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Zoeal Stages of Leptomithrax edwardsii (Crustacea: Decapoda: Majidae)

Described from Laboratory Reared Material

Article in Animal Systematics Evolution and Diversity · July 2012

DOI: 10.5635/ASED.2012.28.3.185

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 Anim. Syst. Evol. Divers. Vol. 28, No. 3: 185-191, July 2012
http://dx.doi.org/10.5635/ASED.2012.28.3.185

Zoeal Stages of Leptomithrax edwardsii

(Crustacea: Decapoda: Majidae) Described
from Laboratory Reared Material
Jung-Ha Kang1, Yong-Seok Lee2, Ji-Eun Jeong2, Hyun-Sook Ko3,*
Biotechnology Research Division, National Fisheries Research and Development (NFRDI),
1

Busan 619-705, Korea

2
Department of Parasitology, College of Medicine and UHRC, Inje University, Busan 614-735, Korea
3
Department of Biological Science, Silla University, Busan 617-736, Korea

Abstract

Zoeas of Leptomithrax edwardsii were reared in the laboratory. Two zoeal stages are described and illustrated.
The first zoeal stage of L. edwardsii is compared with those of seven known species of the family Majidae. It
differs from previous description in the endopodal setation characters of the maxillule and the second maxilliped.
It appears most similar to L. bifidus and Schizophroida simodaensis of the northwestern Pacific. A provisional
key for identifying eight majid zoeas is included.
Keywords: Majidae, spider crab, Leptomithrax edwardsii, zoea, L. bifidus, Schizophroida simodaensis, key,
northwestern Pacific

INTRODUCTION the zoeal stages of L. edwardsii in detail and compare them

with previously described majid zoeas from the northwestern
There are eight species of Majidae Samouelle, 1819 from Pacific.
Korean waters: four species of Majinae [Leptomithrax bifidus
(Ortmann, 1893), L. edwardsii (De Haan, 1835), Maja miersii
Walker, 1887, and M. spinigera (De Haan, 1837)] and four MATERIALS AND METHODS
species of Mithracinae [Micippa cristata (Linnaeus, 1758),
M. philyra (Herbst, 1803), M. platipes Rüppell, 1830, and On 30 December 2011 an ovigerous crab of L. edwardsii was
M. thalia (Herbst, 1803)] (see the Korean Society of System- collected from Gijang (35�19′N, 129�17′E), Busan, Korea.
atic Zoology, 1997; Ng et al., 2008). Of these, L. edwardsii And the larvae hatched in the laboratory on 31 January 2012.
inhabits the sandy and muddy bottoms at depths between A total of 120 zoeas was reared individually at a water tem-
50-150 m with a report distributed to Korea and Japan (Kim, peratures of 15±1� C. Each zoea was held in a plastic well
1973). containing 5-6 mL of sea water for the first and second stages.
Larval descriptions of Majidae are limited to eight species The water was changed daily and each zoea was provided
in the northwestern Pacific: Schizophrys aspera (Milne- with newly hatched Artemia nauplii once a day. Each indi-
Edwards, 1834) by Kurata (1969) and Terada (1981), Schiz- vidual culture was checked daily for exuviae and dead zoeas.
ophroida simodaensis, Sakai, 1933 by Terada (1981), L. bifi- Moults and dead larvae were fixed and preserved in 10%
dus by Kurata (1969) and Terada (1981), L. edwardsii by neutral formalin for later examination. Dissected appendages
Kurata (1969), M. spinigera by Terada (1981), M. philyra by were examined and drawn using a Leitz Laborlux S micro-
Ko (1995), M. thalia by Kurata (1969) and Tiarinia cornig- scope (Swiss) with a camera lucida. Setal counts on appen-
era (Latreille, 1825) by Kurata (1969). However, L. edwardsii dages and measurements were based on the mean of ten
is known for its first zoeal stage with a brief description by specimens for zoeal stage. The sequence of larval description
Kurata (1969) (Table 1). Therefore, in this study, we describe (Clark et al., 1998) is based on the malacostracan somite plan

cc This is an Open Access article distributed under the terms of the Creative *To whom correspondence should be addressed
Commons Attribution Non-Commercial License (http://creativecommons.org/ Tel: 82-51-999-5473, Fax: 82-51-999-5176
licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, E-mail: [email protected]
and reproduction in any medium, provided the original work is properly cited.
pISSN 2234-6953 eISSN 2234-8190 Copyright The Korean Society of Systematic Zoology
Jung-Ha Kang, Yong-Seok Lee, Ji-Eun Jeong, Hyun-Sook Ko

