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The first directly dated cave bear from the Covoli di Velo Cave (Verona Province, Veneto, northern Italy) with some discussion of Italian Alps cave bears
Abstract
Absolute dates of cave bears from Northern Italy are rare. The first radiocarbon date from Covoli di Velo Cave (Verona Province, Veneto, N. Italy) from a cave bear first phalanx is reported; its value is 29,130 ± 0.90 14C yr BP. The date, combined with morphological features of dentition suggest that cave bear populations that lived in Northern Italy were relatively underived compared to other European populations, hinting at patters of migration. Comparison of dental morphology suggest that the Covoli di Velo bear is Ursus spelaeus.
... However, cave bear bones and teeth are not only found in caves, as the name suggests, but also at open air sites in the lowlands, such as Bobenheim-Roxheim in the Upper Rhine Graben. After determining the distinct species of the bear remains, usually radiocarbon dating is applied (Döppes et al., 2016;Rossi et al., 2018;Döppes et al., 2019). When dealing with radiocarbon ages, a closer look at the radiocarbon section of this paper will be helpful as this provides hints on how radiocarbon data should be presented or interpreted. ...
Radiocarbon dating is generally the first choice to date younger Upper Pleistocene organic material. In caves, skeletal remains of animals or humans are often the only preserved datable finds. Routinely, collagen is extracted from bones for radiocarbon dating as it does not readily exchange carbon with its surrounding. However, collagen can also deteriorate and is sometimes not at all or only insufficiently preserved. Apart from the age determination, dietary habits are of special interest and are reconstructed using the stable isotopes of carbon (δ13C) and nitrogen (δ15N). We describe the major challenges of radiocarbon dating and stable isotope analysis of cave bear remains from the Austrian Alps, Franconia (Germany) and fossil finds at a site near Bobenheim-Roxheim (Germany), but also the informative potential of successful application of the methods. We intent to sharpen the knowledge what to look out for to be able to obtain reliable ages of cave bear and other bone material. Moreover, we present two additional scientific methods, luminescence dating and dendroecology as further possibilities to disclose information about sites, their chronology and climatic conditions.
... In order to establish the correct relationship among cave bears, ge netic analyses have been recently addressed by morphometric ana lyses with the result of creating some confusion in the elaboration of an accurate phyletic tree. In Italy the greatest number of contri butions to this debate have been based on morphometric and mor phodynamic analyses, although lately genetic and chronological studies have gained attention, albeit in a lesser number with respect to those from foreign populations of cave bears (PEREGO et al., 2001;ORLANDO et al., 2002;BONA, 2004;QUILES, 2004;ROSSI et al., 2006;, 2011, 2012, 2013ROSSI et al., 2018;, 2008ZUNINO & PAVIA, 2005;MARTINI et al., 2014;TER LATO et al., 2017. The most previously studied caves are those from Lombardy and Veneto in Italy, whereas those from Pied mont (Sambughetto Valstrona, Grotta del Bandito among the most important ones) have been only superficially analysed (e.g. ...
This study is focused on cave bears from the Grotta del Bandito (Cuneo Province, Piedmont Region, Northern Italy), especially their teeth. The morphodynamic analyses have been elaborated and compared with other cave bears from Northern Italy, and other cave bear taxa from particular European sites. The Grotta del Bandito bears are similar in size to other cave bears from Italy, and no differences were observed with the other cave bear species or subspecies (e.g. U. ingressus, U. s. ladinicus, U. s. eremus). Thus their inclusion in the U. gr. spelaeus is the most appropriate classification, and they show a simpler dental surface than the “modern” cave bears (i.e. ingressus). The question of whether a single species of spelaeus lived in North Italy, is also introduced. In fact, the differences in size are probably the result of population variability, and no clear evidence of other spelaeoid species and/or subspecies, i.e. ingressus, s. ladinicus and s. eremus, was discovered.
