人类学学报 ›› 2019, Vol. 38 ›› Issue (03): 446-459.doi: 10.16359/j.cnki.cn11-1963/q.2018.0041cstr: 32091.14.j.cnki.cn11-1963/q.2018.0041
孙承凯1(), 孙小玲2, 周蜜3, 刘立群1, 邢松4,5,*()
收稿日期:
2018-06-22
修回日期:
2018-09-20
出版日期:
2019-08-15
发布日期:
2020-09-10
通讯作者:
邢松
作者简介:
孙承凯,山东博物馆副研究馆员,Email: 基金资助:
SUN Chengkai1(), SUN Xiaoling2, ZHOU Mi3, LIU Liqun1, XING Song4,5,*()
Received:
2018-06-22
Revised:
2018-09-20
Online:
2019-08-15
Published:
2020-09-10
Contact:
XING Song
摘要:
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。
中图分类号:
孙承凯, 孙小玲, 周蜜, 刘立群, 邢松. 山东新泰乌珠台人类牙齿的形态学特征[J]. 人类学学报, 2019, 38(03): 446-459.
SUN Chengkai, SUN Xiaoling, ZHOU Mi, LIU Liqun, XING Song. Morphological studies on Wuzhutai tooth, Xintai of Shandong Province[J]. Acta Anthropologica Sinica, 2019, 38(03): 446-459.
M1 | M2 | M3 | 数据来源 | |||||
---|---|---|---|---|---|---|---|---|
例数 | 地点 | 例数 | 地点 | 例数 | 地点 | |||
东亚早/中更新世直立人 | 19 | Sangiran, 淅川, 周口店 | 12 | Sangiran, 和县, 沂源, 周口店 | 22 | Sangiran, 和县, 陈家窝子, 周口店 | [ | |
尼安德特人 | 8 | Hortus, Krapina, St. Césaire, Vindija | 14 | Arcy GrotteRenne, Arcy Hyene, Hortus, Krapina, La Quina, Regourdou, St. Césaire, Subalyuk, Vindija | 35 | Arcy Hyene, Arcy Sur Cure, Ehringsdorf , Genay (Core D'OR) , Hortus, Krapina, La Quina, Ochoz, Petit Puymoyen, Regourdou, Spy, St. Césaire, Shanidar, Subalyuk, Tabun, Vindija | Wolpoff, personal communication | |
全新世现代人 | 非洲 | 20 | 南非 | 20 | 南非 | 20 | 南非 | - |
东亚 | 20 | 河南淅川;湖北郧县、丹江口 | 20 | 河南淅川;湖北郧县、丹江口 | 88 | 河南淅川;湖北郧县、丹江口、襄阳、房县 | 本文 | |
晚更新世现代人 | 非洲 | - | - | - | - | 2 | Klasies River Mouth | - |
西亚 | - | - | - | - | 9 | Qafzeh, Skhul | Wolpoff, personal communication | |
东亚 | 1 | 广西土博咁前洞 | 8 | 湖北黄龙洞;湖南道县福岩洞;云南马鹿洞;广西宝积岩、扶绥南山洞、崇左智人洞、土博咁前洞、隆林德峨 | 13 | 北京田园洞、湖北黄龙洞;湖南道县福岩洞;贵州老鸭洞;福建清流狐狸洞;云南马鹿洞;广西宝积岩、扶绥南山洞、崇左智人洞、隆林德峨、鸡母岩 | [ | |
欧洲 | 1 | Abri Pataud | 7 | Abri Pataud, Dolní V?stonice, Le Rois, Saint Germain-La Riviere | 16 | Abri Pataud, Dolní V?stonice, Isturitz, Le Rois, Mlade?, Pavlov, Predmostí | Wolpoff, personal communication |
表1 本文所使用的对比标本
Tab.1 The comparative specimens used in the present study
M1 | M2 | M3 | 数据来源 | |||||
---|---|---|---|---|---|---|---|---|
例数 | 地点 | 例数 | 地点 | 例数 | 地点 | |||
东亚早/中更新世直立人 | 19 | Sangiran, 淅川, 周口店 | 12 | Sangiran, 和县, 沂源, 周口店 | 22 | Sangiran, 和县, 陈家窝子, 周口店 | [ | |
尼安德特人 | 8 | Hortus, Krapina, St. Césaire, Vindija | 14 | Arcy GrotteRenne, Arcy Hyene, Hortus, Krapina, La Quina, Regourdou, St. Césaire, Subalyuk, Vindija | 35 | Arcy Hyene, Arcy Sur Cure, Ehringsdorf , Genay (Core D'OR) , Hortus, Krapina, La Quina, Ochoz, Petit Puymoyen, Regourdou, Spy, St. Césaire, Shanidar, Subalyuk, Tabun, Vindija | Wolpoff, personal communication | |
