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Arthropoda Selecta 10 (2): 103136

¿ ARTHROPODA SELECTA, 2001

Decapod Crustaceans of the Southwest Kamchatka Shelf: R/V Professor Levanidov collection in June 1996 Äåñÿòèíîãèå ðàêîîáðàçíûå (Crustacea: Decapoda) þæíîé ÷àñòè çàïàäíî-êàì÷àòñêîãî øåëüôà: ïî ñáîðàì íà ÍÈÑ Ïðîôåññîð Ëåâàíèäîâ â èþíå 1996 ã. V.I. Sokolov Â.È. Ñîêîëîâ
Russian Federal Institute for Marine Fisheries and Oceanography (VNIRO), Verchnyaya Krasnoselskaya Str., 17a, Moscow 107140 Russia. Ðîññèéñêèé Ôåäåðàëüíûé Èíñòèòóò ìîðñêîãî ðûáíîãî õîçÿéñòâà è îêàíîãðàôèè (ÂÍÈÐÎ), óë. Âåðõíÿÿ Êðàñíîñåëüñêàÿ, ä. 17à, Ìîñêâà 107140 Ðîññèÿ.

KEY WORDS: Decapod crustaceans, fauna, Sea of Okhotsk, West Kamchatka, protandric hermaphroditism. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: äåñÿòèíîãèå ðàêîîáðàçíûå, ôàóíà, Îõîòñêîå ìîðå, Çàïàäíàÿ Êàì÷àòêà, ïðîòåðàíäðè÷åñêèé ãåðìàôðîäèòèçì.

ABSTRACT: Thirty two decapod species (ca. 6100 specimens) were collected using a bottom trawl in the southern part of the Kamchatka shelf at depth between 20 and 300 m. Two species, e.g. Argis ochotensis Komai, 1997 and Spirontocaris brevidigitata Kobjakova, 1935 were recorded in the area for the first time, while the latter species was found for the first time in the entire Sea of Okhotsk. New material provide evidence that the latter species and Spirontocaris spinus intermedia are both well separated from Spirontocaris spinus, contrary to earlier synonimization by Hayashi [1977]. The most common species in the region investigated were Chionoecets bairdi, Chionoecetes opilio, Paralithodes camtschaticus, Pagurus trigonocheirus, Labidochirus splendescens, and Hyas coarctatus alutaceus. All these species but Chionoecetes bairdi are widely ranged in the Sea of Okhotsk. Paralithodes camtschaticus and Chionoecetes bairdi were most abundunt species in the shallow waters. Pandalus borealis eous and Chionoecetes opilio dominated in trawl catches in the depth range of 200-300 m to the north of 52?00N. The entire decapod fauna of the West Kamchatka shelf includes 50 species. Remarks on morphological variation, size composition and the presence of ovigerous females are given for most species. External sexual characters in Crangon communis were studied in details in order to clear whether this species is a protandric hermaphrodite. No apparent transitional specimens were recorded. ÐÅÇÞÌÅ: Ïîñëåäíÿÿ ïî âðåìåíè ñâîäêà äàííûõ î äåñÿòèíîãèõ ðàêîîáðàçíûõ Îõîòñêîãî ìîðÿ [Âèíîãðàäîâ, 1947] âêëþ÷àåò 91 âèä è ïîäâèä. Îäíàêî ïî ñëîæèâøèìñÿ â òî âðåìÿ â ÑÑÑÐ óñëîâèÿì èññëåäîâàòåëè â áîëüøèíñòâå ñëó÷àåâ íå èìåëè âîçìîæíîñòè ïóáëèêîâàòü ñâåäåíèÿ î ïîëîæåíèè ñòàíöèé

[Èâàíîâ, 1992] è, ñëåäîâàòåëüíî, òî÷íîì ìåñòîíàõîæäåíèè ðàçëè÷íûõ âèäîâ. ×àñòü ýòèõ ìàòåðèàëîâ â ïîñëåäñòâèè áûëà óòåðÿíà. Ïîýòîìó ìíîãèå äàííûå î ðàñïðîñòðàíåíèè ðàçëè÷íûõ âèäîâ äåñÿòèíîãèõ ðàêîîáðàçíûõ â ðîññèéñêèõ äàëüíåâîñòî÷íûõ ìîðÿõ òðåáóþò äîïîëíåíèÿ è óòî÷íåíèÿ. Èç âñåõ ðàéîíîâ Îõîòñêîãî ìîðÿ êàðöèíîëîãàìè íàèáîëåå àêòèâíî èçó÷àëñÿ øåëüô Çàïàäíîé Êàì÷àòêè. Ïðèñòàëüíîå âíèìàíèå ê ýòîé ÷àñòè ìîðÿ ñâÿçàíî ïðåèìóùåñòâåííî ñ çàïàäíîêàì÷àòñêîé ïîïóëÿöèåé êàì÷àòñêîãî êðàáà, ñèëüíî ýêñïëóàòèðóåìîé íà ïðîòÿæåíèè ïîñëåäíèõ äåñÿòèëåòèé. Ðàñïðåäåëåíèå è áèîëîãèÿ äðóãèõ ïðîìûñëîâûõ âèäîâ ýòîãî ðàéîíà èçó÷àëè â çíà÷èòåëüíî ìåíüøåé ñòåïåíè. ×òî êàñàåòñÿ íåïðîìûñëîâûõ ðàêîîáðàçíûõ òî èõ èçó÷åíèåì â ïîñëåäíèå ãîäû ïðàêòè÷åñêè íå çàíèìàëèñü. Îáñòàíîâêà â çíà÷èòåëüíîé ñòåïåíè îñëîæíÿåòñÿ ïî÷òè ïîëíûì îòñóòñòâèåì íàó÷íî-èññëåäîâàòåëüñêèõ ðàáîò íà ñïåöèàëèçèðîâàííûõ ñóäàõ.  ýòîé ñâÿçè äîâîëüíî ïîëíàÿ áåíòîñíàÿ ñúåìêà þæíîé ÷àñòè çàïàäíîêàì÷àòñêîãî øåëüôà, ïðîâåäåííàÿ ÒÈÍÐÎ-Öåíòðîì ïðè ó÷àñòèè ÂÍÈÐÎ íà ÍÈÑ Ïðîô. Ëåâàíèäîâ â èþëå 1996 ã. ïðåäñòàâëÿåò íåñîìíåííûé èíòåðåñ. Òðèäöàòü äâà âèäà (îêîëî 6100 ýêçåìïëÿðîâ) äåêàïîä áûëî ñîáðàíî â þæíîé ÷àñòè çàïàäíîêàì÷àòñêîãî øåëüôà íà ãëóáèíå îò 20 äî 300 ì. Äâà âèäà Argis ochotensis è Spirontocaris brevidigitata îòìå÷åíû äëÿ çàïàäíîêàì÷àòñêîãî øåëüôà âïåðâûå, ïîñëåäíèé âïåðâûå äëÿ Îõîòñêîãî ìîðÿ â öåëîì. Íà íîâîì ìàòåðèàëå ïîêàçàíî, ÷òî êàê ýòîò ïîñëåäíèé âèä, òàê è Spirontocaris spinus intermedia õîðîøî îáîñîáëåíû îò Spirontocaris spinus, è íå ÿâëÿþòñÿ åãî ñèíîíèìàìè, êàê ðàíåå ïðåäïîëàãàëîñü Hayashi [1977]. Âñÿ ôàóíà äåêàïîä çàïàäíîêàì÷àòñêîãî øåëü-


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V.I. Sokolov distribution pattern of most important commercial species, e.g. blue king crab, Paralithodes platypus (Brandt, 1851), red king crab, and tanner crabs, Chionoecetes bairdi Rathbun, 1902 and Chionocoetes opilio (O. Fabricius, 1788) are out of the scope of this paper.

ôà âêëþ÷àåò 50 âèäîâ. Íàèáîëåå îáû÷íûìè âèäàìè ðåãèîíà áûëèChionoecets bairdi, Chionoecetes opilio, Paralithodes camtschaticus, Pagurus trigonocheirus, Labidochirus splendescens è Hyas coarctatus alutaceus. Âñå ýòè âèäû êðîìå Ch. bairdi øèðîêî ðàñïðîñòðàíåíû â Îõîòñêîì ìîðå. P. camtschaticus è Ch. bairdi áûëè íàèáîëåå ìíîãî÷èñëåííû íà ìåëêîâîäüå. Pandalus borealis eous è Chionoecetes opilio äîìèíèðîâàëî â òðàëîâûõ óëîâàõ íà ãëóáèíå îò 200 äî 300 ì. Äëÿ âñåõ îáíàðóæåííûõ âèäîâ ïðèâåäåíû äàííûå ïî èçìåí÷èâîñòè, ðàçìåðíîìó ñîñòàâó, âñòðå÷àåìîñòè ÿéöåíîñíûõ ñàìîê. Äåòàëüíî èññëåäîâàíû âòîðè÷íûå ïîëîâûå ïðèçíàêè Crangon communis äëÿ âûÿñíåíèÿ âîïðîñà, ÿâëÿåòñÿ ëè äàííûé âèä ïðîòåðàíäðè÷åñêèì ãåðìàôðîäèòîì. Îñîáè ñ ïåðåõîäàìè îò ïðèçíàêîâ ñàìöîâ ê ïðèçíàêàì ñàìîê íå îáíàðóæåíû.

Materials and Methods
Most of 6100 specimens of 32 species of Decapoda were collected during the expedition of the Pacific Research Centre for Marine Fisheries and oceanography (TINRO-Centre, Vladivostok) and the Russian Federal Institute for Marine Fisheries and Oceanology (VNIRO, Moscow) on board R/V Professor Levanidov in the southeastern part of the Sea of Okhotsk (51œ1054œ00N). Ninety eight stations at depth from 20 to 300 m, were sampled between July, 1 and 20 in 1996. Samples were obtained by an otter-trawl (28 m flat bottom trawl) with a mesh size of 45 mm supplied with a panel of 30 mm mesh net lining the cod end, and additionally by a dredge. The dredge had a mesh size of 10 mm and a rectangular mouth opening (120x40cm). All hauls lasted 30 minutes. The stations are listed in the appendix. All decapod specimens were measured and sexed. After that most of specimens of non-commercial species (about 3300 animals) were preserved in buffered 10% seawater formalin solution and later transferred to 70% ethyl alcohol. The carapace length (CL), from the base of eyestalk to the posterior mid-dorsal edge of carapace was measured to the nearest 0.1 mm for the shrimps. The maximum carapace width (CW) was measured to the nearest 1 mm for the crabs. The maximal shield length (SL) from the base of eyestalk to the posterior mid-dorsal point was measured in the hermit crabs. Shrimp specimens were sexed by examination of the endopodite of the first two pair of pleopods. The position of gonopore was used for the sex determination of hermit crabs. Crab specimens were sexed by examination of the abdomen and first two pair of pleopods. All preserved specimens are deposited in the Zoological Museum of Moscow State University (ZMMU). Bottom water temperature was measured with a deep-sea protected reversing thermometer and bathythermograph.

Introduction
Studies on the decapod crustaceans in the Sea of Okhotsk have a rather long history. The first account of the Decapoda from the North-western Pacific including the Sea of Okhotsk was published by Brandt [1851]. Important papers which contain description of new species from the Sea of Okhotsk, regional faunistic records and updates are those by Brajnikov [1907], Rathbun [1924, 1925, 1930, 1932], Kobjakova [1936, 1935], Makarov [1938], McLaughlin [1974], Hayashi [1977]. Up to now the most comprehensive review of the decapod fauna of the Sea of Okhotsk was presented by Vinogradov [1947] who listed 91 species and subspecies. Due to restrictions which existed in the USSR prior to the 1990s, Vinogradovs publication could not contain the data on stations positions [Ivanov, 1992]. Later, considerable part of the station data was lost and this is the reason why distribution data for the Sea of Okhotsk decapods require re-consideration. Within the Sea of Okhotsk, the West Kamchatka shelf remains the most extensively studied area. A persistent attention to this waters is due to a very abundant local stock of red king crab, Paralithodes camtschaticus (Tilesius, 1815) heavily exploited in recent years. The papers by Vinogradov [1945, 1946, 1947, 1969], Rodin [1967, 1969, 1985], and Chekunova [1969] built up the basis of recent studies on population structure, migrations and reproduction of red king crab on the West Kamchatka shelf. Larval biology of several species was treated by Makarov [1966]. Distribution and biology of other species in this area have been poorly studied while recent investigations of non-commercial species are totally lacking. Also special trawl surveys became rare in the 1990s. A detailed benthic survey conducted by the expedition on R/V Professor Levanidov in July 1996 brought an extensive material which extends our knowledge of the decapod fauna in the region. The present study is focused at occurrence, distribution, morphological variability and biological characteristics of mostly non-commercial species of the Decapoda from the shelf of West Kamchatka. Abundance and

List of Decapoda collected by R/V Prof. Levanidov
Family Pandalidae Haworth, 1825 Genus Pandalus Leach, 1814 Pandalus borealis eous Makarov, 1935 Pandalus goniurus Stimpson, 1860 Family Hippolytidae Bate, 1888 Genus Eualus Thallwitz, 1892 Eualus fabricii (Kröyer, 1841) Eualus macilentus (Kröyer, 1842) Eualus pusiolus (Kröyer, 1841) Genus Lebbeus White, 1847 Lebbeus groenlandicus (Fabricius, 1775) Genus Spirontocaris Bate, 1888 Spirontocaris brevidigitata Kobjakova, 1935 Spirontocaris intermedia Makarov in Kobjakova, 1936 Spirontocaris murdochi Rathbun, 1902


