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Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus)
Boreas
Abstract
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.
... The diverse Late Pleistocene cave bear complex evolved from the Middle Pleistocene Ursus deningeri and comprises several morphologically and genetically distinct forms, separated at the species and subspecies levels (Baca et al., 2016;Barlow et al., 2018;Gretzinger et al., 2019;Rabeder et al., 2000). Different lines of evidence indicate that cave bears were herbivorous, with a high amount of high-quality herbaceous plants in their diet (Bocherens, 2019;Pacher & Stuart, 2009;Rabeder et al., 2000;Terlato et al., 2019). Cave bears hibernated in caves, and large numbers of their bones have been found in caves across Europe (Grandal-d'Anglade et al., 2019;Kurtén, 1976). ...
... Cave bear extinction occurred between about 28 to 25 kyr BP across most of its range, broadly coinciding with Greenland stadial 3, the coldest phase of the last glacial (Baca et al., 2016;Pacher & Stuart, 2009;Terlato et al., 2019). Younger dates of 24,200-23,500 cal yr BP for skeletal remains from NE-Italy indicate somewhat longer survival in refugia with more favorable climatic conditions (Terlato et al., 2019). ...
... The ultimate cause(s) of extinction are a matter of debate (Kurtén, 1976;Stuart, 2021). Some authors suggest that vanishing of the cave bear was caused by climatic deterioration and associated decreased productivity of the vegetation, with extinction being preceded by range fragmentation and formation of isolated small subpopulations (Baca et al., 2016;Pacher & Stuart, 2009). Others have concluded that, in combination with climatic deterioration, increasing human pressure was a major factor leading to cave bear extinction (Gretzinger et al., 2019;Stiller et al., 2010;Terlato et al., 2019). ...
The paper discusses the formation of an enamel defect in the crown of a cave bear ( Ursus spelaeus sensu lato) left maxillary second molar (M ² ), based on macroscopic and microscopic analysis. The tooth belongs to a cranium recovered from the Cerovac caves in Croatia that exhibits a partially healed, depressed lesion in the left squama frontalis and a further lesion in the left maxilla associated with loss of the M ¹ . Microscopic inspection demonstrated an accentuated incremental line in both enamel and dentin of the left M ² . It is suggested that in the defect area the outer enamel had been posteruptively lost along the accentuated line in the enamel that constituted a zone of reduced mechanical resistance. Presence of enamel hypoplasia in both M ² indicated that these developmental lesions reflect a systemic stress event during crown formation of the teeth. The underlying cause of this stress is assumed to have been a trauma to the skull that caused the lesion in the left squama frontalis. It is further suggested that a later trauma to the left maxilla had led to the loss of the left M ¹ and the flaking‐off of enamel along the accentuated incremental line in the left M ² . The defect in the left M ² is thus diagnosed as the result of a developmental lesion during crown formation, related to systemic stress due to a skull trauma, followed by posteruptive damage from a second traumatic impact. In addition to reconstructing the formation of the defect in the crown of the left M ² , the paper, for the first time, describes daily and subdaily incremental markings in ursid enamel and provides preliminary information on enamel secretion rate in a cave bear molar.
... Carnivores, despite their further adaptability to various biomes, do not depend directly on a specific habitat like herbivores (Hunter and Turner, 1997). They also experienced a significant population decline and displacement (Varela et al., 2010;Stuart and Lister, 2011), and in some cases extinction (Pacher and Stuart, 2009). This phenomenon has been proposed as being greatly influenced by the displacement and extinction of populations of large herbivores, which formerly constituted their primary food source (Croitor and Brugal, 2010;Stuart and Lister, 2011;Varela et al., 2010). ...
... The carnivores that prevailed and occupied the ecological niche of those extinct species were smaller-sized carnivores and omnivores with broad dietary habits (Sommer and Benecke, 2005;Croitor and Brugal, 2010) such as brown bears (Ursus arctos), which during the end of the Pleistocene decreased in size as a response to the climatic and environmental instability (Marciszak et al., 2015). An ecologically singular member of the carnivore order that was among the first to go extinct was the cave bear (Ursus spelaeus), at about 28 kyr B.P. in western Europe (Pacher and Stuart, 2009;Bocherens et al., 2014;Baca et al., 2016). There are various hypotheses regarding the causes of extinction. ...
... This last theory, which attributes part of the reasons for the extinction to humans, is supported by evidence of cut marks found on cave bear remains throughout Europe during the MIS 3 period (Armand et al., 2004;Münzel and Conard, 2004;Terlato et al., 2019;Blasco et al., 2020). Finally, a well-established theory suggests that their extinction can be attributed to the declining availability of plant-based resources (Pacher and Stuart, 2009;Bocherens, 2019). The cave bear is an exceptional example of an evolutionary adaptation, transitioning from a carnivore-scavenger dominated niche to establishing an ecological similarity with the niche occupied by the large-sized herbivores of the Middle Pleistocene (Croitor and Brugal, 2010). ...
Cave bears (Ursus spelaeus) have been recognized as being predominantly herbivorous. However, RomanianCarpathian cave bears have displayed a flexible diet based on a wide range of δ15N values. In this work, weanalyse 97 lower first molars of Ursus spelaeus and contemporaneous Ursus arctos from different sites in thisregion, using dental microwear analysis to reconstruct their pre-dormancy diets during MIS 3. Newly obtainedradiocarbon dates of the individuals are also reported. The short-term diet of all populations studied is identifiedas omnivorous, with a preference for the consumption of hard objects. For the first time, the widespread presenceof puncture pits on U. spelaeus populations has been observed, which could be linked to the intake of seeded andhard-shelled fruits from temperate broadleaf trees. Hence, the dietary patterns of Carpathian cave bears exhibitsimilarities to those of present-day brown bears inhabiting the same region and suggests that the Carpathiansserved as an ecological refuge for cave bear specimens. This is reinforced by an early 27.1 cal kyr B.P. date froman individual from Ursilor Cave, which sheds light on the potential dietary continuity of Romanian Carpathiancave bear populations even during the final phases leading up to their extinction
... Palaeoclimatological data from: Böse et al. (2012), Gibbard & Head (2009), Gibbard et al. (2005, modified 2007), John (1997), Litt et al. (2007), McMillan (2005) and Zagwijn (1960). Fig. 1 : Une représentation de l'origine et de l'histoire de l'évolution au cours du Pléistocène moyen / supérieur de la lignée arctoïde-spéloïde; La distribution asiatique est représentée en bleu, la distribution européenne en jaune et la distribution européenne et asiatique en vert (Argant, 2009 ;Athen & Pfretzschner, 2005 ;Baca et al., 2016 ;Barlow et al., 2018 ;Baryshnikov, 2008 ;Baryshnikov & Foronova, 2001 ;Baryshnikov & Kalmykov, 2005 ;Baryshnikov & Puzachenko, 2011, 2019Bocherens et al., 2011bBocherens et al., , 2014Bon et al., 2008Bon et al., , 2011Curry, 2010 ;Erdbrink, 1953 ;Hofreiter et al., 2002Knapp et al., 2009 ;Kurtén, 1976 ;Kohn et al., 1995 ;Krause et al., 2008 ;Loreille et al., 2001 ;Madurell-Malapeira et al., 2009 ;Martini et al., 2014 ;Moullé, 1992 ;Münzel et al., 2011 ;Orlando et al., 2002 ;Pacher & Stuart, 2009 ;Rabeder et al., 2000Rabeder et al., , 2002Rabeder et al., , 2004Rabeder et al., , 2010Rabeder & Hofreiter, 2004 ;Sabol et al., 2013 ;Stiller et al., 2009Stiller et al., , 2013Taberlet & Bouvet, 1994 ;Tamako et al., 2001 ;Tchernov & Tsoukala, 1997 ;Terlato et al., 2018 ;Torres Pérez-Hidalgo, 1992 ;Zdansky, 1928). Plusieurs scénarios alternatifs ont été postulés, à savoir : (A) U. etruscus est l'ancêtre de la lignée arctoïde (Argant, 1991 ;Ehrenberg, 1929 ;Erdbrink, 1953 ;Kurtén, 1968) -alternativement (B) U. etruscus s'est éteint sans descendance directe et la lignée arctoïde-spéloïdedérive de cf U. minimus (Mazza & Rustioni 1994). ...
... U. deningeri, the archaic spelaean bear, led rise to the Late Pleistocene spelaean taxon U. spelaeus (Orlando et al., 2002 ;Hofreiter et al., 2002Rabeder & Hofreiter, 2004 ;Knapp et al., 2009) toward the end of Middle Pleistocene (Torres Pérez-Hidalgo, 1992 ;Rabeder et al., 2000). U. spelaeus persisted until the end of the Late Pleistocene c. 27 ka BP (Pacher & Stuart, 2009 ;Bocherens et al., 2014 ;Martini et al., 2014 ;Baca et al., 2016 ;Terlato et al., 2018). ...
... Usually dominating faunal associations of cave deposits (Bocherens et al., 2011b), U. spelaeus remains often amount to large bone accumulations. It is thought that this is the result of U. spelaeus being more dependent on caves for hibernation than other ursid species (Kurtén, 1976 ;Fortes et al., 2016 ;Pérez-Rama et al., 2011a), and thus Late Pleistocene caves of northern Eurasia yield so many remains of U. spelaeus that probably died during their hibernation periods (Kurtén, 1976 ;Pacher & Stuart, 2009 ;Grandal D'Anglade et al., 2019). ...
Bears exhibit marked evolution for Pleistocene Europe. Both lineages are thought to have arisen from etruscan bear U. etruscus in the Early Pleistocene, however their high degree of polymorphism has prevented the establishment of an accepted evolutionary scenario. Isotopic analysis and tooth morphology of fossil brown bear U. arctos suggests that it was an omnivorous opportunist. The deningeri bear U. deningeri represents the spelaean bear of the Middle Pleistocene, sharing certain morphological affinities with brown bear U. arctos (frontal bulge and face; occlusal surface of jugular teeth). Within U. deningeri, several subspecies have been distinguished as evolutionary stages leading to the speciation of the cave bear U. spelaeus, the typical spelaean bear of the Late Pleistocene, which dominates cave fossil deposits. The speloïd lineage might serve as a good chronological marker for Pleistocene stratigraphic levels. There are several morphologically distinct lineages within U. spelaeus "sensu lato", of controversial taxonomic status. Herbivorous feeding habits for U. spelaeus "s.l." have been inferred from morphology (tooth, skull, jaw), demographics, and stable isotope analysis. This dietary difference between brown bears and cave bears shows that ecological competition was probably limited between both types. Paleo-genetic studies suggest that cave bears gradually lowered their reproductive rate (between 52,800 and 27,800 y BP) which led to their extinction at the onset of the last glacial maximum. Climatic changes are the main suggested causes responsible for the extinction of U. spelaeus.
Les ours présentent une évolution marquée pour l'Europe du Pléistocène. On pense que les deux lignées sont issues de l'ours étrusque U. etruscus au Pléistocène inférieur, mais leur degré élevé de polymorphisme a empêché l'établissement d'un scénario évolutif accepté. L'analyse isotopique et la morphologie des dents de l'ours brun fossile U. arctos suggèrent qu'il s'agissait d'un omnivore opportuniste. L'ours de Deninger U. deningeri représente l'ours spéléen du Pléistocène moyen, partageant certaines affi-nités morphologiques avec l'ours brun U. arctos (renflement frontal et face; surface occlusale des dents jugales). Au sein d'Ursus deningeri, plusieurs sous-espèces ont été distinguées comme des stades évolutifs conduisant à la spéciation de l'ours des cavernes U. spelaeus, l'ours spéléen typique du Pléistocène supérieur, qui domine les dépôts fossiles des cavernes. La lignée spéloïde pourrait servir de bon marqueur chronologique pour les niveaux stratigraphiques du Pléistocène. Il existe plusieurs lignées morphologique-ment distinctes au sein de U. spelaeus «sensu lato», de statut taxonomique controversé. Des habitudes alimentaires herbivores de l'U. spelaeus «s.l.» ont été déduits par la morphologie (dent, crâne, mâchoire), la démographie et l'analyse des isotopes stables. Cette différence alimentaire entre les ours bruns et les ours des cavernes montre que la concurrence écologique était probablement limitée entre les deux types. Des études paléogénétiques suggèrent que les ours des cavernes ont progressivement abaissé leur taux de reproduction (entre 52800 et 27800 ans BP), ce qui a conduit à leur extinction au début du dernier maximum glaciaire. Il est suggéré que les changements climatiques sont les causes principales de l'extinction de l'U. spelaeus.
