Journeys Through Shape and Time: Palaeobiology of Cenozoic New Zealand Spissatella and Eucrassatella (Bivalvia, Crassatellidae)
A novel, highly-integrated approach combining morphometric, stratocladistic and sclerochronological methods has been applied to two genera of New Zealand Cenozoic crassatellid bivalve (Family Crassatellidae): Spissatella Finlay, 1926 and Eucrassatella Iredale, 1924. This study builds on previous work on Spissatella that demonstrated their amenability to shape analysis and provided a foundation for evolutionary studies of the group. The taxonomy of these crassatellids has been in need of revision; a number of changes to generic placement having been proposed in recent publications without redescription. These bivalves are character-depauperate and known only from fossil material within New Zealand, making them challenging subjects for the phylogenetic analysis that would, ideally, inform taxonomic revision. Geometric morphometric methods have been used to characterise the morphological variation of the study group in terms of shape. Landmarks/semilandmarks that capture internal hard-part morphology and external shell shape, have been compared with internal landmarks only, outline shape semilandmarks only, and outline shape Fourier transform methods, and are shown to best combine comprehensive coverage of total shell form with high correct reassignment of individuals to taxa in multidimensional morphospace. Procrustes-superimposed landmark/semilandmark configurations have been ordinated using Principal Components Analysis (PCA), and PCA plots have been used to compare the shape variation of each species. The independance in morphospace of Spissatella n. sp. C from S. trailli and S. clifdenensis has been established. Covariation of internal morphology and shell-shape has been interpreted as supporting the interdependance of shell and body/mantle proposed by Stasek (1963). PCA scores have been combined with traditional morphological characters and stratigraphic data to produce a phylogenetic tree using stratocladistics, a form of parsimony-based analysis which seeks to minimise combined morphological and stratigraphic debt. This technique also assesses the placement of taxa in ancestral positions on internal nodes of the tree. Combining discretised morphometric data with stratigraphic and morphological data in a single analysis has been shown to produce a more resolved tree than analyses based only on continuous morphometric data. The new analyses demonstrate paraphyly of both Eucrassatella and Spissatella as previously recognised. A taxonomic revision of the studied taxa has been undertaken, incorporating information from both morphometric and phylogenetic studies. Spissatella subobesa and S. maudensis are referred to Eucrassatella. Spissatella discrepans is synonymised with S. acculta. Triplicitella n. gen. and S.maxwelli n. sp. are described. Oxygen isotope analysis has been employed to show that shell-banding in these species is, on average, likely to have been laid down annually. Using this information, the longitudinal dataset of outlines from Crampton & Maxwell (2000) has been recalibrated to use chronological age rather than size to compare shape across taxa, and investigate heterochrony in twelve pairs of species representing either ancestor-descendant, sister-group or lineage-segment relationships. All of the heterochronic processes sensu Gould (1977), namely progenesis, neoteny, acceleration and hypermorphosis, as well as proportioned dwarfism and proportioned gigantism, are identified as having affected evolution within this clade.