Abstract
Bryozoan colonies often contain many generations of zooids.
This sets up an evolutionary tension between the zooid and the colony levels, because evolutionary change may accumulate among zooids as the colony they are a part of grows asymmetrically.
This within-colony evolution may thwart evolution that occurs among colonies (caused by differential sexual reproduction among colonies.)
In order for bryozoans to be evolutionary successful, this tension between levels of organization must be resolved somehow.
Here I present three strategies that bryozoans have for resolving this tension in favor of the colonial level of evolution.
Big colonies, such as the coral-dwelling cheilostome genus Stylopoma, neutralizes evolution within its colonies by eliminating the heritability of phenotypes between clonal zooids.
This results in no evolutionary accumulation of phenotypes even across many generations of zooids in colonies that grow in response to intense ecological interactions.
Wilbertopora, a Cretaceous cheilostome, and the first genus to evolve avicularia, takes a different strategy. The phenotypic traits of zooids of Wilbertopora are highly heritable-similar to phenotypic traits of solitary animals like humans, cows, and fruit flies. Nevertheless, the variation among a generation of zooids in the multiserial Wilbertopora is structured spatially such that adjacent zooids are sufficiently different from each other to neutralize the accumulation of evolutionary change due to differential growth of the colony. We propose that mutliserial growth and the zone of autogenetic change are both colony-level adaptations that act to neutralize zooid-level evolution by maximizing variability among zooids in all spatial directions. A third strategy is taken by bryozoans that have colonies formed by uniserial growth. These small weedy colonies have low numbers of generations and so never experience a high enough growth rate to permit selection differentials to arise within their colonies. For these small uniserial species, their low growth rates (in terms of numbers of generations) shields them from the evolutionary tension between the zooid-level and the colony-level.
Bryozoan growth forms are therefore evolutionary as well as ecological strategies.