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     

Simulation of dental microwear: Characteristic traces by opal phytoliths give clues to ancient human dietary behavior

In order to further evaluate the process of microwear formation on human dental enamel, microwear was experimentally produced by a chewing simulation with an Academic Center for Dentistry Amsterdam (ACTA) device. For this simulation, several cereal species were processed according to historical milling techniques, the experimental results of which were compared with those obtained from cereals processed after modern techniques, and also with natural microwear on early medieval human molars. Comparison of simulated microwear pits with natural microwear pits showed that the simulation led to traces which matched those found on the historical teeth in terms of both size and shape. Experimentally produced microwear pits were especially characteristic for the cereal species used in the simulations, and both pit morphology and enamel loss were a function of cereal phytolith content. Despite the high variability of phytolith size and shape, certain types are characteristic for certain cereals, which in turn are capable of producing cereal-specific microwear. This experimental approach is likely to further define ancient human dietary behavior, including food processing.     

Dust accumulation in the canopy: A potential cause of dental microwear in primates

Dental microwear researchers consider exogenous grit or dust to be an important cause of microscopic wear on primate teeth. No study to date has examined the accumulation of such abrasives on foods eaten by primates in the forest. This investigation introduces a method to collect dust at various heights in the canopy. Results from dust collection studies conducted at the primate research stations at Ketambe in Indonesia, and Hacienda La Pacifica in Costa Rica indicate that 1) grit collects throughout the canopy in both open country and tropical rain forest environments; and 2) the sizes and concentrations of dust particles accumulated over a fixed period of time differ depending on site location and season of investigation. These results may hold important implications for the interpretation of microwear on primate teeth. © 1995 Wiley-Liss, Inc.     

Tooth chipping can reveal the diet and bite forces of fossil hominins

Mammalian tooth enamel is often chipped, providing clear evidence for localized contacts with large hard food objects. Here, we apply a simple fracture equation to estimate peak bite forces directly from chip size. Many fossil hominins exhibit antemortem chips on their posterior teeth, indicating their use of high bite forces. The inference that these species must have consumed large hard foods such as seeds is supported by the occurrence of similar chips among known modern-day seed predators such as orangutans and peccaries. The existence of tooth chip signatures also provides a way of identifying the consumption of rarely eaten foods that dental microwear and isotopic analysis are unlikely to detect.     

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.     

Relationships between dental microwear and diet in Carnivora (Mammalia) — Implications for the reconstruction of the diet of extinct taxa

Food consumption causes distinct microwear patterns on teeth, especially in mammals that actively masticate food. Here we perform a microwear analysis to assess the relationships between diet and microwear features of diverse Carnivora. Our database includes approximately 230 individuals of 17 extant species having different diets. We analyse both slicing and grinding facets of M1 and m1. The proposed method is reproducible and allows the differentiation, especially on slicing facets, of microwear poles that are significantly distinct from one another. In carnivorans, the microwear features mainly result from their foraging behavior and the proportion of certain food items consumed. We applied our method to extinct taxa such as the amphicyonid Amphicyon major. The results on the m1 slicing facet indicate dietary similarities between this large Miocene predator and the extant red fox; results from the m1 grinding facet do not have equivalent in extant taxa, however.     

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。     

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.     

Form and Function of Early Neolithic Bifacial Stone Tools Reflects Changes in Land Use Practices during the Neolithization Process in the Levant

For many, climate change is no longer recognized as the primary cause of cultural changes in the Near East. Instead, human landscape degradation, population growth, socioeconomic adjustments, and conflict have been proposed as the mechanisms that shaped the Neolithic Revolution. However, as Bar-Yosef noted, even if there is chronological correlation between climate changes and cultural developments, what is important is to understand how Neolithic societies dealt with these improving or deteriorating environments. Changes in bifacial stone tools provide a framework for examining some of these interactions by focusing on changing land use practices during the Neolithization process. The results of microwear analysis of 40 bifacial artifacts from early Pre-Pottery Neolithic (EPPNB) levels at Motza in the Judean hills document changes during the PPNA-PPNB transition at the onset of the Levantine Moist Period (ca. 8000 cal B.C.) when conditions for agriculture improved. EPPNB villagers added heavy-duty axes to a toolkit they had used for carpentry and began to clear forests for fields and grazing lands. Sustainable forest management continued for the duration of the PPN until the cumulative effects of tree-felling and overgrazing seem to have led to landscape degradation at end of the Pre-Pottery Neolithic C (PPNC), when a cold, dry climatic anomaly (6600-6000 cal B.C.) may have accelerated the reduction of woodlands. Early PPNB components at sites like Motza, with data from nearly five millennia of Neolithic occupations, show how complex hunter-gatherers and early food producers were able to establish sustainable resource management systems even as climate changed, population increased, and social relations were redefined.     

