ANNALES · Ser. hist. nat. · 29 · 2019 · 2
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received: 2019-08-22                  DOI 10.19233/ASHN.2019.19
FOOD AND FEEDING HABITS OF MUSTELUS MUSTELUS (LINNAEUS, 
1758) (CHONDRICHTHYES: TRIAKIDAE) ALONG THE WESTERN 
COAST OF LIBYA
Akraam Faraj KARA, Mohamed AL HAJAJI & Hisham GHMATI
Biodiversity department - Marine biology research center, Tajura, Libya
Esmail A. SHAKMAN
Zoology department, Tripoli University, Libya
e-mail: shugmanism@yahoo.com
ABSTRACT
A total of 269 individuals of Mustelus mustelus (Linnaeus, 1758) were provided through monthly collection by 
fishermen from the western coast of Libya (South Mediterranean) between November 2015 and November 2016. 
Each specimen was measured, weighed and dissected to investigate the food and feeding habits. Teleosts were the 
most important prey of M. mustelus, especially in individuals bigger than 90 cm TL. Crustaceans were the second 
most important and abundant prey for small individuals, whereas other prey groups were of minor importance 
and probably constituted just incidentally ingested food. Diet composition showed little seasonal variation: teleosts 
were the most important prey taxon in all seasons, except in autumn. The results indicate that M. mustelus can be 
considered as an opportunistic predator feeding on a wide range of prey items. 
Keywords: Smooth-hound, Diet, South Mediterranean, Libya
ABITUDINI ALIMENTARI DI MUSTELUS MUSTELUS (LINNAEUS, 1758) 
(CHONDRICHTHYES: TRIAKIDAE) LUNGO LA COSTA OCCIDENTALE DELLA LIBIA
SINTESI
Un totale di 269 individui di Mustelus mustelus (Linnaeus, 1758) sono stati forniti attraverso la raccolta mensile 
dei pescatori della costa occidentale della Libia (Mediterraneo meridionale) tra novembre 2015 e novembre 2016. 
Ogni esemplare è stato misurato, pesato e dissezionato per indagare sul tipo di nutrimento e sulle abitudini alimen-
tari. La preda più importante di M. mustelus sono risultati i teleostei, specialmente negli individui di dimensioni 
superiori a 90 cm di lunghezza totale. I crostacei sono risultati la seconda preda più importante e abbondante per i 
piccoli individui, mentre gli altri gruppi di prede sono di minore importanza e probabilmente costituiscono solo cibo 
ingerito casualmente. L’analisi della composizione della dieta ha evidenziato piccole variazioni stagionali, e i taxon 
di teleostei sono risultati le prede più importanti in tutte le stagioni, tranne in autunno. Secondo i risultati dello studio 
M. mustelus può essere considerato un predatore opportunista che si nutre di una vasta gamma di prede.
Parole chiave: palombo, dieta, Mediterraneo meridionale, Libia
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INTRODUCTION
In the Mediterranean, Chondrichthyes species pres-
ently have lower commercial value than Osteoichthyes 
and shellfishes; they only represent about 0.78 % of 
total landings (Cavanagh & Gibson, 2007; Di Francesco, 
2010). But because of general fishing pressure, they 
have become vulnerable since they are often captured 
as bycatch by bottom trawl fleets (Ragnese et al., 
2013). According to the statistics of the Fisheries and 
Agriculture Organization of the United Nations (FAO), 
the Chondrichthyes stock declined in the Mediterranean 
between 1970 and 1985, with landings increasing from 
10,000 to 25,000 tonnes and then slowly decreasing 
back to 10,000 tonnes in 2000. Recently, Bradai et al., 
(2012) reported that landings declined to 7000 tonnes. 
The impact of fishing on Chondrichthyes stocks has be-
come an issue of global concern. Thus, working groups 
and experts, such as the IUCN Sharks Specialist Group, 
have developed an international action plan for the con-
servation and management of shark exploitation, as well 
as lists of species in the Annexes of the Convention on 
International Trade in Endangered Species of Wild Fauna 
and Flora (CITES) (Stevens et al., 2000). Moreover, there 
are many efforts at regional level, such as the Elasmo-
branchs studying group as one of the specific scientific 
groups in GFCM, and at sub-regional level, such as the 
Elasmobranchs working group within the MedSudMed 
project, with Libya as one of the main parties. 