The Cave Bear, Ursus spelaeus (sensu lato), was one of many megafaunal species that became extinct during the Late Pleistocene in Europe. With new data we revisit the debate about the extinction and paleoecology of this species by presenting new chronometric, isotopic and taphonomic evidence from two Palaeolithic cave bear sites in northeastern Italy: Paina Cave and Trene Cave. Two direct radiocarbon dates on well-preserved collagen have yielded ages around 24,200–23,500 cal yr BP, which make them the latest known representatives of the species in Europe. The carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopic values of bone collagen exhibit values similar to those of older cave bears from Swabian Jura and France, suggesting that the feedings preferences of cave bears remained unchanged until the disappearance of this species in Europe. Several bear remains preserved traces of human modification such as cut marks, which enables a reconstruction of the main steps of fur recovery and the butchering process. The broad range of plant types available and the favorable location of Berici Hills may have played an important role in the range expansion of cave bears and their interaction with the Paleolithic hunters settled the same area.
The fossil Fauna from the Steigelfadbalm cave was analysed by Prof. G. Rabeder and C. Rabeder MA from the University of Vienna. The cave bear bones could be attributed to the Ursus ingressus, an are thus the westernmost finds. A few chert and rock Crystal artefacts belongs to the Middle Paleolithic.
For the first time, morphometric variation has been studied in metacarpal and metatarsal bones of all known taxa of cave bears, which belong to different molecular genetic groups (deningeri, kudarensis, spelaeus, and ingressus haplotypes). The examined material involves nearly three thousand specimens from 28 localities of Europe, the Urals, Caucasus, and Central Asia. For comparison we used samples of fossil and recent Ursus arctos, as well as U. etruscus, regarded as a common
ancestor of brown bears and cave bears. Methods of univariate and multivariate statistical analyses of metapodial bones were employed, providing an opportunity to ascertain the degree of sexual dimorphism in different taxa, the degree of morphological difference between taxa, and to define “size” and “shape” morphospaces for concise description of morphological diversity and classification of cave bears. Our study reveals that, on average, sexual dimorphism is more pronounced in U. arctos and U. kudarensis praekudarensis than in cave bears. Sexual dimorphism of bear metapodia is greater than sexual dimorphism of the skull (Baryshnikov and Puzachenko 2011). The contribution of sexual dimorphism to size of the metapodials is close to a contribution associated with morphological disparity between the bears belonging to different taxa. By the example of two chronosubspecies of Kudaro cave bear: U. kudarensis praekudarensis from Middle Pleistocene and U. k. kudarensis from Late Pleistocene, we succeeded in detecting a decrease of sexual dimorphism over time, which suggests that earlier cave bears inherited a pronounced sexual dimorphism from ancestral taxa. Metacarpal and metatarsal bones of cave bears are easily distinguished from those of U. etruscus and U. arctos, simultaneously demonstrating similarity between cave bears from different genetic groups, involving the species U. kudarensis (the basal taxon for all cave bears, including U. deningeri); some peculiarities of these bones are revealed only in the smaller U. rossicus. The examples have shown the presence (U. k. kudarensis) as well as absence (U. deningeri, U. kanivetz ingressus) of evident spatial (geographical) and temporal patterns in metapodial variability. It is determined that taxa can be better differentiated by metacarpals rather than by metatarsals, because the latter proved to be more “conservative” and less variable over time. It is hypothesized that very rapid modification of metapodial bones occurred at an early stage of evolution of this group, which was presumably a result of occupation of a special ecological niche by cave bears. This study reveals that the size and shape of metacarpal and metatarsal bones did not have an observable link with the taxonomic or evolutionary position of cave bears. At the same time, the clear morphological differences between brown bears and cave bears reflect an early evolutionary divergence between “arctoid” and “spelaeoid” lineages.
We report radiocarbon ( ¹⁴ C) dates on bone samples of Ursus ladinicus , a small cave bear species well adapted to a life in the mountains, whose remains were found in Conturines Cave. Located at 2775 m asl in the Dolomites of northern Italy, this cave is by far the highest known cave bear site worldwide. Eleven ¹⁴ C dates obtained by the Belfast and Oxford laboratories on samples showing good collagen preservation yielded consistent ages in excess of 46–50 ka BP. These results show that contrary to the previously held view these cave bear remains are older than Marine Isotope Stage 3, and likely date from a warm climate period with a high treeline, possibly the Last Interglacial.
