FISHERIES SCIENCE

2007; 73: 331340

Maturity and reproduction of goneplacid crab

Carcinoplax vestita (Decapoda, Brachyura)
in Tokyo Bay
Wataru DOI, Than Than LWIN, Masashi YOKOTA, Carlos Augusto STRSSMANN AND
Seiichi WATANABE*
Department of Marine Biosciences, Tokyo University of Marine Science and Technology, Minato,
Tokyo 108-8477, Japan
ABSTRACT:
Sexual maturity, morphological sexual dimorphism, and reproduction of the goneplacid crab Carcinoplax vestita were investigated in Tokyo Bay, Japan, from November 2002 to
October 2003. The puberty molt in males was evidenced by changes in the relative size of the
chelipeds and merus of the walking legs, and was estimated to occur at a size range of 13.20
18.85 mm carapace length. Post-pubertal females were identied by the relative size of the abdomen
and puberty was estimated to occur at a size range of 12.8115.46 mm carapace length. Sexual
dimorphism in C. vestita was observed in all features that showed secondary sexual characteristics.
Monthly changes of gonad index in males, and of seminal receptacle index and occurrence of sperm
plugs in females were synchronized, and indicated that mating was intense in spring. Ovaries began
developing in March. Ovigerous females were found in all months except December, but were clearly
more abundant between August and October. Fecundity ranged from 7800 to 57 000 mature oocytes
per female per batch and was highly correlated with body size. The results suggest that some females
may spawn more than one batch per year.
KEY WORDS: brachyura, Carcinoplax vestita, Goneplacidae, puberty molt, relative growth,
reproduction, sexual maturity, Tokyo Bay.

INTRODUCTION
The goneplacid crab Carcinoplax vestita (Xanthoidea; Goneplacidae) is a small crab species that
reaches approximately 30 mm in carapace width
(CW). It inhabits sandy and muddy bottoms at
depths of 10100 m18 and has been recorded in
Japan, China, Korea, Australia and South Africa.1,2
In Japan, this crab is very abundant in the inner bay
areas of temperate zones, such as Tokyo Bay,36 Ise
Bay7 and Osaka Bay.8 It plays an important role in
the benthic assemblage in these areas, for instance,
as the feed of the mantis shrimp Oratosquilla
oratoria9 and the starspotted-dogsh Mustelusmanazo.10 Although there are 47 species of goneplacid crabs in Japan,1 this family of crabs has
rarely been studied because of its subtidal habitat
and lack of commercial value. The limited literature available on the family Goneplacidae includes
*Corresponding author: Tel: 81-3-5463-0535.
Fax: 81-3-5463-0684. Email: watanabe@kaiyodai.ac.jp
Received 19 May 2006. Accepted 14 November 2006.

descriptions of larval morphology in Lambdophallus anfractus,11 Carcinoplax longimana12 and

Eucrate crenata,12 and of relative growth of the
cheliped in C. longimana.13 Ogawa14 and Doi et al.6
provided preliminary information on the growth
and reproductive pattern of C. vestita and other
crabs caught by trawling in the Seto Inland Sea
and Tokyo Bay, respectively. Because of the small
sample sizes, however, the information provided
by those studies on the reproduction of C. vestita
was limited to the seasonal changes in the presence of ovigerous females. Thus, information on
the reproductive biology, an important parameter
to understand the population dynamics and the
effects of one species on other species sharing
the same habitat, is almost entirely lacking for
goneplacid crabs in general and for C. vestita in
particular.
Information on the reproductive biology of
brachyurans can be obtained through direct observation of the appearance of the gonads and associated reproductive organs, because, as a rule, it
involves the processes of copulation, egg laying