Table 1. Comparison of the first zoeal characteristics of Lepto- lopa was not entirely successful because the megalopa was
mithrax edwardsii described by Kurata (1969) with those obtained unable to extricate themselves from the zoeal exoskeleton.
in the present study
The first zoeal stage is described and illustrated completely.
Kurata (1969) Present study Only the main differences from the first zoea are described
CL ND 1.11 mm for the second zoeal stage.
RDL 1.23 mm 1.56 mm
Carapace
Lateral margin 1 plumose, 3 setae 1 plumose, 4 setae
First zoea (Figs. 1, 2)
Rostral spine Present Present Size: CL, 1.11±0.02 mm; RDL, 1.56±0.03 mm.
Dorsal spine Present Present Chromatophores (Fig. 1). Predominantly yellow pigments
Lateral spine Absent Absent which occurring on carapace, antenna, mandibles, basis of
Antennule ND 3 aesthetascs, 2 setae
Antenna
maxillipeds, pleomeres 1-4, and fork of telson; brown and
Endopod bud Present Present black pigments which occurring behind eye, posterodorsal
Exopod 2 subterminal setae 2 subterminal setae and posterolateral regions of carapace, mandibles, basis of
Maxillule maxillipeds, posterolateral margins of pleomeres 1-4, fork
Endopod 1, 7 setae 1, 2+4 setae
Basial endite ND 7 setae
of telson.
Coxal endite ND 7 setae Carapace (Fig. 2A). Dorsal spine slightly shorter than 1/2 CL,
Maxilla slightly curved posteriorly; rostral spine straight, approxi-
Endopod 5 setae 3+2 setae mately 1/2 length of dorsal spine; lateral spine absent; pair
Basial endite ND 5+4 setae
Coxal endite ND 3+4 setae
of posterodorsal setae present; each ventral margin with 1
Scaphognathite ND 10+1 setae plumose anterior and 4 posterior setae; eyes sessile.
Maxilliped I Antennule (Fig. 2B). Uniramous; endopod absent; exopod
Endopod ND 3, 2, 1, 2, 5 setae with 1 long, stout aesthetasc, 2 shorter, thinner aesthetascs,
Basis ND 2, 2, 2, 3 setae
Maxilliped II
1 long and 1 shorter setae, all terminal.
Endopod 0, 1, 6 setae 0, 7 setae Antenna (Fig. 2C). Biramous; endopod bud present; protopod
Basis ND 1, 1, 1 setae about twice length to rostral spine, spinulate distally; exopod
Abdomen with 2 unequal subterminal setae, its tip shorter than protopod.
Lateral process Somite 2 Somite 2
Telson
Mandibles (Fig. 2D). Asymmetrical; right molar and left
Fork 1 large, 1 large, molar processes each with tooth, confluent with incisor pro-
2 smaller spines 2 smaller spines cess; palp absent.
CL, carapace length; RDL, rotral spine to dorsal spine length; Maxillule (Fig. 2E). Coxal and basial endites both with 7
ND, no data. setae; endopod 2-segmented, proximal segment with 1 seta,
distal segment with 6 (2 subterminal, 4 terminal) setae; exo-
pod seta and epipod absent.
and described from anterior to posterior. Setal armature is Maxilla (Fig. 2F). Coxal endite bilobed, with 3+4 setae;
described from proximal to distal segments and in order of basial endite bilobed, with 5+4 setae; endopod bilobed,
endopod to exopod. The long plumose natatory setae of the with 3+2 setae; exopod (scaphognathite) margin with 10
first and second maxillipeds were drawn truncated. A micro- plumose setae plus 1 distal stout process.
meter was used for measurements: Carapace length (CL) was First maxilliped (Fig. 2G). Coxa without seta; basis with 9
measured from the anterior margin of the eye to the most setae arranged 2+2+2+3; endopod 5-segmented with 3, 2,
posterior carapace margin. Rostral spine to dorsal spine length 1, 2, 5 (1 subterminal, 4 terminal) setae; exopod 2-segmented,
(RDL) was measured from tip of the rostrum to tip of the distal segment with 4 terminal plumose natatory setae.
dorsal spine. The classification follows that of Ng et al. (2008). Second maxilliped (Fig. 2H). Coxa without seta; basis with
The zoeas and spent female were deposited at Silla Univer- 3 setae arranged 1+1+1; endopod 2-segmented, with 0, 7
sity, Korea. (1 proximal, 3 subterminal, 3 terminal) setae; exopod 2-seg-
mented, distal segment with 4 terminal plumose natatory
setae.
RESULTS Third maxilliped (Fig. 2I). Biramous.
Pereiopods (Fig. 2J). Present as buds.
Zoeal development consists of two zoeal stages. The mean Pleon (Fig. 2K). With 5 pleomeres; pleomere 1 with 2 dor-
duration of the first zoeal stage at 15�
C was 16 days. The somedial setae; pleomere 2 with a pair of lateral processes
second zoeal stage survived for 17 days. The moult to mega- directed anteriorly; pleomeres 2-5 each with a pair of pos-

