Marine Environmental Research Centre, Environment Agency-Abu Dhabi, Abu Dhabi, United Arab Emirates
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)
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
Fig. 2. Photomicrograph of transverse section through sagittal otolith
the gonadosomatic index in April and a smaller peak in
of S. canaliculatus showing growth increments; black dots mark November. Patterns in the proportion of sh by maturity
position of opaque bands (scale bar 1 mm) 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
The presence of structural growth increments in transverse
4 sagittal otolith sections of S. canaliculatus extends the
geographic and taxonomic range for which such observations
2
have been made. Moreover, validation of the timing and
annual periodicity of increment formation adds to a growing
0
1 2 3 4 5 6 7 8 9 10 11 12 body of evidence (e.g. Fowler, 1995) dismissing the perception
Month 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
Fig. 5. S. canaliculatus mean monthly gonadosomatic indices (SE
for females) 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,
between April and July, with sh in spawning condition only whereas the translucent zone is formed when there is reduced
being observed during this period (Fig. 6). However, spent sh metabolic activity (Beckman and Wilson, 1995). The depos-
(both males and females) were recorded in January, suggesting ition of growth increments observed here follows this general
that a second spawning period may exist. The mean birth date pattern and the formation of the opaque band appears to be
was estimated as 1 May. There was a signicant (P < 0.05) closely associated with the increase in seawater temperature.
female bias in the overall male to female sex ratio of 1:1.8 and The southern Arabian Gulf exhibits marked seasonal vari-
the female bias was signicant for all but the oldest age ability in oceanographic characteristics with seawater temper-
category (Table 3). The proportion of sh in aggregated size atures exceeding 34C in summer and falling to 21C in the
frequency samples that were below the mean size at rst sexual winter (Sheppard et al., 1992). It is plausible that this extreme
maturity (juvenile retention rate) was 61.0%. temperature uctuation and variables such as productivity and
subsequent food availability are associated with seasonal
growth rate oscillations and the deposition of growth
Mortality increments.
The annual instantaneous rate of total mortality (Z) derived The maximum absolute age for S. canaliculatus established
from the age based catch curve (Fig. 7) was 1.51 year)1 (95% here (7.8 years) was close to the maximum age of 8 years
Population biology of Siganus canaliculatus 57
Table 3
Results of chi-squared goodness-of-t Males Females
tests on sex ratios within age categories Chi-square Chi-square Chi-square
for S. canaliculatus. OBS, observed; Age category OBS EXP OBS EXP Total (males) (females) (total) P-value
EXP, expected.
12 144 204 264 204 408 17.94 17.35 35.30 <0.01
34 61 85 109 85 170 7.06 6.50 13.56 <0.01
57 3 5.5 8 5.5 11 1.64 0.73 2.36 0.12
Total 208 381 589
5.0
been reported to occur between January and April in both the
4.0 Philippines and Singapore (Soh, 1976; Jeyaseelan, 1998), and
Sadovy (1998) reported spawning of S. canaliculatus in Hong
3.0 Kong to occur between March and June. The 4-month main
spawning period determined here (April to July) corresponds
2.0 to the duration of the spawning period in other locations.
Whilst there were no sh recorded in spawning condition
1.0
during the winter months, spent individuals were observed in
0.0 January. Furthermore, there was a small peak in the gonado-
0 1 2 3 4 5 6 7 somatic index in November, suggesting that a second but less
Age (years) well-dened spawning season exists. A second, although less
Fig. 7. Age-based catch curve for S. canaliculatus (y 1.51x + pronounced, spawning period has also been suggested for this
10.68) 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
estimated for the marbled spinefoot (Siganus rivulatus) in the Jaccarini (1990) conclusively identied two major spawning
Mediterranean, which was aged using annuli in scales seasons for S. sutor in the coastal waters o Kenya. The
(Bilecenoglu and Kaya, 2002). The size-at-age relationship reproductive cycle of S. canaliculatus in the southern Arabian
was highly asymptotic in form with the majority of growth Gulf therefore appears to be representative of the general
being achieved by the second year beyond which there was a pattern observed at other locations throughout the Indo-
comparatively small increase in size with age. The reduction in Pacic.
the growth rate coincided with the mean age at sexual Female biased sex ratios are a characteristic (though not
maturity, suggesting a physiological shift from somatic growth diagnostic) feature of protogynous species (Sadovy, 1996).
to reproductive development. Being dioecious, S. canaliculatus would not be expected to
Siganus canaliculatus grows to a mean standard length of have a sex ratio that diered signicantly from unity. The
8 cm in about 3 months, 10 cm in about 412 months and 14 cm apparent female bias could be due to the shing gear
in 78 months (Lavina and Alcala, 1973). For a similar species dierentially targeting the larger females, and/or sex specic
(Siganus sutor), high values of the growth coecient have also habitat preferences with females being more abundant in areas
been recorded: k 0.66 in Mauritius (Jehangeer, 1988), that are shed. It is also possible that errors occurred when
k 0.87 in Kenya (Ntiba and Jaccarini, 1988) and sexing sh in our study, in particular those that were
k 0.65 in the Seychelles (Grandcourt, 2002). The high rate immature, further emphasizing the need for histological
at which the asymptotic size is approached therefore appears diagnosis.
