Decapodid Stage of Neocrangon Communis (Decapoda

ZOOSYSTEMATICA ROSSICA, 25(1): 13–22 25 JUNE 2016

Decapodid stage of Neocrangon communis (Decapoda, Crangonidae) from the eastern part of the Sea of Okhotsk Личинка Neocrangon communis (Decapoda, Crangonidae) декаподитной стадии из восточной части Охотского моря

N.A. SEDOVA* & S.S. GRIGORIEV

Н.А. СЕДОВА, С.С. ГРИГОРЬЕВ

N.A. Sedova, Kamchatka Federal Technical University, Petropavlovsk-Kamchatsky, 683003, Russia. E-mail: [email protected] .S. Grigoriev, Kamchatka Division of Pacific Institute of Geography, Far East Branch, Russian Ac.Sc., Petropavlovsk-Kamchatsky, 683000, Russia. E-mail: [email protected]

For the first time the morphology of the decapodid stage of Neocrangon communis is described in detail. The decapodid can be distinguished from those of the genera Argis, Crangon, and Mesocrangon by the morphology of their telson, antennae, antennulae, and carapace. The main distinguishing features of the decapodid of N. communis were two spines on medial line of the carapace, a short rostrum, relatively wide scaphocerite, characteristic shape and length of the terminal setae on the telson. Drawings of general view and some limbs are presented. Впервые подробно описана личинка декаподитной стадии Neocrangon communis. Ли- чинка этого вида отличалась от соответствующей стадии представителей родов Argis, Crangon и Mesocrangon по морфологии тельсона, антенны, антеннулы и карапакса. Ос- новные признаки N. communis на стадии декаподит: два шипа на медианной линии ка- рапакса, короткий рострум и относительно широкий скафоцерит, характерная форма и длина апикальных щетинок на тельсоне. Приводятся рисунки общего вида и отдельных конечностей. Key words: shrimps, morphological features, development, description, rostrum, telson, carapace, Sea of Okhotsk, Decapoda, Crangonidae, Neocrangon communis Ключевые слова: креветки, морфологические признаки, развитие, описание, рострум, тельсон, карапакс, Охотское море, Decapoda, Crangonidae, Neocrangon communis

INTRODUCTION Representatives of about 20 species and eight genera of shrimps from the fam- Although the larvae of the family ily Crangonidae Haworth, 1825 inhabit the Crangonidae regularly occur in plankton marine waters surrounding the Kamchatka samples collected in the northwestern Pa- Peninsula. Among them, representatives of cific waters, larval stages only for 12 spe- two species of the genus Neocrangon, N. com- cies are described. It is probably because of munis and N. abyssorum occur in the eastern many crangonid species develop without part of the Okhotsk Sea (Sokolov, 2001). metamorphosis or with very short meta- Neocrangon communis Rathbun, 1899 morphosis (except a few genera that have is a Pacific boreal species, one of the most “normal” larval development, e.g. Crangon, common and widespread species of the fam- Philocheras etc.). ily Crangonidae. It occurs from 71°17´N (Chukchi Sea) (Makarov, 1941), through *Corresponding author the Bering Sea to San Diego in North