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and incubation.15 Thus, the copulative cycle can be

determined from the temporal changes in the
occurrence of females with sperm plugs,16 mating
pairs17 and the content of the seminal receptacle,18
whereas the maturation and spawning cycles can
be inferred from the analysis of ovarian development (gonadosomatic index and histology)19 and
the occurrence of ovigerous females. It is also
possible to obtain reproductive information
from the observation of secondary sexual
characteristics.15,2022 Thus, morphological measurement of structures such as the chelipeds,
abdomen and pleopods permits us to identify
subtle changes in their growth patterns that are
associated either with sexual dimorphism or with
the attainment of sexual maturity. A transition in
growth pattern in these structures often occurs at
the puberty molt, so called because it divides the
pre- and post-puberty phases of the life of crustaceans, and this is commonly associated with physiological and functional changes related to sexual
maturity.23
The purpose of the present study was to clarify
the reproductive biology of C. vestita in Tokyo Bay.
This was accomplished by a comprehensive analysis of the onset of puberty, sexual maturation and
spawning of this species using morphological techniques. To the best of our knowledge, this is the rst
report of its kind on the population biology of the
family Goneplacidae.

MATERIALS AND METHODS

Sampling
Samples were collected monthly between November 2002 and October 2003 in the southern part of
Tokyo Bay (Fig. 1) using a small beam trawl. The
sampling location corresponds to an area of Tokyo
Bay where the mantis shrimp O. oratoria is caught
by trawl shing. Sampling was performed once a
month during daytime in six xed stations and one
randomly selected station in the area. The mean

W Doi et al.

depths of stations 16 were 13, 17, 20, 23, 24 and

27 m, respectively. The trawl net had a beam length
of 6 m, a mouth height of 0.5 m and a width of
5.5 m. The cod end mesh size was 18 mm. Towing
was performed for 525 min (mean SD,
15.2 3.6) at a ship speed of 2.53.0 knots
(3.1 0.2). Specimens were preserved in 10%
formaldehyde sea water on board and brought to
the laboratory. Crabs were sexed by the form of
the pleopods and the females were checked for
the presence or absence of attached eggs in the
pleopods and of sperm-plugs in the vaginal tract.
Ovigerous females were separated into those
bearing early developing (non-eyed) eggs and
those with eyed eggs.

Morphological analysis of sexual maturity

and sexual dimorphism
The size of the individuals used in this analysis
ranged from 6.83 to 23.91 mm carapace length (CL)
for males (n = 372) and 7.7423.73 mm CL for
females (n = 340). Carapace length and the following body parts were measured using a vernier
caliper to the nearest 0.01 mm: CW, merus length
of the rst and second walking legs (ML1 and ML2),
propodus length (PL) and height (PH), dactylus
length (DL) of the largest cheliped, and width of the
fth abdominal segment (AW). Small individuals
and the length of the right gonopod (GL) in males
were measured under a stereomicroscope using a
micrometer. Wet body weight (BW) was measured
with an electronic balance to the nearest 0.01 g
only for complete, undamaged individuals.
The relative growth of body parts to CL was
described by either a single regression or a pair of
regression lines. For the two-line tting, data were
divided into pre- and post-puberty phases either
by the method of Knucky24 or using Akaikes information criterion (AIC).25 In the former, frequency
distributions of each measurement per CL class
were constructed and the parameters for the
two normal distributions were estimated by the

Fig. 1 (a,b) Location of the

study area and (c) sampling stations in the southern part of
Tokyo Bay. Lines in (c) indicate
the direction of towing lines.

