The impact of fallback foods on wild ring-tailed lemur biology: A comparison of intact and anthropogenically disturbed habitats

Fallback foods are often viewed as central in shaping primate morphology, and influencing adaptive shifts in hominin and other primate evolution. Here we argue that fruit of the tamarind tree (Tamarindus indica) qualifies as a fallback food of ring-tailed lemurs (Lemur catta) at the Beza Mahafaly Special Reserve (BMSR), Madagascar. Contrary to predictions that fallback foods may select for dental and masticatory morphologies adapted to processing these foods, consumption of tamarind fruit by these lemurs leaves a distinct pattern of dental pathology among ring-tailed lemurs at BMSR. Specifically, the physical and mechanical properties of tamarind fruit likely result in a high frequency of severe tooth wear, and subsequent antemortem tooth loss, in this lemur population. This pattern of dental pathology is amplified among lemurs living in disturbed areas at Beza Mahafaly, resulting from a disproportionate emphasis on challenging tamarind fruit, due to few other fruits being available. This is in part caused by a reduction in ground cover and other plants due to livestock grazing. As such, tamarind trees remain one of the few food resources in many areas. Dental pathologies are also associated with the use of a nonendemic leaf resource Argemone mexicana, an important food during the latter part of the dry season when overall food availability is reduced. Such dental pathologies at Beza Mahafaly, resulting from the use or overemphasis of fallback foods for which they are not biologically adapted, indicate that anthropogenic factors must be considered when examining fallback foods. Am J Phys Anthropol 140:671–686, 2009. © 2009 Wiley-Liss, Inc.     

Interpreting food processing through dietary mechanical properties: a Lemur catta case study

Knowledge of dietary mechanical properties can be informative about physical consequences to consumers during ingestion and mastication. In this article, we examine how Tamarindus indica fruits can affect dental morphology in a population of ring-tailed lemurs (Lemur catta) at Beza Mahafaly special reserve in southwestern Madagascar. Ring-tailed lemurs in tamarind dominated gallery forests exhibit extreme wear and tooth loss on their postcanine dentition that has been related to processing T. indica fruits. We measured and compared mechanical properties of individual food parts in the diet of ring-tailed lemurs in different seasons in 1999-2000, 2008, and 2010. Fracture toughness, hardness, and modulus of foods were measured with a portable mechanical tester. The ripe fruits of T. indica are indeed the toughest and hardest foods ingested by the lemurs. In addition, they are among the largest foods consumed, require high numbers of ingestive bites to process, and are the most frequently eaten by volume. During controlled cutting tests of the ripe fruit shell, multiple runaway side cracks form alongside the cut. Similarly, the lemurs repeatedly bite the ripe shell during feeding and thereby introduce multiple cracks that eventually fragment the shell. Studies of enamel microstructure (e.g., Lucas et al.: BioEssays 30 (2008) 374-385; Campbell et al., 2011) advance the idea that the thin enamel of ring-tailed lemur teeth is susceptible to substantial micro-cracking that rapidly erodes the teeth. We conclude that micro-cracking from repeated loads, in combination with the mechanical and physical properties of the fruit, is primarily responsible for the observed dental damage.     

A comparison of salivary pH in sympatric wild lemurs (Lemur catta and Propithecus verreauxi) at Beza Mahafaly Special Reserve, Madagascar