全新世现代人 | 非洲 | 20 | 南非 | 20 | 南非 | 20 | 南非 | - |
东亚 | 20 | 河南淅川;湖北郧县、丹江口 | 20 | 河南淅川;湖北郧县、丹江口 | 88 | 河南淅川;湖北郧县、丹江口、襄阳、房县 | 本文 | |
晚更新世现代人 | 非洲 | - | - | - | - | 2 | Klasies River Mouth | - |
西亚 | - | - | - | - | 9 | Qafzeh, Skhul | Wolpoff, personal communication | |
东亚 | 1 | 广西土博咁前洞 | 8 | 湖北黄龙洞;湖南道县福岩洞;云南马鹿洞;广西宝积岩、扶绥南山洞、崇左智人洞、土博咁前洞、隆林德峨 | 13 | 北京田园洞、湖北黄龙洞;湖南道县福岩洞;贵州老鸭洞;福建清流狐狸洞;云南马鹿洞;广西宝积岩、扶绥南山洞、崇左智人洞、隆林德峨、鸡母岩 | [ | |
欧洲 | 1 | Abri Pataud | 7 | Abri Pataud, Dolní V?stonice, Le Rois, Saint Germain-La Riviere | 16 | Abri Pataud, Dolní V?stonice, Isturitz, Le Rois, Mlade?, Pavlov, Predmostí | Wolpoff, personal communication |
图1 乌珠台牙齿照片(第1行)、基于micro-CT扫描数据重建的齿冠表面(第2行)、基于micro-CT扫描数据重建的齿质表面(第3行) 从左至右依次为咬合面、颊侧面、近中面、舌侧面、远中面
Fig.1 The Wuzhutai tooth. First row: the photos of the Wuzhutai original tooth; second row: the visual reconstruction of the Wuzhutai M3 outer enamel surface reconstructed from micro-CT scanning; third row: the visual reconstruction of the Wuzhutai EDJ surface reconstructed from micro-CT scanning from left to right: occlusal, buccal, mesial, lingual, and distal view; B: buccal; M: Mesial
图3 乌珠台M3及对比标本的几何形态测量结果 注:图中变形网格及轮廓线代表RW1和RW2正负值两端M3的形状
Fig.3 Geometric morphometric analyses of the crown outline shape of Wuzhutai M3 and other comparative specimens (TPS-grids correspond to the positive-value or negative-value extremes of RW axis). RW:relative warp; B: buccal, M: mesial
东亚早/中更新世直立人 | 尼安德特人 | 晚更新世现代人 | 全新世现代人 | ||||
---|---|---|---|---|---|---|---|
级别 | 东亚 | 欧洲和西亚[ | 西亚和欧洲[ | 东亚 | 非洲和欧洲[ | 东亚 | |
下次小尖 Hypoconulid size | 0 | - | 2(12.5%) | 6(33.3%) | 1(11.1%) | 63(50.0%) | 5(7.1%) |
1 | - | - | 1(5.6%) | - | 3(2.4%) | 1(1.4%) | |
2 | 1(9.1%) | 1(6.3%) | 2(11.1%) | - | 14(11.1%) | 4(5.7%) | |
3 | 4(27.3%) | 7(43.8%) | 6(33.3%) | 3(33.3%) | 17(13.5%) | 14(20.0%) | |
4 | 5(54.5%) | 5(31.3%) | 2(11.1%) | 3(33.3%) | 17(13.5%) | 24(34.3%) | |
5 | 1(9.1%) | 1(6.3%) | 1(5.6%) | 2(22.2%) | 12(9.5%) | 22(31.4%) | |
第6尖 C6 size | 0 | 1(22.2%) | 5(33.3%) | 12(66.7%) | 4(57.1%) | 94(77.7%) | 40(57.1%) |
1 | - | 1(6.7%) | - | - | 9(7.4%) | 2(2.9%) | |
2 | 5(44.4%) | 4(26.7%) | 4(22.2%) | 2(28.6%) | 11(9.1%) | 9(12.9%) | |
3 | 3(33.3%) | 5(33.3%) | 2(11.1%) | 1(14.3%) | 7(5.8%) | 19(27.1%) | |
第7尖 C7 size | 0 | 1(9.1%) | 8(53.3%) | 13(76.5%) | 6(66.7%) | 98(79.0%) | 63(90.0%) |
1 | - | 1(6.7%) | 1(5.9%) | - | 10(8.1%) | 2(2.9%) | |
