On the eruption pattern of the permanent incisors and first permanent molars in Paranthropus

It has been claimed recently that Australopithecus exhibited a pattern of permanent tooth eruption like that of extant great apes, whereas a significantly different pattern was shared by Paranthropus and Homo (Dean, 1985). More particularly, each of the four Paranthropus specimens examined in that study was held to show advanced development and eruption of the permanent incisors relative to the first molar. It is demonstrated here that the eruption sequence that was posited for at least one of these four Paranthropus specimens (SK 61) is clearly erroneous, while the developmental/eruption sequences manifested by the other three specimens would appear to be more ambiguous than was claimed. Another juvenile specimen of Paranthropus (KNM-ER 1820) that was not included in Dean's study also does not necessarily support the eruption pattern that was said to characterize that taxon.     

Enamel-dentine junction (EDJ) morphology distinguishes the lower molars of Australopithecus africanus and Paranthropus robustus

Tooth crown morphology plays a central role in hominin systematics, but the removal of the original outer enamel surface by dental attrition often eliminates from consideration the type of detailed crown morphology that has been shown to discriminate among hominin taxa. This reduces the size of samples available for study. The enamel-dentine junction (EDJ) is the developmental precursor and primary contributor to the morphology of the unworn outer enamel surface, and its morphology is only affected after considerable attrition. In this paper, we explore whether the form of the EDJ can be used to distinguish between the mandibular molars of two southern African fossil hominins: Paranthropus (or Australopithecus) robustus and Australopithecus africanus. After micro-computed tomographic scanning the molar sample, we made high-resolution images of the EDJ and used geometric morphometrics to compare EDJ shape differences between species, in addition to documenting metameric variation along the molar row within each species. Landmarks were collected along the marginal ridge that runs between adjacent dentine horns and around the circumference of the cervix. Our results suggest that the morphology of the EDJ can distinguish lower molars of these southern African hominins, and it can discriminate first, second, and third molars within each taxon. These results confirm previous findings that the EDJ preserves taxonomically valuable shape information in worn teeth. Mean differences in EDJ shape, in particular dentine horn height, crown height, and cervix shape, are more marked between adjacent molars within each taxon than for the same molar between the two taxa.     

Protostylid expression at the enamel-dentine junction and enamel surface of mandibular molars of Paranthropus robustus and Australopithecus africanus

Distinctive expressions and incidences of discrete dental traits at the outer enamel surface (OES) contribute to the diagnoses of many early hominin taxa. Examination of the enamel-dentine junction (EDJ), imaged non-destructively using micro-computed tomography, has elucidated the morphological development of dental traits and improved interpretations of their variability within and among taxa. The OES expressions of one of these dental traits, the protostylid, have been found to differ among African Plio-Pleistocene fossil hominin taxa. In this study protostylid expression is examined at the OES and at the EDJ of Paranthropus robustus (n = 23) and Australopithecus africanus (n = 28) mandibular molars, with the goals of incorporating EDJ morphology into the definition of the protostylid and assessing the relative contribution of the EDJ and enamel cap to its expression in these taxa. The results provide evidence a) of statistically significant taxon-specific patterns of protostylid morphology at the EDJ that are not evident at the OES; b) that in P. robustus, thick enamel reduces the morphological correspondence between the form of the protostylid seen at the EDJ and the OES, and c) that if EDJ images can be obtained, then the protostylid retains its taxonomic value even in worn teeth.     

