Geometric morphometric assessment of the fossil bears of Namur,Belgium: Allometry and ecomorphology

The Namur area in Belgium is useful to study brown (Ursus arctos) and cave bears (Ursus spelaeus) as the assemblage contains little temporal and no geographical variation. Here, we aim to assess ontogenetic allometry within cave bears, as well as ecomorphological differences between adult brown bears (n = 9), adult cave bears (n = 5) and juvenile cave bears (n = 3). Landmarks for 3D digitization of the mandible were chosen based on the taphonomical damage of the specimens. Extant brown bears and extinct Pleistocene brown and cave bears were digitized with a Microscribe G2. Generalized Procrustes superimposition was performed on the coordinates. Allometry was studied using regression analysis. Principal component analysis (PCA) was conducted to assess ecomorphological differences between the groups. 61% of the shape variance within juvenile and adult cave bears was predicted by size (n = 8, p < 0.01). The juvenile cave bears have relatively deep horizontal rami. In adult cave bears, the horizontal ramus is much narrower dorsoventrally. Juvenile cave bears have a small masseteric fossa and a short coronoid process, whereas both are larger, relative to mandible size, in adult cave bears. This made juvenile cave bears likely less effective masticators than fully grown cave bears. In the PCA, principal component (PC) 1 accounts for 45.0% of the total variance and PC2 accounts for 27.6%. Fossil U. arctos from Namur fall within the 95% confidence interval of modern North American U. arctos on both PCs, but are more similar to cave bears than the average extant brown bear. From the similarity of fossil and modern brown bears, it can be deduced that the diet of fossil brown bears was probably also within the range of their modern North American conspecifics, although they might have been more efficient at masticating plant matter.     

Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus)

The cave bear ( Ursus spelaeus ) was one of several spectacular megafaunal species that became extinct in northern Eurasia during the late Quaternary. Vast numbers of their remains have been recovered from many cave sites, almost certainly representing animals that died during winter hibernation. On the evidence of skull anatomy and low δ¹⁵N values of bone collagen, cave bears appear to have been predominantly vegetarian. The diet probably included substantial high quality herbaceous vegetation. In order to address the reasons for the extinction of the cave bear, we have constructed a chronology using only radiocarbon dates produced directly on cave bear material. The date list is largely drawn from the literature, and as far as possible the dates have been audited (screened) for reliability. We also present new dates from our own research, including results from the Urals. U. spelaeus probably disappeared from the Alps and adjacent areas – currently the only region for which there is fairly good evidence – c . 24 000 radiocarbon years BP ( c . 27 800 cal. yr BP), approximately coincident with the start of Greenland Stadial 3 ( c . 27 500 cal. yr BP). Climatic cooling and inferred decreased vegetational productivity were probably responsible for its disappearance from this region. We are investigating the possibility that cave bear survived significantly later elsewhere, for example in southern or eastern Europe.     

Morphometry of upper cheek teeth of cave bears (Carnivora,Ursidae)

Univariate and multivariate statistics were applied to analyse the morphometrical variability of 4920 upper cheek teeth (P4, M1 and M2) of cave bears from 123 geographical sites (180 samples) of different Pliocene – Pleistocene ages. The analysed specimens included those belonging to the big cave bears Ursus kudarensis, U. deningeri, U. spelaeus (three subspecies) and U. kanivetz (including U. ingressus), as well as the small cave bear U. rossicus. The information‐theoretical parameters (Shannon entropy and orderliness (Von Foerster, 1960: On self‐organizing systems and their environments. In Self‐Organizing Systems, 31–50. Pergamon Press, London) were used to estimate tooth diversity in different teeth, different taxa and in selected local chrono‐populations. Multivariate allometry coefficients (Klingenberg, 1996: Multivariate allometry. In Advances in Morphometrics, 23‐49. Plenum Press, New York) were used to describe the relationships of different ‘parts’ of a tooth and to compare allometric patterns amongst species or selected local samples. A multivariate analysis showed a significant overlap of the size/shape parameter ranges in deningeroid and spelaeoid bears within morphological spaces. Within the cave bear lineage, the Deninger's bear has the greatest morphological diversity index (entropy) of all the teeth overall, and the lowest diversity is observed in the final taxon of this lineage – U. kanivetz (=ingressus). The P4 and M2 diversity showed multidirectional correlations with elevation above sea level amongst several ‘local’ populations of Late Pleistocene cave bears. The morphological disparities between the studied taxa are in close agreement with the distances in the available schemes of genetic differentiation based on ancient mitochondrial DNA. The split of U. kudarensis and U. deningeri has a good bootstrap support, which corresponds to the hypothesis about their parallel evolution. The small cave bear U. rossicus is placed between U. arctos and U. deningeri. The phylogenetic signal is more pronounced in the variability of teeth in comparison with other skeletal remains of cave bears (cranium, mandible, or metapodial bones).     