郑州青台遗址新石器时代中晚期人群龋齿的统计与分析

本文通过对郑州青台遗址新石器时代中晚期91例个体、1913枚牙齿罹患龋齿的统计与分析可知,青台人群患龋率为71.43%,龋齿率为13.38%。其中,龋齿率女性高于男性,可能与女性孕期生理变化、食物选择及性别分工等有关。上颌龋齿率高于下颌,臼齿及咬合面为龋齿易患齿类及部位。通过对比可知青台人群显示出较高的龋齿罹患率,暗示该人群饮食中应包含较多的碳水化合物类食物,这可能与新石器时代中晚期黄河中游发达的旱作农业有关。此外,龋齿率在黄河及长江中、下游新石器时代农业人群中的区域性差异可能与龋病病因的复杂性和各地区不同的文化面貌、人群生活方式有关。     

Technical note: An in vitro study of dental microwear formation using the BITE Master II chewing machine

Dental microwear has been used for decades to reconstruct the diets of fossil hominins and bioarchaeological populations. The basic theory has been that hard‐brittle foods (e.g., nuts, bone) require crushing and leave pits as they are pressed between opposing cheek‐tooth surfaces, whereas soft‐tough foods (e.g., grass blades, meat) require shearing and leave scratches as they are dragged along opposing surfaces that slide past one another. However, recent studies have called into question the efficacy of microwear as an indicator of diet. One issue has been the limited number of in vitro studies providing empirical evidence for associations between microwear pattern and chewing behavior. We here describe a new study using a chewing simulator, the BITE Master II, to examine the effects of angle of approach between opposing teeth and food consistency on microwear surface texture. Results indicate that opposing teeth that approach one another: 1) perpendicular to the occlusal plane (crushing) result in pits; 2) parallel to the occlusal plane (shearing) result in striations in the direction of movement; and 3) oblique to the occlusal plane (45°) result in both striations and pits. Results further suggest that different food types and abrasive loads affect the propensity to accumulate microwear features independent of feature shapes. Am J Phys Anthropol 158:769–775, 2015. © 2015 Wiley Periodicals, Inc.     

李家沟遗址的石器工业

本文介绍河南新密李家沟遗址发现的石制品, 并进行简要讨论。该遗址包含了从旧石器时代晚期一直到新石器时代多个文化阶段的遗存。石器工业在不同文化阶段表现出不同的特点。除了旧石器晚期较早阶段的石片石器组合和旧石器时代晚期之末的细石器组合之外, 还在新石器早期文化乃至裴李岗文化阶段的文化遗存中发现数量较多的打制石器。其中最值得关注的是与典型细石器共存的磨刃石锛与陶片。通过对石制品的初步观察可知, 打制石器并不只存在于旧石器时代, 而是延续到新石器早期甚至可能裴李岗文化时期, 打制石器仍然在继续使用。这种情况说明, 李家沟以及中原地区旧、新石器时代的石器工具的变化过渡是一种逐渐变化的过程。     

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.     

First insights into the function of the sawshark rostrum through examination of rostral tooth microwear

Potential roles of the rostrum of sawsharks (Pristiophoridae), including predation and self‐defence, were assessed through a variety of inferential methods. Comparison of microwear on the surface of the rostral teeth of sawsharks and sawfishes (Pristidae) show that microwear patterns are alike and suggest that the elongate rostra in these two elasmobranch families are used for a similar purpose (predation). Raman spectroscopy indicates that the rostral teeth of both sawsharks and sawfishes are composed of hydroxyapatite, but differ in their collagen content. Sawfishes possess collagen throughout their rostral teeth whereas collagen is present only in the centre of the rostral teeth of sawsharks, which may relate to differences in ecological use. The ratio of rostrum length to total length in the common sawshark Pristiophorus cirratus was found to be similar to the largetooth sawfish Pristis pristis but not the knifetooth sawfish Anoxypristis cuspidata. Analysis of the stomach contents of P. cirratus indicates that the diet consists of demersal fishes and crustaceans, with shrimp from the family Pandalidae being the most important dietary component. No prey item showed evidence of wounds inflicted by the rostral teeth. In light of the similarities in microwear patterns, rostral tooth chemistry and diet with sawfishes, it is hypothesised that sawsharks use their rostrum in a similar manner for predation (sensing and capturing prey) and possibly for self‐defence.     

In vivo and in vitro turnover in dental microwear

Given the potential usefulness of dental microwear analyses in interpretations of archaeological and paleontological material, it is surprising how little we know about changes in individual microwear features through time. The purpose of this study was to document the turnover in primate dental microwear through in vivo dental studies of monkeys raised on different diets, and through in vitro studies of the abrasive effects of monkey chow biscuits on isolated monkey teeth. As in previous studies, epoxy replicas were prepared from dental impressions and examined under a scanning electron microscope. Results indicate that, under certain conditions, the turnover in primate dental microwear can be on the order of days, hours, or even minutes. Individual microscopic wear features can be obliterated within 24 hours on the molars of laboratory monkeys, and monkey chow biscuits can easily scratch the enamel of isolated monkey teeth. Monkeys raised on a hard diet showed more rapid turnover in dental microwear than monkeys raised on a soft diet. However, paired-sample tests revealed that, for all animals, the molar shearing facets were being abraded at a significantly slower rate than molar crushing/grinding facets. In light of these results, investigators should make every effort to use large samples in interspecific comparisons of dental microwear involving species with variable diets. Another implication of these results is that changes in dental microwear might be useful indicators of changes in oral behavior over relatively short periods of time.     

