AGE CRITERIA FOR THE AFRICAN ELEPHANT

1. The paper is based on a collection of 385 lower jaws of the African elephant (Loxodonta a. africana Blumenbach) from western Uganda. 2. Thirty age groups are described and illustrated, which are related to the progress of eruption and wear of the six teeth in each side of the lower jaw. 3. Correct identification of individual teeth in the series is essential and several checks are described. Thus, when length is plotted against width for the six teeth the points fall into six well - defined groups. Frequency distributions of laminary indices and lamella numbers support the belief that teeth have been correctly identified, but overlapping, distributions preclude identification on these characters alone. 4. The presence of abnormal seventh molars in four jaws is described. 5. Chronological ages have been assigned to the relative age groups. Information on age and growth of captive animals has been considered and the general form of the growth curve established. Arbitrary estimates of the intervals between successive age groups were made and subsequently checked against growth and seasonal ridges on the roots. These indicate an upper age limit of about 60 years, which is compatible with fuller information on Indian elephants. 6. Growth curves support the validity of the ages assigned. Asymptotic heights at shoulder (∞) are respectively 317 cm and 298 cm for males from Murchison Falls and Queen Elizabeth National Parks. For females no distinction is made (because the sample from the latter area is small) and ∞ is 272 cm. 7. Growth in weight has also been established but shows no significant differences between the two populations. The mean maximum weight of females is 2766 kg (6100 lb), and of males 5450 kg (12000 lb). Growth apparently continues throughout life. 8. Tusk growth is analysed. There is a linear increase in weight of female tusks from eruption at 1–3 years up to the oldest group, with an indicated mean combined weight of 17.7 kg (39 lb) at 60 years. Male tusks show an increasing rate of growth throughout life to a mean combined weight of about 109 kg (240 lb) at 60 years. Big tusks are generally the result of prolonged growth; extremely big tusks probably result from prolonged and above average rate of growth. 9. Seasonal and annual incremental layers on the root 3 of the teeth are briefly described; these give an objective estimate of the chronological intervals between the relative age groups. 10. Field age criteria are presented which are derived from these growth curves. 11. The age at puberty in the female elephant is discussed and compared with earlier conclusions. There is evidence of a retardation in recent years in the Murchison Falls National Park (South bank) population and of a lengthening of the mean calving interval. This considerable depression of the reproductive rate, which is almost certainly density dependent, agrees with the observed lower recruitment in this population. 12. Survivorship curves constructed from material representing 325 natural deaths are presented for the two populations. If estimates of the age at puberty and the reproductive rate are taken into account, the expected differences in recruitment are found. 13. The calculated mean expectation of life is less than 15 years. 14. Mean individual weight is estimated at about 3800 lb.     

EYE LENS WEIGHT AND AGE IN AFRICAN ELEPHANTS

Eye lens dry-weights have been determined for 543 African elephants from three populations in East Africa. When plotted against estimated ages based on tooth replacement and wear criteria they indicate growth curves with rapid initial growth in lens weight, succeeded by a phase of rectilinear growth which apparently persists throughout life.
Parameters for the regressions of lens dry weight on age have been calculated by sex and locality. Confidence limits are fitted and no significant difference in the growth rates can be demonstrated, except for a sex difference in the values for the α intercept.
Variability at age is slightly greater in males than females, but is little greater than i3 indicated by studies on other species using known-age animals. This is taken to confirm the accuracy of the age criteria adopted and leads to conclusions on their precision.
It is suggested that this method might provide an objective check on the accuracy and precision of age estimates in other species.     

On the relative frequencies of hominid maxillary and mandibular teeth and jaws as taphonomic indicators

