Growth in encrusting cheilostome bryozoans: I. Evolutionary trends

Growth of the colony is a basic element of morphological evolution and life history in cheilostome bryozoans. Here we consider the occurrence of different modes of growth in encrusting cheilostomes through geologic time and in well-studied living associations. We assess patterns of zooid formation by direct examination of skeletal characters in species from nearly all diverse fossil assemblages reported from North America and quantify within-assemblage diversities and abundance rankings for fossil encrusting species with different modes of growth. These data document macroevolutionary trends showing a transition from dominance of an apparently primitive mode of budding in the Early Cretaceous to derived modes through the Tertiary. The trends are characterized by their long duration and apparent convergence among systematic subgroups within the Cheilostomata. We then consider the validity of our observations as adaptive trends. Patterns of ecological dominance among living and fossil species indicate that different patterns of zooid formation are important determinants of success of colonies as reflected by their abundance, competitive ability, survivorship, and recovery from injury or predation. The consistency of the long-term trends and evidence for the existence of ecological mechanisms in fossil assemblages suggest a major evolutionary role for biotic interactions.