The Phylogenetic Position of the Tuatara, Sphenodon (Sphenodontida, Amniota), as Indicated by Cladistic Analysis of the Ultrastructure of Spermatozoa

Sphenodon has traditionally been regarded as a little changed survivor of the Permo-Triassic thecodont or eosuchian ‘stem reptiles’ but has alternatively been placed in the Lepidosauria as the plesiomorphic or even apomorphic sister-taxon of the squamates. A cladistic analysis of 16 characters from spermatozoal ultrastructure of Sphenodon and other amniotes unequivocally confirms its exceedingly primitive status. The analysis suggests that monotremes are the sister-group of birds; squamates form the sister-group of a bird + monotreme clade while the three sister-groups successively below the bird + monotreme + squa- mate assemblage are the caiman, the tuatara and the outgroup (turtles). The monotreme + bird couplet, supports the concept of the Haemothermia, but can only be regarded heuristically. The usual concept of mammals as a synapsid-derived outgroup of all other extant amniotes is not substantiated spermatologically. All cladistic analyses made, and a separate consideration of apomorphies, indicate that Sphenodon is spermatologically the most primitive amniote, excepting the Chelonia. It is advanced (apomorphic) for the amniotes in only two of the 16 spermatozoal characters considered. A close, sister-group relationship of Sphenodon with squamates is not endorsed.