Cuspal Enamel Research Articles

Identification of a protein-containing enamel matrix layer which bridges with the dentine–enamel junction of adult human teeth

ObjectiveTo investigate the ultrastructure and chemical composition of the dentine–enamel junction and adjacent enamel of minimally processed third molar tooth sections. DesignUndecalcified human third molar erupted teeth were sectioned and etched with 4% EDTA or 37% phosphoric acid prior to visualization by scanning electron microscopy. Confocal Raman spectroscopy was carried out at 50μm and more than 400μm away from the dentine–enamel junction before and after mild etching. ResultsA novel organic protein-containing enamel matrix layer was identified for the first time using scanning electron microscopy of etched bucco-lingual sections of crowns. This layer resembles a three-dimensional fibrous meshwork that is visually distinct from enamel “tufts”. Previous studies have generally used harsher solvent conditions which likely removed this layer and precluded its prior characterization. The shape of the organic enamel layer generally reflected that of sheath regions of enamel rods and extended from the dentine–enamel junction about 100–400μm into the cuspal enamel. This layer exhibited a Raman CH stretching peak at ∼2931cm−1 characteristic of proteins and this signal correlated directly with the presence and location of the matrix layer as identified by scanning electron microscopy. ConclusionsThe enamel protein layer was most prominent close to the dentine–enamel junction and was largely absent in cuspal enamel >400μm away from the dentine enamel junction. We hypothesize that this protein containing matrix layer could provide an important biomechanical linkage between the enamel and the dentine–enamel junction and by extension, with the dentine, of the adult tooth (246 words).

Comparing the Chronological Distribution of Enamel Hypoplasia in Rogowo, Poland (2nd century ad) Using Two Methods of Defect Timing Estimation

ABSTRACTThe frequency and chronological distribution of enamel hypoplasias were assessed in a Roman Period population of the Wielbark culture from Rogowo, northern Poland, dated to the 2nd century ad. Hypoplasias were recorded on permanent incisors, canines, and premolars of 52 skeletons. The position of linear defects on the crown surface was measured and then converted to the age of occurrence using two methods: a conventional method that employs the chart of enamel development for the permanent teeth, constructed by Massler et al. (1941) in Swärdstedt's (1966) modification, and the method by Goodman & Song (1999), which corrects for hidden cuspal enamel. Hypoplastic defects were found in 48.1% of examined cases. Linear defects [linear enamel hypoplasias (LEH)] were observed in 38.5% of individuals and in 22% of investigated teeth.The chronological distribution of LEH according to the conventional method revealed two peaks of defects: one at 2.6–3.0 years of age and the other at 4.1–4.5 years of age. The method that accounts for hidden cuspal enamel also provided two peaks, but they occurred at later ages: 3.0–4.0 and 4.6–5.0 years of age.The prevalence of hypoplasia in the Rogowo population in comparison with other European populations of the Roman Period seems to be rather low for both individuals and teeth affected. This may indicate advantageous living conditions, which are supported by archaeological data that suggest general well‐being of the Wielbark people. Copyright © 2011 John Wiley & Sons, Ltd.

Human deciduous mandibular molar incremental enamel development

Quantitative studies of incremental markings retained within human enamel have reconstructed the duration and rate (crown and cusp formation times, initiation and completion, daily enamel secretion rates) of permanent tooth development. This approach has provided one way of estimating human age-at-death, and facilitated comparative dental studies of primate evolution. Similar applications from deciduous enamel are inhibited because developmental reconstructions from incremental markings for these teeth are less frequently reported in the literature. This study quantified the duration and rate of enamel development for mesial (protoconid, metaconid) and distal cusps (hypoconid, entoconid) for first (dm1) and second (dm2) deciduous mandibular molars from an archaeological sample of modern human juveniles. Crown formation time can be calculated from the dm1 protoconid because growth initiates and completes in this cusp, and from the dm2 protoconid combined with the final period of hypoconid growth. The dm1 postnatal crown formation time included the time taken for the tubercle of Zuckerkandl to develop, and differed slightly compared to radiographic methods. The majority of dm1 protoconid cuspal (occlusal region) enamel formed before birth. The dm2 entoconid enamel formed mainly after birth. Birth reduced daily enamel secretion rates, changed the visibility of incremental markings, and disrupted enamel growth for 3 to 8 days. Findings presented here can contribute to age-at-death estimates for human infants aged 13-postnatal months or less, and should facilitate comparisons of primate deciduous incremental enamel development in an evolutionary context. Regression equations are included so that cuspal formation time can be estimated from enamel thickness.

