ANNALES · Ser. hist. nat. · 29 · 2019 · 1
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received: 2019-03-08                  DOI 10.19233/ASHN.2019.02
FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, 
SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH NOTES 
ON A RECENT SIGHTING AND INCIDENTAL CAPTURES
Hakan KABASAKAL
Ichthyological Research Society, Tantavi mahallesi, Menteşoğlu caddesi, İdil apt., No: 30, D: 4, Ümraniye, 34764 İstanbul, Turkey
e-mail: kabasakal.hakan@gmail.com
ABSTRACT
Nine angel sharks, Squatina squatina, were recorded in different regions of Turkish waters between 21 March 
2009 and February 2019. Eight specimens were captured by different means of bottom fishing gear (e.g., gill or tram-
mel net, or longline) or harpoon, and 1 specimen was sighted alive. Considering the fishery statistics of angel sharks 
in the last 10 years and biomass and numerical abundance data on the species in Turkish waters as baseline data, an 
alarming reduction in the populations of S. squatina in the mentioned regions has been observed. An extensive study 
providing baseline information on the current spatial distribution patterns, habitat use, abundance and population 
structure of S. squatina in Turkish waters is an urgent necessity. 
Key words: Angelshark, Squatina, protection, baseline shift, rarity, eastern Mediterranean
FINALMENTE PROTETTO! STATO DELLO SQUALO ANGELO, SQUATINA SQUATINA 
(LINNAEUS, 1758) IN MARE TURCO, CON NOTE SU AVVISTAMENTI RECENTI E 
CATTURE ACCIDENTALI
SINTESI
Nove squali angelo, Squatina squatina, sono stati registrati in diverse regioni delle acque turche tra il 21 marzo 
2009 e il febbraio 2019. Otto esemplari sono stati catturati con diversi attrezzi da pesca a fondo o con l’arpione, 
mentre un esemplare è stato avvistato vivo. Considerando le statistiche sulla pesca degli squali angelo negli ultimi 10 
anni e i dati di biomassa e abbondanza numerica delle specie presenti nelle acque turche come dati di riferimento, 
è stata osservata una riduzione allarmante delle popolazioni di S. squatina nelle regioni considerate. Uno studio 
approfondito che fornisca informazioni di base sugli attuali modelli di distribuzione spaziale, sull’uso dell’habitat, e 
sull’abbondanza e la struttura demografica di S. squatina nelle acque turche è una necessità urgente.
Parole chiave: squalo angelo, Squatina, protezione, deviazione dal riferimento, rarità, Mediterraneo orientale
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Hakan KABASAKAL: FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH ..., 17–24
INTRODUCTION
In natural systems, rarity of a species is a common 
phenomenon and most often defined by two attributes: 
the species’ distribution and its abundance (Flather & 
Sieg, 2007). According to Flather and Sieg (2007), a 
species is considered rare if its area of occupancy or its 
number is small when compared to the other species 
that are taxonomically or ecologically comparable. The 
angel shark, Squatina squatina (Linnaeus, 1758), used to 
be reported as abundant in the entire Mediterranean Sea 
(Serena, 2005). Nowadays, it is supposed to be absent or 
even extirpated from several areas in the region (Bradai 
et al., 2012).
Due to the low overall number of records or a 
complete absence of the species in scientific trawl 
surveys conducted since 1958 (Holcer & Lazar, 2017), 
S. squatina is today considered a rare shark species 
throughout the Mediterranean Sea and its rarity has 
been emphasized by several researchers from different 
parts of the region (e.g., Golani, 2006; Serena & Relini, 
2006; Kabasakal & Kabasakal, 2014; Akyol et al., 2015; 
Cavallaro et al., 2015).
