Bull. Natl. Mus. Nat. Sci., Ser. A, 43(4), pp. 129–143, November 22, 2017

Digeneans Parasitic in Freshwater Fishes (Osteichthyes) of Japan. XII. A List of the Papers of the Series, a Key to the Families in Japan, a Parasite-Host List, a Host-Parasite List, Addenda, and Errata

Takeshi SHIMAZU

10486–2 Hotaka-Ariake, Azumino, Nagano 399–8301, Japan E-mail: [email protected]

(Received 16 June 2017; accepted 27 September 2017)

Abstract As a final paper of a series that reviews adult digeneans () parasitic in fresh- water fishes (Osteichthyes) of Japan, this paper presents a list of the papers of the series, a key to the families in Japan, a parasite-host list, a host-parasite list, addenda, and errata. Key words: , freshwater fishes, Japan, review, key to families, parasite-host list, host-par- asite list, addenda, errata.

fishes (Osteichthyes) of Japan. III. Azygiidae and Introduction Bucephalidae. Bulletin of the National Museum of Nature and Science, Series A (Zoology), 40: 167–190. This is the twelfth (final) paper of a series that Shimazu, T. 2015a. Digeneans parasitic in freshwater reviews adult digeneans (Trematoda) parasitic in fishes (Osteichthyes) of Japan. IV. . Bulle- freshwater fishes (Osteichthyes) of Japan tin of the National Museum of Nature and Science, (Shimazu, 2013). This paper deals with a list of Series A (Zoology), 41: 77–103. the papers of the series, a key to the families in Shimazu, T. 2015b. Digeneans parasitic in freshwater Japan, a parasite-host list, a host-parasite list, fishes (Osteichthyes) of Japan. V. Didymozoidae and Isoparorchiidae. Bulletin of the National Museum of addenda, and errata. Nature and Science, Series A (Zoology), 41: 201–216. The Introduction, Materials, and Methods for Shimazu, T. 2016a. Digeneans parasitic in freshwater the series were described in the first paper fishes (Osteichthyes) of Japan. VI. Lissorchiidae. Bul- (Shimazu, 2013). The purpose of the series was letin of the National Museum of Nature and Science, given in the Introduction. While preparing the Series A (Zoology), 42: 1–22. Shimazu, T. 2016b. Digeneans parasitic in freshwater series, I wrote a brief review of the adult digene- fishes (Osteichthyes) of Japan. VII. Allocreadiidae: ans known then from Japanese freshwater fishes Allocreadium. Bulletin of the National Museum of (Shimazu, 2016e). Nature and Science, Series A (Zoology), 42: 55–79. Shimazu, T. 2016c. Digeneans parasitic in freshwater fishes (Osteichthyes) of Japan. VIII. Allocreadiidae, List of the Papers of the Series Crepidostomum. Bulletin of the National Museum of Shimazu, T. 2013. Digeneans parasitic in freshwater Nature and Science, Series A (Zoology), 42: 107–122. fishes (Osteichthyes) of Japan. I. Aporocotylidae, Bive- Shimazu, T. 2016d. Digeneans parasitic in freshwater siculidae and Haploporidae. Bulletin of the National fishes (Osteichthyes) of Japan. IX. Opecoelidae, Ope- Museum of Nature and Science, Series A (Zoology), coelinae. Bulletin of the National Museum of Nature 39: 167–184. and Science, Series A (Zoology), 42: 163–180. Shimazu, T. 2014a. Digeneans parasitic in freshwater Shimazu, T. 2017a. Digeneans parasitic in freshwater fishes (Osteichthyes) of Japan. II. Gorgoderidae and fishes (Osteichthyes) of Japan. X. Opecoelidae, Plagio- Orientocreadiidae. Bulletin of the National Museum of porinae. Bulletin of the National Museum of Nature Nature and Science, Series A (Zoology), 40: 53–78. and Science, Series A (Zoology), 43: 1–28. Shimazu, T. 2014b. Digeneans parasitic in freshwater Shimazu, T. 2017b. Digeneans parasitic in freshwater 130 Takeshi Shimazu

fishes (Osteichthyes) of Japan. XI. Cryptogonimidae fishes (Osteichthyes) of Japan. XII. A key to the fami- and Heterophyidae. Bulletin of the National Museum lies in Japan, a parasite-host list, a host-parasite list, of Nature and Science, Series A (Zoology), 43: 101– addenda, and errata. Bulletin of the National Museum 118. of Nature and Science, Series A (Zoology), 43: 129– Shimazu, T. 2017c. Digeneans parasitic in freshwater 143.

Key to the Families in Japan The known adult digeneans, except Opecoelidae gen. sp. and Allocreadium sp., incertae sedis, of freshwater fishes in Japan are classified into 15 families in 11 superfamilies (see also a parasite-host list below). The following key is designed to facilitate identification of them to family level but not to show phylogenetic relationships. The serial numbers of the papers that deal with the relevant families are shown in the brackets ([ ]) after the respective family names.

Parasitic in circulatory system (heart and blood vessels); oral and ventral suckers absent ...... Family Aporocotylidae [I] Parasitic in lymphatic system (lymphatic vessels of connective tissue); oral and ventral suckers present ...... Family Didymozoidae [V] Parasitic in urinary system (urinary bladder and ureters); oral and ventral sucker present ...... Family Gorgoderidae [II] Parasitic in digestive system (alimentary canal and its associated organs) ...... 1 1.1. Parasitic in air bladder ...... Family Isoparorchiidae [V] 1.2. Parasitic in alimentary canal and its associated organs other than air bladder ...... 2 2.1. Oral and ventral suckers present ...... 3 2.2. Oral and ventral suckers absent ...... 4 3.1. Tegument spinose ...... 5 3.2. Tegument smooth ...... 6 4.1. Mouth anteroterminal ...... Family Bivesiculidae [I] 4.2. Mouth about midventral ...... Family Bucephalidae [III] 5.1. Genital pore lateral to marginal, at about level of ventral sucker ...... Family Lissorchiidae [VI] 5.2. Genital pore median, in forebody ...... 7 6.1. Genital pore submedian, in forebody ...... Family Opecoelidae [IX, X] 6.2. Genital pore median, in forebody ...... 8 7.1. Testis single, in hindbody ...... Family Haploporidae [I] 7.2. Testes two, in hindbody ...... 9 8.1. Cirrus pouch present ...... Family Allocreadiidae [VII, VIII] 8.2. Cirrus pouch absent ...... 10 9.1. Testes submedian, oblique; ventrogenital sac absent ...... Family Orientocreadiidae [II] 9.2. Testes submedian or lateral, almost symmetrical; ventrogenital sac present ...... 11 10.1. Prostatic sac present; sinus sac absent ...... Family Azygiidae [III] 10.2. Prostatic sac absent; sinus sac present ...... Family Derogenidae [IV] 11.1. Ovary multilobulate; gonotyls present ...... Family Cryptogonimidae [XI] 11.2. Ovary entire; gonotyl absent ...... Family Heterophyidae [XI] Digeneans Parasitic in Freshwater Fishes of Japan 131

