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Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea

Authors:
  • Institute for Biological Resources and Marine Biotechnology (IRBIM), National Research Council – CNR, Via Vaccara, 61 – 91026, Mazara del Vallo (Tp), Italy

Abstract

This volume presents a fully illustrated field guide for the identification of the sharks and rays most relevant to the fisheries of the Mediterranean. An extensive literature review was carried out for the preparation of this document. A total of 49 sharks, 34 batoids and 1 chimaera are fully treated. The presence of 5 sharks and 2 batoid included in this field guide, need, however, to be confirmed. The guide includes sections on technical terms and measurements for sharks and batoids, and fully illustrated keys to those orders and families that occur in the region. Each species account includes: at least one annotated illustration of the species highlighting its relevant identification characters; basic information on nomenclature, synonyms and possible misidentifications; FAO, common names; basic information on size, habitat and biology, importance to fisheries, distribution and conservation status. Colour plates for a large number of the species are included as well as two plates showing the egg cases.
FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES
ISSN 1020-6868
FIELD IDENTIFICATION GUIDE TO THE SHARKS AND
RAYS OF THE MEDITERRANEAN AND BLACK SEA
FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES
FIELD IDENTIFICATION GUIDE TO THE SHARKS AND RAYS
OF THE MEDITERRANEAN AND BLACK SEA
by
Fabrizio Serena
Regional Agency for the Protection
of the Environmental in Tuscany
Livorno, Italy
FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
Rome, 2005
PREPARATION OF THIS DOCUMENT
This document was prepared under the coordination of the Species Identification and Data Progrmme of
the Marine Resources Service, Fishery Resources and Environment Division, Fisheries Department,
Food and Agriculture Organization of the United Nations (FAO).
At a meeting of the Sub-Committee on Marine Environment and Ecosystems of the General Fishery
commission for the Mediterranean (GFCM) in February 2001 in Palma de Majorca, some member
countries concerned with the protection of sharks and other cartilaginous species suggested that a field
guide for the identification of cartilaginous species be prepared. At its fourth session in June 2001 in
Athens, the Scientific Advisory Committee (SAC) endorsed the suggestion and asked the COPEMED
Project GCP/REM/O57/SPA to support the preparation of the field guide.
This increasing recognition of the significance of sharks and batoid fishes as ecosystem health
indicators, as well as their particular importance in exploited ecosystems in the Mediterranean, have
been key considerations to promote the preparation of this field guide.
Project managers: M. Lamboeuf, J. Lleonart (FAO, Rome).
Scientific reviser: N. De Angelis (FAO, Rome).
Editorial assistance, page composition and indexing: M. Kautenberger-Longo (FAO, Rome).
Scientific illustrator (for material presented here for the first time):E. D’Antoni (FAO, Rome).
Cover: E. D’Antoni (FAO, Rome).
iii
Serena, F.
Field identification guide to the sharks and rays of the Mediterranean and Black Sea.
FAO Species Identification Guide for Fishery Purposes.
Rome, FAO. 2005. 97p. 11 colour plates + egg cases.
Abstract
This volume presents a fully illustrated field guide for the identification of the sharks and rays
most relevant to the fisheries of the Mediterranean. An extensive literature review was carried
out for the preparation of this document. A total of 49 sharks, 34 batoids and 1 chimaera are fully
treated. The presence of 5 sharks and 2 batoid included in this field guide, need, however, to be
confirmed. The guide includes sections on technical terms and measurements for sharks and
batoids, and fully illustrated keys to those orders and families that occur in the region. Each
species account includes: at least one annotated illustration of the species highlighting its
relevant identification characters; basic information on nomenclature, synonyms and possible
misidentifications; FAO, common names; basic information on size, habitat and biology,
importance to fisheries, distribution and conservation status. Colour plates for a large number of
the species are included as well as two plates showing the egg cases.
ACKNOWLEDGEMENTS
I would especially like to thank Amor El Abed (Institut National des Sciences et Technologies de la Mer,
Salammbô, Tunisie) for his support regarding my efforts to produce this guide and the COPEMED project
which sponsored the publication. Very large thanks to Giulio Relini (University of Genoa, Italy) and my
research colleague Marino Vacchi (ICRAM Rome, Italy) for their encouragement to produce this guide
and for their precious suggestions, useful information and critical reading of the draft. I am grateful to
Leonard J.V. Compagno (Shark Research Center, Iziko Museum of Cape Town, South Africa) for our
discussion in Paris in 2002 and further in Tenerife in 2004 aimed at the definition of the systematic
catalogue structure and to solve taxonomic problems as well as for his critical reading of the systematic
arrangement of the species. Many thanks to Bernard Séret (Museum National d’Histoire Naturelle, Paris
France) for his critical reading of the previous draft, continuous exchange of ideas and finally for his
courtesy. My most sincere thanks go to Pere Oliver (IEO, Barcelona), who has believed in the usefulness
of this field guide since the SAC-SCMEE meeting (Palma de Mallorca, 2001). I appreciate his advice and
encouragement during the preparation of this publication.
Thanks also to: John D. McEachran (Texas A and M University, USA), Kent E. Carpenter (Old Dominion
University - Department of Biological Sciences, Norfolk, Virginia, USA) and Peter Last (CSIRO, Division
of Fisheries Research, Hobart, Tasmania, Australia) for suggestions provided during the Batoids-FAO
meeting in Paris in March 2002; Gian Pietro Gasparini and Mario Astraldi (Istituto di Oceanografia
Fisica-CNR, Lerici, Italy) for advice on the dynamics of the water masses in the Mediterranean Sea. A
very special thanks goes to Claude Millot for his knowledgeable help in updating the information on the
current pattern in the Mediterranean basin and also for letting me use his latest papers, still in press. To
my colleagues Alvaro Juan Abella, Romano Baino, Monica Barone, Enrico Cecchi and Alessandro
Voliani (ARPAT, Livorno, Italy) for their support and suggestions and also to Marco Costantini (WWF,
Italy); to Adib Ali Saad (Marine Sciences Laboratory, Faculty of Agriculture, Tishreen University, Latakia,
Syria) for his collaboration; Giulia Mò (ICRAM Rome, Italy) for helping me with aspects related to the
Conventions for the protection of marine organisms and environment; to Cecilia Mancusi (ARPAT,
Livorno, Italy) for help in the preparation of the single species sheet and in the chapter related to
Conservation and Ian K. Ferguson (The Shark Trust, London, UK). This last chapter has been produced
with the important help of Sarah Fowler, Rachel Cavanagh and Imène Meliane (IUCN Shark Specialist
Group c/o Naturebureau International, UK and Centre de Cooperation pour la Méditerranée) and I also
thank them very much. Sincere thanks to Francesca Ferretti and Ransom Meyers of the Dalhousie
University (Halifax, Nova Scotia, Canada) for their innovative suggestion on the conservation status
analysis of cartilaginous fishes. Thanks also to Ramón Bonfil (International Conservation
Programs-Wildlife Conservation Society, NY, USA), Giuseppe Notarbartolo di Sciara (Tethys Research
Institute, Italy), Daniel Golani (Hebrew University of Jerusalem, Israel) for their important suggestions;
Temel Oguz (Middle East Technical University, Institute of Marine Sciences Erdemli 33731, Icel, Turkey)
for his collaboration; Franco Biagi (European Commission, Fisheries Directorate-General, Brussels,
Belgium) for his support. I acknowledge the helpful comments on an earlier version of the manuscript of
John Stevens (CSIRO Marine Research, Hobart, Tasmania, Australia) and Peter Last given during the
Pre-Conference of Deep Sea Chondrichthyan Fishes in Dunedin, New Zealand in 2003. Many thanks to
Piero Mannini and Caroline Bennett (ADRIAMED-FAO, Rome, Italy) respectively for their collaboration in
particular for providing many useful bibliographical references and helping with manuscript editing.
