Biogeography 2. 3-20. Aug. 28, 2000

Abyssobenthic Shrimps (Crustacea, , Penaeidea and ) from the Northwest Pacific Collected by RV Soyo-Maru

Jung Nyun Kim', Ken-Ichi HayashP, Yutaka Natsukari' and Katsuhiko Yoshida3

I Nagasaki University, 1-14 Bunkyo-machi, Nagasaki 852-8521, Japan (e-mail: [email protected] [J. N. Kim]; [email protected] [Yo Natsukari])

2 National Fisheries University, 2-7-1 Nagata-honmachi, Shimonoseki 759-6595, Japan (e-mail: [email protected])

3 National Research Institute of Fisheries Science, 2-12-4 Fukuura, Kanezawa-ku, Yokohama 236-8648, Japan (e-mail: [email protected])

Abstract. In the summer of seven years from 1987, the following five species of shrimps were collected from the Northwest Pacific at depths of 3,100-6,350 m: Hemipenaells spinidorsalis, Plesiopenaeus annatus, Benthesicymus crellatus, Sclerocrangon zenkevitchi and Neocrangon abyssorulll. Living in great depths these abyssobenthic shrimps have been rarely sampled. B. crenatus and S. zellkevitchi represent the fifth and second record since the original descriptions, respectively. The present material extends known bathymetric range of most species. Brief comments on distribution and color are given for all species.

Key words: abyssobenthic shrimps, Hemipenaeus spillidorsalis, Plesiopenaeus armatus, Benthesicymus crenatus, Sclerocrangon zenkevitchi, Neocrangon abyssorulI!, Decapoda, , Northwest Pacific.

1. Introduction 1881 , Plesiopenaeus armatus (Bate, 1881), Benthesicymus crenatus Bate, 1881 , Sclerocrangon The abyssal zone, occupying the depth range from zenkevitchi Birshtein & Vinogradov, 1953, 3,000 to 6,000 m, comprises a vast biotope that Neocrangon abyssorum (Rathbun, 1902). Of these, extends over two-thirds of the surface of the globe the two crangonid shrimps, S. zenkevitchi and N. (Vinogradova, 1997). However, the taxonomic' and abyssorum, are described in detail because their ecological information of the fauna of this zone is morphological characters and geographic range are incomplete because of the technical difficulties of incompletely known, especially the former represents sampling. the third record of this species. The two penaeideans, During a research of deep-sea environments in the H. spinidorsalis and P. armatus, are cosmopolitan and Northwest Pacific carried out by the RV Soyo-Maru rather well known, while B. crenatus was sporadically of Fisheries Agency of Japan, some rare collected in the Northwest Pacific. The present study abyssobenthic shrimps belonging to Penaeidea and provides the bathymetric range extensions of all Caridea were collected. The following five species species except for S. zenkevitchi. were identified: Hemipenaeus spinidorsalis Bate,

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2. Materials and Methods 3. Systematics

Samples were collected in the summer of 1987. Infraorder Penaeidea 1990-1994, 1996 at II stations in the Northwest Family Aristeidae Pacific (Fig. I). They were captured with the deep-sea Hemipenaeus spinidorsalis Bate, 1881 trap of Soyo-Maru type (Anonymous, 1988) and/or (Figs 2a-c, 8a) the benthos net (Table I). Samples were frozen on board and preserved in 70% buffered methyl alcohol Hemipenaeus spinidorsalis Bate, 1881 : 186; 1888: after photographs were taken in the laboratory. 301, pI. 44, fig. I; Faxon, 1895: 200, pI. 50, fig. 2; Specimens are deposited in the National Fisheries De Man, 1911 : 6 (list); Wasmer, 1972: 259; University, Shimonoseki (NFU). Crosnier, 1978: 80, figs 27b, 28c, d, 29b; 1986: Methods of measurements and terminology mainly 862, fig. 14a; Krygier & Wasmer, 1988: 49 (list), followed Perez Farfante & Kensley (1997) for 68; Perez Farfante & Kensley, 1997: 44, figs 13, penaeidean shrimps, and Butler (1980) and Komai & 14 (no new locality). Takeda (1989) for caridean shrimps. Hemypenaeus (sic) spinidorsalis: Kikuchi & Nemoto, 1986: 52.

Sea of Japan

35 0

0 30 o

250

1200 E 1250 1300 135 0 1400 1450 1500

Fig. I. Sampling stations in the Northwest Pacific carried out by RV Soyo-Maru.

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Table I. List of sampling data of abyssobenthic shrimps collected by the deep-sea trap of Soyo-Maru type, except for four stations with asterisk (*), in which benthos net was operated.

Station Position Depth (m) Data Species 87-Soyo SI. 4 22°59'N,124°IO'E 6350 4 Jun 1987 Benthesicymus crenatus 90-Soyo St. 4 24°IYN,144°4TE 5300 7 Jul 1990 Plesiopenaeus armatus Benthesicymus crenatus 91-Soyo St. I 28°24'N,134°58'E 4950 20 Jul1991 Plesiopenaeus armatus Benthesicymus crenatus 91-Soyo St. 2 30 0 04'N, 134°29'E 4600 23 Jul1991 Plesiopenaeus armatus Benthesicymus crenatus 91-Soyo St. 3 32°09'N, \35°06'E 4620 25 Jul1991 Benthesicymus crenatus *92-Soyo St. 3 41°IYN, 144°02'E 3100-3200 27 Jun 1992 Sclerocrangon zenkevitchi Neocrangon abyssorum *92-Soyo S1. 8 40°ll'N,149°43'E 5000-5200 9 Jul 1992 Benthesicymus crenatus *93-Soyo SI. 3 27°30'N,141°42'E 4100-4200 4,5 Jun 1993 Plesiopenaeus armatus *94-Soyo SI. I 41°29'N,147°03'E 5300-5400 19Jun 1994 Hemipenaeus spinidorsalis 94-Soyo St. 2 42°01 'N, 145°22'E 3900-4000 20 Jul 1994 Sclerocrangon zenkevitchi Neocrangon abyssorum 96-Soyo SI. 2 32°03'N,147°00'E 5800-5900 15 Jun 1996 Plesiopenaeus armatus

