ANNALES · Ser. hist. nat. · 29 · 2019 · 1
1
received: 2018-11-05             DOI 10.19233/ASHN.2019.01
A REVIEW OF SHARK RESEARCH IN TURKISH WATERS
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
Until the last quarter of the 20th century, most of our knowledge on the sharks of the Turkish waters was based 
on elasmobranch-specific chapters in general ichthyological inventory studies and on a limited number of anecdotal 
studies. Since the mid-1990s, however, there has been a remarkable rise in the number and quality of shark-specific 
studies and publications, resulting in a gradual filling of the gaps in our understanding of the sharks of the Turkish 
waters. Based on the publication dates, there were only 2 shark-specific articles published before 1990, 8 scientific 
articles about the sharks of the Turkish waters between 1990 and 2000, and as many as 88 papers dealing with 
several aspects of the sharks of Turkey since, for a total of 96 articles on the sharks of the Turkish waters published 
between 1968 and 2018. Hopefully, future studies by new generations of researchers will provide new data on 
several aspects of the sharks of the Turkish waters, which can deepen our understanding of them and complete their 
life stories.
Key words: shark research, publications, Turkish waters, eastern Mediterranean
REVISIONE DELLA RICERCA SUGLI SQUALI IN ACQUE DELLA TURCHIA
SINTESI
Fino all’ultimo quarto del XX secolo, la maggior parte delle nostre conoscenze sugli squali delle acque della 
Turchia si basava su capitoli specifici dedicati agli elasmobranchi pubblicati in studi generali di inventario ittologico 
e su un numero limitato di studi aneddotici. Dalla metà degli anni 90, tuttavia, c’è stato un notevole aumento del 
numero e della qualità degli studi e delle pubblicazioni specifici sugli squali, con un risultante riempimento graduale 
delle lacune nella comprensione degli squali delle acque turche. Sulla base delle date di pubblicazione, l’autore 
riscontra che solo due articoli specifici sugli squali sono stati pubblicati prima del 1990, 8 articoli scientifici sugli 
squali delle acque turche tra il 1990 e il 2000, e ben 88 articoli riguardanti diversi aspetti degli squali della Turchia 
da allora, per un totale di 96 articoli sugli squali delle acque turche pubblicati tra il 1968 e il 2018. L’autore spera 
che studi futuri di nuove generazioni di ricercatori possano fornire nuovi dati su diversi aspetti riguardanti gli squali 
delle acque turche, al fine di approfondirne la comprensione e completare le loro storie di vita.
Parole chiave: ricerca sugli squali, pubblicazioni, acque turche, Mediterraneo orientale
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
2
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
INTRODUCTION
Despite an anecdotal note on a great white shark, 
Carcharodon carcharias (Linnaeus, 1758), off the coast 
of İzmir (western Turkey) from the 16th century (Bellonii, 
1553), the story of shark research in the seas of Turkey 
started in the early 1920s. Regardless of their remark-
able ecological value, sharks have always been a short 
chapter in the general ichthyological inventory studies 
of Turkish marine waters (e.g., Ninni, 1923; Deveciyan, 
1926; Ayaşlı, 1937; Akşıray, 1987). The first accounts of 
sharks in the seas of Turkey were by Ninni (1923) and 
Deveciyan (1926). Besides being two of the most want-
ed items among collectors of antique books on Turkey’s 
ichthyofauna, these pioneering monumental studies 
also contain the first scientific inventories of the sharks 
of the Turkish waters. The chronology of shark research 
in the mentioned waters can be divided into two distinct 
eras, with the first reaching up to the last quarter of 20th 
century, and the second beginning in almost mid-1990s 
and still continuing. They each have peculiar character-
istics that outline the general trends of shark research in 
the seas of Turkey. While the first era was dominated by 
a clear disregard of sharks in the general ichthyological 
studies and a lack of shark-specific research, the second 
has witnessed a boom in the study of the sharks of the 
seas of Turkey. In the present article, the author reviews 
the contemporary status of shark research in Turkish 
waters in the light of available data.
MATERIAL AND METHODS
The data of the present review were obtained from 
published journals and electronic sources. To extract 
data from electronic sources, a structured Boolean 
search was performed on search engines such as 
Google Scholar, ScienceDirect, etc., with the following 
keywords: “sharks”, “elasmobranchii”, “Turkey”, “Le-
vantine”, “Black, Marmara, Aegean OR Mediterranean 
Seas”, “distribution”, “hexanchiformes”, “lamniformes”, 
“squaliformes”, “carcharhiniformes”. To extract data 
published in journals before 2000 and not accessible 
via internet, a manual search was performed. The col-
lected data were arbitrarily grouped under the following 
headings: ‘distribution, occurrence and first record; 
feeding biology; population dynamics, including studies 
on age, growth and length-weight relationship; repro-
duction, including studies on neonates, gravid females 
and nurseries; morphometry; underwater observations 
and behaviour; miscellaneous, including studies on 
conservation, fishery economy, anthropogenic injuries 
and newspaper portrayals; and general chondrichthyan 
studies including sharks’; and their contents were 
reviewed under the respective shark species. Since 
some of the articles dealt with more than one topic, 
each of these multi-topic articles was considered as a 
single article to prevent the overestimation of the total 
number of publications. The taxonomic nomenclature 
of the shark species follows Serena (2005). The distri-
bution of sharks in Turkish waters was based on Akşıray 
(1987), Bilecenoğlu et al. (2014), Kabasakal (2011b) 
or Kabasakal et al. (2017). The tabulated results of the 
data search are available upon request to the author, for 
further investigation.
RESULTS AND DISCUSSION
Overview of the studies
The data search for the present review revealed 
a remarkable difference between the numbers of 
shark-specific articles published in each of the two eras 
of research on the sharks of the Turkish waters (Fig. 1). 
In the first era, the historical data about sharks in Turkish 
waters were comprised in general ichthyological inven-
tories (Ninni, 1923; Deveciyan, 1926; Ayaşlı, 1937) and 
rare anecdotal notes (e.g., Erazi, 1942; Geldiay & Mater, 
1968), which, however, provided occurrence data of 
some rare (e.g., Oxynotus centrina, Echinorhinus bru-
cus) or questionable (e.g., Lamna nasus) or threatened 
sharks (e.g., Hexanchus griseus, Squatina squatina) in 
the mentioned marine region. These references also 
represent some of the earliest occurrence records of 
sharks in the Levantine Basin, and therefore, provide an 
understanding of the historical distribution of sharks in 
the eastern Mediterranean.  
Based on publication dates, there were only 2 
shark-specific articles published before 1990, 8 scientif-
Fig. 1: Geographical location of Turkish waters in the 
Mediterranean ecosystem. The shaded areas indicate 
the approximate locations of the three possible sharks 
breeding grounds: (1) in the northern Aegean Sea, from 
Gökçeada to Foça, (2) Boncuk Bay in the southern 
Aegean Sea, and (3) in the Bay of İskenderun in the 
eastern Levant.
Sl. 1.: Geografska lega turških voda v Sredozemskem 
morju. Osenčeni predeli označujejo približne lokalitete 
treh možnih območij razmnoževanja morskih psov: 
(1) severno Egejsko morje, v predelu Gökçeada - Foça, 
(2) zaliv Boncuk v južnem Egejskem morju, in (3) zaliv 
İskenderun v vzhodnem Levantu.
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
3
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
ic articles about the sharks of the Turkish waters between 
1990 and 2000, and as many as 88 papers dealing with 
several aspects of the sharks of Turkey since, for a total of 
96 articles on the sharks of the Turkish waters published 
between 1968 and 2018.
With regard to the topics of the publications, the 
largest number of articles (n = 60) dealt with the occur-
rence, distribution and first records of sharks in Turkish 
waters, followed by articles about stomach contents and 
feeding bioecology of sharks (n = 13), reproduction and 
nurseries (n = 13), morphometrics (n = 13), population 
dynamics (n = 11), underwater observations and be-
haviour studies (n = 5), and miscellaneous topics (n = 
5) (Fig. 2). The sharks of the Turkish waters were also 
discussed in general chondrichthyan studies, including 
those about the batoids of the mentioned marine region 
(n = 9) (Fig. 2).
Species accounts
HEXANCHIFORMES
HEXANCHIDAE
Heptranchias perlo (Bonnaterre, 1788)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Hexanchus griseus (Bonnaterre, 1788)
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on H. perlo and H. griseus in Turkish 
waters
The earliest records of the hexanchid sharks, H. 
griseus and Heptranchias perlo, in Turkish waters were 
reported in general ichthyological inventories by Ninni 
(1923), Deveciyan (1926) and Akşıraş (1987); their con-
temporary occurrences in the mentioned marine region 
were confirmed by Kabasakal (1998a, 2004a, 2005, 
2009b, 2013a), Kabasakal and İnce (2008), and Başusta 
(2015). Among these, studies by Kabasakal (2005) and 
Başusta (2015) are particularly worth mentioning. On 
19 November 2004, one male specimen of H. griseus 
of 300 cm TL and weighing 250 kg was captured by 
a commercial gill-netter nearly 3 miles off the coast of 
Amasra. This single capture extends the Mediterranean 
distribution of H. griseus to the Black Sea (Kabasakal, 
2005). Occurrence of neonates of H. perlo in İskenderun 
Bay (north-eastern Mediterranean Sea) was recorded for 
the first time by Başusta (2015). In a recent review of 
large sharks caught by commercial fisheries in Turkish 
waters, H. griseus was the predominant species acco-
unting for 169 specimens and 43.2% of total captures 
between 1990 and August 2015 (Kabasakal et al., 2017). 
Following the capture of newborns of sevengill 
sharks in İskenderun Bay, Başusta (2015) suggested the 
possibility of a breeding ground of H. perlo in the north-
-eastern Mediterranean Sea. In Turkish waters, bony 
fish constitute the main prey of H. griseus (Kabasakal, 
2004a). Available information suggests that H. griseus 
gives birth between October and late February in the 
northern Aegean and Marmara Seas (Kabasakal, 2004a). 