Feeding Ecology in Oligocene Mylodontoid Sloths (Mammalia,Xenarthra) as Revealed by Orthodentine Microwear Analysis

Recently, dental microwear analysis has been successfully employed to xenarthran teeth. Here, we present new data on use wear features on 16 molariforms of Orophodon hapaloides and Octodontotherium grande. These taxa count among the earliest sloths and are known from the Deseadan SALMA (late Oligocene). Modern phylogenetic analyses classify Octodontotherium and Orophodon within Mylodontoidea with whom they share lobate cheek teeth with an outer layer of cementum and a thick layer of orthodentine. Similar target areas of 100μm² were analyzed on the orthodentine surface of each tooth by stereomicroscopic microwear and by SEM microwear. Results were unlike those of extant sloths (stereomicroscopic microwear: Bradypus, Choloepus) and published data from fossil sloths (SEM microwear: Acratocnus, Megalonyx, Megatherium, Thinobadistes); thus, both approaches independently indicate a different feeding ecology for the Oligocene taxa. The unique microwear results suggest that both taxa fed on plant material with low to moderate intrinsic toughness (foliage, twigs) but also proposes intake of tougher food items (e.g., seeds). Frequent gouging of the tooth surfaces can be explained by exogenous influence on microwear, such as possible intake of abrasive grit. We suggest an unspecialized herbivorous diet for Octodontotherium and Orophodon utilizing diverse food resources of their habitat. These interpretations support the reconstruction of (1) Deseadan environments as open habitats with spreading savannas/grasslands and (2) both taxa as wide-muzzled bulk feeders at ground level.     

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.     

Assessing change in diet and biological affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura: A preliminary dental comparison of Feteira II and Bolores

Although the social and political changes accompanying the transition from the Neolithic through Copper Age, between the 4th and 3rd millennia cal BCE, in southwestern Iberia are reasonably well understood, much less is known about whether population movements and dietary changes accompanied these transformations. To address this question, human dental remains from the Middle through Late Neolithic site of Feteira II (3600–2900 cal BCE) and the Late Neolithic site of Bolores (2800–2600 cal BCE) in the Portuguese Estremadura were used to examine diet (microwear) and affinity (dental non-metrics). Microwear features were not found to be significantly different between Feteira II and Bolores, suggesting that the emergence of social complexity during this period did not result in large-scale changes in subsistence practices during the period of use at these sites. Using the Arizona State University Dental Anthropology System and supporting statistics, no significant difference between the samples from Feteira II and Bolores was observed, suggesting that no population replacement occurred between the Middle Neolithic and Late Neolithic/Copper Age. However, at Bolores there is some indication that there may have been demographic exchanges between southern Iberian and North African populations during the Late Neolithic/Copper Age.     

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.     

Potential Bark and Fruit Browsing as Revealed by Stereomicrowear Analysis of the Peculiar Clawed Herbivores Known as Chalicotheres (Perissodactyla,Chalicotherioidea)

This low magnification stereomicrowear study samples a broad range of chalicotheres (Perissodactyla, Chalicotherioidea), including basal chalicotheres and the two chalicotheriid subfamilies Schizotheriinae and Chalicotheriinae, primarily including species from North America and Europe, but also some from Asia. The schizotheriines Moropus, Tylocephalonyx, and Metaschizotherium and the chalicotheriines Anisodon and Chalicotherium are best represented. Paleodiets are interpreted via discriminant analysis, using comparison of microwear variables from fossil chalicothere teeth with those from a database of extant ungulates with known diets. The results suggest that all of the chalicotheres in the study were browsers, with no evidence of significant grass consumption. Basal chalicotheres, like basal equids, seem to have been standard fruit-dominated browsers. Stereomicrowear agrees with mesowear results by Schulz et al. (2007) and Schulz and Fahlke (2009) for Metaschizotherium bavaricum, Metaschizotherium fraasi, Anisodon grande, and Chalicotherium goldfussi in showing a highly abrasive aspect to the diet. In these species, hard food objects such as fibrous fruits, seeds, pits, and nuts may have abraded the teeth (based on high pit counts, the presence of large puncture pits, and many individuals with coarse to hypercoarse scratches). Anisodon grande and C. goldfussi, despite their relatively short, brachydont teeth, show the highest degree of abrasion within the studied sample. Moropus and Tylocephalonyx from North America show somewhat different but also abrasive microwear; in these taxa the resistant foods may have been twigs and bark (large pits common, but gouging more prevalent than puncture pits). A preliminary comparison of stereomicrowear on DP4, the deciduous upper fourth premolar, with that on molars suggests that juveniles consumed similar foods as adults but without the most abrasive elements. Some important methodological differences regarding the scoring of microwear features by different low-magnification microwear methodologies are discussed.