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received: 2019-04-15                 DOI 10.19233/ASHN.2019.03
ADDITIONAL RECORDS OF RARE SHARKS FROM NORTHERN TUNISIA 
(CENTRAL MEDITERRANEAN SEA)
Sihem RAFRAFI-NOUIRA
Unité de Recherches Exploitation des Milieux aquatiques, Institut Supérieur de Pêche et d’Aquaculture de Bizerte, Université de 
Carthage, BP 15, 7080 Menzel Jemil, Tunisia
Youssouph DIATTA & Almamy DIABY
Laboratoire de Biologie marine, Institut fondamental d’Afrique noire, (IFAN Ch. A. Diop), Université Cheikh Anta Diop de Dakar, BP 
206, Dakar, Senegal.
Christian CAPAPÉ
Laboratoire d’Ichtyologie, Université de Montpellier, case 104, 34095 Montpellier cedex 5, France
e-mail: capape@univ-montp2.fr
ABSTRACT
Routine investigative monitoring conducted along the northern Tunisian coast allowed us to collect some speci-
mens of large predatory sharks, such as the great white shark, Carcharodon carcharias (Linnaeus, 1758), the shortfin 
mako Isurus oxyrinchus Rafinesque, 1810, and the common thresher shark Alopias vulpinus (Bonnaterre, 1788). 
These captures indicate that these sharks occur in the Mediterranean Sea, however, further records are needed 
before declaring that viable populations may still be established in the region. On the other hand, the captured 
specimen of smoothback angelshark Squatina oculata Bonaparte, 1840 could probably be considered as one of the 
last ones recorded to date in the Mediterranean Sea. The status of the species remains undetermined between extinct 
and critically endangered.
Key words: Large predatory sharks, distribution, status, central Mediterranean, endangered species
NUOVI RITROVAMENTI DI SQUALI RARI NELLE ACQUE DELLA TUNISIA 
SETTENTRIONALE (MEDITERRANEO CENTRALE)
SINTESI
Il monitoraggio investigativo di routine condotto lungo la costa tunisina settentrionale ci ha permesso di racco-
gliere alcuni esemplari di grandi squali predatori, come il grande squalo bianco, Carcharodon carcharias (Linnaeus, 
1758), il mako Isurus oxyrinchus Rafinesque, 1810, e lo squalo volpe Alopias vulpinus (Bonnaterre, 1788). Queste 
catture indicano che tali squali sono presenti nel mare Mediterraneo; sono tuttavia necessari ulteriori ritrovamenti 
al fine di asserire che nella regione possano essersi stabilite popolazioni vitali. D’altra parte, l’esemplare catturato di 
squalo angelo Squatina oculata Bonaparte, 1840, potrebbe venir considerato come uno degli ultimi ritrovamenti nel 
mare Mediterraneo. Lo stato della specie rimane indeterminato, tra estinto e in pericolo critico.
Parole chiave: grandi squali predatori, distribuzione, stato, Mediterraneo centrale, specie in via di estinzione
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INTRODUCTION
Studies carried out during the last two decades 
showed migrations of elasmobranch species towards 
northern areas, including brackish lagoons (Méjri et 
al., 2004; El Kamel et al., 2009, 2010; Rafrafi-Nouira 
et al., 2015). The species identified among these were 
the blackchin guitarfish Glaucostegus cemiculus (E. 
Geoffroy Saint Hilaire, 1817), the spiny butterfly ray 
Gymnura altavela (Linnaeus, 1758), and the sandbar 
shark Carcharhinus plumbeus (Nardo, 1827). Several 
large specimens of the latter were recorded (Rafrafi-
Nouira et al., 2015; Soufi-Kechaou et al., 2018; Capapé 
et al., 2018a). Since the capture included male and 
female adults, it suggested that a viable population was 
established in an area where the species was formerly 
unknown (Capapé, 1989).
Additionally, these studies enabled the finding 
of species considered to be highly threatened, such 
as the sharpnose sevengill shark Heptranchias perlo 
(Bonnaterre, 1788), a deep-sea species recorded in 
the Eskerkis Bank (El Kamel-Moutalibi et al., 2014), 
which migrated southward to lower than usual depths 
(Capapé et al., 2018b), and a critically endangered 
species, the Maltese skate Leucoraja melitensis (Clark, 
1926), which is the latest specimen recorded to date 
(Ben Amor et al., 2018).
