植食性哺乳动物牙齿磨痕分析方法简介及其在古食性恢复中的应用前景

通过研究古哺乳动物的食性来探讨哺乳动物演化与古生态环境变化之间的关系是目前古生物学研究领域的一个热点,而牙齿磨痕分析是恢复古食性和重建古生态环境的重要手段。牙齿磨痕(dental wear)分析包括微痕(microwear)分析和中痕(mesowear)分析,两种方法均强调食性与牙齿磨痕模式的严格对应,即不同食性的动物具有不同的牙齿磨痕特征模式。近年来,牙齿磨痕分析方法以其简单、快捷和高效等优点已被广泛应用于奇蹄类、偶蹄类、啮齿类、长鼻类和食肉类等哺乳动物的食性研究。但哺乳动物的食性和摄食习性比较复杂,很可能会影响微痕和中痕分析对食性的分辨率。所以,为了获得更加详细的古食性信息和更高的食性分辨率,一方面要对微痕和中痕分析方法进行改进,增添稳定并具有食性识别意义的观测变量,另一方面,需要同时结合微痕和中痕分析,从而获得更加全面的食性信息。虽然牙齿磨痕分析目前主要应用于植食性哺乳动物的食性研究,但其原理对哺乳动物的其它类群也是适用的,随着磨痕分析方法的不断改进和其它类群磨痕数据库的建立,未来的牙齿磨痕分析将可以恢复更多类群的古食性,从而可以更加全面和准确地揭示古食性与古环境信息。     

激光共聚焦显微镜在微痕定量分析中的应用综述

微痕分析是石器、骨器、牙齿等工艺、功能研究的常用方法,但无论是高倍还是低倍等常用方法,受限于体式光学显微镜、扫描电镜等观察方法,微痕分析通常很难进行原位定量分析,因此微痕分析的定量化是目前学术界遇到的主要问题。近年来随着激光共聚焦显微镜（LSCM）在微痕分析中的应用,微痕定量化在西方学术界得到了新的实践和进展。本文主要介绍激光共聚焦显微镜的原理及其在微痕分析中的应用案例,以期促进微痕分析在中国的进一步发展。     

石制品微磨痕分析的实验性研究

以扫描电子显微镜为主要手段,依据Keeley的实验方法,对实验和 “盲测” 的燧石制品进行了微磨痕 (microwear) 的观察分析,验证了该方法的可行性;表明在石制品的加工方式、加工对象与其可能的残留信息——石制品的使用痕迹,如以光泽、条痕、破损疤为主要特征的微磨痕之间所具有的某种实验模式,初步掌握了微磨痕研究的实验方法,取得了一些结果,并予以检验;利用扫描电镜观察 “晶体破损” 与否,可能是鉴定石制品使用或未使用的一种行之有效的方法。     

水洞沟地区白云岩细石叶的微痕实验研究

水洞沟遗址是中国北方旧石器时代晚期的著名遗址,石制品原料以就地取材的白云岩为主。本文利用水洞沟地区的白云岩生产细石叶和石片,进行了29组复制使用实验。对使用痕迹的观察表明:不同加工对象和使用方式导致细石叶上出现不同的微痕特征,细石叶适于装柄使用,以刮、切、削等利用侧刃缘的运动方式效率为高,尖部用于加工硬度较低的动植物效果较好。对比燧石、黑曜岩等不同石料的微痕实验数据可知,不同石料的产品,其使用痕迹存在一定差异。本文对白云岩细石叶的微痕实验,探讨了其适宜的使用方式和加工对象的范围,为分析水洞沟地区出土的白云岩细石叶功能提供了可资参考、对比的实验数据。     

考古标本微磨痕初步研究

从周口店第1地点和马鞍山遗址选取了20件燧石制品,以微磨痕的实验研究为基础,以扫描电子显微镜 (简称电镜) 为主要手段,通过对比分析,尝试了不同遗址间考古标本的微磨痕分析。结果表明,周口店第1地点和马鞍山遗址的石制品的功能都具有多样性;“楔” 的功能见于马鞍山遗址,并为周口店第1地点 “使用石片较多” 的说法提供了微磨痕方面的新证据。     

角页岩雕刻器的微痕实验研究

本文以丁村遗址角页岩为原料打制实验标本,设计展开雕刻器微痕实验,并对其中部分标本按照序贯试验原理进行分阶段实验。实验结果表明,角页岩雕刻器雕刻刃的使用微痕基本特征为:轻度到重度磨圆,片疤数量较少,多小中型片疤,终端形态多为羽状和阶梯状,以分散分布为主,大片疤常单独出现。分阶段实验结果表明,磨圆是以从快到慢速度生成的,片疤在不同阶段发生改变。本文旨在通过对角页岩雕刻器使用微痕的研究,为探讨考古标本功能提供可参照的数据与图像。     

