Crustaceans form a very large group of
arthropodAn arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s, usually treated as a
subphylumIn life, a subphylum is a taxonomic rank intermediate between phylum and superclass. The rank of subdivision in plants and fungi is equivalent to subphylum.Not all phyla are divided into subphyla...
, which includes such familiar animals as
crabTrue crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
s,
lobsterClawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...
s,
crayfishCrayfish, crawfish, or crawdads – members of the superfamilies Astacoidea and Parastacoidea – are freshwater crustaceans resembling small lobsters, to which they are related...
,
shrimpShrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
,
krillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
and
barnacleA barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have...
s. The 50,000 described species range in size from
Stygotantulus stockiStygotantulus stocki is a species of crustacean, living as an ectoparasite on harpacticoid copepods of the families Tisbidae and Canuellidae. It is the smallest arthropod in the world, at a length of less than . The specific epithet stocki commemorates Jan Hendrik Stock, a Dutch carcinologist....
at 0.1 mm (0.00393700787401575 in), to the
Japanese spider crabThe , Macrocheira kaempferi, is a species of marine crab that lives in the waters around Japan. It has the largest leg span of any arthropod, reaching up to and weighing up to . It is the subject of small-scale fishery.-Description:...
with a leg span of up to 12.5 ft (3.8 m) and a mass of 44 lb (20 kg). Like other
arthropodAn arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s, crustaceans have an
exoskeletonAn exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, which they
moultEcdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...
to grow. They are distinguished from other groups of arthropods, such as
insectInsects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s,
myriapodsMyriapoda is a subphylum of arthropods containing millipedes, centipedes, and others. The group contains 13,000 species, all of which are terrestrial...
and
cheliceratesThe subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites...
, by the possession of biramous (two-parted) limbs, and by the nauplius form of the
larvaeCrustaceans 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...
.
Most crustaceans are free-living
aquatic animalAn aquatic animal is an animal, either vertebrate or invertebrate, which lives in water for most or all of its life. It may breathe air or extract its oxygen from that dissolved in water through specialised organs called gills, or directly through its skin. Natural environments and the animals that...
s, but some are
terrestrialTerrestrial animals are animals that live predominantly or entirely on land , as compared with aquatic animals, which live predominantly or entirely in the water , or amphibians, which rely on a combination of aquatic and terrestrial habitats...
(e.g.
woodliceA woodlouse is a crustacean with a rigid, segmented, long exoskeleton and fourteen jointed limbs...
), some are
parasiticParasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
(e.g. fish lice,
tongue wormsPentastomida are an enigmatic group of parasitic invertebrates commonly known as tongue worms due to the resemblance of the species of the genus Linguatula to a vertebrate tongue....
) and some are
sessileIn zoology, sessility is a characteristic of animals which are not able to move about. They are usually permanently attached to a solid substrate of some kind, such as a part of a plant or dead tree trunk, a rock, or the hull of a ship in the case of barnacles. Corals lay down their own...
(e.g.
barnacleA barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have...
s). The group has an extensive fossil record, reaching back to the
CambrianThe Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
, and includes
living fossilLiving fossil is an informal term for any living species which appears similar to a species otherwise only known from fossils and which has no close living relatives, or a group of organisms which have long fossil records...
s such as
Triops cancriformisTriops cancriformis, or tadpole shrimp , is a species of tadpole shrimp found in Europe, the Middle East and Japan....
, which has existed apparently unchanged since the
TriassicThe Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...
period. More than 10 million tons of crustaceans are produced by fishery or farming for human consumption, the majority of it being
shrimpShrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
s and
prawnPrawns are decapod crustaceans of the sub-order Dendrobranchiata. There are 540 extant species, in seven families, and a fossil record extending back to the Devonian...
s.
KrillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
and
copepodCopepods are a group of small crustaceans found in the sea and nearly every freshwater habitat. Some species are planktonic , some are benthic , and some continental species may live in limno-terrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests,...
s are not as widely fished, but may be the animals with the greatest
biomassBiomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...
on the planet, and form a vital part of the food chain. The scientific study of crustaceans is known as carcinology (alternatively, malacostracology, crustaceology or crustalogy), and a
scientistA scientist in a broad sense is one engaging in a systematic activity to acquire knowledge. In a more restricted sense, a scientist is an individual who uses the scientific method. The person may be an expert in one or more areas of science. This article focuses on the more restricted use of the word...
who works in carcinology is a
carcinologist.
