Monthly Archives: November 2009

One chin does not a modern human make

Chinese scientists say that a recently discovered partial jaw from Guangxi challenges the ‘out of Africa’ model of modern human origins, while lending support to the multiregional hypothesis. The 110,000 year-old mandible is described as having a chin that juts “ever so slightly outward.” These scientists assert that the presence of chin shows that there was significant gene flow between populations of modern Homo sapiens and archaic Homo.

Wu Xinzhi of the Chinese Academy of Sciences had the following to say about the find:

    ”The finding was strong evidence to prove the multiregional model, and from this evidence, it was significant to solve the academic dispute between ‘the multiregional mode’ and ‘out of Africa theory’”.

It is interesting to note Xinzhi’s use of the past simple tense to suggest that this is a closed case. Far from it! Palaeoanthropological theory has moved on from the multiregional sensu stricto versus ‘out of Africa’ sensu stricto dichotomy that predominated the discussion during the latter half of the last century. Nevertheless, the question of how much gene flow, if any, took place between modern and archaic Homo is still very much a debated issue.

At this stage you may be wondering why there has been such furore over a chinned jaw. As long ago as 1775, Johann Friedrich Blumenbach commented on the uniqueness of the modern human chin:

    In the animals there is scarcely a particular chin which can be considered as comparable to that of man: and in those men who, as is often said, seem to have something apish in their countenance, this generally resides in a deeply-retreated chin.

The distinctive modern human chin develops through the combination of bone deposition on the inferior part of the jaw and resorption around the alveolar region. In other primates the entire jaw undergoes deposition. The modern human chin is characterised as having a central keel, with hollowed out depressions (known as mental fossae) to either side, together with a protruding inferior portion. This distended mental protuberance and lateral extremities make up the mental trigone, giving the chin the appearance of an inverted T. It is the combination of all these anatomical features that make up the prototypal modern chin. However, chins show great variability, with some modern humans not having any.

This variability is also extends to earlier hominins. The Middle Pleistocene fossils from the Sima de los Huesos have been described as having chins, and even well-developed mental trigones. Among Pleistocene hominins, Neandertals appear to have the most divergent pattern from the modern configuration, universally lacking the inverted T and mental fossae. While it has been argued that the Neandertal mandibles from the Croatian site of Vindija show the development of incipient chins, this has not been borne out by later analyses.

Some of the ‘modern’ Klasies River Mouth mandibles do not have developed mental trigones, midline keel or a thickening of the inferior margin. However, the modern designation of this material is controversial with these fossils showing a mosaic of both archaic and modern features. Similarly, the modern humans from Qafzeh show variable expression of the inverted T and mental fossae, with no indication of these features in the Skhūl specimens. The 700-800,000 year-old Tighenif mandibles show a surprisingly modern configuration complete with central keel, a thickened inferior portion, and the development of a triangular protuberance. The presence of a chin in these specimens could represent a synapomorphy with modern humans.

Based on the archaeological record, it appears that modern humans left Africa some time around 100,000 years ago. Among the oldest undisputed modern human remains in China come from Zhoukoudian Cave at around 35,000 years BP. The possibly earlier fossil from Liujiang is marred with dating problems. In order for the Chinese scientists’ assertion to hold, it would require an even earlier exit from Africa or expansive gene flow between modern humans living in Africa and archaic humans in Asia; claims for which the evidence is currently lacking. Future analyses of the specimens will determine whether these chins have a truly modern form or whether the pattern is more like the non-homologous protruding inferior jaws seen in other archaic specimens. Alternatively, if these specimens end up being the result of convergent evolution it would raise questions about the functional significance of a chin. Finally, if these fossils show a pattern similar to the one seen in the Tighenif fossils it may suggest that they belong to the same clade.

References and further reading
Ahern JC (1993). The Transitional Nature of the Late Neandertal Mandibles from Vindija Cave, Croatia. M.A. thesis. Department of Anthropology, Northern Illinois University.

Blumenbach, JF (1978). The anthropological treatises of Johann Friedrich Blumenbach / translated and edited from the Latin, German, and French originals by Thomas Bendyshe. Boston : Longwood Press.

Hawks, J (2009). It came from Guangxi.

McKenna, P (2009). Chinese challenge to ‘out of Africa’ theory. New Scientist.

Rosas, A. (1995). Seventeen new mandibular specimens from the Atapuerca/Ibeas Middle Pleistocene hominids sample. J. hum. Evol. 28, 533–559.

Schwartz JH, Tattersall I (2000). The human chin revisited: what is it and who has it? J Hum Evol 38:367-409.

Schwartz JH, Tattersall I (2002) The Human Fossil Record, Vol. 1: Craniodental Morphology of Genus Homo (Europe) Wiley-Liss: New York.

Stone R (2009). Signs of Early Homo sapiens in China? Science 326 (5953) p 655.

Bookmark and Share
Above image: Institute of Vertebrate Palaeontology and Palaeoanthropology, Chinese Academy of Sciences.

Full frontal hominins

Modern human skull

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.

Homo erectus skull

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.

Basicranial flexion

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.

References

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).

Bookmark and Share