Crab Spider - Synema Parvulum
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Crabs are
decapod The Decapoda or decapods, from Ancient Greek δεκάς (''dekás''), meaning "ten", and πούς (''poús''), meaning "foot", is a large order of crustaceans within the class Malacostraca, and includes crabs, lobsters, crayfish, shrimp, and p ...
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 of the
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
Brachyura (meaning "short tailed" in
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
), which typically have a very short projecting
tail The tail is the elongated section at the rear end of a bilaterian animal's body; in general, the term refers to a distinct, flexible appendage extending backwards from the midline of the torso. In vertebrate animals that evolution, evolved to los ...
-like
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
, usually hidden entirely under the
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
. Their
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 ...
is often thickened and hard. They generally have five pairs of legs, and they have "
pincer Pincer may refer to: *Pincers (tool) *Pincer (biology), part of an animal *Pincer ligand, a terdentate, often planar molecule that tightly binds a variety of metal ions *Pincer (Go), a move in the game of Go *"Pincers!", an episode of the TV series ...
s" or "
claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or Arthro ...
s" on the ends of the frontmost pair, scientifically termed the ''
chelae A chela ()also called a claw, nipper, or pinceris a pincer-shaped organ at the end of certain limbs of some arthropods. The name comes from Ancient Greek , through Neo-Latin '. The plural form is chelae. Legs bearing a chela are called chelipeds ...
''. They are present in all the world's oceans, in freshwater, and
on land ''Ambient 4: On Land'' is the eighth solo studio album by Brian Eno, released in March 1982 by EG Records. It was the final edition in Eno's Ambient series, which began in 1979 with '' Ambient 1: Music for Airports''. The album was released to ...
, often hiding themselves in small crevices or
burrowing An eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of s ...
into
sediment Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
. Crabs are
omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
s, feeding on a variety of food, including a significant proportion of
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
, as well as
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
and other
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s. Crabs are widely consumed by humans as
food Food is any substance consumed by an organism for Nutrient, nutritional support. Food is usually of plant, animal, or Fungus, fungal origin and contains essential nutrients such as carbohydrates, fats, protein (nutrient), proteins, vitamins, ...
, with over 1.5 million tonnes
caught Caught is a method of dismissing a batsman in cricket. A batsman is out caught if the batsman hits the ball, from a legitimate delivery, with the bat, and the ball is caught by the bowler or a fielder before it hits the ground. If the catch ...
annually. True crabs first appeared in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
during the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period, around 200 million years ago, achieving great diversity by the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period; around 7,000 extant species in 96 families are known. A number of other crustacean groups converged on the
body plan A body plan, (), or ground plan is a set of morphology (biology), morphological phenotypic trait, features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually app ...
of crabs, sometimes closely resembling them; this phenomenon is called
carcinisation Carcinisation (American English: carcinization) is a form of convergent evolution in which non-crab crustaceans evolve a crab-like body plan. The term was introduced into evolutionary biology by Lancelot Alexander Borradaile, who described it a ...
.


Description

Crabs are generally covered with a thick
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 ...
, composed primarily of highly mineralized
chitin Chitin (carbon, C8hydrogen, H13oxygen, O5nitrogen, N)n ( ) is a long-chain polymer of N-Acetylglucosamine, ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is the second most abundant polysaccharide in nature (behind only cell ...
. Behind their pair of
chelae A chela ()also called a claw, nipper, or pinceris a pincer-shaped organ at the end of certain limbs of some arthropods. The name comes from Ancient Greek , through Neo-Latin '. The plural form is chelae. Legs bearing a chela are called chelipeds ...
(claws) are six walking legs and then two swimming legs. The crab breathes through
gills 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 ...
on its underside; gills must be at least moist to work. Crabs vary in size from the
pea crab The pea crab, ''Pinnotheres pisum'', is a small crab in the family Pinnotheridae that lives as a parasite in oysters, clams, mussels, and other species of bivalves. Etymology ''Pinnotheres'' is Greek for "guard of '' Pinna''" and ''pisum'' i ...
, a few millimeters wide, to the
Japanese spider crab The Japanese giant spider crab (''Macrocheira kaempferi'') is a species of marine crab and is the biggest one that lives in the waters around Japan. At around 3.7 meters, it has the largest leg-span of any arthropod. The Japanese name for this s ...
, with a leg span up to . Several other groups of crustaceans with similar appearances – such as king crabs and
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 – are not true crabs, but have evolved features similar to true crabs through a process known as
carcinisation Carcinisation (American English: carcinization) is a form of convergent evolution in which non-crab crustaceans evolve a crab-like body plan. The term was introduced into evolutionary biology by Lancelot Alexander Borradaile, who described it a ...
.