Morphometric measurements of the H. griseus caught in 
the Sea of Marmara were reported by Kabasakal (1998a). 
Based on the specimens captured by commercial fisher-
men in Turkish waters, the length-weight relationship for 
the bluntnose sixgill shark, H. griseus, was calculated as 
logTW=2.76xlogTL-4.6 (r=0.92; n=34; sexes combined; 
Kabasakal, 2006). Analysis of 81 articles related to H. 
griseus and published in major Turkish newspaper and 
internet media between 1974 and 2009 revealed that 
the tenor of the coverage was neutral in 49.3% of the 
articles, 45.6% were negative and only 4.9% were posi-
tive (Kabasakal, 2010c). The effect of anthropogenic and 
fishing-gear induced injuries on the survival and cryptic 
mortality of sharks were examined in two recent studies 
(Kabasakal 2010d, 2017a). Post-release behaviour of 
H. griseus in Saros Bay (north-eastern Aegean Sea) was 
also recorded via underwater videography (Kabasakal, 
2010d).
SQUALIFORMES
ECHINORHINIDAE
Echinorhinus brucus (Bonnaterre, 1788)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on E. brucus in Turkish waters
In October 2002, a bramble shark E. brucus was ima-
ged by means of a ROV camera at a depth of 1,214 m 
in the northern Sea of Marmara (Kabasakal et al., 2005). 
This single recording of the bramble shark shows that E. 
brucus, once thought extinct in Turkish seas, still occurs 
Fig. 2: Numerical distribution of the topics of articles 
on the sharks of the Turkish waters, published between 
1968 and 2018.
Sl. 2: Številčna porazdelitev tematik prispevkov o 
morskih psih turških voda, objavljenih med leti 1968 
in 2018.
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
4
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
in this area. Recent studies provide further records 
confirming the contemporary existence of E. brucus in 
Turkish waters (Kabasakal & Dalyan, 2011; Kabasakal & 
Bilecenoğlu, 2014; Kabasakal, 2017a).
SQUALIDAE
Squalus acanthias Linnaeus, 1758
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Squalus blainvillei (Risso, 1826)
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on S. acanthias and S. blainvillei in 
Turkish waters
Subsequently to the pioneering study of Geldiay 
and Mater (1968), Kutaygil and Bilecik (1977) reported 
on the distribution and abundance of the spurdog, S. 
acanthias, which ranked the second and/or third among 
the demersal fish along the western and central Black 
Sea coasts of Turkey. Contemporary occurrence of the 
congeneric S. blainvillei in Turkish waters was confirmed 
by Kabasakal and Kabasakal (2004), and Başusta et al. 
(2016).
The main prey of S. acanthias and S. blainvillei is 
composed of decapod crustaceans and bony fish (Ka-
basakal 2002d; Demirhan et al., 2007; Özütemiz et al., 
2009). Decapod crustaceans Liocarcinus sp. and the 
commercially important deep-water prawn, P. longiro-
stris, constituted the main prey items recorded in the 
stomach contents of S. blainvillei (Kabasakal, 2002d). 
According to Demirhan et al. (2007), the availability 
of prey is the main criterion for the feeding strategy of 
S. acanthias; thus in winter, the main prey item of the 
spiny dogfish is anchovy (Engraulis encrasicolus). In the 
eastern Mediterranean population of S. acanthias, the 
length at 50% maturity was 87.57 cm for males and 
102.97 cm for females (Demirhan & Seyhan, 2007). On 
the other hand, Yığın and İşmen (2013) estimated the 
mean length at 50% maturity for females and males of 
S. acanthias at 56.4 cm and 52.8 cm, respectively. The 
structural problems of using spines for determining the 
age of the spiny dogfish (S. acanthias) were examined by 
Demirhan et al. (2006), and the authors concluded that 
the percentage of age reading failure was 70% and 37% 
on the first and second spines, respectively. One of the 
previous studies investigated the population structure of 
S. acanthias along the south-eastern Black Sea coast of 
Turkey (Düzgüneş et al., 2006). In this study, a total of 
267 specimens (85 male and 182 female) were collected 
and the mean (±se) length and weight were 88.25±2.157 
cm and 3319±204 g for males, and 92.55±1.73 cm and 
4387±217.6 g for females. The length-weight relation-
ship for the stock was derived as W=0.009*L3.3423 
(r2=0.9607) (Düzgüneş et al., 2006). In a previous study 
carried out in the south-eastern region of the Black Sea, 
Avşar (1996) reported that male specimens of S. acanthi-
as dominated the population, the age of this species in 
the region ranged between 1 and 14 years, and the mean 
annual growth rate was 7.2 cm. The growth parameters 
of S. blainvillei were examined based on the specimens 
captured in Sığacık Bay (Aegean Sea), and the mean 
size for the examined specimens of S. blainvillei was 
21.46+2.21 cm, respectively (Özütemiz et al., 2009).
CENTROPHORIDAE
Centrophorus granulosus (Bloch & Schneider, 1801)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Centrophorus uyato (Rafinesque, 1810)
Distribution in Turkish waters: Marmara Sea (Bi-
lecenoğlu et al., 2014).
Review of studies on C. granulosus and C. uyato in 
Turkish waters
In the early 1990s, the first record of the gulper 
shark, Centrophorus granulosus, in the Sea of Marmara 
was reported by Benli et al. (1993), based on specimens 
caught at a depth of 400 m. Although the congeneric C. 
uyato was also recorded in the Sea of Marmara in the 
mid-1990s (Meriç, 1995), White et al. (2013) suggested 
that C. uyato was not a valid species and that further rese-
arch was necessary to clarify the taxonomic status of this 
species in Turkish waters (Kabasakal & Karhan, 2015).
ETMOPTERIDAE
Etmopterus spinax (Linnaeus, 1758)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Review of studies on E. spinax in Turkish waters
Kabasakal and Ünsal (1999) reported rare occur-
rences of E. spinax caught in deep-water bottom-trawl 
fishery in the northern Aegean Sea, together with the 
morphometric measurements of the examined spe-
cimens. In a previous study, 116 velvet belly lantern 
sharks, E. spinax, were caught at depths between 200 
and 600 m, in Sığacık Bay (Aegean Sea), and the po-
pulation was composed of more females than males, 
at 53.4% and 46.6%, respectively; the maximum total 
lengths for females and males were 20.5 cm and 18.9 
cm, respectively (Bilge et al., 2010). 
OXYNOTIDAE
Oxynotus centrina (Linnaeus, 1758)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on O. centrina in Turkish waters
Based on the chronological order, the first shark-
-specific article on the distribution and occurrence of 
sharks in Turkish waters was published in late 1960s. It 
provided anecdotal data on an angular rough shark (O. 
centrina) caught in the Bay of İzmir (Geldiay & Mater, 
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
5
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
1968). Studies on O. centrina, which is now considered 
a rare and threatened shark in the entire Mediterranean 
Sea, continued in the following decades and provided 
further understanding about the life story, occurrence 
and status of the angular rough shark in Turkish waters 
(Kabasakal, 2009a, 2010a, 2015a; Başusta et al., 2015; 
Yığın et al., 2016). A recent review on the occurrence 
and status of O. centrina in the eastern Mediterranean 
(Kabasakal, 2015a) revealed that between the late 1800s 
and 2012, the highest number of O. centrina specimens 
(72%) was recorded in the Aegean Sea, followed by the 
Sea of Marmara (21.5%). Contemporary occurrence of 
O. centrina in Turkish waters was also confirmed by two 
recent studies (Başusta et al., 2015; Yığın et al., 2016). 
Locomotory and feeding behaviour of the rare angular 
rough shark, O. centrina, were recorded by means of 
diver-operated camera in the northern Sea of Marmara 
(Kabasakal, 2009a). Measurements of O. centrina were 
reported by Yığın et al. (2016), based on a specimen 
caught in Saros Bay (north-eastern Aegean Sea). Based 
on the occurrence of a gravid female in İskenderun 
Bay, Başusta et al. (2015) suggested a possible breeding 
ground of O. centrina in the region.
DALATIIDAE
Dalatias licha (Bonnaterre, 1788)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on D. licha in Turkish waters
Kabasakal and Kabasakal (2002) reported rare occur-
rences of D. licha caught in deep-water bottom-trawl 
fishery in the northern Aegean Sea. Although the main 
prey of D. licha is comprised of decapod crustaceans (Lio-
carcinus sp. and P. longirostris), remains of the sympatric 
blackmouth dogfish (G. melastomus) were also found in 
the stomach contents of the kitefin shark (Kabasakal & 
Kabasakal, 2002). Based on the occurrence of newborn 
specimens of D. licha, Kabasakal and Kabasakal (2002) 
suggested a possible breeding area of the kitefin shark 
over the bathyal grounds of the north-eastern Aegean 
Sea. Morphometric measurements of 5 newborn D. 
licha from the north-eastern Aegean Sea were reported 
by Kabasakal and Kabasakal (2002). Recently, an adult 
female of D. licha, 118 cm TL, got entangled in a tram-
mel net set at a depth of 40 m in Iskenderun Bay (NE 
Mediterranean sea) (Ergüden et al., 2017)
SQUATINIFORMES
SQUATINIDAE
Squatina aculeata Dumeril, in Cuvier, 1817
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Squatina oculata Bonaparte, 1840
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Squatina squatina (Linnaeus, 1758)
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on S. aculeata, S. oculata and S. squa-
tina in Turkish waters
In Turkish waters, the angel shark family Squatinidae 
is represented by 3 species: Squatina aculeata, S. ocula-
ta and S. squatina, and their contemporary occurrences 
have been confirmed by a number of studies (Başusta, 
2002; Başusta et al., 1998; Kabasakal & Kabasakal, 
2004, 2014; Yağlıoğlu et al., 2015). S. squatina is con-
sidered one of the largest sharks in Turkish waters, and, 
historically, it was one of the commercially important 
shark species in Turkish demersal fishery. However, the 
remarkable decrease that has been recorded in angel 
shark populations is alarming, as the survival of the 
species may be threatened. As recent surveys show, S. 
squatina accounts for less than 2 percent of the total 
shark biomass incidentally caught by Turkish fishermen 
(Yağlığlu et al., 2015; Kabasakal et al., 2017 ).