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FISHERIES SCIENCE

maximum likelihood method using solver, a nonlinear optimization tool in Excel 2002 (Microsoft,
Tokyo, Japan).26 The intersection point of the two
normal distributions was then used to divide the
pre- and post-puberty phases. In the AIC method,
the inection point was estimated by stepwise calculation using a value of 0.01 mm.
The ratio of post-pubertal crabs (P) per 1 mm CL
class (l ) was expressed by the logistic curve
P (l) = 100/[1 + e(al + b)]. The parameters a and b
were estimated by the maximum likelihood
method.27 The CL at which 50% of crabs were post
pubertal (CL50%), P (l) = 50 was determined by the
formula CL50% = b/a.
For the relationship between CL and BW, a power
function was tted by the non-linear least square
method using the solver.
Reproductive cycle
The percentages of ovigerous females and of
females with sperm plugs were determined for
each month. The gonads and seminal receptacles
of representative post-pubertal individuals were
removed and dried at 60C for 24 h and then
weighed to the nearest 0.0001 g. Gonad index (GI)
was calculated by the following equation for both
sexes: GI = GW 106/CL3, where GW represents the
dry weight of the ovary or vas deferens in females
and males, respectively. The seminal receptacle
index (SRI) was calculated by the equation:
SRI = SR 107/CL3, where SR denotes the dry
weight of seminal receptacles.
Fecundity
Subsamples of ovarian tissue (range of 0.010.03 g)
were dissected from the anterior, middle and posterior regions of the gonad of females with mature
oocytes, and the number of oocytes in each subsample was counted under a binocular microscope. The subsamples and the remaining ovarian
tissue were then dried at 60C for 24 h and weighed
to the nearest 0.0001 g. Fecundity was determined
by the gravimetric method using the oocyte counts
and weights of ovarian tissue.

333

with changes in the relative size of the chelipeds

and merus of the rst and second walking legs
(Table 1). The frequencies of PL/CL, DL/CL and
PH/CL ratios showed distinct bimodal distributions in the males, with cut-off values between the
two phases of relative growth of 1.24, 0.64 and 0.56
for the three parameters, respectively (Table 1,
Fig. 2). The relative growth of cheliped dimensions
was signicantly faster in post-pubertal males as
compared to pre-pubertal ones (ancova, P < 0.001;
Table 1). Signicant increases in the relative
growth rates of ML1 and ML2 in males after
puberty were detected by AIC analysis. The inection points between growth phases of merus length
were estimated at 13.2 mm CL for both ML1 and
ML2 (Table 1 and Fig. 2).
The logistic equations representing the relationship between the ratio of post-pubertal males, as
inferred from changes in cheliped dimensions, and
CL were as follows:

PL : P (l ) = 100 [1 + e (1.17l +17.55) ]

(R 2 = 0.995),
DL : P (l ) = 100 [1 + e (0.92l +14.49) ] ( R 2 = 0.986 ) ,
PH : P (l ) = 100 [1 + e (0.83l +13.30) ] ( R 2 = 0.982).
The CL50% values estimated from these equations
were 15.04 mm, 15.77 mm and 15.97 mm for PL,
DL, PH, respectively (Fig. 3).

Female sexual maturity

Sexual maturity in females was accompanied only
by a change in abdomen width in proportion to CL,
with the relative growth rate of AW in females
increasing signicantly after the puberty molt
(ancova, P < 0.001; Table 1). Accordingly, prepuberty females had AW/CL ratios 0.42, whereas
post-puberty ones, which included all ovigerous
females, had ratios > 0.42 (Table 1 and Fig. 4).
The relationship between the ratio of postpubertal females to CL, as estimated from the relative size of the abdomen, was described by the
logistic equation:

P (l ) = 100 [1 + e (1.76l + 24.47) ]

(R 2 = 0.998).

The P (l) was 0% and 100% below 11 mm and

above 15 mm CL, respectively, and the CL50% value
was estimated to be 13.93 mm (Fig. 5).

