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Biogeochemical data from well preserved 200 ka collagen and skeletal remains

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Abstract

Data presented herein form part of a pilot study to reconstruct palaeoenvironment using isotopic data from 200 ka collagen and skeletal remains. To date such a study has only been possible on materials of up to 120 ka, due to poor preservation, but amino acid compositional analyses of mammal molar collagen herein indicate excellent preservation of the 200 ka collagen. Samples studied were found in association with faunal evidence for a temperate climate, and oxygen and carbon isotope data support this evidence. Collagen N 15 N values, however, are akin to those typically found in collagen from modern semi-arid to arid environments. Similar data have been reported for the Eemian interglacial, and it is likely that these data reflect specific environmental adaptations, as opposed to aridity. This pilot study emphasises the need for complementary isotopic and stratigraphical analyses in palaeoenvironmental studies. ß 2001 Elsevier Science B.V. All rights reserved.

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... A few co-eval sites do exist (e.g. Stanton Harcourt, Oxfordshire, Jones et al. (2001) for Hutton Cave) however, there is not sufficient equivalent isotope data available to make their inclusion viable. ...
... Although the steppe mammoth sampled from Hutton Cave equally did not produce sufficient collagen for analysis, the frequently-observed elevated d 15 N values in mammoths (eg. Jones et al., 2001) suggests that these megaherbivores did not form part of the diet of either the wolves or the hyaenas from Hutton Cave. This overlapping of wolf and spotted hyaena raises the question of competitive interaction. ...
... The high d 15 N values at Stanton Harcourt could thus potentially represent a suckling rather than a palaeoenvironmental signal. Furthermore, Jones et al. (2001) ruled out aridity as a driver of 15 N elevation based on apparent incompatibility with palaeoenvironmental evidence indicating fully interglacial conditions at the site. However, this contradicts current understanding of the palaeoenvironmental and palaeoclimatic characteristics of MIS 7c-a, which indicate a relatively cool interglacial with mean summer temperatures of þ15 to þ16 C (de Rouffignac et al., 1995;Murton et al., 2001) and higher magnitude insolation variability than any other interglacial of the Middle and Late Pleistocene (Berger et al., 2016). ...
Article
The wolf (Canis lupus L., 1754) has been a major keystone predator in the Palaearctic since the late Middle Pleistocene. Today, wolves display considerable dietary plasticity over their range, characterised by their preferential consumption of large and medium-sized wild ungulates, supplemented by smaller prey, including small mammals, fish and plant foods. However, the origins of this dietary flexibility (arguably the key to the wolf's long persistence) are poorly understood in terms of responses to different drivers over the course of the Pleistocene, including changing climate, environment and competition from other large carnivores. Here, in the first study using direct palaeodietary measurements on British fossil wolves, carnivore competitors and potential prey species, we compare stable isotope (δ¹³C and δ¹⁵N) evidence from three sites representing a late Middle Pleistocene interglacial (Marine Oxygen Isotope Stage [MIS] 7c-a, c.220-190ka BP), the early Devensian (last cold stage, MIS 5a, c.90-80ka BP) and the middle Devensian (MIS 3, c. 60-25ka BP). The results reveal clear patterns of changing wolf prey choice through time. Notwithstanding issues of collagen preservation obscuring some dietary choices in the oldest samples, both small and large prey (hare, horse) were taken by wolves in the MIS 7c-a interglacial, large prey only (reindeer, bison) during MIS 5a and a broader range of large prey items (horse, woolly rhinoceros, bison) during MIS 3. The results also reveal two further important aspects: (1) that where wolves and spotted hyaenas co-existed, they occupied the same dietary niche and the former was not outcompeted by the latter, and (2) that the stable isotope evidence indicates prey choices during MIS 7c-a and MIS 3 that are not in synchrony with palaeodietary reconstructions from previous studies based on wolf cranio-dental morphology. This establishes for the first time a likely lag between changing predatory behaviour and morphological response but is interestingly not seen in the wolves from MIS 5a, where the prey choices are echoed by the cranio-dental morphology.
... Two different bone and tooth fractions potentially record unaltered isotopic signatures: collagen and the carbonate fraction of bioapatite. Bones with well-preserved collagen are exceptionally rare in sites older than the Late Pleistocene, with notable exceptions in the Middle Pleistocene (Jones et al., 2001;Kuitems et al., 2012). Fortunately, preservation of biogenic isotopic signatures has been demonstrated in tooth enamel of the fossils of millions years old (e.g. ...
... Only in a few exceptional cases was it possible to use this approach in older sites (e.g. Bocherens et al., 1999;Jones et al., 2001;Palmqvist et al., 2003;Britton et al., 2012;Kuitems et al., 2012). In mammal fossil remains devoid of collagen, it is still possible to obtain an isotopic signature with biological meaning from carbonate. ...
... The studies of the carbon and oxygen isotopic composition of woolly mammoth tooth enamel in Eastern and Northern Europe reconstructed palaeoenvironment trends during the second half of the late Pleistocene, but did not consider the niche partitioning aspects, as they focus only on one species (Arppe and Karhu, 2010;Arppe et al., 2011). In contrast, two studies included more than one species, Jones et al. (2001) and Britton et al. (2012). Both investigated the carbon and nitrogen isotopic composition of bone and tooth collagen in England (Jones et al., 2001) and Germany (Britton et al., 2012), together with the carbon and oxygen isotopic composition of tooth enamel of proboscideans, bison and horse in the first study (Jones et al., 2001). ...
Article
Species flexibility in diet and habitat and their ability to tolerate a range of unfavourable ecological conditions and survive in unusual habitats accompanied by unexpected faunal components has been determined from various research fields. We present the dietary and environmental reconstructions of interglacial and glacial large mammals from central Germany (Bockstein and Vogelherd caves, Steinheim and Mauer) during the Middle and Late Pleistocene, using carbon and oxygen stable isotope relative abundances in the carbonate fraction of tooth enamel. The same species existed in central Europe during different temperature and ecosystem regimes. It appeared that the species during the glacial periods demonstrated much narrower ranges d 13 C and d 18 O than during the interglacial periods and at the dawn of the species origination. Intriguingly, the early woolly mammoth Mammuthus primigenius fraasi apparently lived in much milder conditions than the late woolly mammoth Mammuthus primigenius primigenius, and shared similar diet and habitat with the straight-tusked forest elephant Palaeoloxodon antiquus. Bovids existed in extremely open habitat at Steinheim, probably during the Saalian glacial, compared to other glacial species. Woolly rhinoceros Coelodonta antiquitatis and the early woolly mammoth came to occupy similar environments only later, during the Weichselian glacial period in Bockstein and Vogelherd caves. A so-called "steppe rhinoceros" Stephanorhinus hemitoechus occurred in the forested habitat, along with the Merck's rhino Stephanorhinus kirchbergensis. The horses appear to have preferred warmer and denser habitat than the woolly rhinoceros during the Pleistocene. The Pleistocene donkeys shared the ecological niches between bovids and horses. These new results demonstrate that stable isotope analyses can be extremely helpful to determine more detailed paleodiet and paleohabitat of extinct species. It appears that the deviation of the inferred palaeoecological patterns from the patterns deduced from modern survivors' ecology increases with increasing age, and species appeared more flexible at their origin.
... Two different bone and tooth fractions potentially record unaltered isotopic signatures: collagen and the carbonate fraction of bioapatite. Bones with well-preserved collagen are exceptionally rare in sites older than the Late Pleistocene, with notable exceptions in the Middle Pleistocene (Jones et al., 2001;Kuitems et al., 2012). Fortunately, preservation of biogenic isotopic signatures has been demonstrated in tooth enamel of the fossils of millions years old (e.g. ...
... Only in a few exceptional cases was it possible to use this approach in older sites (e.g. Bocherens et al., 1999;Jones et al., 2001;Palmqvist et al., 2003;Britton et al., 2012;Kuitems et al., 2012). In mammal fossil remains devoid of collagen, it is still possible to obtain an isotopic signature with biological meaning from carbonate. ...
... The studies of the carbon and oxygen isotopic composition of woolly mammoth tooth enamel in Eastern and Northern Europe reconstructed palaeoenvironment trends during the second half of the late Pleistocene, but did not consider the niche partitioning aspects, as they focus only on one species (Arppe and Karhu, 2010;Arppe et al., 2011). In contrast, two studies included more than one species, Jones et al. (2001) and Britton et al. (2012). Both investigated the carbon and nitrogen isotopic composition of bone and tooth collagen in England (Jones et al., 2001) and Germany (Britton et al., 2012), together with the carbon and oxygen isotopic composition of tooth enamel of proboscideans, bison and horse in the first study (Jones et al., 2001). ...
Article
Species flexibility in diet and habitat and their ability to tolerate a range of unfavourable ecological conditions and survive in unusual habitats accompanied by unexpected faunal components has been determined from various research fields. We present the dietary and environmental reconstructions of interglacial and glacial large mammals from central Germany (Bockstein and Vogelherd caves, Steinheim and Mauer) during the Middle and Late Pleistocene, using carbon and oxygen stable isotope relative abundances in the carbonate fraction of tooth enamel. The same species existed in central Europe during different temperature and ecosystem regimes. It appeared that the species during the glacial periods demonstrated much narrower ranges δ13C and δ18O than during the interglacial periods and at the dawn of the species origination. Intriguingly, the early woolly mammoth Mammuthus primigenius fraasi apparently lived in much milder conditions than the late woolly mammoth Mammuthus primigenius primigenius, and shared similar diet and habitat with the straight-tusked forest elephant Palaeoloxodon antiquus. Bovids existed in extremely open habitat at Steinheim, probably during the Saalian glacial, compared to other glacial species. Woolly rhinoceros Coelodonta antiquitatis and the early woolly mammoth came to occupy similar environments only later, during the Weichselian glacial period in Bockstein and Vogelherd caves. A so-called “steppe rhinoceros” Stephanorhinus hemitoechus occurred in the forested habitat, along with the Merck's rhino Stephanorhinus kirchbergensis. The horses appear to have preferred warmer and denser habitat than the woolly rhinoceros during the Pleistocene. The Pleistocene donkeys shared the ecological niches between bovids and horses. These new results demonstrate that stable isotope analyses can be extremely helpful to determine more detailed paleodiet and paleohabitat of extinct species. It appears that the deviation of the inferred palaeoecological patterns from the patterns deduced from modern survivors' ecology increases with increasing age, and species appeared more flexible at their origin.
... 1); however, since collagen is rarely preserved for more than 10,000 years, these investigations were largely confined to the recent past. Yet, it has been shown that, under the right conditions, bone collagen can survive for 200,000 years or more (Jones et al. 2001). Thus, it has proven possible to analyze the bone collagen of late Pleistocene hominins in certain cases. ...
... After surface cleaning, bone or dentine samples are demineralized in diluted HCl (0.5-1.0 M) for periods ranging from 20 min to 5 days. Performing this step at low temperature (5 C) is one recent innovation that allows collagen to be extracted from very old, fragile samples (Richards and Hedges 1999;Jones et al. 2001). The insoluble residue may then be soaked in 0.125 N NaOH to remove contaminating humic acids; but as this leads to collagen solubilization and decreased extraction yields, it is no longer favored. ...
... Older hominin material is not amenable to this form of analysis as bone collagen is rarely preserved from beyond the late Pleistocene (Jones et al. 2001). However, the carbon isotopes in the bone's mineral component (a biological apatite) can also be used as dietary proxies (Sullivan andKrueger 1981, 1983; Lee-Thorp and van der Merwe 1987). ...
Article
Stable isotope ratio analysis is now regularly used to investigate early hominin diets based on the principle that “you are what you eat.” Analysis of collagen from Neanderthals and anatomically modern humans prior to 20 ka has shown them to be significantly enriched in 15N compared to contemporaneous carnivores and herbivores. This suggests that animal foods were a dominant component of their diets, although it must be borne in mind that collagen δ15N can underemphasize the importance of plant foods. Carbon isotope analysis of the enamel mineral of South African australopiths and early Homo has revealed that these taxa consumed ∼30% C4 foods such as tropical grasses, sedges, or animals that ate these foods. Moreover, the australopiths are characterized by remarkably variable δ13C values. Chimpanzees, in contrast, are nearly pure C3 consumers even in environments with abundant C4 vegetation. These data suggest that when confronted with increasingly open areas, chimpanzees continue to exploit the foods that are most abundant in forest environments, whereas australopiths utilized novel C4 resources in addition to forest foods.
