JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology

Species Diversity, 2oo1, 6,133-164

FreshwaterFishes of the Kuril Islands and AdjacentRegions

Theodore W. Pietsch', Kunio Amaoka2, Duane E. Stevensoni, Erin L. MacDonald', Brian K. Urbaini and Juan Andres L6pez3

i School oj`Aguatic and Ftsher:y Stiences, University of VP(ashington, Box 35510a Seattle Wicshington 98195510a USA E-mail: tuzp@u. washington.edu 2 Fticulty ofFisheries, HbhkaidO Vniversity, Htzkocinte, HOhkaicto, 041-8611 Japan 3 Department of Zoology and Genetics. fowa State Ulriiversity, Ames, fowa 50011-a22a USA

(Received 19 May 2000; Accepted 15 Februarvy 2001)

Fishes were collected in freshwater habitats during six consecutive sum- mers (1994-1999) on the 30 major islands of the Kuril Archipelago as part of a joint U.S., Russian, and Japanese biotic survey and inventory, which has come to be known as the international Kuril Island Project (IKIP). A total of 39,269 specimens, representing eight families, 12 genera, and 28 species (in- cluding two previously undescribed species, 11 new records for the Archipel-

ago, and 33 new records for various islands), was analyzed in the context of

published literature on the freshwater fish faunas of the Kuril Islands and adjacent regions, namely Kamchatka, Hokkaido, and Sakhalin. The geologic history of the Archipelago is reviewed in light of present-day fish distribu- tions. As expected, species diversity, as compared to that of Kamchatka and

Hokkaido, fans off abruptly as one moves centrally from both the northern

and southern ends of the Archipelago, the number of species dropping to

zero on some of the central islands. It appears that differences in island size,

and therefore available habitat, are the primary determinant of diversity, coupled with the secondary effects of former connections between land-

masses and distance from adjacent source biotas.

Key Words: Fishes, freshwater, geologic history, zoogeography, Kuril Is-

lands, Hokkaido, Sakhalin, Kamchatka.

Introduction

In the current renewal of interest in global biodiversity, the complex and threatened fioras and faunas of tropical regions of the world have received consid- erable attention. In contrast, certain temperate and boreal biotas have been ig- nored, despite high biotic diversity, the absence of basic taxonomic miiormation, and the threat of plant and animal extinctions. Of all such northern regions, the is- "biologically lands of the Kuril Archipelago are among the most unknown" and, at the same time, in danger of over-exploitation (Stephan 1974). A chain of more than 56 islands, the system is only slightly smaller than the Hawaiian Islands, covering an area of 15,600km2 and providing 2,409km of coastline (Fig, 1). Stretching 1,2oo

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134 Theodore W. Pietsch et al.

Fig, 1, The islands of the Kuril Archipelago. Note that the name Ushishir incorporates two very closely situated islands (almost touching each other at low tide), Yankicha and Rypon- "The kicha, @ Oxfbrd University Press 1974. Reprinted from Kuril Islands: Russo-Japanese Erontier in the Pacific" by John J. Stephan (1974), by permission of Oxford University Press.

km between Hokkaido, the northernmost island of Japan, and the Russian penin- sula of Kamchatka (from 430 to 510N latitude), the Kurils divide the Sea of Okhotsk from the PacMc Ocean and form the northern extension of an insular arc that orig- inates in the Ryukyu and Mariana archipelagos to the south and continues as the Aleutian Islands to the north and east. All of the Kurils are volcanic in origin, ranging in age from Upper Cretaceous to Late Pleistocene (Table 1); each island has a unique geological and biological history. Substantial epportunities fbr in situ diversification are provided by great distances between the islands and mainland source biotas, and by significant barriers to plant and animal dispersal, such as deep channels between islands, associated with strong ocean currents. Very little has been published on the fauna of the Kuril Islands; general state- ments about the larger mammals, marine birds, and commercially important fishes (salmon and cod) and crustaceans (king crab) are the only exceptions. Brown bears, wolves, squirrels, and snowshoe hares inhabit the 1arger islands. Foxes brought from Japan to Yankicha (Ushishir) fbr commercial breeding are now abundant there. Reindeer are raised on Paramushir. Tens of thousands of sea otters, fur seals, and sea lions once populated the chain, but today only sma}1 colonies exist on the central islands (Stephan 1974; personal observations). The lit- erature records more than 170 species of birds, but some unpublished esttmates

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Fishes of the Kuril Islands 135

are as high as 280 (V. A. Nechaev 1969, and personal communication, 14 May 1993). Island rivers seasonully have heavy runs of salmon, and offtshore waters support abundant populations of cod (Gadus macrocephalus Tilesius, 1810), mackerel (Scomber J'aponicus Houttuyn, 1782), and ocean perch (Sebastes spp,). King crab from Kunashir and Iturup are stil1 harvested in large quantities. Although the general outlines of the fiora and fauna of the Kurils can be sum-

marized, more detailed information is nonexistent or unpublished; what little is published is confined to the Russian and, to a lesser extent, the Japanese scientific

literature. Little systematic collecting has ever been done, and since the close of

World War II, only a few Russian biologists have done any work of significance, With the exception of a few Russian collections (e.g., those of the various institutes ef the Russian Academy of Sciences at Vladivostok, Magadan, St. Petersburg, Moscow, Novosibirsk, and Yuzhno-Sakhalinsk), museum samples of plants and an- imals originating from the Kurils are unavailable and frozen tissues are com- pletely non-existent, The biota is a mix of Japanese, Kamchatkan, and endemic species, but the biodiversity of the islands, relative to each other and to the main-

land, is unknown. Limited coastal regions of several of the larger, more strategi- cally positioned islands have been heavily impacted since the close of World War II by the construction of Russian military installations, but otherwise the islands re-

main undisturbed, Only six islands are currently inhabited. Realizing the scientific value of a detailed biological survey of the islands of the Kuril Archipelago, and with financial support from the U.S. National Science Foundation, we have joined with Japanese scientists, primarily from the Sapporo and Hakodate campuses of Hokkaido University, and Russian scientists from the

Institute of Biology and Soil Sciences, Russian Academy of Sciences, Far East Branch, Vladivostok, to conduct a six-year program designed to survey and inven- tory the 30 major islands of the Kurils (including the five primary isles of the Habo- mai group, see Fig. 1): the International Kuril Island Project (IKIP), focusing on plants, aquatic and terrestrial insects, spiders, freshwater and terrestria] mol-

lusks, freshwater fishes, amphibians, and reptiles. Having completed six field sea- sons (1994-1999) of collecting, on al1 30 major islands, we are now able to provide some general distributional and zoogeographic infoumation. Here we repert specifi

ically on the freshwater fishes. Prior to the work described here, at least 88 species of fishes (in 57 genera and 20 families) had been reported in freshwater habitats of the Kurils and adjacent re- gions (Hokkaido, Sakhalin Island, and Kamchatka), but only 19 species (11 genera and six families) were thought to occur in the Archipelago itself (Taranetz 1937; Kuronuma 1943; Berg 1948, 1949a, 1949b; M. K. Glubokovsky, personal communica- tion, 22 and 29 March 1993; Reshetnikov et al. 1997). Nine of these species were said to occur throughout the Kurils (six anadromous salmonids and osmerids, and three euryhaline gasterosteids), and, in addition to these, one was reported from the northern Kurils [an osmerid, Lb;pomesus oltdus (Pallas, 1814)] and three from the southern Kurils [the osmerids Hblpomesus nipponensis McAllister, 1963 and H jmponicus (Brevoort, 1856), and a gasterosteid, Pungitius pungititts sinensis (Guichenot, 1869)]. Three other species were reported from Iturup [a cyprinid, Tri- bolodon brandti (Dybowski, 1872), and two gobiids, Tridentiger obscurus (Tem- minck and Schlegel, 1845) and Gymnogobius urotaenia (Hilgendorf, 1879)], three from Shikotan [the gobiids Luciogobius guttatus Gil1, 1859, G),mn(rgobius moro-

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ranus (Jordan and Snyder, 1901), and G. castaneus (O'Shaughnessy, 1875)], and two from Kunashir (the gobiids Lucingobitts guttatus and (lp)mnogobius urotaenia). Ac- cording to M. K. Glubokovsky (persenal communication, 22 and 29 March 1993), only a few random collections of non-commercially important freshwater fishes had been made during the past 50 years, and these ef!brts had been confined to only three or four of the larger islands of the chain. The present analysis, based on an examination of some 39,269 specimens collected throughout the Archipelago, indi- cates the presence of 28 species in eight families and 12 genera. Eleven of these species, including two previously undescribed forms (Saruwatari et al, 1997; Stevenson 2000 and in preparation), represent new records for the Archipelago. The collections also represent 33 new records for various islands (see Table 1).

Methods and Materials

Collections of whole specimens of plants and animals, as well as tissue sam- ples for future molecular studies, were made by teams of scientists from Japan, Russia, and the U.S., averaging 30 people for each of the six annual summer expedi- tions completed to date (Table 1), Taxa of major emphasis included vascular plants, aquatic and terrestrial insects, spiders and harvestmen, freshwater and terrestrial

mollusks, freshwater fishes, amphibians, and reptiles, but signMcant collections of lichens, mosses, liverworts, fungi, diatoms, platyhelminths, oligochaetes, am- phipods, pseudoscorpions, mites, decapods, water fleas, centipedes, millipedes, and marine fishes were also made. A research vessel provided by the Russian Academy of Sciences, Far East Branch (the 68.5 m R/V Projlessor Bogorov in 1994, 1995, and 1997; the 75.5m R/V Academic Qparin in 1996, 1998, and 1999), served as the means of transportation to and from the islands, as sleeping quarters, the source of all meals (prepared by a cook and support staff attached to the vessel), and as a float- ing research laboratory while at sea and when on-site. Large rubber infiatables, equipped with 40-horsepower engines, were used to transport equipment and per- sonnel between ship and shore. On some of the larger inhabited islands (Para- mushir, Urup, Iturup, and Kunashir), Russian military vehicles were used to move between distant collecting sites. Collection sites were selected to maximize geo- graphical and habitat diversity, Fishes were collected by seine, dip-net, rotenone, and hook and line; a few specimens were provided by local fishers. To the extent possible, collections were sorted and identMed aboard ship, and field data entered into a computer database. Following each expeditien, the data were downloaded to an IKIP website (available via http://artedi.fish.washington.edu/ikiplhome.html), maintained at the University of Washington to provide easy access to project re- sults and databases. The website provides a fUll-text search intembce to access both locality and taxonomic databases. In addition, the locality database can be ac- cessed via maps that show survey and inventory sites. This map-based browser al- lows the user to click on maps to select an island, then to click on major collection areas to get a list of ai! collection sites for that area, and, finally, te view the ful1 lo- cality record. All ichthyological material is deposited in the University of Washington Fish Collection (UW, see http://artedi.fish.washington.edu). All fish lengths are stan- dard length. Freshwater fishes are defined as those taxa collected by us in non-ma-

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Fishes of the Kuril Islands 137

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rine habitats. The classification follows that of Nelson (1994). The status, spelling, and authorship and dates of Latinized binomials fo11ow those given by Eschmeyer (1998). English common names are those given by Reshetnikov et al. (1997). Japan- ese common names were provided by the second author. No attempt is made to provide fu11 information on the geographic distribution of each species, but rather to indicate occurrences only on the Kurils, and on Hokkaido, Sakhaiin Island, and the Kamchatka Peninsula, For additional geographic data, see Reshetnikov et al. "Million (1997). years before present" is abbreviated MYBP,

Results

Following is a list of the fishes recorded from freshwater habitats of the Kuril Islands and adjacent regions (Hokkaido, Sakhaiin Island, and Kamchatka). Com- mon names are given in the order English, Russian, and Japanese, where applica- ble. Species names in bold are those whose presence on the Kuril Islands was veri- fied by our collections; Kuril Island occurrences fbr the rematning taxa are based on a survey of the available scientific literature. Individual lots collected from each island are listed alphabetically by species in the Appendix. For numbers of new records for various islands, see Table 1. For an analysis of species shared be- tween islands and adjacent regions, see Fig. 2.

