Dusky Grouper
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''Epinephelus marginatus'' (), the dusky grouper, yellowbelly rock cod or yellowbelly grouper, is a species of marine
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
, a
grouper Groupers are a diverse group of marine ray-finned fish in the family Epinephelidae, in the order Perciformes. Groupers were long considered a subfamily of the seabasses in Serranidae, but are now treated as distinct. Not all members of this f ...
from the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Epinephelinae Groupers are a diverse group of marine Actinopterygii, ray-finned fish in the family Epinephelidae, in the order Perciformes. Groupers were long considered a subfamily of the seabasses in Serranidae, but are now treated as distinct. Not all m ...
which is part of the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Serranidae Serranidae is a large family (biology), family of fishes belonging to the order Perciformes. The family contains about 450 species in 65 genera, including the sea basses and the groupers (subfamily Epinephelinae). Although many species are small, ...
, which also includes the anthias and sea basses. This species is the best known grouper species of the
Mediterranean Sea The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern Eur ...
and North Africa coast.


Description

''Epinephelus marginatus'' is a very large, oval-bodied and large-headed fish with a wide mouth which has a protruding lower jaw. The head and upper body are coloured dark reddish brown or greyish, usually with yellowish gold countershading on the ventral surfaces; the base colour is marked by a vertical series of irregular pale greenish yellow or silvery grey or whitish blotching which is normally rather conspicuous on the body and head; the black maxillary streak varies in its markedness; dark brown median fins; distal edges of the anal and caudal fins and also often
pectoral fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish aquatic locomotion, swim. Apart from the tail or caudal fin, fish fins have no direct connection with the vertebral column ...
s have narrow white terminal bands; the
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hi ...
s are black towards their tips while the pectoral fins are dark reddish-brown or grey; the margin of spiny
dorsal fin A dorsal fin is a fin on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...
and basal part of the pectoral fins are often golden yellow in colour. There are eleven spines and 13–16 soft rays in the dorsal fin. This species can grow up to 150 cm in
standard length Fish measurement is the measuring of individual fish and various parts of fish anatomy, their anatomies, for data used in many areas of ichthyology, including Taxonomy (biology), taxonomy and fishery biology. Overall length Standard length (SL) is ...
but is more often 90 cm.


Distribution

''Epinephelus marginatus'' has two disjunct distribution centres, the main one is in the eastern Atlantic from the west coast of Iberia south along the western coast of Africa to the
Cape of Good Hope The Cape of Good Hope ( ) is a rocky headland on the Atlantic Ocean, Atlantic coast of the Cape Peninsula in South Africa. A List of common misconceptions#Geography, common misconception is that the Cape of Good Hope is the southern tip of Afri ...
, extending east into the south-western Indian Ocean, as far as southern
Mozambique Mozambique, officially the Republic of Mozambique, is a country located in Southeast Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west, and Eswatini and South Afr ...
, with doubtful records from
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
and possibly
Oman Oman, officially the Sultanate of Oman, is a country located on the southeastern coast of the Arabian Peninsula in West Asia and the Middle East. It shares land borders with Saudi Arabia, the United Arab Emirates, and Yemen. Oman’s coastline ...
. It is found throughout the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
too. The second population occurs in the south western Atlantic off the coast of South America in southern
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
,
Uruguay Uruguay, officially the Oriental Republic of Uruguay, is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast, while bordering the Río de la Plata to the south and the A ...
and northern
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. In the eastern Atlantic it is not normally found further north than
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
but there have been rare records from the
Bay of Biscay The Bay of Biscay ( ) is a gulf of the northeast Atlantic Ocean located south of the Celtic Sea. It lies along the western coast of France from Point Penmarc'h to the Spanish border, and along the northern coast of Spain, extending westward ...
and in the
English Channel The English Channel, also known as the Channel, is an arm of the Atlantic Ocean that separates Southern England from northern France. It links to the southern part of the North Sea by the Strait of Dover at its northeastern end. It is the busi ...
as far north as northern
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
,
Great Britain Great Britain is an island in the North Atlantic Ocean off the north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
and
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
.


Habitat

''Epinephelus marginatus'' is
demersal The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer o ...
, normally found in and around rocky reefs from surface waters down to as much as 300 metres in depth. It often occurs in the vicinity of beds of ''
Posidonia ''Posidonia'' is a genus of flowering plants. It contains nine species of marine plants ("seagrass"), found in the seas of the Mediterranean and around the south coast of Australia. The APG system (1998) and APG II system (2003) accept this ge ...
'' sea grass. Juveniles are generally found more inshore than the adult fish, even being found in rock pools. Where they are protected, in marine nature reserves and no take zones, both adults and juveniles occur in shallow waters, but the depths at which juveniles are found is always shallower than the preferred depths of adults.


