Development and metamorphosis of two psolid sea cucumbers,Psolus chitonoides andPsolidium bullatum,with a review of reproductive patterns in the family Psolidae (Holothuroidea: Echinodermata)

Reproduction and development of two psolid sea cucumbers in waters of the San Juan Archipelago, Washington, were studied from April 1981 through August 1984. Spawning inPsolus chitonoides Clark andPsolidium bullatum Ohshima in the laboratory and in the field waters of the San Juan Archipelago, Washington, occurred from February through may, with heaviest spawning in April and May. The egg ofPsolus chitonoides was bright red in color and 627µm in diameter;Psolidium bullatum released eggs that were golden yellow in color and 330µm in diameter. Maximum measured fecundities (spawned eggs) forPsolus chitonoides andPsolidium bullatum were 34 700 and 3 074, respectively. Developmental schedules were similar in both species. Early cleavages were equal, radial, and holoblastic; slightly subequal cleavage was common inPsolus chitonoides. The gastrula elongated to form a uniformly ciliated larva and the vestibule first appeared as a ventral crescent-shaped depression, after which the larva decreased in length and formed three posterior ridges that encircled the early doliolaria. Just prior to settlement, there was loss of cilia between the three ridges and on the posterior end of the early doliolaria, leaving three ciliary rings that are characteristic of the dendrochirote doliolaria larva. Vertical swimming ensued, and five primary tentacles protruded through the vestibule, with occasional attachment to the substratum using the primary tentacles. Upon settlement, two primary podia emerged and all external ciliation was lost except among tentacular papillae and on the end of tube feet. Timing of appearance and of loss of ciliary rings suggest that the rings function in vertical swimming and substratum selection behaviors. Metamorphosis involved the following simultaneous events: (1) resorption of the preoral lobe, (2) formation of large, dorsal, calcareous ossicles, (3) flattening of the trivium, and (4) proliferation of ventral tube feet on Ambulacral Radii B and E to form the sole. Reproduction in the family Psolidae is reviewed. The large proportion of brooders in this family appears to be related to small size, the possession of protective armor and sole, and geographic distribution (mainly in Antarctica). Lecithotrophic doliolaria larvae with three ciliary rings are produced by large northern psolids.