Material examined. 94-Soyo St. I; 16' (CL 43.8 sulcus reaching the middorsal line (Fig. 8a) (almost mm); NFU 530-2-2076. invisible in the Indian specimen), (2) 2 simple Coloration. Carapace and abdomen yellowish pinnulae on podobranch of third pereopod (instead of brown, margin of each abdominal somite slightly 6-9 in Crosnier's specimen), and (3) the dorsomedian darker. Rostrum and most appendages dark brown. lobule of the petasma with a wider cincinnuli part, Eye black, with reflecting pigments (Fig. 8a). which is about two-thirds of the mesial margin (Fig. Distribution. Known from the Indo-West Pacific, 2b, c) (nearly half in Crosnier's specimen). eastern Pacific and the South Atlantic (Perez Farfante H. spinidorsalis is readily distinguished from the & Kensley, 1997); 1,930-5,025 m (Crosnier, 1986), congener, H. carpenteri Wood-Mason, 1891 by the 5,300-5,400 m (present study). length of the rostrum. It usually overreaches the eye In the Northwest Pacific, H. spinidorsalis has been in H. spinidorsalis (Fig. 8a), but falling short of the listed once from off the central Japan (33°00'N, distal margin of the cornea in H. carpenteri. 156°30'E) by Kikuchi & Nemoto (1986). The present specimen confirms the distribution of this species in Plesiopenaeus armatus (Bate, 1881) the area and represents bathymetric range extensions (Figs 2d, e, 8b) more deeper. Remarks. Although the long spine on the third Aristeus armatus Bate, 1881: 188; 1888: 312, pis 45, abdominal somite is damaged (Fig. 2a), the present 46. material agrees well with the original and subsequent Aristaeopsis armata: Wood-Mason, 1891: 285. descriptions of Hemipenaeus spinidorsalis from the Arisleopsis armatus: De Man, 1911: 6 (list); South Atlantic and the West Pacific (Bate, 1881; Nakazawa, 1927: 996, fig. 1914 (no new locality); 1888). It, however, differs from Crosnier's (1978) Kubo, 1965: 596, fig. 899 (no new locality). Indian Ocean specimen by having the following Aristaeus (Aristaeopsis) armatus: Alcock, 1901: 41. minor particulars: (1) the weak but long cervical Plesiopeneus armatus: Faxon, 1895: 199.

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b c 5mm lOmm d

d e , lOmm 5mm

r•.

f g lOmm

Fig. 2. Three species of penaeidean shrimps. a-c, Hemipenaeus spinidorsalis Bate, 1881, male (eL 43.8 mm; NFU 530-2-2076) from 94-Soyo St. I (4ID29'N, 147D03'E); d, e, Plesiopenaeus armatus (Bate, 1881), female (CL 67.0 mm; NFU 530-2-2077) from 93-Soyo St. 3 (27D30'N, 14ID42'E); f, g, Benthesicymus crenatus Bate, 1881, male (CL 49.5 mm; NFU 530-2-1446) from 91-Soyo St. I (28°24'N, I 34°58 'E). a, d, second to fourth abdominal somites, lateral; b, right petasma, ventral; c, same, dorsal; e, left second pereopod, lateral; f, third to fifth abdominal somites, lateral; g, right third maxilliped, lateral.

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Plesiopenaeus armatus: Ramadan, 1938: 51; Roberts reaching at most only the midlength of the first & Pequegnat, 1970: 46; Wasmer, 1972: 259; segment in the other four specimens (CL 61.9-71.1 Crosnier & Forest, 1973: 294, fig . 99c-d; Crosnier, mm). Furthermore a small posteroventral spine on the 1978: 92, figs 31d, e, 32d-f, 33b; 1986: 863; Gore, pleura of each of the third to fifth abdominal somites 1985: 126; Kikuchi & Nemoto, 1986: 52; Krygier is usually present (Fig. 2d), but unarmed on either & Wasmer, 1988: 49 (list), 69; Hayashi, 1992: 33, side of the third or fourth somite in the two specimens fig. 20a-d (no new locality); Perez Farfante & (CL 70.8, 73 .3 mm). Kensley, 1997: 50, figs 19,20 (no new locality). The length ratio of the dactylus to the propodus of Aristeus? tridens Smith, 1884: 404, pI. 9, figs 1-6; the fourth pereopod is 0.47-0.54 in the present 1886: 698, pI. 19, figs 2, 2a. specimens versus 0.60 in the Indian specimens of P. Aristeopsis armatus var. tridens: Bouvier, 1908: 62, armatus reported by Crosnier (1978). pI. II, fig. 6; Milne Edwards & Bouvier, 1909: A strong posteromedian spine on each of the third 197, figs 20-27, pI. I, figs 4-7; De Man, 1911: 6 to fifth abdominal somites (Fig. 2d) and a movable (list). meral spine on the second pereopod (Fig. 2e) Aristeopsis tridens: Sund, 1920: 31. distinguish the present species from another member Plesiopenaeus armatus var. tridens: Ramadan, 1938: of the genus, P. eoruscans (Wood-Mason, 1891). 52. Family Material examined. 90-Soyo St. 4; I (CL 73.3 Benthesicymus crenatus Bate, 1881 mm); NFU 530-2-1452. 91-Soyo SI. I; I (CL 61.9 (Figs 2f, g, 8e) mm); NFU 530-2-1447. 91-Soyo SL 2; I (CL 70.8 mm); NFU 530-2-1449. 93-Soyo St. 3; (CL 67.0, Benthesieymus erenatus Bate, 1881: 190; 1888: 329, 71.1 mm); NFU 530-2-2077. 96-Soyo St. 2; I (CL pIs 54, 55; De Man, 1911: 5 (list); Crosnier, 1986: 67.8 mm); NFU 530-2-2078. 851, figs 6d, e, 7d, e, 8f, g; Kikuchi & Nemoto, Coloration. Body crimson entirely; gastric and 1986: 52; 1991: 67, figs 2, 3; Krygier & Wasmer, branchial regions of carapace, and dorsal and ventral 1988: 49 (list), 62. parts of third abdominal somite blackish; sixth Not ?Benthesicymus crenatus: Ohta, 1983: 229, photo abdominal somites somewhat paler. Eye with cornea 57 (= ?Benthesicymus laciniatus Rathbun, 1906). darkly pigmented; distal half of eyestalk dark brown (Fig. 8b). Material examined. 87-Soyo SL 4; 4J' (CL 22.9- Distribution. Known from the Pacific, Indian and 30.2 mm), 3 (CL 32.8-43.8 mm); NFU 530-2-1456. Atlantic Oceans (Perez Farfante & Kensley, 1997); 90-Soyo SI. 4; 2 J' (CL 31.0, 35.2 mm), 3 (eL 752-5,413 m (Crosnier, 1978),4,100-5,900 m (present 29.6-48.0 mm); NFU 530-2-1453. 91-Soyo St. I; I J' study). (CL 49.5 mm); NFU 530-2-1446. 91-Soyo SL 2; I J' P. armatus is widely distributed in three major (CL 51.2 mm); NFU 530-2-1448. 91-Soyo St. 3; I oceans, but it has been uncommon in the Northwest (CL 62.1 mm); NFU 530-2-1450. 92-Soyo SL 8; I Pacific (Bate, 1881; 1888; Kikuchi & Nemoto, 1986). (CL 49.8 mm); NFU 530-2-2079. The present material slightly extends the bathymetric Coloration. Body entirely red; gastric region of range. carapace, and dorsal part of first and second Remarks. As pointed out by Ramadan (1938) and abdominal somites yellowish. Cornea of eye darkly Roberts & Pequegnat (1970), the stylocerite varies in pigmented (Fig. 8e). length also in the present material; it exceeds the Distribution. Known from the Northwest Pacific distal margin of the second segment of the antennule and central Pacific (Perez Farfante & Kensley, 1997); in the two specimens (CL 67.0, 73.3 mm), but 3,530-5,700 m (Kikuchi & Nemoto, 1991), 4,600-