I. FamilyPetromyzontidae 1. GenusLampetraBonnaterre 1. Lampetra tridentata (Riehardson, 1836); Pacific Lamprey, Mitsuba-yatsume (anadromous): Hokkaido, Kamchatka (Lindberg and Legeza 1959; Prokhorov and Grachev 1965; Iwata 2000). 2. Lampetrajtrponica (Martens, 1868); Arctic Lamprey, Kawa-yatsume (anadro- mous): Hokkaido, Sakhalin, Kamchatka (Taranetz 1937; Berg 1948; Schmidt 1950; Lindberg and Krasyukova 1969; Nikoforov et al. 1989, 1994; Reshetnikov et al, 1997; Iwata 2ooO). IKIP records: Iturup. 3. Lampetra kessleri (Anikin, 1905); Siberian Lamprey, Shiberia-yatsume (freshwater): Hokkaido, southern Sakhalin (Berg 1948; Schmidt 1950; Iwata 1990a; Nikoforov et al. 1994; Reshetnikov et al. 1997). IKIP records: Iturup, Kunashir. 4. Lampetra reissneri (Dybowski, 1869); Asiatic Brook Lamprey, Suna-yatsume (freshwater): Hokkaido, Kuril Islands, western Kamchatka (Taranetz 1937; Berg 1948; Lindberg and Legeza 1959; Iwata 1990a; Reshetnikov et al. 1997).

II. FamilyAcipenseridae 2. GenusAcipenserLinnaeus 5. Acipenser medtrostris Ayres, 1854; Green Sturgeon, Ch6zame (anadromous): Hokkaido, Sakhalin (Taranetz 1937; Berg 1948; Schmidt 1950; Nikoforov et al, 1994; Reshetnikov et at. 1997; Amaoka 2000a).

3. Genus Huso Brandt and Ratzeburg 6. Httso dauricus (Georgt, 1775); Kaluga, Dauria-ch6zame (anadromous and fireshwater): Hokkaido (Reshetnikov et aZ. 1997; Amaoka 2000a).

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III. FamilyAnguillidae 4. GenusAnguillaSchrank 7, Anguilla joponica Temminck and Schlegel, 1846; Japanese Eel, Unagi (catadromous): Hokkaido (Tabeta 1990),

IV. FamilyClupeidae 5. GenusClmpeaLinnaeus 8. Clt{pea pallasii Valenciennes, in Cuvier and Valenciennes, 1847; Pacific Her- ring, Nishin (marine, but often entering mouths of rivers; lacustrine forms are also known): Hokkaido, Sakhalin, Kamchatka (Nikoforov et at. 1994; Reshetnikov et al. 1997; Sato 2000).

V. FamilyCyprinidae 6. GenusCarassiusJarocki 9. Ctirassius auratus (Linnaeus, 1758); Goldfish, Kin-buna (freshwater): Hokkaido, Sakhalin, Kamchatka (Taranetz 1937; Nikofbrov et al. 1989, 1994; Tanigmchi 1990).

7, GenusdyprinusLinnaeus 10. Qyprinus carpio Linnaeus, 1758; Carp, Koi (freshwater): Hokkaido, Sakhalin, Kamchatka (Taranetz 1937; Nikoforov et al. 1989, 1994; Hosoya 1990).

8. GenusEIQpichthysBleeker 11. EIQpichthys bantbttsa (Richardson, 1845) (freshwater): Sakhalin (Reshetnikov et al, 1997).

9. GenusGobioCuvier 12. Gohio soldatovi Berg, 1914; Pacific Gudgeon (freshwater): Sakhalin (Nikofbrov et al. 1989; Reshetnikov et al. 1997).

10. GenusHbmibarbusBIeeker 13. Hemibarbus labeo (Pallas, 1776); Amur Barbel, Skin-carp (freshwater): Sakhalin (Reshetnikov et al. 1997).

11. GenusLeuciscusCuvier 14. Leuciscus waleckii (Dybowski, 1869); Amur Ide (freshwater): Sakhalin (Taranetz 1937; Berg 1949a; Nikofbrov et al. 1989; Reshetnikov et al. 1997).

12. GenusPhoxinusAgassiz 15. Phoxinus lagowskii Dybowski, 1869; Lagowski's Minnow, Abura-haya (fresh- water): Sakhalin (Nikoforov et al. 1989; Itai 1990), 16. Phoxinus percnurus sachalinensis (Pallas, 1814); Lake Minnow, Yachi-ugui (freshwater): Hokkaido, Sakhalin (Taranetz 1937; Berg 1949a; Kluchareva 1964; Nikoforov et al. 1989, 1994; Goto 1990b; Reshetnikov et al. 1997). 17. Phoxinus phoxintts (Linnaeus, 1758); Commen Minnow (freshwater): Sakhalin (Berg 1949a; Nikofbrov et al. 1989; Reshetnikov et al. 1997).

13. GenusPseudaspiusDybowski 18. Rseudaspius loptocqphalus (Pallas, 1776); Flathead Asp (freshwater): Sakhalin (Nikoforov et al, 1989).

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14. GenusPseudorasboraBleeker 19. Rseudorasbora pumita pumila Miyadi, 1930; Moroco, Shinai-motsugo (fresh- water): Hokkaido (Uchiyama 1990).

15. GenusRhodeusAgassiz 20. Rhodeus sericeus (Pallas, 1776); Amur Bitterling (freshwater): Sakhalin (Tym and Poronai rivers) (Reshetnikov et at. 1997).

16. Genus7)'ibolodonSauvage 21. Tribolodon brandti (Dybowski, 1872); Pacjfic Redfin, Maruta-ugui (anadro- mous): Hokkaido, Sakhalin, Iturup, Kunashir (Berg 1949a; Kluchareva 1964; Gritsenko 1974; Sakai 1990; Nikoforov et al. 1994; Reshetnikov et al. 1997). IKIP records: Kunashir. Comments. 71ribolodon taczanowskii (Steindachner, 1881) is considered to be a junior synonym of Tribotodon brandti, following the conclusions of Sakai (1990). 22. Tribolodon ezoe Okada and Ikeda, 1937; Redfin, Ezo-ugui (freshwater): Hokkaido, Sakhalin (Berg 1949a; Gritsenko 1974; Kurawaka 1977; Sakai 1990; Nikoforov et al. 1994; Reshetnikov et al. 1997), IKIP records: Iturup, Zelionyi. 23. Tlr"ibotodon hakuensis (GUnther, 1877); Big-scaled Redfin, Ugui (anadromous and freshwater): Hokkaido, Sakhalin (Berg 1949a; Gritsenko 1974; Kurawaka 1977; Nikoforov et al, 1989, 1994; Sakai 1990; Reshetnikov et al. 1997). IKIP records: Iturup, Kunashir.

VI. FamilyCobitidae 17. GenusCObitisLinnaeus 24. Cbbitis lutheri Rendahl, 1935; Luther's Spiny Loach (freshwater): Sakhalin (Reshetnikov et al. 1997).

18. GenusMisgurnusLacepede 25. Misgurnus anguillicaudattts (Cantor, 1842); Amur Mud Loach, Doj6 (fresh- water): Hokkaido, Sakhaiin (Nikoforov et at. 1989, 1994; Saito 1990; Reshet- nikov et al. 1997).

VII, FamilyBalitoridae 19. GenusBarbatulaLinck 26. Barbatuta toni (Dybewski, 1869); Siberian Stone Loach, FukudQj6 (fresh- water): Hokkaido, Sakhalin (Berg 1949a; Kluchareva 1964; Nikoforov et al. 1989, 1994; Sawada 1990; Reshetnikov et al, 1997),

20. GenusLefuaHerzenstein 27. LetLta costata costata (Kessler, 1876); Eight-whiskered Stone Loach (fresh- water): Sakhalin (Reshetnikov et al. 1997). 28. Lefbea costata nikkonis (Jordan and Fewler, 1903); Stone Loach, Ezo-hotokedQj6 (freshwater): Hokkaido (Sawada 1990).

VIII. FamilySiluridae 21. GenusSilurusLinnaeus 29, Silurus asotus Linnaeus, 1758; Amur Catfish, Namazu (freshwater): Hokkaido,

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Sakhalin (Nikoforov et al. 1989; Kebayakawa 1990),

IX, FamilyEsocidae 22. GenuslisoxLinnaeus 30. Esox lttcius Linnaeus, 1758; Northern Pike (freshwater):Kamchatka (Reshet- nikov et al, 1997). 31, Esox reichertii Dybowski, 1869; Amur Pike (freshwater)iSakhalin (Nikoforov et al. 1989; Reshetnikov et at. 1997).

X, FamilyOsmeridae 23. GenusHl)lpomesusGill 32. HZypomesus chishimaensis Saruwatari, L6pez and Pietsch, 1997; Kunashir Smelt, Chishima-wakasagi (anadromous): Iturup, Kunashir, Zelionyi (Saru- watari et al. 1997). IKIP reeords: Iturup, Kunashir, Zelionyi, Tanfilyeva. Comments. This recently described species (Saruwatari et al. 1997) was previ- ously recorded from the southern Kuril Islands as llypomesus nipponensis McAllister, 1963 by Klukanov (1977). 33. HmpomesusJ'mponicus (Brevoort, 1856); Japanese Smelt, Chika (anadromous): Hokkaido, Sakhalin, southern Kuril Islands (Taranetz 1937; Klukanov 1966, 1970b, 1977; Saruwatari et al, 1997; Amaoka 2000b). IKIP records: Shikotan. 34. Ilypomesus nipponensis McAllister, 1963; Wakasagi (anadromous): Hokkaido, Sakhalin (Klukanov 1970b; Nikofbrov et al. 1989, 1994; Saruwatari et al. 1997; Amaoka 2000b). 35. Hypomesus olidus (Pallas, 1814); Pond Smelt, Ishikari-wakasagi (anadro- mous, and land-locked freshwater): Hokkaido, Sakhalin, northern Kuril Is- lands, Kamchatka (Berg 1948; Klukanov 1966, 1970b, 1977; Nikoforov et al. 1989, 1994; Ito 1990; Reshetnikov et al. 1997; Saruwatari et al. 1997). IKIP records: Kamchatka.

24. GenusMallotusCuvier 36. Mallotus villosus (Mtt11er, 1776); Capelin, Karafuto-shishame (marine, but often entering brackish and freshwater): northern coast of Hokkaido, south- eastern Sakhalin, Kamchatka (Berg 1948; Lindberg and Legeza 1965; Reshet- nikov et al. 1997; Amaoka 2000b).

25. GenusOsmerusLinnaeus 37. 0smerus mordax (Mitchil1, 1814); Rainbow Smelt, Kyari-uo (anadromous): Hokkaido, Sakhalin, Kuril Islands, Kamchatka (Berg 1948; Schmidt 1950; Klukanov 1970a; Nikeforov et al. 1989, 1994; Reshetnikev et al. 1997; Amaoka 2000b).

26. GenusPlecoglossusTemminckandSch}egel 38. Plecaglossus altiveJis altivelis Temminck and Schlegel, 1846; Ayu (anadro- mous): western Hokkaido (Kinoshita 2000).

27. GenusSpirinchusJordanandEvermann 39. Spirinchus lanceolatus (Hikita, 1913); Shishamo (anadromous): southeastern Hokkaido (Amaoka 2000b).

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142 Theodore W. Pietsch et at.

XI. FamilySalangidae 28, GenusSalangichtitysBleeker 40. Salangichthys microdon (Bleeker, 1860); Common Icefish, Shira-uo (marine, but often entering brackish and freshwater): Hokkaido, Sakhalin (Berg 1948; Lindberg and Legeza 1965; Reshetnikov et at, 1997; Kinoshita 2000).

XII. FamilySalmonidae 29. GenusBrachympstaxGUnther 41. Bracltympstax lenok (Pallas, 1773); Lenok (freshwater): northwestern Sakhalin (Taranetz 1937; Nikoforov et aL 1989),

30, GenusCOregonusLinnaeus 42. COragonus nasus (Pallas, 1776); Broad Whitefish (euryhaline): Kamchatka (Berg 1948; Reshetnikov et al, 1997; S. Shedko, personal eommunication, 15 September 2000). 43. Coragonus sardinella Valenciennes, in Cuvier and Valenciennes, 1848; Least Cisco (euryhaline): Kamchatka (Berg 1948; Reshetnikov et al. 1997; S. Shedko, personal communication, 15 September 2000). 44. Ct)regonus ussuriensis Berg, 1906; Ussuri Whitefish (euryhaline): northwest- ern Sakhalin (Taranetz 1937; Nikoforov et at. 1989).

31. GenusHuchoGUnther 45. Httcho peri:yi (Brevoort, 1856); Sakhalin Taimen, ItO (anadromous, freshwa- ter): Hokkaido, Sakhalin, southern Kuril Islands (Taranetz 1937; Berg 1948; Kimura 1966; Nikofbrov et al. 1989, 1994; Kawamura 1990; Reshetnikov et al. 1997).