Parasite-Host List Allocreadium sp. of Shimazu (1988) [VII] Gymnogobius opperiens A total of 52 identified and 12 unidentified Allocreadium sp. of Shimazu (2005) [VII] digenean species in 26 genera in 15 families in Tribolodon hakonensis 11 superfamilies, and Opecoelidae gen. sp. and Allocreadium sp. of Shimazu (2008) [VII] Allocreadium sp., incertae sedis, are known at present from freshwater fishes in Japan (see also Allocreadium sp. of Shimazu, Urabe, and Grygier a host-parasite list below). The superfamily, fam- (2011) [VII] ily, generic, and species names of the parasites Tanakia lanceolata and the species names of the hosts are arranged Allocreadium tamoroko Shimazu and Urabe, in alphabetical order. The serial numbers of the 2013 [VII] papers of the present review are given in the elongatus (type brackets ([ ]) after the names of the parasites. host) The type hosts are also indicated in the parenthe- Allocreadium tosai Shimazu, 1988 [VII] ses (( )) after the names of the hosts. Oncorhynchus mykiss percnurus Superfamily Allocreadioidea Looss, 1902 [VII– Salvelinus leucomaenis leucomaenis IX] Tribolodon hakonensis (type host) Family Allocreadiidae Looss, 1902 [VII, VIII] Tribolodon sachalinensis Allocreadium aburahaya Shimazu, 2003 [VII] Allocreadium tribolodontis Shimazu and Hashi- Phoxinus steindachneri (type host) moto, 1999 [VII] Allocreadium brevivitellatum Shimazu, 1992 Tribolodon hakonensis [VII] Tribolodon sachalinensis (type host) Rhynchocypris percnurus (type host) Crepidostomum chaenogobii Yamaguti and Allocreadium gotoi (Hasegawa and Ozaki, 1926) Matumura, 1942 [VIII] Shimazu, 1988 [VII] Cottus amblystomopsis Gnathopogon elongatus elongatus Cottus hangiongensis Gymnogobius opperiens Cottus nozawae Misgurnus anguillicaudatus (type host) Gymnogobius opperiens Allocreadium hasu Ozaki, 1926 [VII] “Chaenogobius annularis urotaenia Hilgen- Gnathopogon elongatus elongatus dorf ” (type host) Nipponocypris temminckii Crepidostomum farionis (Müller, 1780) Lühe, Opsariichthys uncirostris uncirostris (type 1909 [VIII] host) Oncorhynchus masou masou Zacco platypus Oncorhynchus mykiss Allocreadium japonicum Ozaki, 1926 [VII] Salvelinus fontinalis Gasterosteus aculeatus leiurus Salvelinus leucomaenis leucomaenis Nipponocypris temminckii Salvelinus malma malma Phoxinus oxycephalus Crepidostomum metoecus (Braun, 1900) Braun, Tribolodon brandtii 1900 [VIII] Zacco platypus (type host) Barbatula toni Allocreadium shinanoense Shimazu, 2003 [VII] Cottus nozawae Phoxinus steindachneri (type host) Gasterosteus aculeatus Allocreadium sp. of Kataoka and Momma (1934), Gymnogobius castaneus incertae sedis [VII] Gymnogobius urotaenia Plecoglossus altivelis altivelis Oncorchynchus keta 132 Takeshi Shimazu

Oncorchynchus masou masou 1990 [X] Oncorchynchus mykiss Plecoglossus altivelis altivelis (type host) Parahucho perryi Neoplagioporus elongatus (Goto and Ozaki, Pungitius pungitius 1930) Shimazu, 1990 [X] Pungitius tymensis Biwia zezera Salmo trutta Carassius auratus subsp. 1 Salvelinus fontinalis kawamebari Salvelinus leucomaenis leucomaenis Gnathopogon elongatus elongatus Salvelinus malma malma Gymnogobius isaza Family Opecoelidae Ozaki, 1925 [IX, X] Gymnogobius urotaenia Coitocaecum plagiorchis Ozaki, 1926 [IX] Hemibarbus barbus Anguilla japonica Hemibarbus labeo Coreoperca kawamebari Odontobutis obscura Cottus reinii Pseudogobio esocinus esocinus “Gobius similis Gill” Pseudorasbora parva “[Gori]” Pungtungia herzi Gymnogobius isaza Rhinogobius flumineus Gymnogobius urotaenia “Rhinogobius sp.” Misgurnus anguillicaudatus Rhinogobius sp. BW Odontobutis obscura (type host) Rhinogobius sp. OR Rhinogobius flumineus “S. variegatus” “Rhinogobius sp.” “Sarcocheilichthys variegatus (Temm. et Rhinogobius sp. BW Schl.)” “Small GORO” “Sarcocheilichthys variegatus (Temminck et Tachysurus aurantiacus Schlegel)” (type host) Tachysurus nudiceps Sarcocheilichthys variegatus microoculus Tridentiger brevispinis Sarcocheilichthys variegatus variegatus Dimerosaccus oncorhynchi (Eguchi, 1931) Squalidus chankaensis biwae Shimazu, 1980 [IX] Squalidus japonicus japonicus Cottus nozawae Tribolodon hakonensis Cottus pollux Tridentiger brevispinis Liobagrus reinii Neoplagioporus kajika Urabe and Higa, 2006 [X] Odontobutis obscura Cottus pollux (type host) Oncorhynchus masou ishikawae (type host) Nipponocypris temminckii Oncorhynchus masou masou Pseudogobio esocinus esocinus Rhinogobius brunneus Neoplagioporus sp. of Shimazu, Urabe, and Rhinogobius flumineus Grygier (2011) [X] Rhinogobius fluviatilis Odontobutis obscura Rhinogobius nagoyae Neoplagioporus zacconis (Yamaguti, 1934) “Rhinogobius sp.” Shimazu, 1990 [X] Rhinogobius sp. CO Liobagrus reinii Rhinogobius sp. OR Nipponocypris temminckii (type host) Salvelinus leucomaenis leucomaenis Oncorhynchus masou masou Salvelinus leucomaenis pluvius Opsariichthys uncirostris uncirostris Tridentiger brevispinis Pungtungia herzi Neoplagioporus ayu (Takahashi, 1928) Shimazu, Zacco platypus Digeneans Parasitic in Freshwater Fishes of Japan 133