America, and to Peter the Great Bay and All adults of the genus Crangon have only the eastern coast of Honshu Island (Sea of one spine on the median line of carapace, Japan) (Sokolov, 2001). whereas all adults of the genus Mesocrangon Adults of this species occur from 116 to have two spines (Nizyaev et al., 2006). 1537 m of water depth (Komai & Komatsu, Neocrangon communis is a sublitoral- 2009; Butler, 1980). Based on the cited lit- bathyal species. The shrimp generally oc- erature, shrimps of this species were found curs on fine-grained substrate with an ad- in their area in depths be tween 37 and mixture of stones over shelf and upper part 295 m, (water temperature 0.21–2.8 °C and of continental slope (Vinogradov, 1950; salinity 32.66–33.39‰) (Sokolov, 2001). Butler, 1980). In the Okhotsk and Bering Females dominated the catches, but small- Seas the adults were distributed in water size females (CL < 9 mm) were scarce. The layers from 30 to 555 m over bottom depth, scarcity of small females suggests a possible in the Sea of Japan it occurred up to 1450 m, case of protandric hermaphroditism which near California it was caught at 1537 m of has been suggested for some crangonid spe- bottom depth. In the eastern Bering Sea, cies (Frechette et al., 1970; Boddeke et al. the species was obtained at depths of 62 to 1991; Sokolov, 2001). 95 m on mud and sand substrate with water In spite of the regular occurrence of lar- temperatures of 0.5–3.6 °C (Vinogradov, vae of N. communis in plankton samples, 1950; Butler, 1980). This species was caught this is one of the least studied species in the by trawl together with Pandalopsis dispar, northwestern part of the Pacific Ocean. The Pandalus jordani, and P. borealis at bottom descriptions of the larval stages zoea I were depth 128–160 m in Canada (Butler, 1980). given by Makarov (1966), Sedova & Grig- Females of carapace length 9.1–13.9 mm oriev (2015) based on planktonic samples with external eggs were caught in January– from the Sea of Okhotsk. Makarov (1966) February. Fertility of one female was about only provided a short description of the 2200 eggs, with a size of 0.9 × 0.75 mm (Sliz- decapodid of Neocrangon communis. Also kin, 2006). descriptions of the decapodid of a few spe- Larvae of N. communis can be found in cies from the genus Crangon Fabricius 1758, many plankton samples collected in Kam- Argis Kröyer 1842, and Sclerocrangon Sars, chatka waters over bottom depth close to 70 1883 occupying the eastern part of the Sea m from April to October. They often occur of Okhotsk are available (Birshteyn & Vino- during summer in waters over the Western gradov 1953; Kurata, 1964; Makarov, 1966, Kamchatka shelf, and sometimes they were 1968; Squires, 1965; Tesmer & Broad, 1964). caught along the south-eastern coast of Description of the decapodid of Mesocran- Kamchatka Peninsula (Sedova & Grigoriev, gon intermedia and detailed description of 2013). In Avacha Bay (the eastern coast of legs of I–V zoeal stages in N. communis and Kamchatka peninsula) and in the western M. intermedia were presented in our previ- Bering Sea they also occur regularly, but in ous studies (Sedova, 2013; Sedova & Grig- smaller abundances (Sedova & Andronov, oriev, 2014а,b; Sedova & Grigoriev, 2015). 2013; Sedova & Grigoriev, 2013). Maximal Adults of crangonids live burrowing into abundance of the larvae (402 specimens per the substrate. Characteristic protuberance 1 m2 of the sea surface) was found in 1999 in and spikes, as well as a special form of the the north of the western Kamchatka shelf rostrum only appear at postlarval or even (Sedova & Andronov, 2013; Sedova & Gri- juvenile stages (Makarov, 1966). Shrimps goriev, 2013). Late larvae were caught in of the genus Crangon live on soft substrates. fall between 54 and 55°N (Sedova, 2004). Shrimps of the genera Neocrangon and Me- Larval development of N. communis is socrangon live on fine-grained substrate of a normal pelagic nature, not shortened, with an admixture of stones (Slizkin, 2006). with five larval and one decapodid stages.

