ANNALES · Ser. hist. nat. · 30 · 2020 · 1
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received: 2020-05-18                 DOI 10.19233/ASHN.2020.03
CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN 
TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA)
Mohamed Mourad BEN AMOR, Marouène BDIOUI & Khadija OUNIFI-BEN AMOR 
Institut National des Sciences et Technologies de la Mer, port de pêche, 2025 La Goulette, Tunisia
Christian CAPAPÉ
Laboratoire d’Ichtyologie, Université de Montpellier, case 104, 34095 Montpellier cedex 5, France
e-mail: capape@univ-montp2.fr
ABSTRACT
The paper examines three large predatory sharks captured in the north-eastern Tunisian coast and the cen-
tral Mediterranean Sea. These three species are bluntnose sixgill shark Hexanchus griseus (Bonnaterre, 1788), 
smalltooth sand tiger shark Odontaspis ferox (Risso, 1810), and great white shark Carcharodon carcharias 
(Linnaeus, 1758). The distribution of the three species is detailed and their ecological role commented.
Key words: Chondrichthyes, Hexanchus griseus, Odontaspis ferox, Carcharodon carcharias, distribution, 
Tunisian coast 
CATTURE DI GRANDI SPECIE DI SQUALI LUNGO LA COSTA TUNISINA 
NORD-ORIENTALE (MEDITERRANEO CENTRALE)
SINTESI
L’articolo esamina tre grandi squali predatori catturati lungo la costa nord-orientale della Tunisia e nel Mediter-
raneo centrale. Si tratta dello squalo capopiatto Hexanchus griseus (Bonnaterre, 1788), del cagnaccio Odontaspis 
ferox (Risso, 1810) e del grande squalo bianco Carcharodon carcharias (Linnaeus, 1758). Gli autori presentano nel 
dettaglio la distribuzione delle tre specie e commentano il loro ruolo ecologico.
Parole chiave: Chondrichthyes, Hexanchus griseus, Odontaspis ferox, Carcharodon carcharias, distribuzione, 
costa tunisina
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Mohamed Mourad BEN AMOR et al.: CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA), 15–24
INTRODUCTION
The northern Tunisian coast constitutes a transition 
path for fish species between the western and eastern 
Mediterranean Basins, and mainly for large predatory 
sharks (Quignard & Capapé, 1971; Capapé, 1989). 
This opinion has been corroborated by recent reports 
of captures occurring in this same area (Rafrafi-Nou-
ira et al., 2015, 2019; Ben Amor et al., 2016; 2019; 
Capapé et al., 2018).
Investigations regularly conducted concomitantly 
in Tunisian waters in the wake of local assistance of 
experienced fishermen have allowed the collection of 
a shoal of bluntnose sixgill shark Hexanchus griseus 
(Bonnaterre, 1788), a specimen of the rare smalltooth 
sand tiger Odontaspis ferox (Risso, 1810), and a great 
white shark Carcharodon carcharias (Linnaeus, 1758). 
This paper provides information and data about the 
capture of the three species, occurring during com-
mercial surveys off the northern Tunisian coast, and 
some comments about the status of these species in 
the capture areas, and inside and outside the Medi-
terranean Sea.
MATERIAL AND METHODS
Each species is separately presented, including data 
of fishing gear, capture site, depth, nature of bottom 
and, when possible, associated fauna (Fig. 1). The 
specimens were rapidly cut into slices and sold soon 
after landing and it was generally difficult to get for 
each specimen its size, total length (TL) and total body 
weight. Photographs were taken to confirm the authen-
ticity of these captures.
RESULTS 
Bluntnose sixgill shark Hexanchus griseus 
(Bonnaterre, 1788)
This species is distributed worldwide, from the 
Pacific to the Indian Ocean, and off both sides of 
the Atlantic Ocean (Cook & Compagno, 2005). It is 
known throughout the Mediterranean Sea, in both ea-
stern and western basins, and commonly collected in 
certain areas (Capapé et al., 2003, 2004; Kabasakal, 
2006, 2013; Basusta & Basusta, 2015).
Catches of Hexanchus griseus were previously 
cited in northern Tunisian areas at the level of Eskerkis 
Bank (Capapé, 1989; Rafrafi-Nouira et al., 2015) and 
southwards, mainly in the Gulf of Gabès (Bradaï et 
al., 2002). Additionally, single specimens of H. griseus 
were sporadically captured by trawl targeting Euro-
pean pilchard Sardina pilchardus (Walbaum, 1792), 
approximately at a depth of 200 m, in the Gulf of Tunis. 
