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Temporal range: Cambrian–Recent
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Crustacea
Superclass: Multicrustacea
Class: Malacostraca
Latreille, 1802

See text for orders.

Malacostraca (from New Latin; from Ancient Greek μαλακός (malakós) 'soft',and όστρακον (óstrakon) 'shell') 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 other, 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 (rarely 21), and divided into a head, thorax, and abdomen.



The name Malacostraca was coined by a French zoologist Pierre André Latreille in 1802. He was curator of the arthropod collection at the National Museum of Natural History in Paris. [1] The name comes from the Greek roots μαλακός (malakós, meaning "soft") and ὄστρακον (óstrakon, meaning "shell"). [2] The name is misleading, since the shell is only soft immediately after moulting, and is usually hard. [3] Malacostracans are sometimes contrasted with entomostracans, a name applied to all crustaceans outside the Malacostraca, and named after the obsolete taxon Entomostraca. [4]


Leptostraca such as Nebalia bipes retain the primitive condition of having seven abdominal segments. Nebalia bipes.jpg
Leptostraca such as Nebalia bipes retain the primitive condition of having seven abdominal segments.

The class Malacostraca includes about 40,000 species, [5] and "arguably ... contains a greater diversity of body forms than any other class in the animal kingdom". [6] Its members are characterised by the presence of three tagmata (specialized groupings of multiple segments) – a five-segmented head, an eight-segmented thorax and an abdomen with six segments and a telson, except in the Leptostraca, which retain the ancestral condition of seven abdominal segments. [6] Malacostracans have abdominal appendages, a fact that differentiates them from all other major crustacean taxa except Remipedia. [7] Each body segment bears a pair of jointed appendages, although these may be lost secondarily. [8]


The head bears two pairs of antennae, the first of which is often biramous (branching into two parts) and the second pair bear exopods (outer branches) which are often flattened into antennal scales known as scaphocerites. [7] The mouthparts consist of pairs each of mandibles, maxillules (second pair of mouthparts) and maxillae. Usually a pair of stalked compound eyes is present, although in some taxa the eyes are unstalked, reduced or lost. [9] [10]

Up to three thoracic segments may be fused with the head to form a cephalothorax; the associated appendages turn forward and are modified as maxillipeds (accessory mouthparts). [7] A carapace may be absent, present or secondarily lost, and may cover the head, part or all of the thorax and some of the abdomen. [6] It is variable in form and may be fused dorsally with some of the thoracic segments or occasionally be in two parts, hinged dorsally. [9] Typically, each of the thoracic appendages is biramous and the endopods are the better developed of the branches, being used for crawling or grasping. Each endopod consist of seven articulating segments; the coxa, basis, ischium, merus, carpus, propodus and dactylus. In decapods, the claw is formed by the articulation of the dactylus against an outgrowth of the propodus. In some taxa, the exopods are lost and the appendages are uniramous. [7]

There is a clear demarcation between the thorax and the six or seven-segmented abdomen. In most taxa, each abdominal segment except the last carries a pair of biramous pleopods used for swimming, burrowing, gas exchange, creating a current or brooding eggs. The first and second abdominal pleopods may be modified in the male to form gonopods (accessory copulatory appendages). [7] The appendages of the last segment are typically flattened into uropods, which together with the terminal telson, make up the "tail fan". [10] It is the sudden flexion of this tail fan that provides the thrust for the rapid escape response of these crustaceans and the tail fan is also used in steering. [7] In Leptostraca, the appendages on the telson instead form caudal rami (spine-like protrusions). [11]

Internal anatomy

The digestive tract is straight and the foregut consists of a short oesophagus and a two-chambered stomach, the first part of which contains a gizzard-like "gastric mill" for grinding food. The walls of this have chitinous ridges, teeth and calcareous ossicles. The fine particles and soluble material are then moved into the midgut where chemical processing and absorption takes place in one or more pairs of large digestive caeca. The hindgut is concerned with water reclamation and the formation of faeces and the anus is situated at the base of the telson. [12]

Like other crustaceans, malacostracans have an open circulatory system in which the heart pumps blood into the hemocoel (body cavity) where it supplies the needs of the organs for oxygen and nutrients before diffusing back to the heart. [13] The typical respiratory pigment in malacostracans is haemocyanin. [14] Structures that function as kidneys are located near the base of the antennae. A brain exists in the form of ganglia close to the antennae, there are ganglia in each segment and a collection of major ganglia below the oesophagus. [15] Sensory organs include compound eyes (often stalked), ocelli (simple eyes), statocysts and sensory bristles. The naupliar eye is a characteristic of the nauplius larva and consists of four cup-shaped ocelli facing in different directions and able to distinguish between light and darkness. [12]


Grapsus grapsus, a terrestrial crab Sally lightfoot crab.jpg
Grapsus grapsus , a terrestrial crab