Opecoelidae gen. sp. of Shimazu (1990) [X] Tridentiger brevispinis Oncorhynchus keta Rhinogobius sp. OR (type host) Opecoelus ukigori Shimazu, 1988 [IX] Gymnogobius opperiens (type host) Superfamily Bivesiculoidea Yamaguti, 1934 [I] Gymnogobius urotaenia Family Bivesiculidae Yamaguti, 1934 [I] Urorchis acheilognathi Yamaguti, 1934 [X] Bivesicula sp. of Shimazu (1994) [I] Acheilognathus cyanostigma Monopterus albus Acheilognathus rhombeus Acheilognathus tabira tabira Superfamily Bucephaloidea Poche, 1907 [III] Gnathopogon caerulescens Family Bucephalidae Poche, 1907 [III] Oncorhynchus masou subsp. Parabucephalopsis parasiluri Wang, 1985 [III] Phoxinus steindachneri Silurus biwaensis Pseudorasbora parva Silurus lithophilus Sarcocheilichthys variegatus microoculus Prosorhynchoides ozakii (Nagaty, 1937) Margo- Tanakia lanceolata (type host) lis and Author, 1979 [III] Tanakia limbata Silurus asotus Urorchis goro Ozaki, 1927 [X] Silurus biwaensis Barbatula toni Silurus lithophilus Cottus pollux Cottus reinii Superfamily Gorgoderoidea Looss, 1899 [II] Gnathopogon elongatus elongatus Family Gorgoderidae Looss, 1899 [II] Gymnogobius urotaenia Phyllodistomum biringo Shimazu, 2005 [II] Lefua echigonia Gymnogobius breunigii (type host) Rhinogobius flumineus Silurus asotus Rhinogobius kurodai Phyllodistomum carassii Long and Wai, 1958 [II] Rhinogobius sp. OR Carassius auratus grandoculis Tridentiger brevispinis (type host) Phyllodistomum mogurndae Yamaguti, 1934 [II] “Tridentiger obscurus (Temminck & Schle- Gymnogobius urotaenia gel)” Odontobutis obscura (type host) Urorchis imba Ishii, 1935 [X] Rhinogobius sp. OR Pseudorasbora parva (type host) Tachysurus nudiceps Urorchis sp. of Shimazu (1990) [X] Phyllodistomum parasiluri Yamaguti, 1934 [II] Gnathopogon caerulescens Silurus asotus (type host) Odontobutis obscura Silurus lithophilus Tanakia limbata Tachysurus nudiceps Pseudophyllodistomum macrobrachicola (Yama- Superfamily Azygioidea Lühe, 1909 [III] guti, 1934) Cribb, 1987 [II] Family Azygiidae Lühe, 1909 [III] Anguilla japonica Azygia gotoi (Ariake, 1922) Shimazu, 1979 [III] Cottus reinii Anguilla japonica (type host) Gymnogobius urotaenia Azygia perryii Fujita, 1918 [III] “[Kajika]” Parahucho perryi (type host) Odontobutis obscura (type host) Salvelinus leucomaenis leucomaenis Silurus asotus Azygia rhinogobii Shimazu, 2007 [III] Tachysurus nudiceps Gymnogobius urotaenia Tribolodon hakonensis Superfamily Haploporoidea Nicoll, 1914 [I] 134 Takeshi Shimazu

Family Haploporidae Nicoll, 1914 [I] Genarchopsis goppo Ozaki, 1925 [IV] Carassotrema koreanum Park, 1938 [I] Coreoperca kawamebari Carassius auratus langsdorfii “[Gori]” Carassius carassius Gymnogobius petschiliensis* Cyprinus carpio Odontobutis obscura (type host) Tribolodon hakonensis Rhinogobius flumineus* Rhinogobius giurinus* Superfamily Hemiuroidea Looss, 1899 [IV, V] Rhinogobius nagoyae* Family Derogenidae Nicoll, 1910 [IV] Silurus asotus Allogenarchopsis problematica (Faust, 1924) Tridentiger brevispinis* Urabe and Shimazu, 2013 [IV] Genarchopsis sp. 1 of Shimazu (1995) [IV] Acheilognathus rhombeus Tridentiger brevispinis Rhodeus ocellatus ocellatus Genarchopsis sp. 2 of Shimazu (1995) [IV] Tanakia lanceolata (type host) Acanthogobius flavimanus Tanakia limbata Family Didymozoidae Monticelli, 1888 [V] Genarchopsis anguillae Yamaguti, 1938 [IV] Paraphilopinna sp. of Shimazu (2006) [V] Anguilla japonica (type host) Misgurnus anguillicaudatus Gymnogobius urotaenia Philopinna higai Yamaguti, 1936 [V] Genarchopsis chubuensis Shimazu, 2015 [IV] Sarcocheilichthys biwaensis Anguilla japonica “Sarcocheilichthys variegatus (Temm. et Cottus pollux Schleg.)” (type host) “Gobius similis Gill” Sarcocheilichthys variegatus microoculus Gymnogonius castaneus Sarcocheilichthys variegatus variegatus Gymnogobius urotaenia (type host) Philopinna kawamutsu Shimazu, Urabe, and salmoides Grygier, 2011 [V] Odontobutis obscura Nipponocypris temminckii (type host) Rhinogobius flumineus Family Isoparorchiidae Travassos, 1922 [V] Rhinogobius kurodai Isoparorchis eurytremus (Kobayashi, 1915) Rhinogobius sp. OM Travassos, 1922 [V] Rhinogobius sp. OR Silurus asotus (type host) Silurus asotus Silurus biwaensis Tridentiger brevispinis “Pseudobagrus aurantiacus” (type host) Genarchopsis fellicola Shimazu, 1995 [IV] Gymnogobius urotaenia (type host) Superfamily Monorchioidea Odhner, 1911 [VI] Rhinogobius kurodai Family Lissorchiidae Magath, 1917 [VI] Rhinogobius sp. OR Asymphylodora innominata (Faust, 1924) Silurus asotus Shimazu, Urabe, and Grygier, 2011 [VI] Tridentiger brevispinis “[Bote]” Genarchopsis gigi Yamaguti, 1939 [IV] “[Gori]” Anguilla japonica Gymnogobius isaza Cottus reinii Hemibarbus barbus Gymnogobius isaza Odontobutis obscura (type host) Opsariichthys uncirostris uncirostris Opsariichthys uncirostris uncirostris Rhinogobius sp. BW Phoxinus steindachneri Tachysurus nudiceps (type host) Tribolodon hakonensis Tridentiger brevispinis “[Ukikamatsuka?]” Digeneans Parasitic in Freshwater Fishes of Japan 135