Environment

Crabs are found in all of the world's oceans, as well as in
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salt (chemistry), salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include ...
and on
land Land, also known as dry land, ground, or earth, is the solid terrestrial surface of Earth not submerged by the ocean or another body of water. It makes up 29.2% of Earth's surface and includes all continents and islands. Earth's land sur ...
, particularly in
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
regions. About 850
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
are
freshwater crabs Around 1,300 species of freshwater crabs are distributed throughout the tropics and subtropics, divided among eight families. They show direct development and maternal care of a small number of offspring, in contrast to marine crabs, which releas ...
.


Sexual differences

Crabs often show marked
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
. Males often have larger
claws A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tars ...
, a tendency that is particularly pronounced in the
fiddler crab The fiddler crab or calling crab can be one of the hundred species of semiterrestrial marine crabs in the family Ocypodidae. These crabs are well known for their extreme sexual dimorphism, where the male crabs have a major claw significantly l ...
s of the genus ''Uca'' (
Ocypodidae The Ocypodidae is a family of semiterrestrial crabs that includes ghost crabs and fiddler crabs. These crabs are found along tropical and temperate shorelines worldwide. Some genera previously classified within the Ocypodidae family have been r ...
). In fiddler crabs, males have one greatly enlarged claw used for communication, particularly for attracting a mate. Another conspicuous difference is the form of the pleon (
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
); in most male crabs, this is narrow and triangular in form, while females have a broader, rounded abdomen. This is because female crabs brood fertilised eggs on their
pleopod The anatomy of a decapod consists of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment – often called a somite – may possess one pair of appendages, although in various groups these m ...
s.


Life cycle

Crabs attract a mate through chemical (
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s), visual, acoustic, or vibratory means. Pheromones are used by most fully aquatic crabs, while
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
and semiterrestrial crabs often use visual signals, such as
fiddler crab The fiddler crab or calling crab can be one of the hundred species of semiterrestrial marine crabs in the family Ocypodidae. These crabs are well known for their extreme sexual dimorphism, where the male crabs have a major claw significantly l ...
males waving their large claws to attract females. The vast number of brachyuran crabs have
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 intern ...
and mate belly-to-belly. For many aquatic species, mating takes place just after the female has moulted and is still soft. Females can store the
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
for a long time before using it to fertilise their
eggs An egg is an organic vessel in which an embryo begins to develop. Egg, EGG or eggs may also refer to: Biology * Egg cell, the female reproductive cell (gamete) in oogamous organisms Food * Eggs as food Places * Egg, Austria * Egg, Switzerland ...
. When fertilisation has taken place, the eggs are released onto the female's abdomen, below the tail flap, secured with a sticky material. In this location, they are protected during embryonic development. Females carrying eggs are called "berried" since the eggs resemble round berries. When development is complete, the female releases the newly hatched
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
into the water, where they are part of the
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 ...
. The release is often timed with the
tidal Tidal is the adjectival form of tide. Tidal may also refer to: * ''Tidal'' (album), a 1996 album by Fiona Apple * Tidal (king), a king involved in the Battle of the Vale of Siddim * TidalCycles, a live coding environment for music * Tidal (servic ...
and light/dark
diurnal cycle A diurnal cycle (or diel cycle) is any pattern that recurs every 24 hours as a result of one full Earth's rotation, rotation of the planet Earth around its axis. Earth's rotation causes surface diurnal temperature variation, temperature fluctuati ...
. The free-swimming tiny zoea larvae can float and take advantage of water currents. They have a spine, which probably reduces the rate of predation by larger animals. The zoea of most species must find food, but some crabs provide enough
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example bec ...
in the eggs that the larval stages can continue to live off the yolk. Each species has a particular number of zoeal stages, separated by moults, before they change into a
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 ...
stage, which resembles an adult crab, except for having the abdomen (tail) sticking out behind. After one more moult, the crab is a juvenile, living on the bottom rather than floating in the water. This last moult, from megalopa to juvenile, is critical, and it must take place in a habitat that is suitable for the juvenile to survive. Most species of terrestrial crabs must
migrate Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
down to the ocean to release their larvae; in some cases, this entails very extensive migrations. After living for a short time as larvae in the ocean, the juveniles must do this migration in reverse. In many tropical areas with land crabs, these migrations often result in considerable
roadkill Roadkill is a wild animal that has been killed by collision with motor vehicles. Wildlife-vehicle collisions (WVC) have increasingly been the topic of academic research to understand the causes, and how they can be mitigated. History Essenti ...
of migrating crabs. Once crabs have become juveniles, they still have to keep moulting many more times to become adults. They are covered with a hard shell, which would otherwise prevent growth. The moult cycle is coordinated by
hormone A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physio ...
s. When preparing for moult, the old shell is softened and partly eroded away, while the rudimentary beginnings of a new shell form under it. At the time of moulting, the crab takes in a lot of water to expand and crack open the old shell at a line of weakness along the back edge of 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 ...
. The crab must then extract all of itself – including its legs, mouthparts,
eyestalk In anatomy, an eyestalk (sometimes spelled eye stalk and also known as an ommatophore) is a protrusion that extends an eye away from the body, giving the eye a better field of view. It is a common feature in nature and frequently appears in fict ...
s, and even the lining of the front and back of the digestive tract – from the old shell. This is a difficult process that takes many hours, and if a crab gets stuck, it will die. After freeing itself from the old shell (now called an
exuvia In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have molted. The exuviae of an animal can be important to biologists as they can often b ...
), the crab is extremely soft and hides until its new shell has hardened. While the new shell is still soft, the crab can expand it to make room for future growth.


