ANNALES · Ser. hist. nat. · 30 · 2020 · 2
131
received: 2020-07-08                 DOI 10.19233/ASHN.2020.15
REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS 
PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE MEDITERRANEAN: 
HISTORICAL AND RECENT DATA
 
Paraskevi K. KARACHLE, Caterina STAMOULI & Aikaterini DOGRAMMATZI
Hellenic Centre for Marine Research, Institute of Marine Biological Resources and Inland Waters, P.O. Box 712, 19013 Anavyssos Attiki, Greece
e-mail: pkarachle@hcmr.gr
ABSTRACT
This paper presents a thorough literature review of the sharpnose sevengill shark Heptranchias perlo (Bon-
naterre, 1788) in the Mediterranean Sea, creating a database with information on its distribution. Moreover, it 
provides some biological information on a female specimen of this rare and near threatened shark, caught during 
experimental sampling in the Myrtoon Sea, southwestern Aegean archipelago, Greece. The aim of this study is to 
provide a synopsis of the existing information on the species in the Mediterranean Sea for future updates of the 
assessments of the species conservation status in this area.
Key words: elasmobranchs, diet, fecundity, Mediterranean distribution
REVISIONE DELLO SQUALO MANZO HEPTRANCHIAS PERLO (CHONDRICHTHYES: 
HEXANCHIDAE) IN MEDITERRANEO: DATI STORICI E RECENTI
SINTESI
L’articolo presenta una revisione approfondita della letteratura inerente lo squalo manzo Heptranchias perlo 
(Bonnaterre, 1788) nel mare Mediterraneo, creando una banca dati contenente informazioni sulla sua distri-
buzione. Fornisce inoltre alcune informazioni biologiche su un esemplare femmina di questa rara specie, quasi 
minacciata, catturato durante un campionamento sperimentale nel mare Myrtoon, arcipelago dell’Egeo sud-occi-
dentale, in Grecia. Lo scopo di questo studio è fornire una sinossi delle informazioni esistenti sullo squalo manzo 
nel Mediterraneo per futuri aggiornamenti delle valutazioni dello stato di conservazione delle specie in quest’area.
Parole chiave: elasmobranchi, dieta, fecondità, distribuzione in Mediterraneo
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
132
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
INTRODUCTION
The sharpnose sevengill shark Heptranchias 
perlo (Bonnaterre, 1788) is a shark species of cir-
cumglobal distribution, found in tropical and tem-
perate seas excluding the northeast Pacific (Froese 
& Pauly, 2019). In the literature it is often acknowl-
edged as a rare species, and quite a few records of 
its presence have been reported in Mediterranean 
waters (e.g., Papaconstantinou, 2014; Guallart et 
al., 2019a; 2019b and references therein).
The sharpnose sevengill shark is considered a 
“near threatened” species according to the IUCN 
red list, yet this assessment is rather old and needs 
to be updated (Paul & Fowler, 2003). In the Medi-
terranean, the species is “data deficient” (Soldo & 
Bariche, 2016). Indeed, information regarding its 
biology and population status is very scarce, mak-
ing any assessment very difficult. In their analysis of 
data gaps in the biological knowledge concerning 
the Mediterranean, Dimarchopoulou et al. (2017) list 
the species under the least studied ones. Moreover, 
as it is considered a by-catch of deep-sea trawling 
fisheries (Paul & Fowler, 2003), the official capture 
data as presented in the FAO statistics (http://www.
fao.org/fishery/statistics/global-capture-production/
en; accessed 22 June 2020) are almost nonexistent. 
In fact, the only landing reports in the aforemen-
tioned global statistics cite 2 tons per year for the 
years 2008, 2009, 2010 and 2014, and are provided 
solely by Malta. It is a common practice in Mediter-
ranean fisheries (especially in the bottom trawling 
industry) that when shark species are caught, the 
head, fins and internal organs are removed and 
discarded overboard, and quite often the body is 
skinned in order to facilitate its sale. Thus, it is not 
possible to taxonomically identify the “fish” that is 
landed in the fishing markets, which leads to the 
mislabeling of the species (Jacquet & Pauly, 2003; 
Bornatowski et al., 2013), and therefore the land-
ings are underreported.
Due to the sporadic records of the species in 
Mediterranean waters, very little information exists 
on its biology. Apart from the work of Capapé (1980), 
who thoroughly studied the biology of the species 
in Tunisian waters, there is almost no other related 
information. In this work we present additional data 
on the presence of the species in the Myrtoon Sea, 
southwestern Aegean archipelago, Greece, and pro-
vide information on its fecundity, including ovarian 
Fig. 1: (A) The female sharpnose sevengill shark Heptranchias perlo caught in the Myrtoon Sea, Greece, 4 August 
2019 (scale bar = 10 cm); (B) detail of the head where the longline hook can be seen; and (C) the viscera, with 
eggs as seen in the ovary and the stomach of the fish (scale bar=1 cm). 
Sl. 1: (A) Samica morskega psa sedmeroškrgarja Heptranchias perlo, ujeta 4. avgusta 2019 v mirtonskem morju 
(Grčija) (merilo = 10 cm); detajl glave z vidnim parangalskim trnkom: in (C) drobovina, jajca v ovariju in želodec 
ribe (merilo = 1 cm).
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
133
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
follicle (hereafter referred to as “egg”) morphometry, 
and diet. Finally, a literature review is provided to 
summarise the records/information on the species 
in the Mediterranean as retrieved from literature, 
using as a starting point the work by Guallart et al. 
(2019a; 2019b) and elaborating further on additional 
data. The aim of this note is to (a) provide data on 
the species that could help elucidate its biology and 
ecology and fill in the knowledge gaps, (b) clarify the 
species’ distributional status in the Mediterranean, 
and (c) provide a review-report that could be useful 
in future updates of the assessments of the species 
status in the Mediterranean.
MATERIAL AND METHODS
On 4 August 2019 a female sharpnose sevengill 
shark individual (Fig. 1) was caught during the 
experimental Mediterranean Trawl Survey sampling 
(MEDITS; Bertrand et al., 2002), conducted in the 
frame of the Greek National Data Collection Frame-
work (DCF) programme, in the area of Myrtoon 
Sea (coordinates: 37.36833333N, 22.98027778E) 
at a depth of 550 m. When landed the shark was 
already dead and the MEDITS protocol on data 
recording was applied. According to this protocol 
(Spedicato et al., 2019) target species are divided 
into two groups, with H. perlo being included in 
the G1 list; hence, the measurements taken accord-
ing to the protocol were as follows: total number 
of individuals, total weight and individual length, 
and biological parameters including sex, maturity, 
and individual weight. Following the identification, 
the fish was measured for total length with the 
tail in a natural position (TL; in cm) and weighed 
(W; in g), as well as sexed, and the horizontal and 
vertical mouth openings were measured (HMO 
and VMO, respectively; with an accuracy of 0.01 
mm). After dissection the ovaries were removed, the 
eggs were measured (maximum egg diameter [ED], 
in 0.01 mm) using a digital calliper, and weighed 
(egg weight [OW], in g), and the stomach was kept 
in a deep-freezer for further examination. Finally, 
the liver was weighed (LO, in g). In the laboratory, 
stomach contents were examined, identified to the 
lowest possible taxon, and each prey was weighed 
(in 0.001 g). Based on the % weight (% WF) contri-
bution of each prey category, the fractional trophic 
level (TROPH) of the species was estimated using 
TrophLab (Pauly et al., 2000). In order to assess 
the feeding preferences of the species, a literature 
review was performed. Data were extracted and 
tabulated, and TROPHs were estimated.
Furthermore, as the recent review of Guallart et 
al. (2019a, 2019b) on the presence of the species 
in the Mediterranean was missing records from the 
Hellenic Seas, an extensive literature review was 
conducted for the entire Mediterranean basin. For 
this purpose, the aforementioned works were used 
as a basis and further elaborated through Goog-
leSearch. The keywords of “Heptranchias perlo” 
and “Mediterranean” were used, and all the papers 
retrieved were examined and tabulated. In addition, 
a second table was created including secondary 
references of the species (i.e., previous and old 
records of the species within the papers examined, 
but the original references were not available to the 
authors).
