Crustacea

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Crustacea

A highly variable, species-rich group of arthropods that have inhabited marine environments since the beginning of the Cambrian Period. Within the marine realm the crustaceans occupy as diverse a spectrum of habitats as the insects inhabit on land.

The hierarchical rank of the Crustacea is a matter of continuing debate. The Crustacea are regarded as a phylum, distinct from other arthropods, by proponents of the concept of polyphyly in the Arthropoda. Alternatively, they are given the rank of subphylum or superclass by those who view the Arthropoda as a monophyletic taxon.

Species of Crustacea such as the shrimp, prawn, crab, or lobster are familiar. However, there are many more with less common vernacular names such as the water fleas, beach fleas, sand hoppers, fish lice, wood lice, sow bugs, pill bugs, barnacles, scuds, slaters, and krill or whale food. The Crustacea are one of the most difficult animal groups to define because of their great diversity of structure, habit, habitat, and development. No one character or generalization will apply equally well to all.

Crustaceans have segmented, chitin-encased bodies; articulated appendages; mouthparts known as mandibles during some stage of their life, however modified they may be for cutting, chewing, piercing, sucking, or licking; and two pairs of accessory feeding organs, the maxillules and maxillae. One or the other pair is sometimes vestigial or may be lacking. The Crustacea are unique in having two pairs of antennae: the first pair, or antennules, and the second pair, the antennae proper. The latter are almost always functional at some stage of every crustacean's life.

Taxonomy

Not only is there a lack of agreement on the rank of the Crustacea per se, but there is no consensus on hierarchial levels of subordinate taxa. The classification presented here is restricted to extant taxa and is, at best, a compromise among opposing opinions.

  • Superclass Crustacea
  • Class Cephalocarida
  • Class Branchiopoda
  • Order: Anostraca
  • Spinicaudata
  • Laevicaudata
  • Ctenopoda
  • Anomopoda
  • Onychopoda
  • Order: Haplopoda
  • Notostraca
  • Class Remipedia
  • Class Ostracoda
  • Subclass Myodocopa
  • Order: Myodocopida
  • Halocyprida
  • Subclass Podocopa
  • Order: Platyocopida
  • Podocopida
  • Class Maxillopoda
  • Subclass Mystacocarida
  • Subclass Cirripedia
  • Order: Ascothoracica
  • Thoracica
  • Acrothoracica
  • Rhizocephala
  • Subclass Copepoda
  • Order: Calanoida
  • Harpacticoida
  • Cyclopoida
  • Poecilostomatoida
  • Siphonostomatoida
  • Monstrilloida
  • Misophrioida
  • Mormonilloida
  • Subclass Branchiura
  • Subclass Tantulocarida
  • Class Malacostraca
  • Subclass Phyllocarida
  • Order: Leptostraca
  • Subclass Hoplocarida
  • Order: Stomatopoda
  • Subclass Eumalacostraca
  • Superorder Syncarida
  • Order: Bathynellacea
  • Anaspidacea
  • Superorder Peracarida
  • Order: Spelaeogriphacea
  • Mysidacea
  • Mictacea
  • Amphipoda
  • Isopoda
  • Tanaidacea
  • Cumacea
See Branchiopoda, Cephalocarida, Malacostraca, Maxillopoda, Ostracoda, Remipedia

General morphology

The true body segments, the somites or metameres, are usually somewhat compressed or depressed. Each typically includes one pair of biramous appendages. The linear series of somites making up the body of a crustacean are more or less distinctly organized into three regions or tagmata: the head, thorax, and abdomen. Where regional organization of the postcephalic somites is not clearly marked, they collectively form the trunk. The somites are variously fused with one another in diagnostic combinations in different groups of the Crustacea.

Body

A dorsal shield or carapace of variable length arises from the dorsum of the third cephalic somite and covers the cephalon and cephalothorax to varying extent. The carapace reaches its greatest development in the malacostracan Decapoda (shrimps, lobsters, and crabs).

The chitinous cuticle covering the crustacean body is its external skeleton (exoskeleton). The chitin is flexible at the joints, in foliaceous appendages, and throughout the exoskeletons of many small and soft-bodied species, but it is often thickened and stiff in others. It becomes calcified in many species as a result of the deposition of lime salts.

The paired appendages are typically biramous and consist of two branches: the endopod and exopod. The endopod is definitely segmented in the higher Crustacea. The endopods are variously modified to serve a variety of functions and needs such as sensory perception, respiration, locomotion, prehension and comminution of food, cleansing, defense, offense, reproduction, and sex recognition and attraction. If retained in the adult, the exopod may remain leaf- or paddlelike, or become flagellated structures, facilitating swimming or aiding respiration.

Crustacea take up oxygen by means of gills, the general body surface, or special areas of it. Some of the few species that have become more or less terrestrial in their habits have developed modifications of their branchial mechanism such as water-retaining recesses which when sufficiently moist enable them to breathe air. Some sow or pill bugs have special tracheal developments in their abdominal appendages for the same purpose.

