ABSTRACT
Among fossorial mammals, forelimbs are major digging apparatuses for dwelling, sheltering and foraging underground. Forelimb-diggers have independently evolved in many lineages of mammals; thus, the method of digging with forelimbs varies by taxon. Therefore, the reconstruction of digging behaviours in extinct animals leads us to understand the evolutionary process of fossorial adaptation in each lineage. However, no morphological index was found to reconstruct if, or how, extinct taxa dug with forelimbs. In this study, we used the shoulder and elbow muscle moment arms in relation to the out-force lever on the manus as indices of the efficiency of motions. The mechanical advantage of two shoulder motions (medial rotation and retraction) and three elbow motions (extension, flexion and adduction) was measured in 381 extant mammal specimens representing 332 species, 279 genera, 103 families and 24 orders. Assuming that both forelimb-digging and -paddling in water require relatively high-output moment arm efficiency, the studied taxa were categorised into four groups based on the presence or absence of forelimb-digging and -paddling abilities. We found that the efficiencies of all five muscle moment arms in the forelimb-diggers and -paddlers were higher than those of the non-diggers and non-paddlers. Furthermore, among the forelimb-diggers, the taxa that dig compact substrates or frequently burrow tend to emphasise the muscle moment arms compared to the taxa that dig loose substrates or dig less frequently. The comparison among the 53 extant forelimb-diggers revealed that the efficiency marked among the five muscle moment arms reflects the difference in digging strategy: humeral rotation diggers emphasise the shoulder medial rotator and elbow adductor, hook-and-pull diggers emphasise the shoulder retractor and elbow flexor and scratch diggers emphasise the shoulder retractor and elbow extensor. We propose that these indices will be powerful tools for reconstructing the fossorial behaviours of extinct mammals. Applying these indices to extinct taxa, Ceratogaulus, Ernanodon, Metacheiromys and Prozaedyus are capable of more efficient forelimb-digging, and each may have adopted different digging strategies.
