ANNALES · Ser. hist. nat. · 29 · 2019 · 2
205
received: 2019-10-17                  DOI 10.19233/ASHN.2019.20
CAPTURE OF A RARE ENDANGERED SPECIES LEAFSCALE GULPER SHARK 
CENTROPHORUS SQUAMOSUS (CHONDRICHTHYES: SQUALIDAE) FROM 
THE COAST OF SENEGAL (EASTERN TROPICAL ATLANTIC)
Youssouph DIATTA & Almamy DIABY
Laboratoire de Biologie marine, Institut fondamental d’Afrique noire, (IFAN Ch. A. Diop), Université Cheikh Anta Diop de Dakar, BP 
206, Dakar, Senegal
Sihem RAFRAFI-NOUIRA
Unité de Recherches Exploitation des Milieux aquatiques, Institut Supérieur de Pêche et d’Aquaculture de Bizerte, Université de 
Carthage, BP 15, 7080 Menzel Jemil, Tunisia
Christian CAPAPÉ 
Laboratoire d’Ichtyologie, case 104, Université de Montpellier, 34 095 Montpellier cedex 5, France
e-mail: capape@univ-montp2.fr 
ABSTRACT
This note presents a recent record of a specimen of rare shark leafscale gulper shark Centrophorus squamosus. A 
description of the specimen is given, including morphometric measurements, total body weight and dental formulae. 
Comments on this unusual and rare capture off the Senegalese coast are also provided.
Keywords: Squalidae, distribution, conditions of capture, dental formula, coast of Senegal
CATTURA DI UNA SPECIE RARA E MINACCIATA, IL CENTROFORO SQUAME A FOGLIA 
CENTROPHORUS SQUAMOSUS (CHONDRICHTHYES: SQUALIDAE), AL LARGO DELLA 
COSTA DEL SENEGAL (ATLANTICO TROPICALE ORIENTALE)
SINTESI
Questa nota tratta una cattura recente di un esemplare di una rara specie di squali, il centroforo squame a 
foglia, Centrophorus squamosus. Viene fornita una descrizione dell’esemplare, comprese le misurazioni morfo-
metriche, il peso corporeo totale e le formule dentali. Gli autori inoltre discutono questa insolita e rara cattura 
al largo della costa senegalese.
Parole chiave: Squalidae, distribuzione, condizioni di cattura, formula dentale, costa senegalese
ANNALES · Ser. hist. nat. · 29 · 2019 · 2
206
Youssouph DIATTA et al.: CAPTURE OF A RARE ENDANGERED SPECIES LEAFSCALE GULPER SHARK CENTROPHORUS SQUAMOSUS ..., 205–210
INTRODUCTION
The leafscale gulper shark Centrophorus squamosus 
(Bonnaterre, 1788) is widely distributed on both sides 
of the Indian Ocean and off the western Pacific coast 
(White, 2003). The species is known along the eastern 
Atlantic shore from Iceland and the Atlantic Slope to the 
Canary Islands, Senegal, the Faeroes, Madeira and the 
Azores (White, 2003). Southward, C. squamosus occurs 
from Senegal, Gulf of Guinea to the west coast of South 
Africa (McEachran & Branstetter, 1984).
Previously, a total of 37 specimens Centrophorus 
squamosus had been examined, among them two large 
pregnant females carrying near term embryos (Cadenat 
& Blache, 1981). Since then, no specimen had been 
recorded although investigations were continuously and 
regularly conducted in the area (Capapé et al., 1994, 
2001; Diatta et al., 2009; Diatta, 2014). Such lack of 
captures suggested that the species probably no longer 
occurred or, at least, had disappeared from the coast 
of Senegal. Conversely, in the wake of collaboration 
with experienced local fishermen, we were informed 
that a specimen of Centrophorus squamosus had been 
captured and delivered to our laboratory for a thorough 
examination. This new record forms the subject of the 
present paper, which includes a description of the speci-
men and some comments about the rare occurrence of 
this species in Senegalese waters.
MATERIAL AND METHODS
On 19 February 2015, a specimen of Centrophorus 
squamosus was caught by means of gill nets on rocky 
bottoms at a depth of 150-250 m, off the fishing site of 
Ouakam, located 5 km north of Dakar, in the Cape Verde 
Peninsula (17°36’04.20’’ W  and 14°40’39.31’’ N) (Fig. 1).