... During the same �me range, most of the other European megafauna, such as giant deer, woolly rhinoceros, steppe bison or cave lion got ex�nct as well (Hughes et al., 2006;Immel et al., 2015;Kuzmin, 2010;Stuart and Lister, 2011). Cave bears and cave hyena became ex�nct before the end of the glacial period (Baca et al., 2016;Pacher and Stuart, 2009;Rohland et al., 2005;Varela et al., 2010). This ex�nc�ons happened not only in Europe but worldwide (Barnosky et al., 2004;Barnosky and Lindsey, 2010;Faith and Surovell, 2009;Gill et al., 2009;Krantz, 1970;Louys et al., 2007;Mann et al., 2013;Miller et al., 1999;Roberts et al., 2001), even if not completely simultaneously, leading to two main hypotheses of Late Pleistocene megafauna ex�nc�on (Brook and Bowman, 2002). ...
... This hypothesis is cri�cized primarily for the European megafauna (Barnosky et al., 2004;Carotenuto et al., 2018;Řičánková et al., 2018), as this coexisted with Neanderthal and AMH for thousands of years. Nevertheless, humans, especially the AMH, cannot be taken out of responsibility (Haynes, 2018;Kitagawa et al., 2012;Lorenzen et al., 2011;Münzel and Conard, 2004a;Pacher and Stuart, 2009;S�ller et al., 2010;S�ner, 1999). For example, isotopic studies showed that the mammoth was locally over-hunted during the Aurignacian and the Grave�an . ...
... Moreover, we have excluded the cave bear (Ursus spelaeus), which belongs to the order Carnivora, but did not have a carnivorous diet Münzel et al. 2014;Naito et al. 2016;Nelson et al. 1998;Pacher and Stuart 2009). Furthermore, due to their hibernation behavior, it is possible that cave bears died in the cave sites without human influence (Baca et al. 2016;Kitagawa et al. 2012;Münzel and Conard 2004a;Nelson et al. 1998;Pacher and Stuart 2009;Stiner 1999). ...
In many countries and fables, characteristics such as "smart" and "sly" as well as other human-like behavior have been attributed to foxes for a long time. However, is it also smart and sly to invoke foxes to prove human behavior from a long time ago?
Remains of arctic and red foxes (Vulpes lagopus and Vulpes vulpes) are known from almost every European Late Pleistocene archaeological site (about 100 to 13 kyr ago). Of particular interest in archaeological studies are their canines, found in the cave sites of the Swabian Jura (Baden-Württemberg, Germany), originating from Aurignacian (about 42 to 34 kyr ago) and Gravettian (about 34 to 30 kyr ago). The canines were perforated by early modern humans and most likely worn as pendants or ornaments on clothes. Cut marks on fox bones show that fur and meat were important as well.
With my PhD thesis, I would like to explore beyond the obvious evidences the following three questions: How did Neanderthals in the Middle Palaeolithic (about 100 to 42 kyr ago) and modern humans in the Late Palaeolithic (about 42 to 14 kyr ago), hunt and use foxes? Which feeding habits did Late Pleistocene foxes follow and were they influenced by Neanderthals or modern humans, for example by human hunting behavior? Could foxes be used as an indicator of Palaeolithic occupation intensity and human impact on the Late Pleistocene environment?
To answer these questions, I focused on the cave sites of the central Swabian Jura (Ach- and Lonetal) and the Hegau Jura, located at the southwestern edge of the Swabian Jura. To discuss my first question, I used 26 published zooarchaeological reports from twelve sites in the Ach- and Lonetal and re-examined the abundance of foxes over time using Bayesian statistics. I found out that foxes were more abundant in the archaeological record from the Aurignacian period onwards and that this was due to hunting activities by modern humans. Traps were likely used for hunting foxes, some of them baited with food leftovers and set at game passes in the vicinity of the inhabited caves. In contrast, foxes from the Middle Palaeolithic layers entered the caves naturally. To answer my second question, I used the analysis of stable carbon and nitrogen isotopes from the collagen of archaeological fox bones and reconstructed both the diet and the trophic niches. Thereby I could prove three basic feeding behaviors, one of them was influenced by humans. Some foxes from the Upper Palaeolithic showed synanthropic behavior, i.e. an adaptation to humans and their food resources, and this already about 30,000 years before the Neolithic. To address my last question, I used the knowledge gained from the previous two questions. This allowed me to establish a positive correlation between the abundance of foxes and the intensity of human settlement, as well as the occurrence of synanthropic foxes with the human population density and the hunting pressure on the megafauna in that region. Foxes can thus be used as a proxy for human activities and population density. The opportunistic foxes reacted to the environment influenced by human behavior by adapting their feeding strategies already in the early Palaeolithic periods.
... Despite bears typically living nearly three decades, the causes of these deaths remain unclear. Tooth wear suggests a plant-based diet, aligning with the results of the stable isotope analysis reported below and with their herbivorous tendencies as seen in similar species 13,14,[23][24][25][26] . Skeletal remains exhibit extensive chemical corrosion (Fig. 3a-c, e-g), except for teeth. ...
... Habitat fragmentation, likely due to ongoing glaciation, had cascading adverse effects on ecosystems [32][33][34][35] . Cave bears, highly specialised in a vegetarian diet, were particularly vulnerable due to their ecological limitations 13,14,23,26 . Their susceptibility stemmed from their strong dependence on the availability and composition of vegetation 20 . ...
This comprehensive study examines fossil remains from Niedźwiedzia Cave in the Eastern Sudetes, offering detailed insights into the palaeobiology and adversities encountered by the Pleistocene cave bear Ursus spelaeus ingressus. Emphasising habitual cave use for hibernation and a primarily herbivorous diet, the findings attribute mortality to resource scarcity during hibernation and habitat fragmentation amid climate shifts. Taphonomic analysis indicates that the cave was extensively used by successive generations of bears, virtually unexposed to the impact of predators. The study also reveals that alkaline conditions developed in the cave during the post-depositional taphonomic processes. Mortality patterns, notably among juveniles, imply dwindling resources, indicative of environmental instability. Skeletal examination reveals a high incidence of forelimb fractures, indicating risks during activities like digging or confrontations. Palaeopathological evidence unveils vulnerabilities to tuberculosis, abscesses, rickets, and injuries, elucidating mobility challenges. The cave’s silts exhibit a high zinc concentration, potentially derived from successive bear generations consuming zinc-rich plants. This study illuminates the lives of late cave bears, elucidating unique environmental hurdles faced near their species’ end.
... The cave bear (Ursus spelaeus sensu lato) is one of the prominent representatives of the last ice age. During the late Pleistocene, its territory extended throughout most of Europe (Pacher and Stuart, 2009), and it is one of the most studied extinct species due to the abundance of fossil remains in caves. The prevailing opinion is that cave bears died there of natural causes because they did not successfully survive hibernation (Kurtén, 1976;Andrews and Turner, 1992;Stiner et al., 1998;Debeljak, 2002Debeljak, , 2004Debeljak, , 2007Grandal-d'Anglade et al., 2019). ...
... The prevailing opinion is that cave bears died there of natural causes because they did not successfully survive hibernation (Kurtén, 1976;Andrews and Turner, 1992;Stiner et al., 1998;Debeljak, 2002Debeljak, , 2004Debeljak, , 2007Grandal-d'Anglade et al., 2019). The cave bear became extinct during the Last Glacial Maximum as the climate cooled (Hilderbrand et al., 1996;Pacher and Stuart, 2009). ...
In cave bear studies, stable isotope analysis of carbon and nitrogen is typically conducted on bone collagen. However, challenges arise due to biases in the classification of bones into age groups, which can be influenced by various factors that affect bone size. To overcome such issues, we present and evaluate a novel approach that involves sampling dental collagen from different ontogenetic stages of 45 fossil teeth, where only a portion of each tooth formed during the periods of interest was sampled. This approach enabled us to obtain a carbon and nitrogen fingerprint specific to a particular period in a cave bear's life without interference from earlier stages. Stable isotopic analysis revealed a rapid decrease in δ 15 N values from 3 to 15 months of age. After that, the δ 15 N values remain stable. Based on the nitrogen isotope composition (15 N/ 14 N), cubs aged 0-3 months exclusively consumed milk, while the gradual introduction of solid plant food occurs at 5-10 months, resulting in a shift in δ 15 N values. Based on similar δ 15 N values in older age groups, it can be assumed that juveniles did not resume milk suckling after the first hibernation. The δ 13 C values increase until 15 months, followed by a gradual decrease until adulthood, potentially attributable to hibernation. The δ 13 C values also seems to indicate isotopic differences between yearlings with low δ 13 C values that successfully established hibernation and those with high δ 13 C values that failed to hibernate and starved. This novel observation, i.e., decreasing δ 13 C values from the second year of life onward, agrees with predicted isotopic skeletal collagen δ 13 C values calculated from modern brown bear blood plasma. In summary, the ontogenetic sampling strategy facilitated the acquisition of further paleobiological insights into the distinct life periods of cave bears.
... The cave bears (Ursus spelaeus) at Sclayn (Martin et al., 2017a) do not exhibit such a high Ca isotopic variability because their dietary niche was more restricted than for brown bears. Tooth shape and N isotope suggest an herbivorous diet for this taxon (Bocherens et al., 1994;Pacher and Stuart, 2009). In extant brown bears, meat consumption is described as highly variable depending on the resource (Pacher and Stuart, 2009 (Fig. 2), indicating varying Ca dietary sources during the period of bone formation. ...
... Tooth shape and N isotope suggest an herbivorous diet for this taxon (Bocherens et al., 1994;Pacher and Stuart, 2009). In extant brown bears, meat consumption is described as highly variable depending on the resource (Pacher and Stuart, 2009 (Fig. 2), indicating varying Ca dietary sources during the period of bone formation. Indeed, the turnover is different between trabecular and cortical bone. ...
Les outils isotopiques sont couramment utilisés en anthropologie depuis plus de quarante ans car ils apportent des informations complémentaires aux études ostéologiques et archéozoologiques. Le travail entrepris au cours de cette thèse vise à étudier le potentiel que représentent les isotopes du calcium pour la reconstruction paléo-écologique. En effet, la composition isotopique en calcium des mammifères est en majeure partie définie par les apports en calcium de l’alimentation, en particulier les produits laitiers. En se plaçant dans un contexte où le seul produit laitier consommé est le lait maternel, l’impact des autres sources alimentaires sur la composition isotopique en calcium est plus facilement identifiable, les isotopes du calcium deviennent ainsi un indicateur efficace du niveau trophique. Nous présentons dans cette thèse les résultats de l’analyse isotopique du calcium osseux conduite sur dix individus néandertaliens et de la faune de huit sites du Paléolithique moyen. Il est démontré que le choix de l’os comme support de l’analyse isotopique est justifié dans la mesure où celui-ci fait montre d’une résistance suffisante au processus diagenètiques de premier ordre. Les mécanismes du fractionnement isotopique au sein d’un réseau trophique
complet sont discutés sur la base d’observations chez les végétaux et les animaux actuels. L’analyse de la composition isotopique d’ossements néandertaliens provenant de toute la France a permis de mettre en évidence une grande diversité des compositions isotopiques en calcium. Cette diversité est expliquée par une variabilité dans le spectre des proies et par une variabilité de la fraction végétale au sein de l’alimentation. Nos résultats montrent une pluralité des régimes alimentaires chez l’Homme de Néandertal même au sein de groupes géographiquement proches.
Isotopic tools have been used in anthropology for over forty years as they provide
complementary information to osteological an archeozoological studies. The work
undertaken as part of this doctoral thesis aims at studying the potential of calcium isotopes to rebuild paleo-ecology. Indeed, the calcium isotopic composition of mammals is largely defined by the calcium intake of the diet, in particularly dairy products. In a context where the only dairy product consumed is breast milk, the impact of the other food sources on calcium isotopic composition is more easily identifiable and thus calcium isotopes become an effective indicator of trophic level position. In this thesis, we present the results of the isotopic analysis conducted on bones of the remains of ten Neandertal and the fauna from eight archaeological site related to the middle Paleolithic. The choice of bone as a support for the isotopic analysis is justified as it shows sufficient resistance to the first steps of diagenetic processes. Based on the observation of present-day plants and animals, the mechanisms of isotope fractionation within a complete food web are discussed.