RESULTS
Male sexual maturity

Sexual dimorphism

The onset of sexual maturity in males did not cause

any noticeable changes in CW, abdomen width, or
gonopod length relative to CL, but was associated

Sexual dimorphism in relative growth rates to CL

was found in all parameters available for both
sexes (Table 1). The regression parameters were

1
1
2

1
2

1
2

1
1
2

1
1
2

1
1
2

DL

PH

ML1

ML2

AW

GL

NA
NA
1.24
>1.24
NA
0.64
>0.64
NA
0.56
>0.56
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
0.42
>0.42
NA

Bivariate ratios
(DV/CL)

NA

NA
NA
NA

NA
465.670
454.321

NA
415.764
395.273

NA
NA

NA
NA

NA
NA
NA

AIC
6.8323.91
7.7423.73
6.8317.87
13.5223.91
7.7423.73
6.8318.85
13.5223.91
7.7423.73
6.8318.85
13.5223.91
7.7423.73
6.8323.91
6.8313.20
13.2023.91
7.7423.73
6.8323.91
6.8313.20
13.2023.91
7.7423.73
6.8323.91
7.7415.46
12.8123.73
6.8323.91

CL range
1.432
1.409
1.265
1.936
1.101
0.621
1.049
0.590
0.574
0.935
0.430
0.913
0.691
0.960
0.689
0.978
0.775
1.016
0.760
0.360
0.455
0.702
0.453

Slope (b)
-0.298
-0.009
-1.507
-10.238
-0.492
-0.584
-6.238
-0.522
-1.128
-5.817
-0.033
-2.063
0.485
-2.863
0.242
-1.700
0.628
-2.333
0.577
-0.493
-0.972
-3.357
-0.046

Intercept (a)

Parameter

359
309
106
199
262
103
204
259
116
197
270
235
65
170
214
235
65
170
214
337
113
179
356

n
0.995
0.995
0.970
0.943
0.992
0.963
0.926
0.986
0.923
0.882
0.978
0.980
0.944
0.959
0.983
0.949
0.936
0.954
0.983
0.984
0.979
0.954
0.975

*
*
*
*
*
*
*
*
*
*
*
NA
NA
NA
*
NA
NA
NA
*
*
*
*
*

NS

NS

Between
sexes

Signicance

Asterisks indicate the slopes (b) were signicant (t-test; *P < 0.001).
Daggers indicate the slope (b) was signicantly different between both sexes (ANCOVA; P < 0.05; P < 0.001). Non-signicant differences are shown as NS.
AIC, Akaikes information criterion; AW, fth abdominal segment width; CL, carapace length; DL, dactylus length of major cheliped; DV, dependent variables; GL, length of right rst
gonopod (male); ML1 and ML2, merus length of rst and second walking legs; NA, not applicable; PL, propodus length of major cheliped; PH, propodus height of major cheliped.

PL

CW

Dependent
variables

Sex

Regression lines y = a + bx for the morphometric analysis of Carcinoplax vestita

No.
regression
lines

Table 1

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Reproduction of Carcinoplax vestita

335

30
25
20

Propodus height (mm)

20

20
0
1.00

1.20

1.40

1.60

PL / CL

10

First walking leg

15

10

17
14
11

60

Merus length (mm)

Number of individuals

Dactylus length (mm)

40
20
0
0.50 0.58 0.66 0.74 0.82

DL / CL

8
5
2
17 5
14
11

Second walking leg

20

15

10

5
60

10

15

20

25

10

0.40

0.48

0.56 0.64

0.72

PH / CL

8
5
2
5

10

15

20

25

Carapace length (mm)

: Pre-puberty
Number of individuals

11

50

0
15

25

14

10

20

20

100

15

Carapace length (mm)

40

Abdominal width (mm)

Ratio of post-puberty males (%)

60
40

15

20

Fig. 2 Relative growth in propodus length (PL), dactylus length

(DL), propodus height (PH), rst
(ML1) and second (ML2) walking
leg merus length to carapace
length (CL) and bimodal frequency distributions of PL, DL
and PH/CL ratios in male Carcinoplax vestita.

: Post-puberty

Merus length (mm)

Number of individuals

35

Number of individuals

Propodus length (mm)

: Pre-puberty

20

25

Carapace length (mm)

Fig. 3 Relationship between carapace length and percentage of post-puberty males of Carcinoplax vestita as
inferred from the ratio of propodus length (PL, ), dactylus length (DL, ) and propodus height (PH, ) to carapace length. Logistic equations of best t are described
in the text.

