Diverse subphylum of mandibulate arthropods, the body usually divided into three parts: head, thorax, and abdomen. In some crustaceans (e.g. crayfish) the head and thorax may be joined to form the cephalothorax. The head bears two pairs of antennae, one pair of mandibles, and two pairs of maxillae. The limbs are biramous, and are adapted for a wide range of functions. Closely placed setae on the limbs function as filters in filter-feeding species. Respiratory gills are situated on the appendages, but vary greatly in location and number; they are absent only in very small species. In addition to the antennae, sense organs include a pair of compound eyes, and a small, dorsal, median, nauplius eye, comprising three or four closely applied ocelli (clusters of photoreceptors). The nauplius eye, characteristic of crustacean larvae, is absent in many adults; and some groups lack the compound eyes. Mainly marine, but there are many freshwater species, and a relatively small number have invaded the land. Four classes of crustaceans have an important fossil record. The Malacostraca (crabs, lobsters, woodlice, etc., Cambrian to Recent) includes the earliest crustaceans of the subclass Phyllocarida. The Branchiopoda (similar to modern water fleas, Lower Devonian to Recent) are valuable index fossils in non-marine strata. The Cirripedia (barnacles) occur from Upper Silurian to Recent, and the Ostracoda from Lower Cambrian to Recent. The living class Cephalocarida (e.g. Hutchinsonella) is thought to be closest to the ancestral crustacean stock, but the group is without any unequivocal fossil representative. Hexapods (Hexapoda) are believed to be descended from a crustacean, and hexapods and insects are grouped in the clade Pancrustacea.