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Age, growth and reproduction of the white seabream Diplodus sargus sargus
(Sparidae) off eastern coast of Algeria

Article  in  Journal of Applied Ichthyology · February 2013

DOI: 10.1111/j.1439-0426.2012.02057.x

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 Journal of
Applied Ichthyology
J. Appl. Ichthyol. (2012), 1–7
© 2012 Blackwell Verlag, Berlin
ISSN 0175–8659
Age, growth and reproduction of the white seabream Diplodus sargus sargus
(Linneaus, 1758) off the eastern coast of Algeria
By W. Benchalel and M. H. Kara
Marine Bioresources Laboratory, University of Annaba – Badji Mokhtar, Annaba, Alge´ria
Summary
The objective of this study was to determine the basic popu-
lation-specific parameters necessary for fish stock assessment
in the area and to compare these with data from other Medi-
terranean regions. White seabream Diplodus sargus sargus
(n = 449) were caught along the Algerian coast between
December 2005 and December 2006. Total length ranged
from 12.2 to 34.6 cm, representing age classes between 0 and
10 years. Validity of the otolith readings for estimating age
and growth was supported using the back-calculation
method. The von Bertalanffy growth parameters for all fish
were calculated as: TL = 36.3 cm, k = 0.154 year 1, t0 =
0.488 year. The growth performance index (φ) showed a
relatively slow growth of the local population. The length-
weight relationship showed an isometric growth (b = 2.98;
r = 0.98). Subspecies Diplodus sargus sargus was character-
ized as being a proterandric hermaphrodite. Overall ratio of
males to females was 1 : 1.4, with males predominant in the
smaller size intervals and females in the larger ones. The
reproductive season extended from January to May, with a
March–April peak in spawning activity. Fifty per cent matu-
rity of the tested cohort was reached at a total length of
20.2 cm for males and 20.0 cm for females.
Introduction
The white sea bream, Diplodus sargus sargus (Linnaeus
1758), is an important demersal commercial sparid found
throughout the Mediterranean Sea (Fischer et al., 1987),
especially inhabiting rock, sand and seagrass beds (e.g. Posi-
donia oceanica) (Harmelin-Vivien et al., 1995; Sala and Bal-
lesteros, 1997; Vigliola and Harmelin-Vivien, 2001; Louisy,
2005). Commonly occurring at depths of ca 40–180 m (maxi-
mum 420 m; Bauchot, 1987), they congregate in schools of 5
–50 individuals and feed mainly on molluscs (Rosecchi, 1987;
Benchalel et al., 2010). Juveniles often live in coastal lagoons
and estuaries (<2 m) (Quignard and Man-Wai, 1983; Macph-
erson, 1998). Seabream is primarily a species of the artisanal
fishery that uses fixed gear (trammel and gillnets).
Age, growth and reproduction of D. sargus sargus in the
Mediterranean Sea were studied by several authors (El
Maghraby et al., 1981a,b, 1982; Man-Wai and Quignard,
1982, 1984; Quignard and Man-Wai, 1983; Man-Wai, 1985;
Gordoa and Moli, 1997; Mouine et al., 2007); however, their
investigations were performed in the north-western and
south-eastern areas of the Mediterranean. On the North
African coasts, only Mouine et al. (2007) focussed on the
reproductive biology, while Benchalel and Kara (2010) and
Benchalel et al. (2010) studied the morphology and diet of
this species, respectively.
Received: January 10, 2011
Accepted: February 7, 2012
doi: 10.1111/j.1439-0426.2012.02057.x
Part of a broader research programme devoted to coastal
sparids from Algeria (Chaoui et al., 2005, 2006, 2009; Derbal
and Kara, 2006, 2008; Derbal et al., 2007; Benchalel and
Kara, 2010; Benchalel et al., 2010), aim of the present study
was to identify several key parameters of Diplodus sargus sar-
gus needed for stock assessment (sex ratio, spawning cycle,
size at first maturity, age distribution and growth) off the
eastern coast of Algeria while also comparing the data with
studies in other areas of the Mediterranean.
Materials and methods
