Journal of Oceanography, Vol. 57, pp.

535 to 548, 2001

Spatial Distributions of Phyllosoma Larvae (Crustacea:

Decapoda: Palinuridae and Scyllaridae) in Taiwanese
Waters
NARIAKI INOUE1, H IDEO S EKIGUCHI1 * and S HINN-PYNG YEH2
1
Faculty of Bioresources, Mie University, 1515 Kamihama-cho, Tsu, Mie 514-8507, Japan
2
Department of Aquaculture, National Pingtung University of Science and Technology,
Nei-Pu Hsiang, Pingtung, Taiwan 91207

(Received 20 November 2000; in revised form 19 March 2001; accepted 20 March 2001)

Distributions and taxonomy of phyllosoma larvae were examined in Taiwanese wa- Keywords:
ters, based on ichthyoplankton samples collected from May 1990 to July 1995. ⋅ Phyllosoma,
Phyllosoma larvae belonged to the two families Scyllaridae and Palinuridae repre- ⋅ Palinuridae,
senting 6 genera and 13 species. Of the collected phyllosoma larvae, those of Scyllarus ⋅ Scyllaridae,
⋅ spatial distribu-
and Panulirus species were most abundant, forming 90% of total numbers. Early
tions,
stage Scyllarus and Panulirus phyllosoma larvae were abundant in Taiwanese waters.
⋅ Taiwanese waters.
Middle to late stages (except the final stage) of Panulirus phyllosoma larvae were
absent from the waters throughout the year, while those of Scyllarus phyllosoma lar-
vae were collected in the waters. This suggests that all stages of Scyllarus phyllosoma
larvae may be retained in the northern part of the waters around northern Taiwan
while middle to late stages of Panulirus phyllosoma larvae may be flushed out from
the waters, the sub-final and final stages then possibly returning to the waters. An
anticlockwise eddy existed in the waters off northeastern Taiwan, which may be closely
related to flushing out and returning of Panulirus phyllosoma larvae through a much
longer planktonic period.

1. Introduction et al., 1988). However, few studies have been conducted

Phyllosoma larvae, confined to palinurid and
scyllarid lobsters, are plankton with transparent, leaf-like
forms (Phillips and McWilliam, 1986). The palinurid
phyllosoma larvae have a long life span of over 6 months
(Booth and Phillips, 1994), while those of scyllarids have
a comparatively short life span of 2 or 3 months
(Robertson, 1968; Ito and Lucas, 1990). Seventeen
palinurid and 16 scyllarid species have been reported in
Taiwan (Chan and Yu, 1993, 1995; Poupin, 1994; Chan
and Chu, 1996). Of the above lobsters, 21 species have
been described for phyllosoma larvae (Sekiguchi, 1986a).
Three Panulirus species (P. japonicus, P. longipes
and P. stimpsoni) are of economic importance in Taiwan
(Chan and Yu, 1993). Phyllosoma larvae of the first two
have been described (Sekiguchi, 1986a). Previous stud-
ies of the lobsters in Taiwan dealt with the aspects of ecol-
ogy (Deng, 1963), aquaculture (Jong and Lin, 1981), tax-
onomy (Chan and Yu, 1993) and fishery biology (Huang
* Corresponding author. E-mail: sekiguch@bio.mie-u.ac.jp
Copyright © The Oceanographic Society of Japan.
on the population dynamics of the lobsters in Taiwan,
particularly on the larval recruitment processes by which
the populations may form and be maintained in Taiwan-
ese waters.
Phyllosoma samples used in the present study were
collected during the ichthyoplankton surveys conducted
by the Kuroshio Edge Exchange Processes Project
(KEEP). This multidisciplinary study of the
biogeochemical cycle of carbon and associated elements
in the East China Sea revealed many of the sea’s
hydrographic features since 1989 (Chuang et al., 1995).
For example, Tang et al. (1999) reported that an anticlock-
wise eddy with a diameter of <70 km exists in the waters
off northeastern Taiwan (Fig. 1).
The goal of our future research is to clarify the lar-
val recruitment processes of commercially important spe-
cies of palinurid and scyllarid lobsters in Taiwan, par-
ticularly of Panulirus species, in relation to the
hydrographic features of Taiwanese waters. To our
knowledges, however, phyllosoma and puerulus (or nisto)
larvae in Taiwanese waters have not yet been dealt with,
except by Sekiguchi and Saisho (1994). So before begin-

