Approximate evolutionary stasis for bivalve morphology over millions of years: a multivariate, multilineage study
Authors: Stanley Steven M. , Yang Xiangning 
Published: July 2016

We have calculated rates of evolution for 19 lineages of Neogene bivalves over time intervals ranging from 1 ma to 17 ma. Our morphometric comparisons are based on 24 variables, for which we have made more than 43,000 individual measurements normalized for shell size. We initially assessed evolutionary changes in shape for 19 early Pliocene (4 ma old) species of bivalves by comparing these forms to their closest living relatives, which in 12 cases have traditonally been assigned to the same species. To make our study unbiased and comprehensive, we included all species that met certain predetermined guidelines and that offered suitable fossil material for measurement. We compared early Pliocene and Recent populations using (1) all 24 variables treated separately, (2) multivariate distance (Mahalanobis' D), based on the full set of variables, and (3) eigenshapes for shell outlines. For these comparisons, we used as a yardstick the same measures of morphologic distance applied to pairs of geographically separated Recent populations that belong to eight of the living species to which the fossil populations were compared. As it turns out, with minor exceptions, the distribution of morphologic distances between 4 ma old and Recent populations resembled the distribution of distances between conspecific Recent populations.

We calculated net rates of evolution separating pairs of populations that belong to single lineages. For all intervals of time, the distribution of differences between population means for individual variables is remarkably similar to a comparable distribution representing the comparison of pairs of conspecific Recent populations from separate geographic regions. Because morphologic differences between populations do not vary greatly with evolutionary time, measured “rates” of evolution, on the average, decrease with interval of measurement. Because these differences resemble intraspecific variability, however, the rates do not represent significant evolution. Evolution has followed a weak zigzag course, yielding only trivial net trends.

The weak and reversible “trends” that we measured yield net rates averaging less than 10 millidarwins, which is much lower than most rates previously reported for marine invertebrates (average ~200 millidarwins for a 1 ma interval and ~60 millidarwins for a 10 ma interval). We attribute this disparity (1) to the fact that most previously published rates have been calculated when a significant amount of evolution was recognized in advance (often for a poorly documented lineage) and (2) to the fact that most measured variables have represented nothing more than some measure of body size. We conclude that shape, as opposed to size, has been highly stable in bivalve evolution over millions of years and 106–107generations. We conclude that to characterize rates or evolution for any group of organisms, one must employ a large, unbiased sample of measurements for numerous well-documented lineages, and one must segregate data depicting shape from data depicting size.