Illustrated key for the identification of brachyuran zoeal stages (crustacea decapoda) in the plankton of peter the great bay (sea of japan)

Journal of the Marine Biological Association of the United Kingdom, 2009, 89(2), 379 –386.
doi:10.1017/S0025315408002762 Printed in the United Kingdom

#2008 Marine Biological Association of 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

Larval development is one of the most important periods of
the decapod life cycle; recruitment does not occur unless the
larval period is completed. Larval data can be useful in evaluating species diversity in a region and in specifying the reproduction time of brachyuran species. Morphological features of
larvae are complementary characters for crustacean taxonomy
and phylogeny. The study of larvae is important for the
problem of introduced species arriving in ballast waters or
on fouled ships. However, insufficient attention has been
paid to decapod larval development in the Russian Far East
Seas. Many keys exist for the identification of brachyuran
larvae in different regions of the World Ocean (Ingle, 1992;
Paula, 1996; Ba´ez, 1997; Pessani et al., 1998; Anosov, 2000;

Puls, 2001; Santos & Gonza´lez-Gordillo, 2004; Rice &
Tsukimura, 2007), but only one taxonomic guide is known
for the Sea of Japan (Konishi, 1997). There is no comprehensive key for identification of the brachyuran larvae in Russian
waters of the Sea of Japan; however, the species list of this
region considerably differs from that of coastal waters of
Japan. The aim of this work is to provide a key for the identification of brachyuran larvae in Peter the Great Bay.

MATERIALS AND METHODS

The key is based on the accounts previously published by
various authors (Table 1) and new original descriptions of
brachyuran larvae both taken from plankton samples and
reared from ovigerous crab females in the laboratory.
Corresponding author:
E.S. Kornienko
Email:

Zooplankton was sampled in Vostok Bay (inner bay of
Peter the Great Bay, Sea of Japan) between May and
November 2002 using a Norpac net with a ring diameter of
40 cm and a filtering cone made of a 168 mm mesh, and in
Amursky and Ussurijsky Bays (inner bays of Peter the Great
Bay, Sea of Japan) between April and October 2007 using a
Juday net with a ring diameter of 38 cm and a filtering cone
made of a 168 mm mesh (Figure 1).
It is very difficult to identify the larvae (especially the larvae
of congeneric species) using drawings and descriptions of
different authors in more or less detail. So, zoeae of most considered species were obtained under the laboratory conditions.
This material was used for the construction of the key and the
original figures.

Ovigerous crab females were maintained in an aerated seawater aquarium until larvae hatched. After hatching, larvae
were concentrated at the edge of the aquarium using a point-light
source and transferred to 1-l glass vessels with filtered and
UV-sterilized seawater and reared to the megalopal stage. The
density of larvae was about 100 specimens l21. The water in
the vessels was changed daily. The larvae were fed with newly
hatched nauplii of Artemia salina. Very small zoea of varunid
and pinnotherid crabs were reared using nauplii of the rhizocephalan crustacean, Polyascus polygenea, as a food. Earlier, this
method was described in detail (Kornienko & Korn, 2005a).
All larvae were fixed in 4% formaldehyde for light microscopic studies.
The dichotomous identification key is based mainly on
external morphological characters, which are easy to observe
under a stereomicroscope MBS-10 without specimen dissection. When these features are insufficient, the morphology
and setation of appendages has been included. Moreover, new
original figures were provided to make identification easier.
The outlines of the larvae were drawn using a camera lucida
attached to a binocular Ergaval microscope (Carl Zeiss Jena).
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elena s. kornienko and olga m. korn

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

Species

Authors

Dorippidae
Epialtidae
Oregonidae
Pisidae
Cancridae
Cheiragonidae

Paradorippe granulata (De Haan, 1839)
Pugettia quadridens (De Haan, 1837)
Chionoecetes opilio (O. Fabricius, 1788)
Pisoides bidentatus (A. Milne-Edwards, 1873)
Cancer amphioetus Rathbun, 1989
Erimacrus isenbeckii (Brandt, 1848)
Telmessus cheiragonus (Tilesius, 1812)
Charybdis japonica A. Milne-Edwards, 1861
Pinnixa rathbuni Sakai, 1934
Tritodynamia rathbuni Shen, 1932
Pinnaxodes mutuensis Sakai, 1939
Sakaina yokoyai (Glassell, 1933)
Eriocheir japonicus De Haan, 1835
Hemigrapsus sanguineus (De Haan, 1835)
H. penicillatus (De Haan, 1835)
H. longitarsis (Miers, 1879)