... The stability and degradation of the major protein in bone and dentine, collagen, has been extensively studied because of its importance not only in isotopic but also radiocarbon and racemization studies. On geological time-scales, collagen is short-lived compared to fossilized bioapatite, but it is a very robust biomolecule, and it has been repeatedly shown that measurable amounts of collagen can survive under optimal conditions for well over 100 000 years (Jones et al. 2001). Collagen denatures when hydrogen bonds are broken and thereafter fibrils dissolve away relatively quickly, explaining collagen's sensitivity to moisture, temperature and pH conditions.Collins et al. (2002)provided a synthesis of current understanding of the degradation of collagen and other biomolecules in the same special issue of Archaeometry. ...
... Uniform purification procedures are now used, all of which are modifications of the original Longin (1971) method. Chunks or powdered bone/ dentine samples are demineralized in dilute HCl, nowadays at low temperature (5°C; see Richards and Hedges 1999a;Jones et al. 2001), followed by a gelatinization step, then filtration and freeze-drying, before small amounts are combusted and the CO 2 and N 2 fed into a mass spectrometer. Standard deviations of replicate measurements are generally about ±0.1‰ for carbon and ±0.2‰ for nitrogen. ...
... They have done both. Recent progress in extracting good-quality collagen from older material has shown that it can survive under the right conditions for up to 200 000 years (Ambrose 1998;Richards and Hedges 1999a;Jones et al. 2001). This has made it possible to analyse the bone collagen of Late Pleistocene hominins in Europe, where temperatures have been low for most of this time. ...
Article
This review charts the developments and progress made in the application of stable light isotope tools to palaeodietary adaptations from the 1970s onwards. It begins with an outline of the main principles governing the distribution of stable light isotopes in foodwebs and the quality control issues specific to the calcified tissues used in these analyses, and then proceeds to describe the historical landmark studies that have marked major progress, either in their archaeological applications or in enhancing our understanding of the tools. They include the adoption of maize agriculture, marine-focused diets amongst coastal hunter–gatherers, trophic level amongst Glacial-period modern humans and Neanderthals, and the use of savannah resources by early hominins in Africa. Particular attention is given to the progress made in addressing the challenges that have arisen out of these studies, including issues related to the routing of dietary nutrients. I conclude with some firm, and some more speculative, pointers about where the field may be heading in the next decade or so.
... The contaminations include the mineral fraction containing carbonate as well as exogenous organic matter that could have impregnated the bone sample during burial. Although collagen remnants have been identified in fossils as old as dinosaurs (e.g., Wyckoff 1969;Bocherens et al. 1988;Ostrom et al. 1993;Asara et al. 2007), most applications of collagen isotopic biogeochemistry have been restricted to the last 200 kyr (Jones et al. 2001) and to material coming from regions with temperate or cold climates since the rate of collagen loss is temperature dependant (e.g., Holmes et al. 2005). Under warmer climatic conditions, as in Mediterranean areas, and in less protected environments, as in open-air sites, bone is usually less well preserved and collagen is often too altered to yield reliable isotopic signatures. ...
... Paleoclimatological proxies point to a very dry and cold climate during the deposition of this layer (Fizet et al. 1995). Reports of unusually high d 15 N values have also been made for bison bones from England older than 55 ka (Jacobi et al. 2006), and for bison and proboscideans from a 200 ka old English site (Jones et al. 2001). Aridity is often presented as the most likely cause of these positive excursions of d 15 N values (Fizet et al. 1995;Jones et al. 2001), but the biogeochemical behavior of nitrogen under the unique environmental conditions of the Late Pleistocene in Europe is not yet well understood and requires further investigations. ...
... Reports of unusually high d 15 N values have also been made for bison bones from England older than 55 ka (Jacobi et al. 2006), and for bison and proboscideans from a 200 ka old English site (Jones et al. 2001). Aridity is often presented as the most likely cause of these positive excursions of d 15 N values (Fizet et al. 1995;Jones et al. 2001), but the biogeochemical behavior of nitrogen under the unique environmental conditions of the Late Pleistocene in Europe is not yet well understood and requires further investigations. In particular, it would be necessary to consider the possible influence of the herbivore community on the biogeochemistry of nitrogen, since grazing is a factor known to increase the d 15 N values of plants and soils (e.g., Neilson et al. 2002), especially through manuring (e.g., Simpson et al. 1999;Choi et al. 2002;Frank et al. 2004;Dijkstra et al. 2006). ...
Chapter
Full-text available
After reviewing the current knowledge on paleoecological tracking using carbon and nitrogen stable isotopes in fossil bones and teeth, the contribution of this new approach to key questions of Neanderthal diet and ecology is examined. In particular, the determination of ungulate habitat (open environment versus closed forest) is discussed. Thanks to the carbon and nitrogen isotopic differences observed in the main ungulates available as prey to Neanderthal of OIS 3, it is possible to evaluate quantitatively the contribution of different prey in the diet of Neanderthals. The results of this approach suggest that megaherbivores, such as mammoth and woolly rhinoceros, were the most important providers of proteins to the Neanderthal from Saint-Césaire and probably also to one Neanderthal specimen from Spy. In addition, the conclusions of zooarcheological and isotopic dietary determinations are not always in agreement, possibly due to taphonomic biases and site specialization. KeywordsStable isotopes-Carbon-Nitrogen-OIS 3-Prey-Megaherbivores
... Consequently, animal foods will be overrepresented in bone collagen at the expense of low-protein (vegetable) foods, and this bias must be considered when interpreting collagen stable isotope data. Progress in extracting good quality collagen from older material has demonstrated that under the right conditions , bone collagen can survive for up to 200,000 years (Ambrose, 1998; Jones et al., 2001). This has made it possible analyze the bone collagen of Late Pleistocene hominins in certain cases. ...
... Bone collagen is rarely preserved beyond the Late Pleistocene (Jones et al., 2001), so this avenue is not an option for analysis of older hominin material. However, the carbon isotopes in the mineral component can also be used as dietary proxies (Sullivan and Krueger, 1981; Lee-Thorp and van der). ...
Article
Full-text available
Dietary ecology is key to understanding the lifeways and evolutionary pathways of many animals, but determining the diets of long-extinct primates, including early hominins, is a considerable challenge. Although archeological evidence forms a pillar of our understanding of diet and subsistence in the more recent past, for early hominins, the most direct evidence is to be found in the fossils themselves. Here we review the suite of emerging biochemical paleodietary tools based on stable isotope and trace element archives within fossil calcified tissues. We critically assess the contribution of these tools to advancing our understanding of australopith, early Homo, and Neanderthal diets, and placed within the context of contributions of morphological and microwear tools. Perhaps the most significant outcomes to date are the demonstration of high trophic-level diets among Neanderthals in Glacial Europe, and the persistent inclusion of significant amounts of C4 grass-related foods in the diets of both the South African australopiths and Homo in the Pliocene and Pleistocene. These results raise new questions that require improved contextual understanding of these tools from modern ecosystems, but they also clearly show a good deal of promise as quantitative indicators of hominin diets that nicely complement morphological and microwear tools.
... Collagen molecules break-down through time in the burial environment, and the processes of diagenesis, hydrolysis and microbial attack can serve to degrade collagen, resulting in its loss or in the alteration of in vivo stable isotope signatures (Ambrose, 1990;van Klinken, 1999;Collins et al., 2002;Nielsen-Marsh et al., 2007;Smith et al., 2007;Dobberstein et al., 2009). Although the age of deposits can greatly influence the chances of collagen preservation, the rate of protein breakdown largely depends on conditions within the local burial environment Smith et al., 2007) and, where conditions are favourable, well-preserved collagen can be extracted and analysed from Late Pleistocene samples dating to older than 50,000 yr BP (Fizet et al., 1995;Bocherens et al., 1997Bocherens et al., , 1999Bocherens et al., , 2001Jones et al., 2001). ...
... Such preservation is considered rare, and there are very few other published examples of isotope studies on collagen of Table 1 Results of whole bone collagen stable carbon and nitrogen isotope analysis from herbivores from Neumark-Nord 2 (* = > 30 KDa fraction, others were all the b30 to >10 KDa; § denotes samples containing b26.2% C and b 11% N). The stratigraphic sequence (archaeological find level Level 2/2; Horizon A to C) represents b 2000 yr of sediment accumulation (Sier et al., 2011 comparable age or older which meet quality criteria Jones et al., 2001). The breakdown of collagen occurs over time in the burial environment, through chemical degradation (hydrolysis of the peptide bonds in collagen) and microbial attack, which can further weaken the physical structure of collagen and accelerate leaching and biodegradation (Nielsen-Marsh and Jans et al., 2004). ...
Article
Herbivores from the Neumark-Nord 2 archaeological site, Germany, were analysed for bone collagen stable carbon (δ13C) and nitrogen (δ15N) isotope ratios in order to investigate feeding ecology at this early Last Interglacial (Eemian) shallow-lake site. Of 42 faunal samples selected, 23 yielded collagen, demonstrating remarkable preservation for material of this age. The results indicate clear inter-specific differences in δ15N and δ13C values, notably between equids (Equus) and bovids (Bos/Bison), with mean difference Δ15N of + 2‰ measured in the bovids compared to the equids. The potential reasons for these differences are explored, including physiology, herbivore feeding ecology, biogeography and resource partitioning within the local environment. The data are compared to previously published archaeological data, and modern experimental and ecological data, suggesting that these inter-specific differences are not consistent and therefore unlikely to be solely the product of physiology or habitual forage preference. Data from this study are compared to the local vegetation (as reconstructed from pollen profiles), and it is suggested that these trends are likely the result of niche partitioning at the shallow lake site, reflecting the local diversity in vegetational zones. The evidence for resource partitioning amongst Pleistocene herbivore communities at Neumark-Nord 2 and elsewhere is discussed. This study represents one of the largest data sets for collagen of this age, and the implications for our understanding of Late Pleistocene herbivore ecology, local herbivore community behaviour and hominin palaeodietary studies are explored.
... Stable carbon and nitrogen isotope analysis of animal remains are commonly used to reconstruct the palaeodiets and palaeoenvironments of ancient human populations [3,7,9,26,32,33,50,52,53]. Stable isotopic ratios in mammals are dependent on the isotopic composition of plants at the base of the food chain. ...
... Such extreme fractionations are nevertheless rare, at least within any one system, and d 15 N values in biological materials generally range between wÿ10 and C20&. Although stable nitrogen isotopic data are usually limited to studies of animals from the more recent past (within w10 ka due to post-mortem denaturation of biological proteins) they have been used to investigate the diets of ancient hominins, including Neandertals and anatomically modern humans [3,7,26,33,50]. While these studies have provided new insights, there is widespread agreement that the controlling mechanisms for 15 N abundances in terrestrial foodwebs are poorly under- stood [2,51,54] . Importantly, predicted patterns of 15 Nabundances in plants at the base of the food chain are not always consistent. ...
Article
Stable carbon (δ13C) and nitrogen (δ15N) isotope ratios are commonly used to reconstruct palaeodiets and palaeoenvironments. The method is based on our knowledge of isotopic patterns in plants, which are subject to taxonomic and environmental variability. While previous researchers have addressed isotopic variability amongst plants, no studies have looked extensively at a broad suite of taxa over multiple temporal scales from within the savanna biome so as to provide baseline data for palaeodietary and palaeoenvironmental studies. Here we document variations in the isotopic compositions of plants collected over two years from the Kruger National Park, South Africa, with respect to species and anatomical differences, and the influences of geological substrate and spatio-temporal shifts in climate. Results show that environmentally-induced carbon isotopic variations in plants within this region are generally smaller than 2‰, which is lower than what has been previously reported for plants compared across multiple habitat-types. These data suggest that δ13C differences of ∼2‰ or more (or ∼1‰ if the diet is predominantly C4) between animals from a given area reliably indicate real dietary differences. Plant δ15N values vary greatly between different microhabitats (by up to 4‰), responding to a range of environmental influences that may, in turn, significantly influence variation in animal δ15N values.
... Consequently, animal foods will be overrepresented in bone collagen at the expense of low-protein (vegetable) foods, and this bias must be considered when interpreting collagen stable isotope data. Progress in extracting good quality collagen from older material has demonstrated that under the right conditions , bone collagen can survive for up to 200,000 years (Ambrose, 1998; Jones et al., 2001). This has made it possible analyze the bone collagen of Late Pleistocene hominins in certain cases. ...
... Bone collagen is rarely preserved beyond the Late Pleistocene (Jones et al., 2001), so this avenue is not an option for analysis of older hominin material. However, the carbon isotopes in the mineral component can also be used as dietary proxies (Sullivan and Krueger, 1981; Lee-Thorp and van der). ...
