Dental microwear and diet of the Plio-Pleistocene hominin Paranthropus boisei

The Plio-Pleistocene hominin Paranthropus boisei had enormous, flat, thickly enameled cheek teeth, a robust cranium and mandible, and inferred massive, powerful chewing muscles. This specialized morphology, which earned P. boisei the nickname "Nutcracker Man", suggests that this hominin could have consumed very mechanically challenging foods. It has been recently argued, however, that specialized hominin morphology may indicate adaptations for the consumption of occasional fallback foods rather than preferred resources. Dental microwear offers a potential means by which to test this hypothesis in that it reflects actual use rather than genetic adaptation. High microwear surface texture complexity and anisotropy in extant primates can be associated with the consumption of exceptionally hard and tough foods respectively. Here we present the first quantitative analysis of dental microwear for P. boisei. Seven specimens examined preserved unobscured antemortem molar microwear. These all show relatively low complexity and anisotropy values. This suggests that none of the individuals consumed especially hard or tough foods in the days before they died. The apparent discrepancy between microwear and functional anatomy is consistent with the idea that P. boisei presents a hominin example of Liem's Paradox, wherein a highly derived morphology need not reflect a specialized diet.     

Dental microwear texture analysis of hominins recovered by the Olduvai Landscape Paleoanthropology Project, 1995-2007

Dental microwear analysis has proven to be a valuable tool for the reconstruction of aspects of diet in early hominins. That said, sample sizes for some groups are small, decreasing our confidence that results are representative of a given taxon and making it difficult to assess within-species variation. Here we present microwear texture data for several new specimens of Homo habilis and Paranthropus boisei from Olduvai Gorge, bringing sample sizes for these species in line with those published for most other early hominins. These data are added to those published to date, and microwear textures of the enlarged sample of H. habilis (n = 10) and P. boisei (n = 9) are compared with one another and with those of other early hominins. New results confirm that P. boisei does not have microwear patterns expected of a hard-object specialist. Further, the separate texture complexity analyses of early Homo species suggest that Homo erectus ate a broader range of foods, at least in terms of hardness, than did H. habilis, P. boisei, or the “gracile” australopiths studied. Finally, differences in scale of maximum complexity and perhaps textural fill volume between H. habilis and H. erectus are noted, suggesting further possible differences between these species in diet.     

Molar microwear in Praeanthropus afarensis: evidence for dietary stasis through time and under diverse paleoecological conditions

Molar microwear fabrics in extant mammals vary with diet and, more particularly, the physical properties of the items that are consumed. Praeanthropus afarensis is well represented in the fossil record over a prolonged and radiometrically controlled temporal span, and reasonably robust paleoecological reconstructions are available for the various localities from which it is known. We therefore examined molar microwear in this species to determine whether diet varied in relation to time or in response to different ecological conditions. Of more than 70 specimens of Pr. afarensis that contain one or more worn permanent molars, only 19 were found to be suitable for microwear analysis. These derive from eight temporal horizons in the Laetolil Beds and Hadar Formation spanning approximately 400kyr (3.6-3.2Ma). Six paleoecological categories have been reconstructed for these horizons, and these were ranked on the basis of floral cover. None of the microwear variables observed for Pr. afarensis is significantly associated with either temporal or paleoecological rank. Thus, microwear and, by extension, diet does not appear to have altered significantly in Pr. afarensis through time or in response to different paleoecological circumstances. The wear pattern that appears to have characterized Pr. afarensis overlaps extensively that of Gorilla gorilla beringei and differs notably from the fabrics of extant primates (e.g., Cebus apella and Cercocebus albigena) that consume hard objects. The high proportion of scratches on Pr. afarensis molars suggests the inclusion of fine abrasives in or on the food items consumed by those individuals sampled in this study. Although Pr. afarensis may have been morphologically equipped to process hard, brittle items, the microwear data suggest that it did not necessarily do so, even in the face of varying environmental circumstances. Explanatory scenarios that describe Pr. afarensis as part of an evolutionary trajectory involving a more heavily masticated diet with an increased reliance on hard, brittle items need to be reconsidered. However, fallback foods that were consumed during relatively short, albeit critical periods may have exerted sufficient selective pressure to explain the evolution of the comparatively robust Pr. afarensis trophic apparatus. Because it is unlikely that many individuals from such restricted temporal intervals would be sampled in the paleontological record, we suggest that the most productive approach to the elucidation of paleodiet is the integration of genetic (morphological) and epigenetic (microwear and isotopic) lines of evidence.     

