Cancer pagurus
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''Cancer pagurus'', commonly known as the edible crab or brown crab, is a species of crab found in the
North Sea The North Sea lies between Great Britain, Norway, Denmark, Germany, the Netherlands and Belgium. An epeiric sea, epeiric sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the ...
, North
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
, 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 north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ...
. It is a robust crab of a reddish-brown colour, having an oval carapace 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 countries and territories vary depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the ancient Mediterranean ...
, centred on the coasts of the British Isles, with more than 60,000 tonnes caught annually.


Description

The carapace 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 respiratory organ that many 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 respiration on land provided they are ...
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 (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
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 Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and con ...
are situated. The mouthparts comprise three pairs of maxillipeds, behind which are a pair of maxillae, a pair of maxillules, and finally the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
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 Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For int ...
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 found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
ic larva (1 mm) called the
zoea Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...
that develops into a
postlarva Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...
(
megalopa Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...
), 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, where they stay until they reach a carapace 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 might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definit ...
is reached at a carapace width of in females, and in males. Longevity 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 country in Northern Europe, the mainland territory of which comprises the western and northernmost portion of the Scandinavian Peninsula. The remote Arctic island of Jan Mayen and the ...
in the north and
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
in the south, on mixed coarse grounds, mud, and sand from the shallow
sublittoral The littoral zone 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 inundated), to coastal area ...
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 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 inundated), to coastal a ...
. Unconfirmed reports suggest that ''C. pagurus'' may also occur in the Mediterranean Sea and
Black Sea The Black Sea is a marginal mediterranean sea of the Atlantic Ocean 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 bounded by Bulgaria, Georgia, Rom ...
. Adults of ''C. pagurus'' are nocturnal, 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 (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
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 europ ...
'' 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 attack ...
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, alongsi ...
''
Galathea squamifera ''Galathea squamifera'', the black squat lobster, or Montagu's plated lobster, is a species of squat lobster that lives in the north-east Atlantic Ocean and Mediterranean Sea. Description Adults are up to long, with a carapace long; the body ...
'') and
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is estim ...
(including the
gastropods The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. The ...
'' Nucella lapillus'' and ''
Littorina littorea The common periwinkle or winkle (''Littorina littorea'') is a species of small edible whelk or sea snail, a marine gastropod mollusc that has gills and an operculum, and is classified within the family Littorinidae, the periwinkles.Reid, David ...
'', and the bivalves ''
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 sa ...
'', ''
Mytilus edulis The blue mussel (''Mytilus edulis''), also known as the common mussel, is a medium-sized edible marine bivalve mollusc in the family Mytilidae, the mussels. Blue mussels are subject to commercial use and intensive aquaculture. A species with a l ...
'', '' Cerastoderma edule'', ''
Ostrea edulis ''Ostrea edulis'', commonly known as the European flat oyster, is a species of oyster native to Europe. In Britain and Ireland, regional names include Colchester native oyster, mud oyster, or edible oyster. In France, ''Ostrea edulis'' are known ...
'', 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 where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
of ''C. pagurus'' is the octopus, 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 cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsk ...
es, such as the white spot syndrome virus, various
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
that cause dark
lesion A lesion is any damage or abnormal change in the tissue of an organism, usually caused by disease or trauma. ''Lesion'' is derived from the Latin "injury". Lesions may occur in plants as well as animals. Types There is no designated classif ...
s on the
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
, and ''
Hematodinium ''Hematodinium'' is a genus of dinoflagellates. Species in this genus, such as '' Hematodinium perezi'', the type species, are internal parasites of the hemolymph of crustaceans such as the Atlantic blue crab (''Callinectes sapidus'') and Norway ...
''-like dinoflagellates that cause "pink crab disease". Other microscopic
pathogen In biology, a pathogen ( el, πάθος, "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 ...
s include
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
,
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. They were once considered protozoans or pr ...