MATERIAL AND METHODS significantly shorter than the lateral ones (whereas in megalopa of A. dentatа and This study is based on plankton samples A. lar the medial, apical terminal spines are collected during research surveys, per- significantly longer). formed in the eastern part of the Sea of Ok- On the median line of the carapace of hotsk (over the western Kamchatka shelf). our decapodid, as in larvae of the species of Ichthyoplanktonic gear with a mouth di- the genus Argis, there were two spines. But ameter of 80 cm and mesh size 0.56 mm was in contrast, in our decapodid the spines are used. Vertical total haul from bottom to closer together, whereas in Argis the spines surface was carried out with bottom depth are widely separated. 500 m and less, and from 500 m to surface Our decapodid differ from the decapo- in smaller bottom depths. The planktonic did of the genus Crangon by having two samples were fixed in 4% formalin. One lar- spines on the median line of the carapace va of the decapodid in very good condition (versus in Crangon just a single one), and there was caught during one of the surveys by the longer antennal flagella. In addi- (2013.08.05) at a station with coordinates ´ ´ tion, the majority of Crangon species have 56°39 of north latitude and 155°46 of east a longer spine on the scaphocerite, extend- longitude over 25 m of bottom depth. ing beyond the distal edge. This spine in our Classification and morphology of larval decapodid was significantly shorter. stages was made following Tesmer & Broad Furthermore, there is a small exopo- (1964), and Wehrtmann (1991). dite on the first pair of pereiopods in most Measurements of the larva were made species of Crangon (Kurata, 1964; Squires, with an eyepiece micrometer with a preci- 1965; Tesmer & Broad, 1964; Li & Hong, sion of 0.1 mm as follows: carapace length 2003), which was absent in our larvae. (CL) from the anterior tip of the rostrum to Our decapodid differs from the species the posterior median carapace margin; to- M. intermedia by a shorter rostrum, the tal length (TL) from the anterior tip of the structure of the telson, the position of the rostrum to the telson back edge excluding teeth on the carapace, width of the scapho- telson setae. cerite, a smaller body length, and structure of antennula. The total length of the de- RESULTS AND DISCUSSION capodid of M. intermedia from waters near Identification Kamchatka Peninsula was 9.0 mm, whereas our decapodid was much smaller (6.5 mm There was no doubt that the analyzed ТL). Posterior spine on the median line of larva was of the family Crangonidae. Identi- the carapace in our specimen is located clos- fication of our specimen as N. communis was er to the front spine, unlike M. intermedia. based on the morphological features. In the The rostrum in the decapodid of M. inter- following features the present decapodid media reaches the distal edge of the eye, but differs from other species of the family. in our decapodid it was 1.5 times shorter Our decapodid identified as N. commu- (Sedova & Grigoriev, 2014a). Scaphocer- nis differs from the species of the genus Ar- ite of M. intermedia is much narrower. It is gis by body length and morphology of the covered with a series of short setae on the rostrum and telson. The present decapodid outer edge. In our megalopa the outer edge is significantly smaller (in 1.5–2 times) of scaphocerite was naked, and the scapho- than similar staged larvae of the genus Ar- cerite was much wider. gis. The rostrum is longer and straight (but Spine on scaphocerite in our decapodid not dorsally as in Argis). was much shorter than in M. intermedia The medial, apical terminal spines on and other crangonids. Exopodite of anten- the telson in the present megalopawere nula in M. intermedia consists of three seg-

© 2016 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 25(1): 13–22 16 N.A. SEDOVA & S.S. GRIGORIEV. NEOCRANGON COMMUNIS FROM SEA OF OKHOTSK ments, and endopodite of four segments. In oriev, 2014b; Sedova, 2015). The decapo- our decapodid endopodite and exopodite did of C. dalli have a similar body length to were two-segmented and relatively shorter. the last zoea stage (Makarov, 1966). The The telson in M. intermedia is narrower, but decapodid of N. communis, described by both pairs of terminal setae are very long. Makarov (1966), had total body length of Angle setae in M. intermedia are strongly re- approximately 7.0 mm. Larvae of N. commu- duced. Сentral setae in our decapodid were nis of stage V of zoea were caught between in two times shorter than adjacent setae, and 53° and 56°N along Kamchatka Peninsula, angled setae were longer than central ones. where the decapodid of this species were Flagellum in the decapodid of M. intermedia caught too. The structure of the rostrum, are two times longer than that of our de- the position of spines on the carapace, the capodid and consists of more segments. morphology of the antennae and the telson The only other species of the genus in larvae as described by Makarov (1966) Mesocrangon in our waters is M. volkii. corresponded to features of our specimen. Representatives of this species have a short Rostrum in adults of N. communis is tubercle on the carapace instead of the pos- about 20% of the length of the carapace (Fig. terior spine, although the larvae of M. volkii 1c) (Sokolov, 2001), and in our decapodid are unknown. The species was caught in the almost so (Fig. 1a). In adults of N. commu- Bering Sea, but in the Ochotsk Sea it does nis on the median line of the front half of the not occur. carapace there were two spines (Fig. 1c), Our decapodid differ from the species with the frontal spine being smaller. The of the genus Sclerocrangon by numbers spines were close together, like in our de- of spines and by carapace sculpturing. In capodid. This features distinguish the adults adults of the genus Sclerocrangon, three of N. communis from other crangonids. or four spines are present on the carapace, In our previous paper (Sedova & Grig- which is strongly sculptured. The decapo- oriev, 2015) we presented a detailed descrip- did of S. salebrosa have a very long and tion of the last zoea stage of N. communis. sharp rostrum and absent of spines on cara- In some of the larvae through the veils of pace (Makarov, 1968). The decapodid of the telson the next stage was seen (Fig. 1е). other species from this genus that could oc- Two shorter central spines and two pairs of cur in our waters have not been described. long terminal setae were well visible. Morphology of larvae as well as adults Comparing to other crangonids from of Paracrangon echinata are very different our waters the larvae of N. communis have a from other crangonids. Our decapodid dif- shorter spine on scaphocerite (Fig. 1b), and fer from the larvae of P. echinata by spines compared to larvae of M. intermedia they on carapace and teeth on the rostrum. In have a wider scaphocerite. adults of Paracrangon echinata, there are Other species from the genus Neocran- three large spines on the carapace, as well as gon dwell in deep water, so they cannot small and large dorsal teeth on the rostrum. occur in the planktonic samples, although The decapodid of Paracrangon echinata theirlarvae have not been described. Adults have not been described, but we can assume of the genus Neocrangon have one spine on that it will also be very different. carapace, but not two like in our larva. The morphological features to identify the decapodid as N. communis are are as fol- Description lows: Suitable sizes. It is known that the to- The total length of the decapodid was tal body length of larval stage V of zoea 6.5 mm. Eyes were rounded, close to each (last larval stage) of N. communis was 6.0– other. Rostrum slightly pointed, quite 7.5 mm (Makarov, 1966; Sedova & Grig- short, reaching only slightly beyond the

