J. Appl. Ichthyol.
23 (2007), 5359
 2006 The Authors
Journal compilation  2006 Blackwell Verlag, Berlin
ISSN 01758659
Population biology and assessment of the white-spotted spinefoot, Siganus
canaliculatus (Park, 1797), in the southern Arabian Gulf
By E. Grandcourt, T. Al Abdessalaam, F. Francis and A. Al Shamsi
Marine Environmental Research Centre, Environment Agency-Abu Dhabi, Abu Dhabi, United Arab Emirates
Summary
The age, growth, mortality, reproduction and resource status
of Siganus canaliculatus in the southern Arabian Gulf were
investigated using a combination of size frequency, biological
and size-at-age data. Dened structural increments consisting
of alternating translucent and opaque bands in transverse
sections of sagittal otoliths were validated as annuli. The
maximum absolute age estimate was 7.8 years. Parameter
values of the von Bertalany growth function t to size-at-age
data (males and females combined) were: k 1.0,
L 24.8 cm (LF), to )0.1 years. Fish in spawning con-
dition were only observed between April and July although
patterns in gonadosomatic indices suggested a second but less
well dened spawning event in November. The mean sizes and
ages at rst sexual maturity were 21.5 cm LF (1.9 years) for
males and 25.7 cm LF (2.1 years) for females. Fish were fully
recruited to the shery at a size (L100 19.7 cm LF) that was
smaller than the sizes at which sexual maturity was attained.
The annual instantaneous rate of shing mortality
(F 0.85 year)1) (0.261.44 year)1 95% CI) was consider-
ably greater than the target (Fopt 0.33 year)1) and limit
(Flimit 0.44 year)1) biological reference points, indicating
that the stock is overexploited.
Introduction
The white-spotted spinefoot, Siganus canaliculatus, is a mem-
ber of the family Siganidae, commonly known as rabbitsh. It
is distributed throughout the Indo-Pacic from the Arabian
Gulf to the Indo-Malay region, Western Australia and north
to Hong Kong and Taiwan (Randall, 1995). S. canaliculatus
occurs in coastal waters to depths of at least 40 m (Woodland,
1984). Juveniles form schools on algal and seagrass ats,
feeding mainly on lamentous algae. Adults are also schooling
and move into shallows with the rising tide to feed on benthic
plants (Randall et al., 1997).
Siganids are generally regarded as good food shes in
spite of their relatively small size. Some species have been
cultured because of their herbivorous food habits, rapid
growth and economic value (Randall et al., 1997).
S. canaliculatus is a commercially important species which
is caught with a variety of gear, including bottom trawls and
traps, in coastal waters throughout its range (Woodland,
1984). In the Arabian Gulf it is mainly caught using
inter-tidal fence nets and dome-shaped wire traps. During
2002, 149 tonnes of S. canaliculatus were caught in the
waters o the Emirate of Abu Dhabi in the United Arab
Emirates.
U.S. Copyright Clearance Centre Code Statement: 01758659/2006/23010053$15.00/0
Received: October 20, 2005
Accepted: February 15, 2006
doi:10.1111/j.1439-0426.2006.00796.x
Many of the demersal sh populations in the Arabian Gulf
have been heavily exploited and shing eort may be above
optimum levels for some species (Samuel et al., 1987; Siddeek
et al., 1999). The lack of appropriate data on most stocks
underscores the need to assess the regions sheries resources.
In this context, the goal of this study was to evaluate the status
of S. canaliculatus and provide biological reference points and
other pertinent information required for management. Specic
objectives included establishing key demographic parameters
using validated age estimates and identifying characteristics of
the reproductive biology of S. canaliculatus in these waters.
Materials and methods
Study site and sampling protocol
Size frequency data were collected from commercial catches of
S. canaliculatus made o the coast of the Emirate of Abu
Dhabi in the United Arab Emirates (Fig. 1) between October
2003 and September 2004. Fish were selected at random from
landings, lengths were taken using a measuring board and
recorded to the nearest centimetre fork length (LF). Biological
data were collected from specimens purchased from commer-
cial catches between October 2003 and September 2004.
Samples were obtained from 60 sh from a representative size
range during the rst week of each month. Juvenile sh were
sampled with a ne mesh beach seine net. Standard length (Ls),
fork length (LF) and total length (LT) were obtained using a
measuring board and recorded to the nearest millimetre.
