中国人牙齿形态测量分析——近代人群上、下颌前臼齿齿冠轮廓形状及其变异

采用基于标志点的几何形态测量方法对在河南和湖北地区考古遗址出土的291颗近代人群上、下颌前臼齿(P1=72; P2=72; P1=73; P2=74)齿冠轮廓形状(包括外轮廓及内轮廓形状) 进行了分析。结果显示: P1齿冠轮廓基本对称, 颊侧部分向近远中方向突出; 齿尖夹角区呈不规则狭长菱形, 舌侧尖向近中方向偏移。P2齿冠轮廓对称,颊侧部分向近远中方向的突出程度存在明显的个体差异; 齿尖夹角区由于舌侧尖向近中方向偏移而呈不规则菱形。P1和P2内外轮廓形状具有明显的重叠范围。P1齿冠轮廓不对称, 齿冠近中舌侧部分内缩; 齿尖区靠近近中舌侧, 近似方形。P2齿冠轮廓基本对称; 齿尖区靠近齿冠中央, 因近中舌侧尖向近中方向偏移而呈不规则狭长菱形。P1和P2内外轮廓形状重叠范围很小。P1、P2齿冠和齿尖夹角区各个部分所表现出来的变异大小、方式基本相同; 依据齿尖与齿冠基底边缘的距离大小可以将P1分为两种类型, 依据颊侧部分向近远中方向的突出程度可以将P2分为两类。P1、P2变异相对于上颌两颗前臼齿来说较大, 且变异主要集中在齿冠的近远中舌侧、齿尖夹角区的后凹点和舌侧尖;两者的变异大小范围稍有差别; P1和P2主要分为三类, 区别在于颊舌径和近远中径的大小比例、齿尖区后凹点和舌侧尖的位置上。异速生长分析显示所有四颗前臼齿的大小变化对轮廓形状的影响不大。     

贵州毕节韦家洞古人类遗址年代测定及人牙的形态学意义

2007年贵州毕节韦家洞首次发现古人类活动遗物。2016~2018年对遗址做深入考察,发现韦家洞是一个严重破坏的晚更新世古人类遗址,在深度破坏的扰乱层中出土了大量古人类活动证据,包括石制品、动物烧骨和人类牙齿,2018年在洞口和洞内深处发现残留的原生文化层。本文报告了韦家洞原生文化层的¹⁴C年代测定结果,距今1.2~2.7万年前,属于晚更新世末期。同时,观察描述了韦家洞出土的一枚古人类上颌第四前臼齿化石,对牙齿形态测量进行了研究比较,发现韦家洞人牙测量值落在现代中国人变异范围内,牙冠咬合面及釉质-齿质交界面EDJ形态简单,牙根短,根管简单纤细,与现代人相似。牙冠颊舌径大,具有三角形隆起和齿带,牙冠颊侧和舌侧均有垂直沟,近中和远中缘有边缘副结节,牙根中部膨大,这些特征显示了韦家洞人牙形态的原始性。综合比较国内外出土的古人类上颌第四前臼齿的形态特点及其演化趋势,结合遗址年代测定结果,我们认为韦家洞出土的人类上颌第四前臼齿形态特点体现了古人类牙齿演化的镶嵌性,为晚更新世东亚早期现代人的形态变异及其演化增添了新证据。     

禄丰古猿蝴蝶种下第四前臼齿釉质-齿质交界面的三维几何形态

禄丰古猿蝴蝶种 (Lufengpithecus hudienensis) 也称蝴蝶古猿,是重要的早期人科成员,化石产自云南元谋盆地竹棚-小河及雷老两个地点群,其年代为中新世晚期。上世纪八、九十年代的发掘工作共获得幼年个体颅骨1具、残上颌骨10件、残下颌骨17件及1500多枚单个牙齿。受限于当时的技术条件,蝴蝶古猿牙齿内部结构及三维形态一直未有报道和对比研究。本文首次使用高精度CT配合三维几何形态测量方法,对6枚产自小河地点的蝴蝶古猿下颌第四前臼齿的釉质-齿质交界面形态进行了观察和对比,对比材料包括步氏巨猿、猩猩（化石）、大猩猩、黑猩猩及现代人。多变量分析显示,蝴蝶古猿釉质-齿质交界面几何形态接近于本文所涉及的大型猿类对比标本,但并没有表现出与某一特定类群的相似性;咬合面轮廓狭长,前凹尺寸明显小于后凹;整体形态介于齿质尖较高的大猩猩和齿质尖较低、釉质-齿质交界面形态扁平的巨猿、猩猩和黑猩猩之间。本文所观察到的类群之间的异同可能与趋同演化有关,也需要更多数据的进一步验证。将釉质-齿质交界面的三维几何形态和其他牙齿内部结构的信息（如釉质厚度及其三维分布规律等）综合,有助于进一步讨论蝴蝶古猿的分类学、系统发育和食性。     