RESULTS AND DISCUSSION
In this account, information on the biology and 
distribution of the sharpnose sevengill shark Hep-
tranchias perlo in the Mediterranean Sea is compiled. 
The species is considered by-catch in the fisheries 
in Greece (mainly in longlines targeting tuna and 
swordfish, as well as in trawlers), but at times it is 
marketed rather than discarded (e.g., Damalas & 
Vassilopoulou, 2011). In the case of the individual 
examined, there were clear indications that the speci-
men had already been caught some time in the past, 
as a large hook was found in the lower jaw (Fig. 1B), 
like those traditionally used in longline fisheries tar-
geting large pelagic fish (e.g., tuna, swordfish) in the 
country. Thus the fish had been caught and released 
by a professional fisher in the past, a practice that 
was also documented by Megalofonou et al. (2005) 
and Vassilopoulou et al. (2007).
Tab. 1: Measurements taken on a female sharpnose se-
vengill shark Heptranchias perlo, caught in Myrtoon Sea, 
SouthWest Aegean Archipelago, Greece, August 4th 2019.
Tab. 1: Meritve, opravljene na samici morskega psa sed-
meroškrgarja Heptranchias perlo, ujete 4. avgusta 2019 v 
mirtonskem morju (jugozahodno Egejsko morje, Grčija).
Measurement Value
Total length (in cm) 114
Total weight (in g) 6730
Horizontal mouth opening (in mm) 51.99
Vertical mouth opening (in mm) 40.74
Liver weight (in g) 788
Ovary total weight (in g) 425
Number of eggs 22
Mean egg diameter (in mm) ±standard 
error (range)
34.30±0.58
(29.03-38.44)
Mean egg weight (in g)±standard error 
(range)
19.82±0.78
(11-26)
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
134
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Tab. 2: Review of studies on feeding habits of the sharpnose sevengill shark Heptranchias perlo. LT= length type; 
TL=total length; LR=length range; n=number of individuals; F = frequency of occurrence, N= numerical percentage, 
W = percentage by weight; IRI=index of relative importance. 
Tab. 2: Pregled raziskav o prehranjevalnih navadah morskega psa sedmeroškrgarja Heptranchias perlo. LT= tip dol-
žine; TL=celotna dolžina; LR=razpon dolžine; n=število osebkov; F = frekvenca pojavljanja, N= številčni delež, W = 
biomasni delež; IRI=indeks relativne pomembnosti plena. 
Area Date LT
LR 
(cm)
n Method Main prey
contribution 
of prey
[W or (N)]
TROPH SE Reference
Cuba Fish, benthic Crustacea - -
Sierra et al. 
(1994) in Froese 
& Pauly (2019)
coast off 
Namibia
1980-
1984
Myctophidae, 
Cephalopoda
- -
Macpherson 
(1989)
Scilly Isles 8/1999 TL 101 1
F, N, 
W
Cephalopoda 
(Illex coindetii)
100 (100) 4.50 0.37
Henderson & 
Williams (2001)
east coast 
of southern 
Africa
10 F Cephalopoda, fish 4.50 0.48
Bass et al. (1975) 
in Cortés (1999)
South Africa 8
F, N, 
W, IRI
Cephalopoda, fish
53, 47 (50, 
50)
4.50 0.61 Ebert (1990)
South Africa 12
F, N, 
W, IRI
Cephalopoda, fish 4.50 0.61
Barnett et al. 
(2012)
Taiwan TL
63.5-
118.5
28
F, N, 
W, IRI
Fish (Trichiurus lepturus, 
Trachurus japonicus, 
Myctophidae), Crustacea 
(Brachyura, Macrura 
Reptantia), Cephalopoda
98, 1, 1 
(65, 27, 8)
4.49 0.80 Ebert (1990)
Taiwan 36
F, N, 
W, IRI
Fish 4.50 0.80
Barnett et al. 
(2012)
Victoria, 
Australia
2-5/2003 TL
48.5-
90.0
67
F, N, 
W, IRI
Fish (Lepidorhynchus 
denticulatus, Apogonops 
anomalus, Trichiuridae), 
Cepalopoda (Nototodarus 
gouldi), shrimps
86.4, 12.8, 
0.8 (87.2, 
10.7, 2.1)
4.49 0.76 Braccini (2008)
Victoria, 
Australia
2-5/2003 TL
90.1-
136.5
49
F, N, 
W, IRI
Fish (Trichiuridae, 
Gempylidae, Lepidorhynchus 
denticulatus, Paraulopus 
nigripinnis, Centrolophidae), 
Cepalopoda, shrimps
96.9, 2.6, 
0.5 (90, 
8.3, 1.7)
4.50 0.79 Braccini (2008)
Central 
Atlantic
4/8-
17/9/1998
TL
39.0-
116.0
60
F, N, 
W
Fish (Heptranchias perlo, 
Congridae), Cephalopoda 
(Octopoda)
57.5, 28.4 
(45.9, 31.5)
4.34 0.67
Frentzel-Beyme & 
Koster (2002)
Tunisia 9 F
Fish, Crustacea, 
Cephalopoda
4.22 0.69
Capapé (1975a) 
in Cortés (1999)
Tunisia 125 F, N
Fish (Gadiculus argenteus, 
Hoplostethus mediterraneus, 
Trachyrhynchus trachyrhynchus), 
Decapoda (Nephrops 
norvegicus), Cephalopoda 
(Sepietta oweniana)
(72.1, 16.3, 
11.6)
4.33 0.74 Capapé (1980)
Myrtoon 
Sea
4/8/2019 TL 114 1 N, W
Cephalopoda (Octopus 
vulgaris, Loligo vulgaris)
100 (100) 4.50 0.46 present study
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
135
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Following the recommendations of Dimarchopoulou 
et al. (2017) and Karachle & Stergiou (2017), effort was 
made to obtain as much information possible even from 
a single specimen. However, the examination of only 
one individual allows very little potential for analysis, 
virtually just a presentation of morphological measure-
ments (Tab. 1) and a limited insight into its biology. A 
detailed description of the morphology and anatomy of 
the species, as well as various aspects of its biology (e.g., 
morphometric relations, reproduction, feeding) is given 
in Capapé (1980) based on 154 specimens from Tunisian 
waters. To date, the work of Capapé (1980) is the only 
in-depth scientific account on the species in the Mediter-
ranean, covering a wide range of biological aspects.
The sharpnose sevengill shark is a viviparous 
aplacental lecithotrophic species sensu Hamlett et 
al. (2005). According to a visual examination of the 
reproductive system, the specimen was a female (Fig. 
1C), its maturity stage was classified under category 
3b (ICES [2013] maturity scale), i.e., that of mature 
individual. This is also in accordance with the find-
ings of Capapé (1980), who identified the length at 
maturity for the species at 93 cm for males and 105 
cm for females. Overall, 22 eggs were counted and 
measured, exhibiting a mean diameter of 34.30±0.58 
mm and a mean weight of 19.82±0.78 g. These val-
ues, even slightly underweight, are above the limit of 
2.5 cm and 25 g identified by Capapé (1980) as with 
Fig. 2: The sharpnose sevengill shark Heptranchias perlo. (a) Number of publications with referen-
ces to the presence of the species in the Mediterranean; and (b) number of individuals reported in 
the Mediterranean. For more details, see Appendix 1 & 2.
Sl. 2: Morski pes sedmeroškrgar Heptranchias perlo. (a) Število objavljenih del, ki se nanašajo 
na prisotnost te vrste v Sredozemskem morju; in (b) število primerkov, o katerih so poročali v 
Sredozemlju. Za bolj natančne podatke glej Prilogi 1 & 2.
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
136
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
important vitellogenic activity. Taking into consid-
eration the dimensions for the characterization of the 
vitellogenic stage proposed by Correa de Carvalho 
et al. (2020), the female individual examined here 
was in intermediate vitellogenesis (stage III). In the 
Mediterranean Sea, the reported fecundity of the spe-
cies ranges between 6 to 20 eggs (Capapé, 1980 and 
references therein), thus making the number of eggs 
reported herein the highest recorded for the species 
in the basin.
