Reconstructing the behavioral shifts that drove hominin evolution requires knowledge of the timing, magnitude, and direction of anatomical changes over the past ∼6-7 million years. These reconstructions depend on assumptions regarding the morphotype of the Homo-Pan last common ancestor (LCA). However, there is little consensus for the LCA, with proposed models ranging from African ape to orangutan or generalized Miocene ape-like. The ancestral state of the shoulder is of particular interest because it is functionally associated with important behavioral shifts in hominins, such as reduced arboreality, high-speed throwing, and tool use. However, previous morphometric analyses of both living and fossil taxa have yielded contradictory results. Here, we generated a 3D morphospace of ape and human scapular shape to plot evolutionary trajectories, predict ancestral morphologies, and directly test alternative evolutionary hypotheses using the hominin fossil evidence. We show that the most parsimonious model for the evolution of hominin shoulder shape starts with an African apelike ancestral state. We propose that the shoulder evolved gradually along a single morphocline, achieving modern human-like configuration and function within the genus Homo. These data are consistent with a slow, progressive loss of arboreality and increased tool use throughout human evolution.geometric morphometrics | developmental simulation | phylomorphospace | scapula | rotator cuff T he human shoulder exhibits a unique combination of traits for a primate, a fact that has complicated previous attempts to reconstruct both its functional and evolutionary history. Notably, humans are most closely related to knuckle-walking/suspensory chimpanzees and bonobos (Pan or panins) (1, 2), yet morphometric analyses suggest that our shoulders are most similar in shape to that of the highly arboreal, quadrumanous orangutan (Pongo) (3-5). The hominin fossil record is similarly complicated. Scapular remains attributed to Australopithecus afarensis are described as similar to Gorilla (6-8) whereas the more recent Australopithecus sediba (MH2) displays morphometric affinities to both African apes (gorillas, chimpanzees, and bonobos) and Pongo (9). This mix of character states raises the question of whether modern human morphology reflects evolution from a more derived African ape morphology or retention of primitive traits from an earlier ape ancestor.A critical piece of evidence for solving this puzzle is the morphotype of the hominin-panin last common ancestor (LCA) (∼6-7 Mya). Although the fossil record near the hypothesized divergence time with Pan is the most direct means of addressing what the anatomy of the LCA shoulder was like, both hominin and African ape fossils from this time period are rare, fragmentary, and poorly understood (10, 11). Despite these difficulties, competing hypotheses about the LCA make explicit and mutually exclusive predictions of the direction, magnitude, and ordering of character transformations that we should observe in the fo...