Relationship between histological complexity and elemental composition of the cuspal enamels among human deciduous teeth

The purpose of this study is to clear the difference of histology and elemental composition of the cuspal enamels among the human deciduous teeth. The reground surfaces at the cuspal enamels of them were observed under SEM. The contents of seven elements were analyzed quantitatively with EPMA. It is thought that not only the first zone but also the second zone of the inner enamel in the deciduous molar causes the lateral enlargement of the dental caries along the dento-enamel junction because the second zone of the inner enamel in the deciduous molar was more porous than that in the permanent premolar. It is thought that the cuspal enamel in the deciduous teeth is low calcified than that in the permanent teeth because the contents of calcium and phosphorus in the former were significantly lower than those in the latter. It is considered that the cuspal enamel in the deciduous teeth may include less magnesian whitlockite than that in the permanent teeth because the content of magnesium in the former was significantly lower than those in the latter. A relationship was recognized between the complexity level in the histological structure and the elemental composition of the cuspal enamels among the human deciduous teeth.

Role of prism decussation on fatigue crack growth and fracture of human enamel

The role of prism decussation on the crack growth resistance of human enamel is evaluated. Miniature inset compact tension (CT) specimens embodying a section of cuspal enamel were subjected to Mode I cyclic or monotonic loads. Cracks were grown in either the forward (from outer enamel inwards) or reverse (from inner enamel outwards) direction and the responses were compared quantitatively. Results showed that the outer enamel exhibits lower resistance to the inception and growth of cracks. Regardless of the growth direction, the near-threshold region of cyclic extension was typical of “short crack” behavior (i.e. deceleration of growth with an increase in crack length). Cyclic crack growth was more stable in the forward direction and occurred over twice the spatial distance achieved in the reverse direction. In response to the monotonic loads, a rising R-curve response was exhibited by growth in the forward direction only. The total energy absorbed in fracture for the forward direction was more than three times that in the reverse. The rise in crack growth resistance was largely attributed to a combination of mechanisms that included crack bridging, crack bifurcation and crack curving, which were induced by decussation in the inner enamel. An analysis of the responses distinguished that the microstructure of enamel appears optimized for resisting crack growth initiating from damage at the tooth’s surface.

Nothing new under the heavens: MIH in the past?

This was to study an archaeological population of subadult teeth in 17th and 18th century skeletal material from a London (England) cemetery for enamel defects including molar-incisor-hypomineralisation (MIH). Dentitions of 45 sub-adults were examined using standard macroscopic methods and systematically recorded. A total of 557 teeth were examined with a *5 lens and photographed. Ages of the individuals were estimated from their dental crown and root development stages and not from charts that combine tooth eruption with development stages. The dental age of the individual and the approximate age of onset of enamel defects was then calculated on the basis of the chronological sequence of incremental deposition and calcification of the enamel matrix. Affected enamel was graded macroscopically as: - Mild: <30% of the tooth's enamel surface area visibly disrupted (this encompasses the entire range reported in most other studies), Moderate: 31-49% of the tooth's enamel surface area visibly disrupted and Severe: >50% of the tooth's enamel surface area visibly disrupted. Of the total number of individuals 41 (93.2%) showed signs of enamel developmental dysplasia or MIH, 28 of them showing moderate or severe lesions of molars, primary or permanent (63.6% of the sample). Incisors and canines, though surviving much less often, showed episodes of linear hypoplasia. The extensive lesions seen on many of the molars displayed cuspal enamel hypoplasia (CEH). Many of these teeth also exhibited Molar Incisal Hypomineralisation (MIH).