石制品使用微痕多阶段成形轨迹的实验研究

石制品微痕是进行石器功能研究、了解史前人类行为模式的重要考古证据之一。本文在以往实验研究的基础上, 设计并开展多阶段燧石制品"刮骨"微痕实验, 分阶段详细、直观地记录石制品刃缘使用微痕的最初形成与发展动态。实验研究表明, "刮骨"使用微痕迹随着使用时间长度的递增发生复杂的动态变化, 使用微痕的发展与使用时间之间并非简单的正函数关系。片疤破损自开始在一定时间段内呈连续发生状态, 随后停止发展, 是确认石制品是否经过使用的良好指示。磨圆表现出由少到多、由弱及强的逐渐发育过程, 可以反映石制品的使用时间和强度, 也是判断使用微痕的重要参考依据。     

虎头梁遗址尖状器功能的微痕研究

虎头梁遗址位于河北省阳原县泥河湾盆地, 距今1万年左右, 出土的石制品加工精致、器型稳定。本文选择其中代表性的尖状器进行功能研究, 通过微痕分析探讨其使用方式, 并结合其形态特征探讨器物类型与功能之间的关系。对5个亚类型共52件标本的研究表明, 虎头梁尖状器具有专业分工, 并被装柄使用。这种工具的制作和使用特征反映了更新世末期虎头梁人群通过提高工具的效能来应对环境变化的生存策略。     

河南舞阳贾湖遗址出土石器的微痕分析

石器作为人类物质文化的记录者,在史前人类的生产生活中占据着重要地位,对石器功能和使用方式的探讨是考古学关注的问题之一。目前国内石器研究的主要手段是从器物形态出发,进行类型学的研究,针对石器具体功能及使用方式的实证研究较为缺乏,对磨制石器的研究尤为不足。有鉴于此,本研究以中原地区舞阳贾湖遗址出土的磨制石器为研究对象,利用微痕分析和实验考古的方法,对石器的功能、使用方式及加工对象进行分析和判断。结果表明:石镰的主要功能是收割禾本科植物;石刀主要用于加工禾本科植物、木材和兽骨等;石斧的功能主要是加工木材,另外还可能承担加工兽皮和兽骨的工作;石锛的主要功能与木作有关;石凿在形态上有所区别,但其加工对象主要是木材;石铲的功能以掘土为主。微痕分析的结果使我们对贾湖聚落磨制石器的功能有了客观的认识,可以看出石器的功能并不单一,广泛存在着"一器多用"的现象。本研究从石器使用的角度,揭示了史前人类的生产活动和行为特点,同时也为研究先民的生业经济状况提供了重要材料。     

新疆吐鲁番胜金店墓地人骨的牙齿微磨耗

本文使用不同倍率的超景深电子显微镜,对新疆吐鲁番胜金店墓地中出土的13例臼齿的颊侧微磨耗形态进行了观察和研究。通过对臼齿颊侧表面凹坑和条痕形态微磨耗的测量和统计,计算出每例牙齿的微痕数量、均长、均宽、凹坑百分比和L_h/L_v比值,按照性别和年龄分组进行人群内的数据对比分析。研究结果表明,该人群中年龄较大的个体摄入更高比例的植物类食物;两性的食物结构不存在显著性差异,男性摄入植物类食物比例略高,女性的食物结构在不同的年龄阶段有所差异。将胜金店墓地人群与不同生计方式人群的L_h/L_v比值进行人群间的差异性分析,并结合胜金店墓地的随葬品特点推测,胜金店墓地人群的食物结构以肉类食物为主,其生计模式以游牧业为主,兼营种植类经济。     

Ecogeographic variation in Neandertal dietary habits: evidence from occlusal molar microwear texture analysis

In the late Middle and early Late Pleistocene, Neandertals inhabited a wide variety of ecological zones across western Eurasia during both glacial and interglacial times. To elucidate the still poorly understood effects of climatic change on Neandertal subsistence patterns, this study employs dental microwear texture analysis to reconstruct the diets of Neandertal individuals from various sites across their wide temporal and geographic ranges. The results of this study reveal environmentally-driven differences in the diets of Neandertal groups. Significant differences in microwear signatures, correlated with paleoecological conditions, were found among Neandertal groups that lived in open, mixed, and wooded environments. In comparison to recent hunter-gatherer populations with known, yet diverse diets, the occlusal molar microwear signatures of all the Neandertal groups indicate that their diet consisted predominantly of meat. However, the results of this study suggest that plant foods did form an important part of the diet of at least some Neandertal groups (i.e., those that lived in mixed and wooded habitats). Overall, the proportion of plant foods in the Neandertal diet appears to have increased with the increase in tree cover.     

