Platyrrhine dental eruption sequences

To determine dental eruption sequences of extant platyrrhines, 367 mandibles and maxillae of informative juvenile specimens from all 16 genera were scored for presence of permanent teeth including three intermediate eruption stages following Harvati (Am J Phys Anthropol 112 (2000) 69-85). The timing of molar eruption relative to that of the anterior dentition is variable in platyrrhines. Aotus is precocious, with all molars erupting in succession before replacement of any deciduous teeth, while Cebus is delayed in M2-3 eruption relative to I1-2. Callitrichines have a distinct tendency toward delayed canine and premolar development. Platyrrhine eruption sequences presented here show some evidence of conformity to Schultz's Rule, with relatively early replacement of deciduous dentition in "slower"-growing animals. The relationship of dental eruption sequences to degree of folivory, body mass, brain mass, and dietary quality is also examined. The early eruption of molars relative to anterior teeth in Pithecia, Chiropotes, and Cacajao, in comparison to genera such as Ateles, Lagothrix, and Alouatta, showing relatively later eruption of the molars, appears to be consistent with current phylogenetic hypotheses. Schultz (Am J Phys Anthropol 19 (1935) 489-581) postulated early relative molar eruption as the primitive dental eruption schedule for primates. The extremely early molar eruption of Aotus versus Callicebus (where both incisors erupt before M2 and M3, with M3 usually last) may lend support to the status of Aotus as a basal taxon. The early relative molar eruption of the fossil platyrrhine species Branisella boliviana is also consistent with this hypothesis (Takai et al.: Am J Phys Anthropol 111 (2000) 263-281).     

Dental eruption sequence among colobine primates

Dental development is one aspect of growth that is linked to diet and to life history but has not been investigated among colobines since the work of Schultz [1935]. This study establishes the dental eruption sequence for several colobine species and compares it to that of other catarrhines. The mandibles and maxillae of two hundred and four juvenile colobine specimens were scored for presence or absence of permanent teeth and for stages of partial eruption. Eruption was defined as ranging between tooth emergence (any part of a tooth crown above the alveolar margin) and full occlusion, with three intermediate levels manifest between these boundaries. In African colobines, represented by C. guereza, C. angolensis and P. badius, M2 erupts before I2, and in C. angolensis it also erupts before I1. The canine is delayed, erupting after the premolars in females and after M3 in males. Asian colobines show greater diversity in eruption sequences. Nasalis shows no early eruption of the molars and is very similar to Macaca. In Trachypithecus and Pygathrix M(2) erupts before I(2). The canine in Trachypithecus is delayed, erupting after the premolars and, in some males, after M3. In Presbytis M2 erupts before both incisors; M3 erupts before C in both sexes, and often before both premolars. Although the actual timing of eruption is unknown, all colobine species examined except N. larvatus showed some degree of relatively early eruption of M2 and M3. The lack of this tendency in Nasalis sets this genus apart from all other colobines represented in this study. Dental eruption sequence is thought to reflect life history patterns. Early molar eruption in colobines was thought by Schultz (1935) to be a primitive character reflecting shorter life history. Faster growth rates found in folivorous primates have been interpreted as being related to an adaptation to folivory (Leigh 1994), and early eruption of molars may be part of this dietary specialization. The relationships between dental development and both diet and life history are investigated.     

Dental metric assessment of the omo fossils: implications for the phylogenetic position of Australopithecus africanus