Structure
The body of a crustacean is composed of body segments, which are grouped into three regions: the
cephalon or
headIn anatomy, the head of an animal is the rostral part that usually comprises the brain, eyes, ears, nose and mouth . Some very simple animals may not have a head, but many bilaterally symmetric forms do....
, the
thoraxThe thorax is a division of an animal's body that lies between the head and the abdomen.-In tetrapods:...
, and the
pleon or
abdomenIn vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
. The head and thorax may be fused together to form a
cephalothoraxThe cephalothorax is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. The word cephalothorax is derived from the Greek words for head and thorax...
, which may be covered by a single large
carapaceA carapace is a dorsal 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 underside is called the plastron.-Crustaceans:In crustaceans, the...
. The crustacean body is protected by the hard
exoskeletonAn exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
, which must be
moultedEcdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...
for the animal to grow. The shell around each somite can be divided into a dorsal
tergumA tergum is the dorsal portion of an arthropod segment other than the head. The anterior edge is called the base and posterior edge is called the apex or margin. A given tergum may be divided into hardened plates or sclerites commonly referred to as tergites...
, ventral sternum and a lateral pleuron. Various parts of the exoskeleton may be fused together.
Each
somiteA somite is a division of the body of an animal. In vertebrates this is mainly discernible in the embryo stage; in arthropods it is a characteristic of a hypothetical ancestor.- In vertebrates :...
, or body segment can bear a pair of
appendageIn invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s: on the segments of the head, these include two pairs of
antennaeAntennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
, the mandibles and
maxillaThe maxilla is a fusion of two bones along the palatal fissure that form the upper jaw. This is similar to the mandible , which is also a fusion of two halves at the mental symphysis. Sometimes The maxilla (plural: maxillae) is a fusion of two bones along the palatal fissure that form the upper...
e; the thoracic segments bear
legsThe arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments are of Latin origin, and may be confused with terms for bones: coxa , trochanter , femur, tibia, tarsus, ischium, metatarsus, carpus, dactylus ,...
, which may be specialised as pereiopods (walking legs) and maxillipeds (feeding legs). The abdomen bears pleopods, and ends in a
telsonThe telson is the last division of the body of a crustacean. It is not considered a true segment because it does not arise in the embryo from teloblast areas as do real segments. It never carries any appendages, but a forked "tail" called the caudal furca is often present. Together with the...
, which bears the
anusThe anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
, and is often flanked by uropods to form a
tail fanTail fan can refer to:*Tail fan of a decapod*Tail fan of a bird...
. The number and variety of
appendageIn invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s in different crustaceans may be partly responsible for the group's success. Crustacean
appendageIn invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s are typically biramous, meaning they are divided into two parts; this includes the second pair of antennae, but not the first, which is uniramous. It is unclear whether the biramous condition is a derived state which evolved in crustaceans, or whether the second branch of the limb has been lost in all other groups.
TrilobiteTrilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...
s, for instance, also possessed biramous appendages.
The main body cavity is an open circulatory system, where blood is pumped into the haemocoel by a
heartThe heart is a myogenic muscular organ found in all animals with a circulatory system , that is responsible for pumping blood throughout the blood vessels by repeated, rhythmic contractions...
located near the dorsum. Malacostraca have haemocyanin as the oxygen-carrying pigment, while copepods, ostracods, barnacles and branchiopods have haemoglobins. The alimentary canal consists of a straight tube that often has a gizzard-like "gastric mill" for grinding food and a pair of digestive glands that absorb food; this structure goes in a spiral format. Structures that function as kidneys are located near the antennae. A brain exists in the form of ganglia close to the antennae, and a collection of major ganglia is found below the gut.
In many
decapodsThe decapods or Decapoda are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crayfish, crabs, lobsters, prawns and shrimp. Most decapods are scavengers. It is estimated that the order contains nearly 15,000 species in around 2,700 genera, with...
, the first (and sometimes the second) pair of pleopods are specialised in the male for sperm transfer. Many terrestrial crustaceans (such as the
Christmas Island red crabThe Christmas Island red crab, Gecarcoidea natalis, is a species of land crab that is endemic to Christmas Island and the Cocos Islands in the Indian Ocean...
) mate seasonally and return to the sea to release the eggs. Others, such as
woodliceA woodlouse is a crustacean with a rigid, segmented, long exoskeleton and fourteen jointed limbs...
, lay their eggs on land, albeit in damp conditions. In most decapods, the females retain the eggs until they hatch into free-swimming larvae.
