoon 27 Diplozoon nipponicum Oncomiracidium c Diplozoon Diporpa a Diplozoon 29 Oncomiracidium b Diplozoon a Diplozoon 31 Oncomiracidium b Diplozoon 31 Oncomiracidium

16 54 pair Kamegai, S. and Ichihara, A., Studies on Diplozoon nipponicum Goto, No. 32 Ecology & development. Abstracts of ICOPA V, Toronto: a Diplozoon 34 diporpa b Diplozoon 35 diporpae Diplozoon 36 D. nipponicum Diplozoon Diplozoon a Diplozoon 39 clamp b Diplozoon 40 Diporpae Cortisone acetate Diplozoon 41 Diplozoon Diplozoon 42 Diporpae Kamegai, S., Studies on Diplozoon nipponicum Goto, 1891 (43). The gathering phenomenon of diporpae and the effect of cortisone acetate on the union of diporpae. Abstracts of ICOPA VI, Brisbane a Diplozoon b Diplozoon 45 D. nipponicum Clamp a Diplozoon 46 D. nipponicum b Diplozoon 47 larva a Diplozoon 48 D. nipponicum oncomiracidium b Diplozoon 49 D. nipponicum Cortisone acetate a Diplozoon 50 D. nipponicum 1 39(1) b Diplozoon 51 oncomiracidium Kamegai, S., Studies on the genus Diplozoon No. 52. Abnormally increased egg laying of D. nipponicum before the death of experimental host fish Carassius carassius. Abstracts of ICOPA VII Paris: a Diplozoon 53 40(1): b Diplozoon a Diplozoon

17 b Diplozoon a Diplozoon b Diplozoon a Diplozoon b Diplozoon c Diplozoon 62 Diplozoon sp. 43(5): Diplozoon 63 Diplozoon sp. 43(5): Diplozoon 64 Diplozoon sp. 44 : Diplozoon 65 Diplozoon sp. oncomiracidium 44(6): Diplozoon 66 D. nipponicum Diplozoon 67 Diplozoon nipponicum Kamegai, S., Studies on the genus Diplozoon (68). One case report on single adult worm infection of D. nipponicum. Parasitology International, 46 (suppl.): 126.

18 56 A note on Eudiplozoon nipponicum and Diplozoon sp. (Monogenea: Diplozoidae) parasitic on cyprinids in Japan, with a list of the works of Dr. Satoru Kamegai on diplozoids Kazuya Nagasawa Graduate School of Biosphere Science, Hiroshima University Kagamiyama, Higashi-Hiroshima, Hiroshima , Japan Abstract The diplozoid monogenean Eudiplozoon nipponicum (Goto, 1891) is a gill parasite of crucian carps (Carassius spp.) and common carp (Cyprinus carpio) in Japan. This note compiles information on the host species, geographical distribution, life history, maturation and spawning, occurrence related to fish size, pathogenicity, and control of the species based on the literature published in Japan for 126 years from 1891 and Since the 1970 s, an unidentified diplozoid has been reported as Diplozoon sp. from various cyprinids in Japan. This diplozoid is very likely to be Paradilpozoon skrjabini Akhmerov, 1974, another species of diplozoid reported from Japan in The titles of scientific papers, a book, and oral presentations at academic meetings by Dr. Satoru Kamegai on the biology of diplozoids are listed in the appendix. Key words: cyprinids, Diplozoon sp., Eudiplozoon nipponicum, fish parasite, Monogenea, Paradilpozoon skrjabini, Satoru Kamegai

Matsuura et al., NPO

Matsuura et al., NPO 2001 11 2004 12 1969 1996 2004 Matsuura et al., 2000 2000 2002 2005 NPO 9 26 1999 2004 16 53 1998 16 61 1999 2004 7 1 16 62 77 80% 2000 40m 1971 3 7m 1998 55 2000 13 37 1 19 10 2001 1998 1999 2004 38 1986