2 | 2(18.2%) | 3(20.0%) | 3(17.6%) | - | 6(4.8%) | 1(1.4%) | |
3 | 5(45.5%) | 1(6.7%) | - | 1(11.1%) | 7(5.6%) | 2(2.9%) | |
4 | 3(27.3%) | 2(13.3%) | - | 2(22.2%) | 3(2.4%) | 2(2.9%) | |
三角座中央横脊 Middle Trigonid Crest | 0 | 9(75.0%) | 1(5.9%) | 14(93.3%) | 9(90.0%) | 107(89.9%) | 69(98.6%) |
1 | 1(8.3%) | 4(23.5%) | 1(6.7%) | 1(10.0%) | 12(10.1%) | 1(1.4%) | |
2 | 2(16.7%) | 12(70.6%) | - | - | - | - | |
三角座远中横脊 Distal Trigonid Crest | 0 | 6(54.5%) | 10(62.5%) | 10(62.5%) | 8(100.0%) | 90(75.0%) | 65(95.6%) |
1 | 2(18.2%) | 3(18.8%) | 4(25.0%) | - | 21(17.5%) | 3(4.4%) | |
2 | 3(27.3%) | 3(18.8%) | 2(12.5%) | - | 9(7.5%) | - | |
齿沟类型 Groove pattern | 1 | 8(66.7%) | 9(60.0%) | 2(14.3%) | 2(20.0%) | 21(19.1%) | 7(10.0%) |
2 | 4(33.3%) | 6(40.0%) | 12(85.7%) | 8(80.0%) | 89(80.9%) | 63(90.0%) | |
原附尖 Protostylid | 0 | 1(10.0%) | 8(61.5%) | 15(88.2%) | 2(50.0%) | 105(88.2%) | 43(65.2%) |
1 | 1(10.0%) | 3(23.1%) | 1(5.9%) | - | 9(7.6%) | 11(16.7%) | |
2 | - | 2(15.4) | 1(5.9%) | - | - | - | |
3 | 2(20.0%) | - | - | 1(25.0%) | 2(1.7%) | 8(12.1%) | |
4 | 1(10.0%) | - | - | - | 1(0.8%) | 2(3.0%) | |
5 | 1(10.0%) | - | - | - | 1(0.8%) | 1(1.5%) | |
6 | 4(40.0%) | - | - | 1(25.0%) | 1(0.8%) | - | |
7 | - | - | - | - | - | 1(1.5%) |
表2 M3主要形态特征不同级别及出现率(乌珠台牙齿各特征级别使用灰色底纹标记)*
Tab.2 Frequencies of the degrees of expression of M3 main morphological traits (the grey highlighted rows represent the grade characteristic for Wuzhutai tooth)
东亚早/中更新世直立人 | 尼安德特人 | 晚更新世现代人 | 全新世现代人 | ||||
---|---|---|---|---|---|---|---|
级别 | 东亚 | 欧洲和西亚[ | 西亚和欧洲[ | 东亚 | 非洲和欧洲[ | 东亚 | |
下次小尖 Hypoconulid size | 0 | - | 2(12.5%) | 6(33.3%) | 1(11.1%) | 63(50.0%) | 5(7.1%) |
1 | - | - | 1(5.6%) | - | 3(2.4%) | 1(1.4%) | |
2 | 1(9.1%) | 1(6.3%) | 2(11.1%) | - | 14(11.1%) | 4(5.7%) | |
3 | 4(27.3%) | 7(43.8%) | 6(33.3%) | 3(33.3%) | 17(13.5%) | 14(20.0%) | |
4 | 5(54.5%) | 5(31.3%) | 2(11.1%) | 3(33.3%) | 17(13.5%) | 24(34.3%) | |
5 | 1(9.1%) | 1(6.3%) | 1(5.6%) | 2(22.2%) | 12(9.5%) | 22(31.4%) | |
第6尖 C6 size | 0 | 1(22.2%) | 5(33.3%) | 12(66.7%) | 4(57.1%) | 94(77.7%) | 40(57.1%) |
1 | - | 1(6.7%) | - | - | 9(7.4%) | 2(2.9%) | |
2 | 5(44.4%) | 4(26.7%) | 4(22.2%) | 2(28.6%) | 11(9.1%) | 9(12.9%) | |
3 | 3(33.3%) | 5(33.3%) | 2(11.1%) | 1(14.3%) | 7(5.8%) | 19(27.1%) | |
第7尖 C7 size | 0 | 1(9.1%) | 8(53.3%) | 13(76.5%) | 6(66.7%) | 98(79.0%) | 63(90.0%) |
1 | - | 1(6.7%) | 1(5.9%) | - | 10(8.1%) | 2(2.9%) | |
2 | 2(18.2%) | 3(20.0%) | 3(17.6%) | - | 6(4.8%) | 1(1.4%) | |