Asymphylodora japonica Yamaguti, 1938 [VI] Host-Parasite List Cyprinus carpio (type host) Asymphylodora sp. of Shimazu, Urabe, and The family, generic, and species names of the Grygier (2011) [VI] host freshwater fishes in Japan (see also the para- Tridentiger brevispinis site-host list above) are arranged in alphabetical Asymphylotrema monostyloides (Ito, 1960) order. Some host fishes are unidentified to spe- Shimazu, 2016 [VI] cies or vague about species. Each species name Cobitis biwae (type host) is accompanied with its authorship and date and Palaeorchis diplorchis (Yamaguti, 1936) Szidat, its Japanese common name in the brackets ([ ]). 1943 [VI] The serial numbers of the papers of the present Anguilla japonica review are given in the brackets ([ ]) after the Biwia zezera species names of the parasites. Hemibarbus barbus Pseudogobio esocinus esocinus (type host) Family Amblycipitidae Liobagrus reinii Hilgendorf, 1878 [Akaza] Superfamily Opisthorchioidea Looss, 1899 [XI] Dimerosaccus oncorhynchi [IX] Family Cryptogonimidae Ward, 1917 [XI] Neoplagioporus zacconis [X] Exorchis oviformis Kobayashi, 1915 [XI] Family Anguillidae Silurus asotus (type host) Anguilla japonica Temminck and Schlegel, 1846 Family Heterophyidae Leiper, 1909 [XI] [Nihon-unagi] Pseudexorchis major (Hasegawa, 1935) Yama- Azygia gotoi [III] guti, 1938 [XI] Coitocaecum plagiorchis [IX] Silurus asotus (type host) Genarchopsis anguillae [IV] Silurus biwaensis Genarchopsis chubuensis [IV] Genarchopsis gigi [IV] Superfamily Plagiorchioidea Lühe, 1901 [II] Palaeorchis diplorchis [VI] Family Orientocreadiidae Yamaguti, 1958 [II] Pseudophyllodistomum macrobrachicola [II] Orientocreadium chaenogobii Shimazu, 1990 [II] Family Bagridae Gymnogobius castaneus (type host) “Pseudobagrus aurantiacus”† [Gigi] Gymnogobius urotaenia Isoparorchis eurytremus [V] Orientocreadium pseudobagri Yamaguti, 1934 [II] Tachysurus aurantiacus (Temminck and Schle- Silurus lithophilus gel, 1846) [Ariake-gibachi] Tachysurus nudiceps (type host) Coitocaecum plagiorchis [IX] Tachysurus nudiceps (Sauvage, 1883) [Gigi] Superfamily Schistosomatoidea Stiles and Has- Coitocaecum plagiorchis [IX] sall, 1898 [I] Genarchopsis gigi [IV] Family Aporocotylidae Odhner, 1912 [I] Orientocreadium pseudobagri [II] Sanguinicola hasegawai Shimazu, 2013 [I] Phyllodistomum mogurndae [II] Barbatula toni (type host) Phyllodistomum parasiluri [II] Sanguinicola ugui Shimazu, 2007 [I] Pseudophyllodistomum macrobrachicola [II] Tribolodon hakonensis (type host) Family Centrarchidae Sanguinicola sp. of Shimazu (1999) [I] Micropterus salmoides (Lacepède, 1802) [Ōkuchi- Acheilognathus tabira tabira basu] * The parasite found in these hosts from Shi- Genarchopsis chubuensis [IV] koku Region has not yet been identified defini- Family Cobitidae tively (see also a host-parasite list below). Cobitis biwae Jordan and Snyder, 1901 [Shima- 136 Takeshi Shimazu

