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''Cancer pagurus'', commonly known as the edible crab or brown crab, is a species of
crab Crabs are decapod crustaceans of the infraorder Brachyura (meaning "short tailed" in Greek language, Greek), which typically have a very short projecting tail-like abdomen#Arthropoda, abdomen, usually hidden entirely under the Thorax (arthropo ...
found in the
North Sea The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium, and France. A sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Se ...
, North
Atlantic Ocean The Atlantic Ocean is the second largest of the world's five borders of the oceans, oceanic divisions, with an area of about . It covers approximately 17% of Earth#Surface, Earth's surface and about 24% of its water surface area. During the ...
, and perhaps 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 ...
. It is a robust crab of a reddish-brown colour, having an oval
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
with a characteristic "pie crust" edge and black tips to the claws. A mature adult may have a carapace width up to and weigh up to . ''C. pagurus'' is a nocturnal predator, targeting a range of molluscs and crustaceans. It is the subject of the largest crab fishery in
Western Europe Western Europe is the western region of Europe. The region's extent varies depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the Western half of the ancient Mediterranean ...
, centred on the coasts of the Ireland and Britain, with more than 60,000 tonnes caught annually.


Description

The
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
of ''C. pagurus'' adults is a reddish-brown colour, while in young specimens it is purple-brown. It occasionally bears white patches, and is shaped along the front edge into nine rounded lobes, resembling a pie crust. Males typically have a carapace long, and females long, although they may reach up to long in exceptional cases. Carapace width is typically , or exceptionally up to . A fold of the carapace extends ventrally to constitute a branchial chamber where the
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s lie. The first pereiopod is modified into a strong cheliped (claw-bearing leg); the claw's fingers, the dactylus and propodus, are black at the tips. The other pereiopods are covered with rows of short stiff
seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...
e; the dactylus of each is black towards the tip, and ends in a sharp point. From the front, the antennae and antennules are visible. Beside these, the orbits are where the
eyes An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
are situated. The mouthparts comprise three pairs of
maxilliped An appendage (or outgrowth) is an external body part or natural prolongation that protrudes from an organism's body such as an arm or a leg. Protrusions from single-celled bacteria and archaea are known as cell-surface appendages or surface app ...
s, behind which are a pair of maxillae, a pair of maxillules, and finally the
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
s.


Lifecycle

Reproduction occurs in winter; the male stands over the female and forms a cage with his legs protecting her while she moults. Internal fertilisation takes place before the hardening of the new carapace, with the aid of two abdominal appendages (gonopods). After mating, the female retreats to a pit on the sea floor to lay her eggs. Between 250,000 and 3,000,000 fertilised eggs are held under the female's abdomen up to eight months until they hatch. The first developmental stage after hatching is a
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
ic larva (1 mm) called the zoea that develops into a postlarva ( megalopa), and finally a juvenile. The first juvenile stage is characterised by a well-developed abdomen, which in time becomes reduced in size and folded under the sternum. Juveniles settle to the sea floor in the
intertidal zone The intertidal zone or foreshore is the area above water level at low tide and underwater at high tide; in other words, it is the part of the littoral zone within the tidal range. This area can include several types of habitats with various ...
, where they stay until they reach a
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
width of , and then migrate to deeper water. The growth rate in males slows from an increase in carapace width of 10 mm per year before it is 8 years old, to 2 mm per year thereafter. Females grow at about half the rate of males, probably due to the energetic demands of egg laying.
Sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans, it is related to both puberty and adulthood. ''Puberty'' is the biological process of sexual maturation, while ''adulthood'', the condition of being socially recognized ...
is reached at a carapace width of in females, and in males.
Longevity Longevity may refer to especially long-lived members of a population, whereas ''life expectancy'' is defined Statistics, statistically as the average number of years remaining at a given age. For example, a population's life expectancy at birth ...
is typically 25–30 years, although exceptional individuals may live up to 100 years.