Chemical deterioration of teeth is common among modern humans, and has been suggested for some extinct primates. Dental erosion caused by acidic foods may also obscure microwear signals of mechanical food properties. Ring-tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar, display frequent severe tooth wear and subsequent tooth loss. In contrast, sympatric Verreaux's sifaka display far less tooth wear and infrequent tooth loss, despite both species regularly consuming acidic tamarind fruit. We investigated the potential impact of dietary acidity on tooth wear, collecting data on salivary pH from both species, as well as salivary pH from ring-tailed lemurs at Tsimanampesotse National Park, Madagascar. We also collected salivary pH data from ring-tailed lemurs at the Indianapolis Zoo, none of which had eaten for at least 12 hr before data collection. Mean salivary pH for the BMSR ring-tailed lemurs (8.098, n=41, SD=0.550) was significantly more alkaline than Verreaux's sifaka (7.481, n=26, SD=0.458). The mean salivary pH of BMSR (8.098) and Tsimanampesotse (8.080, n=25, SD=0.746) ring-tailed lemurs did not differ significantly. Salivary pH for the Indianapolis Zoo sample (8.125, n=16, SD=0.289) did not differ significantly from either the BMSR or Tsimanampesotse ring-tailed lemurs, but was significantly more alkaline than the BMSR Verreaux's sifaka sample. Regardless of the time between feeding and collection of pH data (from several minutes to nearly 1 hr), salivary pH for each wild lemur was above the "critical" pH of 5.5, below which enamel demineralization occurs. Thus, the high pH of lemur saliva suggests a strong buffering capacity, indicating the impact of acidic foods on dental wear is short-lived, likely having a limited effect. However, tannins in tamarind fruit may increase friction between teeth, thereby increasing attrition and wear in lemurs. These data also suggest that salivary pH varies between lemur species, corresponding to broad dietary categories.     

The impact of dental impairment on ring-tailed lemur food processing performance

During mastication, foods are reduced into particles suitable for swallowing and digestion. Smaller particles possess a greater surface area per unit of volume on which digestive enzymes and bacteria may work than relatively larger particles, and are thus more readily digested. As dental morphology facilitates the breakdown of diets with specific mechanical properties, extensive dental wear and/or tooth loss may impede an individual's ability to break down and exploit foods. We present data demonstrating a relationship between dental impairment and particle size in 43 fecal samples from 33 ring-tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar. All fecal samples were sifted through three sieves of decreasing size (11.2 mm, 4.75 mm, and 1.0 mm). The resulting fraction in each sieve was then weighed and assessed in relation to individual dental impairment status. With increasing wear, the percentage of each sample within the 1.0 mm sieve decreases, whereas that in the 11.2 mm sieve increases with increasing postcanine wear, although these effects are not present when limited to individuals without tooth loss. Individuals with tooth loss also demonstrate larger proportions of fecal material 1.0-4.75 mm in size. Dental impairment results in larger food particles and potentially less efficient utilization of foods. When fecal material was examined by leaf vs. fruit content, individuals with tooth loss demonstrated reduced proportions of fruit in the 1.0 mm and 11.2 mm sieves. These data suggest individuals with tooth loss consume less fruit than those without loss, potentially reflecting a reduced ability to process tamarind fruit, a key fallback resource at BMSR.     

Nanoindentation of lemur enamel: an ecological investigation of mechanical property variations within and between sympatric species

The common morphological metrics of size, shape, and enamel thickness of teeth are believed to reflect the functional requirements of a primate's diet. However, the mechanical and material properties of enamel also contribute to tooth function, yet are rarely studied. Substantial wear and tooth loss previously documented in Lemur catta at the Beza Mahafaly Special Reserve suggests that their dental morphology, structure, and possibly their enamel are not adapted for their current fallback food (the mechanically challenging tamarind fruit). In this study, we investigate the nanomechanical properties, mineralization, and microstructure of the enamel of three sympatric lemur species to provide insight into their dietary functional adaptations. Mechanical properties measured by nanoindentation were compared to measurements of mineral content, prism orientation, prism size, and enamel thickness using electron microscopy. Mechanical properties of all species were similar near the enamel dentin junction and variations correlated with changes in microstructure (e.g., prism size) and mineral content. Severe wear and microcracking within L. catta's enamel were associated with up to a 43% reduction in nanomechanical properties in regions of cracking versus intact enamel. The mechanical and material properties of L. catta's enamel are similar to those of sympatric folivores and suggest that they are not uniquely mechanically adapted to consume the physically challenging tamarind fruit. An understanding of the material and mechanical properties of enamel is required to fully elucidate the functional and ecological adaptations of primate teeth.     