Samples of white seabream were collected monthly from
December 2005 to December 2006 along the Algerian East-
ern coasts (Fig. 1) via the wholesale fish merchants and fish-
mongers in Annaba town from catch of trawls and gill nets
of 45 mm stretched mesh. Total length (TL) was measured
to the nearest mm and total weight (TW) recorded to the
nearest mg. Sex and maturity stages were determined macro-
scopically, and gonad weight (GW) was recorded to the near-
est mg. Stages of maturation were classified as: Stage
1 = inactive (resting) (very thin translucent tubes); Stage
2 = developing (ovaries are swollen with orange tubes, eggs
are visible to the naked eye, testes are white and show some
thickening); Stage 3 = ripe (ovaries are very large, translu-
cent eggs are visible, testes are very large and white, sperm is
found in the testicular tissue and the main sperm duct); and
Stage 4 = post spawning (ovaries are flaccid, bloodshot and
small in size, testes are smaller in size and pinkish-grey in
colour) (Mann and Buxton, 1998). Otoliths sagittae were
removed, cleaned and stored for later analysis.
Age was estimated by interpreting growth rings on 241
otoliths from 98 females (TL: 15.4–34.6 cm; TW: 53.5–560 g)
and 143 males (TL: 11.2–33.0 cm; TW: 19.0–595 g). Whole
otoliths were fully immersed in sesame oil and read using a
compound microscope provided with a micrometer piece
(magnification 94), using reflected light by following standard
techniques. Readings were made by three independent investi-
gators; the otoliths were used in subsequent analyses only
when all three readings were in agreement. Marginal incre-
ment analysis was used to validate annual growth increment
formation (Beamish and McFarlane, 1983). The monthly
mean marginal increment (MI) was calculated using:
MI = (R Rn)/(Rn Rn 1) (R, Rn and Rn 1 are, respec-
tively, the radius, the radius of the last and the next-to-last
growth rings). Existence of a linear relationship was first tested
between the radius of the otolith and the total length. Next,
the age-length relationship was back-calculated using Lee
(1920). Lengths at different ages were compared with the
back-calculation results. Growth parameters of the von
2 W. Benchalel and M. H. Kara
Fig. 1. Map of study area showing geographical limits of Diplodus
sargus sargus fishing sites (▼), eastern coast of Algeria. Arrows indi-
cate geographical limits of fishing zones
Bertalanffy growth equation were estimated (Ricker, 1975):
TL = L∞ (1 e k(t t0)), where L∞ is the ultimate length that an
average fish would achieve if it continued to live and grow, k
is the growth coefficient which determines how fast the fish
approaches L∞, and t0 is the hypothetical age for TL = 0. The
software FISAT II (version 1.2.2) (Gayanilo et al., 1996) was
used to estimate length-weight relationship and growth
parameters. The relation of TW to TL was calculated applying
the exponential regression equation: W = aLb (Ricker, 1973).
The growth performance index Φ = Log (k) + 2Log (L∞)
(Pauly and Munro, 1984) was estimated in order to compare
growth parameter values obtained in the present paper with
those reported for other D. sargus subspecies.
A total of 449 specimens were examined for their reproduc-
tive biology. The numeric proportions of sexes were expressed
as male to female ratio, and tested with a chi-square
test (Dagnelie, 1975). The spawning season was determined by
analysing the temporal evolution in the relative frequency of
maturity stages and the gonado-somatic index (GSI) = (weight
of the gonades/gutted weight of fish 9 100, Anderson and Gu-
treuter, 1983). Monthly values of GSI were compared using a
one-way ANOVA test, completed by a multiple sample compari-
son of means (Dagnelie, 1970). Length at maturity was calcu-
lated by determining the proportion of reproductively active
fish (stage 3). Length at first maturity was estimated as the
length at which 50% of the fish had become mature (Lt50) (Ri-