The discovery of Australopithecus afarensis has led to new interpretations of hominid phylogeny, some of which reject A. africanus as an ancestor of Homo. Analysis of buccolingual tooth crown dimensions in australopithecines and Homo species by Johanson and White (Science 202:321-330, 1979) revealed that the South African gracile australopithecines are intermediate in size between Laetoli/hadar hominids and South African robust hominids. Homo, on the other hand, displays dimensions similar to those of A. afarensis and smaller than those of other australopithecines. These authors conclude, therefore, that A. africanus is derived in the direction of A. robustus and is not an ancestor of the Homo clade. However, there is a considerable time gap (ca. 800,000 years) between the Laetoli/Hadar specimens and the earliest Homo specimens; "gracile" hominids from Omo fit into this chronological gap and are from the same geographic area. Because the early specimens at Omo have been designated A. afarensis and the later specimens classified as Homo habilis, Omo offers a unique opportunity to test hypotheses concerning hominid evolution, especially regarding the phylogenetic status of A. africanus. Comparisons of mean cheek teeth breadths disclosed the significant (P less than or equal to 0.05) differences between the Omo sample and the Laetoli/Hadar fossils (P4, M2, and M3), the Homo fossils (P3, P4, M1, M2, and M1), and A. africanus (M3). Of the several possible interpretations of these data, it appears that the high degree of similarity between the Omo sample and the South African gracile australopithecine material warrants considering the two as geographical variants of A. africanus. The geographic, chronologic, and metric attributes of the Omo sample argue for its lineal affinity with A. afarensis and Homo. In conclusion, a consideration of hominid postcanine dental metrics provides no basis for removing A. africanus from the ancestry of the Homo lineage.     

Hominid molars from a Middle Stone Age level at the Mumba Rock Shelter, Tanzania

Three hominid molars were recovered from a depth of 7.0-7.1 meters in the Mumba Shelter at Lake Eyasi, northern Tanzania. Geological context of the finds and archaeological data indicate that people with a Middle Stone Age technology were using the Mumba locality intermittently whenever retreat of lake waters allowed access to the site. Uranium series dates suggest an age on the order of 130,000 years bp for the teeth and stone tools. Based on morphological analyses, the dental remains probably belonged to one individual and appear to be the crowns of two upper permanent M2s and one lower permanent M2. Crown areas are very small, even in comparison to the variation exhibited by recent African populations. Crown patterns have no archaic features. These teeth are smaller than any verifiable archaic Homo sapiens examples; thus, they may represent early anatomically modern Homo sapiens.     

Schultz's unruly rule: dental developmental sequences and schedules in small-bodied, folivorous lemurs

Schultz's rule (as reconstructed by Smith) states that there is a relationship between the pattern (or relative order) of eruption of molar versus secondary (replacement) teeth and the overall pace (or absolute timing) of growth and maturation. Species with 'fast' life histories (rapid dental development, rapid growth, early sexual maturation, short life spans) are said to exhibit relatively early eruption of the molars and late eruption of the secondary replacement teeth (premolars, canines, incisors), whereas species with 'slow' life histories are said to exhibit relatively late eruption of the molars and early eruption of the secondary dentition. In a recent review, B.H. Smith noted that primates with tooth combs might violate this rule because tooth combs tend to erupt early, regardless of the pace of life history. We show that exceptions to Schultz's rule among lemurs are not limited to the relative timing of eruption of the tooth comb. Rather, among lemurs, some species with extremely accelerated dental development exhibit a pattern of eruption of molars and of secondary teeth in direct opposition to the expectations of Schultz's rule. We focus particularly on the pattern (order) and pace (absolute timing) of dental development and eruption in Avahi and Lepilemur - two relatively small, nocturnal folivores with rapid dental development. These taxa differ markedly in their eruption sequences (the premolars erupt after M2 and M3 in Lepilemur but not Avahi ). We offer an explanation for the failure of Schultz's rule to predict these differences. Schultz's rule presumes that eruption timing is dependent on the size of the jaw and that, therefore, molar crown formation and eruption will be delayed in species with slow-growing jaws. We show that a variety of processes (including developmental imbrication) allows the crowns of permanent teeth to form and to erupt into jaws that might appear to be too small to accommodate them.     