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6,350 m (present study). specimen as B. laciniatus than B. erenatus, because The present material slightly extends the the bathymetric range is clearly separated, shallower bathymetric range downward. This species is than 3,500-4,000 m in B. laeiniatus (e.g., Crosnier, relatively common and widely distributed in the 1986), while deeper in B. erenatus. abyssal zone of the Northwest Pacific (Table 1). Remarks. In the genus Benthesieymus, 10 species Infraorder Caridea have been reported from the Northwest Pacific (Perez Family Fart·ante & Kensley, 1997). B. erenatus is Sclerocrangon zenkevitchi Birshtein & distinguishable from the congeners by having the Vinogradov, 1953 hepatic spine (Fig. 8e), the pectinated posterior (Figs 3-5, 8d. e) margins of the fourth abdominal somite (Fig. 2.1), and an acute subdistal spine on the merus and ischium of Scleroerangon zenkevitehi Birshtein & Vinogradov, the third maxilliped (Fig. 2g). 1953: 218, figs 2-5; Sokolova, 1957: 270 (no new Based on photograph observation, Ohta (1983) locality); 1997: 446 (list), 448 (list). reported B. erenatus with some doubt from Sagami Scleroerangon zenkewitehi (sic): Zarenkov, 1993: 17. Bay at a depth of 2,830 m. As mentioned above, the depth is too shallow to refer his specimen to that Material examined. 92-Soyo St. 3; 1 (CL 24.3 species. It is more reasonable to consider Ohta's mm); NFU 530-2-2080. 94-Soyo St. 2; 12 J' (11.9-

Fig. 3. Scleroerangon zenkevitehi Birshtein & Vinogradov, 1953, ovigerous female (CL 28.0 mm; NFU 530-2-2081) from 94-Soyo St. 2 (42°01 'N, I 45°22'E). Entire in lateral view (top) and dorsal view (bottom).

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20.3 mm), 10 'f. (CL 18.3-29.1 mm), 3 ovig. 'f. (CL part to posterior margin and each with acute spine on 27.3-29.9 mm); NFU S30-2-2081. ventral margin, longitudinal carina of fifth terminating in small spine. Sixth somite 0.4S-0.58 Description of females. Body (Fig. 3) robust, times as long as carapace, dorsally with two slightly depressed dorsoventrally. Integument hard, submedian carinae, each extending beyond posterior moderately sculptured, covered with velvet-like margin as acute spine; lateral surface with carina tomenta. curved inferiorly; posteroventral spine moderately Rostrum (Fig. 3) very long, sharply pointed, strong; posterolateral process strong, sharply pointed slightly compressed laterally, 1.19-2.01 times as long and supported by short carina. Telson (Figs 4n, Sj) as carapace, curved upward, with some bristles on 0.73-0.94 times as long as carapace, tapering into dorsal and ventral margins near base; distinct lateral acute tip, with two dorsal carinae, sulcate medially; carina on basal one-third; postorbital carina continued three pairs of small spines on posterior one-third of from rostral lateral carina curved slightly downward lateral margin (Fig. 4n), but in one specimen (CL 24.3 just behind antennal spine, and extending posteriorly mm) anterior two pairs situated on dorsolateral to midlength of carapace. Carapace (Fig. 3) 1.11-1.51 margin (Fig. Sj). times as long as wide; median carina high, armed with Four posterior thoracic sternites each with two anteriorly curved strong spines, anterior spine subacute median process diminishing in size with small, low tubercle on dorso-basal margin, posteriorly in non-ovigerous specimens; these posterior spine smaller than anterior one; median processes except for that on fifth somite reduced in carina bifurcated posteriorly, forming transverse ovigerous specimens. Anterior five abdominal carina paralleled with posterior margin of carapace sternites in immature specimens each with median and turning anteriorly to join with branchial carina; spine but un armed in mature specimens; sixth sternite branchial carina extending from hepatic spine; always with median tubercle; preanal spine absent. subbranchial carina beginning at inferior margin of Eye (Fig. 4a) with cornea well developed; eyes talk branchiostegal spine, extending near posterior margin provided with dorsal tubercle and small ventral spine. of carapace, eroded with many large pores; ventro- Antennule (Fig. 4b) with peduncle overreaching lateral margin of carapace sharply ridged; antennal midlength of scaphocerite; proximal segment longer spine slender, very sharply pointed, reaching almost than distal two segments combined, with acute spine distal margin of cornea; branchiostegal spine very on ventral margin; stylocerite slightly overreaching strong, flared antero-laterally, reaching near midlength of proxi mal segment, sharply pointed midlength of scaphocerite; hepatic spine moderately distally; outer flagellum more stout than inner strong, followed by deep furrow along inferior part of flagellum. its base; pterygostomian spine minute. Antenna (Fig. 4c) with scaphocerite, 0.S5-0.70 Abdomen (Fig. 3) with well developed times as long as carapace, 2.38-3.39 times as long as dorsomedian carinae on first to fifth somites; median wide, lateral margin straight or slightly concave, with carina on first somite produced anterodorsally as distolateral spine not exceeding distal margin of hooked, acutely pointed spine; that of second somite blade; basicerite bearing blunt lateral spine; truncate or forming light angle anteriorly; that of fifth carpocerite slightly curved laterally, falling short of somite bifurcate posteriorly, each terminating in acute blade of scaphocerite. spine on posterodorsal margin. Pleura of first to third Mouthparts similar to other crangonids as figured somites each with blunt marginal carina and short (Fig. 4d-j); third maxilliped (Fig. 4i, j) depressed carina or ridge on central part, and each with acute dorsoventrally, exceeding scaphocerite by one-third spine on ventral margin; those of fourth and fifth length of ultimate segment; ultimate segment (Fig. 4j) somites each with longitudinal carina from central armed with blunt spine at tip; antepenultimate