M. mustelus, commonly called the smooth-hound, is 
often characterized as a slender shark with a long para-
bolic subangular snout, dorsolateral eyes, an angular 
mouth, pavement teeth with cusps usually obsolete or 
absent, and the second dorsal fin nearly as large as the 
first. The species has a grey-brown back and is white 
underneath (Compagno, 1984). It is widespread in the 
Eastern Atlantic Ocean from the British Isles to South Af-
rica and common in the Mediterranean Sea (Compagno, 
1984; Goosen & Smale, 1997). M. mustelus is more 
common than the other two species of the same genus 
(Serena et al., 2009). This demersal species inhabits 
water depths ranging from shallow to 350 m, but is most 
commonly found in shallow waters, in the depth range 
of 5 to 50 m, on sandy and clay bottoms (De Maddalena 
et al., 2001; Serena, 2005). The catches of this species 
registered a decline of 85% between 1994 and 2006, 
however, there have been no actions undertaken to 
protect or regulate the exploitation of the shark in its 
range areas (Serena et al., 2009). 
Diet composition, which can identify the type of 
food preferred by each species of fish, is considered as 
an indicator of the availability of food in the region. 
In the present study, it showed that in the Gulf of 
Gabes (southern Tunisia, central Mediterranean Sea) M. 
mustelus feeds mainly on crustaceans, fish and cepha-
lopods. Polychaetes, sipunculids and echinoderms are 
occasional preys, and there is no difference between 
the diets of males and females (Saidi et al., 2009). In 
a similar study conducted in Turkey in 2005, the diet 
of this species was heterogeneous and generalized, 
and it was found that crustaceans were consumed 
by most of the individuals (Filiz et al., 2009). Dietary 
studies for M. mustelus from the Adriatic Sea report that 
their diet mainly consists of crustaceans, teleosts and 
cephalopods (Jardas et al., 2007a). The diet composi-
tion analysis from South Africa (Smale & Compagno, 
1997) and Mauritania (Khallahi, 2004) showed similar 
conclusions. In addition, a study comparing the history 
of life and biology of two species of sympatric coastal 
shark, M. mustelus and M. palumbes, off southern Af-
rica was described (Smale & Compagno, 1997). Morte 
et al. (1997) studied the feeding habits of juveniles (total 
length below 75 cm) in the Gulf of Valencia. Another 
study was conducted to describe the diet composition 
of the common smooth-hound in the northern Adriatic, 
linking its feeding habits to the structure of the gastroin-
testinal parasite community and providing new insight 
into the life cycles of the identified parasites (Gračan et 
al., 2014). 
Studies of population dynamics, such as age and 
growth, along with basic information on distribution, 
abundance, feeding and reproduction are essential for 
biologists to understand and predict the trend of popula-
tion growth and the species’ response to fishing pres-
sure. There are no comprehensive studies that examine 
the biological aspects of Chondrichthyes in the Libyan 
coast. Therefore, the present study aims to provide new 
findings on the diet of M. mustelus along the western 
Libyan coast of the southern Mediterranean Sea. 
MATERIAL AND METHODS
Samples of M. mustelus were collected monthly 
from fishermen on the western coast of Libya (Tripoli), 
between November 2015 and November 2016. The 
fish were transferred to the biodiversity laboratory of 
the Marine Biology Research Centre, where the samples 
were sorted and sexed, as well as measured for total 
length (TL) to the nearest centimetre and weighed for 
total weight (TW) to the nearest 10 g (Compagno, 1984). 
Each specimen was dissected with a ventral incision 
from the cloaca to the pectoral girdle in order to expose 
the body cavity. The stomach was removed, weighed 
and preserved in a 10% formaldehyde-seawater solu-
tion. 
Stomach contents were examined as soon as pos-
sible after capture. Preys were identified to the lowest 
possible taxon. The excess liquid was drained off and 
the remaining mass of wet prey was determined to the 
nearest 0.1 g. The importance of prey was evaluated 
using the frequency of occurrence percentage (F %), 
percentage by number (% N) and percentage by wet 
mass (M %) (Pinkas et al., 1971; Cortes, 1997; Morato 
et al., 2003). The index of relative importance (IRI) fol-
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lowing Pinkas et al. (1971) and modified by Hacunda 
(1981) was used: IRI = % F x (% N + % W). This index, 
which integrates the three previous percentages, allows 
a much more accurate interpretation of the diet by 
minimizing the skews caused by each of these percent-
ages. The contribution of each prey in the diet was also 
estimated with the Index of Relative Importance (IRI) 
and its standardized value (% IRI) (Pinkas et al., 1971; 
Cortés 1997). For assessing the diet in relation to shark 
size, the specimens were grouped in four size classes 
according to ontogenetic development: I = newborns, II 
= juveniles, III = subadults and IV = adults.