Whole wet weight was measured using an electronic balance
and recorded to the nearest gram. Fish were sexed by
macroscopic examination of the gonad which was dissected
out and subsequently weighed to 0.1 g using an electronic
balance. The maturity stage was assessed using a developmen-
tal scale ranging from immature (stage I) to spent (stage V).
Sagittal otoliths were extracted, cleaned in water, dried and
stored in culture trays.
Age and growth
Thin transverse sections of sagittae approximately 100200 lm
thick were obtained by mounting the otoliths on glass slides
using thermoplastic glue and grinding to the nucleus with
1200-grit abrasive paper. Sections were examined under low-
power magnication and reected light against a black eld.
Three independent readings were made for each section, with a
minimum period of 2 weeks between each reading. The overall
precision was established using the index of average percent
error (APE) given by Beamish and Fournier (1981). Data were
www.blackwell-synergy.com
54
Fig. 1. Study site showing location of the Emirate of Abu Dhabi in the
southern Arabian Gulf
only included in the subsequent analyses if two or more
readings were in agreement.
In addition to the number of opaque bands observed in
transverse sections, the optical characteristic of the outer
margin (opaque or translucent) was recorded. The proportion
of samples with opaque/translucent margins was calculated for
each month and used to infer the timing and periodicity of
increment formation. The age at which the rst opaque band
formed was calculated as the time between the mean birth
date, derived from reproductive indices, and the time of
formation of opaque bands. Subsequently, the absolute
decimal age was calculated as the age at formation of the rst
band plus the number of opaque bands outside the rst and the
time between the formation of the last band and capture. In
order to establish the relationship of the timing of opaque zone
formation with trends in seawater temperature, data were
converted using the scaling process given in Newman and
Dunk (2002).
Growth was investigated by tting the von Bertalany
growth function (von Bertalany, 1938) to size at age data
using standard nonlinear optimization methods. The model
was t to pooled data and that for each sex separately. The
von Bertalany growth function is dened as follows:
Lt L(1)e)k(t)t0)) where Lt is length at time t, L is the
asymptotic length, k is the growth coecient and t0 is the
hypothetical time at which length is equal to 0. Growth
trajectories were compared between sexes using the analysis
of residual sum of squares (ARSS) method of Zar (1984)
modied by Chen et al. (1992). The rate of increase in size
with age between sexes was compared using a modied t-test
(Sokal and Rohlf, 1995). Dierences in the mean size at age
between sexes were determined using analysis of covariance
(ANCOVA) with loge age as the covariate, loge size the
dependent variable and sex as the xed factor. Parameters
of the length weight relationship were obtained by tting the
power function W a  LbF to length and weight data where:
W is the total wet weight, a is a constant determined
empirically, LF is the fork length and b is close to 3.0 for
species with isometric growth.
Reproduction
The mean size at rst sexual maturity (Lm) was estimated
for both sexes by tting the logistic function to the
proportion of mature sh in 1 cm (LF) size categories. The
E. Grandcourt et al.
mean size at rst sexual maturity was taken as the size at
which 50% of individuals were mature. The same procedure
was used to estimate the mean age at rst sexual maturity
(Am) using the proportion of mature sh in each year class.
Mean monthly gonadosomatic indices (GSI) were calculated
for each sex by expressing the gonad weight as a proportion
of the total body weight. The timing and frequency of
spawning were established by plotting the proportion of sh
by maturity stage and gonadosomatic indices against the
sample period. The mean birth date was estimated from the
period over which there was a sharp decline in the
gonadosomatic index for females. The population sexual
structure was examined using chi-squared goodness-of-t
tests. Independent tests were conducted to determine whe-
ther sex ratios diered signicantly from unity for the whole
sample and age categories within the sample. Juvenile
retention was calculated as the proportion of sh in
aggregated size frequency samples below the mean size at
rst sexual maturity.