535

Fig. 1. Mean circulation pattern at depths of 16, 104 and 200
m in August 1994 (modified from Tang et al., 1999). The
bathymetry is shown with thin depth contours at intervals
of 200 m from 200 to 1000 m.
ning work on the larval recruitment processes, it is first
of all necessary to identify to the species level phyllosoma
and puerulus (or nisto) larvae contained in
ichthyoplankton samples collected in Taiwanese waters,
and then to examine their spatial distribution.
2. Materials and Methods
Ichthyoplankton samples were collected using two
types of ring nets (diameter: 1.3 m and 1.5 m, length: 4.5
m, mesh: 0.5 mm). Samples were collected by surface
and oblique towing during the day or night. The first sam-
pling runs involved a total of 89 surface towings with a
ship speed of 2 kt for 10 min., samples being collected at
57 stations in the northern part of the waters around north-
536 N. Inoue et al.
Fig. 2. Map of the study area and location of sampling sta-
tions. a, b: Sampling stations for surface and oblique tow-
ing, respectively; solid circles: sampling stations in the
waters around northern Taiwan; open circles: sampling sta-
tions in the waters around southwestern Taiwan; dotted lines:
isopleths of 200 m depth.
ern Taiwan from May 1990 to July 1993, covering all
months (Table 1, Fig. 2a). The second runs involved a
total of 195 oblique towings at different depths (15–430
m) from the surface with a ship speed of 2–4 kt for 5
min., samples being collected at 41 and 11 stations in the
northern part of the waters around northern Taiwan and
southwestern Taiwan, respectively, from May 1990 to
May 1995 (Table 2, Fig. 2b). All samples were fixed in
5% formalin seawater immediately after sampling and
preserved in 70% ethanol.
Phyllosoma larvae were sorted from the samples in
the laboratory. Stages of these phyllosoma larvae were
determined according to Braine et al. (1979) and Phillips
and McWilliam (1986). The late stage phyllosoma larvae
 Table 1. Data for sampling phyllosoma larvae by surface towing.

Numerals in the upper table: number of towings; horizontal line: no data.

Numerals in the lower table: number of collected phyllosoma; horizontal line: no data.

were then identified to the species level according to
Sekiguchi (1986a) and Inoue et al. (2000). Based on
morphological features of the late stage phyllosoma lar-
vae identified, we identified the middle stage ones, but
the early stage ones were not identified to the species
level. Body length (BL) from the anterior margin of the
cephalon (forebody) to the posterior end of the hindbody,
cephalic length from the anterior to the posterior margins
of the cephalon, and widths of the cephalic and thorax
were measured to the nearest 0.1 mm. All figures were
drawn with the aid of a drawing tube.
The individual number of phyllosoma larvae col-
lected per net haul was very low, as indicated in Tables 1
and 2. Accordingly, we examined the spatial distributions
of phyllosoma larvae of different species by pooling the
data for the phyllosoma larvae collected by the two types

Spatial Distributions of Phyllosoma Larvae off Northern Taiwan 537

 Table 2. Data for sampling phyllosoma larvae by oblique towing.

Numerals in the upper table: number of towings; horizontal line: no data.

Numerals in the lower table: number of collected phyllosoma; horizontal line: no data.

538 N. Inoue et al.

of nets, and the phyllosoma larvae were counted as indi-
vidual numbers per tow for both types of net. Spatial dis-
tributions of phyllosoma larvae of the two genera
Scyllarus and Panulirus were examined on the basis of
samples collected by surface towing. Identification and
taxonomic descriptions were based on samples collected
by surface and oblique towings.

3. Results
A total of 312 phyllosoma larvae were collected, of
which 123 and 189 specimens were collected by surface
and oblique towing, respectively (Tables 1 and 2). These
phyllosoma larvae belonged to the two families
Scyllaridae and Palinuridae, representing 6 genera and
13 species.
Of the family Scyllaridae, 3 genera and 10 species
were identified (Tables 1 and 2): 7 species of the genus
Scyllarus (S. bicuspidatus, S. cultrifer, S. kitanoviriosus,
S. martensii, Scyllarus sp. c, Scyllarus sp. d, Scyllarus
sp. e), 2 species of the genus Ibacus (I. novemdentatus, I.
ciliatus) and 1 species of the genus Scyllarides
(Scyllarides sp.). Of the family Palinuridae, 3 genera and
3 species were identified (Tables 1 and 2): Panulirus
longipes or P. japonicus, Linuparus sordidus and Puerulus
angulatus. The phyllosoma larvae in stages I to IV of the
Scyllarus and in stages I to VI of the Panulirus were not
identified to the species level.

3.1 Spatial distributions

All stages of the Scyllarus phyllosoma larvae were
collected by surface towing (Table 1). The phyllosoma
larvae in the stages I to IV, which are probably identical
with those of S. kitanoviriosus, and also in the stages V
to VIII stages of S. kitanoviriosus, were mainly found in
the northern part of the waters around northern Taiwan
(Figs. 3a and b). No changes of spatial distributions ac-
cording to larval stages were detected for Scyllarus
phyllosoma larvae. This also holds for those of the other
Scyllarus species.
In contrast to Scyllarus phyllosoma larvae, no stage
VII and VIII phyllosoma larvae were collected of
Panulirus (Table 1). Phyllosoma larvae in stages I to IV
of the Panulirus were found in the northern part of the
waters around northern Taiwan, as in those of the
Scyllarus, while those in stages V and VI of the Panulirus
were found in the offshore waters (Figs. 4a and b). One
specimen of the final stage (stage IX) Panulirus
phyllosoma larvae was found in the coastal waters.
3.2 Phyllosoma larvae of the family Scyllaridae
a) Scyllarus in early to middle stages
Thirty-eight specimens in stages I to IV were col-
lected in the northern part of the waters around northern
Taiwan by surface towing (Table 1 and Fig. 3a): 4 speci-
Fig. 3. Distributions and abundance of phyllosoma larvae of
the Scyllarus found by surface towing. a: Early to middle
stage phyllosoma larvae of the Scyllarus; numerals with
open squares and triangles: individual number of stages I
to II and stages III to IV, respectively. b: Middle to late
stage phyllosoma larvae of Scyllarus kitanoviriosus; numer-
als with open squares and triangles: individual number of
stages V to VI and stages VII to VIII, respectively. Solid
circles: sampling stations. Dotted lines: isopleths of 200 m
depth.
mens in stage I (BL 1.1–1.3 mm), 9 in stage II (BL 1.6–
2.1 mm), 15 in stage III (BL 1.9–3.6 mm) and 10 in stage
IV (BL 2.5–5.0 mm). Most specimens of Scyllarus
phyllosoma larvae in stages V to VIII collected by sur-
face towing were identical with S. kitanoviriosus (Table
1 and Fig. 3b), so most of Scyllarus phyllosoma larvae in
stages I to IV may be identical with S. kitanoviriosus.
b) Scyllarus bicuspidatus
Nine specimens were collected in coastal waters off
Keelung and Tanshui of northern Taiwan and off
Kaohsiung of southwestern Taiwan (Tables 1 and 2): 7

Spatial Distributions of Phyllosoma Larvae off Northern Taiwan 539


Fig. 4. Distributions and abundance of Panulirus phyllosoma
larvae found by surface towing. a: Early to middle stage
phyllosoma larvae of the Panulirus; numerals with open
squares and triangles: individual number of stages I to II
and stages III to IV, respectively. b: Middle to late stage
phyllosoma larvae of the Panulirus; numerals with open
squares and solid squares: individual number of stages V to
VI and stage IX, respectively. Solid circles: sampling sta-
tions. Dotted lines: isopleths of 200 m depth.
specimens in stage V (BL 4.0–5.3 mm) and 2 in stage VI
(BL 6.7 mm and 8.9 mm). Phyllosoma larvae of this spe-
cies have often been reported in Japanese waters (Inoue
et al., 2000).
According to Sekiguchi’s (1986a) key for identifi-
cation, these phyllosoma larvae are identical with
Scyllarus sp. a, though Inoue et al. (2000) made clear
that Scyllarus sp. a, described by Sekiguchi (1986a), is
identical to S. bicuspidatus. The morphology of the
present specimens accords well with that of Scyllarus sp.
b, described by Phillips et al. (1981) and obtained from
the Indian Ocean.
540 N. Inoue et al.
c) Scyllarus cultrifer
Seven specimens of S. cultrifer were collected in
coastal waters off Keelung of northern Taiwan, Kaohsiung
of southwestern Taiwan, and off Iriomote Is. (Tables 1
and 2): 1 specimen in stage V (BL 6.8 mm), 2 in stage VI
(BL 7.2 mm and 10.0 mm), 2 in stage VII (BL 15.8 mm
and 18.4 mm) and 2 in stage VIII (BL 19.0 mm and 22.0
mm).
According to Sekiguchi’s (1986a) key for identifi-
cation, these phyllosoma larvae are identical with S.
bicuspidatus, though Inoue et al. (2000) made clear that
S. bicuspidatus described by Sekiguchi (1986a) is identi-
cal with S. cultrifer. The morphology of the present speci-
mens accords well with that of S. bicuspidatus, described
by Phillips et al. (1981) from the Indian Ocean, of
Scyllarus sp. a, described by Johnson (1971a) from the
South China Sea, and of S. bicuspidatus, described by
Sekiguchi (1990) from Mariana waters. Phyllosoma lar-
vae of this species have also been reported from Japa-
nese waters by Saisho et al. (1983), Sekiguchi (1986b),
and Inoue et al. (2000).
d) Scyllarus kitanoviriosus
Thirty-eight specimens were collected (Tables 1 and
2): 26 specimens in stage V (BL 5.4–7.9 mm), 9 in stage
VI (BL 8.1–9.8 mm), 1 in stage VII (BL 12.7 mm) and 2
in stage VIII (BL 17.3 mm and 19.7 mm). Many
phyllosoma larvae in stages V to VIII were collected in
the northern part of the waters around northern Taiwan
(Fig. 3b).
Higa and Saisho (1983) described late stage
phyllosoma larvae of S. kitanoviriosus which metamor-
phosed to the nisto stage in the laboratory. Most of the
Scyllarus phyllosoma larvae which were collected in the
present study may be identical with this species: Scyllarus
sp. from Japanese waters described by Shojima (1963),
Scyllarus sp. from the Arabian Sea described by Prasad
and Tampi (1960), Scyllarus sp. D from the South China
Sea described by Johnson (1971a) and Scyllarus sp. a from
the Southeastern Indian Ocean described by Phillips et
al. (1981). Phyllosoma larvae of the present species have
also been reported from Japanese waters by Saisho et al.
(1983), Wada et al. (1985) and Inoue et al. (2000).
e) Scyllarus martensii
Sixteen specimens were collected (Tables 1 and 2):
1 specimen in stage V (BL 3.4 mm), 10 in stage VI (BL
4.2–5.1 mm), 3 in stage VII (BL 7.0–9.0 mm) and 2 in
stage VIII (BL 9.4 mm and 10.8 mm). Phyllosoma larvae
in stages V and VI were collected off Keelung and Tanshui
of northern Taiwan, while those in sub-final and final
stages (stages VII and VIII) were collected in the north-
ern part of the waters around northern Taiwan.
Phillips and McWilliam (1986) described phyllosoma
larvae in first to final stages of this species from Austral-
ian waters. Phyllosoma larvae of the present species have
Fig. 5. Phyllosoma larva of Scyllarus sp. d (stage VI). A: ventral view; B: ventral view of cephalon; C: ventral view of abdomen;
D: dorsal view of abdomen; E: second maxilla, first and second maxillipeds; F: distal end of second maxilliped; G: first
maxilla. Scale bar 1.0 mm.
been reported from Japanese waters: Scyllarides sp. by
Shojima (1963) and S. martensii by Inoue et al. (2000).
They have also been reported from South African waters
as Scyllarides sp. by Gurney (1936), from the Indian
Ocean as Scyllarides sp. by Saisho (1966), and from the
South China Sea as S. martensii by Johnson (1971a).
f) Scyllarus sp. c
Four specimens in stage V (BL 6.1–10.7 mm) were
collected in coastal waters off Keelung and Tanshui off
northern Taiwan (Tables 1 and 2).
According to Sekiguchi’s (1986a) key for identifi-
cation, these phyllosoma larvae are identical with S.
cultrifer, though Inoue et al. (2000) made clear that these
are not identical with S. cultrifer phyllosoma. Phyllosoma
larvae of the present species have been reported from the
South China Sea as Scyllarus sp. B by Johnson (1971a),
from Hawaiian waters as Scyllarus sp. by Johnson
(1971b), from South African waters as Scyllarus species
D by Berry (1974), and from the Indian Ocean as S.
cultrifer by Prasad et al. (1975).
g) Scyllarus sp. d
This type of phyllosoma larva is here recorded for
the first time from Taiwanese and Japanese waters. Mor-
phological features of the present specimen are described
below.
Materials: One specimen, with BL of 16.9 mm, was
caught at 22°-34.2′ N, 120°-9.8′ E on November 15, 1994
(Table 2).
Spatial Distributions of Phyllosoma Larvae off Northern Taiwan
Morphological features (Fig. 5): Cephalic shield oblong,
11.5 mm long and 16.5 mm wide, with a low, stout spine
near each base of antennule peduncle on the dorsal sur-
face of cephalon. Postero-lateral margin of cephalic shield
overlapping the bases of a pair of second pereiopod. Tho-
rax 6.7 mm wide. Width ratio of cephalon to thorax 2.46.
Antennule longer than eyestalk, peduncle cylindrical with
2-segments. Antenna shorter than antennule, compressed
and not segmented, bears lateral process (directed later-
ally). First maxilla 2-lobed, anterior lobe with a vestigial
palp and 3 stout terminal spines with minute denticles,
posterior lobe with 2 long terminal setae. Second maxilla
with developed scaphognathite lacking seta. First
maxilliped short, bud-like. Second maxilliped 5-seg-
mented, slender without exopod. Third maxilliped 5-seg-
mented, slender without exopod, third maxilliped and all
pereiopods bear vental-directed coxal spins on the basal
segment. Fifth pereiopod uniramous, bud-like, and 2-seg-
mented. First to fourth pleopods bud-like, not segmented.
Uropod bud-like, not segmented, oval, not reaching be-
yond posterior margin of telson. Telson extends beyond
uropod, bears 2 prominent posteriorly directed lateral
spines.
Remarks: One VI stage phyllosoma larva was collected
in the present study. This phyllosoma larva is closely simi-
lar to that of S. bicuspidatus described by Phillips et al.
(1981), Scyllarus sp. a described by Johnson (1971a), and
S. bicuspidatus described by Sekiguchi (1990). However,
541
Fig. 6. Phyllosoma larva of Scyllarus sp. e (stage VIII, final-stage). A: ventral view; B: ventral view of cephalon; C: ventral view
of abdomen; D: dorsal view of abdomen; E: second maxilla, first and second maxillipeds; F: distal end of second maxilliped;
G: first maxilla. Scale bar 1.0 mm.
the cephalic shield of these phyllosoma larvae are more
slender than the present phyllosoma example. And these
phyllosoma larvae do not have vental-directed coxal
spines on the fifth pereiopod. Further, posterior spines
on the telson of these phyllosoma larvae are sharper than
the present phyllosoma specimen. The present phyllosoma
larva is distinguishable from the previously reported
phyllosoma larvae by (1) width ratios of cephalon to dif-
ferent parts of the body, (2) fifth pereiopod bearing vental-
directed coxal spines on the basal segment, and (3) the
shape of the telson.
h) Scyllarus sp. e
This type of phyllosoma larvae are here recorded for
the first time from Taiwanese and Japanese waters. Mor-
phological features of the present specimens are described
below.
Materials: Fourteen specimens were collected (Tables 1
and 2): 5 specimens in stage V (BL 4.2–5.2 mm), 4 in
stage VI (BL 5.3–5.8 mm), 4 in stage VII (BL 6.8–7.5
mm) and 1 in stage VIII (BL 9.8 mm). Phyllosoma larvae
in stages V and VI were collected in coastal waters off
Keelung and Tanshui of northern Taiwan, while those in
sub-final and final stages (stages VII and VIII) were col-
lected in the northern part of the waters around northern
Taiwan. We described the final stage phyllosoma (stage
VIII) with BL 9.8 mm, collected at 25°-33.3′ N, 122°-00′
E on May 26, 1993.
Morphological features (Fig. 6): Cephalic shield oval,
5.6 mm long and 5.4 mm wide, with a low, stout spine
542 N. Inoue et al.
near each base of antennule peduncle on the dorsal sur-
face of cephalon. Postero-lateral margin of cephalic shield
overlapping the bases of a pair of first pereiopod. Thorax
3.4 mm wide. Width ratio of cephalon to thorax 1.9.
Antennule about 2 times longer than eyestalk, peduncle
cylindrical with 2-segments. Antenna shorter than
antennule, compressed and not segmented, bears lateral
process (directed anteriorly). First maxilla 2-lobed, ante-
rior lobe with a vestigial palp and 3 stout terminal spines
with minute denticles, posterior lobe with 2 long termi-
nal setae. Second maxilla with developed scaphognathite
lacking seta. First maxilliped short, bud-like. Second
maxilliped 5-segmented, slender with bud of exopod.
Third maxilliped 5-segmented, slender without exopod
and vental-directed coxal spins on the basal segment. All
pereiopods bear vental-directed coxal spins on the basal
segment. Fifth pereiopod uniramous, bud-like, and 5-seg-
mented. First to fourth pleopods bud-like, oval, 2-lobed
with shallow cleft on inner margin of endopods. Uropods
enlarged bud-like, segmented, oval, 2-lobed with shal-
low cleft on lateral margin of endopod and exopod, a lit-
tle beyond posterior margin of telson. Telson tapered to-
ward a narrow truncated tip lacking seta.
Remarks: According to Sekiguchi’s (1986a) key for iden-
tification, these phyllosoma larvae are identical with
Scyllarus sp. a. However, the cephalic shield of the present
phyllosoma larvae are more rounded than the Scyllarus
sp. a ones, and eyestalk is much shorter than the Scyllarus
sp. a ones. So, the present phyllosoma larvae are distin-
Fig. 7. Phyllosoma larva of Panulirus japonicus or Panulirus longipes (stage IX, final stage). A: ventral view; b: ventral view of
cephalon; C: ventral view of abdomen; D: dorsal view of abdomen; E: second maxilla, first and second maxillipeds; F: distal
end of second maxilliped, G: first maxilla. Scale bar 1.0 mm.
guishable from Scyllarus sp. a by (1) the shape of the
cephalic shield, and (2) the relative length of the eyestalk.
i) Ibacus ciliatus
Six specimens were collected in the northern part of
the waters around northern Taiwan (Tables 1 and 2): 4
specimens in stage III (BL 6.5–6.8 mm), 1 in stage V (BL
12.5 mm), 1 in stage VI (BL 20.5 mm). Phyllosoma lar-
vae of this species have been reported from Japanese
waters by Shojima (1963) and Tokioka and Harada (1963).
Phyllosoma larvae reared in the laboratory were described
by Dotsu et al. (1966a, b) and Takahasi and Saisho (1978).
j) Ibacus novemdentatus
Four specimens were collected, mainly in coastal
waters off Keelung to the north of Taiwan (Tables 1 and
2): 3 specimens in stage I (BL 2.8–3.1 mm), 1 in stage III
(BL 4.2 mm). Phyllosoma larvae of this species have been
reported from the South China Sea by Johnson (1971a)
and Shojima (1973). Phyllosoma larvae reared in the labo-
ratory were described by Dotsu et al. (1966a, b) and
Takahasi and Saisho (1978).
k) Scyllarides sp.
Four specimens were collected mainly off Keelung
to the north of Taiwan (Tables 1 and 2): 1 specimen in
stage II (BL 2.5 mm), 3 in stage IV (BL 3.3–4.4 mm).
Two adult specimens of the genus Scyllarides (Scy.
haanii and Scy. squammosus) have been reported in the
northern part of the waters around northern Taiwan (Chan
and Yu, 1993). We have several reports for morphologi-
cal features of Scy. squammosus phyllosoma larvae
Spatial Distributions of Phyllosoma Larvae off Northern Taiwan
(Michel, 1968; Johnson, 1971b, 1977; Saisho and Sone,
1971), but none for Scy. haanii phyllosoma ones. Fur-
ther, as all phyllosoma larvae of the genus Scyllarides
collected in this study were in early and middle stages
(stages II and IV), it was impossible to identify these
phyllosoma larvae to the species level.
3.3 Phyllosoma larvae of the family Palinuridae
a) Panulirus in early to middle stages
Forty six specimens were collected by surface tow-
ing (Table 1): 6 specimens in stage I (BL 1.5–1.6 mm),
11 in stage II (BL 2.0–2.5 mm), 18 in stage III (BL 2.2–
4.5 mm), 4 in stage IV (BL 3.6–5.7 mm), 5 in stage V
(BL 6.6–9.7 mm) and 2 in stage VI (BL 9.1 mm and 9.2
mm). Phyllosoma larvae in stages I to IV were collected
in the northern part of the waters around northern Taiwan
(Fig. 4a), while those in stages V and VI were collected
only in offshore areas of the above waters (Fig. 4b). Fur-
ther, phyllosoma larvae in stages VII and VIII were not
collected by surface towing (Table 1).
b) Panulirus japonicus or Panulirus longipes
Little is known about the morphological features of
P. japonicus phyllosoma larvae that were collected at sea.
The features of the present specimen are described be-
low.
Materials: One specimen with BL of 31.0 mm was caught
at 25°-33′ N, 122°-00′ E on May 26, 1993 (Table 1 and
Fig. 4b).
Morphological features (Fig. 7): Cephalic shield oval,
543
Fig. 8. Phyllosoma larva of Linuparus sordidus (stage IV?). A: ventral view; B: ventral view of cephalon; C: ventral view of
abdomen; D: dorsal view of abdomen; E: second maxilla, first and second maxillipeds; F: distal end of second maxilliped;
G: first maxilla. Scale bar 1.0 mm.
12.8 mm long and 7.7 mm wide, with a low, stout spine
near each base of antennule peduncle on the dorsal sur-
face of cephalon. Poslero-lateral margin of cephalic shield
overlapping the bases of a pair of third maxilliped. Tho-
rax 9.5 mm wide. Width ratio of cephalon to thorax 0.81.
Antennule damaged. Antenna elongate, cylindrical with
5-segments, peduncle longer than half of eyestalk. First
maxilla 2-lobed, anterior lobe with a vestigial palp and 3
stout terminal spines with minute denticles, posterior lobe
with 2 long terminal setae. Second maxilla with devel-
oped scaphognathite bearing seta. First maxilliped short,
bud-like. Second maxilliped 5-segmented, slender with
elongate bud of exopod bearing 4 pairs of setae. Third
maxilliped 5-segmented, slender with exopod (found
loose). Third maxilliped and all pereiopods with gill buds
and without subexopodal and coxal spines. Third and
fourth pereiopods with natatorial exopods, first and sec-
ond pereiopods were damaged and that exopods were
found loose. Fifth pereiopod elongate, uniramous with 5-
segments. Abdomen 5-segmented with notch on mid-line
of dorsal surface. First to fourth pleopods bud-like, oval,
2-lobed with shallow cleft on inner margin of endopods.
Uropods enlarged bud-like, segmented, oval, 2-lobed with
shallow cleft on lateral margin of endopod and exopod,
reaching a little beyond posterior margin of telson. Telson
tapered toward a narrow truncated tip lacking seta bud
with shallow cleft on lateral margin.
Remarks: One final stage phyllosoma (stage IX) was
collected in the present study. According to Sekiguchi’s
544 N. Inoue et al.
(1986a) key for identification, the present phyllosoma is
identical with those of P. japonicus group, i.e., P.
japonicus or P. longipes. Late stage phyllosoma larvae of