Finally, thanks also to all colleagues who kindly allowed me to use their pictures: R. Bonfil (ICP-WCS NY,
USA), P.H.F. Bor (Netherlands), M.N. Bradai (INSTM Sfax, Tunisia), G.H. Burgess (MNH, Gainesville, FL,
USA), R. Carlucci (Univ. Bari, Italy), P. Consoli (IST-CNR, Messina, Italy), M. Costantini (WWF, Trieste, Italy),
M. Dalu (ICRAM, Roma, Italy), M. Dicken (Port Elizabeth Museum, Port Elizabeth, South Africa), M. Ducrocq
(IUCN, Mauritania), F. Cigala Fulgosi (Univ. Parma, Italy), F. Hemida (USTHB/ISN, Algiers, Algeria),
S.P. Iglesias and E. Luchetti (MNHN, Concarneau, France), H. Ishihara (Tokyo University of Fisheries,
Japan), R. McAuley (WA Marine Research Laboratory, Department of Fisheries, North Beach, WA, USA),
P. Micarelli (Acquario Mediterraneo dell’Argentario, Porto Santo Stefano, Italy), V. Moore (University of
Dorset, UK), G. Morey, (CSIC/UIB, Mallorca, Spain), P. Psomadakis (ICRAM, Rome, Italy), J. Rey (IEO,
Malaga, Spain); RV DR. FRIDTJOF NANSEN (Research Vessel Staff), B. Séret (MNHN, Paris, France),
R. Silvestri (ARPAT, Livorno, Italy), C. Simpfendorfer (Mote Marine Laboratory, Sarasota, FL, USA), L. Sion
(Univ. Bari, Italy), J. Stafford-Deitsch (The Shark Trust, UK), V. Taylor (USA), N. Ungaro (LBM, Bari, Italy),
M. van Tienhoven (CSIRO, Pretoria, Australia).
iv
TABLE OF CONTENTS
PREPARATION OF THIS DOCUMENT ................................ iii
ACKNOWLEDGEMENTS....................................... iv
INTRODUCTION ............................................ 1
PhysicalandChemicalCharacteristicsoftheRegion........................ 1
GeologyandPhysicalFeatures................................. 1
Oceanographic Features .................................... 3
Biodiversity and Biogeographical Characteristics of the Region ...................3
TheFishery............................................. 8
Fishery Management ....................................... 10
Conservation............................................ 10
Codes for Conservation and Exploitation Status ........................ 11
CLASSIFICATION AND SYSTEMATIC ARRANGEMENT ...................... 13
SHARKS ............................................... 16
Picturekeyofshark-likefishes................................... 15
TechnicalTermsandMeasurements............................... 16
List of Orders, Families and Species Occurring in the Area ....................18
Guide to the Orders and Families of Sharks Occurring in the Area .................19
Order HEXANCHIFORMES – Cow and frilled sharks ..................... 18
HEXANCHIDAE ...................................... 18
OrderSQUALIFORMESDogfishsharks........................... 18
ECHINORHINIDAE .................................... 18
SQUALIDAE........................................ 18
CENTROPHORIDAE.................................... 19
ETMOPTERIDAE ..................................... 19
SOMNIOSIDAE ...................................... 19
OXYNOTIDAE....................................... 19
DALATIIDAE........................................ 21
Order SQUATINIFORMES – Angel sharks ........................... 21
SQUATINIDAE....................................... 21
OrderLAMNIFORMESMackerelsharks........................... 21
ODONTASPIDIDAE .................................... 21
ALOPIIDAE......................................... 22
CETORHINIDAE...................................... 22
LAMNIDAE......................................... 22
v
Order CARCHARHINIFORMES – Ground sharks ....................... 22
SCYLIORHINIDAE..................................... 23
TRIAKIDAE......................................... 23
CARCHARHINIDAE .................................... 23
SPHYRNIDAE....................................... 23
Guide to Families and Species
HEXANCHIDAE ........................................ 24
Heptranchias perlo (Bonnaterre, 1788) .......................... 24
Hexanchus griseus (Bonnaterre, 1788) .......................... 24
Hexanchus nakamurai Teng, 1962............................. 25
ECHINORHINIDAE ...................................... 25
Echinorhinus brucus (Bonnaterre, 1788) ......................... 25
SQUALIDAE.......................................... 26
Squalus acanthias Linnaeus, 1758 ............................. 26
Squalus blainvillei (Risso, 1826).............................. 26
Squalus cf. megalops (Macleay, 1881) ........................... 27
CENTROPHORIDAE ..................................... 27
Centrophorus granulosus (Bloch and Schneider, 1801) .................. 27
Centrophorus uyato (Rafinesque, 1810) .......................... 28
ETMOPTERIDAE....................................... 28
Etmopterus spinax (Linnaeus, 1758) ............................ 28
SOMNIOSIDAE ........................................ 29
Centroscymnus coelolepis Bocage and Capello, 1864 ................... 29
Somniosus (Rhinoscymnus)rostratus (Risso, 1810) ....................29
OXYNOTIDAE......................................... 30
Oxynotus centrina (Linnaeus, 1758) ............................ 30
DALATIIDAE ......................................... 30
Dalatias licha (Bonnaterre, 1788) ............................. 30
SQUATINIDAE......................................... 31
Squatina aculeata Dumeril, in Cuvier, 1817......................... 31
Squatina oculata Bonaparte, 1840 ............................. 31
Squatina squatina (Linnaeus, 1758) ............................ 32
ODONTASPIDIDAE ...................................... 32
Carcharias taurus Rafinesque, 1810 ........................... 32
Odontaspis ferox (Risso, 1810) .............................. 33
vi Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
ALOPIIDAE .......................................... 33
Alopias superciliosus (Lowe, 1839) ............................ 33
Alopias vulpinus (Bonnaterre, 1788)............................ 34
CETORHINIDAE........................................ 34
Cetorhinus maximus (Gunnerus, 1765) .......................... 34
LAMNIDAE........................................... 35
Carcharodon carcharias (Linnaeus, 1758) ........................ 35
Isurus oxyrinchus Rafinesque, 1810 ............................ 35
Isurus paucus Guitart Manday, 1966 ............................ 36
Lamna nasus (Bonnaterre, 1788) ............................. 36
SCYLIORHINIDAE....................................... 37
Galeus atlanticus (Vaillant, 1888) ............................. 37
Galeus melastomus Rafinesque, 1810 ........................... 37
Scyliorhinus canicula (Linnaeus, 1758) .......................... 38
Scyliorhinus stellaris (Linnaeus, 1758) .......................... 38
TRIAKIDAE .......................................... 39
Galeorhinus galeus (Linnaeus, 1758) ........................... 39
Mustelus asterias Cloquet, 1821 .............................. 39
Mustelus mustelus (Linnaeus, 1758) ............................ 40
Mustelus punctulatus Risso, 1826 ............................. 40
CARCHARHINIDAE ...................................... 41
Carcharhinus altimus (Springer, 1950) .......................... 41
Carcharhinus brachyurus (Günther, 1870) ........................ 41
Carcharhinus brevipinna (Müller and Henle, 1839) ....................42
Carcharhinus falciformis (Bibron, in Müller and Henle, 1839) ...............42
Carcharhinus limbatus (Valenciennes, in Müller and Henle, 1839) ............. 43
Carcharhinus melanopterus (Quoy and Gaimard, 1824) .................. 43
Carcharhinus obscurus (Lesueur, 1818) .......................... 44
Carcharhinus plumbeus (Nardo, 1827) .......................... 44
Galeocerdo cuvier (Péron and Lesueur, in Lesueur, 1822) ................. 45
Prionace glauca (Linnaeus, 1758) ............................. 45
Rhizoprionodon acutus (Rüppell, 1837) .......................... 46
SPHYRNIDAE......................................... 46
Sphyrna (Mesozygaena)tudes (Valenciennes, 1822).................... 46
Sphyrna (Sphyrna)lewini (Griffith and Smith, in Cuvier, Griffith and Smith,1834) .....47
Sphyrna (Sphyrna)mokarran (Rüppell, 1837) ....................... 47
Sphyrna (Sphyrna)zygaena (Linnaeus, 1758) ....................... 48
vii
BATOID FISHES ........................................... 49
Picturekeyofskates,raysandguitarfishes ............................ 49
TechnicalTermsandMeasurements............................... 50
List of Families and Species Occurring in the Area ......................... 51
Guide to the Order and Families of Batoid Fishes Occurring in the Area ..............52
OrderRAJIFORMES-Batoids................................. 52
Suborder PRISTOIDEI ................................... 52
PRISTIDAE....................................... 52
Suborder RHINOBATOIDEI ................................ 52
RHINOBATIDAE.................................... 52
Suborder TORPEDINOIDEI ................................ 53
TORPEDINIDAE.................................... 53
Suborder RAJOIDEI .................................... 53
RAJIDAE........................................ 53
Suborder MYLIOBATOIDEI ................................ 54
DASYATIDAE ..................................... 54
GYMNURIDAE ..................................... 54
MYLIOBATIDAE.................................... 55