Mortality
An age length key was developed with size-at-age data
following the method of Ricker (1975), this was used to
convert aggregated length frequency data into an age fre-
quency distribution. The annual instantaneous rate of total
mortality (Z) was subsequently determined using the age based
catch curve method (Beverton and Holt, 1957). The natural
logarithm of the number of sh in each age class was plotted
against the corresponding age and Z (95% CI) was
estimated from the descending slope of the best t line using
least-squares linear regression. Annual instantaneous rate of
natural mortality (M) was estimated using the empirical
equation derived by Rickhter and Efanov (1976) which
estimates M based on the age at which sexual maturation is
attained (tmass) as follows: M [(1.52/tmass)0.72])0.16. The
mean of the age at which 50% maturity was attained (Am)
for males and females (2 years) was used as the input to the
model. The annual instantaneous rate of shing mortality (F)
was calculated by subtracting the natural mortality rate (M)
from the total mortality rate (Z) derived from the age based
catch curve (F Z ) M). The calculation was also made for
the upper and lower 95% condence intervals for the estimate
of Z in order to derive a range of shing mortality rate
estimates. The exploitation rate (E) was calculated as the
proportion of the shing mortality relative to total mortality
(E F/Z).
Fishery assessment
Pooled length frequency samples were converted into a relative
age frequency distribution using parameters of the von
Bertalany growth function following the method of Pauly
(1983). The natural logarithm of the number of sh in each
relative age group divided by the change in relative age was
plotted against the relative age. Backwards extrapolation of
the descending limb was used to estimate probability of
capture data. A selectivity curve was generated by tting the
logistic function to probability of capture and size data which
was used to derive values of the sizes at capture at probabilities
of 0.5 (L50), 0.75 (L75), and the size at which sh were fully
recruited to the shery (L100). Resource status was evaluated
by comparing estimates of the shing mortality rate with target
(Fopt) and limit (Flimit) biological reference points (BRPs)
Population biology of Siganus canaliculatus 55

which were dened as: Fopt 0.5 M and Flimit 2/3 M, 1.0
following Patterson (1992). 0.9 Temp

0.8 % opaque
Results 0.7
0.6

Scaling units
A total of 763 biological samples were collected, ranging in size
from 16.0 to 33.2 cm LF (males) and 16.2 to 36.9 cm LF 0.5
(females). Juvenile sh, for which sex could not be determined, 0.4
ranged in size from 2.4 to 4.4 cm LF. A total of 3323 length
0.3
frequency samples were collected ranging in size from 15.0 to
37.0 cm LF. 0.2
0.1
0.0
Age and growth
0.1
Alternating translucent and opaque bands were observed in 0 1 2 3 4 5 6 7 8 9 10 11 12
transverse sections of sagittal otoliths when viewed with Month
reected light under low power magnication (Fig. 2). A total Fig. 3. Proportion of otoliths with opaque margins by month
of 608 sh was aged with an index of average percent error of (n 601) and monthly sea temperatures o the Emirate of Abu
4.7% for the entire sample. Data from 17 sh representing Dhabi. Values converted to standardized scale to enable comparison of
2.8% of the total sample were excluded from age-associated trends
analyses due to a lack of agreement between readings.
Absolute decimal age estimates ranged from 0.7 to 7.8 years
40
(males) and 0.8 to 7.8 years (females). Absolute age of
juveniles for which sex could not be determined ranged from 35
0.05 to 0.08 years and the age at formation of the rst opaque
30
zone was estimated at 1.0 years.
Fork length (cm)

One growth increment consisting of an opaque and trans- 25

lucent zone was formed on an annual basis with the opaque
20
band being deposited between April and July. There was an
apparent time lag of 1 month from the start of the increase in 15
seawater temperature and the commencement of formation of Females
10
the opaque band. The peak in the proportion of otoliths with Males
opaque outer margins occurred 1 month prior to the peak in 5 Unidentified sex
seawater temperature (Fig. 3).