Tamarind tree seed dispersal by ring-tailed lemurs

In Madagascar, the gallery forests of the south are among the most endangered. Tamarind trees (Tamarindus indica) dominate these riverine forests and are a keystone food resource for ring-tailed lemurs (Lemur catta). At Berenty Reserve, the presence of tamarind trees is declining, and there is little recruitment of young trees. Because mature tamarinds inhibit growth under their crowns, seeds must be dispersed away from adult trees if tree recruitment is to occur. Ring-tailed lemurs are likely seed dispersers; however, because they spend much of their feeding, siesta, and sleeping time in tamarinds, they may defecate a majority of the tamarind seeds under tamarind trees. To determine whether they disperse tamarind seeds away from overhanging tamarind tree crowns, we observed two troops for 10 days each, noted the locations of feeding and defecation, and collected seeds from feces and fruit for germination. We also collected additional data on tamarind seedling recruitment under natural conditions, in which seedling germination was abundant after extensive rain, including under the canopy. However, seedling survival to 1 year was lower when growing under mature tamarind tree crowns than when growing away from an overhanging crown. Despite low fruit abundance averaging two fruits/m³ in tamarind crowns, lemurs fed on tamarind fruit for 32% of their feeding samples. Daily path lengths averaged 1,266 m, and lemurs deposited seeds throughout their ranges. Fifty-eight percent of the 417 recorded lemur defecations were on the ground away from overhanging tamarind tree crowns. Tamarind seeds collected from both fruit and feces germinated. Because lemurs deposited viable seeds on the ground away from overhanging mature tamarind tree crowns, we conclude that ring-tailed lemurs provide tamarind tree seed dispersal services.     

Variation in dental wear and tooth loss among known‐aged,older ring‐tailed lemurs (Lemur catta): a comparison between wild and captive individuals

Tooth wear is generally an age‐related phenomenon, often assumed to occur at similar rates within populations of primates and other mammals, and has been suggested as a correlate of reduced offspring survival among wild lemurs. Few long‐term wild studies have combined detailed study of primate behavior and ecology with dental analyses. Here, we present data on dental wear and tooth loss in older (>10 years old) wild and captive ring‐tailed lemurs (Lemur catta). Among older ring‐tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar (n=6), the percentage of severe dental wear and tooth loss ranges from 6 to 50%. Among these six individuals, the oldest (19 years old) exhibits the second lowest frequency of tooth loss (14%). The majority of captive lemurs at the Indianapolis Zoo (n=7) are older than the oldest BMSR lemur, yet display significantly less overall tooth wear for 19 of 36 tooth positions, with only two individuals exhibiting antemortem tooth loss. Among the captive lemurs, only one lemur (a nearly 29 year old male) has lost more than one tooth. This individual is only missing anterior teeth, in contrast to lemurs at BMSR, where the majority of lost teeth are postcanine teeth associated with processing specific fallback foods. Postcanine teeth also show significantly more overall wear at BMSR than in the captive sample. At BMSR, degree of severe wear and tooth loss varies in same aged, older individuals, likely reflecting differences in microhabitat, and thus the availability and use of different foods. This pattern becomes apparent before “old age,” as seen in individuals as young as 7 years. Among the four “older” female lemurs at BMSR, severe wear and/or tooth loss do not predict offspring survival. Am. J. Primatol. 72:1026–1037, 2010. © 2010 Wiley‐Liss, Inc.     