RHINOPTERIDAE................................... 55
MOBULIDAE...................................... 55
Guide to Families and Species
PRISTIDAE......................................... 56
Pristis pectinata Latham, 1794 ............................. 56
Pristis pristis (Linnaeus, 1758)............................. 56
RHINOBATIDAE...................................... 57
Rhinobatos (Glaucostegus)cemiculus Geoffroy St-Hilaire, 1817 ............57
Rhinobatos (Rhinobatos)rhinobatos (Linnaeus, 1758) ................. 57
TORPEDINIDAE...................................... 58
Torpedo (Tetronarce)nobiliana Bonaparte, 1835 ...................58
Torpedo (Torpedo)marmorata Risso, 1810 ...................... 58
Torpedo (Torpedo)sinuspersici Olfers, 1831 ...................... 59
Torpedo (Torpedo)torpedo (Linnaeus, 1758) ...................... 59
RAJIDAE.......................................... 60
Dipturus batis (Linnaeus, 1758) ............................ 60
Dipturus oxyrinchus (Linnaeus, 1758) ......................... 60
Leucoraja circularis (Couch, 1838) .......................... 61
Leucoraja fullonica (Linnaeus, 1758) ......................... 61
viii
Leucoraja melitensis (Clark, 1926) ........................... 62
Leucoraja naevus (Müller and Henle, 1841)....................... 62
Raja asterias Delaroche, 1809 ............................. 63
Raja brachyura Lafont, 1873 .............................. 63
Raja clavata Linnaeus, 1758 .............................. 64
Raja miraletus Linnaeus, 1758 ............................. 64
Raja montagui Fowler, 1910 .............................. 65
Raja polystigma Regan, 1923 ............................. 65
Raja radula Delaroche, 1809.............................. 66
Raja undulata Lacépède, 1802............................. 66
Rostroraja alba (Lacépède, 1803) ........................... 67
DASYATIDAE ....................................... 67
Dasyatis centroura (Mitchill, 1815) ........................... 67
Dasyatis marmorata Steindachner, 1892 ....................... 68
Dasyatis pastinaca (Linnaeus, 1758).......................... 68
Himantura uarnak (Forsskål, 1775) .......................... 69
Pteroplatytrygon violacea (Bonaparte, 1832) .....................69
Taeniura grabata (Geoffroy St-Hilaire, 1817) ...................... 70
GYMNURIDAE ....................................... 70
Gymnura altavela (Linnaeus, 1758) .......................... 70
MYLIOBATIDAE...................................... 71
Myliobatis aquila (Linnaeus, 1758) .......................... 71
Pteromylaeus bovinus (Geoffroy St-Hilaire, 1817) ...................71
RHINOPTERIDAE..................................... 72
Rhinoptera marginata (Geoffroy St-Hilaire, 1817) ................... 72
MOBULIDAE........................................ 72
Mobula mobular (Bonnaterre, 1788) .......................... 72
CHIMAERAS ............................................. 73
TechnicalTermsandMeasurements............................... 73
Order,FamilyandSpeciesofChimaeraOccurringintheArea................... 73
OrderCHIMAERIFORMES .................................. 73
CHIMAERIDAE..................................... 73-74
Chimaera monstrosa Linnaeus, 1758 ......................... 74
BIBLIOGRAPHY ........................................... 75
INDEX TO SCIENTIFIC AND VERNACULAR NAMES ........................ 85
LIST OF COLOUR PLATES ..................................... 94
ix
INTRODUCTION
After a long period of adaptation starting 450
million years ago, since the Cretaceous (about
100 million years) the cartilaginous fish have not
experienced any further noticeable morphological
or physiological changes. Today the group occupies
almost all aquatic environments: rivers, lakes,
estuaries, coastal lagoons, coastal waters, open
seas and deep seas, extending as far as the
Antarctic convergence. This may be facilitated by
the fact that sharks are mostly predators, and have
adapted to all sort of diets, some being scavengers.
The few species that are plankton-feeders are
characterized by large dimensions, which
discourages predation.
This guide provides a list and brief description of
the species of sharks, batoid fishes and chimaeras
living in the Mediterranean and Black Sea. While
for some species there are no doubts regarding
their presence in the area, for other rare species,
their presence remains doubtful pending further
information.
Commercial fishing samples constitute the main
source of cartilaginous specimens, in particular, for
rare species. Some species, once considered rare,
may become common due to the increased use of
a particular gear or during fishing on new grounds.
Fishing may produce negative effects on
biodiversity: overexploitation of resources may
directly or indirectly lead to the depletion of some
species, especially those that do not have a good
resilience as regards to fishing disturbance, such
as the cartilaginous fish. In these cases it is quite
frequent to observe the disappearance, albeit
locally, of some species.
One major problem using official landing statistics
in any analysis is the difficulty to secure correct
identification of the species of cartilaginous fish, as
they are often grouped in collective codes. Apart
from spotted dog sharks, thorn rays and a few other
species, caught as bycatch, a targeted fishery
aiming at catching cartilaginous species does not
exist in the Mediterranean and Black Sea, and for
the time being, finning is not mentioned either. In
trawl fisheries discard may be a critical aspect
especially for juveniles of some species.
Physical and Chemical
Characteristics of the Region
Geology and Physical Features
The Mediterranean Sea, Black Sea excluded,
constitutes 0.81% (2.514 million km2)ofthetotal
water surface of the planet. Twenty-two different
countries border its coastline. It extends from the
Straits of Gibraltar to the Bosphorus for about
4 000 km. The Mediterranean reaches its maximum
depth (5 121 m) in the Ionian Sea.
The birth of the Mediterranean was caused by the
collision of the African and European continents
about 50–60 million years ago in the area presently
called Gibraltar. During the “Messinian regression”
the Mediterranean basin became completely
closed and the water level descended reaching its
minimum 6 million years ago. About 5.5 million
years ago, communication through the Straits of
Gibraltar opened again and Atlantic waters invaded
the Mediterranean basin. From then on the
Mediterranean Sea began a new life and assumed
the characteristics of a temperate sea.
The Mediterranean Sea can be divided into two
main basins: western and eastern separated by the
Sicily-Tunisia ridge. Within these basins, regional
seas may be defined, connected by channels and
straits (Fig. 1). The eastern basin is characterized
by a great oceanographic variability on the surface
with temperatures of 16°C in winter and up to 29°C
in summer, as opposed to 12° and 23°C in the
western basin and salinities of 39‰ to the east as
opposed to 36‰ in the west.
Currently the Gibraltar threshold, with a maximum
depth of about 320 m and a distance of only 25 km
between the European and African continental
masses, allows the passage of the Atlantic upper
layers with an average temperature of about 15°C.
The slow circulation of water masses, the rate of
exchange of Mediterranean waters and the
consequent poor water replacement, contributes in
keeping the temperature constant year round,
particularly in deep waters.
Introduction 1
2 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
Fig. 1 Mediterranean geography and its main sub-basins
A= Balearic Islands; B= Sardinia; C= Corsica; D= Sicily; E=Malta;F=Crete;G= Rhodes; H=Cyprus
1= Straits of Gilbraltar; 2= Alboran Sea; 3= Catalan Sea; 4= Liguro-Provençal Basin; 5= Tyrrhenian Sea; 6= Sicily Tunisian Ridge; 7= Cape Bon; 8= Ionian Sea; 9= Adriatic
Sea; 10 =PomoPit;11 = Dalmato Garganic Threshold; 12 = Levantine Sea; 13 = Aegean Sea; 14 = Marmara Sea; 15 = Bosphorus; 16 = Black Sea; 17 = Azov Sea; 18 = Suez;
19 =RedSea;20= Atlantic
Compiled by D. Amblàs and J.L. Casamor, GRC en Geociències Marines, Universitat de Barcelona (Spain); after International Ocean Commission, International Hydrographic
Organization and British Oceanographic Data Centre - IOC/IHO/BODC (2003): Centenary Edition of the GEBCO Digital Atlas, published on CD-ROM. Liverpool (UK).
The Black Sea occupies an area of about
465 000 km2and has a maximum depth of 2 245 m.
The Marmara Sea connects the Black Sea with the
Mediterranean. Its salinity is low and does not
exceed 22‰ throughout the whole water column,
however surface waters may have lower values,
down to 16–18‰. In winter the surface
temperature decreases to 3–6°C and often, in
some coastal areas, the water freezes, especially
in the Azov Sea. At depths over 150 m the
temperature is practically constant, about 9°C. The
waters of the southern coasts (Anatolia) show
greater average temperatures and are separated
by an isothermal front of 15°C. One of the main
characteristics of this sea is the complete lack of
dissolved oxygen at depths over 150–200 m; from
this depth to the bottom sulphurous concentration
is very high (Murray, 1991; Oguz, 1992, 1993).
