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.     

Diet-Related Buccal Dental Microwear Patterns in Central African Pygmy Foragers and Bantu-Speaking Farmer and Pastoralist Populations

Pygmy hunter-gatherers from Central Africa have shared a network of socioeconomic interactions with non-Pygmy Bantu speakers since agropastoral lifestyle spread across sub-Saharan Africa. Ethnographic studies have reported that their diets differ in consumption of both animal proteins and starch grains. Hunted meat and gathered plant foods, especially underground storage organs (USOs), are dietary staples for pygmies. However, scarce information exists about forager–farmer interaction and the agricultural products used by pygmies. Since the effects of dietary preferences on teeth in modern and past pygmies remain unknown, we explored dietary history through quantitative analysis of buccal microwear on cheek teeth in well-documented Baka pygmies. We then determined if microwear patterns differ among other Pygmy groups (Aka, Mbuti, and Babongo) and between Bantu-speaking farmer and pastoralist populations from past centuries. The buccal dental microwear patterns of Pygmy hunter-gatherers and non-Pygmy Bantu pastoralists show lower scratch densities, indicative of diets more intensively based on nonabrasive foodstuffs, compared with Bantu farmers, who consume larger amounts of grit from stoneground foods. The Baka pygmies showed microwear patterns similar to those of ancient Aka and Mbuti, suggesting that the mechanical properties of their preferred diets have not significantly changed through time. In contrast, Babongo pygmies showed scratch densities and lengths similar to those of the farmers, consistent with sociocultural contacts and genetic factors. Our findings support that buccal microwear patterns predict dietary habits independent of ecological conditions and reflect the abrasive properties of preferred or fallback foods such as USOs, which may have contributed to the dietary specializations of ancient human populations.     

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.     

Diet of upper paleolithic modern humans: Evidence from microwear texture analysis

This article presents the results of the occlusal molar microwear texture analysis of 32 adult Upper Paleolithic modern humans from a total of 21 European sites dating to marine isotope stages 3 and 2. The occlusal molar microwear textures of these specimens were analyzed with the aim of examining the effects of the climatic, as well as the cultural, changes on the diets of the Upper Paleolithic modern humans. The results of this analysis do not reveal any environmentally driven dietary shifts for the Upper Paleolithic hominins indicating that the climatic and their associated paleoecological changes did not force these humans to significantly alter their diets in order to survive. However, the microwear texture analysis does detect culturally related changes in the Upper Paleolithic humans' diets. Specifically, significant differences in diet were found between the earlier Upper Paleolithic individuals, i.e., those belonging to the Aurignacian and Gravettian contexts, and the later Magdalenian ones, such that the diet of the latter group was more varied and included more abrasive foods compared with those of the former. Am J Phys Anthropol 153:570–581, 2014. © 2014 Wiley Periodicals, Inc.     

Human dental microwear from Ohalo II (22,500-23,500 cal BP), southern Levant

Dietary hardness and abrasiveness are inferred from human dental microwear at Ohalo II, a late Upper Palaeolithic site (22,500-23,500 cal BP) in the southern Levant. Casts of molar grinding facets from two human skeletons were examined with a scanning electron microscope. The size and frequency of microwear was measured, counted, and compared to four prehistoric human groups from successive chronological periods in the same region: pre-pottery Neolithic A, Chalcolithic (this study); Natufian, pre-pottery Neolithic B (Mahoney: Am J Phys Anthropol 130 (2006) 308-319). The Ohalo molars had a high frequency of long narrow scratches, and a few small pits, suggesting a tough abrasive diet that required more shearing rather than compressive force while chewing. These results imply that the diet of the two late Upper Palaeolithic hunter-gatherers did not focus on very hard foods. Aquatic foods with adherent contaminants, as well as grit from plant grinding tools seemed likely causal agents. The size of the pits and scratches on the Ohalo molars were most similar to microwear from the pre-pottery Neolithic A period, though they also compared well to the Chalcolithic period. These results contrasted with the larger pits and scratches from the Natufian hunter-gatherers and pre-pottery Neolithic B farmers, implying that there is no simple increase or decrease in dietary hardness and abrasiveness across the late Upper Palaeolithic to Chalcolithic development in the Southern Levant.     

Neandertal versus Modern Human Dietary Responses to Climatic Fluctuations

The Neandertal lineage developed successfully throughout western Eurasia and effectively survived the harsh and severely changing environments of the alternating glacial/interglacial cycles from the middle of the Pleistocene until Marine Isotope Stage 3. Yet, towards the end of this stage, at the time of deteriorating climatic conditions that eventually led to the Last Glacial Maximum, and soon after modern humans entered western Eurasia, the Neandertals disappeared. Western Eurasia was by then exclusively occupied by modern humans. We use occlusal molar microwear texture analysis to examine aspects of diet in western Eurasian Paleolithic hominins in relation to fluctuations in food supplies that resulted from the oscillating climatic conditions of the Pleistocene. There is demonstrable evidence for differences in behavior that distinguish Upper Paleolithic humans from members of the Neandertal lineage. Specifically, whereas the Neandertals altered their diets in response to changing paleoecological conditions, the diets of Upper Paleolithic humans seem to have been less affected by slight changes in vegetation/climatic conditions but were linked to changes in their technological complexes. The results of this study also indicate differences in resource exploitation strategies between these two hominin groups. We argue that these differences in subsistence strategies, if they had already been established at the time of the first contact between these two hominin taxa, may have given modern humans an advantage over the Neandertals, and may have contributed to the persistence of our species despite habitat-related changes in food availabilities associated with climate fluctuations.     

