Patterns and magnitudes of craniofacial covariation in extant cercopithecids.

The cranium contains almost all of the vertebrate sensory organs and plays an essential role in vertebrate evolution. Research on the primate cranium has revealed that it is both highly integrated and modular, but studies have historically focused on covariance between the neurocranium and facial skeleton rather than on bones specific to special senses such as vision. The goal of this work is to investigate patterns and magnitudes of craniofacial covariation in extant cercopithecids with particular attention to the orbits. This study takes a quantitative approach using data collected from 38 homologous cranial landmarks across 11 genera of cercopithecid monkeys (Cercopithecidae, N = 291). These data demonstrate that both patterns and magnitudes of craniofacial covariation differ across Cercopithecidae at subfamily, tribe, and genus levels, with the strongest integration in the papionins (and specifically Papio) and significantly weaker covariation in the colobines, particularly Presbytis. Orbital height does not covary with other measurements of the cranium to the same degree as other cranial traits in Cercopithecidae and is highly constrained across the family. This study has important implications for our understanding of the evolution and development of morphological diversity in the cercopithecid cranium and evolution of the primate eye. This study also highlights the potential error of broad assumptions about generalizing patterns and magnitudes of modularity and integration across primates. Additionally, these findings reiterate the importance of trait selection for interpreting fossil taxonomy, as craniofacial covariation may impact phenotypes commonly used to differentiate fossil primate species.