JOURNAL OF CRUSTACEAN BIOLOGY, 24(1): 110–120, 2004
ZOEAL STAGES OF CONCHODYTES NIPPONENSIS (DECAPODA:
PALAEMONIDAE) REARED IN THE LABORATORY
Hoi Jeong Yang and Hyun Sook Ko
Department of Life Sciences, Silla University, Busan 617-736, Republic of Korea
(HJY, correspondence): [email protected])
ABSTRACT
Zoeal stages of Conchodytes nipponensis, not previously known from Korea, are described from
laboratory-reared material. This is the first account of the larval development of a species from the genus
Conchodytes and the first larval description of a pontoniid species from Asian waters. The
Gnathophyllidae had been considered as a synonym with the Pontoniinae by Bruce, whereas others
regard both groups as distinct. Morphological comparisons between the first zoea of C. nipponensis and
that of Gnathophyllum americanum indicate that the Gnathophyllidae have great affinity with the
Pontoniinae. According to the viewpoint of some authors, similar larvae do not necessarily signify related
adults. Therefore, the Pontoniinae and the Gnathophyllidae may be distinct.
The palaemonid subfamily Pontoniinae comprises a large number of species with life
histories ranging from free-living to highly
specialized associates of various marine invertebrates. Seven species assigned to Conchodytes are reported from Indo-West Pacific
waters: C. kempi Bruce, 1989; C. maculatus
Bruce, 1989; C. meleagrinae Peters, 1852; C.
monodactylus Holthuis, 1952; C. nipponensis
(De Haan, 1844); C. philippinensis Bruce,
1996; and C. tridacnae Peters, 1852 (see Chace
and Bruce, 1993; Bruce, 1996). One of these,
C. nipponensis, is known to live in commensal
association with pectinid and pinnid bivalve
mollusks and is reported to occur in the
Philippines, eastern Australia, Japan (Chace
and Bruce, 1993), and here for the first time in
Korea. The only other pontoniid species reported from Korea is Periclimenaeus gorgonidarum (Balss, 1913) (The Korean Society of
Systematic Zoology, 1997).
Larvae of the Pontoniinae are poorly documented world-wide, with larval descriptions
known for only nine (about 10.8%) of the 83
genera currently described in the subfamily
(Okuno, 1999). These are represented by 23
species from Atlantic waters, the Mediterranean
Sea, the Red Sea, India, Australia, New
Zealand, and Fiji (Table 1).
The present paper provides the first account
of the larval development of a species assigned
to the genus Conchodytes. The zoeal stages of
C. nipponensis are described and illustrated in
detail, and the morphological characteristics of
the first zoea are compared with those of
Pontonia pinnophylax (Otto, 1821) and Pontonia flavomaculata Heller, 1864. The systematic
relationships between the Pontoniinae and
Gnathophyllidae are discussed.
MATERIALS AND METHODS
On 13 June 2002, one ovigerous female of Conchodytes
nipponensis was collected from fan shells Atrina pectinata
Linnaeus, 1758, in Jagalchi Fish Market, Busan, Korea. The
female released about 250 larvae on 24 June 2003. Two
hundred larvae were reared individually in Corning twelvetissue culture plates, ranging in volume from 6 to 8 mL per
well of aerated seawater of 33.3& in a growth chamber at
258C. Larvae were fed with newly hatched Artemia nauplii
and transferred daily to newly prepared culture plates. Some
specimens in each stage were preserved in 5% neutral
Formalin. Measurements and setal counts were based on ten
specimens for each zoeal stage. Dissected appendages were
examined using a Leitz Laborlux S microscope, and
drawings were made with the aid of a camera lucida. The
chromatophore pattern was determined by observing living
larvae. Body length (BL) was determined from the postorbital margin to the posteromedian margin of the telson,
excluding posterior setae. Carapace length (CL) was
determined from the postorbital margin to the posteromedian
margin of the carapace. The setal armature of the
appendages is described from proximal segment towards
distal segment.
RESULTS
Six zoeal stages were obtained. Four zoeal
stages are described and illustrated. However,
the last two stages could not be described
because only three and four zoeas were
obtained, respectively. The fifth zoea had
pereiopods 1–4 with the exopods only. In the
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YANG AND KO: ZOEAL STAGES OF CONCHODYTES NIPPONENSIS
Table 1. Described larval stages of the species of the Pontoniinae. @ ¼ abbreviated development; Z, L, P, and Y ¼ zoeal,
last, postlarval, and young stages, respectively.
