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Unexpected! New AMS Dating from Austrian Cave Bear Sites

Authors:
  • Naturmuseum St. Gallen

Abstract and Figures

New AMS dating for three Austrian sites were conducted on cave bear bones at the Klaus-Tschira-Archaeometry Center in Mannheim, Germany. In total 14 new dates will be presented. The oldest date is 48 ka BP. The faunal remains from the Schwabenreith Cave, located near Lunz (Lower Austria), only consist of cave bears from the taxa Ursus spelaeus eremus. The basal and top flowstone layers of excavation area 2 yielded U-Th ages of 116±5 ka and 78+30/-23 ka BP, respectively. In the Herdengel Cave, located in the same region, the remains of U. sp. eremus and U. ingressus were found. A basal flowstone layer yielded a U-Th age of 112+12/-11 ka BP. The Brettstein Cave system in the Totes Gebirge (Styria) represents the two cave bear taxa U. sp. eremus and U. ladinicus. Dated cave bear bones were only known to be older than 49 ka BP. The new AMS dates include six bone remains from Schwabenreith Cave dated in the period from 34 ka to 48 ka BP. New dating results from the Herdengel Cave show a very close timespan from 44 ka to 48 ka BP. And finally the bears of the Brettstein Cave represent one of the youngest dated remains (22.5 ka to 35 ka BP) in the Alps.
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ICBS PROCEEDINGS
[26] CRANIUM JUNI 2016
DESCRIPTION OF THE CAVE BEAR
SITES
The Schwabenreith Cave (Austrian cave cadaster no.
1823/32) is located near Lunz am See in the western part of
Lower Austria (Fig.1) at 959 m above sea level. In excava-
tion area 2, a 1.3 m thick bone layer only consisted of cave
bear remains (Ursus spelaeus eremus) (Fig.2). They were an-
alysed from a taphonomic point of view for the rst time by
Pacher (2000). The radiometric data from owstone samples
considered the bear remains to be of an early Wurmian age.
The basal and top owstone layers yielded Uranium-Thori-
um (U-Th) ages of 116±5 ka and 78+30/-23 ka, respectively
(Frank & Rabeder, 1997a). The abundance of cave bear
remains is very high. All skeletal elements are represented.
Despite the density of remains, a taphonomic analysis veri-
es a certain transport of bones. It probably must have taken
place within humid sediment with plasticized consistency.
The Herdengel Cave (Austrian cave cadaster no. 1823/4)
is located near the Schwabenreith Cave (Fig.1) at 878 m
above sea level. Beneath remains of cave bears (U. sp.
eremus and U. ingressus) other pleistocene animals like
cave lion and wolf were found (Frank & Rabeder, 1997b;
Pacher, 2009). Even a Mousterian artifact is documented
UNEXPECTED! NEW AMS DATING FROM
AUSTRIAN CAVE BEAR SITES
DORIS DÖPPES REISS-ENGELHORN-MUSEEN, ZEUGHAUS, C5, 68159 MANNHEIM, GERMANY, DORIS.DOEPPES@MANNHEIM.DE
MARTINA PACHER UNIVERSITY OF VIENNA, INSTITUTE OF PALAEONTOLOGY, PALAEOBIOLOGY-VERTEBRATEPALAEONTOLO-
GY, GEOZENTRUM UZA II, 1090 VIENNA, AUSTRIA, MARTINA.PACHER@UNIVIE.AC.AT
GERNOT RABEDER UNIVERSITY OF VIENNA, INSTITUTE OF PALAEONTOLOGY, PALAEOBIOLOGY-VERTEBRATEPALAEONTOLO-
GY, GEOZENTRUM UZA II, 1090 VIENNA, AUSTRIA, GERNOT.RABEDER@UNIVIE.AC.AT
SUSANNE LINDAUER KLAUS-TSCHIRA-ARCHAEOMETRY CENTER AT THE CURT-ENGELHORN-CENTER FOR ARCHAEOMETRY,
C4, 8, 68159 MANNHEIM, GERMANY, SUSANNE.LINDAUER@CEZ-ARCHAEOMETRIE.DE
