The constrictor dorsalis musculature and basipterygoid articulation in Sphenodon

The constrictor internus dorsalis (CID) trigeminal muscles in vertebrates lie between the braincase and the palatoquadrate bar, and in reptiles they are believed to function in the relative movements between braincase and maxillary segment known as kinesis. In amniote taxa, the presence of a synovial basipterygoid articulation (BPA) correlates with presence of the CID. Previous descriptions of the CID in the tuatara Sphenodon (Rhynchocephalia) are inconsistent regarding presence, size and direction, particularly of the m. protractor pterygoidei. The Sphenodon skull is reported to be akinetic. The CID was investigated in Sphenodon by examination of sectioned embryonic and hatchling material, and by dissection of fixed specimens. Osteological correlates of CID muscles and morphology of the BPA were examined on disarticulated skulls and on CT scan images. The vectors of action of these muscles in relation to the BPA were projected onto CT images. Mm. levator bulbi and levator pterygoidei are found to be similar to most previous descriptions, but m. protractor pterygoidei was found in a different position, lying entirely medial to the palatoquadrate bar. The insertions of mm. levator pterygoidei and protractor pterygoidei are visible on the disarticulated pterygoid bone. The BPA is mobile only by rotation around the horizontal axis of the joints themselves; metakinesis is not possible in the Sphenodon skull. M. protractor pterygoidei appears to either resist or recognize lateral displacement of the BPA. M. levator ptergyoidei is placed to resist dorsal displacement of the braincase at the BPA, or torsion of the braincase around its longitudinal axis. The BPA appears to be a means to direct compressive stress via the base of the braincase and occipital condyle to the cervical spine in Sphenodon, and probably in its direct ancestors. Metakinesis may never have been a feature of the lepidosaur skull.