Article
Dietary ecology is one key to understanding the biology, lifeways, and evolutionary pathways of many animals. Determining the diets of long-extinct hominins, however, is a considerable challenge. Although archaeological evidence forms a pillar of our understanding of diet and subsistence in the more recent past, for early hominins, the most direct evidence is to be found in the fossils themselves. Here we review the suite of emerging biochemical paleodietary tools based on stable isotope and trace element archives within fossil calcified tissues. We critically assess their contribution to advancing our understanding of australopith, early Homo, and Neanderthal diets within the broader context of non-biogeochemical techniques for dietary reconstruction, such as morphology and dental microwear analysis. The most significant outcomes to date are the demonstration of high trophic-level diets among Neanderthals and Late Pleistocene modern humans in Glacial Europe, and the persistent inclusion of C(4) grass-related foods in the diets of Plio-Pleistocene hominins in South Africa. Such studies clearly show the promise of biogeochemical techniques for testing hypotheses about the diets of early hominins. Nevertheless, we argue that more contextual data from modern ecosystem and experimental studies are needed if we are to fully realize their potential.
... Preservation of bones is determined by the ability of the organic and inorganic phases to survive in distinct climatic and pedogenic contexts. The organic phase is primarily composed of collagen, a protein that can remain intact in archaeological contexts up to 100,000 years old (Jones et al., 2001), unless an archaeological site is located in the tropics, or in regions with very hot/dry summers and wet winters. Because bone collagen enables measurement of both stable carbon and nitrogen isotope values, it has become the preferred sampling material for palaeodietary studies; although it primarily provides information about the protein component of consumers' diets (Sullivan and Krueger, 1981;Ambrose and Norr, 1993;Lee-Thorp, 2008). ...
Article
Isotopic analysis has become one of the most popular arenas of archaeological science, in part due to its versatility to uncover intriguing insights from a range of organic and inorganic archaeological materials. However, alongside an increase in popularity, the field has seen the rise of dissemination of publications that do not pass quality control, do not apply robust interpretative frameworks, or do not report data in ways that would make them amenable to critical evaluation or inclusion in large meta-analyses. This paper represents an effort to clarify some of the most pressing weaknesses and misconceptions in ‘traditional’ applications of isotopic techniques in archaeology: measurement of stable carbon, nitrogen, and oxygen isotope values of organic and inorganic materials (bulk bone collagen, bulk tooth dentine, seeds; bulk and incremental tooth enamel, molluscan shells), and strontium isotope ratio analysis of tooth enamel and cremated bone. The discussion centers on three key aspects of research: (1) Selecting samples, with advice on building comparative baselines (or more appropriately ‘baseintervals’) and words of caution on interpreting stable carbon isotope values measured during AMS radiocarbon dating. (2) Handling data, including tips on exploratory data analysis, graphical visualization, and statistical assessment of differences between groups; with particular reference to the Statement on p-values published by the American Statistical Association. (3) Reporting results, with advice on using correct terminology and decimal points, calculating measurement precision and accuracy, and communicating results using effective scientific language. The advice provided in this paper does not cover all aspects of project design and dissemination but will hopefully provide clarification within the above key areas and inspire further discussion of effective and impactful applications of isotopic techniques in archaeology.
... Stable isotope analyses of bone collagen are commonly performed in archaeological (e.g., Schoeninger et al., 1983;Bocherens et al., 2015;Naito et al., 2016;Boethius and Ahlström, 2018) and ecological studies (e.g., Schoeninger and DeNiro, 1984;Matsubayashi et al., 2015;Swift et al., 2018;Han et al., 2019) for the purpose of reconstructing an organism's diet. In humans and other animals, bone collagen generally records long-term isotopic information (Stenhouse and Baxter, 1979;Wild et al., 2000;Hedges et al., 2007) and can be preserved in bone for over 100,000 years (Jones et al., 2001). Therefore, isotopic values obtained from bone collagen can be used to reconstruct the diet and environment of animals that lived in the past. ...
Article
Full-text available
Bone collagen of modern and ancient animals is a useful tissue for isotope analyses because it is stable over time. However, uncertainty regarding metabolic turnover processes of bone collagen can make isotope analysis difficult to correlate with relevant life history information. We used radiocarbon (14 C) dating to examine turnover within cortical bone and to investigate retrospective isotope ratios along the growth direction of mid-shaft femurs of several large mammals with long lifespan , including brown bear (Ursus arctos), sika deer (Cervus nippon), Japanese serow (Capricornis crispus), and Japanese macaque (Macaca fuscata). The individuals examined died between 1971 and 1986, and their bones were thus expected to contain radiocarbon generated by nuclear bomb testing, which led to an atmospheric 14 C spike around 1964. Therefore, 14 C dating could be used to date sections of cortical bone at fine scale. The 14 C ages in the bone sections of all specimens except the Japanese serow specimen showed similar trends; perimedullary bone sections contained younger carbon, which has depleted 14 C after the peak of the 14 C spike, whereas 14 C ages became rapidly older in midcortical sections before gradually becoming younger towards the bone surface. We observed metabolic turnover of collagen driven by bone remodelling in perimedullary bone sections, but we observed no evidence of remodelling in midcortical and pericortical sections. Thus, our results confirm that bone collagen in femoral cortical bone records retrospective isotopic information during skeletal growth of mammals and suggest that entire femoral cortical bone of aged terrestrial mammals represents isotopic values during adolescence rather than an average value from several years prior to death.
... Since bone collagen can be sensitive to degradation, the majority of stable isotope analyses using bone collagen have been carried out on relatively young material with a Late Pleistocene age (usually within the radiocarbon time scale, i.e., up to 50 ka [thousands of years ago]; e.g., Drucker et al., 2003;Richards and Hedges, 2003;Stevens and Hedges, 2004;Szpak et al., 2010). Reliable results from older material have been published but are very rare (e.g., Bocherens et al., 1999;Jones et al., 2001;Palmqvist et al., 2008;Britton et al., 2012). ...
... Since bone collagen can be sensitive to degradation, the majority of stable isotope analyses using bone collagen have been carried out on relatively young material with a Late Pleistocene age (usually within the radiocarbon time scale, i.e., up to 50 ka [thousands of years ago]; e.g., Drucker et al., 2003;Richards and Hedges, 2003;Stevens and Hedges, 2004;Szpak et al., 2010). Reliable results from older material have been published but are very rare (e.g., Bocherens et al., 1999;Jones et al., 2001;Palmqvist et al., 2008;Britton et al., 2012). ...
... This study paved the way for a plethora of applications in the following decades most of which relied on stable isotope abundances in bone collagen. However, since collagen is rarely preserved for very long on geological timescales, most of these studies were confined to the fairly recent past (but see Jones et al. (2001) showing collagen survival for up to 200000 years under optimal preservation conditions). ...
... This study paved the way for a plethora of innovative applications in the following decades that relied on stable isotope abundance data, but these studies tended to use bone collagen. Collagen is a robust tissue, yet it rarely survives beyond 10 000 years, although there is good evidence that it sometimes survives much longer, perhaps as much as 200 000 years (Jones et al., 2001), and some have argued that stable isotopes can be recovered in bone organics from the Cretaceous (Ostrom et al., 1993). Although there is certainly much promise in ancient organics, it is clear that researchers are a long way from being able to retrieve meaningful information from such archives on a regular basis, and it is for this reason that researchers turned their attention to the possibility of getting carbon isotope ratios from the inorganic portion of mineralized tissues. ...
Article
Stable isotopes provide unique insights into the diets of ancient mammals. Carbon and oxygen isotopes are inherited directly from diet and body water and are preserved in tooth enamel for millions of years. The carbon isotopes in enamel apatite provide information about diet with respect to the consumption of plants using the C3 and C4 photosynthetic pathways, which has been an important distinction in many environments over the last 10 My. Carbon and oxygen isotopes have been useful for testing hypotheses about the adoption of C4 agriculture (e.g., maize, millet) by human populations, the allocation of resources among early (>1 Ma) hominins, niche partitioning among ancient communities, and issues of physiology and water use.
... However, collagen is a good repository from which information on dietary habits, nutritional status, approximate postmortem interval, and potentially even geographic provenance can be obtained. Survival rates for bone collagen or tooth-dentine collagen of up to 100 000 years have been repeatedly reported [10] and contextual interpretation of the information contained therein has yielded insights into infant feeding and even the average/maximum age of weaning in medieval societies [11] [12] [13] [14]. ...
... Carbon and nitrogen isotopes of bone collagen are useful proxies for estimating the dietary niches and habitat preferences of extinct mammals (DeNiro and Epstein, 1978;Ambrose and DeNiro, 1986;Jones et al., 2001;Palmqvist et al., 2003;Coltrain et al., 2004;Bocherens et al., 2005;Phillips and Eldridge, 2006). According to their photosynthetic pathways (Edwards and Walker, 1983), terrestrial vegetation comprises C 3 plants (all trees and bushes, temperate shrubs, and grasses adapted to cool, moist climate or high altitude) and C 4 plants (tropical, arid-adapted grasses and some sedges). ...
Article
Biogeochemical (δ13C, δ15N, and δ18O values) and ecomorphological analyses of the early Pleistocene fauna of Venta Mena (Orce, Guadix-Baza basin, SE Spain) provide interesting clues on the physiology, dietary regimes, habitat preferences, and ecological interactions of large mammals. Such inferences are useful in deciphering aspects of paleocommunity structure and predator-prey relationships. Specifically, the hypsodonty index combined with δ13C values allows classifying the ungulates among grazers from open habitat (Equus altidens, Bison sp., Praeovibos sp., Hemitragus albus, Hippopotamus antiquus, and Mammuthus meridionalis), mixed feeders (Soergelia minor and Pseudodama sp.), and browsers from canopy areas (Stephanorhinus sp. and Praemegaceros cf. verticornis). Given that δ13C values indicate that all these herbivores fed exclusively on C3, plants, significant differences in isotopic values between perissodactyls (monogastric, hindgut fermenters) and ruminants (foregut fermenters) reflect differences in digestive efficiency. Values of δ18O indicate the dietary water source of ungulates, revealing that Pseudodama sp., Hemitragus albus, and Soergelia minor obtained a significant fraction of their metabolic water from vegetation. Carnivores show higher δ15N values than herbivores, which records the isotopic enrichment expected with an increase in trophic level. Hippopotamus antiquus and Praeovibos sp. have unexpectedly high δ15N values, suggesting that they predominantly consumed aquatic plants and lichens, respectively. Inferences on predator-prey relationships, derived from the use of linear mixing models, indicate resource partitioning among sympatric predators; saber-tooth Megantereon whitei and jaguar Panthera cf. gombaszoegensis were ambushers in dosed habitat while saber-tooth Homotherium latidens and wild dog Lycaon lycaonoides were coursing predators in open plains. The giant hyena Pachycrocuta brevirostris scavenged the prey of these hypercarnivores.
... However, collagen is a good repository from which information on dietary habits, nutritional status, approximate postmortem interval, and potentially even geographic provenance can be obtained. Survival rates for bone collagen or tooth-dentine collagen of up to 100 000 years have been repeatedly reported [10] and contextual interpretation of the information contained therein has yielded insights into infant feeding and even the average/maximum age of weaning in medieval societies [11][12][13][14]. ...
Chapter
In most of the different body deposition scenarios but for cremation, bone and teeth are typically the human remains that remain intact the longest after death has occurred. In the absence of any distinguishing features or other evidence that could help with positive victim identification, stable isotope profiles or signatures can provide useful information about a person's diet and geographic life trajectory, which can bring focus to an investigation involving unidentified victims of serious crime or mass disasters. No matter what the case circumstance, victim identification is of paramount importance to provide closure for next of kin and to help dealing with legal processes ensuing from the death of a person. In cases of serious crime, identification of the victim is a major stepping stone toward identifying the perpetrator/s.Keywords:bioapatite;bone;carbonate;collagen;diet;forensic intelligence;geographic origin;human provenance;isotope ratio;victim identification;stable isotope profile
... While it is no longer intact collagen, the presence of this material suggests that proteinaceous compounds can survive in the geologic record longer than previously suspected. A review of published isotopic values from well preserved collagen suggests that the oldest specimens containing intact protein in bones and teeth are ~50,000 years old (Bocherens et al. 1994b, Fizet et al. 1995, Bocherens et al. 1996, Bocherens et al. 1997, Iacumin et al. 2000, Richards and Hedges 2003, Coltrain et al. 2004) with rare exceptions beyond this age (Bocherens et al. 1988, Ostrom et al. 1993, Jones et al. 2001. Therefore it has become a common "rule of thumb" that proteinaceous material can only be extracted from bone or tooth specimens less than ~50,000 years old, thus limiting nitrogen isotopic studies of animal tissues to the late Pleistocene. ...
... In cave sites located in areas with temperate climatic conditions, collagen is usually preserved until around 100,000 years ago , but some exceptional cases several times older have been documented (e.g. Jones et al., 2001;Drucker et al., 2014;Kuitems et al., in press). Besides, there is an increasing body of evidence that suggests that waterlogged burial deposits may also be favourable to long term collagen preservation (e.g. ...