186 Anim. Syst. Evol. Divers. 28(3), 185-191

 Zoeal Stages of Leptomithrax edwardsii

Fig. 1. Leptomithrax edwardsii, first stage zoeas, live specimens.

terodorsal setae, pleomeres 3-5 with long posterolateral pro- oped.

cesses, progressing reduced in size; pleopod present as buds Pereiopods (Fig. 3J). More developed, segments differentiated,
without endopods. chela bilobed.
Telson (Fig. 2K, L). Each fork long, covered with setules, Pleon (Fig. 3K). With 6 pleomeres; pleomere 1 with 3 dor-
with 1 stout, 2 shorter spines laterally; each posterior margin somedial setae; pleopod buds more developed, endopods
with 3 serrated setae. present.
Telson (Fig. 3K). Unchanged.
Second zoea (Fig. 3)
Size: CL, 1.43±0.05 mm; RDL, 1.68±0.05 mm.
Carapace (Fig. 3A). Each ventral margin with 1 plumose DISCUSSION
anterior and 7 posterior setae; eyes stalked.
Antennule (Fig. 3B). Biramous, endopod present, exopod Kurata (1969) described only the first zoeal stage; however,
with 1 subterminal and 5 terminal aesthetascs, 2 terminal his description was too brief compared with that of the pre-
short setae. sent study (Table 1). Moreover, he described the endopod of
Antenna (Fig. 3C). Endopod bud approximately 1/2 length the maxillule as having 1, 7 setae, but it does not coincide
of protopod. with the other six known species of Majidae, which have 1,
Mandibles (Fig. 3D). Right molar process with 2 teeth, con- 2+4 setae. Also, he described the endopod of the second
fluent with incisor process; palp present. maxilliped as having 0, 1, 6 setae; however, we found 0, 7
Maxillule (Fig. 3E). Basial endite with 8 setae. Exopod plu- setae as in S. simodaensis.
mose seta now present. In the northwestern Pacific, larval informations of the
Maxilla (Fig. 3F). Basial endite with 5 +5 setae; exopod family Majidae are available for eight species [three species
(scaphognathite) margin with 16 marginal plumose setae. (Micippa philyra, M. thalia, and Tiarinia cornigera) of the
First maxilliped (Fig. 3G). Coxa with 1 seta; exopod distal subfamily Mithracinae and five species (Maja spinigera,
segment with 6 terminal natatory setae. Schizophrys aspera, S. simodaensis, Leptomithrax bifidus,
Second maxilliped (Fig. 3H). Exopod distal segment with 6 and L. edwardsii) of the subfamily Majinae]. The morpho-
terminal natatory setae. logical characteristics of the majid zoeas can be summarized
Third maxilliped (Fig. 3I). Endopod and exopod more devel- as follows (Table 2): Endopod bud of the antenna present;

Anim. Syst. Evol. Divers. 28(3), 185-191 187

Jung-Ha Kang, Yong-Seok Lee, Ji-Eun Jeong, Hyun-Sook Ko

F
E

K
I J

L
G

Fig. 2. Leptomithrax edwardsii, first zoeal stage. A, Lateral view; B, Antennule; C, Antenna; D, Mandibles; E, Maxillule; F, Maxilla;
G, First maxilliped; H, Second maxilliped; I, Third maxilliped; J, Pereiopods; K, Dorsal view of abdomen and telson; L, Fork of telson.
Scale bars: A, K==0.5 mm, B-J, L=
=0.1 mm.

188 Anim. Syst. Evol. Divers. 28(3), 185-191

 Zoeal Stages of Leptomithrax edwardsii

B
C
D

Fig. 3. Leptomithrax edwardsii, second zoeal stage. A, Lateral view; B, Antennule; C, Antenna; D, Mandibles; E, Maxillule; F, Maxilla;
G, First maxilliped; H, Second maxilliped; I, Third maxilliped; J, Pereiopods; K, Dorsal view of abdomen and telson. Scale bars: A,
K=
=0.5 mm, B-J= =0.1 mm.