The latest Ursus spelaeus in Italy,a new contribution to the extinction chronology of the cave bear

The skeleton of a young prime adult cave bear, Ursus spelaeus, was found in Chiostraccio Cave (Siena, Tuscany, central Italy), only slightly buried under rock falls. The specimen was dated yielding a conventional age of 24,030 ± 100 ¹⁴C yr BP (29,200–28,550 cal yr BP), which makes it the latest known representative of the species in Italy. The skeleton was accompanied by the remains of wolf (Canis lupus), wild boar (Sus scrofa), aurochs (Bos primigenius), red deer (Cervus elaphus), roe deer (Capreolus capreolus), bat (Vespertinus murinus), and crow (Corvus monedula). The site seems confirming that the latest Italian U. spelaeus populations shared the risk of intrusion. The association of the cave bear with other animals suggests that the assemblage is an attritional palimpsest of remains of different species not originally associated in life. Cave bears were probably more vegetarian than brown bears and possibly became extinct when plant productivity dropped at the onset of MIS 2. Central and southern Italy may have offered isolated and sheltered refugia for cave bears.     

Late-Quaternary biogeographic scenarios for the brown bear (Ursus arctos), a wild mammal model species

This review provides an up-to-date synthesis of the matrilineal phylogeography of a uniquely well-studied Holarctic mammal, the brown bear. We extend current knowledge by presenting a DNA sequence derived from one of the earliest known fossils of a polar bear (dated to 115 000 years before present), a species that shares a paraphyletic mitochondrial association with brown bears. A molecular clock analysis of 140 mitochondrial DNA sequences, including our new polar bear sequence, provides novel insights into the times of origin for different brown bear clades. We propose a number of regional biogeographic scenarios based on genetic data, divergence time estimates and paleontological records. The case of the brown bear provides an example for researchers working with less well-studied taxa: it shows clearly that phylogeographic models based on patterns of modern genetic variation alone can be substantially improved by including data on historical patterns of genetic diversity in the form of ancient DNA sequences derived from accurately dated samples and by using an approach to divergence-time estimation that suits the data under analysis. Using such approaches it has been possible to (i) establish that the processes shaping modern genetic diversity in brown bears acted recently, within the last three glacial cycles; (ii) distinguish among hypotheses concerning species’ responses to climatic oscillations in accordance with the lack of phylogeographic structure that existed in brown bears prior to the last glacial maximum (LGM); (iii) reassess theories linking monophyletic brown bear populations to particular LGM refuge areas; and (iv) identify vicariance events and track analogous patterns of migration by brown bears out of Eurasia to North America and Japan.     

