Heterochrony and heterotopy in the phylogeny of sea urchins

Abstract

The role of heterochrony is evident in ontogeny and phylogeny of irregular (exocyclic) sea urchins. After metamorphosis, a juvenile passes the stage of regular (endocyclic) sea urchin, in which the periproct is surrounded by plates of the apical system. A shift of the periproct in the area of the fifth interambulacrum occurs in extant taxa at early stages of postlarval development and is accompanied by the reduction of genital plate 5. In some ancient (Jurassic) adult irregular sea urchins, the endocyclic state of the apical system is retained for a long time and the derivative of the fifth genital plate is sometimes observed even in Early Cretaceous species. Considerable transformations in the structure of the lower test surface in members of the order Spatangoida are manifested in changes in the relative positions of plastron plates and ambulacral areas I and V, separation of sternal plates from the labrum, etc. The mechanism of these changes is connected with translocation or “sliding” of sutures of particular plates as a result of nonuniform growth and partial resorption. The study of evolutionary lineages of Cretaceous and Cenozoic sea urchins has shown that the evolution was connected with the directional changes in some morphological characters at late ontogenetic stages. The process was accompanied by either extension, peramorphosis (lineages of the genera Micraster, Infulaster–Hagenowia in the European Province), or the loss of these stages, paedomorphosis (Hemiaster (Bolbaster) lineage, Late Eocene–Middle Miocene of Australia). The phenomena of heterochrony and heterotopy in the development of peripetal, marginal, and lateroanal fascioles in the Late Cretaceous and Paleocene families Hemiasteridae, Schizasteridae, and Paleopneustidae are described. The heterotopy is also illustrated by the example of the development of additional genital pores on ocular plates II and IV of the Middle Jurassic species Pygomalus analis (Disasteroida); its apical system has five pores instead of four. In the Late Cretaceous species Guettaria roccardi (Holasteroida), ocular plates II and IV have two pores each; in the apical system, there are eight genital pores instead of four. In some members of the order Holectypoida, the place of lost genital plate 5 is occupied by a new plate sometimes pierced by a pore, but judging from crystallographic data, it is not homologous to other genital plates. The order Clypeasteroida is characterized by the development of very small pores in both ambulacral and interambulacral fields; they provide passage for numerous accessory tube feet.