In this paper, we report additional and unusual 
records of rare species from northern Tunisian waters, 
among them the smoothback angelshark Squatina 
oculata Bonaparte, 1840, which is affected by drastic 
population decline.
MATERIAL AND METHODS
The four studied species reported in the present 
article were caught in marine waters of northern Tunisia 
and directly landed at the fishing sites of Bizerte and 
Ras Jebel (Fig. 1), where they were examined, measured 
for total length (TL) and weighed for total body weight 
(TBW). They were identified in situ using field guides, 
such as Whitehead et al. (1984), Louisy (2002) and 
Quéro et al. (2003), then delivered to the laboratory for 
confirmation and a thorough scientific analysis. Some 
specimens were measured to the nearest mm (includ-
ing the percentage of total length), and the total body 
mass was weighed to the nearest gram following Com-
pagno (1984a) for sharks, and Capapé & Roux (1980) 
for Squatina oculata. For each specimen, all available 
data related to their capture were given, including fish-
ing gear, depth, type of bottom and, wherever possible, 
associated ichthyofauna. After being carefully measured 
and analysed, each specimen was fixed in 10% buffered 
formaldehyde, preserved in ethanol and deposited in 
the Ichthyological Collection of the Institut Supérieur 
de Pêche et d’Aquaculture de Bizerte, where it was as-
signed a catalogue number.
RESULTS
Great white shark, Carcharodon carcharias (Linnaeus, 
1758)
Carcharodon carcharias is a large shark with 
worldwide distribution, inhabiting temperate waters 
especially, but known to penetrate tropical marine ar-
eas, too. Its occurrence is well documented in the entire 
Mediterranean Sea (De Maddalena & Heim, 2012). 
However, most of these records occurred in the central 
Mediterranean, especially in the Strait of Sicily, where 
several juvenile and adult specimens were recorded. 
Some authors consider this area a nursery ground for the 
species (Saïdi et al., 2005; Maliet et al., 2013; Bradaï & 
Saïdi, 2013). Historical and contemporary occurrences 
of C. carcharias in eastern Mediterranean are also well 
documented (Kabasakal, 2014, 2016). Based on the 
recent captures of newborn, young-of-the-year and 
juvenile specimens, a possible nursery ground for the 
great white shark in the Aegean Sea was also suggested 
(Kabasakal & Gedikoğlu, 2008; Kabasakal et al., 2018).
C. carcharias is known off the Tunisian coast, and 
Bradaï & Saïdi (2013) noted that 59 reliable captures 
had been reported in the region between 1953 and 
2012, with most captures (56%) occurring in the Gulf 
of Gabès, which was considered as a probable nursery 
area for the species. On 12 April 2015, a male measur-
ing 1.8 m approximately was caught by longline in the 
waters surrounding the Cani Rocks (37°25’35.90” N and 
10°08’43.28” E), at a depth of approximately 100 m, on 
rocky bottom. More recently, on 26 December 2018, 
a female was captured by a benthic trawl at the same 
Fig. 1: Map of the northern coast of Tunisia indicating 
(circle) the capture sites of the elasmobranch species 
presented in the paper.
Sl. 1: Zemljevid severno tunizijske obale z označenimi 
(krogec) lokalitetami ulova vrst, ki jih avtorji obrav-
navajo v prispevku.
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depth on sandy-muddy bottom (37° 25’ 50.08” N and 
10°02’ 50. 96 E). It reached 2.80 m in total length and 
weighed 400 kg. As soon as it was landed on deck, the 
specimen was cut into slices to be sold, but it was still 
identified as a great white shark by experienced fisher-
men. Only the head of the specimen was preserved, 
which was more conical, sharper and pointier than 
it usually appears in this species. It was also slightly 
curved, but since no injuries or scars were observed in 
this atypical formation (Fig. 2), this could be considered 
an abnormality that had not been recorded before in any 
shark species. The teeth of both upper and lower jaws 
were huge, triangular, bladelike and serrated, confirm-
ing the identification made by the fishermen. The head 
was deposited in the ichthyological collection of the 
Institut Supérieur de Pêche et d’Aquaculture de Bizerte 
(Tunisia) under catalogue number ISPB-Car-carch -01.