In southern African samples of early hominid remains, maxillary and mandibular teeth (deciduous-plus-permanent) have a virtually equal chance of accumulating in the dolomitic limestone cave deposits, of being preserved therein and recovered therefrom. Thus, of 1066 fossil teeth ofAustralopithecus spp. plusHomo habilis, 51.9 per cent are maxillary and 48.1 per cent mandibular. On the other hand, the East African sample of 847 early hominid, deciduous-plus-permanent teeth, departs more strikingly from a 1∶1 ratio: it comprises 41.0 per cent maxillary and 59.0 per cent mandibular teeth. It is inferred that mandibular teeth have a somewhat better chance of accumulating and being preserved in, and being recovered from, the open, fluvial, lacustrine and deltaic sedimentary environments of the East African sites. The dental proportions are approximately matched by the proportions of jaws. For example, the maxilla: mandible proportions at Koobi fora in northern Kenya are 33.0∶67.0 for teeth and 21.6∶78.4 for jaws. In other words, the preponderance in favour of mandibular remains is somewhat more marked in the case of jaws than of teeth, this distinction doubtless reflecting the more fragile bony structure of the maxilla and the sturdier construction of the mandible. This first study known to the author of the differential distribution of maxillary and mandibular teeth of the Plio-Pleistocene hominids leads the author to hypothesize that, where environmental conditions at the place and time of the death of the hominids have been non-destructive, non-dispersive, relatively mild and protective, maxillae and mandibles may be expected to have been conserved and recovered in approximately equal proportions—and likewise of maxillary and mandibular teeth. On the other hand, the more brutal and destructive the sedimentary environment and other taphonomic influences have been, at the place and time when the hominid individuals died, the more likely it is that the maxillary and mandibular remains of jaws and teeth will deviate from equality of proportions, generally at the expense of the maxillae and upper teeth. Hence, it is proposed that the upper jaw/low jaw ratio (Mx/Mn jaw ratio) and the maxillary teeth/mandibular teeth ratio (Mx/Mn dental ratio) may serve as two useful new gauges of the rigour of palaeo-ecological and taphonomic conditions.     

The mixed dentition and associated skull fragments of a juvenile fossil hominid from Sterkfontein,South Africa

In April–May 1983, the late A.R. Hughes and his field team recovered more than 40 bone fragments and teeth from a single solution pocket of the Sterkfontein Formation. After preparation and reconstruction by JMC, it was recognised that these fragments represent a single juvenile individual (Stw 151), consisting of more than 40 cranial and dental parts, with mixed dentition. It constitutes the most complete set of jaws and teeth of an early hominid child since the Taung child was recovered in 1924. In this paper, the morphological and metrical features of the individual teeth are described. The other associated skull fragments (right ramus of the mandible, left petrous bone, right glenoid region) are also described. Comparisons are made with other South (and East) African fossil hominids. The beautiful preservation simultaneously of most of the deciduous teeth and of the permanent teeth exposed in their crypts allows an accurate analysis of the developmental sequence. A report on the dental developmental status of this juvenile is presented. On the basis of the microanatomical study of the developing permanent teeth, the estimated age at death is 5.2–5.3 years. Reconstructions of the maxillary and mandibular arcades are also offered. The morphological and metrical features of Stw 151 raise the possibility that it may represent a hominid more derived towards an early Homo condition than the rest of the A. africanus sample from Member 4. Am J Phys Anthropol 106:425–465, 1998. © 1998 Wiley-Liss, Inc.     

Age determination and body growth of the Common duiker Sylvicapra grimmia(Mammalia)

Age determination in the Common duiker Sylvicapra grimmia was investigated by analysis of tooth eruption and replacement sequence, incremental lines of tooth cementum and tooth wear in a unique collection of 48 known-age skulls, and also by analysis by post-natal body growth in known-age duiker. In both the mandible and maxilla, permanent molariform teeth were fully erupted and in wear by 26 months of age. There was little variation in the age of eruption and replacement of all molariform teeth, making this a particularly useful feature of the duiker for age determination purposes. In contrast, the variability in eruption of the incisiforms, coupled with the difficulty in distinguishing deciduous incisiforms from the permanent counterparts, placed an unexpected limitation on the use of these teeth. Although the apparent linear relationship between tooth attrition and age has potential for further investigation as an age determination technique, the cementum annuli were not correlated with chronological age. Theoretical Von Bertalanffy equations were used to analyse body growth with age. It was concluded that because the asymptote of growth was reached at such an early age, and because there is so much individual variation in growth, body growth, including horn growth, is of very limited value for age determination. Female duiker were significantly larger than males.     