Ecology
The majority of crustaceans are aquatic, living in either
marineMarine is an umbrella term. As an adjective it is usually applicable to things relating to the sea or ocean, such as marine biology, marine ecology and marine geology...
or
fresh waterFresh Water is the debut album by Australian rock and blues singer Alison McCallum, released in 1972. Rare for an Australian artist at the time, it came in a gatefold sleeve...
environments, but a few groups have
adaptedAn adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....
to life on land, such as
terrestrial crabGecarcinidae is a family of true crabs that are adapted for terrestrial existence, commonly known as land crabs. Similar to all other crabs, land crabs possess a series of gills. In addition, the part of the carapace covering the gills is inflated and equipped with blood vessels. These organs...
s,
terrestrial hermit crabThe genus Coenobita contains the sixteen species of terrestrial hermit crabs. The majority of the species are found in the Indo-Pacific region, with only one species in West Africa, one species occurring along the western Atlantic Ocean, and one species occurring on the Pacific coast of the Americas...
s, and
woodliceA woodlouse is a crustacean with a rigid, segmented, long exoskeleton and fourteen jointed limbs...
. Marine crustaceans are as ubiquitous in the oceans as insects are on land. The majority of crustaceans are also
motileMotility is a biological term which refers to the ability to move spontaneously and actively, consuming energy in the process. Most animals are motile but the term applies to single-celled and simple multicellular organisms, as well as to some mechanisms of fluid flow in multicellular organs, in...
, moving about independently, although a few taxonomic units are parasitic and live attached to their hosts (including sea lice,
fish liceThe family Argulidae contains the carp lice or fish lice – a group of parasitic crustaceans of uncertain position within the Maxillopoda. Although they are thought to be primitive forms, they have no fossil record...
,
whale liceA whale louse is a parasitic crustacean of the family Cyamidae. They are related to the better-known skeleton shrimp, most species of which are found in shallower waters. Whale lice are external parasites, found in skin lesions, genital folds, nostrils and eyes of marine mammals of the order Cetacea...
,
tongue wormsPentastomida are an enigmatic group of parasitic invertebrates commonly known as tongue worms due to the resemblance of the species of the genus Linguatula to a vertebrate tongue....
, and
Cymothoa exiguaCymothoa exigua, or the tongue-eating louse, is a parasitic crustacean of the family Cymothoidae. It tends to be long. This parasite enters through the gills, and then attaches itself at the base of the spotted rose snapper's tongue. It extracts blood through the claws on its front, causing the...
, all of which may be referred to as "crustacean lice"), and adult barnacles live a
sessileIn zoology, sessility is a characteristic of animals which are not able to move about. They are usually permanently attached to a solid substrate of some kind, such as a part of a plant or dead tree trunk, a rock, or the hull of a ship in the case of barnacles. Corals lay down their own...
life – they are attached headfirst to the substrate and cannot move independently. Some branchiurans are able to withstand rapid changes of
salinitySalinity is the saltiness or dissolved salt content of a body of water. It is a general term used to describe the levels of different salts such as sodium chloride, magnesium and calcium sulfates, and bicarbonates...
and will also switch hosts from marine to non-marine species.
KrillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
are the bottom layer and the most important part of the food chain in
AntarcticThe Antarctic is the region around the Earth's South Pole, opposite the Arctic region around the North Pole. The Antarctic comprises the continent of Antarctica and the ice shelves, waters and island territories in the Southern Ocean situated south of the Antarctic Convergence...
animal communities. Some crustaceans are significant invasive species, such as the
Chinese mitten crabThe Chinese mitten crab, Eriocheir sinensis is a medium-sized burrowing crab, named for its furry claws that look like mittens, that is native in the coastal estuaries of eastern Asia from Korea in the north to the Fujian province of China in the south...
and the Asian shore crab.
Life cycle
Mating system
The majority of crustaceans have
separate sexesIn biology, gonochorism or unisexualism describes sexually reproducing species in which individuals have just one of at least two distinct sexes. The term is most often used with animals . The sex of an individual may change during its lifetime, this can for example be found in parrotfish...
, and
reproduce sexuallySexual reproduction is the creation of a new organism by combining the genetic material of two organisms. There are two main processes during sexual reproduction; they are: meiosis, involving the halving of the number of chromosomes; and fertilization, involving the fusion of two gametes and the...
. A small number are
hermaphroditeIn biology, a hermaphrodite is an organism that has reproductive organs normally associated with both male and female sexes.Many taxonomic groups of animals do not have separate sexes. In these groups, hermaphroditism is a normal condition, enabling a form of sexual reproduction in which both...
s, including
barnacleA barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have...
s, remipedes, and
CephalocaridaCephalocarida is a class inside the subphylum Crustacea that comprises only twelve shrimp-like benthic species. They were discovered in 1955 by Howard L. Sanders, and are commonly referred to as horseshoe shrimps. They have been grouped together with the Remipedia in the Xenocarida...