Article
Isotopic tracking of carnivore palaeoecology is a relatively new approach that yielded important results for the study of the non-analogue mammoth steppe biome. After describing the prerequisite to apply this approach and the possible complications, the main achievements will be described for extinct carnivore species such as scimitar-tooth cat Homotherium serum, cave lion Panthera spelaea, giant short-faced bear Arctodus simus, cave bear Ursus spelaeus s.l., as well as for ancient representatives of extant species such as brown bear Ursus arctos and wolf Canis lupus. Isotopic tracking showed that scimitar-tooth cats in Alaska were not specialist proboscidean predators but rather generalist consumers of other large herbivores. The majority of cave lions analysed so far were focused on reindeer, some individuals were specialized on cave bears, especially in contexts of competition with cave hyenas. Giant short-faced bears in Alaska were not pure herbivores and consumed meat from reindeer, muskoxen and possibly other predators, but may have still incorporated plant resources in their menu. In contrast, all cave bear populations studied so far for which a clear dietary reconstruction could be done were virtually pure herbivores, only a few cases are still unclear. Interestingly, brown bears used the opposite extreme of the dietary spectrum when competing with other large bears such as cave bears and giant short-faced bears, i.e. were more carnivorous in Europe and more herbivorous in Alaska. Finally wolves seem to have been outcompeted by hyenas but became dominant predators during the Lateglacial in Europe to the expense of the last cave lions. The results obtained through this approach are also relevant for improving conservation strategies of endangered extant large carnivores.
... This relationship is, however, strongly dependent upon additional factors such as climate, seasonality, diet, metabolism and water turnover (Kohn, Schoeninger and Valley 1998). The relation between atmospheric (surface water) oxygen and phosphate oxygen for elephants is defined by the equation (Jones et al. 2001): To estimate the local range of ∂ 18 O w the ∂ 18 O of modern precipitation is used. Based on the annual mean weighted ∂ 18 O data in the GNIP/ISOHIS database from the International Atomic Energy Agency (IAEA) the ∂ 18 O range of modern precipitation in Africa varies between 0‰ and −4‰ (IAEA/Wmo, ISOHIS database 2004). ...
Article
During the last couple of decades, excavations in Amsterdam, the Netherlands, have uncovered many artefacts from post-medieval contexts made of elephant ivory. Historical and archaeological sources show the import of great quantities of elephant ivory by the Dutch West India Company to the Dutch Republic in the seventeenth and eighteenth centuries. These sources, however, mention only the West African coast, and the Gold Coast in particular, as the source of this ivory and do not shed any light upon the exact locations from which the ivory originated. In order to try to better determine the geographical origins of post-medieval ivory found in Amsterdam, multi-isotope analyses were conducted on twenty-one samples from objects excavated by the Office for Monuments & Archaeology. The results are the first of their kind for West Africa and suggest that ivory was derived from elephants that inhabited different parts of coastal and interior West Africa.
... Since bone collagen can be sensitive to degradation, the majority of stable isotope analyses using bone collagen have been carried out on relatively young material with a Late Pleistocene age (usually within the radiocarbon time scale, i.e., up to 50 ka [thousands of years ago]; e.g., Drucker et al., 2003;Richards and Hedges, 2003;Stevens and Hedges, 2004;Szpak et al., 2010). Reliable results from older material have been published but are very rare (e.g., Bocherens et al., 1999;Jones et al., 2001;Palmqvist et al., 2008;Britton et al., 2012). ...
... In bone, the collagen fibrils are arranged in four main patterns. However, collagen is most often characterized by the only amino acid composition, without characterization of its structure and size (Jones et al., 2001). Moreover, collagen is the main component, but not the only component of the bone organic matrices. ...
Article
Thin sections, scanning electron microscopy (SEM), diffraction X (DRX) and infrared spectrometry (FTIR) have been used to study the structure, mineralogy, crystallinity and bulk composition of fossil rodent long bones extracted from a succession of sedimentary layers in a cave from Morocco (Neolithic - Middle Palaeolithic, El Harhoura 2). The microstructure of fossil bones is well-preserved at this scale of observation, and encrusted deposits are rare. All bones are preserved in apatite, but the crystallinity is modified, as well as the crystallite shape, the organic content and the organic-mineral ratio. No fluor enrichment has been observed. Alone or together, the studied parameters do not show a regular trend from the upper to the lower layers of the cave. The preservation of the fossil bones does not confirm the sequence of arid and humid periods inferred from taphonomic analyses.
... This study paved the way for a plethora of innovative applications in the following decades that relied on stable isotope abundance data, but these studies tended to use bone collagen. Collagen is a robust tissue, yet it rarely survives beyond 10 000 years, although there is good evidence that it sometimes survives much longer, perhaps as much as 200 000 years (Jones et al., 2001), and some have argued that stable isotopes can be recovered in bone organics from the Cretaceous (Ostrom et al., 1993). Although there is certainly much promise in ancient organics, it is clear that researchers are a long way from being able to retrieve meaningful information from such archives on a regular basis, and it is for this reason that researchers turned their attention to the possibility of getting carbon isotope ratios from the inorganic portion of mineralized tissues. ...
... The estimated minimum and maximum spring water elevations (800 and 1930 m) can also be considered as plausible extreme limits of the geographic distribution of modern bear population in Pindos ridge (max. elevation 2637 m).Longinelli, 1984; Genoni et al., 1998; Jones et al., 2001; Hoppe, 2006 values are between À8.0& and À12.3& VSMOW. In both cases, the most negative values of the palaeo-spring waters are much more depleted than the isotopic composition of modern spring water of Greece (Dotsika et al., 2010). ...
Article
The O and C isotopic composition of enamel carbonate hydroxy-apatite in the teeth of certain animals reflects the oxygen isotope composition of the water they ingest. The isotopic composition of meteoric water is well-correlated with mean annual temperature so that there is potential for recovering palaeo-temperature of the regions where the animals lived. Analyses were made on enamel from fossil teeth of Ursus Ingressus from Arkoudospilia Cave in Northern Greece. Analyses were made also on modern teeth of Ursus from different areas in Greece. Oxygen and deuterium isotopic analyses of water were also made. Although the preservation of primary oxygen isotopic composition of enamel carbonate hydroxy-apatite was more difficult to assess, however the isotopic signals seem to have utility for the paleoenvironmental reconstructions of the studied area.
... This assumption does not consider the abovementioned altitude (800 m) as an absolute value, but rather a theoretical minimum of local significance that may provide an arithmetic value for further calculations. Bone carbonate of modern bears from the, the observed difference of 1.9& for modern bear bone apatite isotopic composition corresponds to a hypsometrical difference of approximately 1130 m that is an absolute altitude ofFig. 5. Variation of the oxygen isotopic composition of Grevena area (Pindos) spring waters with altitude.Longinelli, 1984;Genoni et al., 1998;Jones et al., 2001;Hoppe, 2006values are between À8.0& and À12.3& VSMOW. In both cases, the most negative values of the palaeo-spring waters are much more depleted than the isotopic composition of modern spring water of Greece (Dotsika et al., 2010). ...
Article
Bones and teeth samples of Ursus ingressus from Loutra Arideas Cave (Greece) were used to determine the diet of this extinct species and to reconstruct the palaeoclimatic conditions. The age range of the fossil layers is from 32ka BP to a maximum of 38ka BP (radiocarbon dating). The method used was the isotopic analysis of carbonate bioapatite of fossil cave bear (U. ingressus) and brown bear (Ursus arctos) bone and tooth fragments, as well as water samples from springs of the Pindos area, the modern bear habitat. Several possible effects that may affect the isotopic composition of apatite were investigated, including age, sex, tooth type and diagenesis.The comparison of isotopic values of Greek U. ingressus to other Pleistocene Ursidae species in Europe, exhibit more positive and more variable δ13C values. These positive values are attributed mainly to diet, to bear physiology (differences among species) and to climate conditions. The diet of U. ingressus apparently was based mainly on vegetable matter, however with a variable component of animal protein.The variation observed in the δ18O values of U. arctos tooth enamel samples were attributed to environmental conditions, and through the spring waters isotopic gradient for the region of Northeastern Pindos, to differences in the bear’s habitat elevation. Assuming similarity of the metabolism of modern and fossil bear species and incorporating only the effect of drinking water on the isotopic composition of body water, an equation has been derived that could be used to provide palaeoclimatic information.
... Even where conditions are conducive to preservation, collagen has a shorter " shelf-life " than bioapatite since it denatures and dissolves quite quickly on geological time-scales. However, it has been repeatedly shown that measurable amounts of collagen can survive under optimal conditions for over 100,000 yr (Jones et al., 2001). A standard set of quality control measures, involving atomic C:N ratios, complemented by collagen yield and weight percent C and N (Ambrose, 1990; Van Klinken, 1999), is routinely practiced by the stable isotope community and most radiocarbon laboratories. ...
... The bone and tooth microstructure is often well-preserved down to the μm-scale in fossil specimens recording growth marks and other histological features that are often used for life history reconstructions of extinct animals or humans (e.g., Chinsamy-Turan, 2005; Erickson, 2005; Smith, 2008). In contrast, biomolecules such as proteins or DNA are usually only preserved in Holocene or Late Pleistocene skeletal remains (e.g., Hagelberg et al., 1994; Greenwood et al., 1999; Jones et al., 2001; Collins et al., 2002; Adler et al., 2011;) and only rarely reported for pre-Quaternary specimens (e.g., Asara et al., 2007; Schweitzer et al., 2009). The latter finds are, however, controversial. ...
... In temperate latitudes bone collagen persists well into the Pleistocene (Jones et al., 2001;Palmqvist et al., 2003; but until recently it has not been possible to readily access its protein sequence data. The ability to either directly sequence or to peptide fingerprint bone collagen has been transformed by the use of soft-ionization mass spectrometry (Ostrom et al., 2000;Nielsen-Marsh et al., 2002;Ostrom et al., 2006); see below. ...
Article
The last few years have seen an enormous proliferation of ancient biomolecules research, especially in the field of ancient DNA. Ancient DNA studies have been transformed by the advent of next generation sequencing, with the first Pleistocene sample being analysed in 2005, and several complete and draft genomes that have been compiled from ancient DNA to date. At the same time, although less conspicuous, research on ancient proteins has also seen advances, with the time limit for research on ancient biomolecules now extending to over 1 million years. Here we review which effects these developments have on research in Quaternary science. We identify several lines of research that have the potential to profit substantially from these recent developments in ancient biomolecules research. First, the identification of taxa can be made using ancient biomolecules, and in the case of ancient DNA, specimens can even be assigned to specific populations within a species. Second, increasingly large DNA data sets from Pleistocene animals allow the elucidation of ever more precise pictures of the population dynamic processes whereby organisms respond to climate and environmental change. With the accompanying better understanding of process in the Quaternary, past ecologies can also more realistically be interpreted from proxy data sets. The dominant message from this research so far is that the Quaternary saw a great deal more dynamism in populations than had been forecast by conventional palaeoecology. This suggests that reconstructions of past environmental conditions need to be done with caution. Third, ancient DNA can also now be obtained directly from sediments to elucidate the presence of both plant and animal species in an area even in the absence of identifiable fossils, be it macro- or micro-fossils. Finally, the analysis of proteins enables the identification of bone remains to genus and sometimes species level far beyond the survival time of DNA, at least in temperate regions, illustrating that precise data is now forthcoming from seemingly unlikely sources. Together, these approaches allow the study of environmental dynamics throughout a substantial part, and perhaps even the entire Quaternary (the last 2.6 million years).
... Nitrogen is restricted to the organic phase of hard tissue (i.e., bone collagen), and because this has a maximum preservation limit of some 200 kya even under perfect conditions (Jones et al., 2001), nitrogen isotope studies are limited to more recent (i.e., Late Pleistocene and Holocene) hominin specimens. Its most significant application to hominin paleodietary research has been in the study of Neandertal remains, where relatively high 15 N/ 14 N ratios suggest that they derived their dietary protein primarily from animal rather than plant foods (Bocherens et al., 1991(Bocherens et al., , 1999Richards et al., 2000;Richards and Schmitz, 2008). ...