dojō] Carassius auratus grandoculis Temminck and Asymphylotrema monostyloides [VI] Schlegel, 1846 [Nigoro-buna] Misgurnus anguillicaudatus (Cantor, 1842) [Dojō] Phyllodistomum carassii [II] Allocreadium gotoi [VII] Carassius auratus langsdorfii Temminck and Coitocaecum plagiorchis [IX] Schlegel, 1846 [Gin-buna] Paraphilopinna sp. of Shimazu (2006) [V] Carassotrema koreanum [I] Family Cottidae Carassius auratus subsp. 1 [Naga-buna] Cottus amblystomopsis Schmidt, 1904 [Ezo- Neoplagioporus elongatus [X] hana-kajika] Carassius carassius (Linnaeus, 1758)† [Funa] Crepidostomum chaenogobii [VIII] Carassotrema koreanum [I] Cottus hangiongensis Mori, 1930 [Kankyō- Cyprinus carpio Linnaeus, 1758 [Koi] kajika] Asymphylodora japonica [VI] Crepidostomum chaenogobii [VIII] Carassotrema koreanum [I] Cottus nozawae Snyder, 1911 [Hana-kajika] Gnathopogon caerulescens (Sauvage, 1883) Crepidostomum chaenogobii [VIII] [Hon-moroko] Crepidostomum metoecus [VIII] Urorchis acheilognathi [X] Dimerosaccus oncorhynchi [IX] Urorchis sp. of Shimazu (1990) [X] Cottus pollux Günther, 1873 [Kajika] Gnathopogon elongatus elongatus (Temminck Dimerosaccus oncorhynchi [IX] and Schlegel, 1846) [Ta-moroko] Genarchopsis chubuensis [IV] Allocreadium gotoi [VII] Neoplagioporus kajika [X] Allocreadium hasu [VII] Urorchis goro [X] Allocreadium tamoroko [VII] Cottus reinii Hilgendorf, 1879 [Utsusemi-kajika] Neoplagioporus elongatus [X] Coitocaecum plagiorchis [IX] Urorchis goro [X] Genarchopsis gigi [IV] Hemibarbus barbus (Temminck and Schlegel, Pseudophyllodistomum macrobrachicola [II] 1846) [Nigoi] Urorchis goro [X] Asymphylodora innominata [VI] “[Kajika]”† [Utsusemi-kajika (?)] Neoplagioporus elongatus [X] Pseudophyllodistomum macrobrachicola [II] Palaeorchis diplorchis [VI] Family Cyprinidae Hemibarbus labeo (Pallas, 1776) [Kōrai-nigoi] Acheilognathus cyanostigma Jordan and Fowler, Neoplagioporus elongatus [X] 1903 [Ichimonji-tanago] Nipponocypris temminckii (Temminck and Urorchis acheilognathi [X] Schlegel, 1846) [Kawa-mutsu] Acheilognathus rhombeus (Temminck and Schle- Allocreadium hasu [VII] gel, 1846) [Kanehira] Allocreadium japonicum [VII] Allogenarchopsis problematica [IV] Allocreadium sp. of Shimazu (2008) [VII] Urorchis acheilognathi [X] Neoplagioporus kajika [X] Acheilognathus tabira tabira Jordan and Thomp- Neoplagioporus zacconis [X] son, 1914 [Shirohire-tabira] Philopinna kawamutsu [V] Sanguinicola sp. of Shimazu (1999) [I] Opsariichthys uncirostris uncirostris (Temminck Urorchis acheilognathi [X] and Schlegel, 1846) [Hasu] Biwia zezera (Ishikawa, 1895) [Zezera] Allocreadium hasu [VII] Neoplagioporus elongatus [X] Asymphylodora innominata [VI] Palaeorchis diplorchis [VI] Genarchopsis gigi [IV] “[Bote]”† [Tanago-rui] Neoplagioporus zacconis [X] Asymphylodora innominata [VI] Phoxinus steindachneri Sauvage, 1883 [Abura- Digeneans Parasitic in Freshwater Fishes of Japan 137

haya] Urorchis acheilognathi [X] Allocreadium aburahaya [VII] Sarcocheilichthys variegatus variegatus (Tem- Allocreadium shinanoense [VII] minck and Schlegel, 1846) [Kawa-higai] Asymphylodora innominata [VI] Neoplagioporus elongatus [X] Urorchis acheilognathi [X] Philopinna higai [V] Phoxinus oxycephalus (Sauvage and Dabry de Squalidus chankaensis biwae (Jordan and Sny- Thiersant, 1874) [Taka-haya] der, 1900) [Sugo-moroko] Allocreadium japonicum [VII] Neoplagioporus elongatus [X] Pseudogobio esocinus esocinus (Temminck and Squalidus japonicus japonicus Sauvage, 1883 Schlegel, 1846) [Kamatsuka] [Deme-moroko] Neoplagioporus elongatus [X] Neoplagioporus elongatus [X] Neoplagioporus kajika [X] Tanakia lanceolata (Temminck and Schlegel, Palaeorchis diplorchis [VI] 1846) [Yari-tanago] Pseudorasbora parva (Temminck and Schlegel, Allocreadium sp. of Shimazu, Urabe, and 1846) [Motsugo] Grygier (2011) [VII] Neoplagioporus elongatus [X] Allogenarchopsis problematica [IV] Urorchis acheilognathi [X] Urorchis acheilognathi [X] Urorchis imba Ishii, 1935 [X] Tanakia limbata (Temminck and Schlegel, 1846) Pungtungia herzi Herzenstein, 1892 [Mugitsuku] [Aburabote] Neoplagioporus elongatus [X] Allogenarchopsis problematica [IV] Neoplagioporus zacconis [X] Urorchis acheilognathi [X] Rhodeus ocellatus ocellatus (Kner, 1866) Urorchis sp. of Shimazu (1990) [X] [Tairiku-bara-tanago] Tribolodon brandtii (Dybowski, 1872) [Maruta] Allogenarchopsis problematica [IV] Allocreadium japonicum [VII] Rhynchocypris percnurus (Pallas, 1814) [Yachi- Tribolodon hakonensis (Günther, 1877) [Ugui] ugui] Allocreadium tosai [VII] Allocreadium brevivitellatum [VII] Allocreadium tribolodontis [VII] Allocreadium tosai [VII] Allocreadium sp. of Shimazu (2005) [VII] Sarcocheilichthys biwaensis Hosoya, 1982 Asymphylodora innominata [VI] [Abura-higai] Azygia rhinogobii [III] Philopinna higai [V] Carassotrema koreanum [I] “S. variegatus”† [Higai] Neoplagioporus elongatus [X] Neoplagioporus elongatus [X] Sanguinicola ugui [I] “Sarcocheilichthys variegatus (Temm. et Schl.)”† Tribolodon sachalinensis (Nikolskii, 1889) [Ezo- [Higai] ugui] Neoplagioporus elongatus [X] Allocreadium tosai [VII] “Sarcocheilichthys variegatus (Temm. et Sch- Allocreadium tribolodontis [VII] leg.)”† [Higai] “[Ukikamatsuka?]”† [Zunaga-nigoi (?)]) Philopinna higai [V] Asymphylodora innominata [VI] “Sarcocheilichthys variegatus (Temminck et Zacco platypus (Temminck and Schlegel, 1846) Schlegel)”† [Higai] [Oikawa] Neoplagioporus elongatus [X] Allocreadium hasu [VII] Sarcocheilichthys variegatus microoculus Mori, Allocreadium japonicum [VII] 1927 [Biwa-higai] Neoplagioporus zacconis [X] Neoplagioporus elongatus [X] Family Gasterosteidae Philopinna higai [V] Gasterosteus aculeatus leiurus Cuvier in Cuvier 138 Takeshi Shimazu