Book review: Advances in Human Palaeopathology

Severe dental wear and tooth loss is often assumed to impede the processing, breakdown, and energetic conversion of food items, thereby negatively impacting individual health, reproduction, and survival. Ring‐tailed lemurs at the Beza Mahafaly Special Reserve demonstrate exceptionally high frequencies of severe dental wear and antemortem tooth loss, yet often survive multiple years with these impairments. To test the hypothesis that these lemurs mitigate tooth loss through behavioral adjustments, we collected 191 h of observational data from 16 focal subjects, eight without tooth loss and eight with between 3% and 44% loss. These data indicate dentally‐impaired ring‐tailed lemurs show compensatory behaviors consistent with the demands of living in a social group. During early afternoon (12:00–14:30 h) individuals with loss showed trends towards higher frequencies of foraging and grooming, while individuals without loss rested significantly more often. Individuals with >10% loss (n = 7) showed higher frequencies of feeding, foraging, and grooming, and lower frequencies of resting during this period than individuals with <10% loss (n = 9). Individuals with tooth loss maintained relatively higher levels of feeding and foraging throughout the day. These individuals licked tamarind fruit at higher frequencies, likely spending more time softening it before ingestion. These individuals did not demonstrate longer feeding bouts overall, although bouts involving tamarinds were significantly longer. Individuals with marked toothcomb wear engaged in higher rates of certain types of allogrooming, demonstrating that social behaviors are used to compensate for reduced grooming efficiency. These data have implications for interpreting behavioral responses to dental impairment in the fossil record. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

What is dental ecology?

Teeth have long been used as indicators of primate ecology. Early work focused on the links between dental morphology, diet, and behavior, with more recent years emphasizing dental wear, microstructure, development, and biogeochemistry, to understand primate ecology. Our study of Lemur catta at the Beza Mahafaly Special Reserve, Madagascar, has revealed an unusual pattern of severe tooth wear and frequent tooth loss, primarily the result of consuming a fallback food for which these primates are not dentally adapted. Interpreting these data was only possible by combining our areas of expertise (dental anatomy [FC] and primate ecology [MS]). By integrating theoretical, methodological, and applied aspects of both areas of research, we adopted the term "dental ecology"-defined as the broad study of how teeth respond to the environment. Specifically, we view dental ecology as an interpretive framework using teeth as a vehicle for understanding an organism's ecology, which builds upon earlier work, but creates a new synthesis of anatomy and ecology that is only possible with detailed knowledge of living primates. This framework includes (1) identifying patterns of dental pathology and tooth use-wear, within the context of feeding ecology, behavior, habitat variation, and anthropogenic change, (2) assessing ways in which dental development and biogeochemical signals can reflect habitat, environmental change and/or stress, and (3) how dental microstructure and macro-morphology are adapted to, and reflect feeding ecology. Here we define dental ecology, provide a short summary of the development of this perspective, and place our new work into this context.     

Ring-Tailed Lemur Home Ranges Correlate with Food Abundance and Nutritional Content at a Time of Environmental Stress

In order to determine whether ring-tailed lemurs (Lemur catta) adapt their ranging and select an optimal diet at a time of food shortage, we observed two adjacent troops in Berenty Reserve, Madagascar for over 250 h. The troops, created by a recent fission, ranged through closed canopy gallery forest next to a river and open forest away from the river. We conducted the study in September–October, 2000, normally a time of seasonally low resource availability, which was intensified by damage from a previous windstorm and recent drought. To examine the impact of environmental stress, we mapped their ranging patterns, intertroop encounters, feeding patches, siesta trees, and sleeping trees. We then correlated their ranging and feeding behavior with nutritional analyses of leaves and fruit from tamarind trees located in different parts of their ranges. One of the troops, D1A, ranged farther into open forest than previously. However, the range for troop D1B and the closed canopy portion of D1A's range were located in traditional positions for historical troops D and E. Both troops ate significantly more mature leaves from the tamarind trees in the closed canopy forest, where the leaves had significantly higher nutritional content (water and protein) than that of open forest samples. They fed on tamarind fruit significantly more often in the open forest away from the river, where it was more abundant. The lemurs selected a diet that maximized leaf water and protein and ranged where fruit was most abundant but at high energetic costs for troop D1A.     