Decapod Crustaceans of the Southwest Kamchatka Shelf Family Crangonidae Haworth, 1825 Genus Argis Kröyer, 1842 Argis dentata (Rathbun, 1902) Argis lar (Owen, 1839) Argis ovifer (Rathbun, 1902) Argis cf. ochotensis Komai, 1997 Genus Crangon Fabricius, 1798 Crangon septemspinosa Say, 1818 Crangon communis (Rathbun, 1902) Crangon dalli (Rathbun, 1902) Genus Mesocrangon Zarenkov, 1965 Mesocrangon intermedia (Stimpson, 1860) Genus Sclerocrangon G.O. Sars, 1883 Sclerocrangon salebrosa Owen, 1839 Family Paguridae Latreille, 1803 Genus Labidochirus Benedict, 1892 Labidochirus splendescens (Owen, 1839) Genus Pagurus Fabricius, 1775 Pagurus capillatus (Benedict, 1892) Pagurus rathbuni (Benedict, 1892) Pagurus trigonocheirus (Stimpson, 1858) Pagurus brandti (Benedict, 1892) Family Lithodidae Samouelle, 1819 Subfamily Lithodinae Ortmann, 1901 Genus Paralithodes Brandt, 1849 Paralithodes camtschaticus (Tilesius, 1815) Paralithodes platypus Brandt, 1850 Genus Lithodes Latreille, 1806 Lithodes aequispinus Benedict, 1894 Family Majidae Samouelle, 1819 Genus Chionoecetes Kröyer, 1838 Chionoecetes opilio (O. Fabricius, 1788) Chionoecetes bairdi Rathbun, 1902 Genus Hyas Leach, 1814 Hyas coarctatus alutaceus Brandt, 1851. Genus Oregonia Dana, 1851 Oregonia gracilis Dana, 1851 Family Atelecyclidae Ortmann, 1893 Genus Erimacrus Benedict, 1892 Erimacrus isenbeckii (Brandt, 1848) Genus Telmessus White, 1846 Telmessus cheiragonus (Tilesius, 1812)

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them on the carapace posterior to the level of orbital margin, and 1 or 2 toothlike spines near the apex of rostrum (Fig. 1h). The ventral margin of rostrum has 67 teeth. Scaphocerite is 4.15.9 times as long as wide, the distolateral tooth not extending or distinctly overreaching distal margin of lamina (Fig. 1dg). Median projection on the third abdominal somite is more or less prominent (Fig. 1b, c). DISTRIBUTION. North Pacific from the southern part of Chukchi Sea to the Peter the Great Bay (Sea of Japan) and to Puget Sound [Makarov, 1941; Butler, 1980; Komai, 1999]; 5450 m [Butler, 1980]. In the reference area P. goniurus was collected at depth between 52 and 294, on mud and sand, (water temperature 0.212.02oC and salinity 32.633.38 p.p.t.). BIOLOGY. CL of females reaches to 25.5 mm, those of males to 19.1 mm. Specimens in the West Kamchatka region were in general larger, than those recorded by Butler [1964, 1980] near the western coast of North America (CL 16.5 and 13.1 mm respectively). Most of females were ovigerous (79.8%). The percentage of morphologically transitional specimens (15.9%) was relatively low. This species frequently occurred together with Chionoecetes opilio, C. bairdi, Hyas coarctatus, Labidochirus splendescens, Pagurus trigonocheirus, and Crangon communis.

Pandalus borealis eous Makarov, 1935 Figs. 2, 3.
MATERIAL EXAMINED. 118 (CL 8.825.5 mm), 56 transitional (CL 16.527.5 mm), 98 non-ovigerous (CL 9.3 33.8 mm), 19 ovigerous females (CL 2733 mm). LOCALITY. Haul # 6, 8, 19, 24, 26, 31, 32, 36, 37, 43, 45, 49, 55, 56.

Systematics
FAMILY PANDALIDAE HAWORTH, 1825 Genus Pandalus Leach, 1814 Pandalus goniurus Stimpson, 1860 Fig. 1.
MATERIAL EXAMINED. 155 (CL 7.419.1 mm), 183 non-ovigerous (CL 10.228.5 mm), 64 intersexes (CL 9.2 20.3 mm). LOCALITY. Hauls # 6, 9, 10, 17, 18, 19, 20, 30, 33, 34, 37, 39, 40, 47, 48, 49, 50, 52, 53, 54, 55, 58; Dredge # 4, 11, 12, 14, 16, 17, 18, 24, 25, 33, 34, 37, 38, 39, 41, 44.

REMARKS ON MORPHOLOGY. Rostrum is 1.292.27 times as long as carapace, armed with 811 spines: 3 or 4 of

REMARKS ON MORPHOLOGY. Rostrum is 1.392.42 times as long as the carapace length, armed nearly throughout its length with 1524 spines, including 45 spines on the carapace posterior to the level of orbital margin, and a small teeth near the apex of rostrum. The ventral margin of rostrum is armed with 7 or 8 teeth. Rostrum is usually proportionally longer in small specimens than in larger ones (Fig. 3). Scaphocerite is 3.75.8 times as long as wide, the distolateral tooth usually distinctly overreaches the distal margin of lamina, but several specimens have scaphocerite with distolateral tooth not extending to the anterior margin of lamina. Small specimens usually have proportionally slender scaphocerite than larger ones. The third abdominal somite usually has an acute prominent median projection. The median projection of the third somite in few (ca. 10%) specimens in our collection is rounded and relatively small. Makarov [1935] described a new subspecies P. borealis eous based on the differences in the shape of the distal margin of scaphocerite and the relative length of rostrum. Several authors [Kobjakova, 1936, 1937; Vinogradov, 1947, 1950; Zarenkov, 1960] accepted a subspecific status of the Pacific populations of Pandalus borealis. Squires [1992] examined a few specimens from the British Columbia and Newfoundland and recognized the Pacific form as a good species, P. eous. By arguing that Pacific specimens are distinct from the P. borealis in the two particulars (the relative length of rostrum and the shape of the median projection of the third abdominal somite) Komai [1999] accepted the species rank of the Pacific form.


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Fig. 1. Pandalus goniurus Stimpson, 1860: a entire animal in lateral view; b, c dorsal margin of third abdominal somite in lateral view; dg distal part of scaphocerite; h apex of rostrum. a, b, e non-ovigerous (CL 19.8 mm); c non-ovigerous (CL 19.4 mm); d, h transitional (CL 19.5 mm); f (CL 15.4 mm); g transitional (CL 17.8 mm). Scale 1 cm (a), 1 mm (bh). Ðèñ. 1. Pandalus goniurus Stimpson, 1860: a îáùèé âèä ñáîêó; b, ñ ñïèííàÿ ïîâåðõíîñòü òðåòüåãî ñåãìåíòà àáäîìåíà (âèä ñáîêó); dg ôîðìà ïåðåäíåé ÷àñòè ñêàôîöåðèòà; h ôîðìà êîíöà ðîñòðóìà. a, b, e ñàìêà áåç ÿèö (ÄÊ 19,8 ìì); c ñàìêà áåç ÿèö (ÄÊ 19,4 ìì); d, h ïåðåõîäíàÿ îñîáü (ÄÊ 19,5 ìì); f ñàìåö (ÄÊ 15,4 ìì); g ïåðåõîäíàÿ îñîáü (ÄÊ 17,8 ìì). Ìàñøòàá 1 cì (a), 1 ìì (bh).


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Fig. 2. Pandalus borealis eous Makarov, 1935. Non-ovigerous (CL 25 mm). Entire animal in lateral view. Scale 1 cm. Ðèñ. 2. Pandalus borealis eous Makarov, 1935. Îáùèé âèä ñáîêó ñàìêè áåç ÿèö (ÄÊ 25 mm). Ìàñøòàá 1 ñì.

Fig. 3. Carapace length vs. ratio of rostrum length/carapace length of Pandalus borealis eous Makarov, 1935. Ðèñ. 3. Ãðàôèê èçìåíåíèÿ îòíîøåíèÿ äëèíû ðîñòðóìà ê äëèíå êàðàïàêñà ñ óâåëè÷åíèåì ðàçìåðà îñîáè ó Pandalus borealis eous Makarov, 1935.


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Fig. 4. Eualus fabricii (Kröyer, 1841): a entire animal in lateral view; bd shape of rostrum and carapace; e, f of scaphocerite. a, c, e non-ovigerous (CL 7.7 mm); b non-ovigerous (CL 7.8 mm); d, f non-ovigerous Scale 1 cm (ad), 1 mm (e, f). Ðèñ. 4. Eualus fabricii (Kröyer, 1841): a îáùèé âèä ñáîêó; bd ôîðìà ðîñòðóìà è êàðàïàêñà (âèä ñáîêó); ïåðåäíåé ÷àñòè ñêàôîöåðèòà. a, c, e ñàìêà áåç ÿèö (ÄÊ 7,7 ìì); b ñàìêà áåç ÿèö (ÄÊ 7,8 mm); d, f ñàìêà 7,6 ìì). Ìàñøòàá 1 ñì (ad), 1 ìì (e, f).

distal part (CL 7.6 mm). e, f ôîðìà áåç ÿèö (ÄÊ

Sokolov [1997] compared numerous specimens from the North-East Atlantic and the North-West Pacific and failed to find diagnostic characters which are good enough to distinguish between populations from these areas. As shown earlier, the ratio of rostrum length/ carapace length of the Pacific specimens is the same as in the typical form. Thus raising the rank of Pacific populations to a separate species is not sufficiently substantiated. DISTRIBUTION. The Pacific subspecies is widely distributed from the southern part of the Chukchi Sea to Puget Sound, Washington State and to the Peter the Great Bay (Sea of Japan); 101380 m [Vinogradov, 1950; Butler, 1980; Komai, 1999]. In the reference area P. borealis eous was one of the most common species of Decapoda at depth between 200 and 300 m (temperature 0.411.83œC, salinity 32.7433.39 p.p.t.) BIOLOGY. CL in females and males in the West Kamchatka region amount respectively to 33.8 mm and 25.5 mm. Specimens in the West Kamchatka region were in general larger than those recorded by Butler [1964, 1980] near the western coast of North America. Eight size groups determined by modal size were present. Most females (88.9%) were ovigerous. The percentage of transitional specimens (24.8%) was relatively high. Cohabitant species were mostly Chionoecetes opilio, Argis ochotensis, Pagurus trigonocheirus, and Crangon communis.

FAMILY HIPPOLYTIDAE BATE, 1888 Genus Eualus Thallwitz, 1892 Eualus fabricii (Kröyer, 1841) Fig. 4.
MATERIAL. 6 (CL from 7.6 to 10.3 mm) LOCALITY. Haul # 39, 40, 52; Dredge # 17, 18.

REMARKS ON MORPHOLOGY. Rostrum armed with 35 spines, including spines on the carapace posterior to the level of orbital margin (Fig. 4bd). Scaphocerite has the distolateral teeth either not extending beyond or overreaching the distal margin of the lamina (Fig. 4e, f). DISTRIBUTION: North Pacific from the Chukchi Sea to the Sea of Japan and to British Columbia; North Atlantic from the Hudson Bay, Foxe Basin and west Greenland to Cape Cod; 4275 m [Squires, 1990]; up to 630 [Butler, 1980]. In the references area E. fabricii was collected at depth between 51 and 80 m (water temperature 0.441.84œC and salinity 32.57 32.9 p.p.t.). BIOLOGY. All females were mature and had well developed gonads. Other species of decapod taken in the same catches were Chionoecetes opilio, Hyas coarctatus, Pandalus goniurus, Labidochirus splendescens, Neocrangon dalli, Sclerocrangon salebrosa.


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Fig. 5. Eualus macilentus (Kröyer, 1842). Non-ovigerous (CL 10.5 mm). Entire animal in lateral view. Scale 1 cm. Ðèñ. 5. Eualus macilentus (Kröyer, 1842). Îáùèé âèä ñàìêè ñáîêó (ÄÊ 10,5 ìì). Ìàñøòàá 1 cì.

Eualus macilentus (Kröyer, 1842) Fig. 5.
MATERIAL. 3 (CL from 5.0 to 7.8 mm), 10 nonovigerous (CL from 4.0 to 10.5 mm). LOCALITY. Haul #6, 47, 51, 52; Dredge # 25, 33, 37, 39, 40.

Genus Lebbeus White, 1847 Lebbeus groenlandicus (Fabricius, 1775) Fig. 7.
MATERIAL. 2 (CL 13.2 and 13.7 mm). LOCALITY. Haul # 40, 52.

DISTRIBUTION. North Pacific from Chukchi Sea to the Peter the Great Bay (Sea of Japan) and to the Bristol Bay (Alaska) [Vinogradov, 1950]; in the West Atlantic from Hudson Bay, Fox Channel and Greenland to Nova Scotia; 55 540 m [Squires, 1990]. The northernmost record is in the Chukchi Sea at 72?56N [Sivertsen, 1932]. In the references area shrimps were found between 55 and 200 m depths, within a water temperature range of 0.21 1.53?C and a salinity range of 32.6733.59 p.p.t. Cohabitant species included Pandalus goniurus, Hyas coarctatus, Labidochirus splendescens, and Pagurus trigonocheirus.

Eualus pusiolus (Kröyer, 1841) Fig. 6.
MATERIAL. 1 (CL 3.2 mm) LOCALITY. Dredge # 43.

DISTRIBUTION. North Pacific from Bering Sea to British Columbia and Washington in North-West America and to the Sea of Japan in Asia [Butler, 1980; Williams, 1984; Squires, 1990]; Arctic region and North Atlantic from the Barents Sea to Channel Islands and Catalonian coast of Spain (Mediterranean) in Europe, and from Gulf of St. Lawrence to Virginia in North East [Williams, 1984; Squires, 1990]; 0500 m [Holthuis, 1947]. A single specimen was caught at 85 m depth, water temperature was 0.44?C and salinity equaled to 32.7 p.p.t.

VARIABILITY. Squires [1990] noted that the abdominal pleurae of the shrimps from the Eastern Canadian coast have 3, 1, 2, 2, 2, 1 ventral spines on respectively 1st to 6th somites. In the Sea of Okhotsk shrimps have not less than two spines on the pleurae 15 [Vinogradov, 1950]. Our two males have pleurae 15 with 25 spines as it was noted by Vinogradov [1950]. DISTRIBUTION. Southern part of the Chukchi Sea through the Bering Sea to Puget Sound and to Vladivostok (Sea of Japan); Hudson Bay to Greenland and south to Rhode Island; Arctic Canada and Alaska [Squires, 1990]. Depth 2 314 m [Williams & Wigley, 1977; Squires, 1990], up to 518 m [Butler, 1980]. The species usually present in the same catches comprised of Chionoecetes bairdi, Neocrangon dalli, Pandalus goniurus, Hyas coarctatus, Sclerocrangon salebrosa, and Spirontocaris murdochi.

Genus Spirontocaris Bate, 1888 Spirontocaris murdochi Rathbun, 1902 Fig 8.
MATERIAL. 14 (CL 49 mm), 119 non-ovigerous (CL 4.110.2 mm), 3 ovigerous (8.910.3 mm).


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Fig. 6. Eualus pusiolus (Kröyer, 1841). (CL 3.2 mm). Entire animal in lateral view. Scale 1 mm. Ðèñ. 6. Eualus pusiolus (Kröyer, 1841). Îáùèé âèä ñàìöà ñáîêó (ÄÊ 3,2 ìì). Ìàñøòàá 1 ìì.