The diet of the sharpnose sevengill shark has 
been studied in various areas of its distribution, both 
in the Atlantic and Pacific Oceans, as well as in 
the Mediterranean (Tab. 2). In all studies, the prey 
items were the same, with fish and Cephalopoda 
dominating the diet of the species, although in dif-
ferent ratios. It is noteworthy that in the study by 
Frentzel-Beyme & Koster (2002), conducted at the 
Great Meteor Seamount, central-east Atlantic, can-
nibalism was recorded (Tab. 2). In the present study 
the stomach of the shark contained exclusively two 
species of Cephalopoda (one individual of Octopus 
vulgaris [60.1 % WF] and one of Loligo vulgaris [39.9 
% W]) and the estimated TROPH was 4.50±0.46. In 
general, this TROPH value was estimated in the ma-
jority of the diet studies concerning the species (Tab. 
2). The lowest value, however, was estimated based 
on the diet composition of the specimens in Tunisia 
(4.22±0.69 [Capapé, 1975] and 4.33±0.74 [Capapé, 
1980]). These lower values should be attributed to 
the fact that crustaceans contributed more to the diet 
of the species than in other studies, where they were 
negligible or even not recorded. Nevertheless, all 
TROPH values estimated classify the species as an 
apex predator, a carnivore with preference for fish 
and cephalopods (sensu Stergiou & Karpouzi [2002] 
and Karachle & Stergiou [2017]).
Overall, the presence of H. perlo in the Mediter-
ranean Sea basin has been demonstrated in more 
than 90 published papers (Fig. 2a; Appendix 1 & 2). 
Most of the publications are from the eastern and 
western parts of the basin (31 and 28 publications, 
respectively), followed by the central part (21 publi-
cations), whereas only 7 publications make reference 
to the species in the Adriatic (Fig. 2a; Appendix 1 & 
2). However, we recognize that the scarcity of data 
or the absence of records related to specific areas of 
the basin may well indicate a lack of reports rather 
than an actual absence of the species. The number 
of specimens caught was only provided in half of 
the publications, with the sex reported even more 
rarely (Fig. 2b; Appendix 1 & 2). Moreover, cases in 
which the number of specimens recorded exceeded 
5 individuals per report were extremely rare (N>5 
in 8 publications [8.6%]), and there was only one 
publication (Capapé, 1980) with an extraordinary 
number of 120 individuals reported. Records with 
the highest numbers of specimens reported are 
from the East and Central Mediterranean, and this 
should be attributed to the work conducted in the 
Aegean Sea (e.g., Ismen et al., 2007, 2009; Damalas 
& Vassilopoulou, 2011) and in the coasts of Tunisia 
(e.g., El Kamel-Moutalibi et al., 2014; Rafrafi-Nouira 
et al., 2015; Capapé et al., 2018), with respect to 
the morphometry, the establishment of length-weight 
relationships, and the study of the biology of the spe-
cies (Appendix 1).
In conclusion, despite the literature review, the 
authors of the present article strongly believe that 
there might exist more records of the species, mainly 
in unpublished/unavailable survey data. It is therefore 
essential for data, especially those related to species 
that urgently need conservation actions, and even 
more so those collected through public funding, to 
be open and available to the scientific community, 
so that the status of the species can be adequately 
assessed and conservation measures can be based on 
most comprehensive scientific information. It is of 
high priority to investigate and report biological data 
of species when specimens are available, in particu-
lar with those found in small numbers and species 
lacking such information, to fill the gaps in essential 
knowledge and, as a result, improve our conservation 
efforts.
ACKNOWLEDGEMENTS
The sample examined within this research was 
collected during the MEDITS action of the EU Data 
Collection Framework Program.
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
137
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Appendix 1: List of records of Heptranchias perlo in the Mediterranean (literature examined by the authors; 
in blue references also included in Guallart et al., 2019a, b). F=female, M=male, N=number of individuals, 
LWR=length-weight relationship parameters, SEb=standard error of slope b, R2=correlation coefficient.
Priloga 1: Seznam zapisov o pojavljanju vrste Heptranchias perlo v Sredozemskem morju (pregledana literatura s 
strani avtorjev; v modrem so reference, ki so bile že vključene v Guallart s sod., 2019a, b). F=samica, M=samec, 
N=število primerkov, LWR=odnos med dolžino in težo, SEb=standardna napaka naklona b, R2=korelacijski koeficient.
Reference Date locality depth Sex N
length range
(in cm)
weight 
range
(in g)
LWR
comments
min max min max a b SEb R
2
East Mediterranean
Ben-Tuvia 
(1953)
1951-
1953
Israel 250
record from a photo, 
length measurement in 
approximation
Fredj & 
Maurin (1987)
E 
Mediterranean
presence of the species 
(depths from 0 up to >1000 
m)
Labropoulou & 
Papaconstantinou 
(2000)
N Aegean 
Sea
presence of the species in 
the area
Baino et al. 
(2001)
1994-
1999
E Aegean Sea
data from MEDITS 
expedition (biomass index 
1.4 kg/km2)
Filiz & Mater 
(2002)
7/1999-
3/2000
Edremit Bay, 
Gulbahce Bay, 
and Sigacik 
Bay, N Aegean 
Sea, Turkey
presence of the species in 
the area
Golani (2006) Israel
list of cartilagenous species 
in Israeli coasts. H. perlo is 
characterised as "prevalent"
Megalofonou 
et al. (2005)
1998-
2001
Greek seas 1 104
as bycatch in swordfish and 
albacore longlines
Öziç & Yılmaz 
(2006)
7/2003-
3/2004
Gökova Bay, 
Aegean Sea
presence of the species
Saad et al. 
(2006)
2001-
2004
Syrian coasts
Ismen et al. 
(2007)
2/2005-
4/2006
Saros Bay, 
Turkey
28-
370
14 68.6 105 920 3388 0.004 2.927 0.174 0.959
Kabasakal & 
Ince (2008)
15/9/2008
Kömür Cape, 
SW tip of Saroz 
Bay, Turkey
F 1 85 1700
refers to a stranded 
individual
Damalas & 
Vassilopoulou 
(2009, 2011)
1993-
2000
Central 
Aegean Sea
41
research on by-catches and 
discards. CPUE, % weight 
and numbers discarded are 
given in the 2011 paper
Ismen et al. 
(2009)
3/2005-
6/2008
Saros Bay, 
Turkey
5-500 18 68.6 105 920 3388 0.005 2.904 0.958
Ismen et al. 
(2009)
3/2005-
6/2008
Saros Bay, 
Turkey
5-500 M 5 68.6 84 920 1960 0.000 3.558 0.986
Ismen et al. 
(2009)
3/2005-
6/2008
Saros Bay, 
Turkey
5-500 F 13 69.2 105 1170 3388 0.008 2.786 0.957
Damalas & 
Megalofonou 
(2012)
1998–
2001
Antikithyra 
strait, Greece
382 1 104
Güven et al. 
(2012)
10/2009-
12/2010
Antalya Bay, 
Turkey
200-
800
11 31.1 105.3 80.2 3560 0.002 3.080 0.998
Papaconstantinou 
(2014)
Aegean Sea
review of Greek 
ichthyofauna
Lteif (2015) 1-8/2013
South 
Lebanese 
coasts
0-300 1 115 6000
Lteif (2015) 1-8/2013
Central 
Lebanese 
coasts
200-
400
3
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
138
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Ergüden & 
Bayhan (2015) 
in Crocetta et 
al. (2015)
27/6/2014
Mersin Bay, 
Turkey
601 M 1 105 3600
includes morphological 
measurements
Başusta (2016) 4/5/2015
NE 
Mediterranean 
(off Iskederun 
Gulf)
360-
400
F 1 32.3 106.9
includes morphological 
measurements
Başusta (2016) 4/5/2015
NE 
Mediterranean 
(off Iskederun 
Gulf)
360-
400
M 1 32.5 101.8
includes morphological 
measurements
Eronat & 
Özaydın 
(2014)
2008-
2009
Izmir Bay 
and Sığacık 
Bay
F 1 99.6 4382
Alkusairy & 
Saad (2018)
11/2014-
10/2016
Syrian coasts M 27 117
the species is overfished in 
the area
Alkusairy & 
Saad (2018)
11/2014-
10/2016
Syrian coasts F 20 124
female catches (common) 
mainly juveniles. The 
species is overfished in the 
area
Follesa et al. 
