Springer-Verlag 1997

Marine Biology (1997) 128: 5562

J. L. Overton D. J. Macintosh R. S. Thorpe

Multivariate analysis of the mud crab Scylla serrata

(Brachyura: Portunidae) from four locations in Southeast Asia

Received: 2 October 1996 / Accepted: 5 December 1996

Abstract The large, edible mud crab Scylla serrata

(Forskal) exhibits different phenotypes which are recognised by the fishermen of Southeast Asia and are
given local names such as white, green, red or
black crabs on the basis of their colouration. A preliminary study using multivariate techniques was undertaken in order to examine the degree of dissimilarity
between the different morphs of commercially fished
stocks of S. serrata. Twenty-two morphometric and 20
meristic characters were measured on male crabs from
four locations in three countries in Southeast Asia, including Surat Thani in Thailand where black and
white morphs exist sympatrically. Canonical variate
analysis (CVA) on the morphometric data discriminated
three groups with no chain-linking. Surat Thani white
crabs and those from south Vietnam were phenotypically similar and formed one distinct group; a second
group contained crabs from Ranong (Thailand) and
Sarawak which were phenotypically black; the third
group contained black crabs from Surat Thani. CVA
on meristic data confirmed two groupings, but implied
that the Surat Thani black morph may only be a
variant of the Surat Thani/Vietnam white form, rather
than a third species. These findings are discussed in relation to the ecology, fishery management and aquaculture potenial of S. serrata.

Communicated by J.P. Thorpe, Port Erin

J.L. Overton1 (&) D.J. Macintosh1
Institute of Aquaculture, University of Stirling,
Stirling FK9 4LA, Scotland
R.S. Thorpe
School of Biological Sciences, Brambell Building,
University of Wales, Bangor LL57 2UW,
Gwynedd, North Wales, Great Britain
Present address:
1
Centre for Tropical Ecosystems Research,
Department of Genetics and Ecology,
Institute of Biological Sciences, Building 540,
University of Aarhus, D-8000 Aarhus C, Denmark

Introduction
Scylla serrata (Forskal) is the only species of the family
Portunidae that is closely associated with mangrove
environments. Commonly known as the mangrove crab
or mud crab, S. serrata is distributed throughout the
Indo-Pacific region mainly within tropical latitudes, but
it can also be found in more temperate environments as
far north as China and Japan (Macnae 1968).
The wild catch of Scylla is an estimated 10 000 tons
annually (BOBP 1992) and contributes significantly to
the coastal fisheries of many developing countries in
Asia, e.g. Bangladesh, India, Sri Lanka, Indonesia,
Thailand, Vietnam and the Philippines (Ferdouse 1990;
Liong 1993); there is also a small but commercially
valuable S. serrata fishery in Australia producing an
annual catch of about 600 tons (Lee 1992).
There is a high consumer demand for Scylla serrata,
particularly in countries with Chinese communities such
as Malaysia, Singapore, Taiwan and Hong Kong (Liong
1993), where it is regarded as a delicacy, especially the
gravid female or egg crab (Chen 1990); the male is
also appreciated for its large chelae and high meat
content (Harvey 1990). Because of this popularity,
S. serrata has been subjected to heavy, unregulated exploitation, except in Australia which does have a fishing
policy for mud crab (Heasman and Fielder 1977). Many
fishing communities in Southeast Asia use unselective
fishing gear and heavily exploit the available broodstock, including the gravid females. As a result of this
high fishing pressure, the average size of mud crab
caught in Southeast Asian countries appears to be decreasing (Macintosh 1982; Harvey 1990; Overton personal observations 1994). However, any efforts to
manage the crab fisheries are hindered due to the lack of
information on the population dynamics of S. serrata in
Southeast Asia (Macintosh et al. 1993; Tan and Ng
1994).
The present impoverished status of Scylla serrata
stocks in Southeast Asia is also compounded by other