32. GenusOncorhynchusSuckley 46. 0ncorltynchus gorbuscha (Walbaum, 1792); Pink Salmon, Karafutomasu (anadromous): everywhere within region of interest, including Hokkaido, Sakhalin, the Kuril Islands, and Kamchatka (Taranetz 1937; Berg 1948; Schmidt 1950; Lindberg and Legeza 1965; Nikoforov et al. 1989, 1994; Reshet- nikov et al. 1997; Ida 2000). IKIP records: Shumshu, Urup, Iturup, Kunashir, Shikotan. 47. 0ncorhynchus keta (Walbaum, 1792); Chum (Dog) Salmon, Sake, Shiro-zake (anadromous): everywhere within region of interest, including Hokkaido, Sakhalin, the Kuril Islands, and Kamchatka (Taranetz 1937; Berg 1948; Schmldt 1950; Lindberg and Legeza 1959; Nikofbrov et al. 1989, 1994; Reshet- nikov et al. 1997; Ida 2000). IKIP records: Paramushir, Urup, Iturup, Kunashir, Shikotan. 48. 0ncoritynchus kisutch (Walbaum, 1792); Coho (Silver) Salmon, Gin-zake (anadromous): Hokkaido, Sakhalin (Tym River), Kamchatka (Taranetz 1937; Berg 1948; Lindberg and Legeza 1965; Nikofbrov et at. 1994; Reshetnikov et al. 1997; Ida 2000). IKIP records: Kamchatka, Shumshu, Paramushir, Urup, Iturup, Kunashir. 49. 0ncordynchus masou masou (Brevoort, 1856); Cherry (Masu) Salmon, Sakura-masu (anadromous, and land-locked in freshwater): Hokkaido, Sakhalin, western Kamchatka (Taranetz 1937; Berg 1948; Lindberg and Legeza 1965; Nikoforov et al. 1989, 1994; Reshetnikov et al. 1997; Ida 2000). IKIP records: Kunashir.

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50. 0ncorltynchus rnykiss (Walbaum, 1792); Rainbow Trout, Kamchatka Steelhead, Nlji-masu (anadromous, and land-locked in freshwater): northern Kuril Is- lands (?), Kamchatka (Berg 1948; Reshetnikov et al. 1997; Ida 2000). Comments. Introduced and now spawning naturally in Japan, 51. 0ncorh),nchus nerka (Walbaum, 1792); Sockeye Salmon, Beni-zake (anadro- mous, and land-locked in freshwater): Hokkaido, Sakhalin, Iturup (rivers Slavnaya, Raidovaya, Gorelaya, Kurilka, and Kuibyshevka; lakes Krasivoye and Sopochnoye), Kamchatka (Berg 1948; Lindberg and Legeza 1959; Ivankov 1984; Nikofbrov et al. 1994; Reshetnikov et al. 1997; Ida 2ooO). IKIP records: Kamchatka, Shumshu, Paramushir, Onekotan, Iturup. 52. 0ncorhynchus tshaiQ),tscha (Walbaum, 1792); King Salmon, Masunosuke (anadromous, and land-locked in freshwater): Hokkaido, Kamchatka (Reshet- nikov et al. 1997; Ida 2000).

33. GenusProsQpiumJordan 53. Prosopium cylindraceurn (Pennant, 1784); Round Whitefish (freshwater): northern Kamchatka (Reshetnikov et al. 1997).

34. GenusSalvelinusRichardson 54, Salvelinus leucomaenis (Pallas, 1814); East Siberian Char, Kundscha, Ame- masu (anadromous, freshwater): everywhere within region of interest, includ- ing Hokkaido, Sakhalin, the Kuril Islands, and Kamchatka (Taranetz 1937; Berg 1948; Schmidt 1950; Lindberg and Legeza 1965; Nikofbrov et al. 1989, 1994; Reshetnikov et al. 1997; Ida 20oo). IKIP records: Paramushir, Urup, Iturup, Kunashir, Shikotan, Tanfilyeva. 55, Salvelinus malma (Walbaum, 1792); Dolly Varden, Malma, Oshorokoma (anadromous and freshwater): everywhere within region of tnterest, includ- ing Hokkaido, Sakhalin, the Kuril Islands, and Kamchatka (Taranetz 1937; Berg 1948; Lindberg and Legeza 1965; Nikoforov et al. 1989, 1994; Reshetnikov et al. 1997; Ida 2000). IKIP records: Kamchatka, Shumshu, Paramushir, Onekotan, Shiashkotan, Rasshua, Simushir, Urup, Iturup, Kunashir.

35. Genus ThormallusLinck 56. Thymallus arcticus (Pallas, 1776); Arctic Grayling (freshwater): Sakhalin, Kamchatka (Taranetz 1937; Nikoforov et al. 1989; Reshetnikov et al. 1997).

XIII. FamilyGadidae 36. GenusElegin"sFischer 57. Elaginus gracilis (Tilesius, 1810); Saffron Cod, Komai (marine, but often enter- ing mouths of rivers and inhabiting freshwater lakes): Hokkaido, Sakhalin, Kamchatka (Nikoforov et al. 1994; Reshetnikov et al. 1997; Endo 2000),

37. GenusLotaOken 58, Lota lota (Linnaeus, 1758); Burbot (freshwater): Sakhalin (Nikofbrov et al. 1989; Reshetnikov et al. 1997).

XIV, FamilyMugilidae 38. GenusMugilLinnaeus 59. Mugil cqphalus cqphalus Linnaeus, 1758; Striped Mullet,Bora (marine, eury-

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haline): Sakhalin (Berg 1949b; Lindberg and Legeza 1965; Reshetnikov et al. 1997; Seno 2000). 60. M"gil soiay Basilewsky, 1855; Haarder (marine, euryhaline): seuthern Sakhalin (Berg 1949b; Kluchareva 1964; Lindberg and Legeza 1965; Nikoforov et al. 1994).

XV. FamilyGasterosteidae 39. GenusGasterosteusLinnaeus

61. Gasterosteus aculeatus aculeatus Linnaeus, 1758; Threespine Stickleback, Itoyo (euryhaline): everywhere within region of interest, including Hokkaido, Sakhalin, the Kuril Islands, and Kamchatka (Berg 1949b; Nikofbrov et al. 1989, 1994; Hirai 1990; Zuganov 1991; Reshetnikov et al. 1997). IKIP records: Kamchatka, Shumshu, Paramushir, Simushir, Urup, Iturup, Kunashir, Shikotan, Zelionyi, Iurii, Tanfilyeva.

40. GenusPungitiusCoste 62, Pungitius pungitius pungitius (Linnaeus, 1758); Ninespine Stickleback, Ibara-tomiyo (euryhaline): everywhere within region of interest, including Hokkaido, Sakhalin, the Kuril Islands, and Kamchatka (Berg 1949b; Nikofbrov et al. 1989; Takada 1990; Zuganov 1991; Reshetnikov et al. 1997). IKIP records: Kamchatka, Shumshu, Paramushir, Iturup, Kunashir, Shikotan, Polonskogo, Zelionyi, Iurii, Anuehina, Tanfilyeva. 63. Pungitius pungitius sinensis (Guichenot, 1869); Amur Ninespine (Chinese) Stickleback, Tomiyo (freshwater): Hokkaido, Sakhalin, Kuril Islands, western Kamchatka (Berg 1949b; Nikoforov et al. 1989, 1994; Takada 1990; Zuganov 1991; Reshetnikov et al. 1997). IKIP records: Shumshu, Paramushir, Urup, Iturup, Kunashir, Shikotan, Polonskogo, Zelionyi, Iurii, TanfiIyeva, 64. Pungitius tymensis (Nikolsky, 1889); Sakhalin Ninespine (Shortspine) Stick- leback, Ezo-tomiyo (freshwater): Hokkaido, Sakhalin, Kuril Islands (Berg 1949b; Takada 1990; Zuganov 1991; Nikoforov et at. 1994; Reshetnikov et at. 1997). IKIP records: Kunashir, Shikotan.

XVI. FamilyCottidae 41. GenusCottusLinnaeus 65. COttus amblystomqpsis Schmidt, 1904; Sakhalin Sculpin, Ezohana-kajika (freshwater, euryhaline): Hokkaido, Sakhalin (Lutoga, Aleksandrovka, Poronai, and Tym rivers), Kunashir, Iturup (Berg 1949b; Goto 1983, 1990a; Lindberg and Krasyukova 1987; Nikoforov et al, 1994; Reshetnikov et al. 1997). IKIP records: Iturup, Kunashir, Shikotan. 66. Cottus hangiongensis Mori, 1930; Kankyo-kajika (freshwater): Hokkaido (Goto 1990a). IKIP records: Kunashir. 67. COttus nozawae Snyder, 1911; Hana-kajika (freshwater): Hokkaido (Goto 1983, 1990a). 68. COttus poecilopus Heckel, 1837; Spotted Sculpin (euryhaline): Kamchatka (Berg 1949b; Nikolsky 1956; Novikov 1966; Chereshnev 1982; Reshetnikov et at. 1997).

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42. GenusMagalocottusGill 69. Magalocottus platycqphalus (Pallas, 1814); Flathead Sculpin (marine, but often 1997). entering mouths of rivers): Sakhalin, Kamchatka (Reshetnikov et at.

43. GenusMesocottusGracianov 70. Mesocottus haitoj (Dybowski, 1869) (freshwater): rivers of northern Sakhalin (Nikolsky 1956; Nikoforov et al. 1989; Reshetnikov et al. 1997).

44. GenusMyoxocephalusTilesius 71. Myoscocephalus stelleri Tilesius, 1811; Gisu-kajika (marine, but often entering mouths of rivers): Hokkaido, Kamchatka (Yabe 20oo).

XVII. FamilyEleotridae 45. GenusPerccottusDybowski 72. Perccotttts glenii Dybowski, 1877; Amur Sleeper (freshwater):northwestern Sakhalin (Nikoforov et al. 1989).

XVIII. FamilyGobiidae 46. GenusAcanthogobiusGil1 73. AcanthQgobius j7avimanus (Temminck and Schlegel, 1845); Yellowfin Goby, Mahaze (euryhaline): Hokkaido (Lindberg and Krasyukova 1975, 1987; Pinchuk 1978; Hagihara 2000). 74. Acanthcrgobius lactipes (Hilgendorf, 1879); Ashishirohaze (euryhaline): Hokkaido, eastern Sakhalin (Lake Tunaicha) (Berg 1949b; Lindberg and Krasyukova 1975; Pinchuk 1978; Reshetnikov et al, 1997; Hagihara 2000).

47. GenusGlossegobiusGil1 75. Glossogobius otivacetts (Temminck and Schlegel, 1845); Urohaze (euryhaline): Hokkaido, southern Sakhalin (Aniva Bay, Lake Chepisani) (Lindberg and Krasyukova 1975; Iwata 1990b; Reshetnikov et al. 1997).

48. Genus GymnogobiusGill Comments. The recent discovery of the holotype of the type species ef Chaenqgobius (Chaen(rgobiLts annularis Gil1, 1859) has necessitated the reallo- cation of the following six species to the genus (lymnogobius Gil1, 1863 (see Stevenson 2ooO). 76. dymnogobius castaneus (O'Shaughnessy, 1875); Chestnut Goby, Biringo (eu- ryhaline): Hokkaido, Sakhalin, Shikotan (Anama and Notoro bays) (Berg 1949b; Pinchuk 1978, 1981, 1984, 1992; Lindberg and Krasyukova 1975; Sahara 1990; Nikoforov et al. 1994; Reshetnikov et al. 1997). IKIP records: Iturup, Kunashir, Shikotan. 77. GpmnQgobius n. sp. Stevenson, in preparation; Shima-ukigori (euryhaline, but usually in freshwater): Hokkaido, southern Sakhalin (Akihito 1984; Pinchuk 1992; Nikoforov et al. 1994; Stevenson 2ooO and in preparation). IKIP records: Kunashir. Comments. This species was first recognized by Nakanishi (1978a, 1978b) as the "middle reach type" of Chaenogobius annutaris. Subsequent authors have "middle referred to it as the reach type" of C annularis (Ishino et al. 1983), as the "middle reach type" of a urotaenia (Matsumoto et al. 1988), as Chaenogob- ius "sp. 1" (Akihito et al. 1984; Pinchuk 1992; Nikoforov et al. 1994), or as

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146 Theodore W. Pietsch et al.