The comparison between Neandertal from all over France and the faunal assemblage reveal a great diversity of calcium isotopic compositions. This diversity is explained by a variability in the prey spectrum and a variability in the plant fraction within the diet. Our results show a plurality of diets among Neandertals even within geographically close group.
... Climatic cooling and inferred decreased vegetation productivity at the end of the Upper Pleistocene are suggested causes responsible for the extinction of U. spelaeus (Lascu & Puscas, 2002;Orlando et al., 2002;Pacher & Stuart, 2008). Mitochondrial DNA studies by Stiller et al. (2010Stiller et al. ( , 2014 suggest that a decline in the cave bear population occurred between 52,800 and 27,800 BP, which gradually lowered their reproductive rate and led to their extinction. ...
... Radiocarbon dating suggests that the extinction happened c. 27,800 BP, at the onset of the last glacial maximum (Pacher & Stuart, 2009;Münzel et al., 2011;Bocherens et al., 2014). Pacher & Stuart (2008) have argued that it is unlikely that their extinction was brought about by anthropogenic causes. However, other authors argue that climatic changes alone cannot explain the extinction of the cave bear, and that anthropogenic factors also contributed to its extinction (Münzel & Conard, 2004;Germonpré & Hämäläinen, 2007;Stiller et al., 2010;Münzel et al., 2011). ...
An important debate amongst cave bear researchers is the diet and ecoethology of these animals. A great controversy surrounds the phylogenetic resolution of the cave bear lineage. Previous investigations have focussed on teeth, skull and mandibular bones. This study focuses on the morphology of metapodia as an alternative, and for the first time 3D digital methods have been applied on these elements. This study involved 3D scanning of metacarpals and metatarsals belonging to Ursus spelaeus, Ursus arctos and U. deningeri, followed by landmarks assignment onto the surface of virtual bones...
... Macroscopic remains of extinct fauna are common at the entrance, where human activities are often the main accumulating agent, or in the interiors of several European cave systems (e.g., Pacher and Stuart, 2008;Ridush et al., 2013). ...
... No bone and phosphate remains have been found within the sediments below the floatstone and there is no indication that the cave might have been inhabited by any animals at that time. At the present state of research, there is no reliable dates or other evidence to ascertain one specific paleoclimatic event, but it is very tempting to correlate the ending of this phase with speleothem formation to a warmer interglacial stage, when global palaeoclimate proxies and regional reconstructions suggest high speleothem growth rates (Drysdale et al., 2004) b) Stage 2: Last Glacial between ~50 ka and ~25 ka Radiometric dates and the large amount of cave bear remains (Ursus spelaeus and U. arctos) associated with bear footprints, claw marks and nests, indicate that the cave was a lair for hibernating bears which repeatedly, but not necessarily yearly, occupied it during this phase (Fig. 17), before their extinction (about 25 ka, Pacher and Stuart, 2008;Terlato et al., 2018). ...
The Bàsura Cave (Toirano, Savona, NW Italy) hosts important cave bear bone assemblages and a numerous and varied, tracks and traces record left by humans and other producers. An outstanding element of the analysed material is represented by fossil bear fur fragments, which were found in the inner deposits of the cave, and that, to date, are virtually unknown in the cave global record. After analysing and discussing micromorphological features of the inedited material, we integrate and interpret new radiocarbon data, along with taphonomic, sedimentological, geochemical and mineralogical evidences, with the aim of improving our understanding about the nature and chronology of the bear fur-bearing deposit. The bear fur fragments are included in a stratigraphic succession corresponding to a secondary deposit, formed after the dismantling, reworking and redeposition of a former bear-bearing deposit, as a result of short but intensive flooding events that most probably took place at the end of the Last Glacial Maximum. After sediments redeposition, important diagenetic changes have occurred and probably driven by guano deposits, whose pre-existence, in absence of record, is inferred from corrosion features, nutrient concentrations, mineral species identified (REE bearing hydroxyapatite), and claw traces left by bats on the cave ceiling and walls. Diagenetic imprint derived by guano deposits caused mineralization of bear fur fragments by replacement with apatite, which faithfully copied the form and structure of hairs but also of vegetal tissues, phytoliths and pollen found within them. Our study demonstrates for the first time that the bear fur is one of the main vectors in introducing botanical microremains into the interior of the “Old World” caves.
... Tutto ciò può aver favorito l'inserimento nella nicchia ecologica di Ursus spelaeus di nuove specie migratrici più competitive ed adattabili all'ambiente, come ad esempio Ursus ingressus e Ursus arctos. Un'altra possibile teoria è la troppa selettività riguardo ai luoghi dove l'orso delle caverne trascorreva il letargo (Pacher & Stuart, 2008), poiché i reperti di Ursus spelaeus fi nora raccolti sono stati ritrovati quasi esclusivamente in grotte, il che può suggerire l'utilizzo di questi luoghi come unica tipologia di riparo per il periodo di letargo invernale. Un'altra ipotesi interessante è data dagli studi di Diedrich (2006,2012,2013), i quali indicano un'attività di predazione di Ursus spelaeus durante il letargo da parte di altri carnivori, come ad esempio Panthera leo spelaea. ...
... Ursus spelaeus scompare in un periodo corrispondente a Greenland Stadial 3, secondo le datazioni del 14 C (Pacher & Stuart, 2008). Ad esempio in Italia centrale sono stati rinvenuti dei reperti di Ursus spelaeus nella Grotta del Chiostraccio, in Toscana, la cui datazione al 14 C è stata valutata 24,030 ± 100 y BP (Martini et al., 2014). ...
This article gives a perspective of several years of
morphological and morphometrical studies of the paleontological collection of Ursus spelaeus’s remains founds in the
Pocala cave, kept in the Civic Museum of Natural History of Trieste. The aim of this research is to develop a new metrical
approach for the study of the skull and dentition of Ursus spelaeus, based on previous morphological and morphometrical
studies. Goal of the present paper is an easy sexing of the remains and giving a hint of the homogeneity of the population
researched.
... The cave bears (Ursus spelaeus) at Sclayn(Martin 307 et al., 2017) do not exhibit such a high Ca isotopic variability, because their dietary niche was 308 more restricted than for brown bears. Tooth shape and N isotope suggest an herbivorous diet for 309 this taxon(Bocherens et al., 1994;Pacher and Stuart, 2009). In extant brown bears, meat 310 consumption is described as highly variable depending on the resource(Pacher and Stuart, 2009).311Other ...
... Tooth shape and N isotope suggest an herbivorous diet for 309 this taxon(Bocherens et al., 1994;Pacher and Stuart, 2009). In extant brown bears, meat 310 consumption is described as highly variable depending on the resource(Pacher and Stuart, 2009).311Other scavenging behaviors, including necrophagia following hibernation, and bone gnawing,312 which can be associated with direct bone ingestion, are also reported. ...
The calcium isotopic composition (δ44/42Ca) of bone and tooth enamel can be used for dietary reconstructions of extant and extinct mammals. In natural conditions, the δ44/42Ca value of bone and teeth varies according to dietary intake with a constant isotopic offset of about −0.6‰. Owing to the poor conservation of collagen, carbon (C), and nitrogen (N) isotopic compositions of the Regourdou Mousterian site (MIS 5, Dordogne, France) previously failed to provide any paleodietary information. Therefore, to reconstruct the trophic chain, we have measured calcium (Ca) isotopes from fossil bone samples of the fauna from the Regourdou site, as well as from three bone samples of the Regourdou 1 Neandertal specimen. The results show a taxon-dependent patterning of the Ca isotopic compositions: herbivores generally have higher δ44/42Ca values than carnivores. All the δ44/42Ca values of Regourdou 1 are low (<-1.6‰), placing this specimen amid carnivores. Using a bone-muscle Ca isotopic offset determined on extant animals, we further show that the δ44/42Ca value of the Regourdou 1 diet, and that of most carnivores, cannot be accounted for by the consumption of meat only, as plants and meat have indistinguishable δ44/42Ca values. Mass balance calculations indicate that the low δ44/42Ca values of the Neandertal's carnivorous diet are explained by the ingestion of bone marrow containing as little as 1% trabecular bone. Our results show that the Regourdou 1 Neanderthal consumed a mixture of various herbivorous prey, as well as trabecular bone, which probably occurred when marrow was ingested, by accident or intentionally.
... Nucleotide sequences of the obtained fragment of mtDNA correspond to the mtDNA of Ursus deningeri, a cave bear of the Middle Pleistocene which transitioned into Ursus spelaeus sensu lato in the Late Pleistocene, and its descendents Ursus spelaeus sensu stricto and Ursus ingressus which probably disappeared from the Alps and adjacent areas before 28 ka (Pacher & Stuart 2009;Dabney et al. 2013). These cave bear species/subspecies are in complete alignment in the region of cyt b gene amplified in our study. ...
... In both cases, these ages only implicate the limit to when the remnants of bones were transported to the cave. Cave bears lived past MIS 3 (up to ca. 20 ka BP) according to other studies (Rabeder et al. 2004a;Pacher & Stuart 2009). ...
Loose bone fragments coated with calcite precipitate including a loose/broken stalagmite containing small fragments of cemented bones were collected from the Postojna Cave to investigate whether deoxyribonucleic acid (DNA) can be determined. The study is complemented by the Fourier-transform infrared spectroscopy-attenuated total reflectance (FTIR-ATR) analysis in order to determine the alteration of the bones and to test whether this analysis can be used as an indicator of possible DNA preservation. In addition, geochemical analyses were conducted in order to determine whether the associated flow-stone/stalagmite is suitable for elucidating the timing of bone thanatocoenosis and further palaeoenvironmental analyses. The organic matter (collagen) is poorly preserved. However, we succeeded in amplifying a 94 bp long fragment of the cytochrome b (Cyt b) gene of mitochondrial DNA (mtDNA) in polymerase chain reaction (PCR) for one sample, and in sequencing the amplified products. The 70 bp long sequence obtained corresponds to that of the Cyt b of the cave bear (Ursus deningeri or Ursus spelaeus sensu lato). The uranium-thorium dating of the speleothem covering the bones revealed its thanatocoenosis occurred prior to 55 ka, most likely in the late marine isotope stage 4 or early marine isotope stage 3. High porosity and recrystallisation of the flowstone/stalagmite at this part of the cave prevent high-resolution palaeoclimatic interpretation; however, low-resolution stable isotope geochemistry suggests a steppe-like environment during the subsequent growth of the speleothem.
... BP); and a cave bear tooth provided an estimate of 39,420 + 1070/-940 years bp (VERA-2543, 43,467 AE 829 cal. BP) (Fladerer, 1997;Pacher and Stuart, 2009;Rabeder and Kavcik, 2013). We propose that the people responsible for the deposits were intermediate between robust and gracile Homo sapiens, but perhaps closer to the so-called Neanderthals than the so-called modern humans (Bednarik, 2011). ...
... The two lowest of these are superimposed over masked finger grooves, and they are much better preserved than the flutings. This evidence provides the latter with minimum antiquity exceeding the time of extinction of Ursus ingressus in the region (roughly 27.8 to 25 ka ago (Pacher and Stuart, 2009;Baca et al., 2016). One of these superimposed markings is a deep gash where a claw has, with considerable force, cut through the speleothem layer and into the limestone bedrock beneath it. ...
After well over a century of archaeological research in Drachenhöhle, the largest cave bear lair in the Alps, the first Pleistocene rock art in central Europe has been discovered deep in the cave. Two small panels of juvenile finger flutings occur together with cave bear claw marks at the only water source of the area. The site is within a few metres of the cave's large human occupation site, excavated in 1921. It is attributed to the Alpine Palaeolithic or Olschewian, a tradition of montane-adapted people of the Early Upper Palaeolithic of central Europe. The generic phenomena of finger fluting and moonmilk speleothems are discussed to provide a general context for the subject. The cave art is then described and analysed, and the previous claims for “Palaeolithic” age of other central European sites are briefly considered.
... The temporal distribution of this faunal association, as also confirmed by recent radiocarbon datings (Bigagli et al., 2017), corresponds to the MIS3 (Upper Pleistocene), during which numerous climatic oscillations occurred in a colder-thantoday palaeoenvironmental context. Among these remains, the largest number of fossils belong to the cave bear (Ursus spelaeus Rosenmüller, 1794), an extinct species of ursid carnivorans that fed almost exclusively on plants and was widespread in central Europe and western Asia from about 300 ka to about 27 ka (Kurtén, 1976;Pacher & Stuart, 2009). At the Tecchia Preistorica, during an excavation campaign in 2014, two individuals of cave bear were found, here informally named "Cub 1" and "Cub 2". ...
