Crustacean

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Crustaceans

The crustaceans (Crustacea) are a large group of arthropods (55,000 species), usually treated as a subphylum . They include various familiar animals, such as lobsters, crabs, shrimp 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), 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 , and larger crustaceans respire with gills or, as shown by Birgus latro, with abdominal lungs . Both systems (diffusion and gills) were being used by various crustaceans as early as the Middle Cambrian .

In common with other arthropods, crustaceans have a stiff exoskeleton which must be shed to allow the animal to grow (ecdysis). 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 an advanced 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 .

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 & Davis 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 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 , 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.

Geological history

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 Cretacoues Gault clay and the Eocene London clay.

References

General references

External links

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