Segmental mismatch in crustacean appendages: the naupliar antennal exopod of Artemia (Crustacea, Branchiopoda, Anostraca)

Based on traditional techniques and confocal laser scanning microscopy for external morphology, and immunohistochemistry for the muscular system, we describe here the segmental features of the antennal exopod of Artemia nauplii. Two kinds of serial elements are present, i.e. setae (with cuticular folds at their base) and ringlets (serially arranged sclerites separated by joint-like cuticular folds not extending to form complete rings around the appendage). The two series are usually not in register. The cuticular folds of the setae and of the ringlets are also sites of intermediate insertions of the three exopod muscles: as the two tegumentary structures are discordant in periodicity, this is also mirrored in the pattern of muscle insertions on the two sides of the appendage. Similar cases of segmental mismatch are known for the trunk of several arthropods, but segmental mismatch along the appendages has received very little attention. The occurrence of segmental mismatch in the naupliar appendages of both extant and fossil crustaceans is reviewed and it is suggested here to be a primitive feature of the exopods of both second antennae and mandibles. Problems in the interpretation of morphological evidence are discussed, also in relation to development and evolution of segmentation of naupliar appendages.