On 10 June 2019, a shoal of 21 H. griseus was landed 
at the fishing port of Kélibia, located in the north of the 
Cape Bon Peninsula, northeastern Tunisia, by bottom 
longline targeting groupers. The captures occurred in 
the Strait of Sicily between Marettimo Island, close to 
the northwestern Sicilian coast, and the northeastern 
Tunisian coast, at a depth of 700-1000 m approxima-
tely (Ben Amor et al., 2019).
Fig.  1: Map of the central Mediterranean Sea, indicating 
capture sites of the three large sharks presented in this 
paper. Hexanchus griseus: 1. Capture from Ben Amor 
et al. (2019). 2. Capture occurring on 9 April 2020. 3. 
Capture occurring on 24 April 2020. Odontaspis ferox: 
black triangle, around Zembra Island from Capapé 
(1975). 2. Together with Hexanchus griseus. Black star, 
capture site of Carcharodon carcharias. GT = Gulf of 
Tunis. GH = Gulf of Hammamet. GG = Gulf of Gabès.
Sl. 1: Zemljevid obravnavanega območja z označeno 
lokaliteto, kjer so bili ujeti primerki treh velikih mor-
skih psov. Hexanchus griseus: 1. Podatki po Ben Amor 
s sod. (2019). 2. Ulov z dne 9 aprila 2020. 3. Ulov z 
dne 24 aprila 2020. Odontaspis ferox: črni trikotnik, 
okoli otoka Zembra, po Capapé (1975). 2. Skupaj s 
primerkom vrste Hexanchus griseus. Črna zvezdica 
označuje lokaliteto ulova vrste Carcharodon carcha-
rias. GT = Tuniški zaliv. GH = Zaliv Hammamet. GG = 
Zaliv Gabès.
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Mohamed Mourad BEN AMOR et al.: CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA), 15–24
One year later, on 9 April 2020, a second shoal of 
12 H. griseus was collected between the east coast of 
Linosa, an islet close to Lampedusa, and the western 
coast of Malta Island, at 35° 43’ 20’’ N and 13° 9’ 43’’ 
E (see Fig. 1). The specimens were caught by bottom 
longlines, at depths between 400 and 1100 m on rocky 
bottom, together with a specimen of smalltooth sand ti-
ger shark Odontaspis ferox (Risso, 1810), a specimen of 
unidentified torpedo, several Squalus blainvillei (Risso, 
1826), and a specimen of dusky grouper Epinephelus 
marginatus (Lowe, 1834). All captured specimens were 
landed at the fishing port of Kélibia, where they were 
examined and photographed (Fig. 2). The total weight 
of this shoal reached 55 tons.
Additionally, on 24 April 2020, other 2 specimens 
of H. griseus were captured off Pantelleria Island, at 
36° 28’ 20’’ N and 12° 31’ 30’’ E. They were caught by 
bottom longlines, at a depth of 1100 m, on rocky bot-
tom. They were not measured for length, but weighed 
350 kg and 400 kg, respectively.
Finally, on 2 May 2020, 13 specimens of H. griseus, 
2 females and 11 males, were captured 13 miles south-
-east of Pantelleria Island, at 36° 28’ 20’’ N and 12° 
29’ 51’ E. They were caught by bottom longlines, at a 
depth of 800 m on rocky bottom, together with speci-
mens of E. marginatus. Their total body weight reached 
54 tons. Both females measured 3.5 m in total length 
and weighed 700 kg in total body weight, and carried 
120 and 118 fully yolked eggs ready to be ovulated, 
respectively. Each egg was covered by a fine diaphano-
us membrane, and together they were enveloped in a 
single membranous capsule (Fig. 3).
All specimens of H. griseus were identified in situ or 
from photographs provided by Kélibia port authority, 
through the combination of some characteristics, such 
as: body stout, head broad, snout short and blunt, six 
gill slits, a single dorsal fin above fin base, upper jaw 
with 4 rows of front teeth, lower jaw with 6 rows of 
lower blade-like, comb-shaped teeth on each side, 
dorsal surface dark brown, belly beige. This descrip-
tion is in total accordance with Boeseman (1984), 
Compagno (1984), Quéro et al. (2003), and Ebert & 
Stehmann (2013).