东亚中更新世古人类下颔第二臼齿釉质-齿质连接面三维形状和釉质厚度分布

东亚中更新世古人类在头骨、下颌骨、牙齿等解剖部位表现出不同程度的形态多样性,中期成员代表为直立人,而晚期成员的演化地位具有较大争议。为进一步了解东亚中更新世古人类内部的形态变异特点和为东亚中更新世晚期古人类分类提供依据,本文使用微分同胚的表面匹配（Diffeomorphic Surface Matching, DSM）和形态测量图（Morphometric map）对下颌第二臼齿（M₂）釉质-齿质连接面的形状和齿冠侧面釉质厚度分布模式进行了量化分析。结果显示：1）东亚中更新世古人类与晚期人属成员（尼安德特人和现代人）存在较明显的形态差别;2）该时段晚期的东亚古人类相对中期直立人在侧面釉质厚度分布规律上具有独特性,并在釉质-齿质连接面的三维形状上与晚期人属成员更加接近。本文在以往对东亚中更新世古人类牙齿内外结构单个性状研究的基础上,使用三维形态测量方法进一步量化了M₂釉质-齿质连接面三维形状和侧面釉质厚度分布模式两项重要特征的变异特点,这对未来该时段同类型牙齿的形态鉴定以及解决东亚中更新世晚期古人类的分类地位具有一定意义。     

A new molar from the Middle Pleistocene hominid assemblage of Yanhuidong,Tongzi, South China

1988年,贵州省博物馆对桐梓岩灰洞的支洞进行了最后一次发掘。2014年,在洞内第四层堆积物中鉴定出逾2000件的动物牙齿化石,以及一枚古人类上颊齿(编号:TZ-1)。1991年,铀系法测定这些次生堆积物的沉积年代约为距今24万年。本文运用高精度CT(巴黎自然历史博物馆)对TZ-1的釉质齿质界面(EDJ)和牙髓腔几何形态进行了分析。TZ-1的冠面形态有如下特征:次尖小且在远中舌侧不发育,咀嚼面轮廓呈四边形,颊舌径稍大过近中远中径,原尖舌侧齿带发育,齿尖从大到小依次为原尖、后尖、前尖和次尖。TZ-1牙髓腔的髓角与其釉质齿质界面以及釉质表面的形态都具有相关性。TZ-1的形态与M1虽有相似之处,但完全不同于1983年出土于同一层位的另两颗M1;其应被鉴定为dm2,并可被归入中中更新世的中国直立人支系。岩灰洞上臼齿PA 875的形态与建始龙骨洞PA 1279和周口店直立人等古老型直立人相似。岩灰洞另一枚刚萌发的臼齿PA 874具有凸出的次尖和长菱形的外廓,接近于爪哇型直立人Sangiran NG 91-G10;这也是智人和尼安德特人的共有衍征。但PA 874的冠面仍保留了亚洲型的齿带,因而被归入人属未定种。因出土自次生堆积,岩灰洞的三种古人类类型未必曾同时并存,但却揭示了华南地区人类演化进程中的多样面貌。     

河北阳原姜家梁新石器时代人类牙齿形态特征的观察与研究

本文应用牙齿人类学的研究方法,对出自河北阳原姜家梁遗址的新石器时代居民的牙齿形态特征进行了研究。采用“亚利桑那州立大学牙齿人类学系统”进行牙齿形态观察,应用“平均差异度”(MMD)距离公式计算出姜家梁组与其他各对比组的MMD距离系数以估计群体间的相似程度,并根据MMD矩阵对18组人群进行聚类分析和因子分析。本文研究结果表明,姜家梁居民的牙齿形态特征属于中国型牙(Sinodonty),但自身又具有某些独特的特征,如铲形上颌中央门齿和单根上颌第一前臼齿等特征的出现率很高。综合来看,姜家梁组在铲形上颌中央门齿、双铲形上颌中央门齿、单根上颌第一前臼齿和第一臼齿转向皱纹这4项性状上具有很高的出现率;上颌第一臼齿釉质延伸、上颌第三臼齿退化、三根下颌第一臼齿和下颌第二臼齿四尖这四项性状出现率处于中等水平。与同时代的下王岗组和庙子沟组比较而言,姜家梁组与庙子沟组的关系更为密切,有研究表明姜家梁组居民与庙子沟组居民具有较为相近的头骨形态特征,本文研究结果表明两者具有较为接近的牙齿形态特征,这说明在体质特征上相近的人群在头骨形态特征和牙齿形态特征方面的表现可能较为一致,但只通过这两个对比组的比较结果而下此结论还为时过早,尚有待于积累更多的同类型族群的相关牙齿人类学资料验证之。     

