Taxonomic significance of biometric characters and the consequences for classification and biostratigraphy, exemplified through moussoneliform daonellas (Daonella, Bivalvia; Triassic)

The middle Triassic bivalve genusDaonella Mojsisovics, 1874 would be an ideal guide fossil (worldwide distribution, high evolutionary rates); however, the classification of daonellas is difficult. Most daonellas lack distinct characters and have a wide range of morphologic variation. A new taxonomic attempt was necessary to get a coherentDaonella classification.

The new taxonomic concept proposed herein consists of four steps. (1) Biometric characters are defined unambiguously, based on landmarks and pseudolandmarks. (2) The taxonomic significance of biometric characters is analyzed. Several characters depend significantly on growth stage of specimens or on the lithology of fossil localities. In addition, the intercorrelations between biometric characters are studied to exclude weighting in some morphologic features. The study of the taxonomic significance of biometric characters shows, how important a careful selection of taxonomic characters is. (3) Most daonellas are not determinable by univariate or bivariate methods. The principle component analysis (PCA) is applied to detect groups of similar holomorphs in multivariate space. (4) The interpreted groups from PCA are applied to a canonical discriminant function analysis. This analysis helps in deciding whether two groups are sufficiently distinct to be separated as different species.

This new taxonomic concept was applied to moussoneliform daonellas of the middle Triassic from the Southern Alps. Three different forms were used for the study herein:Daonella moussoni (Mérian, 1853) from the Varenna Limestone (Perledo-Varenna Formation, Grigna, Italy),D. aff.moussoni (Mérian)Rieber, 1969 from the lower Meride Limestone (Monte San Giorgio, Switzerland) andD. pseudomoussoni Rieber, 1969 from the Grenzbitumenzone (Monte San Giorgio, Switzerland). The new classification shows thatD. pseudomoussoni is a separate species and distinguishable fromD. moussoni andD. aff.moussoni, which belong to the same species.

A coherent systematics ofDaonella becomes possible on the basis of newly defined taxonomic characters. Daonellas can now be used as relevant guide fossils for the middle Triassic. The presented example demonstrates a biostratigraphic correlation between the lower Meride Limestone at Monte San Giorgio and the Varenna Limestone at Grigna. The stratigraphic distribution ofD. moussoni can be assigned to thegredleri Zone.