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.

The effect of prism orientation on the indentation testing of human molar enamel.

Recent nanoindentation studies have demonstrated that the hardness and Young's modulus of human molar enamel decreases by more than 50% on moving from the occlusal surface to the dentine-enamel junction on cross-sectional samples. Possible sources of these variations are changes in local chemistry, microstructure, and prism orientation. This study investigates the latter source by performing nanoindentation tests at two different orientations relative to the hydroxyapatite prisms: parallel and perpendicular. A single sample volume was tested in order to maintain a constant chemistry and microstructure. The resulting data show very small differences between the two orientations for both hardness and Young's modulus. The 1.5-3.0% difference is significantly less than the standard deviations found within the data set. Thus, the variations in hardness and Young's modulus on cross-sectional samples of human molar are attributed to changes in local chemistry (varying levels of mineralization, organic matter, and water content) and changes in microstructure (varying volume fractions of inorganic crystals and organic matrix). The impact of prism orientation on mechanical properties measured at this scale by nanoindentation appears to be minimal.

Mechanical defenses in leaves eaten by Costa Rican howling monkeys (Alouatta palliata).

Primate species often eat foods of different physical properties. This may have implications for tooth structure and wear in those species. The purpose of this study was to examine the mechanical defenses of leaves eaten by Alouatta palliata from different social groups at Hacienda La Pacifica in Costa Rica. Leaves were sampled from the home-ranges of groups living in different microhabitats. Specimens were collected during the wet and dry seasons from the same tree, same plant part, and same degree of development as those eaten by the monkeys. The toughness of over 300 leaves was estimated using a scissors test on a Darvell mechanical tester. Toughness values were compared between social groups, seasons, and locations on the leaves using ANOVA. Representative samples of leaves were also sun-dried for subsequent scanning electron microscopy and energy dispersive x-ray (EDX) analyses in an attempt to locate silica on the leaves. Both forms of mechanical defense (toughness and silica) were found to be at work in the plants at La Pacifica. Fracture toughness varied significantly by location within single leaves, indicating that measures of fracture toughness must be standardized by location on food items. Monkeys made some food choices based on fracture toughness by avoiding the toughest parts of leaves and consuming the least tough portions. Intergroup and seasonal differences in the toughness of foods suggest that subtle differences in resource availability can have a significant impact on diet and feeding in Alouatta palliata. Intergroup differences in the incidence of silica on leaves raise the possibility of matching differences in the rates and patterns of tooth wear.

Is bruxism severity a predictor of oral splint efficacy in patients with myofascial face pain?

Both the efficacy and mechanism of any effect of oral splint therapy for patients with temporomandibular disorders (TMDs) are a matter of controversy. To address these issues, this study tested the hypothesis that oral splints produce the most marked pain relief for those TMD patients with myofascial face pain (MFP) who also brux (i.e. grind or clench) more than other MFP patients. In a 6-week randomized controlled clinical trial, 52 women with MFP were randomly assigned to receive either a full-coverage hard acrylic splint or a palatal-only splint. Bruxism was assessed both by self-report and by an objective assessment of molar microwear changes over a 2-week period prior to the start of the trial. Tested across multiple outcome measures, results indicated that those receiving the full-coverage splint had marginally better improvement on some pain-related measures than those receiving the palatal splint, but severity of bruxism did not moderate the therapeutic effect of the full-coverage splint. These findings strongly argue against the belief that oral splints reduce MFP by reducing bruxism and raise questions about the importance of bruxism in the maintenance of MFP.

Molar microwear of subfossil lemurs: improving the resolution of dietary inferences.