The specimen was measured to the nearest mil-
limetre and weighed to the nearest gram. Morphological 
measurements and teeth counts on upper and lower 
jaws were carried out following Compagno (1984) and 
are summarized in Table 1. The specimen was fixed in 
buffered formaldehyde and deposited in the Ichthyologi-
cal Collection of the Institute Cheikh Anta Diop of Dakar 
(Senegal), under catalogue number IFAN Centr-squa 01.
RESULTS AND DISCUSSION
The studied specimen was a female, measuring 
1300 mm in total length and weighing 11,200 g in total 
body weight (TWB). It was identified as a Centrophorus 
squamosus based on a combination of main morphologi-
cal characters, such as: snout slightly long, broadly para-
bolic; no anal fin; both dorsal fins with large spines; first 
dorsal lower than the second dorsal, the latter moderately 
large, as high as or higher than first dorsal fin; free rear 
tips of pectoral fins, margins slightly extended (Fig. 2A); 
upper teeth erect at the centre third, but not symmetrical; 
lower teeth with serrated edges (Fig. 2B); denticles on the 
sides of the body closely-set and overlapping, leaf-like, 
with a high median ridge and a small lateral ridge on 
each side (Fig. 2C); colour uniformly dark grey.
The general morphology, the shapes of teeth and 
denticles, the morphometric measurements and the 
dental formula are in total agreement with Bass et al. 
(1976), Cadenat & Blache (1981), Compagno (1984) and 
McEachran & Branstetter (1984), confirming the identi-
fication of this shark, which constitutes the latest known 
record of C. squamosus from the coast of Senegal.
Species of the genus Centrophorus Müller & Henle, 
1817 are generally targeted by fishermen, especially 
for the extraction of liver oil, which is highly appreci-
ated in cosmetology and pharmacology (Diatta, 2014). 
Nowadays, only specimens of gulper shark C. granulo-
sus (Bloch & Schneider, 1801) are commonly captured 
Fig. 1: Map of the coast of Senegal indicating (black 
star) the capture site of the specimen of Centrophorus 
squamosus (ref. IFAN Centr-squa 01), redrawn from 
Diatta et al. (2013).
Sl. 1: Zemljevid senegalske obale z označbo lokalitete, 
kjer je bil primerek (ref. IFAN Centr-squa 01) vrste 
Centrophorus squamosus ujet. Prirejeno po Diatta s sod. 
(2013).
ANNALES · Ser. hist. nat. · 29 · 2019 · 2
207
Youssouph DIATTA et al.: CAPTURE OF A RARE ENDANGERED SPECIES LEAFSCALE GULPER SHARK CENTROPHORUS SQUAMOSUS ..., 205–210
Reference IFAN Centr-squa 01
Sex Female
Total body weight (g) 11200
Measurements mm % TL
Total length 1300 100.0
Fork length 1140 87.7
Precaudal length 1030 79.2
Pre-first dorsal length 43 3.3
Pre-second dorsal length 830 63.8
Head length 290 22.3
Prebranchial length 220 16.9
Prespiracular length 150 11.5
Preorbital length 60 4.6
Prepectoral length 253 19.5
Prepelvic length 780 60.0
Snout-vent lenght 880 67.7
Interdorsal space 310 23.8
Dorsal caudal space 81 6.2
Pectoral-pelvic space 500 38.5
Pelvic-caudal space 700 53.8
Vent-caudal length 460 35.4
Eye length 50 3.8
Eye height 30 2.3
Prenarinal length 45 3.5
Intergill length 60 4.6
First gill slit height 35 2.7
Fifth gill slit height 42 3.2
Pectoral anterior margin 141 10.8
Pectoral inner margin 88 6.8
Pectoral posterior margin 103 7.9
Pectoral height 152 11.7
Dorsal caudal margin 210 16.2
Terminal caudal lobe 122 9.4
Subterminal caudal margin 27 2.1
Preventral caudal margin 168 12.9
First dorsal anterior margin 190 14.6
First dorsal height 80 6.2
First dorsal posterior margin 135 10.4
First dorsal base 205 15.8
First dorsal length 280 21.5
First dorsal inner margin 90 6.9
Second dorsal anterior 
margin
130 10.0
Second dorsal height 42 3.2
Second dorsal posterior 
margin
103 7.9
Second dorsal inner margin 35 2.7
Second dorsal base 125 9.6
Head height 180 13.8
Trunk height 210 16.2
Abdomen height 190 14.6
Tail height 110 8.5
Caudal peduncle height 55 4.2
Pelvic midpoint-second 
dorsal insertion
230 17.7
First dorsal midpoint- pelvic 
origin
270 20.8
Second dorsal insertion-
pelvic insertion
140 10.8
Pelvic posterior margin 
length
80 6.2
Pelvic inner margin length 65 5.0
Pelvic base 87 6.7
Pelvic length 145 11.2
Pelvic anterior margin 130 10.0
Pelvic height 103 7.9
Interorbital space 102 7.8
Head width 240 18.5
Trunk width 300 23.1
Abdomen width 350 26.9
Caudal peduncle width 30 2.3
Mouth length 50 3.8
Mouth width 190 14.6
Internarial space 40 3.1
Dental formulae 16-1-16/13-1-13
Tab 1: Morphometric measurements (in mm and as % TL), total body weight (in gram) and dental formulae 
recorded in the specimen of Centrophorus squamosus (ref. IFAN Centr-squa 01).
