Phenetic discrimination of biometric simpletons: paleobiological implications of morphospecies in the lingulide brachiopod Glottidia

Abstract

The extreme morphological simplicity of lingulide brachiopod shells makes them particularly useful for investigating the species-level taxonomic resolution of the fossil record as well as the relationships between taxonomy, morphological complexity, and evolutionary rates. Lingulides have undergone little change in shell morphology and have had low taxonomic diversity since the Paleozoic. Is this pattern an evolutionary phenomenon or an artifact of the shell's simplicity? Multivariate methods were used to establish morphogroups among seven populations of four extant species of Glottidia. Six characters (three shell dimensions and three internal septa) were measured for 162 specimens from field and museum collections. All populations follow similar allometric trajectories: internal septa display positive allometry and shell dimensions display negative allometry. The allometric pattern may reflect D'Arcy Thompson's Principle of Similitude. Principal component analysis does not reveal any distinct clusters in Glottidia morphospace but suggests that some differences independent from ontogeny exist among the populations. Size-free canonical variate analysis indicates the presence of five size-invariant groups that are statistically distinct. Bootstrap-corrected error rates indicate that four specimens are enough to classify a sample correctly at α = 0.05 and eight specimens at α = 0.01. The groups are consistent with neontological classification with the exception of two populations of G. pyramidata identified by discriminant analysis as two distinct groups. The size-free morphogroups reflect geographic separation rather than ontogenetic or substrate differences among the populations.Despite the morphological simplicity of the shell, size-free multivariate analysis of Glottidia delineates groups that offer taxonomic resolution comparable with the neontological classification. The method offers a promising tool for identifying natural morphogroups on the basis of few morphological characters. Moreover, the agreement between neontological taxonomy and the morphogroups suggests that the size-free approach can be applicable for evaluating the reality of the low diversity and turnover rates observed in the fossil record of lingulide brachiopods (= Family Lingulidae). Assuming that the neontological species of Glottidia are biologically meaningful, this study shows that morphological simplicity of lingulides does not necessarily result in taxonomic underresolution. Our analysis, as well as several previous case studies, suggests that taxonomic diversity and turnover rates do not have to be dependent on the morphological complexity of preservable parts. In many cases, when rigorous quantitative methods are employed, the differences in the rates of morphological evolution may be a real evolutionary phenomenon and not artifacts of morphological complexity.

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