LAMNIFORMES
ODONTASPIDIDAE
Carcharias taurus Rafinesque, 1810
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Odontaspis ferox (Risso, 1810)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Review of studies on C. taurus and O. ferox in Turkish 
waters
There is no specific study available on odontaspidid 
sharks in Turkish waters.
ALOPIIDAE
Alopias superciliosus (Lowe, 1839)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Alopias vulpinus (Bonnaterre, 1788)
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on A. superciliosus and A. vulpinus in 
Turkish waters
Kabasakal (1998b) reported on the incidental captu-
re of a female thresher shark, A. vulpinus, 453 cm TL, by 
a commercial purse-seiner off Şile coast (south-western 
Black Sea), on 8 November 1996, and this anecdotal 
note was followed by further records. Contrary to the 
species’ open water habits, Kabasakal (2007) reported 
on the coastal occurrences of 19 common thresher 
sharks (A. vulpinus) that were incidentally captured by 
coastal stationary netters. Recently, Ergüden et al. (2015) 
reported on a single male thresher shark, 392 cm TL and 
weighing ca. 180 kg, captured incidentally in purse-
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
6
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
-seine fishery in İskenderun Bay; this was the first record 
of A. vulpinus from the north-eastern Mediterranean 
coast of Turkey. The first record of the congeneric bigeye 
thresher shark, A. superciliosus, in Turkish waters dates 
back to the early 2000s (Mater, 2005; Bay of Gökova, 
south-eastern Aegean Sea); a few years later it was 
recorded in the Sea of Marmara (Kabasakal & Karhan, 
2008). The female bigeye thresher shark, 450 cm TL, 
caught on 28 February 2011 off Fethiye coast, is one of 
the largest specimens of A. superciliosus ever recorded 
in the Mediterranean Sea and worldwide (Kabasakal et 
al., 2011). Based on the total number (n = 392) of large 
sharks caught by commercial fishermen between 1990 
and August 2015 in Turkish waters, A. superciliosus and 
A. vulpinus accounted for 2.5% and 9.9% of the total 
catch, respectively (Kabasakal et al., 2017).
CETORHINIDAE
Cetorhinus maximus (Gunnerus, 1765)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Review of studies on C. maximus in Turkish waters
Early writings on the occurrence of basking sharks in 
Turkish waters date back to the 1990s, when an anec-
dotal record of basking shark, Cetorhinus maximus, was 
reported from north-eastern Levantine waters (Kıdeyş, 
1997). It was followed by further records off the Turkish 
coasts, particularly in the Bay of Antalya (Kabasakal, 
2002a, 2004b, 2013b). In January of 2009, an adult male 
basking shark was accidentally caught in a stationary 
net in the waters some 2 nautical miles off Küçükkuyu 
(north Aegean Sea); its total length measured ca. 1,000 
cm (Kabasakal, 2009c). According to Kabasakal (2009c), 
this specimen is the largest well-documented basking 
shark recorded in the Mediterranean to date. In a recent 
review of the status of basking sharks in the eastern 
Mediterranean, Kabasakal (2013c), based on the extre-
mely low number of records off Turkish coast since the 
1950s, emphasized the rarity of C. maximus in Turkish 
waters and the need for a specific monitoring program 
accompanied by zooplankton surveys that would de-
termine the seasonal movements of basking sharks in 
the mentioned region and answer the questions whether 
the occurrence of this species in Turkish waters exhibits 
seasonality and site fidelity or not.
LAMNIDAE
Carcharodon carcharias (Linnaeus, 1758)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on C. carcharias in Turkish waters
In a 16th century record, Bellonii (1553) reported on 
the occurrence of C. carcharias off İzmir coast. Follow-
ing this historical note, this top predator was listed in 
the general ichthyological inventories of Turkish waters 
(Ninni,1923; Deveciyan, 1926; Ayaşlı, 1937; Akşıray, 
1987) When talking about the sharks of the Mediterra-
nean Sea, the great white shark, C. carcharias, has always 
been the “top star” of research efforts and incidental 
captures of this predator have always found room in 
newspaper reports as well. Therefore, a detailed search 
in newspaper archives covering the period between the 
early 1900s and the late 1960s yielded several articles 
about specimens of C. carcharias incidentally captured 
by tuna hand-liners in the Bosphorus Strait during that 
time (Kabasakal, 2003a, b). Further research revealed 
the presence of C. carcharias in Marmara and Turkish 
Aegean waters between the 1960s and the 1990s (Kaba-
sakal 2008, 2011a; Kabasakal & Kabasakal, 2004). Fol-
lowing the capture of two newborns of the great white 
in Edremit Bay (northern Aegean Sea) in the summer 
of 2008, contemporary occurrence of C. carcharias in 
Turkish waters was confirmed (Kabasakal & Gedikoğlu, 
2008) and this incidence was followed by the captures 
of additional specimens (4 newborns and 3 juveniles) 
in the north-eastern Aegean Sea, in the coastal zone 
extending from Foça (central Aegean Sea) to Gökçeada 
(north-eastern Aegean Sea) (Kabasakal, 2014; Kabasakal 
& Kabasakal, 2015; Kabasakal et al., 2009). Among the 
captured newborns, a north Aegean Sea specimen (85 
cm TL) caught by a coastal trammel netter in Edremit 
Bay on 6 July 2011, is possibly the smallest neonate 
great white shark reported from Mediterranean waters 
to date (Kabasakal, 2014). The capture of neonate, 
young-of-the-year and juvenile white sharks suggested 
the presence of a breeding ground of C. carcharias in 
the central and northern coasts of the Turkish Aegean 
Sea (Kabasakal, 2014; Kabasakal & Kabasakal, 2015; 
Kabasakal et al., 2009). In light of the available data, 
C. carcharias is an extant lamnoid shark in the Aegean 
waters off the Turkish coast, but presently not occurring 
in the Sea of Marmara (Kabasakal, 2016a, Kabasakal et 
al., 2018), contrary to the assertions in Bilecenoğlu et al. 
(2014), which are based on historical occurrence data 
of the species.
Isurus oxyrinchus Rafinesque, 1810
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Review of studies on I. oxyrinchus in Turkish waters
A huge female shortfin mako shark, Isurus oxyrinc-
hus, was caught in the late 1950s off Marmaris (south-
-eastern Aegean Sea; Kabasakal & De Maddalena, 
2011). Photographic documentation was used by the 
authors to estimate the total length of this specimen at 
585 cm, which greatly exceeded the previous maximum 
size recorded for the species (445 cm). Although recent 
studies have confirmed the contemporary occurrence 
of I. oxyrinchus in Turkish Aegean and Mediterranean 
waters (Ergüden et al., 2013; Kabasakal 2015b, 2017b; 
Kabasakal & Kabasakal, 2013; Tunçer & Kabasakal, 
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
7
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
2016), this is a rare lamnoid shark in Turkish waters. 
Therefore, the seasonal occurrence of both young and 
adult shortfin mako sharks off Turkey’s Aegean and Me-
diterranean coasts should be monitored to clarify whe-
ther the shortfin mako shark is a resident or a vagrant 
species along the aforementioned coastline. According 
to Kabasakal et al. (2017), I. oxyrinchus accounted for 
5.3% of the total number of large sharks captured by 
commercial fishermen in the 1990–2015 period in 
Turkish waters. Occurrences of I. oxyrinchus newborns 
and youngs-of-the-year off Foça and in İskenderun Bay 
suggest the possibility of a mating region in the northern 
Aegean and north-eastern Mediterranean Seas (Ergüden 
et al., 2013; Kabasakal, 2015b).
Lamna nasus (Bonnaterre, 1788)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on L. nasus in Turkish waters
Occurrence of the porbeagle shark, Lamna nasus, 
in Turkish waters was reported by Deveciyan (1926), 
Akşıray (1987) and Kabasakal (2002b). Kabasakal and 
Kabasakal (2004) reported on a porbeagle shark, 250 cm 
TL, caught off Bozcaada (northern Aegean Sea), on 11 
April 2004. L. nasus is a rare shark in Turkish waters and 
its questionable presence in Marmaric waters requires 
confirmation (Kabasakal & Karhan, 2015).
CARCHARHINIFORMES
SCYLIORHINIDAE
Galeus melastomus Rafinesque, 1810
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Scyliorhinus canicula (Linnaeus, 1758)
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Scyliorhinus stellaris (Linnaeus, 1758)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on G. melastomus, S. canicula and S. 
stellaris in Turkish waters
Contemporary occurrences of cat sharks (family 
Scyliorhinidae), Galeus melastomus, Scyliorhinus cani-
cula and S. stellaris, in Turkish waters were confirmed 
by Başusta et al. (2016), Kabasakal (2002b), Kabasakal 
and Kabasakal (2004), Kabasakal and Karhan (2015), 
Keskin and Karakulak (2006), İşmen et al. (2013), and 
Yağlıoğlu et al. (2015). S. canicula is the most abundant 
cartilaginous fish species caught in demersal fishery in 
Turkish waters (Cihangir et al., 1997; Kabasakal, 2002b; 
Keskin & Karakulak, 2006; İşmen et al., 2013; Yağlıoğlu 
et al., 2015).
