Illustrated Key for the Identification of Brachyuran Zoeal Stages

Journal of the Marine Biological Association of the United Kingdom, 2009, 89(2), 379–386. #2008 Marine Biological Association of the United Kingdom doi:10.1017/S0025315408002762 Printed in the United Kingdom Illustrated key for the identiﬁcation of brachyuran zoeal stages (Crustacea: Decapoda) in the plankton of Peter the Great Bay (Sea of Japan) elena s. kornienko and olga m. korn Institute of Marine Biology, Far East Branch, Russian Academy of Sciences, 17 Pal’chevskogo Street, 690041 Vladivostok, Russia

A dichotomous identification key for brachyuran zoeal stages from Peter the Great Bay (Russian waters of the Sea of Japan) is provided for the first time. The key covers 16 taxa identified to species level and uses only the most conspicuous external characters of larvae that are easy to observe under a stereomicroscope without specimens dissection. The key is based on the accounts by various authors and new original descriptions of larvae obtained both from plankton samples and from laboratory culture. Brief descriptions of larvae of 16 brachyuran species are also included.

Keywords: zoea, Brachyura, crab, identiﬁcation key, plankton, Sea of Japan

Submitted 15 May 2008; accepted 4 August 2008; ﬁrst published online 16 October 2008

INTRODUCTION Zooplankton was sampled in Vostok Bay (inner bay of Peter the Great Bay, Sea of Japan) between May and Larval development is one of the most important periods of November 2002 using a Norpac net with a ring diameter of the decapod life cycle; recruitment does not occur unless the 40 cm and a filtering cone made of a 168 mm mesh, and in larval period is completed. Larval data can be useful in evalu- Amursky and Ussurijsky Bays (inner bays of Peter the Great ating species diversity in a region and in specifying the repro- Bay, Sea of Japan) between April and October 2007 using a duction time of brachyuran species. Morphological features of Juday net with a ring diameter of 38 cm and a filtering cone larvae are complementary characters for crustacean taxonomy made of a 168 mm mesh (Figure 1). and phylogeny. The study of larvae is important for the It is very difficult to identify the larvae (especially the larvae problem of introduced species arriving in ballast waters or of congeneric species) using drawings and descriptions of on fouled ships. However, insufficient attention has been different authors in more or less detail. So, zoeae of most con- paid to decapod larval development in the Russian Far East sidered species were obtained under the laboratory conditions. Seas. Many keys exist for the identification of brachyuran This material was used for the construction of the key and the larvae in different regions of the World Ocean (Ingle, 1992; original figures. Paula, 1996; Baéz, 1997; Pessani et al., 1998; Anosov, 2000; Ovigerous crab females were maintained in an aerated sea- Puls, 2001; Santos & Gonza´lez-Gordillo, 2004; Rice & water aquarium until larvae hatched. After hatching, larvae Tsukimura, 2007), but only one taxonomic guide is known were concentrated at the edge of the aquarium using a point-light for the Sea of Japan (Konishi, 1997). There is no comprehen- source and transferred to 1-l glass vessels with filtered and sive key for identification of the brachyuran larvae in Russian UV-sterilized seawater and reared to the megalopal stage. The waters of the Sea of Japan; however, the species list of this density of larvae was about 100 specimens l21. The water in region considerably differs from that of coastal waters of the vessels was changed daily. The larvae were fed with newly Japan. The aim of this work is to provide a key for the identi- hatched nauplii of Artemia salina. Very small zoea of varunid fication of brachyuran larvae in Peter the Great Bay. and pinnotherid crabs were reared using nauplii of the rhizoce- phalan crustacean, Polyascus polygenea,asafood.Earlier,this method was described in detail (Kornienko & Korn, 2005a). MATERIALS AND METHODS All larvae were fixed in 4% formaldehyde for light micro- scopic studies. The key is based on the accounts previously published by The dichotomous identification key is based mainly on various authors (Table 1) and new original descriptions of external morphological characters, which are easy to observe brachyuran larvae both taken from plankton samples and under a stereomicroscope MBS-10 without specimen dissec- reared from ovigerous crab females in the laboratory. tion. When these features are insufficient, the morphology and setation of appendages has been included. Moreover, new original figures were provided to make identification easier. Corresponding author: E.S. Kornienko The outlines of the larvae were drawn using a camera lucida Email: [email protected] attached to a binocular Ergaval microscope (Carl Zeiss Jena).

