Egyptian Journal of Aquatic Biology & Fisheries

Zoology Department, Faculty of Science,

Ain Shams University, Cairo, Egypt.
ISSN 1110 – 6131
Vol. 28(3): 517 – 539 (2024)
[Link]

Ecology, Anatomy, Reproduction, and Diet of the Atlantic Horse Mackerel,

Trachurus trachurus: A Comprehensive Review

Hanae Nasri1, Rachid Sabbahi2, 3*, Souad Abdellaoui1, Khaoula Kasmi1, Abdelouadoud
Omari1, Khalil Azzaoui3, 4, Reda Melhaoui1, Abdelhafid Chafi1, Belkheir Hammouti3,
Khalid Chaabane1
1
Laboratory for Agricultural Productions Improvement, Biotechnology and Environment, Faculty of
Sciences, University Mohammed First, 60000 Oujda, Morocco
2
Research Team in Science and Technology, Higher School of Technology, Ibn Zohr University, 70000
Laayoune, Morocco
3
Euromed University of Fes, UEMF, Eco-Campus, 30030 Fez, Morocco
4
Laboratory of Organometallic, Molecular Materials and Environment, Faculty of Sciences, Sidi
Mohammed Ben Abdellah University, 30000 Fez, Morocco
*
Corresponding Author: [Link]@[Link]
ARTICLE INFO ABSTRACT
Article History: The Atlantic horse mackerel, Trachurus trachurus, is a semi-pelagic fish
Received: Dec. 7, 2023 species with a broad distribution in the temperate, tropical, and subtropical
Accepted: April 30, 2024 coastal waters of the Atlantic, Mediterranean and Indian Oceans. As a
Online: June 6, 2024 significant species for both commercial and recreational fisheries, it plays a
_______________ pivotal role in the marine food web. This review synthesized current
knowledge of the ecology, reproduction, and growth patterns of T.
Keywords:
Migration,
trachurus, emphasizing its critical biological attributes. Furthermore, it
Nutrition, examined the main threats and conservation challenges this species faces,
Spawning period, alongside the implications for fishery management. The review culminated
Trachurus trachurus with strategic recommendations for future research and monitoring
initiatives aimed at filling the existing knowledge gaps and enhancing
fishery’s sustainability.

INTRODUCTION

Pelagic fish make up the majority of the fish biomass found in pelagic ecosystems
(Fanelli et al., 2023). These species such as small fatty pelagic fish, also called blue fish,
(i.e. horse mackerel, sardines and mackerel) are abundant but hardly exploited (Eymard,
2003). These types of pelagic species live between the surface and the bottom of the
ocean. Small pelagic fish can also be considered continental shelf fish (Sutton, 2013).
They include several hundred species with common characteristics: a dark blue color on
the back and a silver color on the belly, which are supposed to protect them from
predators, an elongated shape, and an often-gregarious lifestyle. Atlantic horse mackerel
(Trachurus trachurus), anchovy (Engraulis encrasicolus), herring (Clupea harengus),
518 Nasri et al., 2024

round sardinella (Sardinella aurita), Sardine (Sardina pilchardus), and flat sardinella
(Sardinella maderensis) play important ecological roles in the Mediterranean Sea marine
ecosystem (Ouled-Cheikh et al., 2022). They also represent significant quantities of fish
catches in the globe (Alheit & Peck, 2019; Hunnam, 2021), and are also the most
coveted species by the feed milling industry for the manufacture of fish oil and fishmeal.
Pelagic fish species account for 25% of all landings worldwide, mostly through
anchovies, sardinellas, sardines, mackerels and herrings (FAO, 2018). Nevertheless,
these small pelagic fish stocks are particularly susceptible to variations in oceanic
climate, leading to significant variations in their abundance and distribution over decades
(Asiedu et al., 2021).
Small pelagic fish are essential species in the marine food web, whose continued
existence is vital to keep the ecosystem in balance. They can control the abundance of
zooplankton they consume (Peck et al., 2021). In fact, the strong fluctuations in their
massive biomasses linked to overexploitation by fishing or environmental variations can
affect ecosystem components both upstream and downstream. Downstream, this group
can exert a top-down control over zooplankton. Conversely, small pelagic fish can exert
upstream control over predators. It is through these ecosystem control roles that small
pelagic fish are crucial to the trophic dynamics of upwelling ecosystems (Chouvelon et
al., 2015).
In addition to key ecological characteristics, small pelagic fish are characterized
by: (i) a wide distribution and dynamics along the coast, in search of optimal conditions
of temperature and food availability; with a typical life cycle of three to four years
(Raybaud et al., 2017; Schickele et al., 2020); (ii) a distribution spreading further
offshore for round sardinella and horse mackerel and closer to the coast in shallow waters
(a depth of less than 50m) in the case of Bonga shade and flat sardinella (Deme et al.,
2019); (iii) a structure organizing juveniles, young fingerlings and adults along a depth
gradient. For example, horse mackerels concentrate their adults at greater depths, moving
away from the coast. Thus, the smallest individuals are generally the most accessible to
fisheries due to their proximity to the coast (Sutton, 2013); and (iv) a gregarious behavior
facilitating their detection and catches contributing to the variability of their abundance
(Kasumyan & Pavlov, 2018).
Among the small pelagic fish, T. trachurus, commonly called the common scad or
the European horse mackerel, is an economically important species that is heavily
consumed around the world (Abaunza et al., 2003; Costa et al., 2021). It is primarily
destined for exploitation and has an extensive presence in the Central East Atlantic and
Mediterranean fisheries. This species is caught for processing into oil and meal and has
been used for human consumption since the 1970s. In Europe, the fish individual is eaten
fresh in such countries as Spain and Portugal. However, nearly 90% of the horse
mackerel fished is exported, primarily frozen, to countries such as Japan and West Africa
 519 Trachurus trachurus: A Review

(Eymard, 2003). Small pelagic fish are key species in the marine food chain, and their
presence is essential for maintaining ecosystem balance. They can regulate the abundance
of the zooplankton they consume (Peck et al., 2021). In Morocco, with a coastline that
stretches over 3,500km long (500km on the Mediterranean and 3,000 km on the Atlantic
coast), the maritime fishing sector plays an important economic and social role in the
Moroccan economy. Horse mackerel is very common throughout the Mediterranean Sea
(El Achi et al., 2021). In Morocco, the latest report from the Department of Maritime
Fisheries indicates that the 2022 pelagic fish catches reached 1,347,813 tons. Of this,
44,320 tons were horse mackerel, which generated a turnover of 185,316,000 Dh in the
Atlantic Zone. In the Mediterranean Zone, horse mackerel catches were reported at 3,245
tons (DPM, 2022).
This work aimed to study the Atlantic horse mackerel based on its biology, its
dynamics, and its feeding behavior, with the objective of adopting a good fishing
management policy for this carangid. This review is the result of an in-depth
bibliographic analysis, which would allow us to establish a solid base of research and
knowledge of T. trachurus.
1. PRESENTATION OF THE CARANGIDAE FAMILY