Brief communication: dental development and enamel thickness in the Lakonis Neanderthal molar.

Developmental and structural affinities between modern human and Neanderthal dental remains continue to be a subject of debate as well as their utility for informing assessments of life history and taxonomy. Excavation of the Middle Paleolithic cave site Lakonis in southern Greece has yielded a lower third molar (LKH 1). Here, we detail the crown development and enamel thickness of the distal cusps of the LKH 1 specimen, which has been classified as a Neanderthal based on the presence of an anterior fovea and mid-trigonid crest. Crown formation was determined using standard histological techniques, and enamel thickness was measured from a virtual plane of section. Developmental differences include thinner cuspal enamel and a lower periodicity than modern humans. Crown formation in the LKH 1 hypoconid is estimated to be 2.6-2.7 years, which is shorter than modern human times. The LKH 1 hypoconid also shows a more rapid overall crown extension rate than modern humans. Relative enamel thickness was approximately half that of a modern human sample mean; enamel on the distal cusps of modern human third molars is extremely thick in absolute and relative terms. These findings are consistent with recent studies that demonstrate differences in crown development, tissue proportions, and enamel thickness between Neanderthals and modern humans. Although overlap in some developmental variables may be found, the results of this and other studies suggest that Neanderthal molars formed in shorter periods of time than modern humans, due in part to thinner enamel and faster crown extension rates.

Variation in modern human premolar enamel formation times: Implications for Neandertals

A recent study demonstrated that variation in enamel cap crown formation in the anterior teeth is greater than that in the molars from two geographically distinct populations: native indigenous southern Africans and northern Europeans. Eighty southern African and 69 northern European premolars (P3 and P4) were analyzed in the present study. Cuspal, lateral, and total enamel formation times were assessed. Although cuspal enamel formation times were not consistently different between the two populations, both lateral and total enamel formation times generally were. Bonferroni-corrected t-tests showed that southern Africans had significantly shorter lateral enamel formation time for five of the six cusps, as well as significantly shorter total enamel formation time for these same cusps. An analysis of covariance performed on the lingual cusps of the upper third and fourth premolars showed that differences in enamel formation times between these populations remained when crown height was statistically controlled. A further goal of this study was to ascertain, based on perikymata counts, what Neandertal periodicities would have to be in order for their teeth to have lateral enamel formation times equivalent to either southern Africans or northern Europeans. To this end, perikymata were counted on 32 Neandertal premolars, and the counts were inserted into regression formulae relating perikymata counts to periodicity for each population and each tooth type. Neandertal enamel formation times could be equivalent to those of southern Africans or northern Europeans only if their hypothetical periodicities fall within the range of periodicities for African apes and modern humans (i.e., 6–12 days). The analysis revealed that both populations could encompass Neandertal timings, with hypothetical periodicities based on the southern African population necessitating a lower range of periodicity (6–8 days) than those based on the northern European population (8–11 days).

Molar crown formation in the Late Miocene Asian hominoids, Sivapithecus parvada and Sivapithecus indicus

During the past decade, studies of enamel development have provided a broad temporal and geographic perspective on evolutionary developmental biology in Miocene hominoids. Here we report some of the first data for molar crown development in one hominoid genus, Sivapithecus. The data are compared to a range of extant and extinct hominoids. Crown formation times (CFTs), daily rates of enamel secretion (DSR), Retzius line number and periodicity, and relative enamel thickness (RET) were calculated in a mandibular first molar of Sivapithecus parvada and a maxillary first molar of Sivapithecus indicus from the Siwalik sequence of Pakistan. A CFT of 2.40 years for the protoconid of S. parvada and 2.25 years for the protocone of S. indicus lie within the range of first molar (M1) formation times for the majority of Miocene hominoids (1.96–2.40 years, excluding Proconsul heseloni), and are similar to an M 1 from Gorilla (2.31 years) and M 1s from Pan (2.22–2.39 years). This is unlike the longer CFTs in modern humans, which appear to be linked with their extended growth period. In contrast to extant great apes and humans, daily rates of enamel secretion are rapid in the Sivapithecus M1s during the early stages of growth, which seems to be a common pattern for most Miocene apes. The rapid accumulation of cuspal enamel in the Sivapithecus molars produced thicker enamel than either Pan or Gorilla in a comparable period of time. Future studies on larger samples of living and fossil hominoids are needed to clarify trends in crown development, which may be better understood in the context of life history strategies coupled with good data on body mass and brain size.