. Some may even change sex during the course of their life.
ParthenogenesisParthenogenesis is a form of asexual reproduction found in females, where growth and development of embryos occur without fertilization by a male...
is also widespread among crustaceans, where viable eggs are produced by a female without needing fertilisation by a male. This occurs in many
brachiopodBrachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...
s, some
ostracodOstracoda is a class of the Crustacea, sometimes known as the seed shrimp because of their appearance. Some 65,000 species have been identified, grouped into several orders....
s, some isopods, and certain "higher" crustaceans, such as the
Marmorkrebs', or marbled crayfish, are parthenogenetic crayfish that were discovered in the pet trade in Germany in the 1990s. are closely related to the "slough crayfish", Procambarus fallax. Procambarus fallax is widely distributed across Florida, but there are no known natural populations of Marmorkrebs...
crayfish.
Eggs
In many groups of crustaceans, the fertilised eggs are simply released into the water column, while others have developed a number of mechanisms for holding on to the eggs until they are ready to hatch. Most
decapodsThe decapods or Decapoda are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crayfish, crabs, lobsters, prawns and shrimp. Most decapods are scavengers. It is estimated that the order contains nearly 15,000 species in around 2,700 genera, with...
carry the eggs attached to the pleopods, while
peracaridsThe superorder Peracarida is a large group of malacostracan crustaceans, having members in marine, freshwater, and terrestrial habitats. They are chiefly defined by the presence of a brood pouch, or marsupium, formed from thin flattened plates borne on the basalmost segments of the legs Other...
,
notostracaThe order Notostraca comprises the single family Triopsidae, containing the tadpole shrimp or shield shrimp. The two genera, Triops and Lepidurus, are considered living fossils, having not changed significantly in outward form since the Triassic. They have a broad, flat carapace, which conceals the...
ns, anostracans, and many isopods form a
brood pouchThe marsupium or brood pouch, is a characteristic feature of Peracarida, including the orders Amphipoda, Isopoda and Cumacea. It is an egg chamber formed by oostegites, which are appendices which are attached to the coxae of the first pereiopods...
from the
carapaceA carapace is a dorsal 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 underside is called the plastron.-Crustaceans:In crustaceans, the...
and thoracic limbs. Female Branchiura do not carry eggs in external ovisacs but attach them in rows to rocks and other objects. Most
leptostracaLeptostraca is an order of small, marine crustaceans. Its members, including the well-studied Nebalia, occur throughout the world's oceans and are usually considered to be filter-feeders. It is the only extant order in the subclass Phyllocarida...
ns and
krillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
carry the eggs between their thoracic limbs; some
copepodCopepods are a group of small crustaceans found in the sea and nearly every freshwater habitat. Some species are planktonic , some are benthic , and some continental species may live in limno-terrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests,...
s carry their eggs in special thin-walled sacs, while others have them attached together in long, tangled strings.
Larvae
Crustaceans exhibit a number of larval forms, of which the earliest and most characteristic is the nauplius. This has three pairs of
appendageIn invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s, all emerging from the young animal's head, and a single naupliar eye. In most groups, there are further larval stages, including the zoea (pl. zoeæ or zoeas). This name was given to it when naturalists believed it to be a separate species. It follows the nauplius stage and precedes the post-larva. Zoea larvae swim with their thoracic
appendageIn invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s, as opposed to nauplii, which use cephalic appendages, and megalopa, which use abdominal appendages for swimming. It often has spikes on its
carapaceA carapace is a dorsal 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 underside is called the plastron.-Crustaceans:In crustaceans, the...
, which may assist these small organisms in maintaining directional swimming. In many
decapodsThe decapods or Decapoda are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crayfish, crabs, lobsters, prawns and shrimp. Most decapods are scavengers. It is estimated that the order contains nearly 15,000 species in around 2,700 genera, with...
, due to their accelerated development, the zoea is the first larval stage. In some cases, the zoea stage is followed by the mysis stage, and in others, by the megalopa stage, depending on the crustacean group involved.
Classification
The name "crustacean" dates from the earliest works to describe the animals, including those of
Pierre BelonPierre Belon was a French naturalist. He is sometimes known as Pierre Belon du Mans, or, in Latin translations of his works, as Petrus Bellonius Cenomanus.Belon was born in 1517 at Soulletiere near Cérans-Foulletourte...
and
Guillaume RondeletGuillaume Rondelet , known also as Rondeletus , was Regus Professor of medicine at the University of Montpellier in southern France and Chancellor of the University between 1556 and his death in 1566. He achieved renown as an anatomist and a naturalist with a particular interest in botany and zoology...