Investigations carried out since 1997 have revealed 
that, in Turkish waters, the main prey items of S. canicula 
are decapod crustaceans (Parapenaeus longirostris, Lio-
carcinus sp., Goneplax rhomboides, Xantho sp. and Mu-
nida sp.) and teleostean fish, while secondary food items 
comprise polychaetes (e.g., Sipinculida) and cephalo-
pods (Cihangir et al., 1997; Kabasakal, 2001, 2002c; 
Filiz & Taşkavak, 2006; Türker Çakır 2006). A previous 
study on the stomach contents and feeding ecologies of 
the S. canicula and thornback ray, Raja clavata, caught in 
the northern Aegean Sea, revealed a clear niche overlap 
between S. canicula (all sizes) and R. clavata (>50 cm 
TL), and competition for similar prey items (Kabasakal, 
2001). In a broader perspective, in Turkish waters, G. 
melastomus preys mainly on crustaceans, teleosteans 
and cephalopods (Kabasakal, 2002c; Özütemiz et al., 
2009), while in the deep zones (>1000 m depth) of the 
Sea of Marmara, the blackmouth catshark feeds mainly 
on crustaceans, Calocaris macandreae and Sergestes 
robustus (Oral, 2010).
In a previous study on the distribution and deter-
minate biological aspects of the lesser spotted dogfish 
in the north-eastern Aegean Sea, Cihangir et al. (1997) 
reported that the reproduction of S. canicula occurs 
year-round, with a relatively low rate of oogenesis in 
winter and early spring. Demersal trawl surveys carried 
out in the northern Aegean Sea showed that seasonal 
biomass of S. canicula in shallow waters (<100 m depth) 
varied from 576 to 2,958 tonnes (Cihangir et al., 1997). 
Filiz and Mater (2002) examined the length-weight rela-
tionships of seven elasmobranch species sampled from 
the north Aegean Sea, and calculated the length-weight 
equations for S. canicula, M. mustelus and S. acanthias 
as follows: WT= 0.0016TL x 3.1804 (r2 = 0.9795), WT= 
0.0008TL x 3.3259 (r2 = 0.9745) and WT= 0.0031TL x 
3.1056 (r2 = 0.9814), respectively. According to Türker-
Çakır et al. (2006), the weight increased allometrically in 
the both sexes of the S. canicula sampled in the northern 
Aegean Sea (b = 2.93). Sexual dimorphism, dentition 
and morphometry of the S. canicula caught in Turkish 
waters were studied by Erdoğan et al. (2004) and Filiz 
and Taşkavak (2006).
Growth parameters of G. melastomus were examined 
based on specimens captured in Sığacık Bay (Aegean 
Sea), and the mean size for the examined specimens was 
14.01+1.96 cm (Özütemiz et al., 2009). The length-wei-
ght relationship parameters of 16 out of the 30 cartilagi-
nous fish species (11 sharks, 18 batoids and 1 chimaera) 
caught in the central Aegean Sea were examined; the 
values of the slope b in the length-weight relationship 
parameters ranged from 2.79 (Torpedo marmorata) to 
3.78 (Scyliorhinus stellaris), a values from 0.0002 (Scyli-
orhinus stellaris) to 0.9713 (Dasyatis pastinaca) (Eronat 
and Özaydın, 2014). 
TRIAKIDAE
Galeorhinus galeus (Linnaeus, 1758)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Mustelus asterias Cloquet, 1821
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
8
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
Distribution in Turkish waters: Black, Marmara, Ae-
gean and Mediterranean Seas (Bilecenoğlu et al., 2014).
Mustelus mustelus (Linnaeus, 1758)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Mustelus punctulatus Risso, 1826
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Review of studies on G. galeus, M. asterias, M. mustelus 
and M. punctulatus in Turkish waters
Contemporary occurrences of triakid sharks (family 
Triakidae), Galeorhinus galeus, Mustelus asterias, M. 
mustelus and M. punctulatus, in Turkish waters have 
been confirmed by a number of studies (Filiz and Ma-
ter, 2002; Kabasakal, 2002b; Kabasakal & Kabasakal, 
2004; Başusta et al., 1998, 2016; Yağlıoğlu et al., 
2015). Two specimens of the starry smoothhound, M. 
asterias, were captured three miles off the coast of Şile 
(south-western Black Sea) on 19 November 2000 at a 
depth of ca. 90 m, and this previous record extended 
the Mediterranean distribution of M. asterias to the 
Black Sea (Eryılmaz et al., 2011). The tope shark, G. 
galeus, is one of the largest shark species occurring in 
Turkish waters, and the recent remarkable decrease in 
its populations is alarming. Between 1990 and 2015, 
G. galeus accounted for less than 2 percent of the total 
shark biomass recorded in the Turkish commercial fish-
eries (Kabasakal et al., 2017). The diet of M. mustelus 
is heterogeneous and generalized, and the main prey 
item found in the stomach contents is decapod crus-
taceans, with teleosteans and cephalopods featuring 
as secondary important food items (Kabasakal, 2002c; 
Filiz, 2009; Özcan & Başusta, 2016). 
CARCHARHINIDAE
Carcharhinus altimus (Springer, 1950)
Distribution in Turkish waters: Mediterranean Sea 
(Bilecenoğlu et al., 2014).
Carcharhinus brevipinna (Müller & Henle, 1839)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Carcharhinus limbatus (Müller & Henle, 1839)
Distribution in Turkish waters: Mediterranean Sea 
(Bilecenoğlu et al., 2014).
Carcharhinus melanopterus (Quoy & Gaimard, 1824)
Distribution in Turkish waters: Mediterranean Sea 
(Akşıray, 1987; Kabasakal, 2011b).
Carcharhinus obscurus (Lesueur, 1818)
Distribution in Turkish waters: Mediterranean Sea 
(Akşıray, 1987; Kabasakal, 2011b).
Carcharhinus plumbeus (Nardo, 1827)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Prionace glauca (Linnaeus, 1758)
Distribution in Turkish waters: Marmara, Aegean and 
Mediterranean Seas (Bilecenoğlu et al., 2014).
Review of studies on Carcharhinus species and P. glauca 
in Turkish waters
Despite historical and contemporary occurrences of 
carcharhinid sharks (family Carcharhinidae) in Turkish 
waters have been noted in general ichthyological or 
chondrichthyan inventories of the mentioned region 
(e.g., Ninni, 1923; Akşıray, 1987; Başusta et al., 1998; 
Kabasakal, 2002b, 2003b; Kabasakal & Kabasakal, 2004; 
Kabasakal et al., 2017; Yağlıoğlu et al., 2015), the paucity 
of species specific studies is obvious. In a previous report 
on the status of the blue shark, Prionace glauca, in the 
northern Aegean Sea, Kabasakal (2010b) emphasized that 
coastal trammel- and gill-netting is a serious threat to the 
survival of the young populations of the species in Edre-
mit Bay. In an extensive survey on the chondrichthyan 
fish of İskenderun Bay (north-eastern Mediterranean 
Sea), Başusta et al. (1998) recorded the bignose shark, 
Carcharhinus altimus, for the first time in Turkish waters 
and confirmed the contemporary presence of the sandbar 
shark, C. plumbeus, as well. Underwater observations 
carried out in Boncuk Bay (south-eastern Aegean Sea) 
and the knowledge gathered from these surveys played a 
critical role in declaring C. plumbeus a protected species 
in Turkish seas (Akça, 2010; Öztürk, 2006). Recent sur-
veys have confirmed the occurrence of the dusky shark, 
C. obscurus, in the Bay of İskenderun (Kabasakal et al., 
2017). The spinner shark, C. brevipinna, is considered a 
rare shark and has only been listed in the general ichthyo-
logical and chondrichthyan inventories of Turkish waters 
(Akşıray, 1987; Kabasakal, 2002b); recently, Filiz and 
Kabasakal (2015) reported on a specimen of this species 
photographed in the Bay of Gökova. The recently disco-
vered photographic evidence suggests that Carcharhinus 
spp. were present in the Sea of Marmara in the 1950s 
(Kabasakal, 2015c), and this evidence also extends the 
known historical distributional range of Carcharhinus 
spp. into Marmaric waters, a northern extension of the 
Mediterranean Basin.
In a pioneering study in which the length-weight 
relationship (LWR) of the sandbar shark, C. plumbeus, 
from the northeastern Mediterranean Sea population 
was examined for the first time, the LWR estimated for 
combined sexes, females and males, were W= 0.01*TL 
(r=0.915, SE =0.174), W= 0.0034*TL (r =0.912, SE 
=0.278) and W= 0.0039*TL (r =0.915, SE =0.231), 
respectively (Başusta, 2016). According to Başusta 
(2016), the type of allometric growth in the examined 
population of C. plumbeus was negative (b < 3) for all 
sexes and for males, and positive (b > 3) for females. 
Seeing that Boncuk Bay (south-eastern Aegean Sea) is 
of critical importance for the reproduction of C. plum-
beus, Öztürk (2006) suggested that the bay should be 
declared a protected area and all fishery should be 
banned in its waters. Underwater observations to record 
the bioecological characteristics of the sandbar shark, 
C. plumbeus, were also carried out in Boncuk Bay 
(south-eastern Aegean Sea) during a 39-day expedition 
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
9
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
in 2009 (Akça, 2010). A study has also been carried out 
which confirmed the year-round aggregations of sandbar 
sharks in the area and provided new and solid evidence 
for banning all fishing activities in this marine protected 
area (Filiz, 2018).
SPHYRNIDAE
Sphyrna (Sphyrna) zygaena (Linnaeus, 1758)
Distribution in Turkish waters: Aegean and Mediter-
ranean Seas (Bilecenoğlu et al., 2014).
Review of studies on S. zygaena in Turkish waters
Our knowledge on hammerhead sharks (family 
Sphyrnidae) in Turkish waters consists of rudimentary 
data. Ulutürk (1987) and Kabasakal and Kabasakal 
(2004) reported rare occurrences of the smooth ham-
merhead shark, Sphyrna zygaena, off Gökçeada coasts 
(northern Aegean Sea), and the species was observed off 
the Kaş Peninsula (western Levantine Basin) in August 
2015 (Kabasakal et al., 2017). Although its occurrence 
was confirmed, S. zygaena is a rare shark in Turkish 
waters (Kabasakal et al., 2017).