-9- Abyssobenthic Shrimps from the Northwest Pacific Collected by RV Soyo-Maru b'

adefgh j ! 5mm

bciklmn J lOmm '----' Imm n

Fig. 4. Sclerocrangon zenkevitchi Birshtein & Vinogradov, 1953, ovigerous female (CL 28.0 mm; NFU 530-2-2081) from 94-Soyo St. 2 (42°01'N, 145°22'E). Appendages dissected from left side. a, eye, lateral; b, antennule, ventral; c, antenna, ventral; d, mandible, external; e, maxillule, external; 1, maxilla, external; g, first maxilliped, external; h, second maxilliped, external; i, third maxilliped, flexor; j, same, detail of distal part of ultimate segment, flexor; k, first pleopod, ventral, exopodal setae omitted; I, second pleopod, ventral, exopodal setae omitted; m, uropod, dorsal; n, tel son, dorsal.

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abcegi lOmm

d{h lmm

J .

j l m n k 5mm lmm 5miIl'

Fig. 5. Sclerocrangon zenkevitchi Birshtein & Vinogradov, 1953. a-i, ovigerous female (CL 28.0 mm; NFU 530-2-2081) from 94-Soyo St. 2 (42°01'N, 145"22'E);j, female (CL 24.3 mm; NFU 530-2- 2080) from 92-Soyo St. 3 (41°15'N, I 44°02'E); k-n, male (CL 19.1 mm; NFU 530-2-2081) from 94-Soyo St. 2 (42°01 'N, 145"22'E). Appendages dissected from left side. a, first pereopod, lateral; b, same, chela, flexor; c, second pereopod, lateral; d, same, fingers, lateral; e, third pereopod, lateral; f, same, distal part of dactylus, lateral; g, fourth pereopod, lateral; h, same, distal part of dactylus, lateral; i, fifth pereopod, lateral; j, telson, dorsal; k, anterior part of carapace and cephalic appendages, dorsal; I, first pleopod, ventral, exopodal setae omitted; m, second pJeopod, ventral, exopodal setae omitted; n, same, endopod and appendix masculina, ventral.

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segment slightly shorter than distal two segments masculina (Fig. 5n) longer than endopod, dorsally combined; coxa with sllbrectangular process on lateral with stout setae. surface; exopod with well developed lash. Coloration. Reddish brown or tan all over body; First pereopod (Fig. 5a, b) slightly overreaching carinae and spines on carapace and abdomen slightly distal margin of antennular peduncle; chela (Fig. 5b) darker. Eye black, with reflecting pigments (Fig. 8d, with palm 2.80-3.42 times as long as wide, cutting e). edge almost transverse and feebly convex; carpus Variation. Females are considerably larger than short, armed with two lateral spines; merus with males: the largest female is CL 29.9 mm and the strong dorsodistal spine. Second pereopod (Fig. 5c, cI) largest male is CL 20.3 mm. chelate, falling short of distal margin of first chela; Proportional length of the rostrum to the carapace chela (Fig. 5c1) with dactylus 0.26-0.38 times as long length tends to decrease with growth, ranging from as palm, fingers with numerous minute spinllies on 2.01 in CL 11.9 mm to 1.19 in CL 29.9 mm. both cutting edges; carpus 1.27-1.97 times as long as The tel son spines are usually situated on the lateral chela; coxa with wing-like process. Third pereopod margin (Fig. 4n), but in one female (CL 24.3 mm; (Fig. 5e, f) slightly more slender than second NFU 530-2-2080) they are on the dorsolateral margin pereopod, reaching distal margin of first pereopod; (Fig. 5). dactylus (Fig. 5f) short, 0.36-0.65 times as long as Distribution. Known from the Northwest Pacific propodus, with fine marginal setae; carpus 1.09-1.62 and the Bering Sea (Zarenkov, 1993); 2,995-4,070 m times as long as distal two segments combined. (Zarenkov, 1993),3, I 00-4,000 m (present study). Fourth pereopod (Fig. 5g, h) moderately stout, failing No abyssobenthic shrimp has been reported from slightly short of distal margin of first pereopod; the Sea of Japan. Although Zarenkov (1993) reported dactylus (Fig. 5h) 0.54-0.71 times as long as S. zenkevilchi from the Kuril Trench and the Sea of propodus, subspatulate, fringed with row of setae Japan, judging from the given sampling data, the along both lateral and mesial margins; carpus 0.57- locality attributed to "the Sea of Japan" should be 1.07 times as long as dactylus. Fifth pereopod (Fig. correctly referred to "the Japan Trench." 5i) similar to fourth pereopod. Remarks. The present material, representing the Pleurobranchs present on fourth to eighth thoracic third record of this poorly known species, agrees well somites. with the original description from the Bering Sea Endopod of first pleopod (Fig. 4k) reaching about (Birshtein & Vinogradov, 1953) in having the midlength of exopod. Second pleopod (Fig. 41) characteristic, very long, unarmed rostrum, two similar to first pleopod but exopod broader than that median spines on the carapace, and one ventral spine of first pleopod. on each pleuron of the anterior five abdominal Uropod (Fig. 4m) with rami almost equal in somites. length, falling short of posterior end of telson. Birshtein & Vinogradov (1953: 220, fig. 3) Eggs large, globular, 2.70 X 3.22 mm in diameter. observed many specimens with very short rostrum Differences in males. Males differ from females and downcurved branchiostegal spine, which were in the following particulars. collected from a comparatively shallower depth of Outer antennular flagellum (Fig. 5k) longer and 2,995 m. These variations are not observed in the stouter than in females, overreaching scaphocerite by present material from around 4,000 m. This fact may distal half. support Birshtein & Vinogradov's (1953) hypothesis First pleopod (Fig. 51) with endopod shorter and that these variations occur especially more frequently sllbrectangular. Second pleopod (Fig. 5m, n) with near the upper frontiers at a depth of about 3,000 m shorter endopod, fringed with setae; appendix than near the lower ones.