"sp. Chaenagobius MR" (Jeon et al. 1997). Prior to 1978, material of this as yet undescribed species was prebably most often misidentified as ChaenQgobius urotaenia (=bymnogobius urotaenia; see Stevenson 2000 and in preparation). 78. (lymnogobius laevis (Steindachner, 1879); Juzukakehaze (euryhaline): Hokkaido, southern Sakhalin (Pinchuk 1978; Akihito 1984; Kishi 1990). IKIP records: Polonskogo, Zelionyi, Tanfilyeva. 79. ([lymncrgobitts macr(rgnathos (Bleeker, 1860); Edohaze (euryhaline): Hokkaido (Iwata 1990b). Comments. Published records of this species from Sakhalin (Berg 1949b; Pinchuk 1978; Lindberg and Krasyukova 1975; Reshetnikov et at, 1997) are all based on specimens of Clymnogobius urotaenia (see below). 80. Gymnogobius mororanus (Jordan and Snyder, 1901); Hebihaze (euryhaline): Hokkaido, Shikotan (Notoro Bay) (Pinchuk 1978; Akihito 1984; Lindberg and Krasyukova 1975). 81. (lymnogobius urotaenia (Hilgendorf, 1879); Ukigori (euryhaline, but usually in freshwater): Hokkaido, southern Sakhalin, Kunashir, Iturup (Berg 1949b; Lindberg and Krasyukova 1975; Pinchuk 1978, 1984; Ishino 1990; Nikofbrov et al. 1994; Reshetnikov et at. 1997). IKIP records: Immp, Kunashir, Shikotan, Zelionyi, Tanfilyeva,

49. GenusLeucopsarionHilgendorf 82. LeucQpsarion petersii Hilgendorf; 1880; Shiro-uo (euryhaline): Hokkaido (Dotsu 1990).

50. GenusLaciogobiusGill 83, Luciogobius guttatus Gill, 1859; Flat-headed Goby, Mimizuhaze (euryhaline): Hokkaido, Kunashir, Shikotan (Berg 1949b; Pinchuk 1978; Lindberg and Krasyukova 1987; Reshetnikov et al. 1997; Aizawa 2000). IKIP records: Kunashir.

51. GenusRhinogobiusGill 84. Rhincrgobius brunneus (Temminck and Schlegel, 1845); Amur Goby, Yoshino- bori (usually in freshwater): Hokkaido, Sakhalin (Berg 1949b; Pinchuk 1978, 1992; Hayashi 1984; Lindberg and Krasyukova 1975; Nikofbrov et al, 1994; Reshetnikov et al. 1997).

52. GenusTridentigerGill 85. Trictentiger bijkesciatus Steindachner, 1881; Shimofuri-shimahaze (euryha- Iine): Hokkaido (Akihito and Sakamoto 1989; Iwata 1990b), 86. Cridentiger kuroiwae brevispinis Katsuyama, Arai and Nakamura, 1972; Numa-chichibu (euryhaline): Hokkaido, southern Sakhalin, Iturup (Zmeika River) (Berg 1949b; Kluchareva 1964; Lindberg and Krasyukova 1975; Akihito 1984; Iwata 1990b; Kishi 1990; Pinchuk 1978, 1992; Nikoforov et al, 1994; Reshet- nikov et al. 1997), IKIP records: Iturup, Kunashir. Comments. Most of the records cited above are based on specimens misiden- tified as Tridentiger obscurus (Temminck and Schlegel, 1845), a species that apparently does not extend farther north than northern Honshu (see Akihito 1984). 87. b"identiger trigonocephalzts (GM, 1859); Shimahaze (euryhaline): Hokkaido

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(Akihito 1984).

XIX. FamilyPleuronectidae 53. Genus Kareius Jordan and Snyder 88. Kttreius bicoloratus (Basilewsky, 1855); Ishigarei (euryhaline): Hokkaido, Sakhalin, Kuril Islands (Sakamoto 2000).

54. GenusLiopsettaGil1 89. Liqpsetta glacialis (Pallas, 1776); Arctic Flounder (euryhaline): Kamchatka (Berg 1949b; Reshetnikov et al. 1997; S. Shedko, personal communication, 15 September 2000). 90. LiQpsetta pinnij;asciatus (Kner, in Steindachner and Kner, 1870); Fleunder, TOgarei (euryhaline): Hokkaido, southern Sakhalin, Kuril Islands, Kam- chatka (Reshetnikov et at. 1997; Sakamoto 2000).

55. GenusPlatichtlt),sGirard 91. Platichttrys stellatus (Pallas, 1788); Starr'y Flounder, Numagarei (marine, but often entering mouths of rivers): Hokkaido, Sakhalin, Kamchatka (Nikofbrov et al. 1994; Reshetnikov et al, 1997; Sakamoto 2ooO). IKIP records: Shumshu, Paramushir, Iturup.

XX. FamilyTetraodontidae 56. GenusTakijUguAbe 92. lakijltgu poecilonotus (Temminck and Schlegel, 1850); Komon-fugu (euryha- line): Hokkaido (Matsuura 2ooO>,

Geologic History of the Kuri1 Islands

The Kuril Archipelago (Fig. 1) represents the central portion of the Kuril-Kam- chatka Island Arc formation, which includes eastern Hokkaido, the Kuril Islands, and southern Kamchatka. It is composed of two main ridges: the Lesser Kuril Ridge and the Greater Kuril Ridge. The Lesser Kuril Ridge includes the Nemuro Peninsula of eastern Hokkaido, the Habomai Island group, and Shikotan, and it continues to the northeast as the submarine Vityaz Ridge. The Greater Kuril Ridge includes the Shiretoko Peninsula of eastern Hokkaido, all of the remaining Kuril Islands from Kunashir north to Shumshu, and the southern tip of the Kamchatka Peninsula. The formation of the Kuril Archipelago apparently began in the Late Creta- ceous (approximately 90 MYBP) when the Okhotsk Terrane of the Kula Plate col- lided with the Siberian continent, thereby creating a subduetion zone along the southeast margin of the Okhotsk Terrane (Ktmura and Tamaki 1985). This subduc- tion zone initiated the formation of the Kuril-Kamchatka Trench and the subse- quent volcanism that created the Academy of Sciences Rise (now located in the central Sea of Okhotsk) and the Lesser Kuril Ridge. Volcanic activity and uplift in the region of the Lesser Kuril Ridge intensified during the Paleocene and Eocene as the Kula-Pacific Ridge was subducted into the Kuril-Kamchatka Trench, and it was probably during this period that the Lesser Kuril Ridge emerged from the sea (Kimura and Tamaki 1985). After the subduction of the Kula-Pacific Ridge, a vol-

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148 Theodore W. Pietsch et al.

canic hiatus ensued, and there is no evidence of further volcanic activity in the

Lesser Kuril Ridge. During the hiatus of the Late Eocene and Oligocene, the

Okhotsk Plate was subsiding, and there is evidence that the Lesser Kuril Ridge may have been submerged during part of this period (Kimura and Tamaki 1985). Late in the Oligocene (approxtmately 30 MYBP) the Okhotsk Terrane began to rotate clockwise and the back-arc basin that now forms the southern Sea of

Okhotsk began to open to the west of the Kuril Arc. Volcanic activity resumed near the Kuril-Kamchatka Trench during this period, but was concentrated in the location of what is now the Greater Kuril Ridge. Consequently the oldest rocks in

the main arc of the Kuril Islands are of Late Oligocene and Early Miocene age (Markhinin 1968; Markov and Khotin 1973; Savostin et al. 1983), The back-arc bastn was fu11y formed by the middle Miocene (Kimura and Tamaki 1985). Although this period marks the beginning of the formation of the primary chain of the present- day Kuril Islands, sediment records indicate that they probably did not emerge above the sea surface until the Early Pliocene (Kimura and Tamaki 1985). During the past 10 million years, the Greater Kuril Ridge has experienced intense volcanic activity and crustal uplift (Markhinin 1968; Yakushko and Nikonov 1983; Gnibidenko 1985). Although most of the islands along this ridge have not been studied closely, there is good evidence that the southern Kuril Islands (Kunashir and Iturup) emerged from the sea during the Pliocene or Early Pleistocene and have been above sea level ever since

and southern end of the Archipelago are relatively shallow. Hokkaido, Kunashir, Shikotan, and the Habomai Island group are presently separated by straits far less than 100m deep, as are Kamchatka, Shumshu, and Paramushir. Depths of no greater than 230m separate Iturup and Kunashir. Therefore, it is highly likely that these southernmost and northernmost islands were connected with each other and to nearby Hokkaido and Kamchatka, respectively, during the sea-level minima of the Late Pleistocene. Additionally, Hokkaido would have been connected to

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Fishes of the Kuril Islands 149

Sakhaiin and to mainland Asia during these periods (Kryvolutskaya 1973). Be- cause the last of these major sea-level regressions occurred 10,OOO-30,ooOyears ago, these islands have probably not been completely isolated for very long. Glaciers covered the northern and central islands during the glacial maxima of the Pleis- tocene, but probably did not extend any farther south than central Iturup, as there are no traces of glaciation on Kunashir or in the Ijesser Kuril Islands (Kryvolut- skaya 1973). In summary, the islands of the Lesser Kuril Ridge were fbmied during the Late Cretaceous and Paleocene, while the islands of the Greater Kuril Ridge began to form during the Late Oligocene and Miocene. The volcanic activity that gave rise

to the Lesser Kuril Ridge ceased during the Eocene, although the elevation of these

islands continues to change due to eustatic crustal movements. The volcanic activ- ity that gave rise to the Greater Kuril Ridge is still in progress, and the elevation of these islands centinues to change with new eruptions and eustatic crustal move-

ments. There is evidence that the Lesser Kuril Ridge emerged from the sea during

the Paleocene or Eocene, but that it has been submerged one or more times since

then. The Greater Kuril Ridge emerged during the Early Pliocene. Some of the Kurll Islands have been above sea level since that time, while others have been formed by more recent volcanic activity. During the glacial sea-level regressions of the Late Pleistocene, Kunashir, the islands of the Lesser Kuril Ridge, and perhaps Iturup were connected to each other and to Hokkaido; and at the northern end, Shumshu and Paramushir were connected to each other and to the Kamchatka Peninsula. At the time of these regressions, glaciers covered most of the islands of the Archipelago, advancing as far south as central Iturup.

Zoogeography

The Kuril Islands are volcanic in origin and therefore expected to be inhabited

by fishes that are salt-tolerant, either anadromous or euryhaline. This is true even for families such as the Cyprinidae (genus Tribolodon), which is usually thought to contain only primary freshwater species. An analysis of the distribution of fishes as recorded by us shows that species diversity falls off abruptly as one moves cen- trally from both the northern and southern ends of the Archipelago, resulting in a complete absence of species on most of the smaller central islands (i.e., Khartmkotan, Chirinkotan, Ekarma, Lovushki, Raikoke, Matua, Ushishir, and Ketoi; see Fig. 2). Setting aside for the moment the islands of the Lesser Kuril Ridge (Shikotan and the isles of the Habomai group), the number of species recorded from Hokkaido and Sakhalin is the same, 64 species, but this number

drops to 21 on Kunashir, the southernmost island of the Greater Kuril Ridge. This large reduction in biodiversity occurs despite the close proximity of Hokkaido and Kunashir (the minimum distance from the tip of Cape Veslo on Kunashir to No- tsuke on the east coast of Hokkaido, a long curved sand-spit, is 17.9km; from Cape Veslo to the nearest solid land on Hokkaide is 26.8km) and the probable connec- tion between these two landrnasses in the relatively recent geological past (i.e., during the sea-level minima of the Late Pleistocene). Similarly, Kamchatka sup- ports at least 33 species, but this number drops to eight and nine on Shumshu and Paramushir, respectively, despite the close physical proximity of these islands to

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150 Theodore W Pietsch et al

IaindiIttktautIIlIUttlHtlgt KMsuPAONSARASIURITKUiysgefi.xq,ZXk,IXI,"ec,:TA,HOSK KM 892112711946'"P.erl/,:rali,:3v,'Ii･･g･2422 et l'"-t/- gM/le//-//llet/S/lt su87 72112Jr86IdeiS!3,ieile,g.88 tt -ttt F --- t4H/'e PA9876 2112697s233''i'ettljit99 .4/Wlt/:Ig '- g ll ON222222 111121,ie:iiginelns9:i,p{,'iaji:2z --t"#Jl frSA11111111 11111ab"oaise11

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' SK239879822rtll227619619611913365la31165478

Fig 2 Numbers of freshwater fish species shared by islands of the Kuril Archipelago (based on IKIP records) and adjacent regions (based on available hterature), arranged from north to south Kamchatka (KM), Shumshu (SU), Paramushir (PA), Onekotan (0N), Shiashkotan (SA), Rasshua (RA), Simushir (SI), Urup (UR), Iturup (IT), Kunashir (KU), Hokkaido (HO), and Sakhalm (SK) The 1ightly shaded area encloses islands of the Lesser Kurd Ridge Shikotan (SH), Polonskogo (PO) Zeliony! (ZE) Iurli (IU), Anuchma (AU), and Tanfilyeva (TA) Num

bers on the upper right are absolute values, those on the lower left are percentages of the

total number af species (92) recorded from the combined region Note that the subspecies of Pungttius are counted as separate species