Anim. Syst. Evol. Divers. 28(3), 185-191 189

Jung-Ha Kang, Yong-Seok Lee, Ji-Eun Jeong, Hyun-Sook Ko

Table 2. Comparison of the first zoeal characteristics in eight species of Majidae from the northwestern Pacific
Micippa Micippa Tiarinia Maja Schizophrys Schizophroida Leptomithrax Leptomithrax
Species
philyra thalia cornigera spinigera aspera simodaensis bifidus edwardsii

CL (mm) 0.80 ND ND 0.91 0.60 0.79 0.68 1.11

RDL (mm) ND 0.92 0.93 1.26 1.22 1.25 0.88 1.56
Carapace
Rostral spine 1/2 CL 1/2 CL Reduced 1/2 CL 1/2 CL 1/5 CL 1/7 CL 1/5 CL
Dorsal spine Absent Absent 1/3 CL 1/4 CL 1/2 CL 1/2 CL 1/4 CL 1/3 CL
Lateral spine Present Present Absent Absent Present Absent Absent Absent
Antenna
Endopod bud Present Present Present Present Present Present Present Present
Exopod spine 2 medial 2 medial 2 submedial 2 subterminal 2 subterminal 2 subterminal 2 subterminal 2 subterminal
Maxillule
Endopod setae 1, 2+4 1, 2+4 ND 1, 2+4 1, 2+4 1, 2+4 1, 2+4 1, 2+4
Maxilla
Endopod setae 3+3 3+3 3 3+2 3+2 ND 3+2 3+2
Maxilliped I
Endopod setae 3, 2, 1, 2, 5 ND ND 3, 2, 1, 2, 5 3, 2, 1, 2, 5 3, 2, 1, 2, 5 3, 2, 1, 2, 5 3, 2, 1, 2, 5
Basis setae 2, 2, 3, 3 ND ND 2, 2, 2, 3 2, 2, 2, 3 2, 2, 2, 3 2, 2, 2, 3 2, 2, 2, 3
Maxilliped II
Endopd setae 0, 1, 4 0, 1, 4 ND 0, 1, 6 0, 1, 6 0, 7 0, 1, 6 0, 7
Basis setae 1, 1, 1 ND ND 1, 1, 1 1, 1, 1 ND 1, 1, 1 1, 1, 1
Abdomen
Lateral process Somites 2, 3 Somites 2, 3 Somite 2 Somites 2, 3 Somites 2, 3 Somite 2 Somite 2 Somite 2
Telson fork
Lateral spine 1 large, 1 large, 1 large 1 large 1 large, 1 large 1 large, 1 large,
2 smaller 1 smaller 2 smaller 2 smaller 2 smaller
Sources Ko (1995) Kurata (1969) Kurata (1969) Terada (1981) Terada (1981) Terada (1981) Terada (1981) Present study

CL, carapace length; RDL, rotral spine to dorsal spine length; ND, no data.

endopod of the maxillule with 1, 2+4 setae; the endopod of ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙M. philyra

the first maxilliped with 3, 2, 1, 1, 5 setae; the basis of the 3. Lateral carapace spine present ∙∙∙∙∙∙∙∙∙∙Schizophrys aspera
second maxilliped with 1, 1, 1 setae; and the telson fork with - Lateral carapace spine absent∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙4
lateral spines. However, the zoeas of Majinae are different 4. Lateral processes on abdominal somites 2, 3 ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙
from those of Mithracinae by having a dorsal carapace spine, ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙Maja spinigera

no lateral carapace spines (except in S. aspera), 2 subterminal - Lateral process on abdominal somite 2 ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙5
setae in the antennal exopod, 3+2 setae in the endopod of 5. Rostral carapace spine reduced or extremely short ∙∙∙∙∙∙∙∙∙∙
the maxilla, 2, 2, 2, 3 setae in the basis of the first maxilliped, ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙Tiarinia cornigera

0, 1, 6 or 0, 7 setae in the endopod of the second maxilliped, - Rostral carapace spine approximately 1/2 length of dorsal
and a lateral process in the abdominal somite 2 (except in carapace spine ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙6
M. spinigera and S. aspera). 6. Endopod of second maxilliped with 0, 1, 6 setae ∙∙∙∙∙∙∙∙∙∙∙∙∙
As shown in Table 2, the first zoea of L. edwardsii appears ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙Leptomithrax bifidus

most similar to those of L. bifidus and S. simodaensis in - Endopod of second maxilliped with 0, 7 setae ∙∙∙∙∙∙∙∙∙∙∙∙∙∙7
Majidae; however, it can be separated from the other species 7. Fork of telson with 1 large lateral spine ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙
on the basis of the endopodal setae of the second maxilliped ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙Schizophroida simodaensis

and the features of telson fork. - Fork of telson with 1 large and 2 smaller lateral spines ∙∙∙∙
∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙L. edwardsii

The following provisional key is provided

for identifying the eight majid zoeas
1. Dorsal carapace spine present ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙2 ACKNOWLEDGMENTS
- Dorsal carapace spine absent ∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙3
2. Tip of exopod longer than that of protopod in antenna ∙∙∙∙∙ This work was supported by the National Fisheries Research
∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙∙Micippa thalia and Development Institute (RP-2012-BT-010).
- Tip of exopod shorter than that of protopod in antenna ∙∙∙∙

190 Anim. Syst. Evol. Divers. 28(3), 185-191

 Zoeal Stages of Leptomithrax edwardsii

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