Oceanographic Features
The distribution of marine organisms in the
environment is clearly related to bottom
characteristics, nutrients abundance and oceano-
graphic conditions. These circumstances are
naturally linked to water masses large movements,
both near the surface and in deep waters, and are
also influenced by meteorological conditions such as
wind intensity, surface temperature and chlorophyle
concentration (Figs 2, 3 and 4).
The superficial layers of Atlantic waters flow in
through the Straits of Gibraltar, progressing over
the entire surface of the Mediterranean basin area.
These waters become warmer and progressively
saltier due to evaporation and their great density as
they sink. Part of the general flow will return to the
Atlantic as intermediate waters, while another part
will mix with deep waters. Heburn (1992) and
Garibaldi and Caddy (1998) distinguish three
different ecological areas based on species
distribution. Three types of water that
fundamentally characterize the balance of the
whole Mediterranean can be suggested:
The Modified Atlantic Water (MAW), which mainly
constitutes the surface water (0–200 m) of the
whole area. It initially flows close to the North
African coasts, from Morocco to Cape Bon and
then splits into three main directions: the first one
constitutes the cyclonic circuit of the Balearic
Islands, the second moves towards the Tyrrhenian
Sea (Astraldi et al., 1999) and the last one towards
the Levantine Sea (Millot, 1999) (Fig. 5).
The Levantine Intermediate Water (LIW)
(200–1 000 m, mainly around 400 m depth) that
constitutes the main component of the returning
flow towards the ocean. This water is mainly
produced in the eastern basin (Lascarotes et al.,
1992, 1993) and, to a lesser extent in the western
basin contributing to the Tyrrhenian movement
(Fig. 6).
The Mediterranean Eastern and Western Deep
Waters (MDW, Mediterranean Deep Water)
(>1 000 m), that are produced respectively in the
Liguro-Provençal area of the basin (Send and
Shott, 1992) and in the Southern Adriatic Sea
(Fig. 7).
The general circulation in the Black Sea consists of
several sub-basin scale gyres. The anticyclonic
coastal eddies appear to play a fundamental role
on the ultimate distribution of the Cold Intermediate
Water (Oguz et al., 1992, 1993; Millot, 2005;
Korataev et al., 2003) (Fig. 5).
In the area of entrance of the Atlantic waters (the
Alboran Sea), there are important phenomena
having repercussions on the entire Mediterranean
basin (Fig. 8). These waters show an almost
permanent anticyclonic gyre in the west and a more
variable circuit in the east, (Allain, 1960; Lanoix,
1974; Heburn and La Violette, 1990; Davies et al.,
1993; Viudez et al., 1996). The main flow is from
Spain to the Algerian coast, commonly named “the
Almeria-Oran jet” (Prieur and Sournia, 1994). After
about 80–100 years the Mediterranean waters
(LIW and MDW) return to the Atlantic Ocean with
different velocity running below the surface
(Bryden and Kinder, 1991) (Fig. 9).
Biodiversity and Biogeographical
Characteristics of the Region
The current level of biodiversity of the
Mediterranean and Black Sea fauna was defined
by the alternation of periods of glaciation and
interglaciation, which brought about dramatic
changes in climatic conditions. Also in recent
times, biodiversity has been enriched both through
internal speciation phenomena and through
species colonization from outside Mediterranean
areas (Golani et al., 2002).
Even though the Mediterranean and Black Seas
represent less than 1% of the total area of world
seas, the fish biodiversity and absolute number of
species are relatively high. In fact, it is possible to
find about 6% of the entire world’s species in this
area (Fredj et al., 1992) and probably the 84
cartilaginous fish species found in the area
represent about 8% of the total number of species
of this group in the world.
Introduction 3
4 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
Fig. 4 Average concentration of chlorophyll (milligrammes per m3)
The Gulf of Lion and Adriatic Sea are richer areas
(Satellite imagery: © OCEAN Project, 2000)
Fig. 3 Average sea surface temperature (°C)
The Gulf of Lion and the northern part of Aegean Sea are colder areas
(Satellite imagery: © OCEAN Project, 2000)
Fig. 2 Average wind speed (metres per second)
The Gulf of Lion and the Aegean Sea close to the island of Rhodes are the most windy areas
(Satellite imagery: © OCEAN Project, 2000)
Introduction 5
Fig. 7 The dynamics of Deep Waters
(<1 000 m depth)
(after Millot (1987), Lacomb et al. (1985) for the western basin. The movements of the deep currents in the eastern basin have
been simplified (Anati, 1977; Roether and Schlitzer, 1991; Millot and Taupier-Letage, 2004)
Fig. 6 The dynamics of the Intermediate Waters (~200–1 000 m)
(from Millot (1987), Robinson and Golnaraghi (1994) for the western basin and Malanotte-Rizzoli et al. (1999);
Millot and Taupier-Letage, 2004 for the eastern basin, modified)
Fig. 5 General circulation of the superficial currents (~0–200 m depth)
(redrawn from Millot (1999) for the western basin also considering information from Heburn and La Violette (1990) and
Tziperman and Malanotte-Rizzoli (1991). The dynamics of the eastern basin come from the information of Robinson and
Golnaraghi (1994) in Malanotte-Rizzoli and Robinson (1994) and Millot and Taupier-Letage, 2004).
6 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
Fig. 8 Noteworthy dynamic superficial waters referring to 200 m depth
(from Lanoix, 1974 in La Violette, 1984)
Fig. 9 Inflow and outflow currents through the Strait of Gibraltar:
a) transversal section, b) longitudinal section.
LIW spreads and mixes but remains a distinct water mass all the way to the Gibraltar Strait (Gascard and Richez,
1985); LIW and MDW probably do not mix completely and flow out of Gibraltar Strait with different velocity (Millot,
1999). The yearly average temperatures and salinity of LIW, MDW and MAW are illustrated respecitvely (Kinder and
Bryden, 1990; Millot, 1999; Robinson and Golnaraghi, 1994; Tintoré et al., 1988; Kinder and Parrilla, 1987). The
renewal of the waters of the entire Mediterranean basin is not known exactly; the eastern circulation is now known to
consist of a single cell encompassing both the Ionian and Levantine basins, with a turnover time extimated at about
125 years. More research is needed to define the western cell (Roether and Schlitzer, 1991).
Two main features can be highlighted concerning
Mediterranean fish communities:
Biodiversity decreases from west to east, probably
due to physical conditions such as the presence of
threshold-strait or canal effects (Gibraltar,
Sicly-Tunisia, Bosphorus, and Suez). The diversity
in number of species shows the same negative
eastward gradient that has been found for nutrients
(Murdoch and Onuf, 1972). Minimum biodiversity
is present in the Adriatic and Black Sea (Fredj and
Maurin, 1987; Garibaldi and Caddy, 1998).
There is a meridionalization from the southern to
the northern coasts (Riera et al., 1995) and a
warming trend in the deep waters of the western
Mediterranean (Berthoux et al., 1990).
There are approximately 1 170 valid species of
cartilaginous fish in the world’s waters; about 50 of
them are chimaeras, 650 batoids and 470 sharks.
In the Mediterranean and Black Sea, 7 orders are
represented by 23 families, 42 genera and a total
number of species of about 47 sharks, 34 batoid
fishes and 1 chimaera. Endemism is low; up to four
species of rays could be considered indigenous.
In the Adriatic Sea, the presence of cartilaginous
fish species is scarce especially in the northern
part. Besides its oceanographic characteristics that
may limit biodiversity, this area was populated
more recently than other parts of the
Mediterranean. This occurred after the sinking of
the Dalmato-Garganic threshold, which was still
above sea level in the Pleistocene. A total of 52
species of cartilaginous fish have been recorded in
the Adriatic Sea. Only 10 species are widely
distributed. Some bathyal species of the group
inhabit exclusively the central and southern parts of
this sea (Jardas, 1984).
In the Black Sea the number of cartilaginous fish
species is less. The Pontic fauna is composed of
Mediterranean species and most of the organisms
present are eurythermic and euryhaline. Twelve
cartilaginous fish species are assumed to live in the
Black Sea (Tortonese, 1969; Bouchot, 1984; Roux
in FNAM, 1984; McEachran and Capapé, in FNAM,
1984; Fredj and Maurin, 1987). Murat et al. (2002)
consider only 8 elasmobranchs along the Turkey
coast of the Black Sea.