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Fishes of the Kuril Islands 151

the mainland and past geological connections (the distance between the tips of Cape Pochtareva on Shumshu and Cape Lopatka on Kamchatka is 11.2km; the minimum width of the Upper Kurilskii Strait between Shumshu and Paramushir is only 1.7km). The abrupt decline in fish diversity at both ends of the chain, as

compared to that of Kamchatka and Hokkaido, is, therefore, not only the result of geological history or distance from these large adjacent source biotas, but more likely due to the great differences in area and therefore habitat available to fishes: in the north, Kamchatka encompasses some 270,033km2, compared to only 388 and 2,053km2 for Shumshu and Paramushir, respectively; in the south, Hokkaido cov- ers 78,084 km2, compared to only 1,490 km2 for Kunashir,

Diversity appears to be size-dependent on the islands of the Lesser Kuril Ridge as well, but it also may be partially due to past conneetions to Hokkaido, Kunashir, and perhaps Iturup. Shikotan (250km2) supports at least 11 species of freshwater fishes, all of which are present on Hokkaido, and all but one (I(Ypomesus japonicus) of which was collected by us on Kunashir; all but two (ftvpomesus joponicus and Pungitius tymensis) appear to be present on Iturup. The isles of the Habomai group each support smaller subsets of the assemblage cornmon to Hokkaido, Kunashir, and Iturup (Fig. 2). The two largest of these isles, Zelionyi (51 km2) and Tanfilyeva (13km2), share at least six species: an osmerid (llypomesus chishimaensis), two gob- iids (Gymncrgobius laevis and Gl,mnctgobius urotaenia), and three gasterosteids. In addition, a cyprinid (Tribolodon ezoe) was collected on Zelionyi, anct a salmonid (Salvelinus leucomaenis) on Tanfilyeva. Polonskogo (11km2) and Iurii (10km2) share at least two species, both ubiquitous gasterosteids, In addition, a gobiid (Gymnogobius laevis) was collected on Polonskogo, and another gasterosteid (Gas- terosteus aculeatus aculeatus) on Iurii. Only the gasterosteid Pungititts pungitius pungitius was collected on Anuchina (2.3 km2), Moving along the chain centrally, away from Kamchatka in the north and Hokkaido in the south, the islands become progressively smaller in area. This re- duction in size, as well as the increasing distance from major source biotas, corre- lates well with a dramatic decrease in fish diversity. From eight and nine species on Shumshu and Paramushir, respectively, the number drops to two on Onekotan and then to one on Shiashkotan and Rasshua, only to increase to two on Simushir and seven on Urup before the relatively high diversity of Iturup (19 species) and Kunashir (21) is reached in the south. The relatively low diversity fbund on Urup as compared to that on Iturup and Kunashir may be size-dependent (450km2, about one-seventh the size of Iturup and a third the size of Kunashir) but could also be due in part to Urup's physical isolation and recent glacial perturbation. Iee cov- ered nearly all of the Kurils during the Late Pleistocene, including Urup but not the southern half of Iturup and Kunashir, at a time when Kunashir and perhaps Iturup as well were connected to Hokkaido (Urup has always been well separated from Iturup; the strait between them today is greater than 800m deep). The close relationship between Hokkaido, Kunashir, and Iturup, and to some extent Shikotan and the larger isles of the Habomai group (Zelionyi and Tanfi- lyeva), is also reflected in species composition. Members of four families and five genera of fireshwater fishes are shared by Hokkaido, Kunashir, and Iturup, to the exclusion of all islands to the north: Petromyzontidae (genus Lainpetra), Cyprinidae (Tribolodon), Cottidae (Cottus), and Gobiidae ((lymnogobius and T)'i- dentiger). Cottus and (lymnogobius are also fbund on Shikotan, while (lymnQgobius

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152 [ilheodore W. Pietsch et al,

is present on Zelionyi and Tanfilyeva. in contrast to the southern end of the chain, there are no sharp faunal breaks among freshwater fishes in the north, The eight and nine species found on

Shumshu and Paramushir, respectively, are found largely throughout the Archi- pelago, as well as on Kamchatka, Hokkaido, and Sakhalin: the anadromous salmonid genera Oncorhynchus and Salvetinus, the euryhaline gasterosteid genera Gasterosteus and Pungitius, and the largely marine pleuronectid Platichtbys stella- tus. The much greater influence from the south is no doubt due to the considerably higher species diversity of Hokkaido, which is in turn the result of a temperate cli- mate (in contrast to subarctic Kamchatka) and immediate connections to even greater species diversity to the south and west.

Acknowledgments

The work described here was supported in part by the International Programs Division and the Biological Sciences Directorate (Biotic Surveys and Inventories Program) of the U.S. National Science Foundation, Grant Nos. DEB-94oo821 and DEB-9505031, Theodore W. Pietsch, principal investigater; the Japan Society for the Promotion of Science, Grant No. BSAR-401, Kunio Amaoka, principal investi- gator; and the Russian Academy of Sciences, Far East Branch, Viktor V. Bogatov, principal investigator. In addition to our great appreciation for the generous fund- ing received from these agencies, we thank the many people who assisted in col- lecting efforts over the past six summers, especially Andrei Balanov, Cynthia A. Catton, Theresa J. Landewe, Wayne A. Palsson, Dmitriy L. Pitruk, Sergei Shedko,

and Mamoru Yabe. We also thank Viktor V. Bogatov, Deputy Chief Academic Sec- retary and Chief of Scientific Investigations, Russian Academy of Sciences Far East Branch, Vladivostok, and Valentina A. Kolesnikova, Assistant Director of In- temational Relations, Institute of Biology and Soil Sciences, Russian Academy of Sciences Far East Branch, Vladivostok, for making the Kuril Island Project possi- ble. Our great appreciation also goes to George B. Elyakov, Vice-President of the Russian Academy of Sciences and Chairman of the Far East Branch, Vladivostok, for his support. Finally, we thank Captain Evgeni N. Marenin and the crews of the R/V Projbssor Bogorov and RfV Akademik Ciparin for taking good care of us while at sea.

Appendix: Material Examined

In the foliowing list of material examined, the island locality (in bold font) is followed by the University of Washington (UW) register number of the particular lot, the number of spec- imens in the lot, the standard length of the specimen (or the range of standard lengths if there is more than one specimen in the lot) in millimeters, and the IKIP field number. Each IKIP

field number consists of four parts: an abbreviation for the particular island, the year the col- lection was made, the initials of the collector, and the station number; fbr example, IT-94

TWP-oo2 means that the specimens in that particular lot were collected on Iturup Island in

1994 by T. W, Pietsch at station O02. In a few cases in which captured specimens were excep-

tional large, only tissue samples were retained. For a fuIl-text search intenhce to locality and

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Fishes of the Kuril Islands 153

taxonomic databases, see the IKIP website accessible via . Cottus amblystomopsis: Iturup (UW 029135, 93, 14 118mm, IT-94-TWP-O02; UW 029193, 19, 71-118mm, IT-94-TWP-O06I UW 029142, 225, 12-35 mm, IT-94-TWP-Oll; UW 029146, 20, 49-110

mm, IT-94-TWI]-O15; UW 029234, 1, 96mm, IT-94-TWP-O16; UW 029242, 18, 55-124mm, IT-94-

TWI)-020; UW 029162, 37, 15-153mm, IT-94-TWP-026; UW 040507, 1, 45mm, IT-98-DJB-060; UW

042196, 3, 84-113mm, IT-98-ELM-O09; UW 042198, 1, 113mm, IT-98-ELM-O15; UW 0405e2, 16, 44-131mm, IT-98-ELM-027; UW 041363, 6, 27-178mm, IT-99-DES-028; UW 043551, 16, 67-120mm, IT-99-DES-022; UW 043599, 12, 66-129mm, IT-99-DES-025), Kunashir (UW 029267, 8, 40-45mm, KU-94-TWP-O05; UW 029268, 5, 56--102mm, KU-94-TWP-O06; UW 029273, 8, 49-106mm, KU-94- TWP-O08; UW 029282, 1, 73mm, KU-94-TWP-O14; UW 029285. 6, 105--110 mm, KU-94-TWP-O15; UW

029287, 1, 95mm, KU-94-TWP-O16; UW 029294, 23, 29-79mm, KU-94-TWP-O19; UW 029298, 16, 39-119mm, KU-94-TWP-020; UW 029299, 3, 43-67mm, KU-94-TWP-021; UW 040108, 2, 82-92mm, KU-95-TWP-O07; UW 040506, 1, 64mm, KU-98-ELM-040; UW 043554, 1, 25mm, KU-99-DES-029), Shikotan (UW 029173, 52, 9-104mm, SH-94-TWP-OOI; UW 029312, 21, 28-96mm, SH-94-TWP-Oe3; UW e29174, 89, 12-111 mm, SH-94-TWP-O09; UW 040505, 4, 47-49 mm, SH-98-ELM-044; UW 040501,

2, 48-54 mm, SH-98-LJW-050; UW 040503, 19, 25-113 mm, SH-98-TJL-O05; UW 042199, 8, 39-84mm, SH-98-TJL-O08; UW 040508, 4, 44-74mm, SH-98-TJL-O09; UW 040500, 1, 101mm, SH-98-TJL-Oll;

UW 040504, 28, 38-52mm, SH-98-TJL-O12; UW 042197, 5, 41-tl7 mm, SH-98-TJL-O16). Cottus hangiongensis: Kunashir (UW 040199, 9, 26-57mm, KU-95-JAIrO08; UW 042249, 1,

1oo mm, KU-97-DES-O04). Gasterosteus aculeatus aculeatus: Iturup (UW 029188, 331, 12-87mm, IT-94-TWI'-OOI; UW 029136, 31, 14-84mm, IT-94-TWP-oo2; UW 029139, 78, 11-71mm, IT-94-TWP-OIO; UW 029196,

14, 15-79mm, IT-94-TWP-Oll; UW 029143, 20, 13-76 mm, IT-94-TWP-O13; UW 029230, 7, 16777mm, IT-94-TWP-O14; UW 02914S, 187, 17-75 mm, IT-94-Twn-O18; UW 029154, 30, 13r85mm, IT-94-TWP- 022; UW 029158, 358, 12m84mm, IT-94-TWP-025; UW 029249, 27, 18-75mm, IT-94-TWP-026; UW 042044, 163, 12-72mm, IT-9&JAL-029; UW 041S82, 1, 81mm, IT-98-AVR-033; UW 041874, 10, 28-65 mm, IT-98-ELM-O12; UW 041880, 6, 45-81mm, IT-98-ELM-O14; UW 041895, 35, 53-83mm, IT-98- ELM-O16; UW 041872, 3, 60-67mrn, IT-98-ELM-028; UW 041887, 1, 80mm, IT-98-ELM-031; UW 041898, 461, 21-81mm, IT-98-ELM.037; UW 041885, 1, 19 mm, IT-98-LJW-069; UW 041360, 19, 43-62 mm, IT-99DES-022; UW 041364, 33, 15-79mm, IT-99-DES-028; UW 043582, 89, 24-72mm, IT-99- DES-021; UW 043602, 19, 39-64mm, IT-99-DES-025; UW 043606, 24, 19-74mm, IT-99-DES-023; UW 043624, 43, 12-82mm, IT-99-DES-027; UW 043703, 4, 12--15mm, IT-99-KLK-044; UW 043723, 291, 18-82 mm, IT-99-DES-026), Iurii (UW 041888, 1, 72 mm, IU-98-TJL-026), Kunashir (UW 029304, 1, 11mm, KU-94-JBB-O12; UW 029165, 16, 14-26mm, KU-94-TWP-OOI; UW 029257, 67, 11-68mm, KU- 94-TWP-oo2; UW 029259, 4, 35-65mm, KU-94-TWP-O03; UW 029260, 15, 17-79mm, KU-94-TWP- O04; UW 029279, 1, 60 mm, KU-94-TWP-O13; UW 029293, 3, 47-88 mm, KU-94-TWP.O19; UW 040192,

95, 20-53mm, KU-95-JAL-O07B; UW 040196, 11, 27-75mm, KU-95-JAL-O08; UW 042041, 33, 18-31 mm, KU-98-JAL-042; UW 041889, 1, 19mm, KU-98-ELM-oo5; UW 041884, 32, 22-62mm, KU-98- ELM-039; UW 041871, 11, 51-76mm, KU-98-ELM-040; UW 04187e, 7, 20-86mm, KU-98-TJL-038;