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Table 1. List of species and sources of descriptions of brachyuran zoeae included in the identiﬁcation key.

Family Species Authors

Dorippidae Paradorippe granulata (De Haan, 1839) Kurata, 1964; Terada, 1981; Quintana, 1987 Epialtidae Pugettia quadridens (De Haan, 1837) Kurata, 1969; Ko, 1998: Kornienko & Korn, 2004 Oregonidae Chionoecetes opilio (O. Fabricius, 1788) Kurata, 1963b; Motoh, 1973; Haynes, 1973, 1981 Pisidae Pisoides bidentatus (A. Milne-Edwards, 1873) Kurata, 1969; Kornienko & Korn, 2007 Cancridae Cancer amphioetus Rathbun, 1989 Iwata & Konishi, 1981 Cheiragonidae Erimacrus isenbeckii (Brandt, 1848) Aikawa, 1937; Kurata, 1963a; Makarov, 1966 Telmessus cheiragonus (Tilesius, 1812) Kurata, 1963a Portunidae Charybdis japonica A. Milne-Edwards, 1861 Yatsuzuka et al., 1984 Pinnotheridae Pinnixa rathbuni Sakai, 1934 Sekiguchi, 1978; Konishi, 1983; Kornienko & Korn, 2005b Tritodynamia rathbuni Shen, 1932 Matsuo, 1998 Pinnaxodes mutuensis Sakai, 1939 Konishi, 1981b Sakaina yokoyai (Glassell, 1933) Not described Varunidae Eriocheir japonicus De Haan, 1835 Morita, 1974; Kim & Hwang, 1990 Hemigrapsus sanguineus (De Haan, 1835) Hwang et al., 1993; Kornienko et al., 2008 H. penicillatus (De Haan, 1835) Hwang & Kim, 1995; Kornienko et al., 2008 H. longitarsis (Miers, 1879) Park & Ko, 2002; Kornienko et al., 2008

The key was constructed for zoea I, but the characters used 1b. Carapace with rostral spine only (Figure 3B) or with in the key do not change or change slightly through the suc- rostral and dorsal spines (Figure 3C), lateral spines cessive zoeal stages, with the exception of peculiar cases. absent ...... 11 Main characters of brachyuran zoea used for the identification 2a. Abdominal somites 3–5 with well developed posterolat- are represented in Figure 2. eral spines; furcal rami with lateral and dorsal spines (Figure 4A) ...... 3 2b. Abdominal somites 3–5 with poorly developed posterolat- RESULTS eral spines (Figure 4B, C); furcal rami without lateral and dorsal spines (Figure 4B) or with lateral spines only (Figure4C)...... 7 Key for the identification of brachyuran zoea I 3a. Furcal rami longer than proximal part of telson; dorsal (interspecific distinctions) and lateral furcal spines short (Figure 5A)...... 4 3b. Furcal rami shorter than proximal part of telson; lateral 1a. Carapace with rostral, dorsal and lateral spines furcal spines long (Figure 5B, C) ...... 6 (Figure 3A) ...... 2 4a. All carapace spines spinulated; dorsal spine straight; posterolateral spines longer than half abdominal somite;eachfurcalramuswiththreespines...... Chionoecetes opilio (Figure 11A)

Fig. 1. Map showing the sampling area. Fig. 2. Main characters used for the identiﬁcation of brachyuran zoea.

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Fig. 5. Telson of zoea in Chionoecetes opilio (A), Telmessus cheiragonus (B) and Erimacrus isenbeckii (C).

4b. All carapace spines smooth; dorsal spine slightly curved; posterolateral spines shorter than half abdominal somite; each furcal ramus with two spines ...... 5 5a. Abdominal somite 2 with a pair of lateral knobs; out- ermost pair of inner setae on posterior telson margin dentated on inside surface (Figure 6A, A0); posterolateral spines very short in zoea I and considerably lengthen in successive stages...... Fig. 3. Carapace of zoea in Eriocheir japonicus (A), Sakaina yokojai (B) and ...... Cancer amphioetus (Figure 11B) Pisoides bidentatus (C).

[Antenna biramous; protopod with two rows of [Lateral spines long, slightly shorter dorsal and spinules, nearly half of rostral spine; exopod with rostral spines. Antenna biramous; protopod spinulated, approximately equal rostral spine; endopod with three terminal setae of unequal length. Lateral knobs on abdominal somites 2–3 long, those of third one reach end of the same somite. Larvae found from April to July.]