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Neocrangon abyssorum (Rathbun, 1902) Description of female. Body (Fig. 6) slender, (Figs 6, 7, 8f) slightly depressed dorsoventrally. Integument thin, almost naked. Crangon abyssorum Rathbun, 1902: 890; 1904: 125, Rostrum (Fig. 6) 0.30 times as long as carapace, fig. 66; Butler, 1980, unnumbered fig.: 112; falling slightly short of anterior margin of cornea, Krygier & Pearcy, 1981: 89. slightly ascending, very narrow, terminating in acute Crangon (Crangon) abyssorum: De Man, 1920: 249 tip. Carapace (Fig. 6) with median carina extending (list). from rostral apex almost to posterior four-fifths of Crago abyssorum: Schmitt, 1921: 97, fig. 65 (no new carapace, bearing two median spines, posterior spine locality). at anterior one-third of carapace, minute anterior Sclerocrangon abyssorum: Birshtein & Vinogradov, spine situated at midpoint between orbital margin and 1951: 359; 1953: 216; Sokolova, 1957: 271 (no posterior spine; gastric region slightly depressed; new locality) ; 1997: 446 (list), 448 (list). postorbital carina distinct, continuous with Crangon (Neocrangon) abyssorum: Zarenkov, 1965: dorsolateral margin of rostrum, extending beyond 1762 (list); Birshtein & Zarenkov, 1970: 422. level of posterior end of median carina; antennal spine Neocrangon abyssorum: Kuris & Carlton, 1977: 554 supported by distinct carina continuous with hepatic (list); Komai, 1991: 81, fig. 8; 1994: 98 (list). spine; hepatic spine moderately strong, supported by Not? Crangon abyssorum: Yokoya, 1933: 34, fig. 14 short carina, situated at level of anterior median spine; (see "Remarks"). branchiostegal spine supported by distinct carina; pterygostomian spine obsolescent. Material examined. 92-Soyo St. 3; 1'f. (CL 11.7 Anterior four abdominal somites (Fig. 6) rounded mm); NFU 530-2-2082. 94-Soyo St. 2; 1'f. (CL 12.2 dorsally; fifth somite with low, blunt carina dorsally; mm); NFU 530-2-2083. pleura of anterior five somites without marginal

Fig. 6. Neocrangon abyssorum (Rathbun, 1902), female (CL 12.2 mm; NFU 530-2-2083) from 94- Soyo St. 2 (42°01'N, 145°22'E). Entire animal in lateral view (top) and dorsal view (bottom).

-13- Abyssobenthic Shrimps from the Northwest Pacific Collected by RV Soyo-Maru a d

cdefg 2mm k fl

J

imo n 2 mm ___ o

abqr 5mm

Fig. 7. Neocrangon abyssorum (Rathbun, 1902), female (eL 12.2 mm; NFU 530-2-2083) from 94- Soyo St. 2 (42°01 'N, Appendages dissected from left side. a, antennule, ventral; b, antenna, ventral; c, mandible, external; d, maxillule, external; e, maxilla, external; f, first maxilliped, external; g, second maxilliped, external; h, third maxilliped, flexor, distal part of ultimate segment missing; i, same, distal part of antepenultimate segment, flexor; j, first pereopod, lateral; k, same, chela, flexor; /, second pereopod, lateral; m, same, fingers, lateral; n,

fourth pereopod, lateral; 0, same, distal part of dactylus, lateral; p, fifth pereopod, lateral; q, first pleopod, ventral; r, second pleopod, ventral.

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Fig. 8. Five abyssobenthic shrimps from the Northwest Pacific. a, Hemipenaeus spinidorsalis Bate, 1881, male (eL 43.8 mm); b, Plesiopenaeus armatus (Bate, 1881), female (eL 71.1 mm); c, Benthesicymus crenatus Bate, 1881, male (eL 49.8 mm); d, Sclerocrangon zenkevitchi Birshtein & Vinogradov, 1953, female (eL 22.3 mm); e, same, male (eL 19.2 mm); f, Neocrangon abyssorum (Rathbun, 1902), female (eL 12.2 mm).