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species (Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.     

Molar microwear and dietary reconstructions of fossil cercopithecoidea from the Plio-Pleistocene deposits of South Africa

The South African Plio-Pleistocene cave deposits have yielded a diverse cercopithecoid fauna. In this study, the possible dietary proclivities of these extinct species are examined using details of molar microwear. Although sample sizes are often small, wear patterns suggest possible temporal changes in the diets of Parapapio jonesi from Makapansgat to Sterkfontein, of Papio robinsoni from Sterkfontein to Swartkrans, and Cercopithecoides williamsi from Makapansgat to Sterkfontein to Swartkrans. However, there does not appear to have been a significant change in the dietary habits of Parapapio broomi over time. The microwear patterns of the two temporally successive congeners, Theropithecus darti and T. oswaldi show no significant differences from one another. The sympatric congeners, Parapapio broomi and Pp. jonesi, have microwear signatures that differ significantly at Makapansgat (Members 3 and 4) but not at Sterkfontein (Member 4). Finally, the microwear analyses suggest that the extinct cercopithecoid species did not necessarily have diets similar to those of their closest living relatives.     

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 primate community of Cachoeira (Brazilian Amazonia): a model to decipher ecological partitioning among extinct species

Dental microwear analysis is conducted on a community of platyrrhine primates from South America. This analysis focuses on the primate community of Cachoeira Porteira (Para, Brazil), in which seven sympatric species occur: Alouatta seniculus, Ateles paniscus, Cebus apella, Chiropotes satanas, Pithecia Pithecia, Saguinus midas, and Saimiri sciureus. Shearing quotients are also calculated for each taxon of this primate community. Dental microwear results indicate significant differences between taxa, but are somewhat insufficient when it comes to discriminating between ecologically similar taxa. The primates of Cachoeira Porteira all incorporate a certain amount of fruit in their diet, entailing a definite amount of inter-specific competition as they must share food resources. Alouatta is the most folivorous taxon of this community, which is corroborated by dental microwear analysis. Ateles, although of a similar size to Alouatta, limits inter-specific competition by incorporating more fruit in its diet. Cebus has a very diverse omnivorous diet, which is highlighted in this study, as it compares to both fruit and leaf eating taxa. In some cases, microwear results need to be supplemented by other methods. For example, dental microwear seems insufficient to distinguish between Pithecia and Chiropotes, which eat foods with similar physical properties. However, other methods (i.e. shearing quotients and body mass) provide enough complimentary information to be able to highlight differences between the two taxa. On the other hand, dental microwear can highlight differences between primates which have similar diets, such as Saimiri and Saguinus. In this case, differences could be due to other exogenous factors.     

Dental microwear evidence for a dietary shift between two nonmaize-reliant prehistoric human populations from Indiana

Several recent studies have demonstrated the efficacy of dental microwear analysis (DMA) for dietary reconstruction among nonhuman primates, early hominids, and prehistoric humans. The current study seeks microwear evidence for a paleobotanically suggested change in the types of foods that were consumed by two temporally distinct populations of the North American eastern woodlands. This particular dietary shift between the Late Archaic and the Early/Middle Woodland periods did not include the introduction of maize or any other tropical cultigen. In contrast, most dietary reconstructions from this area have compared later populations that consumed tropical cultigens (such as maize) with those that did not. High-resolution casts of adult mandibular second molar protoconid phase II wear facets were viewed via a scanning electron microscope at 500x. Photomicrographs of the microwear features were analyzed with specialized software, Microware 2.2 (Ungar [1995] Scanning 17:57-59). Analysis of variance statistical tests were performed, with one variable requiring rank-transformation. A dietary transition is evidenced by a statistically significant increase in the mean number of pits and concomitant decreases in scratch width and scratch length from the Late Archaic to the Early/Middle Woodland. Overall, the diet became harder and less abrasive. The implication here is that dental microwear analysis is able to discern relatively subtle dietary transitions in human populations that may not be as readily accessible by other means of dietary reconstruction.     

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.     