Dental microwear texture analysis of two families of subfossil lemurs from Madagascar

This study employs dental microwear texture analysis to reconstruct the diets of two families of subfossil lemurs from Madagascar, the archaeolemurids and megaladapids. This technique is based on three-dimensional surface measurements utilizing a white-light confocal profiler and scale-sensitive fractal analysis. Data were recorded for six texture variables previously used successfully to distinguish between living primates with known dietary differences. Statistical analyses revealed that the archaeolemurids and megaladapids have overlapping microwear texture signatures, suggesting that the two families occasionally depended on resources with similar mechanical properties. Even so, moderate variation in most attributes is evident, and results suggest potential differences in the foods consumed by the two families. The microwear pattern for the megaladapids indicates a preference for tougher foods, such as many leaves, while that of the archaeolemurids is consistent with the consumption of harder foods. The results also indicate some intraspecific differences among taxa within each family. This evidence suggests that the archaeolemurids and megaladapids, like many living primates, likely consumed a variety of food types.     

Seasonal Mortality Patterns in Primates: Implications for the Interpretation of Dental Microwear

The microscopic traces of use wear on teeth have been extensively studied to provide information that will assist in elucidating the dietary habits of extinct hominin species. 1 - 13 It has been amply documented that dental microwear provides information pertaining to diet for living animals, where there is a strong and consistent association between dental microwear patterns and different types of foods that are chewed. The details of occlusal surface wear patterns are capable of distinguishing among diets when the constituent food items differ in their fracture properties. 14 - 20 For example, the microwear traces left on the teeth of mammals that crush hard, brittle foods such as nuts are generally dominated by pits, whereas traces left on the teeth of mammals that shear tough items such as leaves tend to be characterized by scratches. These microwear features result from and thus record actual chewing events. As such, microwear patterns are expected to be variably ephemeral, as individual features are worn away and replaced or overprinted by others as the tooth wears down in subsequent bouts of mastication. Indeed, it has been demonstrated, both in the laboratory and the wild, that short‐term dietary variation can result in the turnover of microwear. 17 , 21 - 23 Because occlusal microwear potentially reflects an individual's diet for a short time (days, weeks, or months, depending on the nature of the foods being masticated), tooth surfaces sampled at different times will display differences that relate to temporal (for example, seasonal) differences in diet. 24     

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.     

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.     

Similar Associations of Tooth Microwear and Morphology Indicate Similar Diet across Marsupial and Placental Mammals

Low-magnification microwear techniques have been used effectively to infer diets within many unrelated mammalian orders, but the extent to which patterns are comparable among such different groups, including long extinct mammal lineages, is unknown. Microwear patterns between ecologically equivalent placental and marsupial mammals are found to be statistically indistinguishable, indicating that microwear can be used to infer diet across the mammals. Microwear data were compared to body size and molar shearing crest length in order to develop a system to distinguish the diet of mammals. Insectivores and carnivores were difficult to distinguish from herbivores using microwear alone, but combining microwear data with body size estimates and tooth morphology provides robust dietary inferences. This approach is a powerful tool for dietary assessment of fossils from extinct lineages and from museum specimens of living species where field study would be difficult owing to the animal’s behavior, habitat, or conservation status.     

Viewpoints: Diet and dietary adaptations in early hominins: The hard food perspective

Recent biomechanical analyses examining the feeding adaptations of early hominins have yielded results consistent with the hypothesis that hard foods exerted a selection pressure that influenced the evolution of australopith morphology. However, this hypothesis appears inconsistent with recent reconstructions of early hominin diet based on dental microwear and stable isotopes. Thus, it is likely that either the diets of some australopiths included a high proportion of foods these taxa were poorly adapted to consume (i.e., foods that they would not have processed efficiently), or that aspects of what we thought we knew about the functional morphology of teeth must be wrong. Evaluation of these possibilities requires a recognition that analyses based on microwear, isotopes, finite element modeling, and enamel chips and cracks each test different types of hypotheses and allow different types of inferences. Microwear and isotopic analyses are best suited to reconstructing broad dietary patterns, but are limited in their ability to falsify specific hypotheses about morphological adaptation. Conversely, finite element analysis is a tool for evaluating the mechanical basis of form‐function relationships, but says little about the frequency with which specific behaviors were performed or the particular types of food that were consumed. Enamel chip and crack analyses are means of both reconstructing diet and examining biomechanics. We suggest that current evidence is consistent with the hypothesis that certain derived australopith traits are adaptations for consuming hard foods, but that australopiths had generalized diets that could include high proportions of foods that were both compliant and tough. Am J Phys Anthropol 151:339–355, 2013.© 2013 Wiley Periodicals, Inc.     

