On the relationship of dental microwear to dental macrowear

Dental microwear analysts have demonstrated that hard diets leave numerous microscopic pits on occlusal surfaces. The relationship between occlusal pitting and gross macrowear, however, is not well known. The current study seeks to elucidate the relationship between dental microwear and macrowear by determining if microscopically pitted teeth are associated with greater expressions of macrowear. This study examined microwear and macrowear on mandibular second molars from 60 prehistoric adult Native Americans representing three dietary regimes (foraging, mixed economy, and agriculture). Initially, two dental microwear feature variables were studied: percentage of pits and mean scratch width. Standard macrowear scores ranged from 4 to 40. ANOVAs suggested that neither of the microwear variables was affected by age or sex, but age affected macrowear scores. Because of this, the sample had a balanced number of young and old adults (i.e., those below and above skeletal age 35). A Pearson's correlation showed no covariation between scratch width and the percentage of pits. Regression analysis indicated that macrowear was not a function of the percentage of pits. However, a significant positive relationship was found between dental macrowear and scratch width. A post priori test found a significant negative relationship between macrowear and the total number of scratches. It is concluded, then, that wide scratches remove more enamel and dentin than do numerous pits, although both cause dental wear. It is suggested here that the term “abrasive” be used to describe those microwear profiles that lead to heavy macrowear and have relatively wide scratches. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Dental microwear in the orthodentine of the Xenarthra (Mammalia) and its use in reconstructing the palaeodiet of extinct taxa: the case study of Nothrotheriops shastensis (Xenarthra,Tardigrada, Nothrotheriidae)

The utility of orthodentine microwear analysis as a proxy for dietary reconstruction in xenarthrans (tree sloths, armadillos) was quantitatively and statistically accessed via low‐magnification stereomicroscopy. Features such as number of scratches and pits, as well as presence of gouges, hypercoarse scratches, > four large pits, > four cross scratches, and fine, mixed or coarse scratch texture were recorded in 255 teeth from 20 extant xenarthran species. Feature patterns are consistent with scar formation through abrasional (tooth–food) and attritional (tooth–tooth) contact. Number of scratches is the most dietary diagnostic microwear variable for xenarthrans, with herbivorous sloths characterized by > ten scratches and nonherbivorous armadillos by < ten scratches. Discriminant function analysis differentiated arboreal folivores (sloths) and frugivore‐folivores (sloths) both from each other and from fossorial carnivore‐omnivores (armadillos) and insectivores (armadillos). Microwear patterns in carnivore‐omnivores and insectivores are difficult to distinguish between; armadillo microwear may reflect a fossorial lifestyle (grit consumption) rather than primary diet. Cabassous centralis is anomalous in its microwear signal relative to all other insectivores. To test the utility of orthodentine microwear analysis as an indicator of palaeodiet in extinct xenarthrans, microwear in the ground sloth Nothrotheriops shastensis was quantitatively and statistically compared to microwear in extant taxa. Microwear patterns in N. shastensis are most comparable to extant folivores based on scratch number and hierarchical cluster analysis. This strongly supports an herbivorous diet for N. shastensis that is corroborated by multiple independent lines of evidence. Thus, orthodentine microwear analysis can be used to reconstruct diet in extinct xenarthrans. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 201–222.     

MicroWeaR: A new R package for dental microwear analysis

Mastication of dietary items with different mechanical properties leaves distinctive microscopic marks on the surface of tooth enamel. The inspection of such marks (dental microwear analysis) is informative about the dietary habitus in fossil as well as in modern species. Dental microwear analysis relies on the morphology, abundance, direction, and distribution of these microscopic marks. We present a new freely available software implementation, MicroWeaR, that, compared to traditional dental microwear tools, allows more rapid, observer error free, and inexpensive quantification and classification of all the microscopic marks (also including for the first time different subtypes of scars). Classification parameters and graphical rendering of the output are fully settable by the user. MicroWeaR includes functions to (a) sample the marks, (b) classify features into categories as pits or scratches and then into their respective subcategories (large pits, coarse scratches, etc.), (c) generate an output table with summary information, and (d) obtain a visual surface‐map where marks are highlighted. We provide a tutorial to reproduce the steps required to perform microwear analysis and to test tool functionalities. Then, we present two case studies to illustrate how MicroWeaR works. The first regards a Miocene great ape obtained from through environmental scanning electron microscope, and other a Pleistocene cervid acquired by a stereomicroscope.     

