Introduction 13
CLASSIFICATION AND SYSTEMATIC ARRANGEMENT
Considering that the purpose of this document is to Class Chondrichthyes (cartilaginous fishes) provide a simple user-friendly guide for species identification, no reference will be found here to Subclass Holocephali (chimaeras) dichotomy keys for single species. It is important that the classification used in this guide be defined, Order Chimaeriformes (chimaera and silver sharks) as available literature is not always in agreement Subclass Elasmobranchii (sharks) with this presentation. The classification of this group is still under review as no consensus has Superorder Squalomorphi (squalomorph sharks) been found to reconcile different authors’ positions. Order Hexanchiformes (cow and frilled sharks) For more information and further specific details on the taxonomy and biology of cartilaginous fish Order Squaliformes (dogfish sharks) species, refer to Tortonese, 1956; Hureau and Monod 1979; Whitehead et al., 1984; Fischer et al., Order Squatiniformes (angel sharks) 1987; Fredj and Maurin, 1987; Compagno, 1988, 2005; Nelson, 1994; Shirai, 1996; Mould, 1998. The Order Pristiophoriformes (sawsharks) * consultation of FishBase http://www.fishbase.org Order Rajiformes (batoids) (Froese and Pauly, 2000) proved very useful. The most fundamental references are Compagno’s Superorder Galeomorphi (galeomorph sharks) catalogues issued in 1984 and his recent revision partially issued in 2001. Order Heterodontiformes (bullhead sharks) *
This guide follows the systematic organization Order Lamniformes (mackerel sharks) proposed by Compagno (1999, 2001) and the classification reflects a cladogram attempt where a Order Orectolobiformes (carpet sharks) * new concept of cladistic classification is used. For instance, the batoids are raised to the order Order Carcharhiniformes (ground sharks) (Rajiformes) belonging to the superorder of the The species inside the families and in the orders Squalomorphi, even if perhaps a more suitable are mentioned in alphabetical order according to name should be found to indicate both Rajiformes genus. For both orders and families, some and Squaliformes. At the same time the sawsharks summary descriptions with their most significant group is raised to the order Pristiophoriformes. So characteristics are included. The current status is the batoids have been diversely allocated with described for each single species on an individual respect to the previous taxonomic organizations. sheet where, in addition to a drawing, scientific However, even if this new phylogenetic name and more recent synonyms and significant classification is considered valid, for practical misidentifications in some important cases, other reasons sharks and batoids are described synthetic information is given, i.e. FAO common separately in the text. names, maximum size, habitat and biology, Taking only modern sharks into consideration, we methods of capture and, when available, can adopt the following, simplified classification exploitation and conservation status. Arrows are (the orders with no representatives in the superimposed on the drawings to indicate features Mediterranean Sea are indicated by an asterisk): that help in species identification. 14 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
When possible the type of spiral valve which – Raja rondeleti Bougis, 1959 is probably based on characterizes the intestine of cartilaginous fishes is an abnormal specimen of R. fullonica and shown. There are large variations in the anatomy of considered as Leucoraja cfr. fullonica.The the spiral valve, three different types have been taxonomic status of four specimens from French described: columnar spiral with funnels pointing and Italian coasts is doubtful (Séret, pers. comm.). either backward or forward; spiral ring valve and cylindrical (scroll) valve (Compagno, 1988; How to use this guide Hamlett, 1999). Readers are advised to follow these simple steps in order to successfully identify any sharks, batoids Local names are not included considering the large and chimaera found in the region. First, refer to the number of names used in the various picture key of shark-like and batoid fishes then Mediterranean countries. Feeding behaviour is read carefully through the description of key indicated only in some cases. Remarks are characters listed under each order and family. Use indicated for species whose taxonomic status or the illustrations of the families under each order or presences are dubious. suborder only as a secondary aid in making certain For the purpose of this guide, a number of dubious that the right order or suborder has been found. species have not been considered as valid for the Then proceed to narrow down the family of the Mediterranean: specimen using the illustration for each family and key characters annotated in each illustration; make – Carcharhinus leucas (Valenciennes, 1841) is a use of the size data included for each family. Once doubtful species; it is neither recognized by the family has been identified, move to the Compagno nor in this guide. corresponding pages where the species for that family are illustrated. These illustrations and the – Carcharhinus longimanus (Poey, 1865) is a key characters indicated should allow proper doubtful species; it is considered “probable” by identification for all sharks and batoids known for Compagno but is not included in this guide. the area.