Dental microwear variability on buccal tooth enamel surfaces of extant Catarrhini and the Miocene fossil Dryopithecus laietanus (Hominoidea)

Analyses of buccal tooth microwear have been used to trace dietary habits of modern hunter-gatherer populations. In these populations, the average density and length of striations on the buccal surfaces of teeth are significantly cor-related with the abrasive potential of food items consumed. In non-human pri-mates, tooth microwear patterns on both occlusal and buccal wear facets have been thoroughly studied and the results applied to the characterization of dietary habits of fossil species. In this paper, we present inter- and intra-specific buccal microwear variability analyses in extant Cercopithecoidea (Cercopithecus mitis, C. neglectus, Chlorocebus aethiops, Colobus spp., Papio anubis) and Hominoidea (Gorilla gorilla, Pan troglodytes, Pongo pygmaeus). The results are tentatively compared to buccal microwear patterns of the Miocene fossils Dryopithecus and Oreopithecus. Significant differences in striation density and length are found among the fossil taxa studied and the extant primates, suggesting that buccal microwear can be used to identify dietary differences among taxa. The Dryopithecus buccal microwear pattern most closely resembles that of abrasive, tough plant foods consumers, such as the gorilla, in contrast to stud-ies of dental morphology that suggest a softer, frugivorous diet. Results for Oreopithecus were equivocal, but suggest a more abrasive diet than that previously thought.     

牙齿微痕研究在古食性重建中应用的简述

在古食性研究中,牙齿微痕是指动物在咀嚼食物的过程中在牙齿咬合面上产生的微观磨损痕迹。不同食性的动物具有不同的牙齿微痕特征,因此可以通过研究牙齿微痕特征来重建灭绝动物的古食性,为探讨动物演化和古生态环境变化提供重要信息。本文主要介绍牙齿微痕作为一种简单而高效的古食性重建方法在古生物领域中的应用。本文主要内容包括牙齿微痕的发展历史,形成机理与应用,以及近年来被广泛应用的牙齿微痕定量化分析——表面纹理分析法,并在最后浅谈了牙齿微痕研究未来可能研究的方向。     

黄山短尾猴食土行为

从行为生态学角度,依据黄山短尾猴食土行为在年龄、性别、社会等级序位、食土量、食土频率、食土持续时间中的分布和变化规律,结合土壤基本理化性质测定和分析,探讨了黄山短尾猴食土行为。黄山短尾猴对土壤的摄取是寻找和有目的的选择,有些场所是其"喜好"或"常去之处",食土场所通常是以多个体多次取食挖掘而形成的洞穴形式存在。取食土壤颜色黄色或黄棕色,酸性土壤,富含铁、钙、镁等矿物元素,粘土比例较高。社群各年龄段及性别个体均参与食土行为。平均食土行为持续时间在年龄和性别之间无显著差异;平均食土频率和食土量在成年雌性或雄性个体的社会等级序位之间无显著性差异;而成年雌性的平均食土频率和食土量,显著高于其它年龄或性别组,与雌性正值怀孕末期及产仔哺乳期,需要补充大量铁等矿物元素,以维持生理所需和体力消耗有关。典型的植食性动物黄山短尾猴有规律地取食粘土的行为,支持了食土行为具有"食物解毒作用"假说。人工投喂和食土行为的相关性表现为提高了食土频率,可能与粘土能够缓解高热量、低纤维人工食物造成的胃肠不适有关。     

Dietary reconstruction of Spy I using dental microwear texture analysis

Spy I from the Meuse River Basin of Belgium is among the most recent Neandertals. This adult lived at the terminus of Marine Isotope Stage (MIS) 3 in cold steppe environments at the northern edge of the habitable zone for Neandertals where plants were relatively scarce. The dietary proclivities of Spy I are reconstructed using dental microwear texture analysis and compared to 33 Neandertals from western Eurasia, MIS 5 to MIS 3. Spy I has an elevated enamel surface complexity suggesting the consumption of course dietary items such as wild seeds, acorns, nuts, and underground storage organs laden with particles of grit. Unlike the young and old individuals from Hortus with low values for anisotropy, Spy I is closest to the adults from this site suggesting a common pattern of masticatory behavior typified this life cycle stage. Like many other Neandertals, Spy I probably consumed plant foods at appreciable levels, some of which were hard and brittle or poorly processed.     

