Crustacean

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Crustaceans
Temporal range: 511–0 
Ma
 Cambrian to present
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Clade: Pancrustacea
Subphylum: Crustacea
Groups included
Cladistically included but traditionally excluded taxa

Crustaceans are a group of

amphipods and mantis shrimp.[1] The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods (insects and entognathans) emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea.[2] The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans (oligostracans and multicrustaceans).[3]

The 67,000 described species range in size from

branchiopods and copepods
.

Most crustaceans are free-living

shrimp and prawns. Krill and copepods are not as widely fished, but may be the animals with the greatest biomass 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 scientist who works in carcinology is a carcinologist
.

Structure

lady crab, part of the hard exoskeleton
Body structure of a typical crustacean – krill

The body of a crustacean is composed of segments, which are grouped into three regions: the cephalon or head,[5] the pereon or thorax,[6] and the pleon or abdomen.[7] The head and thorax may be fused together to form a cephalothorax,[8] which may be covered by a single large carapace.[9] The crustacean body is protected by the hard exoskeleton, which must be moulted for the animal to grow. The shell around each somite can be divided into a dorsal tergum, ventral sternum and a lateral pleuron. Various parts of the exoskeleton may be fused together.[10]: 289 

Each

maxillipeds (feeding legs).[6] Malacostraca and Remipedia (and the hexapods) have abdominal appendages. All other classes of crustaceans have a limbless abdomen, except from a telson and caudal rami which is present in many groups.[11][12]
The abdomen in malacostracans bears
pleopods,[7] and ends in a telson, which bears the anus, and is often flanked by uropods to form a tail fan.[13] The number and variety of appendages in different crustaceans may be partly responsible for the group's success.[14]

Crustacean

uniramous, the exception being in the Class Malacostraca where the antennules may be generally biramous or even triramous.[15][16] 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. Trilobites, for instance, also possessed biramous appendages.[17]

See also: Hemolymph

The main body cavity is an

haemoglobins.[19] 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.[20] 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.[21]

In many decapods, 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 crab) mate seasonally and return to the sea to release the eggs. Others, such as woodlice, lay their eggs on land, albeit in damp conditions. In most decapods, the females retain the eggs until they hatch into free-swimming larvae.[22]

Ecology

amphipod

Most crustaceans are aquatic, living in either marine or

Eriocheir sinensis,[27] and the Asian shore crab, Hemigrapsus sanguineus.[28] Since the piercing of the Suez Canal, close to 100 species of crustaceans from the Red Sea and the Indo-Pacific realm have established themselves in the eastern Mediterranean sub-basin, with often significant impact on local ecosystems.[29]

Life cycle

Seven round translucent spheres: inside some of them, a pair of compound eyes can be seen.
Eggs of Potamon fluviatile, a freshwater crab
European lobster
, Homarus gammarus

Mating system

Most crustaceans have

Marmorkrebs
crayfish.

Eggs

In many crustaceans, the fertilised eggs are released into the

isopods form a brood pouch from the carapace and thoracic limbs.[30] Female Branchiura do not carry eggs in external ovisacs but attach them in rows to rocks and other objects.[33]: 788  Most leptostracans and krill carry the eggs between their thoracic limbs; some copepods carry their eggs in special thin-walled sacs, while others have them attached together in long, tangled strings.[30]

Larvae

Main article:
Crustacean larvae

Crustaceans exhibit a number of larval forms, of which the earliest and most characteristic is the

post-larva. Zoea larvae swim with their thoracic appendages, as opposed to nauplii, which use cephalic appendages, and megalopa, which use abdominal appendages for swimming. It often has spikes on its carapace, which may assist these small organisms in maintaining directional swimming.[36] In many decapods
, 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.

Providing camouflage against predators, the otherwise black eyes in several forms of swimming larvae are covered by a thin layer of crystalline isoxanthopterin that gives their eyes the same color as the surrounding water, while tiny holes in the layer allow light to reach the retina.[37] As the larvae mature into adults, the layer migrates to a new position behind the retina where it works as a backscattering mirror that increases the intensity of light passing through the eyes, as seen in many nocturnal animals.[38]

DNA repair

In an effort to understand whether DNA repair processes can protect crustaceans against DNA damage, basic research was conducted to elucidate the repair mechanisms used by Penaeus monodon (black tiger shrimp).[39] Repair of DNA double-strand breaks was found to be predominantly carried out by accurate homologous recombinational repair. Another, less accurate process, microhomology-mediated end joining, is also used to repair such breaks. The expression pattern of DNA repair related and DNA damage response genes in the intertidal copepod Tigriopus japonicus was analyzed after ultraviolet irradiation.[40] This study revealed increased expression of proteins associated with the DNA repair processes of non-homologous end joining, homologous recombination, base excision repair and DNA mismatch repair.