Behaviour

Crabs typically walk sideways (hence the term crabwise), because of the articulation of the legs which makes a sidelong gait more efficient. Some crabs walk forward or backward, including raninids, ''
Libinia emarginata ''Libinia emarginata'', the portly spider crab, common spider crab or nine-spined spider crab, is a species of stenohaline crab that lives on the Atlantic coast of North America. Distribution ''Libinia emarginata'' occurs from Nova Scotia to th ...
'' and ''
Mictyris platycheles ''Mictyris platycheles'' is a species of crab found on mudflats on the east coast of Australia from Tasmania and Victoria to Queensland Queensland ( , commonly abbreviated as Qld) is a States and territories of Australia, state in northea ...
''. Some crabs, like the
Portunidae Portunidae is a family of crabs which contains the swimming crabs. Its members include well-known shoreline crabs such as the blue crab (''Callinectes sapidus'') and velvet crab ('' Necora puber''). Description Portunid crabs are characterised b ...
and
Matutidae Matutidae is a family (biology), family of crabs, sometimes called ''moon crabs'', adaptation, adapted for swimming or digging. They differ from the swimming crabs of the family Portunidae in that all five pairs of arthropod leg, legs are flatten ...
, are also capable of swimming, the
Portunidae Portunidae is a family of crabs which contains the swimming crabs. Its members include well-known shoreline crabs such as the blue crab (''Callinectes sapidus'') and velvet crab ('' Necora puber''). Description Portunid crabs are characterised b ...
especially so as their last pair of walking legs are flattened into swimming paddles. Crabs are mostly active animals with complex behaviour patterns such as communicating by drumming or waving their pincers. Crabs tend to be aggressive toward one another, and males often fight to gain access to females. On rocky seashores, where nearly all caves and crevices are occupied, crabs may also fight over hiding holes.
Fiddler crab The fiddler crab or calling crab can be one of the hundred species of semiterrestrial marine crabs in the family Ocypodidae. These crabs are well known for their extreme sexual dimorphism, where the male crabs have a major claw significantly l ...
s (genus ''Uca'') dig burrows in sand or mud, which they use for resting, hiding, and mating, and to defend against intruders. Crabs are
omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
s, feeding primarily on
alga Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular microalgae, suc ...
e, and taking any other food, including
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,
worm Worms are many different distantly related bilateria, bilateral animals that typically have a long cylindrical tube-like body, no limb (anatomy), limbs, and usually no eyes. Worms vary in size from microscopic to over in length for marine ...
s, other
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,
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 ...
,
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 ...
, and
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
, depending on their availability and the crab species. For many crabs, a mixed diet of plant and animal matter results in the fastest growth and greatest fitness. Some species are more specialised in their diets, based in plankton, clams or fish. Some crabs, such as members of
Pinnotheridae Pinnotheridae is a family of tiny soft-bodied crabs that live commensally in the mantles of certain bivalve molluscs and the occasional large gastropod mollusc species in genera such as ''Strombus'' and ''Haliotis''. '' Tunicotheres moseri'' i ...
, live on or inside animals such as
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 ...
, either being
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit f ...
(benign) or
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The ent ...
on their host. Crabs are known to work together to provide food and protection for their family, and during mating season to find a comfortable spot for the female to release her eggs.