RESULTS
A total of 269 stomachs of M. mustelus were exam-
ined, 91 of them were empty (25.27 %). The diet of the 
specimens consisted of five major systematic groups: 
teleosts, crustaceans, cephalopods, nemerteans and 
polychaetes. Teleosts were the most important prey, 
constituting 50.29 % of the total IRI, followed by crusta-
ceans (% IRI = 33.76) (Tab. 1). The relative importance 
of cephalopods, nemerteans and polychaetes was com-
paratively low. 
Diet changes according to ontogenetic development
The diets were calculated and expressed as (% IRI in 
Fig. 1 according to the ontogenetic development of M. 
mustelus specimens from the southern Mediterranean. 
Newborns
In the size class of newborns, crustaceans were the 
most important prey group (% IRI = 68.82), followed by 
teleosts (% IRI = 31.17).
Juveniles
Crustaceans were the most frequent prey (% IRI = 
46.65), followed by teleosts (% IRI = 38.22). Nemerte-
ans (% IRI = 13.78) were the third most important group, 
while cephalopods and polychaetes represented minor 
components of the diet.
Subadults
Teleosts were the dominant prey consumed (% IRI 
= 50.67), followed by crustaceans (% IRI = 28.45) and 
nemerteans (% IRI =19.45). Cephalopods (% IRI = 1.2) 
and polychaetes (% IRI = 0.04) were minor components 
in the diet.
Adults
Teleosts were the most important prey category (% 
IRI = 64.82), followed by crustaceans (% IRI =15.44) 
and cephalopods (% IRI = 12.16), while nemerteans 
represented a minor component (% IRI = 7.22).
Seasonal changes in diet
The diets of M. mustelus in the south Mediterranean 
were also calculated and expressed as % IRI according 
to seasonal changes (Fig. 2): in spring, teleosts were the 
dominant group (% IRI = 64.72), followed by crustaceans 
(% IRI = 15.66) and cephalopods (% IRI =15.17). In 
summer, teleosts were the most frequently captured prey 
category (% IRI = 53.75), followed by crustaceans (% IRI 
= 35.51), while nemerteans and cephalopods were only 
minor components in the diet. In autumn, crustaceans 
constituted the bulk of the diet (% IRI = 45.16), followed 
by teleosts (% IRI = 33.49) and nemerteans (% IRI = 
17.88). In winter, the relative importance of teleosts was 
Tab. 1: Major prey groups in the diet of M. mustelus 
from the western coast of Libya, by percentage number 
(% N), percentage weight (% W), frequency of occur-
rence (% F) and index of relative importance (% IRI).
Tab. 1: Najpomembnejše skupine plena v prehrani nav-
adnega morskega psa ob zahodni libijski obali. Legenda: 
delež števila plena (% N), delež mase plena (% W), 
frekvenca pojavljanja plena (% F) in indeks relativne 
pomembnosti plena (% IRI).
Prey group F % N % W % % IRI
Crustacea 57.62 42.74 14.83 33.76
Teleostei 59.47 30.82 52.26 50.29
Cephalopoda 13.50 5.44 16.46 3.00
Nemertea 34.94 19.94 16.29 12.88
Polychaeta 4.00 1.03 0.14 0.04
Fig. 1: Ontogenetic differences in the diet of M. muste-
lus from the western Libyan coast, expressed as % IRI.
Sl. 1: Ontogenetske razlike v prehrani navadnega mor-
skega psa ob zahodni libijski obali, izražene kot delež 
indeksa relativne pomembnosti plena.  
31,17
38,22
50,67
64,82
68,82
46,65
28,45
15,44
1,33
1,2
12,16
13,78 19,45
7,22
0
10
20
30
40
50
60
70
80
90
100
Newborn juvenile Subadult Adult
%
IR
I
Ontogenetic development
Teleostei Crustace Cephalopoda Nemertea Polycheta
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high (% IRI = 46.1), followed by crustaceans (% IRI = 
39.29), while nemerteans, cephalopods and polychaetes 
were minor components of the diet.