Fig. 1. Neocrangon communis: a, lateral view of body of decapodid; b, anterior part of body of zoea V; c, carapace of adult shrimp (redrawing fragments after Sokolov, 2001); d, telson of megalopa; e, telson of zoea V; f, rostrum of decapodid. Scale bars: 1 mm. middle of the eye (Fig. 1f). Median part dite of antennula two-segmented, with four of carapace raised into a small keel. In the long distal aesthetascs and one very small front part of the carapace there were two one (Fig. 2а). Endopodite two-segmented spines, anterior spine smaller and sharper, with one lateral and four short terminal se- located just behind the eyes, posterior tae. Flagellum of antenna consists of 16 seg- spine larger and located close to the frontal ments. Scaphocerite wide, distal tooth short, spine (Fig. 1а). At the base of the posterior not reaching distal edge of lamella (Fig. 2f). spine one short seta was present. Postorbit- There were a total of 24 setae on the scaph- al spines were small. Pleurae from abdomen ocerite. Basipodite of maxillula with six segments one to five of rounded. A small spines and two lateral setae (Fig. 2c). Coxal anal spine was present. endite of maxillula with three bristles. En- Base of antennula three-segmented, first dopodite with one short bristle. Exopodite segment with a series of short setae. Exopo- of maxilla with 27 bristles (Fig. 2b).

Fig. 2. Neocrangon communis, decapodid; a, antennula; b, maxilla; c, maxillula; d, maxilliped 1; e, maxilliped 2; f, antenna; g, maxilliped 3. Scale bar: 0.5 mm.

Fig. 3. Neocrangon communis, decapodid; a, distal segment of Р1; b, Р1 in total; c, d, second and third pereopods; е, fifth pleopod; f, g, four and fifth pereopods; h, second pleopod. Scale bar: 0.5 mm.

First maxilliped: basis with large epipo- Second maxilliped: basis with small dite; endopod unsegmented with three setae; epipodite. Endopodite four-segmented with exopod with two subterminal setae and four one, one, eight, seven long setae on each seg- terminal plumose natatory setae (Fig. 2d). ment; exopodite with five setae (Fig. 2е).