Distribution and ecology

''C. pagurus'' is abundant throughout the northeast Atlantic as far as
Norway Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of the Kingdom of ...
in the north and
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
in the south, on mixed coarse grounds, mud, and sand from the shallow
sublittoral The littoral zone, also called litoral or nearshore, is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely in ...
to depths around . It is frequently found inhabiting cracks and holes in rocks, but occasionally also in open areas. Smaller specimens may be found under rocks in the
littoral zone The littoral zone, also called litoral or nearshore, is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely flood ...
. Unconfirmed reports suggest that ''C. pagurus'' may also occur in the Mediterranean Sea and
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
. Adults of ''C. pagurus'' are
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, hiding buried in the substrate during the day, but foraging at night up to from their hideouts. Their diet includes a variety of
crustacean 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 ...
s (including the crabs ''
Carcinus maenas ''Carcinus maenas'' is a common littoral crab. It is known by different names around the world. In the British Isles, it is generally referred to as the shore crab or green shore crab. In North America and South Africa, it bears the name Europe ...
'' and '' Pilumnus hirtellus'', the
porcelain crab Porcelain crabs are decapod crustaceans in the widespread family Porcellanidae, which superficially resemble true crabs. They have flattened bodies as an adaptation for living in rock crevices. They are delicate, readily losing limbs when attacke ...
s '' Porcellana platycheles'' and '' Pisidia longicornis'', and the
squat lobster Squat lobsters are dorsoventrally flattened crustaceans with long tails held curled beneath the cephalothorax. They are found in the two superfamilies Galatheoidea and Chirostyloidea, which form part of the decapod infraorder Anomura, alongs ...
'' Galathea squamifera'') and
molluscs 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 ...
(including the
gastropods Gastropods (), commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and from the land. Ther ...
'' Nucella lapillus'' and '' Littorina littorea'', and the
bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
''
Ensis ''Ensis'' is a genus of medium-sized edible saltwater clams, littoral bivalve molluscs in the family Pharidae. ''Ensis'', or razor clams, are known in much of Scotland as spoots, for the spouts of water they eject while burrowing into the san ...
'', ''
Mytilus edulis The blue mussel (''Mytilus edulis''), also known as the common mussel, is a medium-sized edible marine (ocean), marine bivalve mollusc in the family (biology), family Mytilidae, the only extant family in the order (biology), order Mytilida, known ...
'', '' Cerastoderma edule'', '' Ostrea edulis'', and '' Lutraria lutraria''). It may stalk or ambush motile prey, and may dig large pits to reach buried molluscs. The main
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
of ''C. pagurus'' is the
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 ...
, which even attacks them inside the crab pots that fishermen use to trap them.


Diseases

Compared to other commercially important crab species, relatively little is known about diseases of ''C. pagurus''. Its parasites include
virus A virus is a submicroscopic infectious agent that replicates only inside the living Cell (biology), cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Viruses are ...
es, such as the white spot syndrome virus, various
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
that cause dark
lesion A lesion is any damage or abnormal change in the tissue of an organism, usually caused by injury or diseases. The term ''Lesion'' is derived from the Latin meaning "injury". Lesions may occur in both plants and animals. Types There is no de ...
s on the
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
, and '' Hematodinium''-like
dinoflagellate The Dinoflagellates (), also called Dinophytes, are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered protists. Dinoflagellates are mostly marine plankton, but they are also commo ...
s that cause "pink crab disease". Other microscopic
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
s include
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
,
microsporidia Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore.Franzen, C. (2005). How do Microsporidia inva ...
ns, paramyxeans, and
ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
s. ''C. pagurus'' is also targeted by
metazoan Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia (). With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a ho ...
parasites, including trematodes and parasitic
barnacle Barnacles are arthropods of the subclass (taxonomy), subclass Cirripedia in the subphylum Crustacean, Crustacea. They are related to crabs and lobsters, with similar Nauplius (larva), nauplius larvae. Barnacles are exclusively marine invertebra ...
s. A number of sessile animals occasionally settle as
epibiont An epibiont (from the Ancient Greek meaning "living on top of") is an organism that lives on the surface of another living organism, called the basibiont ("living underneath"). The interaction between the two organisms is called epibiosis. An ep ...
s on the exoskeleton of ''C. pagurus'', including barnacles,
sea anemone Sea anemones ( ) are a group of predation, predatory marine invertebrates constituting the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemone ...
s, serpulid
polychaete Polychaeta () is a paraphyletic class of generally marine Annelid, annelid worms, common name, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called c ...
s such as ''
Janua pagenstecheri ''Janua pagenstecheri'' is a species of marine polychaete. It is widely distributed around the British Isles and across north-western Europe, and has been described as "probably the commonest spirorbid in the world". ''Janua pagenstecheri'' liv ...
'',
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns, and saddle oysters.


