^ Py. l\J J-tL ~bivL foists ff fcZJ^I CRUSTACEAN RESEARCH, NO. 28: 62-103, 1999

Revision of the East Asian species of Crangon (Decapoda: Caridea: Crangonidae)

Ken-Ichi Hayashi and Jung Nyun Kim

Abstract.—Based on several type abundant and most important compo- specimens and material collected from nents of the coastal soft bottom communi- various localities, the East Asian species ties, and are a prey or a predator of flat- of Crangon are revised. C. affinis De fishes (Seikai et al., 1993; Mori, 1998; Haan, 1849, C. propinquus Stimpson, Minami, 1998). 1860, C. dalli Rathbun, 1902a, C. Although 12 nominal taxa have been hakodatei, Rathbun, 1902b, C. cassiope reported from East Asian waters (De De Man, 1906, C. amurensis Brashnikov, Man, 1920; Holthuis, 1980), their taxo- 1907 and C. uritai new species are recog- nomic status is very unsettled as species nized. C. affinis is given full specific sta- are easily confused with each other. A tus. A neotype is designated for C. thorough revision of C. affinis De Haan, propinquus. C. septemspinosa forma 1849 and related taxa has long been amurensis and C. septemspinosa forma awaited (Hayashi, 1976; Holthuis, 1980; anivensis are assigned to C. amurensis. Komai, 1994). Very recently, the neotype The taxonomic status of C. dalli, C. of C. affinis was designated as the first hakodatei and C. cassiope is clarified by step of the revision of the East Asian the examination of the type specimens Crangon (Hayashi & Kim, 1998). and/or topotypic specimens. Each species The present study reviews Crangon is diagnosed and illustrated. C. uritai, affinis and related species based on mate- which had been confused with C. affinis, rials collected from various East Asian lo- is fully described and illustrated. Affini- calities, several type specimens and mate- ties and distribution of these seven spe- rial deposited at European and American cies are discussed and a key is provided. museums. The following seven species, including a new one, are recognized: C. affinis De Haan, 1849, C. propinquus Introduction Stimpson, 1860, C. dalli Rathbun, 1902a, Shrimps of the genus Crangon Fabri- C. hakodatei, Rathbun, 1902b, C. cassiope cius, 1798 are characterized by one me- De Man, 1906, C. amurensis Brashnikov, dian gastric spine on the carapace, a 1907 and C. uritai new species. single arthrobranch on the third maxil- Confusion on C. affinis species com- liped and one ventral spine on the merus plex has been partly clarified as a result of the first pereopod (Christoffersen, of the designation of a neotype. The spe- 1988). Nineteen species and subspecies in cific status of C. propinquus Stimpson is the genus have been described (Rathbun, also obscure. The original description is 1904; De Man, 1920; Zarenkov, 1965; short and ambiguous. The types may be Kuris & Carlton, 1977; Holthuis, 1980). no longer extant (Evans, 1967) so a neo- Crangon species commonly occur in type should be established. C. dalli is the littoral and sublittoral areas of cold comparatively well defined by having and temperate regions of the Northern paired longitudinal dorsal carinae on the Hemisphere. They are one of the most sixth abdominal somite, for which it was REVISION OF THE EAST ASIAN CRANGON 63 once assigned to the subgenus amurensis Brashnikov, 1907 Steiracrangon Kinahan, 1862 (Kuris & = Crangon amurensis Brashnikov, 1907 Carlton, 1977). The examination of the 15) Crangon septemspinosa forma anivensis syntypes of C. hakodatei Rathbun, 1902b Brashnikov, 1907 and C. cassiope De Man, 1906 reveals that = Crangon amurensis Brashnikov, 1907 they are valid species. C. septemspinosa, 16) Crangon septemspinosa sensu Derjugin reported from the Russian Far East & Kobjakova, 1935 (Derjugin & Kobjakova, 1935; Kobjakova, = Crangon amurensis Brashnikov, 1907 1936; 1937; 1958a; Vinogradov, 1947), are 17) Crangon crangon sensu Liu, 1955 now assigned to C. amurensis. Crangon = Crangon uritai new species uritai new species, is common in shallow 18) Crangon affinis sensu Liu, 1955 waters and had been previously confused = Crangon hakodatei Rathbun, 1902b with C. affinis. The new species is most closely related to the European C. Materials and Methods crangon (Linnaeus, 1758) in general mor- Specimens examined in this study are phology, but the thoracic sternum is deposited in the institutions indicated by armed with a blunt process in C. uritai the following abbreviations: CBM, Natu- instead of the acute spine found in C. ral History Museum and Institute, Chiba crangon. (with a code of ZC); HUMZ, Laboratory of The species recognized from East Marine Zoology, Faculty of Fisheries, Asian waters are summarized as follows. Hokkaido University (with a code of C); 1) Crangon affinis De Haan, 1849 MNHN, Museum national d'Histoire 2) Crangon propinquus Stimpson, 1860 naturelle, Paris; NFU, National Fisheries 3) Steiracrangon orientalis var. longicauda University, Shimonoseki; NHM, the forma pacifica Czerniavsky, 1884 Natural History Museum, London; PUIZ, = Crangon propinquus Stimpson, 1860 Invertebrate Zoology Laboratory, Depart- 4) Crangon vulgaris sensu Bate, 1888 ment of Marine Biology, Pukyong Na- = Crangon affinis De Haan, 1849 tional University; RMNH, Nationaal 5) Crangon vulgaris var. shidlovskii Natuurhistorisch Museum, Leiden; Ostroumoff, 1896 USNM, National Museum of Natural His- = Crangon propinquus Stimpson, 1860 tory, Smithsonian Institution, Washing- 6) Crangon dalli Rathbun, 1902a ton, D.C. 7) Crangon crangon sensu Rathbun, 1902b The following Crangon species col- = Crangon amurensis Brashnikov, 1907 lected from the other areas were exam- 8) Crangon hakodatei Rathbun, 1902b ined and compared with East Asian spe- 9) Crangon propinquus sensu Rathbun, cies. 1902b Crangon crangon: 2 6 (8.0, 8.0 mm), = Crangon affinis De Haan, 1849 10 9 (6.6-12.0 mm), coll. T. Kikuchi, 10) Crangon alaskensis sensu Rathbun, Vrouwenpolder, Baymouth of 1904 (in part: from Kurile Islands) Oosterschelde (Rhine delta), 29 Oct 1985, = Crangon propinquus Stimpson, 1860 NFU 530-2-1925. 11) Crangon consobrinus De Man, 1906 Crangon crangon: 4 ovig. 9 (12.1-16.8 = Crangon affinis De Haan, 1849 mm), coll. K. Hayashi, market in Paris, 17 12) Crangon cassiope De Man, 1906 Aug 1992, NFU 530-2-1926. 13) Crangon septemspinosa var. propinqua Crangon septemspinosa: 2 8 (7.8, 10.5 sensu Brashnikov, 1907 mm), 14 $ (9.1-14.5 mm), coll. W. H. Dall, = Crangon propinquus Stimpson, 1860 Chamisso Harbor, Eschscholtz Bay, 14) Crangon septemspinosa forma Alaska, 9-15 m, det. Rathbun (1904), 64 K. HAYASHI & J. N. KIM

USNM 23534. Crangon crangon affinis - Ortmann, 1895: 180 Crangon septemspinosa: 2 ovig. $ (in part); Doflein, 1900: 325 (in part). 1 Crangon crangon affinis — Doflein, 1902: 642. (11.3, 11.6 mm), no definite locality, det. Crangon propinquus - Rathbun, 1902b: 42. M. J. Rathbun, MNHN Na 1147. Not Crangon propinquus Stimpson, 1860. Crangon alaskensis: 5 S (8.8-9.2 mm), Crangon consobrinus De Man, 1906: 401 [type 3 $ (10.0-11.0 mm), 39 ovig. 9 (9.5-13.8 locality: Inland Sea of Japan]; 1907: 405, mm), Albatross sta. 3230, Bristol Bay, pi. 31, figs. 16-19; Parisi, 1919: 90. Crangon (Crangon) affinis - De Man, 1920: Alaska, 6 m, det. Rathbun (1904), USNM 249 (list, in part); Kim & Park, 1972: 205 26512. (in part); Kim, 1977: 298, pi. 54, fig. 60, text In the systematic account, species are figs. 127, 128 (in part) (= Crangon uritai arranged chronologically. The diagnosis new species); Kuris & Carlton, 1977: 553 (list); Kim, 1985: 68 (in part). and illustrations are provided for each Crangon (Crangon) consobrina - De Man, species. Crangon uritai, new species, is 1920: 250 (list). given a full description. Some characters Crago affinis - Yasuda, 1956: 7, 8, 64, unnum- are separately given by sex in the diagno- bered fig., text figs. 27, 28; Maekawa, 1961: sis since the members of the genus show 172. Not Crangon affinis - Urita, 1942: 30 (= sexual dimorphism in such characters as Crangon propinquus Stimpson, 1860); Liu, the thickness and length of the antennu- 1955: 60, pi. 22, figs. 5-8 (= Crangon lar flagellum, the length of the hakodatei Rathbun, 1902b); Kubo, 1965: scaphocerite, the sixth abdominal somite 622, fig. 1003 (= Crangon uritai new spe- and the telson and the armature of the cies); Komai et al., 1992: 195 (= Crangon amurensis Brashnikov, 1907 and Crangon thoracic sternum. propinquus Stimpson, 1860). The carapace length (CL) was mea- Not Crangon (Crangon) affinis - Hayashi, sured from the posterior margin of the or- 1976: 15 (= Crangon uritai new species). bit to the mid-dorsal posterior margin of ICrangon (Crangon) affinis - Balss, 1914: 63. the carapace; the rostrum length (RL) ICrangon affinis - Yokoya, 1930: 541; 1933: 32; 1939: 274, fig. 8; Nishimura, 1939: 383; from the rostrum apex to the level of the Sato, 1957: 86 (list); Miyake, 1961a: 9 (list); posterior margin of the orbit; the 1961b: 168 (list); Miyake et al., 1962: 124 scaphocerite length (SL) along the lateral (list); Kurata, 1964: 38, figs. 1-33 (larvae); margin from the distolateral spine to the Funada, 1966: 86 (list); Harada, 1968: 82 posterior end of the lateral margin; and in (list); Kikuchi, 1968: 180 (list); Igarashi, 1969: 9, pi. 9, fig. 26a, b, pi. 17, fig. 53; the first pereopod the longest palm length Kosaka, 1970: 59; Takeda, 1972: 25 (lar- and the broadest palm width were mea- vae); Motoh, 1972: 46, pi. 13, figs. 1-3; sured. The size of specimens is indicated Sakamoto & Hayashi, 1977: 1263; Fujino, by their carapace length (CL) in mm. 1978: 25 (list); Horikoshi et al., 1979: 49 (list); Yamashita & Shiota, 1980: 1; Holthuis, 1980: 148; Kojima & Hanabuchi, Systematics 1981: 45 (list); Ohta, 1983: 230; Ogawa et Crangon affinis De Haan, 1849 al., 1983: 237 (list); Natsukari & Iwasaki, 1987: 1; Seikai et al., 1993: 321. Figs. 1, 2 ICrago affinis - Yamauchi, 1965: 907, figs. 4-6 (larvae). Crangon affinis De Haan, 1849: 183 [type lo- cality: Japan]; Bate, 1888: 484, pi. 86, figs. Type material.—Neotype, ovig. 9 (8.3 1-3; Ortmann, 1890: 531; Fujino & Miyake, 1970: 265, fig. 9; Holthuis & Sakai, 1970: mm), Yokosuka, 5-20 fms, NHM 1888.22 293 (list); Kim, 1976: 144 (in part); (Hayashi & Kim, 1998). Toriyama & Hayashi, 1982: 89; Hayashi & Kim, 1998: 711, figs. 1, 2. Material examined.—Korea. Daedo Crangon vulgaris - Bate, 1888: 484. Not Island, Hadong, 20 m, beam trawl, 25 Feb Crangon vulgaris Fabricius, 1789. 1996, coll. Y. R. An, 1 6 (5.3 mm), 18 $ REVISION OF THE EAST ASIAN CRANGON 65