Fig. 7. Lebbeus groenlandicus (Fabricius, 1775). (CL 13.7 mm). Entire animal in lateral view. Scale 1 cm. Ðèñ. 7. Lebbeus groenlandicus (Fabricius, 1775). Îáùèé âèä ñàìöà ñáîêó (ÄÊ 13,7 ìì). Ìàñøòàá 1 cì.


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Fig. 8. Spirontocaris murdochi Rathbun, 1902: a entire animal in lateral view; bd shape of the rostrum; e distal of scaphocerite. a, e non-ovigerous (CL 9.1 mm); b non-ovigerous (CL 8.8 mm); c non-ovigerous (CL 8.7 mm); non-ovigerous (CL 8 mm). Scale 1 cm (ad), 1 mm (e). Ðèñ. 8. Spirontocaris murdochi Rathbun, 1902: a îáùèé âèä ñáîêó; bd ôîðìà êàðàïàêñà è ðîñòðóìà (âèä ñáîêó); ôîðìà ïåðåäíåé ÷àñòè ñêàôîöåðèòà. a, e ñàìêà áåç ÿèö (ÄÊ 9,1 mm); b ñàìêà áåç ÿèö (ÄÊ 8,8 mm); c ñàìêà áåç (ÄÊ 8,7 mm); d ñàìêà áåç ÿèö (ÄÊ 8 mm). Ìàñøòàá 1 cì (ad), 1 ìì (e).

part d e ÿèö

Fig. 9. Spirontocaris spinus intermedia Makarov, 1941(ac) and Spirontocaris spinus (df): a entire animal in lateral view; b scaphocerite; c antennule; d antennular peduncle; e, f distal part of scaphocerite. ad non-ovigerous (CL 11.7 mm); e (CL 9.5 mm); f non-ovigerous (CL 11 mm). Scale 1 cm (ac), 1 mm (df). Ðèñ. 9. Spirontocaris intermedia Makarov, 1941(ac) and S. spinus (df): a îáùèé âèä S. intermedia ñáîêó; b ôîðìà ñêàôîöåðèòà S. intermedia; c ïåðâàÿ àíòåííà S. intermedia; d ôîðìà ñòåáåëüêà ïåðâîé àíòåííû S. spinus; e, f ôîðìà ïåðåäíåé ÷àñòè ñêàôîöåðèòà S. spinus. ad ñàìêà áåç ÿèö (ÄÊ 11,7 ìì); e ñàìåö (ÄÊ 9,5 ìì); f ñàìêà áåç ÿèö (ÄÊ 11 ìì). Ìàñøòàá 1 cì (ac), 1 ìì (df).


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Barents Seas from the ZMMU collection. The Pacific form is distinct from the Atlantic S. spinus in the following particulars: 1) stylocerite reaches to or is slightly produces beyond the second segment of antennular peduncle in the Pacific specimens (Fig. 9c) while in the S. spinus stylocerite is distinctly overreaching antennular peduncle (Fig 9d); 2) in the Pacific shrimps distolateral tooth of scaphocerite reaches only the distal margin of the blade (Fig. 9b) while in the S. spinus the distolateral tooth distinctly overreaches the blade (Fig. 9e, f). Makarov [1941] used the former character to separate a new subspecies and Vinogradov [1950] utilized the latter in the identification key. These two characters do not vary in all examined specimens. Five specimens of S. spinus from the Atlantic coast of Canada which were studied by Hayashi [1977] had stylocerite not extending beyond the second segment of the antennular peduncle. Squires [1990] noted that S. spinus have stylocerite with attenuate sharp point reaching farther than distal 3rd article. In our material all specimens from the North Atlantic have a long slylocerite, and the specimens from the Sea of Okhotsk always have a short one. Otherwise Makarov [1966] noted, that the larvae of the Pacific form could be distinctly separated from the larvae of the other Spirontocaris species. Therefore, the concept of S. spinus in the North Atlantic and the related Pacific species S. intermedia is rather well substantiated. DISTRIBUTION. From the Chukchi Sea to the Sea of Okhotsk and to Kodjak; 27230 m depth [Vinogradov, 1950]. In the shelf of West Kamchatka shrimps were collected at depth between 150 and 245 m (water temperature between 0.79 and 1.67?C and salinity 32.8733.36 p.p.t.). BIOLOGY. CL in females reaches to 10.9 mm. All examined females were with prolipherated oocytes clearly visible through carapace. Other species of decapod taken in the same catches were Chionoecetes opilio, C. bairdi, Pagururs trigonocheirus, and Argis ochotensis.

LOCALITY. Haul # 30, 34, 38, 39, 40, 42, 47, 51, 52, 53, 54, 58; Dredge # 17, 18, 24, 33, 34, 37, 38, 39, 43, 44, 45.

REMARKS ON MORPHOLOGY. The shapes of rostrum and middorsal spines vary considerably (Fig. 8bd). Rostrum is armed with 1219 irregular teeth on the upper margin and with 36 teeth on the lower margin. The dorsal carina of carapace with 3 or 4 large spines. The anteriormost middorsal spine of carapace sometimes has 12 accessory spinules. The midaxis is straight in all examined specimens, except three females. These three shrimps (CL 89 mm) have midaxis slightly curved upward. The ratio propodus length/ dactylus length varies between 2.3 and 3.1 in females and between 1.4 and 2.5 in males. DISTRIBUTION. Arctic coast of Alaska, the Chukchi Sea through the Bering Sea to the Sea of Japan [Kobjakova, 1937; Vinogradov, 1950; Hayashi, 1977]. In the references area shrimps were found at depth between 51 and 152 m (water temperature 0.241.84?C and salinity 32.5733.59 p.p.t.). BIOLOGY. The percentage of ovigerous females was low (2.4%), but most other females (61%) were with eggs ready to be laid. This species was frequently co-occurred with Hyas coarctatus, Labidochirus splendescens, Pagurus trigonocheirus, Pandalus goniurus, and Argis lar.

Spirontocaris intermedia Makarov in Kobjakova, 1936 Fig. 9ac.
MATERIAL. 6 non-ovigerous (CL 8.110.9 mm). LOCALITY. Haul # 32, 43.

REMARKS ON MORPHOLOGY. The shape of rostrum varies consideraly. In all specimens examined, a large tooth is present on the lower margin of rostrum, which extends anteriorly nearly to the tip of rostral midaxis. Midaxis is straight or slightly curved upward. The middorsal carina of carapace is armed with 3 to 5 large spines. The anteriormost one or two teeth are armed with a few secondary teeth. The upper margin of rostrum with many (1218) small irregular teeth. Stylocerite reaches to or slightly projects beyond the second segment of antennular peduncle (Fig. 9c). The ratio of the propodus length to dactylus length is 3.24.1. DISCUSSION. Makarov [1941] described a new subspecies S. spinus intermedia based on the numerous specimens from the Bering and the Chukchi seas. However, a diagnosis of this subspecies was given in the paper by Kobjakova [1936] with the reference to Makarovs unpublished paper. Russian authors [e.g. Kobjakova, 1937; Vinogradov, 1947, 1950] accepted this taxon. Kobjakova [1937] separated the Pacific subspecies from the typical S. spinus (Sowerby, 1805) by the following characters: 1) low carapace carina; 2) less produced posterodorsal margin of the third abdominal somite; 3) longer rostrum; 4) shape of the second abdominal pleura; 5) longer sixth abdominal somite. Hayashi [1977] did not recognize this subspecies. He examined 17 specimens from the Sea of Japan and the Sea of Okhotsk and 5 specimens of S. spinus from the Atlantic coast of Canada and concluded that most diagnostic characters of the Pacific subspecies which were used by Kobjakova [1937] are poorly expressed and variable. However, the present study indicates that there are another differences between typical S. spinus and the Pacific counterpart, which were discussed neither by Kobjakova [1936, 1937] nor Hayashi [1977]. I examined 223 specimens (60 and 163 ) of S. spinus from the Norwegian and the

Spirontocaris brevidigitata Kobjakova, 1935 Fig. 10.
MATERIAL. 14 non-ovigerous (CL from 6.2 to 12.9 mm), 2 (CL from 5.8 to 6.5 mm). LOCALITY. Haul # 30, 31, 39, 49.

REMARKS ON MORPHOLOGY. The midaxis of rostrum is slightly curved upward (Fig. 10a, b). The middorsal carina of carapace is armed with 4 to 5 large spines. The anteriormost rostral tooth is armed with 1 to 2 secondary teeth. The upper margin of rostrum is armed with 6 to 9 teeth. Sometimes dorsal teeth of rostrum armed with one-two small secondary teeth (Fig. 10a). The ratio of the propodus length to dactylus length ranges 3.0 and 4.12. DISCUSSION. Kobjakova [1935] described S. brevidigitata from the Sea of Japan and Vinogradov [1947, 1950] later accepted this taxon. Hayashi [1977] considered S. brevidigitata as a junior synonym of Spirontocaris spinus. These two species may be separated, however, by the following characters: 1) in S. spinus stylocerite distinctly extends beyond the antennular peduncle (Fig. 9d), while in S. brevidigitata stylocerite only reaches the third segment of antennular peduncle (Fig. 10d); 2) in S. spinus a large tooth is present on the lower margin of rostrum, which extends anteriorly to the tip of the rostrum midaxis, while in S. brevidigitata the midaxis distinctly overreaches the teeth on the lower margin of rostrum (Fig. 10a, b); 3) in S. spinus there are much more small teeth on the upper margin of rostrum than in S. brevidigitata; 4). in


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Fig. 10. Spirontocaris and carapace; c distal d). Ðèñ. 10. Spirontocaris (âèä ñáîêó); c ôîðìà cì (a, b), 1 ìì (c, d).

brevidigitata Kobjakova, 1935, non-ovigerous : a entire animal in lateral view; b shape of rostrum part of scaphocerite; d antennule. a, c, d CL 11.8 mm; b CL 9.3 mm. Scale 1 cm (a, b), 1 mm (c, brevidigitata Kobjakova, 1935, ñàìêà áåç ÿèö: a îáùèé âèä ñáîêó; b ôîðìà ðîñòðóìà è êàðàïàêñà ïåðåäíåãî êðàÿ ñêàôîöåðèòà; d ïåðâàÿ àíòåííà. a, d, c ÄÊ 11,8 ìì; b ÄÊ 9,3 ìì. Ìàñøòàá 1

S. brevidigitata the posterodorsal margin of the third somite is less produced than in S. spinus. S. brevidigitata differs from S. spinus intermedia in the shape of rostrum and the number of teeth on its upper margin. DISTRIBUTION. Sea of Japan [Kobjakova, 1937; Vinogradov, 1950]. Our records of this species in the Sea of Okhotsk considerably widen the range of S. brevidigitata. The bathymetric range of S. brevidigitata ranges between 60 (present study) and 1380 m [Kobjakova, 1937]. Off West Kamchatka the species occurs at the depth range from 60 to 251 m (water temperature 0.971.55?C and salinity 32.3833.23 p.p.t.). BIOLOGY. Most of the collected females (88%) were with prolipherated oocytes. The species cohabitant were Chionoecetes opilio, C. bairdi, Pagururs trigonocheirus, Crangon communis, and Argis ochotensis.

FAMILY CRANGONIDAE HAWORTH, 1825 Genus Argis Kröyer, 1842 Argis dentata (Rathbun, 1902)
MATERIAL. 2 non-ovigerous (CL 18.7 and 19.2 mm), 1 ovigerous (CL 20.8 mm).

REMARKS ON MORPHOLOGY. The ratio of the palm of first pereiopod length to width varies from 3.9 to 4.8. As it was noted by Komai [1997], the armature of the fourth abdominal pleuron is variable. Two females, which were collected near West Kamchatka, have broadly rounded pleurae of the fourth abdominal somite. Fourth abdominal pleurone of the third female bears a small sharp posteroventral tooth. DISTRIBUTION. Arctic Ocean from the Cambridge Bay through the Beaufort Sea and the Bering Sea to San Juan Islands (Washington) in the East Pacific [Butler, 1980; Squires, 1990] and to the southeast coast of Kamchatka and the Sea of Okhotsk in the Western Pacific [Komai, 1997]; from the Canadian Arctic and northwestern Greenland to Nova Scotia in the North Atlantic [Squires, 1990]; subtidal to about 200 m [Komai, 1997]. In the references area the species was found at 20 m depth (water temperature 6.24?C and salinity 32.6 p.p.t).

LOCALITY. Haul # 3

Argis lar (Owen, 1839)
MATERIAL. 51 ovigerous mm). LOCALITY. Dredge # 4, 11, 143 non-ovigerous (CL from 6.1 to 24.3 mm), (CL from 15.8 to 24 mm), 51 (CL 7.014.3 Haul 3, 6, 30, 34, 39, 40, 42, 45, 48, 51, 53, 54; 17, 24, 25, 29, 33, 37, 38, 39, 42, 43, 44, 45.


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Fig. 11. Argis lar (Owen, 1839). Carapace length frequency distributions of females; the number of specimens in parentheses. Ðèñ. 11. Ãèñòîãðàììà ðàçìåðíîãî ñîñòàâà ñàìîê Argis lar (Owen, 1839).  ñêîáêàõ óêàçàíî ÷èñëî îñîáåé.

VARIABILITY. The ratios of the propodus length to width in the 1st pereiopod varies from 3.9 to 4.8 in females and from 2.95 to 4.12 in males. The armature of the fourth abdominal pleuron is variable: in the overwhelming majority of specimens the fourth pleurone is rounded, but in one specimen (CL 9.3 mm) the fourth pleurone has a small posteroventral tooth, in another female (CL 18.8 mm) pleuron was produced on the left side and has a proximo-ventral tooth on the right side. DISTRIBUTION. The North Pacific only: from Point Barrow through the Bering Sea to Sitka and to Peter the Great Bay (Sea of Japan) [Makarov, 1941; Butler, 1980]; 0400 m depth [Vinogradov, 1950]. In the references area the species was common at depth between 51 and 295 m (water temperature 0.211.87?C and salinity 32.5733.39 p.p.t.). BIOLOGY. Large-size females with CL from 18.0 to 23.5 mm dominated in the samples. Three clear-cut size groups with CL modes of respectively 9, 15 and 21 mm can be detected in the carapace length frequency distribution (Fig. 11). The mimimum CL of ovigerous females is 14.8 mm. Ca. 26% of females were ovigerous. Co-occurring species were mostly Pandalus goniurus, Chionoecetes opilio, Hyas coarctatus, Pagurus trigonocheirus, Labidochirus splendescens, and Spirontocaris murdochi.

along the America coast [Butler, 1980; Komai, 1997] and to the Sea of Okhotsk along the Asian coast; 100293 m [Makarov, 1941; Vinogradov, 1947; 1950; Komai, 1997]. In the references area the species was collected at depth between 60 and 147 m (water temperature 1.152.02?C and salinity 32.7333.97 p.p.t.). BIOLOGY. CL in females reaches to 19.3 mm. The species frequently occures with Hyas coarctatus, Pagurus trigonocheirus , Pandalus goniurus , and Paralithodes camtschaticus.