Malacostracans live in a wide range of marine and freshwater habitats, and three orders have terrestrial members: Amphipoda (Talitridae), Isopoda (Oniscidea, the woodlice) and Decapoda (terrestrial hermit crabs, crabs of the families Ocypodidae, Gecarcinidae, and Grapsidae, and terrestrial crayfish). [16] They are abundant in all marine ecosystems, and most species are scavengers, although some, such as the porcelain crabs, are filter feeders, and some, such as mantis shrimps, are carnivores. [10]

Life cycle

Most species of malacostracans have distinct sexes (a phenomenon known as gonochorism), although a few species exhibit hermaphroditism. [10] The female genital openings or gonopores are located on the sixth thoracic segment or its appendages, while the male gonopores are on the eighth segment or its appendages, or in a small number of species, on the seventh. [9] The naupliar larval stages are often reduced and take place before hatching, but where they occur, a metamorphosis usually occurs between the larval and the adult forms. Primitive malacostracans have a free-swimming naupliar larval stage. [9]


Mating behavior has been studied in the freshwater shrimp Caridina ensifera. [17] Multiple paternity, common in the Malacostrica, also occurs in C. ensifera. Reproductive success of sires was found to correlate inversely with their genetic relatedness to the mother. [17] This finding suggests that sperm competition and/or pre- and post-copulatory female choice occurs. Female choice may increase the fitness of progeny by avoiding inbreeding that can lead to expression of homozygous deleterious recessive mutations. [18]


The monophyly of Malacostraca is widely accepted. This is supported by several common morphological traits which are present throughout the group and is confirmed by molecular studies. [19] However, a number of problems make it difficult to determine the relationships between the orders of Malacostraca. These include differences in mutation rates in different lineages, different patterns of evolution being apparent in different sources of data, including convergent evolution, and long branch attraction. [20]

There is less agreement on the status of the subclass Phyllocarida with its single extant order, Leptostraca, depending on whether foliaceous (leaf-like) limbs have a single or multiple origin. Some authors advocate placing Phyllocarida in Phyllopoda, a group used in former classification systems, which would then include branchiopods, cephalocarids and leptostracans. A molecular study by American biologists Trisha Spears and Lawrence Abele concluded that phylogenetic evidence did not support the monophyly of this grouping, and that Phyllocarida should be regarded as a subclass of Malacostraca that had diverged from the main lineage at an early date. [9] [21]

The following cladogram is based on the 2001 phylogenetic analysis of Richter & Scholtz. [22]




















Subclass Phyllocarida

Leptostraca is the only extant order of Phyllocarida, the other two orders, Archaeostraca and Hoplostraca being extinct. Leptostracans are thought to be the most primitive of the malacostracans and date back to the Cambrian period. They range in length from 1 to 4 cm (0.4 to 1.6 in), most being suspension feeders though some are carnivores or scavengers. They have a two part carapace which encloses the head, the whole thorax and part of the abdomen and are the only malacostracans with seven abdominal segments. Three families are known with several genera and about twenty species. They are found worldwide from the intertidal zone to the deep ocean, all but one species being benthic (living on the seabed). [7] [9]

Subclass Hoplocarida

Squilla empusa,
a mantis shrimp Squilla empusa.jpg
Squilla empusa,
a mantis shrimp

Stomatopoda is the only extant order of Hoplocarida, the other two orders, Aeschronectida and Archaeostomatopoda being extinct. Stomatopodans, commonly known as mantis shrimps, range in length from 5 to 36 cm (2 to 14 in) and are predators. They have a dorso-ventrally flattened body and a shield-like carapace and are armed with powerful, raptorial claws normally carried in a folded position. There are about 300 species, most living in tropical and subtropical seas although some live in temperate areas. They are benthic, mostly hiding in cracks and crevices or living in burrows, some emerging to forage while others are ambush predators. [7] [9]

Subclass Eumalacostraca

The Eumalocostraca contains the vast majority of the approximately 40,000 living species of malacostracans and consists of three superorders, Syncarida, Peracarida and Eucarida. Syncaridans are mostly small and found in freshwater and subterranean habitats. Peracaridans are characterised by having a marsupium in which they brood their young. They are found in marine, freshwater and terrestrial habitats and include Amphipoda, Cumacea, Isopoda and Mysida. Eucarida includes lobsters, crabs, shrimps, prawns and krill. [23]

Fossil record

The first malacostracans appeared sometime in the Cambrian, when animals belonging to the Phyllocarida appeared. [24] [25]


The following classification of living malacostracans is based on An Updated Classification of the Recent Crustacea (2001) by the American marine biologists Joel W. Martin, curator of crustaceans at the Natural History Museum of Los Angeles County, and George E. Davies. [26] Extinct orders have been added to this [27] [28] [29] and are indicated by an obelisk (†).