, but the name was not used by some later authors, including Carl Linnaeus, who included crustaceans among the "
ApteraIn the 10th edition of Systema Naturae, Carl Linnaeus classified the arthropods, including insects, arachnids and crustaceans, among his class "Insecta"...
" in his
. The earliest nomenclaturally valid work to use the name "Crustacea" was
Morten Thrane Brünnichthumb|180px|Morten Thrane Brünnich, painting by [[Jens Juel |Jens Juel]], 1799Morten Thrane Brünnich was a Danish zoologist and mineralogist....
's
in 1772, although he also included
cheliceratesThe subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites...
in the group.
The subphylum Crustacea comprises almost 52,000 described
speciesIn biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
, although the number of undescribed species may be 10–100 times higher. Although most crustaceans are small, their morphology varies greatly and they include both the largest arthropod in the world – the
Japanese spider crabThe , Macrocheira kaempferi, is a species of marine crab that lives in the waters around Japan. It has the largest leg span of any arthropod, reaching up to and weighing up to . It is the subject of small-scale fishery.-Description:...
with a leg span of 14 feet (4.3 m) – and the smallest – the 0.1 mm (0.00393700787401575 in) long
Stygotantulus stockiStygotantulus stocki is a species of crustacean, living as an ectoparasite on harpacticoid copepods of the families Tisbidae and Canuellidae. It is the smallest arthropod in the world, at a length of less than . The specific epithet stocki commemorates Jan Hendrik Stock, a Dutch carcinologist....
. Despite their diversity of form, crustaceans are united by the special
larvaA larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
l form known as the nauplius.
The exact relationships of the Crustacea to other taxa are not yet entirely clear. Under the
PancrustaceaPancrustacea is a clade, comprising all crustaceans and hexapods. This grouping is contrary to the Atelocerata hypothesis, in which Myriapoda and Hexapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon is considered well accepted. The clade has...
hypothesis, Crustacea and
HexapodaThe subphylum Hexapoda constitutes the largest grouping of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura . The Collembola are very abundant in terrestrial environments...
(
insectInsects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s and allies) are sister groups. Studies using
DNA sequenceThe sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
s tend to show a paraphyletic Crustacea, with the
insectInsects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s (but not necessarily other hexapods) nested within that
cladeA clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. Although the classification of crustaceans has been quite variable, the system used by Martin and Davis is the most authoritative, and largely supersedes earlier works.
MystacocaridaMystacocarida is a subclass of crustaceans, that form part of the meiobenthos. They are less than long, and live interstitially in the intertidal zones of sandy beaches.-Taxonomy:...
and
BranchiuraThe family Argulidae contains the carp lice or fish lice – a group of parasitic crustaceans of uncertain position within the Maxillopoda. Although they are thought to be primitive forms, they have no fossil record...
, here treated as part of
MaxillopodaMaxillopoda is a diverse class of crustaceans including barnacles, copepods and a number of related animals. It does not appear to be a monophyletic group, and no single character unites all the members.-Description:...
, are sometimes treated as their own classes. Six classes are usually recognised:


| Class | Members | Orders | Photo |
BranchiopodaBranchiopoda is a class of crustaceans. It is the sister group to the remaining crustaceans, and comprises fairy shrimp, clam shrimp, Cladocera, Notostraca and the Devonian Lepidocaris... |
brine shrimpArtemia is a genus of aquatic crustaceans known as brine shrimp. Artemia, the only genus in the family Artemiidae, has changed little externally since the Triassic period...
CladoceraCladocera is an order of small crustaceans commonly called water fleas. Around 620 species have been recognised so far, with many more undescribed. They are ubiquitous in inland aquatic habitats, but rare in the oceans. Most are long, with a down-turned head, and a carapace covering the apparently...
TriopsTriops is a genus of small crustaceans in the order Notostraca . They are considered living fossils, with a fossil record that reaches back to the Carboniferous, .-Relatives and fossil record:... |
AnostracaAnostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are also known as fairy shrimp. They are usually long . Most species have 20 body segments, bearing 11 pairs of leaf-like phyllopodia , and the body lacks a carapace...
Lipostraca NotostracaThe order Notostraca comprises the single family Triopsidae, containing the tadpole shrimp or shield shrimp. The two genera, Triops and Lepidurus, are considered living fossils, having not changed significantly in outward form since the Triassic. They have a broad, flat carapace, which conceals the...