Alleged synapomorphy of the M1/I1 eruption pattern in robust australopithecines and Homo: evidence from high-resolution computed tomography

Ever since Broom and Robinson (1951) published their claim that the eruption pattern of permanent incisors in robust australopithecines was most similar to that of modern man and different from that of gracile australopithecines and apes, the accuracy of this observation has been the subject of periodic debate (e.g., Wallace: Ph.D. thesis, 1972; Dean: Am. J. Phys. Anthropol. 67:251-257, 1985; Grine: Am. J. Phys. Anthropol. 72:353-359, 1987). Part of the problem is that the developing incisors in one of the specimens most crucial to this argument (SK61) are difficult to visualize clearly by conventional radiographic techniques because of the heavy mineralization in the fossil. This study reanalyzes SK 61 by high-resolution computed tomography in order to contribute to the final resolution of its incisor development. Grine's (op. cit.) assessment of the incisors as the deciduous ones, not the permanent ones, is fully confirmed. This fact, in conjunction with the observation that permanent incisor root formation had only just commenced in this specimen, further weakens the argument of M1/I1 eruption pattern synapomorphy between Homo and robust australopithecines.     

New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus

A new complete hallucal metatarsal (SKX 5017) was recovered from the "lower bank" of Member 1 at Swartkrans (ca. 1.8 m.y. BP). The new metatarsal is attributed to Paranthropus robustus, the predominant hominid found in Member 1 (greater than 95% of hominid individuals). SKX 5017 is similar to Olduvai Hominid 8-H from bed I, Olduvai (ca. 1.76 m.y. BP), and both resemble humans most closely among extant hominoids. The base, shaft, and head of SKX 5017 suggest human-like foot posture and a human-like range of extension (= dorsiflexion) at the hallucal metatarsophalangeal joint, while at the same time the distal articular surface indicates that a human-like toe-off mechanism was absent in Paranthropus. The fossil evidence suggests that Homo habilis and Paranthropus may have attained a similar grade of bipedality at roughly 1.8 m.y. BP.     

Changes in the eruption order of the first permanent tooth and their relation to season of birth in Japan

In order to examine the eruption order of the first two permanent teeth, kindergarten children 5 to 6 years old were examined in Hakone, Japan. Among a total of 817 children examined from 1976 to 1984, 349 were determined as I-type children, whose mandibular first incisor erupted earlier than the mandibular first molar, and 183 were as M-type children, whose first molar erupted earlier in the mandible. The mandibular I-type rate, i.e., the proportion of the I-type among a total of I- and M-type children, was 66% (349/532). In 1983-1984, the I-type rate was 70% among boys and 62% among girls, but the overall I-type rate did not differ significantly by the sex of the subject or by the year of examination. The I-type rate varied significantly with the season of subject's birth. Those born in October or November showed a significantly lower I-type rate (33%) than the other subjects (P less than 0.001), in spite of a shift of birth season with low I-type rate toward winter in 1983-1984. The difference of the I-type rate according to birth season suggests that the causes responsible for this change are primarily environmental and act at the prenatal or perinatal stage of life.     

Relationship between enamel formation and eruption rate in rat mandibular incisors

Summary The relationship between the formation of dental enamel and tooth eruption was investigated. Rat mandibular incisor eruption rate was accelerated by maintaining incisors out of occlusion. Rate of eruption, enamel thickness, secretory zone length and matrix breakdown were measured. Eruption rate increased by 120% in experimental teeth but enamel secretion increased by only 90%. There were no obvious differences between control and experimental teeth in final enamel thickness or in the molecular weight distribution of the enamel matrix proteins.     

Age at death of the Neanderthal child from Devil's Tower, Gibraltar and the implications for studies of general growth and development in Neanderthals

This study combines traditional methods of assessing dental developmental status based upon modern human standards with new techniques based upon histological observations in order to reassess the age at death of the Gibraltar child from Devil's Tower. The results indicate that the most likely age of this individual at death was 3 years of age. This result is in agreement with an independent assessment of the age of the temporal bone of this specimen (Tillier, AM [1982] Z. Morphol. Anthropol. 73:125-148) and is concordant with dental developmental ages given for modern humans. Moreover, the fact that this specimen appears at the low end of the age scale for calcification stages in modern humans is also supportive of the findings of Legoux (Legoux, P [1970] Arch. Inst. Paleontol. Hum. Mem. 33:53-87) and Wolpoff (Wolpoff, MH [1979] Am. J. Phys. Anthropol. 50:67-114) that dental eruption schedules in Neanderthals were also accelerated. If the cranial bones from Devil's Tower are associated with the dental material, as we believe, they indicate a remarkably precocious brain growth in this individual, which is consistent with what is known about general growth and development in Neanderthals.     

