The lepidosaurian skull has long been of interest to functional morphologists and evolutionary biologists. Patterns of bone loss and gain, particularly in relation to bars and fenestrae, have led to a variety of hypotheses concerning skull use and kinesis. Of these, one of the most enduring relates to the absence of the lower temporal bar in squamates and the acquisition of streptostyly. We performed a series of computer modeling studies on the skull of Uromastyx hardwickii, an akinetic herbivorous lizard. Multibody dynamic analysis (MDA) was conducted to predict the forces acting on the skull, and the results were transferred to a finite element analysis (FEA) to estimate the pattern of stress distribution. In the FEA, we applied the MDA result to a series of models based on the Uromastyx skull to represent different skull configurations within past and present members of the Lepidosauria. In this comparative study, we found that streptostyly can reduce the joint forces acting on the skull, but loss of the bony attachment between the quadrate and pterygoid decreases skull robusticity. Development of a lower temporal bar apparently provided additional support for an immobile quadrate that could become highly stressed during forceful biting.biomechanics ͉ Lepidosauria ͉ lower temporal bar ͉ streptostyly L epidosauria is composed of 2 subgroups: Rhynchocephalia (Sphenodon and its extinct relatives) and Squamata (lizards, snakes, and amphisbaenians). Several stem taxa (as part of the more inclusive Lepidosauromorpha) are known from fossil deposits of Permian to Jurassic age (Ϸ250 million to 164 million years ago; ref. 1). In relation to lepidosaurian evolution, paleontologists, comparative anatomists, and functional morphologists have focused particularly on changes in cranial morphology, most notably with respect to the evolution of streptostyly (anteroposterior quadrate movement) and cranial kinesis. A longstanding hypothesis held that the ancestral lepidosaur possessed a fully diapsid skull (upper and lower temporal openings) with a complete lower temporal bar (2-6). Loss of that bar supposedly ''freed'' the quadrate, resulting in streptostyly. Elements of this hypothesis persist in the literature, even though it has been demonstrated unequivocally (1, 7-9) that the ancestral lepidosauromorph, the ancestral lepidosaur, and the ancestral rhynchocephalian (7, 8) all had a fixed quadrate (strong quadratepterygoid and squamosal-quadrate joints) but no lower temporal bar (Fig. 1). Indeed, the same seems to have been true for the last common ancestor of lepidosaurs and archosaurs ( Fig. 1) (9). A complete lower temporal bar was developed de novo one or more times within Rhynchocephalia (7,8). The ancestral squamate, on the other hand, inherited a skull without a lower temporal bar but underwent a reduction of the palatoquadrate and its derivatives, as well as a reduction of the connections between those derivatives (quadrate and epipterygoid) and the rest of the skull (quadrate-pterygoid, quadrate-squamosal, and epip...