: Post-puberty

60

40

20

0
0.26 0.30 0.34 0.38 0.42 0.46 0.50 0.54 0.58

AW / CL

2
5

10

15

20

25

Carapace length (mm)

Fig. 4 Relative growth of fth abdominal width (AW) to
carapace length (CL) and bimodal frequency distributions of AW/CL ratios in female Carcinoplax vestita.

FISHERIES SCIENCE

W Doi et al.

14

100

12

Gonad Index

Ratio of post-puberty females (%)

336

50

10
8
6
4
2
0

0
5

10

15

20

Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct

25

Month

Carapace length (mm)

Fig. 5 Relationship between carapace length and percentage of post-puberty females in Carcinoplax vestita
as inferred from ratios of the width of the fth abdominal
segment to the carapace length. The logistic equation of
best t is shown in the text.

Fig. 7 Monthly changes of gonad index in males of Carcinoplax vestita between November 2002 and October
2003. () mean.

Gonad index

100

Body weight (g)

20

15

10

0.1

10
0.01

Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct

Month

0
5

10

15

20

25

Carapace length (mm)

Fig. 6 Relationship between carapace length (CL) and
body weight (BW) in Carcinoplax vestita. () male:
BW = 2.978 10-4 CL3.462 (n = 211, R2 = 0.887); () female:
BW = 9.381 10-4 CL2.991 (n = 186, R2 = 0.919).

generally signicantly different between both

pre- and post-pubertal males and females, the only
exceptions being ML1 and ML2, which only differed between post-pubertal males and females.
A signicant sex difference was found also in the
rate of increase of BW relative to CL, which was
described by power equations for both sexes (likelihood ratio test, P < 0.001; Fig. 6). Thus, the BW of
males increased noticeably faster than that of
females at CL over 15 mm.

Fig. 8 Monthly changes of gonad index in females

of Carcinoplax vestita between November 2002 and
October 2003. Non-ovigerous () and ovigerous females
(with [] and without [] eyed eggs) are indicated. ()
mean.

Reproductive cycle
The gonad index (GI) of post-pubertal males varied
between 0.08 and 13.76 (Fig. 7). Males with high
GI were found in all but the winter months
(DecemberFebruary). The GI of all post-pubertal
non-ovigerous females were consistently low
(01.5) between November and February (Fig. 8),
but showed a bimodal distribution between March
and October. Accordingly, approximately one- to
two-thirds of the females showed values up to
3070, while the remaining females continued
to have low GI. Ovigerous females with eyed eggs
showed higher GI than those with non-eyed eggs
during the spawning period (Fig. 8).

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40

Occurrence of ovigerous
females (%)

Occurrence of females
with sperm plugs (%)

25

337

20
15
10
5
0

35
30
25
20
15
10
5
0

Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct

Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct

Month
Fig. 9 Monthly changes in the occurrence of females of
Carcinoplax vestita with sperm plugs between November 2002 and October 2003.

Month
Fig. 11 Monthly changes in the occurrence of ovigerous females of Carcinoplax vestita between November
2002 and October 2003. Ovigerous females with () and
without () eyed eggs are indicated.

6
15

10

Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct

Month
Fig. 10 Monthly changes in the seminal receptacle
index of female Carcinoplax vestita between November
2002 and October 2003. Non-ovigerous () and ovigerous females (with [] and without [] eyed eggs) are
indicated.

A total of 166 females with sperm plugs were collected during the study and their size ranged from
11.62 to 19.46 mm CL. Females with plugs were
found in December and from April to September,
with the highest frequency in May (21%; Fig. 9).
The seminal receptacle index (SRI) uctuated
between 0.29 and 19.63 (Fig. 10). Females with the
highest SRI values (>10) increased in number from
April to July and then decreased.
Ovigerous females were found in all months
except December (Fig. 11). Their frequency was
low (<3%) between November and July, but
increased abruptly to >20% between August and
October. The CL of ovigerous females ranged from
11.02 to 22.16 with a mean (SD) of 16.14 (2.19)

Number of oocytes (10 4)

Seminal receptacle index

20

5
4
3
2
1
0
13

14

15

16

17

18

19

20

21

22

Carapace length (mm)

Fig. 12 Relationship between carapace length and
number of mature oocytes in the ovaries of Carcinoplaxvestita females.