Tooth microwear and premaxillary shape of an archaic antelope

Extant ungulates can be divided into three dietary categories: browsing feeders, grazing feeders, and mixed feeders. Dietary adaptations can be differentiated in extinct ruminants based upon tooth microwear analysis as well as evaluation of premaxillary morphology. Tooth microwear shows that the extinct bovid Kipsigicerus labidotus from the Miocene of Fort Ternan in Kenya (14 million years old) was most likely a grazing feeder, with mixed-feeder tendencies, while morphologically the premaxilla most closely resembles that of a mixed feeder. Because the paleoenvironment at Fort Ternan was likely to have been forested, as shown by paleosol isotopic studies, grazing in this particular ruminant evolved within a forested environment preceding the origin of savanna. □ Tooth microwear, premaxilla, Miocene, Kenya, bovid, paleodiet.     

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.     

Dental microwear and diet in Venezuelan primates.

Recent microwear analyses have demonstrated that wear patterns can be correlated with dietary differences. However, much of this work has been based on analyses of museum material where dates and locations of collection are not well known. In view of these difficulties, it would be desirable to compare microwear patterns for different genera collected from the same area at the same time. The opportunity to do this was provided by the collections of the Smithsonian Venezuelan Project (Handley, 1976), in which multiple primate genera were collected from the same humid tropical forest sites within the same month. The monkeys represent a wide range of dietary preferences, and include Saimiri, Cebus, Chiropotes, Ateles, Aotus, Pithecia, and Alouatta. As in previous microwear analyses, epoxy replicas were prepared from dental impressions, as described by Rose (1983) and Teaford and Oyen (1989). Two micrographs were taken of facet 9 on an upper second molar of each specimen. Computations and analyses were the same as described by Teaford and Robinson (1989). Results reaffirm previously documented differences in dental microwear between primates that feed on hard objects versus those that do not--with Pithecia and Alouatta at the extremes of a range of microwear patterns including more subtle differences between species with intermediate diets. The subtle microwear differences are by no means easy to document in museum samples. However, additional results suggest that 1) the width of microscopic scratches may be a poor indicator of dietary differences, 2) large and small pits may be formed differently, and 3) there are very few seasonal differences in dental microwear in the primates at these humid tropical forest sites.     

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     

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。     

Differences in Tooth Microwear of Populations of Caribou (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>, Ruminantia,Mammalia) and Implications to Ecology,Migration, Glaciations and Dental Evolution

Tooth microwear was analyzed for a large sample of wild-shot barren-ground caribou (Rangifer tarandus groenlandicus) from the Kaminuriak population of eastern Canada. This sample was compared to the microwear of specimens from three Pleistocene localities in North America (Alaska) and western Europe (Caune de l’Arago in France and Salzgitter in Germany). The results show that the extant samples from eastern Canada have seasonal variation in microwear and presumably in diet. The differences in microwear between the various seasons may reflect a cyclic migration of the population within a year. The extinct population from Alaska has extremely blunt teeth (mesowear), as blunt as those of modern zebras and bison. This observation is corroborated by the lowest number of microwear pits. The findings are untypical, as most typical caribou teeth have sharper apices, and we interpret this as an indication of a local habitat that was different with animals feeding on non-typical vegetation. The combination of Rangifer from Caune de l’Arago and Salzgitter reveals a pattern in microwear variability. The Salzgitter is interglacial and shows a greater diversity of browsing (broad spectrum on average number of pits) than the glacial Caune de l’Arago. The interglacial population from Salzgitter is interesting because it shows several different types of browsing. Collectively all the Rangifer teeth show that diet of a brachydont taxon can vary across most of the dietary morphospace of ungulates as represented by tooth microwear. The three Pleistocene samples exhibit microwear that is different from the extant population in question. This observation implies that the recent diet of Rangifer has changed from the typical caribou diet in the past. This indicates dietary change within a species. This is important because it represents dietary evolution without changes in tooth morphology.     