Species
Anchistus custos (Forsskål, 1775) as
Anchistus inermis
Coralliocaris graminea (Dana, 1852)
Conchodytes nipponensis
(De Haan, 1844)
Harpiliopsis beaupresii (Audouin, 1826) as
Harpilius beaupresii
Philarius gerlachei (Nobili, 1905) as
Harpilius gerlachei
Periclimenaeus bermudensis
(Armstrong 1940) as Periclimenes
( Periclimenaeus) bermudensis
Periclimenaeus wilsoni (Hay, 1917)? as
Periclimenes ( Periclimenaeus)
wilsoni?
Periclimenaeus stylirostris Bruce, 1969
Periclimenes americanus
(Kingsley, 1878)
Periclimenes brevicarpalis (Schenkel,
1902) as Periclimenes ( Ancylocaris)
brevicarpalis
Periclimenes calmani Tattersall, 1921 as
Periclimenes ( Ancylocaris) calmani
Periclimenes diversipes Kemp, 1922 as
Periclimenes ( Ancylocaris) diversipes
Periclimenes grandis (Stimpson, 1860) as
Periclimenes ( Ancylocaris) grandis
Periclimenes grandis (Stimpson, 1860) as
Periclimenes ( Ancylocaris) grandis
Periclimenes indicus (Kemp, 1915) as
Periclimenes ( Periclimenes) indicus
Periclimenes iridescens Lebour, 1949 as
Periclimenes ( Periclimenes) iridescens
Periclimenes longicaudatus (Stimpson,
1860)? as Periclimenes ( Periclimenes)
longicaudatus?
Periclimenes pandionis Holthuis, 1951 as
Periclimenes ( Periclimenes) pandionis
Periclimenes scriptus (Risso, 1822) as
Pontonia scriptus
Periclimenes soror Nobili, 1904 as
Periclimenes ( Periclimenes) soror
Periclimenes yaldwyni Holthuis, 1959
Pontonia flavomaculata Heller, 1864
Pontonia flavomaculata Heller, 1864
Pontonia flavomaculata Heller, 1864
Pontonia pinnophylax (Otto, 1821) as
Pontonia tyrrhena
Pontonia pinnophylax (Otto, 1821)
Pontonia pinnophylax (Otto, 1821)
Pontonia minuta Baker, 1907
Typton spongicola O. G. Costa, 1844
Typton spongicola O. G. Costa, 1844
Typton spongicola O. G. Costa, 1844
Stages
Locality
Reference
Z1
India
Prasad and Tampi, 1957
Z1–2
Z1–4
Ghardaqa, Red Sea
Busan, Korea
Gurney, 1938
present study
Z1–2
Ghardaqa, Red Sea
Gurney, 1938
Z1
Ghardaqa, Red Sea
Gurney, 1938
Z1–2
Bermuda, western Atlantic
Gurney and Lebour, 1941
L, P
Bermuda, western Atlantic
Gurney and Lebour, 1941
P
Z1, 5
Port Essington, Australia
Bermuda, western Atlantic
Bruce and Coombes, 1995
Gurney, 1936
Z1
India
Nayer, 1947
Z1–5
Red Sea
Gurney, 1927
Z1
Ghardaqa, Red Sea
Gurney, 1938
Z1
Ghardaqa, Red Sea
Gurney, 1938
Z1–6, P
India
Pillai, 1950
Z1–5, P
India
Menon, 1949
Z1
Bermuda, western Atlantic
Lebour, 1949a
L, P, Y 1–3 Bermuda, western Atlantic
Gurney and Lebour, 1941
Z1–2
Florida, western Atlantic
Gore et al., 1981
Z1
Naples, Mediterranean
Caroli, 1926
Z1
Nidraunimbota Bay, Suva, Fiji
Wear, 1976
Z8
Z1
Z1
Z1–8, P
Z1
New Zealand
Gulf of Marseilles, Mediterranean
Gulf of Marseilles, Mediterranean
Gulf of Patti, Messina, Mediterranean
Naples, Mediterranean
Packer, 1985
Gourret, 1884, 1888
Bourdillon-Casanova, 1960
Costanzo et al., 1996
Caroli, 1926
Z1–3
Z1–8, P
Z1@
Z1
Z1
L, P
Gulf of Marseilles, Mediterranean
Sicily, Mediterranean
New South Wales, Australia
unknown
Naples, Mediterranean
Plymouth, eastern Atlantic
Bourdillon-Casanova, 1960
Calafiore et al., 1991
Bruce, 1972
Lebour, 1925
Caroli, 1926
Lebour, 1949b
sixth zoea, pereiopods 4–5 were not much
longer than pereiopod 3. The first zoeal stage is
described in detail. For the subsequent stages,
only the main differences from the previous
stage are given.