Article
Determining the diet of an extinct species is paramount in any attempt to reconstruct its paleoecology. Because the distribution and mechanical properties of food items may impact postcranial, cranial, mandibular, and dental morphologies related to their procurement, ingestion, and mastication, these anatomical attributes have been studied intensively. However, while mechanical environments influence skeletal and dental features, it is not clear to what extent they dictate particular morphologies. Although biomechanical explanations have been widely applied to extinct hominins in attempts to retrodict dietary proclivities, morphology may say as much about what they were capable of eating, and perhaps more about phylogenetic history, than about the nature of the diet. Anatomical attributes may establish boundary limits, but direct evidence left by the foods that were actually (rather than hypothetically) consumed is required to reconstruct diet. Dental microwear and the stable light isotope chemistry of tooth enamel provide such evidence, and are especially powerful when used in tandem. We review the foundations for microwear and biogeochemistry in diet reconstruction, and discuss this evidence for six early hominin species (Ardipithecus ramidus, Australopithecus anamensis, Au. afarensis, Au. africanus, Paranthropus robustus, and P. boisei). The dietary signals derived from microwear and isotope chemistry are sometimes at odds with inferences from biomechanical approaches, a potentially disquieting conundrum that is particularly evident for several species.
... In temperate latitudes bone collagen persists well into the Pleistocene (Jones et al., 2001;Palmqvist et al., 2003; but until recently it has not been possible to readily access its protein sequence data. The ability to either directly sequence or to peptide fingerprint bone collagen has been transformed by the use of soft-ionization mass spectrometry (Ostrom et al., 2000;Nielsen-Marsh et al., 2002;Ostrom et al., 2006); see below. ...
... The measurement of carbon-and nitrogen-isotope ratios in an animal's bone collagen provides an indication of aspects of its overall diet for the last few years of life (DeNiro and Epstein, 1978). Apart from a report on bone collagen preserved in Late Cretaceous dinosaurs (Ostrom et al., 1993), original carbon-and nitrogen-isotope compositions have been retrieved from organic residues in fossils as old as 200 ka (Jones et al., 2001), though adequate preservation in such ancient specimens is rare (Bocherens et al., 1996a;Gröcke, 1997a). Thus, the extraction of collagen from 77 out of 105 fossil bone samples of Venta Micena, a locality with an age of~1.5 Ma, constitutes an example of unusual biomolecular preservation. ...
Article
Research into the reconstruction of ancient communities in terms of dietary regimes, habitat preferences and ecological interactions of species has focused predominantly on biogeochemistry or ecomorphology alone and not in combination. The Venta Micena site (Orce, Guadix–Baza basin, SE Spain) has an early Pleistocene vertebrate assemblage with exceptional biomolecular preservation. Collagen was successfully extracted from 77 bone samples of 18 species of large mammals, which allowed analyses of carbon- and nitrogen-isotopes. δ13C, δ15N and δ18O ratios combined with ecomorphological indexes provide interesting clues on the autecology and palaeophysiology of extinct species, which help in deciphering aspects of community trophic structure and predator–prey interactions. Specifically, morphometric ratios (e.g., hypsodonty index and relative length of the lower premolar tooth row; [Palmqvist, P., Gröcke, D.R., Arribas, A., Fariña, R. 2003. Paleoecological reconstruction of a lower Pleistocene large mammals community using biogeochemical (δ13C, δ15N, δ18O, Sr:Zn) and ecomorphological approaches. Paleobiology 29, 204–228.]) allow classifying the ungulates among grazers (Equus altidens, Bison sp., Praeovibos sp., Hemitragus albus, Hippopotamus antiquus, and Mammuthus meridionalis), mized-feeders (Soergelia minor and Pseudodama sp.) and browsers (Stephanorhinus sp. and Praemegaceros cf. verticornis). However, δ13C values reveal that these ungulates consumed exclusively C3 plants and significant differences in isotopic values between perissodactyls (monogastric, hindgut fermenters) and ruminants (foregut fermenters) must reflect physiological differences related to their rates of methane production and digestive efficiency. δ18O ratios allow the interpretation of the dietary water source of these species, suggesting that fallow deer Pseudodama sp., goat H. albus and ovibovine S. minor obtained a significant fraction of their metabolic water from the vegetation consumed. Carnivore species have higher δ15N values than herbivores, which records the isotopic enrichment expected with an increase in trophic level. However, the unexpectedly high δ15N values of hippo H. antiquus and muskoxen Praeovibos sp. suggest that these ungulates predominantly consumed aquatic plants and lichens, respectively. Inferences on predator–prey relationships within this ancient community, derived from the dual linear mixing model, indicate resource partitioning among sympatric predators, suggesting that sabre-tooth Megantereon whitei and jaguar Panthera cf. gombaszoegensis were ambushers of forest environments while sabre-tooth Homotherium latidens and wild dog Lycaon lycaonoides were coursing predators in open habitat. The giant, short-faced hyena Pachycrocuta brevirostris scavenged the prey of these hypercarnivores.
... Bell et al., 1996;Tuross et al., 1989;Trueman et al., 2004) and lasts usually for a few ka to tens of ka (e.g. Kohn and Law, 2006), although under special conditions of preservation it may extend to 200 ka or more ( Jones et al., 2001;Palmqvist et al., 2003;Schweitzer et al., 2005). The early diagenesis of bones is characterized by important changes in protein content, histology, apatite crystallinity, porosity and mineralogy (e.g. ...
Article
Fossil bones and teeth of Late Pleistocene terrestrial mammals from Rhine River gravels (RS) and the North Sea (NS), that have been exposed to chemically and isotopically distinct diagenetic fluids (fresh water versus seawater), were investigated to study the effects of early diagenesis on biogenic apatite. Changes in phosphate oxygen isotopic composition (δ¹⁸OPO4), nitrogen content (wt.% N) and rare earth element (REE) concentrations were measured along profiles within bones that have not been completely fossilized, and in skeletal tissues (bone, dentine, enamel) with different susceptibilities to diagenetic alteration.
... The tissue of interest for the zooarchaeologist is mostly collagen, in bone and dentine, due to its potential for long-term preservation (e.g. [9,10,35]). The actual value of the isotopic shift between the carbon isotopic composition of diet and that of collagen is crucial for interpreting the measured values. ...
Article
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The subsistence patterns of Iron Age and Historical period humans from south-western Turkmenistan have been reconstructed using the carbon and nitrogen isotopic compositions of archaeological faunal and human bones. A qualitative comparison of the isotopic signatures points to a small proportion of ruminant meat and dairy in human diet for both periods. The ranges of proportions of dietary items yielded by a quantitative approach based on concentration dependent mixing models confirm the high proportions of plant food in the average diet, and show little change in the reconstructed diet for both periods. A comparison of results from zooarchaeological and isotopic approaches illustrates their complementarity in subsistence patterns reconstruction.
... Although DNA is more informative, there have already been several reports of the successful sequencing of ancient mammoth and mastodon collagen peptides (Schweitzer et al., 2002; Asara et al., 2007). Collagen is by far the most abundant protein in bone and has long been known to survive well in Quaternary fossils through its use in radiocarbon dating (Longin, 1971) and stable isotope analyses (Jones et al., 2001). The Proboscidea are an iconic order with a rich fossil record which have been a focus of molecular analysis (Rohland et al., 2007; Miller et al., 2008). ...
Article
Near-complete collagen (I) sequences are proposed for elephantid and mammutid taxa, based upon available African elephant genomic data and supported with LC–MALDI-MS/MS and LC–ESI-MS/MS analyses of collagen digests from proboscidean bone. Collagen sequence coverage was investigated from several specimens of two extinct mammoths (Mammuthus trogontherii and Mammuthus primigenius), the extinct American mastodon (Mammut americanum), the extinct straight-tusked elephant (Elephas (Palaeoloxodon) antiquus) and extant Asian (Elephas maximus) and African (Loxodonta africana) elephants and compared between the two ionization techniques used. Two suspected mammoth fossils from the British Middle Pleistocene (Cromerian) deposits of the West Runton Forest Bed were analysed to investigate the potential use of peptide mass spectrometry for fossil identification. Despite the age of the fossils, sufficient peptides were obtained to identify these as elephantid, and sufficient sequence variation to discriminate elephantid and mammutid collagen (I). In-depth LC–MS analyses further failed to identify a peptide that could be used to reliably distinguish between the three genera of elephantids (Elephas, Loxodonta and Mammuthus), an observation consistent with predicted amino acid substitution rates between these species.
... Since the pioneering works done by DeNiro and collaborators (DeNiro and Epstein 1981; DeNiro 1983, 1984), the 15 N/ 14 N isotopic ratio of bone collagen was extensively used to reconstruct extant and past trophic web structures (e.g., Minagawa and Wada 1984; Balter et al. 2002; Post 2002), paleodiet (e.g., Richard et al. 2003), paleoenvironments (e.g., Iacumin et al. 2000; Jones et al. 2001; Stevens and Hedges 2004), and migration pathways (Koch et al. 1995; Hobson 1999). A 15 N-enrichment between an organism and its diet (D 15 N) was systematically documented (e.g., DeNiro and Epstein 1981; Ambrose 2000) with a D 15 N between the lean mass and the diet (D 15 N lm–d ) usually ranging from 3 to 5& vs AIR (e.g., Minagawa and Wada 1984; Schoeninger and DeNiro 1984; Ambrose 2000 ). ...
... Since the pioneering works done by DeNiro and collaborators (DeNiro and Epstein 1981; DeNiro 1983, 1984), the 15 N/ 14 N isotopic ratio of bone collagen was extensively used to reconstruct extant and past trophic web structures (e.g., Minagawa and Wada 1984; Balter et al. 2002; Post 2002), paleodiet (e.g., Richard et al. 2003), paleoenvironments (e.g., Iacumin et al. 2000; Jones et al. 2001; Stevens and Hedges 2004), and migration pathways (Koch et al. 1995; Hobson 1999). A 15 N-enrichment between an organism and its diet (D 15 N) was systematically documented (e.g., DeNiro and Epstein 1981; Ambrose 2000) with a D 15 N between the lean mass and the diet (D 15 N lm–d ) usually ranging from 3 to 5& vs AIR (e.g., Minagawa and Wada 1984; Schoeninger and DeNiro 1984; Ambrose 2000 ). ...
Article
The 15N/14N signature of animal proteins is now commonly used to understand their physiology and quantify the flows of nutrient in trophic webs. These studies assume that animals are predictably 15N-enriched relative to their food, but the isotopic mechanism which accounts for this enrichment remains unknown. We developed a box model of the nitrogen isotope cycle in mammals in order to predict the 15N/14N ratios of body reservoirs as a function of time, N intake and body mass. Results of modeling show that a combination of kinetic isotope fractionation during the N transfer between amines and equilibrium fractionation related to the reversible conversion of N-amine into ammonia is required to account for the well-established approximately 4 per thousand 15N-enrichment of body proteins relative to the diet. This isotopic enrichment observed in proteins is due to the partial recycling of 15N-enriched urea and the urinary excretion of a fraction of the strongly 15N-depleted ammonia reservoir. For a given body mass and diet delta15N, the isotopic compositions are mainly controlled by the N intake. Increase of the urea turnover combined with a decrease of the N intake lead to calculate a delta15N increase of the proteins, in agreement with the observed increase of collagen delta15N of herbivorous animals with aridity. We further show that the low delta15N collagen values of cave bears cannot be attributed to the dormancy periods as it is commonly thought, but inversely to the hyperphagia behavior. This model highlights the need for experimental investigations performed with large mammals in order to improve our understanding of natural variations of delta15N collagen.
