Journal of
Applied Ichthyology
J. Appl. Ichthyol. 29 (2013), 796–800
© 2013 Blackwell Verlag GmbH
ISSN 0175–8659
Reproductive biology of the annular seabream, Diplodus annularis
(Linnaeus, 1758), in the Gulf of Gabes (Central Mediterranean)
By H. Chaouch, O. Hamida-Ben Abdallah, M. Ghorbel and O. Jarboui
Institut National des Sciences et Technologies de la Mer (INSTM), Sfax, Tunisie
Summary
Annular seabream, Diplodus annularis (Linnaeus, 1758), were
caught off the coast of the Gulf of Gabes (Southern Tunisia,
Central Mediterranean) between April 2008 and March 2010
by commercial catches. With a total 2066 specimens the fish
ranged in size from 7.7 cm to 18.5 cm total length and from
7.5 g to 123.3 g in weight. Changes in biological parameters
(weight, length, gonadosomatic index, hepatosomatic index
and condition factor) were examined in order to provide
information on the spawning period and reproductive cycle
in the gulf. Overall sex ratio was 1 : 1.52 in favour of
females. The reproductive season extends from March to
June, with a spawning activity peak in May. Total length
at sexual maturity was 10.5  0.22 cm (males) and
10.6  0.3 cm (females). The results of this study could help
to establish a recommended minimum capture size.
Introduction
The annular seabream, Diplodus annularis (Linnaeus, 1758),
is distributed from the Gulf of Biscay to Gibraltar, around
the Madeira and Canary islands, as well as in the Mediterra-
nean and the Black Sea (Jardas, 1996). The habitat is
restricted mainly to beds of Posidonia. Published studies of
this species concern morphological characteristics (Tortonese,
1975), life history (Paz, 1975), age and growth (Girardin,
1978; Gordoa and Moli, 1997; Pajuelo and Lorenzo, 2001,
2002; Mati
c-Skoko et al., 2006), as well as feeding (Bell and
Harmelin-Vivien, 1983; Rosecchi, 1987; Pita et al., 2002).
The annular seabream D. annularis is an important target
species for Tunisian fisheries and particularly for those in the
Gulf of Gabes. However, the only available studies on this
species were those of Zribi (1974), El Arem (1980) and Bra-
dai (2000). Thus, the study objective was to update some life
history characteristics of the annular seabream, particularly
those related to reproductive biology, for possible consider-
ation in ecologically-based management strategies of the spe-
cies’ fisheries in the study area (Ben Mariem, 1993; Ghorbel
et al., 1997; Lassen and Medley, 2000; Ould, 2002; Cham-
mam, 2004; Hamida-Ben Abdallah et al., 2009).
Materials and methods
Fish samples were collected in the Gulf of Gabes (southeast
of Tunisia, Central Mediterranean), from Cape Kapoudia
(35th parallel) to the Tunisian-Libyan border (33°10′N)
(Fig. 1), from commercial catches at major landing ports.
Catches originated from different types of artisanal fishing
gear (gill nets and trammel nets). The samples were
U.S. Copyright Clearance Centre Code Statement: 0175-8659/2013/2904–796$15.00/0
Received: October 24, 2012
Accepted: December 22, 2012
doi: 10.1111/jai.12162
transferred to the laboratory where they were weighed to the
nearest 0.1 g and measured (total length) to the nearest
0.1 cm. After dissection, the gonads and livers were weighed
to the nearest 0.01 g using a digital balance. Gonads were
examined macroscopically for sex and maturity, with stages
classified after Holden and Raitt (1975): I, immature; II,
resting; III, ripe; IV, ripe and running; V, spent. Sex ratio
was calculated as the percentage of females.
In order to determine the length at first maturity, only
individuals collected during the spawning season were used,
employing the relationship between the length class (L) and
the percentage of mature fish in each length class. This rela-
tionship is described by the logistic function by using the
software FSAS (Saul et al., 1988). The equation used was:
P = 1/(1 + e a(TL TL50). Where, P is the proportion of
mature individuals; TL, total length corresponding to the
proportion (p); a, the constant; and TL50 the total length of
50% mature individuals.
The reproductive period was established by monthly deter-
mination of the gonadosomatic index [GSI = gonadal
weight 9 100/eviscerated weight (g)]. The hepatosomatic
index (HSI) and the condition factor (CF) were also
calculated monthly:
HSI = liver weight (g) 9 100/eviscerated weight (g)
CF 9 eviscerated fish weight 9 100/total length3 (cm)
Differences in the reproductive variables (RGS, RHS and
CF) were statistically analysed (ANOVA, per months, followed
by Tukey’s post hoc test P < 0.05).
Results
Diplodus annularis individuals (n = 2066) examined in this
study ranged from 7.7 to 18.5 cm TL. They were divided
into 817 (39.55%) males and 1249 (60.45%) females. The
observed sex ratio per size class is given in Table 1. Overall
sex ratio was 1:1.52 (v2 = 90.33 > v2Th = 3.84), indicating a
predominance of females in which the percentage increased
progressively with the total length and was clearly higher at
size  16 cm (Fig. 2).