khter and Efanov, 1976). The interval of sexual maturity ‘Im’
is defined as the interval of size corresponding to the pubescent
transformation (Loubens, 1980).
Results
Among the 241 individuals of D. sargus sargus used for age
study, 98 (40.6%) were females and 143 (59.3%) were males.
Comparing successive monthly mean marginal increment val-
ues of otoliths (Fig. 2) using mean comparison tests showed
a significant difference (F = 3.03; P < 0.05) between May and
June. Thus, the rings were considered to be annual incre-
ments. Minimum values of marginal increment were recorded
in June, which is the time of annulus formation for D. sargus
sargus in the study area. The equation expressing the linear
relationship between total length (TL) and the otolith radius
(R) is: TL = 66.9 R 18.1 (r = 0.9; P  0.001). The ordinate
to the origin of this equation (18.1 mm) corresponds to the
theoretical size of the fish at the time of formation of the
first otoliths. Age range of the sampled fish was 1–10 years,
1.5
n = 47
1.3 n = 18
n = 10 n = 22
1.1 n = 26 n = 13
n = 22
n = 17
AM (mm)
0.9
n=6 n = 10
0.7 n = 10 n = 25
n = 15
0.5
0.3
D 2005 J F M A M J J A S O N D 2006
Months
Fig. 2. Monthly evolution of marginal increment for all Diplodus
sargus sargus otoliths. n: monthly sample sizes
with a predominance of age classes of 3–5 years in the catch
(74.2% individuals). The oldest female was estimated at
10 years; the oldest male estimate was 9 years. However, it
has to be taken into account that the size ranges covered
may be limited by the mesh sizes employed by the fishermen.
Back-calculated lengths obtained by Lee’s equation are given
in Table 1. The estimated parameters of the von Bertalanffy
model and the growth performance index Φ are given in
Table 2. Length-weight relationships show isometric growth
for females (Wg = 2.720.10 5 L2:945
t , r = 0.980), males (Wg =
1.900.10 5 L3:009
t , r = 0.980), and both sexes combined
(Wg = 2.240.10 5 L2:987
t , r = 0.980).
A total of 449 fish were examined for their reproductive
state, of which 178 (39.5%) were females (TL = 15.4–
34.6 cm; TW = 51–746 g), and 262 (58.3%) males (TL =
11.2–33.5 cm; TW = 19–600 g), while 173 fish (38.5%) were
immature (TL = 11.2–24.0 cm; TW = 20–191 g) and 9 (2%)
were hermaphrodites (TL = 19.7–24.2 cm, TW = 115–211 g).
The total number of males sampled monthly was greater
than that of females (Fig. 3). Overall sex ratio was 1 : 1.4;
thus deviating from 1 : 1 (v2 = 16.03; P  0.001). Males
predominated in the smaller size classes compared to females;
intersexual individuals were intermediate in size between
males and females (Fig. 4).
Males and females with ripe gonads (stage 2) were recorded
between January and April, being dominant in February; ripe
and running males and females (stage 3) appeared in January
and became dominant in February; spent males and females
(stage 4) were recorded in April (Fig. 5). The highest GSI val-
ues occurred between January and May, with a maximum in
March–April, and a resting period from May to December
(Fig. 6). A one-way ANOVA applied to mean values of GSI indi-
cated their significant heterogeneity (males: F = 13.6; females:
F = 10.4; P  0.001). The curves of the GSI monthly values
showed that D. sargus sargus reproduced only once a year.
Male gonadic activity started in January and reached a maxi-
mum in April. The female gonadosomatic index was the high-
est in March, 1 month earlier than males. The spawning
period starts in March; in May or June the majority of fish are
post-spawners. Size at first maturity (L50) was estimated as
20 cm for females and 20.2 cm for males (Fig. 7). There was
no significant difference in length at 50% maturity between
sexes (tobs = 4.4; P  0.01). According to the otolith reading
of the concerned length classes, this size corresponds to an age
of 4 years. Thus, 39.2% of fish sampled do not reach the first
maturity length.
Age, growth and reproduction of the white seabream 3