The karst complex of “Grotte di Equi” (i.e. the Equi caves) is located at about 300 m a.s.l along the Fagli Valley, close to the village of Equi Terme, in the northern area of the Apuan Alps. This cave system develops in the Metacalcari con Selce Formation (Apuan Unity, Lower Jurassic), consisting
of flint-bearing metacalcilutites and metacalcarenites. Above the entrance to Grotte di Equi, at about 350 m a.s.l., the Tecchia Preistorica opens along the northern slopes of Pizzo d’Uccello. This NEtrending cavity is comprised of two parts: the “Riparo” (i.e., shelter) and the “Grotta” (i.e., cave). The
Tecchia Preistorica represents a paleontological and archeological site of prime importance. Here, several excavations and researches have been performed for over a century (Ghezzo et al., 2014; Ghezzo & Rook, 2014, 2015). The faunal remains found inside the Tecchia Preistorica, most of which lack precise stratigraphic whereabouts, are mostly comprised of terrestrial mammals, including artiodactyls, carnivores, bats, lagomorphs and rodents. The temporal distribution of this faunal association, as also confirmed by recent radiocarbon datings (Bigagli et al., 2017), corresponds to the MIS3 (Upper Pleistocene), during which numerous climatic oscillations occurred in a colder-thantoday palaeoenvironmental context. Among these remains, the largest number of fossils belong to the cave bear (Ursus spelaeus Rosenmüller, 1794), an extinct species of ursid carnivorans that fed almost exclusively on plants and was widespread in central Europe and western Asia from about 300 ka to about 27 ka (Kurtén, 1976; Pacher & Stuart, 2009). At the Tecchia Preistorica, during an excavation campaign in 2014, two individuals of cave bear were found, here informally named “Cub 1” and “Cub 2”. Each skeleton lays on a consolidated silt block. Both specimens are in an excellent state of preservation, despite being partially incomplete and disarticulated. Cub 1, better preserved than Cub 2, lies on his left side. The skull is in anatomical connection with the vertebral column, preserving an undeformed three-dimensional morphology. All the skull bones are discernible, well preserved and almost perfectly articulated (except the nasal bones). All the sutures between the skull bones are still
unfused and the bregmatic or anterior fontanel, between the frontal and parietal bones, is open. The skeleton of Cub 1 has a well-preserved vertebral column, especially in the cervical tract (which is still in anatomical connection with the skull), as well as in the thoracic tract. All the vertebrae are partially ossified and made up of distinct elements probably corresponding to different centers of ossification. The shoulder girdle is completely disarticulated and partially preserved. The rib cage is in an excellent state of conservation and partly disarticulated. Most of the ribs are in anatomical connection with the thoracic vertebrae and parallel to each other. As regards the limbs, only the right humerus and the tibia are preserved, without epiphyses. The skeleton of Cub 2, lying dorsal side-up, is well preserved, although it is less complete and articulated than that of Cub 1. The vertebral column is well preserved and partly disarticulated, lying on the ventral side. Close to the forelimbs, some poorly preserved, displaced vertebrae take their place. The thoracic vertebrae have a subspherical shape and they are partially ossified, consisting of distinct elements corresponding to the different centers of ossification. The shoulder girdle is completely disarticulated and the displaced right scapula is exposed, in dorsal view, above the rib cage. The latter is well preserved and only partially disarticulated, with the ribs in anatomical connection with the vertebral column. As regards the forelimbs, there are two humeri,
two ulnae and two radii. They are not in anatomical connection and the epiphyses are absent. The skeletons of Cub 1 and Cub 2 belong to very young individuals. The vertebrae are not fully ossified and epiphyses of the long bones are absent. The cranial sutures of Cub 1 are not closed and some fontanelles are clearly observable. Furthermore, this specimen does not display any teeth, which usually appear as gems after the third week of life in the present-day brown bear (a close relative of
the cave bear). By comparing the measurements of the scapulae and long bones (humeri and radii) of Cub 1 and Cub 2 with the values reported in literature for other very young individuals of U. spelaeus from various sites of central Europe (Germonpré & Sablin, 2001; Fosse & Cregut-Bonnoure, 2014), an age at death of less than a week is reconstructed for both specimens. The life and death of the two
cave bear cubs of Equi are closely related to the cycles of hibernation and reproduction of U. spelaeus, which various lines of evidence indicate to be similar to those of modern Ursus species. Usually, the pregnant females give birth during hibernation (from November to January) in sheltered dens, as we hypothesize to have happened in the Tecchia Preistorica. The nutritional conditions and body mass of the mother affect the duration of gestation and nursing, as well as the number (from 1 to 4) and size of the cubs (Robbins et al., 2012; López-Alfaro et al., 2013). Since Cub 1 and Cub 2 were found next to each other and in the same stratigraphic horizon, and their body measurements appear to be very similar, an origin of both from the same litter is proposed herein. In modern bears, during the first few weeks after giving birth, the mother is almost completely inactive. The infant mortality rate is very high due to malnutrition, diseases and accidents. Various hypotheses can be proposed for explaining the death of Equi cubs, but it seems reasonable to supposed that they died from malnutrition, due to the critical nutritional conditions and the low content of adipose tissue of the mother.
... The immediate precursor of the cave bear was Deninger's bear (Ursus deningeri von Reichenau, 1904), a species restricted to Pleistocene Europe between 1.8 Myr to 100 kyr (Stuart, 1996;Königswald and Heinrich, 1999). Cave bears disappeared before the onset of the LGM (Pacher and Stuart, 2009;Bocherens et al., 2014;Baca et al., 2016) due to a poorly understood combination of factors, such as competition with and predation by other mammals, climatic deterioration, and loss of genetic diversity. Understanding the mechanisms that led to the extinction of cave bears, one of the first Pleistocene megafaunal species to have suffered this fate, could be a first step toward unraveling the extinction of Quaternary megafaunal species in general. ...
A recent palaeontological excavation in Muierilor Cave, southern Carpathians (Romania), has recovered one of the largest populations of Late Pleistocene wolves (Canis lupus Linnaeus, 1758; MNI =25) documented in Eurasian cave deposits. Many individual nests with wolf bones, including fully articulated skeletons, were found in a remote passage of the cave suggesting a communal behavior similar to that of modern conspecifics. Radiocarbon dates indicate constant use of the cave by these canids throughout the Marine Isotopic Stages 3 (MIS 3; 59‒29 cal ka BP). This deposit affords an opportunity to study the extraordinary resilience and adaptability of C. lupus following the extinction of other large carnivores at the onset of the Last Glacial Maximum (LGM). Using taphonomic, stable isotope and radiocarbon data, we make a case that these canids preyed/scavenged upon the cave bears (Ursus spelaeus Rosenmüller, 1794; MNI =82) that used the same cave for hibernation, along with other Ice Age megafauna such as cave lions (Panthera spelaea Goldfuss, 1810; MNI =4) and cave hyenas (Crocuta spelaea Goldfuss, 1823; MNI =3), which were also found in the same bone assemblage.
... Regarding ecosystem composition and competition, there was a plethora of carnivore species inhabiting western Eurasia at the time of the Neanderthal disappearance and the arrival of AMH populations, coexisting with these human groups for millennia before their eventual extinction (O'Regan et al., 2002;Burger et al., 2004;Sommer and Benecke, 2006;Stuart and Lister, 2007, 2011Pacher and Stuart, 2009;Martini et al., 2014;Mackiewicz et al., 2017). The diverse Late Pleistocene carnivore guild in Eurasia included large taxa, such as cave bear (Ursus spelaeus), brown bear (Ursus arctos), cave lion (Panthera leo spelaea), medium-sized taxa, such as leopard (Panthera pardus), boreal lynx (Lynx lynx); cave hyaena (Crocuta spelaea), spotted hyaena (Crocuta crocuta), wolf (Canis lupus) and cuon (Cuon alpinus), as well as smaller carnivores, such as wildcat (Felis silvestris), lynx (Lynx pardinus), fox (Vulpes vulpes), Arctic fox (Vulpes lagopus), badger (Meles meles), and several mustelid species. ...
The comparative assessment of dietary choices as part of landscape use strategies deployed by Neanderthal and Anatomically Modern Human populations in Eurasia constitutes a fundamental avenue of Palaeolithic research. The increasing number of taphonomic assessments enables a better understanding of what remains were brought to sites by human hunters versus mammalian carnivores or raptors. A zooarchaeological approach can further elucidate the spatio-temporal dynamics of interaction between carnivores and human populations in terms of landscape use and prey choice during this transitional period. To achieve this objective, we conducted an examination of zooarchaeological and taphonomic data, carnivore indices, and other relevant variables across 36 Middle and Early Upper Palaeolithic sites in northern Iberia. These sites encompass 126 archaeological layers dating from 50,000 to 30,000 years ago. Our comprehensive bibliographic meta-analysis reveals that human occupations by both groups at sites across the region were punctuated by episodes influenced by carnivores. This observation implies that human occupations in northern Iberia during Marine Isotope Stage 3 were generally characterised by instability and limited to short periods, often seasonal in nature. From a zooarchaeological perspective, the combined assessment of taxonomic data, species richness and assemblage diversity highlights that the range and proportion of species acquired by these different human groups are similar, although Anatomically Modern Humans engaged in a sustained trend towards increasing dietary diversification even at sites with assemblages heavily dominated by one taxon.
... The cave bear (Ursus spelaeus) went extinct at about 28 kyr B.P. in western Europe. A well-established theory suggests that their extinction can be attributed to the declining availability of plant-based resources (Pacher and Stuart, 2009). The brown bear (U. arctos), smaller-sized and with broad dietary habits, prevailed and occupied the ecological niche of the U. spelaeus after his extinction (Sommer and Benecke, 2005). ...
... El estado de la cuestión ha avanzado desde sus orígenes, en los que se justificaba la elevada acumulación de restos de Ursus spelaeus en cuevas a partir de la caza llevada a cabo por los homininos (Pacher y Stuart, 2008). Este paradigma ya se superó cuando se demostró que las acumulaciones de U. spelaeus se debían a causas naturales y no antrópicas (Koby, 1941;Kosintsev et al., 2021;Patou, 1988;Villaluenga-Martínez et al., 2014;Zunino et al., 2022), puesto que estos osarios kársticos se habían constituido mediante los restos de los animales muertos durante la hibernación (Mackiewicz et al., 2017;Quilès et al., 2006). ...
Resumen: La relación entre los neandertales y el oso cavernario-Ursus spelaeus durante el Pleistoceno Superior ha sido objeto de controversia debido a que la historiografía tradicional sugería ideas relativas a un posible culto hacia esta especie, junto con otras teorías que respaldan la caza masiva de estos animales. Sin embargo, las pruebas que vinculan una interacción entre el Homo neanderthalensis y el Ursus spelaeus en Europa son limitadas y fragmentarias, y proceden
mayormente de Italia, Alemania o Francia, en los alrededores de los Alpes. Hasta la fecha, no se habían encontrado restos de Ursus spelaeus con evidencias de actividad humana en el Paleolítico Medio en la Península Ibérica. No obstante, este artículo presenta y analiza las consecuencias del descubrimiento de una ulna de Ursus spelaeus con marcas de corte, hallada en los niveles musterienses de la cueva del Esquilléu en Cantabria. Este hallazgo, identificado tras un minucioso estudio de la muestra osteológica disponible, representa un nuevo aporte a este debate, ya que amplía el ámbito geográfico y sugiere que las revisiones tafonómicas podrían ser una fuente de nuevas evidencias.
... which had a great impact on vertebrate populations. During this relatively short geological period, significant changes occurred in the ranges of species (Sommer and Nadachowski, 2006;Sommer and Zachos, 2009;Markova et al., 2008;Cooper et al., 2015;Baca et al., 2017), as well as the extinction of many species of large mammals (Stuart and Lister, 2007;Stewart, 2008;Pacher and Stuart, 2009;etc.). Of particular interest is the history of the development of the mammalian fauna on the territory of the Ural region, a unique biogeographic crossroads of northern Eurasia, currently inhabited by representatives of the European, Siberian, and transpalearctic fauna (Bol'shakov et al., 2000). ...