3 | 5(45.5%) | 1(6.7%) | - | 1(11.1%) | 7(5.6%) | 2(2.9%) | |
4 | 3(27.3%) | 2(13.3%) | - | 2(22.2%) | 3(2.4%) | 2(2.9%) | |
三角座中央横脊 Middle Trigonid Crest | 0 | 9(75.0%) | 1(5.9%) | 14(93.3%) | 9(90.0%) | 107(89.9%) | 69(98.6%) |
1 | 1(8.3%) | 4(23.5%) | 1(6.7%) | 1(10.0%) | 12(10.1%) | 1(1.4%) | |
2 | 2(16.7%) | 12(70.6%) | - | - | - | - | |
三角座远中横脊 Distal Trigonid Crest | 0 | 6(54.5%) | 10(62.5%) | 10(62.5%) | 8(100.0%) | 90(75.0%) | 65(95.6%) |
1 | 2(18.2%) | 3(18.8%) | 4(25.0%) | - | 21(17.5%) | 3(4.4%) | |
2 | 3(27.3%) | 3(18.8%) | 2(12.5%) | - | 9(7.5%) | - | |
齿沟类型 Groove pattern | 1 | 8(66.7%) | 9(60.0%) | 2(14.3%) | 2(20.0%) | 21(19.1%) | 7(10.0%) |
2 | 4(33.3%) | 6(40.0%) | 12(85.7%) | 8(80.0%) | 89(80.9%) | 63(90.0%) | |
原附尖 Protostylid | 0 | 1(10.0%) | 8(61.5%) | 15(88.2%) | 2(50.0%) | 105(88.2%) | 43(65.2%) |
1 | 1(10.0%) | 3(23.1%) | 1(5.9%) | - | 9(7.6%) | 11(16.7%) | |
2 | - | 2(15.4) | 1(5.9%) | - | - | - | |
3 | 2(20.0%) | - | - | 1(25.0%) | 2(1.7%) | 8(12.1%) | |
4 | 1(10.0%) | - | - | - | 1(0.8%) | 2(3.0%) | |
5 | 1(10.0%) | - | - | - | 1(0.8%) | 1(1.5%) | |
6 | 4(40.0%) | - | - | 1(25.0%) | 1(0.8%) | - | |
7 | - | - | - | - | - | 1(1.5%) |
尼安德特人1 | 现代人1 | 现代人2 | |
---|---|---|---|
三角座近中横脊 | 30.0% (3/10) | 25.0% (1/4) | 3.9%(2/51) |
三角座中央横脊 | 60.0% (6/10) | 25.0% (1/4) | 0.0%(0/51) |
三角座远中横脊 | 30.0% (3/10) | 0.0% | 3.9%(2/51) |
表3 三角座横脊(近中、中央、远中横脊)在尼安德特人和现代人M3 EDJ上的发生率
Tab.3 Frequencies of trigonid crest (mesial, middle, and distal) on the M3 Enamel-dentine junction (EDJ) of Neanderthals and modern humans
尼安德特人1 | 现代人1 | 现代人2 | |
---|---|---|---|
三角座近中横脊 | 30.0% (3/10) | 25.0% (1/4) | 3.9%(2/51) |
三角座中央横脊 | 60.0% (6/10) | 25.0% (1/4) | 0.0%(0/51) |
三角座远中横脊 | 30.0% (3/10) | 0.0% | 3.9%(2/51) |
乌珠台 | 老鸭洞LYC1 | 尼安德特人 | 化石现代人 | 全新世 现代人 | ||||
---|---|---|---|---|---|---|---|---|
校正前 | 校正后 | 校正前 | 校正后 | |||||
数据来源 | 本文 | 本文 | Xing et al. 2017[ | Xing et al. 2017[ | Smith et al. 2012[ | Olejniczak et al. 2008[ | Smith et al. 2012[ | Smith et al. 2012[ |
例数 | 1 | 1 | 1 | 1 | 6 | 8 | ||
AET (mm) | 1.08 | 1.19 | 1.06 | 1.18 | 1.01 (0.91-1.16) | 0.99 | 1.28 (1.15-1.41) | 1.25 (0.98-1.74) |
RET | 20.22 | 22.28 | 19.74 | 21.97 | 16.8 (14.3-18.3) | 16.55 | 20.5 (19.2-21.9) | 21.7 (17.2-31.8) |
表4 乌珠台M3及对比标本二维釉质厚度比较。AET2D (2D average enamel thickness):二维平均釉质厚度) ;RET2D(2D relative enamel thickness):二维相对釉质厚度 。括号内数值代表范围
Tab.4 Comparisons of AET2D (2D average enamel thickness) and RET2D (2D relative enamel thickness) between Wuzhutai M3 and other comparative specimens (parentheses indicate the range when more than one specimen was reported)