RONNY FRIEDRICH KLAUS-TSCHIRA-ARCHAEOMETRY CENTER AT THE CURT-ENGELHORN-CENTER FOR ARCHAEOMETRY, C4,
8, 68159 MANNHEIM, GERMANY, RONNY.FRIEDRICH@CEZ-ARCHAEOMETRIE.DE
BERND KROMER KLAUS-TSCHIRA-ARCHAEOMETRY CENTER AT THE CURT-ENGELHORN-CENTER FOR ARCHAEOMETRY, C4, 8,
68159 MANNHEIM, GERMANY, BERND.KROMER@CEZ-ARCHAEOMETRIE.DE
WILFRIED ROSENDAHL REISS-ENGELHORN-MUSEEN, ZEUGHAUS, C5, 68159 MANNHEIM, GERMANY, WILFRIED.ROSENDAHL@
MANNHEIM.DE
Abstract
New AMS dating for three Austrian sites were conducted on cave bear bones at the Klaus-Tschira-Archaeom-
etry Center in Mannheim, Germany. In total 14 new dates will be presented. The oldest date is 48 ka BP. The
faunal remains from the Schwabenreith Cave, located near Lunz (Lower Austria), only consist of cave bears from
the taxa Ursus spelaeus eremus. The basal and top owstone layers of excavation area 2 yielded U-Th ages of
116±5 ka and 78+30/-23 ka BP, respectively. In the Herdengel Cave, located in the same region, the remains of
U. sp. eremus and U. ingressus were found. A basal owstone layer yielded a U-Th age of 112+12/-11 ka BP. The
Brettstein Cave system in the Totes Gebirge (Styria) represents the two cave bear taxa U. sp. eremus and U.
ladinicus. Dated cave bear bones were only known to be older than 49 ka BP.
The new AMS dates include six bone remains from Schwabenreith Cave dated in the period from 34 ka to 48 ka
BP. New dating results from the Herdengel Cave show a very close timespan from 44 ka to 48 ka BP. And nally
the bears of the Brettstein Cave represent one of the youngest dated remains (22.5 ka to 35 ka BP) in the Alps.
Samenvatting
Nieuwe AMS-dateringen voor drie Oostenrijkse sites werden uitgevoerd op botten van grottenberen in het
Klaus-Tschira-Archaeometry Center in Mannheim, Duitsland. In totaal worden 14 nieuwe dateringen gepresen-
teerd. De oudste daarvan is 48 ka BP. De faunaresten uit de Schwabenreith grot, nabij Lunz (Neder-Oostenrijk),
bestaan enkel uit grottenberen van de soort Ursus spelaeus eremus. De druipsteenlagen van bodem en plafond
in opgravingsveld 2 gaven U-Th leeftijden van respectievelijk 116±5 ka BP en 78+30/-23 ka BP. In de Herdengel
grot, gelegen in dezelfde regio, werden de resten gevonden van U. sp. eremus en U. ingressus. Een bodemdru-
ipsteenlaag gaf een U-Th leeftijd van 112+12/-11 ka BP. Het Brettstein grottencomplex in het Totes Gebirge
(Stiermarken) vertegenwoordigt de twee grottenbeersoorten U. sp. eremus en U. ladinicus. De enige bekende
gedateerde botten hebben een ouderdom van meer dan 49 ka BP.
De nieuwe AMS-dateringen omvatten zes botresten van de Schwabenreith grot uit de jongere periode van 34
tot 48 ka BP. Nieuwe data van de Herdengel grot geeft een nauwe tijdspanne van 44 tot 48 ka BP. Tot slot ver-
tegenwoordigen de grottenberen uit de Brettstein grot één van de jongst gedateerde resten in de Alpen (22,5
tot 35 ka BP).
ICBS PROCEEDINGS
[26] CRANIUM JUNI 2016
AUTHORS
DORIS DÖPPES
MARTINA PACHER
GERNOT RABEDER
SUSANNE LINDAUER
RONNY FRIEDRICH
BERND KROMER
WILFRIED ROSENDAHL
from the approximately 3 m thick bone layer (Frank &
Rabeder, 1997b). A basal owstone layer yielded a U-Th age
of 112+12/-11 ka and 111+11/-10 ka BP (Leitner-Wild et al.,
1994). Cave bear bones were dated by radiocarbon and U-Th
method from 37+/-0.59 ka to 135+11/-10 ka (Leitner-Wild et
al., 1994; Frank & Rabeder, 1997b).