Laevicaudata Spinicaudata Cyclestherida CladoceraCladocera is an order of small crustaceans commonly called water fleas. Around 620 species have been recognised so far, with many more undescribed. They are ubiquitous in inland aquatic habitats, but rare in the oceans. Most are long, with a down-turned head, and a carapace covering the apparently... |
Daphnia pulexDaphnia pulex is the most common species of water flea. It has a cosmopolitan distribution: the species is found throughout the Americas, Europe and Australia. It is a model species, and was the first crustacean to have its genome sequenced.-Description:... (CladoceraCladocera is an order of small crustaceans commonly called water fleas. Around 620 species have been recognised so far, with many more undescribed. They are ubiquitous in inland aquatic habitats, but rare in the oceans. Most are long, with a down-turned head, and a carapace covering the apparently... ) |
| Remipedia Remipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, the Atlantic Ocean... |
|
Nectiopoda |
|
| Cephalocarida Cephalocarida is a class inside the subphylum Crustacea that comprises only twelve shrimp-like benthic species. They were discovered in 1955 by Howard L. Sanders, and are commonly referred to as horseshoe shrimps. They have been grouped together with the Remipedia in the Xenocarida... |
horseshoe shrimp |
Brachypoda |
|
MaxillopodaMaxillopoda is a diverse class of crustaceans including barnacles, copepods and a number of related animals. It does not appear to be a monophyletic group, and no single character unites all the members.-Description:... |
barnacleA barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have... s copepodCopepods are a group of small crustaceans found in the sea and nearly every freshwater habitat. Some species are planktonic , some are benthic , and some continental species may live in limno-terrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests,... s |
CalanoidaCalanoida is an order of copepods, a kind of zooplankton. They include around 40 families with about 1800 species of both marine and freshwater copepods. Calanoid copepods are dominant in the plankton in many parts of the world's oceans, making up 55%–95% of plankton samples...
Pedunculata SessiliaSessilia is an order of barnacles, comprising the barnacles without stalks, or acorn barnacles. They form a monophyletic group and are probably derived from stalked barnacles. The order is divided into three suborders. Brachylepadomorpha contains a single family, Neobrachylepadidae, while...
c. 20 others |
Chthamalus stellatusChthamalus stellatus, common name Poli's stellate barnacle, is a species of acorn barnacle common on rocky shores in South West England, Ireland, and Southern Europe. It is named after Giuseppe Saverio Poli.-Description:... (SessiliaSessilia is an order of barnacles, comprising the barnacles without stalks, or acorn barnacles. They form a monophyletic group and are probably derived from stalked barnacles. The order is divided into three suborders. Brachylepadomorpha contains a single family, Neobrachylepadidae, while... ) |
| Ostracoda |
ostracodOstracoda is a class of the Crustacea, sometimes known as the seed shrimp because of their appearance. Some 65,000 species have been identified, grouped into several orders.... s |
MyodocopidaThe Myodocopida is one of the two orders within the Myodocopa, in turn a subclass of the Ostracoda. The Myodocopida are distinguished by a worm-like seventh limb, and, usually, a rostrum above an incisur from which the antennae can protrude...
HalocypridaThe Halocyprida is one of the two orders within the Myodocopa, in turn a subclass of the ostracods. Like their relatives in the order Myodocopida, they have a long exopod on the second antenna. However, unlike myodocopids, their fifth appendage is leg-like rather than modified for feeding, their...
Platycopida PodocopidaPodocopida is an order of ostracods in the subclass Podocopa. It is divided into five suborders – Bairdiocopina, Cypridocopina, Cytherocopina, Darwinulocopina and Sigilliocopina. It is the most diverse of the four orders of ostracods, and also has a rich fossil record.... |
CylindroleberididaeCylindroleberididae is a family of ostracods that shows remarkable morphological diversity. The defining feature is the possession of gills: 7–8 leaf-like pairs at the posterior of the body...
|
MalacostracaMalacostraca is the largest of the six classes of crustaceans, containing over 25,000 extant species, divided among 16 orders. Its members display a greater diversity of body forms than any other class of animals, and include crabs, lobsters, shrimp, krill, woodlice, scuds , mantis shrimp and many... |
crabTrue crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax... s lobsterClawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most... s shrimpShrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
krillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
mantis shrimpMantis shrimp or stomatopods are marine crustaceans, the members of the order Stomatopoda. They are neither shrimp nor mantids, but receive their name purely from the physical resemblance to both the terrestrial praying mantis and the shrimp. They may reach in length, although exceptional cases of...
woodliceA woodlouse is a crustacean with a rigid, segmented, long exoskeleton and fourteen jointed limbs...
sandhoppers etc. |
DecapodaThe decapods or Decapoda are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crayfish, crabs, lobsters, prawns and shrimp. Most decapods are scavengers. It is estimated that the order contains nearly 15,000 species in around 2,700 genera, with...