to be a general feature of the growth characteristics for The age-based catch curve used for estimating the annual
members of this family and is one of the main reasons for their instantaneous rate of total mortality (Z) relies on the assump-
mariculture potential (Lam, 1974). Whilst the von Bertalany tions that all of the age groups used in the analyses were
growth function provided a reasonable description of the equally vulnerable to the shing gear and equally abundant at
overall growth trajectory, parameters could have been recruitment. Failure of these assumptions may have intro-
improved by the addition of more juvenile sh between 5 duced errors and Z may have been overestimated if larger sh
and 15 cm LF and samples that were close to the maximum were less vulnerable to the shing gear or, for example, if adult
reported size of 40 cm LT (Randall et al., 1997). sh underwent migrations. A survey of the biomass of
Sizes at rst sexual maturity of 18.5 cm LF and 18.0 cm demersal species in the Arabian Gulf waters of the United
SL (females) have been reported for S. canaliculatus by Arab Emirates showed seasonal changes in the abundance of
Sadovy (1998) and Soh (1976) from Hong Kong and S. canaliculatus, with a dened peak during August (Shallard,
Singapore, respectively. The minimum size at which sh 2003). Therefore, it is possible that the size and age compo-
were observed in spawning condition, 19.4 cm LF for males sitions and associated estimates of Z may have been biased by
and 16.5 cm LF for females, appear to be of the right order ontogenetic or spawning migrations.
by comparison with these results. Still, estimates of the mean Although empirical equations used for estimating the
sizes at rst sexual maturity obtained here would have been natural mortality rate (M) may reect general relationships,
58 E. Grandcourt et al.
individual species may deviate substantially from the normal Chen, Y. D.; Jackson, D.; Harvey, A. H. H., 1992: A comparison of
pattern. Therefore, whilst the estimate of 0.66 year)1 obtained von Bertalany and polynomial functions in modelling sh
growth data. Can. J. Fish. Aquatic. Sci. 49, 12281235.
here compared favourably to that of 0.63 year)1 for a very Fowler, A. J., 1995: Annulus formation in the otoliths of coral reef sh
similar species (S. sutor) in the Indian Ocean (Grandcourt, - a review. In: Recent developments in sh otolith research. D. H.,
2002), this parameter is potentially the greatest source of error Secor, J. M. Dean and S. E. Campana (Eds). University of South
in our assessment. Carolina Press, Colombia, SC, pp. 4563.
As the size at which sh were fully recruited to the shery Grandcourt, E. M., 2002: Demographic characteristics of a selection of
exploited reef sh from the Seychelles: preliminary study. Mar.
(L100 19.7 cm LF) was smaller than the mean size at which Freshwater Res., 53, 123130.
rst sexual maturity was attained for females (Lm 25.7 cm Gulland, J. A., 1970: The sh resources of the ocean. FAO Fish. Tech.
LT), increases in mesh size regulations should be considered by Pap. no. 97., FAO, Rome.
management authorities. This is particularly important given Jehangeer, M. I., 1988: Preliminary estimates of age, growth and
mortality of Siganus sutor from Mauritius. In: Proceedings of the
that 61% of the yield in numbers consisted of sh that were workshop on the assessment of the shery resources in the
below the size at rst sexual maturity. southwest Indian Ocean. M. J. Sanders, P. Sparre and S. C.
Gulland (1970) suggested that in an optimally exploited Venema (Eds). FAO, Rome, pp. 254259.
stock, shing mortality should be about equal to natural Jeyaseelan, M. J. P., 1998: Manual of sh eggs and larvae from Asian
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Lam, T. J., 1974: Siganids: their biology and mariculture potential.
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34 where F M (Beddington and Cooke, 1983). The Lavina, E. M.; Alcala, A. C., 1973: Ecological studies on Philippine
specied precautionary target (Fopt 0.5 * M) and limit siganid shes in southern Negros, Philippines. Abstract (no. MSS/
(Flimit 2/3 * M) values are considered to be appropriate ABS/2/1) submitted to the Marine Sciences Special Symposium,
Hong Kong, 714 December 1973.
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agement objectives which are aimed at stock re-building and deposits and condition of tropical surgeonshes (Teleostei:
resource conservation. The shing mortality rate (F) of Acanthuridae). Mar. Biol. 115, 529536.
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both the target (Fopt 0.33 year)1) and limit (Flimit 0.44 - and other population characteristics of the Red Emperor Snapper,
Lutjanus sebae (Cuvier, 1828), o the Kimberley Coast of North-
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