Gingival eruption sequences of permanent teeth in early hominids.

Gingival eruption of the permanent teeth of Australopithecus, Paranthropus, and early Homo, diagnosed from enamel scratches and facets, is similar save for two sequences: eruption of the canines relative to the premolars which may be sexually dimorphic; and agenesis of M3 with delayed eruption of M2, first seen in Homo at two million years. Gingival eruption sequences are similar also for early and modern Homo, except that in some individuals today M3, M2, M1 and I2 take longer to form and emerge through the gingiva as functioning teeth. Probably, from two million years to the present in the evolutionary history of Homo dental development slowed-down. More and more of ontogeny has been taken over for eruption.     

Periostin is an extracellular matrix protein required for eruption of incisors in mice

A characteristic tooth of rodents, the incisor continuously grows throughout life by the constant formation of dentin and enamel. Continuous eruption of the incisor is accompanied with formation of shear zone, in which the periodontal ligament is remodeled. Although the shear zone plays a role in the remodeling, its molecular biological aspect is barely understood. Here, we show that periostin is essential for formation of the shear zone. Periostin-/- mice showed an eruption disturbance of incisors. Histological observation revealed that deletion of periostin led to disappearance of the shear zone. Electron microscopy revealed that the disappearance of the shear zone resulted from a failure in digestion of collagen fibers in the periostin-/- mice. Furthermore, immunohistochemical analysis using anti-periostin antibodies demonstrated the restricted localization of periostin protein in the shear zone. Periostin is an extracellular matrix protein, and immunoelectron microscopy showed a close association of periostin with collagen fibrils in vivo. These results suggest that periostin functions in the remodeling of collagen matrix in the shear zone.     

Health condition of first permanent molars in year 1977 and 2007 in children in Istria (Croatia)

The aim of this study was to examine differences between health condition of the first permanent molar (M1) in children in 1977 and 2007. The materials for the study consisted of data on the health condition of M1 determined in a study in 1977 (Group I) for children from the district of Buje in Istria. The health condition of M1 was examined again in the same area in 2007 (Group II). The first permanent molar is most frequently affected by caries and represent a good indicator for general caries incidence of children. Study included 709 subjects in Group I (363 boys, 346 girls) and 460 subjects in Group II (242 boys, 218 girls), aged from 6 years and 0 months to 12 years and 5 months. The difference in the frequency of intact, decayed, filled and missingd M1 was examined in both groups. Chi2 test was used to determine the differences between the number of I (intact), D (decayed), F (filled) and M (missing) teeth for each age group in Group I and Group II. In Group I there were 29.3% intact, 48.9% decayed, 17.4% filled and 4.3% missing M1, and in Group II there were 53.0% intact, 22.6% decayed, 22.1% filled and 2.1% missing M1. During the period of 30 years, a significant increase of number of dental surgeries, and thus better preventive and health education, resulted in the significant increase in the number of intact (24.0%) and filled (4.7%) M1, and decrease in the number of decayed (26.3%) and missing (2.2%) M1. From 1977 to 2007, the number of intact M1 in group II increased considerably according to group I, while the number of decayed M1 in group II significant decreased according to group I. These changes were the result of a considerably increased number of dental surgeries.     