DISCUSSION
To understand the biological interactions of organ-
isms in their ecosystem it is necessary to know the 
relationships and differences related to their diets and 
food consumption (Lopez et al., 2009, 2012). Therefore, 
qualitative and quantitative analyses of feeding habits 
are very helpful in determining the level and abundance 
of prey eaten by each species in their habitats (Movillo 
& Bahamonde, 1971; Lopez et al., 2012). Moreover, 
quantitative studies using the same approach can 
estimate the predation and, consequently, reflect the 
important factors and dependencies in predator diets 
(Wootton, 1990; Abrams, 2000; Lopez et al., 2012). In 
this study, the results indicated that M. mustelus feeds on 
benthic and demersal preys, thus determining the feed-
ing behaviour of this species as demersal. There were no 
gender-specific differences or requirements in the diet 
of M. mustelus, which suggests that the foraging habits 
are similar in both sexes or that they encounter the same 
prey items in their environments. In addition, the study 
provided broad support that this species could be an op-
portunistic predator, feeding on a variety of prey items 
(Cappe, 1975; Morte et al., 1997; Smale & Compagno, 
1997; Costantini et al., 2000; Khallahi, 2004; Jardas 
et al., 2007a; Saidi et al., 2009). Its predatory habits 
indicate that M. mustelus feeds mainly on crabs, mantis 
shrimp, shrimp living on sandy and soft sedimentary bot-
tom, pelagic and benthopelagic teleosts, cephalopods, 
nemerteans, and polychaetes (Jardas, 1996; Tortonese, 
1956; Bini, 1967; Branstetter, 1986; King & Clark, 1984; 
Kamura & Hashimoto, 2004; Jardas et al., 2007b; Saidi 
et al., 2009; Filiz, 2009). There were, however, regional 
differences among prey species. The study showed 
that the diet consisted of five major systematic groups: 
crustaceans, teleosts, cephalopods, polychaetes and 
nemerteans. The identified preys belonged to 5 families 
of crustaceans, 5 families of teleosts and 2 families of 
cephalopods, in addition to a few unidentified spe-
cies. Teleosts were the most important prey group of 
M. mustelus on the western coast of Libya, constituting 
(% IRI = 50.29) of the total IRI, Additionally, they were 
the dominant prey by mass (W % = 52.26) and had the 
highest frequency (F % = 59.47). This is probably due to 
the abundance and diversity of fishes along this coast-
line (Tab. 1). Crustaceans were the second most frequent 
prey (% IRI = 33.76), all of them belonging to the order 
of decapods (shrimp, crabs, and Mantis shrimps, as well 
as members of the Penaeidae, Majidae, Portunida, Cara-
pidae and Squillidae, and some unidentified species).
The most consumed teleost species were mainly 
demersal, but there were also many pelagic species 
present (e.g., scombrids, clupeids). Thus, it can be de-
termined that the smooth-hound is the major predator of 
benthic teleosts as well as of some species of the same 
genus, particularly M. palumbes, M. henlei, M. canis, 
and M. antarcticus (Capapé, 1975; Compagno, 1984; 
Simpfendorfer et al., 2001). Most of the cephalopods 
were quantitatively important, for instance, members 
of the Sepiidae and Octopodidae (octopods were 
dominant by mass, and crustaceans were numerically 
important). This finding is in accordance with a study 
carried out by Saidi et al. (2009). While nemerteans 
represented 12.88 % of total IRI, cephalopods (octopods 
and sepia) stood at 3 % and polychaetes at 0.04 % of 
total IRI as minor components of the diet. In contrast, 
Smale and Compagno (1997), Costantini et al. (2000), 
Khallahi (2004), Jardas et al. (2007a), Saidi et al. (2009), 
Filiz (2009) recorded that this species feeds primarily on 
decapod crustaceans and to a lesser extent on teleosts. 
Size-related changes in the diet were considered; the 
data obtained showed that the diet composition of M. 
mustelus changes considerably with its growth (Morte 
et al., 1997; Smale & Compagno, 1997; Jardas et al., 
2007a; Saidi et al., 2009), and there is a slight increase 
in prey diversity with the increase of shark size. These 
changes may be related to altered environmental con-
ditions or to the changing energy requirements of the 
animal. 
The new-born and juvenile sharks under study fed 
mainly on crustaceans (% IRI = 68.82 and % IRI = 46.65, 
respectively), which are considered the most important 
prey group in these size classes. When the sharks in-
crease in length (70 ≤ TL <90 cm), they switch to teleosts 
and cephalopods. The share of crustaceans diminishes 
in importance, while the rates of teleosts, cephalopods 
(Octopus vulgaris and Sepia officinalis), nemerteans and 
polychaetes increase. The studied adult sharks mainly fed 
on teleosts (% IRI = 64.82), followed by cephalopods (% 
IRI = 12.16), while the relative importance of crustaceans 
Fig. 2: Seasonal changes in the diet of M. mustelus from 
the western Libyan coast expressed as % IRI.