-15- Abyssobenthic Shrimps from the Northwest Pacific Collected by RV Soyo-Maru spine; sixth somite 0.74 times as long as carapace, merus with moderately strong dorsodistal spine and 3.21 times as long as posterior height, dorsal surface weak lateral spine, flexor surface unarmed. Second with distinct paired carinae, not extending posterior pereopod (Fig. 7l, m) slender, chelate, falling short of margin; lateral surface with low carina on anterior distal margin of first pereopod; dactylus (Fig. 7m) half, posterolateral process terminating in acute spine. 0.32 times as long as palm, both fingers setose; coxal Telson (Fig. 6) with median dorsal sulcus; distal part process well developed. Third pereopod as slender as broken and missing. second; propodus and dactylus broken and missing. Thoracic and abdominal sternites showing typical Fourth pereopod (Fig. 7n, 0) slender but stouter than condition of non-ovigerous but spawning female. third pereopod, overreaching distal margin of Posterior thoracic sternites deeply concave, without scaphocerite; dactylus (Fig. 70) slightly flattened, median process. Anterior five abdominal sternites 0.60 times as long as propodus, 0.77 times as long as each with median tubercle; sixth somite with preanal carpus, terminating in thin process with several fine spine. setae; propodus setose on extensor surface; merus Eye (Fig. 6) very large, cornea well developed; with scattered setae on both ventral and dorsal mesial surface flat and contiguous with each other. margins. Fifth pereopod (Fig. 7p) equal or slightly Antennule (Fig. 7 a) with peduncle reaching shorter than fourth pereopod; dactylus 0.55 times as proximal one-third length of scaphocerite; proximal long as propodus, 0.69 times as long as carpus; merus segment longer than distal two segments combined, less setose than in fourth pereopod. ventromesial ridge with acute spine; stylocerite rather Pleurobranchs on fourth to eighth thoracic narrow, falcate, terminating in acute tip, slightly somites, apices directed backward. falling short of anterior margin of proximal segment; First pleopod (Fig. 7 q) with endopod reaching outer flagellum slender, slightly overreaching blade of about half of exopod; endopod rounded at distal scaphocerite; inner flagellum slightly longer and more margin. Second pleopod (Fig. 7r) with endopod slender than outer one. nearly reaching distal margin of exopod; endopod Antenna (Fig. 7b) with scaphocerite 0.75 times as strongly curved mesially. long as carapace and 4.50 times as long as wide, Uropod (Fig. 6) with exopod armed with small lateral margin concave proximally, distolateral spine spine just mesial to acute posterolateral tooth; overreaching distal margin of blade; basicerite with endopod slightly overreaching posterior margin of strong lateral spine; carpocerite reaching proximal exopod. two-thirds length of scaphocerite. Coloration. Body entirely purplish brown; Mouthparts similar to other crangonids as figured anterior part of carapace, telson and uropod darker (Fig. 7c-i); third maxilliped (Fig. 7h, i) slender, than the other parts. Eye black, with reflecting overreaching scaphocerite by one-third length of pigments (Fig. 8f>. ultimate segment; ultimate segment shorter than Distribution. Known from the northern North penultimate segment; antepenultimate segment Pacific from southern California to the Pacific coast slightly shorter than distal two segments combined, of Hokkaido, Japan (KomaL 1991); 887-2,975 m ventral surface with two small subdistal spines; coxa (Komai, 1991),3,100-4,000 m (present study). with well developed process on lateral side; exopod This report provides a distinct bathymetric range well developed; arthrobranch absent. extension of the present species. First pereopod (Fig. 7j, k) falling slightly short of Remarks. Neocrangon abyssorum has been blade of scaphocerite; chela (Fig. 7k) with palm 3.50 recorded twice in Japanese waters (Yokoya, 1933; times as long as wide, cutting edge moderately Komai, 1991). As pointed out by Komai (1991), who oblique; dactylus not overreaching base of fixed examined one immature specimen from east of Cape finger when closed; carpus bearing two lateral spines; Erimo, Hokkaido, Yokoya's (1933) Crangon

-16- lung Nyun Kim, Ken-Ichi Hayashi, Yutaka Natsukari I and Katsuhiko Yoshida

abyssorum from the Pacific coast of Japan differs five species agrees well with her opinion. S. from that species. Fortunately Yokoya's (1933) zenkevitchi, which has the narrowest vertical range specimens, one male (CL 7.3 mm) and one juvenile (2,994-4,100 m), shows a rather restricted horizontal (CL 4.4 mm), deposited in Kitakyushu Museum of distribution as mentioned above. The cosmopolitan H. Natural History are reexamined. Although the spinidorsalis and P. armatus have the widest vertical specimens severely damaged, following characters, ranges from about 1,000 or 2,000 m to 5,500-6,000 m. which are distinguished from the present species, are B. crenatus and N. abyssorum, with intermediate retained: more flattened rostrum, the posterior median vertical ranges of 3,530-6,350 m and 887-4,000 m, spine situated nearly at the middle of the carapace, respectively, have a relatively wide distribution in the somewhat longer scaphocerite (0.95 times as long as West Pacific or North Pacific. carapace versus 0.75 times), a posteromarginal spine These distributional pattern was primarily affected on the pleura of the fifth abdominal somite and by surface water characteristics, principally climatic shallower bathymetric range (97-547 m as against zonation (Vinogradova, 1997). In addition to the 887-4,000 m). They certainly represent an physical factors, it seems to depend on the life history undescribed species of Neocrangon. The distinct strategy and the movement ability of the shrimps. The specific status, however, is needed more additional penaeidean shrimps have usually longer larval stages specimens. (Williamson, 1982) and undoubtedly stronger swimming capability (Wasmer, 1972; Gore, 1985; 4. Discussion Hayashi, 1992) than the caridean shrimps. Haynes (1985) estimated that the crangonid caridean shrimps Except for the two cosmopolitan penaeideans, B. have 1-7 zoeal stages. Especially S. zenkevitchi has no crenatus and S. zenkevitchi have been known as the pelagic larvae (Birshtein & Vinogradov, 1953), true abyssal species and N. abyssorum is confirmed though 2 larval stages (Haynes, 1985). Postulated for the first time to inhabit the abyssal zone. The from the larval stages of N. communis (e.g. Haynes, cosmopolitan H. spinidorsalis and P. annatus are 1985), N. abyssorum seems to have about 5 zoeal components of the lower bathyal to abyssal zones. stages. Simultaneously, N. abyssorum may forage into On the basis of the faunistic structure, the abyssal the water column, since it had been collected by a zone of the Northwest Pacific is divided into two midwater trawl at 1,250-1,500 m, a distance of over provinces, the North-Pacific abyssal province and the 1,500 m above the ocean floor (Krygier & Pearcy, West-Pacific abyssal province, with a boundary line 1981). Therefore, N. abyssorum has a wider at about 400N (Vinogradova, 1997). B. crenatus is one distribution than S. zenkevitchi. of the typical examples of the West-Pacific abyssal Three penaeidean shrimps, H. spinidorsalis, P. province. On the other hand, S. zenkevitchi and N. armatus and B. crenatus, have often been reported as abyssorum are the representatives of the North-Pacific benthic or benthic-associate (e.g. Roberts & abyssal province but the former is only recorded from Pequegnat, 1970; Wasmer, 1972; Crosnier, 1978; the western part of that province, such as the western Gore, 1985; Krygier & Wasmer, 1988), but Bering Sea, the Kuril-Kamchatka Trench and the sometimes pelagic (Kikuchi & Nemoto, 1986). Japan Trench. B urkenroad (1937) poi nted out that "the term Vinogradova (1997) suggested that higher taxa benthonic when applied to Decapoda Natantia, can containing a large number of species with a wide refer at most to a vital dependence upon the bottom, vertical range (eurybathic forms) have a wider rather than to an entirely substratal existence." The horizontal distribution than those dominated by present material, which was mostly captured by a species with narrow vertical ranges (stenobathic benthic bait trap (Anonymous, 1988), may support abyssal forms). The distribution pattern of the present that these species are primarily benthic inhabitants of