The dental developmental status of six East African juvenile fossil hominids

Radiographs of five juvenile fossil hominids from Koobi Fora, Kenya are described and presented together with measurements and observations made on the original speciments. Data are also presented for a single specimen from Olduvai Gorge, Tanzania. Four of these specimens are attributed to Paranthropus boisei (KNM ER 812, 1477 1820 and OH 30), and are all of remarkably similar dental developmental status. Conventional age estimates for these specimens of Paranthropus based on the first permanent molar, indicate an age at death of around 2·2 to 3 years. Perikymata counts on permanent lower central incisors of these specimens also indicate an age at death between 2·5 and 3 years. Two specimens attributed to early Homo (KNM ER 820 and 1507), are dentally more mature than specimens of Paranthropus boisei described here being closer to 5 years of age. Differences between the spacing and distribution of perikymata on the surfaces of incisor teeth are now apparent between Homo, Australopithecus. Paranthropus boisei and Paranthropus robustus: these are described in this paper. Details of the dental developmental patterns of these hominids are also discussed in the light of recent publications that have presented data about hominid eruption sequences and fossil hominid growth periods.     

中国早期智人牙齿化石

中国目前较为可靠的早期智人牙齿材料应包括在巢县、许家窑、长阳、周口店新洞和桐梓发现的人类牙齿化石,代表一类与早期组早期智人相当或十分相近的古人类。丁村人类牙齿化石可能是代表晚期组早期智人也可能是代表解剖学上的现代智人。就牙齿材料而言,尚无充分的证据能表明在中国曾同时存在过两种类型的早期智人。     

The eruption pattern of the permanent incisors and first permanent molars in Australopithecus (Paranthropus) robustus

This study aims to reassess the claim that the eruption sequence of the permanent incisor and first permanent molar teeth of Australopithecus (Paranthropus) robustus is identical with that in modern Homo sapiens. Eight fossil hominid mandibles of equivalent dental developmental age were chosen for comparative study. Emphasis has been placed upon the comparative timing of events within the growth period rather than eruption sequence alone. The results of this study indicate that Homo sapiens and Australopithecus (Paranthropus) robustus share the same pattern of permanent molar and incisor eruption and that this is significantly different from the pattern of eruption shared by the great apes, Australopithecus africanus and Australopithecus afarensis.     

广西扶绥南山洞新发现的人类化石和蒙古野驴化石——末次冰期华南气候变化的哺乳动物化石新证据

广西扶绥县南山洞发现人类化石及哺乳动物化石。人类化石包括2枚智人牙齿化石,分别是左下第三臼齿和右下第二臼齿,其形态特征与广西其他山洞发现的智人化石相似,归入晚期智人。哺乳动物化石多为华南大熊猫———剑齿象动物群成员,时代属更新世晚期。该地点新发现的蒙古野驴化石属华南晚更新世动物群的首次发现。南山洞所有化石发现于洞内浅褐色砂质粘土中,堆积物第二层钙板层的铀系年龄为30—40ka。蒙古野驴的出现暗示中国南方大陆可能在末次冰期出现过干冷的气候环境。     

安徽巢湖早期智人的牙齿磨耗和早期智人前部齿工具机能假说

巢湖早期智人的前部齿重度磨耗较大可能是与前部齿行使习惯上的非咀嚼性机能和经常啃咬坚韧的食物有关。如果早期智人前部齿的工具机能被理解为既包括非咀嚼性的也包括咀嚼性的,则早期智人前部齿工具机能假说可用来解释巢湖早期智人的前部齿重度磨耗现象,而巢湖早期智人将是出自东亚的支持这一假说的首例化石材料。     

Pattern and pace of dental eruption in Tarsius

This article uses data on the dental eruption pattern and life history of Tarsius to test the utility of Schultz's rule. Schultz's rule claims a relationship between the relative pattern of eruption and the absolute pace of dental development and life history and may be useful in reconstructing life histories in extinct primates. Here, we document an unusual eruption pattern in Tarsius combining early eruption (relative to molars) of anterior replacement teeth (P2 and incisors) and relatively late eruption of the posterior replacement teeth (C, P3, and P4). This eruption pattern does not accurately predict the "slow" pace of life documented for Tarsius [Roberts: Int J Primatol 15 (1994) 1-28], nor aspects of life history directly associated with dental development as would be expected using Schultz's rule. In Tarsius, the anterior teeth and M1 erupt at an early age and therefore are not only fast in a relative sense but also fast in an absolute sense. This seems to be related to a developmental anomaly in the deciduous precursor teeth, which are essentially skipped. This decoupling among dental eruption pattern, dental eruption pace, and life history pace in Tarsius undermines the assumptions that life histories can accurately be described as "fast" or "slow" and that dental eruption pattern alone can be used to infer overall life history pace. The relatively and absolutely early eruption of the anterior dentition may be due to the utility of these front teeth in early food acquisition rather than with the pace of life history.     

