# The Life Cycle of the Tantulocarida (Crustacea)

G. A. Boxshall, R. J. Lincoln

## Abstract

Four new species of parasitic crustaceans belonging to the class Tantulocarida are described, two of which are placed in a new genus, Onceroxenus. Three of them parasitize deep-sea tanaids, the other, a deep sea asellote. Microdajus langi, originally classified as an epicaridean isopod, is recognized as a tantulocaridan. It is reported from Scottish waters for the first time and from new host species. These records include the shallowest depth, 22 m, known for a tantulocaridan. Cumoniscus kruppi, a parasite of cumaceans, is also recognized as a tantulocaridan. The Tantulocarida now comprises eleven species and five genera, here assigned to the Basipodellidae and two new families, the Deoterthridae and Microdajidae. Several life cycle stages are described and arranged in two developmental sequences. Evidence for a possible third sequence was found. Male development involves a unique type of metamorphosis in which the free-living adult differentiates from a dedifferentiated mass of tissue contained within the expanded trunk of the tantulus larva. Throughout this metamorphosis the male is supplied with nutrients from the host via a tissue connection, the umbilical cord, and the permanently attached larval head. The non-feeding adult male lacks cephalic appendages but possesses two clusters of aesthetascs on its anterior margin. It is free swimming and has six pairs of large thoracopods without endites. The first two thoracic somites are incorporated into the cephalothorax. The abdomen bears a posteriorly directed, median stylet, interpreted as the intromittent organ. It originates on the first abdominal somite. The adult female has a large sac-like trunk attached by the larval head. The larval trunk is sloughed leaving a scar but no complete moult occurs. Eggs develop within the trunk sac and hatch directly at the infective tantulus larval stage. This extreme condensation of early ontogeny is compared with that of other crustaceans and is interpreted as an adaptation to parasitism in situations where a high dispersal ability is not advantageous. In some females the trunk sac forms behind the head but the larval trunk is retained. Small and large females of this type are described, the largest being 737 <latex>$\mu m$</latex> in length. These probably represent females in which sloughing of the larval trunk has failed but it is possible that each may have contained a free-living adult female of comparable size to the adult male. The tantulus larva is described in detail. Scanning electron microscopy reveals that the thoracopodal endites have a complex apical armature, including coupling spines which serve to link the members of a leg pair. Tantulocaridans are permanently attached to their host by the oral disc, presumably by means of an adhesive. In the centre of the disc they make a minute puncture (between 0.5 and 2.0 <latex>$\mu m$</latex> in diameter) through the host integument, probably with the aid of their cephalic stylet. This constitutes their only access to the body fluids of the host. The phylogenetic relationships of the Tantulocarida are discussed. They appear to be related to the barnacles (Thecostraca), both groups possessing a median penis derived from the seventh trunk limb. Their possession of a thorax of six somites and the location of the male gonopores on trunk somite seven suggests an affinity with a larger group containing the Thecostraca and the Copepoda.