Three-rooted mandibular first permanent molars and the question of American Indian origins

Three-rooted mandibular first molars (3RM1) are characteristic of Asian and Asian-derived populations, particularly Aleuts (whose 3RM1 frequency is the highest in the world) and Eskimos. Similarities in the frequency of these teeth between American Indians and contemporary peoples of southeastern Asia indicate a closer relation between these groups than between American Indians and Aleut-Eskimos. Three-rooted mandibular first molar frequency does not differ significantly in males and females except in Aleut-Eskimos. Bilateral asymmetry of 3RM1 is relatively frequent in both sexes and all groups. All American Indian groups examined have a low frequency of 3RM1 pointing to a single Asian origin, except Athabaskan-speaking Arizona Navajos, whose 3RM1 frequency approaches that of Aleut-Eskimos. There is no evidence at present of any significant local microevolution of 3RM1 in two testable prehistoric American Indian groups, although genetic drift had possibly occurred in a few series of 3RM1-deficient southwestern U. S. prehistoric Western Pueblo Indians. No adaptive value can be found for 3RM1 in Indians. In prehistoric western U. S. Indians geographic frequency variation is only slightly greater than the very slight (and non-significant) testable temporal variation. Three migrations from Asia seem best to explain New World 3RM1 frequency variation.     

Indications of habitat association of Australopithecus robustus in the Bloubank Valley, South Africa

Establishing the habitat preferences of early hominin taxa is a necessary, though difficult, requirement for understanding the interaction between environmental change and hominin evolution. The environments typically associated with Australopithecus robustus have been reconstructed as predominantly open grasslands situated within a habitat mosaic that included a more wooded component with a nearby perennial water source. Most studies have concluded that the open grassland component represents the habitat preference of the hominins. In this study we investigate indicators of habitat association of A. robustus that are preserved in the animal paleocommunities represented in a series of fossil cave infills in the Bloubank Valley of South Africa, including Swartkrans, Sterkfontein, Kromdraai, and Coopers. Testing for conditions of isotaphonomy reveals a potential bias relating to depositional matrix and perhaps accumulating agent, though such a bias has not unduly influenced the taxonomic composition the assemblages. Correspondence analysis of census data from modern African nature reserves demonstrates that carnivore predation patterns are indicative of animal communities, which in turn are representative of habitats. As a result, modern census data are used to document patterns of habitat preference of large herbivores, thus allowing assignment of fossil taxa to a series of broadly defined habitat categories. Correspondence analysis of fossil assemblages reveals that the abundance profile of A. robustus is most similar to that of woodland-adapted taxa. In addition, fluctuations in the relative abundance of taxa assigned to the broad habitat categories reveal a significant negative correlation between A. robustus and open grassland-adapted taxa, indicating that the more grassland-adapted taxa there are in a given assemblage, the fewer hominins there tend to be. Thus, it appears that the open grasslands that comprise the majority of the paleoenvironments associated with A. robustus do not necessarily indicate the habitat preference of the hominins. Rather, it would appear that in addition to being dietary generalists, A. robustus were also likely to have been habitat generalists.     

The relationship of handedness to asymmetry in the occlusal morphology of first permanent molars.

Handedness has been shown to be related to a number of systematic asymmetries in body dimensions, dermatoglyphic patterns and cerebral morphology. The aim here was to compare linear and angular tooth crown asymmetries of the permanent molars in healthy right-handed and left-handed subjects. The material comprised 27 children with recorded concordant left-side dominance of hand, eye and foot. The controls were an age- and sex-matched group with right side dominance. The material is based on the Collaborative Perinatal Project where detailed medical records and the dentitions, including accurate dental impressions, of over two thousand American children were examined in the USA in the sixties. Machine vision technique was used to obtain accurate three-dimensional information from the occlusal surfaces of the first permanent upper and lower molars. The directional asymmetry values of angular measurements of mandibular first molars showed evidence of asymmetry of opposite direction between the two examined groups. The results indicate that occlusal morphology of first permanent molars may be affected by handedness, and this tendency is most evident in the angular measurements of the mandibular molars. Fluctuating asymmetry did not differ significantly between the examined groups.     