The huge Brettstein Cave system (“Brettstein Bären-
höhle”, Austrian cave cadaster no. 1625/33), is over 4 km
long and is located in the southern area of the eastern Totes
Gebirge, near Bad Mitterndorf (Fig.1) at 1664 m above sea
level (entrance a). Six excavation areas were installed in
four different cave parts. In none of the six excavation areas
the fossil remains were found in original position (Döppes
et al., 1997). The Pleistocene large mammal fauna consists
mainly of cave bear (U. sp. eremus and U. ladinicus). Cave
lion, wolf, wolverine and ibex are presented by several bones
(Ehrenberg, 1958; Rabeder et al., 2001). The chronological
position of the cave bear remains from the Brettstein Cave
is conrmed by radiocarbon dates older than 37, 41 and 44
ka BP (Döppes, 2000; Pacher, 2003). An AMS-14C dated
bear bone from the Blasloch, at 1623 m above sea level,
nowadays part of the Brettstein Cave system, was dated to
51,300+2,300/-1,800 years BP (Pacher & Stuart, 2009).
METHOD
The development of the accelerator mass spectrometry
(AMS) made it possible not only to signicantly mini-
mize the quantity of samples required, but also to increase
measurement speed and precision of the counting technique
considerably. This method was used to establish the age of
the fossils from the three caves.
Since contamination can occur during soil sedimenta-
tion, datable carbonaceous samples are freed from coarse
impurities and foreign carbon that can distort age. Samples
are pretreated with acid and base steps to remove carbonate
and humic acids. In the case of bone samples, the collagen
– a structural protein – is extracted, ultra-ltered to remove
molecules of chain length lower than 30kDa (potentially
younger proteins taken up by the bone from water), and
freeze-dried. Bone was long considered to be unsuitable
for 14C dating since it is very porous and the bone apatite
is prone to exchange reactions with the groundwater and
surrounding material. However, collagen is hardly prone to
exchanges. In the nal step, organic samples are converted
into carbon dioxide by combustion of the sample material
and subsequently reduced to graphite. Sample sizes in the
mg range are suitable for measurement in an accelerator
mass spectrometer. The graphite sample obtained is sputtered
with cesium ions in order to obtain carbon ions. The ions of
the carbon isotope are separated in the accelerator according
to their different masses. From the measured 14C/12C ratios
the age of the samples can be determined. The measured
13C serves as the control for and correction of fractionation
processes in nature or in the laboratory.
Radiocarbon data is by default reported as conventional
14C age BP. This should not be taken as a calendar age. The
origin of this convention lies in the fact that originally the
data was converted to an age by using the radioactive decay
for age determination assuming constant 14C production,
hence atmospheric 14C level, in the past. Unfortunately, it
turned out that this is incorrect. Radiocarbon is produced
in the atmosphere by extraterrestrial irradiation of which
galactic cosmic rays provide the major component. Variable
shielding by the geomagnetic eld and the magnetic eld
of the protons in the solar wind lead to uctuations in the
atmospheric radiocarbon level. To cope with this, a cal-
ibration curve was established using independent dating
methods such as dendrochronology (until ~10,000 BC),
Uranium-Thorium dating of speleothems and corals, and
varve counting of terrestrial and marine sediments. The
limit of the method is due to the fact that after approxi-
mately 10 half-lives (half-live of 5,730±40 years) only
less than 1 permille of the original 14C remains, hence
no material older than 50,000 years can be dated reliably
with this method (Reimer et al., 2013; Olsson, 2009).
Calibrated ages are usually quoted with a 1-sigma error
range, corresponding to a condence probability of 68.3%.
It rises to 95.5% for 2-sigma. The calibration here was
performed using the programme SWISSCAL 1.0 (L. Wacker,
ETH Zürich) with the INTCAL13 dataset.
Figure 1: (modied after Rabeder et
al., 2011): Important cave bear sites
in Austria with different cave bear
species, and the locations of the three
studied caves
Figuur 1: (naar Rabeder et al.,
2011) Belangrijke Oostenrijkse
vindplaatsen van verschillende soorten
grottenberen en de locaties van de
drie onderzochte grotten
Figure 2: Excavation area 2 of the Schwabenreith Cave
Figuur 2: Opgravingsveld 2 van de Schwabenreith grot
GERNOT RABEDER
GERNOT RABEDER
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[28] CRANIUM JUNI 2016
RESULTS
The Curt Engelhorn-Centre for Archaeometry (CEZA)
received 17 samples of cave bear bones to determine the age
by 14C with the MICADAS Accelerator of their subsidiary
institute Klaus-Tschira-Archaeometry Center. The radiocar-
bon data is shown in Table 1.
