Deciphering the influence of evolutionary legacy and functional constraints on the patella: an example in modern rhinoceroses amongst perissodactyls

In mammals, the patella is the biggest sesamoid bone of the skeleton and is of crucial importance in posture and locomotion, ensuring the role of a pulley for leg extensors while protecting and stabilizing the knee joint. Despite its central biomechanical role, the relation between the shape of the patella and functional factors, such as body mass or locomotor habit, in the light of evolutionary legacy are poorly known. Here, we propose a morphofunctional investigation of the shape variation of the patella among modern rhinoceroses and more generally among perissodactyls, this order of ungulates displaying a broad range of body plan, body mass and locomotor habits, to understand how the shape of this sesamoid bone varies between species and relatively to these functional factors. Our investigation, relying on three dimensional geometric morphometrics and comparative analyses, reveals that, within Rhinocerotidae and between the three perissodactyl families, the shape of the patella strongly follows the phylogenetic affinities rather than variations in body mass. The patellar shape is more conservative than initially expected both within and between rhinoceroses, equids and tapirs. The development of a medial angle, engendering a strong mediolateral asymmetry of the patella, appears convergent in rhinoceroses and equids, while tapirs retain a symmetric bone close to the plesiomorphic condition of the order. This asymmetric patella is likely associated with the presence of a "knee locking" mechanism in both equids and rhinos. The emergence of this condition may be related to a shared locomotor habit (transverse gallop) in both groups. Our investigation underlines unexcepted evolutionary constraints on the shape of a sesamoid bone usually considered as mostly driven by functional factors.