... A significant number of publications deal with the influence of climate and, in particular, Pleistocene glaciations, on species diversity, dynamics in the ranges of individual species, and the process of extinction in mammals (Stuart, 1991;Lister, 2004;Pacher and Stuart, 2009;Kahlke and Lacombat, 2008;Stuart andLister, 2011, 2012;Kahlke, 2014Kahlke, , 2015. The Last Glacial Cycle Maximum (LGM) was not favourable for mammal assemblage in Europe (Tarasov et al., 2000;Arpe et al., 2011;Strandberg et al., 2011;Tzedakis et al., 2013;Simakova and Puzachenko, 2019;Sommer, 2020), especially compared with the MIS3 megainterstade . ...
... The most ancient faunas were found in the unconsolidated Streletsk (MIS 5) and Khanmei (MIS 4) horizons deposits in the Zhilische Sokola (Table 7). The fauna from the Drovatnitsa Grotto includes Ursus kanivetz (Table 7); this species disappeared at the end of MIS 3 (Pacher and Stuart, 2009). This fact permitted to correlate this fauna with the Upper Pleistocene Nev'yansk horizon (MIS 3). ...
... 25-26 ka bp). Not only does this coincide with the cave bear's extinction in the region (roughly 27.8 to 24 ka ago ;Fladerer 1995;Pacher and Stuart 2009;Bocherens et al. 2014;Baca et al. 2016), it also renders the attribution to the Mousterian impracticable. A series of nine more recently secured 14 C AMS dates from cave bear bones ranges from 31.1 ka to >49.9 ka (Rabeder et al. 2005). ...
A review of the history of uranium-series dating of fossil bone and calcite skins related to rock paintings reveals significant limitations to the credibility of many such results. The 'closed system' conditions required do not seem to apply to many ancient faunal remains and may be lacking in many cases also in the types of speleothems frequently used to secure minimum or maximum ages for cave paintings or petroglyphs. The studies comparing 14 C dates with U-Th results from such reprecipitated carbonates, particularly of the Pleistocene, suggest that the latter tend to be much higher. Recent testing of the method implies that the taphonomy of most such deposits is far too complex to allow the determination of age-governed 230 Th/ 234 U ratios. The U concentrations in coeval calcite skins vary significantly on a millimetre scale, and in some cases, apparent ages can be hundreds of times greater than actual ages. Tests also reveal that results obtained by different laboratories from the same samples differ greatly. The lack of reproducibility and testability of such results, combined with the interventional method of obtaining samples, excludes it from sustainable approaches to rock art dating.
... Numerous cave bear claw marks can be seen in the vicinity of the cave spring, and several occur close to panel B. Two of them were superimposed after the top part of finger flutings was masked by the deposit of cutaneous speleothem. This provides a very conservative minimum age for the rock art because the claw marks must precede the time of extinction of Ursus ingressus in the region (about 27.8 to 25 ka ago; Pacher and Stuart 2009;Bocherens et al. 2012;Baca et al. 2016). One of the two superimposed marks is a deep gash that has shattered the speleothem skin with considerable force, which shows that the deposit was already hardened at the time of that event. ...
The recent discovery of the first authentic Pleistocene rock art in central Europe is reported in one of the classical Palaeolithic cave sites investigated for centuries. The Drachen-höhle in Austria has yielded extensive evidence of human habitation in the Würm I/II Inter-stadial 325 m from its entrance, in total darkness. The cave art occurs a few metres from that site and was produced by a child or children, most probably 39±5 ka ago. It can safely be attributed to people of Olschewian or Alpine Palaeolithic tool traditions, which seem to be by Neanderthaloid and intermediate Homo sapiens hominins. These appear to have harvested hibernating cave bears in their lairs. The Drachenhöhle is the largest such hibernation den known in the Alps, having contained the remains of an estimated 30 000 cave bears.
... The art in Rouffi gnac has never been formally dated, but based on stylistic considerations it is considered to be 13-14,000 years old. However, Sharpe and Van Gelder (2006c) have suggested it could be more than 20,000 years old because there are cave bear scratches on top of two of the drawings and some of the fl utings, and cave bears are thought to have died out in this region some 20,000 years ago (Pacher and Stuart 2009;Sharpe and Van Gelder 2006c). Finger fl utings are found in eight chambers of the cave. ...
... Kurtén (1968) concluded that the species entered Europe during the mid-D-Holsteinian (approximately 230 kyBP) but, more recently, fossil remains of putative brown bears dating to approximately 500 kyBP have been identified (Davison et al. 2011). Irrespective of exactly when brown bears appeared in Europe, they evidently co-existed with the cave bear Ursus spelaeus until the Late Pleistocene when the latter became extinct (Pacher and Stuart 2009). In Britain, the cave bear Ursus deningeri is known from early Middle Pleistocene deposits and was replaced by Ursus spelaeus after the Anglian glaciation. ...
This richly illustrated book gives a detailed account of excavations that extended over
ten years at Stanton Harcourt, Oxfordshire, following the discovery of a mammoth tusk
in 1989. More than 1500 vertebrate fossils and a wealth of other biological material
were recorded and recovered, along with 36 stone artefacts attributable to
Neanderthals.
Today the Upper Thames Valley is a region of green pastures and well-managed
farmland, interspersed with pretty villages and intersected by a meandering river.
The discovery in 1989 of a mammoth tusk in river gravels at Stanton Harcourt,
Oxfordshire, revealed the very different ancient past of this landscape. Here, some
200,000 years ago, mammoths, straight-tusked elephants, lions, and other animals
roamed across grasslands with scattered trees, occasionally disturbed by small bands
of Neanderthals.
The pit where the tusk was discovered, destined to become a waste disposal site,
provided a rare opportunity to conduct intensive excavations that extended over a
period of 10 years. This work resulted in the recording and recovery of more than 1500
vertebrate fossils and an abundance of other biological material, including insects,
molluscs, and plant remains, together with 36 stone artefacts attributable to
Neanderthals. The well-preserved plant remains include leaves, nuts, twigs and large
oak logs. Vertebrate remains notably include the most comprehensive known
assemblage of a distinctive small form of the steppe mammoth, Mammuthus
trogontherii, that is characteristic of an interglacial period equated with marine isotope
stage 7 (MIS 7).
Richly illustrated throughout, Mammoths and Neanderthals in the Thames Valley
offers a detailed account of all these finds and will be of interest to Quaternary
specialists and students alike.
... Recent isotopic and genetic analyses reveal complex populational and ecological niche dynamics for brown bears during the end of the Late Pleistocene and during the Holocene before the expansion of human pressure on the western European habitats (Valdiosera et al. 2007(Valdiosera et al. , 2008Ersmark et al. 2019). On the other hand, there is an ecological niche broadening of the brown bears (Ersmark et al. 2019) after the extinction of the cave bears sensu lato (Pacher & Stuart 2009;Baca et al. 2016). Differences in the niche breadth (and hence in the isotopic values and potentially in the mandibular morphology) could have occurred in the brown bears in areas outside the range of Ursus deningeri, such as the southern half of the Iberian Peninsula (Torres 1988) e.g. in Postes cave and other areas in which these two species were competing. ...
Brown bears (Ursus arctos) diverged from the cave bear lineage c. 1.2 million years ago and likely originated in Asia, where the oldest fossils belong to a Middle Pleistocene chronology. Brown bear fossils from the Middle Pleistocene are scarce in the Iberian Peninsula, especially when compared to the cave bear record and they are mainly located in the southern half of the peninsula. The oldest evidence in the Iberian Peninsula is from c. 250 ka, which is 300 ka younger than the oldest European records (˜550 ka; L’Arago). Here we present the brown bear fossil assemblage from Postes cave (Fuentes de León, Extremadura, southern Iberian Peninsula). Postes cave has yielded some of the oldest evidence for brown bears in Iberia with a minimum date of 244 191±2261 a BP (number of identified specimens (NISP) = 4). Additionally, the uppermost part of the Pleistocene deposit has yielded one of the largest Middle Pleistocene brown bear collections in the Iberian Peninsula (NISP = 73) with a chronology roughly between 193 and 244 ka BP, comprising both cranial and postcranial remains and including a complete hemimandible. The Postes mandible fits well within the Middle Pleistocene brown bear range of variation, as it is significantly different in size and shape from the Holocene brown bears, based on geometric morphometric analyses. We show that these differences are due to allometry. This site provides more insight into the Middle Pleistocene morphological variation of brown bears in western Europe and underscores the need for additional Middle Pleistocene fossils in the northern half of Iberia to further test palaeobiogeographical hypotheses of this species’ entrance into the largest of the southern European peninsulas.
... Therefore, U. spelaeus remains within the investigated sequence cannot be used as a precise stratigraphic marker. The earliest appearance of the U. spelaeus dates back to the end of the Middle-Late Pleistocene transition, and it became extinct in central Europe during the LGM, around 24 ka BP (Kurtén, 1958(Kurtén, , 1968Pacher and Stuart, 2009). Within Dinaric karst there are dated cave bear specific sites (e.g., Križna jama) with ages of cave bear thanatocenoses around 47-45 ka and >94 ka BP (Bosák et al., 2012). ...
During archeological excavations in the Lower Cerovačka Cave (Mt. Velebit, Croatia), the test trench penetrated to a depth of 1.8 m. An undisturbed sequence of sediments was exposed. Considering that caves represent highly efficient sediment traps it was possible to recognize changes in the depositional mechanisms during the Pleistocene–Holocene period. Using the multiproxy approach, the mineralogical, petrographic, and biostratigraphic characterization of the cave sediments was performed. Facies analysis revealed several stages in the development of the clastic filling of cave channels. Allochthonous origin of the sediment was assumed. Sedimentation took place under various conditions from pronounced cold and dry climate during Pleistocene stages in the base of the profile, to humid periods with anthropogenic influence during the Holocene at the very top of the profile. Although traditionally these sediments were believed to be of a Pleistocene age, here for the first time a stratigraphic calibration of the profile has been performed based on luminescence dating of detrital cave sediments and radiometric dating of speleothems.
... During the Late Pleistocene, large mammal species, including cave bear (Ursus spelaeus), cave lion (Panthera spelaea), and woolly rhinoceros (Coelodonta antiquitatis) went extinct in northern Eurasia (Pacher and Stuart, 2009;Lister 2011, 2012). Other Pleistocene megafauna, including giant deer (Megaloceros giganteus) and woolly mammoth (Mammuthus primigenius), experienced strong reductions in their distributions during the Late Pleistocene, but only disappeared from the faunal record later, during the Holocene (Stuart et al., 2002;Lister and Stuart, 2019). ...
The woolly rhinoceros (Coelodonta antiquitatis) was a cold-adapted herbivore, widely distributed from western Europe to north-east Siberia during the Late Pleistocene. Previous studies have associated the extinction of the species ∼14,000 calendar years before present to climatic and vegetational changes, suggesting the later survival of populations in north-east Siberia may have related to the later persistence of open vegetation in the region. Here, we analyzed carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotopes and mitochondrial DNA sequences to elucidate the evolutionary ecology of the species. Our dataset comprised 286 woolly rhinoceros isotopic records, including 192 unpublished records, from across the species range, dating from >58,600 to 12,135 ¹⁴C years before present (equivalent to 14,040 calendar years ago). Crucially, we present the first 71 isotopic records available to date of the 15,000 years preceding woolly rhinoceros extinction. The data revealed ecological flexibility and geographic variation in woolly rhinoceros stable isotope compositions across time. In north-east Siberia, we detected stability in δ¹⁵N through time, which could reflect long-term environmental stability, and may have enabled the later survival of the species in the region. To further investigate the paleoecology of woolly rhinoceroses, we compared their isotopic compositions with other contemporary herbivores. Our findings suggested isotopic similarities between woolly rhinoceros and both musk ox (Ovibos moschatus) and saiga (Saiga tatarica), albeit at varying points in time, and possible niche partitioning between woolly rhinoceros and both horse (Equus spp.) and woolly mammoth (Mammuthus primigenius). To provide phylogeographic context to the isotopic data, we compiled and analyzed the 61 published mitochondrial control region sequences. The genetic data showed a lack of geographic structuring; we found three haplogroups with overlapping distributions, all of which showed a signal of expansion during the Last Glacial Maximum. Furthermore, our genetic findings support the notion that environmental stability in Siberia influenced the paleoecology of woolly rhinoceroses in the region. Our study highlights the utility of combining stable isotopic records with ancient DNA to advance our knowledge of the evolutionary ecology of past populations and extinct species.
... bear, a species that was apparently reliant on high-quality plant food which made it 471 vulnerable to decreasing vegetational productivity, goes extinct in Central Europe at around 472 28 ka (Pacher and Stuart 2008). whereas populations in Central Europe remain at a low level. ...