The 14C age is normalized to δ13C = -25 ‰ (Stuiver &
Pollach, 1977). The δ13C value originates from the mea-
surement of the 13C/12C isotope ratios in the accelerator; its
error is reported to approximately 0.5‰. However, the value
can be falsied by isotope separation during preparation and
in the ion source of the accelerator over the original value of
the sample material, and can only be used to correct the frac-
tionation effects. The value is therefore not comparable with
the measurement in a mass spectrometer for stable isotopes
(IRMS) and should not be used for further data interpreta-
tion. Typically, the AMS-derived value is accurate within
2-3‰ compared to the original value. The C/N ratio and
carbon content of the collagen extracted are in the normal
range (Van Klinken, 1999), and the collagen preservation of
the samples is good.
DISCUSSION
The knowledge of the evolution and phylogeny of cave
bears has changed fundamentally in recent years. While it
was assumed that there had been only one cave bear species,
we now know from morphological studies (Rabeder, 1999)
and especially genetic analyses (Rabeder et al. 2004, Stiller
et al., 2010) that the family tree of the cave bear is very
complex and its research still in progress. During the same
geological period (50 – 30 ka BP) at least three species lived
in the Alps: U. sp. eremus, U. ladinicus and U. ingressus
(Rabeder et al. 2004). All are extinct before the coldest phase
of the last ice age (Pacher & Stuart, 2009).
Three dates from the Schwabenreith Cave excavation
area 2 are consistent with the time range from the Herdengel
Cave, in particular those from 46 ka BP to 49 ka BP (includ-
ing errors). Two samples are approximately 10,000 years
younger (range 34,010 – 37,400 years BP).
Another AMS date from excavation area 3, which is
located in another part of the cave, falls at the lower range of
dates from Schwabenreith Cave (52,500+1,900/-2,500 years
BP) (Pacher, 2000).
Meanwhile, both speleothem layers of excavation area 2
have been redated (Christoph Spötl, University of Innsbruck,
personal communication). The initial data were conrmed.
The cave bear bones from this excavation area consistently
dated younger than the speleothem above. Maybe the matrix
of excavation area 2 inuenced the data. The controlling val-
ues of C/N ratio, collagen and carbon of the new AMS data
support taphonomic results. The articulated remains could
have been transported beneath the speleothem. The spatial
distribution of bones and a low inclination of layers indicate
a transport of remains NW-SE or NW-SW. Some of the few
articulated parts of single skeletons in this area have proba-
bly been transported in correct anatomical position (Pacher,
2000). Since there are cave bear bones directly covered by
speleothem, new datings can perhaps clarify the situation.
The stratigraphy of the Herdengel Cave, with a cross
section of nearly 5 m (Fig.3), shows cave bear bones from
2.0 to 3.7 m and below the basal speleothem layer. Con-
Figure 3: (modied after Frank & Rabeder,
1997b) Standardized cross section of the
Herdengel Cave
Figuur 3: (naar Frank & Rabeder, 1997b)
Schematische doorsnede van de Herdengel
grot.
ICBS PROCEEDINGS
[28] CRANIUM JUNI 2016
centrations of bones were found at 3.0 to 3.6 m (layer 4-3).
Layer 5 above contains a mixture of fossils and larger stones.
The basal part of layer 6 contains only few cave bear bones.
The cave bear bones in layer 1 were excavated below the
basal owstone layer that yielded a U-Th age of 112+12/-11
ka and 111+11/-10 ka BP. By applying the uranium series
method (Frank & Rabeder, 1997b) to the Herdengel Cave
cross-section, the evolutionary rate of the cave bears was
determined. Uranium series data from the fossil bones were
partly veried by an independent carbonate speleothem age.
For both, bone layers and the carbonate formation found in
stratigraphic relation, the determined ages correspond to a
normal time sequence. According to the relatively precise
time scale obtained by absolute dating, the evolutionary
mode of the cave bears was determined as gradual (Rabeder,
1999).
The ve new dates from layer 6 to layer 1 show a spread
of approximately 6,000 years (using the minimum and max-
imum range of individual dates). The basal part of layer 6
contains probably reworked cave bear remains, because they
are intermingled with younger fauna elements like marmot,
and the density of nds is low within the sediment. The
sediments consist of a yellow-brownish loam with rubble
and differ clearly from layer 5 to 4 below. An erosion period
is documented in the small layer 5 in-between. The loamy
layer contains still abundant fossil remains, but also a higher
degree of larger stones. Layers 4 and 3 are the richest in cave
bear remains and seem to represent the original cave bear oc-
cupation phase(s). The date from layer 4 is slightly younger
if errors are considered (range 42,600 – 45,660 years BP),
but still overlaps with the range of one of the remaining sam-
ples. Beneath the basal owstone layer – almost of the same
age as the one in the Schwabenreith Cave (excavation area 2)
– bones from layer 1 were dated the same age as bones from
above this speleothem. Maybe the basal speleothem layer
was broken, as is the case in Schwabenreith Cave.