and Valenciennes, 1829 [Hariyo] Crepidostomum metoecus [VIII] Allocreadium japonicum [VII] Genarchopsis anguillae [IV] Gasterosteus aculeatus Linnaeus, 1758 [Itoyo] Genarchopsis chubuensis [IV] Crepidostomum metoecus [VIII] Genarchopsis fellicola [IV] Pungitius pungitius (Linnaeus, 1758) [Ibara- Neoplagioporus elongatus [X] tomiyo] Opecoelus ukigori [IX] Crepidostomum metoecus [VIII] Orientocreadium chaenogobii [II] Pungitius tymensis (Nikolskii, 1889) [Ezo- Phyllodistomum mogurndae [II] tomiyo] Pseudophyllodistomum macrobrachicola [II] Crepidostomum metoecus [VIII] Urorchis goro [X] Family Gobiidae Rhinogobius brunneus (Temminck and Schlegel, Acanthogobius flavimanus (Temminck and 1845) [Kuro-yoshinobori] Schlegel, 1845) [Ma-haze] Dimerosaccus oncorhynchi [IX] Genarchopsis sp. 2 of Shimazu (1995) [IV] Rhinogobius flumineus (Mizuno, 1960) [Kawa- “Chaenogobius annularis urotaenia Hilgendorf”† yoshinobori] [Ukigori] Coitocaecum plagiorchis [IX] Crepidostomum chaenogobii [VIII] Dimerosaccus oncorhynchi [IX] “Gobius similis Gill”† [Yoshinobori] Neoplagioporus elongatus [X] Coitocaecum plagiorchis [IX] Genarchopsis chubuensis [IV] Genarchopsis chubuensis [IV] Genarchopsis goppo§ [IV] Gymnogobius breunigii (Steindachner, 1879) Urorchis goro [X] [Biringo] Rhinogobius fluviatilis Tanaka, 1925 Phyllodistomum biringo [II] [Ō-yoshinobori] Gymnogobius castaneus (O’Shaughnessy, 1875) Dimerosaccus oncorhynchi [IX] [Juzukake-haze] Rhinogobius giurinus (Rutter, 1897) [Gokuraku- Crepidostomum metoecus [VIII] haze] Genarchopsis chubuensis [IV] Genarchopsis goppo§ [IV] Orientocreadium chaenogobii [II] Rhinogobius kurodai (Tanaka, 1908) [Kuroda- Gymnogobius isaza (Tanaka, 1916) [Isaza] haze] Asymphylodora innominata [VI] Genarchopsis chubuensis [IV] Coitocaecum plagiorchis [IX] Genarchopsis fellicola [IV] Genarchopsis gigi [IV] Urorchis goro [X] Neoplagioporus elongatus [X] Rhinogobius nagoyae Jordan and Seale, 1906 Gymnogobius opperiens Stevenson, 2002 [Shima-yoshinobori] [Shima-ukigori] Genarchopsis goppo§ [IV] Allocreadium gotoi [VII] Dimerosaccus oncorhynchi [IX] Allocreadium sp. of Shimazu (1988) [VII] “Rhinogobius sp.”† [Yoshinobori] Crepidostomum chaenogobii [VIII] Coitocaecum plagiorchis [IX] Opecoelus ukigori [IX] Dimerosaccus oncorhynchi [IX] Gymnogobius petschiliensis (Rendahl, 1924) Neoplagioporus elongatus [X] [Sumi-ukigori] Rhinogobius sp. BW* [Biwa-yoshinobori] Genarchopsis goppo§ [IV] Coitocaecum plagiorchis [IX] Gymnogobius urotaenia (Hilgendorf, 1879) Genarchopsis gigi [IV] [Ukigori] Neoplagioporus elongatus [X] Azygia rhinogobii [III] Rhinogobius sp. CO* [Ruri-yoshinobori] Coitocaecum plagiorchis [IX] Dimerosaccus oncorhynchi [IX] Digeneans Parasitic in Freshwater Fishes of Japan 139

Rhinogobius sp. OM* [Ōmi-yoshinobori] Dimerosaccus oncorhynchi [X] Genarchopsis chubuensis [IV] Genarchopsis chubuensis [IV] Rhinogobius sp. OR* [Tō-yoshinobori] Genarchopsis goppo [IV] Azygia rhinogobii [III] Neoplagioporus elongatus [X] Dimerosaccus oncorhynchi [IX] Neoplagioporus sp. of Shimazu, Urabe, and Genarchopsis chubuensis [IV] Grygier (2011) [X] Genarchopsis fellicola [IV] Phyllodistomum mogurndae [II] Neoplagioporus elongatus [X] Pseudophyllodistomum macrobrachicola Phyllodistomum mogurndae [II] [II] Urorchis goro [X] Urorchis sp. of Shimazu (1990) [X] “Small GORO”† [Isaza (?)] Family Percichthyidae Coitocaecum plagiorchis [IX] Coreoperca kawamebari (Temminck and Schle- Tridentiger brevispinis Katsuyama, Arai, and gel, 1843) [Oyanirami] Nakamura, 1972 [Numa-chichibu] Coitocaecum plagiorchis [IX] Asymphylodora sp. of Shimazu, Urabe, and Genarchopsis goppo [IV] Grygier (2011) [VI] Neoplagioporus elongatus [X] Azygia rhinogobii [III] Family Plecoglossidae Coitocaecum plagiorchis [IX] Plecoglossus altivelis altivelis (Temminck and Dimerosaccus oncorhynchi [IX] Schlegel, 1846) [Ayu] Genarchopsis chubuensis [IV] Allocreadium sp. of Kataoka and Momma Genarchopsis fellicola [IV] (1934), incertae sedis [VII] Genarchopsis gigi [IV] Neoplagioporus ayu [X] Genarchopsis goppo§ [IV] Family Salmonidae Genarchopsis sp. 1 of Shimazu (1995) [IV] Oncorhynchus keta (Walbaum, 1792) [Sake] Neoplagioporus elongatus [X] Crepidostomum metoecus [VIII] Urorchis goro [X] Opecoelidae gen. sp. of Shimazu (1990) [X] “Tridentiger obscurus (Temminck & Schlegel)”† Oncorhynchus masou ishikawae Jordan and [Chichibu] McGregor in Jordan and Hubbs, 1925 Urorchis goro [X] [Amago] Family Nemacheilidae Dimerosaccus oncorhynchi [IX] Barbatula toni (Dybowski, 1869) [Fuku-dojō] Oncorhynchus masou masou (Brevoort, 1856) Crepidostomum metoecus [VIII] [Yamame] Sanguinicola hasegawai [I] Crepidostomum farionis [VIII] Urorchis goro [X] Crepidostomum metoecus [VIII] Lefua echigonia Jordan and Richardson, 1907 Dimerosaccus oncorhynchi [IX] [Hotoke-dojō] Neoplagioporus zacconis [X] Urorchis goro [X] Oncorhynchus mykiss (Walbaum, 1792) [Niji- Family Odontobutidae masu] “[Gori]” [Donko] Allocreadium tosai [VII] Asymphylodora innominata [VI] Crepidostomum farionis [VIII] Coitocaecum plagiorchis [IX] Crepidostomum metoecus [VIII] Genarchopsis goppo [IV] Oncorhynchus masou subsp. [Kizaki-masu] Odontobutis obscura (Temminck and Schlegel, Urorchis acheilognathi [X] 1845) [Donko] Parahucho perryi (Brevoort, 1856) [Itou] Asymphylodora innominata [VI] Azygia perryii [III] Coitocaecum plagiorchis [IX] Crepidostomum metoecus [VIII] 140 Takeshi Shimazu