Review of general studies on chondrichthyan fish in 
Turkish waters
In an extensive survey of the chondrichthyan fish of 
İskenderun Bay, the bignose shark, C. altimus, was recor-
ded for the first time in Turkish waters and the presence 
of C. plumbeus, O. centrina and S. oculata in the bay wa-
ters were confirmed as well (Başusta et al., 1998). Based 
on the results of field surveys and a review of available 
data, Kabasakal (2002b) reported on the presence of 39 
shark species in Turkish waters, defining the occurrence 
of 8 of them questionable and requiring confirmation. 
The results of an extensive trawl survey carried out in 
the northern Aegean Sea determined S. canicula as the 
most abundant cartilaginous fish species, with a total 
mass estimated at 2,850.6 kg/nm2 (Keskin & Karakulak, 
2006). Between March 2005 and June 2008, in Saros 
Bay, Turkey, ten shark species were sampled from depths 
ranging from 5 to 500 m for length-weight relationship 
using a commercial trawl vessel. The values of the expo-
nent b of the length-weight relationships ranged from 
2.6816 to 3.6060 (İşmen et al., 2009). In rare occasions, 
selected measurements of lamniform sharks (C. maximus, 
A. superciliosus, A. vulpinus, C. carcharias and I. oxyrin-
chus) were also reported by Kabasakal (2002a, 2017b), 
Kabasakal and Gedikoğlu (2008), Kabasakal and Karhan 
(2008), Kabasakal and Kabasakal (2013), Ergüden et al. 
(2013, 2015) and Tunçer and Kabasakal (2016). 
Between October 2011 and February 2013, the com-
position of by-catch in the beam trawls used in the pink 
shrimp (Parapenaeus longirostris) fishery was surveyed 
seasonally and S. canicula, S. stellaris, S. acanthias and 
O. centrina were recorded (İşmen et al., 2013). Between 
2009 and 2010, the total biomass, species composition, 
depth distribution, seasonal distribution and abundance 
of elasmobranchs caught by commercial bottom-trawlers 
in İskenderun Bay (north-eastern Levantine Sea) were 
also examined (Yağlıoğlu et al., 2015). In this study, M. 
mustelus, S. canicula, S. stellaris, G. melastomus and S. 
squatina represented between 0.45% and 1.7% of the 
total biomass of the examined cartilaginous fish. Single 
or sporadic captures were also recorded for I. oxyrinchus, 
C. altimus, C. plumbeus and O. centrina (Yağlıoğlu et 
al., 2015). In a recent review on cartilaginous fish and 
fisheries along the Mediterranean coast of Turkey, Başusta 
et al. (2016) recorded 32 species from 16 families, and 
concluded that the landings of elasmobranchs were 
reduced from 3,980 tonnes in 2000 to 246.2 tonnes in 
2015. The production and economic values of sharks and 
their relatives in Turkey have been analysed for the last 
34 years, and according to Doğan (2006), the maximum 
catch level was recorded in 1979 (11,125 t), followed by 
a significant decrease after 1989 and reaching the mini-
mum level of 400 t in 2003. The current status of sharks 
and shark fisheries in the waters of the Sea of Marmara 
has been of late extensively reviewed (Kabasakal, 2016b; 
Yığın et al., 2016). Last but not least, in a recent survey of 
shark attacks against humans and boats, 13 shark attacks 
were recorded in Turkey’s waters between 1931 and 
1983, 2 of which fatal (Kabasakal & Gedikoğlu, 2015).  
CONCLUSIONS
Until the last quarter of the 20th century, most of 
our knowledge on the sharks of the Turkish waters was 
based on elasmobranch-specific chapters in general ich-
thyological inventory studies and on a limited number of 
anecdotal studies. Since the mid-1990s, however, there 
has been a remarkable rise in the number and quality 
of shark-specific studies and publications, resulting in 
a gradual filling of the gaps in our understanding of the 
sharks of the Turkish waters. The reference list of the 
present article also serves as a bibliographical archive of 
the shark- and elasmobranch-specific studies related to 
Turkish waters that were published between 1968 and 
2018. The next step in shark- and elasmobranch-spe-
cific studies should involve designing a master plan for 
the management of sharks and their relatives in Turkish 
waters, and implementing the conservatory acts that 
would be based on the available data.
Currently, 36 species of sharks with confirmed 
occurrence are present in Turkish waters (Akşıray, 1987; 
Kabasakal, 2011b; Bilecenoğlu et al., 2014; Kabasakal 
et al., 2017): Hexanchus griseus, Heptranchias perlo, 
Echinorhinus brucus, Squalus acanthias, S. blainvillei, 
Centrophorus granulosus, C. uyato, Etmopterus spinax, 
Oxynotus centrina, Dalatias licha, Squatina aculeata, 
S. oculata, S. squatina, Carcharias taurus, Odontaspis 
ferox, Alopias superciliosus, A. vulpinus, Cetorhinus 
maximus, Carcharodon carcharias, Isurus oxyrinchus, 
Lamna nasus, Galeus melastomus, Scyliorhinus cani-
cula, S. stellaris, Galeorhinus galeus, Mustelus asterias, 
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
10
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
M. mustelus, M. punctulatus, Carcharhinus altimus, C. 
brevipinna, C. limbatus, C. melanopterus, C. obscurus, 
C. plumbeus, Prionace glauca, and Sphyrna zygaena. 
There are also species of questionable occurrence, whi-
ch are included in one of the noteworthy ichthyological 
inventories of Turkish waters (Akşıray, 1987), but require 
confirmation, namely: C. longimanus, S. lewini, S. tudes 
and Somniosus rostratus.
The continuous rise in the number of articles and 
chapters on the sharks of the Turkish waters published 
annually is promising. The seasonal occurrence of neo-
nate and young-of-the-year specimens of several shark 
species in Turkish waters, as well as the presence of 
documented breeding and nursery grounds in the region 
form a solid basis for the implementation of seasonal 
restrictions of commercial fisheries in these grounds. 
Based on the by-catch records of neonates and juvenile 
specimens of some rare sharks, such as D. licha and O. 
centrina, or some large sharks, for example, H. grise-
us, C. carcharias, I. oxyrinchus and C. plumbeus, it is 
possible to speculate that sharks, at least the mentioned 
species, could reproduce in Turkish waters throughout 
the vast coastal zone spreading from the north Aegean 
Sea to the eastern Levant. In light of available and con-
firmed data, there are three possible breeding grounds; 
(1) in the northern Aegean Sea, between Gökçeada and 
Foça, (2) in Boncuk Bay in the southern Aegean Sea, and 
(3) in the Bay of İskenderun, eastern Levant (Fig. 1). To 
allow the survival of shark species, these areas should 
be declared seasonally protected or at least restricted 
zones for commercial fisheries. Currently, Boncuk Bay 
is the only protected area. 
Different geographical populations of the same shark 
species can exhibit different growth parameters, which 
should be taken into account in their management. For 
instance, available data on the parameters of the Aegean 
Sea and Black Sea populations of S. acanthias present a 
clear difference in length at 50% maturity (Demirhan & 
Seyhan, 2007; Yığın & İşmen, 2013) – a critical detail 
that should not be overlooked in the regional manage-
ment of S. acanthias in Turkish waters.
Populations of many large shark species in Turkish 
waters have drastically declined; a recent review of the 
distribution of large sharks captured mostly by com-
mercial fishermen in the 1990–2015 period in Turkish 
waters showed that large sharks accounted for less than 
2 percent of the total captured biomass (Kabasakal et 
al., 2017). According to Kabasakal et al. (2017), large 
sharks in Turkish waters are threatened by multi-para-
meter fishing pressure of demersal and pelagic fisheries 
of artisanal to industrial scale that operate in coastal to 
open waters and in shallow to deep fishing grounds al-
most year-round. Last but not least, fishing-gear induced 
injuries and harsh handling are overlooked contributors 
to the cryptic mortality of sharks in Turkish waters. Ho-
pefully, future studies by new generations of researchers 
will provide new data on several aspects of the sharks of 
the Turkish waters, which can deepen our understanding 
of them and fill in the blanks in their life stories.
ACKNOWLEDGMENTS
The author wishes to thank his colleagues, who sha-
red their experiences and researches on sharks of Turkish 
waters, and two anonymous referees for their valuable 
comments, improving the content of the article. Author 
is grateful to his wife, Özgür, and to his son, Derin, for 
their endless love and patience.
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
11
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
PREGLED RAZISKAV O MORSKIH PSIH V TURŠKIH VODAH
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
Do zadnje četrtine 20. stoletja je bilo poznavanje o morskih psih v turških vodah omejeno na poglavja v specifič-
nih monografijah o morskih psih v splošnih ihtioloških pregledih in manjšem številu anekdotičnih raziskav. Od srede 
devetdesetih let pa je prišlo do občutnega povečanja v številu in kvaliteti specifičnih raziskav in publikacij o morskih 
psih, ki je počasi mašilo vrzel o poznavanju morskih psov v turških vodah. Pred letom 1990 sta bila objavljena le dva 
specifična prispevka o morskih psih, 8 prispevkov med leti 1990 in 2000 ter 88 prispevkov po letu 2000. Skupno 
je bilo objavljeno 96 prispevkov o morskih psih v turških vodah med leti 1968 in 2018. Smiselno je upati, da bodo 
nove generacije raziskovalcev odkrile nove podatke o morskih psih iz turških morij iz vseh vidikov, kar bo poglobilo 
in izpopolnilo znanje o njih in njihovem življenju.
Ključne besede: raziskave morskih psov, objave, turške vode, vzhodno Sredozemlje
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
12
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
REFERENCES
Akça, N. (2010): Underwater observations on the 
bioecology of Carcharhinus plumbeus (Nardo, 1827) 
inhabiting Boncuk Bay (Gökova Gulf). M.Sc. Thesis, 
Department of Biology, Adnan Menderes University, 45 
pp.