The many entrances (a-m) of the Brettstein Cave system
(Fig.4) and the rearrangement of the bones (Rabeder et al.,
2001) are evidence of the shearing action of outowing ice
during the Last Glacial Maximum. With the help of ancient
DNA investigation, the cave bear species Ursus sp. eremus
and Ursus ladinicus can be distinguished. Unfortunately, it is
not possible to differentiate the two cave bear species based
on single bones and tooth elements. Prior to our study, the
cave bear remains were not conrmed by radiocarbon dates,
except an AMS-14C dated bone from the Blasloch, which
was dated to 51,300+2,300/-1,800 years BP.
For the rst time, enough collagen could be obtained for
dating. All three new datings show a younger time span
than the sample from the Blasloch. The dates of 21,970 and
22,510 years BP would be the youngest dated cave bears,
not only in the Alps but throughout its range. The youngest
known specimen of cave bear in the High Alps, based on
audited dates, is from the Lieglloch in the Totes Gebirge
(Styria), with an age of 26,390+/-110 years BP (Pacher &
Stuart, 2009).
CONCLUSION
In general, the cave bears from the Schwabenreith Cave
and the Herdengel Cave have the same main occupation
phase, despite a different stratigraphy and cave morphology.
The Brettstein Cave sample shows the youngest cave bear
site in the Alps. As a result, contradictions arise to the current
opinion, that the plateau of the Totes Gebirge was covered
with ice at that time. Additional analyses are necessary to
clarify this question.
The dated collagen of the Brettstein Cave samples is ex-
amined with the help of ancient DNA analysis to conrm the
bear species. Furthermore, since January 2016, 19 new bone
samples from the Herdengel Cave (layer 6 to layer 1), six
from the Schwabenreith Cave, and four from the Brettstein
Cave (29 in total) have been analysed by the Klaus-Tschi-
ra-Archaeometry Center. With more AMS dated cave bear
bones and ancient DNA analysed samples, we will try to
nd an explanation for what was going on during the Wurm
glacial in these Austrian cave bear sites.
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Verein für Höhlenkunde in Obersteier
03/2009) Map of the Brettstein Cave
system with excavation areas near
the entrances (a, b, c) and Blasloch (h)
Figuur 4: (door Robert Seebacher,
Vereniging voor Grottenkunde in
Obersteier 03/2009) Overzichtskaart
van het Brettstein grottencomplex
met opgravingsvelden nabij de
ingangen (a, b, c) en Blasloch (h)
ICBS PROCEEDINGS
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4.0Lab
no
sample
layer
bone
δ13 C
AMS
age
+/-
C/N
%C
%coll
MAMS-14912 SW 773 F5 or F6 / disturbed Humerus F -27.6 34010 210 3.3 16.2 1.7
MAMS-17791 SW 468 F5 / 150-160 cm Vertebra -21.8 37400 290 2.8 19.3 4.9
MAMS-14911 SW 137 F5 or F6 / disturbed Scapula F -27.3 47350 800 3.3 37.0 7.8
MAMS-17797 SW 793 F5 or F6 / disturbed Calcaneus
F juv
-21.3 47820 800 3.3 37.2 2.1
MAMS-17796 SW 140 F5 / wall Costa F -20.8 48270 930 3.2 32.5 1.0
MAMS-17795 SW 791 disturbed Calcaneus F -18.5 > 49000 2.9 34.8 4.5
MAMS-17793 SW 101 F5 or F6 / direct under
speleothem
Calcaneus juv -21.7 >49000 3.2 29.6 10.6
MAMS-17794 SW 137 F5 or F6 / disturbed Vertebra too little
collagen
MAMS-17792 SW 773 H5 / 222 Costa F no results
MAMS-14899 HD 88 Layer 4 / 290-300 cm F-20.7 44130 1530 3.4 19.3 3.6
MAMS-14900 HD 357 Layer 3 / 330-360 cm F-24.9 48530 840 3.1 20.4 3.0
MAMS-14901 HD 561 Layer 2 / 360-370 cm F-23.6 45460 370 3.1 39.7 2.0
MAMS-14902 HD 85 Layer 1 / 380-390 cm F-24.0 46510 410 n. d. 40.5 4.0
MAMS-17801 BS 189 area 6 / disturbed Metapodium F -26.4 21970 70 3.5 38.9 1.0
MAMS-14893 BS 80 area 4 / disturbed Radius F -20.5 22510 120 3.1 39.2 8.0
MAMS-17800 BS 43 area 3 / 50-60 cm Metapodium -22.5 34820 160 3.4 3.8 2.9
MAMS-24067 BS 196 disturbed Cranium F too little
collagen
Table 1: New AMS dating from cave bear bones from the Schwabenreith Cave, Herdengel Cave and Brettstein Cave
Tabel 1: Nieuwe AMS dateringen van grottenbeerbotten uit de Schwabenreith grot, Herdengel grot en Brettstein grot
Abbreviations / Afkortingen: F: fragment; SW: Schwabenreith Cave; HD: Herdengel Cave; BS: Brettstein Cave, C/N: C/N ratio
... Figueirido et al. 2009;Robu et al. 2013;Jones and DeSantis 2016). Cave bears became extinct in the Alps around 26 ka calBP (Döppes et al. 2016). ...