Kurata, 1964; Terada, 1981; Quintana, 1987
Kurata, 1969; Ko, 1998: Kornienko & Korn, 2004
Kurata, 1963b; Motoh, 1973; Haynes, 1973, 1981
Kurata, 1969; Kornienko & Korn, 2007
Iwata & Konishi, 1981

Aikawa, 1937; Kurata, 1963a; Makarov, 1966
Kurata, 1963a
Yatsuzuka et al., 1984
Sekiguchi, 1978; Konishi, 1983; Kornienko & Korn, 2005b
Matsuo, 1998
Konishi, 1981b
Not described
Morita, 1974; Kim & Hwang, 1990
Hwang et al., 1993; Kornienko et al., 2008
Hwang & Kim, 1995; Kornienko et al., 2008
Park & Ko, 2002; Kornienko et al., 2008

Portunidae
Pinnotheridae

Varunidae

The key was constructed for zoea I, but the characters used
in the key do not change or change slightly through the successive zoeal stages, with the exception of peculiar cases.
Main characters of brachyuran zoea used for the identification
are represented in Figure 2.

RESULTS

Key for the identification of brachyuran zoea I
(interspecific distinctions)
1a. Carapace with rostral, dorsal and lateral spines
(Figure 3A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Fig. 1. Map showing the sampling area.

1b. Carapace with rostral spine only (Figure 3B) or with
rostral and dorsal spines (Figure 3C), lateral spines
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2a. Abdominal somites 3–5 with well developed posterolateral spines; furcal rami with lateral and dorsal spines
(Figure 4A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2b. Abdominal somites 3–5 with poorly developed posterolateral spines (Figure 4B, C); furcal rami without lateral
and dorsal spines (Figure 4B) or with lateral spines only
(Figure 4C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3a. Furcal rami longer than proximal part of telson; dorsal
and lateral furcal spines short (Figure 5A). . . . . . . . . . 4
3b. Furcal rami shorter than proximal part of telson; lateral
furcal spines long (Figure 5B, C) . . . . . . . . . . . . . . . . . 6
4a. All carapace spines spinulated; dorsal spine straight;
posterolateral spines longer than half abdominal
somite; each furcal ramus with three spines. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . Chionoecetes opilio (Figure 11A)

Fig. 2. Main characters used for the identification of brachyuran zoea.


key for brachyuran larvae

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

Fig. 3. Carapace of zoea in Eriocheir japonicus (A), Sakaina yokojai (B) and
Pisoides bidentatus (C).

[Lateral spines long, slightly shorter dorsal and

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
rathbuni (B) and Pisoides bidentatus (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; outermost 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. . . . . . . .
. . . . . . . . . . . . . Cancer amphioetus (Figure 11B)
[Antenna biramous; protopod with two rows of
spinules, nearly half of rostral spine; exopod with

Fig. 6. Abdomen and telson of zoea in Cancer amphioetus (A, A0 ) and
Charybdis japonica (B) (after Yatsuzuka et al., 1984).

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elena s. kornienko and olga m. korn

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
(Figure 6B) . . . . . . . . . . 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
with long posterolateral spines. Each furcal
ramus with three spines: one long lateral spine
and two short dorsal spines. Larvae found in
April and May.]

6b. Abdominal somites 2 and 3 with a pair of lateral knobs;
posterolateral spines of somite 4 shorter than somite 5;
lateral telsonal spine longer than half furcal ramus; each
furcal ramus with four inner setae (Figure 5C) . . . . . . . .
. . . . . . . . . . . . . . . . . Erimacrus isenbeckii (Figure 11E)
[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
with long posterolateral spines. Each furcal
ramus with three spines: long lateral spine and
two short dorsal spines. Larvae found in April
and May.]

Fig. 8. Antenna and carapace of zoea in Tritodynamia rathbuni (A) and
Eriocheir japonicus (B).

