Symmetry of echinoderms: From initial bilaterally-asymmetric metamerism to pentaradiality

Echinoderm radial symmetry has first appeared in the ambulacral system, when the ambulacral channel assumed the shape of a closed ring or a horseshoe with approximated ends, and then spread onto other organ systems. Its origin was a natural consequence of a steady increase of the original asymmetry of bilaterally-asymmetric three-segmented ancestors of echinoderms, culminated by closing of the ancestral linear metamerism into radiality. The evolutionary transformation from a simple pouchlike hydrocoel with one side channel to an elongated hydrocoel (located under the oesophagus) with two side channels, and finally to a nearly closed horseshoeshaped hydrocoel with three radial channels can be reconstructed based on the theca structure and the number of ambulacra in the row Soluta-Cincta-Helicoplacoidea. After the hydrocoel with three outbound ambulacral channels circled around the oesophagus as a horseshoe, it either closed into a ring, or its ends became closely approximated. This has determined the primary triradiate symmetry, which quickly transformed into pentaradial symmetry of the 2-1-2 type as a result of branching of two of the three primary radial channels. This occurred no earlier than the Late Vendian, when the first bilaterians appeared to have begun to acquire body appendages, and no later than the Early Cambrian, when the first skeletal remains of echinoderms entered the fossil record (in the Atdabanian). The 2-1-2 pentaradial symmetry evolved into the true pentaradiality as a result of shifting the timing of tentacle branching to earlier stages of ontogenesis and even spreading of the five tentacle primordia over the ambulacral ring. This occurred during the Ordovician.