Although the occurrence of S. squatina in Turkish 
waters has been noted in historical inventory studies 
of Turkish marine fishes (e.g., Ninni, 1923; Deveci-
yan, 1926; Akşıray, 1987), all of which attributed an 
economic value to the species and mentioned mass 
captures, current statistics indicate a drastic decline in 
annual landings of incidentally captured angel sharks 
in Turkish waters (TUIK, 2017). As in many parts of the 
Mediterranean, populations of S. squatina have been 
severely depleted in Turkish waters as well, which calls 
for the protection of the species.
The present article provides a summary of the cap-
tures and of the single sighting of S. squatina in Turkish 
waters in the last 10 years, and discusses the causes of 
the species’ depletion. These events have eventually 
resulted in the implementation of effective protective 
measures for the species in the mentioned region.
MATERIAL AND METHODS
Data on captures and a single sighting of S. squatina in 
Turkish waters were obtained from the following sources: 
(1) scientific literature; (2) preserved samples in public or 
private collections; (3) newspaper accounts or internet 
news reporting on the capture of angel sharks in the stud-
ied area and (4) visits to fishing ports. As far as popular 
sources are concerned, the validity of the records was 
confirmed through direct contact with the fishermen cited 
by the sources, or else, based on a reliable photograph of 
the specimen accompanying such news. The following 
data were recorded for the examined specimens of angel 
shark: date, depth and locality of capture, total length 
(TL), weight (W), sex and fishing gear.
Commercial landings data on angel sharks from 
2008 to 2017 were used to indicate the trend of annual 
Fig. 1: Maps showing the approximate locations of recent captures and the single sighting of S. squatina, in Turkish 
waters. The numbers correspond to those in Table 1.
Sl. 1: Zemljevidi o recentnih ulovih in opazovanjih navadnega sklata v turških vodah. Številke ustrezajo tistim v 
Tabeli 1.
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Hakan KABASAKAL: FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH ..., 17–24
catches of Squatina species in Turkish waters (TUIK, 
2017). To date, three species of angel shark, S. aculeata 
Dumeril, in Cuvier, 1817, S. oculata Bonaparte, 1840 
and S. squatina have been recorded in Turkish waters 
(Bilecenoğlu et al., 2014). Since no species-specific 
discrimination of angel shark catches is reported in 
TUIK statistics, the annual landings data represent the 
combined data on the landings of three congeneric 
angel sharks in Turkish waters. Raw data and individual 
records of angel sharks are kept in the archive of the 
Ichthyological Research Society.
RESULTS AND DISCUSSION
Nine angel sharks, S. squatina, were recorded in sev-
eral regions of Turkish waters between 21 March 2009 
and February 2019 (Tab. 1; Fig. 1). Eight specimens were 
captured by different means of bottom fishing gear (e.g., 
gill or trammel net, or longline) or harpoon, and 1 speci-
men was sighted alive (see Tab. 1 for specimen-specific 
details). Six specimens were females, 2 were males and 
the sex of 1 angel shark was unknown. The total length 
of the specimens varied from 90 to 174 cm. Five angel 
sharks were captured in Aegean waters, three in the Sea 
of Marmara and 1 specimen was sighted in the south-
eastern part of the region near the northern entrance to 
the Dardanelles Strait (Fig. 1). The angel sharks were 
recorded in very shallow (specimen no. 2 at a depth of 
1.5 m) to quite deep waters (specimen no. 3 at a depth 
of ca. 50 m) of the coastal zone (Tab. 1).
Based on the information obtained from available 
literature, the occurrence of the angel shark, S. squatina, 
in Turkish waters dates back to the early 20th century, 
when Deveciyan (1926) reported on angel sharks landed 
Tab. 1: Old and recent captures or sightings of the angel shark, Squatina squatina, in Turkish waters since 2009. The 
numbers in the “No.” column are same as those in Fig. 1 indicating the approximate localities of capture.
Tab. 1: Stari in novejši zapisi o opazovanju navadnih sklatov, Squatina squatina, v turških vodah od leta 2009. 
Številke v stolpcih so enake kot v Sliki 1 in ponazarjajo približno lokaliteto ulova. 