(2019)
2012-
2015
Aegean Sea 
and Crete, 
Greece
200-
800
data from MEDITS 
expedition (GSAs 22, 23). 
Frequency of occurrence 
is given
Follesa et al. 
(2019)
2012-
2015
Cyprus
200-
800
          
data from MEDITS expedition 
(GSA 25). Frequency of 
occurrence is given
Central Mediterranean
Quignard & 
Capapé (1971) Tunisia
400-
600
from an inventory of species 
in Tunisian waters
Capapé 
(1975b) Tunisia
presence of the species in 
the Tunisian waters
Capapé (1980) Tunisia M 36 0.016 2.502 0.963
the relationship refers to 
juveniles. In the paper it was 
provided in its linear form. 
Capapé (1980) Tunisia M 32 118 0.429 1.178 0.995
the relationship refers to 
adults. In the paper it was 
provided in its linear form
Capapé (1980) Tunisia F 40 0.001 3.447 0.984
the relationship refers to 
juveniles. In the paper it was 
provided in its linear form
Capapé (1980) Tunisia F 12 139 0.347 2.359 0.983
the relationship refers to 
adults. In the paper it was 
provided in its linear form
Bradaï  et al. 
(2002) 4/2/1999
Gulf of 
Gabès, 
Tunisia
M 1 39 138
Bradaï  et al. 
(2002) 19/1/2001
Gulf of 
Gabès, 
Tunisia
80 F 2 69.5 98 688 4000
Bradaï  et al. 
(2002) 1-2/2001
Gulf of 
Gabès, 
Tunisia
M 2 75 1252 1259
Bradaï  et al. 
(2002) 1-2/2001
Gulf of 
Gabès, 
Tunisia
F 3 69.5 98 828 4000
the dimensions of the 3rd 
individual were TL=815, 
W=1450
Bradaï  et al. 
(2002) 1/3/2003
Gulf of 
Gabès, 
Tunisia
F 2 94 100 2300
De Maddalena 
et al. (2002) 26/7/2000
Ganzirri, 
Mesina Strait
70 M 1 85 1610
De Maddalena 
et al. (2002) 6/2000
Linosa 
and Porto 
Empedocle
200 5 70 80
De Maddalena 
et al. (2002) 21/11/1989 Catania 1 95 4000 from the Catania fish market
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
139
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
D'Onghia et al. 
(2003); Politou 
et al. (2003); 
Mytilineou et al. 
(2005)
9/1999, 
4/2000, 
7/2000, 
9/2000
East Ionian 
Sea, Greece
388-
501
first record from the 
Ionian Sea, caught in low 
abundances
Schembri et al. 
(2003)
Malta
review of Matese 
ichthyofauna
Bradaï et al. 
(2006)
Gulf of 
Gabès 
Tunisia
Gristina et al. 
(2006)
autumn 
of 1997-
1998
Strait of 
Sicily
290-
350
mean standardised catch rates 
for the species per sampling 
area and year is provided
Capezzuto 
et al. (2010); 
Maiorano et al. 
(2010)
1985-
2008
NW Ionian 
Sea
322-
345
review of fish caught in 
trawling surveys form 1985 
to 2008
Scacco et al. 
(2010)
2000-
2002
Portopalo di 
Capopassero 
(SE Sicily)
M 8
used for caudal fin shape 
analyses
Scacco et al. 
(2010)
2000-
2002
Portopalo di 
Capopassero 
(SE Sicily)
F 6
used for caudal fin shape 
analyses
Dimech et al. 
(2012)
1/6/2007 Malta
517-
671
biomass (kg/km2) is given
Ragonese et 
al. (2013)
1994-
2009
Central 
Mediterranean
the authors describe the 
species as "rare in Malta 
and Tunisia, common in 
the other zones; sold at the 
market"
El Kamel-
Moutalibi et 
al. (2014)
21/5/2014
Eskerkis 
Bank, 
Tunisia
150-
300
M 1 70 1000
includes morphological 
measurements
El Kamel-
Moutalibi et 
al. (2014)
21/5/2014
Eskerkis 
Bank, 
Tunisia
150-
300
F 1 79 1280
includes morphological 
measurements
El Kamel-
Moutalibi et 
al. (2014)
1/4/2007
Tunisian 
waters
M 1 81 3000
El Kamel-
Moutalibi et 
al. (2014)
15/7/2008
Tunisian 
waters
F 1 110 5000
Papaconstantinou 
(2014)
Ionian Sea
review of Greek 
ichthyofauna
Rafrafi-Nouira 
et al. (2015)
25/9/2014
Cani Rocks, 
Tunisia
56 F 1 99
includes morphological 
measurements
Capapé et al. 
(2018)
24/11/2015
Island of 
Zembra, 
Tunisia
150 M 2 74 84 1300 1735
includes morphological 
measurements
Capapé et al. 
(2018)
24/11/2015
Island of 
Zembra, 
Tunisia
150 F 2 72 112 1150 2255
includes morphological 
measurements
Capapé et al. 
(2018)
2/8/2018
off Bizerte, 
Tunisia
130-
140
M 1 72 1130
includes morphological 
measurements
Capapé et al. 
(2018)
2/8/2018
off Bizerte, 
Tunisia
130-
140
F 1 70 1092
includes morphological 
measurements
Follesa et al. 
(2019)
2012-
2015
Central 
Mediterranean 
(Sicily, South 
Italy, Ionian Sea)
200-
800
          
data from MEDITS 
expedition (GSAs 16, 
19, 20). Frequency of 
occurrence is given
Adriatic Sea
Fredj & 
Maurin (1987)
Adriatic Sea
presence of the species (depths 
from 0 up to >1000 m)
Jukic-Peladic 
et al. (2001); 
Ferretti et al. 
(2013)
1948 Adriatic Sea 2
inventory of species caught 
in scientific surveys
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
140
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Soldo (2006) Adriatic Sea
the author states "often caught 
as bycatch in trawls and by 
deep bottom longlines, but their 
current status in the Adriatic is 
unknown"
Lipej & Dulčić 
(2010)
Adriatic Sea
checklist of the fishes in the 
Adriatic
Follesa et al. 
(2019)
2012-
2015
South 
Adriatic
200-
800
          
data from MEDITS expedition 
(GSA 18). Frequency of 
occurrence is given
West Mediterranean
Tortonese 
(1969)
S. 
Margherita, 
Ligurian Sea
a medium-sized specimen 
from the market of S. 
Margherita (Eastern Riviera)
Fredj & 
Maurin (1987)
W 
Mediterranean
presence of the species (depths 
from 0 up to >1000 m)
Gil de Sola 
Simarro (1994)
1991-
1992
Alboran Sea
200-
500
5 overall weight 21050 g
Lloris et al. 
(1998)
1995
Iberian 
Mediterranean
423-
433
1
data from MEDITS 
expedition
Baino et al. 
(2001)
1994-
1999
Morocco, 
Spain and 
France
data from MEDITS 
expedition (biomass index 
0.7 kg/km2)
Baino et al. 
(2001)
1994-
1999
Tyrrhenian, 
Corsica, 
Sardinia and 
Sicily
data from MEDITS 
expedition (biomass index 
3.9 kg/km2)
Storai (2004) Livorno M 1 98 based on musem material
Storai (2004) Livorno F 2 based on musem material
Serena & 
Relini (2006)
1985-
2004
Northern 
Tuscany
data from GRUND surveys
Serena & 
Relini (2006)
1994-
2004
W 
Mediterranean
data from MEDITS surveys (it 
includes all Mediterranean, 
not specifying the area H. 
perlo was caught)
Dufur et al. 
(2007)
Port Cross 
and Corsica
presence of the species in 
an MPA
Mullas et al. 
(2011)
2008-
2010
Sardenia 600 3
during experimental trawl 
surveys (MEDITS and GRUND)
Bonomo et al. 