... Ursus spelaeus was primarily limited to Europe and may have been an offshoot of early brown bear lineages, which purportedly exhibited metabolic denning and active metabolism similar to modern brown bears and American black bears (Nelson et al. 1998, Fernández-Mosquera 2001. Their specialised vegetative diet (Pacher & Stuart 2009), a reliance on caves as den habitat, climate change (Stiller et al. 2010), and overall decreased ecological plasticity have been postulated to have contributed to speciation and eventual extinction of Ursus spelaeus (Stiller et al. 2010). In Eurasia and North Africa, brown bears underwent multiple range expansions and contractions coinciding with climate events and eventually colonised western North America (Sommer & Benecke 2005). ...
• Through realisation of ecological opportunities, populations and species can experience relaxed selection pressures, facilitating ecological release and leading to rapid speciation and morphological diversification. Behavioural plasticity in response to environmental change contributes to diversification by exposing individuals to novel conditions through their interactions with resources or dispersal to new areas. Despite strong theoretical support, demonstrations of this evolutionary process are rare.
• The family Ursidae is the product of one or more adaptive radiations following the Miocene–Pliocene transition. Denning behaviour associated with over-winter seclusion, fasting, and parturition coinciding with seasonally decreased food availability appears to be paraphyletic in Ursidae phylogenies. However, female bears of all species undergo varying degrees of seclusion and fasting during parturition and early post-natal care, which is not consistent with periods of seasonally decreased food availability. As denning behaviour is tightly linked to fitness through energetics and reproduction, these behaviours are suspected to be under strong selection. Mechanisms responsible for the observed variability among species, populations, and individuals have not been explored.
• In this review, we detail the evolutionary history of extinct and extant Ursidae regarding expression of denning behaviour and as a function of realised ecological opportunities. We compare behaviours across Ursidae and contextualise our results within extant species ecology.
• We demonstrate the role of relaxed extrinsic and intrinsic factors in the expression of metabolic suppression among Ursidae species, and across populations and reproductive groups, through the realisation of ecological opportunities. In doing so, we propose a more refined consideration of and perspective on this behaviour and provide a mechanism for the adaptive radiation in Ursidae.
... Ethological trends of species during the late Pleniglacial in the East European Plain (compiled from Bannikov, 1963;Kurtén B, 1968;Guérin, 1980;Lent, 1988;Kuzmina, 1997;Geist, 1998;van Vuure, 2002;Pucek et al., 2004;Jost and Jost-Tse, 2005;Lister and Bahn, 2007;Pacher and Stuart, 2009;Parrini et al., 2009;Bough, 2012;Julien et al., 2012;Manuel et al., 2020; and Borziac, 2009;Noiret, 2009), the Pontic steppe (Krotova, 1995(Krotova, , 2003(Krotova, , 2013 Romania (Cârciumaru, 1999), Poland (Kozłowski and Kozłowski, 1996;Wilczyński, 2016), the Balkans (Tsanova, 2006) and the Baltic region (Gimbutas, 1956). The Gravettian in Eastern Europe developed from 34.0 to 24.0 ka cal BP (30.0-20.0 ...
The late Pleniglacial was characterized by different palaeoenvironmental and geographical modifications, which affected the diverse living beings that had to adapt to those peculiar conditions. This paper intends to explore how this phenomenon may have impacted the practices of the nomadic human populations who occupied the East European Plain during the last part of the Pleistocene, through their relation to the environments, landscapes, overall territories, and especially with other species in archaeological context. Using both quantitative and qualitative data, we performed an exhaustive review of the faunal record from an extensive set of archaeological recorded contexts ranging between 31.0 and 21.5 ka cal BP (26.0-18.0 ka ¹⁴C BP). We examined the seasons of occupations and the types of activities at each site, the global subsistence economies, as well as the technocultural practices and settlement patterns of the human groups. This research shed new light on the connections amongst different human groups and between humans and other animals, the socio-ecological systems that have been favored during this critical period of human prehistory in the East European Plain.
The first data on the content of isotopes 13C and 15N in the collagen of 16 bones of the Ural cave bear (Ursus (Spelaearctos) kanivetz Verestchagin, 1973) were obtained from the Tayn (Secrets) cave (55°25' N, 57°46' E). The bones date from the middle of MIS 3. The bones of males and females aged about 2 years, about 3 years and older than 4 years were studied. There are no noticeable differences in isotope signatures between individuals of different ages and different sexes. Since the second year of life, cave bears have been eating plant food on their own. The values of δ13C and δ15N of the Ural cave bear are close to the values for U. (S.) spelaeus ingressus.
Learning plays an important role in the ecology and evolution of species. Although learning has been extensively studied in extant populations, studying learning in long extinct species represents a much greater challenge because individuals and thei behaviour cannot be directly observed, and their individual relatedness is more challenging to study using molecular methods. However, an interdisciplinary approach combining dietary behaviour revealed by isotope analysis and relatedness estimated using ancient DNA might offer new insights into the learning behavior of extinct species and populations. In this study, we examine dietary variation in Late Pleistocene cave bears from the Romanian Carpathians using stable isotopes. We find that dietary variation among these cave bears is not associated with genetic relatedness inferred using mitochondrial DNA, nor is it associated with Late Pleistocene large scale-climatic oscillations, since different dietary niches existed contemporaneously. Individual cave bears within the same population therefore appear to have had different dietary preferences. This implies that dietary preference was not determined at birth, and instead developed during the early life stages of the animal, most likely facilitated through learning which was not necessarily acquired from close kin. Combining diet inferred from stable isotopes with genetic relationships inferred from ancient mitogenomes thus reveals aspects of the behavioral ecology of an extinct Pleistocene animal with an unprecedented level of detail.
The 13C and 15N isotope contents in bone collagen were analyzed using bones of the small cave bear Ursus (Spelaearctos) rossicus Borissak, 1930 from localities in the Middle and Southern Urals. The bones date from the last interglacial (MIS 5) and glacial (MIS 3) periods. The bones were from males and females aged 3, 4, and >4 years. Sexual, geographical, and chronological differences in 13C and 15N contents were studied. Notable gender, geographic, and chronological differences were observed between samples. In the Middle Urals, females led a more predatory lifestyle than males during the interglacial period, and the trophic niches of males and females converged due to an increase in herbivory during the transition to the glacial period. In the Southern Urals, males led a more predatory lifestyle than in the Middle Urals during the interglacial period. The extent of changes in δ13C and δ15N values in the Southern Urals during the transition was found to correspond to differences between trophic levels.
First data on the contents of the 13C and 15N isotopes in collagen were obtained for 16 bones of the Ural cave bear Ursus (Spelaearctos) kanivetz Verestchagin, 1973 from the Tayn (Secrets) cave (55°25' N, 57°46' E). The bones are dated to the middle MIS 3 and belonged to males and females of about 2 years, about 3 years, and older than 4 years of age. No considerable difference in isotope signatures was observed between individuals of different ages and different genders. Cave bears were assumed to forage independently on plant food from the second year of life. The δ13C and δ15N values established for the Ural cave bear are close to the values reported for U. (S.) spelaeus ingressus.
In the last decade, the identification of bone fragments by peptide mass fingerprinting or zooarchaeology by mass spectrometry is developing as a powerful tool in Quaternary palaeontology. The sequence of amino acids that make up the bone collagen molecule shows slight variations between taxa, which can be studied by mass spectrometry for taxonomic purposes. This requires reference databases that allow peptide identification. Although the cave bear ( Ursus spelaeus Rosenmüller, 1794) is a common component in many European Pleistocene cave sites, no peptide fingerprint taxonomic study has paid special attention to this species up to now. For peptide markers in Ursidae, the most recent proposal is based on collagen obtained from a modern brown bear sample. In this work we attempt to cover this gap by studying bone collagen of cave and brown bear samples from different origins and different chronology, applying matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry (MALDI TOF). We also performed an in-silico study of ursid bone collagen sequences published in databases. In our results we detected some discrepancies between the peptides obtained from both in silico and MALDI TOF analysis of fossil collagen and those published in the literature, in which we conclude that there are some misidentified peptides. The identification of skeletal remains by means of their peptide fingerprint is proving to be a powerful tool in palaeontology, which will bear greater fruit once the limitations of a technique that is in its initial stages have been overcome.
Исследована ископаемая фауна мелких млекопитающих из отложений пещеры Иманай (53°02′ с.ш., 56°26′ в.д.) (Южный Урал, Россия). В фауне преобладают виды открытых биотопов, узкочерепная полевка (Microtus (Stenocranius) gregalis) является доминирующим видом. Выявлено два типа фауны, которые характеризуют ее состав и структуру в конце позднего плейстоцена и, предположительно, в раннем голоцене. Содоминирующим видом фауны первого типа (нижние и средние отложения пещеры) является степная пеструшка (Lagurus lagurus). Для этого типа характерны значительное преобладание доли (83.0–92.2%) степных видов и низкая доля (1.5–3.9%) лесных видов млекопитающих, что является доказательством распространения в позднеледниковье на территории южной
части Уральских гор преимущественно открытых ландшафтов. В фауне второго типа (верхние отложения пещеры) содоминируют пищухи (Ochotona sp.) и относительно высока доля лесных видов
(14.3–21.4%), что свидетельствует о появлении лесных формаций в конце позднеледниковья или в раннем голоцене на этой территории. Выборки первых нижнекоренных зубов узкочерепных полевок характеризуются высокой долей зубов (>50%) с простыми вариантами строения непарной петли антероконидного отдела (“грегалоидный” морфотип). В выборках зубов степных пеструшек зафиксирована высокая доля (до 51.6%) зубов “транзиенсногo” морфотипа. Среди зубов в нижней
части отложений пещеры обнаружены бескорневые цементные первые нижнекоренные зубы (m1) и третьи верхнекоренные зубы (M3), строение жевательной поверхности которых (широкое слияние треугольников Т4-Т5 и Т2-Т3, соответственно) характерно для древних полевок Microtus (Stenocranius) gregaloides и M. (Terricola) arvalidens из фаун первой половины раннего плейстоцена–второй половины среднего плейстоцена.
New data were obtained for the Chulym River basin in the southeastern part of the West Siberian Plain, one of the understudied parts of Siberia in terms of age and composition of carbon and nitrogen stable isotopes for Late Pleistocene megafauna. The 14C dates from the Sergeevo outcrop, the most complete section of Late Quaternary deposits in the region, are mostly greater than ~30 550 bp. Other localities yielded 14C values in the range from >44 500 to ~19 300 bp. The finite date of ~42 270 bp for the Khozarian steppe elephant (Mammuthus trogontherii chosaricus) from Asino is intriguing because previously it was not detected in the Late Pleistocene of Siberia after the last interglacial (Marine Isotope Stage 5e), ~115 000–130 000 years ago. Stable isotope data show both similarities and differences compared to the pre‐Last Glacial Maximum megafaunal species in other parts of Siberia.
The Sima del Elefante site is located within the Sierra de Atapuerca karst system (Burgos, northern Spain), and forms part of a series of important Early, Middle and Late Pleistocene archaeological complexes that have been dated previously with luminescence techniques (Gran Dolina, Galería Complex, Sima de los Huesos, Galería de las Estatuas). This study focuses on the upper Middle Pleistocene units (TE18 and TE19) at Sima del Elefante, which contain Acheulean and transitional lithic assemblages (Mode 2/3), as well as large and small mammal fossils. Importantly, these uppermost units are associated with a sediment plug located in the cave's interior at Galería Baja, which marks the closure of a significant palaeoentrance to the Atapuerca karst system. Establishing the accumulation history of these related deposits is important for understanding both Lower Palaeolithic technological dynamics via comparisons with similar levels at other Atapuerca sites (i.e., Gran Dolina and Galería Complex), as well as past human occupation patterns and carnivore use of (and accessibility to) the caves. We present single-grain TT-OSL and multi-grain pIR-IR chronologies for the Sima del Elefante upper sequence and the Galería Baja sediment cone, as well as U-series dating results for a stalagmitic crust capping the combined clastic infill sequence. The paired luminescence ages for the upper occupation levels are in agreement with each other and reveal that the host deposits accumulated 576–481 ka for the TE18 stratified scree layers, 266–237 ka for TE19 and 206–250 ka for the Galería Baja upper cone section (weighted mean 2σ age ranges). A concordant U-series age of 202 ± 65 ka is obtained for the overlying stalagmitic crust at Galería Baja. The Sima del Elefante ages are consistent with those previously obtained using U-series and biochronology, confirming that there is an erosional unconformity and complex carbonate deposition phase associated with the upper layers of unit TE18 and that the original cave entrance likely closed by ∼200 ka. Chronological correlation with other Atapuerca sites reveals potential equivalence between TE18 and unit TD8 (at Gran Dolina), and between TE19 and units GIIb–GIIIb (at Galería Complex) and, possibly, TD10.1 and TD10.2 (at Gran Dolina), though more refined dating is required to confirm the latter.