(6.5-8.1 mm), PUIZ 11. - Off Samchonpo, Kagoshima Bay, 19 Sep 1995, coll. Kyo- 15-20 m, collected with C. hakodatei, Maru, 1 6 (6.0 mm), 5 9 (6.2-8.0 mm), 2 trawl, 28 Feb 1997, coll. S. H. Kim, 1 6 ovig. 9 (9.8,10.0 mm), NFU 530-2-1938. - (4.9 mm), 1 9 (9.0 mm), 1 ovig. 9 (8.1 Off Tarumizu, Kagoshima Bay, 200 m, 21 mm), PUIZ 12. - Off Gori, mud, trawl, Nov 1995, coll. Harue-Maru, 1 <$ (6.0 Nov 1987, coll. S. Y. Hong, 4 9 (7.2-8.9 mm), 1 9 (7.7 mm), 3 ovig. 9 (7.0-8.6 mm), PUIZ 13. - Off Gori, trawl, 5 Jan mm), NFU 530-2-1939. 1996, coll. Y. R. An, 2 S (6.8, 7.5 mm), 1 9 Seto Inland Sea. Holotype of C. (7.7 mm), 17 ovig. 9 (7.0-12.0 mm), PUIZ consobrinus De Man, 1906, ovig. 9 (9.9 14. mm), NHM 1907.4.27.13. - Bay of Kobe, Pacific coast of Japan. Tokyo Bay, Challenger sta. 233, 34°39'N, 135°14'E, 8 35°20.00'-35°23.50'N, 139°40.43'- fms, 17 May 1875, referred to C. affinis by 139°46.10'E, 17-31 m, beam trawl, 21 Oct Bate (1888), 8 ovig. 9 (10.0-11.0 mm), 1997, coll. J. Ueda, 3 9 (5.3-5.9 mm), 5 NHM 1888.22. - Challenger sta. 233a, ovig. 9 (5.1-5.9 mm), NFU 530-2-1927. - 34°38'N, 135°1'E, 50 fms, 19 May 1875, Neotype, Yokoska [Yokosuka], 5-20 fms, referred to C. affinis by Bate (1888), 4 coll. HMS Challenger, referred to C. ovig. 9 (9.9-10.9 mm), NHM 1888.22. - vulgaris by Bate (1888), ovig. 9 (8.3 mm), Challenger sta. 233b, 34°18'N, 133°35'E, NHM 1888.22. - Collected with neotype, 1 15 fms, 26 May 1875, referred to C. affinis ovig. 9 (9.1 mm), 1 9 (5.0 mm), NHM by Bate (1888), 1 6 (7.3 mm), 3 ovig. 9 1888.22. - Oosezaki, Suruga Bay, 90-101 (10.0-11.0 mm), NHM 1888.22. - Off m, beam trawl, 22 Sep 1974, 3 9 (4.1-4.6 Osaka City, Osaka Bay, 12 m, mud, small mm), NFU 530-2-1928. - Mikawa Bay, trawl, 22 Feb 1996, coll. K. Ariyama, 2 6 10-20 m, coll. T. Okutani, 3 juvs (3.5-4.1 (6.5, 7.0 mm), 3 9 (10.0-10.5 mm), NFU mm), NFU 530-2-1929. - Ise Bay, 3 6 530-2-1940. - Harimanada Sea, 20-40 m, (6.1-6.6 mm), 4 ovig. 9 (8.6-9.5 mm), small trawl, Jan 1989, coll. K. Yokogawa, NFU 530-2-1930. - Kii Strait, 30 Jan 1 6 (6.8 mm), 3 ovig. 9 (8.9-9.5 mm), 1989, coll. S. Ueno, 2 9 (6.0, 7.3 mm), 2 NFU 530-2-1941. - Off Kawanoe, ovig. 9 (7.7, 7.8 mm), NFU 530-2-1931. - Hiuchinada Sea, 30 m, mud, shrimp Off Tokushima City, Kii Strait, 20 m, 28 trawl, 20 Jul 1995, coll. Y. Hanamura, 1 Apr 1990, coll. Y. Ueda, 3 6 (6.3-6.5 mm), ovig. 9 (11.2 mm), NFU 530-2-2029. - Off 2 9 (9.0, 9.4 mm), NFU 530-2-1932. - Kawanoe, Hiuchinada Sea, 30 m, mud, Tosa Bay, 92 m, beam trawl, 8 Aug 1971, shrimp trawl, 15 Jun 1996, coll. Y. Hana- coll. M. Toriyama, 4 9 (5.1-6.0 mm), NFU mura, 1 9 (11.0 mm), NFU 530-2-2030. - 530-2-1933. - Tosa Bay, 4 Jun 1977, 8 9 Miyakubo, Hiuchinada Sea, 30 m, col- (7.6-9.0 mm), 6 ovig. 9 (7.4-9.0 mm), lected with C. cassiope, sand, shrimp NFU 530-2-1934. - Beppu Bay, cover net, trawl, 13 Mar 1996, coll. Y. Hanamura, 1 21 Jul 1976, coll. M. Toriyama, 2 9 (5.8, 9 (9.6 mm), NFU 530-2-2031. - Touki- 8.7 mm), 3 ovig. 9 (5.6-9.2 mm), NFU jima Island, small sailing trawl, 2 Mar 530-2-1935. - Tarumizu, Kagoshima Bay, 1935, coll. Mukaishima Marine Labora- 19 Dec 1976, coll. Kagoshima Prefectural tory, 2 6 (6.9, 9.0 mm), NFU 530-2-1942. Fisheries Experimental Station, 4 9 (8.1- - Touwa Town, Oshima Island, 5 juvs 10.1 mm), 2 ovig. 9 (8.3, 10.4 mm), NFU (3.0-4.1 mm), NFU 530-2-1943. - Off 530-2-1936. - Kagoshima Bay, Jun 1977, Iwakuni, Jun 1976, coll. N. Takai, 4 ovig. coll. Kagoshima Prefectural Fisheries Ex- 9 (9.1-11.0 mm), NFU 530-2-1944. - Off perimental Station, 1 6 (4.8 mm), 2 9 Odoguchi, Shimonoseki, trawl, 15 May (7.1, 7.5 mm), 1 ovig. 9 (8.3 mm), NFU 1982, coll. K. Hayashi, 1 9 (8.3 mm), 17 530-2-1937. - Nishisakurajima Strait, ovig. 9 (7.1-10.5 mm), NFU 530-2-1949. 66 K. HAYASHI & J. N. KIM

Fig. 1. Crangon affinis De Haan, 1849, ovigerous female (9.5 mm, NFU 350-2-1941) from Harimanada Sea, Seto Inland Sea, Japan; entire animal in lateral view (above) and dorsal view (below). Scale = 2 mm.

NW coast of Kyushu. Genkainada Kakehuchi, Yuya Bay, Yamaguchi Prefec- Sea, 85 m, 26 Mar 1982, coll. H. Misu, 2 ture, shrimp trawl, 16 Nov 1976, coll. K. ovig. 9 (7.4, 8.4 mm), NFU 530-2-1947. - Kojima, 10 9 (4.6-7.4 mm), 1 ovig. 9 (6.9 Dokai Bay, Fukuoka Prefecture, 13 Mar mm), NFU 530-2-1952. - Toyama Bay, 1989, coll. M. Yamada, 1 6 (6.5 mm), 5 9 small trawl, 3 Aug 1976, coll. N. Horii, 5 9 (7.8-10.0 mm), NFU 530-2-1946. - (5.9-8.1 mm), NFU 530-2-1953. - Toyama Oomura Bay, Nagasaki Prefecture, 20 m, Bay, collected with C. uritai, 1 ovig. 9 shrimp trawl, 14 May 1998, coll. Y. (9.7 mm), NFU 530-2-1954. - Off Niigata, Natsukari, 4 ovig. 9 (8.9-1.0 mm), PUIZ 120 m, trawl, 30 Nov 1978, coll. H. Itano, 45. 1 <5 (6.7 mm), 1 9 (8.7 mm), NFU 530-2- East China Sea. trawl, 30 Oct 1976, 1955. coll. Nansei-Maru, 2

Fig. 2. Crangon affinis De Haan, 1849; a, c-h, ovigerous female (8.9 mm, PUIZ 14) from off Gori, Korea; b, female (10.0 mm, NFU 350-2-1936) from Tarumizu, Kagoshima Bay, Japan; i, male (7.5 mm, PUIZ 14) from off Gori, Korea: a, carapace and anterior cephalic appendages, dorsal, left antennule omitted; 6, carapace and cephalothoracic appendages, lateral; c, abdominal somites and telson, dorsal; d, left first pereopod, flexor; e, left third pereopod, lateral; f, left fourth pereo- pod, lateral; g, left fifth pereopod, lateral; h, i, thoracic sternum, ventral, setae omitted. Scales = 2 mm. 68 K. HAYASHI & J. N. KIM somite without mid-dorsal carina, fourth in adults. somite with trace of mid-dorsal carina on Parasites.—The following females posterior half; fifth somite with low but were infected with an unidentified distinct mid-dorsal carina; sixth somite bopyrid: two (7.1, 8.0 mm, PUIZ 11) from flattened or slightly grooved dorsally and Daedo, Hadong, Korea; one (7.8 mm, NFU with median groove ventrally, 0.71-0.87 530-2-1934) from Tosa Bay, Japan; one times as long as carapace in males, 0.58- (8.7 mm, NFU 530-2-1935) from Beppu 0.84 times in females. Telson with shal- Bay, Japan; two (7.1, 7.5 mm, NFU 530-2- low median groove dorsally, 0.94-1.13 1937) from Kagoshima Bay, Japan; one times as long as carapace in males, 0.82- (9.4 mm, NFU 530-2-1951) from Yuya 1.02 times in females. Scaphocerite 0.85- Bay, Japan; and two (9.6 mm, NFU 530-2- 0.96 times as long as carapace in males, 1948; 8.3 mm, NFU 530-2-1949) from off 0.78-0.92 times in females. Third maxil- Shimonoseki, Japan. liped with subdistal clump of 4-6 (mostly Remarks.—Recently the neotype of 5) spines on ventral surface of antepenul- Crangon affinis De Haan, 1849 was desig- timate segment. First pereopod with palm nated by Hayashi & Kim (1998), who re- 3.11-4.05 times longer than broad. Third examined the specimens referred to as pereopod with carpus 1.90-2.60 times Crangon vulgaris and C. affinis by Bate longer than distal two segments combined (1888), C. propinquus by Rathbun (1902b) in adults, 1.53-1.80 times in juveniles. and the holotype of C. consobrinus De Fourth and fifth pereopods moderately Man, 1906. These specimens proved to long and slender; dactylus 1.03-1.46 belong to the same species. times as long as carpus in fourth, 0.94- The following material identified as C. 1.67 times in fifth. Fifth thoracic sternite affinis was reexamined. Specimens from (between coxae of second pereopods) with Tosa Bay (NFU 530-2-1933 & -1934) re- acute median spine in ovigerous females. ported by Toriyama & Hayashi (1982) ac- Size.—Males 4.1-9.0 mm, females tually represent C. affinis. Hayashi's 4.1-11.0 mm, ovigerous females 5.1-12.0 (1976) material from Sado Islands (NFU mm, juveniles 3.0-4.1 mm. 530-2-1999) is referable to C. uritai, new Distribution.—Yellow Sea, northeast- species. Ten specimens from Hokkaido ern part of the East China Sea, coast of (HUMZ-C 380) reported by Komai et al. Honshu and Seto Inland Sea, Japan (De (1992) are found to contain two species, C. Haan, 1849; Bate, 1888; Ortmann, 1890; amurensis and C. propinquus. Rathbun, 1902b; De Man, 1906; 1907; Parisi (1919) reported two species, C. Yasuda, 1956; Fujino & Miyake, 1970; consobrinus De Man from Tokyo Bay and Kim, 1977; Toriyama & Hayashi, 1982; C. crangon Linnaeus from Yokohama and Hayashi & Kim, 1998; present study); 5- Enoshima on the central Pacific coast of 200 m. Japan. Although his material was not ex- Variations.—The rostrum is slightly amined, C. consobrinus is thought to be a variable in direction, from slightly de- Crangon with a carinated abdomen and scending (Fig. 1) to straight (Fig. 26). The C. crangon with the smooth abdomen. third maxilliped, fourth and fifth pereo- The present material indicates that only pods vary in length, reaching to or beyond two species, C. affinis which has a mid- the blade of the scaphocerite (Figs. 1, 26). dorsal carina on the abdomen and C. The carpus of the third pereopod be- uritai new species, which has no carina on comes proportionally longer with growth. the abdomen, are present in the region. The ratio of carpus length/length of the Parisi's (1919) C. consobrinus and C. distal two segments combined changes crangon, therefore, seem to be referable to from 1.53-1.80 in juveniles to 1.90-2.35 C. affinis and C. uritai, respectively. REVISION OF THE EAST ASIAN CRANGON 69