(2011)
1906-
1963
Milazzo, 
Sicily
catches in tuna traps
Ordines et al. 
(2011)
3/2003, 
2/2004
Algeria
300-
505
2
only the presence of 2 
individuals is being reported
Mendoza et al. 
(2014)
2006-
2011
SE Spain, W 
Mediterranean
presentation of a 
vulnerability index
Marongiu et 
al. (2017)
1994-
2015
Sardenia
273-
336
1 80.5
data from MEDITS 
expedition
Agnetta et al. 
(2019)
spring 
2005
Gulfs of 
Castellammare, 
Termini Imerese, 
Sant’Agata and 
Patti
24
fished to be used for stable 
isotope analyses. Average 
lengths are given
Guallart et al. 
(2019b)
26/6/2019
Ibiza 
channel, 
Balearic Sea
F 1 64.3
includes morphological 
measurements
Guallart et al. 
(2019a)
24/2/2018 Balearic Sea 650 F 1  79.6  1590      
references that could not be allocated to a specific Mediterranean Area
Vassilopoulou 
et al. (2007)
Greek Seas
presented in a list of totally 
discarded species
Peristeraki & 
Megalofonou 
(2007)
Greek Seas
presence of the species in 
the Greek Seas
Thessalou-
Legaki & 
Legakis (2005)
 Greek Seas            
presence of the species in 
the Greek Seas, with notes 
on the Greek legislation 
related to the species
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
141
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Appendix 2: List of records of Heptranchias perlo in the Mediterranean, based on secondary references (literature 
not available to the authors). F=female, M=male, N=number of individuals.
Priloga 2: Seznam zapisov o pojavljanju vrste Heptranchias perlo v Sredozemskem morju, ki temeljijo na sekundar-
nih referencah (literatura, ki ni bila dostopna avtorjem). F=samica, M=samec, N=število osebkov.
main Reference secondary Reference date locality Sex N
East Mediterranean
Athanasiou & Boulos (1964) Capapé (1980) Lebanon
Demetropoulos & Neocleous 
(1969)
Hadjichristophorou (2006)
Famagusta Bay and Morphou 
Bay, Cyprus
Cihangir et al. (2002) Çoker & Akyol (2014) Bay of Magusa, East Cyprus
Erhard (1858) Papaconstantinou (2014) Aegean Sea
Heldreich (1978) Papaconstantinou (2014) Aegean Sea
Carus (1893) Papaconstantinou (2014) Aegean Sea
Bertrand et al. (2000) Papaconstantinou (2014) Aegean Sea
BOLD Guallart et al. (2019a,b) 2009 off Cyprus 5
Central Mediterranean
BOLD Guallart et al. (2019a,b) 2007 Malta M 2
BOLD Guallart et al. (2019a,b) 2008 NW Ionian Sea M 1
Adriatic Sea
Soljan (1963) Capapé (1980) N Adriatic (former Yugoslavia)
West Mediterranean
Moreau (1881) Capapé (1980), Guallart et al. (2019a) Nice, Sete
LoBianco (1909) Guallart et al. (2019a) off Naples
Gibert (1913) Guallart et al. (2019a,b) Catalonia, Spain
Lozano Rey (1928) Guallart et al. (2019a,b) 1915 Motril, Alboran 1
Dieuzeide et al. (1953) Capapé (1980) Morocco
Capapé (1975) Capapé (1980) Morocco
Capapé (1977) Capapé (1980), Guallart et al. (2019a) Toulon, France
Barrull & Mate (2002) Guallart et al. (2019a,b) 2000 Alboran 1
Hemida (2005) Guallart et al. (2019a,b) Algeria
Bearez et al. (2017) Guallart et al. (2019a,b) Gulf of Lions
Ramirez (2017) Guallart et al. (2019a) 1995, 2002 Alboran (GSA01) 2
BOLD Guallart et al. (2019a) Mazzara de Vallo F 1
BOLD Guallart et al. (2019a) 2009 off Sardinia F 2
references that could not be allocated to a specific Mediterranean Area
Doderlein (1881) Papaconstantinou (2014) Greek seas
Apostolidis (1883, 1907) Papaconstantinou (2014) Greek seas
Hoffman & Jordan (1892) Papaconstantinou (2014) Greek seas
Belloc (1948) Papaconstantinou (2014) Greek seas
Tortonese (1956) Guallart et al. (2019a) Italian waters
Bini (1960; 1965; 1967) Papaconstantinou (2014) Greek seas
Ondrias (1971) Capapé (1980), Papaconstantinou (2014) Greek seas
Boeseman (1973; 1984) Papaconstantinou (2014) Greek seas
Economidis (1973) Capapé (1980), Papaconstantinou (2014) Greek seas
Fisher et al. (1987) Papaconstantinou (2014) Greek seas
diNatale (1998) Guallart et al. (2019a) Italian waters
Machias et al. (2001) Papaconstantinou (2014) Greek seas
Legakis & Maragou (2009) Papaconstantinou (2014) Greek seas
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
142
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
PREGLED O POJAVLJANJU MORSKEGA PSA SEDMEROŠKRGARJA HEPTRANCHIAS
PERLO (CHONDRICHTHYES: HEXANCHIDAE) V SREDOZEMLJU: ZGODOVINSKI IN 
RECENTNI PODATKI
 
Paraskevi K. KARACHLE, Caterina STAMOULI & Aikaterini DOGRAMMATZI
Hellenic Centre for Marine Research, Institute of Marine Biological Resources and Inland Waters, P.O. Box 712, 19013 Anavyssos Attiki, Greece
e-mail: pkarachle@hcmr.gr
POVZETEK
Avtorji poročajo o pojavljanju morskega psa sedmeroškrgarja Heptranchias perlo (Bonnaterre, 1788) v 
Sredozemskem morju na podlagi celovitega pregleda literature, ki je bil osnova za pripravo podatkovne baze o 
razširjenosti te vrste. Poleg tega v delu omenjajo biološke podatke o samici te redke vrste na robu ogroženosti, 
ki je bila ujeta v okviru eksperimentalnega vzorčenja v mirtonskem morju (jugozahodno Egejsko otočje, Grčija). 
Namen tega prispevka je podati povzetek o razpoložljivih podatkih o tej vrsti v Sredozemskem morju, ki bo 
služil za prihodnja dopolnjevanja ocen glede ohranitvenega statusa vrste v bazenu.
Ključne besede: hrustančnice, prehrana, plodnost, sredozemska razširjenost
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
143
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
REFERENCES
Alkusairy, H. & A. Saad (2018): Species Composi-
tion, Diversity and Length Frequency of By-Catch 
Sharks from the Syrian Coast. Int. J. Zool. Res., 4(1), 
11-21. http://doi.org/10.20431/2454-941X.0401003. 
Baino, R., F. Serena, S. Ragonese, J. Rey & P. Rinelli 
(2001): Catch composition and abundance of elasmo-
branchs based on the MEDITS program. Rapp. Comm. 
Int. Mer. Médit. 36, 234.
Barnett, A., J.M. Braccini, C.A. Awruch & D.A. 
Ebert (2012): An overview on the role of Hexanchi-
formes in marine ecosystems: biology, ecology and 
conservation status of a primitive order of modern 
sharks. J. Fish Biol., 80, 966-990. http://10.1111/j.1095-
8649.2012.03242.x 
Bass, A.J., J.D. D’Aubrey & N. Kistnasamy (1975): 
Sharks of the east coast of southern Africa. 5. The 
families Hexanchidae, Clamydoselachidae, Hetero-
dontidae, Pristiophoridae and Squatinidae. Invest. Rep. 
Oceanogr. Res. Inst., Durban, 43, 1-50. 
Başusta, N. (2016): New records of neonate 
and juvenile sharks (Heptranchias perlo, Squatina 
aculeata, Etmopterus spinax) from the North-eastern 
Mediterranean Sea. Mar. Biodiv., 46, 525–527. http://
doi:10.1007/s12526-015-0391-z. 
Ben-Tuvia, A. (1953): Mediterranean fishes of Israel. 
Government Printing Press, Caesarea, Israel, 40 pp.