Fig. 4. Abdomen and telson of zoea in Chionoecetes opilio (A), Tritodynamia Fig. 6. Abdomen and telson of zoea in Cancer amphioetus (A, A0) and rathbuni (B) and Pisoides bidentatus (C). Charybdis japonica (B) (after Yatsuzuka et al., 1984).

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two terminal setae of unequal length. Larvae found from June to September.] 5b. Abdominal somites 2 and 3 with a pair of lateral knobs; all inner setae on posterior telson margin plumodenticulate; posterolateral spines prominent in all zoeal stages (Figure6B)...... Charybdis japonica (Figure 11C) [Antenna biramous; protopod with two rows of spinules, nearly half of rostral spine; exopod with two terminal setae of unequal length. Larvae not found yet.] 6a. Abdominal somite 2 with a pair of lateral knobs; posterolateral spines of somite 4 shorter somite 5; lateral telsonal spine shorter than half furcal ramus; each furcal ramus with three inner setae (Figure 5B) ...... Telmessus cheiragonus (Figure 11D) [Dorsal spine straight; rostral and dorsal spines spinulated in distal part. Antennal protopod spinulated, shorter than rostral spine; exopod with two nearly equal setae. Abdominal somites 3–5 Fig. 8. Antenna and carapace of zoea in Tritodynamia rathbuni (A) and with long posterolateral spines. Each furcal Eriocheir japonicus (B). ramus with three spines: one long lateral spine and two short dorsal spines. Larvae found in April and May.] 7b. Lateral carapace spines arranged more ventrally than in 6b. Abdominal somites 2 and 3 with a pair of lateral knobs; typical brachyuran zoea; abdominal somites 4–5 not posterolateral spines of somite 4 shorter than somite 5; cylindrical but laterally expanded; telson subrectangular; lateral telsonal spine longer than half furcal ramus; each furcal rami without spines or with minute lateral spines furcal ramus with four inner setae (Figure 5C) ...... (Figure 7D–F)...... 10 ...... Erimacrus isenbeckii (Figure 11E) 8a. Dorsal and rostral spines longer than carapace; dorsal carapace spine straight; antenna uniramous (Figure 8A) . . . [Dorsal spine straight; rostral and dorsal spines ...... Tritodynamia rathbuni (Figure 11F) spinulated in distal part. Antennal protopod spinulated, shorter than rostral spine; exopod with two nearly equal setae. Abdominal somites 3–5 [Antennal protopod with two rows of spinules and with long posterolateral spines. Each furcal with short seta near base. Posterolateral spines on ramus with three spines: long lateral spine and abdominal somites 3–5 highly reduced. Larvae two short dorsal spines. Larvae found in April found from June to September.] and May.] 7a. Lateral carapace spines arranged as in typical brachyuran 8b. Dorsal and rostral spines slightly shorter than carapace; larvae; all abdominal somites cylindrical; telson triangular; dorsal carapace spine slightly curved; antenna biramous furcal rami without spines (Figure 7A, B) ...... 8 (Figure 8B) ...... 9

Fig. 7. Carapace of zoea in Tritodynamia rathbuni (A) and Pinnaxodes mutuensis (C); telson of zoea in Tritodynamia rathbuni (B); Pinnaxodes Fig. 9. Antenna, antennule and abdomen of zoea in Eriocheir japonicus (A) mutuensis (D), Pinnixa rathbuni (E) and Sakaina yokoyai (F). and Hemigrapsus sanguineus (B).

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9a. Abdominal somites 2–4 with a pair of lateral knobs [Rostral spine nearly one-third antenna; dorsal (lateral knobs on somite 4 disappear in successive spine with sparse blunt spinules. Antenna bira- stages); antennule with 2 aesthetascs; antennal exopod mous; protopod and exopod sub-equal. nearly half protopod (Figure 9A)...... Abdominal somites 2 with a pair of lateral ...... Eriocheir japonicus (Figure 11G)

[Antennal exopod as tapering spine with two small unequal subterminal setae. Larvae found from June to September.]