In their ecological study of the Seto In- Balss (1914) considered that C. land Sea, Yasuda (1956) and Maekawa propinquus Stimpson, 1860, C. hakodatei (1961) dealt with two species identified as Rathbun, 1902b, C. consobrinus De Man, Crago affinis and Crago cassiope. The 1906 and C. cassiope De Man, 1906 were present study shows that three species, C. synonymous with C. affinis. His opinion affinis, C. cassiope and C. uritai occur in has been followed by Japanese authors the area. Judging from the locality and (e.g., Yokoya, 1930; 1933; Urita, 1942; his figures (Yasuda, 1956: p. 8, unnum- Igarashi, 1969; Fujino & Miyake, 1970). bered figures), which depict the anterior Crangon affinis, therefore, was listed sub- part of the carapace in dorsal view, sequently without sufficient information Yasuda's (1956) identification of Crago on morphology (Yokoya, 1930; 1933; 1939; affinis was correct. There is still a possi- Nishimura, 1939; Sato, 1957; Miyake, bility that his Crago cassiope consisted of 1961a; 1961b; Miyake et al., 1962; two species, Crangon cassiope and C. Funada, 1966; Harada, 1968; Kikuchi, uritai. Although there is no information 1968; Igarashi, 1969; Kosaka, 1970; on morphology, Maekawa's (1961) identi- Motoh, 1972; Sakamoto & Hayashi, 1977; fication seems to be the same as Yasuda's Fujino, 1978; Horikoshi et al., 1979; (1956). Yamashita & Shiota, 1980; Kojima & Fujino & Miyake (1970) reported Hanabuchi, 1981; Ohta, 1983; Ogawa et Crangon affinis from the East China Sea al., 1983; Natsukari & Iwasaki, 1987; and Yellow Sea. Although they found that Seikai et al., 1993). It is impossible, how- the shape of the rostrum is variable in ever, to identify the species reported by their material, their specimens seem to be these authors without the reexamination referable to the true C. affinis because the of their specimens. Material from the fifth abdominal somite has a dorsal carina same or nearby localities from which the in the shape of a thin line and the sixth authors collected their material, not al- abdominal somite is slightly grooved dor- ways consisted of a single species. Fur- sally and ventrally. thermore, the common species in shallow The given dimensions of Urita's (1942) waters around Japan is not always C. C. affinis from Sakhalin suggest that it is affinis. As some species of Crangon occur probably C. propinquus because of the sympatrically, it is difficult to determine relatively longer scaphocerite (0.80 times the specific status of the previously de- as long as the carapace) and the relatively scribed larvae under the name of Crangon robust palm (2.50 times as long as width) (or Crago) affinis (cf. Kurata, 1964; of the first pereopod of a female. Liu Yamauchi, 1965; Takeda, 1969). (1955) reported C. affinis from northern China. The description and figures, how- Crangon propinquus Stimpson, 1860 ever, show that his species is probably not Figs. 3, 4 C. affinis but C. hakodatei (see "Remarks" of C. hakodatei). There is no doubt that C. Crangon propinquus Stimpson, 1860: 25 [type affinis described by Kubo (1965) repre- locality: northern Japan] (in part); Komai et al., 1992: 195. sents C. uritai (see "Remarks" of C. Steiracrangon orientalis var. longicauda uritai). Crangon affinis reported from Ko- forma pacifica Czerniavsky, 1884: 74 [type rea (Kim & Park, 1972; Kim, 1976; 1977; locality: Sakhalin]. 1985) seems to have been mixed with C. Crangon vulgaris var. shidlovskii OstroumofF, uritai. The figure given by Kim (1977: fig. 1896: 75 [type locality: Vladivostok]. Crangon alaskensis - Rathbun, 1904: 114 (in 127) shows that the sixth abdominal part). somite and telson slightly are grooved Crangon septemspinosa var. propinqua - dorsally. Brashnikov, 1907: 84; Derjugin & 70 K. HAYASHI & J. N. KIM

Kobjakova, 1935: 142; Kobjakova, 1936: 5133. - Odaitou, light collecting, collected 190, 202, 212 (list); 1937: 93, 94 (list); with C. amurensis, referred to C. affinis 1958a: 221, 229, 244; 1958b: 253; 1959: 70; 1967: 238. by Komai et al. (1992), 30 May 1988, 3 6 Crangon (Crangon) alaskensis - De Man, (9.0-10.0 mm), 1 $ (6.0 mm), HUMZ-C 1920: 249 (list, in part). 380. Crangon (Crangon) crangon var. shidlovskii - Russian Far East. Olga Bay, De Man, 1920: 250 (list). subtidal, beach seine, collected with C. Crangon (Crangon) orientalis — De Man, 1920: 250 (list, in part) amurensis, 18 Aug 1994, coll. M. Yabe, 2 Crangon (Crangon) propinquus — De Man, ovig. $ (12.9, 14.6 mm), CBM-ZC 2442. - 1920: 250 (list). Off Kurile Islands, Albatross sta. 3652,14 Crangon affinis - Urita, 1942: 30; Komai et al., fms, referred to C. alaskensis by Rathbun 1992: 195 (in part). Not Crangon affinis De (1904), 4 6 (9.1-10.6 mm), USNM 26511. Haan, 1849. Crangon septemspinosa morpha propinqua - Vinogradov, 1947: 91; 1950: 217, fig. 79. Diagnosis.—Integument partially pu- ICrangon propinquus - Miyake et al., 1962: bescent. Rostrum usually reaching mesial 124. distal margin of eyestalk, 0.15-0.20 times Not Crangon propinquus - Rathbun, 1902b: as long as carapace, directed forward, 42 ( = Crangon affinis De Haan, 1849). rounded distally. Carapace with median gastric spine arising from about 0.25 of Type material.—It may have been lost carapace length. Third abdominal somite (Evans, 1967). The ovigerous female (11.4 without mid-dorsal carina; fourth somite mm; NFU 530-2-1960) from Lake Saro- with faint mid-dorsal carina; fifth somite ma, Hokkaido is designated the neotype with low mid-dorsal carina; sixth somite of C. propinquus (see "Remarks" below). flattened or slightly grooved dorsally, Material examined.—Pacific coast with median groove ventrally, 0.68-0.74 of Japan. Akkeshi Bay, 16 Sep 1990, coll. times as long as carapace in males, 0.62- T. Minami, 1 9 (12.4 mm), 1 ovig. 9 (10.8 0.72 times in females. Telson flattened or mm), NFU 530-2-1957. - Off Katsurakoi, slightly grooved dorsally, 0.93-1.05 times Kushiro, 36 m, 3 Oct 1989, coll. J. Sasaki, as long as carapace in males, 0.87-0.98 1 6 (8.2 mm), 11 juvs (2.0-4.1 mm), PUIZ times in females. Scaphocerite 0.79-0.84 46. - Off Tomakomai, 17 Dec 1984, coll. Y. times as long as carapace in males, 0.74- Hanamura, 1 ovig. $ (15.1 mm), NFU 0.83 times in females. Third maxilliped 530-2-2032. - Off Date City, Uchiura Bay, with subdistal clump of 4-5 (mostly 5) Jun 1982, coll. Y. Hanamura, 5

Fig. 3. Crangon propinquus Stimpson, 1860, neotype, ovigerous female (11.4 mm, NFU 530-2- 1960) from Lake Saroma, Hokkaido, Japan; entire animal in lateral view (above) and dorsal view (below). Scale = 2 mm.

(Stimpson, 1860; Czerniavsky, 1884; latter is C. hakodatei Rathbun. An Ostroumoff, 1896; Brashnikov, 1907; ovigerous female specimen (11.4 mm; Urita, 1942; Vinogradov, 1947; Komai et NFU 530-2-1960) from Lake Saroma, al., 1992; present study); intertidal to 36 Hokkaido, Japan has been selected as the m (Vinogradov, 1947; present study). neotype of C. propinquus. A duplicate Remarks.—Crangon propinquus was specimen (12.2 mm) collected with the described from "prope oras boreales neotype is deposited in CBM (code with Japoniae" by Stimpson (1860). The origi- catalogue no. ZC 5133). nal description is very short and obscure Czerniavsky (1884) described in parts and the types may be no longer Steiracrangon orientalis var. longicauda extant (Evans, 1967). The type specimens forma pacifica based on a single specimen have "abdominis segmento quarto (et from Sakhalin. Ostroumoff (1896), who interdum tertio quoque) in adultis cari- questioned its validity, described a new nato" (Stimpson, 1860). In northern variety, Crangon vulgaris var. Japanese waters, a carina on the fourth shidlovskii, based on three specimens abdominal somite is present in two spe- from Vladivostok and Aniva Bay, cies but the carinated third somite is only Sakhalin. After the examination of the found on one of them. The present mate- type of Czerniavsky's (1884) taxon and rial examined does not show variation of five additional specimens from Tatar the carina on the third somite. There is no Strait and Terpeniya Bay, Sakhalin, doubt, therefore, that Stimpson's descrip- Brashnikov (1907) concluded that tion surely based on two species: one had Czerniavsky's (1884) taxon and a carina on the fourth somite; the other Ostroumoffs (1895) taxon were conspe- had the carinated third and fourth cific with C. septemspinosa var. pro- somites. The former should be referred to pinqua Stimpson, 1860, based on the the true C. propinquus Stimpson and the presence of a low blunt median carina on 72 K. HAYASHI & J. N. KIM

Fig. 4. Crangon propinquus Stimpson, 1860; a-g, neotype, ovigerous female (11.4 mm, NFU 350- 2-1960) from Lake Saroma, Hokkaido, Japan; h, male (10.0 mm, HUMZ-C 380) from off Odaitou, Hokkaido, Japan: a, carapace and anterior cephalic appendages, dorsal, left antennule omitted; b, abdominal somites and telson, dorsal; c, left first pereopod, flexor; d, left third pereopod, lat- eral; e, left fourth pereopod, lateral; f, left fifth pereopod, lateral; g, h, thoracic sternum, ventral, setae omitted. Scales = 2 mm. REVISION OF THE EAST ASIAN CRANGON 73 the fourth and fifth somites in both taxa. (1992) reported C. affinis and C. Although no types of the two Russian propinquus from Odaitou and Akkeshi, varienties were examined, the present Hokkaido, respectively. These specimens study supports Brashnikov's (1907) con- were reexamined. Some of the Odaitou clusion. specimens (HUMZ-C 380) and all of the Derjugin & Kobjakova (1935), Akkeshi specimens (NFU 530-2-1957) Vinogradov (1947, 1950) and Kobjakova proved to belong to C. propinquus. (1958a, 1958b, 1959, 1967) reported C. septemspinosa var. propinqua or C. Crangon dalli Rathbun, 1902 septemspinosa morpha propinqua from Figs. 5, 6 the Sea of Japan and the Sea of Okhotsk. These records from the Russian Far East Crangon dalli Rathbun, 1902a: 889 [type lo- seem to represent C. propinquus, judging cality: off Cape Seniavin, Alaska]; 1904: from the localities and depths (intertidal 119, fig. 60; Brashnikov, 1907: 84, fig. 6; to 35 m). Balss, 1914: 63; Yokoya, 1933: 33; Derjugin & Kobjakova, 1935: 142 (list); Kobjakova, The specimens of C. alaskensis re- 1936: 190, 191, 201, 212, 224; 1937: 93, 94, ported from off the Kurile Islands by 139; Yokoya, 1939: 276; Urita, 1942: 31; Rathbun (1904) have been examined. Vinogradov, 1947: 92; 1950: 217, fig. 80; They proved to be not C. alaskensis Kobjakova, 1955: 150, pi. 37, fig. 1; 1958a: Lockington, 1877 but C. propinquus. C. 221, 229, 244; 1958b: 253; 1959: 70; Igarashi, 1969: 9, pi. 10, fig. 27, pi. 17, fig. propinquus closely resembles the eastern 54; Kim & Park, 1972: 206, pi. 5, fig. 1; North Pacific C. alaskensis in having a Kim, 1976: 145; Kosaka, 1979: 185; relatively longer rostrum (about 0.15- Horikoshi et al., 1979: 49; Butler, 1980: 99, 0.20 times as long as the carapace), the unnumbered fig.; Komai et al., 1992: 195; third maxilliped usually bearing a clump Komai, 1994: 96. Crangon (Crangon) dalli - De Man, 1920: 250 of 5 small spines on the antepenultimate (list). segment, the moderately stout palm of Crago dalli - Makarov, 1941: 135. the first pereopod (about 2.50-3.00 times Crangon ralli (sic) - Zarenkov, 1965: 1762, fig. as long as width) and a thoracic sternite 1. that is provided with an acute spine on Crangon (Neocrangon) dalli - Kim, 1977: 300, pi. 30, fig. 63, text figs. 126, 131, 132. the fifth somite in ovigerous females. C. Crangon (Steiracrangon) dalli - Kuris & propinquus, however, differs from C. Carlton, 1977: 553 (list). alaskensis in having a slightly descending and broader rostrum, a fourth somite that Type material.—Holotype, ovig. 9, is furnished with a faint low mid-dorsal (16.2 mm), Alaska, 30 fms, USNM 25244 carina (versus without a carina) and a (not examined). relatively shorter (0.74-0.83 times as long Material examined.—Korean coast as the carapace versus 0.79-0.88 times) of the Sea of Japan. 36°43'N, 129°31'E, and broader (2.54-2.78 times as long as 110 m, mud, trawl, 2 Nov 1995, coll. J. H. width versus 2.67-3.00 times) scapho- Choi, 1 ovig. 9 (16.6 mm), PUIZ 15. - cerite in females. 36°12'N, 129°29'E, 51m, sandy mud, col- As mentioned above, C. affinis re- lected with C. hakodatei, trawl, 2 Nov ported from Sakhalin by Urita (1942) 1995, coll. J. H. Choi, 1 ovig. 9 (13.0 mm), seems to be referrable to the present spe- PUIZ 16. - 36°56'N, 129°37'E, 100 m, cies. Miyake et al. (1962) listed C. mud, trawl, 5 Nov 1995, coll. J. H. Choi, 2 hakodatei and C. propinquus from 6 (11.2, 12.8 mm), 1 9 (12.7 mm), PUIZ Niigata but neither identification could be 17. - 35°28'N, 129°36'E, 130 m, mud, confirmed with certainly. Komai et al. trawl, 6 Nov 1995, coll. J. H. Choi, 2 9 74 K. HAYASHI & J. N. KIM

Fig. 5. Crangon dalli Rathbun, 1902, ovigerous female (11.5 mm, PUIZ 49) from off Shirito, Kushiro, Hokkaido, Japan; entire animal in lateral view (above) and dorsal view (below). Scale = 2 mm.