Gross enamel hypoplasia in molars from subadults in a 16th–18th century London graveyard

Dental Enamel Hypoplasia has long been used as a common nonspecific stress indicator in teeth from archaeological samples. Most researchers report relatively minor linear and pitted hypoplastic defects on tooth crown surfaces. In this work we report a high prevalence and early age of onset of extensive enamel defects in deciduous and permanent molars in the subadults from the post-medieval cemetery of Broadgate, east central London. Analysis of the dentition of all 45 subadults from the cemetery, using both macroscopic and microscopic methods, reveals disturbed cusp patterns and pitted, abnormal and arrested enamel formation. Forty-one individuals from this group (93.2%) showed some evidence of enamel hypoplasia, 28 of them showing moderate or extensive lesions of molars, deciduous or permanent (63.6% of the sample). Scanning Electron Microscope images reveal many molars with grossly deformed cuspal architecture, multiple extra cusps and large areas of exposed Tomes' process pits, where the ameloblasts have abruptly ceased matrix production, well before normal completion. This indented, rough and poorly mineralized surface facilitates both bacterial adhesion and tooth wear, and when such teeth erupt fully into the mouth they are likely to wear and decay rapidly. We suggest that this complex combination of pitted and plane-form lesions, combined with disruption of cusp pattern and the formation of multiple small cusps, should henceforth be identified as "Cuspal Enamel Hypoplasia."

Enamel microstructure of the hominid KB 5223 from Kromdraai, South Africa

The Plio-Pleistocene site of Kromdraai, South Africa, is well known for the recovery of the holotype of Paranthropus robustus, one of nine individual hominids recovered from this site to date. Among the Kromdraai sample, the specimen KB 5223 comprises several isolated deciduous and permanent lower teeth assigned to Paranthropus, the only recognized genus at this site. However, a more recent analysis of this specimen suggested that it should be classified as Homo. The lower right first permanent molar of KB 5223 had been previously sectioned along the tips of the mesial cusps, exposing its enamel microstructure. Previous studies had indicated differences between Homo and Paranthropus at the microstructural level. A portable confocal scanning microscope was used to describe details of the enamel microstructure of the M1 and I1 of this specimen. Angles formed between the striae of Retzius and the enamel dentine junction (EDJ), daily secretion rates in cuspal enamel of the protoconid and metaconid and crown formation time of the RM1 are provided. The number of perikymata on the right I1 was counted. Results indicate that some features recorded in the KB 5223 molar differ from those of Paranthropus. However, the number of perikymata on the I1 is lower than values so far reported for early Homo but similar to Paranthropus. Crown formation time of KB 5223 M1 was markedly lower than mean values of M1 in H. sapiens, but similar to other early hominids. Daily secretion rates in the cuspal enamel of KB 5223 M1 were higher than in modern humans.