Tab. 1: Morfometrične meritve (v mm in kot % dolžine telesa), celotna telesna teža (v gramih) in zobna formula 
primerka vrste Centrophorus squamosus (ref. IFAN Centr-squa 01).
ANNALES · Ser. hist. nat. · 29 · 2019 · 2
208
Youssouph DIATTA et al.: CAPTURE OF A RARE ENDANGERED SPECIES LEAFSCALE GULPER SHARK CENTROPHORUS SQUAMOSUS ..., 205–210
throughout the Senegalese coast (Diatta et al., 2009). 
Some morphological differences allow distinguishing C. 
squamosus from C. granulosus. C. squamosus displays 
free rear tips of the pectoral fins, broadly angular and 
not reaching past the first dorsal spine, denticles in 
adults exhibit multiple lateral cusps. In C. granulosus, 
the free rear tips of the pectoral fins extend into nar-
row angular lobes, which reach past the first dorsal 
fin origin, and denticles in adults are without cusps, 
broadly rounded posteriorly. Since local fishermen are 
familiar with captures of C. granulosus in the area, they 
were able to distinguish this species from C. squamosus, 
and therefore they could immediately bring it to the 
laboratory for a thorough examination. The rarity of the 
species in the area could also explain why these fisher-
men could express a positive, or at least neutral, attitude 
to this shark species. The specimen was captured over 
continental shelf grounds at a depth of 150 to 200 m, 
which is normally shallower than the depth ranges 
reported by McEachran & Branstetter (1984). According 
to McEachran & Branstetter (1984), C. squamosus is a 
benthic species inhabiting deeper areas, from 400 to 
1875 m, rarely below 1000 m. Although these areas are 
poorly exploited by fishermen, this does not fully ex-
plain the scarcity of the species in commercial catches. 
However, this isolated capture of a single specimen 
suggests that the species is not completely extinct in the 
area, but other records are needed to state if a viable 
population still occurs in the area or if the species is 
dangerously threatened.
ACKNOWLEDGMENTS
The authors wish to thank the fishermen from the 
fishing site of Ouakam, Senegal, who provided the 
specimen, and Dr Mafalda Freitas, Head of the Marine 
Biology Station of Funchal, Madeira Island who enabled 
us to identify the specimen as Centrophorus squamosus.
Fig. 2: A. General morphology of the specimen of Centrophorus squamosus (ref. IFAN Centr-squa 01) caught off the 
coast of Senegal, scale bar = 200 mm. B. Jaw of the same specimen showing upper and lower teeth, scale bar =100 
mm. C. Denticles removed from the dorsal surface of the same specimen, scale bar = 2 mm.
Sl. 2: A. Morfologija primerka vrste Centrophorus squamosus (ref. IFAN Centr-squa 01), ujetega ob obali Senegala, 
merilo = 200 mm. B. Čeljust z zgornjimi in spodnjimi zobmi, merilo =100 mm. C. Dentikli s hrbtne površine 
raziskanega primerka, merilo = 2 mm.  
