Peter S. Ungar

Dental microwear texture analysis shows within-species diet variability in fossil hominins.

Reconstructing the diets of extinct hominins is essential to understanding the paleobiology and evolutionary history of our lineage. Dental microwear, the study of microscopic tooth-wear resulting from use, provides direct evidence of what an individual ate in the past. Unfortunately, established methods of studying microwear are plagued with low repeatability and high observer error. Here we apply an objective, repeatable approach for studying three-dimensional microwear surface texture to extinct South African hominins. Scanning confocal microscopy together with scale-sensitive fractal analysis are used to characterize the complexity and anisotropy of microwear. Results for living primates show that this approach can distinguish among diets characterized by different fracture properties. When applied to hominins, microwear texture analysis indicates that Australopithecus africanus microwear is more anisotropic, but also more variable in anisotropy than Paranthropus robustus. This latter species has more complex microwear textures, but is also more variable in complexity than A. africanus. This suggests that A. africanus ate more tough foods and P. robustus consumed more hard and brittle items, but that both had variable and overlapping diets.

The Diets of Early Hominins

Diet changes are considered key events in human evolution. Most studies of early hominin diets focused on tooth size, shape, and craniomandibular morphology, as well as stone tools and butchered animal bones. However, in recent years, dental microwear and stable isotope analyses have hinted at unexpected diversity and complexity in early hominin diets. Some traditional ideas have held; others, such as an increasing reliance on hard-object feeding and a dichotomy between Australopithecus and Paranthropus, have been challenged. The first known evidence of C4 plant (tropical grasses and sedges) and hard-object (e.g., seeds and nuts) consumption dates to millions of years after the appearance of the earliest probable hominins, and there are no consistent trends in diet change among these species through time.

Diet in Early Homo: A Review of the Evidence and a New Model of Adaptive Versatility

Several recent studies have stressed the role of dietary change in the origin and early evolution of our genus in Africa. Resulting models have been based on nutrition research and analogy to living peoples and nonhuman primates or on archeological and paleoenvironmental evidence. Here we evaluate these models in the context of the hominin fossil record. Inference of diet from fossils is hampered by small samples, unclear form-function relationships, taphonomic factors, and interactions between cultural and natural selection. Nevertheless, craniodental remains of Homo habilis, H. rudolfensis, and H. erectus offer some clues. For example, there appears to be no simple transition from an australopith to a Homo grade of dietary adaptation, or from closed forest plant diets to reliance on more open-country plants or animals. Early Homo species more likely had adaptations for flexible, versatile subsistence strategies that would have served them well in the variable paleoenvironments of the African Plio-Pleistocene.

A solution to the worn tooth conundrum in primate functional anatomy

Worn teeth are a bane to paleobiologists interested in the diets of human ancestors and other fossil primates. Although worn teeth dominate fossil assemblages, their shapes are usually not used to reconstruct the diets of extinct species. The problem is that traditional studies of primate dental functional anatomy have focused on unworn morphology. This has limited most functional analyses to only a few well-represented fossil species. This paper introduces a method to characterize and compare worn occlusal morphology in primates using laser scanning and geographic information systems technologies. A study of variably worn chimpanzee and gorilla molars indicates that differences between these species in tooth shape remain consistent at given stages of wear. Although cusp slope decreases with wear in both taxa, angularity values remain unchanged. These results indicate that African ape teeth wear in a manner that keeps them mechanically efficient for fracturing specific foods. Studies of changes in tooth shape with wear add a new dimension to dental functional anatomy, and offer a more complete picture of dental-dietary adaptations. Also, given how rare unworn teeth are in the fossil record, the ability to include worn specimens in analyses opens the door to reconstructing the diets of many more extinct primate groups, allowing us to better understand the adaptive radiation of our order.

Fruit preferences of four sympatric primate species at Ketambe, northern Sumatra, Indonesia

Recent researchers have drawn attention to fruit preferences in a variety of primates;for instance, in 1991, Davies noted that monogastric primates prefer the flesh of succulent, sugar-rich fruits, while colobines more often consume and digest large seeds of drier fruits. I compare fruit preferences in four sympatric primates— Hylobates lar, Macaca fascicularis, Pongo pygmaeus,and Presbytis thomas—which I studied concurrently at the Ketambe Research Center in northern Sumatra. I collected continuous focal animal data during 40–50 hr per taxon per month for 10 months and recorded fruit species size,pH, and descriptive attributes, including degree of ripeness, hardness, pericarp type, and number of seeds. The langurs prefer dry fruit seeds while the monogastric primates more often consume acidic, succulent fruit flesh. Further, H. lar, M. fascicularis,and P. pygmaeusvary significantly in preferences for fruits vis-à-visthe characteristics examined.