IsopodaIsopods are an order of peracarid crustaceans, including familiar animals such as woodlice and pill bugs. The name Isopoda derives from the Greek roots and...
AmphipodaAmphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. The name amphipoda means "different-footed", and refers to the different forms of appendages, unlike isopods, where all the legs are alike. Of the 7,000 species, 5,500 are classified...
StomatopodaMantis shrimp or stomatopods are marine crustaceans, the members of the order Stomatopoda. They are neither shrimp nor mantids, but receive their name purely from the physical resemblance to both the terrestrial praying mantis and the shrimp. They may reach in length, although exceptional cases of...
c. 12 others |
Gammarus roeseli (Amphipoda) |
Fossil record
Crustaceans have a rich and extensive fossil record, which begins with animals such as
CanadaspisCanadaspis was a Cambrian genus of crustacean or euarthropod, a benthic feeder that moved mainly by walking and possibly used its biramous appendages to stir mud in search of food...
and
PerspicarisPerspicaris is a fossil arthropod from the Cambrian period. It was long and bivalved. The valves, encasing the thorax, were joined together by a dorsal hinge.It is difficult to establish the lifestyle of Perspicaris...
from the Middle Cambrian age
Burgess ShaleThe Burgess Shale Formation, located in the Canadian Rockies of British Columbia, is one of the world's most celebrated fossil fields, and the best of its kind. It is famous for the exceptional preservation of the soft parts of its fossils...
. Most of the major groups of crustaceans appear in the fossil record before the end of the Cambrian, namely the
BranchiopodaBranchiopoda is a class of crustaceans. It is the sister group to the remaining crustaceans, and comprises fairy shrimp, clam shrimp, Cladocera, Notostraca and the Devonian Lepidocaris...
,
MaxillopodaMaxillopoda is a diverse class of crustaceans including barnacles, copepods and a number of related animals. It does not appear to be a monophyletic group, and no single character unites all the members.-Description:...
(including
barnacleA barnacle is a type of arthropod belonging to infraclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosive settings. They are sessile suspension feeders, and have...
s and
tongue wormsPentastomida are an enigmatic group of parasitic invertebrates commonly known as tongue worms due to the resemblance of the species of the genus Linguatula to a vertebrate tongue....
) and
MalacostracaMalacostraca is the largest of the six classes of crustaceans, containing over 25,000 extant species, divided among 16 orders. Its members display a greater diversity of body forms than any other class of animals, and include crabs, lobsters, shrimp, krill, woodlice, scuds , mantis shrimp and many...
; there is some debate as to whether or not Cambrian animals assigned to Ostracoda are truly
ostracodOstracoda is a class of the Crustacea, sometimes known as the seed shrimp because of their appearance. Some 65,000 species have been identified, grouped into several orders....
s, which would otherwise start in the
OrdovicianThe Ordovician is a geologic period and system, the second of six of the Paleozoic Era, and covers the time between 488.3±1.7 to 443.7±1.5 million years ago . It follows the Cambrian Period and is followed by the Silurian Period...
. The only classes to appear later are the
CephalocaridaCephalocarida is a class inside the subphylum Crustacea that comprises only twelve shrimp-like benthic species. They were discovered in 1955 by Howard L. Sanders, and are commonly referred to as horseshoe shrimps. They have been grouped together with the Remipedia in the Xenocarida...
, which have no fossil record, and the
RemipediaRemipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, the Atlantic Ocean...
, which were first described from the fossil
Tesnusocaris goldichi, but do not appear until the
CarboniferousThe Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...
. Most of the early crustaceans are rare, but fossil crustaceans become abundant from the Carboniferous onwards.
Within the Malacostraca, no fossils are known for
krillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
, while both
HoplocaridaHoplocarida is a subclass of crustaceans. The only extant members are the mantis shrimp , but two other orders existed in the Palaeozoic: Aeschronectida and Palaeostomatopoda....
and Phyllopoda contain important groups that are now extinct as well as extant members (Hoplocarida:
mantis shrimpMantis shrimp or stomatopods are marine crustaceans, the members of the order Stomatopoda. They are neither shrimp nor mantids, but receive their name purely from the physical resemblance to both the terrestrial praying mantis and the shrimp. They may reach in length, although exceptional cases of...
are extant, while
AeschronectidaAeschronectida is an extinct order of mantis shrimp-like crustaceans ....
are extinct; Phyllopoda: Canadaspidida are extinct, while
LeptostracaLeptostraca is an order of small, marine crustaceans. Its members, including the well-studied Nebalia, occur throughout the world's oceans and are usually considered to be filter-feeders. It is the only extant order in the subclass Phyllocarida...
are extant).