– Rhinobatos halavi (Forsskål, 1775) was recorded In a few cases the considered area is wider than by Tortonese (1951a) from the Egyptian part of the the Mediterranean basin; it refers to the CLOFNAM Mediterranean Sea but Ben-Tuvia (1966) noted area (Hureau and Monod, 1979): Mediterranean that no specimens of this species were available to and northeastern Atlantic between 30° and 80° of confirm its presence in this sea. Latitude north, -30° and +60° of Longitude, Azores and Madeira Islands included (Whitehead et al., – Torpedo alexandrinsis Mazhar, 1982 and 1984). Torpedo fuscomaculata Peters, 1855 are not Although (Linnaeus, 1758) is considered a valid species. Torpedo alexandrinsis Chimaera monstrosa represented in the sharks cladogram as indicated is known by only five syntypes mentioned in the by Compagno, 2001, the species account is original paper from Alexandria (Egypt) (Séret, inserted at the end of the guide to follow the pers. comm.); therefore its taxonomic status is taxonomic sense. Eventhough there is only one doubtful. The second species recorded Torpedo chimaera species in the Mediterranean Sea, the only once in Alexandria (Egypt) needs to be author thought it was important to illustrate the verified and is probably synonymous of Torpedo technical terms and description of this order. (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). Sharks and Chimaeras - Picture Key of Shark-like Fishes 15
S
*
E
S
S
M
E
*
E
S
R
S
S
M
E
M
E
O
E
R
S
R
M
F
M
M
I
E
O
R
O
R
F
R
N
F
M
O
I
I
I
O
O
L
F
R
H
R
I
F
F
A
I
O
I
R
H
O
U
F
N
B
A
I
C
I H
Q
O
H
T
N P
N
S
S
L
A
C (from Compagno, 2001)
A
E O
M
I
O
U
R
X
A
M
T
T
A
Q E L
R
S
C
I
C S
H
O
E
R
F
R
I
P
J
O
A
R
*
S
E
S
M
E
R
M
O
F R
I
O
T
F
N
I
O R
E
D
A
O
M
R
I
E
H
T
C
E
H
e
g
e
l
k
e
n
k
i
a
i
i
l v
r
l
r
l
-
i
-
a
,
p
w
s
v
w
e
s
a
d
a
l
v
,
s
i
l
s
s a l
,
a
t
e
d n
d
i i
v
o y
l t
e
l
e t
n
s
e
a
a
e
,
y
g
t
t
n
g
e
r
i
n
n
n
t
i
i
o
g t
s
o
l
h
l
a
n e
l
t
s
i t
e
i
t
o
s
n t t
t
r
a
i e
c
u t
c
u
i
y
i
s
o
t
o
e
n
n c
r
n fins not attached to head
f i
s mouth terminal, pectoral
o
s
o attached to sides of head
o n
n mouth ventral, pectoral fins
t
n
o
r
f
d
s n
i
n
e
i
l
y
l
h
e
e
e
f
b
w
o
h
t
h
t
t
n
u
u
o
o
o
r
f
m
m
s
e
n
n
i
i
s
f
p
e
s l
n
a
e
i
n
s
k
i p
r
i
f
s
l
o
-
-
l
d n
y
a
i
f
s
a
e 1
r
r
k ,
o SHARKS AND CHIMAERAS
o
,
i
s
l
n
t
d
d
-
i
e
l
y
s
n
a
s e
r
l
t
l
,
n
n
t
t
i
s
i
a
a t
o
g f
l i
e
n
l
f
l
7
n
s
a
y
y a
r
s
l
r
d l
r
d
o r
i
o
o
o
e
g
t
6
b
d
S
i
b
d 5
E
2
e
H
M
S
I
e
t
h
F
t
n
n
e
n
E
i
i
s
f
K
e
d
I
l
r
)
e
a
L
p
t
-
m
n
n
n
a
a
K
i e
r
f
s
o
R
g
I
l
e
I
n
I
r
o
A
a
p
L
H
n
d
H
e
a
a
A
r
C
l
S
t
s
c H t
N
t
o i
i
F
l
P
l
A
n
a
s
s
O
E
t
R
e
l
l
r
l
l
o
C
i
Y
B
i
a
n
g
g
E
O
(
k O
1 s
L
7 K
i
M
r
O
e
E
S
o
t
t
H
s
A R
a
5
L
U
E
n
E
T
a
K
I
C
S
h
I
L t
i
E
-
P
w
H
K
s
S
R r
I
e
A
F
d
r
H
O
S 16 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
SHARKS TECHNICAL TERMS AND MEASUREMENTS
dorsal-fin spine (if present) st 1 dorsal fin nd 2 dorsal fin precaudal pit spiracle gill slits interdorsal space snout nostril keel
labial caudal peduncle subterminal furrows pelvic fin notch clasper pectoral fin (male sex organ) caudal fin
pectoral-fin length anal fin
head trunk tail
gill slits precaudal tail nostril caudal fin snout anal fin
trunk
preanal ridges
anal opening
pelvic fin pectoral fin (female, no claspers)
underside view
interdorsal-fin ridge
location of intestinal valve
total length (caudal fin depressed to body axis) Sharks - Technical Terms and Measurements 17
terminal lobe posterior tip
dorsal lobe dorsal margin terminal margin
upper origin subterminal margin
subterminal notch
upper postventral margin
postventral notch lower origin lower postventral margin preventral margin ventral tip lower (ventral) lobe
caudal fin