Molar development in common chimpanzees ( Pan troglodytes)

Numerous studies have reported on enamel and dentine development in hominoid molars, although little is known about intraspecific incremental feature variation. Furthermore, a recent histological study suggested that there is little or no time between age at chimpanzee crown completion and age at molar eruption, which is unlikely given that root growth is necessary for tooth eruption. The study presented here redefines growth standards for chimpanzee molar teeth and examines variation in incremental features. The periodicity of Retzius lines in a relatively large sample was found to be 6 or 7 days. The number of Retzius lines and cuspal enamel thickness both vary within a cusp type, among cusps, and among molars, resulting in marked variation in formation time. Daily secretion rate is consistent within analogous cuspal zones (inner, middle, and outer enamel) within and among cusp types and among molar types. Significantly increasing trends are found from inner to outer cuspal enamel (3 to 5 microns/day). Cuspal initiation and completion sequences also vary, although sequences for mandibular molar cusps are more consistent. Cusp-specific formation time ranges from approximately 2 to 3 years, increasing from M1 to M2, and often decreasing from M2 to M3. These times are intermediate between radiographic studies and a previous histological study, although both formation time within cusps and overlap between molars vary considerably. Cusp-specific (coronal) extension rates range from approximately 4 to 9 microns/day, and root extension rates in the first 5 mm of roots range from 3 to 9 microns/day. These rates are greater in M1 than in M2 or M3, and they are greater in mandibular molars than in respective maxillary molars. This significant enlargement of comparative data on nonhuman primate incremental development demonstrates that developmental variation among cusp and molar types should be considered during interpretations and comparisons of small samples of fossil hominins and hominoids.

Variation in enamel development of South African fossil hominids

Dental tissues provide important insights into aspects of hominid palaeobiology that are otherwise difficult to obtain from studies of the bony skeleton. Tooth enamel is formed by ameloblasts, which demonstrate daily secretory rhythms developing tissue-specific structures known as cross striations, and longer period markings called striae of Retzius. These enamel features were studied in the molars of two well known South African hominid species, Australopithecus africanus and Paranthropus robustus. Using newly developed portable confocal microscopy, we have obtained cross striation periodicities (number of cross striations between adjacent striae) for the largest sample of hominid teeth reported to date. These data indicate a mean periodicity of seven days in these small-bodied hominids. Important differences were observed in the inferred mechanisms of enamel development between these taxa. Ameloblasts maintain high rates of differentiation throughout cervical enamel development in P. robustus but not in A. africanus. In our sample, there were fewer lateral striae of Retzius in P. robustus than in A. africanus. In a molar of P. robustus, lateral enamel formed in a much shorter time than cuspal enamel, and the opposite was observed in two molars of A. africanus. In spite of the greater occlusal area and enamel thickness of the molars of both fossil species compared with modern humans, the total crown formation time of these three fossil molars was shorter than the corresponding tooth type in modern humans. Our results provide support for previous conclusions that molar crown formation time was short in Plio-Pleistocene hominids, and strongly suggest the presence of different mechanisms of amelogenesis, and thus tooth development, in these taxa.

The accuracy of histological assessments of dental development and age at death

Histological analyses of dental development have been conducted for several decades despite few studies assessing the accuracy of such methods. Using known-period incremental features, the crown formation time and age at death of five pig-tailed macaques (Macaca nemestrina) were estimated with standard histological techniques and compared with known ages. Estimates of age at death ranged from 8.6% underestimations to 15.0% overestimations, with an average 3.5% overestimate and a 7.2% average absolute difference. Several sources of error were identified relating to preparation quality and section obliquity. These results demonstrate that histological analyses of dental development involving counts and measurements of short- and long-period incremental features may yield accurate estimates, particularly in well-prepared material. Values from oblique sections (or most naturally fractured teeth) should be regarded with caution, as obliquity leads to inflated cuspal enamel formation time and underestimated imbricational formation time. Additionally, Shellis's formula for extension rate and crown formation time estimation was tested, which significantly overestimated crown formation time due to underestimated extension rate. It is suggested that Shellis' method should not be applied to teeth with short, rapid periods of development, and further study is necessary to validate this application in other material.

