How mangabey molar form differs under routine vs. fallback hard-object feeding regimes
Authors: Guatelli-Steinberg Debbie , Renteria Cameron , Grimm Jack R. , Maeret Carpenter Izabela , Arola Dwayne D. , McGraw W. Scott 
Published: December 2023

Background
Components of diet known as fallback foods are argued to be critical in shaping primate dental anatomy. Such foods of low(er) nutritional quality are often non-preferred, mechanically challenging resources that species resort to during ecological crunch periods. An oft-cited example of the importance of dietary fallbacks in shaping primate anatomy is the grey-cheeked mangabey Lophocebus albigena. This species relies upon hard seeds only when softer, preferred resources are not available, a fact which has been linked to its thick dental enamel. Another mangabey species with thick enamel, the sooty mangabey Cercocebus atys, processes a mechanically challenging food year-round. That the two mangabey species are both thickly-enameled suggests that both fallback and routine consumption of hard foods are associated with the same anatomical feature, complicating interpretations of thick enamel in the fossil record. We anticipated that aspects of enamel other than its thickness might differ between Cercocebus atys and Lophocebus albigena. We hypothesized that to function adequately under a dietary regime of routine hard-object feeding, the molars of Cercocebus atys would be more fracture and wear resistant than those of Lophocebus albigena.

Methods
Here we investigated critical fracture loads, nanomechanical properties of enamel, and enamel decussation in Cercocebus atys and Lophocebus albigena. Molars of Cercopithecus, a genus not associated with hard-object feeding, were included for comparison. Critical loads were estimated using measurements from 2D µCT slices of upper and lower molars. Nanomechanical properties (by nanoindentation) and decussation of enamel prisms (by SEM-imaging) in trigon basins of one upper second molar per taxon were compared.

Results
Protocone and protoconid critical fracture loads were significantly greater in Cercocebus atys than Lophocebus albigena and greater in both than in Cercopithecus. Elastic modulus, hardness, and elasticity index in most regions of the crown were greater in Cercocebus atys than in the other two taxa, with the greatest difference in the outer enamel. All taxa had decussated enamel, but that of Cercocebus atys uniquely exhibited a bundle of transversely oriented prisms cervical to the radial enamel. Quantitative comparison of in-plane and out-of-plane prism angles suggests that decussation in trigon basin enamel is more complex in Cercocebus atys than it is in either Lophocebus albigena or Cercopithecus cephus. These findings suggest that Cercocebus atys molars are more fracture and wear resistant than those of Lophocebus albigena and Cercopithecus. Recognition of these differences between Cercocebus atys and Lophocebus albigena molars sharpens our understanding of associations between hard-object feeding and dental anatomy under conditions of routine vs. fallback hard-object feeding and provides a basis for dietary inference in fossil primates, including hominins.