7b. Lateral carapace spines arranged more ventrally than in
typical brachyuran zoea; abdominal somites 4–5 not
cylindrical but laterally expanded; telson subrectangular;
furcal rami without spines or with minute lateral spines
(Figure 7D –F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
8a. Dorsal and rostral spines longer than carapace; dorsal carapace spine straight; antenna uniramous (Figure 8A) . . .
. . . . . . . . . . . . . . . Tritodynamia rathbuni (Figure 11F)
[Antennal protopod with two rows of spinules and
with short seta near base. Posterolateral spines on
abdominal somites 3 – 5 highly reduced. Larvae
found from June to September.]

7a. Lateral carapace spines arranged as in typical brachyuran
larvae; all abdominal somites cylindrical; telson triangular;
furcal rami without spines (Figure 7A, B) . . . . . . . . . . 8

8b. Dorsal and rostral spines slightly shorter than carapace;
dorsal carapace spine slightly curved; antenna biramous
(Figure 8B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Fig. 7. Carapace of zoea in Tritodynamia rathbuni (A) and Pinnaxodes
mutuensis (C); telson of zoea in Tritodynamia rathbuni (B); Pinnaxodes
mutuensis (D), Pinnixa rathbuni (E) and Sakaina yokoyai (F).

Fig. 9. Antenna, antennule and abdomen of zoea in Eriocheir japonicus (A)
and Hemigrapsus sanguineus (B).


key for brachyuran larvae

9a. Abdominal somites 2– 4 with a pair of lateral knobs
(lateral knobs on somite 4 disappear in successive
stages); antennule with 2 aesthetascs; antennal exopod
nearly half protopod (Figure 9A). . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . Eriocheir japonicus (Figure 11G)

[Rostral spine nearly one-third antenna; dorsal
spine with sparse blunt spinules. Antenna biramous; protopod and exopod sub-equal.
Abdominal somites 2 with a pair of lateral

[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
(Figure 7D). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . 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, A 0 ). Larvae
found in July and August.]

12b. Rostral and dorsal spine shorter than carapace, dorsal
spine slightly curved (Figure 10B) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pugettia
quadridens, Pisoides bidentatus (Figure 11M)

Fig. 10. Carapace (A) and telson (A0 ) of zoea in Paradorippe granulata;
carapace of zoea in Pugettia quadridens (B).

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elena s. kornienko and olga m. korn

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).
Larvae found from June to September.]

Most Brachyura from Peter the Great Bay pass through five
zoeal stages. Majiod crabs (Pisoides bidentatus, Pugettia quadridens, and Chionoecetes opilio) have an abbreviated development including only two zoeal stages. Two of three
pinnotherid crabs also undergo an abbreviated development:
Sakaina yokoyai passes through probably three zoeal stages;
Pinnaxodes mutuensis, through four zoeal stages. Four zoeal
stages are known also for Paradorippe granulata.
Different zoeal stages of brachyuran crabs (age distinctions) are easily determined using the number of natatory
setae on the exopods of maxillipeds, the number of setae
along the posterior telsonal margin, the presence of uniramous or biramous pleopod buds and some other morphological features. The same stages in the species with abbreviated
development are more advanced in the number of characters
over those of the species with longer development.

Identification of different zoeal stages of
brachyuran crabs (age distinctions)
Zoea I. Eyes sessile; exopod of maxilliped each with 4 natatory
setae; abdomen consists of 5 somites and telson; posterior
telsonal margin with 3 þ 3 setae (in Paradorippe granulata,

posterior telsonal margin with two setae only in all zoeal
stages).
Zoea II. Eyes stalked; exopod of maxilliped each with 6 natatory setae; in most species abdomen consists of 5 somites and
telson; pleopod buds absent; posterior telsonal margin with
3 þ 3 setae. In Pisoides bidentatus, Pugettia quadridens, and
Chionoecetes opilio, zoea II is the last stage; hence, the sixth
somite is delineated, somites 2–6 with biramous pleopod
buds. In Sakaina yokoyai, pleopod buds uniramous. In