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This represents one of several sections of "A Bibliography Related to Crime Scene Interpretation with Emphases in Geotaphonomic and Forensic Archaeological Field Techniques, Nineteenth Edition" (The complete bibliography is also included at ResearchGate.net.). This is the most recent edition of a bibliography containing resources for multiple areas of crime scene, and particularly outdoor crime scene, investigations. It replaces the prior edition and contains approximately 10,000 additional citations. As an ongoing project, additional references, as encountered, will be added to future editions. Popular and scientific references to the use of stable isotopes in identifying skeletal remains; or, more accurately, identifying geographical ranges in which the decedent may have lived, are the focus of this section. It also includes topics such as Carbon 14 dating and bomb pulse data. Stable isotope analyses may provide investigators clues to the spatial history of unidentified victims. Our bones and teeth, throughout our lives become reservoirs for those chemical elements to which we are exposed. The longer those exposures to the varied concentrations of different elements in different areas of the world, the more likely the victim can be determined as having resided in a particular area. By knowing the areas inhabited by a victim, the more likely investigators will be able to track down his, or her, identity. Unlike radioactive isotopes, stable isotopes never disintegrate. Schwarz, (2007), provides a good example of the forensic value of stable isotopes: "Most of the O atoms in our body come from the water we drink, and is usually isotopically like the precipitation where we live. Therefore, we can often learn where a person lived from the isotopic composition of their teeth and bones. Fortunately, we now have maps showing the distribution of 18O/16O ratios in precipitation falling over North America and Europe which we can use to help trace the place of origin of a murder victim. Even burned remains can be analyzed this way." (Schwarz, 2007:28) Like DNA, stable isotope analyses will continue to be developed and be refined. And like DNA analyses, it may someday be a staple in the forensic scientist's toolbox. Because stable isotope analysis is so dependent on the proper collection of known environmental samples, the researcher is also referred to the section Geoarchaeology and Soil Science. Our culture obviously impacts and reflects where we live and what we consume. For those reasons, the researcher may find useful citations in the section entitled Criminal and Cultural Behavior. That said, crime scene investigators should also remember that other animal species and plant life associated with crime scenes, also reflect stable isotope signatures which may aid in reconstructing crime scene events. (2076 citations)
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Monospecific exploitation of reindeer by Neanderthals is a common behaviour in the Upper Pleistocene of Western Europe. However, reindeer-dominated assemblages have largely been reported from regions of northern Germany and south-western France, with few examples noted in south-eastern France, where faunal assemblages yield most of the time a variety of other large ungulates such as red deer, horse and diverse bovids. Here, we present multi-strand (bio- and eco-) archaeological datasets from the site of Abri du Maras (level 4.1), situated at the mouth of the Ardèche and Rhône rivers, a new example of a reindeer-dominated Neanderthal site in south-eastern France. Dated to the beginning of the MIS 3, the zooarchaeological assemblage is dominated by reindeer (88% of the NISP, representing 16 individuals) but also includes horse, bison, giant deer (Megaloceros giganteus), red deer, ibex and lagomorphs. The combination of zooarchaeological, cementochronological and tooth microwear analyses evidence a single species-dominated spectrum, with catastrophic mortality and repeated autumnal deaths. This integrated approach provides an extensive picture of human subsistence behaviour, pointing to short-term hunting episodes of reindeer herds in an exceptional context of a quasi-exclusive Neanderthal accumulation. The high number of individuals and selective butchery may correspond with a cooperative and planned mass hunting strategy. The multidisciplinary approach undertaken here also incorporating paleontological, charcoal, ecological and isotopic analyses places the archaeological and zooarchaeological data within a broader regional palaeoenvironmental framework, providing valuable landscape-contextual information. The zooarchaeological data suggest a subsistence behaviour different from other Neanderthal reindeer-dominated assemblages often connected with specialised butchery or hunting sites.
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Starting from Ernst Haeckel’s famous definition of ecology, our review considers the premises and the meaning of paleoecological research. Unlike current ecology, paleoecology has to pay more attention when dealing with “real” reconstructions. The concept of uniformitarianism is presented and demonstrates the importance of philosophical constructs in scientific work. Middle-range theory attempts to filter out false conclusions. Abiotic factors have had a strong influence on adaptive evolution; volcanism, tectonics, and climate exemplify this. Subsequently, we address biotic aspects of fossil findings, and in this context we discuss taphonomy, stratigraphic research, and interactions between floral and faunal environment. In a synthesis, we present three cross sections of human evolution at different time horizons (early-middle-late) to exemplify the inevitable multidisciplinarity of paleoecology, and we present some key events that probably altered the direction of radiations among hominids. Obviously, human evolution is not a special kind of evolution; it follows the rules of evolutionary biology, and hence depends undoubtedly on environmental influences.
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Proteins have long been known to persist in Quaternary bone fossils and are often targeted as a source of carbon used in radiocarbon dat-ing and stable isotope analyses for determin-ing provenance and obtaining dietary informa-tion. We have previously reported a technique using the dominant structural protein collagen (type I) as a source of genetic information for species identification in modern and relatively young (Holocene) archaeological samples. We report a systematic investigation of amino acid composition and collagen peptide mass finger-prints (PMF), for a range of samples dating back approximately 1.5 million years. Extrapolation from high temperature experi-mental decomposition rates predict that at a constant 10˚C (the approximate mean annual air temperature in Britain today) it will take between 0.2 and 0.7 Ma for levels of collagen to fall to 1% of their original concentration in an optimal burial environment. Even when the glacial intervals of the British Quaternary are factored into the temperature calculations, the more conservative of these two estimates extends the range for collagen sequencing to the Lower Pleistocene as confirmed by the presence of collagen peptides in bones from the Weybourne Crag (~1.5 Ma). Collagen fin-gerprinting can extend the range of identifi-able taxa present at sites with large assem-blages of fragmentary bone material such as that encountered at the ~900 Ka site at Happisburgh (Norfolk, UK) recently identified as showing signs of the earliest humans in Britain.
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The diagenetic evolution of bones of different mammal species from two French Neolithic sites (around 4,000 BC), Louviers (Eure) and Bercy (Paris), has been studied using different approaches. Global carbon and nitrogen content of bone powders were a good proxy for collagen content; δ13C values of whole bone carbon have been used to detect the presence of humic contaminants; thin sections were investigated to determine the state of preservation of histological structures and to detect staining and microbial alterations. Collagen has been extracted and its yield and C/N ratio have been measured. Both sites are located on the edge of palaeochannels, and bones were recovered from different burial environments. Comparison of the diagenetic alteration in these different depositional environments demonstrates that different mechanisms are involved and lead to very different states of preservation of the histological structures and of the collagen. The results of this study suggest that such sites may be a useful model to understand the early diagenetic alteration stages of much older palaeontological bones deposited in fluvial environments.
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The International Atomic Energy Agency (IAEA), in cooperation with the World Meteorological Organization (WMO), has been conducting a world-wide survey of hydrogen (2H/1H) and oxygen (18O/16O) isotope composition of monthly precipitation since 1961. At present, 72 IAEA/WMO network stations are in operation. Another 82 stations belonging to national organizations continue to send their results to the IAEA for publication. The paper focuses on basic features of spatial and temporal distribution of deuterium and 18O in global precipitation, as derived from the IAEA/WMO isotope database. The internal structure and basic characteristics of this database are discussed in some detail. The existing phenomenological relationships between observed stable isotope composition of precipitation and various climate-related parameters such as local surface air temperature and amount of precipitation are reviewed and critically assessed. Attempts are presented towards revealing interannual fluctuations in the accumulated isotope records and relating them to changes of precipitation amount and the surface air temperature over the past 30 years.
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An isotopic investigation of interglacial Upper Pleistocene mammal bones from layer 4 in Scladina Cave (Sclayn, Belgium) demonstrated good-quality collagen preservation. The extraction protocol had to be modified relative to the usual technique, but the collagen obtained meets the quality criteria requirements for isotopic indigeneity. Carbon isotopic abundances show that the herbivores analysed lived in a forested environment, but the carnivores also consumed prey from more open landscapes. Neanderthal isotopic carbon and nitrogen isotopic compositions suggest that most of his dietary proteins were supplied by herbivore prey from open environments. 1999 Academic Press Une étude isotopique d'os de mammifères du Pléistocène supérieur interglaciaire de la couche 4 de la grotte Scladina (Sclayn, Belgique) a démontré la bonne qualité de conservation du collagène. Le protocole d'extraction a dû être modifié par rapport à la technique usuelle mais le collagène obtenu est conforme aux critères d'indigénéité isotopique. Les abondances isotopiques du carbone montrent que les herbivores analysés vivaient dans un environnement forestier, tandis que les carnivores analysés consommaient aussi des proies de milieux plus ouverts. Les compositions isotopiques en carbone et azote du néandertalien suggèrent que l'essentiel de ses protéines alimentaires étaient fournies par de la viande d'herbivores de milieu ouvert.
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THE isotopic composition of carbon and nitrogen as well as of strontium in animal bone is related to the environment in which the animal lived1-6. It can be assumed that this is also the case for lead isotopes. In theory, therefore, we have a way of pinpointing the origin of elephant ivory, which may be of value in conservation. Here we report that by analysing the isotope ratios of these elements, a clear distinction between several different populations of the African elephant can be made.
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The carbon, nitrogen, and strontium isotope compositions of elephants in Amboseli Park, Kenya, were measured to examine changes in diet and habitat use since the 1960s. Carbon isotope ratios, which reflect the photosynthetic pathway of food plants, record a shift in diet from trees and shrubs to grass. Strontium isotope ratios, which reflect the geologic age of bedrock, document the concentration of elephants within the park. The high isotopic variability produced by behavioral and ecological shifts, if it is representative of other East African elephant populations, may complicate the use of isotopes as indicators of the source region of ivory.
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Although various stages of decomposition of bone collagen have been reported, the molecular breakdown is still not completely understood. By investigating the stable C and N isotopes of individual amino acids, this study sheds light on the biochemical processes of microbial decomposition. Collagen was extracted from sterilized compact bone of modern martens inoculated with a selection of soil bacteria in order to induce decomposition. Furthermore, this study determined the amino acid composition, overall stable carbon and nitrogen isotope ratios and isotope ratios of the separated individual amino acids of the collagen. For the bulk collagen, the carbon δ-values of the carboxyl groups were measured. This study demonstrates that the δ15N-values of the total collagen were increased, while bulk δ13C-values were decreased. With respect to the individual amino acids, the δ15N-values and especially the δ13C-values were decreased. This general isotopic shift of the individual amino acids is not related to the modification of their amounts given by the amino acid profiles of experimentally modified collagen. As expected, the carbon isotope ratios of the carboxyl group surmount the overall value. The observed modifications of the amino acid spectra seem to be the result of a selective breakdown of larger amino acids by soil microorganisms. A probable cause could be the stereo-specific repolymerization of peptides which can create insoluble macromolecules during diagenesis.
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The oxygen isotope composition of tooth enamel phosphate (δP) from cheek teeth in jaws of modern equids is compared with that of fossil equids from Thomson and Burge quarries (Miocene, Nebraska) to determine if paleobiologic and taphonomic signatures are preserved in the δP of fossil teeth. Three distinct patterns of δP variation are found in modern and fossil jaws. Each pattern can be related to the season of birth. An oxygen isotope mass balance model that incorporates seasonality and nursing of foals during tooth enamel mineralization is used to interpret the δP pattern along the toothrow. The range of δP for the same tooth position among "Merychippus" primus jaws from Thomson Quarry is relatively small (2.2‰), and the cheek tooth δP pattern is similar among individual jaws. This is comparable to a modern living population and consistent with a taphonomic interpretation of catastrophic accumulation for the Thomson "M." primus population. The range of δP for the same tooth position among Pseudhipparion retrusum jaws from Burge Quarry is larger (4‰), and the cheek tooth δP patterns are not similar among individual jaws. This is consistent with an attritional accumulation over a long period of time. Such wide ranges in δP limit application to continental paleoclimate reconstruction because of the low signal to noise ratio.
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A summary is given of the geological, faunal and archaeological information obtained during excavations in the Stanton Harcourt Channel Deposits from 1990 to 1995. The channel deposits underlie the ‘cold-climate’ Stanton Harcourt Gravel Member of the Summertown– Radley Terrace Formation. The Channel sediments are attributed to Oxygen Isotope Stage 7, when the Thames was undergoing down-dip migration and eroding the Weymouth Member of the Oxford Clay (Upper Jurassic), the contemporary Jurassic (Corallian) escarpment being near to Stanton Harcourt at that time. Abundant large vertebrate remains have been recovered, mainly from the base of the Channel deposits, where a cobble and boulder bed rests on thin silt or sand horizons or in scour hollows in the clay bedrock. Smaller bones occur throughout the deposits, which are mainly poorly sorted gravels, but especially at erosive horizons. Several palaeolithic artefacts have been found in the same contexts; many of the bones and some of the artefacts appear not to have been transported far. Although the artefacts cannot be linked directly with the bones, a study of them adds to our knowledge of the Middle Pleistocene human settlement of the Upper Thames Valley.It is of interest that mammoth is abundant as part of the interglacial faunal assemblage, and the significance of this is discussed. The environment clearly included substantial areas of open grassland, although there was also some forest in the vicinity. Evidence appears to be accumulating for important faunal and floral differences between particular interglacial events during the British Middle and Late Pleistocene.
Article
Regular intra-tooth variations in the δ18O value of mammalian tooth enamel phosphate (δ18Op) have been considered a potential measure of seasonal changes in continental climate variables since they were first observed. In order to investigate this possibility in more detail, analyses were made of teeth from a number of mammalian herbivores (sheep, cattle, elk, and pigs) that lived over a wide range of geographic locations, ecological settings, and climatic conditions (Iowa, Florida, Wyoming, Iceland, England, Croatia, and the Philippines). The lack of intra-tooth δ18Op variations in teeth of cattle that were given tap water to drink provides strong evidence that the underlying cause of observed intra-tooth variations is primarily a change in the isotopic composition of ingested water. In concert with this interpretation, the range of intra-tooth δ18Op values and their absolute values from each locality mirror observed differences in the range and absolute δ18O values of local precipitation (δ18Opt) and in climate variables. Thus intra-tooth δ18Op values can indeed be considered a qualitative measure of seasonal climate change in continental settings. Quantitative use of intra-tooth δ18Op values as a climate proxy is possible, but is hindered by lack of detailed information on aspects of mammalian physiology, behavior, and perhaps local hydrology that may also play a role in influencing δ18Op. This problem is exemplified by the different range in δ18Op values measured for sheep and cattle from the same locality around York, UK (3.4 vs. 2.6‰, respectively). The observed difference most likely reflects a difference in the relative amount of leaf water ingested by the two species. Future studies of well-constrained samples are required to test physiological models and to develop empirical relations that accurately relate δ18Op to δ18Opt. In addition to their use as indicators of seasonality, intra-tooth variations in δ18Op values provide valuable information for longer-term climate change and paleobiological investigations.