(16.8, 18.0 mm), PUIZ 18. - 35°36'N, Albatross sta. 3288, 15 fms, 10 9 (10.3- 129°30'E, 47 m, mud, collected with C. 13.8 mm), 15 ovig. $ (10.1-13.2 mm), hakodatei, trawl, 18 Mar 1996, coll. J. H. USNM 26623. Choi, 1 $ (12.8 mm), PUIZ 19. - 35°16'N, 129°34'E, 120 m, collected with C. Diagnosis.—Integument pubescent. hakodatei, trawl, 19 Mar 1996, coll. J. H. Rostrum usually reaching distal margin Choi, 1 ovig. $ (13.0 mm), PUIZ 20. of cornea, 0.16-0.22 times as long as cara- Japanese coast of the Sea of Ja- pace, directed forward, rounded distally. pan. Off northeastern Tsushima Island, Carapace with median gastric spine aris- 35°42'N, 130°18'E, 165 m, soft mud, col- ing from about 0.25 of carapace length. lected with C. hakodatei, trawl, 18 Sep Third to fifth abdominal somites without 1995, coll. S. Ueno, 1 6 (12.1 mm), 1 $ median carina; sixth somite with two (12.2 mm), NFU 530-2-1961. blunt submedian carinae dorsally and Pacific coast of Japan. Akkeshi median groove ventrally, 0.67-0.84 times Bay, 30 m, 19 Apr 1990, coll. Y. as long as carapace in males, 0.62-0.80 Hanamura, 1 ovig. $ (14.0 mm), NFU times in females. Telson with median 530-2-2044. - Off Shiriba, Kushiro, groove dorsally, 0.86-0.98 times as long Hokkaido, 57 m, 15 May 1991, coll. J. as carapace in males, 0.87-1.00 times in Sasaki, 2 6 (9.9, 11.1 mm), 2 $ (4.5, 15.2 females. Scaphocerite 0.79-0.94 times as mm), PUIZ 48. - Off Shirito, Kushiro, long as carapace in males, 0.78-0.90 in fe- Hokkaido, 36 m, 29 Jul 1991, coll. J. males. Third maxilliped with subdistal Sasaki, 2 ovig. 9 (11.5, 14.9 mm), PUIZ clump of 5-7 (mostly 6) spines on ventral 49. - Off Konbumori, Kushiro, Hokkaido, surface of antepenultimate segment. 60 m, 29 Jul 1991, coll. J. Sasaki, 3 S First pereopod with palm 2.24-2.91 times (7.9-9.0 mm), PUIZ 50. longer than broad. Third pereopod with Alaska. Off Khondonbine Islands, Al- carpus 1.46-2.10 times longer than distal batross sta. 3284, 25 fms, 10 S (8.1-10.7 two segments combined. Fourth and fifth mm), 1 9 (10.4 mm), 19 ovig. $ (10.4-15.2 pereopods relatively robust; dactylus mm), USNM 26619. - Off Cape Seniavin, 0.92-1.21 times as long as carpus in REVISION OF THE EAST ASIAN CRANGON 75

Fig. 6. Crangon dalli Rathbun, 1902; a-i, ovigerous female (11.5 mm, PUIZ 49) from Shirito, Kushiro, Hokkaido, Japan;./', male (11.1 mm, PUIZ 48) from Shiriba, Kushiro, Hokkaido, Japan: a, carapace and anterior cephalic appendages, dorsal, left antennule omitted; b, left third maxil- liped, ventral; c, same, anterior part of antepenultimate segment, ventral; d, left first pereopod, lateral; e, same, chela, flexor; f, left third pereopod, lateral; g, left fourth pereopod, lateral; h, left fifth pereopod, lateral; ij, thoracic sternum, ventral, setae omitted. Scales = 2 mm. 76 K. HAYASHI & J. N. KIM fourth, 0.92-1.23 times in fifth. Fifth tho- racic sternite with median tubercle in ovigerous females. Crangon hakodatei Rathbun, 1902 Size.—Males 7.9-12.8 mm, females Figs. 7, 8 4.5-18.0 mm, ovigerous females 10.1- Crangon propinquus Stimpson, 1860: 25 (in 16.6 mm. part). Distribution.—Northern North Pacific Crangon hakodatei Rathbun, 1902b: 42, fig. 15 from Puget Sound to the Sea of Japan [type locality: Hakodate, Hokkaido, Ja- pan]; Kim, 1976: 145; Komai et al., 1992: (Rathbun, 1902; Balss, 1914; Yokoya, 195 (list). 1933; 1939; Derjugin & Kobjakova, 1935; Crangon (Crangon) hakodatei - De Man, 1920: Kobjakova, 1936; Makarov, 1941; Urita, 250 (list); Kim, 1977: 297, pi. 30, fig. 61, 1942; Vinogradov, 1950; Kim, 1977; text fig. 129; 1985: 68. Kosaka, 1979; Horikoshi et al., 1979; But- Crago hakodatei - Urita, 1926: 429. ler, 1980; present study); Chukchi Sea, Crangon affinis - Liu, 1955: 60, pi. 22, figs. 5- 8. Not Crangon affinis De Haan, 1849. (Vinogradov, 1950); subtidal to 630 m Crangon nigricauda - Kim & Park, 1972: 206, (Kobjakova, 1936). pi. 5, fig. 2. Not Crangon nigricauda Remarks.—The distinct submedian Stimpson, 1856. carinae on the sixth abdominal somite ?Crangon hakodatei - Miyake et al., 1962: 124 clearly separate Crangon dalli from the (list). other species in East Asian waters. Kinahan (1862) actually divided the ge- Type material.—Syntypes, 1 S (7.8 nus Crangon into two subgenera, mm), 2 9 (8.2, 8.9 mm), Hakodate, Hok- Crangon and Steiracrangon using this kaido, USNM 26156 (see "Material exam- character. Kuris & Carlton (1977) revived ined"). the subgeneric division proposed by Material examined.—Yellow Sea. Kinahan (1862) and they assigned C. Taechon, Korea, winged stow net on an- dalli to the subgenus Steiracrangon. chor, 29 Mar 1995, coll. H. K. Cha, 12 6 Christoffersen (1988) considered that the (8.0-9.9 mm), 2 9 (8.9, 9.0 mm), 4 ovig. $ subgeneric division proposed by Kuris (8.0-12.0 mm), PUIZ 21. - Sikdo Island, and Carlton (1977) was invalid because Puan, Korea, collected with C. cassiope, the definition was based on a symplesio- winged stow net on anchor, 29 Mar 1995, morphic character state and produced coll. H. K. Cha, 2 6 (8.9, 10.0 mm), PUIZ paraphyletic grade-groups in the cladistic 22. - Yongdo Island, Youngkwang, Korea, classification of Crangonidae, such as the winged stow net on anchor, 24 Apr 1995, 'convex group' of Kuris and Carlton coll. H. K. Cha, 1 ovig. $(12.9 mm), PUIZ (1977). The subgenus Steiracrangon is 23. - Yongdo Island, Youngkwang, Korea, not accepted in the present study follow- winged stow net on anchor, 7 Dec 1995,13 ing Christoffersen's (1988) conclusion. 6 (7.0-10.1 mm), 5 $ (10.8-13.0 mm), East Asian specimens agree well with PUIZ 24. - Japanese Fisheries Section the topotypic specimens examined as well 129, 35°50'-36°00'N, 123°50 -124WE, as the original and subsequent descrip- trawl, Autumn 1986, coll. Seikai National tions of C. dalli given by Rathbun (1902a; Fisheries Research Institute, 4 <$ (7.8- 1904) and Butler (1980). Butler (1980) in- 11.0 mm), 6 9(9.0-14.8 mm), 1 ovig. 9 dicated that a slight concavity of the ven- (15.0 mm), NFU 530-2-1962. - Japanese tral margin of the fifth abdominal pleuron Fisheries Section 324, 33°00'-33°50'N, was also diagnostic for this species. The 123°50'-124°00'E, trawl, 29 Sep 1989, present study, however, found that the coll. Seikai National Fisheries Research ventral margin is rather variable from Institute, 2 S (6.0,10.0 mm), 2 9(9.0,11.9 slightly concave to slightly convex. mm), NFU 530-2-2034. - Japanese Fish- REVISION OF THE EAST ASIAN CRANGON 77 eries Section 160, 35°00'-35°50'N, 1978, coll. H. Itou, 5 9 (11.0-15.2 mm), 122°00'-122°50'E, trawl, 29 Jan 1991, NFU 530-2-1964. - Off NW Tsushima Is- coll. Seikai National Fisheries Research land, 33°38'N, 129°10'E, 153-204 m, Institute, 2 9(10.5, 10.6 mm), 1 ovig. 9 trawl, 31 May 1978, 5 9 (9.9-14.1 mm), (12.9 mm), NFU 530-2-2035. - Japanese HUMZ-C 371. - Off NE Tsushima Island, Fisheries Section 159, 35°50'-36°00'N, 35°42'N, 130°18'E, 165 m, soft mud, col- 122°00'-122°50'E, trawl, 15 Sep 1993, lected with C. dalli, trawl, 18 Sep 1995, coll. Seikai National Fisheries Research coll. S. Ueno, 2 9 (10.7, 16.9 mm), NFU Institute, 2 6 (6.9, 9.3 mm), 2 9(13.9,14.0 530-2-1965. - Miwa, Toyama Bay, sta- mm), NFU 530-2-2036. tionary net, 5 Apr 1976, coll. N. Horii, 1 Korean coast of the northeastern ovig. 9 (15.0 mm), NFU 530-2-1966. - East China Sea. Off Samchonpo, 10 m, Miwa, Toyama Bay, stationary net, May trawl, 10 Jan 1997, coll. S. H. Kim, 2 9 1976, 2 9 (14.3, 18.4 mm), NFU 530-2- (15.0, 15.2 mm), PUIZ 25. - Off 1967. - Hijikata, Toyama Bay, 250 m, Samchdnpo, 15-20 m, trawl, collected hand trawl, 19 Apr 1976, coll. N. Horii, 1 with C. affinis, 28 Feb 1997, coll. S. H. S (11.9 mm), NFU 530-2-1968. - Off Kim, 2 6 (8.8, 9.2 mm), 9 9(9.0-12.2 mm), Niigata, 120 m, 30 Nov 1978, coll. H. 2 ovig. 9(10.1, 14.2 mm), PUIZ 26. - Itano, 3 6 (6.0-8.0 mm), 6 9 (6.8-12.8 Hansan Island, 30-40 m, trawl, 22 Jun mm), NFU 530-2-1969. - Off Sakata, 1988, coll. S. Y. Hong, 1 ovig. 9 (12.5 mm), Yamagata Prefecture, 43-45 m, 1 9 (13.2 PUIZ 27. - Hansan Island, 30-40 m, mm), NFU 530-2-1970. trawl, 16 Sep 1988, coll. S. Y. Hong, 2 9 Tsugaru Strait. Hakodate, (5.0, 7.2 mm), PUIZ 28. - Hansan Island, Hokkaido, syntypes of C. hakodatei 30-40 m, trawl, 26 Jun 1992, coll. S. Y. Rathbun, 1902b, 1 S (7.8 mm), 2 9 (8.2, Hong, 4 6 (5.1-7.1 mm), 13 9 (5.0-12.3 8.9 mm), USNM 26156. mm), 4 juvs (3.1-4.1 mm), PUIZ 29. Pacific coast of Japan. Hachinohe, Korean coast of the Sea of Japan. Aomori Prefecture, 10 m, collected with C. 36°12'N, 129°29'E, 51 m, sandy mud, col- uritai, beam trawl, 8 9 (5.9-8.6 mm), lected with C. dalli, trawl, 2 Nov 1995, HUMZ-C 566. coll. J. H. Choi, 3 9 (13.9-14.2 mm), PUIZ Japanese coast of the Sea of 30. - 35°36'N, 129°30'E, 47 m, mud, col- Okhotsk. Off Oumu, Hokkaido, lected with C. dalli, trawl, 18 Mar 1996,1 44°35.66'N, 143°03.71'E, 44 m, 22 Aug 6 (7.9 mm), 1 9 (11.0 mm), PUIZ 31. - 1996, coll. J. Sasaki, 2 S (9.0, 9.0 mm), 1 35°26'N, 129°28'E, 75 m, mud, trawl, 18 9 (11.5 mm), 1 ovig. 9 (16.0 mm), PUIZ Mar 1996, 1 6 (12.5 mm), 1 9 (14.1 mm), 51. PUIZ 32. - 35°16'N, 129°34'E, 120 m, col- lected with C. dalli, trawl, 18 Mar 1996, 2 Diagnosis.—Integument pubescent. 6 (8.8,10.4 mm), 1 9 (10.0 mm), 1 ovig. 9 Rostrum usually reaching distal margin (12.9 mm), PUIZ 33. - Off Gori, trawl, 5 of cornea, 0.17-0.24 times as long as cara- Jan 1996, coll. Y. R. An, 2 6 (8.0, 8.1 mm), pace, directed forward, rounded distally. 2 9 (8.0, 8.1 mm), 1 ovig. 9 (10.0 mm), Carapace with median gastric spine aris- PUIZ 34. - Off Gori, trawl, Jul 1996, 9 6 ing from about 0.20 of carapace length. (7.3-12.2 mm), 9 9 (7.9-12.8 mm), PUIZ Third to fifth abdominal somites with 35. blunt mid-dorsal carina; sixth somite Japanese coast of the Sea of Ja- with median groove dorsally and ven- pan. Tsukumo Bay, Ishikawa Prefecture, trally, 0.62-0.66 times as long as cara- 1 May 1973, 1 6 (5.9 mm), NFU 530-2- pace in males, 0.56-0.66 times in females. 1963. - Off western Tsushima Island, Telson with median groove dorsally, 34°38'N, 129°10'E, 153-204 m, 31 Mar 0.89-0.99 times as long as carapace in 78 K. HAYASHI & J. N. KIM

