Origin of Echinodermata

The common ancestor of Ambulacraria was a mobile bilaterally symmetrical organism. Its body was subdivided into a preoral (proboscis) region, a perioral (collar) region with ciliated tentacles, and a trunk region with a metameric coelom and metameric gill slits. Like in other Bilateria, the preoral and tentacular regions of Ambulacraria are free of Hox gene expression, which begins in the area of the first pair of gill slits in compliance with the colinearity. The axial complex is a synapomorphy of the Ambulacraria clade. A bilaterally symmetrical ancestor of Echinodermata lay on its dorsal side, so its anus was shifted to the ventral side as in present-day echinoderm larvae. This stage of evolution corresponds to the Early Paleozoic bilaterally symmetrical forms, such as Protocinctus, Ctenocystis, and Ctenoimbricata. The common ancestor of echinoderms had a symmetrical tentacular apparatus consisting of five ciliated tentacles on each side of a collar. At the next stage, the ancestors of echinoderms lay on the right side that resulted in the reduction of the tentacles on the right side and the right hydrocoel. This evolution stage includes various Early Paleozoic forms (Cothurnocystis, Dendrocystoides, Syringocrinus, Castericystis, Coleicarpus, Rhenocystis, etc.). The next stage is related to the sedentary lifestyle. During this stage, the mouth and tentacles occupied an apical position, the anus has shifted up, and thus an intestinal loop was formed. The five primary tentacles of the left side of the ancestor of Ambulacraria predestined the formation of pentaradial symmetry of echinoderms. The primary tentacles remained only in Holothuroidea. The secondary ciliary grooves were formed between the primary tentacles. These grooves were accompanied by hydrocoelic canals, which gave rise to the water-vascular (ambulacral) system. The present-day echinoderms are characterized by several metameric rings formed by the derivatives of the left somatocoel. They derive from the coelomic segments of the left side of the trunk of the ambulacrarian ancestor and, thus, the echinoderms retain coelomic metamerism. The attachment of the crinoid larvae by the preoral lobe reflects the ancient method of locomotion of deuterostomes using the proboscis, but, in reality, the Pelmatozoa stalk is homologous to the Pterobranchia stalk, i.e., the posterior end of the body, because the right somatocoel grows into it. During metamorphosis, the internal coelomic complex of the larva is inverted by 180°. The significance of this inversion for the translocation of the anterior genes of the Hox cluster is discussed. The ancestors of Eleutherozoa began to crawl on the oral surface, which led to a shift of the anus to the aboral side. Thus, the ontogeny and phylogeny of echinoderms exhibits a change from bilateral symmetry to dissymmetry, and then the development of pentameric symmetry.