Climate changes that occurred during the Late Pleistocene had profound effects on the distribution of many plant and animal species and influenced the formation of contemporary faunas and floras of Europe. The course and mechanisms of responses of species to past climate changes are now being intensely studied by the use of direct radiocarbon dating and genetic analyses of fossil remains. Here, we review the advances in understanding these processes by the example of four mammal species: woolly mammoth (Mammuthus primigenius), cave bear (Ursus spelaeus s.l.), saiga antelope (Saiga tatarica) and collared lemmings (Dicrostonyx ssp.). The cases discussed here as well as others show that migrations, range shifts and local extinctions were the main responses to climate changes and that the dynamics of these climate-driven processes were much more profound than was previously thought. Each species reacted in its individual manner, which depended on its biology and adaptation abilities to changing environmental and climatic conditions. The most severe changes in European ecosystems that affected the largest number of species took place around 33–31 ka BP, during the Last Glacial Maximum 22–19 ka BP and the Late Glacial warming 15–13 ka BP.
The cave bear (Ursus spelaeus sensu lato) is a typical representative of Pleistocenemegafauna which became extinct at the end of the Last Glacial. Detailed knowledge of cave bear extinction could explain this spectacular ecological transformation. The paper provides a report on the youngest remains of the cave bear dated to 20,930 ± 140 14C years before present (BP). Ancient DNA analyses proved its affiliation to the Ursus ingressus haplotype. Using this record and 205 other dates, we determined, following eight approaches, the extinction time of this mammal at 26,100–24,300 cal. years BP. The time is only slightly earlier, i.e. 27,000–26,100 cal. years BP, when young dates without associated collagen data are excluded. The demise of cave bear falls within the coldest phase of the last glacial period, Greenland Stadial 3. This finding and the significant decrease in the cave bear records with cooling indicate that the drastic climatic changes were responsible for its extinction. Climate deterioration lowered vegetation productivity, on which the cave bear strongly depended as a strict herbivore. The distribution of the last cave bear records in Europe suggests that this animal was vanishing by fragmentation into subpopulations occupying small habitats. One of them was the Kraków-Częstochowa Upland in Poland, where we discovered the latest record of the cave bear and also two other, younger than 25,000 14C years BP. The relatively long survival of this bear in karst regions may result from suitable microclimate and continuous access to water provided by deep aquifers, indicating a refugial role of such regions in the Pleistocene for many species.
The most common methods used for assessing the relative age of a cave bear bone assemblage are the P4/4 index (morphodynamic index of the cave bear fourth premolar), the K-index, and the Index of Plumpness (both used for cave bear’s2nd metatarsal). Preliminary work on these indexes, for Urşilor Cave (NW Romania), has indicated one of the youngest European cave bear populations. As the number of extracted fossil bones from the palaeontological excavation increased recently, a re-assessment of the of the age of the cave bear assemblage is necessary. 206 cave bear fourth lower and upper premolars and 587 metapodials were analyzed. The P4/4 morphodynamic index, the K-index and the Index of Plumpness were calculated for the local MIS 3 cave bear bone assemblage. The results of the three indices have lower values when compared with the previously obtained for the same site and respect the subsequent radiometric ages (ca. 47-39 ky BP). However, the results for P4/4 morphodynamic index, K-index, and Index of Plumpness are of lesser relevance when used to assess the relative age of MIS 3 cave bear bone assemblages. All three methods require caution when applied and interpreted on short time intervals and on smaller geographic areas.
Abstract : In this contribution a morphometric study of the lower carnassial of the Cave Bear (Ursus deningeri and Ursus spelaeus) from several populations of diverse European localities and also different ages is carried out. This study includes a morphological analysis of the deployment of dental cusps and metric comparisons focused on general size and convergence of the cusps.
The morphologic study (by cluster analysis) presents a grouping trend of the populations according to their geographi position first, then to the chronology. This indicates that the expansion of the cave bear happened at a very early time, and that later did not exist great migratory movements that returned to put in contact remote populations.
As for the metric analysis, differences in the degree of convergence of the cusps in the talonid and trigonid are only observed between the oldest sites (more convergent, smaller occlusal surface) and the modern ones (less convergent, larger occlusal surface) independently of their geographic location.
Count rates, representing the rate of 14 C decay, are the basic data obtained in a 14 C laboratory. The conversion of this information into an age or geochemical parameters appears a simple matter at first. However, the path between counting and suitable 14 C data reporting (table 1) causes headaches to many. Minor deflections in pathway, depending on personal interpretations, are possible and give end results that are not always useful for inter-laboratory comparisons. This discussion is an attempt to identify some of these problems and to recommend certain procedures by which reporting ambiguities can be avoided.