No Date Locality
Depth 
(m)
Sex TL (cm) W (kg) Remarks
1 21 March 2009 Bodrum - ♀ 160 ca. 60 Entangled in coastal stationary-nets
2 October 2013 Çanakkale 1.5 - ca. 100 - Harpooned by a freediver in shallow 
water
3 4 January 2014 Mudanya ca. 50 ♀ 174 35
Entangled in a trammel-net, after 
the auction it was sold (Kabasakal & 
Kabasakal, 2014)
4 5 February 2015 Gökova Bay 20 ♀ 156 32.6
Entangled in a trammel-net; it was a 
pregnant female carrying at least four 
developping embryos, which were 
expelled during the handling (Akyol 
et al., 2015)
5 5 March 2015 Gökova Bay - ♀ 160 - Found dead while entangled in a 
stationary-net (unpublished data)
6 December 2015
Dardanelles 
Strait
ca. 30 ♀ ca. 150 -
Sighted by Mr. Mert Gökalp during 
a SCUBA diving expedition in the 
vicinity of the northern entrance of 
the Dardanelles Strait (unpublished 
data) (Fig. 2)
7 21 March 2018 Şarköy ca. 40 ♂ 114 -
Captured by long-lining set for 
sea breams (Sparidae), displayed 
in Beşiktaş fishmarket in İstanbul 
(unpuplished data) (Fig. 3).
8 3 April 2018
Dardanelles 
Strait
ca. 30 ♂ 90 -
Captured by stationary-netting, 
displayed in Beşiktaş fishmarket in 
İstanbul (unpublished data) (Fig. 4).
9 February 2019
Bosphorus 
Strait
20 m ♀ ca. 140 30
Captured by gill-netting around the 
southern entrance of the Bosphorus 
Strait, transported to Edirne city and 
sold (unpublished data)
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Hakan KABASAKAL: FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH ..., 17–24
at the İstanbul fish market. According to his report (De-
veciyan, 1926), angel shark was a regularly captured, 
landed and consumed shark species; however, the 
author did not provide any specific information, such 
as annual catch data, etc., on the species. Kabasakal 
(2002) reported captures of 51 angel sharks in total off 
the coast of Turkey between 1995 and 1999. Kabasakal 
(2003) reported the capture of a female angel shark of 
87 cm TL in the southern Sea of Marmara, in November 
1995. According to Kabasakal and Kabasakal (2004), 5 
males in total were captured off the coast of Gökçeada 
in the northeastern Aegean Sea in October 1996 (3 
specimens) and July 1997 (2 specimens), all by means 
of bottom fishing gear. In the 1950s, S. squatina was a 
popular gamefish among spearfishermen in the Sea of 
Marmara (Kabasakal & Kabasakal, 2014). 
According to Deveciyan (1926), Akşıray (1987) and 
Kocataş et al. (1993), S. squatina is a valuable commer-
cial species of shark in Turkish marine fishery. Deveciyan 
(1926) and Akşıray (1987) in particular pointed out the 
significant landings and high economical value of angel 
sharks captured in Turkish waters in the 20th century. 
However, like many other shark species, populations of 
S. squatina in Turkish waters have drastically declined 
and the rarity or the absence of angel shark from the 
general ichthyological inventory or shark specific 
studies provides alarming evidence that the survival of 
the species is threatened. In two recent surveys on the 
composition of bycatch in pink-shrimp (Parapenaeus 
longirostris) beam trawl fishery in southern and northern 
Sea of Marmara, Bayhan et al. (2006) and Bök et al. 
(2011) reported the absence of S. squatina from both 
marine regions. Karakulak et al. (2000) also reported 
the absence of S. squatina in surveys on demersal fish 
populations of the northern Sea of Marmara. Keskin and 
Eryılmaz (2010) reviewed the demersal fish populations 
of the Sea of Marmara and found S. squatina to be rare in 
the mentioned region, which was further corroborated 
by Kabasakal (2016).
