The taxonomic status of Neanderthals lies at the center of the modern human origins debate. Proponents of the single-origin model often view this group as a distinct species with little or no contribution to the evolution of modern humans. Adherents to the regional continuity model consider Neanderthals a subspecies or population of Homo sapiens, which contributed significantly to the evolution of early modern Europeans. Paleontologists generally agree that fossil species should be equivalent to extant ones in the amount of their morphological variation. E ver since Neanderthals were first discovered in the 19th century, their specific status and contribution to modern human evolution have been debated. The current controversy centers around two contrasting models for modern human evolution (1, 2): the single-origin theory, which frequently views modern humans as a new species arising relatively recently in Africa and replacing indigenous archaic populations around the world; and the regional-continuity model, in which archaic populations contributed to the evolution of modern peoples in each geographic area. Directly tied to these two models is the status of Neanderthals as a distinct species or as a subspecies of Homo sapiens at least partially ancestral to modern Europeans. The morphological differences between Neanderthals and modern humans are well documented; however, their magnitude can be meaningfully assessed only within the broader context of primate species variability. Here we used 12 extant catarrhine species as models of 3D craniofacial shape variation. Our sample comprised 1,089 specimens, including modern humans, African apes, and eight species of Old World monkeys. The morphological distinctiveness of Neanderthals from modern humans, as represented by our data, was then compared to the differences between pairs of model taxa.Although there is consensus that species in paleontology should be equivalent to living ones in the variation that they accommodate (3, 4), there is no absolute degree of difference that defines species boundaries across taxa (5), making the choice of models critical. Recent studies have supported a specific status for Neanderthals based on analogy to chimpanzee (6) and hybridizing Sulawesi macaque (7) craniofacial morphology. These models may not be the most relevant: chimpanzees differ in their geographic distribution and ecology from Pleistocene humans, whereas the Sulawesi macaques are not representative of the full geographic and ecological range of Macaca. Modern humans are the most appropriate phylogenetic model for Neanderthals but can provide only intraspecific measures of variation. As the closest living relatives to modern humans, the African apes are often used as models for hominin species (4, 8). Yet these two genera comprise only three species, all of which are ecologically distinct from Pleistocene humans. Papionin monkeys, although more distantly related, are often considered (9-11) similar to hominins in their ecology and population structure, and their evo...