The Szeleta cave was the first excavated Palaeolithic site in Hungary. The artefacts were scattered along the layer sequence and very few of them were excavated in thin, discrete layers. After the general review of the available documentation and some questions of site formation, four assemblages are analysed in the present paper, all excavated in the so-called hearth levels in the entrance and the main hall of the cave. The variability of the used raw materials and the typological differences makes possible to describe several types of industries, but none of these was a typical Szeletian assemblage.
The lower molar (m1) of cave bears from Late Pleistocene localities of the Urals was studied employing the methods of traditional morphometry and geometric morphometrics. On the basis of the size and shape variation of m1, the small cave bear (Ursus ex gr. savini-rossicus) was found to have been a part of the faunas from the caves Skazka, Viasher, Dynamitnaya, Chudesnitsa, and Chernye Kosti. The small cave bear presence in faunas from the Medvezhya, Makhnevskaya Ledyanaya, Asha 1, Ignat'evskaya, and Barsuchii Dol caves was confirmed as well. The species range of the small cave bear encompassed the Northern, Middle, and Southern Urals in the Late Pleistocene. The ranges of the small cave bear and cave bear (Ursus kanivetz) overlapped from the beginning (marine isotope stage 5e) to the middle (middle marine isotope stage 3) of the Late Pleistocene.
Starting four decades ago, studies have examined the ecology and evolutionary dynamics of populations and species using short mitochondrial DNA fragments and stable isotopes. Through technological and analytical advances, the methods and biomolecules at our disposal have increased significantly to now include lipids, whole genomes, proteomes, and even epigenomes. At an unprecedented resolution, the study of ancient biomolecules has made it possible for us to disentangle the complex processes that shaped the ancient faunal diversity across millennia, with the potential to aid in implicating probable causes of species extinction and how humans impacted the genetics and ecology of wild and domestic species. However, even now, few studies explore interdisciplinary biomolecular approaches to reveal ancient faunal diversity dynamics in relation to environmental and anthropogenic impact. This review will approach how biomolecules have been implemented in a broad variety of topics and species, from the extinct Pleistocene megafauna to ancient wild and domestic stocks, as well as how their future use has the potential to offer an enhanced understanding of drivers of past faunal diversity on Earth.
Before the end of the Last Glacial Maximum (LGM, ∼16.5 ka ago)1 set in motion major shifts in human culture and population structure,2 a consistent change in lithic technology, material culture, settlement pattern, and adaptive strategies is recorded in Southern Europe at ∼18-17 ka ago. In this time frame, the landscape of Northeastern Italy changed considerably, and the retreat of glaciers allowed hunter-gatherers to gradually recolonize the Alps.3-6 Change within this renewed cultural frame (i.e., during the Late Epigravettian phase) is currently associated with migrations favored by warmer climate linked to the Bølling-Allerød onset (14.7 ka ago),7-11 which replaced earlier genetic lineages with ancestry found in an individual who lived ∼14 ka ago at Riparo Villabruna, Italy, and shared among different contexts (Villabruna Cluster).9 Nevertheless, these dynamics and their chronology are still far from being disentangled due to fragmentary evidence for long-distance interactions across Europe.12 Here, we generate new genomic data from a human mandible uncovered at Riparo Tagliente (Veneto, Italy), which we directly dated to 16,980-16,510 cal BP (2σ). This individual, affected by focal osseous dysplasia, is genetically affine to the Villabruna Cluster. Our results therefore backdate by at least 3 ka the diffusion in Southern Europe of a genetic component linked to Balkan/Anatolian refugia, previously believed to have spread during the later Bølling/Allerød event. In light of the new genetic evidence, this population replacement chronologically coincides with the very emergence of major cultural transitions in Southern and Western Europe.
We report a new series of radiocarbon ( ¹⁴ C) dates on the MIS 3 megafauna for a previously poorly studied region of southeastern West Siberia. Some species, like woolly mammoth and woolly rhinoceros, and Pleistocene bison and horse, existed throughout the MIS 3 (ca. 29–59 ka cal BP); cave hyaena is dated to ca. 46,400 cal BP. The very late ¹⁴ C dates on Khozarian steppe elephant ( Mammuthus trogontherii chosaricus ), ca. 45,100–45,400 cal BP, may indicate the survival of this species in Siberia up to MIS 3. More work is needed to confirm or reject this suggestion. Previously, Khozarian steppe elephant was known in Siberia only at the beginning of the Late Pleistocene (MIS 5e).
The sedimentological record in the Račiška pečina cave sediment sequence is one of the best-preserved cave records of palaeoenvironmental changes for the last 3.4 Ma. However, as it is typical for cave terrestrial records, it contains many hiatuses in sedimentation. The section study helped to change the state of knowledge and understanding of the long-lasting deposition characteristics in the caves and provided enormous data on environmental changes over time. In the sequence are by magnetostratigraphy well recorded Pliocene/Pleistocene transition at 2.59 Ma, the Matuyama/Brunhes boundary at 0.773 Ma, and the presence of Olduvai subchron between 1.78 and 1.925 Ma. Records of small mammals from the lower part of the section (a molar of Apodemus cf. atavus and dental fragments of Borsodia sp., and Pliomys sp.) suggest MN17 age, Clethrionomys cf. glareolus from the upper part suggests the Late Early or Middle Pleistocene age. Also worth mentioning are records of snail shells Aegopinella sp. and a troglobiont snail Zospeum sp. In the upper part of the section Ursus ex gr. spelaeus was confirmed in the yellow clay layer older than ∼72 ka, and soot material at the top of the section was radiocarbon dated on ∼11 ka, ∼9 ka, and ∼3 ka. A detailed chronology of the Račiška pečina section based on magnetostratigraphy and isotopic oxygen stratigraphy was created and correlated with palaeontological, U-series, and radiocarbon results. The climatic changes during the growth of the section were at about 2.6–2.5 Ma ago mostly controlled by global Atlantic Ocean factors, while about 0.78 Ma ago by regional Mediterranean Sea factors.
Paleolithic antiquity of parietal art in Ignatievskaya cave, Southern Ural, is supported by its subject (Late Pleistocene animals) as well as by paleontological and palynological data, and 14C dates from cultural layers associated with artistic activity (17.8−16.3 cal ka BP; association is established by finds of ochre in these layers). However, three 14C dates of charcoal motifs yielded younger, Holocene ages (7.4−6.0 cal ka BP). In this study, we constrain the age of parietal art in the cave by 230Th dating of flowstone that brackets the paintings. Flowstone did not form in the cave between c. 78 and 10 ka BP, due to widespread permafrost in northern Eurasia at that time. Our 230Th dates do not support the middle Holocene age of art in Ignatievskaya cave and are consistent with its Upper Paleolithic antiquity instead.
Krk Island is one of the largest Croatian islands, occupying an area of 405.5 km².
Stable isotopes of mammoths and mastodons have the potential to illuminate ecological changes in late Pleistocene landscapes and megafaunal populations as these species approached extinction. The ecological factors at play in this extinction remain unresolved, but isotopes of bone collagen (δ ¹³ C, δ ¹⁵ N) and tooth enamel (δ ¹³ C, δ ¹⁸ O, ⁸⁷ Sr/ ⁸⁶ Sr) from midwestern North America are leveraged to examine ecological and behavioral changes that occurred during the last interglacial-glacial cycle. Both species had significant C3 contributions to their diets and experienced increasing levels of niche overlap as they approached extinction. A subset of mastodons after the last glacial maximum exhibit low δ ¹⁵ N values that may represent expansion into a novel ecological niche, perhaps densely occupied by other herbivores. Stable isotopes from serial and microsampled enamel show increasing seasonality and decreasing temperatures as mammoths transitioned from Marine Isotope Stage (MIS) 5e to glacial conditions (MIS 4, MIS 3, MIS 2). Isotopic variability in enamel suggests mobility patterns and life histories have potentially large impacts on the interpretation of their stable isotope ecology. This study further refines the ecology of midwestern mammoths and mastodons demonstrating increasing seasonality and niche overlap as they responded to landscape changes in the final millennia before extinction.
Americas Redefining the Age of Clovis: Implications for the Peopling of the This copy is for your personal, non-commercial use only. clicking here. colleagues, clients, or customers by , you can order high-quality copies for your If you wish to distribute this article to others here. following the guidelines can be obtained by Permission to republish or repurpose articles or portions of articles): July 7, 2014 www.sciencemag.org (this information is current as of The following resources related to this article are available online at
p>Sex ratios of cave bears (Ursus spelaeus and U. deningeri) are known to vary greatly between sites. A number of explanations have been proposed to account for this variability, involving for example cave size, altitude, and the seasonality of food availability. This contribution presents demographic data from three sites in southern Germany which allow the rejection of these models; the factor(s) behind the variability in sex ratios is (are) still unknown. The data also suggest that, contrary to the case in living brown bears (U. arctos), males in cave bears may have reached a very old age more frequently than females.</p
In a recent INQUA project the event of Pleistocene glaciations has been digitally mapped and the chronology of events reviewed. The onset of the present Ice Age in both hemispheres dates back to the Palaeogene. In Greenland, Iceland, North America and southernmost South America Sizeable ice sheets formed well before 2.6 ka BP hi Alaska and on Tierra del Fuego the ice advanced further than in any later glaciations. Evidence for Early Pleistocene glaciation (2.6-0.78 Ma) has been reported from many parts of the world, but in most cases dating remains problematic, and the size of the glaciers and ice sheets is unknown. A number of Middle Pleistocene glaciations (0.78-0.13 Ma) have beet? identified, mostly correlated with MIS 16, 12 and 6, including the Donian, Elsterian and Saalian of Europe. The extent Of the MIS 6 glaciations is well known. Especially in. Eurasia the e-vent of the Late Pleistocene (0.13 Ma to pref sent) glaciations had to be revised. Major ice advances are reported for 80-100 ka BP c. 70-80 ka BP and c. 20 ka BP, with the earlier glaciations being most extensive in the east. The very different shapes of the ice sheets-Donian vs Elsterian, Early vs Late Weichselian-are as Yet difficult to explain and remain a challenge for climatic modellers.
The cave near Nerubajskoe village is one of several karstic elements in this area. Some of these caves contain Pleistocene material. NORDMANN (1849) was the first to excavate in this area. Only a few new sites have been found since then. The Austrian-Ukrainian research group recovered new Pleistocene material from a cave ruin near Nerubajskoe in 2003. The cave bear belongs to the U. ingressus group with plesiomorphic morphological features. While morphological different from the modern cave bear from the Alpine region, the results of the fossil DNA analyses clearly places them in the same evolutionary line as the findings from the Gamssulzen and Potočka Cave. New Radiocarbon dates place the fauna into the middle part of the Upper Pleistocene.
Sex-ratio determination is essential for palaeontological and palaeoethological researches on fossil taxa, especially for cave bear which shows an important sexual dimorphism. “Classical” methods use uni- and bivariate plots to fix an arbitrary limit between sexes. Therefore, a more robust statistical method, termed “mixture analysis”, has been applied in this study to determine the sex-ratio of the cave bear from Schwabenreith, Gamssulzen (Austrian Alps), Fate, Basura and Badalucco (northwest of Italy). This method has been previously calibrated on the post-cranial material of cave bear from Fate (QUILES & MONCHOT 2004). In order to define sexual dimorphism for the whole skeleton, seven elements (lower canine, lower and upper second molars, scapholunar, navicular, metacarpal 3 and metatarsal 3) have been selected for their reliability and their abundance in studied assemblages. A quantified comparison between results of bivariate plots and mixture analysis highlights the better reliability and rigor of the new method. The latter, however, also offers a standardized and replicable procedure which provides a calculated cut-off point and a misclassification error for each dimension, permitting the analyst to determine precise sexual groups as well as identifying those specimens that cannot be sexed. Cheek teeth can be sexed as reliably as lower canine and post-cranial elements, providing statistically equivalent results. Lower canine, second upper molar and scapholunar are the most effective bones for determining sex-ratio in cave bear.