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 eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...
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, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
parasites, including
trematodes Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host ...
and parasitic barnacles. A number of
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
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 anemones, serpulid
polychaete Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made ...
s such as '' Janua pagenstecheri'',
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
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 a group of islands in the 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 and Outer Hebrides, the Northern Isles, ...
. The fishery is widely dispersed around the British and Irish coasts, and ''C. pagurus'' is thought to be
overfished Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in t ...
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 primarily located in Western Europe. It also comprises of overseas regions and territories in the Americas and the Atlantic, Pacific and Indian Oceans. Its metropolitan area ...
and
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
. 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 political and economic union of member states that are located primarily in Europe. The union has a total area of and an estimated total population of about 447million. The EU has often been de ...
technical regulations and by the
UK government ga, Rialtas a Shoilse gd, Riaghaltas a Mhòrachd , image = HM Government logo.svg , image_size = 220px , image2 = Royal Coat of Arms of the United Kingdom (HM Government).svg , image_size2 = 180px , caption = Royal Arms , date_est ...
. 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 crab" fishery along the coasts of Suffolk,
Norfolk Norfolk () is a ceremonial and non-metropolitan county in East Anglia in England. It borders Lincolnshire to the north-west, Cambridgeshire to the west and south-west, and Suffolk to the south. Its northern and eastern boundaries are the No ...
and
Lincolnshire Lincolnshire (abbreviated Lincs.) is a Counties of England, county in the East Midlands of England, with a long coastline on the North Sea to the east. It borders Norfolk to the south-east, Cambridgeshire to the south, Rutland to the south-we ...
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 county in the East of England. One of the home counties, it borders Suffolk and Cambridgeshire to the north, the North Sea to the east, Hertfordshire to the west, Kent across the estuary of the River Thames to the south, and Grea ...
Kent Kent is a county in South East England and one of the home counties. It borders Greater London to the north-west, Surrey to the west and East Sussex to the south-west, and Essex to the north across the estuary of the River Thames; it faces ...
border, and in the
Irish Sea The Irish Sea or , gv, Y Keayn Yernagh, sco, Erse Sie, gd, Muir Èireann , Ulster-Scots: ''Airish Sea'', cy, Môr Iwerddon . is an extensive body of water that separates the islands of Ireland and Great Britain. It is linked to the Ce ...
south of 55°N. Around
Devon Devon ( , historically known as Devonshire , ) is a ceremonial and non-metropolitan county in South West England. The most populous settlement in Devon is the city of Plymouth, followed by Devon's county town, the city of Exeter. Devo ...
,
Cornwall Cornwall (; kw, Kernow ) is a historic county and ceremonial county in South West England. It is recognised as one of the Celtic nations, and is the homeland of the Cornish people. Cornwall is bordered to the north and west by the Atlantic ...
, and the Isles of Scilly, 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 (crab meat arranged in the cleaned shell, sometimes with decoration of other foodstuffs), soups such as bisque or
bouillabaisse Bouillabaisse (; oc, bolhabaissa, bullabessa ) is a traditional Provençal fish stew originating in the port city of Marseille. The French and English form ''bouillabaisse'' comes from the Provençal Occitan word ''bolhabaissa'', a compound ...
, ''
pâté ''Pâté'' ( , , ) is a paste, pie or loaf filled with a forcemeat. Common forcemeats include ground meat from pork, poultry, fish or beef; fat, vegetables, herbs, spices and either wine or brandy (often cognac or armagnac). It is often ser ...
s, mousses'', and hot ''
soufflé A soufflé is a baked egg-based dish originating in France in the early eighteenth century. Combined with various other ingredients, it can be served as a savory main dish or sweetened as a dessert. The word soufflé is the past participle of t ...
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 in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the ...
, ''Cancer pagurus'' was first described by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
in 1758, in the tenth edition of his '' Systema Naturae'', which marks the starting point of zoological nomenclature. It was chosen to be the
type species In 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 species that contains the biological type specime ...
of the genus ''
Cancer Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal b ...
'' by
Pierre André Latreille Pierre André Latreille (; 29 November 1762 – 6 February 1833) was a French zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained his freedom ...
in 1810. The specific epithet is a
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word, deriving from the grc, πάγουρος (), 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.


References


External links

*
Dressed crab
recipe from
BBC Food BBC Food is a division of the BBC which controls a recipe website part of BBC Online, an online streaming channel, and a former international commercial television channel focusing on food. Website BBC Food is the public service website whic ...
* {{Good article Cancroidea Edible crustaceans Commercial crustaceans Crustaceans of the Atlantic Ocean Crustaceans described in 1758 Taxa named by Carl Linnaeus