Fitting unanchored puzzle pieces in the skeleton: appropriate 3D scapular positions for the quadrupedal support in tetrapods.

Deducing the scapular positions of extinct tetrapod skeletons remains difficult, because the scapulae and rib cage are connected with each other not directly by skeletal joint, but by thoracic muscles. In extant non-testudine quadrupedal tetrapods, the top positions of the scapulae/suprascapulae occur at the anterior portion of the rib cage, above the vertebral column and near the median plane. The adequacy of this position was tested using three-dimensional mechanical models of Felis, Rattus and Chamaeleo that assumed stances on a forelimb on a single side and the hindlimbs. The net moment about the acetabulum generated by the gravity force and the contractive forces of the anti-gravity thoracic muscles, and the resistance of the rib to vertical compression between the downward gravity and upward lifting force from the anti-gravity thoracic muscle depend on the scapular position. The scapular position common among quadrupeds corresponds to the place at which the roll and yaw moments of the uplifted portion of the body are negligible, where the pitch moment is large enough to lift the body, and above the ribs having high strength against vertical compression. These relationships between scapular position and rib cage morphology should allow reliable reconstruction of limb postures of extinct taxa.