9b. Abdominal somites 2–3 with a pair of lateral knobs; antennule with 3 aesthetascs; antennal exopod nearly 2/3 protopod (Figure 9B)...... Hemigrapsus sanguineus, H. penicillatus, H. longitarsis (Figure 11H)

[Antennal exopod as tapering spine with small unequal subterminal setae. Larvae found from June to September.]

10a. Abdominal somites 4–5 expanded; furcal rami shorter than proximal part of telson, with minute lateral spines (Figure7D)...... Pinnaxodes mutuensis (Figure 11I)

[Antenna uniramous; protopod with two rows of spinules. Lateral telsonal margins slightly convex. Larvae found in July.]

10b. Abdominal somite 5 as a horseshoe; furcal rami longer than proximal part of telson, without lateral spines...... Pinnixa rathbuni (Figure 11J)

[Rostral and dorsal carapace spines straight. Antenna uniramous; protopod with two rows of spinules and with short seta near base. Furcal rami without spines (Figure 7E). Larvae found from May to November.]

11a. Carapace with rostral spine only ...... Sakaina yokoyai (Figure 11K)

[Antenna uniramous; protopod with two rows of spinules. Abdominal somites 4–5 as a horseshoe. Telson nearly rectangular; furcal rami shorter than proximal part of telson, without spines; median notch virtually absent (Figure 7F). Larvae found in June–August.]

11b. Carapace with rostral and dorsal spines (Figure 10) ...... 12 12a. Rostral and dorsal spines three times longer than carapace; dorsal spine straight ...... Paradorippe granulata (Figure 11L)

[Dorsal spine with blunt spinules, rostral spine with acute spinules. Antenna biramous; protopod and exopod sub-equal. Abdominal somite 2 with pair of lateral knobs. Telson narrow and long, with a pair of lateral spines; furcal rami twice exceeding proximal part of telson. Posterior telsonal margin with two inner setae only (Figure 10A, A0). Larvae found in July and August.]

12b. Rostral and dorsal spine shorter than carapace, dorsal spine slightly curved (Figure 10B) ...... Pugettia Fig. 10. Carapace (A) and telson (A0) of zoea in Paradorippe granulata; quadridens, Pisoides bidentatus (Figure 11M) carapace of zoea in Pugettia quadridens (B).

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Fig. 11. Lateral view of zoea in Chionoecetes opilio (A), Cancer amphioetus (B), Charybdis japonica (C), Telmessus cheiragonus (D), Erimacrus isenbeckii (E), Tritodynamia rathbuni (F), Eriocheir japonicus (G), Hemigrapsus sanguineus (H), Pinnaxodes mutuensis (I), Pinnixa rathbuni (J), Sakaina yokoyai (K), Paradorippe granulata (L), and Pisoides bidentatus (M).

knobs. Furcal rami with lateral spines (Figure 4C). posterior telsonal margin with two setae only in all zoeal Larvae found from June to September.] stages). Zoea II. Eyes stalked; exopod of maxilliped each with 6 nata- Most Brachyura from Peter the Great Bay pass through ﬁve tory setae; in most species abdomen consists of 5 somites and zoeal stages. Majiod crabs (Pisoides bidentatus, Pugettia quad- telson; pleopod buds absent; posterior telsonal margin with þ ridens, and Chionoecetes opilio) have an abbreviated develop- 3 3setae.InPisoides bidentatus, Pugettia quadridens,and ment including only two zoeal stages. Two of three Chionoecetes opilio, zoea II is the last stage; hence, the sixth pinnotherid crabs also undergo an abbreviated development: somite is delineated, somites 2–6 with biramous pleopod Sakaina yokoyai passes through probably three zoeal stages; buds. In Sakaina yokoyai, pleopod buds uniramous. In Pinnaxodes mutuensis, through four zoeal stages. Four zoeal Charybdis japonica and Chionoecetes opilio, posterior telsonal þ stages are known also for Paradorippe granulata. margin with 4 4 setae. Different zoeal stages of brachyuran crabs (age distinc- Zoea III. Exopod of maxilliped each with 8 natatory setae; tions) are easily determined using the number of natatory the sixth somite delineated, with the exception of Pinnixa setae on the exopods of maxillipeds, the number of setae rathbuni which has 5 somites in all zoeal stages. In Cancer along the posterior telsonal margin, the presence of unira- amphioetus, Trithodynamia rathbuni, Erimacrus isenbeckii, mous or biramous pleopod buds and some other morphologi- Telmessus cheiragonus, Eriocheir japonicus and Hemigrapsus þ cal features. The same stages in the species with abbreviated species, posterior telsonal margin with 4 4 setae; in development are more advanced in the number of characters Charybdis japonica and Pinnaxodes mutuensis, with additional over those of the species with longer development. small unpaired setae. In Sakaina yokoyai, pleopod buds biramous. Zoea IV. Exopod of maxilliped each with 10 natatory setae; Identiﬁcation of different zoeal stages of somites 2–6 with uniramous pleopod buds. In Cancer brachyuran crabs (age distinctions) amphioetus and Tritodynamia rathbuni, posterior telsonal margin with 5 þ 5 setae. Zoea I. Eyes sessile; exopod of maxilliped each with 4 natatory Zoea V. Exopod of maxilliped each with 12 natatory setae; abdomen consists of 5 somites and telson; posterior setae; pleopod buds biramous. In Eriocheir japonicus and telsonal margin with 3 þ 3 setae (in Paradorippe granulata, Hemigrapsus species, posterior telsonal margin with 5 þ 5setae.