-17- Abyssobenthic Shrimps from the Northwest Pacific Collected by RV Soyo-Maru the lower bathyal to abyssal zones. Kamchatka Trench area. In Bogorov V. G. (ed.), Fauna of the Kurile-Kamchatka Trench and its Acknowledgments environment. Trudy Inst. Okeanol., 86: 420-426. (In Russian.) We extend our sincere thanks to Dr J. S. Ho of Bouvier, E. L. 1908. Crustaces Decapodes (Peneides) California State University and anonymous reviewers provenent des compagnes de I'Hironde\le et de la for critical reading and valuable suggestions of the Princesse Alice. (1886-1907). Result. Camp. manuscript. We are also much indebted to Mr A. scient. Monaco, 33: 1-122, pis. 1-16. Anker of Museum national d'Histoire naturelle, Paris Burkenroad, M. D. 1937. Some remarks on the for translations of the Russian literature and Dr K. structure, habits, and distribution of the benthonic Konishi of National Research Institute of sergestid Sicyonella Borradaile (Crustacea, Aquaculture, Nansei for information about some Oecapoda). Ann. Mag. nat. Hist., (10) 19: 505- Ii terature. 514. Butler, T. H. 1980. Shrimps of the Pacific coast of References Canada. Can. Bull. Fish. Aquat. Sci., 202: 1-280. Crosnier, A. 1978. Crustaces Decapodes, Peneides Anonymous. 1988. Shinkaisei soko-uorui no saishuho Aristeidae (Benthesicyminae, Aristeinae, to shinkaiseigyo no houshasei kakushu noushuku Solenocerinae). Faune Madagascar, (46): 1-197. katei no minaoshi [Sampling method of the deep- Crosnier, A. 1986. Crevettes peneides d'eau profonde sea benthic fishes and reconsideration to recoltees dans I' ocean Indien lors des compagnes concentration process of radio nuclei in deep sea Benthedi, Safari I et II, Md 32/Reunion. Bull. fish]. Saishin Gijutsu Jouhou Series, Suisangyou- Mus. natn. Hist. nat. Paris, 4 (7) A: 839-877. hen, 1988: 19-20. Crosnier, A. & Forest, J. 1973. Les crevettes Alcock, A. 1901. A descriptive Catalogue of the profondes de I' Atlantique oriental tropical. Faune Indian deep-sea Crustacea Decapoda Macrura and trop., 19: 1-409. Anomala in the Indian Museum. Being a revised Faxon, W. 1895. The stalk-eyed Crustacea. Reports Account of the deep-sea Species collected by the on an exploration off the west coasts of Mexico, Royal Indian marine Survey Ship Investigator. 1- Central and South America, and off the Galapagos 286, pis 1-3. Calcutta. Islands, in charge of Alexander Agassiz, by the U. Bate, C. S. 1881. On the Penaeidea. Ann. Mag. nat. S. fish commission steamer "Albatross", during Hist., (5) 8: 169-196. 1891, Lieut.-Commander Z. L. Tanner, U. S. N., Bate, C. S. 1888. Report on the Crustacea Macrura commanding. XV. Mem. Mus. compo Zoo I. Harv., dredged by H.M.S. Challenger during the years 18: 1-292, pIs A-K, I-57. 1873-76. Rep. scient. Res. Voy. H.M.S. Gore, R. H. 1985. Abyssobenthic and abyssopelagic Challenger, Zool., 24: 1-942, pis 1-150. penaeoidean shrimps (families Aristeidae and Birshtein, Y. A. & Vinogradov, L. G. 1951. New and Penaeidae) from the Venezuela Basin, Caribbean rare Decapoda in Okhotsk Sea and Kurilian Sea. Crustaceana, 49: 119-138. waters. Dok!. Akad. Nauk SSSR, 79: 357-360. (In Hayashi, K. 1992. Russian.) from Japanese Waters. 300 pp. Seibutsu Birshtein, Y. A. & Vinogradov, L. G. 1953. New data Kenkyusha, Tokyo. (In Japanese.) on the decapod fauna in the Bering Sea. Haynes, E. 1985. Morphological development, Zool. Zh., 32: 215-228. (In Russian.) identification, and biology of larvae of Pandalidae, Birshtein, Y. A. & Zarenkov, N. A. 1970. Bottom Hippolytidae, and Crangonidae (Crustacea, decapods (Crustacea, Decapoda) of the Kurile- Decapoda) of the northern North Pacific Ocean.