apex fin insertion inner margin free rear tip spine posterior margin base anterior margin fin origin height free posterior rear margin tip anterior margin base apex insertion fin origin dorsal fin pectoral fin
preoral length
internasal mouth distance width
head (ventral view) upper eyelid
labial furrow notch
incurrent nictitating aperture lower secondary eyelid lower anterior eyelid nasal flap
labial excurrent posterior fold aperture nasal flap subocular eyes pocket mouth corner nostril 18 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
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 CETORHINIDAE Family HEXANCHIDAE Cetorhinus maximus Heptranchias perlo Family LAMNIDAE Hexanchus griseus Carcharodon carcharias Hexanchus nakamurai Isurus oxyrinchus ? Isurus paucus Order SQUALIFORMES Lamna nasus Family ECHINORHINIDAE Echinorhinus brucus Order CARCHARHINIFORMES Family SQUALIDAE Family SCYLIORHINIDAE Squalus acanthias Galeus atlanticus Squalus blainvillei Galeus melastomus ? Squalus megalops Scyliorhinus canicula Family CENTROPHORIDAE Scyliorhinus stellaris Centrophorus granulosus Family TRIAKIDAE ? Centrophorus uyato Galeorhinus galeus Family ETMOPTERIDAE Mustelus asterias Etmopterus spinax Mustelus mustelus Family SOMNIOSIDAE Mustelus punctulatus Centroscymnus coelolepis Family CARCHARHINIDAE Somniosus (Rhinoscymnus) rostratus Carcharhinus altimus Family OXYNOTIDAE Carcharhinus brachyurus Oxynotus centrina Carcharhinus brevipinna Family DALATIIDAE Carcharhinus falciformis Dalatias licha Carcharhinus limbatus ? Carcharhinus melanopterus Order SQUATINIFORMES Carcharhinus obscurus Family SQUATINIDAE Carcharhinus plumbeus Squatina aculeata ? Galeocerdo cuvier Squatina oculata Prionace glauca Squatina squatina Rhizoprionodon acutus Family SPHYRNIDAE Order LAMNIFORMES Sphyrna (Mesozygaena) tudes Family ODONTASPIDIDAE Sphyrna (Sphyrna) lewini Carcharias taurus Sphyrna (Sphyrna) mokarran Odontaspis ferox Sphyrna (Sphyrna) zygaena Family ALOPIIDAE Alopias superciliosus Alopias vulpinus Sharks - Guide to the Orders and Families Occurring in the Area 19
GUIDE TO THE ORDERS AND FAMILIES OF SHARKS OCCURRING IN THE AREA
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.
HEXANCHIDAE Page 24 Cow sharks single dorsal fin
Three species in the Mediterranean. Mostly demersal, from shallow depths to 1 800 m. Size to 480 cm TL. caudal fin 6–7 gill slits notched distinct lower lobe
Order SQUALIFORMES – Dogfish sharks Five pairs of gill slits; two dorsal fins often with spines on the anterior margin; mouth extending behind front of eyes; no anal fin, caudal peduncle without precaudal pits. Seven families and nine valid species (occurrence of other two species to be confirmed).
ECHINORHINIDAE Page 25 1st dorsal-fin both dorsal fins skin denticles origin behind situated over Bramble sharks conspicuously large pelvic-fin origin pelvic fins 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.
1st and 2nd dorsal fins usually SQUALIDAE Page 26 with spines, no grooves Dogfishes
One genera and two species, plus one dubious. Demersal and pelagic to depths of over 2 000 m. Size to 160 cm TL.
1st dorsal-fin origin behind front of pectoral fin but before pelvic fin 20 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
CENTROPHORIDAE Page 27 dorsal-fin spines with Gulper sharks lateral grooves
Primarily demersal deep-water sharks from depths of 200 to at least 2 400 m. Size to 170 cm TL. Two species in the region, but one questionable.
a notch on caudal fin post ventral margin
ETMOPTERIDAE Page 28
spines on dorsal fins Lantern sharks with lateral grooves
Benthic on shelf and slopes, from depths of 70 to 2 000 m, mostly 200 m. Size to 45 cm TL. One species in the region.
caudal fin with pectoral fin with subterminal notch rounded edge
SOMNIOSIDAE Page 29 dorsal fins with or Sleeper sharks without spines
Benthic on slopes. Size up to 100 cm TL. Two genera and two species in the Mediterranean.