山东滕州西公桥遗址人骨的稳定同位素分析

文章应用稳定同位素C和N分析的方法,研究了山东滕州大汶口文化时期西公桥遗址先民的食谱,试图揭示该遗址先民生活方式、社会经济结构的变化。总体而言,该遗址样品保存较差,骨中骨胶原的含量较低,以C/N摩尔比作为标准,仅有8个样品属于未污染样品,可用作食谱分析。对样品进行稳定同位素C和N分析,结果表明:该遗址先民均为稻、粟混食,但在肉食资源获取上有所不同。以C₃为主的先民,具有较低的^δ15N;而以C₄类为主的先民,^δ15N值较高。在该遗址的发展过程中,人们的食物结构发生了一定的变化。     

Cave bears and paleolithic artifacts in Yarimburgaz Cave,Turkey: Dissecting a palimpsest

This article addresses the taphonomic issues surrounding artifact-bear associations in Yarimburgaz Cave, northwestern Turkey. Our purpose is to evaluate the circumstances of bone assemblage formation in the Middle Pleistocene deposits of the cave, identifying the agencies responsible for the accumulations and the causes of bone damage before and following deposition. Pleistocene species representation, bone damage, body part representation, and mortality data indicate that cave bear remains in Yarimburgaz are unrelated to hominid use of the same site. None of the observations on the cave bear remains contradicts the expectations developed from modern wildlife accounts of modern bear behavior; these accumulations resulted from mortality normally associated with hibernation over many generations of den use. Nonursid carnivores also played roles in bone collection and/or modification. We conclude that at least three distinct biological processes contributed to the formation of the Middle Pleistocene faunas: hibernating bears, bone-collecting carnivores such as wolves, and hominids in descending order of importance. The Yarimburgaz faunas represent palimpsests or overlays of many short-term depositional events, the close spatial associations of which are explained by slow or uneven sedimentation rates inside the cave. Although occupations by hominids appear to have been ephemeral in nature, hominids ultimately discarded nearly 1700 stone artifacts in Yarimburgaz Cave. The hominids did not discard many ungulate bones in the same circumstances. These observations suggest that hominid foraging efforts focused on resources other than large game while at the cave. © 1996 John Wiley & Sons, Inc.     

Holocene brown bear (Ursus arctos L.) from Norwegian caves

Norwegian caves contain stratigraphical information regarding the historical faunal composition valuable in the assessment of postglacial colonization and phylogeographical structure. In some of these limestone caves, brown bear ( Ursus arctos L.) remains have been excavated and radiometrically dated. We present osteology (95 traits defined) from 29 brown bears found within 22 Norwegian caves above 658N. Our data span the time interval 6210–420 ¹⁴C yr BP. The sex of individuals was based on dimorphic canines, while age was determined by cementum analysis and body size estimated from regressions between morphological traits compared to a contemporary reference collection. Five females and 14 males were recognized, while the remaining bears could not be sexed because of their small size, low age or lack of canines. The ages of 26 bears (tooth cementum age and estimates) ranged from juveniles (<0.5 year) to adults (23 years), the majority being old animals. Both sexes have used these caves for over-wintering lairs, as seen in several caves, where denning is also suggested based on the observations of adult females and cubs. There were no signs of predator or human transported bear remains, suggesting natural mortality. The age of the bears suggests that young or old bears may have died from insufficient nutrient storage, environmental harshness, or may have drowned in spring or autumnal flooding. We conclude that Norwegian brown bears have used these caves sporadically for thousands of years, which suggests that caves are not a preferred site for the brown bear during hibernation or denning.     