In addition, on 11 April 2019, a second female of C. 
carcharias was captured by a bottom trawler in the very 
same area, measuring 2 m in TL and weighing 250 kg 
according to the information given by fishermen. It was 
sold immediately, but the fishermen provided us with 
a photograph, based on which we could confirm the 
identification of the species (Fig. 3).
Shortfin mako Isurus oxyrinchus Rafinesque, 1810
Isurus oxyrinchus is a pelagic, coastal and oceanic 
shark that can be found in tropical and temperate wa-
ters worldwide ranging from the surface to depths of 
400 m. It is well known in the eastern Atlantic and 
throughout the Mediterranean Sea (Quéro, 1984). The 
species was first recorded in the northern Tunisian area 
Fig. 2: Snout of Carcharodon carcharias (Ref. ISPB-Car-
carch-01), scale bar = 200 mm.
Sl. 2: Gobec velikega belega morskega volka (Ref. ISPB-
Car-carch-01), merilo = 200 mm.
Fig. 3: The Carcharodon carcharias specimen captured 
on 11 April 2019, scale bar = 200 mm.
Sl. 3: Primerek velikega belega morskega volka, ujet 11. 
aprila 2019, merilo = 200 mm.
Fig. 4: Isurus oxyrinchus (Ref. FSB-Isu-oxy-01). A. Gen-
eral morphology, scale bar: 150 mm. B. Snout lateral 
view, scale bar = 40 mm. Snout ventral view, scale bar 
= 30 mm. 
Sl. 4: Isurus oxyrinchus (Ref. FSB-Isu-oxy-01). A. Mor-
fologija, merilo: 150 mm. B. stranski pogled na gobec, 
merilo = 40 mm. Pogled na gobec od spodaj, merilo = 
30 mm. 
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by Vinciguerra (1884). The shortfin shark is sporadi-
cally captured throughout the Tunisian coast. (Capapé, 
1989; Bradaï et al., 2002). Additionally, Capapé (1980) 
noted that I. oxyrinchus is caught off the northern Cape 
Bon Peninsula in spring and early summer, and follow-
ing Bradaï et al. (2002), it rarely occurs in southern 
regions.
Recently, investigations conducted in the same study 
area have allowed us to record the capture of several mako 
specimens. On 15 May 2015 and 7 July 2015, two males 
were caught by longline at a depth of approximately 150 
m. They measured 1300 mm and 800 mm and weighed 
24 kg and 4.2 kg, respectively. Several other specimens 
were collected during June 2018 by bottom trawlers ac-
cording to the information provided by fishermen. The 
smallest specimen was delivered to our laboratory for 
examination and measurements (Fig. 4, Tab. 1).
All specimens were identified as Isurus oxyrinchus 
based on the combination of the following characters: 
body fusiform, snout very pointed; first dorsal with apex 
acutely pointed, pectoral fins shorter than head fins and 
falcate, origin of anal fin below the middle of second 
dorsal fin base, teeth very slender, alike in both jaws. 
Such description is in total accordance with Cadenat & 
Blache (1981), Compagno (1984b) and Quéro (1984). 
Isurus oxyrinchus’ congeneric species, the longfin mako 
I. paucus Guitart Manday, 1966, occurs off the Algerian 
coast (Hemida & Capapé, 2008). It differs from I. oxyrin-
chus in pectoral fins being as long as the head, anal fin 
originating below the posterior end of the second dorsal 
fin base, and teeth widening towards the base.
The smallest specimen was deposited in the Ichthyo-
logical Collection of the Faculté des Sciences de Bizerte 
(Tunisia) under catalogue number FSB-Isu-oxy-01 (see 
Tab. 1). All these captures suggest that the species is 
not extinct in this area and that the presence of a viable 
population remains a hypothesis that cannot be totally 
ruled out.
Common thresher shark Alopias vulpinus (Bonnaterre, 
1788)
Alopias vulpinus is a large global coastal and pe-
lagic shark occurring in depths up to 650 m (Compagno, 
1984b). The species is present in the entire Mediterrane-
an, but commoner in the western basin (Capapé, 1989). 
The species seems to be rather abundant off the Algerian 
coast, where it is caught as target or bycatch in pelagic 
and coastal commercial and small-scale longline, purse 
seine and gillnet fisheries. It is locally used for human 
consumption, especially the meat and fins, despite the 
fact that international regulations prohibit retention. 
