Biochronology of South African hominin-bearing sites: A reassessment using cercopithecid primates.

Despite recent advances in chronometric techniques (e.g., Uranium-Lead [U-Pb], cosmogenic nuclides, electron spin resonance spectroscopy [ESR]), considerable uncertainty remains regarding the age of many Plio-Pleistocene hominin sites, including several in South Africa. Consequently, biochronology remains important in assessments of Plio-Pleistocene geochronology and provides direct age estimates of the fossils themselves. Historically, cercopithecid monkeys have been among the most useful taxa for biochronology of early hominins because they are widely present and abundant in the African Plio-Pleistocene record. The last major studies using cercopithecids were published over 30 y ago. Since then, new hominin sites have been discovered, radiometric age estimates have been refined, and many changes have occurred in cercopithecid taxonomy and systematics. Thus, a biochronological reassessment using cercopithecids is long overdue. Here, we provide just such a revision based on our recent study of every major cercopithecid collection from African Plio-Pleistocene sites. In addition to correlations based on shared faunal elements, we present an analysis based on the dentition of the abundant cercopithecid Theropithecus oswaldi, which increases in size in a manner that is strongly correlated with geological age (r2 ∼0.83), thereby providing a highly accurate age-estimation tool not previously utilized. In combination with paleomagnetic and U-Pb data, our results provide revised age estimates and suggest that there are no hominin sites in South Africa significantly older than ∼2.8 Ma. Where conflicting age estimates exist, we suggest that additional data are needed and recall that faunal estimates have ultimately proved reliable in the past (e.g., the age of the KBS Tuff).

Nutritional contributions of insects to primate diets: implications for primate evolution.

Insects and other invertebrates form a portion of many living and extinct primate diets. We review the nutritional profiles of insects in comparison with other dietary items, and discuss insect nutrients in relation to the nutritional needs of living primates. We find that insects are incorporated into some primate diets as staple foods whereby they are the majority of food intake. They can also be incorporated as complements to other foods in the diet, providing protein in a diet otherwise dominated by gums and/or fruits, or be incorporated as supplements to likely provide an essential nutrient that is not available in the typical diet. During times when they are very abundant, such as in insect outbreaks, insects can serve as replacements to the usual foods eaten by primates. Nutritionally, insects are high in protein and fat compared with typical dietary items like fruit and vegetation. However, insects are small in size and for larger primates (>1 kg) it is usually nutritionally profitable only to consume insects when they are available in large quantities. In small quantities, they may serve to provide important vitamins and fatty acids typically unavailable in primate diets. In a brief analysis, we found that soft-bodied insects are higher in fat though similar in chitin and protein than hard-bodied insects. In the fossil record, primates can be defined as soft- or hard-bodied insect feeders based on dental morphology. The differences in the nutritional composition of insects may have implications for understanding early primate evolution and ecology.

Cercopithecoid cervical vertebral morphology and implications for the presence of Theropithecus in early Pleistocene Europe.

While it is recognized that the overall configuration of the vertebral column, as well as the size and shape of individual vertebrae, differ within and between primate taxa, relatively little is known about the degree to which vertebral morphology reflects a phylogenetic signal or the degree to which vertebral elements can be used in accurate taxonomic classification. Isolated vertebrae are occasionally found in fossil assemblages, and proper taxonomic identification is necessary to make inferences about the animal's biology and place it in a broader phylogenetic and evolutionary context. Recently, three large primate cervical vertebrae (C3, C5, and C6) from Pirro Nord, Italy (early Pleistocene, late Villafranchian) were attributed to the genus Theropithecus based on size comparisons with extant cercopithecoid primates (Rook et al., 2004, J. Hum. Evol. 47, 267-277). These fossils were suggested to indicate an early dispersal of this genus out of Africa around 1.6-1.3 Ma possibly co-incident with early Pleistocene dispersals of Homo. Because of the potential importance of these fossils for interpreting Theropithecus evolution and the relatively few morphological data on primate cervical vertebral morphology, we examined the size and shape of cervical vertebrae in a large sample of extant cercopithecoid taxa (n=106). Specifically, we evaluated whether subfamily and genus level assignments can be made on the basis of isolated cervical elements. Discriminant analyses reveal that scaled shape variables are good discriminators of taxonomic affinity at the subfamily level but are poor discriminators at the genus level. Least-squares regressions show that raw linear dimensions of cervical vertebral morphology are good predictors of body mass in the extant sample. Our regression results produce a likely body mass estimate of 22-38 kg for the Pirro Nord cervical vertebrae. Based on these regression estimates, the poor ability to discriminate cervical vertebrae at the genus level, and paleoenvironmental reconstructions of Pirro Nord, it is unlikely that the Pirro Nord fossils can be confidently attributed to the genus Theropithecus. These findings have important implications for recent interpretations of the nature of Theropithecus dispersal out of Africa.