Age-related variability in occlusal wear planes

Investigation of two populations of 136 individuals shows several patterns of occlusal wear plane change which are positively correlated with age. For individuals up to the age of 18, there is a characteristic pattern in which the occlusal wear planes of the mandibular teeth are lingually sloped and those of the maxillary dentition buccally sloped, with the exception of the maxillary premolars, which are also lingually sloped. The long axes of the mandibular teeth give them a lingual orientation relative to the maxillary teeth, and the long axes of the maxillary molars, by contrast, are buccally oriented. In the 18-30 age range for all sexes, the mandibular M1 becomes buccally sloped on its occlusal surface while the occlusal wear plane on the maxillary M1 becomes lingually sloped. Later age changes indicate a trend for the mandibular premolars to become buccally sloped, while the wear planes of the maxillary premolars remain lingually sloped. There is a corresponding tendency for the maxillary and mandibular second molars to undergo changes in the initial orientation of the occlusal wear planes.     

Reciprocal Allogrooming in Dam-reared and Hand-reared Impala Fawns

Among adult females and males of African antelope impala are unique in their performance of reciprocal allogrooming. The occurrence of this behaviour in neonatal impala fawns was explored in a free-ranging impala herd at the San Diego Wild Animal Park where 5 dam-reared fawns were observed from birth through 10 weeks of age. One-way maternal grooming and reciprocal allogrooming with the dam and non dam partners emerged as distinct behavioural systems. Maternal grooming, directed mostly to the anogenital area, was typical of that seen in other ungulates, and sharply declined over the first two weeks. Reciprocal allogrooming, characterized by alternate exchanges of grooming bouts with a partner in the same manner as in adults, was seen as early as 3–8 d after birth. All fawns were grooming with unrelated adult females by the end of the second week. By week 2 virtually every measure of reciprocal allogrooming by fawns (grooming delivered per hour, reciprocity, and percent of encounters initiated) was as high as for adults. The appearance of this reciprocal allogrooming pattern, especially at such an early age, appears to be unique among ungulates, and possibly mammals in general. Three hand-reared impala fawns, deprived of the opportunity to interact with older herdmates, but having access to impala fawns and heterospecific fawns, were observed from 1–3 mo of age. The hand-reared impala showed no alteration in the occurrence of reciprocal allogrooming behaviour compared with the dam-reared control fawns, indicating that allogrooming experience with older animals was not required for the appearance of reciprocal allogrooming at an early age. Interestingly, hand-reared fawns persisted in grooming heterospecific fawns despite the fact that heterospecifics rarely reciprocated grooming. We postulate that the strong predisposition for impala young to groom others may be related to the threat of tick infestation in the impala's ecotone habitat.     

Phylogeography and conservation of impala and greater kudu

The phylogeography of the bush habituated African bovid species impala (Aepyceros melampus) and greater kudu (Tragelaphus strepsiceros) is investigated using mitochondrial DNA (mtDNA) markers. Combined analysis of individual lineages, relationships and population genetics suggest a colonization process from Southern Africa toward Eastern regions in the greater kudu. Results are less clear for the impala, although remaining consistent with a similar pattern of historical dispersion. The study reveals a similar pattern, that is a marked divergence of lineages from South-western Africa relative to other regions. This pattern is opposed to previously published findings in other African bovid species. In the impala, the genetically isolated region is consistent with morphology because it is recognized as the subspecies A. m. petersi, the black-faced impala. In contrast, the similar split of South-western mitochondrial lineages was not expected in the greater kudu on the basis of morphology. Both species show a significant population genetic differentiation. Beyond their phylogeographical value, our results should raise conservation concerns about South-western populations of both species. The black-faced impala is categorized as vulnerable and our data show indications of hybridization with common impala A. m. melampus. The previously unrecognized genetic status of the South-western kudus could also imply conservation regulations.     