Influence of diet on the distribution of nitrogen isotopes in animals

The influence of diet on the distribution of nitrogen isotopes in animals was investigated by analyzing animals grown in the laboratory on diets of constant nitrogen isotopic composition.The isotopic composition of the nitrogen in an animal reflects the nitrogen isotopic composition of its diet. The δ₁₅N values of the whole bodies of animals are usually more positive than those of their diets. Different individuals of a species raised on the same diet can have significantly different δ¹⁵N values. The variability of the relationship between the δ¹⁵N values of animals and their diets is greater for different species raised on the same diet than for the same species raised on different diets. Different tissues of mice are also enriched in ¹⁵N relative to the diet, with the difference between the δ¹⁵N values of a tissue and the diet depending on both the kind of tissue and the diet involved. The δ¹⁵N values ofcollagen and chitin. biochemical components that are often preserved in fossil animal remains, are also related to the δ¹⁵N value of the diet.The dependence of the δ¹⁵N values of whole animals and their tissues and biochemical components on the δ¹⁵N value of diet indicates that the isotopic composition of animal nitrogen can be used to obtain information about an animal's diet if its potential food sources had different δ¹⁵N values. The nitrogen isotopic method of dietary analysis probably can be used to estimate the relative use of legumes vs non-legumes or of aquatic vs terrestrial organisms as food sources for extant and fossil animals. However, the method probably will not be applicable in those modern ecosystems in which the use of chemical fertilizers has influenced the distribution of nitrogen isotopes in food sources.The isotopic method of dietary analysis was used to reconstruct changes in the diet of the human population that occupied the Tehuacan Valley of Mexico over a 7000 yr span. Variations in the δ¹⁵C and δ¹⁵N values of bone collagen suggest that C₄ and/or CAM plants (presumably mostly corn) and legumes (presumably mostly beans) were introduced into the diet much earlier than suggested by conventional archaeological analysis.     

Sources of carbon isotope variation in kangaroo bone collagen and tooth enamel

The stable carbon isotopic composition (expressed as δ¹³C) of herbivore remains is commonly used to reconstruct past changes in the relative abundance of C₄ versus C₃ grass biomass (C₄ relative abundance). However, the strength of the relationship between herbivore δ¹³C and C₄ relative abundance in extant ecosystems has not been thoroughly examined. We determined sources of variation in δ¹³C of bone collagen and tooth enamel of kangaroos (Macropus spp.) collected throughout Australia by measuring δ¹³C of bone collagen (779 individuals) and tooth enamel (694 individuals). An index of seasonal water availability, i.e. the distribution of rainfall in the C₄ versus C₃ growing seasons, was used as a proxy for C₄ relative abundance, and this variable explained a large proportion of the variation in both collagen δ¹³C (68%) and enamel δ¹³C (68%). These figures increased to 78% and 77%, respectively, when differences between kangaroo species were accounted for. Vegetation characteristics, such as woodiness and the presence of an open forest canopy, had no effect on collagen or enamel δ¹³C. While there was no relationship between collagen δ¹³C and kangaroo age at death, tooth enamel produced later in life, following weaning, was enriched in ¹³C by 3.5‰ relative to enamel produced prior to weaning. From the observed relationships between seasonal water availability and collagen and enamel δ¹³C, enrichment factors (ε^∗) for collagen-diet and enamel-diet (post-weaning) were estimated to be 5.2‰ ± 0.5 (95% CI) and 11.7‰ ± 0.6 (95% CI), respectively. The findings of this study confirm that at a continental scale, collagen and enamel δ¹³C of a group of large herbivores closely reflect C₄ relative abundance. This validates a fundamental assumption underpinning the use of isotopic analysis of herbivore remains to reconstruct changes in C₄ relative abundance.     

The first record of “spelaeoid” bears in Arctic Siberia

For a long time, “spelaeoid” (cave-bear-like) bears, Ursus (Spelearctos) spp., were believed to be almost purely European animals. Their geographic range has recently been extended to the east, in southern Siberia, Transbaikalia, Kirghizia, Mongolia and Korea. Two unexpected new findings, presented here in detail, significantly change existing views on the distribution of cave bears; both were found in North-Eastern Siberia, far beyond the Arctic Circle, more than 1500 km to the north-east of the previously accepted range.One of the fossils is a mandible, found near the town of Cherskiy at 68.73°N, 161.38°E. The analysis of local geology and accompanying mammal fossils suggests that it comes from the Olyorian Fauna (Early to early Middle Pleistocene). Morphologically, the Cherskiy mandible is closest to Ursus savini, a small middle Pleistocene cave bear from the British Cromer Forest-bed Formation, but differs in having a slightly more advanced dentition, and thus it is described as a new subspecies Ursus savini nordostensis. Another newly recognized fossil of the “spelaeoid” bear is an astragalus found at the Oskhordokh site at 67.54°N, 135.67°E, on a large gravel bar on the right bank of the Adycha River. This specimen is attributed to Ursus cf. deningeri.The paper also presents an interesting example of the interaction between classical and “molecular” palaeontology.The new finds significantly change existing ideas on the ecology and evolution of cave bears, some of the most remarkable members of the extinct Pleistocene megafauna.     