Dental topography indicates ecological contraction of lemur communities

Understanding the paleoecology of extinct subfossil lemurs requires reconstruction of dietary preferences. Tooth morphology is strongly correlated with diet in living primates and is appropriate for inferring dietary ecology. Recently, dental topographic analysis has shown great promise in reconstructing diet from molar tooth form. Compared with traditionally used shearing metrics, dental topography is better suited for the extraordinary diversity of tooth form among subfossil lemurs and has been shown to be less sensitive to phylogenetic sources of shape variation. Specifically, we computed orientation patch counts rotated (OPCR) and Dirichlet normal energy (DNE) of molar teeth belonging to 14 species of subfossil lemurs and compared these values to those of an extant lemur sample. The two metrics succeeded in separating species in a manner that provides insights into both food processing and diet. We used them to examine the changes in lemur community ecology in Southern and Southwestern Madagascar that accompanied the extinction of giant lemurs. We show that the poverty of Madagascar's frugivore community is a long-standing phenomenon and that extinction of large-bodied lemurs in the South and Southwest resulted not merely in a loss of guild elements but also, most likely, in changes in the ecology of extant lemurs.     

Functional ecology and evolution of hominoid molar enamel thickness: Pan troglodytes schweinfurthii and Pongo pygmaeus wurmbii

The divergent molar characteristics of Pan troglodytes and Pongo pygmaeus provide an instructive paradigm for examining the adaptive form-function relationship between molar enamel thickness and food hardness. Although both species exhibit a categorical preference for ripe fruit over other food objects, the thick enamel and crenulated occlusal surface of Pongo molar teeth predict a diet that is more resistant to deformation (hard) and fracture (tough) than the diet of Pan. We confirm these predictions with behavioral observations of Pan troglodytes schweinfurthii and Pongo pygmaeus wurmbii in the wild and describe the mechanical properties of foods utilized during periods when preferred foods are scarce. Such fallback foods may have exerted a selective pressure on tooth evolution, particularly molar enamel thinness, which is interpreted as a functional adaptation to seasonal folivory and a derived character trait within the hominoid clade. The thick enamel and crenulated occlusal surface of Pongo molars is interpreted as a functional adaptation to the routine consumption of relatively tough and hard foods. We discuss the implications of these interpretations for inferring the diet of hominin species, which possessed varying degrees of thick molar enamel. These data, which are among the first reported for hominoid primates, fill an important empirical void for evaluating the mechanical plausibility of putative hominin food objects.     

Microhabitat Variation and Its Effects on Dietary Composition and Intragroup Feeding Interactions Between Adult Female <Emphasis Type="Italic">Lemur catta</Emphasis> During the Dry Season at Beza Mahafaly Special Reserve,Southwestern Madagascar

We compared diets and female feeding agonism in 2 groups of ring-tailed lemurs living in markedly diverse microhabitats in and near the Beza Mahafaly Reserve, Madagascar during mid-to-late gestation periods and height of the dry season to test predictions concerning differences in resource availability, preferred foods, female rank, and degree and frequency of feeding agonism in relation to usurpability and monopolization of food resources. Quadrat sampling in disturbed forest habitat revealed a greater number of plant species than in the gallery forest home range area, but females in both groups consumed nearly equal numbers of actual food plant species. Higher-ranking females in disturbed forest consumed human food scraps from the researchers camp significantly more often than low-ranking females did, while there was no rank effect for consumption of any food type between females in the gallery forest group. Higher rates of female feeding agonism in both groups occurred around usurpable fruit and leaves, as well as over monopolizable human food scraps in the disturbed forest group. There is no association between degree of agonism and food type in either group, and rate of feeding agonism is similar for both groups. The most highly contested food items came in large packages and were high in nutrients: beneficial foods for gestating females in the height of dry season in southwestern Madagascar.     

Dental pathologies in ten free-ranging chimpanzees from Gombe National Park, Tanzania

The dental remains of ten adult chimpanzees from Gombe National Park, Tanzania, were examined for enamel attrition, caries, abscesses, periodontal disease, and tooth loss. Age was the underlying factor in the development of dental pathology, in that enamel wear was present to some extent in all ten but was uniformly severe only in the three for whom estimated age at death was 39-43 years. In turn, enamel wear appears to have been the direct cause of abscess development, periodontal disease, and tooth loss. Periodontal disease was commonly expressed as alveolar resorption, particularly around the premolars and molars. This involvement was variable in all except the two youngest. Some interesting wear patterns were evident in the form of deep grooves in the upper incisors and dramatic notching of the lower canines. These patterns, and enamel attrition in general, were attributed to normal mastication and to various stripping activities. Only one carious lesion was observed, in a male with an estimated age of 26 years. An accurate assessment of the actual prevalence of caries was obscured by enamel wear and tooth loss in the older individuals.     