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Fish. Bull. U.S., 83: 253-288. new species, Crangon handi, and new genus, Kikuchi, T. & Nemoto, T. 1986. List of pelagic Lissocrangon, of crangonid shrimps (Crustacea: shrimps (Crustacea, Decapoda) from the western Cm'idea) from the California coast, with notes on North Pacific. Bull. biogeogr. Soc. Japan, 41: 51- adaptation in body shape and coloration. BioI. 59. Bull., 153: 540-559. Kikuchi, T. & Nemoto, T. 1991. Deep-sea shrimps of Man, 1. G. de. 1911. Family Penaeidae. The the genus Benthesicymus (Decapoda: Decapoda of the Siboga Expedition. Part I. Siboga Dendrobranchiata) from the western North Pacific. Exped., 39a: 1- I 3 I. 1. Crust. BioI., 11: 64-89. Man, 1. G. de. 1920. Decapoda of the Siboga Komai, T. 1991. Deep-sea decapod crustaceans from Expedition. IV. Families Pasiphaeidae, the Pacific coast of eastern Hokkaido, northern Stylodactylidae, Hoplophoridae, Nematocarcinidae, Japan (Crustacea, Decapoda, Penaeidea and Thalassocaridae, Pandalidae, Psalidopodidae, Caridea). Kita-Nihon Teigyobu Kaihou, (24): 55- Gnatophyl1idae, Processidae, Crangonidae and 96. Glyphocrangonidae. Siboga Exped. 39a3: 1-3 I 8. Komai, T. 1994. Nihonkai no rikutanasei koebirui Milne Edwards, A. & Bouvier, E. L. 1909. Les (Tarabaebika, Moebika, Ebijakoka) no Peneides et Stenopides. Reports on the results of bunruigakuteki gaiyou [Taxonomic synopsis of dredging, under the supervision of Alexander Caridea (Pandalidae, Hippo1ytidae, Crangonidae) Agassiz, in the Gulf of Mexico (1877-78), in the occurred on continental shelf of the Sea of JapanJ. Caribbean Sea (1878-79), and along the Atlantic Contr. Fish. Res. Japan Sea Block, 31: 81-107. (In coast of the United States (1880), by the U. S. Japanese.) coast survey steamer "Blake", Lieut.-Com. C. D. Komai, T. & Takeda, M. 1989. Sclerocrangon Sigsbee, U. S. N., and Commander J. R. Bartlett, unidentata, a new crangonid shrimp from the U. S. N., commanding. XLIV. Mem. Mus., compo Pacific coast of Honshu, Japan (Crustacea: Zool. Harv., 27: 177-274, pIs. 1-9. Decapoda). Bull. biogeogr. Soc. Japan, 44: 77-84. N akazawa, T. 1927. Decapoda. In Illustration of Krygier, E. E. & Pearcy, W. G. 1981. Vertical Japanese Zoology: 992-1124. Tokyo. (In distribution and biology of pelagic decapod Japanese.) crustaceans off Oregon. J. Crust. BioI., I: 70-95. Ohta, S. 1983. Photographic census of large-sized Krygier, E. E. & Wasmer, R. A. 1988. Zoogeography benthic organisms in the bathyal zone of Suruga of pelagic shrimps (Natantia: Penaeidea and Bay, central Japan. Bull. Ocean Res. Inst., Univ. Caridea) in the North Pacific Ocean (with Tokyo, 15: 1-244. synopses and key to the species of the Subarctic Perez Farfante, I. & Kensley, B. 1997. Penaeoid and and Transitional Zones). In Nemoto T. and Pearcy sergestoid shrimps and prawns of the world-keys W. G. (eds), The biology of the Subarctic Pacific. and diagnoses for the families and genera. Mem. Proceedings of the Japan-United States of America Mus. natn. Hist. nat. Paris, 175: 1-233. seminar on the biology of micronekton of Ramadan, M . M. 1938. Crustacea: Penaeidae. John Subarctic Pacific, part I. Bull. Ocean Res. Inst., Murray Exped., 1933-34. Scient. Rep., 5(3): 35- Univ. Tokyo, 26: 43-98. 76. Kubo, I. 1965. Macrura. In Okada Y. K., Uchida S., Rathbun, M. J. 1902. Description of new decapod Uchida T. and others, New illustrated crustaceans from the west coast of North America. Encyclopedia of the Fauna of Japan. part 2: 592- Proc. U. S. natn. Mus., 24: 885-905. 629, figs 891- 1031. Hokuryukan, Tokyo. (In Rathbun, M. J. 1904. Decapod crustaceans of the Japanese.) northwest coast of North America. Harriman Kuris, A. M. & Carlton, J. T. 1977. Description of a Alaska Exped., 10: 1-21 O.

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Rathbun, M. J. 1906. The Brachyura and Macrura of abyssal and hadal zones. Adv. mar. BioI., 32: 325- the Hawaiian Islands. Bull. U. S. Fish Commn, 23: 383. 827-930, pIs 1-24. Wasmer, R. A. 1972. New records for four deep-sea Roberts, T. W. & Pequegnat, W. E. 1970. Deep-water shrimps from the northeastern Pacific. Pacif. Sci., decapod shrimps of the family Penaeidae. In: 26: 259-263. Pequegnat W. E. and Chace F. A., Jf. (eds.), Williamson, D. 1. 1982. Larval morphology and Contributions on the Biology of the Gulf of diversity. In Abele L. G. (ed.), The Biology of Mexico. Texas A & M Uni v. Oceanogr. Studies, 1: Crustacea, vol. 2 (Embryology, Morphology, and 21-57. Genetics) : 43-100 Academic Press, London. Schmitt, W. 1921. The marine decapod Crustacea of Wood-Mason, J. 1891. Phylum Appendiculata. California with special reference to the decapod Branch Arthropoda. Class Crustacea. In Wood- Crustacea collected by the United States bureau of Mason J. and Alcock A. (eds.), Natural History fisheries steamer "Albatross" in connection with Notes from H. M. Indian marine Survey Steamer the biological survey of San Francisco Bay during "Investigator" , Commander R. F. Hoskyn, R. N., the years 1912-1913. Univ. Calif. PubIs Zool., 23: commanding. Series II, no. 1. On the results of 1-470, pIs I-50. deep-sea dredging during the season 1890-1891. Smith, S. 1. 1884. Report on the decapod Crustacea of Ann. Mag. nat. Hist., (6) 8: 269-286. "Albatross" dredging off the east coast of the Yokoya, Y. 1933. On the distribution of decapod United States in 1883. Rep. U. S. Fish Commn 10: crustaceans inhabiting the continental shelf around 345-426, pIs 1-10. (Not seen.) Japan, chiefly based upon the materials collected Smith, S. I. 1886. Report on the decapod Crustacea of by S. S. Soyo-Maru; during the year 1923-1930. 1. "Albatross" dredging off the east coast of the ColI. Agric. Tokyo Univ., 12: 1-222. United States during the summer and autumn of Zarenkov, N. A. 1965. Revision of the genera 1884. Rep. U. S. Fish Commn (appendix) 21: 605- Crangon Fabricius and Sclerocrangon G. O. Sars 705, pis 1-20. (Not seen.) (Decapod a, Crustacea). Zoo I. Zh., 44: 1761-1775. Sokolova, M. N. 1957. The nourishment of some (In Russian with English summary.) species of the Crangonidae of Far Eastern seas. Zarenkov, N. A. 1993. On the new records of the Trudy Inst. Okeanol., 23: 269-285. (In Russian.) deep-sea crangonid shrimp Sclerocrangon Sokolova, M. N. 1997. Trophic structure of abyssal zenkewitchi Birshtein et Winogradow, 1953 macrobenthos. Adv. mar. BioI., 32: 427-525. (Crustacea Decapoda Crangonidae) in the Kuril Sund, O. 1920. Penaeides and stenopides. Rep. scient. Trench and the Sea of Japan. Arthropoda selecta, Res. "Michael Sars" North Atlantic Deep-Sea 2: 17. (In Russian with English abstract.) Exped., 1910,3: 1-36, pIs 1,2. Vinogradova, N. G. 1997. Zoogeography of the (Accepted January 21, 2(00)

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