"Untying the knot": The primitive orofacial muscle architecture in the gorilla (Gorilla gorilla) as a key to the evolution of hominin facial movement.

The anatomical underpinnings of primate facial expressions are essential to exploring their evolution. Traditionally, it has been accepted that the primate face exhibits a "scala natura" morphocline, ranging from primitive to derived characteristics. At the primitive end, the face consists of undifferentiated muscular sheets, while at the derived end there is greater complexity with more muscles and insertion points. Among these, the role of the human modiolus ("knoten" in German) has been emphasized. Recent studies have challenged this view by revealing significant complexity in the faces of several non-human primates, thereby rejecting the linear notion of facial evolution. However, our knowledge of the facial architecture in gorillas, the second closest living relatives to modern humans, remains a significant gap in the literature. Here, we present new findings based on dissection and histological analysis of one gorilla craniofacial specimen, alongside 30 human hemifaces. Our results indicate that while the number and overall arrangement of facial muscles in the gorilla are comparable to those of chimpanzees and modern humans, several orofacial features distinguish the gorilla's anatomy from that of hominins. Among these are the absence of a modiolus, the continuity of muscular fibers over the region of the mouth corner, the flat (uncurving) sheet of the orbicularis oris muscle, and the insertion of direct labial tractors both anterior and posterior to it. Collectively, the anatomical characteristics observed in the gorilla suggest that the complex anatomy of the hominin face should be considered synapomorphic (shared-derived) within the Pan-Homo clade.