乌珠台 | 老鸭洞LYC1 | 尼安德特人 | 化石现代人 | 全新世 现代人 | ||||
---|---|---|---|---|---|---|---|---|
校正前 | 校正后 | 校正前 | 校正后 | |||||
数据来源 | 本文 | 本文 | Xing et al. 2017[ | Xing et al. 2017[ | Smith et al. 2012[ | Olejniczak et al. 2008[ | Smith et al. 2012[ | Smith et al. 2012[ |
例数 | 1 | 1 | 1 | 1 | 6 | 8 | ||
AET (mm) | 1.08 | 1.19 | 1.06 | 1.18 | 1.01 (0.91-1.16) | 0.99 | 1.28 (1.15-1.41) | 1.25 (0.98-1.74) |
RET | 20.22 | 22.28 | 19.74 | 21.97 | 16.8 (14.3-18.3) | 16.55 | 20.5 (19.2-21.9) | 21.7 (17.2-31.8) |
乌珠台 (未校正) | 老鸭洞LYC1(未校正) | 尼安德特人 | 全新世现代人 | |||
---|---|---|---|---|---|---|
数据来源 | 本文 | 本文 | Pan et al. 2016[ | Olejniczak et al. 2008[ | Pan et al. 2016[ | Smith et al. 2012[ |
例数 | 1 | 1 | 5 | 6 | 4 | 9 |
AET 3D(mm) | 1.28 | 1.40 | 1.38 (1.24-1.54) | 1.06 | 1.38 (1.20-1.47) | 1.45 |
RET3D | 21.33 | 24.65 | 21.24 (19.11-23.73) | 15.43 | 22.06 (20.52-23.74) | 23.79 |
表5 乌珠台M3及对比标本三维釉质厚度比较。AET3D (3D average enamel thickness):三维平均釉质厚度) ;RET3D (3D relative enamel thickness):三维相对釉质厚度 。括号内数值代表范围
Tab.5 Comparisons of AET3D (3D average enamel thickness) and RET3D (3D relative enamel thickness) between Wuzhutai M3 and other comparative specimens (parentheses indicate the range when more than one specimen was reported)
乌珠台 (未校正) | 老鸭洞LYC1(未校正) | 尼安德特人 | 全新世现代人 | |||
---|---|---|---|---|---|---|
数据来源 | 本文 | 本文 | Pan et al. 2016[ | Olejniczak et al. 2008[ | Pan et al. 2016[ | Smith et al. 2012[ |
例数 | 1 | 1 | 5 | 6 | 4 | 9 |
AET 3D(mm) | 1.28 | 1.40 | 1.38 (1.24-1.54) | 1.06 | 1.38 (1.20-1.47) | 1.45 |
RET3D | 21.33 | 24.65 | 21.24 (19.11-23.73) | 15.43 | 22.06 (20.52-23.74) | 23.79 |
图4 乌珠台M3及对比标本三维釉质厚度分布模式 对比标本包括和县直立人(PA831),尼安德特人(Krapina D7,下载自http://www.nespos.org)和3个全新世地点的现代人(YYM114, NC114, ML81)。如对比标本为右侧,将其镜像为左侧,便于与乌珠台标本进行对比
Fig.4 The distribution patterns of 3D enamel thickness in Wuzhutai M3 and other five specimens The comparative specimens include H. erectus (PA831), Neanderthal (Krapina D7,downloaded from NESPOS), and recent modern humans (YYM114, NC114, ML81). The photos were mirror-imaged to facilitate the visual comparisons if they are right M3. The images were not scaled;O: occlusal; B: buccal; M: mesial; L: lingual; D: dista
图5 乌珠台M3及对比标本齿冠尺寸对比散点图。MD:近远中径;BL:颊舌径
Fig.5 Bi-variable plots of the crown sizes of Wuzhutai M3 and the comparative specimens (MD: mesiodistal dimension; BL: buccolingual dimension)
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