Callocystites fresti sp. nov. is characterized by an elongate theca, protuberant periproct and raised ambulacral facets. Ambulacral branching in Callocystites is isotomous, but left- and right-handed branches occur because the ambulacral groove branches in the same direction twice, interrupting the regularly alternating branching to brachioles. Branching in Strobilocystites does not interrupt the alternation of brachioles and lateral food grooves always pass adoral to the first brachiole of each branch. Sphaerocystites branched as in Strobilocystites. Other morphological features suggest Callocystites, Sphaerocystites and Strobilocystites are not closely related. Callocystites has open infra-lateral and lateral circlets and five ambulacra; Sphaerocystites and Strobilocystites have closed/modified circlets and four ambulacra. In Callocystites and Sphaerocystites the B and D ambulacra differ from the others and they have one hydropore; in Strobilocystites all four ambulacra are the same and the hydropore is double. All three genera have large, globular thecae. Ambulacral branching was a response to increased food-gathering requirements. The following are phylogenetically informative callocystitid characters: truly open vs closed/modified plate circlets; number of pectinirhombs; number of ambulacra; presence vs absence of the ‘B D different’ pattern of primary brachioles; single vs double hydropore.
In echinoderms with five ambulacra in a 2–1–2 pattern ambulacral homologies are unambiguous: the single amb is A (III). In echinoderms without this pattern or with less than five ambulacra the hydropore or a single gonopore is best for orientation. Both always occur in the CD (V-I) interambulacrum. The position of the periproct is unreliable. Different patterns and numbers of ambulacra in Palaeozoic echinoderms render a single developmental sequence improbable. Cystoids (blastozoans) had radial water vessels in their ambulacra and added new ambulacral plates terminally. Functional morphological considerations suggest tube feet were essential for food gathering. Copyright © 2014 John Wiley & Sons, Ltd.