Growth models in fossil arborescent cheilostome bryozoans

Abstract
Cheilostome bryozoans that grew as rigidly erect arborescent colonies dominate many bryozoan-rich assemblages of Tertiary age, in which they are found most commonly as small dissociated fragments. The regularity with which branching and branch thickening occur in intact colonies of living species provides a basis for quantitative reconstruction of these growth processes in fossils. We propose models to describe branch thickening, develop methods to extend both thickening and branching models to fossils, investigate the thickening and branching properties of four Paleocene and five Oligocene species and compare the properties of these fossils to those of nine living species.The properties investigated are largely mutually independent and species specific irrespective of geologic age and have similar numerical ranges among different assemblages of coeval species. Species are evenly distributed across the range of possible morphologies between observed extremes, without obvious gaps. Statistically significant trends through time are identified in gradients of branch thickening, which have implications for the resistance of colonies to mechanical stress, and in angles of bifurcation, that are important in the way growing colonies occupy space.