Sexual dimorphism of the human mandible and its association with dental development

The present study investigates whether the human mandible is sexually dimorphic during early postnatal development and whether early dimorphic features persist during subsequent ontogeny. We also examine whether mandibular dimorphism is linked to dimorphism of dental development. Dense CT-derived mandibular meshes of 84 females and 75 males, ranging from birth to adulthood, were analyzed using geometric morphometric methods. On the basis of the specimen's chronological ages and mineralization stages of the deciduous and permanent teeth, we compute dental age as proxy for dental development by the additive conjoint measurement method. By birth, males have, on average, more advanced age-specific shapes than females. However, sex differences decrease quickly as females catch up via a different association between shape and size. This leads to an almost complete reduction of sexual dimorphism between the ages of 4 and 14. From puberty to adulthood, males are characterized by allometric shape changes while the shape of the female mandible continues to change even after size has ceased to increase. Dimorphism of dental maturation becomes visible only at puberty. Sexual dimorphism, concentrated at the ramus and the mental region during the earliest ontogenetic stages and again at adulthood, is not associated with the development of the teeth. At puberty there is a simultaneous peak in size increase, shape development, and dental maturation likely controlled by the surge of sex hormones with a dimorphic onset age. We argue that the infant and adult dimorphism of the mental region may be associated with the development of supralaryngeal structures.     

Faster or slower: has growth of eastern Baltic cod changed?

Recent environmental changes have influenced the ecology and biology of eastern Baltic cod. Declining somatic condition, maturation at smaller size and restricted size distribution of the population suggest that growth rates have decreased between the early 2000s and the 2010s. Extensive age estimation problems have until now precluded testing of this hypothesis. This study presents evidence for a decrease in somatic growth rate of Baltic cod. Temporal patterns of growth, condition and maturation were analysed based on two complementary analyses: length frequency mode progression derived from DATRAS bottom trawl survey data and known-age samples, where size at age was back-calculated from daily otolith growth patterns. In the known-age samples, growth was positively related to somatic condition at capture with maturity dependent differences. Immature individuals had experienced significantly lower growth and were in lower condition at capture than mature individuals. Growth rates in the known-age samples were estimated at 9.5, 7.8 and 5.7?cm per year for age classes 1, 2 and 3 respectively. Growth between age 2 and 3 decreased significantly from 8.8?cm in the 1997 year class to 7.6?cm in the 2010 year class. While the 2001 and 2004 known-age samples were representative for the population, the 2013 sample was biased towards individuals with a higher condition and growth. Complementary length frequency analysis following the length mode of fish from age 2 to age 3 confirmed growth estimates from the early 2000s, while suggesting a 37.5% lower growth in 2013 compared with 2005.     

Geratodontology and horn growth of the impala (Aepyceros melampus)

Examination of the jaws of an impala population showed advanced wear on mandibular M1 compared with other ungulates which have been examined. This could lead to an erroneous interpretation of age if based upon mandibular tooth wear alone. Explanations are offered for this pattern in terms of the apparent pattern of wear of the impala molar teeth. Suggestions are also put forward for a method of determining specific age, from a conceptual wear model, when only extreme parameters are known. Horn growth in the male is also described.     

AGE DETERMINATION IN MANATEES USING GROWTH-LAYER-GROUP COUNTS IN BONE

Growth layers were observed in histological preparations of bones of known-age, known minimum-age, and tetracycline-marked free-ranging and captive Florida manatees ( Trichechus manatus latirostris ), substantiating earlier preliminary findings of other studies. Detailed analysis of 17 new case histories showed that growth-layer group (GLG) counts in the periotic bone were consistent with known age, or time since tetracycline administration, but were less reliable in other bones. GLG counts were also made in periotic bones of 1,196 Florida manatees of unknown age found dead from 1974 through 1991. These counts were conducted in order to assess variability and to determine relationships among estimated age, size, sex, and degree of bone resorption. Resorption can interfere with accuracy of GLG counts. This effect does not occur until ages greater than about 15 yr and body lengths greater than 300 cm are attained. GLGs were also observed in periotic bones of Antillean manatees ( Trichechus manatus manatus ) but were not validated against known-age specimens. Use of GLG counts in the periotic bone is suitable for application to studies of population dynamics and other age-related aspects of manatee biology.     