Table 1
Diplodus sargus sargus back-calculated sizes-at-age, eastern coast of Algeria sampling campaign December 2005 to December 2006

Age 1 2 3 4 5 6 7 8 9 10

Females
n 98 98 95 61 33 16 10 8 6 2
M (cm) 10.4 14.2 17.3. 19.9 22.4. 24.3 25.8 26.9 27.9 30.1
SD 13.1 17.6 16.7 15.1 14.7 12.3 18.2 18.9 10.8 22.1
Males
n 143 139 119 79 43 19 13 7 2 –
M (cm) 10.4 14.0 17.3 19.7 22.2 24.2 25.6 27.0 28.3 –
SD 14.2 17.2 17.5 15.7 13.9 10.8 15.4 20.6 22.3 –
All fish
n 241 237 214 140 76 35 23 15 9 2
M (m) 10.4 14.1 17.3 19.8 22.7 24.5 25.7 27.4 28.7 30.1
SD 13.8 17.4 17.1 15.4 14.5 12.3 16.7 19.8 18.6 22.1

n, number of fish; M, mean length by age class; SD, standard deviation.

Table 2
Comparison of von Bertalanffy growth parameters and growth performance indexes (Φ), Diplodus sargus subspecies in different areas: all, all
fish; m, males; f, females; L∞, asymptotic length; k, growth rate; t0, theoretical age at length zero; Φ, growth performance index

Source Location Subspecies Method L∞ k t0 Φ

Man-Wai and Quignard (1982) NW Mediterranean D. sargus sargus Otoliths 46.70 0.12 1.63 2.42
Martinez-Pastor and Cantabrian Sea D. sargus sargus Otoliths all 48.4 0.18 0.58 2.63
Villegas-Cuadros (1996) m 52.9 0.13 3.73 2.56
f 57.6 0.10 5.33 2.52
Gordoa and Moli (1997) NW Mediterranean D. sargus sargus Otoliths 41.70 0.25 0.08 2.64
Mann and Buxton (1997) South Africa D. sargus capensis Otoliths 30.94 0.25 1.05 2.38
Pajuelo and Lorenzo (2002) Canary Islands D. sargus cadenati Otoliths all 47.3 0.14 1.97 2.49
m 45.9 0.14 1.91 2.46
f 49.4 0.15 2.05 2.56
Pajuelo and Lorenzo (2004) Canary Islands D. sargus cadenati Otoliths all 46.7 0.14 2.48 2.48
m 44.7 0.14 1.89 2.45
f 49.9 0.12 2.23 2.47
Abecasis et al. (2008) S. Portugal D. sargus Otoliths 40.93 0.18 1.28 2.49
Scales 39.55 0.15 1.87 2.37
Present study E. Algeria D. sargus sargus Otoliths all 36.39 0.15 0.49 2.30
m 35.14 0.16 0.43 2.29
f 35.41 0.16 0.60 2.30

Males Females Hermaphrodites

n = 34 n = 15 n = 49 n = 71 n = 40 n = 38 n = 29 n = 17 n = 34 n = 44 n = 7 n = 40 n = 31
100

90
80

70
Frequency (%)

60

50
40

30
20
10
0
D 2005 J F M A M J J A S O N D 2006
Months
Fig. 3. Seasonal variation in sex ratio of Diplodus sargus sargus during study period (December 2005–December 2006). n: monthly sampling
sizes

Discussion otoliths with hyaline edges observed at the end of the winter
Otolith marginal increment values suggest that only one and the beginning of spring could be related to physiological
growth ring is formed in June. The high percentage of stress during the spawning season (January–May).
4 W. Benchalel and M. H. Kara

Males Females Hermaphrodites

n=5 4 3 9 15 11 16 37 34 77 31 14 31 32 16 26 8 17 6 15 12 4 3 2
100
90
80
70
Frequency (%) 60
50
40
30
20
10
0
[11–12]
[12–13]
[13–14]
[14–15]
[15–16]
[16–17]
[17–18]
[18–19]
[19–20]
[20–21]
[21–22]
[22–23]
[23–24]
[24–25]
[25–26]
[26–27]
[27–28]
[28–29]
[29–30]
[30–31]
[31–32]
[32–33]
[33–34]
[34–35]
Length interval (cm)
Fig. 4. Sex ratio of Diplodus sargus sargus as a function of total length. n: monthly sampling sizes

Stage 1 Stage 2 Stage 3 Stage 4

(a) n = 18 11 28 47 21 15 15 11 21 18 4 13 22
100

90

80

70
Frequency (%)

60

50

40

30

20

10

0
D J F M A M J J A S O N D
2005 2006
Months

Stage 1 Stage 2 Stage 3 Stage 4

(b) n = 13 4 17 23 19 22 14 6 13 16 3 17 9
100

90

80

70
Frequency (%)

60

50

40

30

20

10

0
D J F M A M J J A S O N D
2005 2006
Months
Fig. 5. Maturity stages of Diplodus sargus sargus: (a) males; (b) females. Stage 1: inactive (resting); Stage 2: developing; Stage 3: ripe; Stage
4: post-spawning. n: monthly sampling sizes

