Marine Biology 71,187-192 (1982)
MARINE BIOLOGY
Factors Influencing Pot Catches and Population Estimates of the Portunid
Crab Scylla serrata
M. J. Williams * and B. J. Hill **
Queensland Department of Primary Industries, Fisheries Research Branch; P.O. Box 344, Fortitude Valley, Queensland 4006, Australia
Abstract
Crab pots were used to sample a population of Scylla ser-
rata (Forskal) in an estuarine area in Queensland, Austra-
lia. Pots were laid 100 m apart at fixed positions for 4 d
each month for 1 yr (April 1980-June 1981). Data from re-
capture of tagged crabs showed that males larger than
140 m m carapace width and females larger than 150 m m
had a higher capture probability than did smaller crabs;
thus size-frequency distributions based on crab-pot cap-
tures are biased. Spacing trials showed that pots positioned
50 m apart fished competitively but that there was no dif-
ference in catch between those placed 100 and 200 m apart.
Catch distribution indicated that the presence of a crab in
a pot reduced the probability of further captures. Tem-
perature and incidence of recently moulted crabs account-
ed for 66% of variation in monthly catches. Catch-per-unit-
effort (CPUE) data can be used as a measure of relative
abundance of adults if allowance is made for temperature
and the incidence of moulting. The capture-mark-release-
recapture method for making population estimates was
tested but it is concluded that, because of cost and bias in
collecting techniques, it cannot be recommended as a tech-
nique for studying S. serrata.
Introduction
The large portunid crab Scylla serrata occurs throughout
much of the inshore regions of the Indo-West Pacific
Ocean. Although in many areas the crabs are fished com-
mercially, there is little information on population dynam-
ics and especially on techniques which could be used for
* Present address: South Pacific Commission, P.O. Box D5,
Noum~a Cedex, New Caledonia
** Present address: C.S.I.R.O., Division of Fisheries, P.O. Box
120, Cleveland, Queensland 4163, Australia
9 Springer-Verlag 1982
estimation of parameters such as population structure and
density. The most practical method of sampling S. serrata
is by means of baited pots. This method relies upon a feed-
ing response by the crabs. Feeding by decapod crustaceans
is affected both by environmental factors such as tempera-
ture, and physiological factors such as moult condition. It
is desirable, therefore, that where a sampling technique
such as baited pots or traps are used, a second non-biased
method should be employed for determining the catcha-
bility of the crabs. For example, Morgan (1974) compared
diver catches with pot catches of rock lobsters (Panulirus
cygnus). Morrissy and Caputi (1981) drained or seined
ponds to obtain the entire population of marron (Cherax
tenuimanus) in order to estimate catchability of marron in
baited drop nets. In the case of S. serrata, it is not possible
to collect the crabs using divers because of the high turbidi-
ty in the muddy estuarine areas which are their main habi-
tat. S. serrata are seldom captured in trawls, apparently be-
cause they can bury or avoid trawls. Thus, at present there
appears to be no method which would provide unbiased
samples of the population. We therefore decided to direct
attention to factors influencing the catch rate and catch
composition of crab pots in order to determine their suit-
ability as a sampling technique for S. serrata. The capture-
mark-release-recapture (CMRR) method for population
estimation was also tested to determine whether this tech-
nique could be applied successfully to S. serrata.
Materials and Methods
The study site was in Pumicestone Passage (27~ 153~
a 30 km long and 1 to 5 km wide shallow channel between
Bribie Island and the Queensland mainland. The southern
end of the Passage opens into Moreton Bay, while the
northern end opens directly into the sea. The substrate in
the study area was soft mud, and the shore from the mid-
tide level upwards was covered with mangroves - chiefly
Avicennia marina. The shore had a gentle slope and, at low
0025-3162/82/0071/0187/$ 01.20
188 M.J. Williams and B. J. Hill: Pot Catches and Population Estimates of Scylla
tide, about 600 m of mud was exposed below the mangrove
line. A 36 ha permanent plot was marked out into
100 X 100 m squares in April 1980 using 49 stakes. The up-
per row of stakes was just outside of and parallel to the
mangrove line. In June 1980, the plot was extended down-
wards by two more rows to give a total of 63 stakes enclos-
ing an area of 48 ha. The lowest row was at a depth of
about 300 mm at low tide. Scylla serrata (Forskal) were
captured by means of crab pots baited with fish (usually
Mugil cephalus). Two types of pots were used in approxi-
mately equal numbers, firstly a collapsible pot 240 mm
high made of a rectangular steel frame (900 x 600 mm),
covered with 38ram mesh nylon net. The second pot