The Mediterranean Sea comprises several
sub-basins characterized by more or less widely
diverging oceanographic conditions and faunistic
features. Bearing this situation in mind, and also
considering the proximity of the Mediterranea Sea
to the Atlantic Ocean, strictly speaking the
Mediterranean fauna can be defined as the fauna
of a single, well-known, well-defined basin, and in a
wider sense the term also covers the forms existing
in the adjacent part of the Atlantic, between
Portugal and Mauritania including the Azores,
Canary and Madeira islands (Tortonese, 1989).
For the last 5.5 million years, the Straits of Gibraltar
have never constituted a rigid boundary, and there
is, therefore a reciprocal influence between the
Atlantic Ocean and the Mediterranean Sea. Infact,
the classical statement of Ekman (1953) considers
the Atlantic-Mediterranean area as a single
faunistic unit, and divides it into three areas:
Lusitanic, Mauretanic and Mediterranean Region.
Even if considering only the Mediterranean basin, the
distribution of cartilaginous fish species is not
homogeneous. This phenomenon is often linked to
the typology of the sea bottoms or to the chemical
and physical characteristics of the different
sub-basins. Basically, the bathymetry delimits three
distinct ecological areas, which can be used to
categorize species distribution patterns and hence
habitat preference. Obviously, species could belong
to more than one category or to all of them (Garibaldi
and Caddy, 1998): 1) those living over the shelf
(0–200 m); 2) demersal on the slope, oceanic and
mesopelagic species within the water column over
the 200–1 000 m depth; and 3) oceanic, mesopelagic
and bathypelagic species occupying waters over
1 000 m depth.
The superficial Atlantic current, which comes
through the Straits of Gibraltar, is of crucial impor-
tance for Mediterranean Sea life as it facilitates the
immigration of oceanic species. In a very synthetic
way we can state the following biodiversity consid-
erations on several biogeographical areas of the
Mediterranean basin:
The Alboran Sea is rich in Atlantic species.
The northwestern area of the Mediterranean,
including the Catalan, Ligurian and north
Tyrrhenian seas, is characterized by the presence
of Atlantic boreal elements.
The central zone, that includes waters around the
Balearic Islands, Corsica and Sardinia and the
northern coasts of Sicily, shows specific Mediterra-
nean or Atlantic-Mediterranean characteristics; many
subtropical species are found in this area.
The Tunisian and Libyan coasts, characterized by
the presence of rare tropical Atlantic species, are
the southernmost areas and are closest to
subtropical in their characteristics.
The eastern region, that includes the coasts of
Egypt, Israel, Lebanon and Syria, is inhabited by
many species coming from the Red Sea.
Introduction 7
The Adriatic Sea that does not reach great
depths (50–60 m in average), apart from the
“Pomo Pit” (south Adriatic Sea) where depth
reaches about 1 200 m; the most abundant
marine organisms in this area are prevalently
species of Atlantic-boreal origin.
The Black Sea is characterized by species of
Sarmatic origin.
The Fishery
In all the seas of the world, the cartilaginous fish
species are exploited for their fins, skin, jaws or
meat (Vannuccini, 1999). Sometimes they are
directly targeted by commercial and recreational
fisheries while in other cases they are incidentally
caught as bycatch. In many areas of the world a
decline in cartilaginous fish species landings has
been observed while fishing effort has generally
increased. This especially applies to fisheries
targeting shark fins. Moreover, most countries
report shark statistics without distinction between
species or, worse still, the species are not recorded
at all. As a result, it is impossible to identify the
species in multispecific fishery and hence estimate
and monitor fishing mortality.
Because of their life history characteristics, sharks
and rays are especially susceptible to over-
exploitation and it is very difficult to restore depleted
populations. Very often species have restricted
distribution, small population size, dependence on
mating, spawning, nursery and breeding grounds or
specific habitats. Well- documented cases of
collapsed shark fisheries have been reported
(Musick et al., 2000). In such cases a sudden
collapse of yields can occur and consequently the
local extinction of a particular species.
No marine fish is yet known to have been driven to
biological extinction due to fishing (Musick, 1999) but
regional stocks of some species can be considered
to have disappeared, such as Squatina sp. (Vacchi
and Notarbartolo di Sciara, 2000). The assumption
that marine fish are not vulnerable to extinction
because they live in open seas, where their
movements are unlimited, is unfounded. Sharks also
constitute a bycatch in open sea fisheries targeting
highly migratory species such as tunas.
The fishing methods used to catch cartilaginous
fish species in the Mediterranean are highly varied:
the two most efficient gears for sharks are gillnets
and longlines, while a frequent method for catching
batoids in general and some smaller sharks like
smoothhound, catsharks, etc. is the bottom trawl.
This fishing gear is probably responsible for a large
amount of bycatch and discard of cartilaginous fish
throughout the world (Bonfil, 2002).
Bottom trawl fishing activity is commonly
performed throughout the Mediterranean area. The
cartilaginous fish species most frequently caught
with these gears are Galeus melastomus,
Scyliorhinus canicula,Etmopterus spinax,Raja
clavata,Squalus acanthias (Bertrand et al., 2000;
Relini et al., 2000; Baino et al., 2001; Serena et al.,
2005). Some species such as the starry ray (Raja
asterias) are constantly captured as bycatch and in
large amounts in several Italian fisheries,
especially in the Tyrrhenian Sea. The fishing gear
employed is a modified beam trawl targeting sole
(Minervini et al., 1985; Serena and Abella, 1999;
Abella and Serena, 2005).
Even in the Adriatic Sea, accessory catches of
many species of cartilaginous fish species are
carried out with set gears and the most frequent
species caught are Squalus acanthias,
Mustelus spp., Raja spp., Torpe d o spp.,
Scyliorhinus spp., Galeorhinus galeus,Alopias
vulpinus and juveniles of Carcharhinus plumbeus
(Costantini et al., 2000).
Large individuals of Hexanchus griseus as well as
those of Galeus melastomus,Centrophorus
granulosus, etc. are captured with bottom longlines
targeting hake. The drifting longlines set near the
surface, targeting tuna and swordfish, also capture
Prionace glauca,Pteroplatytrygon violacea,
Alopias vulpinus,Isurus oxyrinchus,Lamna nasus,
Sphyrna (Sphyrna) zygaena,Hexanchus griseus,
Carcharhinus spp., Mobula mobular,etc.(Fleming
and Papageorgiou, 1997; Kabasakal, 1998;
Hemida, 1998; De Metrio et al., 2000; Garibaldi
and Orsi Relini, 2000; Orsi Relini et al., 2000).
Driftnets are largely used to catch cartilaginous fish
species, and in the recent past they were
extensively utilized throughout the Mediterranean.
Fortunately nowadays their use is prohibited in
European countries. It is advisable that the use of
this gear be prohibited, and this should be
extended to all Mediterranean countries in order to
find a definitive solution to the problem. The main
species caught with driftnets are Prionace glauca,
Alopias vulpinus,Isurus oxyrinchus,Lamna nasus,
Carcharhinus spp., Cetorhinus maximus,
Sphyrna spp., Mobula mobular,Pteroplatytrygon
violacea (De Metrio et al., 2000).
Occasionally, species such as Prionace glauca,
Cetorhinus maximus,Sphyrna spp. and Mobula
mobular are caught with purse seines (Notarbartolo
di Sciara and Serena, 1988). In such cases the
8 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
catches have to be considered incidental rather than
accessory, the same applies to some artisanal
fisheries (Serena et al., 1999a, b). No official practice
of “finning” has been reported in the Mediterranean
Sea so far, but mortality through discarding from
trawls, gillnets, purse seines and longlines is
significant (De Metrio et al., 1984). However, in the
past the fishing activity with the greatest incidence in
cartilaginous fish species catches was tuna traps.
Some years ago, these fishing structures were widely
distributed all around the Mediterranean area. In
countries such as Spain, France and Turkey, and
particularly along the Italian coasts, the use of the
tuna trap was due to the presence of the prevailing
migration routes of tuna, directed towards not only
the rich waters of the Liguro-Provençal basin but also
in the Adriatic Sea (FAO, 1985). Between 1890 and
1914, there were 37–54 tuna traps (Cushing, 1988) in
Italy. Today only a few units are still present, mainly
concentrated on the major Italian islands. The
large-sized cartilaginous fish species more
commonly present in catches were Alopias vulpinus,
Cetorhinus maximus,Sphyrna mokarran,Prionace
glauca,Mobula mobular and sometimes
Carcharodon carcharias (Boero and Carli; 1979,
Vacchi et al., 2002).
