crustacean

 

(krŭstā'shən) , primarily aquatic arthropod of the subphylum Crustacea. Most of the 44,000 crustacean species are marine, but there are many freshwater forms. The few groups that inhabit terrestrial areas have not been particularly successful in an evolutionary sense; most require very humid environments in order to survive.

Types of Crustaceans

Crustaceans can be divided according to size into two main groups. The larger group, which is divided into the classes Branchiopoda, Copepoda, Ostracoda, and Cirripedia, includes the familiar shrimp, crayfish, lobsters, and crabs. The smaller group includes species that are either microscopic or range up to a few inches (about 5 cm) in size. Most of the smaller marine forms can be found in plankton (see marine biology) and thereby occupy an important position in the marine food chain. Other copepods supply food for small fish, and still others exist as parasites on the skin and gills of fish. Best known of the smaller freshwater crustaceans are members of the genus Daphnia (water fleas), the fairy shrimp (a phyllopod that swims inverted), and Cyclops (a copepod). The order Isopoda includes the only large group of truly terrestrial crustaceans. Known as wood lice, sow bugs, or pillbugs, these small animals can be found under the bark of trees, beneath stones and rocks, and in other damp places. When disturbed they curl up armadillolike, withdrawing into the exoskeleton.

Crustacean Anatomy

All crustaceans have bilaterally symmetrical bodies covered with a chitinous exoskeleton, which may be thick and calcareous (as in the crayfish) or delicate and transparent (as in water fleas). Since it does not grow, the exoskeleton must be periodically molted when the animal undergoes metamorphosis (typically from free-swimming larva to adult) or simply outgrows its shell. The free-swimming larva characteristic of crustaceans, called a nauplius larva, has an unsegmented body, a median eye, and three pairs of appendages.

Like other arthropods, adult crustaceans have segmented bodies and jointed legs; the segments are usually grouped into a recognizable head, thorax, and abdomen. In the majority of larger crustaceans the head and thorax are fused into a cephalothorax, which is protected by a large shieldlike area of the exoskeleton called the carapace. The head bears two pairs of antennae, usually one median eye and two lateral eyes, and three pairs of biting mouthparts—the mandibles and the two pairs of maxillae. Crustacean appendages have undergone extensive adaptation for various tasks such as swimming, sensory reception, and walking. Many species have the first pair of thoracic appendages modified into claws and pincers. The gills are generally attached at the bases of the thoracic appendages, and the beating of the appendages creates a flow of water over the gills that facilitates respiration. Reproduction is sexual, and in most forms the sexes are separate. In many species the eggs are brooded beneath the abdominal segments of the female.

Classification

Crustaceans constitute the subphylum Crustacea of the phylum Arthropoda.

Crustacea Fossil range: Cambrian to recent Abludomelita obtusata, an amphipod Scientific classification Kingdom: Animalia Phylum: Arthropoda Subphylum: Crustacea Brünnich, 1772 Classes & Subclasses Branchiopoda Phyllopoda Sarsostraca Remipedia Cephalocarida Maxillopoda Thecostraca Tantulocarida Branchiura Pentastomida Mystacocarida Copepoda Ostracoda Myodocopa Podocopa Malacostraca Phyllocarida Hoplocarida Eumalacostraca

The nauplius larva of a dendrobranchiate

Porcellio scaber, the common rough woodlouse, a terrestrial crustacean

Pollicipes polymerus, the gooseneck barnacle

Glyphea pseudastacus, a fossil glypheoid

The crustaceans (Crustacea) are a large group of arthropods, comprising approximately 52,000 described species ^[1], and are usually treated as a subphylum ^[2]. They include various familiar animals, such as lobsters, crabs, shrimp, crayfish and barnacles. The majority are aquatic, living in either fresh water or marine environments, but a few groups have adapted to terrestrial life, such as terrestrial crabs, terrestrial hermit crabs and woodlice. The majority are motile, moving about independently, although a few taxa 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 head-first to the substrate and cannot move independently.

The scientific study of crustaceans is known as carcinology. Other names for carcinology are malacostracology, crustaceology and crustalogy, and a scientist who works in carcinology is a carcinologist, crustaceologist or crustalogist.