Ursus spelaeus Rosenmüller, 1794 had a wide regional colonization in the Alps. Several cave bear fossils belonging to U. spelaeus were collected in the Buse di Bernardo cave (Tesero, Trento province, North Italy). With the purpose of placing these bears in the evolutionary frame of the "Italian" and European speleians, several morphological, morphometric and morphodynamic investigations on the fossils from this cave have been carried out. The most important conclusions of this work are : 1) there are not elements that may induce to include the Buse di Bernardo bears into the new species of cave bears, namely U. ingressus, U. spelaeus ladinicus and U. eremus, or even in U. deningeri and U. arctos. 2) U. deningeri Von Reichenau, 1906 and other primitive forms of U. spelaeus are absent. According with the morphodynamic index, the cave bears from this cave are not at the top of the evolutionary sequences, and this can indicate they never reached the top of the evolutionary step before their extinction. The morphometric and the morphodynamic analyses showed that these bears are, from the evolutionary point of view, very "similar" to those from other Italian caves.
Tischoferhöhle and Pendling-Bärenhöhle near Kufstein, Tyrol, are among the only locations where remains of cave bear, Ursus spelaeus-group, were found in the western part of Austria. One sample from each site was radiocarbon-dated four decades ago to ca. 28 14C ka BP. Here we report that attempts to date additional samples from Pendling-Bärenhöhle have failed due to the lack of collagen, casting doubts on the validity of the original measurement. We also unsuccessfully tried to date flowstone clasts embedded in the bone-bearing sediment to provide maximum constraints on the age of this sediment. Ten cave bear bones from Tischoferhöhle showing good collagen preservation were radiocarbon-dated to 31.1–39.9 14C ka BP, again pointing towards an age underestimation by the original radiocarbon-dated sample from this site. These new dates from Tischoferhöhle are therefore currently the only reliable cave bear dates in western Austria and constrain the interval of cave occupation to 44.3–33.5 cal ka BP. We re-calibrate and re-evaluate dates of alpine cave bear samples in the context of available palaeoclimate information from the greater alpine region covering the transition into the Last Glacial Maximum, eventually leading to the demise of this megafauna.
This article describes the metapodial bones from Križna jama in Slovenia, which are housed in the collections of the Institute of Palaeontology, University of Vienna, and at the Museum of Natural History, both Vienna. These metapodial bones are very big and plump and are superseded only by those from Ajdovska jama, which is also situated in Slovenia. The bears are attributable to the species Ursus ingressus Rabeder et al., 2004 and it is astonishing that they are only superseded in size by a population of Ursus ladinicus Rabeder et al., 2004. A suitable explanation for this behaviour can be found in different modes of responses to environmental conditions, see Rabeder et al. (2008).
L’évolution de l’ours des cavernes est caractérisée par des tendances particulières, incluant l’augmentation de la taille, la complexification progressive de la surface des dents et le renforcement progressif des os des métapodes. D’importants indicateurs de niveau évolutif sont l’indice morphodynamique de P4/4 et l’indice de la rondeur des os des métapodes. Ce n’est que récemment que l’analyse morphologique et morphométrique a été complétée par l’analyse génétique – particulièrement du mtDNA. En conséquence, des nouveaux taxons ont été proposés sur des critères génétique, et le rang taxonomique exact (espèces ou sous-espèces ?) de certains d’entre eux est encore en discussion. Cependant, le nouveau modèle évolutif présente certains problèmes car les seules données génétiques ne sont pas suffisantes pour une distinction spécifique, et les données morphométriques et morphodynamiques ne justifient pas une distinction spécifique entre les populations d’ours des cavernes. L’analyse morphométrique effectuée sur de nombreux ossements métapodes appartenant à différentes populations italiennes (Buco dell’Orso, Covoli Velo and S. Donà di Lamon) et européennes semble confirmer, dans l’ensemble, un niveau de diversité pas plus élevé que celui des sous-espèces, ce qui permet tout au plus l’identification de certaines tendances évolutives locales, comme on le suppose pour les populations ayant vécu dans la péninsule italienne.