First Zoea (Fig. 1).—Duration. 5–7 days.
BL. 1.82 (1.70–1.85) mm; CL. 0.24 (0.20–
0.25) mm.
Carapace (Fig. 1A, B). Flattened dorsoventrally; rostrum unarmed, reaching one-fourth
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 1, 2004
Fig. 1. First zoea of Conchodytes nipponensis (De Haan, 1844). A, dorsal view; B, carapace, lateral view; C, abdomen,
lateral view; D, antennule; E, antenna; F, mandibles; G, maxillule; H, maxilla; I, first maxilliped; J, second maxilliped; K,
third maxilliped; L, pereiopods 1–2. Scale bars ¼ 0.1 mm.
length of peduncle of antennule; anterior
dorsomedian papilla and pterygostomian spine
present; anteroventral or posteroventral denticles absent; antennal spine absent; eyes sessile.
Antennule (Fig. 1D). Peduncle unsegmented; inner flagellum with long plumose seta;
outer flagellum with 4 aesthetascs and plumose
seta.
YANG AND KO: ZOEAL STAGES OF CONCHODYTES NIPPONENSIS
Antenna (Fig. 1E). Peduncle with basal spine;
endopod rod-like, with long terminal plumose
seta and strong spine; scale 6-segmented, with
11 marginal setae, distolateral spine, and inner
papilla.
Mandibles (Fig. 1F). Palps absent. Left and
right mandibles with spine between molar and
incisor processes.
Maxillule (Fig. 1G). Coxal endite with subterminal seta and 4 terminal setae; basial endite
with 5 cuspidate setae; endopod segmented,
with terminal spine and terminal seta.
Maxilla (Fig. 1H). Coxal endite with 4 setae;
basial endite bilobed, each with 2 setae;
endopod with fine hairs on basal lobe and
terminal seta; scaphognathite with 5 marginal
plumose setae.
First maxilliped (Fig. 1I). Basis with 4 setae,
distolateral margin protuberant; endopod 3segmented, with 0, 1, 3þ1 setae; exopod with
4 terminal natatory setae symmetrically disposed in 2 pairs.
Second maxilliped (Fig. 1J). Coxa unarmed;
basis with tubercle and 2 setae; endopod 3segmented, with 0, 2, 4þ1 setae; exopod with 2
short subterminal setae and 4 terminal natatory
setae symmetrically disposed in 2 pairs.
Third maxilliped (Fig. 1K). Coxa unarmed;
basis with tubercle and 2 setae; endopod 3segmented, with 2, 2, 4þ1 setae; exopod with
short subterminal seta and 4 terminal natatory
setae symmetrically disposed in 2 pairs.
Pereiopods 1–2 (Fig. 1L). Rudimentary,
biramous as bud.
Abdomen (Fig. 1A, C). Flattened dorsoventrally; composed of 5 somites; abdominal
somite 6 not differentiated; junction of thorax
and abdomen flexed; somite 3 conspicuously
hump-backed; somite 1 with lateromedian setae;
somite 3 with 2 pairs of posterodorsal setae as
well as pair of marginal setae; somite 4 with tuft
of dorsal setae as well as pairs of marginal setae;
somite 5 with pair of posterodorsal setae; all
somites without spines; pleopods absent.
Telson (Fig. 1A). Subtriangular, with shallow
posteromedian concavity and 7þ7 posterior
setae; outermost 2 pairs of setae plumose only
on inner side; bases of all setae except outermost with row of minute spinules; anal spine
absent; uropods absent.
Chromatophores. Orange chromatophores interspersed with yellow present on superolateral
margin of each eye, carapace, labrum, basis of
third maxilliped, endopods of second and third
maxillipeds, exopods of second and third
maxillipeds, rudiments of pereiopods 1 and 2,
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abdominal somites 1–5 laterally, and abdominal
somites 3–5 dorsally. Yellow chromatophores
present on peduncle of antennule, scale of
antenna, and telson.
Second Zoea (Fig. 2).—Duration. 5–7 days.