Dental Microwear Texture Analysis of Varswater Bovids and Early Pliocene Paleoenvironments of Langebaanweg, Western Cape Province, South Africa

The extensive early Pliocene mammalian assemblages at Langebaanweg hold the potential to provide important information about paleoenvironments of the southwestern tip of Africa, an area that today consititutes the Fynbos Biome. We here add to a growing body of literature on the paleoenviornments of the site with an examination of dental microwear textures of bovids from the Varswater Formation. Microwear texture analysis is a new, automated and repeatable approach that measures whole surfaces in three dimensions without observer error. A study of extant ruminants indicates that grazers have more anisotropic microwear surface textures, whereas browsers have more complex microwear surface textures. Fossil bovids recovered from the Muishond Fontein Pelletal Phosphorite Member vary in their microwear textures, with some taxa falling within the extant browser range, some closer to extant grazers, and others in between. These results are consistent with scenarios suggesting mosaic habitats including fynbos vegetation, some (probably C3) grasses, and woodland elements when these fossils were accumulated.

Dental microwear texture and anthropoid diets

Dental microwear has long been used as evidence concerning the diets of extinct species. Here, we present a comparative baseline series of dental microwear textures for a sample of 21 anthropoid primate species displaying interspecific and intraspecific dietary variability. Four dental microwear texture variables (complexity, anisotropy, textural fill volume, and heterogeneity) were computed based on scale-sensitive fractal analysis and high-resolution three-dimensional renderings of microwear surfaces collected using a white-light confocal profiler. The purpose of this analysis was to assess the extent to which these variables reflect variation in diet. Significant contrasts between species with diets known to include foods with differing material properties are clearly evident for all four microwear texture variables. In particular, species that consume more tough foods, such as leaves, tended to have high levels of anisotropy and low texture complexity. The converse was true for species including hard and brittle items in their diets either as staples or as fallback foods. These results reaffirm the utility of dental microwear texture analysis as an important tool in making dietary inferences based on fossil primate samples.

Molar microwear textures and the diets of Australopithecus anamensis and Australopithecus afarensis

Many researchers have suggested that Australopithecus anamensis and Australopithecus afarensis were among the earliest hominins to have diets that included hard, brittle items. Here we examine dental microwear textures of these hominins for evidence of this. The molars of three Au. anamensis and 19 Au. afarensis specimens examined preserve unobscured antemortem microwear. Microwear textures of these individuals closely resemble those of Paranthropus boisei, having lower complexity values than Australopithecus africanus and especially Paranthropus robustus. The microwear texture complexity values for Au. anamensis and Au. afarensis are similar to those of the grass-eating Theropithecus gelada and folivorous Alouatta palliata and Trachypithecus cristatus. This implies that these Au. anamensis and Au. afarensis individuals did not have diets dominated by hard, brittle foods shortly before their deaths. On the other hand, microwear texture anisotropy values for these taxa are lower on average than those of Theropithecus, Alouatta or Trachypithecus. This suggests that the fossil taxa did not have diets dominated by tough foods either, or if they did that directions of tooth–tooth movement were less constrained than in higher cusped and sharper crested extant primate grass eaters and folivores.

The diet of Australopithecus sediba

Specimens of Australopithecus sediba from the site of Malapa, South Africa (dating from approximately 2 million years (Myr) ago) present a mix of primitive and derived traits that align the taxon with other Australopithecus species and with early Homo. Although much of the available cranial and postcranial material of Au. sediba has been described, its feeding ecology has not been investigated. Here we present results from the first extraction of plant phytoliths from dental calculus of an early hominin. We also consider stable carbon isotope and dental microwear texture data for Au. sediba in light of new palaeoenvironmental evidence. The two individuals examined consumed an almost exclusive C3 diet that probably included harder foods, and both dicotyledons (for example, tree leaves, fruits, wood and bark) and monocotyledons (for example, grasses and sedges). Like Ardipithecus ramidus (approximately 4.4 Myr ago) and modern savanna chimpanzees, Au. sediba consumed C3 foods in preference to widely available C4 resources. The inferred consumption of C3 monocotyledons, and wood or bark, increases the known variety of early hominin foods. The overall dietary pattern of these two individuals contrasts with available data for other hominins in the region and elsewhere.

Dental Evidence for Diet in Some Miocene Catarrhines with Comments on the Effects of Phylogeny on the Interpretation of Adaptation

Studies of the dental anatomy of Miocene catarrhines have concentrated on either phylogenetic or adaptive interpretations. Most investigations of systematics or dental function have been considered more mutually exclusive than reciprocally illuminating. In this chapter, we attempt to develop a balanced view of the two together. Data presented here suggest that functional inferences require consideration of the phylogenetic affinities of the groups being compared. While the way a character functions is independent of its phylogenetic polarity, the way in which that character manifests itself may depend, in part, on phylogenetic considerations.