Columnal ontogeny in articulate crinoids and its implications for their phylogeny

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Abstract
Detailed documentation of columnal ontogeny in several Lower Jurassic isocrinids reveals a number of features which are of phylogenetic significance. Columnal proportions are determined by two largely independent growth vectors. Columnal diameter is primarily a function of cup size of the time of formation, though it may subsequently undergo considerable increase through lateral accretion. Columnal height is largely independent of cup size except for a gradual increase with the growth of the whole animal. Columnal diameter increases at a much greater rate than columnal height and so columnals are relatively lower in large individuals. Heterochronous variation in vertical growth rate or the extent of lateral accretion in columnals may significantly alter their shape. The smallest isocrinid columnals found (0.35 mm diameter) have synarthrial articula. The transformation to a typical pentaradiate symplexy, via a triradiate stage, occurs at a diameter of about 0.75 mm. Synarthrial articula are unknown in millericrinid columnals, which instead have symplectial articula at small diameters (0.45 mm). The presence of synarthrial articula in the stems of juvenile isocrinids, and their absence in millericrinids, supports previous hypotheses concerning derivation of bourgueticrinids from an isocrinid, rather than a millericrinid, ancestor.