UW 041879, 1, 72mm, KU-98-TJL-039; UW 041876, 4, 21-27mm, KU-98-TJL-040; UW 041890, 3, 73-83mm, KU-98-TJL-041), Paramushir (UW 042025, 6, 75-94mm, PA-96-JAL-O05; UW 042022, 2, 82-83mm, PA-96-JAL-O06; UW 042028, 10, 68-94mm, PA-96-JAL-O07; UW 042222, 97, 38-62mm, PA-97-DES-e15; UW 042247, 10, 75-90mm, PA-97-DES-051; UW 042251, 5, 78-87mm, PA-97-TWP- 037; UW 043649, 3, 71-85mm, PA-99-CAC-O14), Shikotan (UW 029322, 2, 14-16mm, SH-94-EMS- O09; UW 029309, 30, 70-81mm, SH-94-TWP-O03; UW 029318, 2, 18mm, SH-94-TWP-OIO; UW 041875, 4, 51-77mm, SH-98-ELM-041; UW 041886, 2, 70-79mm, SH-98-TJL-O04; UW 041873, 27,

25-90 mm, SH-98-TJL-O17; UW 041892, 29, 18-76mm, SH-98-TJL-O19; UW 041881, 1, 82mm, SH-98-

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TJL-020), Shumshu (UW 042235, 4, 70-81mm, SU-97-DES-021; UW 042220, 5, 74-82mm, SU-97- DES-022i UW 042239, 4, 73-83mm, SU-97-DES-024; UW 042265, 4, 20-80mm, SU-97-TWP-028; UW

043557, 61, 67-86mm, SU-99-CAC-O19; UW 043664, 12, 65-87mm, SU-99-CAC-O17i UW 043709, 24, 63-83 mm, SU-99-CAC-O18), Simushir (UW 040209, 11, 10-14 mm, SI-95-PO-046; UW 040165, 84, 70-76mm, SI-95-TWP-032; UW 043647, 16, 51-70mm, SI-99-DES-O15), Tanfilyeva (UW 041877, 6, 44-81mm, TA-98-TJL-e23; UW 041893. 10, 71 77mm, TA-98-TJL-e24; UW 041883, 32, 41-82mm, TA-98-TJL-025), Vrup (UW 040145, 13, 18-76mm, UR-95-TWP-024; UW 040212, 69, 17 83 mm, UR- 95-TWP-055; UW 042011, 8, 20 48 mm, UR-96-JAL-025), Zelionyi (UW 029186, 109, 14-82mm, ZE- 94-TWP-OOI; UW 029177, 66, 11-22mm, ZE-94-TWP-Oe2; UW 029328, 1, 16mm, ZE-94-TWP-O05;

UW 029330, 2, 15mm, ZE-94-TWP-oo6; UW 029333, 210, 13-37mm, ZE-94-TWI)-O07; UW 041894, 17,

21-83mm, ZE-98-DES-O14; UW 041878, 8, 31-79mm, ZE-98-DES-O15; UW 041891, 19, 40-77mm, ZE-98-DES-O16), Kamchatka (UW 043667, 3, 65-78mm, KM-99-CAC-Oll). Gymnogobius n. sp.: Kunashir (UW 029290, 110, 48-77mm, KU-94-TWP-O17; UW 040195, 292, 17 80mm, KU-95-JAL-O08i UW 040100, 7, 49-80mm, KU-95-TWP-OOIi UW 040101, 7, 47-60

mm, KU-95-TWI]-O02; UW 040103, 10, 45-55mm, KU-95-TWP-O04; UW 040104, 23, 48-56mm, KU- 95-TWP-O05; UW 040106, 11, 42-61mm, KU-95-TWP-O061 UW 040109, 4, 50-71mm, KU-95-TWP-

O07; UW 040111, 11, 50-70mm, KU-95-TWP-O08; UW 040113, 20, 53-68mm, KU-95-TWP-O09; UW 040115, 10, 46-77mm, KU-95-TWP-OIO; UW 040116, 17, 44-71mm, KU-95-TWP-Oll; UW 040118, 10,

56-68mm, KU-95-TWP-O12; UW 042ooO, 9, 51-70mm, KU-96-JAL-033i UW 042049, 20, 54-71mm, KU-96-JAL-034; UW 042036, 17, 48-70mm, KU-96-JAL-035i UW 042034, 9, 48-75mm, KU-96-JAL- 036; UW 0420e7, 16, 52 77mm. KU-96-JAL-037; UW 042oo9, 24, 5U-76mm, KU-96-JAL-038; UW

042001, 8, 51-70mm, KU-96-JAL-039; UW 042045, 6, 55-67mm, KU-96-JAL-040; UW 042003, 11, 56- 72mm, KU-96-JAL-041; UW 042223, 17, 45-67mm, KU-97-DESOOI; UW 042226, 1, 64mm, KU-97-

DES-O02; UW 042225, 8, 52-71 mrn, KU-97-DES-O03; UW 042229, 7, 47-70mm, KU-97-DES-O04; UW 040532, 20, 45 76mm, KU-98-ELM-OOII UW 040512, 14, 51-77mm, KU-98-ELM-O02; UW 041387, 8,

44-72mm, KU-99-DES-032; UW 043542, 69, 42 75mm, KU-99-DES-030; UW e43553, 67, 47-79mm,

KU-99-DES-029; UW 043701, 2, 54-65mm, KU-99-NM-O17). (lymnogobius castaneas: Iturup (UW 029160, 1401, 14-63mm, IT-94-TWP-025; UW

029163, 15, 31-49mm, IT-94-TWP-026; UW 043545, 1, 47mm, IT-99-DES-022; UW 043604, 4, 41-48

mm, IT-99-DES-025; UW 043605, 32, 38 55mm, IT-99-DES-024; UW 043613, l2, 35.49mm, IT-99- DES-021; UW 043716, 710, 34-51mm, IT-99-DES-023), Kunashir (UW 029281, 68, 2an6mm, KU- 94-TWI)-O13i UW 040524, 502, 22-46mm, KU-98-TJL-038; UW 040523, 15, 31--48mm, KU-98-TJL- 039; UW 040534, 62, 29-43mm, KU-98-TJL-040i UW 040533, 390, 23-46mm, KU-98-TJL-041; UW

043543, 6, 28 31mm. KU-99-DES-030), Shikotan (UW 029324, 3, 27-30mm, SH-94-EMS-oo9; UW

029175, 109, 22-35mm, SH-94-TWP-oo9; UW 029319, 302, 12- 32mm, SH-94-TWP-OIO; UW 040525,

429, 19-58mm, SH-98-TJL-O06; UW 040527, 21, 41-56mm, SH-98-TJL-O17; UW 040517, 1, 51mm, SH-98-TJL-O19).

(lymnogobius laevis: Polonskogo (UW 040531, 10, 17-27mm, PO-98-TJL-029>, Tanfi- lyeva (UW 040522, 602, 28 62 mm, TA-98-TJL-023; UW 040530, 15, 37-49mm, TA-98-TJL-024; UW --52 040520, 9, 35 mm, TA-98-TJL-025), Zelionyi (UW 029325, 211, 22-62mm, ZE-94-TWP-OOI; UW

029179, 53, 9-40 mm, ZE-94-TWP-O02; UW 029183, 580, 20-56mm, ZE-94-TWP-O05; UW 029332, 257,

19-45mm, ZE-94-TWP-O06; UW 029184, 202, 19-50mm, ZE-94-TWP-O07; UW 040526, 35, 21 51 mm,

ZE-98-DES-O14; UW 040521, 501, 22-60mm, ZE-98-DES-O15; UW 040519, 49, 30.52mm, ZE-98-DES- O16).

Gymnogobius urotaenia: Iturup (UW 029190, 2, 55-61mm, IT-94-TWP-OOI; UW 029141, 32, 25-112mm, IT-94-TWP-OIO; UW 029232, 3, 50-51mm, IT-94-TWP-O141 UW 029150, 184, 35-71

mm, IT-94-TWP-O18; UW 028208, 4, 66-83mm, IT-94-TWI)-026i UW 040509, 49, 43-96mm, IT-98-

ELM-O16; UW 040528, 36, 36-46mm, IT-98-ELM-O17; UW 043546, 1, 57mm, IT-99-DES-022; UW

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043600, 36, 56 105mm, IT-99-DES-025; UW 043710, 78, 49-95mm, IT-99-DES-023), Kunashir (UW

029254, 5, 39-93mm, KU-94-TWP-OOI; UW 029169, 5, 48-54mm, KU-94-TWI)-O031 UW 029262, 1, 67

mm, KU-94-TWP-Oe4; UW 029172, 2663, 18-40mm, KU-94-TW?-024; UW 029303, 78, 23-:l3mm,

KU-94-TWP-025; UW 040194, 4, 17-28mm, KU-95-JAL-O07B; UW 042042, 11, 21 100mm, KU-96-

JAL-042; UW 040529, 13, 25-92mm, KU-98-ELM-039; UW 040514, 3, 64-92mm, KU-98-ELM-040; UW 040513, 1, 48mm, KU-98-TJL-038; UW 0437oo, 2, 47-75mm, KU-99-NM-O17), Shikotan (UW

029311, 1, 94mm, SH-94-TWP-O03i UW 027488, 1, 44mm, SH-94-TWP-O091 UW 040515, 4, 82-97

mm, SH-98-TJL-O08; UW 040516, 2, 43-46mm, SH-98-TJL-O17; UW 040536, 2, 80-98mm, SH-98- TJL-O17), Tanfilyeva (UW 040518, 71, 60-117mm, TA-98-TJL-023; UW 040511, 9, 89-122 mm, TA- 98-TJL-024; UW 040510, 23, 51-126mm, TA-98-TJL-025), Zelionyi (UW 027494, 8, 50-86mm, ZE- 94-TWP-OOI). flrypomesus chishintaensis: Iturup (UW 029228, 6, 24-92mm, IT-94-TWP-Oll; UW 029239, 4, 48-67mm, IT-94-TWPO18; UW 029161, 64, 45-90 mm, IT-94-TWP-025; UW 029253, 5, 60-79 mm,

IT-94-TWP-026; UW 041860, 2, 93-96mm, IT-98-ELM-O12i UW 041858, 1, 85mm, IT-98-ET.M-O13;

UW 041861, 370, 47-82mm, IT-98-ELM-O16; UW 041367, 1, 75mm, IT-99-DES-028; UW 043603, 7,

71-81mm, IT.99-DES-025: UW 043612, 1, 64mm, IT-99-DES-024; UW 043719, 244, 51-100mm, IT- 99-DES-021), Kunashir (UW 029256, 8, 53-78mrn, KU-94-TWP-OOI; UW 041700, 1, 64mm, KU-94- TWP-oo1; UW 029264, 1, 70mm, KU-94-TWP-O041 UW 040190, 6, 49-75mm, KU-95-JAL-O07B; UW

041859, 28, 65 89mm, KU-98-ELM-e39; UW 041866, 5, 38-48mm, KU-98-TJL-038; UW 041862, 322, 41-121mm, KU-98-TJL-039; UW 041863, 67, 35-48mm, KU-98-TJL-0401 UW 041868, 3, 37--42mm, KU-98-TJL-e41; UW 043697, 7, 66 99 mm, KU-99-NM-O17), Tanfilyeva (UW 041865, 9, 41-102 mm, TA-98-TJL-024; UW 041864, 5, 60-103mm, TA-98-TJL-025), Zelionyi (UW 029327, 696, 20-64mm, ZE-94-TWP-OOI; UW 041867, 22, 29-80mm, ZE-98-DES-O14; UW 041869, 392, 28-129mm, ZE-98- DES-O16). dvpomesusjaponicus: Shikotan (UW 041857, 9, 74-136 mm, SH-98-TJL-O15), Hlypomesus olidus: Kamchatka (UW 043724, 749, 61-104mm, KM-99-CAC-Oll), Lampetra joponicum: Iturup (UW 029244, 1, 149 mm, IT-94-TWP-021). Lampetra kessleri: Iturup (UW 029308, 1, 85 mm, IT-94-RG-O08; UW 029191, 7, 12-112mm, IT-94-TWP-O02; UW 029241, 4, 85-139mm, ]rl"-94-TWP-O19; UW 029246, 3, 31 -165 mrn, IT-94-TWP-

022; UW 042494, 54, 35-196mm, IT-98-ELM-032; UW 041365, 10, 53-132mm, IT-99-DES-028), Ku-

nashir (UW 029284, 1, l35 mm, KU-94-TWP-O14). Luciogobius guttatus: Kunashir (UW 043544, 3, 42-53 mm, KU-99-DES-e30), Oncorhynchus gorbuscha; Iturup (UW 042493, 1, 395 mm, IT-98-EIiM-O14; UW 042491, 7, 45-85mm, IT-98-ELM-031), Kunashir (UW e29266, 45, 55-74mm, KU-94-TWP-O04), Shikotan (UW 042492, 1, 415mm, SH-98-TJL-OIO), Shumshu (UW 042291, 1, 60mm, SU-97-TWP-024; UW 043623, 1, 385 mm, SU-99-DES-O03; UW 043682, 1, 55 mm, SU-99-CAC-O19), Urup (UW 040202, 1,

tissue only, UR-95-JAL-O05; UW 040142, 3, tissue only, UR-95-TWP-023; UW 040102, 2, tissue

only, UR-95-TWP-055). Oncorhynchus keta: Iturup (UW 029199, 12, 29-41mm, IT-94-TWP-Oll; UW 029151, 6, 34-40 mm, IT-94-TWP-O18; UW 029156, 14, 31-tllmm, IT-94-TWP-022), Kunashir (UW 029272, 69,

32-48mm, KU-94-TWP-O07i UW 029275, 2, 42-44mm, KU-94-TWP-oo8; UW 042404, 3, 30-37mm, KU-94-TWP-O19; UW 042403, 1, 39 mm, KU-94-TWP-e21), Paramushir (UW 042019, 79, 28--68

mm, PA-96-JAL-O06; UW 042016, 49, 28-49mm, PA-96-JAL-O07i UW 042256, 9, 40-51mm, PA-97-

DES-050; UW 042246, 31, 35-82mm, PA-97-DES-051; UW 042253, 2, 39-50mm, PA-97-TWP-e37),

Shikotan (UW 042431, 1, 30mm, SH-94-TWP-oo3). Urup (UW 040126, 10, 26-44 mm, UR-95-TWP- O16; UW 040130, 123, 29-61 mm, UR-95-TWP-O17; UW 040132, 8, 30-37mm, UR-95-TWP-O18; UW

040134, 21, 32-45 mm, UR-95-TWP-O19).

