Biogeography a key influence on distal forelimb variation in horses through the Cenozoic.

Locomotion in terrestrial tetrapods is reliant on interactions between distal limb bones (e.g. metapodials and phalanges). The metapodial-phalangeal joint in horse (Equidae) limbs is highly specialized, facilitating vital functions (shock absorption; elastic recoil). While joint shape has changed throughout horse evolution, potential drivers of these modifications have not been quantitatively assessed. Here, I examine the morphology of the forelimb metacarpophalangeal (MCP) joint of horses and their extinct kin (palaeotheres) using geometric morphometrics and disparity analyses, within a phylogenetic context. I also develop a novel alignment protocol that explores the magnitude of shape change through time, correlated against body mass and diet. MCP shape was poorly correlated with mass or diet proxies, although significant temporal correlations were detected at 0-1 Myr intervals. A clear division was recovered between New and Old World hipparionin MCP morphologies. Significant changes in MCP disparity and high rates of shape divergence were observed during the Great American Biotic Interchange, with the MCP joint becoming broad and robust in two separate monodactyl lineages, possibly exhibiting novel locomotor behaviour. This large-scale study of MCP joint shape demonstrates the apparent capacity for horses to rapidly change their distal limb morphology to overcome discrete locomotor challenges in new habitats.