Bertrand, J.A., L. Gil de Sola, C. Papaconstantinou, 
G. Relini & A. Souplet (2002): The general specifica-
tions of the MEDITS surveys. Sci. Mar., 66(Suppl 2), 
9−17.
Bonomo, F., P. Battaglia, V. Esposito, P. Perzia, F. 
Andaloro & T. Romeo (2011): Trend di catture di specie 
pelagiche nella tonnara del tono di Milazzo dal 1906 
al 1963. Biol. Mar. Mediterr., 18(1), 334-335.
Bornatowski, H., R.R. Braga & J.R.S. Vitule (2013): 
Shark Mislabeling Threatens Biodiversity. Science, 
340(6135), 923. http://10.1126/science.340.6135.923-a.
Braccini, J.M. (2008): Feeding ecology of two 
high-order predators from south-eastern Australia: 
the coastal broadnose and the deepwater sharpnose 
sevengill sharks. Mar. Ecol. Prog. Ser., 371, 273–284.
Bradaï, M.N., B. Saïdi, M. Ghorbel, A. Bouaïn, O. 
Guélorget & C. Capapé (2002): Observations sur les 
requins du golfe de Gabès (Tunisie méridionale, Médi-
terranée central. Mésogée, 60, 61-77.
Bradaï, M.N., B. Saïdi, S. Enajjar & A. Bouaïn 
(2006): The Gulf of Gabès: a spot for the Mediterranean 
elasmobranches. In: Basusta, N., C. Keskin, F. Serena 
& B. Seret (Eds.),  The proceedings of the workshop 
on Mediterranean cartilaginous fish with emphasis on 
southern and eastern Mediterranean).Turkish Marine 
Research Foundation, Istanbul-Turkey, pp. 107–117.
Capapé, C. (1975a): Observations sur le regime 
alimentaire de 29 Selaciens pleurotremes des cotes 
tunisiennes. Arch. Instit. Pasteur Tunis, 52, 395-414.
Capapé, C. (1975b): Sélaciens nouveaux et rares le 
long des côtes tunisiennes. Premières observations bi-
ologiques. Arch. Instit Pasteur Tunis., 52(1–2), 107–128.
Capapé, C. (1980): Nouvelle description de Hep-
tranchias perlo (Bonnaterre, 1788) (Pisces, Pleurotramata, 
Hexanchidae). Données sur la biologie de la reproduc-
tion et le régime alimentaire des spécimens des côtes tu-
nisiennes. Bull. Office nat. Pêches Tunisie, 4(2), 231-264.
Capapé, C., S. Rafrafi-Nouira, K. Ounifi-Ben Amor 
& M.M. Ben Amor (2018): Unusual and substantiated 
records of Heptranchias perlo (Chondrichthyes: Hex-
anchidae) in the N-E Tunisian waters (Central Mediter-
ranean Sea). Thalassia Sal., 40, 17-24. http://10.1285/
i15910725v40p17.
Capezzuto, F., R. Carlucci, P. Maiorano, L. Sion, 
D. Battista, A. Giove, A. Indennidate, A. Tursi & G. 
D’Onghia (2010): The bathyal benthopelagic fauna in 
the north-western Ionian Sea: structure, patterns and 
interactions, Chem. Ecol., 26, 199-217. http://10.1080
/02757541003639188. 
Çoker, T. & O. Akyol (2014): An overview on the fish 
diversity in the coasts of Turkish Republic of Northern 
Cyprus (Mediterranean). Ege J. Fish. Aqua. Sci., 31(2), 
113-118. http://doi: 10.12714/egejfas.2014.31.02.08.
Compagno, L.J.V. (1984): FAO Species Catalogue. 
Vol. 4. Sharks of the world. An annotated and illus-
trated catalogue of shark species known to date. Part 
1 - Hexanchiformes to Lamniformes. FAO Fish. Synop., 
125(4/1), Rome, FAO. 249 pp.
Correa de Carvalho, A.P, B. de Sousa Rangel & A. 
Ferreira de Amorim (2020): Insights into the repro-
ductive biology of Sharpnose sevengill shark (Hep-
tranchias perlo) in the Western South Atlantic. In: da 
Silva, E. (Ed.), Tópicos Multidisciplinares em Ciências 
Biológicas 2, Capítulo 6. Atena Editora, Ponta Grossa, 
Paraná, Brasil, pp. 45-60. https://doi.org/10.22533/
at.ed.2892027076
Cortés, E. (1999): Standardized diet compositions 
and trophic levels of sharks. ICES J. Mar. Sci., 56, 707-
717.
Crocetta, F., D. Agius, P. Balistreri, M. Bariche, Y. 
Bayhan, M. Çakir, S. Ciriaco, M. Corsini-Foka, A. Dei-
dun, R. El Zrelli, D. Ergüden, J. Evans, M. Ghelia, M. 
Giavasi, P. Kleitou, G. Kondylatos, L. Lipej, C. Mifsud, 
Y. Özvarol, A. Pagano, P. Portelli, D. Poursanidis, L. 
Rabaoui, P. Schembri, E. Taşkin, F. Tiralongo & A. Ze-
netos (2015): New Mediterranean Biodiversity Records 
(October 2015). Medit. Mar. Sci., 16(3), 682-702. 
doi:http://dx.doi.org/10.12681/mms.1477.
D’Onghia, G., F. Mastrototaro, A. Matarrese, C.Y. 
Politou & Ch. Mytilineou (2003): Biodiversity from 
the upper slope demersal community of the eastern 
Mediterranean: preliminary comparison between two 
areas with and without trawl fishing. J. Northw. Atl. 
Fish. Sci., 31, 263-273. http:// 10.2960/J.v31.a20.
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
144
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Damalas, D. & V. Vassilopoulou (2009): Chondrich-
thyan by-catches and discards in the bottom trawl fish-
ery of the Central Aegean Sea (Eastern Mediterranean). 
Proceedings of 9th Symposium on Oceanography & 
Fisheries, 2, 935-940.
Damalas, D. & V. Vassilopoulou (2011): Chondrich-
thyan by-catch and discards in the demersal trawl fish-
ery of the central Aegean Sea (Eastern Mediterranean). 
Fish. Res., 108, 142–152.
De Maddalena, A., A. Celona, M. Zuffa & A. Va-
nadia (2002): Su alcune catture di notidano cinereo, 
Heptranchias perlo (Bonnaterre, 1788), nelle acque 
della Sicilia (On the capture of some sharpnose sev-
engill sharks, Heptranchias perlo (Bonnaterre, 1788), 
in Sicilian waters). Thalassia Sal., 26, 39–44. http:// 
10.1285/i15910725v26p39.
Dimarchopoulou, D., K.I. Stergiou & A.C. Tsikliras 
(2017): Gap analysis on the biology of Mediterranean 
marine fishes. PLoS ONE, 12, e0175949. https://doi.
org/10.1371/journal.pone.0175949. 
Dimech, M., M.J. Kaiser, S. Ragonese & P. Schembri 
(2012): Ecosystem effects of fishing on the continental 
slope in the Central Mediterranean. Mar. Ecol. Prog. 
Ser., 449, 41–54. http://doi:10.3354/meps09475.
Ebert, D.A. (1990): The taxonomy, biogeography 
and biology of cow and frilled sharks (Chondrichthyes: 
Hexanchiformes). PhD thesis, Rhodes University, Gra-
hamstown, 308 pp. 
El Kamel-Moutalibi, O., N. Mnasri-Sioudi, S. Raf-
rafi-Nouira, M. Boumaïza, C. Reynaud & C. Capapé 
(2014): Additional records of a rare elasmobranch 
species, sharpnose seven-gill shark Heptranchias 
perlo (Hexanchidae) off the northern Tunisian coast 
(central Mediterranean). Ann. Ser. Hist. Nat., 24(2), 
99-106.
Eronat, E.G.T. & O. Özaydın (2014): Length-weight 
relationship of cartilaginous fish species from Central 
Aegean Sea (Izmir Bay and Sığacık Bay). Ege J. Fish. 
Aqua. Sci., 31(3), 119-125.http://doi: 10.12714/egej-
fas.2014.31.3.01.
