Ontogenetic regulatory mechanisms and evolution of mellitid lunules (Echinoidea, Clypeasteroida)

Abstract

Lunules have evolved independently in several groups of clypeasteroids, including the Rotulidae, Astriclypeidae, Mellitidae and Scutasteridae. In this paper, only the monophyletic assemblage Monophorasteridae plus Mellitidae is considered. Lunules result from modifications of the growth patterns of test plates which bring about changes in relative growth in specific directions. It is unnecessary to postulate resorption of the test. Ambulacral lunules, which are known to have hydrodynamic functions, originated as part of a series of changes: 1) bifurcation of food grooves, 2) formation of pressure drainage channels, 3) lobulation of the ambitus (as inMonophoraster), 4) complete lunule formation. The anal lunule shares the hydrodynamic function but arose separately, as a developmental aberration. The position of the periproct is highly variable but it is most commonly located at the junction of suture lines, at plate corners. InScutellait is located between the first and second post-basicoronal plates. InMonophorasterthe periproct is located further forward, between the first post-basicoronal plates, and the small anal lunule occupies the junction point between first and second post-basicoronals. It is hypothesized that the anal lunule originated as an all-or-nothing event following the forward migration of the periproct and failure to resume normal plate growth at the sutural junction point between first and second post-basicoronal plates. Its walls were derived from paired interambulacral supports. The hypothesis is discussed in connection with the ontogenetic formation of the anal lunule in living mellitid sand dollars.