Biology

''Epinephelus marginatus'' adults are solitary and territorial, preferring areas with a rocky substrate but both adults and juveniles will enter brackish waters, such as estuaries. Their main food is
mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
s,
crustacea Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
ns, and
octopus An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like oth ...
es. But as they grow larger other fish form an increasingly important part of their diet, with reef fish being preferred. ''E. marginatus'' is a protogynous hermaphrodite, meaning that all fish begin adult life as females but as they grow larger and older they develop into males. They attain sexual maturity at quite a late age, females begin to breed when they are around five years of age, and then between their 9th and 16th years they change into males, most commonly at 12. The fish start to transform into males at a length of 65 cm, although most change sex when they are between 80 and 90 cm in total length. In some populations the presence of large female fish suggests that not all females change sex. During the breeding season small clusters of a few tens of individuals form at specific spawning sites, an exception to their normally solitary existence. Known sites where ''E. marginatus'' traditionally gather to spawn include the Medes Islands
Marine Reserve A marine reserve is a type of marine protected area (MPA). An MPA is a section of the ocean where a government has placed limits on human activity. A marine reserve is a marine protected area in which removing or destroying natural or cultural ...
in
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
, off
Lampedusa Lampedusa ( , , ; ; ) is the largest island of the Italian Pelagie Islands in the Mediterranean Sea. The ''comune'' of Lampedusa e Linosa is part of the Sicilian province of Agrigento which also includes the smaller islands of Linosa and Lamp ...
in
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
and Port-Cros National Park in
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
, all in the Mediterranean; fishermen in Brazil suspect there are aggregations off the coast of Santa Catarina but so far none has been definitely found. In the Mediterranean spawning lasts from June to September, mating is polygynous and the spawning clusters normally have seven females to each male. Off Brazil ''E. marginatus'' reproduces in the early summer, between November and December. While spawning, the dominant males set up territories and aggressively defend them from neighbouring males and smaller females. They are reported to live for up to 50 years.


Taxonomy

''Epinephelus marginatus'' was first formally described as ''Serranus marginatus'' in 1834 by the English
botanist Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
,
ichthyologist Ichthyology is the branch of zoology devoted to the study of fish, including bony fish (Osteichthyes), cartilaginous fish (Chondrichthyes), and jawless fish (Agnatha). According to FishBase, 35,800 species of fish had been described as of March 2 ...
,
malacologist Malacology, from Ancient Greek μαλακός (''malakós''), meaning "soft", and λόγος (''lógos''), meaning "study", is the branch of invertebrate zoology that deals with the study of the Mollusca (molluscs or mollusks), the second-largest ...
and clergyman
Richard Thomas Lowe Richard Thomas Lowe (1802–1874) was an English botanist, ichthyologist, malacologist, and clergyman. In 1825 he graduated from Christ's College, Cambridge, and in the same year he took holy orders. In 1832 he became a clergyman in the Madeira ...
(1802–1874) with the type locality given as "off
Madeira Madeira ( ; ), officially the Autonomous Region of Madeira (), is an autonomous Regions of Portugal, autonomous region of Portugal. It is an archipelago situated in the North Atlantic Ocean, in the region of Macaronesia, just under north of ...
". The name ''Perca gigas'' was coined by the Danish
zoologist Zoology ( , ) is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one ...
Morten Thrane Brünnich Morten Thrane Brünnich (30 September 1737 – 19 September 1827) was a Danish zoologist and mineralogist. Biography Brünnich was born in Copenhagen, the son of a portrait painter. He studied oriental languages and theology, but soon became i ...
(1737–1827) in 1768 but was unused and some authorities are of the view that this name should be suppressed by the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
.


Fishing

''Epinephelus marginatus'' is a popular food fish and is caught across its range by commercial fishermen while large adult fish are targeted as trophies by spear-fishing, and is readily taken by anglers. The slow growth rate of this species and its particular mode of reproduction make it vulnerable to over-exploitation, for example the targeting of large males by spear fishers may skew the sex ratio even further and affect reproductive productivity. There have been attempts to grow and breed this fish in
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
in Italy. In some countries the dusky grouper is considered a delicacy. Referring to its preference among restaurant guests, the Spanish say 'De la mar el mero y de la tierra el carnero' (From the sea the dusky grouper, from the land the lamb).


Conservation

''Epinephelus marginatus'' catch declined by 88% in seven countries between 1990–2001, these countries forming a significant part of its overall distribution. In other regions, such as West Africa, where this species is heavily exploited, there is little data about the status of this fish. For these reasons, the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the stat ...
has assessed E. marginatus as being Vulnerable, citing a suspected reduction in the population size reduction in excess 50% over the last three generations and where the causes of the decline continue. Conservation measures have included a spearfishing ban for ten years in France and bag limits in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. In addition, a number of Marine Protected Areas have been established to protect the habitat of ''E. marginatus''. In
Turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
recommended actions have included no take zones along the Aegean and Mediterranean Sea coasts and a total fisheries ban for a minimum of 3–5 years.


References


External resources

* {{Taxonbar, from=Q192167 marginatus Fish described in 1834 Taxonomy articles created by Polbot