Maxillipeds were of a typical structure the decapodid of this species differs from (Fig. 2d, e, g). First maxilliped: basis with the corresponding stages of the genera Ar- large epipodite; endopod unsegmented with gis, Crangon, Mesocrangon by morphol- three setae; exopod with two subterminal ogy of the telson, antennae, antennulae and setae and four terminal plumose natatory carapace. The main distinguishing features setae (Fig. 2d). Second maxilliped: basis of the decapodid of N. communis were two with small epipodite. spines on the median line of the carapace, a Endopodite four-segmented with one, short rostrum, relatively wide scaphocerite, one, eight, seven long setae on each segment characteristic shape and length of the ter- correspondingly; exopodite with five setae minal setae on the telson. (Fig. 2е). In previous studies, we observed that lar- Third maxilliped: coxa with epipodite val development of M. intermedia is closed (Fig. 2g). Basipodite with five setae; en- to N. communis (Sedova, 2013; Sedova & dopod four-segmented with numerous Grigoriev, 2014b; 2015). Our decapodid is short setae placed as shown; exopod two- different from M. intermedia decapodid but segmented, with two subterminal and four is similar to the one described by Makarov terminal setae. (1966), although somewhat smaller. Ac- Pereopods are uniramous, consist of five cording to our data, the average body length to seven segments (Fig. 3a–d, f, g). Subchel- of the zoea V of N. communis was 7.2 mm. A la on the first pair of pereopods was well- reduction in the body length of the decapo- developed, and that is characteristic for all did stage can be due to the sharp shorten- representatives of the family. In the second ing of the rostrum. At the last stage of zoea, pair of pereopods the claw was very small, length of the rostrum was an average of 45% narrow. The gills are located at the base of of the length of the carapace, and on stage of pereopods. All five pairs of pleopods are decapodid, only 20% (Figs 1а–с). well developed, with long swimming setae, Our decapodid at this stage more differs and one seta on one segmented flagellum from the species of the genus Crangon than (Fig. 3e, h). from the species of the genus Mesocrangon. Telson was almost rectangular, with Nevertheless, our decapodid differs from two pairs of lateral spines. There are three the species of the genus Argis less than that pairs of long setae on the end of the telson. of Crangon, but more than from Mesocran- Medial setae were shorter, on average al- gon. most twice as short. Endopodite of uropods slightly longer then exopodite. Uropods ACKNOWLEDGEMENTS shorter then telson. Spine on uropods was We thank the crews of the research vessels short (Fig. 1d). as well as the scientific staff of the Kamchatka A few larvae with a total body length Institute of Fisheries and Oceanography for col- 6.5–7.0 mm similar that the one described lection of plankton samples and presenting them herein, were caught over the western Kam- to us for study. chatka shelf in 1999 and 2003 (1999.07.26, latitude 53°N bottom depth 25 m; and REFERENCES 2003.09.27, latitude 53°20´N, bottom depth about 30 m), but here these larvae are not Birstein J.A. & Vinogradow L.G. 1953. New data on the fauna of decapod Crustacea of the Ber- considered. ing Sea. Zoologicheskiy Zhurnal, 32(3): 215– 228. (In Russian, with English summary). CONCLUSION Boddeke R., Bosschieter J.R. & Goudswaard P.C. 1991. Sex change, mating and sperm Detailed studies of a caught larva iden- transfer in Crangon crangon (L.). In: Bauer tified as Neocrangon communis showed that R.T. & Martin J.W. (Eds). Crustacean Sex-