Model for the examination of evolutionary trends in tooth development

Through the use of serial computerized tomography (C-t) scans, two distinct developmental stages can be identified in mature teeth. C-t scans thus provide a non-destructive method for assessing growth within individual teeth, as well as for comparison of the development of modern and fossil teeth. The second deciduous molar (DM2) and first permanent molar (M1) resemble one another morphologically, despite differences in size and developmental rates. Thus, they provide an excellent model for studying variation in growth within an individual. To test the C-t method, we first examined a recent archaeological sample and then examined teeth from Skhul I. Serial C-t scans were used to compare two distinct developmental stages represented by the dentine-enamel junction (DEJ) and outer enamel surface (OES), respectively, in mandibular DM2 and M1 of 31 archaeological specimens. The difference in form and size between these two surfaces in and between teeth was calculated from intercusp distances measured at the DEJ and OES using the form distance matrix. Intercusp distances at the DEJ and OES of these teeth were then compared to their counterparts in the DM2 and M1 of Skhul I, taken here as representative of early anatomically modern Homo sapiens sapiens. Form differences between paired DM2 and M1 at the DEJ were smaller than those at the OES, supporting the hypothesis that differences between the two teeth increase throughout development. The increase in intercusp distances from the DEJ to OES was found to reflect the angulation of cusps relative to one another, rather than enamel thickness. Form differences between the Skhul DM2 and M1 were smaller than those observed in the recent series, and the recent M1 differed more than the DM2 from its fossil counterpart. The similarities found between the Skhul permanent and deciduous teeth and the recent DM2, may reflect a similar growth pattern. This would contribute to earlier crown completion in the fossil M1. Am. J. Phys. Anthropol. 102:283–294, 1997 © 1997 Wiley-Liss, Inc.     

Dental Ontogeny in Macroscelides proboscideus (Afrotheria) and Erinaceus europaeus (Lipotyphla)

We investigated the state of dental eruption in specimens of Macroscelides proboscideus and Erinaceus europaeus of known age. When M. proboscideus reaches adult size and sexual maturity, few or none of its replaced permanent cheek teeth have erupted. The approximate sequence of upper tooth eruption is P1, [I3, C, M1], [I1–2], M2, P4, [P2, P3]. Chronologically, E. europaeus erupts its molars and most premolars prior to M. proboscideus; but its first two upper incisors erupt after those of M. proboscideus, and its canines erupt around the same time. The approximate sequence of upper tooth eruption in E. europaeus is [M1, M2, P2, I3], C, M3, P4, P3, I2, I1. Unlike M. proboscideus, E. europaeus does not reach adult size until all permanent teeth except for the anterior incisors have erupted. While not unique among mammals, the attainment of adult body size prior to complete eruption of the permanent cheek teeth is particularly common among macroscelidids and other afrotherians.     

Foot bones from Omo: implications for hominid evolution

We reanalyze a hominid talus and calcaneus from Omo dating to 2.2 mya and 2.36 mya, respectively. Although both specimens occur at different localities and times, both tarsals articulate well together, suggesting a single taxon on the basis of size and function. We attribute these foot bones to early Homo on the basis of their morphology. The more modern-like tarsal morphology of these Omo foot bones makes them very similar to a talus from Koobi Fora (KNM-ER 813), a specimen attributed to Homo rudolfensis or Homo erectus. Although the Omo tarsals are a million years younger than the oldest known foot bones from Hadar, both localities demonstrate anatomical differences representing two distinct morphological patterns. Although all known hominid tarsals demonstrate clear bipedal features, the tarsal features noted below suggest that biomechanical changes did occur over time, and that certain features are associated with different hominid lineages (especially the robust australopithecines).     