In this study we use molar microwear analyses to examine the trophic distinctions among various taxa of Malagasy subfossil lemurs. High resolution casts of the teeth of Megaladapis, Archaeolemur, Palaeopropithecus, Babakotia, and Hadropithecus were examined under a scanning electron microscope. Megaladapis was undoubtedly a browsing folivore, but there are significant differences between species of this genus. However, dietary specialists appear to be the exception; for example, Palaeopropithecus and Babakotia probably supplemented their leaf-eating with substantial amounts of seed-predation, much like modern indrids. Hadropithecus was decidedly not like the modern gelada baboon, but probably did feed on hard objects. Evidence from microwear and coprolites suggests that Archaeolemur probably had an eclectic diet that differed regionally and perhaps seasonally. Substantial trophic diversity within Madgascar's primate community was diminished by the late Quaternary extinctions of the large-bodied species (>9 kg).

Error rates in dental microwear quantification using scanning electron microscopy.

There is a degree of correlation between dietary habits and dental microwear in extant primates, and this has enabled inferences to be made about prehistoric diets. Several techniques have been used to quantify microwear, but the comparability of results derived from each has not been demonstrated. Moreover, neither intra- nor interobserver error rates in microwear quantification have been documented to date. We here assess intra- and interobserver error using Microware 4.0, and evaluate intertechnique comparability using the three methods that have been most widely employed in the field. This study documents an overall intraobserver error rate of about 7%, and an overall interobserver error rate of some 9%. Both intra- and interobserver error appears to be influenced substantially by the nature of the micrograph being measured. In no instance did the results obtained by different observers using Microware 4.0 differ significantly, and there was a reasonable degree of interobserver consistency in the rank ordering of micrographs in relation to any given parameter. The results obtained through the use of different quantitative techniques differed significantly, with an overall intertechnique error rate of approximately 19%. Several variables, including differences in magnification factor, scanning electron microscope kV settings, and specimen-detector relationships undoubtedly contribute to the differences among the three methods, but we were not able to assess their relative importance. Microwear quantification permits distinctions between broad dietary categories, but the margin of intra- and interobserver error should be taken into account when defining pattern differences between populations (or species) or when documenting seasonally mitigated differences within a taxon. In view of the error introduced by the use of different methods, we suggest that a consistent technique, such as offered by the Microware software package, be adopted by current researchers to establish a common microwear database.

Nanoindentation mapping of the mechanical properties of human molar tooth enamel.

The mechanical behavior of dental enamel has been the subject of many investigations. Initial studies assumed that it was a more or less homogeneous material with uniform mechanical properties. Now it is generally recognized that the mechanical response of enamel depends upon location, chemical composition, and prism orientation. This study used nanoindentation to map out the properties of dental enamel over the axial cross-section of a maxillary second molar (M(2)). Local variations in mechanical characteristics were correlated with changes in chemical content and microstructure across the entire depth and span of a sample. Microprobe techniques were used to examine changes in chemical composition and scanning electron microscopy was used to examine the microstructure. The range of hardness (H) and Young's modulus (E) observed over an individual tooth was found to be far greater than previously reported. At the enamel surface H>6GPa and E>115GPa, while at the enamel-dentine junction H<3GPa and E<70GPa. These variations corresponded to the changes in chemistry, microstructure, and prism alignment but showed the strongest correlations with changes in the average chemistry of enamel. For example, the concentrations of the constituents of hydroxyapatite (P(2)O(5) and CaO) were highest at the hard occlusal surface and decreased on moving toward the softer enamel-dentine junction. Na(2)O and MgO showed the opposite trend. The mechanical properties of the enamel were also found to differ from the lingual to the buccal side of the molar. At the occlusal surface the enamel was harder and stiffer on the lingual side than on the buccal side. The interior enamel, however, was softer and more compliant on the lingual than on the buccal side, a variation that also correlated with differences in average chemistry and might be related to differences in function.

A review of interproximal wear grooves on fossil hominin teeth with new evidence from Olduvai Gorge.

Interproximal (approximal) grooves at the cementum-enamel junction of premolar and molar teeth have been observed in a broad range of human ancestors and related extinct species from 1.84 million years ago to the present. Many hypotheses have been presented to explain the aetiology of these grooves, though their form and positioning are most consistent with tooth-picking behaviours. This paper reviews occurrences of interproximal grooves in the cheek teeth of modern and fossil humans, evaluates hypotheses on their cause, and reports on a previously undescribed groove found in OH 60, a molar tooth from Olduvai Gorge. This specimen is among the earliest to show such grooving, and is most likely attributable to Homo erectus. It is concluded that, because interproximal grooves have been observed only on Homo teeth, they probably reflect a behaviour or behaviours unique to that genus.