Fig. 7. Crangon hakodatei Rathbun, 1902, ovigerous female (12.9 mm, PUIZ 33) from the Korean coast of the Sea of Japan; entire animal in lateral view (above) and dorsal view (below). Scale = 2 mm. males, 0.88-0.92 times in females. long as carapace; Figs. 7, 8a). In a female Scaphocerite 0.74-0.83 times as long as (14.1 mm, NFU 530-2-1962) from the Yel- carapace in males, 0.69-0.82 times in fe- low Sea, it exceptionally does not reach males. Third maxilliped with subdistal the distal margin of the cornea (0.12 clump of 4-6 (mostly 5) spines on ventral times as long as carapace; Fig. 86). surface of antepenultimate segment. The antepenultimate segment of the First pereopod with palm 2.32-2.93 times third maxilliped mostly bears a subdistal longer than broad. Third pereopod with clump of 4-6 (mostly 5) spines on the ven- carpus 1.56-2.18 times longer than distal tral surface. In two females (10.0 mm, two segments combined. Fourth and fifth NFU 530-2-1969; 18.4 mm, NFU 530-2- pereopods relatively robust; dactylus 1967) from off Niigata and Miwa, Toyama 0.83-1.10 times as long as carpus in Bay, respectively, it exceptionally bears a fourth, 0.91-1.12 times in fifth. Fifth tho- subdistal clump of 9 spines on the left racic sternite with acute median spine in side. ovigerous females. Remarks.—Three of the eight syn- Size.—Males 5.1-12.5 mm, females types of Crangon hakodatei, that consist 5.0-18.4 mm, ovigerous females 8.0-16.0 of one male and two females from Hako- mm, juveniles 3.1-4.1 mm. date, Hokkaido, were reexamined. Unfor- Distribution.—Yellow Sea, northern tunately, they were not in good condition. part of the East China Sea, Sea of Japan, The cephalothorax was detached from the northern part of the Pacific coast of Japan abdomen in all three specimens. The male and southern part of the Sea of Okhotsk and smaller female were missing several (Rathbun, 1902b; Urita, 1926; Liu, 1955; cephalothoracic appendages and the Kim & Park, 1972; Kim, 1976; 1977; 1985; larger female was mutilated. These types, present study); 10-250 m. however, retain the abdominal features Abnormality.—The rostrum usually well, which are the most important in the reaches or slightly overreaches the distal species identification. margin of the cornea (0.17-0.24 times as C. hakodatei is readily separated from REVISION OF THE EAST ASIAN CRANGON 79

35) from off Gori, Korea: a, carapace and anterior cephalic appendages, dorsal, left antennule omitted; 6, anterior part of carapace and eyes, dorsal; c, abdominal somites and telson, dorsal; d, left first pereopod, flexor; e, left third pereopod, lateral; f, left fourth pereopod, lateral; g, left fifth pereopod, lateral; h, i, thoracic sternum, ventral, setae omitted. Scales = 2 mm. 80 K. HAYASHI & J. N. KIM other species by a high, blunt median dor- coll. K. Yokogawa, 3 6 (5.9-6.6 mm), 1 9 sal carina on the third to fifth abdominal (7.6 mm), 3 ovig. 9 (8.7-10.2 mm), NFU somites. 530-2-1971. - Bisan Strait, small trawl, As mentioned in the account for C. 21 Jan 1989, coll. K. Yokogawa, 2 6 (6.2, propinquus, there is little doubt that the 7.1 mm), 2 ovig. 9 (9.7, 11.7 mm), NFU types of C. propinquus contained two spe- 530-2-1972. - Bisan Strait, small trawl, cies, C. propinquus and C. hakodatei. 11 May 1989, coll. K. Yokogawa, 2 6 (6.2, Urita (1926) and Kim (1977) recog- 7.1 mm), 2 ovig. 9 (9.7, 11.7 mm), NFU nized that C. hakodatei was distinct from 530-2-1973. - Miyakubo, Hiuchinada Sea, other related species in having the me- 30 m, sand, shrimp trawl, 21 Dec 1995, dian carina on the third to fifth abdominal coll. Y. Hanamura, 1 6 (6.5 mm), 2 9 somites. Liu's (1955) C. affinis from (7.7, 8.0 mm), NFU 530-2-2037. - northern China is referred to C. hakodatei Miyakubo, Hiuchinada Sea, 30 m, sand, because he clearly mentioned the pres- collected with C. affinis, shrimp trawl, 13 ence of the median carina on the third to Mar 1996, coll. Y. Hanamura, 2 S (7.0, 8.4 fifth abdominal somites. C. nigricauda mm), 4 9 (9.6-11.2 mm), 2 ovig. 9 (9.5, from Korea (Kim & Park, 1972) was later 10.0 mm), NFU 530-2-2038. - Touyo City, reidentified as C. hakodatei by Kim Hiuchinada Sea, 1.5-2 m, fine sand, col- (1976). Miyake et al. (1962) recorded C. lected with C. uritai, 1 Jul 1998, coll. Y. hakodatei from Niigata, but they did not Hanamura, 1 9 (9.9 mm), NFU 530-2- substantiate the specific status. 2039. Japanese coast of the Sea of Ja- pan. Yoshimi, Shimonoseki, 1 m, skin Crangon cassiope De Man, 1906 diving, 3 May 1989, 1 9 (14.3 mm), NFU Figs. 9, 10 530-2-1974. - Yuya Bay, Yamaguchi Pre- Crangon cassiope De Man, 1906: 402 [type lo- fecture, 2 m, collected with C. uritai, cality: Inland Sea of Japan]; 1907: 406, pi. shrimp trawl, 9 Jun 1983, coll. K. Kojima, 32, figs. 20-25; Liu, 1955: 59, pi. 22, figs. 1- 1 ovig. 9 (12.2 mm), NFU 530-2-1975. - 4. Tango Sea, Kyoto Prefecture, 4-7 m, 18 Crangon (Crangon) cassiope - De Man, 1920: Jun 1971, 1

Fig. 9. Crangon cassiope De Man, 1906, ovigerous female (14.3 mm, NFU 350-2-1974) from Yoshimi, Shimonoseki, Japan; entire animal in dorsal view. Scale = 2 mm.

Diagnosis.—Integument naked. Ros- Size.—Males 4.5-8.4 mm, females trum usually not extending beyond me- 4.8-14.3 mm, ovigerous females 7.0-12.2 sial distal margin of eyestalk, 0.13-0.19 mm, juveniles 2.1-4.0 mm. times as long as carapace, directed for- Distribution.—Yellow Sea, northern ward, rounded distally. Carapace with part of the East China Sea, central and median gastric spine arising about 0.20 of southern the Sea of Japan, Tsugaru carapace length. Third to fifth abdominal Strait and Seto Inland Sea (De Man, somites without median carina; sixth 1906; 1907; Liu, 1955; present study); somite not grooved dorsally or ventrally, 1.5-30 m. 0.66-0.79 times as long as carapace in Parasites.—The following specimens males, 0.62-0.75 times in females. Telson from the Seto Inland Sea were infected without median groove dorsally, 1.03- with an unidentified bopyrid: one female 1.08 times as long as carapace in males, (9.9 mm, NFU 530-2-2039) from Touyo 0.79-0.94 times in females. Scaphocerite City, Hiuchinada Sea; one male (8.4 mm, 0.77-0.87 times as long as carapace in NFU 530-2-2038) from Miyakubo, males, 0.67-0.79 times in females. Third Hiuchinada Sea. maxilliped with subdistal clump of 4-6 Remarks.—The syntypes of Crangon (mostly 6) spines on ventral surface of an- cassiope are two ovigerous females from tepenultimate segment. First pereopod the Seto Inland Sea. The smaller speci- with palm 2.04-2.73 times longer than men is still in good condition. The other broad. Third pereopod with carpus 1.29- specimen is damaged: the cephalothorax 1.60 times longer than distal two seg- is detached from the abdomen, the distal ments combined. Fourth and fifth pereo- parts of both antennular flagella are pods moderately short and robust; missing and the left antenna, third maxil- dactylus 0.66-0.90 times as long as car- liped and fifth pereopod had been re- pus in fourth, 0.54-0.90 times in fifth. moved, probably for examination. Fifth thoracic sternite with median tu- Crangon cassiope is a good species that bercle in ovigerous females. is distinguished from the other species of 82 K. HAYASHI & J. N. KIM

Fig. 10. Crangon cassiope De Man, 1906; a-i, ovigerous female (7.0 mm, PUIZ 36) from Puan, Korea; j, male (5.6 mm, PUIZ 36) from Puan, Korea: a, carapace and anterior cephalic append- ages, dorsal, left antennule omitted; b, carapace and cephalothoracic appendages, lateral; c, ab- dominal somites and telson, dorsal; d, sternite of sixth abdominal somite, ventral; e, left first pereopod, flexor; f, left third pereopod, lateral; g, left fourth pereopod, lateral; h, left fifth pereo- pod, lateral; i,j, thoracic sternum, ventral, setae omitted. Scales = 2 mm. REVISION OF THE EAST ASIAN CRANGON 83 the East Asian Crangon by the rounded pan. Odaitou, Sea of Okhotsk coast of sternite of the sixth abdominal somite, a Hokkaido, light collecting, 30 Apr 1988, 4 tubercle on the thoracic sternum in the 9 (7.6-12.0 mm), 2 ovig. 9 (11.7, 11.8 ovigerous female, the smallest palm mm), NFU 530-2-1984. - Odaitou, Sea of length/width ratio of the first pereopod Okhotsk coast of Hokkaido, light collect- and the smallest dactylus/carpus of the ing, collected with C. propinquus, referred fourth and fifth pereopods (see Tables 1 to C. affinis by Komai et al. (1992), 30 and 2). It has been treated as a junior syn- May 1988, 3 6 (6.8-7.4 mm), 1 9 (11.7 onym of C. affinis, however (cf. Balss, mm), 2 ovig. 9 (9.3, 12.5 mm), HUMZ-C 1914; Yokoya, 1930; Urita, 1941; 380. - Furen Lake, Nemuro Bay, 26 Jul Igarashi, 1969; Fujino & Miyake, 1970). 1991, coll. Y. Hanamura, 4 6 (5.1-6.1 Liu (1955) examined material from the mm), 5 9 (4.9-5.4 mm), NFU 530-2-2040. coast of northern China and recognized - Hakodate, Tsugaru Strait, coll. Alba- that C. cassiope was separated from other tross, referred to C. crangon by Rathbun congeners of East Asia by having rounded (1902b), 6 S (7.0-8.0 mm), 13 9 (9.0-13.7 abdominal somites and the sixth somite mm), USNM 28506. - Oohama Beach, without median groove on the ventral sur- Ishikari Bay, intertidal, fine sand, 27 Aug face. 1989, coll. Y. Hanamura, 1 $ (6.3 mm), 3 As mentioned above, Crago cassiope 9 (4.7-7.7 mm), NFU 530-2-2041. from the Seto Inland Sea reported by Russian Far East. Srednya Bay, 1 m, Yasuda (1956) and Maekawa (1961) Phyllospadix bed, beach seine, 24 Aug seems to be Crangon cassiope or C. uritai 1994, coll. M. Yabe and K. Ikeya, 3 6 (5.3- since both are known from the same re- 6.1 mm), 7 9 (6.0-9.5 mm), 2 ovig. 9 (9.0, gion and share a similar shape of the an- 9.4 mm), NFU 530-2-1979. - Srednya terior part of body. Bay, 42°38.6'N, 131°12.4'E, seagrass bed, beach seine, 27 Aug 1994, coll. M. Yabe, 2 Crangon amurensis Brashnikov, 1907 9 (10.0, 13.0 mm), NFU 530-2-1980. - Figs. 11, 12 Ostrovok Falsiuy Cape, 42°27.2'N, 131°47.0'E, beach seine, collected with C. Crangon crangon - Rathbun, 1902b: 42. Not Crangon crangon Linnaeus, 1758. uritai, 30 Aug 1994, coll. M. Yabe, 4 9 Crangon septemspinosa forma amurensis (10.0-13.3 mm), NFU 530-2-1981. - Brashnikov, 1907: 76 [type locality: Amur Tzoitsy Bay, sandy and rocky bottom, River Estuary, Russian Far East]. beach seine, collected with C. cassiope Crangon septemspinosa forma anivensis Brashnikov, 1907: 76 [type locality: Amur and C. uritai, 29 Aug 1994, coll. M. Yabe, Bay, Sakhalin]; 2 6 (4.8, 5.9 mm), 3 9 (6.0-8.5 mm), NFU Crangon septemspinosa - Derjugin & 530-2-1982. - Olga Bay, subtidal, beach Kobjakova, 1935: 142; Kobjakova, 1936: seine, collected with C. propinquus, 18 190, 191, 202, 212 (list); 1937: 138; 1958a: Aug 1994, coll. M. Yabe, 1 9 (10.1 mm), 1 244 (list); Vinogradov, 1947: 91; 1950: 217, fig. 78. Not Crangon septemspinosa Say, ovig. 9 (10.9 mm), CBM-ZC 5135. - 1818. Piltum Bay, Sakhalin, 7 m, beam trawl, Crangon affinis - Komai et al., 1992: 195 (in 22 Jul 1995, coll. M. Yabe, 1 6 (9.5 mm), 3 part). 9 (9.8-15.8 mm), 1 ovig. 9 (15.0 mm), NFU 530-2-1983. Type material.—The type may be de- posited at the Zoological Institute, Rus- Diagnosis.—Integument naked. Ros- sian Academy of Science, St. Petersburg, trum usually not extending beyond me- but it was not located. sial distal margin of eyestalk, 0.13-0.19 Material examined.—Hokkaido, Ja- times as long as carapace, directed for- Fig. 11. Crangon amurensis Brashnikov, 1907, ovigerous female (10.1 mm, CBM-ZC 5135) from Olga Bay, Russian Far East; entire animal in lateral view (above) and dorsal view (below). Scale = 2 mm. ward, rounded distally. Carapace with males. median gastric spine arising from about Size.—Males 4.8-9.5 mm, females 0.20 of carapace length. Third to fifth ab- 4.7-15.8 mm, ovigerous females 9.0-15.0 dominal somites without median carina; mm. sixth somite flattened or slightly grooved Distribution.—Northern part of the dorsally and with median groove ven- Sea of Japan, Tsugaru Strait and Sea of trally, 0.73-0.82 times as long as cara- Okhotsk (Rathbun, 1902b; Brashnikov, pace in males, 0.65-0.77 times in females. 1907; Kobjakova, 1936; Vinogradov, 1947; Telson slightly grooved dorsally, 0.88- present study); intertidal-27 m (Vino- 1.06 times as long as carapace in males, gradov, 1947; present study). 0.86-1.01 times in females. Scaphocerite Remarks.—Brashnikov (1907) de- 0.80-0.91 times as long as carapace in scribed two new forms, C. septemspinosa males, 0.71-0.81 times in females. Third forma amurensis and C. septemspinosa maxilliped with subdistal clump of 3-5 forma anivensis from the Russian Far (mostly 4) spines on ventral surface of an- East. Brashnikov's (1907) two new forms tepenultimate segment. First pereopod have the following common features: the with palm 2.56-3.14 times longer than fourth and fifth abdominal somites are broad. Third pereopod with carpus 1.60- smooth and convex on the dorsal surface, 1.84 times longer than distal two seg- the rostrum is relatively short (0.12-0.14 ments combined. Fourth and fifth pereo- times of the carapace) and the scapho- pods relatively robust; dactylus 0.88-1.11 cerite has a narrow lamina and long times as long as carpus in fourth, 0.86- distolateral spine. He also considered 1.13 times in fifth. Fifth thoracic sternite that C. septemspinosa forma amurensis with acute median spine in ovigerous fe- differs from C. septemspinosa forma REVISION OF THE EAST ASIAN CRANGON 85