56

factors. These include habitat destruction due to extensive mangrove clearance; declining quality of the coastal
environment (Hong and San 1993; Liong 1993) and a
recent increasing interest in soft-shell crab farming in
addition to the more conventional culture methods (i.e.
rearing crabs in ponds for one to several months). Since
there is still no commercial-scale hatchery production of
S. serrata, all forms of mud crab culture depend on an
already limited natural seed supply (Harvey 1990; Liong
1993).
Progress towards effective stock management of
Scylla serrata, and the development of hatchery and
growout techniques for crab farming, are constrained by
many factors, including the uncertain taxonomic status
of S. serrata. Many reports have described more than
one variety of S. serrata in local populations; for example, different colour and size forms have been recognised by fishermen and dealers in Malaysia, Thailand,
India, the Philippines and Australia (Estampador 1949;
Macintosh 1982; Radhakrishnan and Samuel 1982;
BOBP 1992; Lee 1992). Use of common names such as
black crab, golden-backed crab and green crab
(Ferdouse 1990; Harvey 1990) demonstrates that there is
a wide variation in the morphology of S. serrata. Furthermore in some markets certain morphs of S. serrata
command a higher price than others; for example boo
khao or white crab in Thailand is consistently more
valuable than the black variety or boo dum (Thongkum 1988; Overton personal observation 1996).
This variation in the phenotype of the mud crab has
led to much debate with respect to its taxonomy (Alcock
1899; Estampador 1949; Serene 1952; Stephenson and
Campbell 1959; Kathirval and Srinivasagam 1992). The
taxonomy of Scylla serrata continues to be confusing
and has been further complicated by more recent descriptions based on general morphological features
(Radhakrishnan and Samuel 1982; Joel and Raj 1983).
Moreover, these studies have been carried out on a local
basis, without taking into account the crabs morphological degree of variation on a wider geographical scale.
This paper reports on the results of multivariate
analysis of morphometric and meristic data for Scylla
collected from four commercial crab-fishing sites in
Southeast Asia.

Materials and methods

Study sites
Scylla serrata (Forskal) were collected from four locations in
Southeast Asia where there are significant crab landings by local
fishermen: Klong Ngao in Ranong Province (953 to 957 N;
9831 to 9837 E : Site A), southwest Thailand; Ban Don Bay in
Surat Thani Province (911 to 921 N; 9914 to 9940 E : Site B),
western Gulf of Thailand; Can Gio near Ho Chi Minh City, south
Vietnam (1022 N; 10715 E : Site C); Sematan in Sarawak, East
Malaysia (151 N; 10947 E : Site D). These locations are illustrated in Fig. 1. The sites were chosen for their degree of geographical separation as well for their importance as mud crab
fishing-areas. All four study areas were located in coastal man-

groves colonising fine alluvial sediments. The Ranong, Can Gio

and Sematan sites are extensive mangrove delta formations with
many interconnecting creeks. Surat Thani has more coastal development, but the crab fishery is centred around Ban Don Bay which
still retains fringing mangroves. Care was taken to ensure that the
crabs were of local origin by actually going on crab-collecting trips
with the local people. The crabs were collected principally with
small baited traps or gill nets.
Crab collection and measurement
In order to eliminate the effects of variation due to sexual dimorphism, only male crabs were used in this study. Thirty males were
collected from each site except for Surat Thani (Site B), where two
morphs are common; at this site, an extra 20 crabs were collected of
the second morph type. This provided five groups of crab for
measurement. To minimise ontogenic influences on body shape, the
crabs collected were approximately the same size (200 g) and on
dissection were noted to have mature testes. Only healthy individuals were selected, with all limbs intact.
Twenty-two morphometric characters were measured on each
individual to the nearest 0.01 mm using digital callipers (Mitutoyo
Absolute Digimatic, Tokyo). These characters were measured from
the carapace, abdomen, both chelae and pereiopods from the right
side of the crab, as illustrated in Fig. 2. Twenty meristic characters
were also recorded from each individual. These included spination
along the anterolateral carapace and the spination and dentition
from both chelipeds (Fig. 3).
Data analysis
Morphometric data
The 22 characters measured on each crab from the five groups were
analysed by canonical variate analysis (CVA) using the computer
software BMDP-7M (BMDP Statistical Software Inc., Cork, Ireland).
To remove the effect of size, MGPCA (multiple group principle-components analysis); (Thorpe 1988) was used to discover the
possible size vector involved (in this instance the first vector for all
22 characters). Comparable size in and size out CVA analyses
were run; in the former, the MGPCA size vector is included while
in the latter it is excluded. Comparison of these analyses indicate
the influence of size on the relationships among groups.
Meristic data
The meristic data were also analysed using CVA. In order to
confirm that the CVA had not been perturbed by heteroscedasticity, principal component/co-ordinate analysis (PCA) was performed on the group means for all the characters (Thorpe 1980).
Values from the first two component scores (normalised vectors
from PCA) and scaled eigenvectors from principal co-ordinate
analysis (PCDA) were then plotted in order to confirm the position
of the group centroids from the CVA ordination plot.