The Carangidae family consists of a group of seawater bony fish belonging to the
order Perciformes, which includes the perch-like fishes. The Carangidae family is one of
the most diverse and abundant families in tropical and subtropical seas, comprising about
33 genera and 146 species, with a wide variety of shapes and colors. They are
poikilothermic animals, meaning that their body temperature varies with the ambient
temperature (Mukherjee et al., 2018). The first fossils of this family date back to the
early Tertiary era (Berg, 1958).
The body of the carangids is usually slightly compressed laterally, either naked or
covered with small and smooth scales that are often detached and scattered in fish traps.
The head is large and flattened, with a prominent ridge on the back of the skull. The
mouth is protractile, meaning that it can be extended forward. The eyes and the gills are
located on the sides of the head. The lateral line is a sensory organ that runs along the
body, usually straight and only curved at the front. Some species have bony plates or
scutes along the lateral line (Letaconnoux, 1951; Gherram, 2019).
Fins are the appendages that help a fish swim and balance. Carangids have two
dorsal fins, with the first one being short and tall, with spiny rays. On the other hand, the
second is longer and lower, with soft rays. The anal fin is like the second dorsal fin but
located on the opposite side of the body. The pectoral fins are positioned on the body
sides, near the gills. The pelvic fins are on the lower side of the body, near the anus. The
caudal fin is located at the end of the tail and can have different shapes depending on the
species. The fins can have different colors and patterns, which can help identify the
species.
520 Nasri et al., 2024

2. CHARACTERISTICS OF TRACHURUS GENUS

The genus Trachurus comprises 15 recognized species of fish (Karaiskou et al.,

2003). These species are distinguished from other carangids by having two lateral lines,
which are sensory organs that run along the body. In the Mediterranean Sea, three species
of Trachurus are found: T. trachurus, the Mediterranean scad or the Mediterranean horse
mackerel, T. mediterraneus (Steindachner, 1868), and the blue jack mackerel or the
painted comber, T. picturatus (Bowodich, 1825) (Fig. 1).
The most evident feature distinguishing all three species from one another comes
from the length of the accessory lateral line, which is a branch of the main lateral line
running along the dorsal side of the body. In T. trachurus, the accessory lateral line
almost reaches the end of the second dorsal fin; in T. mediterraneus, it terminates just
beneath the first dorsal fin, and in T. picturatus, it has an intermediate length (Zohra,
2011).

Fig. 1. Morphological characteristics of Trachurus genus (Lloris & Rucabado, 1998)

 521 Trachurus trachurus: A Review

3. MORPHOLOGICAL DESCRIPTION OF T. TRACHURUS

Trachurus trachurus is a fish species that has a fusiform body shape, meaning that
it is tapered at both ends. The body is pearly on the sides and darker on the back. The
eyes are large and have fatty eyelids that cover most of the eye. The nostrils are small and
narrow and located on either side of the head (Fig. 2). A small black spot is usually
visible at the tip of the operculum. The first branchial arch features gill rakers on the
upper and lower parts. Gill rakers are bony projections that help filter food from the
water. The mouth is tubular, long, oblique, and can be extended forward. The upper jaw
is broad and has a large bone called the maxilla. The lower jaw is prominent and
protrudes beyond the upper jaw. Teeth are tiny and arranged in a row on both jaws. These
features make this fish species a predator. It is mainly a saltwater fish, but it can also
tolerate brackish water. Moreover, it lives in warm waters (Quéro, 2003).
The Atlantic horse mackerel has 23 vertebrae, which are the bones that make up
the backbone. The vertebrae are adapted to aquatic life and support the fins and the lateral
line. The fins are the appendages that help the fish swim and balance. T. trachurus has
two dorsal fins, which are on top of the body. The first dorsal fin is short and tall, with
eight spiny rays. The second dorsal fin is longer and lower, with one spiny ray and
numerous soft rays. The soft rays are flexible and support the fin membrane. The second
dorsal fin extends from the middle of the back to almost the tail (Meunier & Ramzu,
2006). The pectoral fins are located on the sides of the body, close to the gills. They reach
and cover the lateral line, which is a sensory organ that runs along the body. Pelvic fins
are found on the lower side of the body, close to the anus. They are also called the ventral
fins. The anal fin is almost symmetrical to the second dorsal fin but is positioned on the
opposite side of the body. It has two small spines at the front, which are separated from
the rest of the fin. The caudal fin is at the end of the tail and is very forked. The caudal
peduncle is the slender, thin part of the body that connects the tail to the trunk. The
maximum weight of the fish is 2kg, and its size can reach 60 and 70cm in fork length, i.e.
the length between the tip of the snout and the end of the middle rays of the caudal fin.
Fork length is generally between 15 and 30cm (Eymard, 2003; Charef-Belifa, 2009).
The color of the fish varies depending on the age and the environment. The upper
part of the body and the top of the head are dark, almost black or grayish to greenish blue.
The lower part of the body and the belly are paler, whitish to silvery. The young fish,
unlike the adults, are silvery gray with green reflections (Rahmani, 2020).
4. INTERNAL ANATOMY

The internal structure of fish consists of a skeleton, a circulatory system, and a

digestive system. The skeleton is made up of cartilage or bone, and includes the skull, the
vertebrae, and the fins. The circulatory system is simple and consists of a heart, blood
522 Nasri et al., 2024

vessels, and blood. The digestive system comprises the mouth, pharynx, esophagus,
stomach, intestine, liver, spleen, and pancreas.
The mouth of T. trachurus is tubular, long, oblique, and can be extended forward.
The upper jaw is broad and has a large bone called the maxilla. The lower jaw is
prominent and protrudes above the upper jaw. The teeth are short and arranged in a single
row on both jaws. These features make this fish species a predator. The eyes are large and
have fatty eyelids that cover most of the eye. The nostrils are small and narrow, and
located on either side of the head. A small black spot is usually visible at the tip of the
gill cover. The gills are located on the sides of the head and help the fish breathe in water.
The first gill arch has gill rakers on both the upper and lower parts. The gill rakers are
bony projections that help filter food from the water.
The esophagus is a tubular structure that connects the mouth to the stomach. The
stomach is internally wrinkled in the longitudinal direction and can expand or contract
depending on the season and the reproductive state of the fish. During sexual maturity,
the stomach is almost empty and very small. During sexual rest, on the other hand, it is
capable of a great expansion. At the lower and anterior part of the stomach, there is a
pocket that contains a variable number of pyloric caeca. These caeca are long, thin-
walled, and joined together by a thin membrane called the mesentery. They help digest
and absorb food (Letaconnoux, 1951).
The intestine is a tube that connects the stomach to the anus. It is folded back on
itself twice forming a loop. The intestine absorbs nutrients and water from the food. The
liver is a small organ that is pressed against the stomach and the pyloric caeca. It
produces bile, which helps digest fats. The spleen is a dark red organ that is located
between the folds of the intestine and the pyloric caeca behind the liver. It helps filter
blood and fight infections. The pancreas is a diffuse organ that is hidden by fat tissue. It
covers the entire gastrointestinal system. Moreover, it produces enzymes and hormones
that help digest and regulate blood sugar (Letaconnoux, 1951).
Fish have two rather specific organs: the gills and the swim bladder. The gills are
used for respiration and exchange gases with the water. The swim bladder is a large organ
that occupies the entire abdominal cavity. It is filled with gas and helps the fish adjust its
buoyancy and depth. In general, the swim bladder is terminated by two horns that are
separated by the first haemal processes of the caudal trunk. The haemal processes are
bony projections that protect the blood vessels of the tail (Dorson et al., 2018).
The reproductive system consists of a pair of glands that are in the abdominal
cavity under the swim bladder in its posterior region (Dorson et al., 2018) (Fig. 2). They
are richly supplied with blood vessels and have the shape of two cords. Their size and
color vary with the age of the fish and its sexual maturity. The glands produce gametes,
which are the reproductive cells. The male and female reproductive glands, testes and
ovaries, produce sperm and eggs, respectively. The testes are milky white, and the ovaries
 523 Trachurus trachurus: A Review