山东新泰乌珠台人类牙齿的形态学特征

近些年在东亚发现的晚更新世现代人化石及其研究使得关于该地区现代人起源问题更加复杂,更多该时段人类标本的研究有助于对其有更清晰的认识。1966年在山东新泰乌珠台发现一枚古人类下颌臼齿,吴新智和宗冠福(1973)对其进行了报道,而后再无详细研究。本文将使用牙齿非测量性状的半定量化(分级)、齿冠外轮廓形状的几何形态测量、基于显微断层扫描(Micro-computed tomography 或micro-CT)的釉质厚度、釉质厚度分布规律和齿质表面三维结构复原等方法对乌珠台人类牙齿进行综合研究,进而对东亚晚更新世古人类牙齿形态特征变异有一个进一步的了解。结果显示,乌珠台人类牙齿的形态特征基本与现代人接近,但其所表现出的三角座横脊、Y型齿沟排列、原附尖在现代人中出现率较低,而更多发现在直立人或尼安德特人中。相对于东亚其他晚更新世现代人,乌珠台M3所表现出的特征组合具有特殊性,增加了东亚晚更新世现代人的牙齿形态特征多样性。未来研究可尝试测定乌珠台人类牙齿的绝对年代,以更好的将其归入到现代人演化序列中去。     

人类牙齿齿冠和齿根分离两种技术方法对牙釉质厚度测量的影响

在基于计算机断层扫描技术（CT）和虚拟图像处理技术的灵长类牙齿测量学研究中,经常需要分离三维虚拟模型的齿冠和齿根,再进行后续测量工作,如计算机辅助的生物力学分析、釉质厚度测量等。而分离齿冠和齿根这一步骤,目前有多种方法,如,1)根据齿颈线切分齿冠,或2)人工建立基底平面切分齿冠。为了评估这两种不同的处理方式对后续的牙齿测量学上的影响,本文使用三维方法测量了82例化石和现代人类下颌后部牙齿的釉质厚度,包括南方古猿、早期人属、尼安德特人和现代人。使用配对t检验对比发现,两种方法得到的釉质厚度数值上没有显著差别,但随后进行的种间比较发现,使用基底平面切分齿冠的方法比较费时,更依赖于测量者的人工操作,并且可能弱化了物种间前臼齿绝对釉质厚度的差异,造成系统误差。其原因是对于前臼齿和前部牙齿等齿颈线形状不规则的标本,基底平面难以建立或误差较大。在未来对釉质厚度的种间差异的研究中,特别对齿颈线形状不规则的标本（如人类前部牙齿及猩猩、黑猩猩的牙齿等）,本文推荐使用齿颈线分离齿冠和齿根,测量和计算齿颈线之上的釉质厚度。釉质厚度有一定的分类学、功能形态学和系统发育学意义。本文积累了一批可供未来对比研究的原始数据,并且发现尼安德特人前臼齿的相对釉质厚度显著小于现代人,这与前人利用臼齿、犬齿所做的对比研究结果相同,支持了尼安德特人拥有较薄的相对釉质厚度这一观点。     

懒猴的牙齿形态观察

本文就云南所产懒猴的牙齿形态作初步观察。恒齿式为(2.1.3.3)/(2.1.3.3)=36;上门齿为异形齿,中门齿比侧门齿阔大,形态亦不同;上犬齿有性别差异,下犬齿特化为门齿形;前臼齿无论齿数和齿尖分布都与猕猴、金丝猴和长臂猿有较大差别,但齿尖分布却与树鼩有某些相似之点;下颌第1前臼齿犬齿化;在臼齿的齿尖分布方面,与猕猴、金丝猴相似,上颌M~3次尖架消失,只具3个齿尖。     

