A cystoid with two left facets: the significance of Tetracystis in the evolution and classification of the Callocystitidae (Echinodermata, Glyptocystitoida)

Abstract

Revision of some North American Silurian callocystitid cystoids shows that Tetracystis Schuchert has the first two brachioles to the left in ambulacra B and D (B D different pattern), whereas Troosticystis Paul and Lovenicystis Regéll have only the first brachiole to the left in all four ambulacra (B–E the same pattern). ‘Apiocystites’ elegans Hall is transferred to Tetracystis because it has the B D different pattern. Lovenicystis kopfi sp. nov. is characterized by four short ambulacra. Salirocystis gen. nov. has the B D different pattern and two hydropores. All callocystitids that lack ambulacrum A are assigned to a new subfamily, the Tetracystinae; those with five ambulacra are referred to informally as ‘lepadocystines’. The previously recognized subfamily Callocystitinae is reduced in rank to a tribe within the ‘lepadocystines’. Preliminary character analysis suggests that the loss of ambulacrum A was the most significant evolutionary event within the family Callocystitidae, but has been overlooked in taxonomy. Character analysis also shows that all ‘lepadocystines’ had the B D different pattern of primary brachioles, which is retained in four tetracystine genera, Lepocrinites Conrad, Sphaerocystites Hall, Tetracystis and Salirocystis. Since Salirocystis has the plesiomorphic B D different pattern plus the apomorphic double hydropore, one cannot define a monophyletic taxon characterized by the B–E the same pattern plus one characterized by a double hydropore. Both alternatives are equally parsimonious. Here a new monophyletic tribe Lipsanocystini is defined characterized by a double hydropore. The B–E the same pattern evolved independently twice. Among current characters the loss of ambulacrum A, development of the B–E the same pattern and development of two hydropores are the most significant evolutionary events. Number of periproct border plates, length and structure of ambulacra are less significant. Copyright © 2014 John Wiley & Sons, Ltd.

Origin and significance of Lovén's Law in echinoderms

“…Sumrall (2008) explained the ‘BD different pattern’ as arising because the first left facet in the B and D ambulacra represented the first facet of the combined B + C and D + E ambulacra, whereas the second left facet represented the first facet of the separate B and D ambulacra. Paul (2015b, p. 626–627) criticized this interpretation on two grounds. First, that the two different ambulacra under Lovén's Law were the A and D ambulacra of Carpenter.…”