Fig. 12. Crangon amurensis Brashnikov, 1907; a-g, ovigerous female (10.1 mm, CBM-ZC 5135) from Olga Bay, Russian Far East; h, male (9.5 mm, NFU 530-2-1983) from Piltum Bay, Sakhalin, Russia: a, carapace and anterior cephalic appendages, dorsal, left antennule omitted; b, abdomi- nal somites and telson, dorsal; c, left first pereopod, flexor; d, left third pereopod, lateral; e, left fourth pereopod, lateral; f, left fifth pereopod, lateral; g, h, thoracic sternum, ventral, setae omit- ted. Scales = 2 mm. 86 K. HAYASHI & J. N. KIM anivensis in having a longer chela of the Moreover, C. amurensis differs from C. first pereopod (0.50-0.55 times versus septemspinosa in having a more stout 0.49 times as long as the carapace) and a palm of the first pereopod: 2.56-3.14 shorter sixth abdominal somite (0.64- times as long as broad in C. amurensis, 0.67 times versus 0.73 times as long as while it is 2.80-3.28 times in C. the carapace) in females. septemspinosa. Although the type specimens of C. Derjugin & Kobjakova (1935), septemspinosa forma amurensis and C. Kobjakova (1937) and Vinogradov (1947, septemspinosa forma anivensis were not 1950) followed Brashnikov's (1907) iden- examined, the present specimens from tification and reported C. septemspinosa various localities, including those re- from various localities of the Russian Far ferred to C. crangon by Rathbun (1902b), East. Although the specimens reported by agree well with Brashnikov's (1907) spe- Russian authors were not available, C. cies. The characters of Brashnikov's septemspinosa of these authors seems to (1907) two forms are also almost included be conspecific with C. amurensis because in the range of the following individual of similar localities and depth (4-27 m). variation of the present material: the Komai et al. (1992) referred ten speci- chela of the first pereopod is 0.47-0.56 mens from Odaitou, Hokkaido to C. times as long as the carapace and the affinis (HUMZ-C 380). The specimens, sixth abdominal somite is 0.65-0.77 times which were reexamined, consisted of two as long as the carapace in females. The species: six specimens of the present spe- two forms, therefore, should be placed un- cies and four specimens of C. propinquus. der the name of C. amurensis, which has a page precedence over C. anivensis. Crangon amurensis is closely related Crangon uritai new species to the European C. crangon and Alaskan Figs. 13-16 C. septemspinosa in that they all lack any sculpture on the abdomen and have a Crago sp. - Urita, 1926: 429. relatively short rostrum. C. amurensis, Crangon crangon - Parisi, 1919: 90; Liu, 1955: 58, pi. 21, figs. 1-10. Not Crangon crangon however, is distinguished from C. Linnaeus, 1758. crangon and C. septemspinosa by having Crangon affinis - Kubo, 1965: 622, fig. 1003. an intermediate position between them in Not Crangon affinis De Haan, 1849. the shape of the scaphocerite. In C. Crangon (Crangon) affinis - Hayashi, 1976: amurensis, the distolateral spine is 15; Kim, 1977: 298, pi. 54, fig. 60, text figs. shorter or longer than the distal width of 127, 128 (in part). Not Crangon affinis De Haan, 1849. the blade. On the other hand, the ? Crago cassiope - Yasuda, 1956: 8, 65, unnum- distolateral spine is as long as or longer bered fig., fig. 29; Maekawa, 1961: 172. than the distal width of the blade in C. septemspinosa, while it is usually shorter Material examined.—Holotype: in C. crangon. The distal portion of the Dadaepo, Pusan, Korea, 1 m, sand, hand blade is wider in C. amurensis than in C. net, 22 Jul 1996, coll. M. H. Kim, ovig. 9 septemspinosa and narrower than in C. (7.1mm), PUIZ 38. crangon. Furthermore, the scaphocerite is Paratypes: Korea. Sinjin Island, most slender in C. septemspinosa, broad- Taean, 4 m, SCUBA, 3 Aug 1996, coll. J. est in C. crangon and intermediate in C. H. Choi, 1 6 (3.9 mm), 2 9 (4.1, 5.0 mm), 4 amurensis. The ratios of length/width of juvs (2.0-3.5 mm), PUIZ 39. - Mokpo, 23 the scaphocerite are as follows: 2.84-2.95 Apr 1988, coll. S. Y. Hong, 2 ovig. 9 (8.2, in C. septemspinosa, 2.31-2.48 in C. 9.0 mm), NFU 530-2-2043. - Youngsan crangon and 2.36-2.83 in C. amurensis. River Estuary, trawl, 23 Apr 1988, coll. S. REVISION OF THE EAST ASIAN CRANGON 87

Y. Hong, 1 ovig. 9 (9.0 mm), PUIZ 40. - NFU 530-2-1990. Nakdong River Estuary, sandy mud, Seto Inland Sea. Touyo City, shrimp trawl, Jun 1988, coll. S. Y. Hong, 4 Hiuchinada Sea, 1.5-2 m, fine sand, col- ovig. 9 (6.8-7.6 mm), PUIZ 41. - lected with C. cassiope, 1 Jul 1998, coll. Y. Nakdong River Estuary, sandy mud, Hanamura, 3 S (3.1-3.6 mm), 1 9 (3.5 shrimp trawl, 22 Nov 1988, coll. S. Y. mm), 2 ovig. 9 (7.0, 8.9 mm), NFU 530-2- Hong, 2 9 (6.9, 7.0 mm), PUIZ 42. - Col- 2042. lected with holotype, 2

Fig. 13. Crangon uritai new species, paratype, ovigerous female (6.5 mm, PUIZ 43) from Dadaepo, Pusan, Korea; entire animal in lateral view (above) and dorsal view (below). Scale = 2 mm.

42°27.2'N, 131°47.0'E, beach seine, col- males. Third maxilliped with subdistal lected with C. amurensis, 30 Aug 1994, clump of 3-4 (mostly 4) spines on ventral coll. M. Yabe, 1 ovig. 9 (7.0 mm), NFU surface of antepenultimate segment. 530-2-2002. - Tzoitsy Bay, sandy and First pereopod with palm 2.65-3.33 times rocky bottom, collected with C. cassiope longer than broad. Third pereopod with and C. amurensis, beach seine, 29 Aug carpus 1.43-2.05 times longer than distal 1994, coll. M. Yabe, 1 9 (5.5 mm), 1 ovig. two segments combined. Fourth and fifth $ (6.0 mm), NFU 530-2-2003. pereopods relatively robust; dactylus 0.89-1.19 times as long as carpus in Diagnosis.—Integument naked. Ros- fourth, 0.88-1.13 times in fifth. Fifth tho- trum usually not extending beyond me- racic sternite with median tubercle in sial distal margin of eyestalk, 0.12-0.20 ovigerous females. times as long as carapace, directed for- ward or slightly descending, rounded dis- Description of females.—Body de- tally. Carapace with median gastric spine pressed (Fig. 13). Integument naked, not arising from anterior about 0.20 of cara- particularly firm. pace length. Third to fifth abdominal Rostrum usually not extending beyond somites without median carina; sixth mesial distal margin of eyestalk, 0.12- somite with median groove ventrally, but 0.20 times as long as carapace, triangu- without submedian carinae or median lar, straight or slightly descending, con- groove dorsally, 0.74-0.78 times as long cave dorsally, rounded at apex, with fine as carapace in males, 0.64-0.73 times in marginal setae (Fig. 14a). Carapace with females. Telson without median groove median gastric spine on anterior about dorsally, 0.87-1.02 times as long as cara- 0.20 length of carapace; hepatic spine pace in males, 0.81-0.91 times in females. supported by short carina; moderately Scaphocerite 0.74-0.90 times as long as strong antennal, strong branchiostegal carapace in males, 0.64-0.77 times in fe- and weak pterygostomial spines present; REVISION OF THE EAST ASIAN CRANGON 89