was a commercial design, it had a circular base of approxi-
mately 1 m diam, was 300 mm high and made of a steel
frame covered with wire mesh having 40 mm diam open-
ings. Both types of pots had two entrance funnels
(minimum opening 200 • 80 mm) at opposite ends. There
was no difference in the size and sex composition of crabs
caught in the different types of pots. Sampling was carried
out on the study site each month from April 1980 to June
1981, inclusive, with the exception of June 1980 and May
1981. On each occasion a baited trap was laid at every
stake and lifted and rebaited every 24 h for 4 successive
days, thus, after the first 2 mo, each sampling was based on
252 pot days. The catch from each pot was recorded sepa-
rately, with carapace width, sex and approximate moult
stage being noted for each crab. Crabs in Stages A2 to Ca in
Drach's (1939) scheme were recorded as having moulted
recently. The crabs were then tagged with individually
numbered Floy FD67B tags in the manner described by
Hill (1975). After tagging, each crab was released alongside
its capture site. Water temperature and salinity were re-
corded on each sampling day.
Effect of Size and Sex on Capture Probability
The effect of size and sex on capture probability was es-
timated by comparing the recapture frequency between
10 mm size classes of males and females. Months in which
recapture rates of all tagged crabs exceeded 20% were com-
bined to provide a sample sufficiently large to allow break-
down into sex and size classes. If it is assumed that the cap-
ture process does not affect probability of subsequent re-
capture, then the probability of recapture is an index of
probability of initial capture.
Pot Spacing Trials
A series of trials on the effects of trap spacing on catch
rates was conducted in February and March 1981 on the
grid. The configurations used were: 50 m spacing - 49 pots
in a 7 • 7 grid (2 replications); 100 m spacing - 63 pots in a
7• grid (8 replications); 200m spacing - 2 0 pots in a
5 x 4 grid (8 replications). Each test consisted of laying the
pots at the particular spacing and checking the catch after
24 h.
Frequency Distribution of Pot Catches
A power curve V=arn b was fitted to the mean (m) and
variance (V) of the pot catch frequencies (crabs pot -1) to
test for spacing patterns of crabs in pots. Taylor (1961) pro-
posed that the constant b was indicative of spacing when
the power curve was applied to quadrat counts of organ-
isms. In the present study, the catch of each pot was taken
as a "quadrat count".
Population Estimates
The population size within each month was estimated us-
ing the non-parametric model developed by Burnham and
Overton (1978, 1979). This model assumes that the popu-
lation is closed, that probability of capture may vary
among individuals but is constant over the sampling period
for any one individual, and that captures are independent
events. A test described by Burnham and Overton (1978)
was used for estimating day to day variation in capture
probabilities of individuals. Burnham and Overton (1978)
also provided a test for closure on the population and,
although this test does not detect immigration and emi-
gration during the middle stages of the experiment, in the
present study where t (number of capture times) is only 4,
the "middle stages" are confined to Days 2 and 3. Lack of
equal catchability among individuals, at least with respect
to size, precluded the use of many other population esti-
mation techniques such as the Jolly-Seber stochastic model
(Jolly, 1965; Seber, 1973) and the Manly-Parr model
(Manly and Parr, 1968). In the present study, population
estimates for a sampling period were not considered if the
probability of recapture within the 4 d period was less than
0.10 (as recommended by Otis et al., 1978 on the basis of
simulation studies), and/or the test for closure failed.
Burnham and Overton (1978) and Otis et al. (1978) have
found that the jacknife estimator used in this non-para-
metric method for estimating population size is quite
robust for day to day variation in capture probabilities of
individual animals provided that closure holds. The popu-
lation estimate for each month was based solely upon re-
capture data for crabs marked in that month, i.e., over a
4 d period. Crabs marked in previous months were not
used in population estimates.
Results
Factors Affecting Catches
Since population size estimates would refer only to that
part of the Scylla serrata population which was actively
feeding, we decided to use catch rather than catchability,
as the latter parameter requires a knowledge of the whole
population present in a given area. Only the months from
July 1980 to June 1981 were used since effort was constant
over this period. Salinity values (range 24 to 35%0) were not
M. J. Williams and B. J. Hill: Pot Catches and Population Estimates of Scylla 189

