Origin and significance of Lovén's Law in echinoderms

Lovén’s Law described the position of larger basicoronal ambulacral plates in echinoids. The smaller basicoronal plates form first in ontogeny. We restate Lovén’s Law to describe the position of first ambulacral plates using Carpenter’s ambulacra and left (L) or right (R) as: AR, BL, CR, DL, ER, with EA the pair of ambulacra both identical and adjacent. First ambulacral plates of the Cambrian edrioasteroid, Walcottidiscus, code identically. The transition from a tri-radiate 1-1-1 pattern to the 2-1-2 ambulacral pattern of Walcottidiscus and other pentaradiate Paleozoic echinoderms results in Lovén’s Law. This provides an hypothesis for the origin of Lovén’s Law and predicts its widespread occurrence among echinoderm classes. The ‘BD different’ pattern of primary brachioles in pentaradiate glyptocystitoid Rhombifera results from subterminal branching of the ambulacra. The ontogenetic sequence was triradiate, then lateral ambulacra bifurcated, and finally second brachioles developed. Positions of second brachioles of pentaradiate glyptocystitoids code as AR, BL, CR, DL, ER. Other examples of Lovén’s Law occur in Diploporita, Ophiuroidea, Edrioasteroidea, Edrioblastoidea, Holothuroidea, Ophiocistioidea, and Bothriocidaris. Our working hypothesis is that Lovén’s Law arose with pentaradiality. An objective test requires a reliable independent landmark across all classes, which does not exist currently. Our hypothesis is only parsimonious if Lovén’s Law arose just once in echinoderm evolution and is widespread within the phylum. So far, both conditions appear to be met, but edrioasteroid first ambulacral cover plates commonly exhibit an alternative Lovén-like pattern of AL, BL, CR, DL, ER, with AB the identical adjacent ambulacra.