全新世中国北方人群第一臼齿齿冠面积和齿尖相对面积的测量分析

通过对532枚中国北方地区新石器时代到青铜时代和铁器时代人类上、下颌骨第一臼齿(M1和M1)齿冠面积和齿尖面积的测量和分析, 得到以下关于中国北方近代人群牙齿尺寸的准确数据和变异情况的结果: 两性M1和M1齿冠面积存在显著性差异, 男性齿冠面积大于女性齿冠面积, 齿尖相对面积不存在性别差异; 从新石器时代到青铜时代男性M1和M1分别缩小了1.16%和4.96%, 女性分别缩小了5.68%和6.11%,青铜时代到铁器时代齿冠面积无明显变化; M1颊侧尖相对基底面积大于舌侧尖, 形成齿尖大小差异的原因很可能是在人类演化过程中, 齿冠面积整体缩小的趋势下, 为使咬合面最大化使用, 各齿尖朝着更利于扩大咬合面基底面积的方式进化的结果; 世界各地区近代人群M1和M1各齿尖大小顺序基本一致, 其中M1为原尖>前尖>后尖>次尖, M1下原尖最大, 下次小尖最小, 其他三个齿尖面积非常接近, 不存在明显大小变化。     

The relation of canine size to pattern and extent of attrition in baboons

For most genera of animals the association between dental morphology and diet is such that inferences concerning the diet can be made on the basis of the dentition alone. The canine is the one tooth that appears exempt from such generalisation and exhibits a wide range of variability of size and shape in all groups. In order to determine the effect of dimorphism of the canine on the dental apparatus, male and female specimens of Theropithecus and Papio baboons were examined. Occlusal relationships and dental pathology for 21 Theropithecus and 40 Papio skulls were recorded, and crown height measurements obtained for maxillary incisors, buccal and lingual molar cusps. Attrition was the most common and severe cause of abscess formation in older animals: the pattern of attrition differed in adult males and females, the latter showing more wear of the incisor and lingual molar cusps. Partial correlations for incisor, buccal and lingual molar cuspal crown height show a significant correlation between incisor and lingual cusp attrition in juveniles of both sexes and adult females (p < 0.01). Adult males show no correlation of attrition. It is proposed that the correlation of incisor and lingual molar cusp attrition is associated with anterior-posterior grinding movements, such as take place during incision, that the large canine present in the male limits such movements, the sharp blade-like canine being employed as a cutting tool. This use of the canine by reducing functional demands on other teeth, prolongs the utility of the dental apparatus, and hence the lifespan of its possessor.     

Sexual dimorphism of cusp dimensions in human maxillary molars

Cusp dimensions of human maxillary molars were compared between males and females to determine whether the later-developed, distal cusps displayed greater sexual dimorphism than the earlier-developed, mesial cusps, and whether the later-forming second molar displayed greater sexual dimorphism than the first molar. First and second permanent molar crowns (M1 and M2) were measured indirectly, using dental casts obtained from 117 Japanese (65 males and 52 females). Measurements included maximum mesiodistal and buccolingual crown diameters and the diameters of the four main cusps: the paracone, protocone, metacone, and hypocone. Mean values of crown dimensions were larger in males than in females for both M1 and M2, but the sexual difference in protocone diameter of M1 was not significant. The protocone in M1 showed the least amount of sexual dimorphism, followed by the metacone, hypocone, and paracone, while in M2, the percentage sexual dimorphism corresponded to the order of cusp formation: paracone, protocone, metacone, and hypocone. With the exception of the paracone diameter, M2 showed greater sexual dimorphism than M1. Sexual dimorphism was not always greater in the later-developed, distal cusps of M1 or M2, but the protocone, the most important cusp in terms of occlusal function, displayed the least dimorphism in M1.     

Cusp size, sexual dimorphism, and heritability of cusp size in twins.