0
The size at age relationship was asymptotic in form with the 0 1 2 3 4 5 6 7 8
majority of growth being achieved by the second year beyond Age (years)
which there was little increase in size with age (Fig. 4). The
Fig. 4. Von Bertalany growth function t to S. canaliculatus size-at-
analysis of residual sums of squares suggested dierential age data (females dashed line and males solid line)
growth characteristics between sexes (P < 0.05, F 3.12,
d.f. 664). The results of the modied t-test revealed that the
rate of increase in size with age was signicantly greater for Table 1
males (P < 0.05, t 2.38, d.f. 579) whilst the ANCOVA Parameters of the von Bertalany growth function, coecients of
indicated that females grew to a signicantly greater mean size determination (r2) and sample sizes (n) by sex for S. canaliculatus
at age (P < 0.05, F 4.12, d.f. 1). Parameters of the von
Bertalany growth function for each sex and pooled size at age Parameter All Males Females
data are presented in Table 1, and the age length key is given k 1.0 1.2 0.9
in Table 2. The length weight relationship (y 2)5*x3.0) L cm (LF) 24.8 23.5 25.5
provided a good t to length and weight data (r2 0.99). to (years) )0.1 )0.1 )0.1
r2 0.80 0.91 0.85
n 627 205 375

Fig. 2. Photomicrograph of transverse section through sagittal otolith
of S. canaliculatus showing growth increments; black dots mark
position of opaque bands (scale bar 1 mm)
Reproduction
Mean size at rst sexual maturity (Lm) was 21.5 cm LF for
males and 25.7 cm LF for females. The mean age at rst sexual
maturity (Am) was 1.9 years for males and 2.1 years for
females. Minimum sizes at which sh were observed in
spawning condition were 19.4 cm LF for males and 16.5 cm
LF for females. The gonadosomatic index for both males and
females increased between February and April and decreased
from April to July (Fig. 5). There was a pronounced peak in
the gonadosomatic index in April and a smaller peak in
November. Patterns in the proportion of sh by maturity
development stage also suggested that spawning took place
56 E. Grandcourt et al.

Table 2 100
Age length key for S. canaliculatus in the southern Arabian Gulf 90
80
Age class
70

Proportion (%)
LF (cm) 0+ 1 2 3 4 5 6 7 Total 60
50
16 3 5 8
40
17 16 14 30
18 1 35 16 2 54 30
19 4 30 18 3 55 20
20 1 31 15 16 1 64 10
21 28 34 16 2 80
22 23 24 18 3 68 0
Jan Feb Mar Apr May Jun Ju l Aug Sep Oct Nov Dec
23 18 26 32 4 1 81 Month
24 5 22 17 4 48 Stage I Stage II Stage III Stage IV Stage V
25 1 15 13 4 3 36 Fig. 6. Proportion of S. canaliculatus by maturity stage and month for
26 1 10 6 3 20 males and females combined
27 8 8 4 20
28 4 4
29 1 2 1 4
30 3 2 1 6 CI 0.922.10). Annual instantaneous rate of natural mortality
31 1 1 2 4 (M) derived from the Rickhter and Efanov (1976) equation
32 2 1 1 4 was estimated at 0.66 year)1. Annual instantaneous rate of
33 2 2
shing mortality (F) was 0.85 year)1 (95% CI 0.261.44) and
34 1 1
35 0 the exploitation rate (E) was 0.56.
36 1 1
n 6 196 208 142 27 5 1 5 590
Mean LF (cm) 19.4 20.8 22.1 23.7 25.0 28.0 31.7 Fishery assessment
SD (cm) 0.6 2.7 2.8 2.8 2.6 3.7 3.0
Values of the sizes where the probability of capture was 50%
(L50) and 75% (L75) were 15.0 and 17.1 cm (LF), respectively.
Fish were fully recruited to the shery at a size (L100
16 19.7 cm LF) that was smaller than the mean size at which rst
Females sexual maturity was attained for females (Lm 25.7 cm LT).
14 Males The annual instantaneous rate of shing mortality
(F 0.85 year)1) (95% CI 0.261.44) was considerably
Gonadosomatic index (%)

12
greater than the target (Fopt 0.33 year)1) and limit
10 (Flimit 0.44 year)1) biological reference points, indicating
that the stock is overexploited.
8

6 Discussion
4 sagittal otolith sections of S. canaliculatus extends the
2
0
1 2 3 4 5 6 7 8
Month
Fig. 5. S. canaliculatus mean monthly gonadosomatic indices (SE
for females)
between April and July, with sh in spawning condition only
being observed during this period (Fig. 6). However, spent sh
(both males and females) were recorded in January, suggesting
that a second spawning period may exist. The mean birth date
was estimated as 1 May. There was a signicant (P < 0.05)
female bias in the overall male to female sex ratio of 1:1.8 and
the female bias was signicant for all but the oldest age
category (Table 3). The proportion of sh in aggregated size
frequency samples that were below the mean size at rst sexual
maturity (juvenile retention rate) was 61.0%.