Argis cf. ochotensis Komai, 1997
MATERIAL. 11 ovigerous (CL 24.128.2 mm), 34 nonovigerous (CL 9.327.8 mm), 12 (CL 6.526.2 mm). LOCALITY. Haul # 9, 31, 32, 37, 43, 45, 49, 56.

Argis ovifer (Rathbun, 1902)
MATERIAL. 5 non-ovigerous (CL 16.719.3 mm) LOCALITY. Haul # 17, 30, 33.

VARIABILITY. The ratios of the palm of first pereiopod length to width varies between 3.3 to 3.6. DISTRIBUTION. From the North Pacific only: from the Bering Sea to the Queen Charlotte Sound, British Columbia

REMARKS ON MORPHOLOGY. The ratio of the palm of the first pereiopod length to width ranges between 4.13 and 5.38 in females and from 3.75 and 5.14 in males. The number of articles composing the outer flagellum varies from 15 to 25 in females and 1922 in males. The ratio of the scaphocerite length to width varies from 2.41 to 3.36 in females and from 2 to 2.63 in males. DISCUSSION. Our specimens resemble both A. ochotensis and A. toyamaensis [Yokoya, 1933] by the follow characters: 1) the posterodorsal margin of the fifth abdominal somite produced posteriorly in form of an acute tooth; 2) carapace with two median spines; 3) the submedian carinae on the sixth abdominal somite nearly parallel; 4) the propodus of the female 1st pereiopod being more than 4 times as long as wide; 5) rostrum with an acute or subacute apex; 6) the propodi of the 3rd to the 5th pereiopods lacking extensor lateral ridge, and 7) scaphocerite being than 2.4 times as long as wide. The present


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Fig. 12. Crangon septemspinosa Say, 1818. Non-ovigerous (CL 12.8 mm): a entire animal in lateral view; b carapace in lateral view; c carapace in dorsal view; d telson and sixth abdominal segment in dorsal view. Scale 1 cm. Ðèñ. 12. Crangon septemspinosa Say, 1818. Ñàìêà áåç ÿèö (ÄÊ 12,8 ìì): a îáùèé âèä ñáîêó; b ôîðìà êàðàïàêñà (âèä ñáîêó); c ôîðìà êàðàïàêñà (âèä ñâåðõó); d ôîðìà òåëüñîíà è øåñòîãî ñåãìåíòà àáäîìåíà (âèä ñâåðõó). Ìàñøòàá 1 cì.

material differ from A. toyamaensis by a long stylocerite (in A. toyamaensis the stylocerite does not reach to the middle of the proximal segment of antennular peduncle, while in our specimens it is produced beyond it) and by the coloration of living specimens (in our specimens the body is generally light brown, without tints on the pleural margins of the first to fifth abdominal somite which are usually present in A. toyamaensis [Komai, 1997]). The sculpture of the body integument is relatively stronger in A. toyamaensis than in our specimens. DISTRIBUTION. Sea of Okhotsk; depth greater than 600 m [Komai, 1997]. In the references area the shrimps were found at depth between 150 and 295 m (water temperature 0.791.81?C and salinity 32.8733.36 p.p.t.). BIOLOGY. CL in females reaches up to 28.2 mm, in males it amounts up to 26.2 mm. Large females dominated in the samples. Ca. 25% of females were ovigerous. Common co-occuring species were Chionoecetes opilio, Labidochirus splendescens, Pagurus trigonocheirus, Pandalus borealis, and Crangon communis.

dov, 1950]. Distributed in the depth range of 027 m in North Pacific [Vinogradov, 1950] and between 0 and 90 m in North Atlantic [Squires, 1990]. Our single specimen was collected at depth 22 m (water temperature 4.72?C, salinity 32.21 p.p.t.).

Crangon communis (Rathbun, 1902) Fig. 1316.
MATERIAL. 5 2 (CL 5.114.9 mm), 12 ovigerous (CL 11.513.8 mm), 102 non-ovigerous (CL 816.4 mm). LOCALITY. Trawl # 6, 18, 30, 31, 36, 37, 38, 39, 42, 43, 45, 47, 49, 50, 51, 54, 55, 56, 58; Dredge # 9, 14, 16, 17, 24, 25, 27, 28, 29, 34, 35, 38, 39, 43, 44.

Genus Crangon Fabricius, 1798 Crangon septemspinosa Say, 1818 Fig. 12.
MATERIAL. 1 non-ovigerous (CL 12.8 mm) LOCALITY. Dredge # 20

DISTRIBUTION. From the Gulf of St. Lawrence to east Florida in North Atlantic; from the Arctic coast of Alaska to Shumagin Islands and from the northern part of the Sea of Okhotsk to Peter the Great Bay in the North Pacific [Vinogra-

VARIABILITY. Rostrum usually extends beyond eyes (Fig. 13b, c), but in a 30% of specimens it does not reach eyes (Fig. 13a). Most of our specimens have unarmed pleurae of the 4th abdominal somite. In three specimens 4th pleura has proximo-ventral tooth. DISTRIBUTION. From 71?17N (Chukchi Sea) [Makarov, 1941], through Bering Sea to San Diego in North America and to Peter the Great Bay and Eastern coast of Honshu Island (Sea of Japan); 161537 m depth [Butler, 1980]. In the references area shrimps were found at depth between 37 and 295 m, (water temperature 0.212.8?C and salinity 32.6633.39 p.p.t.). BIOLOGY. Females dominated in the catches. Small-size females (CL <9 mm) were scarce (Fig. 16b). The size distributions of males and females show that males dominated in small-size groups and females prevailed in large-size groups (Fig. 16). The scarcity of small females suggests a possible case of protandric hermaphroditism which was supposed for


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Fig. 13. Crangon communis (Rathbun, 1902). Non-ovigerous (CL 13.7 mm): a entire animal in lateral view; b carapace in dorsal view; c dactylus of fifth pereiopod. Scale 1 cm (a, b), 1 mm (c). Ðèñ. 13. Crangon communis (Rathbun, 1902). Ñàìêà áåç ÿèö (ÄÊ 13,7 ìì): a îáùèé âèä ñáîêó; b ôîðìà êàðàïàêñà (âèä ñâåðõó); c ôîðìà äàêòèëþñà ïÿòîãî ïåðåîïîäà. Ìàñøòàá 1 cì (a, b), 1 ìì (c).

Fig. 14. First (a, c, e) and second endopodite (b, d, f) and a. interna on the second endopodite of females Crangon communis (Rathbun, 1902): a, b non-ovigerous (CL 8.7 mm); c, d non-ovigerous (CL 13.7 mm); eg ovigerous (CL 12.8 mm). Scale 1 mm. Ðèñ. 14. Ôîðìà ýíäîïîäèòîâ ïåðâîé (a, c, e) è âòîðîé ïàðû ïëåîïîä (b, d, f) è a. interna íà âòîðîì ýíäîïîäèòå ñàìîê Crangon communis (Rathbun, 1902): a, b ñàìêà áåç ÿèö (ÄÊ 8,7 ìì); c, d ñàìêà áåç ÿèö (ÄÊ 13,7 ìì); eg ñàìêà ñ ÿéöàìè íà ïëåîïîäàõ (ÄÊ 12,8 ìì). Ìàñøòàá 1 ìì.

Fig. 15. First endopodite (a, c) and a. masculina (b, d, e) of males Crangon communis (Rathbun, 1902): a, b CL 8.5 mm; c, d CL 9.7 mm; e CL 14 mm. Scale 1 mm. Ðèñ. 15. Ôîðìà ýíäîïîäèòà ïåðâûõ ïëåîïîä (a, c) è a. masculina (b, d, e) ó ñàìöîâ Crangon communis (Rathbun, 1902): a, b ÄÊ 8,5 ìì; c, d ÄÊ 9,7 ìì; e ÄÊ 14 ìì. Ìàñøòàá 1 ìì.


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Fig. 16. Carapace length frequency distribution of males (a) and females (b) Crangon communis (Rathbun, 1902) and size specific sex ratio of shrimp samples at July 1996. Ðèñ. 16. Ãèñòîãðàììû ðàçìåðíîãî ñîñòàâà ñàìöîâ (a) è ñàìîê (b) Crangon communis (Rathbun, 1902) è êðèâûå äîëè îñîáåé ðàçíûõ ïîëîâ â ïðîáàõ â èþëå 1996 ã.


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Fig. 17. Crangon dalli (Rathbun, 1902): a d anterior part of carapace in dorsal view. a, (ac), 1 mm (d). Ðèñ. 17. Crangon dalli (Rathbun, 1902): a êàðàïàêñà (âèä ñâåðõó); d ôîðìà êàðàïàêñà áåç ÿèö (ÄÊ 13,5 ìì). Ìàñøòàá 1 cì (ac), 1

entire animal in lateral view; bñ carapace in lateral (b) and dorsal (c) views; c ovigerous (CL 10.3 mm); b, d non-ovigerous (CL 13.5 mm). Scale 1 cm îáùèé âèä ñáîêó; b ôîðìà êàðàïàêñà (âèä ñáîêó); c ôîðìà ïåðåäíåé ÷àñòè (âèä ñâåðõó). a, c ñàìêà ñ ÿéöàìè íà ïëåîïîäàõ (ÄÊ 10,3 ìì); b, d ñàìêà ìì (d).

some crangonid species [Fréchette et al., 1970; Boddeke et al. 1991]. The graph of size specific sex ratio (Fig. 16c) shows the rapid decrease in the percentage of males in the classes greater than 10 mm and scarcity of females below this size. Figures 14 and 15 show the development of secondary sexual characters of females and males respectively. The endopodite of the first pleopod gradually increases in size throughout the life of females (Fig. 14a, c, e). Small females with CL 89 mm have only rudiments of setae on the first endopodite. Ovigerous females have endopodite of first pleopod with long and feathery setae adapted for the attachment of eggs (Fig. 14e). Appendix interna on the endopodite of the second pleopod of a small female looks like a small projection without any setae (Fig. 14b), a. interna of large females bearing few short (Fig. 14d) or long setae (Fig. 14g). Endopodite of the first pleopod of males is relatively short and broad (Fig. 15 a, c). Appendix masculina of males with CL 710 mm bears long and stout setae (Fig. 15b, d). Several large males (cl 13.514 mm) have a. masculina with short, reduced setae (Fig. 15e). However no apparent morphological intersexes (transitional specimens) were recorded. Therefore the differences in size distribution of males and females may have an explanation other than protandry, e.g. different growth rates of males and females or differences of distribution of males and females, and the question

must be left to more detailed surveys, and to histological investigation of the gonads. The percentage of ovigerous females was rather low (10.5%). Cohabitant species were Chionoecetes bairdi, Hyas coarctatus, and A. lar.

Crangon dalli (Rathbun, 1902) Fig. 17, 18.
MATERIAL. 9 9 (CL 7.411.1 mm), 75 ovigerous (CL 8.514.5 mm), 61 non-ovigerous (CL 7.313.4 mm) LOCALITY. Haul # 3, 13, 39, 40, 52, 53; Dredge # 1, 2, 7, 8, 9, 10, 19, 20, 23, 25, 33, 37, 45.

REMARKS ON MORPHOLOGY. Ca. 70% of specimens have the rostrum extending beyond the base of cornea (Fig. 17d); the rostrum in other shrimps is not reaching of the basis of cornea (Fig. 17c). The ratio of the scaphocerite length to width ranges betwen 2.5 and 3.9 in both sexes. DISTRIBUTION. From the Chukchi Sea, through the Bering Sea to Puget Sound and Peter the Great Bay; 3630 m depth [Butler, 1980]. In the West Kamchatka area C. dalli was most common shrimp species at depth between 19 and 60 m. Only twice this shrimps were found deeper (85 and 100 m). Bottom water temperature in the area of collection ranged between 0.44 and 6.24?C and salinity was in the range of 32.56 to 33.59 p.p.t.


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Fig. 18. Carapace length frequency distribution of Crangon dalli: a ; b . Ðèñ. 18. Ãèñòîãðàììà ðàçìåðíîãî ñîñòàâà Crangon dalli: a ñàìêè; b ñàìöû.

BIOLOGY. The maximum CL in females was 14.5 mm, in males it reached 11.1 mm. The minimum length at maturity in females was 8.3 mm. Large-size females dominated in samples (Fig. 18). The majority of large-size females with CL more than 10 mm (ca. 60%) were ovigerous. Ca. 37% of ovigerous females had developed embryos (eyed eggs), other ovigerous females were with newly released eggs. Only one clear-cut size group with a mode at 9.5 mm was detected in carapace length frequency distribution of males (Fig. 18a). In females several modal groups can be detected (Fig. 18a). The species commonly present together withCrangon dalli were Chionoecetes bairdi, Hyas coarctatus, and Argis lar.

DISTRIBUTION. From St. Lawrence Island (Bering Sea) to the Peter the Great Bay (Sea of Japan) and Yokohama; 15 400 m depth [Vinogradov, 1950]. In the references area the shrimps were collected at 51 191 m depth (usually 5185 m depth), at water temperature 0.341.73?C and salinity 32.5733.59 p.p.t. BIOLOGY. Males were found only at one station at 56 m depth. Ca. 25% of females had proliferated oocytes well visible under carapace. The present species frequently occurred together with Pandalus goniurus, Chionoecetes opilio, Labidochirus splendescens, and Argis lar.

LOCALITY. Haul # 34, 39, 40, 42, 51, 52, 53, 54; Dredge # 14, 33, 37, 38.

Genus Mesocrangon Zarenkov, 1965 Mesocrangon intermedia (Stimpson, 1860) Fig. 19.
MATERIAL. 2 (CL 4.1 and 4.9 mm), 25 non-ovigerous (CL 7.19.1 mm)

Genus Sclerocrangon G.O. Sars, 1883 Sclerocrangon salebrosa Owen, 1839 Fig. 20.
MATERIAL. 26 (CL 621.5 mm), 1 ovigerous (CL 30 mm), 23 non-ovigerous (CL 11.830.3 mm).