... Radiocarbon dating was done on 28 cave bear bones from nine sites at the Curt-Engelhorn-Centre Archaeometry (CEZA) employing standard sample pre-treatment methods for bone samples, combustion of the samples using an Elemental Analyzer and radiocarbon-determination using a MICADAS AMS (Accelerator Mass Spectrometry) system (see Döppes et al. 2016). Samples with lab codes including 'MAMS' as an identifier are analyzed in that lab and are calibrated by SwissCal1.0 ...
... Brettstein (Brettstein bear cave). This cave is part of the Brettstein cave system located on the karst plateau of Totes Gebirge (Döppes et al. 2016). The most recent radiocarbon dates indicate that the plateau was still ice-free at the time around 26,400 years BP. ...
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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.
... 14 C a BP (VERA 0061 -Pacher, 2000). In a later review, Pacher and Stuart (2009: Döppes et al. (2016Döppes et al. ( , 2018 reported seven AMS radiocarbon analyses. Five of them ranged from as young as 34.0 ± 0.2 to 48.3 ± 0.9k 14 C a BP and two were reported as >49k 14 C a BP. ...
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The cave bear was a prominent member of the Upper Pleistocene fauna in Eurasia. While breakthroughs were recently achieved with respect to its phylogeny using ancient DNA techniques, it is still challenging to date cave bear fossils beyond the radiocarbon age range. Without an accurate and precise chronological framework, however, key questions regarding the palaeoecology cannot be addressed, such as the extent to which large climate swings during the last glacial affected the habitat and possibly even conditioned the final extinction of this mammal. Key to constraining the age of cave bear fossils older than the lower limit of radiocarbon dating is to date interlayered speleothems using 230Th/U. Here we report new results from one such site in the Eastern European Alps (Schwabenreith Cave), which yielded the highest density of bones of cave bear ( Ursus spelaeus eremus). Although dating of the flowstones overlying this fossiliferous succession was partly compromised by diagenetic alteration, the 230Th/U dates indicate that the bear hibernated in this cave after about 113 ka and before about 109 ka. This time interval coincides with the equivalent of Greenland Stadial 25, suggesting possible climate control on the cave bear's habitat and behaviour.
... Our main purpose is to explore patterns of RA across maxillary teeth in the different species/subspecies of the cave bear complex and by extension their inferred feeding behaviours from this ecomorphological indicator. (Ehrenberg 1929;Abel and Kyrle 1931;Kadlec et al. 2001;Döppes and Rosendahl 2009;Döppes et al. 2011Döppes et al. , 2016Döppes et al. , 2018Pérez-Rama et al. 2011;Diedrich 2012;Horacek et al. 2012;Frischauf et al. 2014;Fortes et al. 2016;Kavcik-Graumann et al. 2016;Spötl et al. 2018;Nagel et al. 2018(in press)). As sexual dimorphism among cave bears is well reported (e.g., Kurtén 1955;Grandal d'Anglade and López-González 2005), we sexed our specimens using a protocol detailed in the Supplementary information. ...