BL. 1.96 (1.95–1.98) mm; CL. 0.29 (0.28–
0.30) mm.
Carapace (Fig. 2A). Supraorbital spine present; eyes stalked.
Antennule (Fig. 2B). Peduncle swollen basally, 2-segmented: first segment unarmed; second with 2 terminal setae.
Antenna (Fig. 2C). Endopod with long
plumose and 2 short terminal setae.
Mandibles (Fig. 2D). Left mandible with 2
spines between molar and incisor processes.
Maxillule (Fig. 2E). Basial endite with 6
cuspidate setae.
Maxilla (Fig. 2F). Scaphognathite with 7
marginal plumose setae.
First maxilliped (Fig. 2G). Unchanged.
Second maxilliped (Fig. 2H). Unchanged.
Third maxilliped (Fig. 2I). Endopod 4segmented, with 2, 1, 2, 4þ1 setae.
Pereiopod 1 (Fig. 2J). Coxa unarmed; basis
with 2 setae and tubercle; endopod 4-segmented, with 2, 1, 2, 2 setae; exopod with 2 short
subterminal setae and 4 terminal natatory setae.
Pereiopod 2 (Fig. 2K). Coxa unarmed; basis
with 2 setae and tubercle; endopod 4-segmented,
with 2, 1, 2, 2 setae; exopod with 2 short setae
and 4 terminal natatory setae.
Pereiopods 3–4 (Fig. 2L). Uniramous.
Abdomen (Fig. 2A). Somite 6 with 2 pairs of
dorsal setae.
Telson (Fig. 2A). Posterior margin with 8þ8
setae; outermost pair of setae plumose only on
inner side; uropods visible.
Third Zoea (Fig. 3).—Duration. 4–6 days.
BL. 2.09 (2.08–2.10) mm; CL. 0.34 (0.33–
0.35) mm.
Carapace (Fig. 3A, B). Anteroventral margin
bifid.
Antennule (Fig. 3C). Peduncle 2-segmented,
each with 7 plumose setae.
Antenna (Fig. 3D). Scale 4-segmented, with
12 plumose setae and terminal spine; endopod
3-segmented: first and second segments unarmed; third with 4 terminal setae.
Mandibles (Fig. 3E). Left and right mandibles
with 3 spines between molar and incisor
processes.
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Fig. 2. Second zoea of Conchodytes nipponensis (De Haan, 1844). A, dorsal view; B, antennule; C, antenna; D, mandibles;
E, maxillule; F, maxilla; G, first maxilliped; H, second maxilliped; I, third maxilliped; J, pereiopod 1; K, pereiopod 2; L,
pereiopods 3–4. Scale bars ¼ 0.1 mm.
YANG AND KO: ZOEAL STAGES OF CONCHODYTES NIPPONENSIS
115
Fig. 3. Third zoea of Conchodytes nipponensis (De Haan, 1844). A, dorsal view; B, carapace, lateral view; C, antennule;
D, antenna; E, mandibles; F, maxillule; G, maxilla; H, first maxilliped; I, second maxilliped; J, third maxilliped; K, pereiopod
1; L, pereiopod 2; M, pereiopods 3–4; N, telson and uropods. Scale bars ¼ 0.1 mm.
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 1, 2004
Maxillule (Fig. 3F). Basial endite with 7
cuspidate setae.
Maxilla (Fig. 3G). Unchanged.
First maxilliped (Fig. 3H). Unchanged.
Second maxilliped (Fig. 3I). Unchanged.
Third maxilliped (Fig.3J). Endopod 4-segmented, with 2, 1, 3, 4þ1 setae.
Pereiopod 1 (Fig. 3K). Endopod 4-segmented, with 2, 1, 3, 2 setae.
Pereiopod 2 (Fig. 3L). Endopod 4-segmented,
with 2, 1, 3, 2 setae.
Pereiopods 3-5 (Fig. 3M). Pereiopod 3
biramous as bud. Pereiopods 4 and 5 uniramous.
Abdomen (Fig. 3A). Composed of 6 somites.
Telson and uropods (Fig. 3N). Telson with
7þ7 posterior setae and pair of lateral spines.
Uropods free: endopod rudimentary; exopod
with 6 marginal plumose setae.
Fourth Zoea (Fig. 4).—Duration. 5–7 days.
BL. 2.17 (2.13–2.25) mm; CL. 0.35 (0.33–
0.38) mm.
Carapace (Fig. 4A). Unchanged.