Diet and the evolution of the earliest human ancestors.

Over the past decade, discussions of the evolution of the earliest human ancestors have focused on the locomotion of the australopithecines. Recent discoveries in a broad range of disciplines have raised important questions about the influence of ecological factors in early human evolution. Here we trace the cranial and dental traits of the early australopithecines through time, to show that between 4.4 million and 2.3 million years ago, the dietary capabilities of the earliest hominids changed dramatically, leaving them well suited for life in a variety of habitats and able to cope with significant changes in resource availability associated with long-term and short-term climatic fluctuations.

Dental microwear and microstructure in early oligocene primates from the Fayum, Egypt: implications for diet.

Textbook descriptions usually portray the Fayum anthropoideans as frugivores, with Parapithecus grangeri including a folivorous component in its diet and Apidium a component of hard-object feeding. Recent work with modern mammals has shown that analyses of both dental microwear and dental microstructure may yield insights into diet and tooth use. The purpose of this study was to combine these two techniques to gain a better perspective on the paleobiology of the Fayum higher primates. Dental microwear analyses involved the use of high resolution epoxy casts of Aegyptopithecus, Parapithecus, and Apidium housed in the Duke University Primate Center. Scanning electron micrographs were taken at x500, and all microwear features in each micrograph were digitized. For microstructure analyses, molar teeth were sectioned in a variety of planes, lightly etched, and photographed in the SEM. Results of the dental microwear analyses indicate that the three Fayum anthropoideans all clustered with modern primate frugivores but that there were also significant differences between Aegyptopithecus and the other two Fayum genera. By contrast, dental microstructure analyses showed important differences between Apidium and the other two genera. The reason for these differences probably lies in a combination of body size and dietary differences, with Aegyptopithecus occasionally feeding on hard objects and Apidium maximizing wear resistance through a unique emphasis of radial (rather than decussating) enamel.

Preliminary examination of non-occlusal dental microwear in anthropoids: implications for the study of fossil primates.

Most studies of microscopic wear on non-human primate teeth have focused on the occlusal surfaces of molars. Recent analyses of the buccal surfaces of human cheek teeth have demonstrated an association between diet and dental microwear on the these surfaces as well. In the current study, we examine microwear on both the buccal and lingual surfaces of non-human primate molars to assess the potential of these surfaces to reveal information concerning anthropoid feeding behaviors. We compare frequency of microwear occurrence in 12 extant and 11 fossil anthropoid species. Among the living primates, the occurrence of microwear on nonocclusal surfaces appears to relate to both diet and degree of terrestriality. The implications of this research for the inference of feeding behaviors and substrate use in fossil cercopithecoids are discussed.

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.

Significance of silica in leaves to long-tailed macaques (Macaca fascicularis).

Leaves of two plant species eaten by Macaca fascicularis in Bukit Timah Nature Reserve, Singapore, were collected and colour-tested. Leaves matching those eaten by M. fascicularis were examined by energy-dispersive X-ray micro-analysis. The leaves of Streblus elongatus (Moraceae) and Gluta wallichii (Anacardiaceae), together forming 19.6% of the leaf diet of the macaques, contained silica. In G. wallichii, this in the base of hairs that project from the underside of the leaf, whereas S. elongatus leaves have short sharp siliceous trichomes which are densely packed on the undersurface of leaf veins. We predict from an indentation analysis that chewing on the latter species could cause dental microwear at low occlusal forces. The leaves are reportedly common in the diet of three other primate species in peninsular Malaysia and the finding could have general significance for studies of dental wear.

Radiographic estimation of long bone cross-sectional geometric properties.