Analysis of Cave Bear Metapodial Bones from Ajdovska jama (Slovenia). — The analysis of metapodial bones from Ajdovska jama has shown that these bears, alongside with some other peculiarities, possessed astonishingly long metapodials, even in comparison with the biggest member of the cave bear group, U. ingressus. This is even more interesting in the light of the specific attribution of these bears, which turned out to belong to U. ladinicus, which was believed to be one of the smallest members of the cave bear group. Moreover this demonstrates the immense variability of this species in dependence of its habitats altitude. Izvleček Analiza metapodijev jamskega medveda iz Ajdovske jame (Slovenija). — Analiza metapodijev iz Ajdovske jame je pokazala, da imajo ti medvedi, poleg nekaterih drugih posebnosti, nenavadno dolge dlančnice in nartnice celo v primerjavi z osebki vrste Ursus ingressus – največjimi predstavniki skupine jamskih medvedov. Ta okoliščina je toliko bolj zanimiva, če pomislimo, da so pripisali medve-de iz Ajdovske jame vrsti Ursus ladinicus. Do nedavnega so osebki te vrste veljali za telesno najmanjše zastopnike skupine jamskih medvedov. To kaže na veliko spremen-ljivost te vrste v odvisnosti od nadmorske višine njihovega bivalnega okolja. Ključne besede: Ursus ladinicus, metapodiji, Ajdovska jama, Slovenija Zusammenfassung Eine Analyse der Höhlenbären-Metapodien aus der Aj-dovska jama (Slowenien). — Die Analyse der Metapodien aus der Ajdovska jama hat gezeigt, dass diese Bären, neben einigen anderen Eigenheiten, außergewöhnlich lange Mittelhand-und Mittelfußknochen haben, die auch den Vergleich mit dem größten Vertreter der Höhlenbären-Gruppe, U. ingressus, nicht zu scheuen brauchen. Dieser Umstand ist noch interessanter, wenn man bedenkt, dass diese Bären zu U. ladinicus gestellt werden, der bislang als einer der kleinsten Vertreter der Höhlenbären-Gruppe gegolten hat. Darüber hinaus zeigt das die immense Variabilität dieser Art in Abhängigkeit von der Seehöhe ihres Habitats.
Morphological and molecular data suggest the existence of several taxa of cave bears, which were found to belong to three major mitochondrial haplogroups: kudarensis (kudarensis), spelaeus (ladinicus, eremus, spelaeus), and ingressus (uralensis, ingressus, kanivetz). An analysis of craniometrical variability was carried out based on 20 measurements of 279 skulls from 40 European, Ural, and Caucasian localities, in order to investigate morphological similarity/dissimilarity of these taxa in a multivariate approach. The craniometrical analysis divides cave bears into two groups: small cave bears and large cave bears. The group of small bears consists of rossicus/uralensis and ladinicus. In some skull proportions, these taxa display intermediate position between brown and cave bears, i.e. presumably possessing archaic characters of their common ancestor. The group of large bears includes taxa with more specialized skulls. An early radiation within this group is demonstrated by kudarensis which probably ranged across Asia, and deningeri which occupied Europe east to the Ural Mountains. In its craniometrical characters, kanivetz from the Late Pleistocene of the Urals resembles deningeri. Other taxa of large cave bears (spelaeus, ingressus and eremus) reveal further evolution of cranial characters, being similar in the skull proportions. The level of difference between spelaeus and ingressus does not exceed that between subspecies of the recent brown bear, such as Ursus arctos beringianus and U. a. piscator. The examined isolated population of large cave bears from Volga River region (Zhiguli Hills) is similar to ingressus. Thus, based on the craniometrical data, the following species of cave bear are recognized: Ursus kudarensis (with the subspecies U. k. praekudarensis and U. k. kudarensis), U. deningeri (several subspecies), U. rossicus (with subspecies U. r. rossicus and U. r. uralensis), U. ladinicus, U. spelaeus (with subspecies U. s. spelaeus, U. s. eremus, U. s. ingressus, and, provisionally, U. s. kanivetz,).