Sl. 2: Sezonske razlike v prehrani navadnega morskega 
psa ob zahodni libijski obali, izražene kot delež indeksa 
relativne pomembnosti plena.
64,72
53,75
33,49
46,1
15,66 35,51
45,16
39,29
4,42
0,73
3,26
3,9
15,17 9,98 17,88 5,4
0,01 0,18
0,03
0
10
20
30
40
50
60
70
80
90
100
      Spring       Summer      Autumn       Winter
%
 IR
I
Season
Teleostei Crustacea Cephalopoda Nemertea Polycheta
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decreased with the increase of shark size, to a minimal 
level of % IRI = 15.44 in large specimens. However, the 
wide dietary diversity in larger specimens may reflect the 
ability of large individuals to use a wider range of habitat 
resources, also on the trophic level, due to their increased 
morphological adaptation. The high presence of teleosts 
in the stomachs of M. mustelus can be explained by 
the fact that the species is a relatively active and fast 
predator, agile in swimming and manoeuvring, moving 
from the bottom to the centre of the water column. The 
second factor to be perhaps considered is the mechanical 
damage incurred by bony fish during fishing operations 
involving trawling and use of other fishing tools. How-
ever, these species become vulnerable to larger predatory 
fishes in the surrounding medium, including M. mustelus. 
Cartilaginous fishes play an important role in protecting 
and cleaning the marine environment. 
Little seasonal variation in the diet of smooth-hound 
was noticed within the study area. Values of the index 
of relative importance suggested that teleosts dominate 
the diet composition in all seasons, except autumn. The 
highest percentage was measured in spring, (64.72 of 
the total % IRI), followed by summer (% IRI = 53.75). 
Increased teleost consumption during these months co-
incides with the spawning period in many fishes, which 
may be present in high densities. In autumn, crustaceans 
constituted the bulk of the diet (% IRI = 45.16) as the 
most frequent prey (F % = 85.9), followed by teleost (% 
IRI = 33.49) and nemerteans (% IRI = 17.88). Cephalo-
pods and polychaetes were present in stomach contents 
during all seasons but in smaller quantities. 
CONCLUSIONS
As this study has shown, the diet composition of M. 
mustelus inhabiting the western Libyan coast displays 
little seasonal variation: teleosts are the most important 
prey taxon in all seasons, except in autumn. The results 
indicate that M. mustelus can be considered as an op-
portunistic predator feeding on a wide range of prey 
items.
ACKNOWLEDGEMENTS
The authors express their thanks to the marine biol-
ogy research center to support this work. We also would 
like to thank the fishermen and fishermen’s union who 
have helped us in this work. 
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PREHRANJEVALNE NAVADE NAVADNEGA MORSKEGA PSA, MUSTELUS MUSTELUS 
(LINNAEUS, 1758) (CHONDRICHTHYES: TRIAKIDAE), 
VZDOLŽ ZAHODNE OBALE LIBIJE
Akraam Faraj KARA, Mohamed AL HAJAJI & Hisham GHMATI
Biodiversity department - Marine biology research center, Tajura, Libya
Esmail A. SHAKMAN
Zoology department, Tripoli University, Libya
e-mail: shugmanism@yahoo.com
POVZETEK
Med novembrom 2015 in 2016 so ribiči tekom mesečnega izlova ujeli 269 primerkov vrste Mustelus mustelus 
(Linnaeus, 1758) ob zahodni obali Libije (južno Sredozemsko morje). Vsak primerek so avtorji izmerili, stehtali in 
secirali, da bi ugotovili prehranjevalne navade. Najbolj pomembna skupina plena so bile ribe kostnice, še posebej pri 
primerkih navadnih morskih psov, ki so merili več kot 90 cm v dolžino telesa. Raki so bili druga najpomembnejša sku-
pina plena, še posebej pri manjših primerkih morskih psov, medtem ko so bile druge skupine plena manj pomembne 
in jih avtorji smatrajo kot slučajno ulovljen plen. Sestava prehrane kaže majhno sezonsko variabilnost: kostnice 
so bile najpomembnejši plen v vseh letnih časih, razen jeseni. Rezultati kažejo, da je M. mustelus oportunističen 
plenilec, ki se prehranjuje s širokim izborom plena. 
Ključne besede: navadni morski pes, prehrana, južno Sredozemsko morje, Libija
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