Antennule (Fig. 4B). Peduncle 2-segmented,
with 12 and 7 plumose setae.
Antenna (Fig. 4C). Scale with 14 plumose
setae and terminal spine.
Mandibles (Fig. 4D). Unchanged.
Maxillule (Fig. 4E). Unchanged.
Maxilla (Fig. 4F). Unchanged.
First maxilliped (Fig. 4G). Unchanged.
Second maxilliped (Fig. 4H). Unchanged.
Third maxilliped (Fig. 4I). Unchanged.
Pereiopod 1 (Fig. 4J). Unchanged.
Pereiopod 2 (Fig. 4K). Endopod 4-segmented,
with 2, 1, 4, 2 setae.
Pereiopod 3 (Fig. 4L). Coxa unarmed; basis
with 2 setae and tubercle; endopod 5-segmented,
with 0, 0, 0, 2, 2 setae; exopod with short
subterminal seta and 4 terminal natatory setae.
Pereiopods 4–5 (Fig. 4M). Pereiopod 4
biramous. Pereiopod 5 unchanged.
Abdomen (Fig. 4A). Unchanged.
Telson and uropods (Fig. 4N). Telson
narrower, with 4þ4 marginal setae and pair of
lateral spines. Uropods developed: endopod
with 10 setae; exopod with 14 setae and
outermost tooth.
DISCUSSION
Of the descriptions of the pontoniid larvae
known so far, only three have been detailed:
P. flavomaculata described by Costanzo et al.
(1996), P. pinnophylax described by Calafiore et
al. (1991), and Periclimenaeus stylirostris
Bruce, 1969, described by Bruce and Coombes
(1995). However, the description of P. stylirostris, collected from Trepang Bay, Australia,
includes only the postlarva. Useful comparison
of early zoeas, therefore, is possible only
between C. nipponensis, P. flavomaculata,
and P. pinnophylax. The first zoea of the three
species can be distinguished from each other by
the length of the rostrum, the setation and
number of distal segments of the scale of the
antenna, the setation of the coxal endite of the
maxillule and the basial endite of the maxilla,
and the presence or absence of a tubercle on the
bases of the second and third maxillipeds (Table
2).
Gurney and Lebour (1941), in their revision
of Gurney’s (1936, 1938) grouping, separated
the larvae of the Pontoniinae into two groups,
ancylocaris and mesocaris, by the shape of the
body and the relative length of the pereiopods
4–5 to the pereiopod 3. The larvae of C.
nipponensis have the body double flexed at the
junction of the thorax and the abdominal somite,
and the abdominal somite 3. Furthermore,
pereiopods 4–5 are not much longer than
pereiopod 3 in the sixth zoea of C. nipponensis.
These characteristics reveal that the larvae of C.
nipponensis belong to Gurney and Lebour’s
(1941) mesocaris group. In the larvae of P.
flavomaculata and P. pinnophylax, Calafiore et
al. (1991) and Costanzo et al. (1996) provided
no available information on the shape of the
body and the relative lengths of pereiopods 4–5
to pereiopod 3. Therefore, it is unknown
whether P. flavomaculata and P. pinnophylax
belong to mesocaris group. However, other
characteristics such as the scale of the antenna
with inner papilla in the first zoea, the distolateral margin of the basis of the first maxilliped
protuberant, the dactyl of the endopod of the
second maxilliped with fringe of spinules, and
pereiopod 4 with the exopod from the sixth zoea
onward indicate that the two species probably
belong to the mesocaris group.
The systematic relationships between the
Pontoniinae and the Gnathophyllidae are confusing. The four genera Gnathophylloides,
Gnathophyllum, Levicaris, and Pycnocaris
were regarded as members of the palaemonid
subfamily Pontoniinae until Holthuis (1955)
transferred them to the separate family Gnathophyllidae. On the other hand, in his work on the
first zoea of
Gnathophyllum americanum
Guérin, 1856, Bruce (1986) regarded the
Gnathophyllidae as belonging to the Palae-
YANG AND KO: ZOEAL STAGES OF CONCHODYTES NIPPONENSIS
117
Fig. 4. Fourth zoea of Conchodytes nipponensis (De Haan, 1844). A, dorsal view; B, antennule; C, antenna; D, mandibles;
E, maxillule; F, maxilla; G, first maxilliped; H, second maxilliped; I, third maxilliped; J, pereiopod 1; K, pereiopod 2; L,
pereiopod 3; M, pereiopods 4–5; N, telson and uropods. Scale bars ¼ 0.1 mm.
