Population genetics of four species of Ordovician bryozoans: stereology and jackknifed analysis of variance

Abstract

Collections of four Ordovician bryozoan species from habitats of differing levels of taxonomic diversity were examined in order to assess the effect of environmental conditions on intraspecific and interspecific morphologic patterns. Up to 20 morphometric characteristics were measured for each species from populations in both high-diversity and low-diversity settings. Character means and the levels of character variability differ between populations in different diversity zones. Across species, mean character values differ significantly in 41 percent of the total comparisons, permitting intraspecific and interspecific multivariate discrimination of populations from the two diversity zones.The partitioning of clonal morphologic variability into within-colony (nonheritable) and among-colony (potentially heritable) components reveals that habitat conditions favoring high diversities appear to promote a greater proportion of potentially heritable genetic variation both within- and among-species than do low-diversity settings wherein a larger percentage of variation is ecophenotypic, or nonheritable, plasticity. Although the actual percentages vary from species to species, this pattern is obtained for representatives of four different trepostome bryozoan families and therefore is robust. Variability of this sort, if unrecognized, could lead to taxonomic oversplitting along ecologic gradients. These local, facultative adjustments to environmental conditions may be indicative of processes involved in the reactions of populations to selective pressures.Analyses such as this, coupled with examinations of character covariances, may permit the recognition not only of environmentally-induced morphologic plasticity and the proportions of potentially heritable and nonheritable variability, but also provide evidence for intraspecific changes in the rates and timing of developmental events (i.e., across-population or facultative heterochrony).