(n = 269). Females bearing eyed eggs were found

only between May and October.
Fecundity
The size of the 31 females used for the fecundity
analysis ranged from 13.12 to 22.16 mm CL
(mean SD, 16.85 2.39). Fecundity (F ) varied
between
7800
and
57 000
(mean
of
25 000 11 000, Fig. 12) and was highly correlated
to CL as follows:

F = 9.20CL2.78

(n = 31, R 2 = 0.81).

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DISCUSSION
Sexual maturity and secondary sex
characteristics
The secondary sexual development of the chelipeds in male crabs is often associated with combat,
display and courtship of potential mates.20,21,28 It is
also known that the rst and second walking legs
are used to hold and carry the female during preand post-copulatory guarding.29 Thus, the relative
growth of cheliped and merus on and after the
puberty molt in crabs is closely related to functional sexual maturity. This study revealed clear
changes in the relative size of the chelipeds and
merus of the rst and second walking legs in males
of C. vestita and showed that both structures grow
faster after puberty. In contrast, we did not detect
any changes in gonopod morphology, even though
this feature is considered the best external indicator of puberty and gonadal development in the
males of some species (e.g. Eriphia smithii30 and
Panopeus austrobesus31). The secondary sexual
development of the chelipeds seems to be a
common feature among crabs of the superfamily
Xanthoidea (e.g. C. longimana,13 E. smithii,30
P. austrobesus31 and Platyxanthus patagonicus32)
although their dimensions do not change with
maturity in Trapezia ferruginea.33 Likewise,
elongation of the merus of the rst (ML1) and
second (ML2) walking legs has been recorded in
males of the majid crab Tiarinia cornigera.18 Interestingly, the cheliped and merus appear to grow
by different processes in males of C. vestita. For
instance, the truncated, bimodal distribution of
PL/CL, DL/CL and PH/CL ratios observed in crabs
of 13.5218.85 mm CL suggests that the relative
size of the chelipeds increases abruptly at the
puberty molt. This is not the case with merus elongation, which, as suggested by the presence of a
clear inexion point in the ML1/CL and ML2/CL
relationships, seems to occur gradually with
each molt.
A growth phase transition in the cheliped
dimensions of males was observed between 13.52
and 18.85 mm CL and it was estimated that 50%
of the crabs attained sexual maturity by 15.04
15.97 mm CL. The relatively wide range of sizes
between the smallest (13.52) and largest (18.85)
pubertal individuals could be related to differences in birth dates (see section on reproductive
cycle), period of growth, and consequently on the
physiological readiness to attain sexual maturity
at appropriate times (e.g. reproductive season).
The morphological analysis of ML1 and ML2 provided an estimated size for puberty for males of
approximately 13.20 mm CL. Thus, the size for

W Doi et al.

sexual maturity as inferred from merus growth

agrees well with the lower value based on the
changes in cheliped dimensions, and is probably
the closest estimate of the actual size at which
most males begin gonadal development. This
assumption is supported by the results of the GI
analysis, which showed that males more than
13.00 mm CL can have GI values comparable to
larger males during the reproductive season.
Although we did not examine the gonads by histology, we feel condent that increased GI values
accurately reect the onset of gonadal development and the state of physiological maturity in
males of C. vestita, as is generally found in other
crabs.
Allometric growth associated with sexual maturity in C. vestita females was observed only in the
abdomen, whose width increased after the puberty
molt. This pattern of relative growth agrees with
the general pattern described for other female
brachyurans,20 including the related xanthoid
species P. austrobesus,31 P. patagonicus,32 Leptodius
exaratus34 and E. smithii.30 According to the
changes in AW proportion, female C. vestita
attained sexual maturation between 12.81 and
15.46 mm CL, with an estimated CL50% of 13.93 mm
CL. Nevertheless, the smallest ovigerous and
plugged females found had 11.02 and 11.62 mm
CL, respectively. This discrepancy might be
attributed to the large sample size used in this
study, which increases the probability of nding
exceptional individuals. It may be also that this
kind of secondary sex characteristic only appears
some time after the attainment of physiological
maturity.