Results
Morphometric analysis
Canonical variate analysis on the raw, untransformed
data for the five Scylla serrata groups indicated three
main clusters. This is illustrated in the scatter plot of the
first two canonical variates shown in Fig. 4a. Individuals
from Ranong (Site A) and Sarawak (Site D) were shown
to be very similar in morphology, and phenotypically

57
Fig. 1 Scylla serrata. Locations
of crab collection sites in
Southeast Asia

were designated as the black type. One of the two

Surat Thani (Site B) morphs and the samples from
Vietnam (Site C) also formed a discrete cluster and were
designated as the white type. The other morph from
Surat Thani physically resembled the Ranong and
Sarawak individuals but formed a separate, third cluster,
with its centroid equidistant between the other two
clusters (Fig. 4a). This group exhibited a larger degree of
scatter between the individuals indicating more withingroup variance than illustrated by the other four groups.
No chain-linking was discovered between the three
clusters even though one of the individuals from Sarawak was reclassified as belonging to the Surat Thani
black morph group and vice versa.
CVA on the MGPCA scores showed that 30% of the
between-group variance was attributable to the sixteenth
MGPCA vector. This revealed that the main characters
contributing to the variance were: frontal length (between the orbital sockets); right and left antero-lateral
lengths of the carapace; right dactylus width, right
propodus length; right and left carpus lengths, and right

merus width. The mean, range and standard deviation

for these characters is presented in Table 1. The size
character, i.e. carapace width, has been included to show
the uniformity of size ranges between the five groups.
The MGPCA analysis also revealed the first eigenvector to be the size vector, where each coefficient had a
similar sign and magnitude. When this vector was
omitted to form the size out analysis, it was found
that the ordinance plot (Fig. 4b) was tantamount to that
of the size in plot, indicating that growth-dependent
size was not having an effect on the racial expression of
the five groups studied.
Meristic analysis
The resultant ordinance plot derived from the meristic
data on the same five groups illustrated a different pattern from that of the morphometric data. The scatter
formed two clusters rather than the three suggested
by the morphometric data (Fig. 4c). The Ranong and

58
Fig. 2 Scylla serrata. Illustration of 22 characters forming
data for multivariate analysis.
a carapace; b abdomen;
c anterior view of cheliped,
d posterior view of cheliped
(both right and left chelipeds
measured); e third right pereiopod; f fifth right pereiopod (AL
abdominal length; CL carpus
length; CW carpus width; DL
dactyl length; DW dactyl width;
FL frontal length; ICL internal
carapace length; ICW internal
carapace width; LC left anterolateral length of carapace;
LPL lower paddle length; LPW
lower paddle width; ML merus
width; MW merus width; PL
propodus length; 3PML third
pereiopod merus length;
3PMW third pereiopod merus
width; 3PTL third pereiopod
total length; PW propodus
width; RC right anterolateral
length of carapace; TPL total
length of swimming leg; UPL
upper paddle length; UPW upper paddle width)

Sarawak groups formed the same pattern as before, but

the Surat Thani white morph and the individuals from
Vietnam had a larger within-group variance than in the
morphometric analysis.
Canonical variate analysis on the meristic data also
indicated a high level of congruence between the Surat
Thani black morph and the Surat Thani white/Vietnam group, implying that the black morph collected
from Surat Thani could be a variant of the white
type rather than a third morph/species. Principal component/co-ordinate analysis on the character means
plotted for all five groups produced the same result as
the CVA analysis, thus reinforcing the two morph/
species theory.

Discussion
Multivariate analysis of morphometric characters has
been shown to be a rapid and effective technique in
providing an insight into the discrimination of many
animal species, including various types of Crustacea;
moreover, the hard, well-defined body parts of crustaceans facilitate the collection of accurate data. For example, multivariate analysis of morphometric data was
used to show interspecific variation between two previously undescribed species of Procambarus spp. crayfish
discovered in a Mexican cave, confirming them as two
sympatric species (Allegrucci et al. 1992). Similar studies
revealed evidence of two species from four colour forms
of Liopetrolisthes mitra, a porcellanid commensal crab

59

Fig. 3 Scylla serrata. Details of spines and dentition (teeth) on

carapace (a) and both right and left chelipeds (b) forming meristic data
in multivariate analysis

