Mandibular premolar and second molar root morphological variation in modern humans: What root number can tell us about tooth morphogenesis

This investigation of modern human mandibular premolar root variation tests the hypothesis that population-specific mandibular single-rooted premolar root size can predict a predisposition to root morphological complexity. Mandibular postcanines were examined and quantified from dental radiographs in a globally spread sample of 1,615 modern humans. Multirooted premolars and a fused molar root phenotype were investigated as probes into greater than, and less than, the normative root number. Twelve questions were addressed concerning root structure of mandibular premolars and second molars. A direct correlation was found between single-rooted mandibular premolar size and the predisposition to multirootedness. This correlation infers the following: 1) that postcanine primordia size during root formation predisposes to the development of more, or less, than the normative postcanine root number; and 2) that the epigenetic effect of tooth primordium size per se influences the induction of interradicular processes, which divides the root during its development. This simple developmental model helps explain the following observations: 1) population-specific variation in postcanine root number; 2) sexual dimorphism for multirooted mandibular premolar prevalence; 3) why microdont teeth are single-rooted; 4) the hierarchy of developmental canalization of interradicular processes; 5) megadont-hominin to late-hominin mandibular premolar root number transition; and 6) the fluctuation of mandibular premolar root number in primate evolutionary history.     

Subocclusal dental morphology of sahelanthropus tchadensis and the evolution of teeth in hominins

The evolution of the teeth in hominins is characterized by, among other characters, major changes in root morphology. However, little is known of the evolution from a plesiomorphic, ape‐like root morphology to the crown hominin morphology. Here we present a study of the root morphology of the Miocene Chadian hominin Sahelanthropus tchadensis and its comparison to other hominins. The morphology of the whole lower dentition (I₁–M₃) was investigated and described. The comparison with the species Ardipithecus kaddaba and Ardipithecus ramidus indicates a global homogeneity of root morphology in early hominins. This morphology, characterized notably by a reduction of the size and number of the roots of premolars, is a composite between an ape‐like morphology and the later hominin morphology. Trends for root evolution in hominins are proposed, including the transition from a basal hominoid to extant Homo sapiens. This study also illustrates the low association between the evolution of tooth root morphology and the evolution of crowns in hominins. Am J Phys Anthropol 153:116–123, 2014. © 2013 Wiley Periodicals, Inc.     

A geometric morphometric analysis of hominin upper premolars. Shape variation and morphological integration

This paper continues the series of articles initiated in 2006 that analyse hominin dental crown morphology by means of geometric morphometric techniques. The detailed study of both upper premolar occlusal morphologies in a comprehensive sample of hominin fossils, including those coming from the Gran Dolina-TD6 and Sima de los Huesos sites from Atapuerca, Spain, complement previous works on lower first and second premolars and upper first molars. A morphological gradient consisting of the change from asymmetric to symmetric upper premolars and a marked reduction of the lingual cusp in recent Homo species has been observed in both premolars. Although percentages of correct classification based on upper premolar morphologies are not very high, significant morphological differences between Neanderthals (and European middle Pleistocene fossils) and modern humans have been identified, especially in upper second premolars. The study of morphological integration between premolar morphologies reveals significant correlations that are weaker between upper premolars than between lower ones and significant correlations between antagonists. These results have important implications for understanding the genetic and functional factors underlying dental phenotypic variation and covariation.     

Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia

The hominin fossils of Dmanisi, Republic of Georgia, present an ideal means of assessing levels of skeletal size and shape variation in a fossil hypodigm belonging to the genus Homo because they have been recovered from a spatially and temporally restricted context. We compare variation in mandible size and shape at Dmanisi to that of extant hominoids and extinct hominins. We use height and breadth measurements of the mandibular corpus at the first molar and the symphysis to assess size, and analyze shape based on size-adjusted (using a geometric mean) versions of these four variables. We compare size and shape variation at Dmanisi relative to all possible pairs of individuals within each comparative taxon using an exact resampling procedure of the ratio of D2600 to D211 and the average Euclidean distance (AED) between D2600 and D211, respectively. Comparisons to extant hominoids were conducted at both the specific and subspecific taxonomic levels and to extinct hominins by adopting both a more, and less speciose, hominin taxonomy. Results indicate that the pattern of variation for the Dmanisi hominins does not resemble that of any living species: they exhibit significantly more size variation when compared to modern humans, and they have significantly more corpus shape variation and size variation in corpus heights and overall mandible size than any extant ape species. When compared to fossil hominins they are also more dimorphic in size (although this result is influenced by the taxonomic hypothesis applied to the hominin fossil record). These results highlight the need to re-examine expectations of levels of sexual dimorphism in members of the genus Homo and to account for marked size and shape variation between D2600 and D211 under the prevailing view of a single hominin species at Dmanisi.     