The rarity of S. squatina in Turkish waters was also 
emphasised by Kabasakal and Kabasakal (2014) and 
Akyol et al. (2015). According to Başusta et al. (2016), 
the paucity of publications on S. squatina (less than 5) is 
conspicuous among the studies on cartilaginous fishes 
occurring in Turkish Mediterranean waters. Recently, 
Filiz et al. (2018) estimated the biomass and numerical 
Fig. 2: A live specimen of S. squatina (specimen no. 6 in 
Table 1) sighted in the Dardanelles Strait at a depth of 
30 m (Photo: Mert Gökalp).
Sl. 2: Opaženi primerek navadnega sklata (primerek št. 
6 v Tabeli 1), opažen v prelivu Dardanel na globini 30 
m (Foto: Mert Gökalp).
Fig. 3: A S. squatina incidentally captured off Şarköy 
coast (southwestern Sea of Marmara; specimen no. 7 
in Tab. 1), on display at the fishmonger’s (Photo: Hakan 
Kabasakal).
Sl. 3: Naključno ujeti primerek navadnega sklata ob 
obali Şarköy (jugozahodno Marmarsko morje; primerek 
št. 7 v Tabeli 1), razkazan na ribji tržnici (Foto: Hakan 
Kabasakal).
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Hakan KABASAKAL: FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH ..., 17–24
abundance of S. squatina at 13 kg/km2 and 5 individu-
als/km2 based on the bycatch composition of bottom 
trawl fishery in southern Aegean Sea. Since no species-
specific discrimination of angel shark catches is reported 
in TUIK statistics, the annual landings data represent 
the combined data on the landings of three congeneric 
angel sharks, S. aculeata, S. oculata and S. squatina, 
captured in Turkish waters. A dramatic decline in the 
annual landings of Squatina species is clear from the 
fishery statistics for the 2008–2017 period (TUIK, 2017; 
Fig. 5). While in 2008 the annual landings of Squatina 
species by commercial fishermen amounted to 34 tons, 
in 2017 they came nearly 1 ton (Fig. 5). Although the 
landings rose to 17 tons in 2013, the declining trend in 
the annual landings during the 2008–2017 period did 
not change (Fig. 5).
The first records of S. squatina in Turkish waters date 
back to the early 20th century (Ninni, 1923; Deveciyan, 
1926). The occurrence of S. squatina in Turkish waters 
was also reported in later studies (e.g., Roux, 1984; 
Akşıray, 1987). Both in the earliest and later studies, 
S. squatina was reported as an abundant shark popula-
tion sustaining a fishery based on it (Deveciyan, 1926; 
Akşıray, 1987). Furthermore, the maximum total lengths 
reported for the species ranged between 200 cm (Akşıray, 
1987) and 250 cm (Roux, 1984). Unfortunately, there are 
no studies reporting about the population dynamics and 
abundance of S. squatina in Turkish waters during the 
last century, however, recent data show drastic declines 
in terms of biomass, numerical abundance and annual 
landings (TUIK, 2017; Filiz et al., 2018; Fig. 5). Also, the 
maximum total length of angel sharks captured in recent 
years (174 cm TL in the present study; specimen no: 3, 
Tab. 1) is remarkably below the historically reported 
sizes (200 to 250 cm TL).
Based on the limited historical and contemporary 
data on biomass, numerical abundance, annual landings 
and maximum total length of the S. squatina observed in 
Turkish waters, the ‘shifting baseline syndrome’ outlined 
by Pauly (1995) is dramatically present among young 
generations of fishermen, a common phenomenon that 
was also highlighted by Holcer and Lazar (2017) for the 
angel sharks from the Adriatic Sea. Due to the absence 
of baseline data indicating the status of S. squatina in 
Turkish waters during the last century, we cannot carry 
Fig. 5: Annual landings of Squatina species by Turkish 
fishermen in the 2008–2017 period. The graph based 
on the annual landings data of Squatina species is ex-
tracted from fisheries statistics (TUIK, 2017).