Incisor microwear of Sumatran anthropoid primates

Several studies have suggested that incisor microwear reflects diet and feeding adaptations of anthropoids. However, such studies have been largely qualitative, and interpretations have relied on anecdotal references to diet and tooth use reported in the socioecology literature. The current study relates incisor microwear in four anthropoid primates to specific ingestive behaviors and food types. Central incisor casts of wild-shot museum specimens of Hylobates lar, Macaca fascicularis, Pongo pygmaeus, and Presbytis thomasi were examined by scanning electron microscopy, and analyzed using a semiautomated image analysis procedure. Microwear patterns were used to generate predictions regarding diet and anterior tooth use. These predictions were evaluated using data collected during a 1 year study of feeding behavior of these same taxa in the wild (Ungar, 1992, 1994a, b). Results suggest that (1) enamel prism relief is associated with the effectiveness of etching reagents in foods, (2) dental calculus buildup results from a lack of incisor use and perhaps the ingestion of sugar-rich foods, (3) striation density varies with degree of anterior tooth use in the ingestion of abrasive food items, (4) striation breadth is proposed to relate to the ratio of exogenous grit to phytoliths consumed; and (5) preferred striation orientation indicates the direction that food items are pulled across the incisors during ingestion. It is concluded that incisor microwear studies can contribute to the understanding of diets and feeding behaviors of extinct primates. © 1994 Wiley-Liss, Inc.     

Dental microwear texture analysis: technical considerations

Dental microwear analysis is commonly used to infer aspects of diet in extinct primates. Conventional methods of microwear analysis have usually been limited to two-dimensional imaging studies using a scanning electron microscope and the identification of apparent individual features. These methods have proved time-consuming and prone to subjectivity and observer error. Here we describe a new methodological approach to microwear: dental microwear texture analysis, based on three-dimensional surface measurements taken using white-light confocal microscopy and scale-sensitive fractal analysis. Surface parameters for complexity, scale of maximum complexity, anisotropy, heterogeneity, and textural fill volume offer repeatable, quantitative characterizations of three-dimensional surfaces, free of observer measurement error. Some results are presented to illustrate how these parameters distinguish extant primates with different diets. In this case, microwear surfaces of Cebus apella and Lophocebus albigena, which consume some harder food items, have higher average values for complexity than do folivores or soft fruit eaters.     

Testing hypotheses of dietary reconstruction from buccal dental microwear in Australopithecus afarensis

A recent study of occlusal microwear in Australopithecus afarensis described this species as an opportunistic dweller, living in both forested and open environments and greatly relying on fallback resources and using fewer food-processing activities than previously suggested. In the present study, analysis of buccal microwear variability in a sample of A. afarensis specimens (n = 75 teeth) showed no significant correlations with the ecological shift that took place around 3.5 Ma in Africa. These results are consistent with the occlusal microwear data available. In fact, significant correlations between buccal and occlusal microwear variables were found. However, comparison of the buccal microwear patterns showed clear similarities between A. afarensis and those hominoid species living in somewhat open environments, especially the Cameroon gorillas. A diet based mainly on succulent fruits and seasonal fallback resources would be consistent with the buccal microwear patterns observed.     

Dental microwear texture analysis and diet in the Dmanisi hominins

Reconstructions of foraging behavior and diet are central to our understanding of fossil hominin ecology and evolution. Current hypotheses for the evolution of the genus Homo invoke a change in foraging behavior to include higher quality foods. Recent microwear texture analyses of fossil hominin teeth have suggested that the evolution of Homo erectus may have been marked by a transition to a more variable diet. In this study, we used microwear texture analysis to examine the occlusal surface of 2 molars from Dmanisi, a 1.8 million year old fossil hominin site in the Republic of Georgia. The Dmanisi molars were characterized by a moderate degree of surface complexity (Asfc), low textural fill volume (Tfv), and a relatively low scale of maximum complexity (Smc), similar to specimens of early African H. erectus. While caution must be used in drawing conclusions from this small sample (n = 2), these results are consistent with continuity in diet as H. erectus expanded into Eurasia.     