A Late Weichselian find of polar bear (Ursus maritimus Phipps) from Denmark and reflections on the paleoenvironment

The mandible of a polar bear (Ursus maritimus Phipps) found in about 1920 at Kjul Å, North Jutland, and described by Nordmann & Degerbol in 1930. has been ^l4C dated to 11.100 ± 160 B.P. It is so far the only find of polar bear in Denmark. Comparison with recent ¹⁴C datings of Swedish and Norwegian polar bears shows that the Danish specimen was a member of a southern Scandinavian Late Weichselian population. The contemporaneous Zirphaeu sea deposits can be regarded as the boreal-arctic shallow water equivalent of the arctic Upper Saxicava sand deposits from northern Jutland. The polar bear mandible, however, was deposited on land, as was the metacarpal bone of a brown bear ( Ursus arctos ) from the nearby Nr. Lyngby locality of Allerød age. The overall picture of the Late Weichselian mammal fauna in Denmark shows a mixed composition of different ecotypes. Their sympatric occurrence points at a unique environment not comparable to any now existing, and probably related to the very low latitude of the Weichselian ice sheet.     

Patterns of faunal extinction and paleoclimatic change from mid-Holocene mammoth and polar bear remains, Pribilof Islands, Alaska

Qagnaxˆ Cave, a lava tube cave on St. Paul Island in the Pribilofs, has recently produced a mid-Holocene vertebrate faunal assemblage including woolly mammoth, polar bear, caribou, and Arctic fox. Several dates on the mammoth remains converge on 5700 ¹⁴C yr BP. These dates, ~ 2300 yr younger than mammoth dates previously published from the Pribilof Islands, make these the youngest remains of proboscideans, and of non-extinct Quaternary megafauna, recovered from North America. Persistence of mammoths on the Pribilofs is most parsimoniously explained by the isolation of the Pribilofs and the lack of human presence in pre-Russian contact times, but an additional factor may have been the local existence of high-quality forage in the form of grasses enriched by nutrients derived from local Holocene tephras. This interpretation is reinforced by stable carbon and nitrogen isotope values obtained from the mammoth remains. The endpoint of mammoth survival in the Pribilofs is unknown, but maybe coterminous with the arrival of polar bears whose remains in the cave date to the Neoglacial cold period of ~ 4500 to 3500 ¹⁴C yr BP. The polar bear record corroborates a widespread cooling of the Bering Sea region at that time.     

A Pilot Study of Trophic Level and Human Origins at the Xiaoshuangqiao Site, China (ca. 1400 BC) Using δD Values of Collagen

We present here a pilot study to examine trophic level effects and migration patterns at the middle Shang Dynasty site of Xiaoshuangqiao in Henan Province using δD results combined with δ13C and δ15N values. A total of 33 specimens (17 humans, 7 cattle, 5 pigs, 3 sheep, 1 dog) of bone collagen were isotopically analyzed for δ13C, δ15N, and dD values. A strong positive correlation (R2 = 0.94) between mean δ15N and δD values of herbivores (cattle and sheep), omnivores (pig), carnivores (dog) and humans was observed. The δD results were found to increase by ~10‰ to 20‰ from herbivores to omnivores to carnivorous, evidence that collagen δD results are a useful indicator for the study of trophic levels and dietary patterns at archaeological sites. The δD results were also used to examine the origins of two different groups of individuals buried at Xiaoshuangqiao. Individuals buried in sacrificial pits of district V had mean δD values (?47.0 ± 2.9‰, n = 11) that were significantly (p = 0.049) elevated compared to the people buried in the stratigraphy of district IX (?51.3 ± 3.3‰, n = 3), indicating that they were ingesting water from different locations. In addition, the D values of the people buried in the stratigraphy were similar to the pigs (?54.5 ± 4.2‰, n = 5) at Xiaoshuangqiao, suggesting that they were most probably of the local population, and that the individuals buried in the sacrificial pits were most possibly from the coast and prisoners of the Dongyi (“东夷”) people. Thus, δD results have the potential to examine human origins and migration patterns and should be increasingly used in conjunction with δ13C and δ15N values at archaeological sites.     