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Variation in the concentration of the stable isotopes of hydrogen, oxygen and carbon have been tested in continental records as tools for quantitative or semi-quantitative paleoclimatological studies. Among the different methods, the potential use of oxygen isotopes in mammal bone and tooth phosphate has recently been recognized. Measurements carried out on fossil mammal bones of Holocene age and their paleoclimatological interpretation corresponded well with paleontological and paleobotanical records. In the case of considerably older fossils, diagenetic processes may change the primary oxygen isotopic composition of phosphate. Fossil horse bones and teeth (principally of the species Equus stenonis) were studied to ascertain how far back in time fossil mammal remains may be considered reliable material for paleoclimatological studies. The samples come from 13 different locations in southeastern Spain, their age ranging from Maspinian (late Pleistocene) to Middle Villafranchian (Pliocene). Apart from the variations of the δ18O(PO43−) values which may be related to climatic changes, it is apparent that the isotopic composition of bones and teeth from the same deposit are frequently rather different from one another. The isotopic differences range from a few tenths of one % to several %. This suggests a strong influence of taphonomy over the measured isotopic values of fossils in each deposit. Time can be considered neither the only variable nor the most important one responsible for the change of the primary isotopic composition of fossils. Under these conditions it is rather difficult to establish lower or upper limits for the age of fossils to be studied for reliable paleoclimatological information since the limits are directly related to taphonomic history. This, in turn, is related to local environmental conditions and not only to the age of fossils.
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About eighty specimens from ten different species of mammals, collected from different areas under different climatic and environmental conditions, were measured for the oxygen isotopic composition of their bone and tooth phosphate. The equations relating these values to the mean oxygen isotopic composition of local meteoric water were also derived. The same equation can be used for goats, roe-bucks, and mouflons, despite the biological differences among these species. Measurements were made on about fourty different specimens of rabbit and hare from Europe, Africa, and Canada, but in this case the data obtained clearly show no direct relationship between the oxygen isotopic composition of local meteoric water and the isotopic composition of the skeletal phosphate. However, there seems to be an inverse relationship between the relative humidity of the studied areas and the delta(18)O(PO43-) of the skeletal phosphate, thus suggesting the use of fossil bones of these mammal species as recorders of palaeoenvironmental relative humidity. Finally, a new equation was derived for the isotopic scale for horses, on the basis of all the previous data and of a few newly obtained results
Article
Carbon isotopic abundances in Alaskan, Albertan and Russian mammoths indicate a diet of C3‐plants. The relatively high nitrogen isotopic abundances suggest arid conditions in Alberta and Alaska during the last ice age, and similar conditions in Russia. Nitrogen isotopic abundances are higher in mammoths relative to coeval herbivores, which may be due to differences in protein content of their diet. Oxygen isotopic abundances are similar in mammoths and in modern mammals from the same areas. The characteristic isotopic signature of mammoth ivory allows distinction from elephant or marine mammal ivory.
Article
The 13C/12C isotope ratios in animal1 and human2 bone can be used as indicators of diet, more recently it was shown that the 15N/14N ratios of animals and humans are similarly determined by the food they eat3–5. Specifically, the stable carbon isotope composition reflects the proportion of C3 and C4 plants at the base of the food chain1,2, while both 15N and 13C reveal the difference between a marine and terrestrial diet in modern as well as archaeological contexts5–7. Here we present data for human and animal bones from southern Africa which only partly conform to previously recognized patterns for 15N/14N ratios. Prehistoric human bones from a particular coastal region of South Africa show 15N/14N ratios consistent with the marine and terrestrial diets indicated by the 13C/12C ratios, but bones of both prehistoric humans and modern wild animals from a larger part of the subcontinent show variations in 15N/14N ratios which cannot be ascribed to known variations in diet. It appears that, in some environments, nitrogen isotope studies must also take into account the possible influence of the climate.
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RECENT international efforts to conserve the African elephant Loxodonta africana prompted us to seek an appropriate method for determining the area from which individual tusks were derived. Trace element analysis of ivory has indicated the potential of chemical analysis for source identification1, but recent isotopic studies of African mammals2–5 suggest another approach. Stable carbon isotope ratios (13C/12C) in elephant bone collagen clearly reflect the mixture of C3 foliage and C>4 grasses in the diet, and are directly proportional to the density of C3 browse2. Furthermore, nitrogen isotope ratios (15N/14N) in bone collagen of African mammals are related to rainfall or water stress3–5. Strontium isotopes (87Sr/86Sr) in bone or ivory can be expected to reflect local geology6,7. Here we report on a study of ivory and bone samples from different regions of Africa demonstrating the feasibility of trivariate isotopic analysis to identify the area in which an elephant lived, thus providing a potentially powerful tool for the control of illegal trading in ivory.
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Since its introduction in 19771, stable isotope analysis of bone collagen has been widely used to reconstruct aspects of prehistoric human and animal diets2–11. This method of dietary analysis is based on two well-established observations, and on an assumption that has never been tested. The first observation is that bone collagen 13C/12C and 15N/14N ratios reflect the corresponding isotope ratio of an animal's diet1–5,12. The second is that groups of foods have characteristically different 13C/12C and/or 15N/14N ratios13,14. Taken together, the two observations indicate that the isotope ratios of collagen in the bones of a living animal reflect the amounts of these groups of foods that the animal ate. Thus, it has been possible to use fresh bone collagen 13C/12C ratios to determine the relative consumption of C3 and C4 plants15–17, while 13C/12C and 15N/14N ratios have been used to distinguish between the use of marine and terrestrial foods14. The 15N/14N ratios of fresh bone collagen probably also reflect the use of leguminous and non-leguminous plants as food5, but this has not yet been demonstrated. Prehistoric consumption of these same groups of foods has been reconstructed from isotope ratios of collagen extracted from fossil bone1–11. Implicit in the application of the isotopic method to prehistoric material is the assumption that bone collagen isotope ratios have not been modified by postmortem processes. Here I present the first examination of the validity of this assumption. The results show that postmortem alteration of bone collagen isotope ratios does occur, but that it is possible to identify prehistoric bones whose collagen has not undergone such alteration.
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THE correlation between the oxygen isotope stratigraphy of the oceans and continental stratigraphic records is one of the main challenges for Quaternary research. Most continental classifications, however, are based on techniques and hypotheses that predate recent advances. Nevertheless, the deep-sea oxygen isotope global stratigraphic framework has been extended to the continents by means of some long sequences1; but these are exceptional and regional geology is unsatisfactorily classified for correlation with the oxygen isotope signal. Because successive geological events produced similar evidence (homotaxis), geochronometric dating is essential for correlation, but available methods are highly site-, regional- or sample-specific. Fortunately non-marine molluscs are ubiquitous in time and space, and here we use the time-dependent epimerization of L-isoleucine in these fossils to subdivide the Pleistocene of the British Isles, and to identify more events than recognized by the existing classification2. By calibrating the relative aminostratigraphic scale with independent dating methods we have set up a geochronology which is the basis for land–sea correlations back to oxygen isotope stage 15.
Article
The oxygen isotopic composition of phosphate (δ18Op) and structural carbonate (δ18Oc) of hydroxylapatite was determined in 31 bone and tooth samples of modern mammals from different countries. These two variables are highly correlate (r2 = 0.98) and the calculated best fit of linear regression is very similar to the equation calculated from the phosphate and carbonate palaeotemperature equations [1,2]. According to previous measurements [3–6] on fossils of different ages from different areas it seems quite improbable to find isotopically altered skeletal remains showing a good correlation between δ18Op and δ18Oc, as is the case with modern samples. It therefore seems possible, at least in some cases, to use these measurements for monitoring fossil bone and tooth diagenetic alteration. When a set of points lie on the equilibrium line or close to it, the δ18O values could be considered close to the original values. In contrast, when the points lie to the left or to the right of this line this probably means that the values are diagenetically modified, due to interaction with meteoric water or18O-enriched water, respectively.
Article
Collagen has been extracted and analysed for δ15N from modern kangaroos, macropods (Macropus eugenii, M. antilopinus, M. fuliginosus, M. rufus, M. robustus), in Western Australia and South Australia. Comparison of nitrogen-isotope ratios with annual rainfall for the respective areas produced a good negative correlation for macropods, but not for other marsupials. This may be related to metabolic adaptations in macropods to compete against water loss. Hence, the correlation between δ15N and annual rainfall may be species-specific. Nitrogen-isotope analyses on collagenic material extracted from fossil macropods (Macropus spp.) have been determined from two Late Pleistocene deposits in South Australia. Corroboration with spore-pollen and sedimentological data confirm that δ15N of collagenic material can be used as a proxy for palaeoprecipitation levels in Australia. This has great potential for understanding palaeoclimatic changes, especially drought periods, in Australia during the Pleistocene.
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In chapter 2 the isotopic fractionation of water in some simple condensation-evaporation processes are considered quantitatively on the basis of the fractionation factors given in section 1.2. The condensation temperature is an important parameter, which has got some glaciological applications. The temperature effect (the δ's decreasing with temperature) together with varying evaporation and exchange appear in the “amount effect” as high δ's in sparse rain. The relative deuterium-oxygen-18 fractionation is not quite simple. If the relative deviations from the standard water (S.M.O.W.) are called δD and δ18, the best linear approximation is δD = 8 δ18.Chapter 3 gives some qualitative considerations on non-equilibrium (fast) processes. Kinetic effects have heavy bearings upon the effective fractionation factors. Such effects have only been demonstrated clearly in evaporation processes, but may also influence condensation processes. The quantity d = δD −8 δ18 is used as an index for non-equilibrium conditions.The stable isotope data from the world wide I.A.E.A.-W.M.O. precipitation survey are discussed in chapter 4. The unweighted mean annual composition of rain at tropical island stations fits the line δD = 4.6 δ18 indicating a first stage equilibrium condensation from vapour evaporated in a non-equilibrium process. Regional characteristics appear in the weighted means.The Northern hemisphere continental stations, except African and Near East, fit the line δD = 8.0 δ18 + 10 as far as the weighted means are concerned (δD = 8.1 δ18 + 11 for the unweighted) corresponding to an equilibrium Rayleigh condensation from vapour, evaporated in a non-equilibrium process from S.M.O.W. The departure from equilibrium vapour seems even higher in the rest of the investigated part of the world.At most stations the δD and varies linearily with δ18 with a slope close to 8, only at two stations higher than 8, at several lower than 8 (mainly connected with relatively dry climates).Considerable variations in the isotopic composition of monthly precipitation occur at most stations. At low latitudes the amount effect accounts for the variations, whereas seasonal variation at high latitudes is ascribed to the temperature effect. Tokyo is an example of a mid latitude station influenced by both effects.Some possible hydrological applications are outlined in chapter 5.
Article
Data are presented for the 15N/14N ratios of 140 indigenous terrestrial plants from a wide variety of natural habitats in South Africa and Namibia. Over much of the area, from high-rainfall mountains to arid deserts, the 15N values of plants lie typically in the range -1 to +6; with no evident differences between C3 plants and C4 grasses. There is a slight correlation between 15N and aridity, but this is less marked than the correlation between the 15N values of animal bones and aridity. At coastal or saline sites, however, the mean 15N values for plants are higher than those at nearby inland or non-saline sites-e.g.: arid Namib coast (10 higher than inland Namib); wet Natal beach (5 higher than inland Natal); saline soils 500 km from coast (4 higher than non-saline soils). High values were also found at one site where there were no marked coastal or saline influences. These environmental effects on the isotopic composition of plants will extend upwards to the animals and humans they support. They therefore have important consequences for the use of nitrogen isotope data in the study of the dietary habits and trophic structures of modern and prehistoric communities.
Article
Human bone collagen stable isotope analysis of humans from coastal Mesolithic sites in Scotland, Denmark, France and Portugal indicates the importance of marine foods in the diet. We define the expected human δ13C and δ15N values of 100% marine and 100% terrestrial diets and conclude that at most sites isotope variability is due to differing proportions of these defined marine and terrestrial diets, rather than due to differences in the actual types of marine and terrestrial foods exploited. By comparing the European human values with marine faunal values, and values of marine-diet humans from North America, we propose that the marine component of human diet in the Late European Mesolithic was based mainly on marine fish, with only minor contributions from shellfish or marine mammals.