Functional implications of primate enamel thickness.

Recent evolutionary interpretations of Hominoidea have postulated functional relationships between tooth form, diet and masticatory biomechanics. A major consideration is the durability of the tooth under certain dietary conditions. Teeth with low cusps and thicker enamel are able to withstand heavy mastication of abrasive food bolus for a longer period. When comparisons are made between species of higher primates the variables of tooth size, cusp morphology, and enamel thickness appear to be related but until now no systematic analysis has been made to determine the functional relevance of several dental dimensions. This study provides data gained from comparisons of dentition of nine species of primates. Histological sections were made of the post canine teeth and 21 dimensions were compared. The relevant dimensions identified serve to withstand dental wear. The distribution of thicker enamel corresponded to the observed wear planes. Humans had thicker enamel than pongids while the macaque had the thinnest. These preliminary results tend to support theories which explain low, thick, enameled cusps in hominids.     

Berenty 2006: Census of <Emphasis Type="Italic">Propithecus verreauxi</Emphasis> and Possible Evidence of Population Stress

We provide a survey of a folivorous lemur, Propithecus verreauxi (sifaka), in the Berenty Reserve, southern Madagascar. Higher densities of folivores in small patches occur in either high-quality food areas or in less disturbed refugia. The skewness of sex ratio, which in lemurs is often male-biased, can be critically exacerbated in population stress. We predicted that sifaka would show higher densities in areas where protein-rich food is abundant (prediction 1a) and in refugium areas (prediction 1b). Owing to increased competition by brown and ring-tailed lemurs and decreased food production by tamarind trees, we expected an extremely male-biased sex ratio (prediction 2). In November–December 2006, we counted and sexed 206 adult/subadult sifaka (49 groups) during daily walks in different forest zones (Ankoba secondary forest, to the north, Malaza gallery/scrub areas, and spiny forest, to the south). Sifaka may have decreased in the gallery forest to concentrate in Ankoba (in a sort of out-of-Malaza). The area contains protein-rich food (prediction 1a confirmed). Sifaka are proportionally more concentrated in the spiny area than in the degraded scrub forest (prediction 1b confirmed). The sex ratio is extremely male biased, possibly due to either high sifaka density, in Ankoba, or food availability reduction, in Malaza (prediction 2 confirmed). The sifaka population seems to be under stress: researchers need to resume demographic studies, interrupted in Berenty in the mid-1980s, to preserve in situ a species that is difficult to protect ex situ.     

Dento-alveolar lesions and nutritional habits of a Roman Imperial age population (1st–4th c. AD): Quadrella (Molise, Italy)

The study of teeth is very important in archaeoanthropology for reconstruction of the nutritional habits and living conditions of past populations. We have analysed dental lesions of pathological (caries, abscesses and ante mortem tooth loss) and non-pathological origin (calculus), linear enamel hypoplasia and tooth wear in 67 adults from the Roman Imperial age necropolis (1st-4th c. AD) of Quadrella (Molise, Italy). The high frequency of caries (likely giving rise to the abscesses and ante mortem tooth loss), the abundant calculus and the low frequency of heavy wear are probably due to a limited use of hard fibrous foods and a high consumption of carbohydrates. The high frequency of linear enamel hypoplasia suggests metabolic problems during growth. Comparison of these data with those for two coeval Italian necropoleis near Rome (Latium), Isola Sacra and Lucus Feroniae, indicates poorer living conditions in the Quadrella population.     