(Weber and Galleguillos 1991), but the insignificance of

racial variation between different populations of the
mitten crab Eriocheir spp., from Southern China (Li
et al. 1993).
Canonical variate analysis has often been stated as
being a powerful multivariate technique in the identification of unknown species (Pimentel 1979; Quicke 1993).
Although there is evidence that CVA may give misleading results if the data do not conform to the mathematical assumptions (i.e. that there is homoscedasticity
between the within-group covariance matrices), the ordination of the population under analysis can be checked
by using principal component analysis/principal co-ordinates analysis (PCA/PCOA; Thorpe 1983). Moreover,
MGPCA on the raw data collected for Scylla serrata
indicated that size was not affecting the results.
The results show that CVA can discriminate between
different groups of Scylla serrata regardless of their
geographical origin. All 22 morphometric characters
were shown to have some contribution to this variance,
however, larger contributions were attributed to the
frontal length (the measurement between the two orbital
sockets), the right and left antero-lateral lengths, and a
significant proportion of the characters from the right
chelae. In a previous study, Chayarat and Kaew-ridh
(1984) demonstrated, using regression analysis, that
frontal length is wider in the white morph than in
other morphs from Ranong which they described as
red and green.

Fig. 4 Scylla serrata. Plots of first two canonical variables from

populations collected from four sites in Southeast Asia: Ranong,
Thailand (Site A), Surat Thani, Thailand (ST Site B), Can Gio,
Vietnam (Site C), and Sematan, Sarawak (Site D). Shown are
canonical variate analyses on morphometric data (a), on multiplegroup principle-component analysis (MGPCA) scores with size vector
removed (b), and on meristic data (c)

96.92
(90.67104.26)
4.12

96.00
(88.93104.96)
5.03

93.32
(87.01104.91)
8.01

94.68
(82.18102.47)
6.27

97.05
(82.48114.38)
8.21

ST black
mean
(range)
SD

Ranong
mean
(range)
SD

Vietnam
mean
(range)
SD

Sarawak
mean
(range)
SD

ICW

Character

Thailand
ST white
mean
(range)
SD

Group

33.19
(28.1437.70)
2.41

28.86
(25.1033.29)
1.96

33.13
(29.4637.61)
1.89

31.82
(28.4043.56)
1.90

29.29
(25.7131.34)
1.46

FL

45.09
(38.1054.10)
3.99

43.64
(38.7448.01)
3.00

44.75
(40.3249.38)
2.51

44.03
(40.4848.55)
2.44

45.02
(41.4249.00)
2.04

RC

44.89
(37.6948.88)
3.93

44.50
(38.4647.97)
3.04

43.67
(40.4448.51)
2.38

44.17
(40.0249.72)
2.83

45.35
(42.2150.31)
2.04

LC

11.00
(7.6816.81)
2.27

10.03
(7.4712.05)
1.27

12.45
(8.0818.25)
2.23

9.64
(7.5612.95)
1.57

10.13
(7.0613.44)
1.12

RDW

67.84
(53.1692.17)
10.26

60.68
(50.7770.67)
5.52

69.98
(57.3984.50)
7.52

61.50
(54.8570.71)
4.89

61.59
(45.7967.88)
4.47

RPL

29.62
(22.8040.04)
4.27

25.70
(21.1828.65)
2.05

31.18
(27.0335.43)
2.91

25.91
(20.1232.66)
3.37

26.34
(20.2328.59)
1.86

RCL

28.83
(22.5938.52)
4.60

25.12
(21.8328.44)
1.99

28.90
(24.1835.23)
3.23

26.86
(22.0129.82)
2.27

26.12
(22.5829.58)
1.55

LCL

23.69
(19.1828.58)
2.88

20.60
(17.3223.27)
1.52

24.36
(20.3928.74)
2.19

21.56
(18.3323.93)
1.59

21.23
(16.1923.09)
1.56

RMW

Table 1 Scylla serrata. Means, standard deviations (SD) and ranges for morphometric characters contributing significantly to between-group variance using canonical variate
analysis on multiple-group principle-component analysis scores for five groups of mud crab collected from four locations in Southeast Asia (ICW internal carapace width; FL
frontal length; RC right anterolateral length of carapace; LC left anterolateral length of carapace; RDW right dactylus width; RPL right propodus length; RCL right carpus
length; LCL left carpus length; RMW right merus width; ST Surat Thani)