are orange yellow. Usually, the testes and ovaries are symmetrical and elongated organs
that span the length of the general cavity from the spot behind the anus to the height of
the liver. They compress all the other organs, and the stomach is almost always
completely empty. When fish reach three years old, they can reproduce sexually. The
males and females have different reproductive seasons. The females are sexually active
year-round, while the males have a shorter reproductive season. The fish are
gonochoristic, meaning that they have separate sexes and oviparous, meaning that they
lay eggs. The eggs are fertilized externally in the water (Prolonge-Chevalier, 2007;
Zohra, 2011).

A B

Fig. 2. Reproductive system of Trachus trachurus showing: A. Male reproductive glands

(testes), and B. Female reproductive glands (ovaries)
5. GEOGRAPHIC DISTRIBUTION

All oceanic waters in temperate, moderate subtropical and tropical zones are
home to Trachurus species along their coasts (Leitão et al., 2018). T. trachurus is an
adaptable species that can live in different oceans and seas. In the Atlantic Ocean, this
species ranges from Iceland to Senegal in the eastern part and from Argentina to Canada
in the western part (Abaunza et al., 2008). It is also found in the Norwegian Sea, the
English Channel, and the North Sea (Bektas & Belduz, 2009). It inhabits the continental
shelf and the continental slope from 10 to 500m depth (Giacomo et al., 2018). In
contrast, in the Mediterranean Sea, this species is very popular and widespread and can
be present in the Sea of Marmara, and occasionally in the Black Sea (Gherram et al.,
2018). It is also present along the Algerian and Moroccan coasts of the Mediterranean
Sea (Eymard, 2003; Abattouy et al., 2014) (Fig. 3). T. trachurus is found in the eastern
and western parts from Japan to Australia and from California to Chile (Froese & Pauly,
2020). Moreover, in the Indian Ocean, T. trachurus is found in the western and southern
parts from South Africa to India and from Madagascar to Australia (Froese & Pauly,
2020).
524 Nasri et al., 2024

Fig. 3. Geographic distribution of Trachurus trachurus (Fishbase, 2019)

6. SPECIES HABITATS

Trachurus trachurus hunts in large schools in the water column (D’Elia et al.,
2014). It is generally present on sandy seabeds at depths of 100 to 200m but can
sometimes reach depths of 600m or more. It also lives in the pelagic zone and sometimes
near the surface, depending on environmental conditions (Sutton, 2013). Therefore, it
can be considered a semi-pelagic species, because it moves between the surface and the
bottom, or between different water layers (Quéro, 2003; Jardas et al., 2004). Its habitat
preferences are poorly understood, but they may be influenced by a variety of
hydrographic features such as temperature. In addition, it can adapt to brackish water
(Santic et al., 2002). Its activity is relatively reduced outside the warm season.
Moreover, it is a migratory fish species and an excellent swimmer. It migrates and
approaches the coasts in summer, which makes it more abundant and easier to catch. It
returns to the open sea in winter at depths of around 400m, where it is less accessible to
fisheries. Its fishing also decreases as the spawning period ends, as the schools become
more dispersed (Abaunza et al., 2003; Rumolo et al., 2017). Furthermore, it is a species
with pelagic behavior, whose larvae and juveniles often congregate under any floating
object, which may provide them with protection from predators. The larvae live at an
average depth of 20m between March and September (Michael, 2002). When they reach
4mm long, they have four to five spines on the outer row, and approximately five on the
inner row of the gill cover. This number increases as they grow. These larvae have many
dark pigment cells on the jaw, head, and throughout the abdomen (Russel, 1976).
 525 Trachurus trachurus: A Review

7. MIGRATION AND BATHYMETRY

The Atlantic horse mackerel is a highly migratory fish. It can travel long
distances depending on its needs, such as feeding, spawning, and predator avoidance.
The distribution of fishing catches and data obtained from surveys provide indications of
the migration routes of different groups (Gherram et al., 2018). Three types of
migration are known for this species:
- Nychthemeral migration (Motility)
This type of migration is part of the species' daily behavior, influenced by a day-
night cycle. Large schools of horse mackerel are found close to the bottom and in the
middle of the water during the day; while at night, they scatter and form a layer just
above the seabed (Macer, 1977). The migration of this fish species can be mainly
affected by the water temperature. This species generally occupies continental seas to a
depth of 200m. However, some specimens have been recorded at depths of 500m
(Rumolo et al., 2017).
- Seasonal migration
This type of migration is related to the reproductive cycle of the species, which
varies according to the geographic location and the water temperature. After spawning,
horse mackerels migrate north along the slope of the continental margin during June and
July. From July to September, they remain off the Norwegian coast and migrate to the
central North Sea in October and November. Subsequently, they begin a return migration
to the western area, passing through Scotland and the Shetlands (Gherram et al., 2018).
During the month of June, the North Sea horse mackerel migrate northeast from the
English Channel along the Dutch coast where they spawn in July, heading to the west
coast of Denmark in August. In the third and fourth quarters, horse mackerels are
abundant in the southern and southeastern parts of the North Sea; however in October,
they have been observed to cross the channel in western and southern directions (Mahe
et al., 2007). It is possible that the western and North Sea spawning groups mix in the
central North Sea during the third and fourth quarters, and in the Western Channel
mainly in the fourth quarter. Little information is available on the migratory behavior of
horse mackerel in the Bay of Biscay. The presence of T. trachurus was also observed
throughout the year along the Algerian coast as well as the Atlantic coasts of Portugal
and Spain (Gherram et al., 2018).
- Trophic migration
This type of migration is related to the species' feeding behavior that depends on
the availability and distribution of prey. T. trachurus lives near the bottom and also
throughout the water column. In spring, the fish disperses much more widely, migrating
northward from the southern Black Sea to feed and reproduce as water temperature rises
526 Nasri et al., 2024

(Polonsky, 1965). This species heads south in autumn at temperatures below 10°C. It
withdraws from feeding areas in the southern Norwegian and North Seas and migrates to
wintering grounds further south (Zohra, 2011). Moreover, it migrates to the English
Channel and along the continental margin in the Bay of Biscay and the Celtic Sea
(Macer, 1977). T. trachurus from the Romanian Black Sea coast overwinters in the Sea
of Marmara (Smith-Vaniz et al., 2015). The North Sea stock appears in April in the
southern North Sea, reaching the west coast in July and southern Norway in August.
Parts of the western stock can reach Trondheim Fjord in July- August (Iversen et al.,
2002). T. trachurus has been observed in the company of the lion's mane jellyfish,
Cyanea capillata L., and various other jellyfish. This recent discovery marks a new
association for this species.
8. ECOLOGICAL CHARACTERISTICS