Fig. 2: Shoal of Hexanchus griseus landed at the fishing 
site of Kélibia.
Sl. 2: Jata morskih psov šesteroškrgarjev, ujetih na loka-
liteti Kélibia.
Fig. 3: Fully yolked eggs removed from a female of 
Hexanchus griseus: 1. Each egg was covered by a fine 
diaphanous membrane. 2. All eggs were enveloped 
together in one single membranous capsule.
Sl. 3: Z rumenjakom bogato jajce, odstranjeno iz 
samice vrste Hexanchus griseus: 1. Vsako jajce je pre-
krito s fino presevno membrano. 2. Vsa jajca so skupaj 
združena v eni membranski kapsuli.
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Smalltooth sand tiger shark Odontaspis ferox 
(Risso, 1810)
This shark was captured together with a shoal of 
H. griseus in similar conditions (see above). It was 
identified by the combination of the following main 
morphological characters: a bulky body with a long 
conical snout; eye small more than 4 times in snout, 
without nictitating eyelids; mouth long, extending 
behind the eyes (Fig. 4); teeth moderately large, each 
with a prominent narrow cusp and two or more pairs 
of lateral cusplets (Fig. 5); second dorsal fin origin 
above or slightly behind anal fin origin; anal fin and 
second dorsal fin smaller than first dorsal fin, caudal 
fin asymmetrical, with a strong lower lobe. Colour 
medium grey, sometimes with darker reddish spots 
scattered along the body, pelvic and anal fins very 
dark grey, lower edge of caudal fin almost black. These 
morphological characters are in total agreement with 
Compagno (1984). and Ebert & Stehmann (2013). 
The smalltooth sand tiger shark, Odontaspis ferox, 
has a cosmopolitan distribution in warm temperate 
and tropical waters, and although essentially demer-
sal, it has also been captured pelagically in mid-ocean 
(Compagno, 1984). The species often occurs inshore 
at steeply shelving coastal and insular locations, and 
in the Southern Hemisphere, most O. ferox were cau-
ght by trawl on the continental slope down to 880 m 
(Fergusson et al., 2008).
Due to its wide distribution, the species has been 
reported in oceans and seas throughout the world 
(Ebert & Stehmann, 2013). It is sporadically captured 
off the western Atlantic coast from North America 
(Sheehan, 1998) to Brazil (Menni et al., 1995). Off the 
eastern Atlantic coast, O. ferox is recorded off France 
(Quéro et al., 2003), Portugal (Carneiro et al., 2014) 
and southward of the Strait of Gibraltar from Morocco, 
without information about the capture site according 
to Collignon & Aloncle (1972), to Mauritania (Ebert & 
Stehmann, 2013). It is recorded in waters surrounding 
islands such as the Azores (Barcelos et al., 2018), Ma-
deira (Maul, 1955), the Canaries (Barría et al., 2018) 
and Cape Verde (Wirtz et al., 2013).
The first Mediterranean record of O. ferox 
occurred off Nice, southern France (Risso, 1810), 
subsequently, it was also reported by Bougis (1959) 
and Granier (1964); conversely, Capapé (1977) and 
Capapé et al. (2000) did not find the species in the 
area. Barrull & Mate (2002) considered the species 
as present off the Spanish coast. Tortonese (1956) 
Fig. 4: A. Hexanchus griseus. B. Odontaspis ferox. landed at the fishing site of Kélibia.
Sl. 4: A. Hexanchus griseus. B. Odontaspis ferox, iz ribiške lokalitete Kélibia.
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Mohamed Mourad BEN AMOR et al.: CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA), 15–24
noted the capture of a large female, 370 cm TL, from 
the Gulf of Genoa, southwards, two other specimens 
were reported by Vanni (1992), while Vacchi & Sere-
na (1997) and Sperone et al. (2012) reported captures 
from the Sicily Strait. Soldo & Jardas (2000) added 
that O. ferox is sporadically caught in the Adriatic 
Sea. Eastwards, O. ferox is present in the Aegean 
Sea (Ondrias, 1971), furtherly Kabasakal & Bayri 
(2019) summarized several captures of specimens 
that occurred in Turkish waters, while Akbora et al. 
(2019) reported other captures from Cyprus Island. 
Additionally, O. ferox is also recorded in the Levant 
Basin (Mouneimne, 1977; Golani, 2005; Bariche & 
Fricke, 2020).
