Comparative morphology of the humerus in forward-burrowing frogs

Burrowing is one of the many locomotor modes of frogs (order Anura) and is found within many clades. Burrowing is generally categorized into two groups: forward-burrowing and backward-burrowing. While forward-burrowing is more rare than backward-burrowing, we show that it has evolved independently at least eight times across anurans and is correlated with distinct features of the external and internal anatomy. The shape of the humerus is especially important for forward-burrowing, as many species use their forelimbs for digging. Using X-ray computed tomography data, we characterize shape variation in the humerus, including three-dimensional (3D) morphometrics, assess the morphology of muscles related to this variation in the humerus, and discuss the mechanical and evolutionary consequences of our results. We show that the humeri of most forward-burrowing frogs are morphologically distinct from those of non-forward-burrowers, including features such as a curved and thick diaphysis, the presence of a pronounced ventral crest, and relatively large epicondyles and humeral head. Our findings also suggest that pectoral muscle anatomy differs substantially among burrowing modes in frogs. This work provides a framework for predicting locomotor modes in taxa for which the natural history is poorly known as well as extinct taxa.