correlated with catches ( r = 0.09, n = 44). Daily water tem- were in the range 23 ~ to 29 ~ and recapture rates were
perature (7) was positively correlated with daily catches greater than 20%. In April 1981 the recapture rate dropped
(R=0.56, n = 4 4 ) . The percentage of recently moulted to 14% ( T = 2 5 ~ suggesting less frequent feeding pos-
crabs (M) was negatively correlated with catch (r=0.43, sibly due to late inter-moult condition among most of the
n = 4 4 ) . Neither mean size nor proportion o f large population.
(> 150 ram) crabs were correlated with catch. Large crabs
dominated the lowest catches (in June 1980), suggesting
that they were less affected by low temperature than were Effect of Size and Sex on Capture Probability
smaller crabs. Temperature (~ and percentage of re-
cently moulted crabs were fitted in combination in a re- The recapture percentage for each size class o f male and
gression model to explain the variation in catch. The mod- female Scylla serrata is given in Fig. 2. Sample sizes in
el, catch = - 2 3 . 1 8 9 + 3 . 6 8 8 T-1.199 M explained 66% of each class varied between 37 and 106 crabs, but in only 4
the variation in catch numbers for the period July 1980 to cases were there less than 50 individuals. The probability
June 1981, inclusive. Fig. 1 shows catch, T and M plotted of being recaptured increased in an approximately linear
against time. Day to day variation in catches was large in manner with size for males up to 130 m m and for females
some months, the coefficient of variation of daily catches up to 140 ram. Recapture probabilities for males 140 m m
varying from 3.8 to 42.8, this probably accounts for much of and larger and for females 150 m m and larger were in ex-
the residual variation in catches after the model was fitted. cess of 30% except for males of 170 m m and larger which
The proportion of tagged crabs recaptured within any one had a 23% change of recapture in the months studied.
month may be used as an indicator of the level of feeding
of those individuals which have positive vulnerability to
capture, i.e., those which are active and feeding to some ex-
Table 1. Scylla serrata, Effort (pot days), number of crabs caught
tent. Less than 10% o f marked crabs were recaptured in and tagged, and percentage of tagged crabs recaptured in study
months in which temperatures were below 20 ~ (May, Ju- area in each 4 d sampling period between April 1980 and June
ly, August 1980 and June 1981) (Table 1). In September, 1981. Population estimates and standard errors of the estimates
the recapture rate was only 6%, even though the tempera- are given for months in which recapture exceeded 10%
ture was 23 ~ At this time a large proportion of crabs
Date Effort No. % re- Population SE
were in late intermoult or pre-rnoult condition. In October, captured captured estimate
during the peak of the moulting period, few crabs were
caught, but the recapture rate of 13% indicated that vulner- April 196 295 26 438 22
able crabs were feeding more frequently than in the pre- May 196 124 6 - -
July 252 88 3 - -
vious months. From N o v e m b e r to March, temperatures
August 252 217 6 - -
September 252 232 6 - --
October 252 100 13 180 15
40
< November 252 228 29 339 20
~g ~o December 252 178 26 304 19
January 252 199 22 314 19
~ ao
February 252 312 21 505 24
10 March 252 319 26 501 24
April 252 300 14 571 26
June 252 191 3 - -