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Fig. 19. Mesocrangon intermedia (Stimpson, 1860): a entire animal in lateral view; b entire animal in dorsal view; c anterior part of carapace in dorsal view; d sixth abdominal segment in dorsal view. a, b non-ovigerous (CL 7.1 mm); c non-ovigerous (CL 7.4 mm); d non-ovigerous female (CL 8.8 mm). Scale 1 cm (a, b), 1 mm (c, d). Ðèñ. 19. Mesocrangon intermedia (Stimpson, 1860): a îáùèé âèä ñáîêó; b îáùèé âèä ñâåðõó; c ïåðåäíÿÿ ÷àñòü êàðàïàêñà (âèä ñâåðõó); d ôîðìà øåñòîãî ñåãìåíòà àáäîìåíà (âèä ñâåðõó). a, b ñàìêà áåç ÿèö (ÄÊ 7,1 ìì); c ñàìêà áåç ÿèö (ÄÊ 7,4 ìì); d ñàìêà áåç ÿèö (ÄÊ 8,8 ìì). Ìàñøòàá 1 cì (a, b), 1 ìì (c, d).

Fig. 20. Sclerocrangon salebrosa Owen, 1839. Non-ovigerous (CL 20.8 mm): a entire animal in lateral view; b carapace in dorsal view. Scale 1 cm. Ðèñ. 20. Sclerocrangon salebrosa Owen, 1839. Ñàìêà áåç ÿèö (ÄÊ 20,8 ìì): a îáùèé âèä ñáîêó; b ôîðìà êàðàïàêñà (âèä ñâåðõó). Ìàñøòàá 1 cì.


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Fig. 21. Labidochirus splendescens (Owen, 1839). (SL 10.3 mm): a carapace; b right chela and carpus in dorsal view; c left chela and carpus in dorsal view; d dactylus, propodus and carpus of left third pereiopod; e telson. Scale 1 cm (ad), 1 mm (e). Ðèñ. 21. Labidochirus splendescens (Owen, 1839). Ñàìåö (ÄÙ 10,3 ìì): a ôîðìà ùèòêà; b ôîðìà êëåøíè è êàðïóñà (âèä ñâåðõó); c ôîðìà ëåâîé êëåøíè è êàðïóñà (âèä ñâåðõó); d äàêòèëþñ, ïðîïîäóñ è êàðïóñ òðåòüåãî ïåðåîïîäà; e ôîðìà òåëüñîíà. Ìàñøòàá 1 cì (ad), 1 ìì (e). LOCALITY. Haul # 39, 40, 52, 53, 55; Dredge # 33, 37, 45.

DISTRIBUTION. From the western Bering Sea to Peter the Great Bay; 10250 m depth [Vinogradov, 1950]. In the references area the shrimps were collected at depth from 51 to 100 m (water temperature 0.441.31?C and salinity 32.57 33.59 p.p.t.). Common co-habitant species were Chionoecetes opilio, Hyas coarctatus, Labidochirus splendescens, Argis lar, and Spirontocaris murdochi.

FAMILY PAGURIDAE LATREILLE, 1803 Genus Labidochirus Benedict, 1892 Labidochirus splendescens (Owen, 1839) Fig. 21, 22.
MATERIAL. 173 (SL 2.313.8 mm), 15 ovigerous (CL 68.4 mm), 69 non-ovigerous (2.412.3 mm). LOCALITY. Haul # 1, 5, 9, 11, 17, 18, 19, 20, 24, 29, 30, 34, 37, 38, 39, 40, 41, 42, 43, 47, 49, 50, 51, 52, 53, 54; Dredge # 6, 11, 13, 14, 16, 17, 18, 24, 25, 27, 29, 33, 34, 35, 38, 39, 43, 44, 45.

REMARKS ON MORPHOLOGY. Rostrum terminates in 2 to 4 small denticles (Fig. 21a, 22b, c). Ocular peduncles

are short, less than the half SL in large specimens (Fig. 21a). In small animals the ocular peduncles are relatively long, about or slightly more than one-half SL (Fig. 22a). The shape of the right chelae varies: dactyli in small specimens usually are subequal to the palm length (Fig. 22d, e), in larger males the dactyli ussually comprises three fourths of the palm length (Fig. 21b). DISTRIBUTION. Arctic Ocean (Chukchi Sea) from the Kolyma River to Point Barrow [Makarov, 1941]; from the Bering Sea to the Sea of Okhotsk and the Sea of Japan along the Asian coast and from the Gulf of Alaska to Washington and Puget Sound along the American coast; subtidal to 412 m [McLaughlin, 1974]. In the West Kamchatka area L. splendescens is one of the most common species. It was found between 30 and 294 m depth (water temperature 0.214.93?C and salinity 32.56 33.37 p.p.t.). BIOLOGY. The fraction of ovigerous females (ca. 16%) was relatively low. Size (SL) of ovigerous females varies from 6 to 10 mm. In the catches this species was commonly found together with Chionoecetes opilio, Hyas coarctatus, Pagurus trigonocheirus, Crangon communis, Pandalus goniurus, and Argis lar.


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Fig. 22. Labidochirus splendescens (Owen, 1839): a carapace; b, c anterior part of shield; df right chela in dorsal view. a (SL 2.9 mm); b (SL 11.3 mm); c ovigerous (SL 7.8 mm); d (SL 7 mm); e (SL 8.3 mm); f (SL 10.7 mm). Scale 1 mm (a), 1 cm (bf). Ðèñ. 22. Labidochirus splendescens (Owen, 1839): a ôîðìà ùèòêà (âèä ñâåðõó); b, c ôîðìà ïåðåäíåé ÷àñòè ùèòêà (âèä ñâåðõó); df ôîðìà ëåâîé êëåøíè (âèä ñâåðõó). a ñàìåö (ÄÙ 2,9 ìì); b ñàìåö (ÄÙ 11,3 ìì); c ñàìêà ñ ÿéöàìè (ÄÙ 7,8 ìì); d ñàìåö (ÄÙ 7 ìì); e ñàìåö (ÄÙ 8,3 ìì); f ñàìåö (ÄÙ 10,7 ìì). Ìàñøòàá 1 ìì (a), 1 cì (bf).

Fig. 23. Pagurus capillatus (Benedict, 1892): a shield; be right chela in dorsal view; fh left chela in dorsal view; i telson. a, b, f, i non-ovigerous (SL 8.2 mm); c, g non-ovigerous (SL 12.4 mm); d ovigerous (SL 10.4 mm); e ovigerous (SL 10 mm); h (SL 14.3 mm). Scale 1 cm (ah), 1 mm (i). Ðèñ. 23. Pagurus capillatus (Benedict, 1892): a ôîðìà ùèòêà (âèä ñâåðõó); bd ôîðìà ïðàâîé êëåøíè (âèä ñâåðõó); f-h - ôîðìà ëåâîé êëåøíè (âèä ñâåðõó); i ôîðìà òåëüñîíà. a, b, f, i ñàìêà áåç ÿèö (ÄÙ 8,2 ìì); c, g ñàìêà áåç ÿèö (ÄÙ 12,4 ìì); d ñàìêà ñ ÿéöàìè (ÄÙ 10,4 ìì); e ñàìêà ñ ÿéöàìè (ÄÙ 10 ìì); h ñàìåö (ÄÙ 14,3 ìì). Ìàñøòàá 1 cì (ah), 1 ìì (i).


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Fig. 24. Pagurus rathbuni (Benedict, 1892): a shield; b right chela and carpus in dorsal view; c left chela and carpus in dorsal view; d left chela in lateral view; e right chela in dorsal view; f left chela in dorsal view; g, h telson. ad (SL 10.8 mm); e (SL 8.1 mm); f (SL 12.5 mm); g (SL 8.1 mm); h (SL 11.5 mm). Scale 1 cm (af), 1 mm (g, h). Ðèñ. 24. Pagurus rathbuni (Benedict, 1892): a ôîðìà ùèòêà (âèä ñâåðõó); b ôîðìà ïðàâîé êëåøíè è êàðïóñà (âèä ñâåðõó); c ôîðìà ëåâîé êëåøíè è êàðïóñà (âèä ñâåðõó); d ôîðìà ëåâîé êëåøíè (âèä ñáîêó); e ôîðìà ïðàâîé êëåøíè (âèä ñâåðõó); f ôîðìà ëåâîé êëåøíè (âèä ñâåðõó); g, h ôîðìà òåëüñîíà. ad ñàìåö (ÄÙ 10,8 ìì); e ñàìåö (ÄÙ 8,1 ìì); f ñàìåö (ÄÙ 12,5 ìì); g ñàìåö (ÄÙ 8,1 ìì); h ñàìåö (ÄÙ 11,5 ìì). Ìàñøòàá 1 cì (af), 1 ìì (g, h).

Genus Pagurus Fabricius, 1775 Pagurus capillatus (Benedict, 1892) Fig. 23.
MATERIAL. 20 (SL 6.715.7 mm), 6 ovigerous (SL 7.110.8 mm), 8 non-ovigerous (SL 6.711.5 mm). LOCALITY. Haul # 1, 3, 13, 15, 16; Dredge # 19, 20.

REMARKS ON MORPHOLOGY. The palm of the right chela varies in shape from prominent oval to nearly triangular (Fig. 23be). The left cheliped is moderately short (Fig. 23fh). Distribution. From southern part of the Chukchi Sea, through the Bering Sea southward to the Sea of Japan and North Korea in Asian coast and from Bering Sea to California in American coast [Makarov, 1938, McLaughlin, 1974]; 4 432 m depth [McLaughlin, 1974]. In the references area the present species occurred between 19 and 43 m depth (water temperature 2.546.24?C and salinity 32.3132.65 p.p.t.). Other decapod crustacean species occasionally co-occurring with P. capillatus were Chionoecetes bairdi and Crangon dalli.

DISTRIBUTION. In the Arctic Ocean from the mouth of Kolyma River to Point Barrow [Makarov, 1938], southward through the Bering Sea to the Peter the Great Bay [Vinogradov, 1950] and Niigata [Miyake et al., 1962]; 9 to 210 m [McLaughlin, 1974]. P. rathbuni was collected by R/V Prof. Levanidov only at depth between 200 and 300 m depth (water temperature 1.221.83?C and salinity 33.1533.39 p.p.t.). Thus the known vertical range of P. rathbuni is extended to 300 meters. Frequently co-occuring species were Chionoecetes opilio, Pandalus goniurus, and Crangon communis.

LOCALITY. Haul # 6, 20, 36, 49.

Pagurus trigonocheirus (Stimpson, 1858) Fig. 25, 26.
MATERIAL. 155 (SL 1.419.8 mm), 2 ovigerous (SL 13.2 and 13.8 mm), 123 non-ovigerous (SL 1.713.6 mm). LOCALITY. Haul # 3, 6, 9, 11, 12, 16, 17, 18, 19, 24, 31, 33, 37, 38, 39, 40, 41, 42, 43, 47, 50, 52, 53, 54, 56; Dredge # 3, 10, 11, 12, 13, 14, 15, 16, 17, 24, 25, 28, 29, 33, 34, 37, 39, 42, 43, 44.

Pagurus rathbuni (Benedict, 1892) Fig. 24.
MATERIAL. 11 (SL 7.112.2 mm), 1 non-ovigerous (SL 6.9 mm).

REMARKS ON MORPHOLOGY. Variability in the most important characters of P. trigonocheirus in the reference area was discussed by Sokolov [1998]. The ratio of the right cheliped palm width to length varies from 0.87 to 1.63 in males and from 0.9 to 1.75 in females. There is a weak


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Fig. 25. Pagurus trigonocheirus (Stimpson, 1858). (SL 13.3 mm): a shield; b right chela; c left chela; d telson. Scale 1 cm (b, c), 1 mm (a, d). Ðèñ. 25. Pagurus trigonocheirus (Stimpson, 1858). Ñàìåö (ÄÙ 13,3 ìì): a ôîðìà ùèòêà (âèä ñâåðõó); b ôîðìà ïðàâîé êëåøíè; c ôîðìà ëåâîé êëåøíè; d ôîðìà òåëüñîíà. Ìàñøòàá 1 cì (b, c), 1 ìì (a, d).

negative correlation between SL and palm width/length ratio (Fig. 26). DISTRIBUTION. From the southern Chukchi Sea [Makarov, 1938] southward to Korea and Japan [Vinogradov, 1947] and from Point Barrow to the Pribilov Islands and the Aleutians Islands [McLaughlin, 1974]; subtidal to 182 m [Vinogradov, 1947; McLaughlin, 1974]. In the references area the species was collected at the depth between 20 and 250 m (water temperature 0.216.24?C and salinity 32.5733.59 p.p.t.). Our records indicate a deeper occurence of P. trigonocheirus than known previously (180 m). BIOLOGY. Large-size males (SL 1018 mm) and smallsize females (SL 1.77 mm) dominated in the samples. The percentage of ovigerous females was low (1.6%). Other species present in the same catches were Chionoecetes bairdi, C. opilio, Hyas coarctatus, Labidochirus splendescens, Crangon communis, Pandalus goniurus, and Argis lar.

Pagurus brandti (Benedict, 1892) Fig. 26, 27.
MATERIAL. 35 (SL 6.318.9 mm), 7 non-ovigerous (SL 3.416.1 mm). LOCALITY. Haul # 10, 20, 30, 38, 48, 49, 55, 56; Dredge # 4, 34, 43.

REMARKS ON MORPHOLOGY. The ratio of the right cheliped palm width to length varies from 1.68 to 2 in males and from 1.63 to 1.92 in females. DISCUSSION. Pagurus brandti is distinguished from P. trigonocheirus by the differences in the shape of the left chela and the morphology of telson [McLaughlin, 1974; Ivanov, 1979a], and by larval morphology [Ivanov, 1979b]. Besides of this, P. trigonocheirus is separated from P. brandti by having a more slender and longer palm of the right cheliped; in fact, there is no overlap between these


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Fig. 26. Shield length vs. ratio of right cheliped palm width/length in Pagurus trigonocheirus and P. brandti: a ; b . Ðèñ. 26. Ãðàôèêè èçìåíåíèÿ îòíîøåíèÿ äëèíû êëåøíè ê åå øèðèíå ñ óâåëè÷åíèåì ðàçìåðîâ ó Pagurus trigonocheirus è P. brandti: a ñàìöû; b ñàìêè.

species in the right palm width to length ratio throughout the size range (Fig. 26). DISTRIBUTION. Chukchi Sea, Bering Sea [McLaughlin, 1974; Ivanov, 1979a], Sea of Okhotsk; depth 59294 m [Sokolov, 1998]. In the West Kamchatka area the species was collected the water temperature range from 0.24 to 1.73?C, salinity ranged from 32.6 to 33.38 p.p.t. Other species present in the catches along with P. brandti were Chionoecetes opilio, Pandalus goniurus, Crangon communis, and Labidochirus splendescens.

