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Testing evolutionary constraint hypotheses with early Paleozoic gastropods

The evolution of higher taxa among early Paleozoic gastropods is similar to that among early metazoans as a whole, as higher taxa diversified rapidly and early. There are two issues pertinent to this pattern. First, were greater morphologic changes concentrated in the early phases of evolution? Second, does the pattern better fit models of increasing phylogenetic constraints or increasing ecologic restrictions? This paper presents a phylogeny-based method designed to test whether amounts of morphologic evolution decreased over time. It also explores whether the data better fits models of increasing phylogenetic (i.e., developmental or genetic) constraint or increasing ecologic restriction. Two metrics of morphologic separation (i.e., the morphologic difference between sister-species) are used: (1) Euclidean distance in morphospace and (2) transition magnitude. The latter metric is calculated by a multivariate analysis of sister-species contrasts, which determines both types and magnitudes of morphologic transitions. The advantage of using transition magnitudes is that it balances the effects of transitions that either affect more morphometric characters or occur more frequently. Both metrics indicate that larger morphologic separations between sister-species were concentrated early in gastropod evolution. Among gastropods, gross shell morphology often reflects basic trophic strategy and function whereas basic internal anatomy does not. Transition magnitudes can be broken down into transitions associated with differences in basic trophic strategies and shell functional biology (“external”), and those associated with differences in basic internal anatomy (“internal”). Internal transition magnitudes show a highly significant decrease over time (p < 10–04) whereas external transition magnitudes show a much less significant decrease over time (p < 0.10) and no significant decrease after the earliest Ordovician (p ≅ 0.50). The results therefore suggest that increasing phylogenetic constraints played a greater role in the early evolution of gastropods than did increasing ecologic ones.