60

61

The discrimination due to the measurements of the

right chelae indicates a variance due to handedness.
The group means of these characters suggest that the
right chelae is enlarged in the black morph type
whereas the chelae of the white morph were found to
be bilaterally symmetrical. In a study of handedness
in the brachyuran crab Cancer productus, phenotypic
plasticity due to environmental induction was demonstrated whereby claw size was found to increase in order
to eat hard-shelled diets (e.g. molluscs). There is a suggestion that bilaterally symmetrical species develop a
behavioural preference for one chelae, allowing it to
increase in size in such a manner that the induction of
this slight asymmetry becomes inheritable (Smith and
Palmer 1994). Thus, behavioural stimulation of handedness, especially with respect to dietary preferences,
merits closer examination in Scylla serrata.
Based on the measured characteristics, crabs from the
four locations show three discrete groupings with no
evidence of chain-linking. The white morph from
Surat Thani and the crabs originating from Vietnam
formed their own group, which was consistent with their
phenotypic likeness. Similarly, the Ranong and Sarawak
groups also formed their own cluster, while the third
grouping comprised of black morph crabs from Surat
Thani.
Even though this analysis indicates that groups of
Scylla serrata can be discriminated racially (irrespective
of geographical distribution), it is hard to know how
much of this observed variation is due to direct environmental induction (phenomic plasticity) and what is
genetically controlled. Heterogeneous coastal environments would be expected to have a significant influence
on the phenotypic expression of S. serrata; however two
well discriminated morphs were found in the same location (Surat Thani), suggesting that there is more than
environmental induction influencing the expression of
these two apparently sympatric morphs. Environmental
temperature is one of the main variables known to affect
morphometric features. The four locations have only a
9 latitudinal range, and all lie within a warm equatorial
region where the dominant climatic feature in the coastal
belt is the monsoons rather than temperature fluctuation. The average monthly temperatures at Ranong and
Can Gio, the most northerly sites, each vary by <4 C.
Moreover, the two sites furthest away from each other,
i.e. Ranong and Sematan, formed their own separate
group on the basis of morphology even though they
were the most geographically distant sites.
Different morphs of Scylla serrata have been described as inhabiting different niches within the mangrove coastal zone. The white type has been described
as being more subtidal and less likely to burrow than the
black type (Estampador 1949). However, in the case
of Surat Thani the two morphs occupy the same geographical area (Ban Don Bay), suggesting other barriers
to prevent interspecific matings. Segregation of morphs

has been noted in Pulicat Lake by Joel and Raj (1983)

where tandems were observed between male and female
S. serrata of the same morph type.
The separation of the Surat Thani black morph
into a third group was surprising, as phenotypically it
resembled the populations collected from Ranong and
Sarawak that were also of the black type. It could be
that the CVA discriminated between the predetermined
groups that are actually part of a cline. Thorpe (1983)
indicated that because CVA can discriminate between
predetermined categories, it does not mean that the
variation is absolute. A more extensive sampling programme around the eastern seaboard of Thailand may
help answer this question.
The meristic analysis conducted in this experiment
provides evidence for only two distinct morphs and not
three as illustrated by CVA on the morphometric data.
Meristically, the black morph from Surat Thani is
closely associated with the white Surat Thani group
and the group collected from Vietnam. Most of the
between-group variance underlying the discrimination is
attributable to the nodule found on the inner lateral
propodus. However meristics seem to be a weak form of
identification between species of Crustacea where spination and dentition on the chelae are the paramount
characters used. Ranges of spines have been indicated
for many species of Brachyura (e.g. Crane 1975). Spines
are often broken or are subject to ontogenic variation,
and increase in number as the animal gets larger or
older. Dentition is often worn down with usage, which
makes it difficult to count the teeth with any certainty.
Obviously morphometric data cannot be expected to
give all the answers alone. The fact that phenotypic expression is controlled by many external factors, e.g. diet
and habitat, indicates the need for an understanding of
the level of endogenous intervention in the observed
phenotypic variation of Scylla serrata. The taxonomy
and biology of S. serrata merits further study, since the
presence of more than one species or morph has major
implications for both the fisheries management and
aquaculture development of this crab. In addition, there
is a growing risk of genetic contamination of S. serrata
stocks [already a problem for certain other farmed
aquatic species, e.g. salmon (Altukhov and Salmenkova
1987)], due to the large-scale transportation of live mud
crabs not only from one area to another to support
aquaculture, but also between consumer and producer
countries (Ferdouse 1990; BOBP 1992).
Acknowledgements This study was funded by Contract No. TS3CT92-1052 awarded to the University of Stirling by the European
Community Science and Technology for Development Programme
(STD-3). We are also grateful to the Natural Environment Research Council (NERC) for scholarship funding to J.L Overton.
We particularly wish to thank staff of the Royal Thai Forestry
Department and Department of Fisheries, Thailand; Forest Department of the Ho Chi Minh City Agricultural Service, Vietnam;
and the Department of Agriculture, Sarawak; for their kind assistance during the fieldwork in Southeast Asia.

62

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