Salmo trutta Linnaeus, 1758 [Burauntorauto] Bivesicula sp. of Shimazu (1994) [I] Crepidostomum metoecus [VIII] * BW, CO, OM, and OR indicate four types, Salvelinus fontinalis (Mitchill, 1814) [Kawa- or unidentified species, of Rhinogobius. masu] † The exact species names of these fishes are Crepidostomum farionis [VIII] vague at present. Crepidostomum metoecus [VIII] § This parasite from Shikoku Region has not Salvelinus leucomaenis leucomaenis (Pallas, yet been identified definitively (see also the para- 1814) [Ezo-iwana] site-host list above). Allocreadium tosai [VII] Azygia perryii [III] Addenda Crepidostomum farionis [VIII] Crepidostomum metoecus [VIII] II (Shimazu, 2014a). Phyllodistomum sp. Dimerosaccus oncorhynchi [IX] Urabe et al. (2015) found cercariae of Phyllo- Salvelinus leucomaenis pluvius (Hilgendorf, distomum sp. in Nodularia douglasiae (Bivalvia, 1876) [Nikkō-iwana] Unionidae) and immature worms of this species Dimerosaccus oncorhynchi [IX] in Cyprinus carpio from the Yodo River, Osaka Salvelinus malma malma (Walbaum, 1792) Prefecture. They made a morphological and [Oshorokoma] molecular study of the species, but they failed to Crepidostomum farionis [VIII] identify it definitively to species level. Crepidostomum metoecus [VIII] Family Siluridae IV (Shimazu, 2015a). Genarchopsis: a new cys- Silurus asotus Linnaeus, 1758 [Namazu] tophorous cercaria Exorchis oviformis [XI] A new cystophorous cercaria was found in Genarchopsis chubuensis [IV] collected by Akifumi Genarchopsis fellicola [IV] Ohtaka in an irrigation canal in Nozaki, Genarchopsis goppo [IV] Ishikawa, Hirosaki City, Aomori Prefecture, on Isoparorchis eurytremus [V] 16 July 2015. This cercaria is morphologically Phyllodistomum biringo [II] similar to Cercaria longicerca Ito, 1953 but Phyllodistomum parasiluri [II] molecularly different from it (Misako Urabe, Prosorhynchoides ozakii [III] 2015, personal communication). Pseudexorchis major [XI] Pseudophyllodistomum macrobrachicola [II] VII (Shimazu, 2016b). Allocreadium gotoi: Silurus biwaensis (Tomoda, 1961) [Biwako-ō- molecular identification of the cercaria and namazu] new locality records Isoparorchis eurytremus [V] 1) Misako Urabe successfully sequenced the Parabucephalopsis parasiluri [III] cytochrome c oxidase subunit I gene of the mito- Prosorhynchoides ozakii [III] chondrial DNA (COI mtDNA) of adults of A. Pseudexorchis major [XI] gotoi found in the intestine of Misgurnus anguil- Silurus lithophilus (Tomoda, 1961) [Iwatoko- licaudatus on 28 August 2015 and cercariae namazu] found in Pisidium cinereum nikkoense on 1 Orientocreadium pseudobagri [II] November 2016 both from the small river at Parabucephalopsis parasiluri [III] Midori, Iiyama City, Nagano Prefecture (see Phyllodistomum parasiluri [II] Shimazu, 2016b). Partial COI sequences (889 bp) Prosorhynchoides ozakii [III] determined were identical between the adult and Family Synbranchidae the cercaria. Consequently, the cercaria is defini- Monopterus albus (Zuiew, 1793) [Taunagi] tively identified as A. gotoi, which supports Digeneans Parasitic in Freshwater Fishes of Japan 141

Shimazu’s (2002, 2016b) morphological identifi- guti, 1934.” cation of the cercaria. She also sequenced the p. 178, right column, between line 4 and line 5 large subunit region of the ribosomal RNA gene from top. Insert “Genus Carassotrema Park, (28S rDNA) of adults, and the entire sequence 1938.” (1274 bp) of the 28S was determined. The Gen- p. 178, right column, line 11 from top. Delete Bank accession numbers of the sequences of the “(?).” adult deposited by her are: COI sequence, p. 178, right column, line 13 from top. Read LC215273; and 28S sequence, LC215274. “auratus langsdorfii Temminck and Schlegel, 2) Adults were found in the intestine of M. 1846” for “sp.” anguillicaudatus collected by Akifumi Ohtaka p. 178, right column, line 20 from top. Read from an irrigation canal at Nozaki, Ishikawa, “Asahikita” for “Kamikita-kita.” Hirosaki City, Aomori Prefecture, on 16 August p. 178, right column, lines 4–5 from bottom. 2016 (MPM Coll. No. 21297); and fished in the Delete “(Temminck and Schlegel, 1846) [sic, Ishikari River at Ishikari City (obtained by now Carassius sp.].” Mitsu hiko Asakawa at a local fish market), p. 181, left column, line 14 from top. Read Hokkaido, on 5 October 2016 (MPM Coll. No. “auratus langsdorfii” for “sp.” 21298). p. 181, right column, line 6 from top. Read “[Cercaria D]” for “Cercaria D.” VIII (Shimazu, 2016c). Crepidostomum: life p. 181, right column, line 8 from top. Read cycle “Kumamoto” for “Kumamato.” Moravec (2004) found a progenetic metacer- caria of C. farionis in the body cavity of an II (Shimazu, 2014a) unidentified caddis-fly larva (Trichoptera) from p. 53, right column, lines 2–3 from bottom; p. 54, the trout brook Vydří potok in the Šumava left column, line 5 from bottom; p. 62, right Mountains, present Czech Republic, in 1962. column, line 23 from top; and p. 64, right col- Accordingly, it may well be that progenetic umn, line 4 from top. Read “Asahikita” for metacercariae may occur as well in C. metoecus. “Kamikita-kita.” p. 57, right column, line 10 from top; and p. 59, IX (Shimazu, 2016d). Dimerosaccus oncorhyn- left column, line 15 from top. Insert “OR” after chi: molecular studies “sp.” Bray et al. (2016) and Fayton and Andres p. 70, right column, line 13 from top. Read (2016) also have recently demonstrated that D. “1875” for “1880.” oncorhynchi belongs to the subfamily Opecoeli- p. 76, left column, line 21 from top. Read “Mizu- nae in their molecular studies. moto” for “Mizuno.”