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.     

陕西彬县晚中新世猪类化石

本文记述了陕西彬县发现的晚中新世哺乳动物群中的猪类化石,共两属两种.双齿尖河猪 (Dicoryphochoerus binxianensis),为—新种其性质较印度 Chinji 组的 D. chisholmi 进步,而比我国蓝田组的 D. medius 又略显得原始;弓颌猪 (Chleuastochoerus stehlini) 是我国特有的一个属种,是北方三趾马动物群中的成员.     

新发现的和县直立人牙齿化石

本文对一颗新发现的安徽和县直立人上第三前臼齿的尺寸、形态特征、齿冠外轮廓形状和齿尖排列样式进行了观测, 并与世界范围内相关古人类标本进行了对比。结果发现, 新发现的和县直立人P3齿冠尺寸较大, 同一地点还发现了另外一颗尺寸较大的P3, 两颗牙齿基本是目前发现的中国直立人和早期智人标本中齿冠尺寸最大的, 显示其较为原始的一面。在齿冠外轮廓形状和齿尖排列样式上, 新发现的和县直立人标本表现出与世界各地其他直立人的一致性, 但齿冠颊侧面显著发育的近中纵向沟将这颗牙齿归入到了亚洲直立人的变异范围内, 与非洲和Dmanisi直立人区别。中国直立人P3在齿冠尺寸、横脊发育与否、颊侧面近远中纵向沟发育程度、齿根数目、齿冠外轮廓形状和齿尖排列样式这些在人类演化过程中具有明显演化变化趋势的特征上表现出较大的变异, 和县这颗牙齿处于中国直立人总体变异范围内较原始的一侧。与周口店直立人相比,新发现的和县直立人标本和同一地点发现的另一颗P3部分形态特征较原始, 这与和县人头骨形态特征相对周口店直立人进步的趋势相反。新发现的这颗牙齿及其他和县直立人化石对探讨东亚地区直立人的起源和地区性差异具有重要意义。     

Postnatal Cranial Development in Papionin Primates: An Alternative Model for Hominin Evolutionary Development

The evolution of hominin growth and life history has long been a subject of intensive research, but it is only recently that paleoanthropologists have considered the ontogenetic basis of human morphological evolution. To date, most human EvoDevo studies have focused on developmental patterns in extant African apes and humans. However, the Old World monkey tribe Papionini, a diverse clade whose members resemble hominins in their ecology and population structure, has been proposed as an alternative model for human craniofacial evolution. This paper reviews prior studies of papionin development and socioecology and presents new analyses of juvenile shape variation and ontogeny to address fundamental questions concerning primate cranial development, including: (1) When are cranial shape differences between species established? (2) How do epigenetic influences modulate early-arising pattern differences? (3) How much do postnatal developmental trajectories vary? (4) What is the impact of developmental variation on adult cranial shape? and, (5) What role do environmental factors play in establishing adult cranial form? Results of this inquiry suggest that species differences in cranial morphology arise during prenatal or earliest postnatal development. This is true even for late-arising features that develop under the influence of epigenetic factors such as mechanical loading. Papionins largely retain a shared, ancestral pattern of ontogenetic shape change, but large size and sexual dimorphism are associated with divergent developmental trajectories, suggesting differences in cranial integration. Developmental simulation studies indicate that postnatal ontogenetic variation has a limited influence on adult cranial morphology, leaving early morphogenesis as the primary determinant of cranial shape. The ability of social factors to influence craniofacial development in Mandrillus suggests a possible role for phentotypic plasticity in the diversification of primate cranial form. The implications of these findings for taxonomic attribution of juvenile fossils, the developmental basis of early hominin characters, and hominin cranial diversity are discussed.     