Oncorltynchus kisutch: Iturup (UW 042406, 1, 40mm, IT-94-TWP-O02), Kunashir (UW

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156 Theodore W. Pietsch et al.

042250, 92, 30-95mm, KU-97-DES-O04), Paramushir (UW 042017, 2, 52-67mm, PA-96-JAL-O07;

UW 042260, 36, 25-48mm, PA-97-DES-0331 UW 042276, 2, 80-88mm, PA-97-DES-035; UW 042274,

1, 54mm, PA-97-NM-074; UW 042252, 15, 30-58mm, PA-97-TWP-037; UW 043666, 6, 47-83mm, PA-99-CAC-O14i UW 043675, 38, 25-41mm, PA-99-CAC-O15), Shumshu (UW 042237, 10, 33-56 mm, SU-97-DES-021; UW 042238, 2, 32-37mm, SU-97-DES-022i UW 042270, 69, 28-66mm, SU-97-

DES-024; UW 043610, 190, 30-El8mm, SU-99-CAC-O17; UW 04368e, 3e, 32-66mm, SU-99-CAC-O19;

UW 043712, 80, 29-88mm, SU-99-CAC-O18), Urup (UW 040204, 6, 40-61mm, UR-95-PO-089; UW 040129, 7, 48-87mm, UR-95-TWP-O17; UW 040146, 30, 44-67mm, UR-95-TWP-024), Kamchatka

(UW 043641, 132, 31-89 mm, KM-99-CAC-Ol1). Oncorhynchus inasou masou: Kunashir (UW 029277, 1, 103 mm, KU-94-TWP-Oe9). Oncorhptnchus nerka: Iturup (UW 042430, 2, 30-45mm, IT-94-TWP-O18>, Onekotan (UW 043622, l, 442mm, ON-99-CAC-O04), Paramushir (UW 043674, 4, 39-45mm, PA-99-CAC-O15), Shumshu (UW 042266, 1, 54mm, SU-97-TWP-028; UW 043609, 65, 27-47mm, SU-99-CAC-O17; UW 043683, 33, 28-51 mm, SU-99-CAC-O19; UW 043713, 403, 28-50mm, SU-99-CAC-O18>, Kamchatka

(UW 043640, 2, 35-37 mm, KM-99-CAC-Oll). Platichthys stellatas: Iturup (UW 043598, 11, 32-49mm, IT-99-DES-021), Paramushir (UW 043671, 1, 54 mm, PA-99-CAC-O15), Shumshu (UW 043536, 1, 156 mm, SU-99-CAC-O19), Pungitius pungitius pungitius: Anuchina (UW 042178, 2, 19-47mm, AU-98-LJW-064), Iturup (UW 029144, 47, 21.56mm, IT-94-TWP-O13i UW 029231, 59, 19-53mm, IT-94-TWP-O14; UW

042424, 73, 38-72 mm, IT-94-TWP-O18; UW 029240, 1, 32mm, IT-94-TWP-O19; UW 042425, 4, 40-65

mm, IT-94-TWP-025; UW 042159, 8, 41-60mm, IT-98-ELM-O16; UW 043601, 4, 42-67mm, IT-99-

DES-025; UW 043715, 8, 44-60mm, IT-99-DES-023), Iurii (UW 042144, 6S, 23-69mm, IU-98-TJL- 026), Kunashir

UW 042170, 18, 22-36mm, KU-98-ELM-O06), Paramushir (UW 042015, 37, 18-61mm, PA-96-JAL- O03; UW 042021, 11, 35-50mm, PA-96-JAL-O06; UW 042027, 4, 47-63mm, PA-96-JAL-O07; UW

042243, 99, 35-54mm, PA-97-DES-O15; UW 042244, 69, 15-59rnm, PA-97-DES-042; UW 042248, 36, 43-66mm, PA-97-DES-051), Polonskogo (UW 042139, 41, 29-64 mm, PO-98-TJL-029; UW 042145, 71, 28-62mm, PO-98-TJL-030; UW 042158, 625, 23m64mm, PO-98-TJL-031; UW 042176, 26, 25-46

mm, PO-98-TJL-033; UW 042173, 1, 30 mm, PO-98-TJL-036; UW 042167, 1, 40mm, PO-98-TJL-037), Shikotan (UW 042157, 575, 23-65 mm, SH-98-ELM-041; UW 042168, 2, 33-51 mm, SH-98-ELM-042; UW 042194, 1, 47mm, SH-98-TJL-O05; UW 042147, 28, 35-55mm, SH-98-TJL-oo6; UW 042151, 112, 28-67mm, SH-98-TJL-O17; UW 042153, 288, 28-63mm, SH-98-TJL-O19; UW 042169, 9, 44-59rnm, SH-98-TJL-020), Shumshu (UW 042236, 1, 54mm, SU-97-DES-021; UW 042221, 210, 36-66mm, SU-97-DES-022; UW 042241, 77, 41-66mm, SU-97-DES-024; UW 042267, 15, 43-63mm, SU-97-TWP- 028; UW 043556, 20, 49.75mm, SU-99-CAC-O19; UW 043580, 161, 34-62mm, SU-99-CAC-O17; UW 043708, 314, 35-66 mm, SU-99-CAC-O18), Tanfilyeva (UW 042150, 2, 28-43 mm, TA-98-LJW-065; UW 042148, 26, 43-62mm, TA-98-TJL-023; UW 042155, 165, 24-76mm, TA-98-TJL-024; UW 041899, 111, 46-69 mm, TA-98-TJL-e25), Zelionyi (UW 029181, 48, 12-48mm, ZE-94-TWP-O03; UW 028132,

13, 37-54mm, ZE-94-TWP-O05; UW 042142, 11, 34-57mm, ZE-98-DES-O14; UW 042186, 34, 2EH56

rnm, ZE-98-DES-O15), Kamchatka (UW 043714, 30, 37-64 mm, KM-99-CAC-O07; UW 043717, 100,

33-68 mm, KM-99-CAC-Oll).

Pungitius pungitius sinensis: Iturup (UW 029189, 363, 15-64mm, IT-94-TWP-OOI; UW 029137, 161, 17-55mm, IT-94-TWP-oo2; UW 029140, 448, 35-56mm, IT-94-TWP-Ole; UW e29197, 7,

20L51mm, IT-94-TWP-Oll; UW 027243, 133, 18-62mm, IT-94-TWP-O13; UW 042429, 6, 22m53mm,

IT-94-TWP-O13; UW 026845, 157, 12-56mm, IT-94-TWP-O14; UW 042423, 401, 38-72mm, IT-94-

TWP-O18; UW 029155, 152, 15-56mm, IT-94-TWP-022; UW 029159, 191, 23-63mm, IT-94-TWP-025;

UW 029250, 32, 23-63mm, IT-94-TWP-026; UW 042043, 15, 16-52mm, IT-96-JAL-029; UW 042192,

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Fishes of the Kuril Islands 157

1, 36 mm, IT-98-DJB-022; UW 042166, 38, 28-53mm, IT-98-ELM-OIO; UW 042184, 25, 27-59mm, IT- 98-ELM-Oll; UW 042162. 334, 18-68mm, IT-98-ELM-O12; UW 042160, 671, 40 70mm, IT-98-ELM- O16; UW 042190, 10, 50-68mm, IT-98-ELM-031; UW 042180, 66, 15-6] mm, IT-98-ELM-032; UW 042161, 273, 36-63mm, IT-98-ELM-037; UW 043523, 172, 22-52mm, IT-99-DES-028; UW 043547, 9, 45-62mm, IT-99-DES-022; UW 043581, 60, 42-67mm, IT-99-DES-025; UW 043583, 5, 53r58mm, IT- 99-DES-021; UW 043611, 4, 46-62mm, IT-99-DES-024; UW 043625, 16, 16-49mm, IT-99-DES-027; UW 043702, 2, 12-19mm, IT-99-KLK-044; UW 043720, 153, 24-68mm, IT-99-DES-023; UW 043721, 433, 24-66mm, IT-99-DES-e26), Iurii (UW 042181, 5, 23-51mm, IU-98-TJL-026), Kunashir (UW 029166, 79, 18-58mm, KU-94-TWP-OOI; UW 028207, 11, 19-31mm, KU-94-TWP-O02; UW 029168, 101, 3254mm, KU-94-TWP-oo3; UW 029261, 4, 51-56mm, KU-94-TWP-O04; UW 042426, 3, 31-43 mm, KUT94-TWP-O13; UW 040191, 41, 21-45 mm, KU-95-JAL-O07B; UW 040197, 47, 35-56mm, KU- 95-JAL-O08I UW 042040, 120, 19-56mm, KU-96-JAL-042; UW 042175, 14, 22-54mm, KU-98-ELM- O04; UW 042171, 3, 25-31mm, KU-98-ELM-O06; UW 042182, 54, 18-52mm, KU-98-ELM-039; UW 042164, 441, 2062mm, KU-98-TJL-038i UW 042165, 169, 36-66mm, KU-98-TJL-039; UW 042163, 220, 25.61mm, KU-98-TJL-040; UW 042183, 31, 29-69 mm, KU-98-TJL-041; UW 043698, 1, 50mm, KU-99-NM-O17; UW 043711, 44, 23-62mm, KU-99-CAC-Oll), Paramushir (UW 042428, 8, 33-45 mm, PA-96-JAL-O06; UW 042427, 3, 39-40mm, PA-96-JAL-O07; UW 042242, 55, 38-58 mm, PA-97- DES-O15; UW e43642, 38, 29-47mm, PA-99-CAC-O14; UW 043646, 1, 51mm, PA-99-CAC-O13; UW 043672, 1, 30mm, PA-99-CAC-O15), Polonskogo (UW 042140, 1, 63mm, PO-98-TJL-029; UW 042146, 3, 47-50mm, PO-98-TJL-030), Shikotan (UW 02931e, 3, 39-50mm, SH-94-TWP-O03; UW 042191, 2, 22-24mm, SH-98-EMS-043; UW a42179, 8, 18 43mm, SH-98-LJW-052; UW 042188, 4, 38-48mm, SH-98-LJW-063; UW 0421g5, 50, 26-58mm, SH-98-TJL-oo4; UW 042193, 3, 44-56mm, SH-98-TJL-O05; UW 042152, 10, 42-63mm, SH-98-TJL-O17i UW 042154, 28, 57-78mm, SH-98-TJL- O19), Shumshu (UW 042240, 3, 35-37mm, SU-97-DES-024; UW 042268, 8, 47-55mm, SU-97-TWP- 028; UW 043555, 1, 51mm, SU-99-CAC-O19I UW 043564, 187, 30-57mm, SU-99-CAC-O17; UW 043706, 690, 34-68mm, SU-99-CAC-O18), Tanfilyeva (UW 042149, 3, 43-64mm, TA-98-TJL-023; UW 042156, 19, 37-67mm, TA-98-TJL-024; UW 042138, 13, 50-76 mm, TA-98-TJL-025), Urup (UW 040208, 7, 19-45mm, UR-95-EMS-Oll; UW 040189, 562, 16-50mm, UR-95-JAL-O06; UW 04e207, 5, 47-52mm, UR-95-PO-106; UW 040206, 42, 12-38 mm, UR-95-PO-114; UW 040211, 15, 17-52mm, UR- 95-SL-O18; UW 040119, 4, 26-55mm, UR-95-TWI)-O17; UW 040137, 1, 48mm, UR-95-TWP-021; UW 040215, 4, 23-38mm, UR-95-VAT-063; UW 042033, 27, 16-60mm, UR-9&PO-067), Zelionyi (UW