Contribution of stable isotopes to the understanding of the unsaturated zone of a carbonate aquifer (Nerja Cave, southern Spain)

We analysed the stable isotopes (¹⁸O and ²H) of rainwater and drip water within a cave (Nerja Cave) located in the unsaturated zone of a carbonate aquifer. Rainfall is more abundant and presents lower isotopic content in winter, while the volume of drip water is greater and its isotopic content is lower in summer. The flow analysis of ¹⁸O through the unsaturated zone confirms the seasonal lag between rainfall and the appearance of drip water in the cave and reveals that the unsaturated zone of the aquifer, in the sector of the cave, behaves like an inertial system with a strong capability to modulate the input signal. To cite this article: F. Carrasco et al., C. R. Geoscience 338 (2006).     

The Inka mummy from Mount Aconcagua: Decoding the geographic origin of the “Messenger to the Deities” by means of stable carbon,nitrogen, and sulfur isotope analysis

Human sacrifices were a ritual practice during the expansion of the Inka Empire. Nevertheless, the homeland of the victims has never been clearly determined. A mummy recently found in a mediterranean environment in the Andes of Argentina provides a unique opportunity to shed some light on the matter. His clothing and funerary trousseau give no evidence regarding his ethnic filiation, geographic origin, nor whether he belonged to the local population or to the Inka elite. Therefore, a decodification of his geographic origin was attempted. This was done through the reconstruction of his palaeodiet: A diet based on continental products would indicate a local origin whereas one including marine food would suggest ties with the Pacific littoral and, indirectly, a foreign origin. Carbon, nitrogen, and sulfur isotope ratios of the mummy's bone collagen and hair were compared with values from archaeological populations from littoral and altiplanic environments of South America. The carbon and nitrogen isotopic values of bone collagen were ambiguous tracers, indicating either a continental diet with a high percentage of maize, one including a certain amount of marine products, or a combination of both. The carbon and sulfur isotopic values of hair were conclusive, indicating a clear continental origin of the diet, at least during the period of a year and a half before death. © 1999 John Wiley & Sons, Inc.     

A test of uranium-series dating of fossil tooth enamel: results from Tournal Cave,France

A series of well preserved mammal bones and horse teeth was analyzed from archaeological levels of Tournal Cave (Magdalenian, Aurignacian, and Mousterain) to test the hypothesis that well-crystallized enamel behaves more as a closed system than does whole bone. The isotopic composition of bones and tooth enamels from this deposit meet criteria for confidence, and gave no reasons to suspect contamination or open-system behavior. Two samples for which²³¹Pa could be analyzed showed internal concordance with the respective²³⁰Th ages. In spite of the favourable isotopic criteria, however, comparison of the U-series ages of the bones and the tooth enamel with stratigraphic position and¹⁴C control indicated the dates were not meaningful. In general, both bones and tooth enamels gave ages too young, although some were clearly too old. Neither group showed any systematic increase of age with stratigraphic depth. Tooth enamel, therefore, shows no advantage over bone for U-series dating for this site. In Tournal cave both bones and enamel are apparently open to U, which is probably cycling as a consequences of post-depositional groundwater movement.     