En este trabajo se realiza un estudio métrico y morfológico de la dentición inferior de U. etruscus Cuv.,
U. deningeri V. REICH., U. spelaeus Ros-HEIN., y U. arctos LIN.
El cuarto premolar inferior de las especies espéleas desarrolla formas progresivamente más anchas y con
el paracónido cada vez más complejo. En U. arctos el premolar recuerda al de U. etruscus, aunque es más ano
cho y el protocónido no tiene aristas. La población de morfologías correspondiente a U. deningeri oscila entre
las formas arcaicas, etruscoides, hasta las hiperespéleas.
En el primer y segundo premolares de la línea evolutiva espeloide hay una verticalización progresiva de las
cúspides (paracónido-protocónido), que además se hacen cada vez más complicadas (metacónido y entocónido).
En la línea arctoide (U. etruscus - U. arctos) aparece un fenómeno similar, aunque no tan marcado.
Como ocurre en los molariformes superiores es imposible establecer a priori la existencia de dimorfismo
sexual, pero si se estudia una amplia serie de molares todavía implantados en mandíbulas, susceptibles de ser se·
xados, se puede concluir que existe un dimorfismo sexual incompleto, pudiendo afirmarse que los dientes más pe.
queños pertenecen a hembras y los más robustos a machos, siendo imposible establecer el sexado de los más
frecuentes tamaños intermedios.
El análisis métrico muestra un aumento progresivo, en ambas líneas evolutivas, con abundantes medidas
que permiten separar las diversas especies.
Tal y como se vio en los molariformes superiores, aparece una alometría negativa en la anchura, en U. deningeri
y U. spelaeus, y el aumento de anchura se centra ~n la zona del talónido, mientras que se acorta el trigónido.
En este trabajo se realiza una revisión de la dentición de leche (decidual) de los Ursidos, aunque el principal
objetivo es el análisis de la fórmula dentaria y de la morfología y biometrfa de la dentición superior de los
grandes osos del Pleistoceno: U. etruscus G. CUVIER, U. deningeri VON REICH, U. deningeri VON REICH, U. spelaeus
Ros-HEIN. y U. arctos LINN.
En lo concerniente a la fórmula dentaria, se evidencia una reducción progresiva de la presencia de los tres
primeros premolares, que están presentes en la especie ancestral U. etruscus, faltando en la más evolucionada,
U. spelaeus. En U. deningeri pueden aparecer el primero y/o el tercero, pero nunca el segundo; algo similar ocurre
en U. arctos. Del análisis morfológico Se puede deducir que existe una gran diferencia morfológica entre U.
etruscus y U. deningeri, este último muy cercano a U. spelaeus, aunque abundantes caracteres morfológicos los diferencian.
La morfología de la dentición superior de U. arctos es muy parecida a la de U. etruscus, pero con un desarrollo
más complejo de las cúspides que además se hacen más verticales.
El análisis métrico prueba la existencia de dimorfismo sexual en el canino y tercer incisivo superiores, no
siendo posible establecerlo en los premolares y molares superiores.
Hay una excelente separación métrica entre las poblaciones de medidas correspondientes a U. deningeri y
U. spelaeus, que a su vez se diferencian fuertemente de la correspondiente a U. etruscus y U. arctos. El enorme
aumento de longitud del cuarto premolar y molares superiores de los representantes de la línea espeloide, U. deningeri
y U. spelaeus, no fue acompañado por un aumento armónico en anchura, de forma que estos dientes resultan
relativamente más estrechos. En las dos líneas evolutivas, arctoide y espeloide, hay un aumento progresivo
de la altura de las cúspides.
En este trabajo se realiza un estudio biométrico comparado de los huesos del miembro pelviano, tarso,
metatarso y dedos de U. deningeri, U. spelaeus y U. arctos.
Las medidas de los huesos largos revelan la existencia de dimorfismo sexual, que no se pone tan claramente
de manifiesto en las referentes a huesos del tarso, metatarso y dedos. Hay una neta separación morfológica
entre los huesos de las distintas especies: los huesos de oso de las cavernas están pesadamente construidos,
mientras que en los de oso pardo permanecen muchas de las características de su remoto ancestro U. etruscus
G. CUVIER U. deningeri muestra características intermedias. El aumento de robustez, característico de las especies . espéleas, fue acompañado por un notable crecimiento de las áreas de las facetas articulares de los huesos del tarso.