FAMILY LITHODIDAE SAMOUELLE, 1819 Subfamily Lithodinae Ortmann, 1901 Genus Paralithodes Brandt, 1849 Paralithodes camtschaticus (Tilesius, 1815)
LOCALITY. Haul # 1, 3, 4, 5, 6, 10, 11, 13, 14, 15, 16, 17, 18, 21, 22, 23, 25, 26, 27, 28, 29, 30, 31, 33, 34, 37, 38, 39, 40,


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Fig. 27. Pagurus brandti (Benedict, 1892). (SL 11.5 mm): a shield; b left chela and carpus in dorsal view; c right chela and carpus in dorsal view; d telson. Scale 1 cm (ac), 1 mm (d). Ðèñ. 27. Pagurus brandti (Benedict, 1892). Ñàìåö (ÄÙ 11.5 ìì): a ôîðìà ùèòêà; b ôîðìà ïðàâîé êëåøíè è êàðïóñà; c ôîðìà ëåâîé êëåøíè è êàðïóñà; d ôîðìà òåëüñîíà. Ìàñøòàá 1 cì (ac), 1 ìì (d).

DISTRIBUTION. From the eastern Olutorsky Bay [Ivanov, 2000] and the Penzhinskaya Guba (Sea of Okhotsk) to the East coast of Korea and Honshu Island in the northwestern Pacific, from Norton Sound to Queen Charlotte Islands in the northeastern Pacific [Vinogradov, 1946]; 4510 m [Klitin, 1996]. P. camtschaticus has been introduced in the North Atlantic in the 1960s and now this species is common in Barents Sea [Orlov and Ivanov, 1978; Kuzmin & Olsen, 1994; Kuzmin et al., 1996]. In the references area crabs were collected at depth from 19 to 295 m (water temperature 0.216.24?C and salinity

41, 42, 43, 44, 45, 46, 48, 49, 50, 51, 52, 53, 54, 55, 56, 57, 58; Dredges # 9, 10, 11, 20, 39, 44.

32.233.39 p.p.t.). The species was most abundant at depth between 19 and 50 m. BIOLOGY. Large-size males with CW 110210 mm dominated in the trawl catches. Female crabs were rare (4% of collected specimens). Ca. 95 % of females were ovigerous.

Paralithodes platypus Brandt, 1850
MATERIAL. 19 (CW 105155 mm), 1 non-ovigerous (CW 104 mm). LOCALITY. Haul # 10, 17, 18, 30, 38, 41, 47, 48, 51, 52, 53.


Decapod Crustaceans of the Southwest Kamchatka Shelf
DISTRIBUTION. From the Bering Straight and the northern Part of the Sea of Okhotsk to Peter the Great Bay along the Asian coast and from Point Barrow to northern British Columbia [Ivanov, 2001]; 12500 m [Makarov, 1941]. In the references area this species was found at depth between 52 and 114 m (water temperature 0.212.02?C and salinity 32.5733.15 p.p.t.) Other species present in the same catches were Chionoecetes bairdi , Hyas coarctatus, Labidochirus splendescens , Pagurus trigonocheirus , Pandalus goniurus, and Paralithodes camtschaticus.

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Genus Lithodes Latreille, 1806 Lithodes aequispinus Benedict, 1894
MATERIAL. 97 (CW 101170 mm), 55 ovigerous (CW 101150 mm), 15 non-ovigerous (CW 86 110 mm). LOCALITY. Haul # 26, 32, 33, 34, 36, 43, 44, 45, 46, 47, 49, 55.

DISTRIBUTION. From the Bering Sea to the Sea of Okhotsk and Pacific coast of Northern Hokkaido [Takeda, 1982] and to the Aleutian Islands and southern British Columbia [Jamieson and Sloan, 1985]; 100800 m. In the references area the species was collected at depth between 100 and 300 m (water temperature 0.211.81?C and salinity 32.6733.39 p.p.t.). BIOLOGY. Large-size males with CW 130160 mm dominated in catches. Females were recorded rather frequently (in ca. 42% of catches). Specimens with orange colored eggs dominated among recorded ovigerous females. This species frequently occurred together with Chionoecetes opilio, Hyas coarctatus, Pandalus borealis, Labidochirus splendescens, and Paralithodes camtschaticus.

Fig. 28. Hyas coarctatus alutaceus, males. Shape of carapace in dorsal (CW 8.1 mm); b (CW 17.5 mm); c (CW 43.9 mm); d mm). Scale 1cm. Ðèñ. 28. Hyas coarctatus alutaceus. Ôîðìà êàðàïàêñà (âèä ñâåðõó): a (ØÊ 8,1 ìì); b ñàìåö (ØÊ 17,5 ìì); c ñàìåö (ØÊ 43,9 ìì); d (ØÊ 49,2 ìì). Ìàñøòàá 1 cì.

view: a (CW 49.2 ñàìåö ñàìåö

FAMILY MAJIDAE SAMOUELLE, 1819 Genus Chionoecetes Kröyer, 1838 Chionoecetes opilio (O. Fabricius, 1788)
MATERIAL. 593 (CW 2.8135 (CW 6075 mm), 159 non-ovigerous LOCALITY. Haul # 6, 8, 9, 19, 20, 21, 33, 34, 36, 37, 38, 39, 43, 44, 45, 46, 47, 55, 56; Dredge # 7, 10, 11, 13, 14, 16, 17, 35, 37, 38, 42, 43, 44, 45. mm), 5 ovigerous (CW 12.180 mm). 24, 26, 27, 30, 31, 32, 48, 49, 50, 51, 52, 53, 24, 26, 27, 29, 33, 34,

REMARKS ON MORPHOLOGY. Variability of the most principal characters of C. opilio in the references area was

discussed by Sokolov[2001]. Sex and size compositions of C. opilio based on materials of R/V Prof. Levanidov were described earlier [Ivanov & Sokolov, 1997]. DISTRIBUTION. From the Wrangel Island to Point Barrow in the Arctic Ocean; from the Bering Sea to the Sea of Okhotsk and the Sea of Japan, and to British Columbia in the North Pacific [Makarov, 1941; Fedoseev & Slizkin, 1988]; from Greenland to the Gulf of Maine [Squires, 1990] in the Western Atlantic and in the Barents Sea in the Eastern Atlantic [Kuzmin et al., 1996]; 71000 m [Vinogradov, 1950]. In the references area these crabs were common at depth between 20 and 300 m (most frequently at 100300 m depth), at water temperature ranging from 0.21 to 3.94?C and salinity between 32.2 and 33.39 p.p.t.


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dominated in catches) south of 53?N, and only a few specimens were found at north of 53?N. Water temperature in the place of collection varied from 0.34 to 6.24?C, salinity was in the range of 32.2 to 33.37 p.p.t. Other decapod species obtained in the same catches wereParalithodes camtschaticus, Hyas coarctatus, Labidochirus splendescens, Pagurus trigonocheirus, Pandalus goniurus.

Genus Hyas Leach, 1814 Hyas coarctatus alutaceus Brandt, 1851 Fig. 2830.
MATERIAL. 189 (CW 2.884.2 mm), 46 ovigerous (CW 16.753.5 mm), 112 non-ovigerous (CW 347.5 mm). LOCALITY. Houl # 12, 14, 15, 17, 18, 23, 29, 30, 34, 40, 41, 42, 43, 52, 53, 58; Dredge # 8, 12, 13, 17, 24, 28, 29, 33, 34, 35, 39, 44.

REMARKS ON MORPHOLOGY. The Pacific subspecies alutaceus differs from the nominal Atlantic subspecies in the following characters: 1) in the Pacific subspecies carapace is wider and more rounded posteriorly, with fewer and smaller tubercles or ridges; 2) rostrum is shorter and wider in the Pacific specimens than in the Atlantic population (CL 7.19.3 times greater than the rostrum length in H. coarctatus alutaceus, while in the nominal subspecies it is only 4.56.4 times greater) [Squires, 1990]. The shape of rostrum and carapace of males and females varies being strongly related to size/ age (Fig. 28, 29ac). Largesize males and females, as a rule, have a relatively shorter rostrum and wider carapace than small specimens. The ratio of Fig. 29. Hyas coarctatus alutaceus: ac shape of female carapace in dorsal carapace length to rostrum length ranges view; d, e right chela. a non-ovigerous (CW 28.1 mm); b non-ovigerous from 3.4 to 6.7 in the small crabs with CW (CW 15.7 mm); c non-ovigerous (CW 15.7 mm); d (CW 63.2 mm); 2.810.0 mm; and from 3.7 to 13.0 in the e non-ovigerous (CW 36.2 mm). Scale 1 cm. males and females of CW 1030 mm; it Ðèñ. 29. Hyas coarctatus alutaceus: ac ôîðìà êàðàïàêñà (âèä ñâåðõó); d, e ôîðìà êëåøíè. a ñàìêà áåç ÿèö (ØÊ 28,1 ìì); b ñàìêà áåç ÿèö (ØÊ falls between 5 and 14 in the specimens 15,7 ìì); d ñàìêà áåç ÿèö (ØÊ 15,7 ìì); d ñàìåö (ØÊ 63,2 ìì); e ñàìêà with CW 3050 mm, and between 7 and 15 áåç ÿèö (ØÊ 36,2 ìì). Ìàñøòàá 1 cì. in almost all large-size males with CW greater than 60 mm (Fig. 30c). The number and size of the tubercles and the ridges on the The decapod species present in the same catches with C. dorsal surface of the carapace varies in different specimens. opilio were Pagururs trigonocheirus, Labidochirus splendeMature males can be well separated from the females and scens, Crangon communis, Hyas coarctatus, and Argis lar. immature males by the shape of chela (Fig. 29d, e). DISTRIBUTION. North Pacific and adjacent areas of the Chionoecetes bairdi Rathbun, 1902 Arctic Ocean only [Squires, 1990]. From the Bennet Island to the Beaufort Sea in the Arctic Ocean; from the Bering Sea to LOCALITY. Haul # 1, 2, 3, 4, 5, 6, 8, 11, 12, 13, 14, 15, 16, the La Perouse Strait in the North Pacific; subtidal to 1650 m 17, 19, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 34, 35, 38, [Vinogradov, 1950]. 39, 40, 41, 42, 43, 44, 47, 48, 49, 51, 52, 53, 54; Dredge # 1, 7, H. coarcatatus alutaceus is one of the most common 8, 9, 10, 11, 14, 15, 17, 18, 19, 20, 24, 25, 33, 35, 37, 39, 43. species in the West Kamchatka area. This species was found DISTRIBUTION. From the southern Bering Sea to the at depth between 21 and 155 m (water temperature 0.34 Aleutian Islands and Oregon, and from the Cape Navarin 5.18?C and salinity 32.3533.21 p.p.t.). [Slizkin, 1982, 1990] to 59?50N in the northeastern Sea of BIOLOGY. Small-size specimens with CW<25 mm domOkhotsk [Sokolov, 2001]; 20415 m [Slizkin, 1982]. inated in dredge catches. Four modal groups with mean CW In the references area C. bairdi were common at depth of respectively 8, 30, 32, 43, and 62 mm were detected in the between 20 and 300 m (at 40100 m depth this species


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Fig. 30. Hyas coarctatus alutaceus: a relationship between chela length and carapace length in large-clawed and smallclawed males; b carapace width frequency distributions of males and females; c carapace width vs. ratio of carapace length/ rostrum length. Ðèñ. 30. Hyas coarctatus alutaceus: a îòíîøåíèå äëèíû êàðàïàêñà è äëèíû êëåøíè â ëîãàðèôìè÷åñêîé çàâèñèìîñòè; b ãðàôèêè ðàçìåðíîãî ñîñòàâà ñàìöîâ è ñàìîê; c ãðàôèê èçìåíåíèÿ îòíîøåíèÿ äëèíû ðîñòðóìà ê äëèíå êàðàïàêñà ñ óâåëè÷åíèåì øèðèíû êàðàïàêñà.


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carapace width frequency distributions (Fig. 30b). The percentage of ovigerous females was relatively low (29%). Crabs with bright orange eggs dominated amongst ovigerous females. Other ovigerous females (about 15%) were with developed embryos. Small-clawed and largeclawed males were recorded in catches (Fig. 30a). Large-clawed males dominated among largesize specimens with CW exceeding 55 mm. The minimum size (CW) of large-clawed males was 29.7 mm, the maximum size of small-clawed males was 53.5 mm. The relation between chelae length (Y) and CW (X) for largeclawed males (morphometrically mature) and small-clawed (adolescent) males can be described as follows: lnY=1.0154lnX 0.8415, R 2=0.8932 and lnY= 1.0197lnX1.2396, R2=0.9836, respectively. The proportion of large-clawed males in catches was 14.3%. The species occured together with Chionoecetes opilio, C. bairdi, Pandalus goniurus, Pagurus trigonocheirus , Labidochirus splendescens , Argis lar , and Crangon communis.

V.I. Sokolov

Genus Oregonia Dana, 1851 Oregonia gracilis Dana, 1851 Fig. 31.
MATERIAL. 1 2 (CW 8.1 29.4 mm), 3 ovigerous (CW 19.224.5 mm), 6 non-ovigerous (CW 6.320.7 mm). LOCALITY. Houl # 18, 29, 30; Dredge # 4, 10, 11, 17, 18, 24.

DISTRIBUTION. From the Commander Islands to Inubo Zaki (Japan) and Chefoo (China) and from the Nunivak Island to California [Vinogradov, 1950], but no records in the western Sea of Okhotsk and in the Primorie Region (northwestern Sea of Japan) are known; subtidal to 390 m depth [Vinogradov, 1950]. Crabs were collected by R/V Prof. Levanidov at depth from 51 to 180 m (water temperature 1.222.15?C and salinity 32.7233.08 p.p.t.). BIOLOGY. Five large-clawed (Fig. 31b) and seven smallclawed (Fig. 31d) males were caught. Ca. 30% of females had orange eggs. Other decapod species present in catches together with this species were Hyas coarctatus, Chionoecetes bairdi, Labidochirus splendescens, Pagurus trigonocheirus, and Pandalus goniurus.