Lemur habitat and dental senescence in Ranomafana National Park, Madagascar

Not only can teeth provide clues about diet, but they also can be indicators of habitat quality. Conspecific groups living in different habitats with different kinds of foods may exhibit different rates of dental attrition because their teeth are less well adapted to some foods than to others. Ecological disequilibrium describes the situation in which animals live in habitats to which they are relatively poorly adapted. We test whether dental senescence, the wear-related decrease in dental functionality that is associated with decreased survival of infants born to older Propithecus edwardsi females, can be explained by ecological disequilibrium. Specifically, we compare the rates of dental wear in sifaka groups living in nearby habitats that differ in the degree of anthropogenically induced disturbance. We hypothesize that sifakas living in disturbed areas have an unusual rate of tooth wear compared to those living in a more pristine area, and that dental senescence is a consequence of an atypically high wear rate in a degraded habitat. To test whether habitat quality affects tooth wear more generally, we compare rates of use-wear in two subsets of Microcebus rufus living in either relatively undisturbed or disturbed habitats. Contrary to our predictions, we did not detect different rates of tooth wear in disturbed versus undisturbed habitats for either species and consider that reproductively detrimental dental senescence in P. edwardsi females is unlikely to be a pathological consequence of ecological disequilibrium.     

Tooth wear and dental pathology of the Bronze-Iron Age people in Xinjiang, Northwest China: Implications for their diet and lifestyle

Tooth wear and dental pathology related to diet and lifestyle were investigated in the human dental remains unearthed from three archaeological sites of Bronze Age and Iron Age in Xinjiang of northwest China, and in comparative samples from two Neolithic sites in Henan and Shanxi in central China along the Yellow River.Our results indicate that the average tooth wear on most tooth types in the three Xinjiang sites was close to those of the Neolithic samples from central China. The variation within the Xinjiang samples was also explored. Some special wear patterns such as severe wear on the first molar and relatively heavy wear of anterior teeth were observed on the specimens from the Xinjiang sites. Obvious differences in caries and antemortem tooth loss were found between Xinjiang and comparative samples with higher frequencies of caries observed in samples from central China and higher antemortem tooth loss in samples from the Xinjiang sites. Strongly developed exostoses (tori) were also identified on mandibles and maxillae of the specimens from Xinjiang.The authors believe that the differences in tooth wear and dental pathology between Xinjiang and central China were caused by differences in diet and lifestyle. Food of a harder texture was consumed by the people who lived in Xinjiang than by the people in Henan and Shanxi of central China. The higher occurrence of heavily worn anterior teeth and some other special wear patterns, antemortem tooth loss and presence of exostoses on jaw bones in Xinjiang suggest that the people in Xinjiang lived in a relatively harsh environment, frequently gnawing hard objects, or using teeth as some kind of tools. All these activities put masticatory organs under a heavy load. The differences in caries frequencies between the frontier and central areas of China indicate that food richer in carbohydrates was consumed by the people in the central areas.It is proposed that about 3000-2000 years BP in many areas of frontier Xinjiang, people mainly relied on the type of hunter-gatherer economy with agriculture playing a smaller role in their lifestyle.     

Food Intake and Dietary Overlap in Native Lemur catta and Propithecus verreauxi and Introduced Eulemur fulvus at Berenty,Southern Madagascar

The introduction of Eulemur fulvus in 1975 into the Berenty Reserve and their recent attainment of population densities comparable to those of Lemur catta led us to analyze food partitioning among the 3 large prosimian species in the gallery forest. We assessed the diets of ring-tailed lemurs (Lemur catta), brown lemurs (Eulemur fulvus) and sifakas (Propithecus verreauxi) from food intake measurements during 3 successive short-term studies. All species exhibited marked seasonal changes in their major food categories. Dietary overlap was very high between ring-tailed lemurs and brown lemurs during 2 of 3 seasons, including the middle of the dry season. During the latter period, Eulemur appeared to compensate for a low quality diet by increasing the amount of food eaten. In contrast, Lemur fed on lower amounts of food and seemed more efficient at coping with fibrous plant materials. There is low dietary overlap of Lemur catta and Eulemur fulvus versus Propithecus, which exhibit by far the highest dietary diversity of the 3 species. We discuss sustainable coexistence among them, based on respective dietary adaptations and potential for dietary flexibility.