Cougars’ key to survival through the Late Pleistocene extinction: insights from dental microwear texture analysis

Cougars (Puma concolor) are one of only two large cats in North America to have survived the Late Pleistocene extinction (LPE), yet the specific key(s) to their relative success remains unknown. Here, we compare the dental microwear textures of Pleistocene cougars with sympatric felids from the La Brea Tar Pits in southern California that went extinct at the LPE (Panthera atrox and Smilodon fatalis), to clarify potential dietary factors that led to the cougar''s persistence through the LPE. We further assess whether the physical properties of food consumed have changed over time when compared with modern cougars in southern California. Using dental microwear texture analysis (DMTA), which quantifies surface features in three dimensions, we find that modern and Pleistocene cougars are not significantly different from modern African lions in any DMTA attributes, suggesting moderate durophagy (i.e. bone processing). Pleistocene cougars from La Brea have significantly greater complexity and textural fill volume than Panthera atrox (inferred to have primarily consumed flesh from fresh kills) and significantly greater variance in complexity values than S. fatalis. Ultimately, these results suggest that cougars already used or adopted a more generalized dietary strategy during the Pleistocene that may have been key to their subsequent success.     

Technical note: Dental microwear textures of "Phase I" and "Phase II" facets

The power stroke of mastication has been traditionally divided into two parts, one which precedes centric occlusion, and the other which follows it-"Phase I" and "Phase II," respectively. Recent studies of primate mastication have called into question the role of Phase II in food processing, as they have found little muscle activity or accompanying bone strain following centric occlusion. That said, many researchers today look to Phase II facets to relate diet to patterns of dental microwear. This suggests the need to reevaluate microwear patterns on Phase I facets. Here we use texture analysis to compare and contrast microwear on facets representing both phases in three primate species with differing diets (Alouatta palliata, Cebus apella, and Lophocebus albigena). Results reaffirm that microwear patterns on Phase II facets better distinguish taxa with differing diets than do those on Phase I facets. Further, differences in microwear textures between facet types for a given taxon may themselves reflect diet. Some possible explanations for differences in microwear textures between facet types are proposed.     

New primate first metatarsals from the Paleogene of Egypt and the origin of the anthropoid big toe

The specialized grasping feet of primates, and in particular the nature of the hallucal grasping capabilities of living strepsirrhines and tarsiers (i.e., ‘prosimians’), have played central roles in the study of primate origins. Prior comparative studies of first metatarsal (Mt1) morphology have documented specialized characters in living prosimians that are indicative of a more abducted hallux, which in turn is often inferred to be related to an increased ability for powerful grasping. These include a well-developed peroneal process and a greater angle of the proximal articular surface relative to the long axis of the diaphysis. Although known Mt1s of fossil prosimians share these characters with living non-anthropoid primates, Mt1 morphology in the earliest crown group anthropoids is not well known. Here we describe two Mt1s from the Fayum Depression of Egypt - one from the latest Eocene (from the ∼34 Ma Quarry L-41), and one from the later early Oligocene (from the ∼29-30 Ma Quarry M) - and compare them with a sample of extant and fossil primate Mt1s. Multivariate analyses of Mt1 shape variables indicate that the Fayum specimens are most similar to those of crown group anthropoids, and likely belong to the stem catarrhines Catopithecus and Aegyptopithecus specifically, based on analyses of size. Also, phylogenetic analyses with 16 newly defined Mt1 characters support the hypotheses that “prosimian”-like Mt1 features evolved along the primate stem lineage, while crown anthropoid Mt1 morphology and function is derived among primates, and likely differed from that of basal stem anthropoids. The derived loss of powerful hallucal grasping as reflected in the Mt1 morphology of crown anthropoids may reflect long-term selection for improved navigation of large-diameter, more horizontal branches at the expense of movement in smaller, more variably inclined branches in the arboreal environment.     

Incisor microwear of Sumatran anthropoid primates

Several studies have suggested that incisor microwear reflects diet and feeding adaptations of anthropoids. However, such studies have been largely qualitative, and interpretations have relied on anecdotal references to diet and tooth use reported in the socioecology literature. The current study relates incisor microwear in four anthropoid primates to specific ingestive behaviors and food types. Central incisor casts of wild-shot museum specimens of Hylobates lar, Macaca fascicularis, Pongo pygmaeus, and Presbytis thomasi were examined by scanning electron microscopy, and analyzed using a semiautomated image analysis procedure. Microwear patterns were used to generate predictions regarding diet and anterior tooth use. These predictions were evaluated using data collected during a 1 year study of feeding behavior of these same taxa in the wild (Ungar, 1992, 1994a, b). Results suggest that (1) enamel prism relief is associated with the effectiveness of etching reagents in foods, (2) dental calculus buildup results from a lack of incisor use and perhaps the ingestion of sugar-rich foods, (3) striation density varies with degree of anterior tooth use in the ingestion of abrasive food items, (4) striation breadth is proposed to relate to the ratio of exogenous grit to phytoliths consumed; and (5) preferred striation orientation indicates the direction that food items are pulled across the incisors during ingestion. It is concluded that incisor microwear studies can contribute to the understanding of diets and feeding behaviors of extinct primates. © 1994 Wiley-Liss, Inc.     