4(

3,=

"< 3(

~
350
CO
25
a 30(]
20
25e " " - , , , , ,

z 15 ~ .x~~ x ~
u. 200 C~
I
(.) ,,=, 1o
,~ 150 Q.
5
100

o 5o ~<100 120 140 160 >~180

CARAPACE WIDTH (ram)
A M J J A S O N D J F M A M J Fig. 2. Scylla serrata. Recapture percentage by size class and sex
MONTH
for all months in which total recapture rate exceeded 207o. Circles
Fig. 1. Scylla serrata. Total catch, temperature and percentage of = males; crosses = females. Largest size category for males =
recently moulted crabs for each month (July 1980-June 1981) 170 to 179 ram, for females 180 to 189 mm
190 M.J. Williams and B.J. Hill: Pot Catches and Population Estimates of Scylla

Pot Spacing Trials sults in bias in many methods of sampling, but especially
in methods relying upon feeding responses. In the present
Pots spaced 50 m apart caught fewer crabs (mean 1.0, SD
study, catches of Scylla serrata in pots varied both in the
0.06 crabs pot -~) than those spaced 100 m apart (mean 1.3,
short term (day to day) and in the longer term (month to
SD 0.15 crabs pot-l), suggesting that they were competing
month), even though pots were placed at the same site on
for crabs. This difference is significant (Student's t-test --
each day of an experiment. The main features of the longer
2.29) at the 5% level. There was no significant difference
term variation were depressed catches in June-July
between catches in pots spaced I00 and 200 m apart (mean
and again in October and to a lesser extent in
1.4, SD 0.34 crabs pot -1) (Student's t-test = 0.59). The
December-January. June and July are the coolest months
greater variability around the mean for catches from pots
in the study area and the lower temperatures could account
spaced at 200 m is probably due to the lower sample size at
for the lower catch. Hill (1980) found that both feeding and
this test distance. Mean distance between pot of capture
activity of S. serrata decreased markedly at temperatures
and pot of recapture (100m grid) was 253.6m (SD
below 20 ~ and Morrissy (1975) suggested that the active
145.9 m) for 129 crabs which were recaptured 24 h after in-
range of bait odour would be reduced at low temperatures.
itial trapping during the warm months (water temperatures
Subadult and adult S. serrata in the study area showed a
exceeding 20 ~ and overall recapture rate of crabs exceed-
high incidence of moulting in October and December, and
ing 20%). This is a minimum estimate of distance moved
decapods in premoult and during moult have been shown
over 24 h, since most crabs appear to move offthe intertid-
to cease feeding (Passano, 1960; Chittleborough, 1975).
al flat to the lower channel at low tide.
Since capture in pots relies upon a feeding response, any
factor which reduces feeding, such as low temperature or
Frequency Distribution of Pot Catches moulting, will result in decreased pot captures. A regres-
sion model fitted to temperature and moulting data ac-
The frequency distribution of crabs pot -1 for each month
counted for 66% of the long-term variability in catches.
was strongly skewed to the left because most pots captured
Mathematical relationships between catchability or the
none or 1 crab and only a minority caught more than 1
logarithm of catchability and combinations of temperature,
crab. The power curve relationship was V=0.9212 m 06396.
moult factors, salinity and relative size have been derived
The value of b was significantly less than 1 (p< 0.05,
for the rock lobster Panulirus cygnus by Morgan (1974) and
Student's t-test) indicating that the distribution of crabs in
for the marron Cherax tenuimanus by Morrissy and Caputi
pots was approaching a regular rather than a random
(1981). Catchability (q) is a preferred variable to catch in
(b = 1) or an aggregated (b > 1) distribution.
such explanatory regressions, since the population size is a
major variable affecting catches. In the case of S. serrata,
Population Estimates however, population numbers and hence q could be found
for only a limited number of months and, in these months,
Data from May to September in 1980 and June 1981 were
the population estimates were highly positively correlated
rejected for the purpose of population estimates because of
with catch. Since tag recaptures showed little evidence for
the low probability of capture (<0.10) (Table 1). In all
movement out of the vicinity of the grid over long periods,
cases for which population estimates could be made and
large fluctuations in catch numbers are due mainly to
using the maximum likelihood test given by Burnham and
changes in activity and feeding rather than large changes
Overton (1978), there was no significant (p < 0.05) variabil-
in population numbers.
ity in capture probabilities for individuals over the 4 d of
The distance which pots are laid apart can influence
each sampling period. Estimated population numbers
their catch, Sinoda and Kobayasi (1969) for example,
(Table 1) were highly correlated with number of crabs
found that when traps were laid in lines, those spaced 33 m
caught during the sampling period (r=0.96, n = 7), prob-
apart caught fewer crabs (Chionoecetes japonica) than
ably because the estimates relate only to that part of the
those placed further apart. Caddy (1979) estimated that the
whole population which is actively feeding at the time of
effective volume of influence of bait was approximately
trapping. In all cases, the adjusted third-order jacknife es-
1 824 ma(equivalent to a circle of radius 34.5 m if dispersal