Ferretti, F., G.C. Osio, C.J. Jenkins, A.A. Rosenberg 
& H.K. Lotze (2013): Long-term change in a meso-
predator community in response to prolonged and het-
erogeneous human impact. Sci. Rep., 3, 1057. http:// 
10.1038/srep01057.
Filiz, H. & S. Mater (2002): A preliminary study on 
length-weight relationships for seven elasmobranch 
species from north Aegean Sea, Turkey. J. Fish. Aquat. 
Sci., 19(3-4), 401-409.
Follesa, M.C., M.F. Marongiu, W. Zupa, A. Bellodi, 
A. Cau, R. Cannas, F. Colloca, M. Djurovic, I. Isajlovic, 
A. Jadaud, C. Manfredi, A. Mulas, P. Peristeraki, C. 
Porcu, S. Ramirez-Amaro, F. Salmerón Jiménez, F. 
Serena, L. Sion, I. Thasitis, A. Cau & P. Carbonara 
(2019): Spatial variability of Chondrichthyes in the 
northern Mediterranean. Sci. Mar., 83S1, 81-100. 
http:// 10.3989/scimar.04998.23A.
Fredj, G. & C. Maurin (1987): Les poissons dans 
la banque de données MEDIFAUNE. Application à 
l’étude des caractéristiques de la faune ichtyologique 
méditerranéenne. Cybium, 11(3), 31-139.
Frentzel-Beyme, B.Z. & F.W. Koster (2002): On the 
biology of the sharpnose sevengill shark, Heptranchias 
perlo, from the Great Meteor Seamount (central eastern 
Atlantic), in Vacchi, M., La Mesa, G., Serena, F., Seret, 
B. (Eds.), Proceedings of the 4th European Elasmobranch 
Association Meeting, Societe Francaise d’Ichtyologie, 
Livorno, pp. 77-96.
Froese, R. & D. Pauly (Eds) (2019): FishBase. World 
Wide Web electronic publication. www.fishbase.org, 
version (12/2019). (Accessed on June 22nd 2020).
Gil de Sola Simarro, L. (1994): Ictiofauna demer-
sal de la plataforma continental del mar de Alborán 
(Mediterráneo suroccidental ibérico). Bol. Inst. Esp. 
Oceanogr., 10(1), 63-79.
Golani, D. (2006): Cartilaginous fishes of the Medi-
terranean Coast of Israel. In: Basusta, N., C. Keskin, 
F. Serena & B. Seret (Eds.),  The proceedings of the 
workshop on Mediterranean cartilaginous fish with 
emphasis on southern and eastern Mediterranean).
Turkish Marine Research Foundation, Istanbul-Turkey, 
pp. 95–100.
Gristina, M., T. Bahri, F. Fiorentino & G. Garofalo 
(2006): Comparison of demersal fish assemblages in 
three areas of the Strait of Sicily under different trawl-
ing pressure. Fish. Res., 81, 60–71. http:// 10.1016/j.
fishres.2006.05.010.
Guallart, J., G. Morey & À. Bartolí (2019a): New 
record of a sharpnose sevengill shark Heptranchias 
perlo (Elasmobranchii, Hexanchidae) from the Balearic 
Sea, western Mediterranean Sea. J. Fish Biol., 94, 526-
531. https://doi.org/10.1111/jfb.13905. 
Guallart, J., G. Morey & À. Bartolí (2019b): Another 
recent record of the uncommon sharpnose sevengill 
shark Heptranchias perlo in the Balearic Sea (Western 
Mediterranean). Proceedings of the 23rd Annual confer-
ence of the European Elasmobranch Association, 16-18 
October 2019, Rende, Italy, 123 pp.
Güven, O., T. Kebapçıoğlu & M.C. Deval (2012): 
Length-weight relationships of sharks in Antalya Bay, 
eastern Mediterranean. J. Appl. Ichthyol., 28, 278-279. 
http://10.1111/j.1439-0426.2011.01823.x
Hadjichristophorou, M. (2006): Chondrichthyes in 
Cyprus. In: Basusta, N., C. Keskin, F. Serena & B. Seret 
(Eds.),  The proceedings of the workshop on Mediter-
ranean cartilaginous fish with emphasis on southern 
and eastern Mediterranean).Turkish Marine Research 
Foundation, Istanbul-Turkey, pp. 162-168.
Hamlett, W.C., G. Kormanik, M. Storrie, B. Stevens 
& T.I. Walker (2005): Chondrichthyan parity, lecitho-
trophy and matrotrophy. In: Hamlett WC (ed). Repro-
ductive biology and phylogeny of chondrichthyes: 
sharks, rays and chimaeras. Science Publishers, INC., 
Enfield, pp. 395-434
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
145
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Henderson, A.C. & R.S. Williams (2001): A new 
record of the sharpnose seven-gill shark Heptranchias 
perlo, from the north-east Atlantic. J. Mar. Biol. As-
soc. U.K., 81 (4), 707–708. https://doi.org/10.1017/
S0025315401004453 
ICES (2013): Report of the workshop on Sexual 
Maturity Staging of Elasmobranchs (WKMSEL), 11–14 
December 2012, Lisbon, Portugal. ICES CM 2012/
ACOM: 59, 66 pp. http://www.ices.dk/sites/pub/
Publication%20Reports/Expert%20Group%20Report/
acom/2012/WKMSEL/WKMSEL%202012.pdf 
Ismen, A., O. Ozen, U. Altınağaç, U. Ozekinci & A. 
Ayaz (2007): Weight–length relationships of 63 fish spe-
cies in Saros Bay, Turkey. J. Appl. Ichthyol., 23, 707-708. 
https://doi.org/10.1111/j.1439-0426.2007.00872.x. 
Ismen, A., C.C. Yığın, U. Altınağaç & A. Ayaz 
(2009): Length-weight relationships for ten shark 
species from Saros Bay (North Aegean Sea). J. Appl. 
Ichthyol., 25, 109-112. https://doi.org/10.1111/j.1439-
0426.2009.01263.x. 
Jacquet, J.L. & D. Pauly (2003): Trade secrets: Renam-
ing and mislabeling of seafood. Mar. Policy, 32, 309-
318. https://doi.org/10.1016/j.marpol.2007.06.007. 
Jukic-Peladic, S., N. Vrgoc, S. Šifner, C. Piccinetti, 
G. Piccinetti-Manfrin, G. Marano & N. Ungaro (2001): 
Long-term changes in demersal resources of the Adri-
atic Sea: Comparison between trawl surveys carried 
out in 1948 and 1998. Fish. Res., 53, 95-104. https://
doi.org/10.1016/S0165-7836(00)00232-0.
Kabasakal, H. & P. Ince (2008): Note on a sharp-
nose sevengill shark, Heptranchias perlo (Bonnaterre, 1788) 
(Chondrichthyes: Hexanchidae), stranded in Saroz Bay (NE 
Aegean Sea, Turkey). Annales Ser. Hist. Nat., 18(2), 173-176.
Karachle, P.K. & K.I. Stergiou (2017): An update on 
the feeding habits of fish in the Mediterranean. Medit. 
Mar. Sci., 18(1), 43-52. http://dx.doi.org/10.12681/
mms.1968. 
Labropoulou, M. & C. Papaconstantinou (2000): Com-
munity structure of deep-sea demersal fish in the North 
Aegean Sea (Northeastern Mediterranean). Hydrobio-
logia, 440, 281-296. http:// 10.1023/A:1004199917299.
Lipej, L. & J. Dulčić (2010): Checklist of the Adri-
atic Sea fishes. Zootaxa, 2589, 1-92. http:// 10.11646/
zootaxa.2589.1.1.
Lloris, D., L. Gil de Sola & J. Rucabado (1998): 
Ichthyofauna caught during the MEDITS–ES cruise 
(1994–1997) in the Iberian western Mediterranean. 
In: Plouzané, France (Eds.) Demersal resources in the 
Mediterranean. Ifremer, 26, 221–228.
Lteif, M. (2015): Biology, distribution and diversity 
of cartilaginous fish species along the Lebanese coast, 
eastern Mediterranean. Ecology, environment. PhD 
Thesis, Université de Perpignan, 262 pp.