Measurements of tooth wear among Australian Aborigines: I. Serial loss of the enamel crown

The dental casts made from Aboriginal children during the course of a longitudinal growth study in Central Australia provided material for analyzing tooth wear under known environmental conditions. The wear facets produced on the occlusal surfaces were clearly preserved on the dental stone casts and recorded the progress of enamel attrition from ages 6 to 18. These casts were photographed and traced by electronic planimetric methods that automatically recorded the location and size of wear facets on the first and second permanent molars. These areas of worn tooth surface were compared to the total tooth surface. The worn surface was regressed on age to calculate wear rates of each tooth. Discriminant analyses were also performed to determine the significance of dental attrition differences between the sexes at each age group. The total wear on each tooth was highly correlated with age as expected but females wore their teeth at a significantly higher rate than males. The mandibular molars wore more rapidly than maxillary teeth in both sexes. The discriminant analysis successfully grouped 91% of the cases according to age and sex. Pattern of wear, the location, and size of wear facets also differed between age groups and sex. The questions of why there is a difference between male and female wear or why there is greater wear on one arch or arch region have no ready answers. The differing rates and pattern of dental wear do suggest that arch shape and growth rates may be the answer though it has yet to be tested. However, the occlusal surface wear is useful for age estimation in a population and provides a record of shifting masticatory forces during growth.     

The Old Ladies of the Seed Harvester ant Pogonomyrmex Rugosus: Foraging Performed by Two Groups of Workers

We examined temporal polyethism in Pogonomyrmex rugosus, predicting a pattern of decreasing age from foragers to nest maintenance workers to individuals that were recruited to harvest a temporary food source. Nest maintenance workers were younger than foragers, as indicated by their heavier mass and lower mandibular wear. In contrast, recruited foragers were similar in mass to foragers but they displayed higher mandibular wear, suggesting that they were at least as old as foragers. Longevity estimates for marked individuals of these two latter task groups showed mixed results. Higher mandibular wear of recruited foragers suggests that they did not follow the normal sequence for temporal polyethism, but rather that they functioned as seed-millers, which should more quickly abrade their dentition. This would be the first demonstration of specialist milling individuals in a monomorphic seed-harvester ant.     

Tooth wear and compensatory modification of the anterior dentoalveolar complex in humans

In populations living in environments where teeth wear severely, some compensatory modification of the dentoalveolar complex is thought to occur during life whereby functional occlusion is maintained as tooth substance is lost by wear. This study investigates one aspect of this modification process: Changes in the anterior dentoalveolar complex that are accompanied with wear were examined in a series of Japanese skeletal samples. In the prehistoric Japanese hunter-gatherer population heavy wear occurs over the entire dentition. The following changes were demonstrated to have occurred in the anterior segment of the dentition accompanied by wear on the anterior teeth: The anterior teeth tip lingually with wear up to a nearly upright position to fill in interproximal spaces that would have been generated by wear, and to maintain contact relations between adjacent teeth. At the same time, the anterior surface of the maxillary alveolar process also inclines lingually to a certain extent. The amount of lingual tipping is greater in the maxillary anterior teeth than in their mandibular antagonists. It is because of this discrepancy that, with age, the horizontal component of the overlap between maxillary and mandibular anterior teeth decreases, and their bite form changes from scissor bite to edge-to-edge bite. Lesser degrees of lingual tipping of the anterior teeth were also detected in the prehistoric agriculturists and historic Japanese populations. The variation in the degree of lingual tipping observed among the samples is explained by inter-population variation in severity and pattern of tooth wear. This and other evidence suggests that mechanisms that compensate for wear in the anterior dentition may be characteristic of all living human populations, independently of the degree of wear severity endured in their environments.     

Interproximal grooving in the Atapuerca-SH hominid dentitions

The dental sample recovered from the Sima de los Huesos (SH) Middle Pleistocene cave site of the Sierra de Atapuerca (Spain) includes 296 specimens. Interproximal wear grooves have been observed in 20 maxillary and mandibular posterior teeth belonging to at least five of the 32 individuals identified so far in the SH hypodigm. Interproximal grooving affected only the adults, and at an age between 25 and 40 years. The appearance, morphology, and location pattern of the SH wear grooves are similar to those reported in other fossil hominids and in more recent human populations. Two alternative proposals, the toothpicking and the fiber or sinew processing hypotheses, compete for explaining the formation of this anomalous wear. The characteristics observed in the wear grooves of the SH teeth are compatible only with the habitual probing of interdental spaces by means of hard and inflexible objects. Dietary grit may also have contributed to the abrasion of the root walls during the motion of the dental probes. Am. J. Phys. Anthropol. 102:369–376, 1997. © 1997 Wiley-Liss, Inc.     