- Nutrition
Figuring out how fish feed in their natural habitats is an essential step toward
understanding their biology and ecology. Their feeding habits reflect not only predatory
activities but also explain variations in their growth, foraging, and feeding behavior,
migrations, and even certain aspects of reproduction. In addition, data on food
composition is utilized to predict trophic levels, which is vital for managing fisheries and
quantifying the effect of fishing on ecosystems. Information on the place of fish in food
webs is also helpful in understanding how marine ecosystems function and how they are
influenced by natural or human interventions. T. trachurus is a voracious carnivorous
fish (benthopelagic). Its diet is based on pelagic prey (copepods and teleosts) and benthic
prey (primally crustaceans). It is a very active predator, moving from the bottom to the
surface, where it rises to hunt, particularly during the first part of the night, which is its
principal activity period. This was confirmed by several studies (Jardas et al., 2004;
Šantić et al., 2005; Bayhan & Sever, 2009; Villegas-Ríos, 2009; Bayhan et al., 2013;
Shawket et al., 2015; Koç & Erdoğan, 2019).
The food of T. trachurus is varied and includes four or five major groups or
taxonomic units of prey (crustaceans, mollusks, echinoderms and fish, teleosts, etc.). Its
diet includes a wide variety of fish, such as juvenile mugilid species, blue whiting
(Micromesistius poutassou), and S. pilchardus. The species also feeds on small
crustaceans, which constitute the most diverse group, such as euphausiids, mysidaceans,
amphipods, copepods, decapods, isopods, cumaceans, small swimming crustaceans,
brachyurans and macrourids. Additionally, it consumes molluscs (bivalves, scaphopods,
gastropods and cephalopods, squids), vertebrates (teleosts and teleost eggs), and
echinoderms. However, the food of T. trachurus is based primally on small swimming
crustaceans, and euphausiids constitute the dominant fraction of its diet. Several authors
have highlighted the diversity of its diet (Bayhan & Sever, 2009).
 527 Trachurus trachurus: A Review

On the other hand, the feeding strategy developed by this species varies
depending on its stage of growth. In fact, the size of the predator influences its food
composition. The species would tend to adapt its feeding habits according to its size and
therefore according to its age. Juveniles prefer euphausiids, while adults tend to be
piscivorous. Therefore, the food differs according to the individual's size, and they tend
to eat larger and larger prey as they grow (Koç & Erdoğan, 2019).
- Diet by length classes
The study of the diet shows that the principal preys of T. trachurus are
crustaceans (Koç & Erdoğan, 2019). In the first two classes (1 and 2), for a fish length
lower than 20cm, teleosts (anchovies, sardines) are the most favored prey group. The diet
does not change much with the seasons, and small animals with shells are the main food
for the smallest two classes of sizes. When they grow bigger (class 3), they prefer and
mostly eat young fish (with bones).

To sum up, the smallest group of sizes likes to eat small animals with shells; the
next group chooses small shrimp-like animals, and the largest group mainly hunts young
fish (with bones). This means that the food they eat depends on how big they are. We can
say that this fish has a very adaptable diet, and it can find food in any environment. This
is consistent with literature data showing that the frequency of feeding is higher in
smaller individuals, and that this is thought to be associated with prey size, as smaller
prey are digested faster than larger fish prey (Bayhan & Sever, 2009).

Research into the feeding biology of horse mackerel in Morocco reveals that
stomach content analysis indicates a diversified diet for T. trachurus. This species feeds
primarily on crustaceans, with Teleostei classified as complementary prey. A total of 21
prey categories were identified in the stomach contents. Furthermore, diet fluctuations
were found to be associated with environmental factors, including upwelling, temperature
variations, chlorophyll concentration levels, and dissolved oxygen levels (El Achi et al.,
2021; Maroua et al., 2023).
- Eating habits
After the spawning in summer, the fat content of T. trachurus becomes very low.
However, in August and September, its energy content increases rapidly due to intensive
feeding. The formation of opaque zones in the otoliths is therefore to be expected during
periods of intensive feeding, allowing rapid growth and increased calcification (Zohra,
2011). This fish stops feeding in part once water temperature goes under 10°C. When the
water temperature reaches 8- 9°C, it stops feeding and migrates for the winter. After
overwintering, the energy consumed by gonad development (in winter and spring) and
spawning will result in the formation of a transparent zone in the otoliths (Abaunza et
al., 2008). In the North Sea (except in July), T. trachurus feeds mostly on fish, whiting
being the most significant prey, followed by other gadids and herring. Further south,
528 Nasri et al., 2024

invertebrates make up the bulk of the fish's diet, including crabs, decapods, and other
crustaceans (Dahl & Kirkegaard, 1986).
However, Dahl and Kirkegaard (1986) reported a clear diurnal feeding pattern
in the eastern North Sea, but with more intense feeding in the morning than at night. A
change in food preferences with age has been demonstrated; small individuals (less than
20- 24cm) feed mainly on crustaceans, gobiids, and haddock, while larger specimens
prefer herring. In the English Channel, adult horse mackerels apparently feed 70% of the
time on crustaceans and only 17% of the time on fish; these proportions vary monthly
(Macer, 1977). However, T. trachurus is a planktivorous fish for which crustaceans
(euphausiids and copepods) are the main prey. Porumb (1979) reports a different diet
between the two sexes: females feed mainly on the Mediterranean sand eel,
Gymnammodytes cicerellus (Rafinesque) (benthic fish), while males feed on the sand
smelt, Atherina mochon pontica (Eichwald), and wandering polychaetes (Koç &
Erdoğan, 2019). Regarding samples from northwestern Spain, Cabo (1950) suggested
that juveniles are planktivorous, while adults are mainly piscivorous. In the southern Bay
of Biscay, T. trachurus shows seasonal differences: it feeds on crustaceans in spring,
while in autumn, fish over 30cm start to eat other fish (blue whiting, gobiids, anchovies),
which accounts for 45% of the food volume in this length range. A diurnal feeding
pattern has also been described, with feeding peaking around midday in spring (for fish
over 30cm) and at sunrise in autumn. In Portuguese waters, T. trachurus feeds mostly on
zooplankton, especially euphausiids and copepods. It is not until they are 20cm long that
fish become their main prey (Abaunza et al., 2003).
9. REPRODUCTION