Sexual dimorphism of the canine roots in theMacaca fuscata

In situ radiographic analysis of the maxillary canines ofMacaca fuscata was conducted on 88 specimens in 44 individuals (23 dry skulls and 21 live animals) in order to examine the number of roots. The left canines were then extracted from ten female skulls for measurement, further radiographic examination, and visual morphological observation. The results showed a clear sexual dimorphism in root morphology: all male canines were clearly distinguished as single-rooted from the radiograph, whereas more than 40% of the female canines were double-rooted. Variation was also found among the single-rooted female canines, in that some of these teeth appeared to have a bifurcated canal. This sexual dimorphism in the number of maxillary canine roots and the individual variation found among the females in root and canal morphology are previously unreported for this species. No observations were attempted on mandibular canines, however, because of the incomplete nature of the sample.     

山西垣曲先炭兽类一新种

本文主要报道1983年9月,在山西省垣曲县寨里发现的一件较完整的先炭兽类老年个体上颌骨.带有颇为完整的左、右齿列. I~1-P~2 各齿前、后均有较长的齿隙. P~1、P~2、P~3 前的齿隙颌骨上,保留有乳前臼齿的齿槽.这块标本在大小及形态特征上,与目前所知先炭兽属 Anthracokeryx 中任何种都有相当大的差异.它代表了该属中的一个新种.     

Dietary correlates of temporomandibular joint morphology in the great apes

Behavioral observations of great apes have consistently identified differences in feeding behavior among species, and these differences have been linked to variation in masticatory form. As the point at which the mandible and cranium articulate, the temporomandibular joint (TMJ) is an important component of the masticatory apparatus. Forces are transmitted between the mandible and cranium via the TMJ, and this joint helps govern mandibular range of motion. This study examined the extent to which TMJ form covaries with feeding behavior in the great apes by testing a series of biomechanical hypotheses relating to specific components of joint shape using linear measurements extracted from three‐dimensional coordinate data. Results of these analyses found that taxa differ significantly in TMJ shape, particularly in the mandibular fossa. Chimpanzees have relatively more anteroposteriorly elongated joint surfaces, whereas gorillas tend to have relatively anteroposteriorly compressed joints. Orangutans were most commonly intermediate in form between Pan and Gorilla, perhaps reflecting a trade‐off between jaw gape and load resistance capabilities. Importantly, much of the observed variation among taxa reflects differences in morphologies that facilitate gape over force production. These data therefore continue to emphasize the unclear relationship between mandibular loading and bony morphology, but highlight the need for further data regarding food material properties, jaw gape, and ingestive/food processing behaviors. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

Geometric morphometric analysis of the crown morphology of the lower first premolar of hominins, with special attention to Pleistocene Homo

This article is the third of a series that explores hominin dental crown morphology by means of geometric morphometrics. After the analysis of the lower second premolar and the upper first molar crown shapes, we apply the same technique to lower first premolar morphology. Our results show a clear distinction between the morphology seen in earlier hominin taxa such as Australopithecus and African early Homo, as well as Asian H. erectus, and more recent groups such as European H. heidelbergensis, H. neanderthalensis, and H. sapiens. The morphology of the earlier hominins includes an asymmetrical outline, a conspicuous talonid, and an occlusal polygon that tends to be large. The morphology of the recent hominins includes a symmetrical outline and a reduced or absent talonid. Within this later group, premolars belonging to H. heidelbergensis and H. neanderthalensis tend to possess a small and mesiolingually-displaced occlusal polygon, whereas H. sapiens specimens usually present expanded and centered occlusal polygons in an almost circular outline. The morphological differences among Paranthropus, Australopithecus, and African early Homo as studied here are small and evolutionarily less significant compared to the differences between the earlier and later homin taxa. In contrast to the lower second premolar and the upper first molar crown, the inclusion of a larger hominin sample of lower first premolars reveals a large allometric component.     