Fig. 14. Crangon uritai new species, holotype, ovigerous female (7.1 mm, PUIZ 38) from Dadaepo, Pusan, Korea. Appendages dissected from left side: a, carapace and anterior cephalic appendages, dorsal, left antennule omitted; b, abdominal somites and telson, dorsal; c, sternite of sixth abdominal somite, ventral; d, telson and uropods, dorsal; e, antennule, ventral, flagella and setae omitted; f, antenna, ventral, flagellum and setae omitted; g, mandible, external; h, maxillule, external; i, maxilla, external;,;, first maxilliped, external; k, second maxilliped, exter- nal; I, third maxilliped, flexor. Scales = 1 mm. 90 K. HAYASHI & J. N. KIM postorbital carina defined, overreaching rounded distally, with several setae on midlength of carapace; ventral margin of distal margin; distal endite curved in- carapace forming weak lobe at base of sec- ward, distal margin widely subtruncate ond pereopod (Figs. 13,14a). with 5 strong spines and sparse setae; All abdominal somites rounded dor- palp slightly curved, pointed distally, sally (Figs. 13, 146). Sixth somite 0.64- with some apical setae (Fig. 14h). Maxilla 0.73 times as long as carapace; posterolat- with vestigial endites; palp spatulate, eral process pointed; sternum grooved, with several apical setae; scaphognathite with anal spine (Fig. 14c). Telson rounded with subtriangular anterior lobe, poste- dorsally, gradually tapering to sharp rior lobe narrower and rounded, with pos- point, 0.81-0.91 times as long as cara- terior setae slightly longer than mesial pace, with 3 pairs of lateral spines, setae (Fig. 14i). First maxilliped with anteriormost pair situated at anterior endite completely reduced; endopod elon- 0.60 of telson; posterior margin with 2 gate, reaching beyond anterior margin of pairs of spines, outer pair longer than in- epipod with long lateral plumose setae; ner (Figs. 13, 146, d). exopod longer than endopod, with moder- Eye with cornea well developed, hav- ately curved lash; epipod large, distinctly ing small dorsal tubercle; eyestalk with bilobed, each lobe subtriangular (Fig. distoventral spine (Fig. 14a). 14/). Second maxilliped with endopod Antennule depressed, with peduncle compressed; dactylus attached diagonally reaching near midlength of scaphocerite. to propodus, with 3 strong spines on Proximal segment longer than distal two proximal half of mesial margin and dense segments combined, with two subdistal setae on distomesial margin; propodus spines on mesial margin; ventromedial with 2 strong spines and several setae on ridge with acute spine. Stylocerite wide, subdistomesial margin; exopod with well subtriangular at base; lateral margin developed lash; epipod well developed, slightly rounded, slightly sulcate later- sickle-shaped (Fig. 14k). Third maxilliped ally, terminating in acute apex, falling reaching beyond distolateral spine of slightly short of anterior margin of proxi- scaphocerite; ultimate segment slightly mal segment. Second and distal segments longer than penultimate; antepenulti- subequal in length. Outer flagellum slen- mate segment with subdistal clump of 3- der, falling slightly short of blade of 4 (mostly 4) spines on ventral surface; scaphocerite. Inner flagellum more slen- exopod well developed, with moderately der than outer flagellum, reaching beyond curved lash; epipod rudimentary (Fig. blade of scaphocerite (Fig. 14a, e). 14Z); arthrobranch present. Antenna with scaphocerite 0.64-0.77 First pereopod subchelate, nearly times as long as carapace, lateral margin reaching blade of scaphocerite; palm mod- slightly convex, distolateral spine slightly erately stout, 2.65-3.33 times as long as curved inward, exceeding rounded blade. broad, cutting edge moderately oblique; Basicerite with distolateral spine. movable finger not overreaching base of Carpocerite reaching midlength of fixed finger when closed; carpus short, scaphocerite (Fig. 14a, /). Flagellum as with two distolateral spines; merus with long as body. strong spine on middle of mesial margin Mouthparts similar to typical and weak spine on dorsodistal margin crangonid form. Mandible simple, com- (Figs. 15a, 166). Second pereopod slender, posed only of strongly curved molar pro- chelate; movable finger about 0.40 times cess, divided distally into 4 teeth (Fig. as long as palm, both fingers setose (Fig. 14g). Maxillule with proximal endite 156). Third pereopod slightly more slen- REVISION OF THE EAST ASIAN CRANGON 91

Dadaepo, Pusan, Korea. Appendages dissected from left side: a, first pereopod, flexor; b, second pereopod, lateral; c, third pereopod, lateral; d, fourth pereopod, lateral; e, fifth pereopod, lateral; f, thoracic sternum, ventral, setae omitted; g, first pleopod, ventral; h, second pleopod, ventral. Scales = 1 mm. 92 K. HAYASHI & J. N. KIM der than second, reaching distal margin of exopod increasing from about 0.20 (4.4 palm of first pereopod; carpus 1.43-2.05 mm) to 0.50 (9.0 mm); setation longer in times as long as dactylus and propodus ovigerous females than in non-ovigerous combined, becoming longer with growth females, but naked in immature speci- (Fig. 15c). Fourth pereopod more robust mens (Fig. 15g). Second pleopod with than third, reaching two-thirds length of endopod tapering distally, strongly scaphocerite; dactylus subspatulate, mod- curved laterally, about 0.25 of exopod erately curved, 0.66-0.91 times as long as length (Fig. 15h). Protopods with mesial propodus, 0.89-1.19 times as long as car- margin slightly concave medially, bearing pus; flexor margin of propodus setose; setae at disto- and proximo-lateral cor- merus with row of setae on both flexor ners in large specimens (Fig. 15g,h). Ap- and extensor margins (Fig. 15c?). Fifth pendix interna absent on all pleopods. pereopod similar to fourth, reaching Uropod reaching posterior end of tel- midlength of scaphocerite; dactylus 0.70- son; exopod slightly shorter than 0.95 times as long as propodus, 0.88-1.13 endopod, with small spine just mesial to times as long as carpus; merus less setose acute posterolateral spine (Fig. 15c/). than that of fourth (Fig. 15e). Dimorphism.—Males differ from fe- Pleurobranchs on fourth to eighth tho- males as follows. racic somites inclined anteriorly. Body more slender than in females. Branchial formula as follows: Sixth abdominal somite 0.74-0.78 times Maxillipeds Pereopods as long as carapace. Telson 0.87-1.02 1 2 3 1 2 3 4 5 times as long as carapace. Outer antennu- lar flagellum more stout and longer than Pleurobranchs - - - 1 1 1 1 1 in females, exceeding distal end of Arthrobranchs - - r _ _ _ _ _ Podobranchs scaphocerite (Fig. 16a). Scaphocerite Epipods 1 1 r — — — — — 0.74-0.90 times as long as carapace (Fig. Exopods 1 1 1 16a). Thoracic sternum convex with moder- r: rudimentary. ately strong median spine on fifth somite Sculpture and armament of thoracic and weak median spines on sixth to sternum varying greatly with growth; in eighth somites; transverse sutures sepa- juveniles fifth to eighth thoracic somites rating sixth to eighth somites distinct convex each with median spine, trans- (Fig. 16c). Abdominal sternites each with verse sutures separating sixth to eighth acute median spine. somites distinct; in mature but non- First pleopod tapering distally; ovigerous specimens fifth to eighth endopod curved laterally, about 0.20 of somites slightly convex each with sub- exopod length (Fig. 16d). Second pleopod acute median tubercle; in ovigerous fe- with endopod longer than appendix males posterior part of thoracic somites masculina with numerous mesial and concave, with median tubercle on fifth marginal setae (Fig. 16e, /). Protopods somite only, transverse sutures indicat- with mesial medial margin slightly con- ing sixth to eighth somites very obscure vex (Fig. 16d, e). (Fig. 15/). Abdominal sternites each with Size.—Males 3.1-9.9 mm, females acute median spine in immature speci- 3.4-11.9 mm, ovigerous females 4.5-12.5 mens. mm, juveniles 2.0-3.5 mm. First pleopod with endopod changing Distribution.—Yellow Sea, northern in relative length and setation with part of the East China Sea, central and growth; comparative length against southern part of the Sea of Japan, Seto REVISION OF THE EAST ASIAN CRANGON 93

a

±L

Fig. 16. Crangon uritai new species; a, par a type, male (7.0 mm, HUMZ-C 566) from Hachinohe, Japan; b, paratype, ovigerous female (11.4 mm, NFU 350-2-1997) from Yuya Bay, Japan; c-f, paratype, male (4.7 mm, PUIZ 43) from Dadaepo, Pusan, Korea: a, carapace and anterior cepha- lic appendages, dorsal; b, left first chela, flexor; c, thoracic sternum, ventral, setae omitted; d, left first pleopod, ventral; e, left second pleopod, ventral; f, same, endopod and appendix masculina. Scales = 1 mm.

Inland Sea, central and southern Pacific 166). coasts of Japan, (Urita, 1926; Liu, 1955; The carpus of third pereopod becomes present study); intertidal to 24 m. proportionally longer with growth. The Variations.—The palm of the first ratio of carpus length/length of the distal pereopod varies from having a moderately two segments combined changes from oblique cutting edge and straight flexor 1.43 in 4.7 mm to 2.05 in 11.3 mm. margin (Fig. 15a) to a more oblique cut- Parasites.—A female (9.5 mm, HUMZ- ting edge and concave flexor margin (Fig. C 99) from Miyako Bay infested by an uni- 94 K. HAYASHI & J. N. KIM dentified bopyrid. A female (NFU 530-2-1999) from Etymology.—This species is named for Mano Bay, Sado Island, which was re- the late Mr. Tomoye Urita, who first rec- ferred to C. affinis by Hayashi (1976), ognized the present form as distinct from proved to be C. uritai. As mentioned other congeners. above, Yasuda's (1956) and Maekawa's Remarks.—The present new species is (1961) Crago cassiope from the Seto In- very similar to Crangon crangon in the land Sea and Kim's (1977) Crangon absence of a median carina on the ab- affinis from Korea appear to have con- dominal somites, the presence of a me- sisted of one or more species, including dian groove on the ventral surface of the the present new species. sixth abdominal somite and having a rela- tively broad scaphocerite. It differs from the European species by the following Discussion morphological features. In the ovigerous Differentiation of species females, the fifth thoracic sternite is The seven species of the genus armed with a blunt process in C. uritai, Crangon recognized here have long been instead of an acute spine in C. crangon. confused with each other (Holthuis, The third maxilliped distinctly over- 1980). Most of them have been mis- reaches the distolateral spine of the identified as C. affinis. The reliable scaphocerite in C. uritai, while it only characters to differentiate the seven spe- reaches the distal margin of the blade in cies are discussed in Tables 1 and 2. C. crangon. The antepenultimate seg- The Atlantic C. crangon, and Alaskan ment of the third maxilliped is usually C. septemspinosa and C. alaskensis were armed with a subdistal clump of four once reported from East Asian waters spines on the ventral surface in C. uritai, (Rathbun, 1902b; 1904; Brashnikov, instead of three spines in C. crangon. 1907; Liu, 1955). They are clearly differ- As mentioned in the account for C. ent from the East Asian congeners. Al- affinis above, C. crangon reported by though C. dalli is known from both sides Parisi (1919) from Yokohama and Eno- of the North Pacific, the remaining spe- shima, Japan seems to be referable to the cies have their characteristic features in present new species. Urita (1926) com- East Asian waters. pared Crago sp. from Tsingtao, China The rostrum shape is useful in the rec- with Crangon affinis and C. cassiope. Al- ognition of some species, although it though the description is brief, his Crago shows some variation in some of the spe- sp. surely represents Crangon uritai. He cies (Figs. 1, 2b, 8a, b). In C. affinis (Fig. mentioned the absence of a mid-dorsal 2a) it usually reaches the distal margin of abdominal carina and the presence of a the cornea and has a sub acutely pointed ventral median groove on the sixth ab- apex, while C. cassiope (Fig. 10a), C. dominal somite. Liu's (1955) description amurensis (Fig. 12a) and C. uritai (Figs. and figures of C. crangon from northern 14a, 16a) have a shorter rostrum that China quite agree with the present spe- usually does not reach the mesial distal cies, especially in the presence of a blunt margin of the eyestalk as well as a median process on the fifth thoracic stern- rounded apex. The rostrum is similar in ite in ovigerous females. Kubo's (1965) il- C. propinquus (Fig. 4a), C. dalli (Fig. 6a) lustration of C. affinis also appears very and C. hakodatei (Fig. 8a). It usually similar to this species. His account was reaches the distal margin of the cornea rather general, but the illustration may and the apex is rounded. have been based on a specimen of C. The sculpture on the abdominal uritai. somites and telson is most important in REVISION OF THE EAST ASIAN CRANGON 95 species recognition. Crangon dalli (Fig. 5) males than in females in all species. The is the unique species in having two dis- comparison between scaphocerite length tinct dorsal submedian carinae on the and carapace length is also useful in spe- sixth somite. In C. hakodatei (Fig. 8c), the cies recognition (Table 2): it is the highest third to fifth somites have a blunt mid- in C. affinis and C. dalli, the lowest in C. dorsal carina and the sixth somite and tel- cassiope and C. uritai and intermediate in son have a median groove. All abdominal the other three species. somites and telson of both C. cassiope As indicated by previous authors (Fig. 10c) and C. uritai (Fig. 146) are (Ortmann, 1895; Rathbun, 1902a; 1902b; rounded dorsally. Crangon cassiope (Fig. 1904; Kuris & Carlton, 1977; Butler, 10c0 is unique in not having a grooved 1980), proportions of the palm of the first sternite in the sixth somite. Crangon pereopod provide less discrete but notable affinis (Fig. 2c) and C. propinquus (Fig. differences in species recognition (Table 46) have a low but distinct mid-dorsal ca- 2). The palm is most robust in C. cassiope rina on the fifth somite and a shallow dor- (Fig. lOe) and most slender in C. affinis sal median groove on the sixth somite and (Fig. 2d), while it is intermediate in the telson. The fourth somite in C. affinis has other five species (Figs. 4c, 6e, 8d, 12c, a trace of a mid-dorsal carina on the pos- 15a). terior half (Fig. 2c), while a faint mid-dor- The comparative length of each seg- sal carina is present in C. propinquus ment of the last three pereopods is useful (Fig. 46). The anterior five somites are in the recognition of some species (Table rounded dorsally and the sixth somite and 2). The ratio of dactylus length to carpus telson are flattened or slightly grooved in length in the fourth and fifth pereopods C. amurensis (Fig. 126). displays more noticeable differences than The thoracic sternum shows a specific the comparative length of the carpus of character in ovigerous females (Table 1). the third pereopod (Table 2). These ambu- Crangon dalli (Fig. 6i), C. cassiope (Fig. latory pereopods are the longest in C. lOi) and C. uritai (Fig. 15f) have a median affinis (Fig. 2e, f, g) and shortest in C. blunt process on the fifth thoracic stern- cassiope (Fig. 10f,g, h). The other five spe- ite, while C. affinis, C. propinquus, C. cies are intermediate with similar lengths hakodatei and C. amurensis have an (Figs. 4d, e, f, 6f, g, h, 8e, f, g, 12d, e, f, 15c, acute spine (Figs. 2h, 4g, 8h, 12g). Al- d, e). though the length of the spine is rather The largest specimens in all speices variable, it is usually the longest in C. are always females. The maximum size of amurensis (Fig. 12g), the shortest in C. females is distinctive in some species propinquus (Fig. 4g) and intermediate in (Table 1). The largest females are found C. affinis (Fig. 2h) and C. hakodatei (Fig. in C. dalli and C. hakodatei (more than 8h). The males and nonovigerous females 18.0 mm in CL), while the smallest belong usually have a moderately long median to C. affinis and C. uritai (less than 12.5 spine on the fifth sternite and a short me- mm in CL). Females of C. propinquus, C. dian spine or subacute tubercle each on cassiope and C. amurensis are intermedi- sixth to eighth sternites. These spines are ate in size, 15.9 mm, 14.3 mm and 15.8 usually longer in C. affinis (Fig. 2i), C. mm in CL, respectively. propinquus (Fig. 4h), C. hakodatei (Fig. The ventral surface of the antepenulti- 8i) and C. amurensis (Fig. 12/) than in C. mate segment of the third maxilliped is dalli (Fig. 6j), C. cassiope (Fig. 10/) and C. armed with a subdistal clump of spines uritai (Fig. 16c), though slightly variable (Figs. 66, c, 141,). The number of spines is in length. useful in the recognition of some species, The scaphocerite is always longer in though the character is somewhat vari- 05

Table 1. Comparison of the distinctive characters of seven species of the genus Crangon from East Asia.