Fig. 31. Oregonia gracilis Dana, 1851: a carapace in dorsal view; b anterior part of carapace; c right cheliped in dorsal view; d right chela; e left chela. a, c non-ovigerous (CW 14.8 mm); b non-ovigerous (CW 22.8 mm); d (CW 8.1 mm); e (CW 26.7 mm). Scale 1 cm (a, e), 1 mm (bd). Ðèñ. 31. Oregonia gracilis Dana, 1851: a ôîðìà êàðàïàêñà è ðîñòðóìà (âèä ñâåðõó); b ôîðìà ïåðåäíåé ÷àñòè êàðàïàêñà; c ôîðìà ïðàâîãî ïåðâîãî ïåðåîïîäà; d ôîðìà êëåøíè; e ôîðìà êëåøíè. a, c ñàìêà áåç ÿèö (ØÊ 14,8 ìì); b ñàìêà áåç ÿèö (ØÊ 22,8 ìì); d ñàìåö (ØÊ 8,1 ìì); e ñàìåö (ØÊ 26,7 ìì). Ìàñøòàá 1 cì (a, e), 1 ìì (bd).

FAMILY ATELECYCLIDAE ORTMANN, 1893 Genus Erimacrus Benedict, 1892 Erimacrus isenbeckii (Brandt, 1848)
MATERIAL. 125 (CW 51145 mm); 5 non-ovigerous (CW 41118 mm). LOCALITY. Haul # 1, 2, 12, 13, 14, 15, 16, 17, 21, 22, 23, 24, 27, 28, 29, 41, 42, 53; Dredge # 20.


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DISTRIBUTION. Sea of Japan (from 50?N to Southern part of Honshu Island and Southern part of Korea) [Armetta and Stevens, 1987; Ivanov, 2001], Sea of Okhotsk (in SouthEast Sakhalin from Aniva Bay to 49?30N [Ivanov, 2001]; Kuril Islands; in the West Kamchatka north to Cape Khayryuzova [Vinogradov, 1947]); in the Eastern Kamchatka from Cape Lopatka to 54?30N [Ivanov, in press], and in eastern Bering Sea from St. Matthew Island along Alaska Peninsula to Attu Island [Armetta and Stevens, 1987]; 0350 m depth [Vinogradov, 1950]. The species was common at depth from 19 to 82 m (water temperature 0.344.97?C and salinity 32.232.73 p.p.t.). Other species occasionally present in the catches together with this species wereParalithodes camtschaticus, Chionoecetes bairdi, Hyas coarctatus, Pagurus trigonocheirus, and Labidochirus splendescens.

131

Genus Telmessus White, 1846 Telmessus cheiragonus (Tilesius, 1812)
MATERIAL. 46 (CW 4180 mm), 1 ovigerous (CW 53 mm). LOCALITY. Haul # 14, 15, 21, 27.

corded in this region in the deep waters (deeper than 307 m). The following eight species: Pandalus hypsinotus Brandt, 1851; Pandalopsis lamelligera (Brandt, 1851); Spirontocaris prionota (Stimpson, 1864), S. arcuata Rathbun, 1902; Lebbeus shrencki (Brashnicov, 1907), Eualus suckleyi (Stimpson, 1864); Pagurus undosus (Benedict, 1892) and Hapalogaster grebnitzkii Schalfeew, 1892 rarely occur in the southern part of West Kamchatka shelf [Vinogradov, 1947]. These species were as a rule recorded north to the study area. Although Argis crassa (Rathbun, 1902), Eualus townsendi (Rathbun, 1902), Heptacarpus flexus (Rathbun, 1902) and Sclerocrangon boreas (Phipps, 1774) were reported to be rather common species in this part of Sea of Okhotsk [Vinogradov, 1947] but they were not found in the present survey.

Distribution of decapod species in the Southern part of West Kamchatka shelf
Amongst 33 species and subspecies recorded in the present study the most common species were Chionoecets bairdi , Chionoecetes opilio , Paralithodes camtschaticus, Pagurus trigonocheirus, Labidochirus splendescens, and Hyas coarctatus alutaceus. All these species but Chionoecetes bairdi are widely ranged in the Sea of Okhotsk. The latter species was recorded there only recently [Slizkin, 1982]. Paralithodes camtschaticus and Chionoecetes bairdi were most abundant species at the shallow depth. Pandalus borealis eous and Chionoecetes opilio dominated at the depth from 200 to 300 m to the north of 52?00N. The assemblage of the most common species collected by the R/V Prof. Levanidov is subdivided into following groups according to the depth of occurence (Fig. 32): 1) shallow water species, e.g. Telmessus cheiragonus, Erimacrus isenbekii and Crangon dalli, which recorded at the depth from 20 to 100 m, but were numerous at depth 2025 m; 2) species occurred at depth from 20 to 200 m being most common between 60 and 80 m, e.g. Spirontocaris murdochi, Hyas coarctatus alutaceus, Mesocrangon intermedia, and Pagurus trigonocheirus; 3) widely distributed species most common at depth from 50 to 100 m, e.g. Chionoecetes bairdi, Paralithodes camtschaticus, Argis lar, Labidochirus splendescens, Pandalus goniurus, and Crangon communis; 4) deep sea species most abundant deeper than 200 m, e.g. Chionocetes opilio, Pandalus borealis eous, Lithodes aequispinus, and Argis ochotensis. Some species of the second and the third groups (Hyas coarctatus alutaceus, Paralithodes camtschaticus) which occurred at most station within the depth range of 6070 m (Fig. 32a), were most abundant at shallow water stations (Fig. 32b). Large-size males of Paralithodes camtschaticus dominated in the trawls catches and juvenile specimens of Hyas coarctatus alutaceus dominated in dredge samples at depth from 20 to 25 m. Mature males

DISTRIBUTION. From the nothern Bering Sea to California and to the North Korea and the Hokkaido Island [Makarov, 1941; Vinogradov, 1950]; depth 050 m [Vinogradov, 1950]. In the reference area the species was collected at depth between 19 and 21 m (water temperature 3.925.18?C and salinity 32.232.52 p.p.t.). Common co-occuring species were Chionoecetes bairdi, Paralithodes camtschaticus, Hyas coarctatus, and Erimacrus isenbeckii.

Decapod fauna of the West Kamchatka shelf
Vinogradov [1947] recorded 91 decapod taxa in the Sea of Okhotsk. Since that the following eight decapod species were found in this region: Paralomis multispina (Benedict, 1894) and Lithodes couesi Benedict, 1894 [Nizyaev, 1992]; Chionoecetes bairdi [Slizkin, 1982]; Pandalopsis coccinata Urita, 1941 [Komai, 1994]; Argis ochotensis Komai, 1997 [Komai, 1997]; Lebbeus vinogradowi Zarenkov, 1960 [Zarenkov, 1960]; Pagurus brandti [Sokolov, 1998]; and Spirontocaris brevidigitata Kobjakova, 1935 (this study). On the other side Hayashi [1977] synonymized Spirontocaris makarovi (Kobjakova, 1936) and Hetairus brevipes Kobjakova, 1936 with respectively Spirontocaris ochotensis (Brandt, 1851) andLebbeus unalaskensis (Rathbun, 1902). Therefore 97 decapod species are known for the Sea of Okhotsk and 50 species are recorded for the West Kamchatka shelf [Kobjakova, 1958; this study]. The entire collection of R/V Prof. Levanidov consists of 18 genera and 33 species/ subspecies. Another 17 taxa which were recorded in this region earlier were not found by this expedition. Heptacarpus camtschaticus (Stimpson, 1860), Spirontocaris ochotensis, and Dermaturus mandtii Brandt, 1850 occur in the Sea of Okhotsk at shallow depth (030 m). On the other hand Argis robusta (Kobjakova, 1935) were re-


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Fig. 32. Distribution of most common species according to depth: a depth at which species were most common; b depth at which species were most numerous. Ðèñ. 32. Ðàñïðåäåëåíèå ïî ãëóáèíàì íàèáîëåå ìàññîâûõ âèäîâ: a ãëóáèíà íà êîòîðîé âèä íàèáîëåå ÷àñòî âñòðå÷àëñÿ; b ãëóáèíà, íà êîòîðîé âèä áûë íàèáîëåå ìíîãî÷èñëåíåí.

and females of Hyas coarctatus alutaceus usually occurred at depth between 60 and 100 m. The following groups of the most common species may be defined according to temperature preference (Fig. 33): 1) the species which were found in the narrow temperature range being most abundant at water temperature 1.31.8?C, e.g. Argis ochotensis, Pandalus borealis eous, Pandalus goniurus, and Lithodes aequispinus; 2) relatively cold-water species (Spirontocaris murdochi, Argis lar, Mesocrangon intermedia, Crangon communis), which usually occurred at water temperature less than 1?C; 3) species occurring within a wide temperature ranges, most common and numerous at temperature between 1 and 3?C (Chionoecetes opilio, Pagurus trigonocheirus, Crangon dalli, Hyas coarctatus alutaceus);

4) relatively warm-water species (Paralithodes camtschaticus, Erimacrus isenbeckii, Telmessus cheiragonus). Juvenile specimens of Hyas coarctatus alutaceus were most abundant in dredge catches at water temperature 5?C, but mature specimens of this species usually occurred at water temperature 12?C. Labidochirus splendescens was common in relatively warm water (temperature between 3 and 3.2?C) but the most abundant catches of this secies were obtained at water temperature 1.5?C.
ACKNOWLEDGEMENTS I wish to express my particular gratitude to Drs. V.A. Spiridonov (Moscow State University) and B.G. Ivanov (VNIRO) for their valuable comments and stylistic help. I am indebted to the crew members and the scientific team of the


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Fig. 33. Distribution of most common species according to bottom water temperature: a temperature at which species were most common; b temperature at which species were most numerous. Ðèñ. 33. Ðàñïðåäåëåíèå ïî òåìïåðàòóðå íàèáîëåå îáû÷íûõ âèäîâ: a òåìïåðàòóðà, ïðè êîòîðîé âèä íàèáîëåå ÷àñòî âñòðå÷àëñÿ; b òåìïåðàòóðà, ïðè êîòîðîé âèä áûë íàèáîëåå ìíîã÷èñëåíåí.

TINRO-Centre for their cooperation on board R/V Prof. Levanidiv. The expedition was organized by the TINROCentre (Vladivostok), the preparation of this paper was supported by the project # 99-04-4853 of the Russian Foundation for Basic Research.

References
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Nauka. 162 pp. [in Russian]. Makarov V.V. 1938. [Crustacés Décapodes anomures] // Fauna SSSR (n. ser # 16). Rakoobraznye [Crustaceans]. Vol.10. No.3. Leningrad: Acad. Sci. USSR Publ. House. 324 pp. [in Russian with English summary]. Makarov W.W. 1941. [The Decapod Crustacea of the Bering and the Chukchees Seas] // Issledovanija dalnevostochnikh morei SSSR. Vol.1. P.111163 [in Russian with English summary]. McLaughlin P.A. 1974. The hermit crabs (Crustacea, Decapoda, Paguridae) of Northwestern North America // Zool. Verh. Vol.130. 396 pp. Miyake S., Sakai K., Nishikawa S. 1962. A fauna list of the decapod Crustacea from the coasts washed by the Tsushima warm current // Rec. oceanogr. Works Japan. Spec. no. 6. P.121131. Nizyaev S.A. 1992. [Distribution and abundance of deep-water crabs in the Sea of Okhotsk] // Ivanov B.G. (ed.). Rybokhoziaistvennye issledovaniaya morskikh bespozvonochnykh [Fishery researches of marine invertebrates]. Moscow: VNIRO. P.2637 [in Russian]. Orlov Y.I., Ivanov B.G. 1978. On the introduction of the Kamchatka king crab Paralithodes camtschatica (Decapoda Anomura: Lithodidae) into the Barents Sea // Mar. Biol. Vol.48. P.373375. Rathbun M.J. 1924. New species and subspecies of Spider Crabs // Proc. U. S. Nat. Mus. Bull. Vol.64. No.14. 25 pp. Rathbun M.J. 1925. The spider crabs of America // Bull. U. S. Nat. Mus. No.129. 613 pp. Rathbun M.J. 1930. The Cancroid Crabs of America // Ibid. No.152. 593 pp, 283 pls. Rathbun M.J. 1932. Preliminary descriptions of new species of Japanese crabs // Proc. Biol. Soc. Wash. Vol.45. P.2838. Rodin V.E. 1967. [Some features of biology and peculiarities of distribution of Kamchatka crab concentrations off West Kamchatka coast] // Izvestiya TINRO. Vol.61. P.243254 [in Russian]. Rodin V.E. 1969. [Distribution of Kamchatka crab off Western Kamchatka] // Trudy VNIRO. Vol.65. P.368377 [in Russian]. Rodin V.E. 1985. [Spatial and functional structure of king crab population] // Izvestiya TINRO. Vol.110. P.8697 [in Russian]. Slizkin A.G. 1982. [Distribution of snow crab (Chionoecetes gen.) and their habitat conditions in the North Pacific] // Ibid. Vol.106. P.2633 [in Russian]. Slizkin A.G. 1990. Tanner Crabs (Chionoecetes opilio, C. bairdi) of the Northwest Pacific: Distribution, Biological Peculiarities, and Population Structure // Proc. Internat. Symp. on King and Tanner Crabs, November 2830, 1989, Anchorage, Alaska USA. Alaska Sea Grant College Program Rep. # 9004. P.2733. Sokolov V.I. 1997. [Variability of deep sea prawn Pandalus borealis (Crustacea, Decapoda, Pandalidae)] // Zool. Zhurnal. Vol.76. No.3. P.281286 [in Russian with English summary]. Sokolov V.I. 1998. New data about distribution and variability of hermit crabs Pagurus brandti and P. trigonocheirus in the Sea of Okhotsk // Ibid. Vol.77. No.5. P.558564 [in Russian with English summary]. Sokolov V.I. 2001. [New data on the distribution C. bairdi with the notes on the morphological variability (Crustacea, Decapoda, Majidae)] // Ibid. Vol.80. No.1. P.3238 [in Russian with English summary]. Squires J.H. 1990. Decapoda Crustacea of the Atlantic Coast of Canada // Canadian Bull. of Fish. Aq. Sci. No.221. P.532. Takeda M. 1982. Keys to the Japanese and foreign crustaceans fully illustrated in colors. 1st Edition. Tokyo: Hokuryukan Publisher. 285 pp. Vinogradov L.G. 1945. [The year cycle in the life and migrations of the Paralithodes camtschatica in the northern part of the West Kamchatka shelf] // Byull. TINRO [Pacif. Sci. Ist. Fish. Oceanogr.]. No.19. P.154 [in Russian]. Vinogradov L.G. 1946. [The Geographical Distribution of the Kamchatka Crab Paralithodes camtschatica (Til.)] // Ibid. No.22. P.195232. [in Russian].