- Reproductive cycle of teleosts

Fish reproduction is a complex process. It is regulated by hormones and depends
heavily on environmental conditions, in particular temperature and photoperiod that
affect the timing of gonad maturation, the development of secondary sexual features, and
reproductive behavior. These changes lead to a reproductive period during the year,
when environmental conditions are favorable for an optimal juvenile development
(Waldron & Kerstan, 2001).
The extreme diversity of fish is manifested in their modes of reproduction and
development. Most fish are single-sex, but about 10% of species are hermaphrodites,
meaning that they have both male and female reproductive organs (Prolonge-Chevalier,
2007).
Fish reproduce according to two methods of fertilization (Gaillard, 2006):
 External fertilization: Oviparous females release the eggs into the external
environment. They are then fertilized by males by the release of sperm. In this
 529 Trachurus trachurus: A Review

type of fertilization, there is no mating. It is the specific recognition that exists

between sperm and eggs that enables fertilization.
 Internal fertilization: Male sperm are deposited inside (or near) the female
reproductive system, and their union takes place within the female. Three main
modes of reproduction have been identified (Bruslé & Quignard, 2004):
o Oviparity: A few species of teleosts perform intra-ovarian self-
fertilization and lay eggs.
o Ovoviviparity: The embryo is only housed in a female genital cavity,
without any trophic relationship with the mother. This simple bond
implies an exclusive use of its own yolk reserves.
o Viviparity: The embryo receives, in the female genital tract, maternal
nourishment other than that supplied by its yolk sac.
The reproductive cycle entails a series of physiological and behavioral processes
influenced by a range of abiotic and biotic environmental factors. Understanding the
timing and duration of egg-laying is crucial for comprehending population dynamics.
Various indices have been employed to determine the breeding period. Rahmani (2020)
used the gonadosomatic ratio, histological analysis of female ovaries, monitoring of
sexual maturity phases, and egg quantity.
- Breeding, spawning period and sexual maturity
Seasonal gonadal changes (gametogenesis) in teleosts follow successive stages.
However, the rates of these changes vary greatly between species. Gametogenesis
involves both oogenesis (oocyte growth) and spermatogenesis (growth of male germ
cells) (Abaunza et al., 2003). Spermatogenesis is divided into three stages:
spermatocytogenesis, meiosis, and spermiogenesis. Three types of reproduction are
defined (Abaunza et al., 2003):
 Synchronous reproduction: a rare type in which all germ cells are at the same
stage. This type of reproduction is found in fish that spawn once in their lifetime
and then die (semelparous species).
 Group-synchronous reproduction: in this type of reproduction, there are two
populations of germ cells that develop at the same time; the first is at an early
stage, and the second is at a late stage.
 Asynchronous reproduction: this type of reproduction is the most common. All
stages occur in the gonads. T. trachurus has this type of reproduction.
Trachurus trachurus spawning period varies from one region to another; this
phenomenon is affected by certain climatic factors like temperature and physicochemical
parameters (e.g., salinity). Indeed, a slight temperature rise can induce spawning
(Lévêque, 2006). This species has a reproductive strategy characterized by a gonochoric
and oviparous state, with an extended spawning season (Gordo et al., 2008). T.
trachurus spawning period involves a migration to the coast in spring and to the open sea
in autumn. Atlantic horse mackerel has a high reproduction rate (Murua & Saborido-
530 Nasri et al., 2024

Rey, 2003). Spherical and smooth eggs are released in water. When hatched, larvae
barely measure 5mm. Females spawn their eggs at temperatures ranging from 18 to
21°C. Eggs are spawned in batches and they are pelagic, their diameter varies between
0.9 and 1.1mm (Artüz, 2000). Sexual maturity is attained at two years for females and
around three years for males. The sexes are separated, and fecundation takes place
outside the body. Males reach maturity at a shorter length compared with females and
seem to be sexually active almost all the time, whereas females are only sexually active
in spring and summer, with a peak in June- July (Mahe et al., 2007). Oviposition is
preceded by a fasting period that accompanies sexual maturity (swelling of the ovaries
and testes) (Mouneimne, 1978).
The eggs are pelagic, spherical in shape, with a fat droplet. The diameter of the
egg varies from 0.92 to 1.02mm, while that of the fat droplet varies from 0.22 to
0.26mm. The envelope is very thin, transparent, and cohesive; the yolk is vascular-
lobular. The body length of newly hatched pre-larvae averages 2mm. The large ovoid
yolk sac protrudes from the head, and the fat droplet is located near its anterior border
(Key, 2015). The larvae, soon after hatching, have a much reduced yolk, with the droplet
being carried forward, and the intestine extending far back, while the fins are large and
pigmented (Letaconnoux, 1951; Artüz, 2000).
Trachurus trachurus grows rapidly during its first year, and then progressively
slows down with age (Letaconnoux, 1951; Nasri et al., 2021). Its lifespan is very long
(around 30 years), with a maximum length reaching 70cm (Overko & Mylnikov, 1979).
Determining the age of fish is essential for the use of several fishery models to study
growth. This is generally done using bone pieces, vertebrae, head bones, or more
classically, scales, and sagittate otoliths (Wright et al., 2002).
Research on T. trachurus reproduction in Morocco is limited. A thesis by Nasri
(2022) revealed two distinct Mediterranean spawning seasons: late winter to early spring
(February- April) and summer (July- September). Concurrent macroscopic and
histological studies have delineated varied reproductive timelines across the
Mediterranean:
• Bay of Béni Saf: February to July, with a peak in June (Rahmani et al., 2020).
• Bay of Oran: January to May (Gherram et al., 2018).
• Aegean Sea: April to August (Aydin & Erdoğan, 2018).
• Gulf of Skikda: December to April (Azzouz et al., 2019).
CONCLUSION

This review provides a deeper insight into the reproductive biology of T.

trachurus. It is a semi-pelagic migratory species that is widespread along the coasts, with
three migration modes, diel, seasonal, and trophic. The feeding strategy developed by
this species varies depending on its stage of growth; it is a very active predator with a
very flexible diet. This fish demonstrates considerable plasticity in its diet and can feed
 531 Trachurus trachurus: A Review

in any environment. Its diet comprises small crustaceans, mollusks, vertebrates, and
echinoderms. T. trachurus has an asynchronous reproduction. The spawning period
differs from region to region; this phenomenon is influenced by climatic conditions like
temperature and physicochemical parameters. Its reproduction rate is high. Sexual
maturity is attained at the age of two years for females, and around three years for males,
who have permanent sexual activity, while females are active in spring and summer.
However, this species faces several threats and challenges for its conservation
and management, such as overfishing, climate change, habitat degradation, and pollution.
Overfishing is one of the main causes of the decline of T. trachurus stocks, especially in
the Atlantic and the Mediterranean, where the fishing effort and catch have exceeded the
sustainable levels (El Mghazli et al., 2022). In turn, climate change can affect the
distribution, abundance, growth, reproduction, and survival of this species by modifying
water temperature, salinity, oxygen, currents, and food availability (Albo-Puigserver et
al., 2022). Moreover, habitat degradation can result from the destruction or alteration of
the coastal and marine ecosystems that provide shelter, food, and nursery grounds for
this species, such as seagrass beds, coral reefs, and mangroves (Rogers et al., 2019).
Pollution can also harm the health and quality of this species, by exposing it to
contaminants, such as heavy metals, pesticides, plastics, and oil spills, which can
accumulate in its tissues and affect its growth, reproduction, and survival (Köker et al.,
2021; Maaghloud et al., 2021).
It is therefore essential to implement effective conservation and management
actions for T. trachurus, on the basis of the latest scientific information and the
precautionary approach, to guarantee its long-term sustainability and resilience. These
measures include setting catch limits and quotas, regulating fishing gear and methods,
monitoring and enforcing compliance, creating marine protected areas, reducing
greenhouse gas emissions, restoring and conserving habitats, as well as preventing and
reducing pollution.
REFERENCES

Abattouy, N.; López, A.V.; Maldonado, J.L.; Benajiba, M.H. and Martín-Sánchez,
J. (2014). Epidemiology and molecular identification of Anisakis pegreffii (Nematoda:
Anisakidae) in the horse mackerel Trachurus trachurus from northern Morocco. J.
Helminthol., 88: 257–263.