The timing of eruption of the primary dentition in Nigerian children

The objectives of this study were to clarify the eruption time and sequence for primary teeth in Nigerian children. It also investigated the effect of sex and socioeconomic status on the timing and sequence of eruption. A random sample of 1,657 children from ages of 3-40 months were examined--921 (55.6%) males and 736 (44.4%) females. The age of eruption of the teeth was estimated using probit regression. The results show that there was no effect of sex, socioeconomic status or breastfeeding status on the timing of eruption and pattern of teeth eruption in Nigerian children. Left and right teeth had similar eruption times. Eruption times of the lateral incisor, canine, and molars were similar for upper and lower teeth. Interpopulation studies however showed that though the sequence of eruption of primary teeth in Nigerian population is similar to that of their peers in other compared populations, there are observable sex differences in the timing of tooth eruption.     

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.     

Dental formulae and dental eruption patterns in Parapithecidae (Primates, Anthropoidea)

The eruption sequence for the lower teeth of Apidium phiomense based on 18 juvenile specimens is dP3, dP4, M1, M2, P2, P4, (P3, M3), C. Only five specimens of Parapithecus grangeri show developing lower teeth. P2, M1, and M2 all erupted before P3 and P4; C and M3 were the last cheek teeth to erupt. Late eruption of the lower canines in parapithecids is a possible shared derived resemblance linking these species with Anthropoidea and Adapidae and distinguishing both from Omomyidae, Tarsiidae, and tooth-combed lemurs. Late eruption of M3 in parapithecids is a shared derived resemblance with Anthropoidea alone. The lower dental formula of Apidium phiomense is confirmed as 2 X 1 X 3 X 3 by additional specimens which show the incisors. Based in part on tooth socket counts, the deciduous lower dental formula was 2 X 1 X 3. New specimens of Parapithecus grangeri now demonstrate an adult mandibular dental formula of 0 X 1 X 3 X 3 (not 2 X 1 X 3 X 3 as previously thought) and a juvenile formula of 1 X 1 X 3. The number of incisors possessed by Parapithecus fraasi is again open to debate. Material is insufficient to judge whether this species had a pair of incisors in each lower jaw quadrant, by analogy with Apidium, or had undergone reduction to just one incisor. In any event, the presence of two incisors in another parapithecid Apidium shows anterior tooth reduction of Parapithecus grangeri occurred independent of, and should not be considered a shared derived similarity with, Tarsiidae, as was once thought.     

Human Life History Evolution Explains Dissociation between the Timing of Tooth Eruption and Peak Rates of Root Growth

We explored the relationship between growth in tooth root length and the modern human extended period of childhood. Tooth roots provide support to counter chewing forces and so it is advantageous to grow roots quickly to allow teeth to erupt into function as early as possible. Growth in tooth root length occurs with a characteristic spurt or peak in rate sometime between tooth crown completion and root apex closure. Here we show that in Pan troglodytes the peak in root growth rate coincides with the period of time teeth are erupting into function. However, the timing of peak root velocity in modern humans occurs earlier than expected and coincides better with estimates for tooth eruption times in Homo erectus. With more time to grow longer roots prior to eruption and smaller teeth that now require less support at the time they come into function, the root growth spurt no longer confers any advantage in modern humans. We suggest that a prolonged life history schedule eventually neutralised this adaptation some time after the appearance of Homo erectus. The root spurt persists in modern humans as an intrinsic marker event that shows selection operated, not primarily on tooth tissue growth, but on the process of tooth eruption. This demonstrates the overarching influence of life history evolution on several aspects of dental development. These new insights into tooth root growth now provide an additional line of enquiry that may contribute to future studies of more recent life history and dietary adaptations within the genus Homo.     

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.