El análisis métrico, realizado mediante histogramas y representaciones semilogarítmicas y bivariadas, revela que
existe una buena separación entre las especies. Se incluyen tablas de medidas.
The IntCal04 and Marine04 radiocarbon calibration curves have been updated from 12 cal kBP (cal kBP is here defined as thousands of calibrated years before AD 1950), and extended to 50 cal kBP, utilizing newly available data sets that meet the IntCal Working Group criteria for pristine corals and other carbonates and for quantification of uncertainty in both the 14C and calendar timescales as established in 2002. No change was made to the curves from 0–12 cal kBP. The curves were constructed using a Markov chain Monte Carlo (MCMC) implementation of the random walk model used for IntCal04 and Marine04. The new curves were ratified at the 20th International Radiocarbon Conference in June 2009 and are available in the Supplemental Material at www.radiocarbon.org.
Morphological and metrical differences between several alpine cave bear associations of the same age are large enough to suspect more than one evolutionary line. The contemporaneous age of several morphologically distinct forms is confirmed by radiocarbon dating. Moreover analyses of fossil DNA indicate at least three different lineages within the cavebear group. For two of these groups, both morphological and genetic data show evidence of reproductive isolation. Thus, we suggest that Ursus spelaeus comprised at least two different species. Ursus ingressus n. sp. occurred in the eastern parts of the Alpine region and in the Dinarids of Slovenia and Croatia. For two other morphological forms, subspecific status with Ursus spelaeus ladinicus n. ssp. in the Dolomites and Ursus spelaeus eremus n. ssp. in the Totes Gebirge is suggested, as they are genetically relatively close and no data about their reproductive relationship is yet available.
Cave bears (Ursus spelaeus) existed in Europe and western Asia until the end of the last glaciation some 10,000 years ago. To investigate the genetic
diversity, population history, and relationship among different cave bear populations, we have determined mitochondrial DNA
sequences from 12 cave bears that range in age from about 26,500 to at least 49,000 years and originate from nine caves. The
samples include one individual from the type specimen population, as well as two small-sized high-Alpine bears. The results
show that about 49,000 years ago, the mtDNA diversity among cave bears was about 1.8-fold lower than the current species-wide
diversity of brown bears (Ursus arctos). However, the current brown bear mtDNA gene pool consists of three clades, and cave bear mtDNA diversity is similar to the
diversity observed within each of these clades. The results also show that geographically separated populations of the high-Alpine
cave bear form were polyphyletic with respect to their mtDNA. This suggests that small size may have been an ancestral trait
in cave bears and that large size evolved at least twice independently.
Cave bears have disappeared from the Alps from different altitudes at different times. The temporal progression of the HDEL (Height Dependent Extinction Line) – a compilation of the geologically most recent radiocarbon dates per altitude level – is not consistent with the general cooling of the temperatures from about 45 ka BP. The cave bear sites of the Northern Alps with the most recent radiocarbon ages are not situated in the lowlands but in caves in altitudes of 1,500 m to 1,700 m above sea level (a.s.l.).
Cave bears fed almost exclusively on herbs and leaves. It was assumed that with the general cooling in the OIS 3 since about 45 ka BP also the migration of the alpine elements into the lowlands took place. It could be recognized that the populations in the lower situated cave bear site became earlier extinct than the cave bear population in the higher altitudes.
With new radiocarbon dates, done at the Curt-Engelhorn-Center Archaeometry at the Reiss-Engelhorn-Museen in Mannheim (Germany), the HDEL can be determined much more precisely and the causes of gradual extinction are also better understood.
W F Libby's new dating method from the 1940s, based on experience in physics and chemistry, opened possibilities to check and revise chronologies built on other principles than radioactive decay. Libby's method initially implied collaboration with archaeologists to demonstrate that it worked but also with physicists to improve the technique to measure low β – activities. Chemists, geophysicists, botanists, physiologists, statisticians, and other researchers have contributed to a prosperous interdisciplinary development. Some pitfalls were not recognized from the beginning, although issues such as contamination problems were foreseen by Libby. Pretreatment of samples was discussed very early by de Vries and collaborators, among others. This subject has not yet been abandoned. Closely related to pretreatment is the choice of fraction to be dated and chemicals to be used, especially for accelerator mass spectrometry (AMS) measurements. Calibration against tree rings and comparison with dates obtained using other methods as well as intercomparison projects are partly history but still very actual. The impact by man and climate is also studied since the early days of the method. Also, the carbon cycle has been of great interest. The tools for measurements and statistical analysis have been improved during these first 3 or 4 decades, allowing interpretations not possible earlier. δ 13 C determinations are mostly very important and useful, but sometimes they have been misleading in discussions of the origin of carbon, especially for human tissues—the metabolism was not yet fully understood. The history and development of the method can only be illustrated by selected examples in a survey like this.