Akşıray, F. (1987): Türkiye Deniz Balıkları Ve Tayin 
Anahtarı, 2nd edn. Publication No. 3490. Istanbul: 
Istanbul University, 811 pp.
Avşar, D. (1996): Sex, age and growth of the spurdog 
(Squalus acanthias Linnaeus, 1758) in the southeastern 
Black Sea. Yugoslav Journal of Operations Research, 6, 
295-304.
Ayaşlı, S. (1937): Boğaziçi Balıkları. Istanbul: Cum-
huriyet Matbaası.
Başusta, N. (2002): Occurrence of a sawback angel-
shark (Squatina aculeata Cuvier, 1829) off the eastern 
Mediterranean coast of Turkey. Turk. J. Vet. Anim. Sci., 
26, 1177-1179.
Başusta, N. (2015): New records of neonate and 
juvenile sharks (Heptranchias perlo, Squatina aculeata, 
Etmopterus spinax) from the north-eastern Mediterrane-
an Sea. Mar. Biodiv., DOI 10.1007/s12526-015-0391-z
Başusta, N. (2016): Length-weight relationship of 
sandbar shark Carcharhinus plumbeus (Nardo, 1827) 
in Iskenderun Bay (north-eastern Mediterranean Sea). 
Rapp. Comm. int. Mer Médit., 41, 318.
Başusta, N., Ü. Erdem & C. Çevik (1998): An in-
vestigation on chondrichthyes in İskenderun Bay. Celal 
Bayar University Journal of Science, 1, 63-69.
Başusta N., C. Turan & A. Başusta (2015): New 
records of gravid female and adult male of the angular 
rough shark, Oxynotus centrina (Oxynotidae) from the 
northeastern Mediterranean. J. Black Sea / Mediterrane-
an Environment, 21, 92-95.
Başusta, N., A. Başusta & E.Ö. Özbek (2016): Carti-
laginous fishes and fisheries in the Mediterranean coast 
of Turkey.  In: Turan, C., Salihoğlu, B., Özgür Özbek, E., 
Öztürk, B. (Eds.) The Turkish Part of the Mediterranean 
Sea; Marine Biodiversity, Fisheries, Conservation and 
Governance. Turkish Marine Research Foundation (TU-
DAV), Publication No: 43, Istanbul, TURKEY. 248-274 
pp.
Bellonii, P. (1553): De aquatilibus, Librio duo. Cum 
eiconibus ad viuam ipforum effigiem, quoad eius fieri 
potuit, expreffis. Paris: Cum priuilegio Regis.
Benli, H. A., B. Cihangir & K. C. Bizsel (1993): A 
new record for the Sea of Marmara; (Family: Squalidae) 
Centrophorus granulosus (Bloch & Schneider, 1801. Tr. 
J. of Zoology, 17, 133-135.
Bilecenoğlu M., M. Kaya, B. Cihangir & E. Çiçek 
(2014): An updated checklist of the marine fishes of 
Turkey. Turk J Zool, 38, 901-929.
Bilge, G., H. Filiz & A. N. Tarkan (2010): Length-we-
ight relationship of velvet belly lantern shark Etmopterus 
spinax (Linnaeus, 1758) in Sığacık Bay (Aegean Sea). 
Istanbul University Journal of Fisheries and Aquatic 
Sciences, 25, 1-8.
Cihangir, B., A. Ünlüoğlu & E. M. Tıraşın (1997): 
Distribution and some biological aspects of the lesser 
spotted dogfish (Chondrichthyes, Scyliorhinus canicula, 
Linnaeus, 1758) from the northern Aegean Sea. Mediter-
raneran Fisheries Congress, 9-11 April 1997, İzmir, pp. 
585-603.
Demirhan S. A. & K. Seyhan (2007: Maturity and 
fecundity of spiny dogfish (Squalus acanthias L., 1758) 
in the eastern Black Sea. Turk J Zool., 31, 301-308.
Demirhan, S. A., H. Öğüt, S. Engin, N. Başusta & 
E. Genç (2006): Difficulties in age readings from dorsal 
spines of spiny dogfish Squalus acanthias L., 1758. 
In: Başusta, N., Keskin, Ç., Serena, F., Seret, B. (Eds.), 
2006. “The Proceedings of the Workshop on Mediter-
ranean Cartilaginous Fish with Emphasis on Southern 
and Eastern Mediterranean” Turkish Marine Research 
Foundation. Istanbul- TURKEY. Publication Number 23, 
pp. 28-34.
Demirhan, S.A., K. Seyhan & N. Başusta (2007): 
Dietary overlap in spiny dogfish (Squalus acanthias) and 
thornback ray (Raja clavata) in the southeastern Black 
Sea. Ekoloji, 16, 1-8.
Deveciyan, K. (1926): Peĉhe et Pećheries en Turquie. 
İstanbul: Imprimerie de l’Administration de la Dette 
Publique Ottomane.
Doğan, K. (2006): The production and economic 
importance of sharks in Turkey. In: Başusta, N., Keskin, 
Ç., Serena, F., Seret, B. (Eds.), 2006. “The Proceedings 
of the Workshop on Mediterranean Cartilaginous Fish 
with Emphasis on Southern and Eastern Mediterranean” 
Turkish Marine Research Foundation. Istanbul- TURKEY. 
Publication Number 23, pp. 79-87.
Düzgüneş, E., I. Okumuş, I. Feyzioğlu & N. Sivri 
(2006): Population parameters of spiny dogfish, Squalus 
acanthias from the Turkish Black Sea coast and its com-
mercial exploitation in Turkey. In: Başusta, N., Keskin, 
Ç., Serena, F., Seret, B. (Eds.), 2006. “The Proceedings 
of the Workshop on Mediterranean Cartilaginous Fish 
with Emphasis on Southern and Eastern Mediterranean” 
Turkish Marine Research Foundation. Istanbul- TURKEY. 
Publication Number 23, pp. 1-9.
Erazi, R.A.R. (1942): Marine fishes found in the Sea 
of Marmara and in the Bosphorus. Revue de la Faculte 
´ des Sciences de l’Université d’Istanbul B 7, 103–115.
Erdoğan, Z. Z., H. Torcu Koç, D. Türker Çakır, V. 
Nerlović & J. Dulčić (2004): Sexual dimorphism in the 
small-spotted catshark, Scyliorhinus canicula (L., 1758), 
from the Edremit Bay (Turkey). Annales ser. hist nat., 14, 
165-170.
Ergüden, D., M. Gürlek & C. Turan (2013): A young 
Isurus oxyrinchus Rafinesque, 1810 (Chondrichthyes: 
Lamnidae) individual captured from Iskenderun Bay, 
Turkey. Medit. Mar. Sci., 14, 463-480.
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
13
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
Ergüden, D., M. Gürlek & C. Turan (2015): Occur-
rence of the thresher Alopias vulpinus (Bonnaterre, 
1788) from the northeastern Mediterranean coast of 
Turkey. Biharean Biologist, 9, 76-77.
Ergüden, D., M. Çekiç, S. Alagöz Ergüden, A. Altun 
& N. Uygur (2017): Occurrence of adult female kitefin 
shark Dalatias licha (Bonnaterre, 1788) in Iskenderun 
Bay (Eastern Mediterranean, Turkey). Comm. J. Biol., 1, 
60-62.
Eronat, E. G. T. & O. Özaydın (2014): Length-weight 
relationship of cartilaginous fish species from central 
Aegean Sea (İzmir Bay and Sığacık Bay). Ege J Fish Aqua 
Sci., 31, 119-125.
Eryılmaz, L., E. Yemişken & C. Dalyan (2011): The 
first documented record of genus Mustelus (Chondri-
chthyes: Triakidae) in the Black Sea. Turkish Journal of 
Fisheries and Aquatic Sciences, 11, 157-160.
Filiz, H. (2009): Diet composition of smooth-hound, 
Mustelus mustelus (Linnaeus, 1758) in Aegean Sea, 
Turkey. Belg. J. Zool., 139, 81-84.
Filiz, H. (2018): Year-round aggregation of sandbar 
shark, Carcharhinus plumbeus (Nardo, 1827), in Boncuk 
Cove in the southern Aegean Sea, Turkey (Carcharhini-
formes: Carcharhinidae). Zoology in the Middle East, 
http://dx.doi.org/10.1080/09397140.2018.1540148.
Filiz, H. & H. Kabasakal (2015): Photographic record 
of the spinner shark, Carcharhinus brevipinna (Müller 
& Henle, 1839), in Gökova Bay (south Aegean Sea, 
Turkey). Annales ser. hist. nat., 25, 123-128.
Filiz, H. & S. Mater (2002): A Preliminary Study on 
Length-Weight Relationships for Seven Elasmobranch 
Species from North Aegean Sea, Turkey. E.U. Journal of 
Fisheries & Aquatic Sciences, 19, 401-409.
Filiz, H. & E. Taşkavak (2006): Sexual dimorphism 
in the head, mouth and body morphology of the lesser-
-spotted dogfish, Scyliorhinus canicula, from Turkey. In: 
Başusta, N., Keskin, Ç., Serena, F., Seret, B. (Eds.), 2006. 
“The Proceedings of the Workshop on Mediterranean 
Cartilaginous Fish with Emphasis on Southern and Eastern 
Mediterranean” Turkish Marine Research Foundation. 
Istanbul- TURKEY. Publication Number 23, pp. 48-59.
Geldiay, R. & S. Mater (1968): Two species of fish 
found in the Aegean Sea (Oxynotus centrina L. and 
Ophisurus serpens L.). Scientific Reports of the Faculty 
of Science, Ege University, No. 52, 10 pp.
İşmen, A., C. Ç. Yığın, U. Altınağaç & A. Ayaz (2009): 
Length-weight relationships for ten shark species from 
Saros Bay (north Aegean Sea). J. Appl. Ichthyol. 25 
(Suppl. 1), 109–112.