Structure of crustaceans

Crustaceans have three distinct body parts: head, thorax, and abdomen (or pleon), although the head and thorax may fuse to form a cephalothorax. The head bears two pairs of antennae, one pair of compound eyes and three pairs of mouthparts. The thorax and pleon bear a number of lateral appendages, including the gills, and the tail ends with a telson. Smaller crustaceans respire through their body surface by diffusion ^[3], and larger crustaceans respire with gills or, as shown by Birgus latro, with abdominal lungs ^[4]. Both systems (diffusion and gills) were being used by various crustaceans as early as the Middle Cambrian ^[5].

In common with other arthropods, crustaceans have a stiff exoskeleton which must be shed to allow the animal to grow (ecdysis or moulting). Various parts of the exoskeleton may be fused together; this is particularly noticeable in the carapace, the thick dorsal shield seen on many crustaceans. 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 uniramous. There is some doubt whether this is a derived state, as had been traditionally assumed, or whether it may be a primitive state, with the branching of the limbs being lost in all extant arthoropod groups except the crustaceans. One piece of evidence supporting the latter view is the biramous nature of trilobite limbs ^[6].

Despite their diversity of form, crustaceans are united by the special larval form known as the nauplius.

Although a few are hermaphroditic, most crustaceans have separate sexes, which are distinguished by appendages on the abdomen called swimmerets or, more technically, pleopods. 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 many decapods, the eggs are retained by the females until they hatch into free-swimming larvae.

Taxonomy

Although the classification of crustaceans has been quite variable, the system used by Martin and Davis ^[1] is the most authoritative, and largely supersedes earlier works.

Six classes of crustaceans are generally recognised:

• Branchiopoda — including brine shrimp (Artemia) and Triops (Notostraca)
• Remipedia — a small class restricted to deep caves connected to salt water, called anchialine caves
• Cephalocarida — horseshoe shrimp
• Maxillopoda — various groups, including barnacles and copepods. It contains Mystacocarida and Branchiura, which are sometimes treated as their own classes.
• Ostracoda — small animals with bivalve shells
• Malacostraca — the largest class, with the largest and most familiar animals, such as crabs, lobsters, shrimp, krill and woodlice.

The exact relationships of the Crustacea to other taxa are not yet entirely clear. Under the Pancrustacea hypothesis ^[7], Crustacea and Hexapoda (insects and allies) are sister groups. Studies using DNA sequences tend to show a paraphyletic Crustacea, with the insects (but not necessarily other hexapods) nested within that clade.

Fossil record

Those crustaceans that have hard exoskeletons reinforced with calcium carbonate, such as crabs and lobsters, tend to preserve well as fossils, but many crustaceans have only thin exoskeletons. Most of the fossils known are from coral reef or shallow sea-floor environments, but many crustaceans live in open seas, on deep sea-floors or in burrows. Crustaceans tend, therefore, to be rarer in the fossil record than trilobites. Some crustaceans are reasonably common in Cretaceous and Caenozoic rocks, but barnacles have a particularly poor fossil record, with very few specimens from before the Mesozoic era.

The Late Jurassic lithographic limestone of Solnhofen, Bavaria, which are famous as the home of Archaeopteryx, are relatively rich in decapod crustaceans, such as Eryon (an eryonoid), Aeger (a prawn) or Pseudastacus (a lobster). The "lobster bed" of the Greensand formation from the Cretaceous period which occurs at Atherfield on the Isle of Wight contains many well preserved examples of the small glypheoid lobster Mecochirus magna. Crabs have been found at a number of sites, such as the Cretaceous Gault clay and the Eocene London clay.

Consumption

Many crustaceans are consumed by humans, and nearly 10,000,000 tons were produced in 2005 ^[8]. The vast majority of this output is of decapod crustaceans: crabs, lobsters, shrimp and prawns. Over 70% by weight of all crustaceans caught for consumption are shrimp and prawns, and over 80% is produced in Asia, with China alone producing nearly half the world's total. Non-decapod crustaceans are not widely consumed, with only 130,000 tons of krill being caught, despite krill having one of the greatest biomasses on the planet.

References

General references

• Frederick Schram (1986). Crustacea. Oxford University Press, 620 pp. ISBN 0-19-503742-1. 

External links

• Crustacea.net, an online resource on the biology of crustaceans

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