Article
We have analysed human and animal collagen samples from three geographically and temporally distinct cemeteries at the Dakhleh Oasis, Egypt. All sites display strikingly high average values of δ15N: Kellis 1 (Late Ptolemaic–Early Roman period) 18·0 per mil Kellis 2 (Romano-Christian period) 18·0 per mil, and ‘ein Tirghi (Roman period) 17·0 per mil. Rainfall at Dakhleh is essentially zero. The δ15N values for humans and animals lie on the respective quasi-linear relationship between rainfall and δ15N found by Heaton et al . (1986). Data from Dakhleh and other sites suggest that a single linear trend describes the rainfall-δ15N relationship in a wide range of sites. This correlation is believed to be due to a combination of two effects: excretion of excess15N-depleted urea in order to increase osmolality of urine (Ambrose & DeNiro, 1986 a , b) and15N-enrichment in arid-region plants, as a result of15N-enrichment in soils. Higher δ15N values in human consumers were acquired through consumption of animal-derived protein. High δ15N in desert soils may be caused by volatilization of isotopically light ammonia formed during bacterial activity, an effect which increases near to the soil surface.
Article
Phosphate δ18O values (δ18Op) of modern elephant bone and teeth are found to vary linearly according to the δ18O of local meteoric water (δ18Omw), a parameter with close ties to regional and local climatic conditions. Enamel, dentine, cementum and bone separates from individual fossil elephant specimens, of Late Pleistocene age, have δOp values which vary by up to elephant specimens. The larger spread in the δ18Op values of the various skeletal phases for the fossil samples is interpreted as evidence for post-depositional alterantion of primary δ18Op signatures of some, if not all, of these fossil skeletal components.In the fossil samples investigated, enamel and dentine phases have systematically lower δ18O values than associated bone and cementum phases. Differential re-equilibration with soil waters, either by the wholesale replacement of primary apatite or by processes of isotopic exchange, is suggested as a mechanism to account for the observed spread in the fossil phosphate δ18O data.
Article
An isotopic investigation of upper Pleistocene mammal bones and teeth from Scladina cave (Sclayn, Belgium) demonstrated the very good quality of collagen preservation. A preliminary screening of the samples used the amount of nitrogen in whole bone and dentine in order to estimate the preserved amount of collagen before starting the extraction process. The isotopic abundances of fossil specimens from still-extant species are consistent with their trophic position. Moreover, the15N isotopic abundance is higher in dentine than in bone in bears and hyenas, a phenomenon already observed in modern specimens. These results demonstrate that the isotopic compositions of samples from Scladina cave can be interpreted in ecological terms. Mammoths exhibit a high15N isotopic abundance relative to other herbivores, as was the case in Siberian and Alaskan samples. These results suggest distinctive dietary adaptations in herbivores living in the mammoth steppe. Cave bears are clearly isotopically different from coeval brown bears, suggesting an ecological separation between species, with a pure vegetarian diet for cave bear and an omnivorous diet for brown bear.
Article
In order to establish baseline nitrogen isotope data for certain African ecosystems, we have measured the of some 300 marine and terrestrial organisms. The majority of these specimens come from the southwestern Cape, and were chosen to represent a cross-section of the foods important in prehistoric diets in the region. δ15N analyses of 78 Holocene human skeletons from the same area are interpreted in the light of these results. Additional terrestrial animal samples were collected from the northern and eastern Cape and from Botswana and Malawi. They represent a wide range of climatic and environmental zones, from semi-desert to sub-tropical swamps. The patterning in the values for marine organisms is consistent with previously published data; that for terrestrial organisms, however, is more complex than recent studies have indicated. Our data confirm the proposal that animal δ15N values vary with rainfall: high δ15N values for herbivores occur in areas receiving less than 400 mm of rain per annum. We critically examine a recently proposed model explaining this phenomenon, and suggest some additional mechanisms which should be considered. In such arid areas, nitrogen isotope ratios cannot be used as indicators, but may provide some indication of the trophic level of the food consumed. Dietary studies on human populations can only be undertaken with a thorough appreciation of the isotopic ecology of the relevant foodweb.
Article
Theoretical predictions and measured isotope variations indicate that diet and physiological adaptation have a significant impact on animals δ18O and cannot be ignored. A generalized model is therefore developed for the prediction of animal body water and phosphate δ18O to incorporate these factors quantitatively. Application of the model reproduces most published compositions and compositional trends for mammals and birds. A moderate dependence of animal δ18O on humidity is predicted for drought-tolerant animals, and the correlation between humidity and North American deer bone composition as corrected for local meteoric water is predicted within the scatter of the data. In contrast to an observed strong correlation between kangaroo δ18O and humidity (), the predicted humidity dependence is only 1.3 – 1.7‰/10% r.h., and it is inferred that drinking water in hot dry areas of Australia is enriched in 18O over rainwater. Differences in physiology and water turnover readily explain the observed differences in δ18O for several herbivore genera in East Africa, excepting antelopes. Antelope models are more sensitive to biological fractionations, and adjustments to the flux of transcutaneous water vapor within experimentally measured ranges allows their δ18O values to be matched. Models of the seasonal changes of forage composition for two regions with dissimilar climates show that significant seasonal variations in animal isotope composition are expected, and that animals with different physiologies and diets track climate differently. Analysis of different genera with disparate sensitivities to surface water and humidity will allow the most accurate quantification of past climate changes.
Article
A suite of skeletons and frozen carcasses of upper Pleistocene mammals from Yakutia (Sakha Republic, Russia) has been analyzed for their stable isotopic abundances in carbon and nitrogen. Results from bone collagen and soft tissues have been compared. The samples studied belong to herbivorous (mammoths, woolly rhinoceros, horse, bison, muskox) and carnivorous (wolf, lion) species. Bone collagen of herbivorous and carnivorous modern mammals from the same region have been analyzed for comparison.The bone samples exhibit a very good preservation of collagen. The isotopic enrichment between herbivorous and carnivorous species is similar in modern and Pleistocene specimens, except for the mammoths, which show more negative δ13C values and higher δ15N values of collagen relative to other herbivorous species. A similar isotopic pattern can be seen in upper Pleistocene mammoths from Alaska, and this pattern suggests a paleobiological significance.
Article
Oxygen isotopic, elemental, and X-ray data are presented for a suite of 24 fossil horse teeth from Nebraska ranging in age from 18.2 to 8.5 Ma, to test the use of δ18O of enamel phosphate (δ18OP04) as a quantitative record of continental climate. Modern equid teeth were analyzed to estimate a relationship between δ8OP04 and environment water. Multiple samples of seven different fossil species from Burge Quarry, a ∼ 12 Ma attritional fossil deposit, indicate the diagenetic overprints exist but can be detected by decreased P concentration and increased crystallinity relative to modern enamel. Isotopic variation for the pristine samples from Burge Quarry is ±1.5% (1σ, n=9), which may represent the resolution of the procedure within a stratigraphic horizon. There are no apparent correlations with body size, hypsodonty, or phylogeny. A range of 7%0 in δ18OP04 occurs over the 10 m.y. interval. A trend towards depleted δ18OP04 of about 4% corresponds to a depletion of up to 6%0 in δ18O of precipitation between 18.2 and 8.5 Ma, but the range of variation of Burge is large relative to the climate signal. Our results demonstrate that δ18OP04 should be useful in quantitatively reconstructing Cenozoic continental paleoclimate on 106-year timescales. Isotopic variation due to taphonomic bias and the terrestrial rock record will likely obscure higher-order climate signals.
Article
Correlations between mean annual temperature (MAT) and the weighted average oxygen isotope composition of yearly precipitation (δ18Opt) are well-known, but the utility of modern relations to make reliable estimates of temperature change over geological time is uncertain. This question has been addressed by using seasonal subsets of the global data base of temperature and isotopic measurements to represent two different climate modes. A comparison of middle- to high-latitude δ18Opt/temperature relations for each climate mode reveals (1) a significant offset between them, and (2) a difference in the strength of their correlations. The offset in relations is due to differences in temperature and water vapor budget in the tropics, and can lead to serious underestimates of temperature change. Differences in the strength of correlations arise from the influence of climate mode-specific, non-temperature factors on δ18Opt. The overall result is that no single relation can be used in all cases to make unambiguous temperature estimates using a temporal record of δ18Opt values. One way to overcome these problems is to reconstruct δ18Opt/temperature relations for the time periods being investigated. If an appropriate proxy for δ18Opt is available, it may also be possible to estimate temperature without relying on δ18Opt/temperature relations. A promising alternative to these options is to use records of δ18Opt to test predictions of global climate models, an approach that may allow a reliable and more complete reconstruction to be made of climate change over geologic time.
Article
Thesis (Ph. D.)--University of London, 1998.
Article
THE cause of extinction of the woolly mammoth, Mammuthus primigenius (Blumenbach), is still debated. A major environmental change at the Pleistocene-Holocene boundary, hunting by early man, or both together are among the main explanations that have been suggested. But hardly anyone has doubted that mammoths had become extinct everywhere by around 9,500 years before present (BP). We report here new discoveries on Wrangel Island in the Arctic Ocean that force this view to be revised. Along with normal-sized mammoth fossils dating to the end of the Pleistocene, numerous teeth of dwarf mammoth dated 7,000-4,000 yr BP have been found there. The island is thought to have become separated from the mainland by 12,000 yr BP. Survival of a mammoth population may be explained by local topography and climatic features, which permitted relictual preservation of communities of steppe plants. We interpret the dwarfing of the Wrangel mammoths as a result of the insularity effect, combined with a response to the general trend towards unfavourable environment in the Holocene.
Article
Methods for the quantitative derivatization of amino acids with phenylisothiocyanate and for the separation and quantitation of the resulting phenylthiocarbamyl derivatives by reverse-phase high-performance liquid chromatography are described. Phenylthiocarbamylation of amino acids proceeds smoothly in 5 to 10 min at room temperature. Coupling solvents, reagent, and some byproducts are removed by rotary evaporation under high vacuum, and the phenylthiocarbamyl derivatives are dissolved in 0.05 M ammonium acetate, pH 6.8, for injection onto the octyl or octadecylsilyl reverse-phase column. Columns are equilibrated with the same solvent and the effluent stream is monitored continuously at 254 nm for detection of the amino acid derivatives. Elution of all of the phenylthiocarbamyl amino acids is achieved in about 30 min utilizing gradients of increasing concentrations of ammonium acetate and acetonitrile or methanol. This approach to amino acid analysis offers select advantages, both with respect to methods which employ reverse-phase separation of prederivatized samples and to the classical ion-exchange procedure. All amino acids, including proline, are converted quantitatively to phenylthiocarbamyl compounds and these are stable enough to eliminate any need for in-line derivatization. Furthermore, results comparable in sensitivity and precision to those obtained by state-of-the-art ion-exchange analyzers may be generated with equipment that need not be dedicated to a single application.
Tracing the diets of fossil animals using stable isotopes
  • P L Koch
  • M L Fogel
  • N Tuross
P.L. Koch, M.L. Fogel, N. Tuross, Tracing the diets of fossil animals using stable isotopes, in: K. Lajtha, R.H. Michener (Eds), Stable Isotopes in Ecology and Environ-mental Science, Blackwell Scienti¢c Publications, 1994.
Lister, Palaeontology; mammoths in miniature
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Evolution and taxonomy of Eurasian mam-moths The Proboscidea: Trends, Evolution and Palaeoecology
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A.M. Lister, Evolution and taxonomy of Eurasian mam-moths, in: J. Shoshani, P. Tassy (Eds.), The Proboscidea: Trends, Evolution and Palaeoecology, Oxford University Press, 1996, pp. 203^218.
Quaternary £uvial deposits and palaeontology at Stanton Harcourt, Oxfordshire The Quaternary of the South Midlands and the Welsh Marches
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K. Scott, C. Buckingham, Quaternary £uvial deposits and palaeontology at Stanton Harcourt, Oxfordshire, in: S.G. Lewis, D. Maddy (Eds.), The Quaternary of the South Midlands and the Welsh Marches, Quaternary Research Association, 1997.
Young, Modeling N 18 O in proboscidean phosphate
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A.M. Jones, P.L. Koch, E.D. Young, Modeling N 18 O in proboscidean phosphate, in preparation.
A preliminary report on the Stanton Harcourt Channel Deposits (Oxfordshire, England); geological context, vertebrate remains and Palaeolithic stone artefacts
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Isotopic patterns in modern global precipitation
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Modeling δ18O in proboscidean phosphate
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Quaternary fluvial deposits and palaeontology at Stanton Harcourt
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Palaeontology; mammoths in miniature
  • Lister