Mortality
The annual instantaneous rate of total mortality (Z) derived
from the age based catch curve (Fig. 7) was 1.51 year)1 (95%
The presence of structural growth increments in transverse
geographic and taxonomic range for which such observations
have been made. Moreover, validation of the timing and
annual periodicity of increment formation adds to a growing
9 10 11 12 body of evidence (e.g. Fowler, 1995) dismissing the perception
that annuli do not form in the otoliths of reef sh due to a lack
of seasonality in the tropics. Whilst the mechanisms of growth
increment formation are poorly understood, the deposition of
the opaque zone in tropical species generally occurs during the
summer in association with periods of accelerated growth,
whereas the translucent zone is formed when there is reduced
metabolic activity (Beckman and Wilson, 1995). The depos-
ition of growth increments observed here follows this general
pattern and the formation of the opaque band appears to be
closely associated with the increase in seawater temperature.
The southern Arabian Gulf exhibits marked seasonal vari-
ability in oceanographic characteristics with seawater temper-
atures exceeding 34C in summer and falling to 21C in the
winter (Sheppard et al., 1992). It is plausible that this extreme
temperature uctuation and variables such as productivity and
subsequent food availability are associated with seasonal
growth rate oscillations and the deposition of growth
increments.
The maximum absolute age for S. canaliculatus established
here (7.8 years) was close to the maximum age of 8 years
Population biology of Siganus canaliculatus
Table 3
Results of chi-squared goodness-of-t Males
tests on sex ratios within age categories
for S. canaliculatus. OBS, observed; Age category OBS
EXP, expected.
12 144
34 61
57 3 5.5
Total 208
8.0
r 2 = 0.96
7.0
n = 3312
6.0
ln Frequency
5.0
4.0
3.0
2.0
1.0
0.0
0 1 2 3 4 5 6 7
Age (years)
Fig. 7. Age-based catch curve for S. canaliculatus (y 1.51x +
10.68)
estimated for the marbled spinefoot (Siganus rivulatus) in the
Mediterranean, which was aged using annuli in scales
(Bilecenoglu and Kaya, 2002). The size-at-age relationship
was highly asymptotic in form with the majority of growth
being achieved by the second year beyond which there was a
comparatively small increase in size with age. The reduction in
the growth rate coincided with the mean age at sexual
maturity, suggesting a physiological shift from somatic growth
to reproductive development.
Siganus canaliculatus grows to a mean standard length of
8 cm in about 3 months, 10 cm in about 412 months and 14 cm
in 78 months (Lavina and Alcala, 1973). For a similar species
(Siganus sutor), high values of the growth coecient have also
been recorded: k 0.66 in Mauritius (Jehangeer, 1988),
k 0.87 in Kenya (Ntiba and Jaccarini, 1988) and
k 0.65 in the Seychelles (Grandcourt, 2002). The high rate
at which the asymptotic size is approached therefore appears
to be a general feature of the growth characteristics for
members of this family and is one of the main reasons for their
mariculture potential (Lam, 1974). Whilst the von Bertalany
growth function provided a reasonable description of the
overall growth trajectory, parameters could have been
improved by the addition of more juvenile sh between 5
and 15 cm LF and samples that were close to the maximum
reported size of 40 cm LT (Randall et al., 1997).
Sizes at rst sexual maturity of 18.5 cm LF and 18.0 cm
SL (females) have been reported for S. canaliculatus by
Sadovy (1998) and Soh (1976) from Hong Kong and
Singapore, respectively. The minimum size at which sh
were observed in spawning condition, 19.4 cm LF for males
and 16.5 cm LF for females, appear to be of the right order
by comparison with these results. Still, estimates of the mean
sizes at rst sexual maturity obtained here would have been
57
Females
Chi-square Chi-square Chi-square
EXP OBS EXP Total (males) (females) (total) P-value
204 264 204 408 17.94 17.35 35.30 <0.01
85 109 85 170 7.06 6.50 13.56 <0.01
8 5.5 11 1.64 0.73 2.36 0.12
381 589
improved by histological diagnosis of the maturity develop-
ment stage.
The dened spawning period of S. canaliculatus supports the
contention that seasonal reproductive cycles are common
among tropical shes (Robertson, 1990; Montgomery and
Galzin, 1993; Sadovy, 1996). Spawning for this species has
been reported to occur between January and April in both the
Philippines and Singapore (Soh, 1976; Jeyaseelan, 1998), and
Sadovy (1998) reported spawning of S. canaliculatus in Hong
Kong to occur between March and June. The 4-month main
spawning period determined here (April to July) corresponds
to the duration of the spawning period in other locations.