Fishery

''C. pagurus'' is heavily exploited commercially throughout its range, being the most commercially important crab species in Western Europe. The crabs are caught using crab pots (similar to lobster pots), also known as creels, which are placed offshore and baited. The catch of ''C. pagurus'' has increased steadily, rising from 26,000 tonnes in 1978 to 60,000 t in 2007, of which more than 70% was caught around the
British Isles The British Isles are an archipelago in the Atlantic Ocean, North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner Hebrides, Inner and Outer Hebr ...
. The fishery is widely dispersed around the British and Irish coasts, and ''C. pagurus'' is thought to be overfished across much of this area. Most of the edible crabs caught by the British fleet are exported live for sale 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 ...
and
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 ...
. A number of legal restrictions apply to the catching of ''C. pagurus''. Catching "berried" crabs (females carrying eggs) is illegal, but since ovigerous females remain in pits dug in the sediment and do not feed, fishing pressure does not affect the supply of larvae.
Minimum landing size The minimum landing size (MLS) is the smallest fish measurement at which it is legal to keep or sell a fish. The MLS depends on the species of fish. Sizes also vary around the world, as they are legal definitions which are defined by the local regu ...
s (MLSs) for ''C. pagurus'' are set by both the
European Union The European Union (EU) is a supranational union, supranational political union, political and economic union of Member state of the European Union, member states that are Geography of the European Union, located primarily in Europe. The u ...
technical regulations and by the
UK government His Majesty's Government, abbreviated to HM Government or otherwise UK Government, is the central government, central executive authority of the United Kingdom of Great Britain and Northern Ireland.
. Different minimum sizes are employed in different geographical areas, to reflect differences in the crab's growth rate across its range. In particular, the "
Cromer Cromer ( ) is a coastal town and civil parish on the north coast of the North Norfolk district of the county of Norfolk, England. It is north of Norwich, northwest of North Walsham and east of Sheringham on the North Sea coastline. The local ...
crab" fishery along the coasts of
Suffolk Suffolk ( ) is a ceremonial county in the East of England and East Anglia. It is bordered by Norfolk to the north, the North Sea to the east, Essex to the south, and Cambridgeshire to the west. Ipswich is the largest settlement and the county ...
,
Norfolk Norfolk ( ) is a Ceremonial counties of England, ceremonial county in England, located in East Anglia and officially part of the East of England region. It borders Lincolnshire and The Wash to the north-west, the North Sea to the north and eas ...
and
Lincolnshire Lincolnshire (), abbreviated ''Lincs'', is a Ceremonial counties of England, ceremonial county in the East Midlands and Yorkshire and the Humber regions of England. It is bordered by the East Riding of Yorkshire across the Humber estuary to th ...
is subject to an MLS of , rather than the MLS in most of the species' range. An intermediate value of is used in the rest of the North Sea between the 56°N and the
Essex Essex ( ) is a Ceremonial counties of England, ceremonial county in the East of England, and one of the home counties. It is bordered by Cambridgeshire and Suffolk to the north, the North Sea to the east, Kent across the Thames Estuary to the ...
Kent Kent is a Ceremonial counties of England, ceremonial county in South East England. It is bordered by Essex across the Thames Estuary to the north, the Strait of Dover to the south-east, East Sussex to the south-west, Surrey to the west, and Gr ...
border, and in the
Irish Sea The Irish Sea is a body of water that separates the islands of Ireland and Great Britain. It is linked to the Celtic Sea in the south by St George's Channel and to the Inner Seas off the West Coast of Scotland in the north by the North Ch ...
south of 55°N. Around
Devon Devon ( ; historically also known as Devonshire , ) is a ceremonial county in South West England. It is bordered by the Bristol Channel to the north, Somerset and Dorset to the east, the English Channel to the south, and Cornwall to the west ...
,
Cornwall Cornwall (; or ) is a Ceremonial counties of England, ceremonial county in South West England. It is also one of the Celtic nations and the homeland of the Cornish people. The county is bordered by the Atlantic Ocean to the north and west, ...
, and the
Isles of Scilly The Isles of Scilly ( ; ) are a small archipelago off the southwestern tip of Cornwall, England. One of the islands, St Agnes, Isles of Scilly, St Agnes, is over farther south than the most southerly point of the Great Britain, British mainla ...
, the MLS for males is different () from females (). The Norwegian catch is 8,500 tons annually, compared to 20,000 tons in the United Kingdom, 13,000 tons in Ireland, 8,500 tons in France, and a total 45,000 tons globally. Recent studies have shown that edible crabs are negatively affected by electromagnetic fields emitted from sub-sea power cables around offshore wind farms.