CumaceaCumacea is an order of small marine crustaceans, occasionally called hooded shrimp. Their unique appearance and uniform body plan makes them easy to distinguish from other crustaceans.-Anatomy:...
and
IsopodaIsopods are an order of peracarid crustaceans, including familiar animals such as woodlice and pill bugs. The name Isopoda derives from the Greek roots and...
are both known from the
CarboniferousThe Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...
, as are the first true mantis shrimp. In the
DecapodaThe decapods or Decapoda are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crayfish, crabs, lobsters, prawns and shrimp. Most decapods are scavengers. It is estimated that the order contains nearly 15,000 species in around 2,700 genera, with...
,
prawnPrawns are decapod crustaceans of the sub-order Dendrobranchiata. There are 540 extant species, in seven families, and a fossil record extending back to the Devonian...
s and
polychelidsPolychelida is a group of decapod crustaceans. Fossil representatives are known dating from as far back as the Upper Triassic. A total of 38 extant species, all in the family Polychelidae, and 55 fossil species have been described....
appear in the Triassic, and
shrimpShrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
and
crabTrue crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
s appear in the
JurassicThe Jurassic is a geologic period and system that extends from about Mya to Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...
; however, the great radiation of crustaceans occurred in the
CretaceousThe Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
, particularly in crabs, and may have been driven by the
adaptive radiationIn evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...
of their main predators, bony fish. The first true
lobsterClawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...
s also appear in the Cretaceous.
Consumption by humans
Many crustaceans are consumed by humans, and nearly 10,700,000
tonThe ton is a unit of measure. It has a long history and has acquired a number of meanings and uses over the years. It is used principally as a unit of weight, and as a unit of volume. It can also be used as a measure of energy, for truck classification, or as a colloquial term.It is derived from...
s were produced in 2007; the vast majority of this output is of
decapod crustaceansThe decapods or Decapoda are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crayfish, crabs, lobsters, prawns and shrimp. Most decapods are scavengers. It is estimated that the order contains nearly 15,000 species in around 2,700 genera, with...
:
crabTrue crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
s,
lobsterClawed lobsters comprise a family of large marine crustaceans. Highly prized as seafood, lobsters are economically important, and are often one of the most profitable commodities in coastal areas they populate.Though several groups of crustaceans are known as lobsters, the clawed lobsters are most...
s,
shrimpShrimp are swimming, decapod crustaceans classified in the infraorder Caridea, found widely around the world in both fresh and salt water. Adult shrimp are filter feeding benthic animals living close to the bottom. They can live in schools and can swim rapidly backwards. Shrimp are an important...
, and
prawnPrawns are decapod crustaceans of the sub-order Dendrobranchiata. There are 540 extant species, in seven families, and a fossil record extending back to the Devonian...
s. Over 60% by weight of all crustaceans caught for consumption are shrimp and prawns, and nearly 80% is produced in Asia, with China alone producing nearly half the world's total. Non-decapod crustaceans are not widely consumed, with only 118,000 tons of
krillKrill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world...
being caught, despite krill having one of the greatest
biomassesBiomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...
on the planet.
External links
- Crustacea.net, an online resource on the biology of crustaceans
- Crustacea: Natural History Museum of Los Angeles County
The Natural History Museum of Los Angeles County opened in Exposition Park, Los Angeles, California, USA in 1913 as the Museum of History, Science, and Art. The moving force behind it was a museum association founded in 1910. Its distinctive main building, with fitted marble walls and domed and...
- Crustacea: Tree of Life Web Project
The Tree of Life Web Project is an ongoing Internet project providing information about the diversity and phylogeny of life on Earth. This collaborative peer reviewed project began in 1995, and is written by biologists from around the world....
- The Crustacean Society
- Natural History Collections: Crustacea: University of Edinburgh
The University of Edinburgh, founded in 1583, is a public research university located in Edinburgh, the capital of Scotland, and a UNESCO World Heritage Site. The university is deeply embedded in the fabric of the city, with many of the buildings in the historic Old Town belonging to the university...
- Crustaceans (Crustacea) on the shore of Singapore
- Crustacea(crabs, lobsters, shrimps, prawns, barnacles): Biodiversity Explorer