Reproductive system

The sexes are separate in most Crustacea and usually can be differentiated from each other by secondary sex characters. Chief among these characters are the size and shape of the body, appendages, or both, and placement of the genital apertures. Hermaphroditism is the rule in the Cephalocarida, Remipedia, some ostracods, sessile Cirripedia (barnacles), in isolated cases in other crustaceans, and in certain parasitic forms. Parthenogenesis (eggs developing and hatching without prior fertilization) occurs frequently in some of the lower crustaceans that have what might be called an alternation of generations. The parthenogenetic generations alternate with a generation produced by fertilized eggs. See Cirripedia

The eggs of most crustaceans are carried attached to the female until hatched. Some females develop brood pouches in which the young are retained for a time. A nutrient secretion which sustains the young until they are released is produced in some species having a brood chamber. Penaeid shrimp and a few of the lower Crustacea deposit their eggs in the medium in which they live, in some cases attaching them to aquatic vegetation.

Development

The nauplius larva is characteristic of Crustacea. This first larval stage is common in the lower forms, but in many of the higher forms it occurs during development in the egg, and the young are hatched as a different and more advanced larva or, as in many Malacostraca, in a form similar to the adult. Life histories vary from the simple to the complex within the different groups of Crustacea.

Molting (ecdysis)

This process involves several steps: (1) preparation, which includes some degree of resorption of the old cuticle; (2) the formation of a new, temporarily soft and thin one within it; (3) the accumulation and storing of calcium in the midgut gland or as lenticular deposits (gastroliths). The preparatory period is less complicated in the thinly chitinous forms. The actual molt follows. The old shell or cuticle splits at predetermined places, permitting the crustacean within, already enclosed in new but still soft exoskeleton, to withdraw. A temporary absorption of water enables the animal to split or crack its housing. Upon withdrawal of the entire animal, absorption of water again rapidly takes place with a pronounced increase in body size. The tender new cuticle is reinforced rapidly by the resorbed chitin, and hardened by whatever reserves of calcium the animal may have stored, supplemented and extended by the far more plentiful supplies in solution in the sea which may be absorbed or ingested by the growing crustacean. Molting takes place quite frequently in the larval stages when growth is rapid, but becomes less frequent as the animal ages. In many species there is a terminal molt at maturity.

Autotomy and regeneration

The mechanisms of autotomy and regeneration are developed in the crustaceans to minimize injury or loss to an enemy. When an appendage is broken, it is cast off or broken at the fracture or breaking plane. This sacrifice often enables the victim to escape. Even more remarkable is the fact that crustaceans, by voluntary muscular contraction, can part with a limb which may be injured. Crustacea also have the ability to regenerate lost parts. Although the regenerated parts are not always the same size as the original in the first molt after injury, increase in size in successive molts soon restores a lost limb to virtually its former appearance.

Bionomics and economics

Crustacea are ubiquitous. They live at almost all depths and levels of the sea, in fresh waters at elevations up to 12,000 ft (3658 m), in melted snow water, in the deepest of the sea's abysses more than 6 mi (9 km) down, and in waters of 0°C (32°F) temperature. Some species live on land, although most must descend to salt water areas again to spawn their young. Some live in strongly alkaline waters and others in salt water which is at the saturation point, still others in hot springs and hydrothermal vents with temperatures in excess of 55°C (131°F).

Crustacea are of all sizes, ranging from copepods 0.01 in. (0.25 mm) long to huge spider crabs of Japan, which span 12 ft (3.7 m) from tip to tip of the laterally extended legs. The American lobster, the heaviest so far known, tops all crustaceans at 44½ lb (20 kg).

Most crustaceans are omnivorous and essentially scavengers. Many are filter feeders and screen particulate life, plankton, and organic detritus from the waters in which they live; others are largely carnivorous, still others vegetarian. Among the vegetarians are the grazers of the ocean meadows which convert the microscopic plant life (diatoms) into flesh and food for larger animals which in turn are harvested as food for humans.

Crustacea

 

a class of aquatic animals of the phylum Arthropoda. The Crustacea include the subclasses Branchiopoda, Cephalocarida, Maxillopoda, Ostracoda, and Malacostraca. There are approximately 20,000 species. The body of a crustacean ranges from a fraction of a millimeter to 80 cm in length and consists of a head, thorax, and segmented abdomen. The abdomen is covered with a chitinous cuticle, which often contains lime and forms a carapace. The head consists of a preoral lobe and four somites, the first of which fuses with the preoral lobe to form the primary head, or protocephalon; the three posterior somites form the maxillary part of the head, or the gnathocephalon. In some crustaceans (the orders Anostraca, Mysidacea, Euphausiacea, Decapoda, and Stomatopoda) the protocephalon and gnathocephalon are separate; in others they fuse into a compound head, known as the syncephalon. The head has two pairs of antennae (antennules and antennae), upper jaws (mandibles), and two pairs of lower jaws (maxillae). The antennae serve as sense organs and, sometimes, as organs of locomotion. The remaining appendages participate in the gathering and grinding of food.