The coevolution of male and female genitalia in a mammal: A quantitative genetic insight

Male genitalia are among the most phenotypically diverse morphological traits, and sexual selection is widely accepted as being responsible for their evolutionary divergence. Studies of house mice suggest that the shape of the baculum (penis bone) affects male reproductive fitness and experimentally imposed postmating sexual selection has been shown to drive divergence in baculum shape across generations. Much less is known of the morphology of female genitalia and its coevolution with male genitalia. In light of this, we used a paternal half-sibling design to explore patterns of additive genetic variation and covariation underlying baculum shape and female vaginal tract size in house mice (Mus musculus domesticus). We applied a landmark-based morphometrics approach to measure baculum size and shape in males and the length of the vaginal tract and width of the cervix in females. Our results reveal significant additive genetic variation in house mouse baculum morphology and cervix width, as well as evidence for genetic covariation between male and female genital measures. Our data thereby provide novel insight into the potential for the coevolutionary divergence of male and female genital traits in a mammal.     

Symmetry of non-carious cervical lesions in canines and premolars

Objectives: Clinically non‐carious cervical lesions (NCCLs) are frequently seen. The aim of this study was to investigate the relationship between the shape and symmetry of NCCLs, wear of cuSPS and triangular ridge, and the curvature of the tooth root. Methods: One hundred and twenty‐nine extracted human upper canine teeth and 274 extracted human upper premolar teeth with NCCLs were used in this study. The specimens were studied using photographs and three‐dimensional scanning. Results: Asymmetric NCCL was observed in 69.0% of the canines and 44.5% of the premolars. Wear of cusp and lingual ridges was observed in 82.9% and 93.0% of the canines, respectively. Wear of the buccal cusp and buccal triangular ridge was observed in 85.4% and 89.8% of the premolars, respectively. On the other hand, the wear of lingual cusp and lingual triangular ridge was observed in 89.1% and 93.8% of the premolars, respectively. The curvature of the root was observed in 48.1% of the canines and 43.4% of the premolars. Conclusions: There was no relationship between the symmetry of NCCLs, and the wear of cuSPS and triangular ridges for either canines or premolars. Although there was a relationship (p < 0.05) between the symmetry of NCCL and the curvature of the root in the canines, no relationship was observed between the symmetry of NCCL and the curvature of the root in the premolars.     

Characteristic high- and low-frequency dental traits in sub-Saharan African populations

In an earlier investigation (Irish [1993] Biological Affinities of Late Pleistocene Through Modern African Aboriginal Populations: The Dental Evidence [Ann Arbor: University Microfilms]), biological affinities of 32 sub-Saharan and North African dental samples were estimated using comparative analyses of 36 dental morphological traits. Marked dental homogeneity was revealed among samples within each of the two geographic regions, but significant interregional differences were noted. Assuming dental phenetic expression approximates or is an estimate of genetic variation, the present study of 976 sub-Saharan-affiliated Africans indicates they are not closely related to other world groups; they are characterized by numerous morphologically complex crown and root traits. Turner ([1984] Acta Anthropogenetica 8:23–78; [1985] in R Kirk and E Szathmary (eds.): Out of Asia: Peopling the Americas and the Pacific [Canberra: The Journal of Pacific History], pp. 31–78; [1990] Am. J. Phys. Anthropol. 82:295–318; [1992] Persp. Hum. Biol. 2/Archaeol. Oceania 27:120–127; [1992] in T Akaszawa, K Aoki, and T Kimura (eds.): The Evolution and Dispersal of Modern Humans in Asia [Tokyo: Hokusen-Sha Publishing Co.], pp. 415–438) reports that Northeast Asian/New World sinodonts also have complex teeth relative to Europeans, Southeast Asian sundadonts, Australian/Tasmanians, and Melanesians. However, sinodonty is characterized by UI1 winging, UI1 shoveling, UI1 double shoveling, one-rooted UP1, UM1 enamel extension, M3 agenesis, and three-rooted LM1. Sub-Saharan peoples exhibit very low frequencies of these features. It is proposed that the collection of dental traits which best differentiate sub-Saharan Africans from other worldwide samples includes high frequencies of the Bushman Canine, two-rooted UP1, UM1 Carabelli's trait, three-rooted UM2, LM2 Y-groove pattern, LM1 cusp 7, LP1 Tome's root, two-rooted LM2, UM3 presence, and very low incidences of UI1 double shoveling and UM1 enamel extension. This suite of diagnostic traits is termed the sub-Saharan African dental complex. Am J Phys Anthropol 102:455–467, 1997. © 1997 Wiley-Liss, Inc.     