Nevertheless, because these rings are present in juveniles as January. The high percentage of otoliths with an opaque ring
well as adults, they should also be related to the lowest val- during summer, which was attributed to a fast growth
ues of sea temperature with an annual minimum of 11°C in period, could be due to temporal variation in the food
Age, growth and reproduction of the white seabream 5

10
Males Females
9
8

7
6
GSI (%) 5

4
3

2
1

0
D2005 J F M A M J J A S O N D2006
Months
Fig. 6. Monthly variation in Diplodus sargus sargus gonadosomatic index (GSI), males (April peak, n = 21) and females (March peak,
n = 23), eastern coast of Algeria. Error bars: standard deviations

100 may result in the deposition of additional rings in the

Males Females otoliths, by which the ages might be overestimated. Age esti-
mation in South African sparids has moreover proved to be
75 difficult as a result of the phenomenon of stacking of growth
zones towards the otolith margin, particularly in older fish
% mature

(Buxton and Clarke, 1989, 1991, 1992; Smale and Punt,

50 20 cm 1991; Mann, 1992).
20.2 cm The von Bertalanffy parameters calculated in this study
show that white seabream is a relatively slow-growing fish
25
compared to other similar subspecies (Man-Wai and Quig-
nard, 1982; Martinez-Pastor and Villegas-Cuadros, 1996;
0
Gordoa and Moli, 1997; Abecasis et al., 2008) such as
D. sargus capensis (Mann and Buxton, 1997) and D. sargus
[11–12]
[12–13]
[13–14]
[14–15]
[15–16]
[16–17]
[17–18]
[18–19]
[19–20]
[20–21]
[21–22]
[22–23]
[23–24]
[24–25]
[25–26]
[26–27]
[27–28]
[28–29]
[29–30]
[30–31]
[31–32]
[32–33]
[33–34]
[34–35]

cadenati (Pajuelo and Lorenzo, 2002) (Table 2). Similarly,

Length interval (cm)
Fig. 7. Diplodus sargus sargus male and female sexual maturity
curves as a function of total length. Arrows indicate total length at
which 50% of fish are sexually mature
abundance (Benchalel et al., 2010) and the high sea tempera-
ture (24°C), as was also observed with D. cadenati (Pajuelo,
1997). Different findings have been recorded in studies on
other sparids in South Africa (Hecht and Baird, 1977;
Nepgen and De, 1977; Buxton and Clarke, 1986; Pulfrich
and Griffiths, 1988; Mann, 1992; Buxton, 1993). Marginal
increment analyses indicate that the opaque deposition,
which is representative of discontinuous growth, coincides
with the peak in gonad maturation.
The highest age estimated in this study was 10 years for
females and 9 years for males. These values compare well
with those reported for other subspecies in the Mediterra-
nean and the Cantabrian Sea. Gordoa and Moli (1997) and
Martinez-Pastor and Villegas-Cuadros (1996) examined 10-
and 12-year-old specimens of D. sargus sargus in the Medi-
terranean and D. sargus cadenati of the Canary Islands. This
indicates that the species has a relatively long life span
(12 years) (Pajuelo and Lorenzo, 2004). However, for D. sar-
gus capensis in South African waters of the Indian Ocean, a
maximum of 21 years was reported (Mann and Buxton,
1997). This difference may be due to insufficient and biased
sampling due to the limited artisanal fishing gear employed;
also overfishing may be a major reason that certain areas
always catch smaller fish. Otherwise, strong upwelling events
the growth performance index (φ) is the lowest compared to
those from other areas (Table 2) showing a relatively slow
growth in the local population.
The allometry coefficient of the length-weight relationship
for all fish: b = 2.98; P  0,01; r = 0.98) indicates isometric
growth. The same conclusion (b = 3.05) was reached by
Mouine et al. (2007) in the Gulf of Tunis, Man-Wai and
Quignard (1982) in the Gulf of Lion (b = 3.09), and Morato
et al. (2001) in the Azores (b = 3.05, females; b = 3.03, males
and 3.18 for both). However, variations may occur through
the influence of fishing gear, sample size and size ranges cov-
ered in the studies.
Hermaphrodites (2.0%) are few when compared to those
observed in the Gulf of Tunis (6.4%) for the same species
(Mouine et al., 2007) and in northern Spain (5.6%) (Marti-
nez-Pastor and Villegas-Cuadros, 1996). Combining the mac-
roscopic appearance of gonads and histological sexing,
hermaphrodites accounted for 23% of D. sargus along the
Egyptian coasts (El Maghraby et al., 1981a,b, 1982) and
50% in the Gulf of Lions (Man-Wai, 1985).
The presence of individuals with well-formed testes and res-
idues of degenerated ovaries (between 19.7 and 24.2 cm) and
the predominance of females in larger sizes generally confirm
a protandrous development of the local population. However,
the presence of males in the larger size classes could confirm
that sex inversion does not occur in all fish. Sexual succession
takes place mainly between 2 and 5 years of age. In the
Mediterranean Sea, a rudimentary hermaphrodism with a
facultative protandry was also observed (D’Ancona, 1949;
Tortonèse, 1965; Lissia-Frau, 1968; Man-Wai, 1985). Even in
6 W. Benchalel and M. H. Kara

