Forelimb posture has been a controversial aspect of reconstructing locomotor behaviour in extinct quadrupedal tetrapods. This is partly owing to the qualitative and subjective nature of typical methods, which focus on bony articulations that are often ambiguous and unvalidated postural indicators. Here we outline a new, quantitatively based forelimb posture index that is applicable to a majority of extant tetrapods. By determining the degree of elbow joint adduction/abduction mobility in several tetrapods, the carpal flexor muscles were determined to also play a role as elbow adductors. Such adduction may play a major role during the stance phase in sprawling postures. This role is different from those of upright/sagittal and sloth-like creeping postures, which, respectively, depend more on elbow extensors and flexors. Our measurements of elbow muscle moment arms in 318 extant tetrapod skeletons (Lissamphibia, Synapsida and Reptilia: 33 major clades and 263 genera) revealed that sprawling, sagittal and creeping tetrapods, respectively, emphasize elbow adductor, extensor and flexor muscles. Furthermore, scansorial and non-scansorial taxa, respectively, emphasize flexors and extensors. Thus, forelimb postures of extinct tetrapods can be qualitatively classified based on our quantitative index. Using this method, we find that Triceratops (Ceratopsidae), Anhanguera (Pterosauria) and desmostylian mammals are categorized as upright/sagittally locomoting taxa.

Some tetrapods hang upside down from tree branches when moving horizontally. The ability to walk in quadrupedal suspension has been acquired independently in at least 14 mammalian lineages. During the stance (supportive) phase of quadrupedal suspension, the elbow joint flexor muscles (not the extensors as in upright vertebrates moving overground) are expected to contract to maintain the flexed limb posture. Therefore muscular control in inverted, suspended quadrupeds may require changes of muscle control, and even morphologies, to conditions opposite to those in upright animals. However, the relationships between musculoskeletal morphologies and elbow joint postures during the stance phase in suspended quadrupeds have not been investigated. Our analysis comparing postures and skeletal morphologies in Choloepus (Pilosa), Pteropus (Chiroptera), Nycticebus (Primates) and Cynocephalus (Dermoptera) revealed that the elbow joints of these animals were kept at flexed angles of 70-100°during the stance phase of quadrupedal suspension. At these joint angles the moment arms of the elbow joint flexors were roughly maximized, optimizing that component of antigravity support. Our additional measurements from various mammalian species show that suspended quadrupeds have relatively small extensor ⁄ flexor ratios in both muscle masses and maximum moment arms. Thus, in contrast to the pattern in normal terrestrial quadrupeds, suspended quadrupeds emphasize flexor over extensor muscles for body support. This condition has evolved independently multiple times, attendant with a loss or reduction of the ability to move in normal upright postures.

Reconstruction of limb posture is a challenging task in assessing functional morphology and biomechanics of extinct tetrapods, mainly because of the wide range of motions possible at each limb joint and because of our poor knowledge of the relationship between posture and musculoskeletal structure, even in the extant taxa. This is especially true for extinct mammals such as the desmostylian taxa Desmostylus and Paleoparadoxia. This study presents a procedure that how the elbow joint angles of extinct quadruped mammals can be inferred from osteological characteristics. A survey of 67 dried skeletons and 113 step cycles of 32 extant genera, representing 25 families and 13 orders, showed that the olecranon of the ulna and the shaft of the humerus were oriented approximately perpendicular to each other during the stance phase. At this angle, the major extensor muscles maximize their torque at the elbow joint. Based on this survey, I suggest that olecranon orientation can be used for inferring the elbow joint angles of quadruped mammals with prominent olecranons, regardless of taxon, body size, and locomotor guild. By estimating the elbow joint angle, it is inferred that Desmostylus would have had more upright forelimbs than Paleoparadoxia, because their elbow joint angles during the stance phase were approximately 165 degrees and 130 degrees , respectively. Difference in elbow joint angles between these two genera suggests possible differences in stance and gait of these two mammals.