Determining the role that ecological and developmental constraints play in controlling disparity: examples from the crinoid and blastozoan fossil record

It is widely believed that morphological constraints are responsible for the observed pattern of decreasing major morphological innovation in both the Metazoa and Metaphytes over geological time. This is readily seen as the decreasing trend of origination of higher taxa: phyla, classes, and orders. Currently, there are two competing evolutionary hypotheses that have been proposed to explain this phenomenon: (1) the empty ecospace hypothesis and (2) the developmental constraint hypothesis. To distinguish between hypotheses 1 and 2, the change of morphological innovation before and after several mass extinction events was measured in the Crinoidea and Blastozoa. Mass extinction intervals provided a means in which to remove ecospace limiting constraints and allow the developmental constraint hypothesis to be thoroughly tested. Within the Crinoidea, disparity was measured before and after three mass extinctions. Within the Blastozoa, disparity was measured before and after two mass extinctions. For each taxon, three suites of characters were analyzed: ecological, nonecological, or developmental and a combination of the two previous suites plus 50 additional characters. Four different measures of disparity were used to analyze each character suite. In the majority of the cases investigated, disparity rebounds to comparable levels or in some cases higher levels in both the Crinoidea and Blastozoa. The results indicate that developmental constraints are not responsible for the decrease in disparity throughout the geologic range of the taxa. The more likely scenario is that increasingly structured ecological guilds have made it much more difficult to allow large increases in disparity.