Off the Maghreb shore, Dieuzeide et al. (1953) 
noted the occurrence of the species in the Algerian 
coast, which was furtherly confirmed (Hemida, pers. 
com., in Barrull & Mate, 2002). Capapé (1975) repor-
ted the capture of a female measuring 247 cm TL and 
weighing 70 kg, off Zembra Island, located in the Gulf 
of Tunis, northeastern Tunisia. The present capture 
(Fig. 5) constitutes a new record for the central Me-
diterranean Sea, where Schembri et al. (2003) noted 
that specimens may be collected annually in small 
numbers around Malta Islands; among them adult 
females of up to ca 360 cm TL were captured during 
years 1998 and 1999 in the area.
Great white shark Carcharodon carcharias 
(Linnaeus, 1758)
This large shark has worldwide distribution, 
especially in temperate waters, and its occurrence is 
well documented throughout the Mediterranean Sea, 
especially in the Strait of Sicily, where several juveni-
le and adult specimens were recorded (Quéro, 1984; 
Fergusson 1996, 2002; Saïdi et al., 2005; Maliet et al., 
2013). C. carcharias is known off the Tunisian coast, 
like other large and dangerous elasmobranch species 
(Capapé et al., 1975), and it appears that more than 
60 reliable captures of the species should be taken 
into consideration from Tunisian waters (Zaouali et 
al., 2020).
On 28 April 2020, a specimen was caught by drift 
longline baited with clupeid and scombrid species. 
The capture occurred off Sousse, a city located in 
eastern Tunisia, at 35° 01’ 01 N and 12° 11’ 11’’ E, at a 
depth of 30-50 m, on soft bottom. The specimen was 
a female measuring 232 cm TL and weighing 90 kg. 
It was identified based on the combination of main 
morphological characters, such as: body very large, 
fusiform, snout rather blunt, flattened above, origin of 
first dorsal fin slightly posterior to inner corner of pec-
toral fin, a single keel on caudal base, teeth triangular, 
coarsely serrate, blade-like; colour greyish-brown or 
blue above, belly white (Fig. 6).
DISCUSSION
The captures of large sharks reported in the present 
paper indicate that these species are not totally extinct 
in the Mediterranean and some viable populations 
continue to develop and reproduce in this sea. Such is 
for instance the case for H. griseus and C. carcharias.
Conversely, such pattern is not evident for O. ferox, 
but it cannot be totally ruled out seeing that some 
specimens were recently captured in the eastern Me-
diterranean (Kabasakal & Bayri, 2019; Akbora et al., 
2019; Bariche & Fricke, 2020).
About the specimens caught from Malta Islands, 
Schembri et al. (2003) noted that such discoveries 
suggest that adult females reach annually selected 
sites, possibly for reproduction, which makes O. ferox 
especially vulnerable during aggregations when they 
fall prey to overfishing and spearfishing more easily. 
The recent captures of O. ferox from both central and 
eastern Mediterranean suggest that the species is not 
totally extinct in this sea. However, the presence of a 
viable population in some areas is yet uncertain, parti-
ally due to the lack of information about all traits of its 
reproductive biology.
These two new captures of shoals of Hexanchus 
griseus from the Tunisian coast confirm once again 
that the species is not facing a drastic decline despite 
its K-selective traits (Ebert, 1986; Capapé et al., 
Fig. 5: Head and mouth of Odontaspis ferox showing 
the teeth (black arrow).
Sl. 5: Glava in usta vrste Odontaspis ferox z zobmi 
(črna puščica).
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Mohamed Mourad BEN AMOR et al.: CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA), 15–24
2004). Also, there have been several records reported 
throughout the Mediterranean during several deca-
des, suggesting that a viable population of H. griseus 
is at present probably established in this sea. The 
captures of the two shoals probably occurred during 
the shark’s reproductive period as indicated by the 
presence of large males and females, probably adults. 
Such hypothesis is confirmed by the fact that three 
females carried fully yolked oocytes. The record of 
first such specimen was provided by Ounifi-Ben Amor 
et al. (2017), and two are described in the present 
paper. The numbers of these oocytes, 85, 118 and 
120 revealed higher levels of ovarian fecundity than 
is generally recorded in viviparous sharks. H. griseus 
could be classified as a relatively prolific elasmo-
branch species and such phenomenon explains its 
abundance in some areas, Tunisian and Turkish waters 
as best instances. Each oocyte is protected by a fine 
membrane, and all oocytes are enveloped together 
in an external membrane. This prevented them from 
being scattered while handled by fishermen, which 
unfortunately, can happen, too (see Ounifi-Ben Amor 
et al., 2017). 