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DISCUSSION REFERENCES

According to Adrianov & Kussakin (1998), 20 brachyuran Adrianov A.V. and Kussakin O.G. (1998) A check-list of biota of the Peter species belonging to 8 families and 16 genera inhabit Peter the Great Bay, the Sea of Japan. Vladivostok: Dalnauka. the Great Bay (Russian waters of the Sea of Japan). We Aikawa H. (1937) Further notes on brachyuran larvae. Records of found larvae of only 16 species from 9 families and 14 Oceanographic Works in Japan 9, 87–162. genera occurring in the plankton of Peter the Great Bay Anosov S.E. (2000) Keys to the identification of brachyuran larvae of the (Table 1). To date, we have not found both adults and Black Sea. Crustaceana 73, 1239–1246. larvae of Goetice depressus (de Haan, 1835), Helice tridens de Haan, 1833, and Paradromia japonica (Henderson, Baéz P. (1997) Key to the families of decapod crustacean larvae collected 1888). According to Vassilenko (1990), adult specimens of off northern Chile during an EI Ninõ event. Investigaciones Marinas Pinnixa tumida Stimpson, 1858 are found only in Possyet Valparaı´so 25, 167–176. Bay (eastern Peter the Great Bay). Brachyuran larvae occur Haynes E. (1973) Description of prezoeae and stage I zoeae of in Peter the Great Bay from April to November. Chionoecetes bairdi and C. opilio (Oxyrhyncha, Oregoninae). Fishery Zoea of each species represented in this key has been Bulletin 71, 769–775. previously described. Only the larvae of Sakaina yokoyai Haynes E. (1981) Description of stage II zoeae of snow crab, Chionoecetes are not known yet; however, the morphology of closely bairdi, (Oxyrhyncha, Majidae) from plankton of Lower Cook Inlet, related species, S. japonica, has been described (Konishi, Alaska. Fishery Bulletin 79, 177–182. 1981a). Both adults and larvae of S. yokoyai are rarely Hwang S.G. and Kim C.H. (1995) Zoeal stages and megalopa of found in Peter the Great Bay, so we failed to obtain the Hemigrapsus penicillatus (De Haan, 1835) (Decapoda, Brachyura, total series of their larval stages. Nevertheless, we believe Grapsidae) reared in the laboratory. Korean Journal of Systematic that S. yokoyai, like S. japonica, passes through three zoeal Zoology 11, 389–409. stages. Hwang S.G., Lee C. and Kim C.H. (1993) Complete larval development Frequently, the distinction between larvae of congeneric of Hemigrapsus sanguineus (Decapoda, Brachyura, Grapsidae) reared species is based on slight differences. The larvae of congene- in laboratory. Korean Journal of Systematic Zoology 9, 69–86. ric species, described on the basis of specimens hatched in the laboratory from ovigerous females, can be identified Ingle R.W. (1992) Larval stages of northeastern Atlantic crabs. An illustrated key. London: Chapman & Hall. only to the generic level when collected in nature. In our case, the specific identification of three Hemigrapsus Iwata F. and Konishi K. (1981) Larval development in laboratory of species (H. sanguineus, H. penicillatus and H. longitarsis)is Cancer amphioetus Rathbun, in comparison with those of seven very difficult. Zoea I and zoea II of these species are nearly other species of Cancer (Decapoda, Brachyura). Publications of the identical. The larvae are distinguishable only from zoea III. Seto