X (Shimazu, 2017a). Neoplagioporus elongatus: IV (Shimazu, 2015a) life cycle p. 77, right column, line 2 from bottom. Read Yano and Urabe (2017) studied the life cycle “Cercaria cystophora C” for “[Cercaria cys- of N. elongatus in the Uji River, Kyoto Prefec- tophora C].” ture (see also Shimazu, 2017a). p. 85, right column, line 18 from top. Insert “of” after “was.” Errata V (Shimazu, 2015b) I (Shimazu, 2013) p. 209, left column, line 21 from bottom; and p. p. 176, left column, between line 14 and line 15 214, left column, line 14 from top. Read “[Cer- from bottom. Insert “Genus Bivesicula Yama- caria XIV, or U]” for “Cercaria XIV, or [U].” 142 Takeshi Shimazu

VI (Shimazu, 2016a) right column, line 10 from bottom. Read p. 1, right column, line 8 from bottom; and p. 6, “[Gori]” for “Gori.” left column, lines 15 and 36 from top. Read p. 173, right column, lines 5–6 from top. Read “[Cercaria H]” for “Cercaria H.” “on dorsal side of” for “around dorsal.” p. 1, right column, line 7 from bottom; and p. 6, left column, lines 19–20 and 37 from top. Acknowledgments Read “[Cercaria VIII, or Shin]” for “Cercaria VIII, or [Shin].” I am grateful to Prof. Akifumi Ohtaka (Faculty p. 1, right column, line 6 from bottom; and p. 6, of Education, Hirosaki University, Hirosaki) for left column, lines 9–10 from bottom. Read collecting the snails and fish examined, Prof. “[Cercariaeum A]” for “Cercariaeum A.” Mitsuhiko Asakawa (School of Veterinary Medi- p. 2, left column, line 1 from top; and p. 6, left cine, Rakuno Gakuen University, Ebetsu) for column, line 13 from bottom. Read “[Cercaria obtaining the fish examined, Dr. František T]” for “Cercaria T.” Moravec (Institute of Parasitology, Branišovská, p. 8, right column, line 20 from bottom. Read Czech Republic) for informing me on the finding “[Metacercaria IV]” for “Metacercaria IV.” of a progenetic metacercaria of Crepidostomum p. 8, right column, line 19 from bottom; and p. farionis, Prof. Misako Urabe (School of Environ- 11, right column, line 2 from bottom. Read mental Science, University of Shiga Prefecture, “[Cercaria A]” for “Cercaria A.” Hikone) for sequencing the samples of Allo- p. 8, right column, line 18 from bottom; and p. creadium gotoi, and Dr. Thomas H. Cribb 12, left column, line 3 from top. Read “[Cer- (School of Biological Sciences, The University caria B]” for “Cercaria B.” of Queensland, Brisbane, Australia) for review- p. 17, left column, line 15 from top. Read “1943” ing a draft of the manuscript. Finally, I thank my for “1942.” wife Tomiko Shimazu for serving as assistant to me in the field for many years. VII (Shimazu, 2016b) p. 59, right column, line 5 from bottom; and p. References 60, left column, lines 17–18 from top. Read “Koumi” for “Komi.” Bray, R. A., T. H. Cribb, D. T. J. Littlewood and A. Wae- schenbach, 2016. The molecular phylogeny of the dige- p. 62, right column, line 22 from bottom. Insert nean family Opecoelidae Ozaki, 1925 and the value of “subunit” between “oxidase” and “I.” morphological characters, with the erection of a new p. 64, right column, line 3 from top. Read subfamily. Folia Parasitologica, 63: 013. “Cuvier in Cuvier and Valenciennes, 1829” for Fayton, T. J. and Andres, M. J. 2016. New species of Pla- “Linnaeus, 1758.” gioporus Stafford, 1904 (Digenea: Opecoelidae) from California, with an amendment of the genus and a phy- p. 64, right column, line 10 from top. Insert logeny of freshwater plagioporines of the Holarctic. “(Cyprinidae)” between “1872)” and Systematic Parasitology, 93: 731–748. “(Shimazu.” Moravec, F. 2004. Metazoan Parasites of Salmonid Fishes p. 68, right column, between line 5 and line 6 of Europe. 510 pp. Academia, Praha. from top. Insert “Life cycle. Not known.” Shimazu, T. 2016e. Japan. pp. 260–261. In Scholz, T., V. p. 72, left column, line 2 from bottom; and p. 72, V. Besprozvannykh, T. E. Boutorina, A. Choudhury, T. H. Cribb, A. V. Ermolenko, A. Faltýnková, M. B. right column, line 4–5 from top. Read “Asa- Shedko, T. Shimazu, and N. J. Smit, Trematode diver- hikita” for “Kamikita-kita.” sity in freshwater fishes of the Globe I: ‘Old World.’ Systematic Parasitology, 93: 257–269. (Electronic sup- IX (Shimazu, 2016d) plementary material, doi:10.1007/s11230-016-9630-3) p. 164, left column, line 1 from bottom; p. 166, Urabe, M., R. Ishibashi and K. Uehara 2015. The life cycle and molecular phylogeny of a gorgoderid trematode right column, line 9 from bottom; and p. 168, Digeneans Parasitic in Freshwater Fishes of Japan 143

recorded from the mussel Nodularia douglasiae in the porus elongatus (Goto and Ozaki, 1930) (Opecoelidae: Yodo River, Japan. Parasitology International, 64: 26–32. Plagioporinae), with notes on potential second interme- Yano, A. and M. Urabe 2017. Larval stages of Neoplagio- diate hosts. Parasitology International, 66: 181–185.