The thorax consists of eight somites in Malacostraca and an indefinite number of somites in other crustaceans. Sometimes the four anterior thoracic somites are fused with the head, with their limbs converted to maxillipeds. The remaining thoracic limbs serve for locomotion and often bear gills. The head and thorax in some crustaceans are covered with a common carapace, which in the Ostracoda is in the form of a bivalve shell. The abdomen in most Malacostraca consists of six somites with legs; in other crustaceans, the abdominal somites vary in number and are limbless. The limbs of crustaceans are biramous: each limb consists of a two- or three-segmented part, two articulated branches (an exopodite and an endopodite), and a gilled appendage (epipodite). One of the branches of the limb often does not develop.

The nervous system of crustaceans resembles a ventral nerve cord. Vision is provided by a pair of compound eyes or, less commonly, by an unpaired, or median, eye. The organs of equilibrium are statocysts. The alimentary canal usually has a masticatory stomach and a “liver,” which opens into the midgut. The circulatory system is lacunar, and the muscular heart lies on the dorsal side in a pericardium. The organs of respiration are gills, which are situated on the limbs or on the sides of the body. In some cases, respiration is cutaneous. The excretory organs are coelomoducts that have been modified into antennal and maxillary glands, which open at the bases of the antennae and maxillae. In most adult Malacostraca the antennal glands function, whereas in their larvae the maxillary glands function. In the other subclasses, this is reversed. In the majority of crustaceans the sexes are separate, but many Cirripedia, which are sessile, are hermaphrodites. Fertilization is external: the males attach spermatophores near the genital openings of the females.

Most crustaceans are characterized by a larva—the nauplius—which has three pairs of segmented appendages (antennules, antennae, and mandibles); the antennae and mandibles are biramous. In some species the nauplius emerges from the egg and leads a natatorial life; in others its development proceeds under the protection of the egg membranes. After the nauplius stage there are a series of other larval stages; with each, the number of somites and their paired extremities gradually increases. Some crustaceans (Cirripedia, Mysidacea, Cumacea, Isopoda, Amphipoda, and many Decapoda) are characterized by direct development, that is, an individual differing little from the adult emerges from the egg.

Most crustaceans inhabit seas, making up the principal mass of plankton and, sometimes, a significant part of the benthos. In freshwaters, crustaceans also predominate in the plankton. Only a few species have adapted to terrestrial life (woodlice, beach fleas, some tropical decapods). All Ascothoracica and some Copepoda and Cirripedia are parasites. Planktonic crustaceans feed on bacteria, with a few feeding on unicellular organisms and detritus. Benthic crustaceans feed on plant or animal substances. Amphipoda eat animal carcasses, thus promoting the purification of bodies of water.

The origin of crustaceans is not clear. According to one hypothesis, they are descended from ancient extinct arthropods—trilobites; another theory identifies annelids, independently of trilobites, as the ancestors of crustaceans. In the fossil state, crustaceans have been traced to the Cambrian. Fossil Phyllopoda and Ostracoda are especially numerous; the latter serve as index fossils in stratigraphy. Many Decapoda are used by man as food. They are commercially valuable and have been bred (king crab, lobsters, spiny lobsters, shrimp, crayfish). Crustaceans also serve as food for many commercially valuable fishes, such as herring and the Pacific sardine. Some parasitic Copepoda bring serious harm to fishes. A number of crustaceans destroy wooden structures in the sea, and Cirripedia attach themselves in large numbers to the bottom of ocean vessels. Eriocheir sinensis, which appeared recently in Europe, digs burrows, destroying embankments and dams. It also tears fishing nets and spoils the fish caught in them. Some crustaceans are intermediate hosts of parasitic worms (broad tapeworm, Dracunculus medinensis, Acanthocephala).

REFERENCES

Birshtein, Ia. A. “Vysshie raki (Malacostraca).” In Zhizn’ presnykh vod SSSR, vol. 1. Moscow-Leningrad, 1940.
Bol’shoi praktikum po zoologii bespozvonochnykh, part 2. Moscow, 1946.
Iashnov, V. A. “Klass Crustacea—rakoobraznye.” In Opredelitel’ fauny i flory severnykh morei SSSR. Moscow, 1948.
Gur’ianova, E. F. Bokoplavy morei SSSR i sopredel’nykh vod. Moscow-Leningrad, 1951.
Ivanov, A. V. Promyslovye vodnye bespozvonochnye. Moscow, 1955.
Beklemishev, V. N. Osnovy sravnitel’noi anatomii bespozvonochnykh, 3rd ed., vols. 1-2. Moscow, 1964.
Zhizn’ zhivotnykh, vol. 2. Moscow, 1968.
Dogel’, V. A. Zoologiia bespozvonochnykh, 6th ed. Moscow, 1974.

A. V. IVANOV

Crustacea

[krə′stā·shə]
(invertebrate zoology)
A class of arthropod animals in the subphylum Mandibulata having jointed feet and mandibles, two pairs of antennae, and segmented, chitin-encased bodies.
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