|From left to right and from top to bottom: Grapsus grapsus (a crab), Homarus gammarus (a lobster), Procambarus clarkii (a crayfish), Lysmata amboinensis (a shrimp), Euphausia superba (a krill), Hemilepistus reaumuri (a woodlouse), Calanoida (a copepod), and Lepas anatifera (a barnacle)|
|Cladistically included but traditionally excluded taxa|
Crustaceans (Crustacea // ) form a large, diverse arthropod taxon which includes such animals as crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata; because of recent molecular studies it is now well accepted that the crustacean group is paraphyletic, and comprises all animals in the clade Pancrustacea other than hexapods. Some crustaceans (Remipedia, Cephalocarida, Malacostraca) are more closely related to insects and the other hexapods than they are to certain other crustaceans.
The 67,000 described species range in size from Stygotantulus stocki at 0.1 mm (0.004 in), to the Japanese spider crab with a leg span of up to 3.8 m (12.5 ft) and a mass of 20 kg (44 lb). Like other arthropods, crustaceans have an exoskeleton, which they moult to grow. They are distinguished from other groups of arthropods, such as insects, myriapods and chelicerates, by the possession of biramous (two-parted) limbs, and by their larval forms, such as the nauplius stage of branchiopods and copepods.
Most crustaceans are free-living aquatic animals, but some are terrestrial (e.g. woodlice), some are parasitic (e.g. Rhizocephala, fish lice, tongue worms) and some are sessile (e.g. barnacles). The group has an extensive fossil record, reaching back to the Cambrian, and includes living fossils such as Triops cancriformis , which has existed apparently unchanged since the Triassic period. More than 7.9 million tons of crustaceans per year are produced by fishery or farming for human consumption, most of it being 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.
The body of a crustacean is composed of segments, which are grouped into three regions: the cephalon or head, : 289the pereon or thorax, and the pleon or abdomen. The head and thorax may be fused together to form a cephalothorax, which may be covered by a single large carapace. 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.
Each somite, or body segment can bear a pair of appendages: on the segments of the head, these include two pairs of antennae, the mandibles and maxillae;the thoracic segments bear legs, which may be specialised as pereiopods (walking legs) and maxillipeds (feeding legs). The abdomen bears pleopods, and ends in a telson, which bears the anus, and is often flanked by uropods to form a tail fan. The number and variety of appendages in different crustaceans may be partly responsible for the group's success.
Crustacean appendages are typically biramous, meaning they are divided into two parts; this includes the second pair of antennae, but not the first, which is usually uniramous, the exception being in the Class Malacostraca where the antennules may be generally biramous or even triramous.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.
The main body cavity is an open circulatory system, where blood is pumped into the haemocoel by a heart 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 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.
Most crustaceans are aquatic, living in either marine or freshwater environments, but a few groups have adapted to life on land, such as terrestrial crabs, terrestrial hermit crabs, and woodlice. Marine crustaceans are as ubiquitous in the oceans as insects are on land. : 672 Krill are the bottom layer and the most important part of the food chain in Antarctic animal communities. : 64 Some crustaceans are significant invasive species, such as the Chinese mitten crab, Eriocheir sinensis , and the Asian shore crab, Hemigrapsus sanguineus .Most crustaceans are also motile, moving about independently, although a few taxonomic units are parasitic and live attached to their hosts (including sea lice, fish lice, whale lice, tongue worms, and Cymothoa exigua , all of which may be referred to as "crustacean lice"), and adult barnacles live a sessile life – they are attached headfirst to the substrate and cannot move independently. Some branchiurans are able to withstand rapid changes of salinity and will also switch hosts from marine to non-marine species.
Most crustaceans have separate sexes, and reproduce sexually. In fact, a recent study explains how the male crustaceans,T. Californicus decide which females to mate with by dietary differences, preferring when the females are algae-fed instead of yeast-fed.A small number are hermaphrodites, including barnacles, remipedes, and Cephalocarida. Some may even change sex during the course of their life. Parthenogenesis is also widespread among crustaceans, where viable eggs are produced by a female without needing fertilisation by a male. This occurs in many branchiopods, some ostracods, some isopods, and certain "higher" crustaceans, such as the Marmorkrebs crayfish.
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 decapods carry the eggs attached to the pleopods, while peracarids, notostracans, anostracans, and many isopods form a brood pouch from the carapace and thoracic limbs. : 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.Female Branchiura do not carry eggs in external ovisacs but attach them in rows to rocks and other objects.
Crustaceans exhibit a number of larval forms, of which the earliest and most characteristic is the nauplius. This has three pairs of appendages, 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 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. 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.