The first sexual maturity among the males and females of
D. annularis was determined from 851 individuals (348 males
and 503 females). The change in the proportion of the
mature individuals showed at a size <9 cm TL that no indi-
vidual was mature; at TL > 14 cm, all observed individuals
were mature. The proportions of the mature individuals are
grouped in Table 2. The results obtained by the application
of the logistic function showed that the length corresponding
to a proportion of 50% sexually mature individuals was
10.5  0.2 cm and 10.6  0.3 cm TL, respectively, for males
Reproductive biology of Diplodus annularis 797

Fig. 1. Geographical position of Gulf of Gabes, Tunisia

Table 1 Table 2
Diplodus annularis sex ratio in relation to fish size, Gulf of Gabes Variation in proportions of mature individuals (%) according to size,
(April 2008–March 2010) Diplodus annularis males and females, Gulf of Gabes (April 2008–
June 2008; March 2009/2010–June 2009)
TL (cm) Males Females Total % (F) % (M) v2Cal
Males Females
8 27 28 55 50.9 49 0.018
10 160 178 338 52.6 47.3 0.959 Mature Total Mature
12 450 665 1115 59.6 40.3 41.457 TL (cm) Total (n) Mat. fish (%) (n) Mat. fish (%)
14 158 332 490 67.7 32.2 61.788
 16 22 46 68 67.6 32.3 8.470 8 7 0 0 8 0 0
9 19 6 0.32 19 7 0.37
10 28 14 0.50 31 16 0.52
Males Females 11 59 37 0.63 71 41 0.58
12 114 80 0.7 119 81 0.68
100
13 61 49 0.8 122 86 0.70
90
14 31 27 0.87 82 66 0.80
80 15 22 22 1.00 32 31 0.96
70 16 4 4 1.00 12 12 1.00
Frequency (%)

60 17 3 3 1.00 6 6 1.00
50 18 1.00 1 1 1.00
40
Mat, mature individuals; % mat, percentage of mature individuals.
30
20
10 showed that liver is the organ in which lipid reserves are
0 accumulated.
8 10 12 14 ≥16
TL (cm)
Fig. 2. Frequency of Diplodus annularis females (n = 1249) and Discussion
males (n = 817) in relation to size classes (total length, TL), Gulf of
Gabes (April 2008–March 2010) In the Sparidae family, both forms of sequential hermaphro-
ditism (protogyny and protandry) have been found, as well
as rudimentary hermaphroditism leading to secondary go-
and females (Fig. 3). No significant difference in length at nochorism, which is characteristic of all species of the family
first maturity was found between sexes (Buxton and Garratt, 1990). Also, Micale and Perdichizzi
(v2 = 2.35 < v2Th = 16.91 for males; v2 = 4.56 < v2Th = 18.30 (1994) noted that sex inversion in a single individual is a very
for females). common phenomenon in the family Sparidae.
As shown for females (Fig. 4), the monthly variations in Diplodus is one of the genus in the family Sparidae in
the somatic index showed a stage of pre-maturation in which hermaphroditism has been reported (Buxton and
March, followed by a fast maturation, a phase marked by a Garratt, 1990). Historically, Diplodus annularis has been
considerable increase of the GSI from April to May (ANOVA, described as protandrous (Tortonese, 1975; Bauchot and
Tukey’s post hoc, P < 0.05, dl = 622.00). The spawning Hureau, 1986) and as a rudimentary hermaphrodite (Bini,
phase manifested by a fast fall in the GSI occurs in May– 1968). Recently, Pajuelo and Lorenzo (2001) determined
June. The last phase (sexual rest) extends from July to Feb- D. annularis of the Canary Islands as protandrous. There is
ruary. For males (Fig. 5), the reproductive season much confusion regarding the reproductive style expressed
extends from March to June, with maximal gonadal activity by annular sea bream. In this study, we noted the absence of
in May–June. The GSI follows the same pattern in both hermaphrodite individuals having both male and female
sexes, showing higher values for females than for males. gonad tissues, based on macroscopic evaluation of the
The analysis of the monthly evolution of the hepatosomat- gonads and suggesting that D. annularis in the Gulf of Gabes
ic index (HSI) and condition factor (CF) for both sexes is a gonochoric species. In the same area, Bradai (2000)
798
Fig. 3. Size at 50% sexual maturity, Diplodus annularis males
(n = 348) and females (n = 503), Gulf of Gabes (April 2008–June
2008; March 2009/2010–June 2009). Data points = observed values;
lines = 50% maturity values
Fig. 4. Monthly variations in gonadosomatic index (GSI), hepatoso-
matic index (HSI), and condition factor (CF), Diplodus annularis
females, Gulf of Gabes. Data points = means; bars = standard devi-
ation
determined rudimentary hermaphroditism for this species.