Spines on Armature of Largest size Dorsal surface antepenultimate segment thoracic sternum Species in CL (mm) RL/CL Abdominal somites of telson of 3rd maxilliped of ovig. female

C. affinis 12.0 9 0.20-0.29 3rd rounded dorsally grooved 4-6, mostly 5 acute spine 9.0 6 4th carinate dorsally on posterior half 5th carinate dorsally 6th grooved dorsally, ventrally C. propinquus 15.9 9 0.15-0.20 3rd rounded dorsally grooved 4-5, mostly 5 acute spine 10.6 6 4th faintly carinate dorsally 5th carinate dorsally 6th grooved dorsally, ventrally C. dalli 18.0 9 0.16-0.22 3rd to 5th rounded dorsally grooved 5-7, mostly 6 tubercle 12.8 6 6th with 2 blunt submedian carinae dorsally C. hakodatei 18.4 9 0.17-0.24 3rd to 5th carinate dorsally grooved 4—6, mostly 5 acute spine 12.5 6 6th grooved dorsally, ventrally C. cassiope 14.3 9 0.13-0.19 3rd to 5th rounded dorsally rounded 4—6, mostly 6 tubercle 7.1 6 6th rounded dorsally, ventrally C. amurensis 15.8 9 0.13-0.19 3rd to 5th rounded dorsally rounded 3-5, mostly 4 acute spine 9.5 6 6th flattened or grooved dorsally, and grooved ventrally C. uritai 12.5 9 0.12-0.20 3rd to 5th rounded dorsally rounded 3—4, mostly 4 tubercle 9.9 6 6th rounded dorsally, and grooved ventrally Table 2. Meristic characters of seven species of Crangon from East Asia. The ranges with the mean and number of individuals examined in parentheses are given for each species.

Length of carpus/distal Dactylus/carpus Dactylus/carpus First palm two segments combined length of length of Species SL/CL length/width of 3rd pereopod 4th pereopod 5th pereopod

C. affinis 0.78-0.92 (0.86; 17) 9 3.11^.05 1.90-2.60 1.03-1.46 0.94-1.67 0.85-0.96 (0.90; 8) 8 (3.52; 14) (2.11; 14) (1.18; 15) (1.16; 16) W ra 25 CO C. propinquus 0.74-0.83 (0.80; 11) 9 2.46-3.00 1.41-1.87 0.80-1.03 0.82-1.13 I—I 0.79-0.84 (0.82; 5) 8 (2.76; 14) (1.75; 10) (0.94; 9) (0.97; 10) O O C. dalli 0.78-0.90 (0.84; 45) 9 2.24-2.91 1.46-2.10 0.92-1.21 0.92-1.23 "3 ffi 0.79-0.94 (0.85; 18) 8 (2.59; 33) (1.78; 34) (1.02; 23) (1.04; 23) M M C. hakodatei 0.69-0.82 (0.77; 19) 9 2.32-2.93 1.56-2.18 0.83-1.10 0.91-1.12 0.74-0.83 (0.80; 9) 8 (2.62; 18) (1.79; 18) (1.00; 14) (0.99; 14)

C. cassiope 0.67-0.79 (0.71; 14) 9 2.04-2.73 1.29-1.60 0.66-0.90 0.54-0.90 0.77-0.87 (0.82; 10) 8 (2.46; 15) (1.44; 11) (0.79; 15) (0.71; 14)

C. amurensis 0.71-0.81 (0.76; 22) 9 2.56-3.14 1.60-1.84 0.88-1.11 0.86-1.13 0.80-0.91 (0.84; 8) 8 (2.94; 22) (1.74; 14) (0.99; 20) (0.99; 20)

C. uritai 0.64-0.77 (0.67; 36) 9 2.65-3.33 1.43-2.05 0.89-1.19 0.88-1.13 0.74-0.90 (0.81; 13) 8 (2.91; 29) (1.68; 21) (1.06; 27) (1.05; 27)

CD 98 K. HAYASHI & J. N. KIM able and it overlaps in some cases (Table 6 1): 4-6, mostly 5, in C. affinis and C. 5. Sixth abdominal somite and telson flat- hakodatei; 4-5, mostly 5, in C. tened or slightly grooved dorsally. propinquus; 5-7, mostly 6, in C. dalli; 4- Thoracic sternum with acute median 6, mostly 6 in C. cassiope; 3-5, mostly 4, in spine in ovigerous females. [Palm of C. amurensis; and 3-4, mostly 4, in C. first pereopod 2.56-3.14 times as long uritai. as broad] Key to the East Asian species of C. amurensis Brashnikov, 1907 Crangon [Adult] 6. Sixth abdominal somite rounded ven- trally. Palm of first pereopod 2.04-2.73 1. Sixth abdominal somite with 2 distinct times as long as broad. Dactylus of dorsal submedian carinae fourth and fifth pereopods 0.54-0.90 C. dalli Rathbun, 1902 times as long as carpus 1. Sixth abdominal somite without dis- C. cassiope De Man, 1906 tinct submedian carinae 2 6. Six abdominal somite grooved ven- 2. Fifth abdominal somite with mid-dor- trally. Palm of first pereopod 2.65-3.33 sal carina 3 times as long as broad. Dactylus of fourth and fifth pereopods 0.88-1.19 2. Fifth abdominal somites without mid- times as long as carpus dorsal carina 5 C. uritai new species 3. Third to fifth abdominal somites with blunt mid-dorsal carina. [Palm of first Biogeography pereopod 2.32-2.93 times as long as Vertical distribution.—Crangon pro- broad. Carpus of third pereopod 1.56- pinquus, C. cassiope, C. amurensis and C. 2.18 times as long as distal two seg- uritai seem to be restricted to depths less ments combined] than about 40 m. C. amurensis and C. C. hakodatei Rathbun, 1902 uritai even occur in the intertidal zone. C. 3. Third abdominal somite without mid- affinis and C. hakodatei are inhabitants dorsal carina 4 of the continental shelf from sublittoral 4. Fourth abdominal somite with faint depths to about 200 m. C. dalli has the mid-dorsal carina. Palm of first pereo- broadest bathymetric range and extends pod 2.46-3.00 times as long as broad. to the greatest depth (3-630 m). Carpus of third pereopod 1.41-1.87 Geographical distribution. —Crangon times as long as distal two segments hakodatei has a relatively wide distribu- combined tion: the southern Sea of Okhotsk, the C. propinquus Stimpson, 1860 northern Pacific coast of Japan, the cen- tral and southern parts of the Sea of Ja- 4. Fourth abdominal somite with trace of pan, the Yellow Sea and the northern mid-dorsal carina on posterior half. East China Sea. C. dalli has the widest Palm of first pereopod 3.11-4.05 times distribution in the northern North Pa- as long as broad. Carpus of third cific, from Puget Sound to the Sea of Ja- pereopod 1.90-2.35 times as long as pan and the Chukchi Sea (Butler, 1980). distal two segments combined The remaining five species show a more C. affinis De Haan, 1849 limited distribution. C. propinquus and C. 5. Sixth abdominal somite and telson amurensis have a restricted distribution rounded dorsally. Thoracic sternum in the Sea of Okhotsk, the northern part with tubercle in ovigerous females of the Sea of Japan and the northern Pa- REVISION OF THE EAST ASIAN CRANGON 99

cific coast of Japan. C. uritai and C. (ed.), Dekapoden, part 7 in volume 2 of cassiope show a similar distribution in the Beitrage zur Naturgeschichte Ostasiens. Abhandlungen der Bayerischen Akademie Yellow Sea, the northern East China Sea, der Wissenschaften, Miinchen, 2 (supple- the central and southern parts of the Sea ment), 10: 1-101, 1 pi. of Japan and the Seto Inland Sea, but Bate, C. S., 1888. Report on the Crustacea there have been no records of C. cassiope Macrura dredged by H.M.S. Challenger from the Pacific coast of Japan. C. affinis during the years 1873-76. Report on the is restricted to the Yellow Sea (Fujino & Scientific Results of the Voyage of H.M.S. Challenger during the Years 1873-76, Zo- Miyake, 1970), the northeastern East ology, 24: xc + 942 pp., pis. 1-157. China Sea and the coast of Honshu, Ja- Brashnikov, V., 1907. Materiali po fauni pan. Russkikh vostochnikh morei, sovrannie shkhnoju "Storoz" vi 1899-1902 gg. [Mate- rials on the fauna of Russian Eastern Sea Acknowledgments collected by the schooner "Storoz" during We wish to express our sincere grati- the year 1899-1902]. Zapiski tude to many collegues for the loan of type Imperatorskoi Akademii Nauki, po phiziki- material or donation of specimens, on matematichekomu otdilenileniju, 20: 1- which the present paper is based: K. 185. Butler, T. H., 1980. Shrimps of the Pacific Amaoka, Y. R. An, K. Ariyama, H. K. Cha, coast of Canada. Canadian Bulletin of J. H. Choi, P. F. Clark, A. Crosnier, S. Fisheries and Aquatic Science, 202: 1-280. Goshima, T. Hamano, Y. Hanamura, N. Christoffersen, M. L., 1988. Genealogy and N. Ho, L. B. Holthuis, S. Y. Hong, N. phylogenetic classification of the world Horii, K. Ikeya, H. Itano, H. Itou, T. Crangonidae (Crustacea, Caridea), with a new species and new records for the south Kikuchi, M. H. Kim, S. H. Kim, K. western Atlantic. Revista Nordestina de Kojima, T. Komai, R. B. Manning, H. Biologi'a, 6: 43-59. Misu, H. Mukai, Y. Natsukari, K. Ogawa, Czerniavsky, V., 1884. Crustacea Decapoda J. Ohtomi, T. Okutani, H. Sando, J. Pontica littoralia. Materialia ad Sasaki, N. Takai, M. Toriumi, M. Zoographiam Ponticam comparatam. II. Travavaux de la Societe de l'Universite de Toriyama, J. Ueda, T. Ueno, Y. Ueta, M. Kharkov, 13 (supplment): 1-268, pis. 1-7. Yabe, H. Yamada, M. Yamada and K. Derjugin, K. M., & Kobjakova, Z., 1935. Zur Yokogawa. We are also much indebted to Decapodenfauna des Japanischen Meeres. K. Baba, P. Castro and L. B. Holthuis for Zoologicher Anzeiger, 112: 141-147. providing old references and A. L. Doflein, F., 1900. Die dekapoden Krebse der arktischen Meere. In: F. Romer and F. Rybakov and A. Anker for translations of Schaudinn, (eds.), Fauna Arctica. Eine the Russian references. We extend our Zusammenstellung der arktischen sincere thanks to P. Castro, T. Komai and Tierfrmen, mit besonderer C. L. McLay for kindly reviewing the Berscksichtigung, des Spitzbergen- manuscript. The senior author would par- Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nordliche ticularly like to thank L. B. Holthuis, Eismeer im Jahre 1898, 1: 313-362. C.H.J. M. Fransen and P. F. Clark for pro- Doflein, F., 1902. Ostasiatische Dekapoden. viding facilities to examine Japanese Abhandlungen der Bayerischen Akademie specimens deposited in their museums. der Wissenschaften, Miinchen, 21: 613- Part of this work was supported by a 670, pis. 1-6. Evans, A. C., 1967. Syntypes of Decapoda de- Grant from the Mikimoto Fund for Ma- scribed by William Stimpson and James rine Ecology. Dana in the collections of the British Mu- seum (Natural History). Journal of Natu- ral History, 1: 399^11. Literature Cited Fabricius, J. C., 1798. Supplementum Balss, H., 1914. Ostasiatische Decapoden II. Entomologiae Systematicae, pp. 1-572. Die Natantia und Reptantia. In: F. Doflein, [Not seen] 100 K. HAYASHI & J. N. KIM

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