The 19981999 direct dating of two Neandertal specimens from level G1 of Vindija Cave in Croatia to 28,000 and 29,000 radiocarbon (14 C) years ago has led to interpretations concerning the late survival of Neandertals in south-central Europe, patterns of interaction between Neandertals and in-dispersing early modern humans in Europe, and complex biocultural scenarios for the earlier phases of the Upper Paleolithic. Given improvements, particularly in sample pretreatment techniques for bone radiocarbon samples, especially ultrafiltration of collagen samples, these Vindija G 1 Neandertal fossils are redated to 32,000 –33,000 14 C years ago and possibly earlier. These results and the recent redat-ing of a number of purportedly old modern human skeletal remains in Europe to younger time periods highlight the importance of fine chronological control when studying this biocultural time period and the tenuous nature of monolithic scenarios for the establishment of modern humans and earlier phases of the Upper Paleolithic in Europe.
The Dupont collections curated at the Royal Belgian Institute of Natural Sciences contain a large bone assemblage from Chamber B, horizon 4 of the third cave of Goyet (Belgium). This assemblage dates from the Weichselian (AMS date: 35.470 + 780-710 years BP). Ninety eight percent of the remains are from Ursus spelaeus. The presence of several skeletons and articulated body parts, and the low frequency of gnawing traces and charriage-à-sec indicate that the carcasses were only slightly disturbed. Based on the eruption sequence of the teeth, cubs perished at or soon after birth, at an age of two and a half months and at an age of six-seven months old, when they were leaving the cave at the end of the winter rest. In Belgium, the timing and length of the dormancy period and the birth season varied according to the climatic conditions of the ice age. At Goyet, Chamber B, horizon 4, female cave bears probably began their winter rest in October, gave birth in November or December and left the cave in June.
The family Ursidae appears to be a sensitive mammal group, promptly reflecting large but also
small scale changes in global environmental conditions. Although the Greek ursid record is extremely incomplete,
especially regarding the late Miocene-early Pliocene period, it is evident that the main evolutionary
and ecological trends of the family are roughly depicted. The presence of Ursidae in Greece, and the
southern Balkans in general, is highly controlled by palaeoecological factors concerning primarily climatic
and vegetational changes.
I examined the relationship of diets to skull morphology of extant northern bears and used this information to speculate on diets of the recently extinct cave (Ursus spelaeus) and short-faced (Arctodus simus) bears. Analyses relied upon published skull measurements and food habits of Asiatic (U. thibetanus) and American (U. americanus) black bears, polar bears (U. maritimus), various subspecies of brown bears (U. arctos), and the giant panda (Ailuropoda melanoleuca). Principal components analysis showed major trends in skull morphology related to size, crushing force, and snout shape. Giant pandas, short-faced bears, cave bears, and polar bears exhibited extreme features along these gradients. Diets of brown bears in colder, often non-forested environments were distinguished by large volumes of roots, foliage, and vertebrates, while diets of the 2 black bear species and brown bears occupying broadleaf forests contained greater volumes of mast and invertebrates and overlapped considerably. Fractions of fibrous foods in feces (foliage and roots) were strongly related to skull morphology (R2 = 0.97). Based on this relationship, feces of cave and short-faced bears were predicted to consist almost wholly of foliage, roots, or both. I hypothesized that cave bears specialized in root grubbing. In contrast, based upon body proportions and features of the ursid digestive tract, I hypothesized that skull features associated with crushing force facilitated a carnivorous rather than herbivorous diet for short-faced bears.
The volume contains summaries of facts, theories, and unsolved problems pertaining to the unexplained extinction of dozens of genera of mostly large terrestrial mammals, which occurred ca. 13,000 calendar years ago in North America and about 1,000 years later in South America. Another equally mysterious wave of extinctions affected large Caribbean islands around 5,000 years ago. The coupling of these extinctions with the earliest appearance of human beings has led to the suggestion that foraging humans are to blame, although major climatic shifts were also taking place in the Americas during some of the extinctions. The last published volume with similar (but not identical) themes -- Extinctions in Near Time-- appeared in 1999; since then a great deal of innovative, exciting new research has been done but has not yet been compiled and summarized. Different chapters in this volume provide in-depth resumés of the chronology of the extinctions in North and South America, the possible insights into animal ecology provided by studies of stable isotopes and anatomical/physiological characteristics such as growth increments in mammoth and mastodont tusks, the clues from taphonomic research about large-mammal biology, the applications of dating methods to the extinctions debate, and archeological controversies concerning human hunting of large mammals.
As recently as 11,000 years ago "near time" to geologists mammoths, mastodons, gomphotheres, ground sloths, giant armadillos, native camels and horses, the dire wolf, and many other large mammals roamed North America. In what has become one of science's greatest riddles, these large animals vanished in North and South America around the time humans arrived at the end of the last great ice age. Part paleontological adventure and part memoir, "Twilight of the Mammoths "presents in detail internationally renowned paleoecologist Paul Martin's widely discussed and debated "overkill" hypothesis to explain these mysterious megafauna extinctions. Taking us from Rampart Cave in the Grand Canyon, where he finds himself "chest deep in sloth dung," to other important fossil sites in Arizona and Chile, Martin's engaging book, written for a wide audience, uncovers our rich evolutionary legacy and shows why he has come to believe that the earliest Americans literally hunted these animals to death. As he discusses the discoveries that brought him to this hypothesis, Martin relates many colorful stories and gives a rich overview of the field of paleontology as well as his own fascinating career. He explores the ramifications of the overkill hypothesis for similar extinctions worldwide and examines other explanations for the extinctions, including climate change. Martin's visionary thinking about our missing megafauna offers inspiration and a challenge for today's conservation efforts as he speculates on what we might do to remedy this situation both in our thinking about what is "natural" and in the natural world itself."
Many forms are represented only by isolated bone fragments, and comparatively little has been accomplished in the way of functional morphology and detailed description of the extinct beasts. Consequently interpretations are speculative and subject to revision. Observations imply that Australian fauna was (and still is) less able to withstand the impact of ecological change in the presence of man, even with the small numbers of men that were present 50 000-30 000 yr ago. -from Author
During and after the last glaciation (i.e., during the last 100,000 years), many large-bodied vertebrate species (the so-called Pleistocene megafauna) died out. This megafaunal extinction reduced by more than half the number of large-bodied mammals in the world, including large herbivores, predators, and scavengers. Australia lost 86% of its Pleistocene megafauna, mostly between 50,000 and 40,000 years ago, about the time that people arrived there. South America lost 80% of its megafauna, mostly during the late glacial interval. North America lost 73% of its megafauna, also during the late glacial interval. The timing of New World human habitation is controversial, but the most widely accepted view is that most of the New World megafaunal extinctions took place just as people arrived in the Americas.
A harvest of 300 radiocarbon dates on extinct elephants (Proboscidea) from the northern parts of the New and Old Worlds has revealed a striking difference. While catastrophic in North America, elephant extinction was gradual in Eurasia (Stuart 1991), where straight-tusked elephants ( Palaeoloxodon antiquus ) vanished 50 millennia or more before woolly mammoths ( Mammuthus primigenius ). The range of the woolly mammoths started shrinking before 20 ka ago (Vartanyan et al. 1995). By 12 ka bp, the beasts were very scarce or absent in western Europe. Until the dating of Wrangel Island tusks and teeth (Vartanyan, Garrutt and Sher 1993), mammoths appeared to make their last stand on the Arctic coast of Siberia ca. 10 ka bp. The Wrangel Island find of dwarf mammoths by Sergy Vartanyan, V. E. Garrut and Andrei Sher (1993) stretched the extinction chronology of mammoths another 6 ka, into the time of the pharaohs.
This list contains most of the unpublished radiocarbon dates measured between 1961 and 1968. Analyses of the C 14 content of groundwater are excluded and a number of geologic dates will be included in a later list. Certain earlier analyses performed by H. de Vries and H. de Waard are included, as well as a few results obtained since W. G. Mook assumed responsibility for the laboratory in 1969, to complete series. Descriptions and comments are mainly based on information supplied by the submitters or contained in the publications cited. However, our own interpretation is often given in the comments. Sections I A to D2, II B1, and B2 were mainly prepared by the first author and the rest by the second.
Recently, an article was published in this journal, discussing evidence for a solar flare cause of faunal extinction during the Late Pleistocene (LaViolette 2011). The article is based on the hypothesis that an increase in atmospheric radiocarbon concentration might have been produced by a giant solar proton event (SPE). This proposed SPE would deliver a lethal radiation dose of at least 3–6 Sv to the surface of the Earth, causing termination of the Pleistocene megafauna.
The cave discovered in 1996 in the Laubenstein area (Chiemgau / Bavarian Alps) contained mainly rests of Ursus spelaeus and Ursus arctos. This cave is the first alpine cave bear bearing cave in Germany. A test pit was dug and numerous special analyses, for example collagen isotopy and absolute datations, were done on the bones. The collagen isotopy confirms the results of other research groups. The datations place the cave bear occupation of the cave in the Hengelo-Denekamp Interstadial complex (OIS 3).
This chapter presents last occurrence dates and associated data for 255 extinct mammal species that are currently known or suspected to have died out during the Holocene. This includes species known to have survived into the historical period, and those represented in Late Quaternary subfossil deposits known or believed to be Holocene in age, or from island systems which humans did not reach until the Holocene.
The lineage of the cave bear Ursus deningeri --> U. spelaeus is well known, but to draw a limit in this evolution is not easy. The real difficulty is met when this cave bear lineage has to be linked to its ancestors... Dating physical methods, paleoclimatic and paleo environmental data, works on teeth and bones morphology, progress in paleogenetics are means our disposal to give essential information to phylogeny. A large collaboration is needed. Practical information are given about a possible discussion group.
The late Pleistocene vertebrate record of Africa contributes to our understanding of human evolution and the development of modern biotic environments. This article explores this record, paying particular attention to terrestrial mammals. Because the origins of physically and behaviorally modern humans occurred in Africa during the late Pleistocene, much of what we know about vertebrates from this time comes from archaeological research. Most of the mammal species found in Africa during the late Pleistocene were also found here historically; only a few African species went extinct during the late Pleistocene. However, many changes in species’ ranges, abundance, and associations made late Pleistocene faunal communities look different from today's communities. As climate changed, so did vegetation, creating unique habitats, many of which do not exist today. Each species’ unique responses to these changes caused the formation of mammal communities that do not have analogs today, meaning that species are often found together in fossil assemblages that were not found together historically. Often fossil species are found in places where today their descendants live hundreds of kilometers away, indicating that their past ranges had either shifted or expanded.
More than 12,000 years ago, in one of the greatest triumphs of prehistory, humans colonized North America, a continent that was then truly a new world. Just when and how they did so has been one of the most perplexing and controversial questions in archaeology. This dazzling, cutting-edge synthesis, written for a wide audience by an archaeologist who has long been at the center of these debates, tells the scientific story of the first Americans: where they came from, when they arrived, and how they met the challenges of moving across the vast, unknown landscapes of Ice Age North America. David J. Meltzer pulls together the latest ideas from archaeology, geology, linguistics, skeletal biology, genetics, and other fields to trace the breakthroughs that have revolutionized our understanding in recent years. Among many other topics, he explores disputes over the hemisphere's oldest and most controversial sites and considers how the first Americans coped with changing global climates. He also confronts some radical claims: that the Americas were colonized from Europe or that a crashing comet obliterated the Pleistocene megafauna. Full of entertaining descriptions of on-site encounters, personalities, and controversies, this is a compelling behind-the-scenes account of how science is illuminating our past.
Huge numbers of prehistoric vertebrate extinctions and large-scale range contractions have been documented throughout the Holocene. Evidence for direct human involvement in these extinctions and population shifts is not confounded by other factors and remains relatively undisputed. The Holocene has the potential to act as an ideal study system for investigating the long-term dynamics of anthropogenically mediated extinctions at a global scale, but it remains uncertain whether most prehistoric Holocene extinction events occurred as a result of direct overkill or indirect factors such as habitat destruction. This chapter reviews data on global patterns of mammal and bird species extinctions to provide an assessment of patterns of prehistoric human impact across space and time since the end of the last glaciation. Whereas continental mammals and bird extinctions were relatively minor in comparison to Late Pleistocene megafaunal extinctions, insular faunas have experienced massive-scale extinction events of varying complexity over the past few thousand years.