Cookery

Around one-third of the weight of an adult edible crab is meat, of which one-third is white meat from the claws (see declawing of crabs), and two-thirds is white and brown meat from the body. As food, male edible crabs are referred to as cocks and females as hens. Cocks have more sweet white meat; hens have more rich brown meat. Dishes include dressed crab (crab meat arranged in the cleaned shell, sometimes with decoration of other foodstuffs), soups such as bisque or
bouillabaisse Bouillabaisse ( , , ; ) is a traditional Cuisine of Provence, Provençal fish soup originating in the port city of Marseille. The word is originally a compound of the two Provençal verbs ('to boiling, boil') and ('to reduce heat', i.e. 'sim ...
, ''
pâté Pâté ( , , ) is a forcemeat. Originally, the dish was cooked in a pastry case; in more recent times it is more usually cooked without pastry in a terrine. Various ingredients are used, which may include meat from pork, poultry, fish or bee ...
s,
mousse A mousse (, ; ) is a soft prepared food that incorporates air bubbles to give it a light and airy texture. Depending on preparation techniques, it can range from light and fluffy to creamy and thick. A mousse may be sweet or savory. as early ...
s'', and hot ''
soufflé A soufflé () is a baked egg dish originating in France in the early 18th century. Combined with various other ingredients, it can be served as a savoury main dish or sweetened as a dessert. The word ''soufflé'' is the past participle of the Fr ...
s''.


Taxonomy and systematics

According to the rules of the
International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted Convention (norm), convention in zoology that rules the formal scientific name, scientific naming of organisms treated as animals. It is also informally known as the I ...
, ''Cancer pagurus'' was first described by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in 1758, in the tenth edition of his ''
Systema Naturae ' (originally in Latin written ' with the Orthographic ligature, ligature æ) is one of the major works of the Sweden, Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the syste ...
'', which marks the starting point of zoological nomenclature. It was chosen to be the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the genus ''
Cancer Cancer is a group of diseases involving Cell growth#Disorders, abnormal cell growth with the potential to Invasion (cancer), invade or Metastasis, spread to other parts of the body. These contrast with benign tumors, which do not spread. Po ...
'' by
Pierre André Latreille Pierre André Latreille (; 29 November 1762 – 6 February 1833) was a French zoology, zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained hi ...
in 1810. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
is a
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word, deriving from the (), which, alongside (), was used to refer to edible marine crabs; neither classical term can be confidently assigned to a particular species. Although the genus ''Cancer'' formerly included most crabs, it has since been restricted to eight species. Within that set of closely related species, the closest relative of ''C. pagurus'' is the Jonah crab, ''C. borealis'', from the east coast of
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
.


References


External links

*
Dressed crab
recipe from BBC Food * {{Authority control Cancroidea Edible crustaceans Commercial crustaceans Crabs of the Atlantic Ocean Crustaceans described in 1758 Taxa named by Carl Linnaeus