Sl. 5: Letni ulov različnih vrst iz rodu Squatina, ki so jih 
ujeli turški ribiči v obdobju 2008-2017. Graf temelji na 
letnih ulovih sklatov, pridobljenih iz  statističnih ribiških 
podatkih (TUIK, 2017).  
Fig. 4: A S. squatina incidentally captured in Darda-
nelles Strait (specimen no. 8 in Tab. 1), on display at the 
fishmonger’s (Photo: Hakan Kabasakal).
Sl. 4: Navadni sklat naključno ujet v prelivu Dardanel 
(primerek št. 8 v Tabeli 1), razkazan na ribji tržnici 
(Foto: Hakan Kabasakal).
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Hakan KABASAKAL: FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH ..., 17–24
out a reasonable comparison between the historical and 
contemporary statuses of the species. However, based 
on available data it is possible to speculate that the 
abundance of S. squatina in demersal catch composi-
tion significantly declined over time, with the maximum 
total length of the species decreasing (Deveciyan, 1926; 
Roux, 1984; Akşıray, 1987; TUIK, 2017; Filiz et al., 
2018). According to Meyers et al. (2017), in regions 
where baseline information on the current spatial distri-
bution patterns, habitat use, abundance and population 
structure of the angel shark is lacking, urgent conserva-
tion policies should be implemented. Therefore, the 
inclusion of S. squatina, as well as the congeneric angel 
sharks, S. aculeata and S. oculata, in the list of marine 
protected species, which was declared in the Official 
Gazzette in 2018 (Resmi Gazete, 2018), was a correct 
decision with respect to the status of angel shark in Turk-
ish seas, where the baseline data of squatinid sharks is 
inadequate.
Except for the single live sighting, the angel sharks 
examined in the present study were incidentally cap-
tured specimens (Tab. 1); however, bycatch pressure on 
this rare shark by untargeted fisheries can also become 
a serious threat to its future survival. In an extensive 
study on the reproductive biology of S. squatina in the 
Mediterranean Sea, Capapé et al. (1990) reported that 
males and females mature between the total lengths of 
80–132 cm and 128–169 cm, respectively. According 
to Capapé et al. (1990), males and females of over 80 
cm and 128 cm in TL, respectively, can be considered 
as reproducing adult angel sharks. Recently, a pregnant 
female angel shark of 156 cm TL got entangled in a 
trammel net in Gökova Bay (southern Aegean Sea) and 
expelled its developing embryos during handling (Akyol 
et al., 2015). Based on the sizes of the reproducing 
adults of angel sharks reported by Capapé et al. (1990) 
and Akyol et al. (2015), it can be concluded that all of 
the specimens examined in the present study were ma-
ture males and females (Tab. 1). Incidental captures of 
reproducing adults create another threat to the survival 
of S. squatina. Fishing-gear induced injuries, keeping 
sharks on the deck in rough conditions and subjecting 
them to harsh handling before releasing them into the 
sea negatively impact the post-release survival rate of 
specimens (see Kabasakal, 2010 for relevant references), 
and cannot be ruled out as causes of mortality in the 
incidentally captured specimens of S. squatina in the 
present study.
CONCLUSIONS
Although the protective legislation concerning angel 
sharks applies to all 3 species, S. aculeata, S. oculata and 
S. squatina (Resmi Gazete, 2018), it is still very early 
to observe any positive results of the recent enactment 
of this legal protection on the angel shark populations 
occurring in Turkish waters. Since a “shifting baseline” 
effect is possibly present among younger fishermen, as 
reported by Holcer & Lazar (2017), the latter can lack 
incentive to act upon catching angel sharks, not recog-
nising that as important. Due to life history parameters 
of S. squatina, such as low fecundity, long reproductive 
cycle (2 year) and the large size at which females reach 
sexual maturity (128 cm, which equals 51.2% of the 
reported maximum total length, 250 cm) (Roux, 1984; 
Capapé et al., 1990), the species is considered as highly 
vulnerable to targeted and untargeted fisheries alike. 
