Quaternary International xxx (2013) 1e16

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Quaternary International
journal homepage: www.elsevier.com/locate/quaint

Reconstructing the evolution of cave systems as a key to

understanding the taphonomy of fossil accumulations: The case
of Urşilor Cave (Western Carpathians, Romania)
Silviu Constantin a, *, Marius Robu b, Cristian-Mihai Munteanu b, Alexandru Petculescu b,
guşin a, Dirk Hoffmann d,
Marius Vlaicu b, Ionuţ Mirea a, Marius Kenesz c, Virgil Dra
Valentina Anechitei , Alida Timar-Gabor , Relu-Dumitru Roban g, Cristian G. Panaiotu g
e, f e, f

a ” Institute of Speleology, Center for Geochronology and Paleoclimate, str. Frumoasa

The “Emil Racoviţ a  31, 010986 Bucharest, Romania
b ” Institute of Speleology, Calea 13 Septembrie, 13, 050711 Bucharest, Romania
The “Emil Racoviţ a
c ” Institute of Speleology, Cluj Branch, str. Clinicilor 5, 400006 Cluj-Napoca, Romania
The “Emil Racoviţ a
d
CENIEH, Paseo Sierra de Atapuerca s/n, 09002 Burgos, Spain
e
Faculty of Environmental Science and Engineering, Babeş-Bolyai University, Fântânele 30, 400294 Cluj Napoca, Romania
f
Interdisciplinary Research Institute on Bio-Nano-Science, Babeş-Bolyai University, Treboniu Laurean 42, 400271 Cluj-Napoca, Romania
g
Faculty of Geology and Geophysics, University of Bucharest, N. Bâlcescu 1, 010041 Bucharest, Romania

a r t i c l e i n f o a b s t r a c t

Article history: The Urşilor Cave (NW Romania) is a famous cave bear paleontological site hosting an important Late
Available online xxx Pleistocene faunal assemblage and subject to systematic excavation works. To better understand the
origin of fossil assemblages, the sedimentary history of the cave must be reconstructed. We conducted a
series of investigations on various cave deposits which included sedimentology and grain-size analyses,
U-series dating of speleothems, OSL dating of sediments, and AMS radiocarbon dating of fossil remains.
The results allowed for the identification of several major chronological controls for the evolution of the
cave during the last 300,000 years. Five evolutionary stages or key-moments were dated, and a tentative
speleogenetic scenario is presented. The combined numerical dates and sedimentological study show
that the evolution of the cave was more complex than previously thought. In particular, the fossil
accumulation was related to a succession of rapid flooding events at w47e40 ka. Alternating deposi-
tional and erosional phases have occurred since at least 210 ka complicating the sediment structures. The
results suggest that the upper and lower levels of the cave may have been functioned occasionally as
hydrologically separated karst systems and that the animal populations from what is now a single cave
system may not necessarily be synchronous. This case study shows that reconstructing sedimentary
history of a given cave is crucial for the correct understanding of its thanatocenosis.
Ó 2013 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction hydraulic regimes (Sasowsky and Mylroie, 2004). The associations

Cave deposits are largely recognized as holding the potential of
valuable palaeoenvironmental and paleoclimatic archives (see, for
example, Ford and Williams, 2007; White, 2007). Speleothems are
currently considered as one of the most reliable cave archives for
reconstructing paleoclimate evolution at regional scale (Fairchild
and Baker, 2012). Fluviatile or lacustrine sediments from caves
are also physical and chemical recorders of past climate and
* Corresponding author.
E-mail address: Silviu.Constantin@iser.ro (S. Constantin).
of fossil remains preserved in caves, especially those of micro-
mammals, have been long considered as a paleoclimate proxy.
Moldovan et al. (2011) have shown that even invertebrate fossil
associations from cave sediments may be used as another paleo-
environmental proxy.
The advance of direct and indirect dating techniques (e.g.
Couchoud, 2006; Grün, 2006; Berger et al., 2008) has allowed for a
better understanding of both the timing and the palae-
oenvironmental significance of large fossil accumulations and/or
archaeological sites. However, especially in deep-cave settings, a
key-requirement for understanding the taphonomy of a paleonto-
logical deposits is the correct interpretation of the evolution of the
given cave system. Often (e.g. Zupan Hajna et al., 2008; Trinkaus

1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved.
http://dx.doi.org/10.1016/j.quaint.2013.10.012

Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
2 S. Constantin et al. / Quaternary International xxx (2013) 1e16
et al., 2013), this can only be achieved by the combined study of a
wide range of cave archives and/or the analysis of coeval proxies.
The paleontological excavation of the sediment deposit from the
Scientific Reserve in Urşilor (¼Bears) Cave (Robu et al., 2011) has
revealed the presence of an extensive bone bed including mostly
cave bear remains but also two cave lions and at least one hyaena
and one mustelid. The taphonomic analysis has revealed that the
vertebrate remains may have been accumulated via multiple
mechanisms such as pitfall entrapment, natural death (including
drowning) or fluvial transport. As direct radiocarbon dating has
yielded results that are close to or exceed the limits of the method,
the dating of complementary cave deposits and the understanding
of the speleogenetic scenario are crucial for taphonomical
interpretations.
With only a few exceptions (Constantin, 2003; Petculescu and
Murariu, 2009; Häuselmann et al., 2010; Trinkaus et al., 2013)
most studies on the evolution of karst systems from the Romanian
Carpathians rely only on stratigraphical and geomorphological
evidence. Urşilor Cave is no exception, with the earlier attempts to
reconstruct the evolution of the cave and the sedimentation
mechanisms and chronology (Hadnagy, 1977; Terzea, 1978, 1989;
Valenaş, 1979; Jurcsák et al., 1981) relying mostly on the analysis
of the mineralogy of sediments, cave morphology, and a partial
paleontological record.
This study focuses on the establishment of a chronological and
evolutionary framework for the Urşilor Cave in the last w300,000
years. It integrates the study of relevant cave deposits (cave sedi-
ments, speleothems, and fossil bones) which were directly or
indirectly dated by means of radiocarbon, U-series, and optically
stimulated luminescence (OSL). It also integrates new, detailed,
geomorphological and topographical surveys of the cave that are
relevant to its speleogenesis. The aim of the study was to provide a
solid chronological and paleoevolutive basis for the taphonomical
interpretation of the fossil accumulation from the Scientific Reserve
of Urşilor Cave.
2. Background
Urşilor Cave is located on the western slope of the Apuseni
Mountains (Western Carpathians, Romania), on the left side of the
Craiasa Valley, at 491 m a.s.l (Fig. 1). The cave was discovered in
1975 by blasting in a local marble quarry. Owing to both its rich and
complex Upper Pleistocene thanatocenosis, and the abundance of
speleothems, the cave was immediately protected, studied and
surveyed over two years. Since 1980, the upper part of the cave was
set up for tourism, becoming the most important show cave of
Romania.
The cave is carved in the Upper Jurassic (Tithonian) recrystallized
limestones of the Bihor Autochthonous unit, which is partially
overlain by the Arieşeni and Codru Nappes, respectively (V alenaş,
1979) (Fig. 1). The cave comprises two distinct levels, separated by
a 17 m high steep slope, and consists of 1500 m of large and hori-
zontal passages. The upper level, hydrologically inactive, consists of
five main passages (the Access Passage, The Candlesticks Passage,
The Twisted Passage, The Passage of the Pits, and the “Emil Racoviţa”
Passage, see Fig. 2) and a large chamber (The Intersection Chamber,
w10  15  25 m), totaling c. 1000 m of galleries. The two artificial
entrances correspond broadly with two former natural entrances
that were sealed by sediments and flowstone precipitation.
In the final part of the “Emil Racoviţa ” Passage, a smaller,
ascending passage leads to a breakdown cone formed by blocks of
greywacke and flowstone fragments embedded in a clay matrix.
The overall morphology of this cave sector suggests the presence of
a local fault with the debris originating from the surrounding non-
karst deposits. This passage ends with a 17 m-drop (the Shaft)
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
(Fig. S1) that leads to a small chamber (the Excavation Chamber)
and the Scientific Reserve.
The Scientific Reserve is the lower level of Urşilor Cave (w500 m in
length) and includes a generally large horizontal passage (10  15 m),
with several small side-passages and fossil meanders (Fig. S2). A small
stream (1e10 l/s) flows through the lower level and sinks into an
impenetrable sump in the Excavation Chamber. Along the lower
passage, the stream has deposited alluvial sediments consisting of
fine sands, silts, and clays which often support stalagmite formations
on their top. Currently the stream meanders through w0.5e3 m-high
alluvial terraces. In the Excavation Chamber, it sinks and flows
through an accumulation of sediments that exceeds 200 m3 of a mix
of sandy-clayish alluvia and fossil bone beds.
Both cave levels host important cave bear bone assemblages
(both on the surface and within the sediment, scattered or fully
articulated specimens) and cave bear ichnofabrics (nests, scratch
marks, footprints and fur imprints). However, most articulated
skeletons and ichnofabrics are found in the Lower Level whereas in
the Upper Level the fossil remains are more scattered and/or
covered by massive speleothem formations.
The genesis of the cave system has been largely interpreted by
corroborating observations on the overall cave morphology and
correlations of the different cave levels with the local river terraces
(Valenaş, 1979; Rusu, 1981; Rusu and Racoviţa , 1981). In broad
terms, the evolution of the cave was thought to include the
following stages:
(1) an initial karst drainage towards the Cr aiasa Valley led to
the formation of the Upper Level passages. This stage in-
cludes successive abandonments of older passages, in pace
with the deepening of the base level. The Twisted Passage
was abandoned in favour of a new resurgence via the
Candlesticks Passage which was, in turn, abandoned for a
new drainage via the Access Passage.
(2) the capture of the subterranean stream towards the end of the
“Emil Racoviţa” Passage. A new drainage was established to-
wards the Peştera de la Chişcau, a small (148 m) resurgence
cave located w40 m below the current entrance of Urşilor
Cave (Figs. 1 and 2). The greatest part of the passages between
the sink and Peştera de la Chişc au are impenetrable and
thought to be generally small in size. This may have caused
periodic accumulations of water upstream the sink and the
deposition of alluvial material throughout the Lower Level.
(3) during the Holocene, the massive speleothem deposition
throughout the Upper Level has sealed the former cave en-
trances and largely covered the old sediments, and bone
remains. The Holocene age of the typical “candlestick” sta-
lagmites from the Upper Level was also indicated by subse-
quent studies (Onac et al., 2002; Constantin S. and Onac B.P.,
unpublished data). No other age controls existed for the
previous evolutionary stages.
3. Studied profiles
3.1. The Excavation Chamber
The new paleontological excavation at Urşilor Cave is located at
the northern (downstream) end of the Excavation Chamber (Fig. 3).
The Excavation Chamber (w20 m  6e10 m) reaches a maximum
height of w20 m directly below the Shaft. At its southern end, the
underground stream sinks laterally at the base of the left wall. The
small passage is impenetrable for explorers but it is evident that the
stream flows at the limit between the limestone wall and the
sediments accumulated in the chamber. The stream may be
 S. Constantin et al. / Quaternary International xxx (2013) 1e16 3