Variation in modern human enamel formation times

Most of what we know about the timing of human enamel formation comes from radiographic studies on children of known age. Here, we present new longitudinal data derived from a histological analysis of tooth enamel. Two samples, one from southern Africa and one from northern Europe, contained all anterior and molar tooth types. Two further samples contained only one tooth type: canines from a medieval Danish sample and third molars from a modern North American sample. Data were collected on 326 molars and 352 anterior teeth. Each tooth was sectioned and prepared for polarized light microscopy. We used daily enamel cross striations to determine cuspal enamel formation time, recorded the periodicity of long-period striae in the lateral enamel, and used this value to calculate enamel formation times for each decile of crown length. We present data that reveal some of the processes whereby differences in enamel formation times arise between our samples. Mean cuspal enamel formation times were similar in southern African and northern European anterior teeth, but differed in certain molar cusps. All the southern African anterior teeth completed enamel formation earlier. The greatest difference in mean chronological age at enamel completion was 5.2 vs. 6.2 years of age in lower canines. However, enamel completion times in the molar teeth showed few differences between the samples, with mean times for the longest forming cusps all falling between 3.0 years and 3.45 years. Our data suggest fewer differences between samples and smaller ranges of variation than in many radiographic studies and present a more realistic picture of worldwide variation in enamel formation times.

Dental development and life history in Anapithecus hernyaki

The sample of Anapithecus from Rudabánya, Hungary, is remarkable in preserving a large number of immature individuals. We used perikymata counts, measurements of root length and cuspal enamel thickness, and observations of the sequence of tooth germs that cross match specific developmental stages in Anapithecus to construct the first composite picture and time scale for dental development in a pliopithecoid (Catarrhini, Primates). We conclude that the age of eruption of M1 in Anapithecus was similar to various macaque species (∼1.45 months), but that M2 and M3 emergence were close to 2.2 and 3.2 years, respectively (both earlier than expected for similarly sized cercopithecoids). There may have been little difference in individual tooth formation times between cercopithecoids and Anapithecus, but the degree of molar overlap during M1, M2, and M3 crown development, which is extreme in Anapithecus, is fundamentally different. Overall dental development in Anapithecus was very rapid. Old World monkeys appear derived in lacking significant molar overlap, and hominoids may be derived in having longer tooth formation times, both resulting in longer overall dental development times. This is consistent with the general conclusion that the Pliopithecoidea is an outgroup to the Cercopithecoidea and the Hominoidea. On the other hand, rapid dental formation in Anapithecus may be an apomorphy indicative of an unusually rapid life history or unique pressures related to diet and maturation. Folivory and/or predation pressure may be responsible for generating selection to more rapidly erupt permanent teeth and possibly attain adult body masses in Anapithecus. Whatever the case, Anapithecus, with an M3 emergence of ∼3.2 years, is dramatically faster than any extant catarrhine of similar body mass. This represents yet another unusual attribute of this poorly known fossil catarrhine.

An examination of dental development in Graecopithecus freybergi (= Ouranopithecus macedoniensis)

This study examined enamel thickness and dental development in Graecopithecus freybergi (= Ouranopithecus macedoniensis), a late Miocene hominoid from Greece. Comparative emphasis was placed on Proconsul, Afropithecus, Dryopithecus, Lufengpithecus, and Gigantopithecus, fossil apes that vary in enamel thickness and patterns of development. In addition, comparisons were made with Paranthropus to investigate reported similarities in enamel thickness. Several sections of a right lower third molar were generated, from which enamel thickness and aspects of the enamel and dentine microstructure were determined. Data from parallel sections shed light on the effects of section obliquity, which may influence determination of both enamel thickness and crown formation time. Graecopithecus has relatively thick enamel, greater than any fossil ape but less than Paranthropus, with which it does show similarity in prism path and Hunter–Schreger band morphology. Aspects of enamel microstructure, including the periodicity and daily secretion rate, are similar to most extant and fossil apes, especially Afropithecus. Total crown formation time was estimated to be 3.5 years, which is greater than published values for modern Homo, similar to Pan, and less than Gigantopithecus. Data on dentine secretion and extension rates suggest that coronal dentine formation was relatively slow, but comparative data are very limited. Graecopithecus shares a crown formation pattern with several thick-enamelled hominoids, in which cuspal enamel makes up a very large portion of crown area, is formed by a large cell cohort, and is formed in less than half of the total time of formation. In Paranthropus, this pattern appears to be even more extreme, which may result in thicker enamel formed in an even shorter time. Developmental similarities between Paranthropus and Graecopithecus are interpreted to be parallelisms due to similarities in the mechanical demands of their diets.