Overall measures of mandibular molars reflect the combined size contributions of the component cusps and ridges. Until now, the size hierarchy of primary and permanent mandibular molar cusps remained unclear. This paper utilizes the relative plane surface areas (basal area dimensions) of the individual molar cusps, as assays of cusp size to demonstrate cusp size variations within populations, antimere cuspal variations, sexual dimorphism, and, the heritability of cusp size. Duplicate dental casts from 199 pairs of like-sexed twins provide the raw dats. Defined anatomic landmarks on the occlusal surfaces were reduced to X-Y rectangular coordinates prior to the computation of the basal areas dimensions. The results establish a cusp size hierarchy specific for molar type, i.e., five-cusped molars with a distal fovea and distal marginal ridge (5fd), five-cusped molars without a distal fovea and without a distal marginal ridge (5o), and four-cusped molars (4c). Sexual dimorphism in cusp size is apparent in 5fd molar cusped but not in 5o molar cusps. However, males have a significantly higher frequency of 5fd molars. Females have a higher frequency of smaller 5o and 4c molars which have fewer crown components. Moreover, female 5o molars have cusps as large as or larger than 5o male molor cusps. Right-side-left-side differences exist between antimere cusps based on relatively low correlations. The mirroring of molor types occurs infrequently. When observed, most intrapair differences for cusp size, using F-ratios, indicate a low component of hereditary variability.     

Study of the cuspal ridges of the upper first molars in a modern Japanese population

Materials used were dental casts of the upper first molars of modern Japanese subjects, comprising 29 males and 25 females. Their molar occlusal surfaces were photographed by moiré contourography using the standard trigonal plane. The ridges of a cusp, comprising a central ridge and mesial and distal accessory ridges, were identified from the patterns of the moiré fringes. The central ridge was observed in all cusps except for the hypocone in both sexes. Frequencies of the mesial and distal accessory ridges of trigonal cusps were over 90% except for the distal accessory ridge of the metacone, and those of the hypocone were under 25% in both sexes. These values were generally higher in males than in females, especially for the distal accessory ridge of the metacone. The running pattern of the cuspal ridges showed little difference between sexes. The oblique ridge which was higher than the central groove formed a saddle-like structure. This ridge was observed in all materials, but its heights and structural components varied remarkably. In this study, the distal accessory ridge of the metacone was found to be incorporated into the oblique ridge in about 13% of cases. Variability in the running pattern of the ridges within a single cusp was highest in the hypocone and lowest in both the paracone and protocone. The results obtained are considered to represent the stability or reductive tendency of cusps in the upper first molars.     

Geometric morphometric 3D shape analysis and covariation of human mandibular and maxillary first molars

Dental casts of 160 Greek subjects (80 males, 80 females) were scanned by a structured‐light scanner. The upper and lower right first molar occlusal surface 3D meshes were processed using geometric morphometric methods. A total of 265 and 274 curve and surface sliding semilandmarks were placed on the upper and lower molar surfaces, respectively. Principal component analysis and partial least square analysis were performed to assess shape parameters. Molars tended to vary between an elongated and a more square form. The first two principal components (PCs), comprising almost 1/3 of molar shape variation, were related to mesiodistal–buccolingual ratios and relative cusp position. Distal cusps displayed the greatest shape variability. Molars of males were larger than those of females (2.8 and 3.2% for upper and lower molars respectively), but no shape dimorphism was observed. Upper and lower molar sizes were significantly correlated (r² = 0.689). Allometry was observed for both teeth. Larger lower molars were associated with shorter cusps, expansion of the distal cusp, and constriction of the mesial cusps (predicted variance 3.25%). Upper molars displayed weaker allometry (predicted variance 1.59%). Upper and lower molar shape covariation proved significant (RV = 17.26%, P < 0.0001). The main parameter of molar covariation in partial least square axis 1, contributing to 30% of total covariation, was cusp height, in contrast to the primary variability traits exhibited by PC1 and PC2. The aim of this study was to evaluate shape variation and covariation, including allometry and sexual dimorphism, of maxillary and mandibular first permanent molar occlusal surfaces. Am J Phys Anthropol 152:186–196, 2013. © 2013 Wiley Periodicals, Inc.     

Dental trait expression at the enamel-dentine junction of lower molars in extant and fossil hominoids

Discrete dental traits are used as proxies for biological relatedness among modern human populations and for alpha taxonomy and phylogeny reconstruction within the hominin clade. We present a comparison of the expression of lower molar dental traits (cusp 6, cusp 7, trigonid crest pattern, and protostylid) at the enamel-dentine junction (EDJ) in a variety of extant and fossil hominoid taxa, in order to assess the contribution of the EDJ to the morphology of these traits at the outer enamel surface (OES). Molars (n=44) were imaged nondestructively using high-resolution microCT, and three-dimensional surface models of the EDJ and OES were created to compare trait expression at each surface. Our results indicate that these dental traits originate at the EDJ, and that the EDJ is primarily responsible for their degree of expression at the OES. Importantly, variable trait morphology at the EDJ (often not easily recognizable at the OES) indicates that different developmental processes can produce traits that appear similar at the enamel surface, suggesting caution in intra- and intertaxonomic comparisons. The results also highlight the importance of the EDJ for understanding the morphological development of discrete traits, and for establishing graded scales of variation to compare trait frequency among groups for the purpose of taxonomic and/or phylogenetic analysis. Finally, this study demonstrates that imaging the EDJ of both worn and unworn fossil hominin teeth provides a novel source of information about tooth development and variation in crown morphology.     