Hominin lower second premolar morphology: evolutionary inferences through geometric morphometric analysis

Mandibular premolars are increasingly used in taxon-specific diagnostic analyses of hominins. Among the principal difficulties in these evaluations is the absence of discrete, discernible, and comparable anatomical structures for rigorous quantitative assessment. Previous research has addressed either internal crown surface features (such as cusps and fossae) or the morphology of the crown outline. In the present paper, we integrate both types of information in the examination of morphological variation of lower P4s (n = 96) among various fossil hominin species with an emphasis on genus Homo. We use a set of 34 2D landmarks combining coordinate data from four classical dental landmarks on the occlusal surface and 30 sliding semilandmarks of the crown outline. Our results indicate that external shape variation is closely related to the configuration of the occlusal morphological features and influenced by dental size. The external and internal shapes of P4 are polymorphic but still useful in depicting a primitive-derived gradient. The primitive pattern seems to have been an asymmetrical contour with a mesially displaced metaconid, development of a bulging talonid, and a broad occlusal polygon. The trend toward dental reduction during the Pleistocene produced different morphological variants with a reduced occlusal polygon and decreased lingual occlusal surface in later Homo species. Homo heidelbergensis/neanderthalensis have fixed plesiomorphic traits in high percentages, whereas in modern humans a symmetrical outline with a centered metaconid and talonid reduction evolved.     

Discrimination of extant Pan species and subspecies using the enamel–dentine junction morphology of lower molars

Previous research has demonstrated that species and subspecies of extant chimpanzees and bonobos can be distinguished on the basis of the shape of their molar crowns. Thus, there is potential for fossil taxa, particularly fossil hominins, to be distinguished at similar taxonomic levels using molar crown morphology. Unfortunately, due to occlusal attrition, the original crown morphology is often absent in fossil teeth, and this has limited the amount of shape information used to discriminate hominin molars. The enamel–dentine junction (EDJ) of molar teeth preserves considerable shape information, particularly in regard to the original shape of the crown, and remains present through the early stages of attrition. In this study, we investigate whether the shape of the EDJ of lower first and second molars can distinguish species and subspecies of extant Pan. Micro‐computed tomography was employed to non‐destructively image the EDJ, and geometric morphometric analytical methods were used to compare EDJ shape among samples of Pan paniscus (N = 17), Pan troglodytes troglodytes (N = 13), and Pan troglodytes verus (N = 18). Discriminant analysis indicates that EDJ morphology distinguishes among extant Pan species and subspecies with a high degree of reliability. The morphological differences in EDJ shape among the taxa are subtle and relate to the relative height and position of the dentine horns, the height of the dentine crown, and the shape of the crown base, but their existence supports the inclusion of EDJ shape (particularly those aspects of shape in the vertical dimension) in the systematic analysis of fossil hominin lower molars. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Genetic and environmental variations in quantitative characters in fishes: a comparative analysis of monoclonal triploid and bisexual tetraploid spined loaches (<Emphasis Type="Italic">Cobitis</Emphasis>, Cobitidae)

The heritability estimates of 25 external morphometric characters and 23 craniometric indices are obtained by use of variances in monoclonal all-female triploids and bisexual tetraploids of spined loaches (genus Cobitis, Cobitidae) collected from the same breeding biotope. Most of studied traits demonstrate low heritability confirming previous conclusion on the similarity between external morphometric characters and craniological indices in relative effects of genetic and environmental components in their total phenotypic variation. Low heritability estimates in most of external morphological traits correspond to their low diagnostic value in Cobitis species. As a whole, in spite of certain deviations, studies on clonal forms do not refute the concept on higher heritability estimates in diagnostically significant traits in comparison with traits without diagnostic values in the same taxonomic group. Low heritability in most morphometric traits more probably is resulted from their low additive genetic variation caused by strong selection of evolutionary developed specific body shape in spined loaches, because strong selection should reduce the genetic variance in body proportions to minimal size. Sex differences observed in heritability estimates should be interpreted as a result of linkage of several additive genes controlling these traits to sex chromosomes. A few characters demonstrating high heritability estimates up to 0.492–0.580 are of great interest for taxonomic and phylogenetic studies in genus Cobitis and related taxa.