Observations were carried out from 1999 to 2002 in the 
great fish market of Algiers, where all captures occurring 
throughout the Algerian coast are landed. A total of 194 
specimens were examined. Conversely, recent observa-
tions carried out in the area and information provided 
by fishermen indicated a drastic decline of captures of 
the species (Hemida et al., in press).
Quignard & Capapé (1971) noted that A. vulpi-
nus occurs throughout the Tunisian coast, but more 
Tab. 1: Morphometric measurements (mm and %TL) of 
the specimen of Isurus oxyrinchus (Ref. FSB-Isu-oxy-01).
Tab. 1: Morfometrične meritve (mm in %TL) primerka 
vrste Isurus oxyrinchus (Ref. FSB-Isu-oxy-01).
Reference FSB-Isu-oxy-01 
Sex Female
Morphometric measurements mm % TL 
Total length (TL) 880 100
Head length 270 30.7
Prebranchial length 190 21.6
Preorale length 60 6.8
Prenasal length 45 5.1
Intergill length 80 9.1
Eye with 22 2.5
Eye height 18 2.0
Internasal length 4 0.5
Mouth width 85 9.7
Distance between snout 1st gill 210 23.9
Precaudal length 670 76.1
Pelvic fin length 45 5.1
Pelvic height 45 5.1
Pelvic inner margin 20 2.3
Pelvic anterior margin 30 3.4
Pelvic posterior margin 30 3.4
First dorsal height 55 6.3
First dorsal base 75 8.5
First dorsal inner margin 25 2.8
First dorsal posterior margin 43 4.9
First dorsal anterior margin 80 9.1
Second dorsal height 13 1.5
Second dorsal base 15 1.7
Second dorsal inner margin 10 1.1
Second dorsal posterior margin 10 1.1
Second dorsal anterior margin 23 2.6
Pectoral inner margin 35 4.0
Pectoral anterior margin 160 18.2
Pectoral posterior margin 90 10.2
Pectoral fin length 50 5.7
Total body mass (g) 4200
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Tab. 2: Morphometric measurements (mm and %TL) of 
the specimen of Alopias vulpinus (Ref. ISPB-Alo-vul-01).
Tab. 2: Morfometrične meritve (mm in %TL) primerka 
vrste Alopias vulpinus (Ref. ISPB-Alo-vul-01).
Reference ISPB- Alo-vul-01
Sex Male
Morphometric measurements mm % TL 
Total length (TL) 3775 100.00
Precaudal length 1840 48.74
Pre first dorsal length 830 21.99
Pre first dorsal length 160 4.24
Pre pectoral length 530 14.04
Head length 580 15.36
Prebranchial length 430 11.39
Preorale length 120 3.18
Pre-pelvic length 1350 35.76
Distance between 210 5.56
Eye with 35 0.93
Eye height 50 1.32
Mouth width 170 4.50
Pectoral fin length 335 8.87
Pectoral inner margin 480 12.72
Pectoral anterior margin 605 16.03
Pectoral posterior margin 200 5.30
Anal fin length 30 0.79
First dorsal posterior margin 290 7.68
First dorsal base 230 6.09
First dorsal inner margin 260 6.89
First dorsal anterior margin 345 9.14
Second dorsal height 30 0.79
Second dorsal base 35 0.93
Second dorsal inner margin 20 0.53
Second dorsal anterior margin 40 1.06
Second dorsal posterior margin 220 5.83
Tail anterior margin 1935 51.26
Total body mass (kg) 120
frequently in northern areas. However, Bradaï et al. 
(2002) reported the capture of 3 specimens in the Gulf 
of Gabès in the south. Routine monitoring of elasmo-
branch species in the area of Bizerte-Ras Jebel offered 
us the opportunity to collect 4 specimens: on 14 May 
2015, two specimens were captured measuring 4 m 
and 3.8 m, and weighing 130 kg and 120 kg, respec-
tively. Morphometric measurements were recorded for 
the smallest A. vulpinus and summarized in Table 2, 
its head was recovered and deposited in the Institut 
Supérieur de Pêche et d’Aquaculture de Bizerte (Tu-
nisia), under catalogue number ISPB-Alo-vul-01 (Fig. 