Evolution

The earliest unambiguous crab
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s date from the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
, with the oldest being '' Eocarcinus'' from the early
Pliensbachian The Pliensbachian is an age of the geologic timescale and stage in the stratigraphic column. It is part of the Early or Lower Jurassic Epoch or Series and spans the time between 192.9 ±0.3 Ma and 184.2 ±0.3 Ma (million years ago). The Plie ...
of Britain, which likely represents a stem-group lineage, as it lacks several key morphological features that define modern crabs. Most Jurassic crabs are only known from
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
(top half of the body)
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 ...
s, making it difficult to determine their relationships. Crabs radiated in the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
, corresponding with an increase in reef habitats, though they would decline at the end of the Jurassic as the result of the decline of reef ecosystems. Crabs increased in diversity through the Cretaceous and represented the dominant group of
decapods The Decapoda or decapods, from Ancient Greek δεκάς (''dekás''), meaning "ten", and πούς (''poús''), meaning "foot", is a large order (biology), order of crustaceans within the class Malacostraca, and includes crabs, lobsters, crayfis ...
by the end of the period. The crab
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
Brachyura belongs to the group
Reptantia Reptantia is a clade of decapod crustaceans named in 1880 which includes lobsters, crabs and many other well-known crustaceans. Classification In older classifications, Reptantia was one of the two suborders of Decapoda alongside Natantia, w ...
, which consists of the walking/crawling decapods (
lobsters Lobsters are malacostracans decapod crustaceans of the family Nephropidae or its synonym Homaridae. They have long bodies with muscular tails and live in crevices or burrows on the sea floor. Three of their five pairs of legs have claws, in ...
and crabs). Brachyura is the
sister clade In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to the infraorder
Anomura Anomura (sometimes Anomala) is a group of decapod crustaceans, including hermit crabs and others. Although the names of many anomurans include the word ''crab'', all true crabs are in the sister group to the Anomura, the Brachyura (the two groups ...
, which contains the hermit crabs and relatives. The
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below shows Brachyura's placement within the larger
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Decapoda The Decapoda or decapods, from Ancient Greek δεκάς (''dekás''), meaning "ten", and πούς (''poús''), meaning "foot", is a large order of crustaceans within the class Malacostraca, and includes crabs, lobsters, crayfish, shrimp, a ...
, from analysis by Wolfe ''et al.'', 2019. Brachyura is separated into several sections, with the basal
Dromiacea Dromiacea is a group of crabs, ranked as a section. It contains 240 extant and nearly 300 extinct species. Dromiacea is the most basal grouping of Brachyura crabs, diverging the earliest in the evolutionary history, around the Late Triassic o ...
diverging the earliest in the evolutionary history, around the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
or
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
. The group consisting of
Raninoida Raninoida is a taxonomic section of the crabs, containing a single superfamily, Raninoidea. This group of crabs is unlike most, with the abdomen not being folded under the thorax. It comprises 46 extant species, and nearly 200 species known o ...
and
Cyclodorippoida Cyclodorippoida is a group of crabs, ranked as a section. It contains the single superfamily Cyclodorippoidea, which holds three families, Cyclodorippidae, Cymonomidae and Phyllotymolinidae. Below is a cladogram A cladogram (from Greek ...
split off next, during the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period. The remaining
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Eubrachyura then divided during the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period into Heterotremata and Thoracotremata. A summary of the high-level internal relationships within Brachyura can be shown in the
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below: There is a no consensus on the relationships of the subsequent superfamily (taxonomy), superfamilies and family (biology), families. The proposed
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below is from analysis by Tsang ''et al'', 2014:


Classification

The
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
Brachyura contains approximately 7,000 species in 98 family (biology), families, as many as the remainder of the
Decapoda The Decapoda or decapods, from Ancient Greek δεκάς (''dekás''), meaning "ten", and πούς (''poús''), meaning "foot", is a large order of crustaceans within the class Malacostraca, and includes crabs, lobsters, crayfish, shrimp, a ...
. The evolution of crabs is characterized by an increasingly robust body, and a reduction in the
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
. Although many other groups have undergone similar processes,
carcinisation Carcinisation (American English: carcinization) is a form of convergent evolution in which non-crab crustaceans evolve a crab-like body plan. The term was introduced into evolutionary biology by Lancelot Alexander Borradaile, who described it a ...
is most advanced in crabs. The telson is no longer functional in crabs, and the uropods are absent, having probably evolved into small devices for holding the reduced abdomen tight against the Sternum (arthropod), sternum. In most decapods, the gonopores (sexual openings) are found on the legs. Since crabs use their first two pairs of
pleopod The anatomy of a decapod consists of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment – often called a somite – may possess one pair of appendages, although in various groups these m ...
s (abdominal appendages) for sperm transfer, this arrangement has changed. As the male abdomen evolved into a slimmer shape, the gonopores have moved toward the midline, away from the legs, and onto the sternum. A similar change occurred, independently, with the female gonopores. The movement of the female gonopore to the sternum defines the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Eubrachyura, and the later change in the position of the male gonopore defines the Thoracotremata. It is still a subject of debate whether a Monophyly, monophyletic group is formed by those crabs where the female, but not male, gonopores are situated on the sternum.


Families

Numbers of extant and extinct (†) species are given in brackets. The superfamily Eocarcinoidea, containing '' Eocarcinus'' and ''Platykotta'', was formerly thought to contain the oldest crabs; it is now considered part of the
Anomura Anomura (sometimes Anomala) is a group of decapod crustaceans, including hermit crabs and others. Although the names of many anomurans include the word ''crab'', all true crabs are in the sister group to the Anomura, the Brachyura (the two groups ...
. * Section †Callichimaeroida ** †Callichimaeroidea (1†) * Section
Dromiacea Dromiacea is a group of crabs, ranked as a section. It contains 240 extant and nearly 300 extinct species. Dromiacea is the most basal grouping of Brachyura crabs, diverging the earliest in the evolutionary history, around the Late Triassic o ...
** †Dakoticancroidea (6†) ** Dromioidea (147, 85†) ** Glaessneropsoidea (45†) ** Homolodromioidea (24, 107†) ** Homoloidea (73, 49†) * Section
Raninoida Raninoida is a taxonomic section of the crabs, containing a single superfamily, Raninoidea. This group of crabs is unlike most, with the abdomen not being folded under the thorax. It comprises 46 extant species, and nearly 200 species known o ...
(46, 196†) * Section
Cyclodorippoida Cyclodorippoida is a group of crabs, ranked as a section. It contains the single superfamily Cyclodorippoidea, which holds three families, Cyclodorippidae, Cymonomidae and Phyllotymolinidae. Below is a cladogram A cladogram (from Greek ...
(99, 27†) * Section Eubrachyura ** Subsection Heterotremata *** Aethroidea (37, 44†) *** Bellioidea (7) *** Bythograeoidea (14) *** Calappoidea (101, 71†) *** Cancroidea (57, 81†) *** Carpilioidea (4, 104†) *** Cheiragonoidea (3, 13†) *** Corystoidea (10, 5†) *** †Componocancroidea (1†) *** Dairoidea (4, 8†) *** Dorippoidea (101, 73†) *** Eriphioidea (67, 14†) *** Gecarcinucoidea (349) *** Goneplacoidea (182, 94†) *** Hexapodoidea (21, 25†) *** Leucosioidea (488, 113†) *** Majoidea (980, 89†) *** Orithyioidea (1) *** Palicoidea (63, 6†) *** Parthenopoidea (144, 36†) *** Pilumnoidea (405, 47†) *** Portunoidea (455, 200†) *** Potamoidea (662, 8†) *** Pseudothelphusoidea (276) *** Pseudozioidea (22, 6†) *** Retroplumoidea (10, 27†) *** Trapezioidea (58, 10†) *** Trichodactyloidea (50) *** Xanthoidea (736, 134†) ** Subsection Thoracotremata *** Cryptochiroidea (46) *** Grapsoidea (493, 28†) *** Ocypodoidea (304, 14†) *** Pinnotheroidea (304, 13†) Recent studies have found the following superfamily (taxonomy), superfamilies and family (biology), families to not be monophyletic, but rather paraphyletic or polyphyletic: * The Thoracotremata superfamily Grapsoidea is polyphyletic * The Thoracotremata superfamily Ocypodoidea is polyphyletic * The Heterotremata superfamily Calappoidea is polyphyletic * The Heterotremata superfamily Eriphioidea is polyphyletic * The Heterotremata superfamily Goneplacoidea is polyphyletic * The Heterotremata superfamily Potamoidea is paraphyletic with respect to Gecarcinucoidea, which is resolved by placing Gecarcinucidae within Potamoidea * The Majoidea families Epialtidae, Mithracidae and Majidae are polyphyletic with respect to each other * The Dromioidea family Dromiidae may be paraphyletic with respect to Dynomenidae * The Homoloidea family Homolidae is paraphyletic with respect to Latreilliidae * The Xanthoidea family Xanthidae is paraphyletic with respect to Panopeidae