Brief communication: Contributions of enamel‐dentine junction shape and enamel deposition to primate molar crown complexity

Molar crown morphology varies among primates from relatively simple in some taxa to more complex in others, with such variability having both functional and taxonomic significance. In addition to the primary cusps, crown surface complexity derives from the presence of crests, cuspules, and crenulations. Developmentally, this complexity results from the deposition of an enamel cap over a basement membrane (the morphology of which is preserved as the enamel‐dentine junction, or EDJ, in fully formed teeth). However, the relative contribution of the enamel cap and the EDJ to molar crown complexity is poorly characterized. In this study we examine the complexity of the EDJ and enamel surface of a broad sample of primate (including fossil hominin) lower molars through the application of micro‐computed tomography and dental topographic analysis. Surface complexity of the EDJ and outer enamel surface (OES) is quantified by first mapping, and then summing, the total number of discrete surface orientation patches. We investigate the relative contribution of the EDJ and enamel cap to crown complexity by assessing the correlation in patch counts between the EDJ and OES within taxa and within individual teeth. We identify three patterns of EDJ/OES complexity which demonstrate that both crown patterning early in development and the subsequent deposition of the enamel cap contribute to overall crown complexity in primates. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Der Bau der Canini bei den Paulchoffatiidae (Multituberculata; Ober-Jura)

Canines are preserved among Multituberculata only in the upper jaw of the Paulchoffatiidae, the Pinheirodontidae and the North American genusGliodon Engelmann &; Callison, 1999. They resemble the anterior premolars (p^1–3) in the morphology of their crown, but they differ from them by the presence of only one root. In the present paper, 126 isolated canines of the Paulchoffatiidae from the Guimarota coal-pit in Portugal are treated. They show a wide morphological variation, from bicuspid to pentacuspid expression, and they can be grouped into 18 morphological units (Tab. 1). The bicuspid canines (11 specimens) show one buccal and one lingual cusp, arranged side by side, the latter normally being larger than the former. In the tricuspid canines (32 specimens) the cusps (one buccal and two lingual) are arranged in a triangle, with a great variation in the position of the cusps to each other. Tetracuspid canines (69 specimens) are the dominating group. Two buccal and two lingual cusps are present, which differ markedly in their largeness and their position to each other. The teeth differ also much in the shape of their crown. The pentacuspid canines (14 specimens) show two buccal and three lingual cusps. The variation in the arrangement of the cusps and in the shape of the crown is similar as in the tetracuspid specimens. One or two small cuspules can be present at the anterior border of the crown additionally to the main cusps in the canines. Tricuspid and tetracuspid canines are present also in the skulls of the Paulchoffatiidae, whereas bicuspid and pentacuspid canines are known only as isolated specimens. In the Pinheirodontidae the canines are tricuspid or tetracuspid, inGlirodon they are unicuspid. The Paulchoffatiidae and Pinheirodontidae — Paulchoffatiid line sensuKielan-Jaworowska &; Hurum, 2001 — are characterized by increasing premolarization of the upper canines. With that they differ markedly from all other multituberculates where the canini become reduced.     

A quantitative investigation of irregular cuspules in human maxillary permanent molars

The frequency of occurrence of anomalous cusps or tubercles on human upper first molars was investigated in seven racial populations using moiré contourography, which permits the three-dimensional measurement of minute cusps. Tubercles on the mesial marginal ridge were more frequently found in Mongoloid populations (Japanese and Eskimo) than in others. The frequency of the protoconule was high in Eskimos and Negroids (Bantu and San). The lingual paracone tubercle (mesial cusp) showed a particularly high frequency in Australian aborigines. The metaconule was rare or absent in all of the populations. Caucasoid groups (Dutch White and Asiatic Indian) showed generally low frequencies of all these abnormal tubercles, especially the distal accessory cusp (C5). Racial differences in the frequencies of occurrence may offer a key to understanding the adaptive significance of these traits and human microevolution. Confusion in nomenclature for upper molar tubercles is also discussed.