e29178, 2, 17-38mm, ZE-94-TWP-O02; UW 028141, 5, 39-54mm, ZE-94-TWP-O03; UW 029182, 53, 26-65mm, ZE-94-TWP-O05; UW 042143, 16, 33-60mm, ZE-9B-DES-O14; UW 042!87, 12, 40-77mm, ZE-98-DES-O15). Pungitius tymensis: Kunashir (UW 043699, 2, 51-59mm, KU-99-NM-O17), Shikotan (UW 042189, 1, 54mm, SH-98-LJW-063). Salvelinus leucomaenis: Iturup (UW 029138, 17, 74-14e mm, IT-94-TWP-O02; UW 029195, 13, 44-134mm, IT-94-TWP-O06; UW 029235, 1. 127mm, IT-94-TWI)-O16; UW 029152, 1, 77mm, IT- 94-TWP-O19; UW 042416, 1, 105mm, IT-94-TWP-O19; UW 029164, 13, 112-151mm, IT-94-TWP-e26; UW 040121, 4, 129-162mm, IT-95-TWP-O14; UW 042231, 1, 71 mm, IT-97-DES-oo5; UW 042263, 1, 58 mm, IT-97-DES-oo6; UW 042448, 4, 49-84mm, IT-98-ELM-O09; UW 042475, 1, 121 mm, IT-98-ELM- O19; UW 042468, 3, 109-･127mm, IT-98-ELM-023; UW e42443, 1, 165mm, IT-98-ELM-024; UW 042455, 1, 120mm, IT-98-ELM-025; UW 042458, 1, 118mm, IT-98-ELM-026; UW 042466, 2, 82-112 mm, IT-98-ELM-027; UW 042469, 5, 83-114mm, IT-98-ELM-029; UW 042437, 5, 93-216mm, IT-98- ELM-034; UW 042488, 7, 68-102mm, IT-98-ELM-038; UW 041366, 5, 79-349mm, IT-99-DES-028; UW 043552, 19, 92-133mm, IT-99-DES-022), Kunashir (UW 029265, 3, 5g- 144 mm, KU-94-TWP- O04; UW 029269, 1, 145mm, KU-94-TWP-oo6; UW 029271, 6, 42-62mm, KU-94-TWP-O07; UW 029276, 2, 103-181 mm, KU-94-TWP-O09; UW 029283, 10, 48-61mm, KU-94-TWP-O14; UW 029288, 2,

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158 Theodere W. Pietsch et al.

103-122mm, KU-94-TWP-O16; UW 029292, 3, B9-114mm, KU-94-TWP-O17; UW 029297, 4, 52-113 mm, KU-94-TWP-020i UW 029300, 2, 43-93mm, KU-94-TWP-021; UW 029302, 15, 75-106mm, KU- 94-TWP-022; UW 040105, 2, 82-119mm, KU-95-TWP-oo6; UW 040107, 1, 75mm, KU-95-TWP-O07; UW 040110, 2, 87 100mm, KU-95-TWP-O08; UW 040112, 1, 77mm, KU-95-TWP-O09; UW 040114, 6, 42-93mm, KU-95-TWP-OIO; UW 040117, 2, 45-65mm, KU-95-TWP-O12; UW 042035, 2, 74-79mm, KU-96-JAL-036; UW 042008, 2, 76m82mm, KU-96-JAL-037; UW 042010, 4, 78-104mm, KU-96-JAL-

038; UW 042002, 3, 58-74mm, KU-96-JAL-039; UW 042433, 4, 72"112mm, KU-96-JAL-040; UW 042004, 3, 46-85mm, KU-96-JAL-041i UW 042227, 1, 64 mm, KU-97-DES-O02; UW 042440, 25, 51-194 mm, KU-98-ELM-OOI; UW 042449, 4, 87-107mm, KU-98-ELM-O02; UW 042435, 8, 57-204mm, KU- 98-ELM-O03; UW 042451, 18, 69 103mm, KU-98-ELM-040; UW 042454, 2, 78-126mm, KU-98-TJL-

039; UW 041377, 4, 42--224mm, KU-99-DES-031i UW 041386, 5, 66-113mm, KU-99-DES-032; UW 043541, 1, 184mm, KU-99-DES-030; UW 043548, 47, 48-98mm, KU-99-DES-029), Paramushir (UW 043643, 1, 114mm, PA-99-CAC-O16; UW 043648, 2, 116-132mm, PA-99-CAC-O14), Shikotan (UW 029314, 1, l32mm, SH-94-TWP-O03; UW 029316, 4, 29-110mm, SH-94-TWP-O07; UW e29176, 2, 73- 100mm, SH-94-TWP-O09i UW 042487, 58, 70-205mm, SH-98-ELM-044; UW 042463, 1, 85mm, SH- 98-NM-027; UW 042483, 9, 97-203mm, SH-98-TJL-oo5; UW 042447, 2, 89-98mm, SH-98-TJL-oo8; UW 042477, 6, 90-123mm, SH-98-TJL-O09; UW 042442, 20, 50-129 mm, SH-98-TJL-OIO; UW 042446, 28, 70-163mm, SH-98-TJL-Oll; UW 042467, 4, 75-121 mm, SH-98-TJL-O12; UW 042489, 2, 140-l68 mm, SH-98-TJL-O15; UW 042438, 18, 74-148mm, SH-98-TJL-O16; UW 042474, 6, 72-119mm, SH-98- TJL-O18), Tanfilyeva (UW 042472, 1, 214mm, TA-98-TJr]-024), Urup (UW 040164, 2, tissue only, UR-95-JAL-O05; UW 040128, 5, 80-118 mm, UR-95-TWP-O17; UW 042414, 2, 71-78 mm, UR-95-TWP-

O19; UW 040144, 1, 195 mm, UR-95-TWP024; UW 042047, 2, 89-97mm, UR-96-JAL-023; UW 042006,

4, 77-143 mm, UR-96-JAL-027). Salvelinus malma: Iturup (UW 029307, 1, 92mm, IT-94-JBB-046; UW 029194, 19, 75-217 mm, IT-94-TWP-O06; UW 029147, 11, 94-121mm, IT-94-TVVP-O151 UW 029236, 5, 106-136mm, IT- 94-TWP-O16; UW 029153, 15, 58-223mm, IT-94-TWP-O19; UW 029248, 1, 130mm, IT-94-TWP-025;

UW 029252, 1, 127 mm, IT-94-TWP-026; UW 040120, 10, 100-2oo mm, IT-95-TWP-O14; UW 042e26, 43, 58--185 mm, IT-96-JAIr022; UW 042230, 1, 96 mm, IT-97-DES-O05; UW 042262, 3, 72-88mm, IT-

97-DES-O06; UW 042479, 18, 68-144mm, IT-98-ELM-O15; UW 042486, 48, 69-140mm, IT-98-ELM- O19i UW 042439, 24, 68T137mm, IT-98-ELM-020; UW 042457, 9, 95-136mrr1, IT-98-ELM-021; UW

042478, 40, 51-103 mm, IT-98-ELM-022; UW 042476, 3, 102-121 mm, IT-98-ELM-024; UW 042445, 60, 32-137mm, IT-98-ELM-025; UW 042482, 19, 90-124mm, IT-98-ELM-026; UW 042484, 25, 8a-172

mm, IT-98-ELM-027; UW 042481, 23, 60-162mm, IT-98-ELM-029; UW 042490, 786, 27-171mm, IT-

98-ELM-030; UW 042436, 1, 229mm, IT-98-ET.M-031; UW 042480, 19, 75-202 mm, IT-98-ELM034; UW 042444, 32, 83-186mm, IT-98-ELM-038; UW 043633, 2, 63-77mm, IT-99-DES-026), Kunashir

(UW 029270, 2, 114-135mm, KU-94-TWP-O06; UW 029278, 3, 108-190mm, KU-94-TWP-O12; UW 029291, 1, 83mm, KU-94-TWP-O17i UW 042005, 1, 84 mm, KU-96-JAL-041; UW 042228, 1, 98mm,

KU-97-DES-O02i UW 042441, 1, 118mm, KU-98-ELM-OOI; UW 042470, 9, 10-110mm, KU-98-ELM- O03; UW 042473, 5, 69-113mm, KU-98-ELM-040; UW 041388, 2, 92-111mm, KU-99-DES-032; UW

043549, 2, 79-80mm, KU-99-DES-029), Onekotan (UW 042029, 93, 28 180mm, ON-96-JAL-OIO; UW 043681, 12, 53-133mm, ON-99-CAC-O04; UW 043722, 58, 35-163 mm, 0N-99-CAC-O03), Para- mushir (UW 042014, 1, 119mm, PA-96-JAL-O03; UW 042030, 15, 58 -143mm, PA-96-JAL-O04; UW 042020, 1, 102 mm, PA-96-JAL-O06i UW 042018, 2, 92-114mrn, PA-96-JAL-oo7; UW 042272, 2, 92L94

mm, PA.97-DES-025; UW 042258, 16, 68-128mm, PA-97-DES-036; UW 042245, 1, 215mm, PA-97-

DES-051; UW 042264, 3, 89-133mm, PA-97-NM-066i UW 043662, 21, 62-127 mm, PA-99-CAC-O16i UW 043665, 2, 75-98 mm, PA-99-CAC-e14), Rasshua (UW 040153, 7, 90-135mm, RAS-95-TWP- 028), Shiashkotan (UW 042032, 17, 68-158mm, SA-96-JAL-O18), Simushir (UW 040167, 145, 54-240mm, SI-95-TWP-033; UW 043626, 84, 48-237mm, SI-99-DES-O16; UW 043639, 252, 25r280

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Fishes of the Kuril Islands 159

mm, SI-99-DES-O17; UW 043655, 57, 28-111 mm, SI-99-DES-O15), Shumshu (UW 041398, 5, 33-126 mm, SU-99-CAC-O19; UW 043627, 21, 45-134 mm, SU-99-DES-oo3; UW 043676, 2, 102-111 mm, SU- 47, 77-117mm, UR- 99-CAC-O17), Urup (UW 040127, 4, 87-152mm, UR-95-TWP-O17; UW 040133, 95-TWP-O19i UW 040138, 272, 70-145mm, UR-95-TWP-021; UW 040143, 20, 60-247mm, UR-95- TWI)-024; UW 042046, 13, 87-208mm, UR-96-JAL-023; UW 042405, 2, 64-88mm, UR-96-JAL-023; UW 040123, 2, 103-106mm, UR-95-TWP-055), Kamchatka (UW 043670, 8, 75-119mm, KM-99- CAC-Oll). Tribolodon brandti: Kunashir (UW 029167, 23, 36-90mm, KU-94-TWP-O03). Tribolodon ezoe: Iturup (UW 029299, 6, 148-173mm. IT-94-TWP-O12; UW 029251, 4, 71-96 mm, IT-94-TWP-026), Zelionyi (UW 029326, 14, 74-104mm, ZE-94TWP-oo1; UW 042498, 1, 193 mm, ZE-98-DES-O14; UW 042495, 20, 86- 157mm, ZE-98-DES-O16). Tribolodon hakuensis: Iturup (UW 041361, 14, 56-183mm, IT-99-DES-022; UW 041362, 51, 28-318mm, IT-99-DES028), Kunashir (UW 042496, 2, 54-56mm, KU-98-TJI]-040; UW 042497, 31,

44-66 mm, KU-98-TJL-038). Tridentiger kuroiwae brevispinis: Iturup (UW 029145, 48, 25-76mm, IT-94-Twu-O13; UW 029233, 62, 4479mm, IT-94-TWP-O14; UW 040538, 1, 54mm, IT-98-ELM-O16; UW 040537, 13, 48-62mm, IT-98-ELM-O17), Kunashir (UW 029255, 2, 46-50 mm, KU-94-TWP-OOI; UW 029170, 102, 26-79mm, KU-94-TWP-O03; UW 029263, 5, 43-55mm, KU-94-TWP-O04; UW 040193, 3, 34-37 mm, KU-95-JAL-O07B; UW 040535, 46, 26-85mm, KU-98-ELM-039).

References

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