Human consumption


Fisheries

Crabs make up 20% of all marine
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 caught, farmed, and consumed worldwide, amounting to 1.5 million tonnes annually. One species, ''Portunus trituberculatus'', accounts for one-fifth of that total. Other commercially important taxon, taxa include ''Portunus pelagicus'', several species in the genus ''Chionoecetes'', the blue crab (''Callinectes sapidus''), Charybdis (crab), ''Charybdis'' spp., ''Edible crab, Cancer pagurus'', the Dungeness crab (''Metacarcinus magister''), and ''Scylla serrata'', each of which yields more than 20,000 tonnes annually. In some crab species, meat is harvested by manually twisting and Declawing of crabs, pulling off one or both claws and returning the live crab to the water in the knowledge that the crab may survive and regenerate the claws.


Crabs as food

Crabs are prepared and eaten as a dish in many different ways all over the world. Some species can be eaten whole, including the shell, as soft-shell crabs; with other species, just the claws or legs are eaten. The latter is particularly common for larger crabs, such as the snow crab. In many cultures, the roe of the female crab is also eaten, which usually appears orange or yellow in fertile crabs. This is popular in Southeast Asian cultures, some Mediterranean and Northern European cultures, and on the East, Chesapeake Bay, Chesapeake, and Gulf Coasts of the United States. In some regions, spices improve the culinary experience. In Southeast Asia and the Indosphere, spice mix, masala crab and chilli crab are examples of heavily spiced dishes. In the Chesapeake Bay region, blue crab is often steamed with Old Bay Seasoning. Alaskan king crab or snow crab legs are usually simply boiled and served with garlic or lemon butter. For the British dish Cancer pagurus#Cookery, dressed crab, the crab meat is extracted and placed inside the hard shell. One American way to prepare crab meat is by extracting it and adding varying amounts of binders, such as egg white, cracker meal, mayonnaise, or mustard, creating a crab cake. Crabs can also be made into a bisque (food), bisque, a global dish of French origin which in its authentic form includes in the broth the pulverized shells of the shellfish from which it is made. Crab stick, Imitation crab, also called surimi, is made from minced fish meat that is crafted and colored to resemble crab meat. While it is sometimes disdained among some elements of the culinary industry as an unacceptably low-quality substitute for real crab, this does not hinder its popularity, especially as a sushi ingredient in Japan and South Korea, and in home cooking, where cost is often a chief concern.Daniel P. Puzo (February 14, 1985
Imitation Crab Draws Criticisms
''Los Angeles Times''
Indeed, surimi is an important source of protein in most East and Southeast Asian cultures, appearing in staple ingredients such as fish balls and fish cake.


Pain

Whether crustaceans as a whole experience pain or not is a scientific debate that has ethical implications for crab dish preparation. Crabs are often boiled alive as part of the cooking process.


Cultural influences

Both the constellation Cancer (constellation), Cancer and the astrological sign Cancer (astrology), Cancer are named after the crab, and depicted as a crab. William Parsons, 3rd Earl of Rosse drew the Crab Nebula in 1848 and noticed its similarity to the animal; the Crab Pulsar lies at the centre of the nebula. The Moche (culture), Moche people of ancient Peru worshipped nature, especially the sea, and often depicted crabs in their art. In Greek mythology, Karkinos was a crab that came to the aid of the Lernaean Hydra as it battled Heracles. One of Rudyard Kipling's ''Just So Stories'', ''The Crab that Played with the Sea'', tells the story of a gigantic crab who made the waters of the sea go up and down, like the tides. In Malay people, Malay mythology (as related by Hugh Clifford (colonial administrator), Hugh Clifford to Walter William Skeat (anthropologist), Walter William Skeat), ocean tides are believed to be caused by water rushing in and out of a hole in the Navel of the Seas (''Pusat Tasek''), where "there sits a gigantic crab which twice a day gets out in order to search for food". The Kapsiki people of North Cameroon use the way crabs handle objects for divination. The term ''crab mentality'' is derived from a type of detrimental social behavior observed in crabs.


Explanatory notes


References


External links

* {{Authority control Crabs, Articles containing video clips Commercial crustaceans Edible crustaceans Extant Jurassic first appearances