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.     

Direct Comparisons of 2D and 3D Dental Microwear Proxies in Extant Herbivorous and Carnivorous Mammals

The analysis of dental microwear is commonly used by paleontologists and anthropologists to clarify the diets of extinct species, including herbivorous and carnivorous mammals. Currently, there are numerous methods employed to quantify dental microwear, varying in the types of microscopes used, magnifications, and the characterization of wear in both two dimensions and three dimensions. Results from dental microwear studies utilizing different methods are not directly comparable and human quantification of wear features (e.g., pits and scratches) introduces interobserver error, with higher error being produced by less experienced individuals. Dental microwear texture analysis (DMTA), which analyzes microwear features in three dimensions, alleviates some of the problems surrounding two-dimensional microwear methods by reducing observer bias. Here, we assess the accuracy and comparability within and between 2D and 3D dental microwear analyses in herbivorous and carnivorous mammals at the same magnification. Specifically, we compare observer-generated 2D microwear data from photosimulations of the identical scanned areas of DMTA in extant African bovids and carnivorans using a scanning white light confocal microscope at 100x magnification. Using this magnification, dental microwear features quantified in 2D were able to separate grazing and frugivorous bovids using scratch frequency; however, DMTA variables were better able to discriminate between disparate dietary niches in both carnivorous and herbivorous mammals. Further, results demonstrate significant interobserver differences in 2D microwear data, with the microwear index remaining the least variable between experienced observers, consistent with prior research. Overall, our results highlight the importance of reducing observer error and analyzing dental microwear in three dimensions in order to consistently interpret diets accurately.     

Brief communication: intertooth and intrafacet dental microwear variation in an archaeological sample of modern humans from the Jordan Valley

Dental microwear was recorded in a Bronze-Iron Age (3570-3000 BP) sample of modern humans recovered from Tell es-Sa'idiyeh in the Jordan Valley. Microwear patterns were compared between mandibular molars, and between the upper and lower part of facet 9. The comparison revealed a greater frequency of pits and shorter scratches on the second and third molars, compared to the first. Pit frequency also increased on the lower part of the facet on the first molar, compared to the upper part. These results support previous calls for standardization when selecting a molar type for a diet-microwear study. Otherwise the microwear variations along the tooth row could mask any diet-microwear correlations. The results also suggest that there may be a need to choose a consistent location on a facet in order to enhance comparability among studies.     

Molar Microwear of Narrow-Headed Vole (<Emphasis Type="Italic">Microtus gregalis</Emphasis> Pall., 1779) Depending on the Feed Abrasiveness

In the narrow-headed vole, enamel microwear of the first mandibular molar (of the protoconid and entoconid anterior enamel wall) was studied under the laboratory conditions and at the fixed feed composition. The classic parameters and the area of the enamel prism lesion were taken into account. The enamel lesion patterns caused by the tooth–tooth and tooth–food interactions have been determined. Differences were found between the voles kept on feed with different abrasive properties, as well as between the lingual and buccal conids of the first mandibular molar. In the Microtus species, the ratio of micro-lesions (pits and scratches) did not depend on the abrasive properties of the feed consumed. The extent of preservation of the enamel contour anterior edge depended on the feed composition and could be used as an indicator for indirect evaluation of the Microtus species diet.     

Dietary Abrasiveness Is Associated with Variability of Microwear and Dental Surface Texture in Rabbits

Dental microwear and 3D surface texture analyses are useful in reconstructing herbivore diets, with scratches usually interpreted as indicators of grass dominated diets and pits as indicators of browse. We conducted feeding experiments with four groups of rabbits (Oryctolagus cuniculus) each fed a different uniform, pelleted diet (lucerne, lucerne & oats, grass & oats, grass). The lowest silica content was measured in the lucerne and the highest in the grass diet. After 25 weeks of exposure to the diets, dental castings were made of the rabbit''s lower molars. Occlusal surfaces were then investigated using dental microwear and 3D areal surface texture analysis. In terms of traditional microwear, we found our hypothesis supported, as the grass group showed a high proportion of (long) “scratches” and the lucerne group a high proportion of “pits”. Regardless of the uniform diets, variability of microwear and surface textures was higher when silica content was low. A high variability in microwear and texture analysis thus need not represent dietary diversity, but can also be related to a uniform, low-abrasion diet. The uniformity or variability of microwear/texture analysis results thus might represent varying degrees of abrasion and attrition rather than a variety of diet items per se.     