Abaunza, P.; Gordo, L.; Karlou-Riga, C.; Murta, A.; Eltink, A.; Santamaría, M.G.;
Zimmermann, C.; Hammer, C.; Lucio, P. and Iversen, S.A. (2003). Growth and
reproduction of horse mackerel, Trachurus trachurus (Carangidae). Rev. Fish Biol. Fish.,
13: 27–61.
532 Nasri et al., 2024

Abaunza, P.; Murta, A.G.; Campbell, N.; Cimmaruta, R.; Comesaña, A.S.; Dahle,
G.; Santamaría, M.G.; Gordo, L.S.; Iversen, S.A. and MacKenzie, K. (2008). Stock
identity of horse mackerel (Trachurus trachurus) in the Northeast Atlantic and
Mediterranean Sea: Integrating the results from different stock identification approaches.
Fish. Res., 89: 196–209.

Albo-Puigserver, M.; Bueno-Pardo, J.; Pinto, M.; Monteiro, J.N.; Ovelheiro, A.;
Teodósio, M.A. and Leitão, F. (2022). Ecological sensitivity and vulnerability of fishing
fleet landings to climate change across regions. Sci. Rep., 12: 17360.
[Link]

Alheit, J. and Peck, M.A. (2019). Drivers of dynamics of small pelagic fish resources:
biology, management and human factors. Mar. Ecol. Prog., Ser., 617–618: 1–6.

Artüz, M.L. (2000). The egg and larval growth stages of Horse mackerel Trachurus
trachurus (Linnaeus, 1758) from the Sea of Marmara. Fisheries Advisory Commission
Technical Paper, Nr 202.

Asiedu, B.; Okpei, P.; Nunoo, F.K.E. and Failler, P. (2021). A fishery in distress: An
analysis of the small pelagic fishery of Ghana. Mar. Policy., 129: 104500.
[Link]

Aydin, G.U. and Erdoğan, (2018). Some reproductive characteristics of Trachurus

trachurus, (Linneaus, 1758) from Edremit Bay (Northern Aegean Sea, Turkey). Balıkesir
Üniv. Fen Bilim. Enst. derg., 20: 164–176. [Link]

Azzouz, S.; Mezedjri, L. and Tahar, A. (2019). Reproductive cycle of the pelagic fish
Saurel Trachurus trachurus (Linnaeus, 1758) (Perciformes Carangidae) Caught in the
Gulf of Skikda (Algerian East Coast). Biodivers. J., 10: 13–20.

Bayhan, B. and Sever, T.M. (2009). Food and feeding habits of the Atlantic horse
mackerel, Trachurus trachurus, from the Aegean Sea (Osteichthyes: Carangidae). Zool.
Middle East, 46: 47–54. [Link]

Bayhan, B.; Sever, T.M. and Kara, A. (2013). Diet composition of the Mediterranean
horse mackerel, Trachurus mediterraneus (Steindachner, 1868) (Osteichthyes:
Carangidae), from the Aegean Sea. Belg. J. Zool., 143: 15–22.

Bektas, Y. and Belduz, A.O. (2009). Morphological variation among Atlantic horse
mackerel, Trachurus trachurus populations from Turkish coastal waters. J. Anim. Vet.
Adv., 8: 511–517.

Berg, L.S. (1958). System der Rezenten und Fossilen fischartigen und Fische. Deutscher
Verlag Wissenschaften, Berlin, 310 pp.
 533 Trachurus trachurus: A Review

Bruslé, J. and Quignard, J.P. (2004). Les poissons et leur environnement:

ecophysiologie et comportements adaptifs. Éditions Tec & Doc, Lavoisier, 1522 pp.

Cabo, F.L. (1950). El jurel o chicharro (Trachurus Trachurus). PhD thesis, Universidad
Complutense de Madrid, 132 pp.

Campbell, N. (2005). The myxosporean parasitofauna of the Atlantic horse mackerel,

Trachurus trachurus (L.) in the North-East Atlantic Ocean and Mediterranean Sea. Acta
Parasitol., 50: 97–101.

Charef-Belifa, Z.E. (2009). Contribution à l’étude de la croissance de saurel, Trachurus

trachurus (Linné, 1758) pêche à Oran, par lecture d’otolithes et distribution des
fréquences de taille. Mémoire de Magister, Université d’Oran, 78 pp.

Chouvelon, T.; Violamer, L.; Dessier, A.; Bustamante, P.; Mornet, F.; Pignon-
Mussaud, C. and Dupuy, C. (2015). Small pelagic fish feeding patterns in relation to
food resource variability: an isotopic investigation for Sardina pilchardus and Engraulis
encrasicolus from the Bay of Biscay (north-east Atlantic). Mar. Biol., 162: 15–37.
[Link]

Costa, F.; Coelho, J.P.; Baptista, J.; Martinho, F.; Pereira, E. and Pardal, M.A.
(2021). Lifelong mercury bioaccumulation in Atlantic horse mackerel (Trachurus
trachurus) and the potential risks to human consumption. Mar. Pollut. Bull., 173: 113015.
[Link]

Dahl, K. and Kirkegaard, E. (1986). Stomach contents of mackerel, horse mackerel and
whiting in the eastern part of the North Sea in July 1985. Danmarks Fiskeri-og
Havundersøgelser, 17 pp.

D’Elia, M ; Patti, B.; Bonanno, A.; Fontana, I.; Giacalone, G.; Basilone, G. and
Fernandes, P.G. (2014). Analysis of backscatter properties and application of
classification procedures for the identification of small pelagic fish species in the Central
Mediterranean. Fish. Res., 149: 33–42.

Deme, M.; Cheikh, A. and Baboucar, B. (2019). Importance economique, sociale et

écologique des petits pélagiques au sénegal, en mauritanie et en guinée bissau. Rapport
technique du PRCM, 76 pp.

Dorson, M.; de Kinkelin, P. and Prunet, P. (2018). Eléments d’Ichtyologie. Santé des
poissons, [Link]

El Achi, A.; Nafia, M.; Manchih, K.; Baali, A. and Moncef, M. (2021) Diet of the
horse mackerel (Trachurus trachurus) in the North Atlantic of Morocco. AACL Bioflux,
14: 2554-2569.
534 Nasri et al., 2024

El Mghazli, H.; Znari, M. and Mounir, A. (2022). Stock Discrimination in the Horse
Mackerel Trachurus trachurus (Teleostei: Carangidae) off the Moroccan Atlantic Coastal
Waters using a Morphometric-Meristic Analysis. Thalassas, 38: 171–181.
[Link]

Eymard, S. (2003). Mise en évidence et suivi de l’oxydation des lipides au cours de la

conservation et de la transformation du chinchard (Trachurus trachurus): choix des
procédés (PhD Thesis). Université de Nantes.

Fanelli, E.; Da Ros, Z.; Menicucci, S.; Malavolti, S.; Biagiotti, I.; Canduci, G.; De
Felice, A. and Leonori, I. (2023). The pelagic food web of the Western Adriatic Sea: a
focus on the role of small pelagics. Sci Rep., 13: 14554. [Link]
023-40665-w

FAO. (2018). The state of world fisheries and aquaculture 2018 - Meeting the sustainable
development goals. FAO, Rome.