Whilst there were no sh recorded in spawning condition
during the winter months, spent individuals were observed in
January. Furthermore, there was a small peak in the gonado-
somatic index in November, suggesting that a second but less
well-dened spawning season exists. A second, although less
pronounced, spawning period has also been suggested for this
species in Singapore, the Philippines and Palau based on the
presence of juveniles and fry during two distinct periods of the
year (Lavina and Alcala, 1973). Additionally, Ntiba and
Jaccarini (1990) conclusively identied two major spawning
seasons for S. sutor in the coastal waters o Kenya. The
reproductive cycle of S. canaliculatus in the southern Arabian
Gulf therefore appears to be representative of the general
pattern observed at other locations throughout the Indo-
Pacic.
Female biased sex ratios are a characteristic (though not
diagnostic) feature of protogynous species (Sadovy, 1996).
Being dioecious, S. canaliculatus would not be expected to
have a sex ratio that diered signicantly from unity. The
apparent female bias could be due to the shing gear
dierentially targeting the larger females, and/or sex specic
habitat preferences with females being more abundant in areas
that are shed. It is also possible that errors occurred when
sexing sh in our study, in particular those that were
immature, further emphasizing the need for histological
diagnosis.
The age-based catch curve used for estimating the annual
instantaneous rate of total mortality (Z) relies on the assump-
tions that all of the age groups used in the analyses were
equally vulnerable to the shing gear and equally abundant at
recruitment. Failure of these assumptions may have intro-
duced errors and Z may have been overestimated if larger sh
were less vulnerable to the shing gear or, for example, if adult
sh underwent migrations. A survey of the biomass of
demersal species in the Arabian Gulf waters of the United
Arab Emirates showed seasonal changes in the abundance of
S. canaliculatus, with a dened peak during August (Shallard,
2003). Therefore, it is possible that the size and age compo-
sitions and associated estimates of Z may have been biased by
ontogenetic or spawning migrations.
Although empirical equations used for estimating the
natural mortality rate (M) may reect general relationships,
58
individual species may deviate substantially from the normal
pattern. Therefore, whilst the estimate of 0.66 year)1 obtained
here compared favourably to that of 0.63 year)1 for a very
similar species (S. sutor) in the Indian Ocean (Grandcourt,
2002), this parameter is potentially the greatest source of error
in our assessment.
As the size at which sh were fully recruited to the shery
(L100 19.7 cm LF) was smaller than the mean size at which
rst sexual maturity was attained for females (Lm 25.7 cm
LT), increases in mesh size regulations should be considered by
management authorities. This is particularly important given
that 61% of the yield in numbers consisted of sh that were
below the size at rst sexual maturity.
Gulland (1970) suggested that in an optimally exploited
stock, shing mortality should be about equal to natural
mortality, resulting in an exploitation rate of 0.5 year)1.
However, potential yields may be over estimated by a factor of
34 where F M (Beddington and Cooke, 1983). The
specied precautionary target (Fopt 0.5 * M) and limit
(Flimit 2/3 * M) values are considered to be appropriate
biological reference points, in particular given current man-
agement objectives which are aimed at stock re-building and
resource conservation. The shing mortality rate (F) of
0.85 year)1 (95% CI 0.261.44) was substantially greater than
both the target (Fopt 0.33 year)1) and limit (Flimit 0.44 -
year)1) biological reference points. This demonstrates that
growth overshing is occurring and suggests that reductions in
shing eort will be required in order to achieve resource
management objectives.
This study has established the demographic characteristics,
reproductive biology and status of S. canaliculatus in the
southern Arabian Gulf, and demonstrates the utility of
structural increments in sagittal otoliths as a foundation for
age based approaches to the assessment of tropical sheries
resources. The results suggest that the shery is overexploited
and that management interventions in the form of eort
restrictions and increases in mesh size regulations are required.
Acknowledgements
We are very grateful to Sultan Al Ali and Khaled Al Ali for
assistance with data collection. The Abu Dhabi Fishermens
Cooperative Society facilitated size frequency sampling. This
study was funded by the Government of the United Arab
Emirates through the Environment AgencyAbu Dhabi and
forms part of the activities of the Fish Landings and
Population Dynamics Project (no. 02-23-0008-05).
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Authors address: Edwin Grandcourt, Marine Environmental
Research Centre, Environment Agency-Abu
Dhabi, PO Box 45553, Abu Dhabi, United Arab
Emirates.
E-mail:egrandcourt@erwda.gov.ae