Non-occlusal dental microwear variability in a sample of Middle and Late Pleistocene human populations from Europe and the Near East

Non-occlusal, buccal tooth microwear variability has been studied in 68 fossil humans from Europe and the Near East. The microwear patterns observed suggest that a major shift in human dietary habits and food processing techniques might have taken place in the transition from the Middle to the Late Pleistocene populations. Differences in microwear density, average length, and orientation of striations indicate that Middle Pleistocene humans had more abrasive dietary habits than Late Pleistocene populations. Both dietary and cultural factors might be responsible for the differences observed. In addition, the Middle Paleolithic Neanderthal specimens studied show a highly heterogeneous pattern of microwear when compared to the other samples considered, which is inconsistent with a hypothesis of all Neanderthals having a strictly carnivorous diet. The high density of striations observed in the buccal surfaces of several Neanderthal teeth might be indicative of the inclusion of plant foods in their diet. The buccal microwear variability observed in the Neanderthals is compatible with an overall exploitation of both plant and meat foods on the basis of food availability. A preliminary analysis of the relationship between buccal microwear density and climatic conditions prevailing in Europe during the Late Pleistocene has been attempted. Cold climatic conditions, as indicated by oxygen isotope stage data, seem to be responsible for higher densities of microwear features, whereas warmer periods could correspond to a reduced pattern of scratch density. Such a relationship would be indicative of less abrasive dietary habits, perhaps more meat dependent, during warmer periods.     

黑斑羚粪便中碳同位素揭示的食性变化

利用稳定碳同位素数据(δ13C)分析了南非克鲁格国家公园混食性黑斑羚(Aepyceros melampus)时间和空间尺度上的食性变化,验证了两个假说,即有蹄类食性变化是由生境中木本植物与草本植物的相对配比导致;降雨控制有蹄类生态。结果表明:黑斑羚的食性涵盖了精食者-粗食者采食谱系,且食性中木本与草本比例在不同月间、季节、年度和区域间存在很大变化。栖息于开放性热带稀树草原和草原中的黑斑羚通常采食比生境中更高比例的草本,但在时间尺度上并不恒定。在克鲁格北部的一个区域(Punda Maria) ,黑斑羚采食的草本比克鲁格国家公园中其它任何区域都多。与其它生境相比,在河边的黑斑羚采食草本数量更少,尤其是在食性空间变化更为明显的旱季。因此,我们的数据不支持有蹄类食性组成变化是由生境中木本与草本比例不同造成的假说,食性与降雨量间也无明显的关系。我们的结果支持草本中蛋白含量增加引起黑斑羚采食比例的增加这一模型。粪便中氮含量在时间和空间上的变化很小,揭示在可利用食物中,无论木本还是草本,黑斑羚进行选择采食以保证最好的食物质量。基于这些结果,我们认为更具体的食物选择和可利用性最适采食理论能够更好地解释这种生态学变化。     

Dental microwear features as an indicator for plant food in early hominids: A preliminary study of enamel

Studies of dental microwear have shown qualitative and quantitative differences related with specific diets of mammals. Here we report a classification of the features of microwear and the results of a comparative study of the characteristics of wear produced by plant food. Species which are mainly folivorous or herbivorous show common features useful for providing information about the diet of early hominids at Garusi and Laetoli (Tanzania), Hadar (Ethiopia) and Olduvai (Tanzania).     

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。     

Dental use wear in extinct lemurs: evidence of diet and niche differentiation

A new technique for molar use-wear analysis is applied to samples of all 16 species of extinct lemurs with known dentitions, as well as to a large comparative sample of extant primates. This technique, which relies on the light refractive properties of wear pits and scratches as seen under a standard stereoscopic microscope, has shown itself to be effective in distinguishing the diets of ungulates and extant primates. We draw dietary inferences for each of the 16 extinct lemur species in our database. There is a strong phylogenetic signal, with the Palaeopropithecidae showing use-wear signatures similar to those of the Indriidae; extinct lemurids (Pachylemur spp.) showing striking similarities to extant lemurids (except Hapalemur spp.); and Megaladapis showing similarities to Lepilemur spp. Only the Archaeolemuridae have dietary signatures unlike those of any extant lemurs, with the partial exception of Daubentonia. We conclude that the Archaeolemuridae were hard-object feeders; the Palaeopropithecidae were seed predators, consuming a mixed diet of foliage and fruit to varying degrees; Pachylemur was a fruit-dominated mixed feeder, but not a seed predator; and all Megaladapis were leaf browsers. There is no molar use wear evidence that any of the extinct lemurs relied on terrestrial foods (C4 grasses, tubers, rhizomes). This has possible implications for the role of the disappearance of wooded habitats in the extinction of lemurs.     

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.     

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.