Fig. 1. Karst map in the Urşilor Cave area (simplified, after Valenaş, 1979; base geological map after Bordea et al., 1975). Legend: 1. Quaternary alluvial deposits; 2. Upper Miocene
(Pannonian) sediments; 3. Karst forming deposits: J3 ¼ Late Jurassic (massive limestones, partially recrystallized); T3cr ¼ Carnian limestones; T2lneT3cr ¼ Upper Ladinian-Lower
Carnian black limestones; T2an-ln ¼ dolomites; 4. Impervious (non-karst) deposits: J1e2 and T1 ¼ quartzitic sandstones; P ¼ Permian feldsparic sandstones; 5. Cave passages; 6. a:
Cave entrance; b: Pothole; 7. Sinkhole; 8. Cliff; 9. Karst spring; 10. Overthrust; 11. Surface rivers: a. perennial; b. temporary; 12. Settlement. Inset: Map of Romania with the location
of the Urşilor Cave indicated by the arrow.

reached through a narrow passage at the northern end of the
chamber but becomes again impenetrable after w50 m. The walls
of the chamber are largely covered by massive flowstone forma-
tions; a thin film of clay covers the walls up to w4 m in height,
indicating a former stagnant water level.
The floor of the chamber is covered by brown clay with both
scattered cave bear remains and partially articulated skeletons.
Toward the walls, some of these remains are encrusted in calcite.
The thickness of the sediment deposit can only be assessed in the
southern end, where it forms a 1.8 m-high step (Fig. 3).

Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
 4
j.quaint.2013.10.012
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of

S. Constantin et al. / Quaternary International xxx (2013) 1e16

Fig. 2. Map and long profile of the Urşilor Cave, with the location of the profiles and sampling points discussed in text. Rectangles (a)e(c) are shown in detail with the location of sampling points and obtained ages. Legend:
squares ¼ UeTh dates; circles ¼ OSL dates; diamonds ¼ AMS 14C dates. All figures rounded to one significant decimal for simplicity. Ages in square brackets are considered as chronologically unreliable, but significant for the
, 1981. All other cave passages were re-surveyed by our team.
evolutionary stage. Show cave sector redrawn after Rusu and Racoviţa
 S. Constantin et al. / Quaternary International xxx (2013) 1e16
Fig. 3. Sketch of the Excavation Chamber with the position of the excavation trench
and the U/Th dated calcite samples.
The excavation is still in progress and the detailed paleonto-
logical and taphonomical results will be presented elsewhere. Here
we will only summarize the stratigraphy of the excavation and the
surrounding deposits that were sampled for dating purposes as well
as for sedimentological, geochemical, and rock-magnetic studies.
The excavation trench is “L” shaped, has a total area of 7 m2 and
a current depth of w1.6 m without reaching the bedrock (Fig. S3).
Eight distinct sediment layers (labeled L1-8) were found, with L1,
L3, and L5 being very rich in cave bear remains while the remaining
layers are sterile. The bone beds extend across the whole section
while the lower sterile ones were thicker along the A1-D1 transect
and become thinner towards the D4 square, where they lie over a
flowstone terrace. The transects reveal a channel facies that cor-
responds to the general direction of flow of the subterranean
stream (A1eD1) and a cave terrace facies towards the sides (D1e
D4) consisting of interbedded cave sediments and flowstone above
L5 (Fig. 4).
The excavation yielded a large number of cave bear bone re-
mains (both articulated and scattered), as well as one articulated
cave lion skeleton (Fig. S4). Twelve cave bear bone fragments were
sampled for radiocarbon dating. In addition, sediments from two
layers (L4; 100 cm and L5; 150 cm) were sampled for OSL dating.
In the northern side of the Excavation Chamber, a flowstone
pavement suspended w0.5 m above the floor (Fig. S5) indicates
deposition on top of a former alluvial terrace, subsequently
removed by erosion. A flowstone sample was collected from this
speleothem for U-series dating. In addition, we collected two other
samples from the flowstone that covers both the wall and the top of
the Shaft (Fig. 3).
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
5
In the southern end of the chamber, a massive stalagmite has its
basis engulfed in sediments but has a lateral calcite overgrowth on
top of those sediments. The sediments were sampled (Profile PU4)
for sedimentological, geochemical, and rockmagnetic in-
vestigations. Profile PU4 (Fig. S6) consists of a 2.3 m thick suc-
cession of brown, laminated, poorly sorted muds, sandy muds and
muddy sands. Sediment samples were taken at every 5 cm. In
addition, we took a sample of the calcite sinter overlying them and
one sediment sample for OSL dating from a depth of 115 cm.
3.2. Subterranean terraces profiles
Along the subterranean stream, two terrace profiles were also
sampled for sediments (Fig. 2c). Such cut-in-fill terraces are com-
mon all along the Lower Level and they are typically formed by a
mixture of brown, laminated, clay- and silt-size sediments. The top
surfaces of the terraces are generally flat and their elevation with
respect to the river bed matches that of the infill deposit in the
Excavation Chamber to gradually decrease upstream. Scattered
fossil remains are frequent and cave bear traces (footprints,
scratches, hibernation nests, etc.) are common on terrace top sur-
faces, terrace bluffs, and passage walls. The most remarkable finds
are those of two cave bear articulated skeletons (an adult and a cub)
, 1981) and one of an artic-
(Jurcsák et al., 1981; Rusu and Racoviţa
ulated cave lion skeleton, all three lying on top of the left-bank
terrace deposits at w170e200 m upstream of the Excavation (see
Fig. 2c; Figs. S8 and S9) and thus postdating the last depositional
event. The presence of undisturbed skeletons on top of alluvial
terraces together with the good preservation of footprints and
scratches on terrace faces suggests that any subsequent cave floods
must have been of lesser magnitude than those responsible for the
deposition of those terraces.
The PU2 Profile (Fig. 2c; Fig. S10) is located w65 m from the
Excavation, on a left-side, cut-bank terrace. The profile is 2.5 m high
and consists of brown, laminated, poorly sorted muds and sandy
muds with coarser units at the base of the profile. The top of the
profile consists of a flat surface that preserves ichnofabrics (floor
scratch marks, cave bear nests, partially preserved footprints), and
its elevation corresponds to that of the neighboring terrace surface
where the fully articulated cave bear skeletons can be found. As in
the case of PU4, sediment samples were collected at 5 cm intervals.
In addition, three sediment samples were taken for OSL measure-
ments: at 70 cm, 155 cm, and 270 cm from the terrace top,
respectively.
The PU1 Profile (Fig. 2c; Fig. S11) is located w135 m upstream
from the Excavation, at the point bar of an underground meander,
in the right-side wall of the passage. The thickness of the deposits
is 1.55 m, slightly lower than that of the opposite cut-terrace. The
deposits consist of brownish, laminated, poorly sorted muds. The
top of the sequence consists of the same flat surface with cave
bear nests, bones and scratch marks. The profile’s position is
opposite to the terrace where the cave bears and lion articulated
skeletons are found. The top levels can be considered coeval
although (owing to local cave morphology) the sediment stack on
the left bank is w0.5 m higher. Sediment samples were collected
along the profile at 5 cm intervals. One sediment sample for OSL
dating was collected at 75 cm below the top. In addition, four bone
samples for radiocarbon dating were collected from the articu-
lated skeletons of cave bears and the lion resting on top of the
opposite terrace.
3.3. Additional samples (the Upper Passage)
Additional bone samples were collected for radiocarbon dating
from two significant locations in the Upper Passage:
6 S. Constantin et al. / Quaternary International xxx (2013) 1e16