Patterns of dental wear and the role of the canine in Cercopithecinae.

The importance of dental wear patterns in understanding masticatory functions in primates has long been appreciated. However, studies of wear patterns among populations of nonhuman primates are few. The purpose of this investigation is to establish the developmental aspects of dental wear in the Cercopithecinae and to describe certain relevant morphological traits. Studies were made of dental casts from 200 primate specimens of Macaca nemestrina, Macaca mulatta, and Papio cynocephalus. These casts were taken at four-month intervals, beginning at two years of age and continuing over a period of six to seven years. The wear pattern starts with the rounding and eventual flattening of the protoconid and protocone of the erupted first molars. Once this stage is reached, the hypoconid and metaconid of the mandibular, and the hypocone and paracone of the maxillary molars are rounded and eventually flattened. This pattern is maintained until the cusp tips are removed and the dentin exposed, however, the entoconid and metacone are not subjected to significant wear at this stage. Analysis of these dental casts and museum specimens has provided data on the development of dental wear during the maturation of these primates. The distribution of forces acting upon the teeth produce diagnostic patterns of wear, which provide evidence of the force location and magnitude. In examining the data, the hypothesis of canine guidance and its limitation of mandibular motion was evaluated. Specimens whose canines were removed demonstrate that the canines play no significant role in the development or maintenance of dental wear planes.     

Cheek tooth erosion in male babirusa (genus Babyrousa)

The wear on the occlusal surfaces of male babirusa cheek teeth was evaluated in 53 skulls of Babyrousa babyrussa from Buru and the Sula Islands and 87 skulls of B. celebensis from Sulawesi, Indonesia. Based on the comparative lengths of their continually growing maxillary canine teeth, the skulls were divided into five ‘age categories’ (A–E). Numerical and symbolic codes representing tooth wear were applied to each pillar (cusp region) of the mandibular and maxillary permanent third and fourth premolar teeth, and the first, second and third permanent molar teeth. There was no significant difference between the tooth wear patters of skulls in groups A and B, or in groups C and D, and so these were amalgamated. There was close correspondence in wear patterns between each side of the mouth in both species and in each age group. The wear patterns of the mandibular and maxillary teeth, although not identical, were very similar, as were the wear patterns of both species. In group A + B for both species tooth wear was relatively slight, with the M1 teeth experiencing most relative wear. There was almost no wear of the M3 teeth. In group C + D substantial wear of upper and lower M1 was evident. In group E more widespread wear of the cheek teeth was seen, with increased severity of M1 tooth wear, yet there was comparatively much less M2 and M3 tooth wear. The pattern of cheek tooth wear of the Babyrousa spp. was different from that shown by Sus scrofa. Differences in diet selection and processing were highlighted as potential contributing factors. The pattern of cheek tooth wear in male babirusa was not adequate for use to monitor their age.     

Development of the helicoidal plane

In this study of thebelicoidal occlusal plane the relationships between tooth wear, the transverse slopes of mandibular molars and dental arch breadths were examined in 74 pre-contemporary Australian Aboriginal skulls. With increasing age and tooth wear the orientation of the mandibular occlusal surfaces increased towards the buccal. The differential occlusal orientation from first to third molars, present at eruption, tended to increase progressively with tooth wear. These functionally induced changes, together with regional differences in relative breadths of the maxillary and mandibular dental arches, are important in the development of abelicoidal occlusal plane.     

Nursing behavior and early development of impala at San Diego Wild Animal Park

Because impala are commonly exhibited and handreared in zoos and their natural nursing behavior had not previously been studied, we examined nursing and early development in five impala calves housed in a large, naturalistic enclosure. Calves were observed for the first 5 weeks of life during 12-hr continuous watches and 20-min focal animal samples. Total daily suckling time decreased while time grazing and feeding on concentrates increased from 1–5 weeks of age. An increasing proportion of nursing bouts were terminated by the dam as calves matured, with dams terminating almost 70% of bouts during the first week. Suckling success and maternal grooming also decreased after week I, suggesting that impala mothers cut back early on nursing and grooming of offspring. Rapid decline in mother-young spatial proximity and a concomitant increase in calf association with age-mates over time suggests that the mother-young bond is weak and ephemeral in impala. An example of the implications of our results for improving handrearing programs would be that newborn calves should be started on a daylight feeding schedule of one bottle-feeding every three hours, and that they not be allowed to gorge themselves at any one of these feedings.     

