ALBERT

Description: Many cheilostome bryozoans of diverse phylogenetic origin grow as erect, arborescent colonies with branches of modified planar form composed of two layers of zooids back to back. Regular branching enables a growing colony to expand in surface area, and hence in the number of zooids that feed, reproduce, and perform other vital functions, at an accelerating rate. During growth, branches first all diverge, then increasingly converge, and in late stages of growth begin to interfere with each other's growth and function. Interference can set limits to the width and thickness of branches and hence to the number and size of zooids.Simulation of growth using a 3–dimensional mathematical model shows that a narrow range of possible values of branching angles minimizes branch interference in late growth stages. These values are prevalent in fossil and modern species. Branch spacing at later growth stages is correlated with the distance between branches at first crossing, providing room for feeding organs of the two facing layers of zooids to protrude and function. Interbranch distances dwindle as branches increasingly converge, so emphasis on minimizing interference at a late stage sets a practical limit to growth beyond that stage. To gain this long-term benefit requires adhering to a regular pattern throughout growth. The considerable variation in branching properties in fossil and modern species, and a variability in spacing inherent in the growth pattern itself, limit the amount of usable interbranch space. Despite a higher intraspecific variability, branching properties are as distinctive interspecifically as zooidal properties, and variability is randomly distributed through the colony. A small reduction in variability between fossil and modern species suggests that increasing regularity may provide a selective advantage in the utilization of interbranch space.

Description: 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.