The reasons are still unknown as to why the same fish spe-
cies exhibit different sexual strategies in different areas. Van
der Walt and Mann (1998) thought that some sparids fall
into the category of partial protandrous hermaphroditism. In
that case, sex can be expressed as caused by behavioural or
demographic alterations within a fish’s social system (Shap-
iro, 1989).
H. Chaouch et al.
Fig. 5. Monthly variations in gonadosomatic index (GSI), hepatoso-
matic index (HSI), and condition factor (CF), Diplodus annularis
males, Gulf of Gabes. Data points = means; bars = standard devia-
tion
The overall ratio of males to females was 1 : 1.52 in
favour of females. The predominance of females has also
been observed for this species in the same area (1 : 1.86)
(Bradai, 2000), but was unbalanced in favour of males
(1 : 0.79) in the coastal waters of the Canary Islands (Mati
c-
Skoko et al., 2006).
The size at first maturity was 10.5 for males and 10.6 cm
for females. These results are in accordance with those of
Bradai (2000) (10.4 cm) for the species in the Gulf of Gabes.
Pajuelo and Lorenzo (2001) found that males from the Can-
ary Islands reach sexual maturity at a smaller size (10.3 cm)
than females (12.8 cm). However, Santos et al. (1998) found
size at first maturity to be 13.4 cm. The sizes at first maturity
estimated in the present study are slightly smaller than the
current minimum legal landing size in Tunisia
(MLS = 11 cm). However, a minimum size at capture of
about 12.5 cm is generally recommended, allowing at least
75–80% of the population to reproduce. A revision of the
current legal size should be considered, with is a 2 cm safety
margin from size at first maturity.
In this study the reproductive season of the annular sea
bream extended from March to June, with maximal gonadal
activity in May. This developmental pattern confirms the
results of Bradai (2000) for this species in the Gulf of Gabes.
In Tunisian waters, Zribi (1974) identified the spawning per-
iod to be between May and July. In the Gulf of Tunis, El
Arem (1980) indicated that the spawning period took place
in April–May. It is also interesting to compare our results
with those of authors who reported from different areas of
the Mediterranean (Table 3). Data in the literature showed
some variation in the spawning time for D. annularis. When
considering all areas, spawning occurred over a prolonged
period, January–September. According to Wootton (1990),
temperature appears to be the most important environmental
factor influencing fish reproduction. Temperature is a funda-
mental physical regulatory factor in the lives of fishes; this
effect is expressed particularly strongly in the control of all
reproductive processes, from gamete development and matu-
ration, ovulation and spermiation, spawning and hatching,
to larval and juvenile development and survival. In reproduc-
tively mature adults, temperature is generally considered
to be a secondary cue to the photoperiod in phasing
reproductive seasonality but has a major role in synchroniz-
ing the final stages of reproductive maturity, as well as in
truncating reproductive episodes (Pankhurst and Porter,
2003). The effects of temperature can be expressed differentially,
Reproductive biology of Diplodus annularis
Table 3
Spawning periods of Diplodus annularis in different areas of the Mediterranean Sea
Spawning period Area
April–June Atlantic and western Mediterranean
February–April Eastern Mediterranean
April–July Italian coasts
April–August Italian coasts
May–August Northern Adriatic
Entire summer Adriatic Sea
End of April end of August Eastern Adriatic coast
April–June Northern Adriatic
April–August Izmir Bay, Aegean Sea
April–August Algarve (South Portugal)
May–June Gulf of Cadiz (Spain)
May–July Gulf of Lion (France)
May–June Gulf of Tunis
April–May Gulf of Tunis
May–June Gulf of Tunis
May–June Gulf of Gabes (Tunisia)
depending on when the spawning normally occurs in
the annual thermal cycle, with rising spring temperatures
required to cue maturation in spring and early summer spaw-
ners (Stacey, 1984; Scott and Pankhurst, 1992; Shimizu,
2003), but with the elevated temperatures delaying onset of
maturation and ovulation in autumn-spawning species
(Pankhurst and King, 2010).
In this study, the sexual cycle periods were determined by
analysing the monthly evolution of the gonadosomatic index.
GSI is often considered as a surrogate of reproductive effort
(Jakobsen et al., 2009) and can be used to assess the effects
of fishing on the life-history strategies of fish (Gunderson,
1997). As the GSI tends to increase with fish length (Shatu-
novskii and Ruban, 2009), intense fishing activity and the
removal of larger individuals may reduce the reproductive
potential of a stock (Sadovy, 2001; Conover and Munch,
2002). Reproductive effort, as approximated by a GSI
increase, may be increased in areas or periods of higher fish-
ing mortality (Hutchings, 2002).
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Author’s address: Houda Chaouch, Institut National des Sciences et
Technologiesde la Mer (INSTM), BP 1035 – 3018
Sfax, Tunisia.
E-mail: houdachaouch@yahoo.fr