Marine Biological Laboratory 26, 369–391. Zoea III–V of Hemigrapsus species differ in the number of Kim C.H. and Hwang S.G. (1990) The complete larval development of dorsomedial setae on the abdominal somite I and in the Eriocheir japonicus De Haan (Crustacea, Brachyura, Grapsidae) number of setae on the posterodorsal arch. It is pertinent reared in the laboratory. Korean Journal of Zoology 33, 411–427. to note that among the three Hemigrapsus species, zoea Ko H.S. (1998) Zoeal development of three species of Pugettia (Decapoda: and megalopa of H. penicillatus differ greatly (Kornienko Majidae), with a key to the known zoeas of the subfamily Epialtinae. et al., 2008). Despite the great similarity of larvae in Journal of Crustacean Biology 18, 499–510. varunid crabs of the genera Hemigrapsus and Eriocheir, the Konishi K. (1981a) A description of laboratory-reared larvae of the pin- latter possess a number of distinctive features in all develop- notherid crab Sakaina japonica Sere`ne (Decapoda, Brachyura). mental stages (Kornienko & Korn, 2005a; Kornienko et al., Journal of the Faculty of Science Hokkaido University Series VI. 2008). Zoology 22, 165–176. It was recently shown that zoea of Pisoides bidentatus and Konishi K. (1981b) A description of laboratory-reared larvae of the com- Pugettia quadridens belonging to different families are also mensal crab Pinnaxodes mutuensis Sakai (Decapoda, Brachyura) from nearly identical, with the exception of a more intensive Hokkaido, Japan. Annotations Zoologicae Japonensis 54, 213–229. coloration of the latter. Some differences appear only in the megalopal stage. Based on the larval similarity Konishi K. (1983) Larvae of the pinnotherid crabs (Crustacea, Brachyura) (Kornienko & Korn, 2007) and high genetic identity of found in the plankton of Oshoro Bay, Hokkaido. Journal of the Faculty of Science Hokkaido University Series VI. Zoology 23, 266–295. adults (Zaslavskaya et al., 2007) these two species should be assigned to one genus. Konishi K. (1997) Phylum Arthropoda. Class Malacostraca. Order The larvae of the other ten species belonging to different Decapoda. In Chihara M. and Murano M. (eds) An illustrated guide families and genera are easy to identify in the plankton. The to marine plankton in Japan. Tokyo: Tokai University Press, pp. features used in the key were chosen to enable identification 1439–1519. of zoea to the species level and do not reflect any systematic Kornienko E.S. and Korn O.M. (2004) Morphological features of the arrangement of decapod families. larvae of spider crab Pugettia quadridens (Decapoda: Majidae) from the northwestern Sea of Japan. Russian Journal of Marine Biology 30, 402–413. ACKNOWLEDGEMENTS Kornienko E.S. and Korn O.M. (2005a) Rearing under laboratory conditions and morphological peculiarities of the larvae of Japan mitten The project was supported by the Far East Branch of the crab Eriocheir japonicus (De Haan). Izvestiya TINRO 143, 35–51. Russian Academy of Sciences (grant no. 06-III-A-06-164) Kornienko E.S. and Korn O.M. (2005b) Morphological peculiarities of and by the Russian Foundation for Fundamental Researches larvae in Pinnixa rathbuni (Decapoda, Pinnotheridae) from the (grant no. 08-04-00929). Vostok Bay (the Sea of Japan). Zoologicheskii Zhurnal 84, 778–794.