Larger antelopes are sensitive to heat stress throughout all seasons but smaller antelopes only during summer in an African semi-arid environment

Heat stress can limit the activity time budget of ungulates due to hyperthermia, which is relevant for African antelopes in ecosystems where temperature routinely increases above 40 °C. Body size influences this thermal sensitivity as large bodied ungulates have a lower surface area to volume ratio than smaller ungulates, and therefore a reduced heat dissipation capacity. We tested whether the activity pattern during the day of three antelope species of different body size—eland, blue wildebeest and impala—is negatively correlated with the pattern of black globe temperature (BGT) during the day of the ten hottest days and each season in a South African semi-arid ecosystem. Furthermore, we tested whether the larger bodied eland and wildebeest are less active than the smaller impala during the hottest days and seasons. Our results show that indeed BGT was negatively correlated with the diurnal activity of eland, wildebeest and impala, particularly during summer. During spring, only the activity of the larger bodied eland and wildebeest was negatively influenced by BGT, but not for the smallest of the three species, the impala. We argue that spring, with its high heat stress, coupled with poor forage and water availability, could be critical for survival of these large African antelopes. Our study contributes to understanding how endothermic animals can cope with extreme climatic conditions, which are expected to occur more frequently due to climate change.     

Age and individual foraging behavior predict tooth wear in Amboseli baboons

Teeth represent an essential component of the foraging apparatus for any mammal, and tooth wear can have significant implications for survival and reproduction. This study focuses on tooth wear in wild baboons in Amboseli, southern Kenya. We obtained mandibular and maxillary tooth impressions from 95 baboons and analyzed digital images of replicas made from these impressions. We measured tooth wear as the percent dentine exposure (PDE, the percent of the occlusal surface on which dentine was exposed), and we examined the relationship of PDE to age, behavior, and life history variables. We found that PDE increased significantly with age for both sexes in all three molar types. In females, we also tested the hypotheses that long‐term patterns of feeding behavior, social dominance rank, and one measure of maternal investment (the cumulative number of months that a female had dependent infants during her lifetime) would predict tooth wear when we controlled for age. The hypothesis that feeding behavior predicted tooth wear was supported. The percent of feeding time spent consuming grass corms predicted PDE when controlling for age. However, PDE was not associated with social dominance rank or maternal investment. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Bush selection along foraging pathways by sympatric impala and greater kudu

In order to identify the selection mechanism of two sympatric African browsers, we analysed encounter rates and selection of bushes along foraging pathways. We monitored the tracks, left overnight, by kudu and impala on an experimental plot of natural Acacia nilotica and Dichrostachys cinerea in the highveld of Zimbabwe, and recorded the number of bushes attacked in each category. Both ungulates were selective for the bush categories, but kudu were consistently more selective than impala, and showed a higher preference for the larger A. nilotica and D. cinerea bushes, which had a significantly greater number of bites which were not reachable by impala. For both kudu and impala, the probability of attacking larger bushes increased significantly with the proportion of large bushes encountered along the foraging pathways, whereas the consumption of smaller bushes was apparently unpredictable. For the most abundant food item (medium D. cinerea), the probability of attack by impala along a pathway decreased with increasing proportions of larger bushes in the experimental area, but was also dependent on impala group size and season. In addition, we found that encounter rates with larger bushes were significantly higher for kudu than for impala. Experimentally reducing the availability of the larger bushes had little effect on both impala and kudu during the following rainy season. However, during the following cool dry season, kudu showed an increased selectivity with a strong preference for the remaining large bushes (large A. nilotica), followed by a sharp decrease in selectivity in the hot dry season when they also fed from significant numbers of medium trees. Impala had little reaction to the experimental changes in the availability of bush categories in either season. We suggest that both kudu and impala selected bushes on the basis of the potential number of bites they can provide, and this resulted in different search strategies. Kudu focussed on the larger bushes which have a larger number of twigs which are out of reach of impala and kudu also probably directed their path preferentially towards the few larger bushes to maximize encounter rates with this favoured bush category. These differences in bush selection process lead to a low overlap in resource use between the two browsers in this type of savanna.     