Charybdis japonica and Chionoecetes opilio, posterior telsonal
margin with 4 þ 4 setae.
Zoea III. Exopod of maxilliped each with 8 natatory setae;
the sixth somite delineated, with the exception of Pinnixa
rathbuni which has 5 somites in all zoeal stages. In Cancer
amphioetus, Trithodynamia rathbuni, Erimacrus isenbeckii,
Telmessus cheiragonus, Eriocheir japonicus and Hemigrapsus
species, posterior telsonal margin with 4 þ 4 setae; in
Charybdis japonica and Pinnaxodes mutuensis, with additional
small unpaired setae. In Sakaina yokoyai, pleopod buds
biramous.
Zoea IV. Exopod of maxilliped each with 10 natatory setae;
somites 2–6 with uniramous pleopod buds. In Cancer
amphioetus and Tritodynamia rathbuni, posterior telsonal
margin with 5 þ 5 setae.
Zoea V. Exopod of maxilliped each with 12 natatory
setae; pleopod buds biramous. In Eriocheir japonicus and
Hemigrapsus species, posterior telsonal margin with 5 þ 5 setae.


key for brachyuran larvae

DISCUSSION

REFERENCES

According to Adrianov & Kussakin (1998), 20 brachyuran
species belonging to 8 families and 16 genera inhabit Peter
the Great Bay (Russian waters of the Sea of Japan). We
found larvae of only 16 species from 9 families and 14

genera occurring in the plankton of Peter the Great Bay
(Table 1). To date, we have not found both adults and
larvae of Goetice depressus (de Haan, 1835), Helice tridens
de Haan, 1833, and Paradromia japonica (Henderson,
1888). According to Vassilenko (1990), adult specimens of
Pinnixa tumida Stimpson, 1858 are found only in Possyet
Bay (eastern Peter the Great Bay). Brachyuran larvae occur
in Peter the Great Bay from April to November.
Zoea of each species represented in this key has been
previously described. Only the larvae of Sakaina yokoyai
are not known yet; however, the morphology of closely
related species, S. japonica, has been described (Konishi,
1981a). Both adults and larvae of S. yokoyai are rarely
found in Peter the Great Bay, so we failed to obtain the
total series of their larval stages. Nevertheless, we believe
that S. yokoyai, like S. japonica, passes through three zoeal
stages.
Frequently, the distinction between larvae of congeneric
species is based on slight differences. The larvae of congeneric species, described on the basis of specimens hatched in
the laboratory from ovigerous females, can be identified
only to the generic level when collected in nature. In our
case, the specific identification of three Hemigrapsus
species (H. sanguineus, H. penicillatus and H. longitarsis) is
very difficult. Zoea I and zoea II of these species are nearly
identical. The larvae are distinguishable only from zoea III.
Zoea III –V of Hemigrapsus species differ in the number of
dorsomedial setae on the abdominal somite I and in the
number of setae on the posterodorsal arch. It is pertinent
to note that among the three Hemigrapsus species, zoea
and megalopa of H. penicillatus differ greatly (Kornienko

et al., 2008). Despite the great similarity of larvae in
varunid crabs of the genera Hemigrapsus and Eriocheir, the
latter possess a number of distinctive features in all developmental stages (Kornienko & Korn, 2005a; Kornienko et al.,
2008).
It was recently shown that zoea of Pisoides bidentatus and
Pugettia quadridens belonging to different families are also
nearly identical, with the exception of a more intensive
coloration of the latter. Some differences appear only in
the megalopal stage. Based on the larval similarity
(Kornienko & Korn, 2007) and high genetic identity of
adults (Zaslavskaya et al., 2007) these two species should
be assigned to one genus.
The larvae of the other ten species belonging to different
families and genera are easy to identify in the plankton. The
features used in the key were chosen to enable identification
of zoea to the species level and do not reflect any systematic
arrangement of decapod families.

Adrianov A.V. and Kussakin O.G. (1998) A check-list of biota of the Peter
the Great Bay, the Sea of Japan. Vladivostok: Dalnauka.

ACKNOWLEDGEMENTS

The project was supported by the Far East Branch of the
Russian Academy of Sciences (grant no. 06-III-A-06-164)
and by the Russian Foundation for Fundamental Researches
(grant no. 08-04-00929).

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Correspondence should be addressed to:
E.S. Kornienko
Institute of Marine Biology
Far East Branch, Russian Academy of Sciences
17 Pal’chevskogo Street, 690041 Vladivostok, Russia
email: