2006 larval molting and growth of the japanese spiny lobster panulirus japonicus under laboratory conditions

Blackwell Publishing AsiaMelbourne, AustraliaFISFisheries Science0919 92682006 Blackwell Science Asia Pty LtdAugust 2006724767773Original ArticleLarval development of
Panulirus japonicus
H Matsuda and T Takenouchi

FISHERIES SCIENCE

2006; 72: 767–773

Larval molting and growth of the Japanese spiny lobster
Panulirus japonicus under laboratory conditions
Hirokazu MATSUDA* AND Taisuke TAKENOUCHI
Fisheries Research Division, Mie Prefectural Science and Technology Promotion Center, 3564-3 Hamajima,
Shima Mie 517-0404, Japan

ABSTRACT: Ten newly hatched phyllosoma of Panulirus japonicus were cultured individually to
monitor body length (BL) and intermolt period, and 2000 were cultured in groups to sample specimens
for measurement of body weight. Phyllosoma were fed with Artemia and mussel gonad; the culture
seawater temperature was 24–26°C. The individually cultured phyllosoma showed an increment in
body length by the first molt of approximately 0.5 mm, and the molt increment increased to approximately 1 mm at 5 mm BL; it was constant to 15 mm BL. Thereafter, the molt increment increased
exponentially. The duration of the first instar was 6–7 days. Instar duration increased with development
up to approximately 2 weeks at the 20th instar (∼16 mm BL) and then became constant. Of the 10 larvae reared individually, five metamorphosed to the puerulus stage. The entire phyllosoma life ranged
from 245–326 days (mean 289.0 days), and the number of instars ranged from 22–29 (mean 26.2).
Body length in the final instar ranged from 28.50–33.10 mm (mean 30.280 mm). For the phyllosoma
cultured in groups, relationships between BL and wet/dry body weights (WW/DW, mg) were
expressed as exponential equations: WW = 0.0686BL2.2023 and DW = 0.0209BL2.1905.
KEY WORDS: Body weight, growth, Panulirus japonicus, phyllosoma, rearing.

INTRODUCTION
The Japanese spiny lobster Panulirus japonicus
occurs in shallow rocky areas of the north-western
Pacific.1 In Japan, the lobster plays an important

role in coastal fisheries because of its high economic value.2 Hence, the ecology, stock management and aquaculture technology have long
been studied, and some measures to manage
and enhance the natural population have been
proposed.2,3
Panulirus japonicus has a pelagic larval stage
(phyllosoma) in its early life history as do scyllarids
and other palinurids. The phyllosoma stage has
been the target of research because an understanding of the variation in recruitment to the fishing
ground is crucial for establishment of stock-management strategies for the lobster.4–6 The number of
phyllosoma that had been caught in the ocean,
however, was too small, and little information
was available on their development, distribution
and subsequent recruitment. Recently, Yoshimura

*Corresponding author: Tel: 81-5-9953-0130.
Fax: 81-5-9953-2225. Email:
Received 24 October 2005. Accepted 26 January 2006.

et al.7 successfully collected many later-stage phyllosoma off the south coast of Kyusyu Island and
demonstrated that the late-stage larvae were dispersed widely in and south of the Kuroshio Current. In addition, Sekiguchi8 and Sekiguchi and
Inoue9 advanced hypotheses on larval recruitment
processes by re-examining plankton samples collected to survey the distribution of ichthyoplankton. These studies provide important information
to elucidate the recruitment processes.
Research on larval culture of P. japonicus has
been conducted both to produce a large number of
juveniles for use in aquaculture, and to ascertain
ecologic and ethological aspects of larvae.10–12 Thus
far, outlines of phyllosomal development have
been reported from larval cultures in the laboratory. Under laboratory conditions, the entire length
of the phyllosoma stage ranged from 231–417 days

(n = 136, 24–27°C) and the total number of molts
from hatch to the puerulus stage ranged from 20–
31.11 The molt increment was constant at about
1 mm up to approximately 18 mm body length
(BL), and then this increment increased exponentially.10 However, exhaustive observations on larval
development throughout the phyllosoma life have
not yet been made; a better understanding of larval
development is necessary to interpret behavioral


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H Matsuda and T Takenouchi

traits in the ocean and to determine adequate
culturing conditions, which would alter as larvae
develop.
In the present study, phyllosoma of P. japonicus
were cultured individually from hatch to the puerulus stage in order to monitor their growth. Phyllosoma were also mass-cultured to obtain samples
for measurements of body weight. The present
paper reports the growth and the change in body
weight under laboratory conditions for P. japonicus
phyllosoma, from hatch to the puerulus stage.
MATERIALS AND METHODS
Individual culture
An ovigerous female of P. japonicus was collected
off Shima (34°17′Ν, 136°49′E), Mie Prefecture, on 5
June 1997 using tangle nets. On the day of capture,

the lobster was transferred to Mie Prefectural
Science and Technology Promotion Center. It was
maintained at ambient temperature (20–24°C) in a
flow-through tank until hatching occurred. Seawater was supplied after it was filtered through sand.
The lobster was fed daily with the mussel Mytilus
galloprovincialis and frozen krill.
Hatching occurred on 20 July 1997. Of several
hundred thousand newly hatched phyllosoma, 10
were used for individual culture. The culture methods were similar to those used for larval Panulirus
longipes.13 Each larva was kept in a 120-mL glass
cup until 100 days after hatching, after which each
was cultured in a 400-mL glass cup. The first to
fourth instar larvae were fed Artemia nauplii
(∼0.6 mm BL) exclusively. From the fifth instar
onward, they were fed Artemia cultured with the
diatom Phaeodactylum tricornutum and finely
minced mussel gonads. The size of Artemia given
gradually increased up to approximately 6 mm BL
as the phyllosoma developed; accordingly, the
density of Artemia decreased from 2–0.3 individual/mL. The size of the pieces of mussel gonad
given also increased (from ∼1–4 mm3) as the larvae
grew. About 10 pieces of mussel gonad were placed
in each culturing vessel. Artemia and mussel gonad
were replaced daily during culture.
The seawater temperature was maintained at
26.0°C until 130 days after hatching; it was gradually decreased to 24.0°C over 2 weeks and then
maintained at 24.0°C until the end of the culture,
according to Matsuda and Yamakawa.10 The larvae
were exposed to a constant artificial light–dark
cycle (lighting from 08:30–20:30 hours) by fullspectrum fluorescent bulbs equipped with an electric timer. The light intensity during the light phase

was about 5 μE/m2 per s.

Fig. 1 Body length (BL) of Panulirus japonicus phyllosoma, from the anterior margin of the cephalic shield
between the eyestalks to the posterior end of the
abdomen.

During the individual culture, the intermolt
period and BL were monitored regularly for each
phyllosoma and puerulus. Body length of the phyllosoma refers to the length from the anterior margin of the cephalic shield between the eyestalks to
the posterior end of the abdomen (Fig. 1). Body
length of the puerulus was measured from
between the supraorbital horns to the posterior
margin of the telson. Measurements of BL were
made every 2–7 days after molting. To measure BL,
each larva was transferred from the culture vessel
to a 12-cm diameter laboratory dish with a small
amount of seawater using a small ladle. Body
length was then measured with a profile projector
(V-12A, Nikon, Tokyo, Japan) after the sea water
had been poured out by tipping the dish. Finally,
the larva was returned to the same vessel. During
the measurement process, special care was taken


Larval development of Panulirus japonicus

FISHERIES SCIENCE

to avoid damaging the larva. The larval culture
continued until all of the larvae had either died or

metamorphosed to the puerulus stage.

As the wet and dry weights increased linearly
with increasing days after hatching during the
first to fifth phyllosoma instar, the relationships
between days after hatching (D) and wet/dry body
weights (WW/DW, mg) could be described by:

Mass culture
A total of 2000 phyllosoma hatched on 27 July 1998
from a female lobster that had been captured on 8
June 1998 off Shima were used in the mass culture.
They were placed in two 40-L specialized acrylic
hemispherical tanks11 and cultured in a flowthrough system at a water flow rate of 20–80 L/h.
The seawater temperature was controlled at a similar temperature as for the individual culture using
a unit to supply constant-temperature seawater
(Aquatron-portable APS-206A, Koito Industries,
Yokohama, Japan). Food items were also similar to
those used in the individual culture. The density of
Artemia fed was lower in the mass culture than in
the individual culture: 1.0 individual/mL with nauplii and 0.1 individual/mL with larger Artemia.
Approximately 50–200 pieces of minced mussel
gonad were prepared and fed once daily for each
group tank. Uneaten foods were removed before
feeding. Artemia were removed by changing
screens with 0.25-mm mesh attached around a
drainpipe, to those with 1- or 3-mm mesh that
ensured escape of Artemia. Uneaten mussel
gonads were siphoned. The cultures were transferred to clean tanks twice per week.
Until the sixth instar, all larvae molted within 2–

4 days and distinct morphological changes were
observed with ecdyses. This enabled sampling,
from the first to fifth instar, of larvae for measurements of body weight 2–4 times within each instar.
For the first to fifth instar, 10–50 larvae were collected in triplicate at each sampling, then the total
wet weight of each group was measured and converted to individual weight. Beyond the fifth instar
it was impossible to determine the number of the
instar for each larva because of the large individual
differences in BL, intermolt period and morphology, and larvae were consequently collected once
or twice every 2 weeks. From the sixth instar
onward, larval weights were measured individually.
The larvae were placed in sea water without food
for 2 h prior to measurement; the larvae were then
placed on filter paper and rinsed with 3.5% ammonium formate to remove saline matter.14 They were
transferred to a preweighed aluminum sheet with
tweezers, and the wet body weight was measured.
Dry body weight was measured after the samples
were dried at 60°C for 24 h and placed in a desiccator at an ambient temperature for 1–2 h. Wet and
dry body weights were measured with an autobalance (1712MP8, Sartorius, Göttingen, Germany).

769

WW or DW = aD + b
where a and b are constants. However, because
the difference in BL between individuals became
larger beyond the fifth instar and it was no longer
appropriate to describe the relationships between
days after hatching and body weights, increases in
body weights were expressed relative to BL by the
following power function:
WW or DW = c × BLd

where c and d are constants. The constants a, b,
c and d can be estimated by the least-squares
method using MS Excel (Microsoft, Redmond, WA,
USA).
The survival rate (Su) in the mass culture was
calculated by excluding the number of larvae sampled for measurements of body weight as follows:
n -1

Su (%) = ’ Si,
i =0

where Si represents the survival rate from the ith to
the (i + 1)th sampling.
RESULTS
Individual culture
Of the 10 phyllosoma cultured, five died in the
course of the culture and five reached the puerulus
stage. Mortalities were found on days 75, 126, 129,
239 and 318 after hatch (Fig. 2). The causes of mortality were bacterial disease (n = 3), a symptom of
which was cloudiness of the mid-gut gland and
hind-gut, and complications in molting (n = 2). The
five phyllosoma that reached the puerulus stage
metamorphosed after 22–29 molts (mean 26.2
molts). Their phyllosoma lifetimes ranged from
245–326 days (mean 289.0 days), and BL in the final
phyllosoma instar ranged from 28.50–33.10 mm
(mean 30.280 mm).
The BL of phyllosoma was 1.55 ± 0.01 mm
(mean ± SD, n = 10) for the first instar. It then
increased linearly with moltings: 7.69 ± 0.39 mm

for the 10th instar (n = 10), 16.30 ± 2.54 mm for the
20th instar (n = 7) and 20.95 ± 3.16 mm for the 25th
instar (n = 4) (Fig. 3). There were three phases in
the relationship between BL before a molt and
increment in BL by the next molt (phases A, B and
C, Fig. 4). The molt increment increased gradually


FISHERIES SCIENCE

770

H Matsuda and T Takenouchi

1.5

100

1.4

60
40

Metamorphoses to the
puerulus stage

20
0
0

50

100

150

200

250

300

350

Days after hatching

Growth index (BLn+1/BLn)

Survival rate (%)

80

1.3

1.2

1.1

Fig. 2 Survival of Panulirus japonicus phyllosoma in
the individual culture with a still-water system.

1.0
0

Body length (mm)

40

10

35

20

30

Body length (BLn, mm)

30
25

Fig. 5 Growth index of the increase in body length from
one instar (n) to the next (n + 1) for Panulirus japonicus
phyllosoma cultured individually in a still-water system.
Solid lines are drawn through the data points as visual
aids.

20
15
10

5
0
1

3

5

7

9

11

13

15

17

19

21

23

25

27

29

Instar

Molt increment in body length (mm)

20
15
10
5
0

Phase C

6

Intermolt period (days)

25

Fig. 3 Changes in body length with development of
Panulirus japonicus phyllosoma cultured individually in
a still-water system. Means (᭹) and ranges (vertical bars)
of body length are shown.

1

3

5

7

9

11

13

15

17

19

21

23

25

27

Instar

5

Phase B

4

Fig. 6 Changes in intermolt period with development
of Panulirus japonicus phyllosoma cultured individually
in a still-water system. Means (᭹) and ranges (vertical
bars) of body length are shown.

Phase A

3
2
1
0
0

5

10

15

20

25

30

35

Body length (mm)

Fig. 4 Relationship between body length (BL) before a
molt and increment in BL by the next molt for Panulirus
japonicus phyllosoma cultured individually in a stillwater system. Three phases (A, B and C) were recognized. Solid lines are drawn through the data points as
visual aids.

from approximately 0.5 mm (at ∼1.5 mm BL at
hatch) to about 1 mm (at ∼5 mm BL, phase A), then
it was constant at approximately 1 mm (until
∼15 mm BL, phase B). Beyond approximately

15 mm BL, it increased exponentially and at the
same time the variability in molt increment
became larger as the larvae grew (phase C). The
growth index of increase in BL from one instar (n)
to the next (n + 1), calculated as BL(n+1)/BLn, ranged
initially from 1.29 to 1.41 (mean 1.348, n = 10); it
decreased with development up to approximately
15 mm BL, and then increased gradually (Fig. 5).
The instar duration increased from approximately
1–2 weeks until the 20th instar; thereafter it
became constant at 2 weeks (Fig. 6).
Figure 7 shows the relationships between BL
and days after hatching for the two animals with


Larval development of Panulirus japonicus

FISHERIES SCIENCE

1.2

Body weight (WW, DW; mg)

Metamorphoses to the puerulus
stage

35

Body length (mm)

30
25
20
15

771

5th instar

1.0

WW = 0.0213D + 0.1310

4th instar

0.8

R2 = 0.9861

3rd instar
2nd instar

0.6

1st instar

0.4

DW = 0.0076D + 0.0259
R2 = 0.9807

0.2
0.0

Body length of the puerulus stage

0

10

10

20

30

40

Days after hatching (D)

5

Fig. 9 Relationships between days after hatching and
(᭺) dry (DW), and (᭹) wet body weight (WW) for Panulirus japonicus phyllosoma from the first to fifth instar
under laboratory conditions.

0
1

50

100 150 200 250 300

Days after hatching

Survival rate (%)

100
80
60
40

120

Body weight (WW, DW; mg)

Fig. 7 Growth of the two larval Panulirus japonicus
with the shortest (black line) and the longest (gray line)
larval life among the five larvae that reached the puerulus stage.

WW = 0.0686BL2.2023

100

2

R = 0.9966
80
60
DW = 0.0209BL2.1905

40

2

R = 0.9946
20
0

20

0

5

10

15

20

25

30

Body length (BL; mm)

0
0

100

200

300

400

Days after hatching
Fig. 8 Survival of Panulirus japonicus phyllosoma in
the mass culture for obtaining samples for measurements of body weight.

the shortest and longest phyllosoma life, respectively, among the five larvae that reached the puerulus stage. Their growth rates were similar to each
other until approximately 180 days after hatching,
at which point the difference in BL between the
two larvae widened, resulting in a disparity of
3 months in larval life length.
Mass culture
The phyllosoma cultured in groups suffered from a
disease that showed necrosis on the pereiopod and
antennule from 2 weeks after hatching; the disease

continued until the end of the culture. Hence,
the survival rate decreased gradually to 45% at
100 days and 35% at 200 days after hatching
(Fig. 8). In the mass culture, two phyllosoma metamorphosed to the puerulus stage.

Fig. 10 Relationships between body length and (᭺) dry
(DW), and (᭹) wet body weight (WW) throughout the
entire phyllosomal life of Panulirus japonicus under
laboratory conditions.

For the first to fifth instar larvae, WW and DW
increased linearly as the days after hatching progressed (Fig. 9). The relationships between WW or
DW and days after hatching (D) can be expressed
by the following linear equations:
WW = 0.0213D + 0.1310 (R2 = 0.9861)
DW = 0.0076D + 0.0259 (R2 = 0.9807)
From the data throughout the entire phyllosoma
phase, WW and DW increased exponentially as BL
increased (Fig. 10). The relationships between WW
or DW and BL were expressed as the exponential
equations below:
WW = 0.0686BL2.2023 (R2 = 0.9966)
DW = 0.0209BL2.1905 (R2 = 0.9946)
The moisture content of WW (%), calculated from
the equation (WW − DW)/WW × 100, was approxi-


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mately 80% for newly hatched larvae. It decreased
to 63–70% at approximately 5 mm BL and then
increased to 68–75% at approximately 10 mm BL.
Thereafter, it was constant at approximately 70%.
DISCUSSION
Several reports on the length of the phyllosoma
lifetime of P. japonicus grown in the laboratory
have been published. Yamakawa et al.15 successfully obtained a puerulus from about 1000 newly
hatched larvae and reported that the puerulus had
a phyllosoma stage that lasted 307 days (24–26°C).
Kittaka and Kimura16 also noted that two phyllosoma metamorphosed to the puerulus stage 340
and 391 days after hatching (24–28°C). Sekine
et al.11 produced 325 pueruli in the laboratory for
several years and reported that the phyllosoma
stages ranged from 231–417 days (mean
319.4 days) (24–27°C). In the individual culture
of the present study, five larvae reached the puerulus stage 245–326 days after hatching (mean
289.0 days), with shorter phyllosoma phases than
those reported in other studies. This is probably
related to the high survival rate in the present
study. The survival rate from hatch to puerulus
stage was 50% in the present study, while other
studies reported survival rates under 10%. The high
survival rate in this study indicates that the individual culture using small glass cups provided
better conditions for the larvae than other studies
did, resulting in their fast growth.
P. japonicus phyllosoma reached the puerulus
stage after 22–29 molts (n = 5, mean 26.2 molts) in
the present study. The variability in the number of

instars as well as in the length of larval life was
caused mainly by differences in molt increment
and BL of the final instar between individuals. The
animal that possessed the smallest number of
phyllosoma instars (22) gained a 4.95-mm molt
increment at 23.50 mm BL, and it metamorphosed
to the puerulus stage from the final phyllosoma
instar of 28.45 mm BL by the next molt. The animal
that had the greatest number of phyllosoma instars
(29) had molt increments of 2.40 and 2.60 mm
at BL of 25.50 and 27.90 mm, respectively, and
reached the puerulus stage from the final phyllosoma instar of 33.05 mm. The fact that all larvae
were cultured individually with the same methods
suggested that these variabilities were induced by
the growth potential of each larva. Once a larva
showed a small molt increment, in many cases it
grew with small molt increments for several subsequent instars. Although it is not clear what determines the growth potential, variation in the length
of life and the number of instars could be charac-

H Matsuda and T Takenouchi

teristic of P. japonicus phyllosoma. This finding is
indicated by the fact that wild pueruli can be collected on the coast of Japan from April to December, which is a long period compared with the
relatively short period (from July to August) when
hatching occurs.17,18
During the entire phyllosoma life of P. japonicus,
some changes in the behavior and optimal growth
conditions have been recognized. Matsuda et al.12
observed diel timing of molting in P. japonicus
phyllosoma and reported that the timing of molting varied with individuals until around 1 month

after hatch (4–5 mm BL), at which point individual
differences became small and all larvae molted
synchronously around dawn. The larval response
to light changes at approximately 4–5 mm BL.
Newly hatched larvae display strong positive phototaxis, which gradually disappears with development, and beyond 4–5 mm BL they generally show
negative phototaxis.19 The rearing temperature for
high survival and rapid growth of P. japonicus phyllosoma shifted from 26 to 24°C at 15 mm BL.9
Moreover, molt death occurs more frequently in
larger (BL > ∼18 mm) than in smaller larvae.20
These changes occur at around 4–5 and 15–18 mm
BL. In the present study, the trend in the relationship between BL before a molt and the increment
in BL by the next molt changed at approximately 5
and 15 mm BL, and the trend in the growth index of
increase in BL from one instar to the next also
altered at approximately 15 mm BL, indicating that
the boundaries at 5 and 15 mm appear to have biological implications for P. japonicus phyllosoma.
Accordingly, the three phases (partitioned at 5 and
15 mm BL) can be defined as early, middle and late
phases in the long phyllosoma life. Optimal culture
conditions should be determined for each stage.
Body weight has been measured for larvae of
several decapod crustaceans.14,21–23 Larvae of many
decapod crustaceans have unique and complex
body forms, and the body forms change drastically
with molts, which make it impossible to measure
BL throughout the entire larval stage in the same
way. Therefore, the increase of BL was generally
examined not in relation to changes in BL but
instead with increasing days after hatch or number
of instars. The length of phyllosoma life of

P. japonicus, however, was extremely long (245–
326 days in the present study), and the number of
instars was large (22–29). Also, differences between
individuals were large. These reasons prevented
expressing the relationship between body weight
and days after hatching or number of instar, except
for the early stages. Unlike other decapod crustaceans, the body form of the phyllosoma underwent
no change throughout the entire phase, and BL
increased linearly with molts (Fig. 3), suggesting


Larval development of Panulirus japonicus

FISHERIES SCIENCE

that it is appropriate to express increases in body
weight in relation to BL. The relationships between
dry/wet body weights and BL could be well
described using exponential equations with high
coefficients of determination (R2 = 0.9966 and
0.9946, respectively). The exponential equations
have exponents of about 2 (2.2023 and 2.1905,
respectively), indicating that phyllosoma growth is
directed not toward an increase in body volume
but toward spreading the surface of their dorsoventrally flattened body. The flattened body form of
the phyllosoma is considered to be advantageous
for passive horizontal transport with currents.24 It
is likely that a body surface that increases with
development is greatly beneficial for oceanic larval
dispersal.

ACKNOWLEDGMENTS
We thank Professor M. Tanaka, Kyoto University,
and Associate Professor T. Yamakawa, University of
Tokyo, for helpful and constructive criticism during the preparation of our original manuscript.
Thanks to the staff of our laboratory for help during
larval culture. This research was financially supported in part by a grant from the Ministry of
Agriculture, Forestry and Fisheries of Japan.

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