Dentition and age determination of the giraffe Giraffa camelopardalis

Thirteen tooth eruption stages and their corresponding chronological ages are descri bed from a series of giraffe jaws. These can be used for age determination in giraffes with immature dentition. Significant correlations of the lingual crown height ( r =0-957; P < 0001) and lingual occlusal surface width ( r =0-959; P < 0001) with the number of dark staining incremental lines in the cementum of thin decalcified sections of the maxillary first molar were found. The regression equations derived from these relationships provide a further method for determining the age of a giraffe. A composite plate showing maxillary first molar wear patterns provides a means of roughly assigning an age to a particular specimen. Thin sections of undecalcified teeth, mandible measurements, various other indices of tooth wear and eye lens mass were investigated and found unsuitable for age determination in this     

African elephant age determination from teeth: validation from known individuals

We assess tooth‐based age criteria for African elephants developed by Laws in relation to known‐aged individuals in the Amboseli elephant population. Laws’s technique remains a robust and useful mechanism for age determination, although we suggest revisions to the oldest age categories. Blind age assignment to jaws of unknown sex using the Laws criteria resulted in misclassification of M4 and M5; measures overlapped too much to differentiate these teeth by sex. Sexes could be reliably distinguished after age 30 or XIX in tooth category by two measures: mandible thickness and width of the ascending ramus, but individuals of the same known age differed in tooth wear and progression rates. Such variation needs to be incorporated in the error assigned to tooth age categories. Ages at death of found jaws (n = 266) were similar to results of survival analysis from all demographic data (n = 2455), excluding calves whose jaws decompose because of weathering and scavengers. Jaw‐based models of age at death need correction for the inability to detect this early mortality, which artificially extends mean longevity by up to 6 years.     

Sexual dimorphism in mandibular growth of French-Canadian children 6 to 10 years of age

Polynomial regression is used to model the mandibular growth of 28 girls and 26 boys who were followed longitudinally from 6 to 10 years of age. The pooled-within individual designs indicate that ramus height follows a linear pattern of size increase; corpus and total mandibular lengths display curvilineal, decelerating, patterns of growth over the age range. Multivariate analyses of variance reveal significant sex differences in size, favoring boys, for the two length measures at 6 years of age. Growth velocity for corpus length is also significantly greater in boys than in girls. Sexual dimorphism in the growth of total mandibular length is more complex, including differences in velocity and deceleration. Ramus height shows no significant pattern of variation between boys and girls for either size or growth velocity.     

Tooth size variation related to age in Amboseli baboons

We measured the molar size from a single population of wild baboons from Amboseli (Kenya), both females (n=57) and males (n=50). All the females were of known age; the males represented a mix of known-age individuals (n=31) and individuals with ages estimated to within 2 years (n=19). The results showed a significant reduction in the mesiodistal length of teeth in both sexes as a function of age. Overall patterns of age-related change in tooth size did not change whether we included or excluded the individuals of estimated age, but patterns of statistical significance changed as a result of changed sample sizes. Our results demonstrate that tooth length is directly related to age due to interproximal wearing caused by M2 and M3 compression loads. Dental studies in primates, including both fossil and extant species, are mostly based on specimens obtained from osteological collections of varying origins, for which the age at death of each individual in the sample is not known. Researchers should take into account the phenomenon of interproximal attrition leading to reduced tooth size when measuring tooth length for ondontometric purposes.     

AGE CRITERIA AND VITAL STATISTICS OF A BLACK RHINOCEROS POPULATION

Tsavo National Park, in Kenya, probably contains the largest population of black rhinoceros (Diceros bicornis (L.)) left in existence. Large-scale damage of the vegetation initiated by elephants and aggravated by fire has changed considerable areas of the park. The ecology of the black rhinoceros in this changing environment has been studied; the present status and population structure is considered here, including the development of detailed ageing criteria for the species, an analysis of natural mortality and survivorship, and a record of the structure of the living populations within the major habitat types. Crania and mandibles were collected from 506 rhinoceros found dead in all areas of the park. This material was divided into 20 relative age classes based on dental characteristics. Crude chronological ages, based on an estimate of the maximum expectation of life and the examination of seven known-age dental records of captive animals, were assigned to each age class. These crude ages were then refined by examination of 16 dental records of known-age wild rhinoceros, and a chronological age scale established. A survivorship curve of the population was thus constructed. Annual mortality during the first and second year of life is about 16%, and the indicated mean annual mortality from 5–25 y is 9.8%. A theoretical model of the population structure is shown, and analysis of the annual mortality and recruitment at birth suggests that the population was stable during the 1960's. Assuming the data represent a stable population the mean expectation of life at birth is 8.4 y. Thirteen major habitat types are described. The characteristics of the rhinoceros population within each habitat type were established both from ground studies and aerial observations. Nearly 700 rhinoceros were identified and catalogued on the ground, in sample areas selected for intensive study. Population structures, cow: calf ratios, and recruitment appear to be average in most habitat types. The analysis of recruitment at birth and mortality during the first year of life, both from computations from the survivorship curve and from the structure of the living populations, support the validity of the ageing criteria, and further suggest that the population was stable during the 1960's. Finally, the relationship between the elephant and the rhinoceros in the changing environment is discussed with reference to recent findings in elephant ecology. In conclusion it is recommended that population reduction of the Tsavo elephants should be initiated.     