pectoral fin with inner with or without edge broadly rounded slight lateral ridges
OXYNOTIDAE Page 30
st 1 dorsal-fin origin near high, sail-shaped dorsal fins Rough sharks front of pectoral-fin origin Body very high and compressed, triangular in cross-section with longitudinal ridges, one species. Demersal from depths of 60 to 660 m. Size up to 150 cm TL.
dermal ridges between pelvic and pectoral-fins bases Sharks - Guide to the Orders and Families Occurring in the Area 21
DALATIIDAE Page 30 no spines on dorsal fins Kitefin sharks (Liche sharks)
Benthic to mesopelagic, primarily on slopes at depths of 300 to 600 m. Size to 180 cm TL. One species present in the area. pectoral fin with caudal fin with rounded edge subterminal notch
Order SQUATINIFORMES – Angel sharks Five pairs of gill slits; eyes dorsal; two spineless dorsal fins; pectoral fins greatly expanded along sides of head as a free triangular lobe; no anal fin. Only one family and three species in the Mediterranean.
SQUATINIDAE Page 31 enlarged pectoral fins anterior edge of pectoral fin not attached pelvic fins Angel sharks 2 spineless dorsal fins Three species occuring in the area. On continental slope and shelf between depths of 30 and 500 m. Size up to 190 cm TL.
mouth terminal anal fin absent 5 pairs of gill slits spiracle dorsal view ventral view of head
Order LAMNIFORMES – Mackerel sharks Five pairs of gill slits; two dorsal fins without spines; anal fin present; no movable nictitating eyelid; mouth strongly arched and extending behind front of eyes; ring-type intestinal valve. Four families reported in the Mediterranean. ring-type intestinal valve
ODONTASPIDIDAE Page 32
Sand tiger sharks no lateral keel
Two species in the Mediterranean area. Neritic, in shallow water down to around 200 m. Size to 320 cm TL.
all gill openings in front of pectoral fin caudal lower lobe much shorter than upper 22 Field Identification Guide to the Sharks and Rays of the Mediterranean and Black Sea
ALOPIIDAE Page 33 Thresher sharks precaudal pits present Two species reported in the Mediterranean. Oceanic and coastal, to depths of 500 m. Size to 610 cm TL. extremely elongate upper lobe of caudal fin
no keels on caudal peduncle
CETORHINIDAE Page 34
strong keels on Basking sharks caudal peduncle
Coastal and on the continental shelf. A filter feeder, gills slits elongated. Size to about 1 000 cm TL. One species in the area.
5 extremely long gill slits nearly encircling the head
LAMNIDAE Page 35
strong lateral keel Mackerel sharks and precaudal pits Three species present in the Mediterranean. The presence of a fourth species, Isurus paucus, in the area needs confirmation. Coastal and epipelagic, from the surface to depths of around 1 200 m. Size probably up to 700 cm TL. Gillrakers absent. gill slits long not encircling head upper and lower caudal lobe almost equal in length
Order CARCHARHINIFORMES – Ground sharks Five pairs of gill slits, gillrakers absent; two dorsal fins without spines; anal fin present; movable nictitating eyelid; mouth arched and extending behind anterior edge of eyes; intestinal valve of scroll or spiral type. Four families in the Mediterranean.
intestinal valve of intestinal valve of spiral type scroll type nictitating eyelid Sharks - Guide to the Orders and Families Occurring in the Area 23
base of 1st dorsal fin over SCYLIORHINIDAE Page 37 or behind pelvic-fin base precaudal pit absent Catsharks
Four species and two genera. Mostly demersal from shallow inshore waters to 700 m. Size to 120 cm TL. Spiral type intestinal valve.
base of 1st dorsal fin well TRIAKIDAE Page 39 ahead of pelvic-fin base
Hound sharks eyes horizontally precaudal pit absent oval Four species and two genera in the Mediterranean. From inshore shallow depth to around 400 m. Size to about 200 cm TL. Spiral Intestinal valve. ventral lobe from poorly to well developed
CARCHARHINIDAE Page 41 2nd dorsal fin much less than precaudal pit st Requiem sharks 2/3 height of 1 dorsal fin present
Nine species and three genera. Other two eyes round species and one genus are doubtful. The presence of Galeocerdo cuvier needs confirmation. Neritic and oceanic pelagic to depth of 600 m. Size to 400 cm TL.
SPHYRNIDAE Page 46
Hammerhead sharks 1st dorsal fin much larger than the 2nd and its base anterior to origin of pelvic fin Four species in the Mediterranean. Coastal and oceanic, from intertidal and surface waters to at least 260 m of depth. Size to 600 cm TL. Scroll type intestinal valve.
no precaudal keel
head laterally expanded in “hammer” form
underside of head