DENTITION AND AGEING OF THE HIPPOPOTAMUS

I. Summary The present paper is based on the examination of lower jaws from some 3000 individuals of Hippopotamus amphibius L. which were collected in the course of cropping operations in the Queen Elizabeth Park, Uganda between 1961 and 1966. It extends an earlier study by W. M. Longhurst, who described 20 relative age groups based on tooth replacement and wear. The age groups, slightly modified, are described and illustrated here and should serve as a useful field guide. Mean chronological ages from 0–43 years have been allocated to the groups. Checks which confirm the validity of the ages allotted are presented and discussed. These include correlation with the mandibular age group of six known-age animals; findings on the age-related incidence of rinderpest-neutralising antibodies; the orderly progress of fusion of the mandibular symphysis and rate of loss of the first premolar. Growth layers present in the teeth are discussed. The growth of the eye lens is also described. After an initial phase of rapid growth the lens continues to grow throughout life and follows a rectilinear pattern from an estimated age of eight years onwards. This is expected from the findings of similar studies on other species for which known-age specimens are available. Variability of lens dry weight at age is relatively small and indicates that the estimated ages are reasonably precise. Growth of the mandible is analysed and is not inconsistent with the ages allotted. A marked sex difference in mandible weight at age allows the sex of found jaws to be determined at ages above eight years. The growth of the teeth is described. Both canines and incisors show well-marked sex differences in growth rate and size. The post-canine teeth do not show sex differences. Cycles of growth, wear and resorption of these teeth are discussed and it is concluded that mechanical senescence of the teeth is a major factor in mortality at ages above 30 years. Growth in body length is briefly discussed and provides further confirmation of the validity and precision of the age criteria. Growth equations are presented. Finally a survivorship curve derived from the ageing of 207 jaws assumed to represent natural deaths is presented. Population models are constructed from the survivorship curves by calculation of estimated natality rates (obtained by applying data on age at first breeding and annual pregnancy rate to the survivorship data) and life tables are constructed. The shape of the survivorship curve and the percentage recruitment agree with expectation and provide further evidence of the consistency of the age criteria.     

Effects of Relatedness,Dominance, Age,and Association on Reciprocal Allogrooming by Captive Impala

Wild impala display a highly reciprocal allogrooming system that, by virtue of its frequency and high degree of reciprocity, is unique among ungulates. A herd of 35 free-ranging captive impala provided opportunity to examine the degree of reciprocity of allogrooming exchanges and the influence of relatedness, dominance, age and association on partner preferences and distribution of grooming between allogrooming partners. As in wild impala, the exchange of allogrooming bouts in the captive impala was highly reciprocal regardless of partners. Kinship and dominance had no influence on partner preference or distribution of grooming between partners. Although mothers showed a significant preference to allogroom with their unweaned offspring, this preference practically disappeared with older offspring. Age-mates (no greater than 6 mo apart) tended to associate with one another and spatial proximity was positively correlated with grooming partner preference. It was not clear whether impala actively sought out age-mates for grooming, or randomly chose grooming partners from nearby age-mates. The failure to find a role for kinship and dominance is counter to what has generally been found in most Old World terrestrial primate studies. The absence of pronounced social influences, coupled with the known effectiveness of grooming in removing ectoparasites, suggest that a utilitarian role, especially removal of ticks, is an important function of the impala reciprocal allogrooming system.     

