Partitioning phenotypic variation: genotypic, environmental and residual components from bryozoan skeletal morphology

Using morphometric studies of colonial (clonal) organisms such as Bryozoa grown as replicates in controlled laboratory experiments, phenotypic variation (hard part morphology) can be partitioned into its genotypic and environmental (ecophenotypic) components. The interaction between these, i.e. different genotypic responses to the same environmental change, can also be recognized. Palaeobiological studies are inherently constrained by species concepts based on morphotypes (preserved morphological phenotype). Uncertainties associated with fossil species concepts restrict the deductive resolution potential of fossil taxa in discussions of the broader biological questions of species evolution, ecology, biogeography and phylogeny. The relationship between species-level morphological variation and genetic variation in modern taxa is central to evaluating the viability of fossil morphotypes as biological species. Results from a preliminary study of three genotypes of Electra pilosa L., grown as replicate colonies in tanks comprising different microenvironmental conditions, allow for direct evaluation of morphospecies concepts. Numerical analyses (Cluster Analysis, Principal Component Analysis and Two-way ANOVA) of six morphometric characters demonstrated a strong Genotype control over zooid morphology and limited environmental (Tank) effects with minimal environmental differences among tanks. No significant Genotype by Tank (=environment) interactions were found for any characters. These results demonstrate that it is possible to quantify the extent to which a given trait is plastically expressed in different environments. The strong degree of correlation between morphology and genetics for this species is encouraging for the use of morphotypes as proxies for biological species.