The impact of cranial plasticity on the reconstruction of human population history

Palaeoanthropologists are interested in reconstructing the evolutionary history of our species from fossil remains. Like other taxonomists they are primarily interested in identifying shared derived traits, otherwise known as synapomorphies. A synapomorphic trait is shared by some members of a taxa and not by others, since the former inherited this trait from a common ancestor. For instance, the retention of a clavicle (collar bone) is one of the more cited synapomorphic traits common to all primates.

A homoplasy is a trait that is present in two or more taxa but that has not been derived through common ancestry but rather through convergence, parallelism, or reversal. The wings of insects, birds and bats are homoplasies, since they arose through convergent evolution. Thus, homoplasies and synapomorphies may be identical in appearance but are distinguished by whether or not they arose through common ancestry. As a result, it can often difficult to pry apart traits which are synapomorphies from those which are homoplasies. A subset of homoplasies are termed homoiologies. Lycett and Collard (2005) define homoiologies as:

“… phylogenetically misleading resemblances among a group of taxa that can be ascribed to phenotypic plasticity. That is, homoiologies are homoplasies that result from the expression by a genotype of different phenotypes in response to different environmental conditions.”

They arise primarily from nonheritable epigenetic responses to mechanical stimuli. The "homoiology hypothesis" (Lieberman 1995) was derived from the well known fact that bone shape and size can be modified by mechanical loading. As such, homoiologies are expected to have greater influence upon the more plastic regions of a phenotype.

The homoiology hypothesis makes two testable predictions:

1. Traits subject to biomechanical stress should exhibit higher within-taxon variability due to the increased plasticity.

2. These traits should be less reliable for reconstructing phylogeny.

Previous studies have examined the homoiology hypothesis in various primate species (Collard and Wood 2007, 2000). These studies found that the regions of the skull associated with mastication indeed exhibited higher within-taxon variability but they were as reliable in reconstructing phylogenetic relationships as other regions of cranium not directly associated with masticatory function. Since these studies looked at interspecific studies, it was suggested that maybe homoiology was a greater problem for intraspecific studies.

A paper (in press) in the Journal of Human Evolution by Noreen von Cramon-Taubadel tests the homoiology hypothesis in an intraspecific study of human populations. Areas of the skull related with mastication would be expected to be under greater biomechanical stress, and as such be more affected by homoiology. She divided the skull into zones thought to be related to mastication (zygotemporal and palatomaxilla regions) and zones relatively unaffected by mastication (the upper face, cranial vault and basicranium). She tested the predictions of the homoiology hypothesis by comparing craniometric data with matched molecular data for 12 modern human populations.

Like previous interspecific studies, regions of the skull related to mastication show great variability (as predicted by the homoiology hypothesis) but these regions were no less reliable at reconstructing phylogenies (at variance with the homoiology hypothesis). It is worth noting that if biomechanical stress affects all individuals in the same way then these characters will not confound the phylogenetic analysis.

These findings mean that the homoiology hypothesis is flawed in at least some of its premises. The results of this and previous studies suggest that within-taxon variability should not be used to assertion the usefulness of cranial traits for determining phylogenetic relationships. Moreover, it does not appear to hold that homoiologies are any more problematic in determining intraspecific evolutionary relationships as interspecific ones. Finally, even though regions of the skull related to mastication are more variable than non-masticatory regions, they do not seem to be any less reliable for the reconstruction of phylogenies.

References
Collard and Wood. Hominin homoiology: An assessment of the impact of phenotypic plasticity on phylogenetic analyses of humans and their fossil relatives. Journal of human evolution (2007) vol. 52 (5) pp. 573-584.

Collard and Wood. How reliable are human phylogenetic hypotheses?. Proc. Natl. Acad. Sci. U.S.A. (2000) vol. 97 (9) pp. 5003-6.

Cramon-Taubadel. Revisiting the homoiology hypothesis: the impact of phenotypic plasticity on the reconstruction of human population history from craniometric data. Journal of Human Evolution (2009) pp. 1-12.

Lieberman. Testing hypotheses about recent human evolution from skulls: integrating morphology, function, development, and phylogeny. Curr. Anthropol. (1995) 36, 159–197.

Lycett and Collard. Do homoiologies impede phylogenetic analyses of the fossil hominids? An assessment based on extant papionin craniodental morphology. Journal of human evolution (2005) vol. 49 (5) pp. 618-642.

Wood and Lieberman. Craniodental variation in Paranthropus boisei: a developmental and functional perspective. American Journal of Physical Anthropology (2001) vol. 116 (1) pp. 13-25.



Above photo by wauter de tuinkabouter under creative commons license.