A fossilized human mandibular fragment from Kangatotha, Kenya, East Africa

Working west of Lake Rudolf in 1965 Professor Bryan Patterson found at Kangatotha, among other human specimens yet to be described, a fragment of the corpus of a human mandible bearing three molar teeth. Its C-14 date is 2835 B.C . The mandible is stout, heavily mineralized, and closely similar to others from Ishango on the west shore of Lake Edward found by de Heinzelin earlier. The three teeth are large and also similar to those from Ishango. Cranial and postcranial bones of the Ishango people show them to have been Negroes, and the jaws and teeth fit the same classification. The individual from Kangatotha was also indubitably a Negro.     

Association of relatively delayed emergence of mandibular molars with molar reduction and molar position

Among 234 children examined annually from age three to 20 years at the Burlington Growth Centre, there was statistically significant cooccurrence of early and late emergence sequences of the permanent first and second molars relative to the central incisors and second premolars in the same jaw and in both jaws. Alternatively, mandibular molar delay was not accompanied by corresponding maxillary molar delay, and the mandibular molars emerged later than the maxillary molars. This was strongly associated with Angle Class II malocclusion, indicating a relationship between relative time of emergence and relative position of opposing molars. Delay of the mandibular molar relative to the successional teeth or maxillary molars was associated with increased frequency of four cusped first and second molars and agenesis of third molars, indicating a tendency for co-occurrence of delay in timing of molar emergence with reduction in structure of the molars. These relationships were evident even though emergences were affected by early loss of a deciduous second molar which increased M1I1 and M2P2 sequences by earlier emergence of M1 and delayed emergence of P2.     

Third molar agenesis in human prehistoric populations of the Canary Islands

The occurrence of third molar agenesis was recorded in a sample of 1,492 maxillary and 1,718 mandibular arches belonging to the prehistoric settlers of the Gan Canaria, Tenerife, and La Gomera Islands (Canary Islands). There were significant sex differences only in the Tenerife sample for the maxilla, the incidence in females being higher than in males. In the Gran Canaria sample, the total frequency (male and female combined) of third molar agenesis (individual count method) was 8.7% for the maxilla and 9.3% for the mandible. In the Tenerife and La Gomera samples, the frequencies were 11.1% and 10.7% for the maxilla and 14.6% and 13.3% for the mandible. In the Tenerife sample, the differences between both jaws were statistically significant. The incidence of missing third molars in the mandible was significantly higher in Tenerife than in Gran Canaria, but the other sample differences were statistically nonsignificant. Bilateral absence of third molars was observed in about two-thirds of the specimens examined. Some correlation between both jaws for the occurrence of third molar agenesis was found. The hypotheses that have been proposed in order to explain third molar agenesis in man are discussed. It is suggested that the loss of the third molar in Homo sapiens could be produced by a heterochronic phenomenon of postdisplacement, as a consequence of the phylogenetic tendency toward the delay of the onset of the third molar formation, and that the genetic factors responsible for the absence of these teeth could be related to the general process of delay in tooth formation.     

Formation of the permanent dentition in Arikara Indians: timing differences that affect dental age assessments

This report concerns one problem encountered with application of American white dental formation standards to age assessment of sub-adults of archaeological context. Dental ages for eight mandibular permanent teeth and maxillary central and lateral incisors of Arikara Indian immature skeletons were determined according to degree of crown or root mineralization. Ages assigned to the various teeth of the same individual were compared. They showed similarities as well as patterned differences. First premolar, second premolar, and mandibular incisor ages closely approximated one another. In relation to this complex, dental ages for maxillary incisors and mandibular second molars were older by 0.5 to 1.1 years. Developmental ages assigned to individuals on the basis of third molars showed relative advancement by more than 2 years. The systematic occurrence of these observations reflects more than just individual variability; it shows the presence of population differences in tooth-formation timing. Timing differences complicate assessment of dental ages needed for growth or demographic studies.