Significant bycatch mainly constituted by Alopias
vulpinus and Prionace glauca is caught by fishing
performed with small swordfish-driftnets targeting
swordfish, carried out mainly in the southern part of
the Mediterranean by Italy, Malta, Morocco,
Tunisia, and others. Very few and geographically
localized fishing activities can be considered,
targeting species of this group in the
Mediterranean. Traditionally Hexanchus griseus is
caught with bottom longlines in the Ligurian Sea
(Aldebert, 1997) and also along the southern Italian
coasts. In this area, drifting surface longlines,
called “stese” are also utilized in spring for the
catch of large individuals of Prionace glauca.
These are short lines with hooks that are set near
the surface. In the northern Adriatic, gillnets are
utilized to catch Mustelus mustelus,Mustelus
punctulatus,Squalus acanthias,Scyliorhinus
stellaris,Myliobatis aquila and Galeorhinus
galeus during winter and spring; and Prionace
glauca,Pteromylaeus bovinus and Alopias
vulpinus during summer (Costantini et al., 2000).
Important catches of Carcharhinidae species
(Carcharhinus brachyurus,Carcharhinus brevipinna,
Carcharhinus falciformis,Carcharhinus obscurus,
Carcharhinus plumbeus and Carcharhinus altimus)are
also made by offshore pelagic longline fishery operating
from ports in the east of Algeria (Hemida and Labidi,
2000). The recent FAO-COPEMED-MBRC report by
Lamboeuf (2000) analyses the artisanal fishery in Libya
showing some examples of fisheries targeting
cartilaginous fish such as Carcharhinidae, Lamnidae,
Rhinobathos and Squatina squatina caught by fixed
gillnet, bottom set and drifting longlines.
Finally, we cannot neglect the role of recreational
fishing that has recently grown in popularity
causing concern. Following the development that
occurred in the United States and in Australia, the
number of angler associations has also notably
increased in the Mediterranean, mainly in the
northern Adriatic (Bianchi et al., 1997) and in the
Tyrrhenian but also in other countries such as
France and Spain. The targets of game fishing are
essentially Alopias vulpinus,Prionace glauca and
Hexanchus griseus. However, juveniles compose
most of the catch and, sometimes, they are
recently born individuals. Currently, there are no
specific laws or a suitable control aimed at the
protection of any cartilaginous fish. This may soon
lead to a rarefaction of the populations of the two
above-mentioned species as has already occurred
in Cornwall, United Kingdom (Vas, 1995).
Cartilaginous fish catches in the 1970–2002 period
represent only 1.1% of the total landings in
Mediterranean ports (FAO, 2000a). The most
important landings of this group occurred in the
Ionian and Black seas each one with 30% of the total
Mediterranean catches; Sardinian, Adriatic and
Balearic waters show catches of 12%, 8% and 7%,
respectively of the Mediterranean total.
The catches during the last 30 years (Fig. 10) show
an increasing trend from 10 000 to 25 000 tonnes
attained in 1985 and since then a regular decrease
to 15 000 tonnes to present (FAO, 2000b). This is
mainly due to the Turkish and Italian catches of
sharks and rays in the Black and Ionian seas,
respectively. Unfortunately it cannot be stated
whether these variations are real or if they are
simply due to changes in recording procedures (i.e.
in some years at least part of them were reported
as sharks and in others generically as marine
fishes).
Introduction 9
0
5,000
10,000
15,000
20,000
25,000
30,000
1970 1975 1980 1985 1990 1995 2000 2005
Fig. 10 Mediterranean and Black Seas trend of
cartilaginous catches in the last 30 years
Fishery Management
The life history strategy of cartilaginous fish
species suggests the need for conservative
management of a balanced population and a
compatible fishery activity. However, most shark
fisheries are completely unmonitored and
unmanaged (Shotton, 1999). About 50% of the
estimated global catch of cartilaginous fish species
is gathered as bycatch and these are not
mentioned in official fishery statistics (Stevens
et al., 2000).
In general, the contribution of cartilaginous fish to
the market is low because of their scarce economic
value. Consequently they are also a low priority for
research and management if compared to bony
fishes, although recently, certain products such as
shark fins have become important in the trade.
Cartilaginous fish resources management needs
particular attention, above all in order to maintain
biodiversity and ecosystem structures. The
interactions between species in marine
ecosystems, as well as the impact of the removal
of top predators on other marine organisms on the
functionality of the whole ecosystem are poorly
understood. Basically, besides an adequate
identification guide, we need to know the
abundance of each species, life history
characteristics, fishing effort, catches, discards,
etc. In the case of the Mediterranean, many
fisheries belonging to different countries are taking
place and the resources are often shared between
nations. This situation requires cooperative
management at the intergovernmental level and a
precautionary approach for their exploitation (FAO,
1995a, 1995b, 1996).
Finally, for the successful management of
cartilaginous fish species we must simultaneously
consider biological and fishery information. The
choice of suitable mathematical models for stock
assessment of these resources is not easy and it
has to take into consideration the long life span and
late maturation of many sharks and rays.
Therefore, the effects of changes in fishing effort
and other management measures will be apparent
only many years later.
Conservation
The cartilaginous fish species belong to an ancient,
conservative taxonomic group that was formerly
very abundant in the world’s oceans. Despite the
evolutionary success of the species which have
survived until the present day, some are now
threatened with extinction, regionally or globally,
often as a result of human activities. The main
reason for this is that a K-selected life-history traits
characterize many species; they grow slowly,
mature at a relatively late age, have only a few
young with low natural mortality rates, and their
populations increase very slowly (Hamlett, 1999,
2005). As apex predators occupying the top of the
marine food web, many cartilaginous fish species
are also naturally rare compared with other fish
species but have a vital role in maintaining the
balance of marine ecosystems. Their biological
characters make them susceptible to population
depletion as a result of anthropogenic activity,
including unsustainable fisheries supplying local
demand or international trade, bycatch, habitat
modification and persecution (particularly of
species perceived as dangerous to man).
The IUCN–World Conservation Union Species
Survival Commission’s Shark Specialist Group is
currently assessing global- and regional-extinction
risks for all species of sharks and their relatives,
including Mediterranean populations. This list is
updated annually as new information becomes
available and may be consulted on www.redlist.org.
The Red List has no legal standing, but is widely
used to monitor changes in the status of
biodiversity and to set conservation and
management priorities. Regional networks of
experts are involved in assessing and reassessing
the Red List status of species, drawing upon
information collected by stock assessment and
other research projects within the region (Fowler
et al., in press).
Recognition of the threatened status of sharks and
their relatives has been recognized through the
addition of several species to national, regional and
international species and fisheries conservation
and management instruments. A number of
species of sharks and rays are listed in the
Appendices of the Barcelona Convention for the
protection of the marine and coastal environment
of the Mediterranean (Protocol concerning
Specially Protected Areas and Biological Diversity)
and the Bern Convention for the Conservation of
European Wildlife and Natural Habitats, which
specifically cover populations in the
Mediterranean. Other species are listed in the
Convention on Migratory Species, the Convention
on International Trade in Endangered Species
(CITES), and the UN Fish Stocks Agreement.
Since some of these lists change fairly regularly
readers are advised to consult the appropriate
Convention web pages for the most up-to-date
information.
10 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
It is important to note that the biological
vulnerability of sharks1/, recognized in the FAO
International Plan of Action for the Conservation
and Management of Sharks (FAO IPOA-Sharks,
1998) means that it is important to monitor the
status of all species and to ensure that appropriate
management measures are introduced in order to
guarantee the sustainable use of all shark stocks,
not only those which are listed in the Conventions
or the IUCN Red List of Threatened Species.
Concerns over the sustainability of shark fisheries
led to the development and adoption in 1999 of the
FAO IPOA-Sharks, 1998, elaborated within the
framework of the Code of Conduct for Responsible
Fisheries (FAO, 2000). This voluntary plan urges
states to carry out a regular assessment of the
status of shark stocks subject to fishing, in order to
determine if there is a need for development of a
shark plan, and to adopt a national plan of action
(Shark-plan) for conservation and management of
shark stocks (if their vessels conduct directed
fisheries for sharks or if their vessels regularly
catch sharks in non-directed fisheries). It also
recognizes the importance of international
collaboration for the sustainable management of
transboundary, straddling, highly migratory and
high seas shark stocks, including, where
appropriate, the development of subregional or
regional shark plans.