Sexual dimorphism
As expected, sexual dimorphism in C. vestita was
observed in all measurements that showed secondary sexual characteristics. However, while in
the PL, DL, PH and AW the sexual dimorphism
occurred before the puberty molt, differences
between the ML1 and ML2 of males and females
became evident only after the onset of sexual
maturity. Whether and when sexual dimorphism
occurs varies greatly with the measurement and
the species.13,3032 The BW also showed sexual
dimorphism, with weight increasing relatively
faster in males than in females, a phenomenon
probably caused by the enlargement of the male
cheliped. A small but statistically signicant
dimorphism was observed in the relative growth
in CW but its ecological meaning remains
uncertain.

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Reproductive period
Ovarian development began in March, whereas
mating appeared to be largely concentrated
around May, as inferred from GI data, occurrence
of females with sperm plugs, and monthly changes
of SRI. The spawning activity of C. vestita was
largely concentrated in autumn (AugustOctober),
as inferred from the occurrence of ovigerous
females, but some of the females also spawned in
another months. The duration of the ovigerous
period is highly variable among temperate
xanthoid crabs: 3 months for Rhithropanopeus
harrisii tridentatus,35 4 months for L. exaratus34
and E. smithii,16 5 months for Neopanope sayi,36
8 months for P. patagonicus,32 Panopeus herbstii
(intense for 2 months)37 and Eurypanopeus
depressus,37 and 10 months for Lophopanopeus
bellus bellus.38 Thus, the pattern of occurrence of
ovigerous females in the present species, that is, a
well-dened, main period within a protracted
period of limited activity, coincides with that of
P. herbstii.37 We surmised that some females are
able to spawn more than once per year in Tokyo
Bay because ovigerous females sometimes had
high GI values and because those with eyed eggs
had higher GI values than those with non-eyed
eggs. Ongoing studies on the length of the incubation time should help clarify the number of spawnings made by each female during one reproductive
season.

339

in the cheliped and abdomen, and signicant, but

less pronounced, in the merus of the rst and
second walking legs and carapace, and that allometric growth in these structures provides an estimate of the size at sexual maturity in C. vestita.
Thus, C. vestita follows the typical pattern of development of secondary sexual characteristics and
sex dimorphism as for other brachyurans.21 From
this study, the reproductive cycle of C. vestita in
Tokyo Bay can be surmised as follows. Ovaries
begin to develop and mating occurs in spring,
whereas spawning is largely concentrated from
late summer to autumn. Some females may be able
to spawn consecutively during the same reproductive season, but an accurate number of spawnings
per female and their interval are still unclear.
Ongoing studies should help elucidate these points
and provide an integrative view of the life history of
C. vestita.

ACKNOWLEDGMENTS
We are grateful to Dr S. Segawa (Tokyo University
of Marine Science and Technology) for critical
reading of the manuscript. We also thank Mr M.
Koyama (Shiba Branch, Yokohama City Fisheries
Cooperative Association), Dr K. Kodama (National
Institute for Environmental Studies), and Mr R.
Yamada (University of Tokyo) for their help in
sampling.

Fecundity
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24.00 mm CL or 18.0031.00 mm CW). Fecundity
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crabs from the same group.
CONCLUSIONS
This study demonstrates that secondary sexual
characteristics and sexual dimorphism are marked

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