IV. The genus Spirontocaris Bate // J. Shimonoseki University of Fish. Vol.25. No.3. P.155186. Holthuis L. B., 1947. The Decapoda of the Siboga Expedition. Part IX. The Hippolytidae and Rhynchocinetidae collected by the Siboga and Snellius Expeditions with remarks on other species // Siboga-Expedite, Monographie XXXIXa8 de, 100 pp. Ivanov B.G. 1979a. [New data on hermit crabs of the North Pacific. 1. Notes on Pagurus brandti and P. trigonocheirus (Crustacea, Decapoda, Paguridae)] // Zool. Zhurnal. Vol.58. No.6. P.816823 [in Russian, with English summary]. Ivanov B.G. 1979b. [New data on hermit crabs of the North Pacific. 2. The first larval stages of some species reared in the laboratory (Crustacea, Decapoda, Paguridae)] // Ibid. Vol.58. No.7. P.977986 [in Russian, with English summary]. Ivanov B.G. 1994. [Fishery hydrobiology: heritage, problems, perspective] // Rybnoe Khoziaistvo. No.5. P.4346, No.6. P.3037 [in Russian]. Ivanov B.G. 2001 [Decapod crustaceans (Crustacea Decapoda) of the North Pacific as a pool for introduction into the Atlantic: the introduction is feasible but is there reason?] // Ivanov B.G. (ed.). Issledovaniya kommercheskikh dejapod i vodoroslei. Trudy VNIRO. P.5285 [in Russian, with English summary]. Ivanov B.G., Sokolov V.I., 1997. [The snow crab Chionoecetes opilio in the Sea of Okhotsk and the Bering Sea (Crustacea Decapoda Brachyura Majidae)] // Arthropoda Selecta. Vol.6. No.3/4. P.6386 [in Russian, with English summary]. Jamieson G.S. and N.A. Sloan. 1985. King crabs in British Columbia // Proc. Internat. King Crab Symp., Anchorage, Alaska, USA, January 2224, 1985. Univ. of Alaska, Alaska Sea Grant Rep. No.85-12. P.4962. Klitin A.K. 1996. [Kamchatka crab of shelf area of Sakhalin Island (Literature revue, history of fisheries, spatial and functional structure of population] // Vestnik Sakhalinskogo Muzeya. No.3. P.324342 [in Russian]. Kobjakova Z.I. 1936. [Zoogeographical survey of the Decapoda fauna from the Okhotsk and Japanese seas] // Trudy Leningradskogo Obshestva Estestvoispitatelei. Vol.65. P.185 228 [in Russian, with German summary]. Kobjakova Z.I. 1937. [Systematic review of the Decapoda of the Okhotsk and Japanese Seas] // Uchenye Zapiski Leningradskogo Universiteta. Vol.15. P.93154 [in Russian, with German summary]. Kobjakova Z.I. 1958. [Decapod crustaceans of the Southern Kuril Islands] // Issledovanija dalnevostochnikh morei USSR. Vol.5. P.220248 [in Russian]. Komai T. 1994. Deep-sea shrimps of the genus Pandalopsis (Decapoda: Caridea: Pandalidae) from the Pacific coast of Eastern Hokkaido, Japan, with the description of two new species // J. Crustacean Biol. Vol.14. No.3. P.538559. Komai T. 1997. Revision of Argis and related species (Decapoda: Caridea: Crangonidae), with description of new species from the Ochotsk Sea // Ibid. Vol.17. No.1. P.135161. Komai T. 1999. A revision of the genus Pandalus (Crustacea: Decapoda: Caridea: Pandalidae) // J. Natural History. Vol.33. P.12651372. Kuzmin S., Olsen S. 1994. Barents Sea king crab (Paralithodes camtschatica): The transplantation experiments were successful // Internat. Council Explor. Sea (I.C.E.S.), Shellfish Committee C.M. 1994/K:12, 12 pp, 7 figs. Kuzmin S., Olsen S., Gerasimova O. 1996. Barents Sea king crab (Paralithodes camtschaticus): Transplantation experiments were successful // High latitude crabs: biology, management, and economics. Proc. Internat. Symp. on biology, management, and economics of crabs from high latitude habitats, Anchorage, Alaska, USA, October 1113, 1995. Univ. Alaska Sea Grant College Program Rep. No. 96-02. P.649663. Kuzmin S.A, Akhtarin S.M., Menis D.T. 1998. [The first findings of snow crab Chionoecetes opilio (Decapoda, Majidae) in the Barents Sea] // Zool. Zhurnal. Vol.77. No.4. P.489491 [in Russian, with English summary]. Makarov R.R. 1966. [Larvae of the shrimps, hermits and crabs of Western Kamchatka Shelf and their distribution]. Moscow:


Decapod Crustaceans of the Southwest Kamchatka Shelf
Vinogradov L.G. 1947. [Decapod crustaceans of the Okhotsk Sea] // Izvestiya TINRO. Vol.25. P.67124 [in Russian]. Vinogradov L.G. 1950. [Classification of shrimps, prawn and crabs from Far East Seas] // Izvestiya TINRO. Vol.33. P.179 358 [in Russian]. Vinogradov L.G. 1969. [On the recruitment mechanisms in the stock of Kamchatka crab (Paralithodes camtschatica) off Western Kamchatka in the Okhotsk Sea] // Trudy VNIRO [All Union Res. Inst. Mar. Fish. Oceanogr.]. Vol.65. P.337344

135

[in Russian]. Zarenkov N.A. 1960. [Note about some decapod Crustacea of the Okhotsk and Bering Seas] // Trudy Instituta Okeanologii. Vol.34. P.343350 [in Russian]. Williams A.B. 1984. Shrimps, lobsters and crabs of the eastern United States, Maine of Florida. Smiths. Inst. Press, Washington, DC. 550 pp. Williams A.B., Wigley L. 1977. Distribution of Decapod Crustacea off Northeastern United States Based on Specimens at the Northeast Fisheries Center, Woods Hole, Massachusetts // NOAA Technical Rep. NMFS. No.407. 44 pp.

Attachment. Ïðèëîæåíèå.
1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41.

Haul ##

01.07.96 01.07.96 02.07.96 02.07.96 02.07.96 02.07.96 03.07.96 03.07.96 04.07.96 04.07.96 04.07.96 04.07.96 05.07.96 05.07.96 05.07.96 05.07.96 05.07.96 06.07.96 06.07.96 06.07.96 07.07.96 08.07.96 08.07.96 09.07.96 09.07.96 10.07.96 10.07.96 12.07.96 12.07.96 13.07.96 13.07.96 13.07.96 13.07.96 14.07.96 14.07.96 15.07.96 15.07.96 15.07.96 15.07.96 16.07.96 16.07.96

Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth

30 m 80 m 20 m 40 m 69 m 200 m 100 m 300 m 250 m 152 m 84 m 50 m 31 m 21 m 19 m 43 m 60 m 101 m 217 m 294 m 21 m 31 m 52 m 82 m 151 m 247 m 20 m 47 m 61 m 99 m 199 m 245 m 147 m 81 m 52 m 300 m 200 m 114 m 60 m 51 m 53 m

51œ17 51œ14 51œ33 51œ32 51œ31 51œ31 51œ31 51œ33 51œ47 51œ45 51œ45 51œ45 51œ45 51œ45 52œ00 52œ00 52œ00 52œ00 51œ59 52œ20 52œ15 52œ15 52œ15 52œ15 52œ15 52œ15 52œ30 52œ30 52œ30 52œ30 52œ30 52œ45 52œ45 52œ45 52œ45 53œ00 53œ00 53œ00 53œ00 53œ00 53œ15



7N156œ313E 7N156œ296E 7N156œ273E 1N-156œ221E 5N156œ175E 7N156œ111E 3N156œ313E 7N156œ074E 5N155œ507E N155œ57E 8N156œ029E 8N156œ171E N156œ24E N156œ26E N156œ25E N156œ18E N156œ08E N155œ44E N155œ20E N154œ548E N156œ228E N156œ18E N156œ02E N155œ375E N155œ20E N154œ56E N156œ14E N155œ55E N155œ41E N155œ18E N154œ57E N154œ43E N155œ05E N155œ22E N155œ39E N154œ25E N154œ43E N155œ09E N155œ36E N155œ45E N155œ42E

T=4.93œ T=2.148œ T=6.24œ T=4.31œ T=3.12œ T=1.225œ T=1.87œ T=1.63œ T=1.54œ T=1.08œ T=1.42œ T=4.22œ T=4.57œ T=4.97œ T=5.18œ T=2.54œ T=2.025œ T=1.22œ T=1.58œ T=1.79œ T=3.35œ T=2.18œ T=1.83œ T=1.69œ T=1.26œ T=1.58œ T=3.92œ T=1.76œ T=1.8œ T=1.55œ T=1.4œ T=1.67œ T=1.15œ T=1.47œ T=1.33œ T=1.83œ T=1.32œ T=0.66œ T=0.97œ T=1.31œ T=0.86œ

S=32.56 S=32.83 S=32.6 S=32.65 S=32.74 S=33.15 S=32.9 S=33.32 S=33.3 S=33.15 S=32.93 S=32.7 S=32.65 S=32.52 S=32.35 S=32.62 S=32.73 S=33.08 S=33.37 S=33.38 S=32.5 S=32.56 S=32.95 S=33.1 S=33.29 S=32.2 S=32.6 S=32.72 S=32.98 S=33.23 S=33.36 S=32.97 S=32.82 S=32.55 S=33.39 S=33.13 S=32.77 S=32.57 S=32.58 S=32.59


136 Haul ##
4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 . . . . . . . . . . . . . . . . .

V.I. Sokolov

16.07.96 16.07.96 17.07.96 18.07.96 18.07.96 18.07.96 19.07.96 19.07.96 20.07.96 20.07.96 20.07.96 20.07.96 21.07.96 21.07.96 21.07.96 22.07.96 23.07.96

Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth Depth

78 m 150 m 251 m 295 m 195 m 100 m 59 m 251 m 151 m 85 m 85 m 52 m 61 m 100 m 200 m 52 m 152 m 19 m 37 m 76 m 180 m 154 m 72 m 52 m 28 m 37 m 51 m 77 m 110 m 155 m 191 m 192 m 125 m 80 m 54 m 32 m 22 m 50 m 80 m 100 m 165 m 178 m 113 m 79 m 56 m 95 m 190 m 55 m 70 m 94 m 152 m 166 m 127 m 85 m 65 m 53 m

53œ15 53œ15 53œ15 53œ30 53œ30 53œ30 53œ30 53œ45 53œ45 53œ16 53œ45 53œ45 54œ00 54œ00 54œ00 53œ01 51œ45



N155œ26 N154œ51 N154œ30 N154œ24 N154œ34 N155œ10 N155œ32 N154œ23 N154œ43 N155œ16 N155œ16 N155œ31 N155œ22 N155œ01 N154œ28 N155œ43 N155œ56



E E E E E E E E E E E E E E E E E

T=0.34œ T=0.79œ T=1.59œ T=1.81œ T=1.28œ T=0.21œ T=0.54œ T=1.52œ T=1.06œ T=0.44œ T=0.44œ T=0.73œ T=0.74œ T=0.41œ T=1.31œ T=1.1œ T=1.73œ T=4.67œ T=4.67œ T=2.02œ T=1.3œ T=1.7œ T=2.97œ T=3.94œ T=4.67œ T=2.8œ T=2.15œ T=1.87œ T=1.27œ T=1.31œ T=1.5œ T=1.3œ T=1.39œ T=1.84œ T=1.83œ T=2.39œ T=4.72œ T=1.42œ T=1.47œ T=1.53œ T=1.23œ T=1.01œ T=0.43œ T=0.44œ T=0.75œ T=0.24œ T=1.18œ T=0.88œ T=0.53œ T=0.3œ T=1.03œ T=1.13œ T=0.44œ T=0.44œ T=0.67œ T=0.83œ

S=32.64 S=32.87 S=33.27 S=33.39 S=33.11 S=32.67 S=32.62 S=33.22 S=32.91 S=32.68 S=32.64 S=32.66 S=32.74 S=33.13 S=32.59 S=33.09 S=32.61 S=32.62 S=32.92 S=33.03 S=32.9 S=32.24 S=32.6 S=32.65 S=32.68 S=32.73 S=32.86 S=33.15 S=33.21 S=33.34 S=33.21 S=32.99 S=32.9 S=32.75 S=32.56 S=32.31 S=32.64 S=32.82 S=32.93 S=33.1 S=32.98 S=32.74 S=32.63 S=33.59 S=32.65 S=33.06 S=32.61 S=32.66 S=32.67 S=32.89 S=32.98 S=32.75 S=32.7 S=32.68 S=32.64

1 2 3 4 5 6 7 8 9 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 3 3 3 3 3 3 4 4 4 4 4 4

Dredge ##
. . . . . . . . . 0 1 2 3 4 5 6 7 8 9 0 3 4 5 6 7 8 9 3 4 5 7 8 9 0 1 2 3 4 5

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

01.07.96 01.07.96 01.07.96 01.07.96 03.07.96 03.07.96 03.07.96 03.07.96 07.07.96 07.07.96 07.07.96 07.07.96 07.07.96 07.07.96 09.07.96 09.07.96 09.07.96 09.07.96 09.07.96 09.07.96 14.07.96 14.07.96 14.07.96 14.07.96 17.07.96 17.07.96 17.07.96 17.07.96 17.07.96 18.07.96 19.07.96 19.07.96 19.07.96 19.07.96 21.07.96 21.07.96 21.07.96 22.07.96 22.07.96

51œ10N156œ41E 51œ092N156œ379E 51œ10N156œ26E 51œ103N156œ165E 51œ36N156œ095E 51œ36N156œ14E 51œ35N156œ18E 51œ35N156œ25E 52œ00N156œ20E 52œ00N156œ10E 52œ00N155œ51E 52œ00N155œ40E 52œ00N155œ378E 52œ00N155œ28E 52œ20N155œ04E 52œ20N155œ185E 52œ20N155œ345E 52œ20N155œ545E 52œ20N156œ13E 52œ20N156œ19E 52o45N155œ42E 52œ45N155œ23E 52œ45N155œ08E 52œ45N154œ57E 53œ10N154œ45E 53œ10N155œ10E 53œ10N155œ245E 53œ25N155œ458E 53œ25N155œ15E 53œ25N154œ38E 53œ35N155œ35E 53œ35N155œ25E 53œ35N155œ115E 53œ35N154œ436E 54œ00N154œ356E 54œ00N154œ52E 54œ00N155œ06E 54œ00N155œ20E 54œ00N155œ30E