Odontodactylus scyllarus (Hoplocarida: Stomatopoda) Odontodactylus scyllarus3.jpg
Odontodactylus scyllarus (Hoplocarida: Stomatopoda)
Porcellio scaber and Oniscus asellus (Peracarida: Isopoda) Porcellio scaber and Oniscus asellus - Zalne20070205.jpg
Porcellio scaber and Oniscus asellus (Peracarida: Isopoda)
Cancer pagurus (Eucarida: Decapoda) Cancer pagurus.jpg
Cancer pagurus (Eucarida: Decapoda)

Class Malacostraca Latreille, 1802

Related Research Articles

Branchiopoda Class of crustaceans

Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca, Notostraca and the Devonian Lepidocaris. They are mostly small, freshwater animals that feed on plankton and detritus.

Dendrobranchiata Suborder of prawns

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.

Decapoda Order of crustaceans

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.

Mysida Small, shrimp-like crustacean

Mysida is an order of small, shrimp-like crustaceans in the malacostracan superorder Peracarida. Their common name opossum shrimps stems from the presence of a brood pouch or "marsupium" in females. The fact that the larvae are reared in this pouch and are not free-swimming characterises the order. The mysid's head bears a pair of stalked eyes and two pairs of antennae. The thorax consists of eight segments each bearing branching limbs, the whole concealed beneath a protective carapace and the abdomen has six segments and usually further small limbs.

Isopoda Order of arthropods

Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, and five pairs of branching appendages on the abdomen that are used in respiration. Females brood their young in a pouch under their thorax.

Cephalocarida Class of crustaceans

The Cephalocarida are a class in the subphylum Crustacea comprising only 12 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. Although a second family, Lightiellidae, is sometimes used, all cephalocaridans are generally considered to belong in just one family: Hutchinsoniellidae. Though no fossil record of cephalocaridans has been found, most specialists believe them to be primitive among crustaceans.

Leptostraca Extinct order of crustaceans

Leptostraca 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. They are believed to represent the most primitive members of their class, the Malacostraca, and first appear in the fossil record during the Cambrian period.

Eucarida Superorder of crustaceans

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.

Phyllocarida Subclass of crustaceans

Phyllocarida is a subclass of crustaceans, comprising the extant order Leptostraca and the extinct orders Hymenostraca and Archaeostraca.

Peracarida Order of crustaceans

The 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 (oostegites) borne on the basalmost segments of the legs. Peracarida is one of the largest crustacean taxa and includes about 12,000 species. Most members are less than 2 cm (0.8 in) in length, but the largest is probably the giant isopod which can reach 76 cm (30 in). The earliest known perecaridian was Oxyuropoda ligioides, a fossil of which has been found dating to the Late Devonian of Ireland.

Eumalacostraca Subclass of crustaceans

Eumalacostraca is a subclass of crustaceans, containing almost all living malacostracans, or about 40,000 described species. The remaining subclasses are the Phyllocarida and possibly the Hoplocarida. Eumalacostracans have 19 segments. This arrangement is known as the "caridoid facies", a term coined by William Thomas Calman in 1909. The thoracic limbs are jointed and used for swimming or walking. The common ancestor is thought to have had a carapace, and most living species possess one, but it has been lost in some subgroups.

Cumacea Order of crustacean

Cumacea is an order of small marine crustaceans of the superorder Peracarida, occasionally called hooded shrimp or comma shrimp. Their unique appearance and uniform body plan makes them easy to distinguish from other crustaceans. They live in soft-bottoms such as mud and sand, mostly in the marine environment. There are more than 1,500 species of cumaceans formally described. The species diversity of Cumacea increases with depth.

<i>Caridina serratirostris</i> Species of crustacean

Caridina serratirostris is a species of freshwater shrimp that lives in the Indo-west Pacific region, from Madagascar to Fiji, including northern Queensland, Australia, which may be a different subspecies. Its common name in the aquarium trade, "ninja shrimp", comes from its ability to quickly change colour and disappear into its surroundings like a ninja. Adults grow to a length of 25–35 millimetres (1.0–1.4 in).

Crustacean larva Crustacean larval and immature stages between hatching and adult form

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.

<i>Hymenodora</i> Genus of crustaceans

Hymenodora is a genus of shrimp, containing four species, which collectively have a southern circumpolar distribution.

Crustacean Subphylum of arthropods

Crustaceans 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. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustaceans. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

Multicrustacea Superclass of crustaceans

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.

<i>Acanthosquilla derijardi</i> Crustacean from the Indo-Pacific region

Acanthosquilla derijardi is a species of stomatopod crustacean. Its distribution is widespread throughout the Indo-West Pacific. The species was initially described by the American carcinologist Raymond B. Manning in 1970. Its junior synonym, A. sirindhorn, was named in 1995 in honor of Princess Sirindhorn of Thailand.


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