Buccal dental microwear variability in extant African Hominoidea: taxonomy versus ecology

Buccal microwear patterns on teeth are good indicators of the abrasiveness of foodstuffs and have been used to trace the dietary habits of fossil species, including primates and hominids. However, few studies have addressed the variability of this microwear. The abrasiveness of dietary components depends not only on the hardness of the particles ingested, but also on the presence of dust and other exogenous elements introduced during food processing. These elements are responsible for the microwear typology observed on the enamel surfaces of primate teeth. Here we analyzed the variability of buccal microwear patterns in African Great Apes (Gorilla gorilla and Pan troglodytes), using tooth molds obtained from the original specimens held in several osteological collections. Our results suggest that ecological adaptations at subspecies or population level account for differences in microwear patterns, which are attributed to habitat and ecological conditions within populations rather than differences between species. The findings from studies on the variability of buccal dental microwear in extant species will contribute to a better understanding of extinct hominids’ diet and ecology.     

Premolar microwear and tooth use in Australopithecus afarensis

The mandibular third premolar (P₃) of Australopithecus afarensis is notable for extensive morphological variability (e.g., metaconid presence/absence, closure of the anterior fovea, root number) and temporal trends in crown length and shape change over its 700 Ka time range. Hominins preceding A. afarensis have unicuspid, mesiodistally elongated P₃s with smaller talonids, and subsequent australopiths have bicuspid, more symmetrically-shaped P₃ crowns with expanded talonids. For these features, A. afarensis is intermediate and, thus, evinces the incipient stages of P₃ molarization. Here, we examine A. afarensis P₃ Phase II microwear and compare it with that of Australopithecus africanus and Cercocebus atys, an extant hard-object specialist, to assess whether the role of the P₃ in food processing changed over time in A. afarensis. Premolar Phase II microwear textures are also compared with those of the molars to look for evidence of functional differentiation along the tooth row (i.e., that foods with different mechanical properties were processed by separate regions of the postcanine battery). Microwear textures were also examined along the mesial protoconid crest, the site of occlusion with the maxillary canine, of the A. afarensis P₃ and compared with the same region in Pan troglodytes to determine whether microwear can be useful for identifying changes in the occlusal relationship between the P₃ and maxillary canine in early Australopithecus. Finally, temporal trends in P₃ Phase II and mesial microwear are considered. Results indicate that 1) both the P₃ and molar Phase II facets of A. afarensis have less complex microwear textures than in A. africanus or C. atys; 2) A. afarensis P₃ and molar Phase II textures differ, though not to the extent seen in taxa that eat hard and tough items; 3) microwear along the A. afarensis mesial protoconid crest is clearly distinct from that of the P. troglodytes, indicating that there is no honing equivalent in A. afarensis; and 4) there is little evidence of change over time in A. afarensis P₃ microwear on either the mesial or Phase II facet. In sum, these results provide no evidence that A. afarensis routinely loaded either its premolars or molars to process hard objects or that A. afarensis P₃ function changed over time.     

Dental microwear texture analysis: technical considerations

Dental microwear analysis is commonly used to infer aspects of diet in extinct primates. Conventional methods of microwear analysis have usually been limited to two-dimensional imaging studies using a scanning electron microscope and the identification of apparent individual features. These methods have proved time-consuming and prone to subjectivity and observer error. Here we describe a new methodological approach to microwear: dental microwear texture analysis, based on three-dimensional surface measurements taken using white-light confocal microscopy and scale-sensitive fractal analysis. Surface parameters for complexity, scale of maximum complexity, anisotropy, heterogeneity, and textural fill volume offer repeatable, quantitative characterizations of three-dimensional surfaces, free of observer measurement error. Some results are presented to illustrate how these parameters distinguish extant primates with different diets. In this case, microwear surfaces of Cebus apella and Lophocebus albigena, which consume some harder food items, have higher average values for complexity than do folivores or soft fruit eaters.