İşmen, A., C. Ç. Yığın, H. İnceoğlu, M. Arslan, B. 
Daban, S. Kale, E. Kocabaş & M, Şirin (2013): Chond-
richthyan bycatches in the beam trawl shrimp fishery of 
the Marmara Sea. Rapp. Comm. int. Mer Médit., 40, 487.
Kabasakal, H. (1998a): The first record of the blunt-
nose six-gill shark [Hexanchus griseus (Bonnaterre, 
1788)] in the Sea of Marmara. Acta Adriat., 39, 67-70.
Kabasakal, H. (1998b): A note on the occurrence of 
the thresher sharl, Alopias vulpinus from south-western 
Black Sea. J. Mar. Biol. Ass. U. K., 78, 685-686.
Kabasakal, H. (2001): Preliminary data on the fee-
ding ecology of some selachians from the north-eastern 
Aegean Sea. Acta Adriat., 42, 15-24.
Kabasakal, H. (2002a): Capture of a female basking 
shark, Cetorhinus maximus (Gunnerus, 1765), from 
southern Turkey. Annales ser. hist. nat., 12, 31-34.
Kabasakal, H. (2002b): Elasmobranch species of the 
seas of Turkey. Annales ser. hist. nat., 12, 15-22.
Kabasakal, H. (2002c): Cephalopods in the stomach 
contents of four elasmobranch species from the northern 
Aegean Sea. Acta Adriat., 43, 17-24.
Kabasakal, H. (2002d): Stomach contents of the 
longnose spurdog, Squalus blainvillei (Risso, 1826) from 
the north-eastern Aegean Sea. Annales ser. hist. nat., 12, 
173-176.
Kabasakal, H. (2003a): Historical records of the gre-
at white shark, Carcharodon carcharias (Linnaeus, 1758) 
(Lamniformes: Lamnidae), from the Sea of Marmara. 
Annales ser. hist. nat., 13, 173-180.
Kabasakal, H. (2003b): Historical and contemporary 
records of sharks from the Sea of Marmara, Turkey. 
Annales ser. hist. nat., 13, 1-12.
Kabasakal, H. (2004a): Preliminary observations on the 
reproductive biology and diet of the bluntnose sixgill shark, 
Hexanchus griseus (Bonnaterre, 1788) (Chondrichthyes: 
Hexanchidae), in Turkish seas. Acta Adriat., 45, 187-196.
Kabasakal, H. (2004b): Cetorhinus maximus (Gun-
nerus, 1765) (Lamniformes, Cetorhinidae) in the Gulf of 
Antalya in 1987: A summery of the previous records of 
the species from Turkish coastal waters in the Mediterra-
nean. Annales ser. hist. nat., 14, 29-34.
Kabasakal, H. (2005): Pontic occurrence of the 
bluntnose sixgill shark, Hexanchus griseus (Bonnaterre, 
1788) (Chondrichthyes: Hexanchidae). Annales ser. hist. 
nat., 15, 65-68.
Kabasakal, H. (2006): Distribution and biology of the 
bluntnose sixgill shark, Hexanchus griseus (Bonnaterre, 
1788) (Chondrichthyes: Hexanchidae), from Turkish 
waters. Annales ser. hist. nat., 16, 29-36.
Kabasakal, H. (2007): Incidental captures of thresher 
sharks (Lamniformes: Alopiidae) from Turkish coastal 
waters. Annales ser. hist. nat., 17, 23.28.
Kabasakal, H. (2008): Two recent records of the great 
white sharks, Carcharodon carcharias (Linnaeus, 1758) 
(Chondrichthyes: Lamnidae), caught in Turkish waters. 
Annales ser. hist. nat., 18, 11-16.
Kabasakal, H. (2009a): Observations on a rare shark, 
Oxynotus centrina (Chondrichthyes: Oxynotidae), in the 
Sea of Marmara (north-western Turkey). Pan-American 
Journal of Aquatic Sciences, 4, 609-612.
Kabasakal, H. (2009b): On the occurrence of the blunt-
nose sixgill shark, Hexanchus griseus (Chondrichthyes: 
Hexanchidae), in the Sea of Marmara. Marine Biodiversity 
Records. doi:10.1017/S1755267209001018, Vol. 2, e110.
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
14
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
Kabasakal, H. (2009c): On the capture of a large 
basking shark Cetorhinus maximus (Chondrichthyes: Ce-
torhinidae) in the Bay of Edremit (northeastern Aegean 
Sea). Annales ser. hist. nat., 19, 11-16.
Kabasakal, H. (2010a): Historical and contemporary 
records of the angular rough shark Oxynotus centrina 
(Chondrichthyes: Oxynotidae) in Turkish waters. Med. 
Mar. Sci., 11, 361-367.
Kabasakal, H. (2010b): On the occurrence of 
the blue shark, Prionace glauca (Chondrichthyes: 
Carcharhinidae), off Turkish coast of northern Aege-
an Sea. Marine Biodiversity Records. doi:10.1017/
S1755267210000266, Vol. 3, e31.
Kabasakal, H. (2010c): A review of newspaper and 
internet portrayals of the sixgill shark, Hexanchus griseus 
(Bonnaterre, 1788) (Chondrichthyes: Hexanchidae), 
caught in Turkish waters between 1974-2009. Annales 
ser. hist. nat., 20, 175-180.
Kabasakal, H. (2010d): Post-release behavior and 
anthropogenic injuries of the bluntnose sixgill shark, 
Hexanchus griseus (Bonnaterre, 1788) (Chondrichthyes: 
Hexanchidae) in Turkish waters. Annales ser. hist. nat., 
20, 39-46.
Kabasakal, H. (2011a): On an old record of Carc-
harodon carcharias (Chondrichthyes: Lamnidae) from 
the Bosphoric waters. Marine Biodiversity Records. 
doi:10.1017/S1755267211000613, Vol. 4, e61.
Kabasakal, H. (2011b): Türk Sularında Köpekbalıkları 
(Sharks of Turkish Waters). 4Deniz Yayınları, İstanbul.
Kabasakal, H. (2013a): Bluntnose sixgill shark, 
Hexanchus griseus (Chondrichthyes: Hexanchidae), 
caught by commercial fishing vessels in the seas of 
Turkey between 1967 and 2013. Annales ser. hist. nat., 
23, 33-48.
Kabasakal, H. (2013b): Two young basking sharks, 
Cetorhinus maximus (Gunnerus, 1765), caught in the 
Levantine basin off the Turkish coast (eastern Mediterra-
nean Sea). Medit. Mar. Sci., 14, 463-480.
Kabasakal, H. (2013c): Rare but present: Status of 
basking shark, Cetorhinus maximus (Gunnerus, 1765) in 
eastern Mediterranean. Annales ser. hist. nat., 23, 127-132.
Kabasakal, H. (2014): The status of the great 
white shark (Carcharodon carcharias) in Turkey’s 
waters. Marine Biodiversity Records. doi:10.1017/
S1755267214000980, Vol. 7, e109.
Kabasakal, H. (2015a): Occurrence of the angular 
rough shark, Oxynotus centrina (Chondrichthyes: Oxy-
notidae) in the eastern Mediterranean. Annales ser. hist. 
nat., 25, 1-10.
Kabasakal, H. (2015b): Occurrence of shortfin mako 
shark, Isurus oxyrinchus Rafinesque, 1810, off Turkey’s 
coast. Marine Biodiversity Records. doi:10.1017/
S1755267215001104, Vol. 8, e134.
Kabasakal, H. (2015c): Historical occurrence of 
Carcharhinus spp. in the Sea of Marmara during the 
1950s. Marine Biodiversity Records. doi:10.1017/
S1755267215000305, Vol. 8, e48.
Kabasakal, H. (2016a): Historical dispersal of the 
great white shark, Carcharodon carcharias, and bluefin 
tuna, Thunnus thynnus, in Turkish waters: Decline of a 
predator in response to the loss of its prey. Annales ser. 
hist. nat., 26, 213-220.
Kabasakal, H. (2016b): Status of cartilaginous fishes 
in the Sea of Marmara. In: Özsoy, E., Çağatay, M.N., 
Balkıs, N., Balkıs, N., Öztürk, B. (Eds.) (2016). The Sea of 
Marmara; Marine Biodiversity, Fisheries, Conservation 
and Governance. Turkish Marine Research Foundation 
(TUDAV), Publication No: 42, Istanbul, TURKEY, 903-
918 pp.
Kabasakal, H. (2017a): Remarks on incidental captu-
res of deep-sea sharks in Marmaric shelf waters. Annales 
ser. hist. nat., 27, 37-144.
Kabasakal, H. (2017b): Notes on historical and 
contemporary catches of lamniform sharks in Turkish 
waters. Annales ser. hist. nat., 27, 51-58.
Kabasakal, H. & M. Bilecenoğlu (2014): Not disap-
peared, just rare! Status of the bramble shark, Echinorhi-
nus brucus (Elasmobranchii: Echinorhinidae) in the seas 
of Turkey. Annales ser. hist. nat., 24, 93-98.
Kabasakal, H. & C. Dalyan (2011): Recent records of 
the bramble shark, Echinorhinus brucus (Chondrichthyes: 
Echinorhinidae), from the Sea of Marmara. Marine Bio-
diversity Records. doi:10.1017/S1755267211000108, 
Vol. 4, e12
Kabasakal, H. & A. De Maddalena (2011): A huge 
shortfin mako shark Isurus oxyrinchus Rafinesque, 1810 
(Chondrichthyes: Lamnidae) from the waters of Marma-
ris, Turkey. Annales ser. hist. nat., 21, 21-24.
Kabasakal, H. & S. Ö. Gedikoğlu (2008): Two 
new-born great white sharks, Carcharodon carcharias 
(Linnaeus, 1758) (Lamniformes; Lamnidae) from Turkish 
waters of the northern Aegean Sea. Acta Adriat., 49, 
125-135.