Fishbase. (2019). World Wide Web electronic publication. [Link].

Gaillard, S. (2006). Détermination et différenciation sexuelles chez les poissons:" le sexe

des esturgeons": SPACne et “opeintre” devient “protéine”. Thèse de doctorat, Toulon,
199 pp.

Gherram, M. (2019). Ecobiologie de trois taxons de Saurel, Trachurus trachurus (L,

1758), Trachurus mediterraneus (S, 1868) et Trachurus picturatus (B, 1825) de la baie
d’Oran : dynamique de population et diversité génétique. Thèse de Doctorat, Université
d’Oran, 252 pp.

Gherram, M.; Talet, A.B.; Dalouche, F. and Ayad, S.M.E.A.A. (2018). Study of
reproductive aspects of Trachurus trachurus (Linnaeus, 1758) from western coast of
Algeria. Indian J Mar Sci., 47: 2469–2476.

Giacomo, M.; Germana, G.; Samia, F.; Okbi, R.; Othman, J.; Bachra, C.; Luca, C.;
Angelo, B.; Simona, G.; Gualtiero, B.; Roberta, M.; Valentina, L.; Michele, G.;
Francesco, C. and Fabio, F. (2018). Biomass HotSpot distribution model and spatial
interaction of two exploited species of horse mackerel in the south-central Mediterranean
Sea. Hydrobiologia, 821: 135–150. [Link]

Gordo, L.S.; Costa, A.; Abaunza, P.; Lucio, P. Eltink, A. and Figueiredo, I. (2008).
Determinate versus indeterminate fecundity in horse mackerel. Fish. Res., 89: 181–185.

Hunnam, K. (2021). The biology and ecology of tropical marine sardines and herrings in
Indo-West Pacific fisheries: a review. Rev. Fish Biol. Fish., 31: 449–484.
[Link]
 535 Trachurus trachurus: A Review

Iversen, S.A.; Skogen, D.M. and Svendsen, E. (2002). Availability of horse mackerel
(Trachurus trachurus) in the north-eastern North Sea, predicted by the transport of
Atlantic water: Horse mackerel catches. Fish. Oceanogr., 11: 245–250.

Jardas, I.; Šantić, M and Pallaoro, A. (2004). Diet composition and feeding intensity
of horse mackerel, Trachurus trachurus (Osteichthyes: Carangidae) in the eastern
Adriatic. Mar. Biol., 144: 1051–1056.

Karaiskou, N.; Apostolidis, A.P.; Triantafyllidis, A.; Kouvatsi, A. and

Triantaphyllidis, C. (2003). Genetic Identification and Phylogeny of Three Species of
the Genus Trachurus Based on Mitochondrial DNA Analysis. Mar. Biotechnol., 5: 493–
504. [Link]

Kasumyan, A.O. and Pavlov, D.S. (2018). Evolution of schooling behavior in fish. J.
Ichthyol., 58: 670–678. [Link]

Köker L.; Aydın F ; Gaygusuz Ö.; Akçaalan R.; Çamur D.; İlter H ; Ayoğlu F N ;
Altın A ; Topbaş M and Albay M. (2021). Heavy Metal Concentrations in Trachurus
Mediterraneus and Merlangius Merlangus Captured from Marmara Sea, urkey and
Associated Health Risks. Environ. Manag., 67(3): 522-531. doi: 10.1007/s00267-020-
01352-y.

CrEC’hriou, R ; Marinaro, J.Y., and Planes, S. (2015). Advance in identification of

pelagic eggs of Mediterranean teleostean fish: development of a new identification key.
Vie Milieu, 65(1): 47-61.

Koç, H.T. and Erdoğan, (2019). Feeding Habits of the Mediterranean Horse
Mackerel, Trachurus mediterraneus (Steindachner, 1868) in the Sea of Marmara
(Bandırma Bay, Turkey). Nat. Eng. Sci., 4: 182–193.

Leitão, F.; Maharaj, R.R.; Vieira, V.M.N.C.S.; Teodósio, A. and Cheung, W.W.L.
(2018). The effect of regional sea surface temperature rise on fisheries along the
Portuguese Iberian Atlantic coast. Aquatic Conservation: Mar. Freshw. Ecosyst., 28,
1351–1359.

Letaconnoux, R. (1951). Contribution à l’étude des espèces du genre Trachurus et

spécialement du Trachurus trachurus (Linné 1758), Mem. Off. Sci. Techn. Peche Marit.,
15: 1–67.

Lévêque, C. (2006). Les poissons des eaux continentales africaines: diversité, écologie,
utilisation par l’homme, IRD Editions, Paris. 521 pp.

Lloris, D. and Rucabado, J. (1998). Guide d’identification des ressources marines

vivantes du Maroc. FAO, Rome, 263 pp.
536 Nasri et al., 2024

Maaghloud, H.; Houssa, R.; Bellali, F.; El Bouqdaoui, K.; Ouansafi, S.; Loulad, S.
and Fahde, A. (2021). Microplastic ingestion by Atlantic horse mackerel (Trachurus
trachurus) in the North and central Moroccan Atlantic coast between Larache (35°30′N)
and Boujdour (26°30′N), Environ Pollut., 288: 117781,
[Link]

Macer, C.T. (1977). Some aspects of the biology of the horse mackerel [Trachurus
trachurus (L.)] in waters around Britain. J. Fish Biol., 10, 51–62.

Mahe, K.; Delpech, J.P. and Carpentier, A. (2007). Synthèse bibliographique des
principales espèces de Manche orientale et du golfe de Gascogne, Rapport. Convention
Ifremer-Ministère de l’Industrie n°2006-0000708.

Maroua, B.; Said, B.; Chaima, A.; Chaik, A. and Mohammed, E. (2023). Effect of the
Impact of Environmental Factors on the Diet of Horse Mackerel (Trachurus trachurus)
from the Coasts of the Moroccan. In: Kacprzyk, J., Ezziyyani, M., Balas, V.E. (eds)
International Conference on Advanced Intelligent Systems for Sustainable Development.
AI2SD 2022. Lecture Notes in Networks and Systems, vol 713. Springer, Cham.

Meunier, F.-J. and Ramzu, M.-Y. (2006). La régionalisation morphofonctionnelle de

l’axe vertébral chez les Téléostéens en relation avec le mode de nage. Comptes Rendus
Palevol, Cent ans après Marey : Aspects de la morphologie fontionnelle aujourd’hui. CR
Palevol., 5: 499–507. [Link]

Michael, C. (2002). Contribution à l’étude de la biologie d’un poisson côtier le Saurel

Trachurus trachurus (LINNE, 1758): Anatomie et histologie du tube digestif, Mémoires
de Magistère, Université de Annaba-Badji Mokhtar, 99 pp.

Mouneimne, N. (1978). Poissons des cotes du Liban (Mediterranee orientale): biologie et

pêche, Thèse de Doctorat d’Etat ès-Sciences Naturelles, Université Pierre et Marie Curie,
France.

Mukherjee, M.; Karna, S.K.; Suresh, V.R.; Manna, R.K.; Panda, D. and Apurba, R.
(2018). New records of Carangids (Perciformes: Carangidae) from Chilika Lagoon, east
coast of India. FishTaxa, 2: 226–231.