Such abundance could be correlated to food sources 
and feeding aggregation, or, like in the present cases, 
to sharks apparently schooling to revisit sites with good 
environmental conditions and availability of food and 
in search of favourable nursery grounds where females 
could lay and protect their brood. Such patterns cannot 
be totally ruled out.
The flesh of H. griseus is not appreciated for 
consumption by Tunisian people, it is probably 
ichthyosarcotoxic due to its richness in oil and rather 
dangerous to health (see Capapé et al., 1975). The 
shark was not targeted by fishermen, but due to the 
fact that the country is facing economic difficulties, 
these captures are considered an opportunity by fis-
hermen and consequently the sharks are not released 
Fig. 6: Carcharodon carcharias landed at the fishing site of Sousse. A. Head. B. Snout showing teeth.
Sl. 6: Carcharodon carcharias iz ribiške lokalitete Sousse. A. Glava. B. Gobec z zobmi.
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Mohamed Mourad BEN AMOR et al.: CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA), 15–24
back into the sea, which may contribute to a possible 
depletion of stocks. A strong monitoring of the species 
should therefore be conducted to avoid a drastic de-
cline of the species in the area.
Boldrocchi et al. (2017) noted that between 476 
and 2015, 628 reliable records of C. carcharias were 
reported in the Mediterranean Sea, and informed us 
(Boldrocchi, 2020, in letteris) that no other record 
occurred in this sea since 2015. The two records of C. 
carcharias reported by Rafrafi et al. (2015, 2019) from 
the northeastern Tunisian coast and the present record 
are probably the last ones known to date locally and 
in this sea. Such records mean that Tunisian waters 
are a hotspot for conservation and reproduction of 
this species. The specimen caught off Sousse is pro-
bably a juvenile specimen (see Compagno, 1984) and 
confirms the hypothesis that Tunisian waters could be 
a nursery ground for the species. Despite a number 
of white sharks caught in Tunisian waters and their 
relative abundance, their role in the conservation 
and reproduction of the species were completely 
dismissed by Moro et al. (2019), which is a rather 
incomprehensible and inadmissible opinion.
Following Ferretti et al. (2008) it appears, accor-
ding to the evidence available, that large predatory 
sharks in the Mediterranean Sea are generally decli-
ning in abundance, diversity and range. Conversely, 
the data herein presented indicate that may not be the 
case in certain Mediterranean areas.
ACKNOWLEDGEMENTS
The authors are grateful to Mr Raouf Galii, captain 
and owner of the long-liner boat “Nassim-Hasan”, for 
providing information about the captures of Hexan-
chus griseus and Odontaspis ferox. They wish also 
thank two anonymous referees for their helpful and 
useful comments allowing to improve the scientifc 
quality of the present article.
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Mohamed Mourad BEN AMOR et al.: CAPTURES OF LARGE SHARK SPECIES FROM THE NORTHEASTERN TUNISIAN COAST (CENTRAL MEDITERRANEAN SEA), 15–24
ULOVI VELIKIH MORSKIH PSOV OB SEVEROVZHODNI TUNIZIJSKI OBALI 
(OSREDNJE SREDOZEMSKO MORJE)
Mohamed Mourad BEN AMOR, Marouène BDIOUI & Khadija OUNIFI-BEN AMOR 
Institut National des Sciences et Technologies de la Mer, port de pêche, 2025 La Goulette, Tunisia
Christian CAPAPÉ
Laboratoire d’Ichtyologie, Université de Montpellier, case 104, 34095 Montpellier cedex 5, France
e-mail: capape@univ-montp2.fr
POVZETEK
Avtorji poročajo o treh vrstah velikih morskih psov, ujetih ob severovzhodni tunizijski obali. Gre za primerke 
morskega psa šesteroškrgarja Hexanchus griseus (Bonnaterre, 1788), morskega bika Odontaspis ferox (Risso, 
1810), in belega morskega volka Carcharodon carcharias (Linnaeus, 1758). Razpravljajo o razširjenosti vrst in 
njihovi ekološki vlogi. 
Ključne besede: Chondrichthyes, Hexanchus griseus, Odontaspis ferox, Carcharodon carcharias, razširjenost, 
tunizijska obala
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