P. longipes have been described by Michel (1969),
Johnson (1971a), Prasad et al. (1975), Sekiguchi (1990),
and Matsuda and Yamakawa (2000), while those of P.
japonicus have been described by Murano (1971) and
Inoue (1981). Based on the morphological features re-
ferred to in the above studies, it is impossible to identify
the present phyllosoma.
c) Linuparus sordidus
Little is known about morphological features of
Linuparus phyllosoma larvae. This type of phyllosoma
larva is here recorded for the first time from Taiwanese
and Japanese waters. The features of the present speci-
mens are described below.
Materials: Three specimens were collected in the present
study (Tables 1 and 2): 1 specimen in stage III? (BL 2.7
mm), 2 in stage IV? (BL 3.4 mm and 4.8 mm). We have
described a specimen in the stage IV? with BL 4.8 mm,
collected at 26°-00′ N, 122°-54′ E on June 20, 1990.
Morphological features (Fig. 8): Cephalic shield oval,
3.5 mm long and 2.2 mm wide, with a low, stout spine
near each base of antennule peduncle on the dorsal sur-
face of cephalon. Postero-lateral margin of cephalic shield
overlapping the bases of a pair of third maxilliped. Tho-
rax 2.1 mm wide. Width ratio of cephalon to thorax 1.05.
Antennule longer than eyestalk, peduncle cylindrical with
2-segments. Antenna shorter than antennule, not seg-
mented, bears lateral process (directed anteriorly). First
maxilla 2-lobed, anterior lobe with a vestigial palp and 2
stout terminal spines with minute denticles, posterior lobe
with 2 long terminal setae. Second maxilla with devel-
oped 4 long setae. First maxilliped short, bud-like. Sec-
ond maxilliped 5-segmented, with bud of exopod. Third
maxilliped 5-segmented, slender with exopod. Third
maxilliped and all pereiopods with vental-directed coxal
spines on basal segment. First to third pereiopods with
natatorial exopods, damaged. Fourth pereiopod not seg-
mented with bud-like exopod (non-setose). Fifth
pereiopod bud-like. Abdomen without pleopods and
uropods.
Remarks: These phyllosoma larvae belong to the family
Palinuridae since they bear setose exopod on the third
maxilliped. Five genera (Justitia, Linuparus, Palinustus,
Panulirus and Puerulus) have been reported for the
Palinuridae in Taiwanese waters (Chan and Yu, 1993),
and morphological features of phyllosoma larvae of the
above 5 genera have been described by Sekiguchi (1986a).
Phyllosoma larvae collected in the present study have
setose exopod on the third maxilliped and antennae bear
lateral process directed anteriorly (Fig. 8). Judging from
these morphological features, these phyllosoma larvae do
not belong to Justitia, Palinustus, Panulirus, and
Puerulus, but to Linuparus. Two species of the Linuparus
(L. trigonus and L. sordidus) have been reported in Tai-
wanese waters (Chan and Yu, 1993). Johnson (1971a)
described middle stage phyllosoma larvae of Linuparus
sp. from the South China Sea, which Sekiguchi (1986a)
identified as L. trigonus. Further, Ikeda and Oka (1974)
described first stage phyllosoma larvae of L. trigonus
reared in the laboratory. However, morphological features
of L. sordidus phyllosoma larvae have not yet been re-
ported. The present phyllosoma larvae are distinguish-
able from Johnson’s (1971a) phyllosoma as follows: the
cephalic shield of Johnson’s (1971a) phyllosoma is al-
most triangular while the present phyllosoma is oval. The
present phyllosoma larvae may be identical with L.
sordidus.
d) Puerulus angulatus
One specimen in stage V with BL of 6.4 mm was
collected in coastal waters off Tanshui, northern Taiwan
(Table 2). Phyllosoma larvae of this species have been
reported from the South China Sea by Johnson (1968,
1971a), from the Indian Ocean by Prasad et al. (1975),
from the western North Pacific adjacent to Taiwan
(Sekiguchi and Saisho, 1994) and from Tongan waters
by Sekiguchi et al. (1996).
4. Discussion
The Kuroshio Current (KC) enters the East China
Sea (ECS) to the northeast of Taiwan. It is separated into
two parts, the mainstream and its branch, due to the block-
Spatial Distributions of Phyllosoma Larvae off Northern Taiwan
ing action of the shelf of the southern ECS, which runs
east-west (Chern et al., 1990; Hseuh et al., 1992, 1993;
Tang and Yang, 1993; Tang et al., 1999). The main stream
flows along the shelf break and then turns to the east,
while the branch current flows to the northwest, carrying
the saline and warm Kuroshio water onto the shelf of
southern ECS. An anticlockwise circulation of <70 km in
diameter was observed southwest of the branch current
(Fig. 1), coinciding with the observed cold dome/eddy.
This cold eddy, which marks the upwelling of the intruded
subsurface Kuroshio water, has been frequently observed
over the shelf break of the ECS to the north of Taiwan
(Lin et al., 1992; Liu et al., 1992a, b; Yeh, 1992, 1994).
In contrast to the eastward flow of the mainstream, exist-
ing as a strong flow in the upper 200 m depth, there is a
southwestward to southeastward countercurrent that usu-
ally spans over the whole water column just offshore of
the northern tip of Taiwan, which is identified as a part of
the eddy (Fig. 1) (Tang et al., 1999). Although this eddy
and the countercurrent exist throughout the year, it is sig-