ANNALES · Ser. hist. nat. · 29 · 2019 · 2
209
Youssouph DIATTA et al.: CAPTURE OF A RARE ENDANGERED SPECIES LEAFSCALE GULPER SHARK CENTROPHORUS SQUAMOSUS ..., 205–210
ULOV REDKEGA IN OGROŽENEGA MORSKEGA PSA CENTROPHORUS SQUAMOSUS 
(CHONDRICHTHYES: SQUALIDAE) IZ OBALNIH VODA SENEGALA 
(VZHODNI TROPSKI ATLANTIK)
Youssouph DIATTA & Almamy DIABY
Laboratoire de Biologie marine, Institut fondamental d’Afrique noire, (IFAN Ch. A. Diop), Université Cheikh Anta Diop de Dakar, BP 
206, Dakar, Senegal
Sihem RAFRAFI-NOUIRA
Unité de Recherches Exploitation des Milieux aquatiques, Institut Supérieur de Pêche et d’Aquaculture de Bizerte, Université de 
Carthage, BP 15, 7080 Menzel Jemil, Tunisia
Christian CAPAPÉ 
Laboratoire d’Ichtyologie, case 104, Université de Montpellier, 34 095 Montpellier cedex 5, France
e-mail: capape@univ-montp2.fr 
POVZETEK
Avtorji poročajo o ulovu primerka redkega morskega psa vrste Centrophorus squamosus. Navajajo opis vrste, 
morfometrične meritve, telesno težo in zobno formulo. Poleg tega razpravljajo o tem redkem in nenavadnem ulov 
ob obali Senegala. 
Ključne besede: Squalidae, razširjenost, podatki o ulovu, zobna formula, senegalska obala
ANNALES · Ser. hist. nat. · 29 · 2019 · 2
210
Youssouph DIATTA et al.: CAPTURE OF A RARE ENDANGERED SPECIES LEAFSCALE GULPER SHARK CENTROPHORUS SQUAMOSUS ..., 205–210
REFERENCES
Bass, A.J., J.D D’Aubrey & N. Kistnasamy (1976): 
Sharks of the east coast of southern Africa. Part V: The 
Families Oxynotidae, Chlamydoselachidae, Squalidae, 
Dalatiidae and Echinorhinidae. Invest. Rep. Oceanogr. 
Res. Inst. Durban, 45, 1-103.
Cadenat, J. & J. Blache (1981): Requins de Méditer-
ranée et d’Atlantique (plus particulièrement de la côte 
occidentale d’Afrique). Faune tropicale, ORSTOM, 21, 
1-330.
Capapé, C., M. Diop & M. N’Dao (1994): Observa-
tions sur dix-sept espèces de Sélaciens d’intérêt économ-
ique capturées dans la région marine de Dakar-Ouakam 
(Sénégal, Atlantique tropical oriental). Bull. Inst. fond. Afr. 
noire Cheikh Anta Diop, Dakar, 1994, sér. A, 47, 87-102.
Capapé, C., A. Gueye-Ndiaye, Y. Diatta, M. Diop & 
A.A. Seck (2001): Observations on six elasmobranch 
species recorded from off the coast of Senegal (eastern 
tropical Atlantic). Acta Adriat., 42(1), 89-102.
Compagno, L.J.V. (1984): FAO Species Catalogue, 
vol. 4, Sharks of the World. An annotated and illustrated 
catalogue of shark species known to date. FAO Fisher-
ies Synopsis, 125, vol. 4, part 1 (non carcharhinoids): 
viii+1–250 pp.
Diatta, Y., A. Gueye-Ndiaye, P. Ndiaye, S. Ndaw, C. 
Reynaud & C. Capapé (2009): Production of elasmo-
branch species off the coast of Senegal (eastern tropical 
Atlantic). Elasmovisor, 4, 22-25.
Diatta, Y., C. Reynaud & C. Capapé (2013): First 
case of albinism recorded in striped panray, Zanobatus 
schoenleinii (Chondrichthyes: Platyrhinidae) from the 
coast of Senegal (Eastern Tropical Atlantic). J. Ichthyol., 
53(11), 1007-1012.
Diatta, Y. (2014): Contribution to the knowledge of 
the bioecology of 26 elasmobranch species from the 
coast of Senegal Thesis, Dakar, University Cheikh Anta 
Diopof Dakar, Senegal, 402 pp.
Mc Eachran, J.D. & S. Branstetter (1984): Squalidae. 
In: P.J.P. Whitehead, M.L. Bauchot, J.C. Hureau., J. 
Nielsen J.& E. Tortonese. (Editors), pp. 128-147. Fishes 
of the North-western Atlantic and the Mediterranean, 
Vol I, UNESCO, Paris.
White, W.T. (2003): Centrophorus squamosus. The 
IUCN Red List of Threatened Species. https://www.
iucnredlist.org/species/41871/10581731. Accessed 
on 16 October 2019.