During locomotion over land, vertebrates' limb bones are exposed to loads. Like most biological structures, limb bones have a capacity to withstand greater loads than they usually experience, termed a safety factor (SF). How diverse are limb-bone SFs, and what factors correlate with such variation? We have examined these questions from two perspectives. First, we evaluated locomotor SF for the femur in diverse lineages, including salamanders, frogs, turtles, lizards, crocodilians, and marsupials (opossums). Comparisons with values for hind-limb elements in running birds and eutherian mammals indicate phylogenetic diversity in limb-bone SF. A high SF (∼7) is primitive for tetrapods, but low magnitudes of load and elevated strength of bones contribute to different degrees across lineages; moreover, birds and eutherians appear to have evolved lower SFs independently. Second, we tested the hypothesis that SFs would be similar across limb bones within a taxon by comparing data from the humerus and femur of alligators. Both in bending and in torsion, we found a higher SF for the humerus than for the femur. Such a "mixed chain" of different SFs across elements has been predicted if bones have differing variabilities in load, different costs to maintain, or high SF values in general. Although variability in load is similar for the humerus and femur, a high SF may be less costly for the humerus because it is smaller than the femur. The high SFs of alligators also might facilitate differences in SF among their limb bones. Beyond these specific findings, however, a more general implication of our results is that evaluations of the diversity of limb-bone SFs can provide important perspective to direct future research. In particular, more complete understanding of variation in SF could provide insight into factors that promoted the evolutionary radiation of terrestrial locomotor function in vertebrates.

SUMMARYSalamanders are often used as representatives of the basal tetrapod body plan in functional studies, but little is known about the loads experienced by their limb bones during locomotion. Although salamanders' slow walking speeds might lead to low locomotor forces and limb bone stresses similar to those of non-avian reptiles, their highly sprawled posture combined with relatively small limb bones could produce elevated limb bone stresses closer to those of avian and mammalian species. This study evaluates the loads on the femur of the tiger salamander (Ambystoma tigrinum) during terrestrial locomotion using threedimensional measurements of the ground reaction force (GRF) and hindlimb kinematics, as well as anatomical measurements of the femur and hindlimb muscles. At peak stress (29.8±2.0% stance), the net GRF magnitude averaged 0.42body weights and was directed nearly vertically for the middle 20-40% of the contact interval, essentially perpendicular to the femur. Although torsional shear stresses were significant (4.1±0.3MPa), bending stresses experienced by the femur were low compared with other vertebrate lineages (tensile: 14.9±0.8MPa; compressive: -18.9±1.0MPa), and mechanical property tests indicated yield strengths that were fairly standard for tetrapods (157.1±3.7MPa). Femoral bending safety factors (10.5) were considerably higher than values typical for birds and mammals, and closer to the elevated values calculated for reptilian species. These results suggest that high limb bone safety factors may have an ancient evolutionary history, though the underlying cause of high safety factors (e.g. low limb bone loads, high bone strength or a combination of the two) may vary among lineages.

SUMMARYSkeletal elements are usually able to withstand several times their usual load before they yield, and this ratio is known as the bone's safety factor. Limited studies on amphibians and non-avian reptiles have shown that they have much higher limb bone safety factors than birds and mammals. It has been hypothesized that this difference is related to the difference in posture between upright birds and mammals and sprawling ectotherms; however, limb bone loading data from a wider range of sprawling species are needed in order to determine whether the higher safety factors seen in amphibians and non-avian reptiles are ancestral or derived conditions. Tegus (family Teiidae) are an ideal lineage with which to expand sampling of limb bone loading mechanics for sprawling taxa, particularly for lizards, because they are from a different clade than previously sampled iguanas and exhibit different foraging and locomotor habits (actively foraging carnivore versus burst-activity herbivore). We evaluated the mechanics of locomotor loading for the femur of the Argentine black and white tegu (Tupinambus merianae) using threedimensional measurements of the ground reaction force and hindlimb kinematics, in vivo bone strains and femoral mechanical properties. Peak bending stresses experienced by the femur were low (tensile: 10.4±1.1MPa; compressive: -17.4±0.9MPa) and comparable to those in other reptiles, with moderate shear stresses and strains also present. Analyses of peak femoral stresses and strains led to estimated safety factor ranges of 8.8-18.6 in bending and 7.8-17.5 in torsion, both substantially higher than typical for birds and mammals but similar to other sprawling tetrapods. These results broaden the range of reptilian and amphibian taxa in which high femoral safety factors have been evaluated and further indicate a trend for the independent evolution of lower limb bone safety factors in endothermic taxa.