Macpherson, E. (1989): Influence of geographical 
distribution, body size and diet on population density 
of benthic fishes off Namibia (South West Africa). Mar. 
Ecol. Prog. Ser., 50, 295-299.
Maiorano, P., L. Sion, R. Carlucci, F. Capezzuto, 
A. Giove, G. Costantino, M. Panza, G. D’Onghia & A. 
Tursi (2010): The demersal faunal assemblage of the 
north-western Ionian Sea (central Mediterranean): cur-
rent knowledge and perspectives, J. Chem. Ecol., 26, 
219-240. http:// 10.1080/02757541003693987.
Marongiu, M.F., C. Porcu, A. Bellodi, R. Cannas, A. 
Cau, D. Cuccu, A. Mulas & M.C. Follesa (2017): Tempo-
ral dynamics of demersal chondrichthyan species in the 
central western Mediterranean Sea: The case study in 
Sardinia Island. Fish. Res., 193, 81-94. http:// 10.1016/j.
fishres.2017.04.001.
Megalofonou, P., D. Damalas & C. Yannopoulos 
(2005): Composition and abundance of pelagic shark 
by-catch in the eastern Mediterranean Sea. Cybium, 
29(2), 135–140.
Mendoza, M., D. Garrido & J.M. Bellido (2014): Fac-
tors affecting the fishing impact on cartilaginous fishes 
in southeastern Spain (western Mediterranean Sea). Sci. 
Mar., 78, 67-76. http:// 10.3989/scimar.04025.21A.
Mulas, A., A. Bellodi, A.I. Cau, A. Gastoni, I. Locci 
& M.C. Follesa (2011): Trophic interactions among 
chondrichthyans in the Central-Western Mediterranean 
Sea. Biol. Mar. Mediterr., 18 (1), 81-84. http:// 10.3354/
meps11188.
Mytilineou, Ch., C.Y. Politou, C. Papaconstantinou, 
S. Kavadas, G. D’Onghia & L. Sion (2005): Deep-water 
fish fauna in the Eastern Ionian Sea. Belg. J. Zool., 135(2), 
229-233.
Ordines, F., E. Massutí, J. Moranta, A. Quetglas, B. 
Guijarro & K. Fliti (2011): Balearic Islands vs Algeria: 
two nearby western Mediterranean elasmobranch 
assemblages with different oceanographic scenarios 
and fishing histories. Sci. Mar., 75(4), 707-717. http:// 
10.3989/scimar.2011.75n4707.
Öziç, F. & F. Yılmaz (2006): An investigation of 
demersal fishes of Gökova Bay in Aegean Sea. Ekoloji, 
15(58), 16-20.
Papaconstantinou, C. (2014): Fauna Graeciae. An 
updated checklist of the fishes in the Hellenic Seas, 
Monographs on Marine Sciences 7, Institute of Marine 
Biological Resources Publisher,. HCMR, Athens, Greece, 
340 pp.
Paul, L. & S. Fowler, (SSG Australia & Oceania 
Regional Workshop, March 2003) (2003): Heptranchias 
perlo. The IUCN Red List of Threatened Species 2003: 
e.T41823A10572878. https://dx.doi.org/10.2305/IUCN.
UK.2003.RLTS.T41823A10572878.en (Downloaded on 
22 June 2020).
Pauly, D., R. Froese, P. Sa-a, M.L. Palomares, V. 
Christensen & J. Rius (2000): Trophlab manual. Manila: 
ICLARM, 4 pp.
Politou, C.Y., S. Kavadas, C. Mytilineou, A. Tursi, 
R. Carluci & G. Lembo (2003): Fisheries resources in 
the deep waters of the Eastern Mediterranean (Greek 
Ionian Sea) J. Northw. Atl. Fish. Sci., 31, 35-46. http:// 
10.2960/J.v31.a3.
ANNALES · Ser. hist. nat. · 30 · 2020 · 2
146
Paraskevi K. KARACHLE et al.: REVIEW OF THE SHARPNOSE SEVENGILL SHARK HEPTRANCHIAS PERLO (CHONDRICHTHYES: HEXANCHIDAE) IN THE ..., 131–146
Rafrafi-Nouira, S., O. El Kamel-Moutalibi, C. 
Reynaud, M. Boumaïza & C. Capapé (2015): Addi-
tional and unusual captures of elasmobranch species 
from the northern coast of Tunisia (central Mediter-
ranean). J. Ichthyol., 55(6), 836-848. http://10.1134/
S0032945215060181.
Ragonese, S., S. Vitali, M. Dimech & S. Maz-
zola (2013): Abundances of demersal sharks and 
chimaera from 1994-2009 scientific surveys in the 
central Mediterranean Sea. PLoS One, 8(9), e74865. 
https://doi. org/10.1371/journal.pone.0074865.
Saad, A., M. Ali & B. Seret (2006): Shark exploi-
tation and conservation in Syria. In: Basusta, N., C. 
Keskin, F. Serena & B. Seret (Eds.), The proceedings of 
the workshop on Mediterranean cartilaginous fish with 
emphasis on southern and eastern Mediterranean). 
Turkish Marine Research Foundation, Istanbul-Turkey, 
pp. 135–148.
Scacco, U., G. La Mesa & M. Vacchi (2010): Body 
morphometrics, swimming diversity and niche in 
demersal sharks: a comparative case study from the 
Mediterranean Sea. Sci. Mar., 74(1), 37-53. http:// 
10.3989/scimar.2010.74n1037.
Serena, F. & G. Relini (2006): Use of scientific 
campaigns (trawl surveys) for the knowledge of the 
sensitive habitats. A review of the MEDITS, GRUND 
and APHIA data with special attention to the Ital-
ian seas. In: Basusta, N., C. Keskin, F. Serena & B. 
Seret (Eds.), The proceedings of the workshop on 
Mediterranean cartilaginous fish with emphasis 
on southern and eastern Mediterranean). Turkish 
Marine Research Foundation, Istanbul-Turkey, pp. 
135–148.
Sierra, L.M., R. Claro & O.A. Popova (1994): Alimen-
tacion y relaciones tróficas. In: Rodolfo, C. (Ed.), Ecología 
de los Peces Marinos de Cuba. Instituto de Oceanología 
Academia de Ciencias de Cuba and Centro de Investiga-
ciones de Quintana Roo, Mexico, pp. 263-284.
Soldo, A. (2006): Status of the sharks in the Adri-
atic. In: Basusta, N., C. Keskin, F. Serena & B. Seret 
(Eds.): The proceedings of the workshop on Mediter-
ranean cartilaginous fish with emphasis on southern 
and eastern Mediterranean). Turkish Marine Research 
Foundation, Istanbul-Turkey, pp. 128–134.
Soldo, A. & M. Bariche (2016): Heptranchias 
perlo. The IUCN Red List of Threatened Species 2016: 
e.T41823A16527717. https://www.iucnredlist.org/spe-
cies/41823/16527717 (Downloaded on 22 June 2020)
Spedicato M.T., E. Massuti, B. Merigot, G. Tserpes, 
A. Jadaud & G. Relini (2019): The MEDITS trawl survey 
specifications in an ecosystem approach to fishery 
management. Sci. Mar., 83(S1), 9-20. https://doi.
org/10.3989/scimar.04915.11X.
Stergiou, K.I. & V.S. Karpouzi (2002): Feeding hab-
its and trophic levels of Mediterranean fish. Rev. Fish 
Biol. Fish., 11, 217-254.
Tortonese, E. (1969): The Squaliforms of the Ligurian 
Sea: a revised list with notes. Isr. J. Ecol. Evol., 18, 233-
236. https://doi.org/10.1080/00212210.1969.10688286.
Vassilopoulou, V., A. Machias & K. Tsagarakis (2007): 
By-catch and discards in multi-species fisheries and their 
impact in the Hellenic waters. In: Papaconstantinou, C., 
A. Zenetos, V. Vassilopoulou & G. Tserpes (Eds), State of 
Hellenic Fisheries. Hellenic Centre for Marine Research, 
Institute of Marine Biological Resources Publisher, Athens, 
Greece, pp. 251-260. http://www.fao.org/fishery/statistics/
global-capture-production/en; accessed June 22nd 2020.