Dietary adaptations in an ungulate community from the late Pliocene of Greece

The dietary morphological methods of mesowear and microwear were applied to ungulates of the late Pliocene fauna of Sésklo (Thessaly, Greece). The results provide evidence for the predominance of open grassland in the area, as the most common species, Equus stenonis, was a strict grazer. The rare cervid cf. Croizetoceros ramosus was the only browser. The antelopes (genera Gazella and Gazellospira) yielded discrepant microwear and mesowear results. This is interpreted as an indication of regional or seasonal dietary resource differentiation, inferring that the antelopes were probably mixed feeders that grazed occasionally or periodically.     

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.     

Seasonal or ecological differences in diet and molar microwear in Cebus nigrivittatus

Dental microwear analyses have raised new hopes and questions for functional morphologists. One hope is that analyses will allow insights into subtle dietary differences of extinct species. One major question is whether seasonal and/or habitat differences in dental microwear are reliably detectable. The extensive collections of Cebus nigrivittatus obtained by the Smithsonian Venezuelan Project allowed us to examine seasonal and habitat differences in dental microwear. Specimens were collected from three distinct ecological life zones that are distinguished by both the amount of rainfall and its seasonability. Environmental variation is generally correlated with variation in resource availability which, in C. nigrivittatus, affects diet. Published field studies indicate that these animals depend more on dry hard fruit and chitinous invertebrates during drier times and succulent fruits and caterpillars during wetter times of the year. As in previous microwear analyses, epoxy replicas were prepared from dental impressions, and the replicas were examined under a scanning electron microscope. Two micrographs were taken of facet 9 on M² of each specimen. Mean values for the proportion of pits (vs. scratches), pit wdith, and scratch width were computed for each of 62 individuals and compared between ecological zones and collecting dates by using a multiple comparison test. Results indicate that, while seasonal differences in molar microwear in C. nigrivittatus are perhaps reliably detectable, 1) they are small in magnitude, 2) they are only detectable in certain ecological life zones, and 3) they are not of the order of magnitude that will obscure major interspecific differences in molar microwear such as those between C. apella and C. nigrivittatus.     

Microwear and morphology: functional relationships between human dental microwear and the mandible

Microscopic pits and scratches form on teeth during chewing, but the extent to which their formation is influenced by mandibular morphology is unknown. Digitized micrographs of the base of facet nine of the first, second, and third mandibular molar were used to record microwear features from an archaeological sample of modern humans recovered from Semna South in northern Sudan (n=38; 100 BC to AD 350). Microwear patterns of the molar row are correlated with mandibular corpus width and depth, and with mandibular length. Variations in shear and compression at the base of facet nine during chewing were inferred. It may be that some correlations between microwear and mandibular morphology are predictable, reflecting similar aspects of masticatory loading, though the full extent of the relationship remains to be resolved.     

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.     

The Bovidae of Dorn-Dürkheim 1, Germany (Turolian age)

Three bovid species are present at Dorn-Dürkheim 1. The overwhelmingly abundant species is a boselaphine, Miotragocerus sp., with smaller and less advanced teeth than Tragoportax amalthea from Pikermi, Greece. Miotragocerus was present in the latest middle Miocene and Vallesian of western and central Europe and survived into the Turolian. The other two bovid species remain enigmatic and of uncertain tribal affiliation. Each is represented by very few teeth, none of them associated. One species is larger and the other smaller than the Miotragocerus sp. Mesowear analysis of M²s and M³s was used to investigate the dietary regime of Miotragocerus sp. from Dorn-Dürkheim 1. Miotragocerus was found to be linked to extant browsing ungulates close to the transition to mixed feeders. The percentage of abrasive food components like grass in the diet of this species was probably close to 10%. This ratio suggests Miotragocerus is intermediate between the two hipparionine horse species of this early Turolian (MN11) palaeoenvironment.