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 .The earliest nomenclaturally valid work to use the name "Crustacea" was Morten Thrane Brünnich's Zoologiæ Fundamenta in 1772, although he also included chelicerates in the group.
The subphylum Crustacea comprises almost 67,000 described species, 1⁄10 to 1⁄100 of the total number as most species remain as yet undiscovered. Although most crustaceans are small, their morphology varies greatly and includes both the largest arthropod in the world – the Japanese spider crab with a leg span of 3.7 metres (12 ft) – and the smallest, the 100-micrometre-long (0.004 in) Stygotantulus stocki . Despite their diversity of form, crustaceans are united by the special larval form known as the nauplius.which is thought to be just
The exact relationships of the Crustacea to other taxa are not completely settled as of April 2012 [update] . Studies based on morphology led to the Pancrustacea hypothesis, in which Crustacea and Hexapoda (insects and allies) are sister groups. More recent studies using DNA sequences suggest that Crustacea is paraphyletic, with the hexapods nested within a larger Pancrustacea clade.
Although the classification of crustaceans has been quite variable, the system used by Martin and Davislargely supersedes earlier works. Mystacocarida and Branchiura, here treated as part of Maxillopoda, are sometimes treated as their own classes. Six classes are usually recognised:
|Ostracoda||Seed Shrimp|| Myodocopida |
|Mystacocarida||Mystococaridans|| Ctenocheilocaris |
|Pentastomida||Tongue Worms|| Cephalobaenida |
Armillifer armillatus (Porocephalida)
|Branchiura||Fish Lice|| Arguloida |
Argulus ambystoma (Arguloida)
|Thecostraca|| facetotecta |
| Facetotecta |
Perforatus perforatus (Cirripedia)
|Copepoda||Copepods|| Calanoida |
|Malacostraca|| Mantis Shrimp |
| Stomatopoda |
Ocypode ceratophthalma (Decapoda)
|Cephalocarida||Horseshoe Shrimp||Brachypoda|| |
|Branchiopoda|| Fairy Shrimp |
| Anostraca |
Lepidurus arcticus (Notostraca)
|Hexapoda|| Springtails |
| Collembola |
Mantispa styriaca (Neuroptera)
Crustaceans have a rich and extensive fossil record, which begins with animals such as Canadaspis and Perspicaris from the Middle Cambrian age Burgess Shale.Most of the major groups of crustaceans appear in the fossil record before the end of the Cambrian, namely the Branchiopoda, Maxillopoda (including barnacles and tongue worms) and Malacostraca; there is some debate as to whether or not Cambrian animals assigned to Ostracoda are truly ostracods, which would otherwise start in the Ordovician. The only classes to appear later are the Cephalocarida, which have no fossil record, and the Remipedia, which were first described from the fossil Tesnusocaris goldichi , but do not appear until the Carboniferous. Most of the early crustaceans are rare, but fossil crustaceans become abundant from the Carboniferous period onwards.
Within the Malacostraca, no fossils are known for krill,while both Hoplocarida and Phyllopoda contain important groups that are now extinct as well as extant members (Hoplocarida: mantis shrimp are extant, while Aeschronectida are extinct; Phyllopoda: Canadaspidida are extinct, while Leptostraca are extant ). Cumacea and Isopoda are both known from the Carboniferous, as are the first true mantis shrimp. In the Decapoda, prawns and polychelids appear in the Triassic, and shrimp and crabs appear in the Jurassic;. 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.
However, the great radiation of crustaceans occurred in the Cretaceous, particularly in crabs, and may have been driven by the adaptive radiation of their main predators, bony fish.The first true lobsters also appear in the Cretaceous.
Many crustaceans are consumed by humans, and nearly 10,700,000 tons were produced in 2007; the vast majority of this output is of decapod crustaceans: crabs, lobsters, shrimp, crawfish, and prawns. 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 krill being caught, despite krill having one of the greatest biomasses on the planet.
Dendrobranchiata is a suborder of decapods, commonly known as prawns. There are 540 extant species in seven families, and a fossil record extending back to the Devonian. They differ from related animals, such as Caridea and Stenopodidea, by the branching form of the gills and by the fact that they do not brood their eggs, but release them directly into the water. They may reach a length of over 330 millimetres (13 in) and a mass of 450 grams (1.0 lb), and are widely fished and farmed for human consumption.
Natantia is an obsolete taxon of decapod crustaceans, comprising those families that move predominantly by swimming – the shrimp, prawns (Dendrobranchiata) and boxer shrimp. The remaining Decapoda were placed in the Reptantia, and consisted of crabs, lobsters and other large animals that move chiefly by walking along the bottom. The division between Natantia and Reptantia was replaced in 1963, when Martin Burkenroad erected the suborder Pleocyemata for those animals that brood their eggs on the pleopods, leaving Dendrobranchiata for the prawns. Under this system, Natantia is a paraphyletic group. Burkenroad's primary division of Decapoda into Dendrobranchiata and Pleocyemata has since been corroborated by molecular analyses.
Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen), usually hidden entirely under the thorax. They live in all the world's oceans, in fresh water, and on land, are generally covered with a thick exoskeleton, and have a single pair of pincers. Several other groups of crustaceans with similar appearances – such as king crabs and porcelain crabs are not true crabs, but have evolved features similar to true crabs through a process known as carcinisation.
Malacostraca is the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, amphipods, mantis shrimp and many others, less familiar animals. They are abundant in all marine environments and have colonised freshwater and terrestrial habitats. They are segmented animals, united by a common body plan comprising 20 body segments, and divided into a head, thorax, and abdomen.
The Decapoda or decapods are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crabs, lobsters, crayfish, shrimp and prawns. Most decapods are scavengers. The order is estimated to contain nearly 15,000 species in around 2,700 genera, with around 3,300 fossil species. Nearly half of these species are crabs, with the shrimp and Anomura including hermit crabs, porcelain crabs, squat lobsters making up the bulk of the remainder. The earliest fossil decapod is the Devonian Palaeopalaemon.
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, and the Atlantic Ocean. The first described remipede was the fossil Tesnusocaris goldichi. Since 1979, at least seventeen living species have been identified in subtropical regions around the world.
Reptantia is a clade of decapod crustaceans named in 1880 which includes lobsters, crabs and many other well-known crustaceans.
Pleocyemata is a suborder of decapod crustaceans, erected by Martin Burkenroad in 1963. Burkenroad's classification replaced the earlier sub-orders of Natantia and Reptantia with the monophyletic groups Dendrobranchiata (prawns) and Pleocyemata. Pleocyemata contains all the members of the Reptantia, as well as the Stenopodidea, and Caridea, which contains the true shrimp.
The Achelata is an infra-order of the decapod crustaceans, holding the spiny lobsters, slipper lobsters and their fossil relatives.
Anomura 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 Stenopodidea or boxer shrimps are a small group of decapod crustaceans. Often confused with Caridea shrimp or Dendrobranchiata prawns, they are neither, belonging to their own group.
The decapod crustacean, such as a crab, lobster, shrimp or prawn, is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in various groups these may be reduced or missing. They are, from head to tail:
Eucarida is a superorder of the Malacostraca, a class of the crustacean subphylum, comprising the decapods, krill, Amphionides and Angustidontida. They are characterised by having the carapace fused to all thoracic segments, and by the possession of stalked eyes.
Pancrustacea 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, with most studies recovering Hexapoda within Crustacea. The clade has also been called Tetraconata, referring to having four cone cells in the ommatidia. That name is preferred by some scientists as a means of avoiding confusion with the use of "pan-" to indicate a clade that includes a crown group and all of its stem group representatives.
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 larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults, more-so than where the larvae are planktonic, and thereby easily caught.
Phylogeny of Malacostraca is the evolutionary relationships of the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members display a great diversity of body forms. Although the class Malacostraca is united by a number of well-defined and documented features, which were recognised a century ago by William Thomas Calman in 1904, the phylogenetic relationship of the orders which compose this class is unclear due to the vast diversity present in their morphology. Molecular studies have attempted to infer the phylogeny of this clade, resulting in phylogenies which have a limited amount of morphological support. To resolve a well-supported eumalacostracan phylogeny and obtain a robust tree, it will be necessary to look beyond the most commonly utilized sources of data.
Shrimp are decapod crustaceans with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata. More narrow definitions may be restricted to Caridea, to smaller species of either group or to only the marine species. Under a broader definition, shrimp may be synonymous with prawn, covering stalk-eyed swimming crustaceans with long, narrow muscular tails (abdomens), long whiskers (antennae), and slender legs. Any small crustacean which resembles a shrimp tends to be called one. They swim forward by paddling with swimmerets on the underside of their abdomens, although their escape response is typically repeated flicks with the tail driving them backwards very quickly. Crabs and lobsters have strong walking legs, whereas shrimp have thin, fragile legs which they use primarily for perching.
The clade Multicrustacea constitutes the largest superclass of crustaceans, containing approximately four-fifths of all described crustacean species, including crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, barnacles, copepods, amphipods, mantis shrimp and others. The largest branch of multicrustacea is the class Malacostraca.
Powers, M., Hill, G., Weaver, R., & Goymann, W. (2020). An experimental test of mate choice for red carotenoid coloration in the marine copepod Tigriopus californicus. Ethology., 126(3), 344–352. https://doi.org/10.1111/eth.12976
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