Taking the fishery statistics of Squatina species during 
the last 10 years (TUIK, 2017) and the biomass and nu-
merical abundance data on the species in Turkish waters 
(Filiz et al., 2018) as baseline data, we should point out 
an alarming reduction in the populations of S. squatina 
in the mentioned region. An extensive study providing 
baseline information on the current spatial distribu-
tion patterns, habitat use, abundance and population 
structure of S. squatina in Turkish waters is therefore an 
urgent necessity. Given the Red List status of S. squatina 
(critically endangered; Cavanagh & Gibson, 2007) and 
its rarity (Golani, 2006; Serena & Relini, 2006; Kabasakal 
& Kabasakal, 2014; Akyol et al., 2015; Cavallaro et al., 
2015), the proposed study should employ non-invasive 
methods, such as capture and release of live specimens 
after satellite tagging and underwater imaging, perhaps 
with the contribution of citizen scientist divers. Last but 
not least, although S. aculeata, S. oculata and S. squatina 
are currently under full protection in Turkish seas (Resmi 
Gazete, 2018), it would be reasonable to expect cryptic 
mortality of the released angel sharks due to fishing-gear 
induced injuries and harsh handling before release. To 
prevent this and insure the safety of both fishermen and 
captured sharks, training of fishermen on which species 
of sharks are endangered and under protection and how 
these species should be kept alive on deck and handled 
before release, is critical. 
ACKNOWLEDGMENTS
The author wishes to thank his colleagues who share 
their information on angel sharks, and two anonymous 
referees for their valuable comments improving the con-
tent of the article. Very special thank goes to Mr. Mert 
Gökalp, an Istanbul based underwater videographer and 
photographer, for sharing the photo of the angelshark 
(specimens No. 6 in Tab. 1). Author is grateful to his 
wife, Özgür, and to his son, Derin, for their endless love 
and patience.
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Hakan KABASAKAL: FINALLY UNDER PROTECTION! STATUS OF THE ANGEL SHARK, SQUATINA SQUATINA (LINNAEUS, 1758) IN TURKISH SEAS, WITH ..., 17–24
KONČNO ZAVAROVAN! STATUS NAVADNEGA SKLATA, SQUATINA SQUATINA 
(LINNAEUS, 1758) V TURŠKIH MORJIH S POSEBNIM POUDARKOM NA NOVEJŠA 
OPAZOVANJA IN NAKLJUČNE ULOVE
Hakan KABASAKAL
Ichthyological Research Society, Tantavi mahallesi, Menteşoğlu caddesi, İdil apt., No: 30, D: 4, Ümraniye, 34764 İstanbul, Turkey
e-mail: kabasakal.hakan@gmail.com
POVZETEK
Devet navadnih sklatov, Squatina squatina, je bilo popisanih v različnih predelih turških voda med 21. marcem 
2009 in februarjem 2019. Osem primerkov je bilo ujetih z različnimi ribiškimi orodji (npr. zabodne in trislojne mreže, 
ribolov s parangali) ali harpuno, en primerek pa je bil opažen. Upoštevaje ribiško statistiko v zvezi s sklati v zadnjih 
desetih letih in podatke o abundanci in biomasi o vrsti v turških morjih kot temeljne podatke, je razviden veliki upad 
populacij sklatov v omenjenih okoljih. Zato je potrebno čimprej pripraviti obsežno raziskavo o vzorcih prostorske 
razširjenosti, izbiri habitata, abundanci in populacijski strukturi navadnega sklata v turških morjih. 
Ključne besede: sklat, Squatina, varovanje, bazični odmik, redkost, vzhodno Sredozemlje
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