Cave bear occupation in Schwabenreith Cave,Austria, during the early last glacial: constraints from 230Th/U‐dated speleothems

The cave bear was a prominent member of the Upper Pleistocene fauna in Eurasia. While breakthroughs were recently achieved with respect to its phylogeny using ancient DNA techniques, it is still challenging to date cave bear fossils beyond the radiocarbon age range. Without an accurate and precise chronological framework, however, key questions regarding the palaeoecology cannot be addressed, such as the extent to which large climate swings during the last glacial affected the habitat and possibly even conditioned the final extinction of this mammal. Key to constraining the age of cave bear fossils older than the lower limit of radiocarbon dating is to date interlayered speleothems using ²³⁰Th/U. Here we report new results from one such site in the Eastern European Alps (Schwabenreith Cave), which yielded the highest density of bones of cave bear (Ursus spelaeus eremus). Although dating of the flowstones overlying this fossiliferous succession was partly compromised by diagenetic alteration, the ²³⁰Th/U dates indicate that the bear hibernated in this cave after about 113 ka and before about 109 ka. This time interval coincides with the equivalent of Greenland Stadial 25, suggesting possible climate control on the cave bear's habitat and behaviour. © 2019 The Authors. Journal of Quaternary Science Published by John Wiley & Sons Ltd     

Diagenesis and the reconstruction of paleoenvironments: A method to restore original δO values of carbonate and phosphate from fossil tooth enamel

Intra-tooth δ¹⁸O variations within the carbonate (δ¹⁸Oc) and phosphate (δ¹⁸Op) components of tooth apatite were measured for Miocene and Pliocene hypsodont mammals from Afghanistan, Greece and Chad in order to evaluate the resistance of enamel to diagenetic alteration. Application of water-apatite interaction models suggest that the different kinetic behaviours of the phosphate-water and carbonate-water systems can be used to detect subtle oxygen isotope disequilibria in fossil enamel when intra-individual variations are considered. Selective alteration of the oxygen isotope composition from the carbonate component of Afghan and Greek enamels suggests inorganic isotopic exchange processes. Microbially-induced isotopic exchange for phosphate is demonstrated for the first time in enamel samples from Chad, in association with extensive recrystallization. In Chad, δ¹⁸Op values were derived from partial isotopic exchange with fossil groundwater during early diagenesis. Mass balance calculations using average carbonate content in enamel as a proxy for recrystallization, and the lowest δ¹⁸Op value of dentine as a proxy for the isotopic composition of the diagenetic fluid, indicate that diagenesis can alter δ¹⁸Op by as much as 3‰ in some enamel samples. This diagenetic alteration is also responsible for a decrease in intra-individual variations of up to 1‰ in affected specimens. The effects of diagenesis on δ¹⁸Op values of fossil enamel are not systematic, however, and can only be estimated if sequential δ¹⁸Op and δ¹⁸Oc analyses are performed on fossil enamel and dentine. Reconstruction of large temporal- or spatial-scale paleoclimates based on δ¹⁸Op analyses from mammalian teeth cannot be considered valid if enamel has been affected by bacterial activity or if the data cannot be corrected for diagenetic effects.     

Additive effects of climate change and human hunting explain population decline and extinction in cave bears

Cave bears (Ursus spelaeus) are an iconic component of the European late Quaternary Ice Age megafauna. Recent demographic analyses based on cave bear mtDNA sequences and refined radiocarbon dating indicate that cave bear population size and genetic diversity started to decline some 50 kilo years ago (kya). Hence, neither the coldest phase of the last glaciation (started some 24 kya), nor the colonization of Europe by Palaeolithic hunters (started some 45 kya) coincides with the beginning of population decline. Here, we reconstructed cave bear climatic niche evolution through time. Then, we performed spatially explicit population viability analyses to assess cave bear demographics through time in response to climatic changes, human effects on bear survival and their combination. We found that climate change was responsible for a 10‐fold decrease in cave bear population size after 40 kya. However, climate change on its own could not explain U. spelaeus extinction at 24 kya. Additional negative effects consistent with human population expansion are required to explain both U. spelaeus' retreat from eastern Europe since 40 kya and its final extinction.