A radiographic and histological study of modern human lower first permanent molar root growth during the supraosseous eruptive phase

There is increasing focus on the relationship between root growth and the eruptive process in studies of primate dental development, and the first permanent molar (M1) is regarded as a key tooth in many of these comparative studies. In this study of modern human M(1)s, histological and radiographic data were compared. Rates of root extension were determined histologically in 20 M(1)s from individuals of known sex using data for daily incremental markings and the orientation of accentuated lines in root dentine. Mean values at the mesiobuccal enamel cervix were 4.3-5.4 microm per day and then rose to a maximum of 6.7-8.4 microm per day during the first 5mm of root growth before gradually declining again to 2.8-3.6 microm per day towards apical closure. A sample of 101 orthopantomograms of children, where M(1)s were between the stages of alveolar eruption and complete eruption, were then used to determine total mesial tooth height and mesial and distal root lengths at four successive stages of eruption. At complete eruption, mean values for mesial and distal root lengths were 8-10mm, respectively. Expressed as a percentage total of mesial tooth height these averaged 45.6-56.2%. Maximum rates of M(1) eruption occur just prior to gingival emergence but did not coincide with maximum rates of root extension in this study. These results emphasise that rates of eruption and rates of root growth do not follow the same pattern of change during the supraosseous eruptive phase. They highlight the need for greater consideration of the role of the eruptive process in explaining differences in gingival emergence times in comparative studies of modern humans and fossil hominins.     

Milk molars or extra premolars in Mesotheriinae (Mesotheriidae,Notoungulata): New insights into an old controversy

A recently recovered specimen of Mesotheriinae (Mesotheriidae, Notoungulata) from the late Miocene-early Pliocene of La Rioja Province (Argentina), CRILAR Pv 433, corresponds to an individual with three upper premolars, which appears to be an “anomaly” among mesotheriines. The detailed study of this specimen, however, brings up an old controversy on the interpretation of different mesotheriine specimens with three upper or two lower premolars. After being described as different taxa, these were later considered to be juvenile representatives of other known species. The three upper or two lower teeth were interpreted as the milk molars DP2–4 and dp3–4, respectively, which would be replaced in adult life by two upper (P3–4) and one lower (p4) permanent premolars. The new material leads us to set up a different interpretation. In our opinion, all these specimens actually preserve the permanent dentition, corresponding to different ontogenetic stages of more or less young individuals. This consideration implies the necessity of a deep systematic revision of the whole subfamily, keeping in mind this new point of view and the ontogenetic variation within a species. Therefore, the presence of P2/p3 is not enough to define a different taxon at this moment. Pending this taxonomic revision, the dental morphology of CRILAR Pv 433 resembles both Typotheriopsis (e.g., upper premolars with one labial sulcus) and Pseudotypotherium (e.g., P4 with lingual groove, wide median lobe of M3) as these two late Miocene genera are currently characterized. Furthermore, P2/p3 could be expelled soon in the lifetime of individuals, and even the presence of P2/p3 could be a variable character within the same taxon; if so, this might reflect an evolutionary trend to the loss of a dental element within mesotheriines, but the revision of a large sample is necessary to support or reject these hypotheses.     

The effect of rate of eruption on periodontal ligament glycosylaminoglycan content and enamel formation in the rat incisor

The rate of eruption of rat mandibular incisors was either increased by cutting one tooth out of occlusion or eliminated by means of pinning. The effects of such changes in eruption rate on the sulphated glycosylaminoglycan content of the periodontal ligaments was analysed. The length of the enamel secretory zone and the composition of the developing enamel matrix protein was also compared. Sulphated glycosylaminoglycan content of the periodontal ligament increased fourfold (P<0.001) during accelerated eruption but decreased to a corresponding extent (P<0.001) in the absence of eruption, when compared with controls. The length of the enamel secretory zone was also significantly reduced in the immobilised teeth, although the protein content was similar compared with controls. The results demonstrate the differential response to varied eruption rates of the periodontal ligament and enamel, particularly in respect of the extracellular matrix. The data are consistent with the view that the ground substance of the periodontal ligament plays a role in the generation of the eruptive force.