5). On 5 December 2018, a third specimen measuring 
3.0 m in TL and weighing 80 kg was captured. A fourth 
specimen was captured on 4 April 2019 measuring 4.0 
m and weighing 150 kg. 
The four specimens were captured by longlines 
at a depth of approximately 100 m, on muddy-sandy 
bottoms. They were identified as A. vulpinus following 
the main characters: short snout, eyes rather small and 
not reaching the dorsal surface of the head, first dorsal 
fin closer to pectoral than pelvic fins. A related thresher 
shark, A. superciliosus, differs from A. vulpinus in hav-
ing larger eyes, which reach the dorsal surface of the 
head, and the first dorsal closer to pelvic than pectoral 
fins (Quéro et al., 2003). It is the first time that so many 
specimens were captured in such a short time, though 
according to fishermen such captures are rather com-
mon in the area, which may indicate a viable population 
inhabiting the area.
Fig. 5: Alopias vulpinus (Ref. ISPB-Alo-vul-01). A. Gen-
eral morphology without tail, scale bar = 300 mm. B. 
Head, lateral view, scale bar = 100 mm. C. Jaws, lateral 
view, scale bar = 30 mm. D. Snout, lateral view, scale 
bar = 200 mm.
Sl. 5: Alopias vulpinus (Ref. ISPB-Alo-vul-01). A. Mor-
fologija morskega psa brez repa, merilo = 300 mm. B. 
Glava s strani, merilo = 100 mm. C. čeljusti s strani, 
merilo = 30 mm. D. Gobec s strani, merilo = 200 mm.
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Smoothback angelshark Squatina oculata Bonaparte, 
1840
Squatina oculata is known in the eastern Atlantic, 
south of the Strait of Gibraltar, from Morocco (Lloris 
& Rucabado, 1998) to Angola (Capapé et al., 2002). 
The species is rather abundant off the coast of Senegal, 
where it plays an important role in craft fishery. There 
the sharks are exploited for their fins, meat and the 
oil extracted from their livers for therapeutic purposes 
(Capapé et al., 2002).
Roux (1984) wrongly noted that S. oculata is only 
present in the western Mediterranean Basin, however, 
the species is indeed unknown off the Mediterranean 
coast of France (Capapé et al., 2000). Tortonese (1956) 
noted its occurrence in Italian waters, and Zava et al. 
(2016) collected 4 juvenile specimens from the Strait of 
Sicily. Ergüden et al. (2019) found a female S. oculata 
in Turkish waters and eastwards the species is also re-
corded in the Levant Basin (Golani, 2005).
S. oculata used to be caught quite abundantly off 
the Tunisian coast, especially in northern areas. The 
captured specimens were used to provide information 
about the species’ reproductive biology (Capapé et 
al., 1991). Additionally, Bradaï et al. (2002) noted the 
captures of some specimens southwards, in the Gulf of 
Gabès. Later, 4 adult specimens, 2 males and 2 females, 
were caught in the study area, their average total length 
was 1020 mm (Mnasri, 2008). On 26 December 2018, 
a female was captured in our study area, reaching 1300 
mm in TL and weighing 5 kg. It was recorded based on 
a photograph taken by a fisherman (Fig. 6), which was 
helpful in identifying the specimen as S. oculata owing 
to the lack of the median line of spines, the hind tips 
of pelvic fins not reaching the level of first dorsal fins, 
and the specimen displaying a greyish colour with white 
spots on dorsal surface and dark bars on tail. This was 
probably one of the latest Mediterranean specimens 
found to date and confirmed the scarcity of the species in 
this sea. Similarly, its closely related congeneric species 
S. squatina (Linnaeus, 1758) has also been subjected to 
fishing pressure and has, following Giusto & Ragonese 
(2014), almost disappeared not only from Sicilian wa-
ters, but other areas as well.
DISCUSSION
Previous and new discoveries of the mentioned 
elasmobranch species suggest that the northern Tunisian 
Fig. 6: Smoothback angelshark Squatina oculata, general morphology, scale bar = 200 mm.
Sl. 6: Pegasti sklat Squatina oculata, morfologija, merilo = 200 mm.