Sex differences in tooth wear ofMacaca fuscata,the Arashiyama-A troop in Texas

The incidence of tooth wear was studied in a wild troop ofM. fuscata, that had previously been transplanted from Arashiyama, Japan, to Texas. This study was undertaken to determine differences of attrition between males and females, and between maxillary and mandibular dentitions. Contrary to other findings, the rate of wear was not found to be an expression of sex difference, but seemed rather related to function. The following observations may suffice as examples: The mandibular third premolars function as a honing surface for the maxillary canines, and experience greater wear over time in males due to their proximity to smaller canines which leave their neighbors more vulnerable to wear. The degree of attrition intensity is neither the same for males and females, nor the maxillary and mandibular dentitions. Certain maxillary and mandibular teeth “pair up”; although all “pairs” are identical in males and females, they rank differently in the degree of wear experienced. Overall, females express greater attrition in the maxillary, and males in the mandibular dentitions.     

Canine role in dental wear patterns: Macaca nemestrina

A study of variables and patterns in dental wear among 30 individuals in a colony of Macaca nemestrina shows that consideration of age and sex are crucial for understanding differential wear degrees on molar occlusal planes. With advanced age, this species of non-human primate undergoes obliteration of initial morphological characteristics through the gradual erosion of enamel. Wear gradients are differential from PM1-M3 in both sexes. It appears that there is a functional relationship between degrees of occlusal plane wear and the degree of wear on the canines, and that females show a greater degree of wear changes relative to males of equivalent age because of initial differences in canine length and robusticity due to sexual dimorphism. It appears that there is a direct relationship between occlusal wear plane changes and the degree of wear on the canines, with advanced differential wear showing up in individuals in whom years of maxillary canine honing, canine damage, and the normal wear of mastication has reduced dimensions of unworn permanent canines. Other considerations included in this study are the honing functions of the deciduous first mandibular molars and molar cusp height relative to canine function.     

Mineralization and wear of mandibular first molars in red deer (Cervus elaphus) of known age

The degree of dental wear was measured in 41 mandibular first molars from red deer of known age (range 2–18 years). The calculated wear indices were positively correlated with age ( r = 0.780, P < 0.001). In the same sample, significant ( P < 0.001) differences in hardness (determined by microhardness testing) were observed between enamel and coronal dentine as well as between outer and inner enamel and dentine, respectively. In animals belonging to the same age group (three-year-olds, n = 11), a negative correlation ( r = -0.830, P = 0.0016) was found between wear index and maximum mean enamel hardness. In a sample of five specimens from this age group, a negative correlation ( r = -0.918, P = 0.028) between maximum calcium concentration of enamel (measured by electron microprobing) and wear index was also recorded. We concluded that the intensity of enamel mineralization had a decisive effect on the degree of dental wear. Measuring of enamel hardness and subsequent modification of wear scores or indices is therefore recommended for future studies on the relationship between age and the degree of dental attrition in deer.     

Reciprocal Allogrooming in Wild Impala Lambs

Adult impala engage in a form of reciprocal allogrooming distinguished by a high degree of reciprocity and ***lack of influence of dominance or relatedness on partner preference or distribution of grooming between partners. A previous study on reciprocal allogrooming of captive newborn impala lambs in a zoological park found that the allogrooming emerged as early as the first week after birth and was identical in structure and reciprocity to allogrooming in adults. Because these findings of apparently unique allogrooming behavior of newborn impala could have been a reflection of the effects of being born and raised in a small, stable captive herd, it was necessary to investigate reciprocal allogrooming in newborn impala in the wild. The emergence, reciprocity, rate, and partner distribution of reciprocal allogrooming in wild newborn impala were observed at two study sites: a national park in Zimbabwe and a game farm in South Africa. Maternal one-way grooming between mother and newborn emerged as distinct from reciprocal allogrooming and rapidly declined after week 1 postpartum. Reciprocal allogrooming by lambs was first seen between 5 and 8 d postpartum, and as soon as the behavior occurred it was the same basic pattern as seen in adult impala. The reciprocity index for lambs was near 0.5, indicating that lambs delivered as much grooming during an encounter as the partner. Lambs were grooming frequently with non-mother adults and other lambs by week 1 or 2; by week 3 and onward the allogrooming rate of lambs was more than twice that of their mothers, as predicted by the body size principle of the programmed grooming model. The strong predisposition of neonatal impala lambs to deliver reciprocal allogrooming as early as the first or second week postpartum would appear to reflect a genetically acquired adaptation to the threat of tick infestation in their natural habitat.