The European Union considers that sharks are fish
species whose conservation falls within the domain
of the Common Fishery Policy, therefore their
management should be addressed by measures
dictated by the EC for implementation within EU
countries. The European Plan of Action,
announced at the FAO Committee on Fisheries
(COFI) meeting held in Rome in February 2001, is
still to be released. Meanwhile, two meetings of the
ad hoc Elasmobranch’s Working Group of the
Scientific, Technical and Economic Committee for
Fisheries-Subgroup on review of stocks (STECF-
SGRST) have been held (2002 and 2003) in order
to address elasmobranch fisheries with a view to
preparing a Community Plan of Action as
requested by the FAO-IPOA Sharks. The draft
Italian National Shark Action Plan recognizes the
need for regional cooperation in addition to national
action for Mediterranean shark species. This was
the starting point for Italy’s active involvement
within the relevant international and regional
organizations, such as FAO and UNEP-MAP 2003.
This stimulated the formulation of a Mediterranean
Action Plan for the conservation and management
of cartilaginous fishes, proposed during meetings
of the GFCM-SAC working group on the
environment in 2001 and 2002, and accepted by
the National Focal Points to the SPA Protocol in
2001. The approved Action Plan was scheduled for
adoption in November 2003. It strongly
recommended that the representative parties grant
urgent legal protection status to a list of priority
species assessed as Critically Endangered or as
Endangered by the IUCN at the Mediterranean
level and urges assessment of the extinction risk to
species, such as hammerhead sharks and
guitarfishes, for which data are lacking. The
representative parties are also asked to develop
management programmes for sustainable fisheries
for a number of commercially important species, to
identify and protect critical habitats and to develop
research, monitoring and training programmes.
Although legal instruments for the conservation of
some cartilaginous-fish species in the
Mediterranean have been in place for over eight
years, implementation has not yet followed. For
example, species listed under Appendix III of both
the Barcelona and the Bern Conventions, which
call for the regulation of their exploitation, have
continued to decline without any management
during this period. There is now a critical need for a
concerted action and synergy of both fisheries and
environmental agencies throughout the region to
ensure the conservation and sustainable use of
this vulnerable group, and hence the maintenance
of the stability of the Mediterranean ecosystem.
Such action should stem from the frameworks of all
those institutions whose mandate involves
environmental and fisheries policies within the
Mediterranean basin and the application of the
ecosystem approach and precautionary principle.
It should also be mentioned that an Action Plan for
the Conservation of Cartilaginous Fishes in the
Mediterranean has been proposed (UNEP MAP
RAC/SPA, 2003).
Codes for Conservation and Exploitation Status
With the aim of assigning status categories
regarding the overall human utilization of sharks,
the FAO (Castro et al., 1999) allocated sharks
species into two main groups: “exploited” and “not
exploited” species. The group “exploited species”
is successively divided into five numerical
categories. These categories and criteria for
inclusion are explained as follows. Unfortunately
batoids are not yet considered in the FAO status
evaluation.
Introduction 11
1/ The term “shark” is used here in the sense of the Convention on International Trade in Endangered Species (CITES) and the FAO
International Plan of Action for the Conservation and Management of Sharks (IPOA-SHARKS).
A. Not exploited species: species that are not
currently targeted by fisheries and are not normally
found as bycatch of any fisheries.
B. Exploited species: species that are directly
exploited by fisheries or caught as bycatch.
Category 1: exploited species that cannot be
placed in any of the subsequent categories
because of lack of data.
Category 2: species pursued in directed
fisheries and/or regularly found in bycatch,
whose catches have not decreased historically,
probably due to their high reproductive potential.
Category 3: species that are exploited by
directed fisheries or bycatch and, due to a
limited reproductive potential and/or their life
history characteristics, are especially vulnerable
to overfishing and/or are being fished in their
nursery areas.
Category 4: species that show substantial
historical declines in catches and/or have
become locally extinct.
Category 5: species that have become rare
throughout the ranges where they were formerly
abundant, based on historical records, catch
statistics or experts’ reports.
The conservation and exploited status of the
Mediterranean Chondrichthyans fish have been
discussed in several meetings, UNEP-RAC/SPAin
Rome, December 2002 (UNEP 2003), STECF in
Brussels, July 2003, (STECF 2003) and
IUCN-SSG in San Marino, September 2003 (IUCN
2003). Some other information proceeding from
elaborations of the data gathered during national
and international surveys (GRUND, MEDITS,
project N° 97/50 DG XIV/C1, etc.) (Relini et al.,
2000; Bertrand et al., 2000; Megalofonou et al.,
2000; Baino et al., 2001).
Some species as Chimaera monstrosa,Galeus
melastomus,Raja miraletus and Raja clavata,are
referredtoashavinga“stablebiomass”insome
areas from the exploitation point of view (Abello
and Serena, 2005; Serena et al., 2003).
12 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
CLASSIFICATION AND SYSTEMATIC ARRANGEMENT
Considering that the purpose of this document is to
provide a simple user-friendly guide for species
identification, no reference will be found here to
dichotomy keys for single species. It is important
that the classification used in this guide be defined,
as available literature is not always in agreement
with this presentation. The classification of this
group is still under review as no consensus has
been found to reconcile different authors’ positions.
For more information and further specific details on
the taxonomy and biology of cartilaginous fish
species, refer to Tortonese, 1956; Hureau and
Monod 1979; Whitehead et al., 1984; Fischer et al.,
1987; Fredj and Maurin, 1987; Compagno, 1988,
2005; Nelson, 1994; Shirai, 1996; Mould, 1998. The
consultation of FishBase http://www.fishbase.org
(Froese and Pauly, 2000) proved very useful. The
most fundamental references are Compagno’s
catalogues issued in 1984 and his recent revision
partially issued in 2001.
This guide follows the systematic organization
proposed by Compagno (1999, 2001) and the
classification reflects a cladogram attempt where a
new concept of cladistic classification is used. For
instance, the batoids are raised to the order
(Rajiformes) belonging to the superorder of the
Squalomorphi, even if perhaps a more suitable
name should be found to indicate both Rajiformes
and Squaliformes. At the same time the sawsharks
group is raised to the order Pristiophoriformes. So
the batoids have been diversely allocated with
respect to the previous taxonomic organizations.
However, even if this new phylogenetic
classification is considered valid, for practical
reasons sharks and batoids are described
separately in the text.
Taking only modern sharks into consideration, we
can adopt the following, simplified classification
(the orders with no representatives in the
Mediterranean Sea are indicated by an asterisk):
Class Chondrichthyes (cartilaginous fishes)
Subclass Holocephali (chimaeras)
Order Chimaeriformes (chimaera and silver sharks)
Subclass Elasmobranchii (sharks)
Superorder Squalomorphi (squalomorph sharks)
Order Hexanchiformes (cow and frilled sharks)
Order Squaliformes (dogfish sharks)
Order Squatiniformes (angel sharks)
Order Pristiophoriformes (sawsharks) *
Order Rajiformes (batoids)
Superorder Galeomorphi (galeomorph sharks)
Order Heterodontiformes (bullhead sharks) *
Order Lamniformes (mackerel sharks)
Order Orectolobiformes (carpet sharks) *
Order Carcharhiniformes (ground sharks)
The species inside the families and in the orders
are mentioned in alphabetical order according to
genus. For both orders and families, some
summary descriptions with their most significant
characteristics are included. The current status is
described for each single species on an individual
sheet where, in addition to a drawing, scientific
name and more recent synonyms and significant
misidentifications in some important cases, other
synthetic information is given, i.e. FAO common
names, maximum size, habitat and biology,
methods of capture and, when available,
exploitation and conservation status. Arrows are
superimposed on the drawings to indicate features
that help in species identification.
Introduction 13
When possible the type of spiral valve which
characterizes the intestine of cartilaginous fishes is
shown. There are large variations in the anatomy of
the spiral valve, three different types have been
described: columnar spiral with funnels pointing
either backward or forward; spiral ring valve and
cylindrical (scroll) valve (Compagno, 1988;
Hamlett, 1999).
Local names are not included considering the large
number of names used in the various
Mediterranean countries. Feeding behaviour is
indicated only in some cases. Remarks are
indicated for species whose taxonomic status or
presences are dubious.
For the purpose of this guide, a number of dubious
species have not been considered as valid for the
Mediterranean:
Carcharhinus leucas (Valenciennes, 1841) is a
doubtful species; it is neither recognized by
Compagno nor in this guide.
Carcharhinus longimanus (Poey, 1865) is a
doubtful species; it is considered “probable” by
Compagno but is not included in this guide.