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coast could be a hotspot for large sharks. Such phenom-
enon could be explained by the geographical location 
of the Tunisian waters in the central Mediterranean Sea, 
an inevitable passageway between western and eastern 
basins at a halfway between Atlantic and sub-tropical 
marine influences. Additionally, the richness of marine 
biodiversity sustains the hypothesis of nursery areas, 
especially for viviparous species, which look for the 
best environmental conditions before laying their litters 
in the wild (Rafrafi-Nouira, 2016). Biological rich-
ness minimizes the pressure of inter- and intraspecific 
competition for food between sharks considered as top 
predators (Stergiou & Karpouzi, 2002). 
On the other hand, the warming of the Mediter-
ranean waters encourages the occurrence of species 
in areas where they were previously unknown or rare. 
The best instance of this phenomenon is Carcharhinus 
plumbeus, abundantly captured in the Gulf of Gabès 
and at present in northern areas as well (Capapé et al., 
2018a). Captures of C. plumbeus are equally numerous 
off the Algerian coast (Hemida et al., 2002), the sharks 
probably migrating along Maghreb shores from west to 
east. Similar patterns could also be taken into considera-
tion in relation to the recent abundance of Isurus oxy-
rinchus and Alopias vulpinus off the northern Tunisian 
coast. Information provided by experienced fishermen 
indicates that both species are regularly captured there. 
Since large sharks are locally targeted due to their 
high economical value, they are not discarded at sea after 
the capture. They are landed at fishing sites, where they 
are immediately sold, despite recommendations from 
Tunisian authorities. Thus, it appears that large sharks 
have not disappeared from Tunisian waters completely, 
therefore the opinion of Ferretti et al. (2008) about the 
loss of large sharks in the area is partially contrasted. 
However, due to their K-selected characteristics (Camhi 
et al., 1998), most elasmobranch species are considered 
to be threatened. Still, the actual relative abundance of 
large sharks captured from the northern Tunisian coast 
shows their populations have not experienced a drastic 
decline in this area.
The captures of few S. oculata along the northern Tu-
nisian coast remain questionable. Were they occasional 
or is a viable population of this species still present? For 
fishermen, captures of squatinid species are considered 
a rather amazing event. Recent findings of S. oculata 
(Mnasri 2008; Zava et al., 2016; Ergüden et al., 2019; 
this study) suggest that the species is critically endan-
gered, following Morey (2019), yet not extirpated. In 
any case, further records are needed before it is possible 
to confirm or refute the presence of a viable population 
of this species in the central Mediterranean.
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DODATNI ZAPISI O POJAVLJANJU REDKIH MORSKIH PSOV IZ SEVERNE TUNIZIJE 
(OSREDNJE SREDOZEMSKO MORJE)
Sihem RAFRAFI-NOUIRA
Unité de Recherches Exploitation des Milieux aquatiques, Institut Supérieur de Pêche et d’Aquaculture de Bizerte, Université de 
Carthage, BP 15, 7080 Menzel Jemil, Tunisia
Youssouph DIATTA & Almamy DIABY
Laboratoire de Biologie marine, Institut fondamental d’Afrique noire, (IFAN Ch. A. Diop), Université Cheikh Anta Diop de Dakar, BP 
206, Dakar, Senegal.
Christian CAPAPÉ
Laboratoire d’Ichtyologie, Université de Montpellier, case 104, 34095 Montpellier cedex 5, France
e-mail: capape@univ-montp2.fr
POVZETEK 
Na podlagi rutinskega monitoringa vzdolž severne tunizijske obale so bili pridobljeni podatki o primerkih neka-
terih večjih plenilskih vrst kot so beli morski volk, Carcharodon carcharias (Linnaeus, 1758), atlantski mako Isurus 
oxyrinchus Rafinesque, 1810, in navadna morska lisica Alopias vulpinus (Bonnaterre, 1788). Ti ulovi kažejo, da 
se ti morski psi še vedno pojavljajo v Sredozemskem morju, vendar so za oceno prisotnosti viabilne populacije 
na raziskanem predelu potrebni še dodatni podatki. Ujeti primerek pegastega sklata Squatina oculata Bonaparte, 
1840 pa je verjetno eden izmed zadnjih primerkov te vrste v Sredozemskem morju. Status te vrste je tako še vedno 
nejasen, saj ni jasno ali je vrsta kritično ogrožena ali morda izumrla.
Ključne besede: veliki plenilski morski psi, razširjenost, status, osrednje Sredozemsko morje, ogrožene vrste
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