Table 3
Spawning period and duration of Diplodus sargus as reported in various areas of the Mediterranean Sea, Atlantic and Indian oceans
Spawning
duration
Source Location Latitude Spawning period (months)

Man-Wai (1985) Gulf of Lion, France 43°N April–May 3

Joubert, 1981; Kwazulu-Natal, So. 30°S May–November 7
Africa
Martinez-Pastor and Asturias, Spain 43°N April–June 3
Villegas-Cuadros (1996)
Morato et al. (2003) Azores, Portugal 38°N March–June 4
F.J.O. Almada, unpubl. data Lisbon, Portugal 38°N February–May 4
University of Açores, unpubl. data Algarve, Portugal 37°N January–May 5
Mouine et al. (2007) Gulf of Tunis,Tunisia 36°N March–May 3
El Maghraby et al. (1982) Egyptian coasts 31°N January–April 4
Abou-Seedo et al. (1990) Kuwait 29°N November–March 5
Mann and Buxton (1998) South-east, So. Africa 34°S August–March 8
Mann and Buxton (1998) Eastern Cape, So. Africa 34°S June–December 7
Present study East Algerian coast 37°N March–May 3

captivity an optional protandry is argued (Micale and Perdi-
chizzi, 1994). However, for all others subspecies (D. sargus
capensis of South Africa, D. sargus kotschyi of the Persian
Gulf and D. sargus cadenati of northern Spain and the
Azores), the observation of both primary and secondary
females is explained by the digyny (Buxton and Garratt,
1990; Mann and Buxton, 1998).
Reproduction takes place between January and May, and
spawning extends from March to May with a peak in March–
April. In the Gulf of Tunis (36°N), reproduction takes place in
the same period (Mouine et al., 2007) and it extends for a
month (March–June) in the Azores, Portugal (38°N) (Morato
et al., 2003). As the latitude decreases, the spawning period
begins earlier and the duration increases from 3 to 5 months,
as shown in Table 3. These regional dissimilarities have
already been noted and correlated with changes in thermal
regimes (Morato et al., 2003). The size at first maturity (L50) is
estimated as 20 cm at an average age of 4 years; thus among
the 449 sampled fish 39.20% do not reach the first maturity
length, which is probably related to the method of fishing. The
size at first maturity is close to that reported in the Gulf of
Tunis (21 cm at 4 years) by Mouine et al. (2007) and in the
north of Spain (21 cm) by Martinez-Pastor and Villegas-
Cuadros (1996). This size is lower than in South Africa
(24.3 cm) (Mann and Buxton, 1998) and in the Lion Gulf
(23 cm, females; 20 cm, males) (Man-Wai and Quignard,
1984) but is higher than in the Azores (16.7 cm) (Morato
et al., 2003) and Egypt (18 cm) (El Maghraby et al., 1982).
Acknowledgements
The authors are thankful to the Algerian Ministry of Supe-
rior Education and Scientific Research, which financed this
study within the framework of the National Research Fund
(NRF).
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Author’s address: W. Benchalel, Marine Bioresources Laboratory,
University of Annaba – Badji Mokhtar, BP 230
Oued Kouba, Annaba 23003, Algéria.
E-mail: wafachalel@yahoo.fr