Full frontal hominins
November 5, 2009Posted by on
One of the most visually striking differences between modern humans and other hominins is the shape of the forehead. The frontal bone of the forehead serves two primary functions: it houses the frontal lobes of the brain in the anterior cranial fossa and also forms the orbital roof. When the orbits are positioned anterior to the frontal lobes, a supraorbital torus or brow ridge, forms in order to bridge the gap. This is particularly the case in archaic members of the genus Homo, whose brain cases are positioned well behind their faces.
The incredible brow ridges of Homo erectusis perhaps this species most salient physical feature. They possess a flattened forehead with a bar-like brow ridge over the eye sockets. The supraorbital torus is continuous and thickened laterally, which in turn is associated with a pinching of the orbital breadth behind the eye sockets, known as postorbital constriction. In H. erectus, the supraorbital torus is separated from the frontal squama by a depression called the posttoral sulcus. While most Erectines conform to this general bauplan, there is a lot of regional variation in the exact form of the torus.
Neandertals are characterised by their long, large, low and wide skull. They have a double-arched browridge above the orbits, which angles backward on the sides of the face. It is depressed along the middle by the presence of a supraglabellar fossa. Compared to H. erectus, Neandertals have a more vertical and rounded forehead, with a less pronounced supraorbital torus.
Modern humans have a vertical forehead, due to in no small part to the expansion of the front part of the brain. Unlike in other hominins, the frontal lobes sit directly above the orbits, negating the need for a supraorbital torus. Instead, we tend to have relatively lightly developed superciliary arches. In present day populations, large supraorbitals are generally seen in individuals that have both robust and narrow skulls. Supraorbital ridges can also occur in cases of neurodevelopmental disorders, such as microcephaly, in which case normal orbital size is combined with smaller cerebral size. The presence of a supraorbital torus in the hominin Homo floresiensis was one of the traits that some researchers used to suggest that these dwarf humans were in fact microcephalic Homo sapiens.
Modern adult humans have the most flexed basicranium of any mammal. This is due largely to us having a more vertically oriented sphenoid bone. A more flexed cranial base repositions the face directly below the anterior cranial fossa, while a more extended cranial base results in greater facial prognathism. In turn, the combination of an extended cranial base and facial forwardness influences the development of the supraorbital region. Early modern human skulls, such as Skhūl V and Dar es-Soltan, have prominent brow ridges. The development of large supraorbitals in these specimens results from greater cranial base angulation. In this regard, the development of the supraorbital region in some early modern humans does not result from neuro-orbital disjunction like in archaic humans, but primarily because of their more extended cranial base.
While much has been written about the non-metric variation of the frontal in hominins, there is little in the way of metric analyses, due to the bone’s lack of cranial landmarks. Sheela Athreya recently carried out a quantitative study of the frontal bones of various Pleistocene hominins. She collected outlines along the sagittal and parasagittal planes of the bone. Based on her analyses, specimens were classified as either Early Pleistocene, Homo erectus, Middle Pleistocene, Neandertal or anatomically modern Homo sapiens.
The highest classification accuracy was along the midsagittal plane, with a success rate of a mere 68%. In other words, using this technique almost one-third of specimens were misclassified. A well-seasoned palaeoanthropologist would have a much higher success rate using only non-metric traits. The key to identifying which species a particular frontal bone comes from involves looking at the totality of features along the entire length of the torus and surrounding bone. It is likely that if each of the curves were combined in a multivariate analysis they would have yielded a much higher classificatory success rate. Linear measurements along a curve only capture two dimensions of the frontal form, thereby losing a lot of information contained in the third dimension. A better approach would be to digitise a three-dimensional dense point cloud along the entire bone and to analyse the region using geometric morphometrics. However, such equipment is expensive and not available in most anthropology departments.
Perhaps the most important outcome of this study was that it quantitatively confirmed some of the general characteristics of the frontal form of Homo, that have been previously described qualitatively. These include the fact that most of the variation in the frontal bone between Pleistocene groups is along the midsagittal plane. The study additionally found Homo erectus to differ from all other groups in the projection of the glabellar region. Finally, it identified modern humans as differing from all other groups in the curvature of the forehead, as well as the prominence of the lateral supraorbital torus. This confirms what many palaeoanthropologists have been saying for a long time – the lack of a supraorbital torus in modern humans is a uniquely derived feature.
Athreya, S. A comparative study of frontal bone morphology among Pleistocene hominin fossil groups, J Hum Evol (2009), doi:10.1016/j.jhevol.2009.09.003.
Lahr, MM. The Evolution of Modern Human Diversity : A Study on Cranial Variation . Cambridge; New York : Cambridge University Press, 1996.
Lieberman, Daniel E, Osbjorn M Pearson, and Kenneth M Mowbray. “Basicranial Influence on Overall Cranial Shape.” Journal of Human Evolution 38 (2000): doi:10.1006/jhev.1999.0335.
Martin RD, MacLarnon AM, Phillips JL, Dussebieux L, Williams PR, Dobyns WV. 2006a. Comment on ‘‘The brain of LB1, Homo floresiensis.’’ Science 312:999b.
Trinkaus. Modern Human versus Neandertal Evolutionary Distinctiveness. Current Anthropology (2006) vol. 47 (4) pp. 597-620.
Trinkaus. European early modern humans and the fate of the Neandertals. Proceedings of the National Academy of Sciences (2007) 104 (18) pp. 7367-7372.
Above photos modified from originals by missmareck and arnybo under creative commons license.
Image of lateral dissected skull by dollinjune14, via deviantART (modified from original).