Cave bear remains are known from 36 caves and other karst features, and from one open-air site in
Serbia. The sites vary greatly by their morphology and size, position, altitude, stratigraphy and diversity
of the fossil fauna. The cave bear stands out as a dominant species at the majority of the sites; at a
number of sites it is the single species found. There are only few sites where herbivores’ or predators’
remains exceed those of the cave bears. The majority of cave bear remains originate from the Late
Pleistocene deposits. Previously, they have been identified as Ursus spelaeus.
We present a morphology and metric study of cave bears from three sites: two recently discovered
localities of Middle Pleistocene age, and the Risova�ca cave, which may be regarded as the reference site
for the Late Pleistocene mammals in the region based on a large number of collected remains and
published data. The remains of the Middle Pleistocene spelaeoid bears are described from Mala Balanica
cave (Eastern Serbia) and an open-air site at Ruma brickyard (Northern Serbia) for the first time.
Considering the size and morphology of cheek teeth and metapodial bones, these cave bears are identified
as Ursus spelaeus deningeroides. The cheek teeth morphology and size, as well as metapodial bones’
metric, relate the cave bear from the Risova�ca cave to the advanced evolutive type of the cave bear
established in Central and Eastern Europe Ursus spelaeus ingressus.
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.
Buse di Bernardo" e dal "Covelo di Rio Malo
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1919, I Mammiferi quaternari della regione veneta: Memorie Istituto di Geologia dell'Università di Padova
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Extrait d'une lettre de M. l'abbé Fortis datée de Vérone le 24 Septembre 1785 à M. le Comte de Cassini de l'Académie des Sciences sur différentes pétrifications
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Fortis, A., 1786, Extrait d'une lettre de M. l'abbé Fortis datée de Vérone le 24 Septembre 1785 à M. le Comte de Cassini de l'Académie des Sciences sur différentes pétrifications: Journal Physique, v. 28, no 1, p. 161-168.
The Cave Bear Incisors from Križna jama (Slovenia): Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften
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Frischauf, C., 2014, The Cave Bear Incisors from Križna jama (Slovenia): Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, v. 21, p. 75-83.
Genetic analysis of a cave bear-morphothype specimen from Prepoštská cave, Slovakia featuring carnivorous-like isotopic signature: Aragonit
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The latest Ursus spelaeus in Italy
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Osteologia degli Orsi fossili nel Veronese, con un saggio sopra le principali caverne del Distretto di Tregnago: Naturwissenschaftliche Abhandlungen
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Ancient DNA and the population genetics of cave bears (Ursus spelaeus) through space and time
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Ursus spelaeus from Grotta Sopra Fontana Marella
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Morphometrical analyses of cave bear populations from Barová cave in moravian karst
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Republic): Aragonit, v. 22, no 1, p. 20-21.
Analyse morphodynamique de l'ours des caverne (Carnivora, Ursidae) de cinq sites du pourtour Méditerranéen : Cahiers scientifique du Muséum d'Histoire naturelle de Lyon Hors Série
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Metrics and Evolutionary Level of Teeth of the Bears from Križna jama (Slovenia): Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften
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Rabeder, G., 2014, Metrics and Evolutionary Level of Teeth of the Bears from Križna jama (Slovenia): Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, v. 21, p. 57-63.
The chronological position of the cave bears from Conturines cave: Abstract, 2 nd International Cave Bears Symposium Alta Badia
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Studio morfometrico e morfodinamico di resti craniali, dentali e mandibolari di Ursus spelaeus dalla Grotta del Buco del Frate e dall'Altopiano di Cariadeghe (Brescia) nel quadro evolutivo degli orsi delle caverne: Natura Bresciana, Annali del Museo Civico di Storia Naturale Brescia
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