Because of their biomechanical significance, cross-sectional geometric properties of long bone diaphyses (areas, second moments of area) have been increasingly used in a number of form/function studies, e.g., to reconstruct body mass or locomotor mode in fossil primates or to elucidate allometric scaling relationships among extant taxa. In the present study, we test whether these biomechanical section properties can be adequately estimated using biplanar radiographs, as compared to calculations of the same properties from computer digitization of cross-sectional images. We are particularly interested in smaller animals, since the limb bone cortices of these animals may not be resolvable using other alternative noninvasive techniques (computed tomography). The test sample includes limb bones of small (25-5,000 g) relatively generalized quadrupedal mammals--mice, six species of squirrels, and Macaca fascicularis. Results indicate that biplanar radiographs are reasonable substitutes for digitized cross-sectional images for deriving areas and second moments of area of midshaft femora and humeri of mammals in this size range. Potential application to a variety of questions relating to mechanical loading patterns in such animals is diverse.

Partial skeleton of Proconsul nyanzae from Mfangano Island, Kenya.

A partial skeleton attributed to Proconsul nyanzae (KNM-MW 13142) is described. The fossils were found at a site on Mfangano Island, Kenya, which dates to 17.9 +/- .1 million years ago. KNM-MW 13142 consists of six partial vertebrae (T12-S1), a nearly complete hipbone, most of the right femur and left femoral shaft, a fragmentary tibia and fibula, and a nearly complete talus and calcaneus. This skeleton provides the first pelvic fossil known for any East African Miocene hominoid. The new Proconsul specimen is compared to a large sample of extant anthropoids to determine its functional and phylogenetic affinities. In most aspects of its anatomy, KNM-MW 13142 closely resembles nonhominoid anthropoids. This individual had a long, flexible spine, narrow torso, and habitually pronograde posture, features characteristic of most extant monkeys. Evidence of spinal musculature suggests a generalized condition intermediate between that of cercopithecoids and hylobatids. The hindlimb of KNM-MW 13142 exhibits relatively mobile hip and ankle joints, with structural properties of the femur like those of hominoids. This mix of features implies a pattern of posture and locomotion that is unlike that of any extant primate. Many aspects of the Proconsul nyanzae locomotor skeleton may represent the primitive catarrhine condition.

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.

Dental microwear in live, wild-trapped Alouatta palliata from Costa Rica.

One problem with dental microwear analyses of museum material is that investigators can never be sure of the diets of the animals in question. An obvious solution to this problem is to work with live animals. Recent work with laboratory primates has shown that high resolution dental impressions can be obtained from live animals. The purpose of this study was to use similar methods to begin to document rates and patterns of dental microwear for primates in the wild. Thirty-three Alouatta palliata were captured during the wet season at Hacienda La Pacifica near Canas, Costa Rica. Dental impressions were taken and epoxy casts of the teeth were prepared using the methods of Teaford and Oyen (1989a). Scanning electron micrographs were taken of the left mandibular second molars at magnifications of 200x and 500x. Lower magnification images were used to calculate rates of wear, and higher magnification images were used to measure the size and shape of microwear features. Results indicate that, while basic patterns of dental microwear are similar in museum samples and samples of live, wild-trapped animals of the same species, ecological differences between collection locales may lead to significant intraspecific differences in dental microwear. More importantly, rates of microwear provide the first direct evidence of differences in molar use between monkeys and humans.

A new approach to the study of tooth wear.

Human tooth wear occurs so slowly that traditionally it has needed months or years to be measurable. This study showed that microscopic changes in wear patterns on human teeth could be detected in a matter of days and could be used as indicators of rates of wear. Thus, daily or weekly changes in rates of wear can be documented for specific locations on teeth. For instance, through this new approach, rates of wear of human teeth were shown to be significantly slower than rates of wear of the teeth of laboratory monkeys raised on hard or soft diets. Similar techniques may ultimately be used to monitor subtle changes in tooth use--including those associated with growth and development and those occurring in response to various dental clinical procedures.