Enamel thickness and development in a third permanent molar of Gigantopithecus blacki

A ground section was prepared from a lower right M3 attributed to Gigantopithecus blacki as close as possible to axial plane of the mesial cusps. Daily cross striations were imaged, measured and counted in each cusp using polarised light microscopy. Long-period striae of Retzius were counted in the lateral enamel and their periodicity determined from counts and measurements of daily cross striations between adjacent striae. Cross striation spacings in the cusps were between 3.8 μm at the enamel dentine junction and 6 μm close to the enamel surface. Cuspal enamel formation times were long (800 days in the protoconid and 620 days in the metaconid). Linear enamel thickness was as much as 3.75 mm in the protoconid. There were 63 and 61 long-period striae of Retzius in the mesial aspects of the lateral enamel and the periodicity was 11 days. Lateral enamel formation took 1493 and 1291 days and when summed with cuspal enamel formation times totalled 4 years in the protoconid and 3.5 in the metaconid. Relative enamel thickness was 23, calculated through the mesial cusps. This falls short of that in the so-called ‘thick hyper-thick’ enamel described in ‘robust’ australopithecines to which Gigantopithecus blacki has previously been compared in both its dental and mandibular morphology. With respect to enamel thickness, therefore, Gigantopithecus blacki falls squarely among an increasingly large number of Miocene hominoids that can all be described as having ‘thick enamel’.

Age at first molar emergence in early Miocene Afropithecus turkanensis and life-history evolution in the Hominoidea.

Among primates, age at first molar emergence is correlated with a variety of life history traits. Age at first molar emergence can therefore be used to broadly infer the life histories of fossil primate species. One method of determining age at first molar emergence is to determine the age at death of fossil individuals that were in the process of erupting their first molars. This was done for an infant partial mandible of Afropithecus turkanensis (KNM-MO 26) from the approximately 17.5 Ma site of Moruorot in Kenya. A range of estimates of age at death was calculated for this individual using the permanent lateral incisor germ preserved in its crypt, by combining the number and periodicity of lateral enamel perikymata with estimates of the duration of cuspal enamel formation and the duration of the postnatal delay in the inception of crown mineralization. Perikymata periodicity was determined using daily cross striations between adjacent Retzius lines in thin sections of two A. turkanensis molars from the nearby site of Kalodirr. Based on the position of the KNM-MO 26 M(1)in relation to the mandibular alveolar margin, it had not yet undergone gingival emergence. The projected time to gingival emergence was estimated based on radiographic studies of M(1)eruption in extant baboons and chimpanzees. The estimates of age at M(1)emergence in KNM-MO 26 range from 28.2 to 43.5 months, using minimum and average values from extant great apes and humans for the estimated growth parameters. Even the absolute minimum value is well outside the ranges of extant large Old World monkeys for which there are data (12.5 to <25 months), but is within the range of chimpanzees (25.7 to 48.0 months). It is inferred, therefore, that A. turkanensis had a life history profile broadly like that of Pan. This is additional evidence to that provided by Sivapithecus parvada (Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, 1997, 173) that the prolonged life histories characteristic of extant apes were achieved early in the evolutionary history of the group. However, it is unclear at present whether life-history prolongation in apes represents the primitive catarrhine pace of life history extended through phyletic increase in body mass, or whether it is derived with respect to a primitive, size-adjusted life history that was broadly intermediate between those of extant hominoids and cercopithecoids. Life history evolution in primates as a whole may have occurred largely through a series of grade-shifts, with the establishment of fundamental life-history profiles early in the histories of major higher taxa. These may have included shifts that were largely body mass dependent, as well as those that occurred in the absence of significant changes in body mass.

Enamel thickness, microstructure and development in Afropithecus turkanensis.

Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.