Classification and phylogeny

Copepods, from Ernst Haeckel's 1904 work Kunstformen der Natur
Decapods, from Ernst Haeckel's 1904 work Kunstformen der Natur

The name "crustacean" dates from the earliest works to describe the animals, including those of Pierre Belon and Guillaume Rondelet, but the name was not used by some later authors, including Carl Linnaeus, who included crustaceans among the "Aptera" in his Systema Naturae.[41] The earliest nomenclatural valid work to use the name "Crustacea" was Morten Thrane Brünnich's Zoologiæ Fundamenta in 1772,[42] although he also included chelicerates in the group.[41]

The subphylum Crustacea comprises almost 67,000 described

nauplius
.

The exact relationships of the Crustacea to other taxa are not completely settled as of April 2012. Studies based on morphology led to the

paraphyletic, with the hexapods nested within a larger Pancrustacea clade.[48][49]

The traditional classification of Crustacea based on morphology recognised four to six classes.[50] Bowman and Abele (1982) recognised 652 extant families and 38 orders, organised into six classes: Branchiopoda, Remipedia, Cephalocarida,

Branchiura and Pentastomida).[56][57]

Class Members Orders Photo
Ostracoda
Seed shrimp
Platycopida
Podocopida
 A translucent, sculptured shell conceals a small animal. Some of its appendages extend beyond the shell.
Cylindroleberididae
(Myodocopida)
Mystacocarida
Mystococaridans
Mystococarida
 A line drawing of a dorsal view of a small animal with many segments and appendages.
Ctenocheilocaris galvarini
Branchiura and Pentastomida

may be recognised as classes)
fish lice
Reighardiida
Arguloida
 A translucent, sculptured shell conceals a small animal. Some of its appendages extend beyond the shell.
Armillifer armillatus
(Porocephalida)
Thecostraca
Barnacles
Cryptophialida
Lithoglyptida

etc.
Cirripedia
)
Copepoda
 Copepods
Gelyelloida
Harpacticoida
Misophrioida

etc.		 A translucent, sculptured shell conceals a small animal. Some of its appendages extend beyond the shell.
Cylindroleberididae
(Calanoida)
Tantulocarida Tantulocaridians Tantulus larva (Microdajus sp.)
Microdajus sp.
Malacostraca
Hooded shrimp
Amphipods

etc.
Euphausiacea
Isopoda
Cumacea
Amphipoda

etc.
Ocypode ceratophthalma
(Decapoda
)
Cephalocarida
Horseshoe shrimp
Brachypoda
Hutchinsoniella macracantha
Branchiopoda
Tadpole shrimp
Clam shrimp
Spinicaudata

etc.		 A microscopic, transparent, oval animal against a black background. The head has a large eye, antennae, and comes to a pointed beak. The rest of the animal is smooth round and fat, culminating in a pointed tail. The internal anatomy is apparent.
Lepidurus arcticus
(Notostraca)
Remipedia Remipedes
Nectiopoda
†Enantiopoda
Speleonectes tanumekes
Hexapoda Springtails
Proturans
Diplurans
Insects
Coleoptera
Neuroptera
Hymenoptera

etc.		 A translucent, sculptured shell conceals a small animal. Some of its appendages extend beyond the shell.
Mantispa styriaca
(Neuroptera)

The following cladogram shows the updated relationships between the different extant groups of the paraphyletic Crustacea in relation to the class Hexapoda.[53]

Pancrustacea Crustacea

According to this diagram, the Hexapoda are deep in the Crustacea tree, and any of the Hexapoda is distinctly closer to e.g. a Multicrustacean than an Oligostracan is.

Fossil record

Eryma mandelslohi, a fossil decapod from the Jurassic of Bissingen an der Teck, Germany

Crustaceans have a rich and extensive

Carboniferous period onwards.[58]

Norway lobsters
on sale at a Spanish market

Within the Malacostraca, no fossils are known for

Canadaspidida are extinct, while Leptostraca are extant[59]). Cumacea and Isopoda are both known from the Carboniferous,[65][66] as are the first true mantis shrimp.[67] In the Decapoda, prawns and polychelids appear in the Triassic,[68][69] and shrimp and crabs appear in the Jurassic.[70][71] The fossil burrow Ophiomorpha is attributed to ghost shrimps, whereas the fossil burrow Camborygma is attributed to crayfishes. The Permian–Triassic deposits of Nurra preserve the oldest (Permian: Roadian) fluvial burrows ascribed to ghost shrimps (Decapoda: Axiidea, Gebiidea) and crayfishes (Decapoda: Astacidea, Parastacidea), respectively.[72]

However, the great radiation of crustaceans occurred in the

bony fish.[71] The first true lobsters also appear in the Cretaceous.[73]

Consumption by humans

Many crustaceans are consumed by humans, and nearly 10,700,000 

crawfish, and prawns.[74] 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.[74] Non-decapod crustaceans are not widely consumed, with only 118,000 tons of krill being caught,[74] despite krill having one of the greatest biomasses on the planet.[75]

See also

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Sources

External links

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