timator was chosen as the best estimate of population size.
is restricted to within 0.5 m of the bottom). If this estimate
Standard errors of population estimates were low, with co-
is applied to our grid system, then traps 100 m apart do not
efficients of variation (CV) between 0.05 and 0.08. The val- compete for crabs. In the present study, pots placed 50 m
ues of CV of the population estimate in the present study apart appear to fish competitively but there was no consis-
are comparable to values obtained by Otis et al. (1978) in
tent improvement in catches with pots 200 m apart as com-
simulation studies using Burnham and Overton's (1978)
pared to those 100 m apart. In the case of units fishing non-
model. competitively, CPUE may be expressed as catch per pot for
comparison of relative catch numbers; however, for the
Discussion purposes of estimating population size, sampling units
should fish competitively (Morrissy, 1975).
Activity and particularly feeding activity levels can vary The frequency distribution of catch of Scylla serrata
both between individuals and on a temporal basis. This re- pot -1 suggested that entries to pots were not random
M. J. Williams and B.J. Hill: Pot Catches and Population Estimates of Scylla
events. Crabs tended to space regularly over the pots, in-
dicating that the presence of a crab in a pot reduced the
probability of further crabs entering that pot. This could be
expected, since S. serrata display agonistic behaviour
towards conspecifics. Miller (1980) noted frequent ago-
nistic encounters between crabs inside traps and those out-
side for Cancer productus, C. irroratus and Hyas araneus
and he suggested that such encounters reduced entry of
more crabs to a trap.
The present study has shown differential vulnerability
to capture among Scylla serrata of different sizes, since
subadult crabs were less vulnerable to capture in pots than
were adults. The size at which vulnerability increased
markedly was 130 m m for males and 140 m m for females.
Heasman (1980) found that while recently moulted females
less than 150 m m never entered commercial crab pots,
some recently moulted males as small as 120 m m did enter
pots. Large crabs of both sexes were found in all moult
stages in pots. Thus, at least with respect to the moult
stage, there is a difference between the sexes of subadult
crabs with respect to entry into crab pots. Behavioural dif-
ferences between the sexes and among size classes and
moult stages probably provide the mechanism for unequal
vulnerability of individual crabs to pot capture.
As far as commercial fishermen are concerned, pots
provide a simple and effective means of capturing crabs
which is enhanced by their bias toward large individuals.
The present study has shown however that catches of Scyl-
la serrata taken in crab pots are inadequate in representing
the relative size and sex composition of the total popu-
lation although they do provide a representative sample of
the adult (> 150 mm) portion. With the aid of tagging
studies some of the bias in pot catches can be determined.
A more accurate size-frequency distribution during sum-
mer months may be derived for crabs larger than 100 m m
by extrapolations based on recapture rates and the sample
size composition (Fig. 3). The derived composition differs
markedly from the sample composition and indicates that,
220 MALES FEMALES
2O0
i
180
16(]
ra 140
co
i the mud crab Scylla serrata (Forskal) in Moreton Bay,
<
(z 120
0 versity of Queensland 1980
100 ~ --
"0 iiiiiiiii!iiiiii-
20 iiiiiiiii!i!!iiiiiii!!
100 1t0 1'~0 1;30 1JJO150 1(i0 lJ/O ' I~IQ 110 120 1;~0 14.0 1]~0 160 1"}0 180 '
CARAPACE WIDTH (mm)
Fig. 3. Scylla serrata. Actual sample size-frequency distribution
collected from pots (stippled bars) and predicted population size-
frequency (open plus stippled bars) for crabs in summer months
191
contrary to the pot capture data, there were far more sub-
adult than adult crabs present in the study area. Fig. 3 il-
lustrates that accurate data on population composition
cannot be obtained solely from pot catches. Pot catches
can, however, be used as a measure of relative abundance
of adult crabs provided that allowance is made for tem-
perature and the incidence of moulting. Because popu-
lation structure of captured crabs varies depending upon
where the sample is taken, it would be advisable for pots to
be placed at the same position at each comparative sam-
pling.
The high correlation between population estimates de-
rived from CMRR experiments and sample catch nmnbers
suggests that CMRR experiments give little added in-
formation on relative population size for the extra effort re-
quired. CMRR experiments however, do indicate the ab-
solute size of that part of the population vulnerable to pot
capture and, in addition, marking of crabs provides valu-
able data on the degree of bias in the capture method.
Since this method is extremely time-consuming and there-
fore costly, its use as a standard method for studying popu-
lations of Scylla serrata cannot be recommended.
Acknowledgements. This work formed part of a program on
Scylla serrata funded by the Australian Fishing Industry
Research Committee. The assistance of P. Dutton, S. Hy-
land, P. Tucker and C. Matilda is gratefully acknowledged.
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Date of final manuscript acceptance: August 24, 1982.
Communicated by G. F. Humphrey, Sydney