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The morphology of both crowns and tooth-roots reflects dietary specialisation in mammalian carnivores. In this article, we analyse the tooth-root morphology of maxillary teeth from CT scans of living bears (Ursus arctos, Ursus americanus, Ursus maritimus, Ursus thibetanus, Melursus ursinus, Helarctos malayanus, Tremarctos ornatus and Ailuropoda melanoleuca) in order to make inferences about the diet and feeding behaviour of the extinct cave bear (Ursus spelaeus sensu lato). Specifically, we investigate two major mitochondrial clades of extinct cave bears recognized by previous authors: Ursus ingressus and Ursus spelaeus (U. spelaeus spelaeus, U. spelaeus ladinicus, U. spelaeus eremus). Our results indicate a close association between tooth-root surface area and feeding behaviour in all living bear species. Tooth-root surface area values of cave bears suggest that they relied more on vegetative matter than living brown bears (Ursus arctos) but subtle differences between these species/subspecies could also indicate different feeding strategies among the members of cave bear complex.
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The Middle Pleistocene – Late Pleistocene transition of European large mammal's fauna (Proboscidea, Artiodactyla, Perissodactyla, Carnivora, Hystrix and Castor) assemblages has been studied in 18 European regional faunal assemblages. This study is based on the data yielded from 423 palaeontological sites (758 localities) dated within interval of MIS 6–MIS 4. All the data was aggregated by 9 time intervals (time scale). For ten bioregions, we have been able to obtain descriptive models of evolution of their faunal assemblages. It allowed detecting common rules of changes in large mammals' fauna composition in Europe on the whole as well as changes in the distribution of individual species and their groups within the regions. We have studied the changes in biodiversity parameters (Shannon index, index of self-organization) and Mourelle–Ezcurra species turnover index within MIS 6–MIS 4 time interval. The evolution of European fauna was compared for MIS 6–MIS 5 transition and MIS 2–MIS 1 transition as well as influence of change in global temperature on these transitions was described. In addition, we have showed the correlation between modern species richness with the species richness in MIS 6, MIS 5 and MIS 4 and proposed the hypothesis of historical succession of European bioregions.
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The variability of lower 6776 cheek teeth of spelaeoid (Ursus spelaeus, U. kanivetz, U. deningeri, U. kudarensis, U. savini and U. rossicus) and arctoid lineages (extant and extinct U. arctos) recovered from 176 palaeontological localities was analysed by methods of univariate and multivariate statistics. For comparison, teeth of U. minimus and U. etruscus species, which are ancestral taxa for cave bears and brown bear, were studied. The performed analyses indicate great variability of tooth crown in lower cheek teeth; no distinct general trend was identified in the changes, except for an increase in p4 roundness and in m3 size from U. minimus to big cave bears. Centroids of U. minimus and U. etruscus are at a great distance from those of cave bears; U. arctos occupies an intermediate position between the two groups. Molar characteristics in general are close in U. savini and U. rossicus. The bear species are differentiated in the morphospaces by morphometric variations in the lower cheek teeth but less markedly than in the morphospaces of the upper cheek teeth. A test of the inhibitory developmental cascade model showed that a linear relationship between lower molar dimensions became clear only after premolar p4 was included in the model. Evolutionary trends were found in the tooth changes with time (differently expressed in individual species and in different teeth) depending on environments, in particular on the elevation of the locality above the sea level. The allometric pattern differentiation does not always coincide with phylogenetic relationships between taxa, being most often revealed at the subspecies level. There is a dominant covariance between the upper and lower rows of teeth in accordance with the occlusion scheme. It may vary in manifestation depending on the particular species.
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Univariate and multivariate statistics were applied to analyse the morphometrical variability of 4920 upper cheek teeth (P4, M1 and M2) of cave bears from 123 geographical sites (180 samples) of different Pliocene – Pleistocene ages. The analysed specimens included those belonging to the big cave bears Ursus kudarensis, U. deningeri, U. spelaeus (three subspecies) and U. kanivetz (including U. ingressus), as well as the small cave bear U. rossicus. The information‐theoretical parameters (Shannon entropy and orderliness (Von Foerster, 1960: On self‐organizing systems and their environments. In Self‐Organizing Systems, 31–50. Pergamon Press, London) were used to estimate tooth diversity in different teeth, different taxa and in selected local chrono‐populations. Multivariate allometry coefficients (Klingenberg, 1996: Multivariate allometry. In Advances in Morphometrics, 23‐49. Plenum Press, New York) were used to describe the relationships of different ‘parts’ of a tooth and to compare allometric patterns amongst species or selected local samples. A multivariate analysis showed a significant overlap of the size/shape parameter ranges in deningeroid and spelaeoid bears within morphological spaces. Within the cave bear lineage, the Deninger's bear has the greatest morphological diversity index (entropy) of all the teeth overall, and the lowest diversity is observed in the final taxon of this lineage – U. kanivetz (=ingressus). The P4 and M2 diversity showed multidirectional correlations with elevation above sea level amongst several ‘local’ populations of Late Pleistocene cave bears. The morphological disparities between the studied taxa are in close agreement with the distances in the available schemes of genetic differentiation based on ancient mitochondrial DNA. The split of U. kudarensis and U. deningeri has a good bootstrap support, which corresponds to the hypothesis about their parallel evolution. The small cave bear U. rossicus is placed between U. arctos and U. deningeri. The phylogenetic signal is more pronounced in the variability of teeth in comparison with other skeletal remains of cave bears (cranium, mandible, or metapodial bones).