The assessment of jaw movement direction from dental microwear

In order to resolve several controversies about unusual jaw movements such as thegosis and orthal retraction, information about direction of jaw movement is essential. Ryan (1979a,b) proposed that asymmetry of microwear striations might provide such data. In vivo occurrences of asymmetric striations on chimpanzee molars are presented and analyzed. Results suggest that either thegosis contacts commonly occur on chimpanzee molars or that Ryan's model, as developed in vitro, does not accurately apply to naturally occurring molar microwear.     

Microwear and morphology: functional relationships between human dental microwear and the mandible

Microscopic pits and scratches form on teeth during chewing, but the extent to which their formation is influenced by mandibular morphology is unknown. Digitized micrographs of the base of facet nine of the first, second, and third mandibular molar were used to record microwear features from an archaeological sample of modern humans recovered from Semna South in northern Sudan (n=38; 100 BC to AD 350). Microwear patterns of the molar row are correlated with mandibular corpus width and depth, and with mandibular length. Variations in shear and compression at the base of facet nine during chewing were inferred. It may be that some correlations between microwear and mandibular morphology are predictable, reflecting similar aspects of masticatory loading, though the full extent of the relationship remains to be resolved.     

Terrestrial foraging and dental microwear inPapio ursinus

Dental microwear of ten wild-shot chacma baboons (Papio urinus) form Northwest and Northern Privinces, South Africa was examined by scanning electron microscopy. All specimens were collected during the dry season, during which these primates exploit hypogeous (underground) food items, including tubers and corms. The microwear fabric of thisP. ursinus sample is characterized by high pitting frequencies and large microwear features. It differs significantly from those displayed by other terrestrially foraging papionins of the genusTheropithecus. Exogenous grit is hypothesized to be largely responsible for the observedP. ursinus wear pattern, which resembles the microwear profiles of durophagous primates. It is suggested that large microwear features and a high incidence of enamel pitting, which are generally held to represent a microwear “signature” of durophagy, may not always be indicative of hard-object feeding in anthropoid primates.     

Dietary divergence in space and time – Lessons from the dwarf-goat Myotragus balearicus (Pleisto-Holocene,Mallorca, Spain)

Newly colonised, isolated habitats, like islands, provide diverse niches to be filled and are prone to facilitate ecological separation which might lead to an adaptive radiation. Examples of such radiations can be found in the Mediterranean for the genera Candiacervus (Crete), Nesogoral (Sardinia) and Hoplitomeryx (Gargano). A different strategy to cope with limited resources on islands is generalism. We test whether populations of the endemic bovid Myotragus balearicus from two sites and Pleistocene as well as Holocene levels on Mallorca island displays ecological separation indicated by diet, or whether the species shifted its dietary trait towards generalism. We expect to find either: (1) dietary divergence in space and time (between sites and stratigraphic levels), which would indicate niche partitioning and/or a shift in dietary traits due to environmental influences; or (2) dietary congruence in a less specialised, generalistic dietary strategy in space and time which would indicate a flexible trait to cope with instable resource availability. We compare individuals from a fossil assemblage at a northern site and one assemblage from the eastern coast in terms of their dietary traits. Traits are reconstructed using dental dietary proxies, complementary in time scale and resolution. (1) 3D-dental topometry and (2) enamel surface texture analysis. Data suggest that individuals from both assemblages of M. balearicus behaved as variable browse dominated intermediate feeders. We thus conclude that the observed variability relates to a shift towards generalism as a subsistence strategy. We consider hypsodonty the pre-adaptation for this life style that enabled M. balearicus to exploit almost any food source in its energetically restricted island habitat.     

Quantitative differences in dental microwear between primate species with different diets and a comment on the presumed diet of Sivapithecus

Studies of dental microwear have been used to relate tooth form to function in a variety of recent and extinct mammals. Probably the most important aspect of microwear analysis is the possibility of using it to deduce the diet of extinct animals. Such deductions must be based on comparative studies of modern species with known diets, but to date, only qualitative studies have been attempted and all have been based on small samples. Here we report quantitative differences in dental microwear between primate species that are known to have different diets. Occlusal facets with different functions have previously been shown to exhibit different microwear patterns. However, the differences between facets of one species are shown to be far less than those between homologous facets of different species. Study of seven species of extant primates shows that enamel microwear can be used to distinguish between those with a mainly frugivorous diet and those with a mainly folivorous one. Microwear can also distinguish hard-object feeders from soft-fruit eaters. The microwear of Miocene Sivapithecus indicus cannot be distinguished statistically from that of the chimpanzee, but it is different from that of the other species. On this evidence S. indicus was not a hard-object feeder and the adaptive significance of its thick molar enamel is at present unknown.