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Table 2. Morphological differences between the first zoea of three species of the Pontoniinae.
C. nipponensis (present study)
Rostrum
Antenna scale
P. flavomaculata (Costanzo et al., 1996)
,1/2AP
6-segmented
(11 marginal setae, inner papilla,
distolateral spine)
P. pinnophylax (Calafiore et al., 1991)
¼1/2AP*
5-segmented
(12 marginal setae, inner papilla)
¼1/2AP*
5-segmented
(12 marginal setae, inner papilla)
Maxillule
coxal endite
1þ4 setae
1þ5 setae
1þ5 setae
Maxilla
basial endite
2þ2 setae
1þ2 setae
2þ2 setae
Maxilliped II
basis
2 setae, tubercle
2 setae
2 setae
Maxilliped III
basis
2 setae, tubercle
2 setae
2 setae
* ¼ data from figure; AP ¼ peduncle of antennule.
monidae and synonymized the former with the
Pontoniinae. Subsequently, despite larval similarity, Chace and Bruce (1993) concluded
tentatively that the unique mouthparts of the
Gnathophyllidae discarded the possibility of the
synonomy.
The presence or absence of posterior dorsomedian papilla on the carapace and outer seta or
lobule of the base of the maxillule, the number
of the endites of the maxilla, and the arrangement of distal natatory setae of the exopods of
the maxillipeds are important characters separating the first zoea of carideans at family level
(Gurney, 1938; Yokoya, 1957; Albornoz and
Wehrtmann, 1997). The first zoea of C.
nipponensis has a strong resemblance to that
of
G. americanum: (1) carapace without
posterior dorsomedian papilla; (2) base of
maxillule without outer seta or lobule, endopod
of maxillule segmented; (3) maxilla with three
endites; (4) bases of second and third maxillipeds with tubercle and two setae; and (5)
exopods of maxillipeds with four distal natatory
setae, symmetrically disposed in two pairs.
These similarities indicate that the Gnathophyllidae have great affinity with the Pontoniinae.
Moreover, the only differences that can be
observed in the first zoea of C. nipponensis and
Table 3. Morphological difference of the first zoea of C. nipponensis and G. americanum.
Rostrum
Antennule
outer flagellum
Antenna scale
C. nipponensis (present study)
G. americanum (Bruce, 1986)
,1/2AP
J1/2AP
4 aesthetascs, plumose seta
3 aesthetascs, plumose seta
6-segmented
(11 plumose setae, distolateral
spine, inner papilla)
4-segmented
(9 plumose setae, inner papilla)
Maxillule
coxal endite
1þ4 setae
0þ3 setae
Maxilla
basial endite
coxal endite
2þ2 setae
4 setae
1þ1 setae
3 setae
Maxilliped I
endopod
basis
3-segmented
4 setae
1-segmented
3 setae
Maxilliped II
endopod
0, 2, 4þ1 setae
0, 2, 4þ0 setae
Maxilliped III
endopod
2, 2, 4þ1 setae
2, 2, 3þ1 setae
AP ¼ peduncle of antennule.
YANG AND KO: ZOEAL STAGES OF CONCHODYTES NIPPONENSIS
G. americanum are usually found at the generic
or specific level of caridean larvae (Table 3).
Recently, Williamson and Rice (1996) and
Williamson (2001) proposed that similar larvae
do not necessarily signify related adults, and
therefore, the Pontoniinae and the Gnathophyllidae may be distinct. Further evidence from
molecular sequencing data as well as other
larvae of other gnathophyllid genera including
Gnathophylletum, which was recently erected
by d’Udekem d’Acoz (2001), will be necessary
to establish the systematic relationships between
the two groups.
ACKNOWLEDGEMENTS
The authors sincerely thank Dr. D. I. Williamson of Port
Erin Marine Laboratory, Isle of Man, U. K., for his critical
reading and suggestion of the manuscript. Drs K.-I. Hayashi,
J. N. Kim, J. Okuno, and Y. Hanamura are acknowledged
for providing valuable literature. This work was supported
by a grant No. R04-2001-00022 from the Korea Science and
Engineering Foundation.
LITERATURE CITED
Albornoz, L., and I. S. Wehrtmann. 1997. Descripción y
clave de los primeros estadı́os larvales de camarones
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RECEIVED: 18 February 2003.
ACCEPTED: 27 August 2003.