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Based on the radiocarbon dating it is very likely that the cave bear fauna from Ajdovska jama can be attributed to the Middle Wurmian warm-period or to the transgression from Early- to Middle Wurmian period. According to the results from the morphodynamic and genetic analyses the bears from Ajdovska jama seem to belong to Ursus ladinicus.
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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.
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The IntCal09 and Marine09 radiocarbon calibration curves have been revised utilizing newly available and updated data sets from 14C measurements on tree rings, plant macrofossils, speleothems, corals, and foraminifera. The calibration curves were derived from the data using the random walk model (RWM) used to generate IntCal09 and Marine09, which has been revised to account for additional uncertainties and error structures. The new curves were ratified at the 21st International Radiocarbon conference in July 2012 and are available as Supplemental Material at www.radiocarbon.org. The database can be accessed at http://intcal.qub.ac.uk/intcal13/. © 2013 by the Arizona Board of Regents on behalf of the University of Arizona.
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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.
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The causes of the late Pleistocene megafaunal extinctions are still enigmatic. Although the fossil record can provide approximations for when a species went extinct, the timing of its disappearance alone cannot resolve the causes and mode of the decline preceding its extinction. However, ancient DNA analyses can reveal population size changes over time and narrow down potential causes of extinction. Here, we present an ancient DNA study comparing late Pleistocene population dynamics of two closely related species, cave and brown bears. We found that the decline of cave bears started approximately 25,000 years before their extinction, whereas brown bear population size remained constant. We conclude that neither the effects of climate change nor human hunting alone can be responsible for the decline of the cave bear and suggest that a complex of factors including human competition for cave sites lead to the cave bear's extinction.
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The quality of bone collagen extracts is central to the14C dating and isotope palaeodietary analysis of bone. The intactness and purity of the extracted gelatin (“collagen”) is strongly dependent on the extent of diagenetic degradation, contamination and the type of extraction method. Possible chemical, elemental and isotopic parameters for the assessment of “collagen” quality are discussed. The most important distinction that can be made is the one between contaminated bone (mostly from temperate zones), and bone low in collagen content (mostly from arid and tropical zones). The latter shows more variability in all quality parameters than the former. The natural level of contamination is mostly so low that stable isotopic measurements are not impaired, although14C measurements can be. It is concluded that there is no unequivocal way to detect natural levels of contamination with the discussed parameters, although their use can identify many cases. In low “collagen” bone, the parameters can identify the great majority of problematic samples: although deviations in these parameters do not necessarily mean isotopic alterations, the increased background found in these samples makes most samples unusable.
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Middle and Late Pleistocene sediments in many caves in Central and South Europe contain large numbers of bones and teeth of the cave bear (Ursus spelaeus). The cave bear differs in many characteristics from the recent brown bear and shows a rapid evolution especially in the changes of the teeth due to adaptation to pure herbivorous nutrition. The shifts of the morphotype frequencies of the fourth premolar from the upper jaw were used as a measure of the evolution. The uranium series method is the only suitable tool for the absolute age determination of the fossil bones with ages beyond the time range accessible to the radiocarbon method. By applying this method to the Herdengel cave profile the evolutionary rate of the cave bears was determined. Uranium series data from the fossil bones were partly verified by an independent carbonate speleothem age. For both, bone layers and carbonate formation found in stratigraphic relation, the determined ages correspond to a normal time sequence. According to the relatively fine time scale obtained by absolute dating, the evolutionary mode of the cave bears was determined as gradual. The main novelty of this study is the dating of the successive layers of the Herdengel cave and the determination of evolutionary stages of the cave bear in them.
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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.