nificant only in summer and disappears above 150 m depth
in winter.
Many specimens of Scyllarus phyllosoma larvae in
first to final stages (stages I to VIII), most of them being
identical with S. kitanoviriosus, were collected in the
northern part of the waters around northern Taiwan (Ta-
ble 1 and Figs. 3a and b). Two alternative explanations
are possible for the spatial distributions of the Scyllarus
phyllosoma larvae: first, the phyllosoma larvae are re-
leased in the northern part of the waters around northern
Taiwan and are retained within the water through their
larval life. Second, they are released elsewhere and later
transported into the northern part of the waters around
northern Taiwan. According to Robertson (1968) and Ito
and Lucas (1990), who cultured S. americanus and S.
demani from eggs to juveniles, it took less than 20 days
from egg hatching to stage IV for the above two species.
Judging from this short period for egg hatching to stage
IV, it is assumed that the Scyllarus phyllosoma larvae in
stages I to IV may be released in the northern part of the
waters around northern Taiwan, being retained within the
waters through the larval life, although we have little in-
formation about the occurrence and abundance of juve-
niles and adults of the Scyllarus species in Taiwanese
waters, except the report by Chan and Yu (1986, 1993).
Many specimens of Panulirus phyllosoma larvae in
stages I to IV were collected in the northern part of the
waters around northern Taiwan, as in Scyllarus
phyllosoma larvae (Figs. 3a and 4a). According to Inoue
(1981), Kittaka and Kimura (1989) and Matsuda and
Yamakawa (2000), who succeeded in culturing P.
japonicus and P. longipes from eggs to juveniles, it took
less then 14 days from egg hatching to stage II for the
above two species. Judging from this short period for egg
545
hatching to stage II, it is assumed that Panulirus
phyllosoma larvae in stages I to II may be released within
the northern part of the waters around northern Taiwan,
as in Scyllarus phyllosoma larvae. Since juveniles and
adults of three Panulirus species (P. japonicus, P. longipes
and P. stimpsoni), which are target species for fisheries,
are common and abundant in the northern part of the
waters around northern Taiwan (Chan and Yu, 1993), we
may deduce that most Panulirus phyllosoma larvae in
stages I to VI collected in the northern part of the waters
around northern Taiwan may be identical with one of the
above three species.
In contrast to Scyllarus phyllosoma larvae and stages
I to VI Panulirus larvae, the Panulirus phyllosoma lar-
vae in stages VII and VIII were not collected in the north-
ern part of the waters around northern Taiwan, despite
sampling being done throughout the year. This suggests
that Panulirus phyllosoma larvae in VII and VIII stages
may not be retained within the northern part of the wa-
ters around northern Taiwan, and those in the final stage
(stage IX) may be transported into the water from else-
where.
Two alternative explanations are possible for differ-
ences in spatial distribution between phyllosoma larvae
of the Scyllarus and the Panulirus: first, Scyllarus
phyllosoma larvae of all stages and Panulirus ones in
stages I to VI may be found in shallower depths, so these
may be retained within the anticlockwise eddy off north-
eastern Taiwan, as indicated in Fig. 1. On the other hand,
Panulirus larvae in stages VII and VIII may be found at
deeper depths, so these may flush out elsewhere due to
the eastward flow in the northern part of the water around
northern Taiwan. However, this explanation is not sup-
ported in this case, based on Rimmer and Phillips (1979)
and Phillips et al. (1981), who reported that Scyllarus and
Panulirus phyllosoma larvae in middle to late stages are
mainly found during daytime at depths of 80–100 m and
50–120 m, respectively. Alternatively, Panulirus
phyllosoma larvae have a much longer larval period of 6
months or more (Kittaka and Kimura, 1989; Booth and
Phillips, 1994; Matsuda and Yamakawa, 2000) while
Scyllarus larvae have a shorter period of at most 3 months
(Robertson, 1968; Sekiguchi, 1986b; Ito and Lucas, 1990),
so that Panulirus larvae in stages VII and VIII may flush
out elsewhere due to the eastward flow in the northern
part of the waters around northern Taiwan, and those in
the final stage (stage IX) may be transported into the
northern part of the waters around northern Taiwan,
though no detailed mechanism for this is immediately
apparent. We have some reports indicating that Scyllarus
phyllosoma larvae were found in abundance within the
coastal waters while only a few Panulirus larvae were
found (Rimmer and Phillips, 1979; Phillips et al., 1981;
Sekiguchi, 1986b; Inoue et al., 2000). Furthermore,
546 N. Inoue et al.
Sekiguchi (1986b) and Inoue et al. (2000) suggest that
Scyllarus phyllosoma larvae may be retained within Japa-
nese coastal water north of the Kuroshio throughout their
larval life, while Panulirus larvae in middle stages may
be transported into the Kuroshio-Counter Current gyre.
Acknowledgements
The authors would like to express their hearty thanks
to the captain and crew of the vessels for the sampling on
board.
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