Murua, H. and Saborido-Rey, F. (2003). Female reproductive strategies of marine fish

species of the North Atlantic. J. Northwest Atl. Fish. Sci., 33, 23–31.

DPM, Departement de la Pêche Maritime. (2022). La mer en chiffres. Rapport,

Morocco, 59 p. [Link]
a97b5b7cce06/Mer+en+Chiffres+DPM+[Link]?MOD=AJPERES
 537 Trachurus trachurus: A Review

Nasri, H.; Abdellaoui, S.; Omari, A.; Kada, O.; Chafi, A.; Hammouti, B. and
Chaabane, K. (2021). Length-weight relationship and condition factor of Trachurus
trachurus found in the central-east region of the Moroccan Mediterranean. Indones. J.
Sci. Technol., 6: 457–468.

Nasri, H. (2022). Etude de la reproduction et la croissance du Saurel Trachurus

trachurus (Linné, 1758) pêché dans la zone Orientale de la Méditerranée Marocaine, PhD
Thesis, Mohammed First University, Morocco

Ouled-Cheikh, J.; Giménez, J.; Albo-Puigserver, M.; Navarro, J.; Fernández-

Corredor, E., Bellido, E.; Grazia Pennino M. and Coll M. (2022) Trophic importance
of small pelagic fish to marine predators of the Mediterranean Sea. Mar. Ecol. Prog. Ser.,
696: 169–184. [Link]

Overko, S.M. and Mylnikov, N.I. (1979). biologie et pèche du Chinchard (Trachurus
trachurus L.) de l’Atlantique Centre-Est. Atlant-NIRO, Kaliningrad, URSS.

Peck, M.A.; Alheit, J.; Bertrand, A.; Catalán, I.A.; Garrido, S.; Moyano, M.;
Rykaczewski, R.R.; Takasuka, A. and Van Der Lingen, C.D. (2021). Small pelagic
fish in the new millennium: A bottom-up view of global research effort. Prog. Oceanogr.,
191: 102494.

Pikitch, E.; Boersma, P.D.; Boyd, I.; Conover, D.; Cury, P.; Essington, T.; Heppell,
S.; Houde, E.; Mangel, M. and Pauly, D. (2012). Little fish, big impact: managing a
crucial link in ocean food webs. Lenfest Ocean Program. Washington, DC, 108 pp.

Polonsky, A.S. (1965). The horse mackerel of the Eastern Atlantic and its fishery. Rybn.
Khoz., 41: 8–10.

Prolonge-Chevalier, C. (2007). Etude histologique du développement sexuel de l’apron

du Rhône Zingel asper L., Percidé endémique menacé d’extinction. PhD thesis, EPHE,
Lyon.

Quéro, J.-C. (2003). Guide des poissons de l’Atlantique européen. Les Guides du
Naturaliste, Delachaux et Niestlé, Lonay/Paris.

Rahmani, K.; Koudache, F.; Mouedden, N. E. R.; Talet, L. B. and Flower, R. (2020).
Spawning period, size at first sexual maturity and sex ratio of the atlantic horse mackerel
Trachurus trachurus from béni-saf bay (western coast of Algeria, southwestern
mediterranean sea). Ann. Ser. Hist. Nat., 30-1: 43–52.

Raybaud, V.; Bacha, M.; Amara, R. and Beaugrand, G. (2017). Forecasting climate-
driven changes in the geographical range of the European anchovy (Engraulis
encrasicolus). J. Mar. Sci., 74: 1288–1299. [Link]
538 Nasri et al., 2024

Robison, B.H. (2004). Deep pelagic biology. J. Exp. Mar. Biol. Ecol., 300: 253–272.

Rogers, L.A.; Griffin, R.; Young, T.; Fuller, E.; St. Martin K. and Pinsky, M.L.
(2019). Shifting habitats expose fishing communities to risk under climate change. Nat.
Clim. Change, 9: 512–516. [Link]

Rumolo, P.; Basilone, G.; Fanelli, E.; Barra, M.; Calabrò, M.; Genovese, S.;
Gherardi, S.; Ferreri, R.; Mazzola, S. and Bonanno, A. (2017). Linking spatial
distribution and feeding behavior of Atlantic horse mackerel (Trachurus trachurus) in the
Strait of Sicily (Central Mediterranean Sea). J. Sea Res., 121: 47–58.
[Link]

Russel, F.S., (1976). The Planktonic Stages of British Marine Fishes, Academic Pres Inc.
Ltd., London 524 pp.

Šantić, M ; Jardas, I. and Pallaoro, A. (2005). Feeding habits of horse mackerel,

Trachurus trachurus (Linneaus, 1758), from the central Adriatic Sea. J. Appl. Ichthyol.,
21: 125–130.

Santic, M.; Jardas, I., and Pallaoro, A. (2002). Age, growth, and mortality rate of horse
mackerel, Trachurus trachurus (L.), living in the eastern central Adriatic. Period. Biol.,
104: 165–173.

Schickele, A.; Leroy, B.; Beaugrand, G.; Goberville, E.; Hattab, T.; Francour,
P.,and Raybaud, V. (2020). Modelling European small pelagic fish distribution:
Methodological insights. Ecol. Model., 416: 108902.
[Link]

Shawket, N.; Youssir, S.; El Halouani, H.; Elmadhi, Y.; El Kharrim, K. and
Belghyti, D. (2015). Description des habitudes alimentaires du chinchard Trachurus
trachurus de l’atlantique nord marocain. Eur. Sci. J., 11: 294–304.

Smith-Vaniz, W.F.; Sidibe, A., Nunoo, F.; Lindeman, K.; Williams, A.B.; Quartey,
R., Camara, K.; Carpenter, K.E.; Montiero, V. and de Morais, L. (2015). Trachurus
trachurus. IUCN red list of threatened species Version 2015: e. T198647A43157137
2015–4.

Sutton, T.T. (2013). Vertical ecology of the pelagic ocean: classical patterns and new
perspectives. J. Fish Biol., 83: 1508–1527. [Link]

Villegas-Ríos, D. (2009). Associative behaviour between Trachurus trachurus

(Teleostei: Carangidae) and the hydromedusae Aequorea forskalea: a new relationship
reported from Galician waters. Mar. Biodivers. Rec., 2: e129.
[Link]
 539 Trachurus trachurus: A Review

Waldron, M.E. and Kerstan, M. (2001). Age validation in horse mackerel (Trachurus
trachurus) otoliths. J. Mar. Sci., 58, 806–813.

Wright, P.J.; Panfili, J.; Folkvord, A.; Mosegaard, H. and Meunier, F.J. (2002).
Manual of fish sclerochronology, Ifremer-IRD coedition, Brest, France, 464 pp.

Zohra, M.T.F. (2011). Contribution à l’étude de la biologie de la reproduction d’un petit

pélagique le saurel Trachurus trachurus: Spermatogenèse, Condition, RGS, RHS.
Mémoire Master, Univirsité Oran, 95 pp.
Copyright of Egyptian Journal of Aquatic Biology & Fisheries is the property of Egyptian
Society for Fisheries Development & Human Health and its content may not be copied or
emailed to multiple sites or posted to a listserv without the copyright holder's express written
permission. However, users may print, download, or email articles for individual use.