Kabasakal, H. & S. Ö. Gedikoğlu (2015): Shark at-
tacks against humans and boats in Turkey’s waters in the 
twentieth century. Annales ser. hist. nat., 25, 115-122.
Kabasakal, H. & P. İnce (2008): Note on a sharpnose 
sevengill shark, Heptranchias perlo (Bonnaterre, 1788) 
(Chondrichthyes: Hexanchidae), stranded in Saroz Bay (NE 
Aegean Sea, Turkey). Annales ser. hist. nat., 18, 173-176.
Kabasakal, H. & E. Kabasakal (2002): Morphome-
trics of young kitefin sharks, Dalatias licha (Bonnaterre, 
1788), from northeastern Aegean Sea, with notes on its 
biology. Annales ser. hist. nat., 12, 161-166.
Kabasakal, H. & E. Kabasakal (2004): Sharks captu-
red by commercial fishing vessels off the coast of Turkey 
in the northern Aegean Sea. Annales ser. hist. nat., 14, 
171-180.
Kabasakal, H. & Ö. Kabasakal (2013): First record 
of a shortfin mako shark, Isurus oxyrinchus Rafinesque, 
1810 (Chondrichthyes: Lamnidae) from the Bay of Saroz 
(NE Aegean Sea). Annales ser. hist. nat., 23, 27-32.
Kabasakal, H. & Ö. Kabasakal (2014): Status of 
angelshark, Squatina squatina (Elasmobranchii: Squati-
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
15
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
niformes: Squatinidae) in the Sea of Marmara. Annales 
ser. hist. nat., 24, 41-46.
Kabasakal, H. & Ö. Kabasakal (2015): Recent record 
of the great white shark, Carcharodon carcharias (Linna-
eus, 1758), from central Aegean Sea off Turkey’s coast. 
Annales ser. hist. nat., 25, 11-14.
Kabasakal, H. & S. Ü. Karhan (2008): On the 
occurrence of the bigeye thresher shark, Alopias 
superciliosus (Chondrichthyes: Alopiidae), in Turkish 
waters. Marine Biodiversity Records. doi:10.1017/
S1755267207007452, Vol. 1, e69.
Kabasakal, H. & S. Ü. Karhan (2015): Shark biodiver-
sity in the Sea of Marmara: departures and arrivals over 
a century. Marine Biodiversity Records. doi:10.1017/
S1755267215000342, Vol. 8, e59.
Kabasakal, H. & N. Ünsal (1999): Observations on 
Etmopterus spinax (Pisces: Squalidae), from the north-
-eastern Aegean Sea. Bilješke – Notes, 81, 12 pp.
Kabasakal, H., M. İ. Öz, S. Ü. Karhan, Z. Çaylarbaşı 
& U. Tural (2005): Photographic evidence of the occur-
rence of bramble shark, Echinorhinus brucus (Bonnater-
re, 1788) (Squaliformes: Echinorhinidae) from the Sea of 
Marmara. Annales ser. hist. nat., 15, 51-56.
Kabasakal, H., A. Yarmaz & S. Ö. Gedikoğlu (2009): 
Two juvenile great white sharks, Carcharodon carchari-
as (Linnaeus, 1758) (Chondrichthyes; Lamnidae), caught 
in the Northeastern Aegean Sea. Annales ser. hist. nat., 
19, 127-134.
Kabasakal, H., C. Dalyan & A. Yurtsever (2011): 
Additional records of the bigeye thresher shark Alopias 
superciliosus (Lowe, 1839) (Chondrichthyes: Lamnifor-
mes: Alopiidae) from Turkish waters. Annales ser. hist. 
nat., 21, 143-148.
Kabasakal, H., S.Ü. Karhan & S. Sakınan (2017): Revi-
ew of the distribution of large sharks in the seas of Turkey 
(Eastern Mediterranean). Cah. Biol. Mar., 58, 219-228.
Kabasakal, H., E. Bayrı & E. Ataç (2018): Recent 
records of the great white shark, Carcharodon carcha-
rias (Linnaeus, 1758) (Chondrichthyes: Lamnidae), in 
Turkish waters (eastern Mediterranean). Annales ser. 
hist. nat., 28, 93-98.
Keskin, Ç. & F. S. Karakulak (2006): Preliminary 
results on depth distribution of cartilaginous fish in the 
north Aegean Sea and their fishing potential in Summer 
2001. In: Başusta, N., Keskin, Ç., Serena, F., Seret, B. 
(Eds.), 2006. “The Proceedings of the Workshop on 
Mediterranean Cartilaginous Fish with Emphasis on 
Southern and Eastern Mediterranean” Turkish Marine 
Research Foundation. Istanbul- TURKEY. Publication 
Number 23, pp. 69-78.
Kıdeyş, A. E. (1997): Occurrence of the basking 
shark, Cetorhinus maximus in the northern Levantine, 
the eastern Mediterranean. Mediterranean Fisheries 
Congress, 9-11 April 1997, Izmir, Turkey, 4 pp.
Kutaygil, N. & N. Bilecik (1977): Recherches sur 
le Squalus acanthias L. du littoral Anatolien de la Mer 
Noire. Rapp. Comm. int. Mer Médit., 24, 80-83.
Mater, S. (2005): Denizlerimizde yeni bir köpekbalığı, 
Alopias superciliosus (Lowe, 1841). Aquaculture, 1, 10.
Meriç, N. (1995): A study on existence of some fishes 
on the continental slope of the Sea of Marmara. Tr. J. of 
Zoology, 19, 191-198.
Ninni, E. (1923): Primo contributo allo studio dei 
pesci e della pesca nelle acque dell’Impero Ottomano. 
Venezia: Premiate Officine Grafiche Carlo Ferrari.
Oral, M. (2010): A preliminary study on feeding 
habits of black-mouth catshark Galeus melastomus, 
Rafinesque, 1810 sampled from the Sea of Marmara. 
Marmara Denizi 2010 Sempozyumu, 25-26 Eylül 2010, 
İstanbul, pp. 312-316.
Özcan, E.İ. & N. Başusta (2016): Digestive system 
contents of the Mustelus mustelus (Linnaeus, 1758) in-
habiting northeastern Mediterranean. Fırat Unv. Journal 
of Science, 28, 7-12.
Öztürk, B. (2006): Save the sandbar sharks of Bon-
cuk Bay, Turkey. In: Başusta, N., Keskin, Ç., Serena, F., 
Seret, B. (Eds.), 2006. “The Proceedings of the Workshop 
on Mediterranean Cartilaginous Fish with Emphasis on 
Southern and Eastern Mediterranean” Turkish Marine 
Research Foundation. Istanbul- TURKEY. Publication 
Number 23, pp. 42-47.
Özütemiz Ş., M. Kaya & O. Özaydın (2009): Growth 
and feeding characteristics of two shark species [Galeus 
melastomus Rafinesque, 1810 and Squalus blainvillei 
(Risso, 1826)] from Sığacık Bay (Aegean Sea). E. U. 
Journal of Fisheries & Aquatic Sciences, 26, 211-217.
Serena F. (2005): Field identification guide to the 
sharks and rays of the Mediterranean and Black Sea. 
FAO Species Identification Guide for Fishery Purposes.
Rome, FAO. 97 pp.
Tunçer, S. & H. Kabasakal (2016): Capture of a juve-
nile shortfin mako shark, Isurus oxyrinchus Rafinesque, 
1810 (Chondrichthyes: Lamnidae) in the Bay of Edremit, 
northern Aegean Sea (Turkey). Annales ser. hist. nat., 26, 
31-36.
Türker Çakır, D., H. Torcu Koç & Z. Erdoğan (2006): 
Some biological aspects of the lesser spotted dogfish 
Scyliorhinus canicula (Linnaeus, 1758) in Edremit Bay 
(the northern Aegean Sea) Turkey. In: Başusta, N., Keskin, 
Ç., Serena, F., Seret, B. (Eds.), 2006. “The Proceedings 
of the Workshop on Mediterranean Cartilaginous Fish 
with Emphasis on Southern and Eastern Mediterranean” 
Turkish Marine Research Foundation. Istanbul- TURKEY. 
Publication Number 23, pp. 17-27.
Ulutürk T. (1987): Fish fauna, background radioac-
tivity of the Gökçeada marine environment. Journal of 
Aquatic Products, University of Istanbul, 1, 95–119.
White W.T., D.A. Ebert, G.J.P. Naylor, H-C. Ho, P. 
Clerkin, A. Veríssimo & Cotton C.F. (2013): Revision of 
the genus Centrophorus (Squaliformes: Centrophoridae) 
Part 1 – Redescription of Centrophorus granulosus (Blo-
ch & Schneider), a senior synonym of C. acus Garman 
and C. niaukang Teng. Zootaxa, 3752, 035–072.
ANNALES · Ser. hist. nat. · 29 · 2019 · 1
16
Hakan KABASAKAL: A REVIEW OF SHARK RESEARCH IN TURKISH WATERS, 1–16
Yağlıoğlu, D., T. Deniz, M. Gürlek, D. Ergüden & C. 
Turan (2015): Elasmobranch bycatch in a bottom trawl 
fishery in the Iskenderun Bay, northeastern Mediterrane-
an. Cah. Biol. Mar., 56, 237-243.
Yığın C. Ç. & A. İşmen (2013): Reproductive biology 
of spiny dogfish Squalus acanthias, in the north Aege-
an Sea. Turk. J. Fish. Aquat. Sci., 13: 169-177. DOI: 
10.4194/1303-2712-v13_1_20.
Yığın, C. Ç., A. İşmen & U. Önal (2016): Occurrence 
of a rare shark, Oxynotus centrina (Chondrichthyes: 
Oxynotidae), from Saros Bay, North Aegean Sea, Turkey. 
J. Black Sea / Mediterranean Environment, 22, 103-109.