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Kornienko E.S. and Korn O.M. (2007) The larvae of the spider crab Puls A.L. (2001) Arthropoda: Decapoda. In Shanks A.L. (ed.) A guide to Pisoides bidentatus (A. Milne-Edwards, 1873) (Decapoda: Majoidea: the identification of the larval invertebrates of the Pacific Northwest. Pisidae) reared under laboratory conditions. Journal of Plankton Oregon: Oregon State University Press, pp. 179–250. Research 29, 605–617. Quintana R. (1987) Later zoeal and early postlarval stages of three dorip- Kornienko E.S., Korn O.M. and Kashenko S.D. (2008) Comparative pid species from Japan (Brachyura: Dorippidae: Dorippinae). morphology of larvae of coastal crabs (Crustacea: Decapoda: Publications of the Seto Marine Biological Laboratory 32, 233–274. Varunidae). Russian Journal of Marine Biology 34, 77–93. Rice A. and Tsukimura B. (2007) A key to the identification of bra- Kurata H. (1963a) Larvae of Decapoda Crustacea of Hokkaido. 1. chyuran zoeae of the San Francisco Bay estuary. Journal of Atelecyclidae (Atelecyclinae). Bulletin of the Hokkaido Region Crustacean Biology 27, 74–79. Fisheries Research Laboratory 27, 13–24. Kurata H. (1963b) Larvae of Decapoda Crustacea of Hokkaido. 2. Majidae dos Santos A. and Gonza´lez-Gordillo J.I. (2004) Illustrated key for the (Pisinae). Bulletin of the Hokkaido Region Fisheries Research identification of the Pleocyemata (Crustacea: Decapoda) zoeal stages, Laboratory 27, 25–31. from the coastal region of south-western Europe. Journal of the Marine Biological Association of the United Kingdom 84, 205–227. Kurata H. (1964) Larvae of decapod Crustacea of Hokkaido 8. Dorippidae (Brachyura). Bulletin of the Hokkaido Region Fisheries Research Sekiguchi H. (1978) Larvae of pinnotherid crab, Pinnixa rathbuni Sakai. Laboratory 29, 71–74. Proceedings of the Japanese Society of Systematic Zoology 15, 36–46. Kurata H. (1969) Larvae of Decapoda Brachyura of Arasaki, Sagami Terada M. (1981) Zoeal development of three species of crab in the Bay—IV. Majidae. Bulletin of the Tokai Region Fisheries Research subfamily Dorippinae. Zoological Magazine 90, 21–32. Laboratory 57, 81–127. Makarov R.R. (1966) The larvae of Macrura, Anomura, and Brachyura of Vassilenko S.V. (1990) On taxonomy and ecology of commensal crabs of the west Kamchatka shelf and their distribution. Moscow: Nauka. family Pinnotheridae (Crustacea, Decapoda, Brachyura) of the Sea Okhotsk and northern part of the Japan Sea. Proceedings of the Matsuo M. (1998) Larval development of two pinnotherid crabs, Zoological Institute, Leningrad 218, 75–95. Asthenognathus inaequipes Stimpson, 1858 and Trithodynamia rath- bunae Shen, 1932 (Crustacea, Brachyura), under laboratory con- Yatsuzuka K., Sakai K. and Del Rio Roman N. (1984) The larvae and ditions. Crustacean Research 27, 122–149. juvenile crabs of Japanese Portunidae (Crustacea Brachyura). III. Charybdis japonica A. Milne-Edwards. Reports of the Usa Marine Morita T. (1974) Morphological observation on the development of larva Biological Institute, Kochi University 6, 23–40. of Eriocheir japonica De Haan. Zoological Magazine 83, 24–81. Motoh H. (1973) Laboratory-reared zoeae and megalopae of Zuwai crab and from the Sea of Japan. Bulletin of the Japanese Society of Scientific Zaslavskaya N.I., Kornienko E.S. and Korn O.M. (2007) Genetic differ- Fisheries 39, 1223–1230. ences between two spider crabs Pisoides bidentatus (A. Milne- Paula J. (1996) A key and bibliography for the identification of zoeal Edwards, 1873) and Pugettia quadridens (de Haan, 1839) stages of brachyuran crabs (Crustacea, Decapoda, Brachyura) from (Decapoda: Brachyura: Majoidea) from the Sea of Japan. Biochemical the Atlantic coast of Europe. Journal of Plankton Research 18, 17–27. Systematics and Ecology 35, 750–756. Park Y.S. and Ko H.S. (2002) Complete larval development of Hemigrapsus longitarsis (Miers, 1879) (Crustacea, Decapoda, Correspondence should be addressed to: Grapsidae), with a key to the known grapsid zoeas of Korea. Korean E.S. Kornienko Journal of Biological Sciences 6, 107–123. Institute of Marine Biology Pessani D., Burri R. and Salton L. (1998) A key for the identification of Far East Branch, Russian Academy of Sciences the known larval stages of the Mediterranean Brachyura. Invertebrate 17 Pal’chevskogo Street, 690041 Vladivostok, Russia Reproduction and Development 33, 191–199. email: [email protected]