Rhinobatos halavi (Forsskål, 1775) was recorded
by Tortonese (1951a) from the Egyptian part of the
Mediterranean Sea but Ben-Tuvia (1966) noted
that no specimens of this species were available to
confirm its presence in this sea.
Torpedo alexandrinsis Mazhar, 1982 and
Torpedo fuscomaculata Peters, 1855 are not
considered a valid species. Torpedo alexandrinsis
is known by only five syntypes mentioned in the
original paper from Alexandria (Egypt) (Séret,
pers. comm.); therefore its taxonomic status is
doubtful. The second Torpedo species recorded
only once in Alexandria (Egypt) needs to be
verified and is probably synonymous of Torpedo
(Torpedo) marmorata (Séret, pers. comm.).
The species Raja africana Capapé, 1977,
previously defined as dubious by Compagno
(1999), is now indicated as not a valid species
(syntypes lost).
Raja rondeleti Bougis, 1959 is probably based on
an abnormal specimen of R. fullonica and
considered as Leucoraja cfr. fullonica.The
taxonomic status of four specimens from French
and Italian coasts is doubtful (Séret, pers. comm.).
How to use this guide
Readers are advised to follow these simple steps in
order to successfully identify any sharks, batoids
and chimaera found in the region. First, refer to the
picture key of shark-like and batoid fishes then
read carefully through the description of key
characters listed under each order and family. Use
the illustrations of the families under each order or
suborder only as a secondary aid in making certain
that the right order or suborder has been found.
Then proceed to narrow down the family of the
specimen using the illustration for each family and
key characters annotated in each illustration; make
use of the size data included for each family. Once
the family has been identified, move to the
corresponding pages where the species for that
family are illustrated. These illustrations and the
key characters indicated should allow proper
identification for all sharks and batoids known for
the area.
In a few cases the considered area is wider than
the Mediterranean basin; it refers to the CLOFNAM
area (Hureau and Monod, 1979): Mediterranean
and northeastern Atlantic between 30° and 80° of
Latitude north, -30° and +60° of Longitude, Azores
and Madeira Islands included (Whitehead et al.,
1984).
Although Chimaera monstrosa (Linnaeus, 1758) is
represented in the sharks cladogram as indicated
by Compagno, 2001, the species account is
inserted at the end of the guide to follow the
taxonomic sense. Eventhough there is only one
chimaera species in the Mediterranean Sea, the
author thought it was important to illustrate the
technical terms and description of this order.
14 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
Sharks and Chimaeras - Picture Key of Shark-like Fishes 15
HOLOCEPHALI
SHARK-LIKE
FISHES
6or7gillslits,1dorsalfin
mouth ventral, pectoral fins
attached to sides of head
ELASMOBRANCHII
CHIMAERIFORMES
RAJIFORMES
HEXANCHIFORMES
PRISTIOPHORIFORMES*
HETERODONTIFORMES*
LAMNIFORMES
CARCHARHINIFORMES
SQUALIFORMES
SQUATINIFORMES
ORECTOLOBIFORMES*
snoutshort,notsaw-like
mouth terminal, pectoral
fins not attached to head
nictitatingeyelids,spiral
orscrollintestinalvalve
dorsal-finspines
nofinspines
mouthwellinfrontofeyes
OrderswithanasteriskarenotrepresentedintheMediterranean
PICTUREKEYOFSHARK-LIKEFISHES
(notacladogram)
snoutelongated,saw-like
1gillslit
noanalfin
5to7gillslits
analfinpresent
5gillslits,
2dorsalfins
bodyflattened,ray-like
nonictitatingeyelids,ring
intestinalvalve
mouthbehind
frontofeyes
bodynotray-like
SHARKS AND CHIMAERAS
(from Compagno, 2001)
SHARKS
TECHNICAL TERMS AND MEASUREMENTS
16 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
dorsal-fin spine
(if present)
pectoral fin
pelvic fin
clasper
(male sex organ) caudal fin
pectoral-fin length
tail
head trunk
snout
1st dorsal fin
interdorsal space
2nd dorsal fin
keel
gill slits
subterminal
notch
caudal
peduncle
anal fin
precaudal pit
nostril
spiracle
labial
furrows
total length (caudal fin depressed to body axis)
location of
intestinal valve
interdorsal-fin ridge
snout
gill slits
pectoral fin
anal opening
pelvic fin
(female, no claspers)
preanal ridges
caudal fin
anal fin
nostril
trunk
underside view
precaudal tail
Sharks - Technical Terms and Measurements 17
upper origin
dorsal margin
terminal lobe
preventral margin
ventral tip
lower postventral margin
postventral notch
upper postventral margin
subterminal notch
subterminal margin
terminal margin
posterior tip
caudal fin
lower origin
lower (ventral) lobe
dorsal lobe
apex
inner margin
fin insertion
anterior margin
fin origin
posterior
margin
free
rear
tip
pectoral fin
base
excurrent
aperture
nostril
posterior
nasal flap
anterior
nasal flap
incurrent
aperture
labial furrow
mouth corner
labial
fold
upper eyelid
secondary
lower
eyelid
nictitating
lower
eyelid
subocular
pocket
eyes
notch
spine
anterior
margin
fin origin
apex
dorsal fin
free
rear
tip
insertion
base
posterior
margin
height
internasal
distance
preoral
length
mouth
width
head (ventral view)
LIST OF ORDERS, FAMILIES AND SPECIES
OCCURRING IN THE AREA
A question mark (?) before the scientific name indicates that presence in the area needs confirmation.
Order HEXANCHIFORMES
Family HEXANCHIDAE
Heptranchias perlo
Hexanchus griseus
Hexanchus nakamurai
Order SQUALIFORMES
Family ECHINORHINIDAE
Echinorhinus brucus
Family SQUALIDAE
Squalus acanthias
Squalus blainvillei
?Squalus megalops
Family CENTROPHORIDAE
Centrophorus granulosus
?Centrophorus uyato
Family ETMOPTERIDAE
Etmopterus spinax
Family SOMNIOSIDAE
Centroscymnus coelolepis
Somniosus (Rhinoscymnus)rostratus
Family OXYNOTIDAE
Oxynotus centrina
Family DALATIIDAE
Dalatias licha
Order SQUATINIFORMES
Family SQUATINIDAE
Squatina aculeata
Squatina oculata
Squatina squatina
Order LAMNIFORMES
Family ODONTASPIDIDAE
Carcharias taurus
Odontaspis ferox
Family ALOPIIDAE
Alopias superciliosus
Alopias vulpinus
Family CETORHINIDAE
Cetorhinus maximus
Family LAMNIDAE
Carcharodon carcharias
Isurus oxyrinchus
?Isurus paucus
Lamna nasus
Order CARCHARHINIFORMES
Family SCYLIORHINIDAE
Galeus atlanticus
Galeus melastomus
Scyliorhinus canicula
Scyliorhinus stellaris
Family TRIAKIDAE
Galeorhinus galeus
Mustelus asterias
Mustelus mustelus
Mustelus punctulatus
Family CARCHARHINIDAE
Carcharhinus altimus
Carcharhinus brachyurus
Carcharhinus brevipinna
Carcharhinus falciformis
Carcharhinus limbatus
?Carcharhinus melanopterus
Carcharhinus obscurus
Carcharhinus plumbeus
?Galeocerdo cuvier
Prionace glauca
Rhizoprionodon acutus
Family SPHYRNIDAE
Sphyrna (Mesozygaena)tudes
Sphyrna (Sphyrna)lewini
Sphyrna (Sphyrna)mokarran
Sphyrna (Sphyrna)zygaena
18 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
GUIDE TO THE ORDERS AND FAMILIES OF SHARKS
OCCURRING IN THE AREA
HEXANCHIDAE Page 24
Cow sharks
Three species in the Mediterranean. Mostly
demersal, from shallow depths to 1 800 m.
Size to 480 cm TL.
ECHINORHINIDAE Page 25
Bramble sharks
One genus with one species. Demersal
sluggish sharks, mostly in cold and deep
waters to at least 100 m; occasionally found in
the upper shelf. Size to 270 cm TL.
SQUALIDAE Page 26
Dogfishes
One genera and two species, plus one
dubious. Demersal and pelagic to depths of
over 2 000 m. Size to 160 cm TL.
Sharks - Guide to the Orders and Families Occurring in the Area 19
Order HEXANCHIFORMES – Cow sharks
Six or seven pairs of gill slits; single dorsal fin without spines; anal fin present; eyes without nictitating
eyelid; spiracle present but small.
6–7 gill slits
single dorsal fin
distinct lower lobe
caudal fin
notched
Order SQUALIFORMES – D