ual Biology: 181–182. New York: Columbia larvae over the western Kamchatka shelf University Press. in 1999 and 2001]. Problems of Fisheries, 5, Butler T.H. 1980. Shrimps of the pacific coast 2(18): 193–205. (In Russian). of Canada. Canadian bulletin of fisheries and Sedova N.A. 2013. Morphological character- aquatic sciences, 202: 1–280. istics of Neocrangon communis and Meso- Frechette J., Corrivault G.W. & Couture R. crangon intermedia (Decapoda, Crango- 1970. Hermaphroditisme proterandrique nidae) from the northwestern Pacific. In: chez une crevette de la familie des crango- Chereshnev I.A. (Ed.). Chteniya pamyati nides, Argis dentata Rathbun. Naturaliste ca- akademika K.V. Simakova. Tezisy dokla- nadien, 97(6): 805–822. dov Vserossiiskoy nauchnoy konferentsii Komai T. & Komatsu H. 2009. Deep-sea (Magadan, 26–28 noyabrya 2013 g.) [Read- Shrimps and Lobsters (Crustacea: Decapo- ings in memory of academician K.V. Sima- da) from Northern Japan, Collected during kov: Materials of All-Russian scientific con- the Project “Research on Deep-sea Fauna ference (Magadan, 26–28 November 2013)]: and Pollutants off Pacific Coast of Northern 169–170. Magadan. (In Russian). Japan”. In: Fujita T. (Ed.). Deep-sea Fauna Sedova N.A. & Andronov P.U. 2013. Qualita- and Pollutants off Pacific Coast of Northern tive composition and horizontal distribution Japan. National museum of Nature and Sci- of larvae of shrimps in the northwestern Ber- ence Monographs, Tokyo, 39: 495–580. ing Sea. Bulletin of the North-East Scientific Kurata H. 1964. Larvae of decapod crustacea Center, Russia Academy of Sciences Far East of Hokkaido. 4. Crangonidae and Glypho- Branch, 1: 30–38. (In Russian). crangonidae. Bulletin of Hokkaido Regional Sedova N.A. & Grigoriev S.S. 2013. Distri- Fisheries Research Laboratory, 28: 35–50. (In bution of shrimp larvae near south-eastern Japanese). coast of Kamchatka during spring 2009. Bul- Li H.Y. & Hong S.Y. 2003. Larval development letin of the North-East Scientific Center, Rus- Crangon hakodatei Rathbun (Decapoda: sia Academy of Sciences Far East Branch, 3: Crangonidae) reared in the laboratory. Ox- 77–86. (In Russian). ford Journals Life Sciences Journal of Plank- Sedova N.A. & Grigoriev S.S. 2014a. Mega- ton Research, 25: 1367–1381. lopa of Mesocrangon intermedia (Decapoda, Makarov R.R. 1941. The Decapod Crustacea of Crangonidae) from the eastern part of the the Bering and the Chukchees Seas. Issledo- sea of Okhotsk. Zoosystematica Rossica, vanija dalnevostochnikh morei SSSR, 1: 111– 23(2): 189–197. 163 (In Russian with English summary). Sedova N.A. & Grigoriev S.S. 2014b. System- Makarov R.R. 1966. Lichinki krevetok, rakov atic Position of Neocrangon communis (De- otshelnikov i krabov Zapadno-Kamchatskogo capoda, Crangonidae) Based on the Features shel’fa i ikh raspredelenie [Larvae of shrimps, of Larval Morphology. Zootaxa, 3827(4): hermit crabs and crabs from western Kam- 559–575. chatka shelf and their distribution]. Moscow: Sedova N.A. & Grigoriev S.S. 2015. Features Nauka. 164 p. (In Russian). of larval morphology of Mesocrangon inter- Makarov R.R. 1968. On the Larval Develop- media and Neocrangon communis (Decapoda, ment of the Genus Sclerocrangon G. O. Sars Crangonidae) from the Northwestern Pacif- (Caridea, Crangonidae). Crustaceana. Sup- ic. Zoologicheskiy Zhurnal, 4(3): 215–228. (In plement 2, Studies on Decapod Larval De- Russian, with English summary). velopment: 27–37. Slizkin A.G. 2006. Atlas-opredelitel krabov i kre- Nizyaev S.А., Bukin S.D. & Klitin А. К. 2006. vetok dalnevostochnykh morey [Atlas-guide Rukovodstvo po isucheniyu promyslovykh ra- of crabs and shrimps of far East seas of Rus- koobrasnykh dalnevostochnykh morey [Manual sia] Vladivostok: TINRO-centre. 216 p. (In for studying on commercial crustaceans of the Russian). Russian Far-Eastern seas]. Yuzhno-Sakha- Sokolov V.I. 2001. Decapod Crustaceans of the linsk: Sakhalin Research Institute of Fisheries Southwest Kamchatka Shelf: R/V “Professor and Oceanography. 114 p. (In Russian). Levanidov” collection in June 1996. Arthropo- Sedova N.A. 2004. Raspredeleniye lichinok kre- da Selecta, 10(2): 103–136. vetok v raione Zapadno-Kamchatskogo shel’fa Squires H. J. 1965. Larvae and megalopa of Ar- v 1999 i 2001 godu [Distribution of shrimp gis dentata (Crustacea: Decapoda) from Un-

gava Bay. Journal Fisheries Research Board of TINRO, Vladivostok, 33: 180–356. (In Rus- Canada, 22(1): 69–82. sian). Tesmer C.A. & Broad A.C. 1964. The larval de- Wehrtmann I.S. 1991. How important are star- velopment of Crangon septemspinosa (Say) vation periods in early larval development (Crustacea: Decapoda). The Ohio Journal of for survival of Crangon septemspinosa? Ma- Science, 64(4): 239–250. rine Ecology Progress Series, 73: 183–190. Vinogradov L.G. 1950. Key to the shrimps, crayfishes and crabs of the Far East. Izvestia Received 5 October 2015 / Accepted 29 March 2016 Scientific editor: M.E. Daneliya