1) The Passage of the Pits is an intricate network of horizontal
passages and shafts starting in the “Emil Racoviţ
a” Passage and
descending to w3 m above the current level of the underground
stream (Fig. 2b; Fig. S12). Based on the phreatic morphology of
the passages, the sediment fill and the overall topography of the
cave, this passage is considered to have functioned as an over-
flow of the waters that sank in the Excavation Chamber.
Numerous cave bear remains, including one partially articulated
skeleton, have been found in this passage and three samples
were taken for dating.
2) An old test-pit excavated in the Access Passage during the initial
research in the late ‘970s (Fig. 2a; Fig. S13) was reopened and
two cave bear bone samples were taken from the freshly cleaned
wall of the pit at depths of 0.12 and 1.1 m, respectively. The
sediments here are much coarser that in the lower level and
they consists of limestone pebbles and clay indicating a highly
dynamic depositional environment.
4. Analytical methods
14
4.1. AMS C dating
Most of the AMS 14C dating was done at the Poznan Radio-
carbon Laboratory (Poland) (8 samples) and Centro di Datazione
e Diagnostica (CEDAD) (University of Salento, Italy) (11 samples).
One sample was dated at the Vienna Environmental Research
Accelerator (VERA) and two samples were dated at the Oxford
Radiocarbon Accelerator Unit (ORAU) and reported by Stuart and
Lister (2011). The analytical methods for collagen extraction,
graphitization, and measurement of 14C/12C and 13C/12C ratios are

Fig. 4. Stratigraphic section of the excavation trench in Urşilor Cave along the D1eD4 transect (top) and photograph showing the main identified levels (bottom). Note that L2, L6,
and L7 are sterile levels (laminated clays) and L8 consists of flowstone. L4 is a debris cone with sediments originating from the upper part of the Shaft which explains the apparent
OSL age reversal with respect to L5. Also note the discrepancy between the OSL ages which are indicative for the time of sediment input into the cave and the radiocarbon ages of
fossil remains (Fig. 7) which date the depositional events.

Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
 S. Constantin et al. / Quaternary International xxx (2013) 1e16
described in detail on the websites of the respective laboratories.
Basically all laboratories use the collagen extraction procedure
originally described by Longin (1971), although with some
modifications. At the Poznan Laboratory, collagen extraction was
performed using the method described by Pietrowska and Goslar
(2002) and the extracted collagen is then purified on Elkay 8 mm
filters, and ultrafiltered on Vivaspin 15 MWCO 30 kD filters
(Bronk Ramsey et al., 2004). Content of 14C in carbon samples
was measured following the protocol described by Goslar et al.
(2004). A preliminary step includes a check for collagen degra-
dation by measuring %C and %N in whole bone using a Flash
EA1112 Series analyser (Thermo Scientific). The samples are
regarded suitable for collagen dating if nitrogen content in bone
is not lower than 1%. The CEDAD laboratory does not apply an
ultrafiltration step (Calcagnile et al., 2004, 2005) and, at the time
of analysis, it did not measure either %C or %N. The Oxford and
VERA collagen extraction procedures are similar, although
treatment procedures are slightly different (Higham and Wild,
2013). None of these laboratories measures %C and %N in
extracted collagen as a routine, and hence the atomic C:N ratio
cannot be calculated.
4.2. U-Th dating
Speleothem calcite subsamples were hand drilled, weighed,
placed in clean Teflon beakers and covered with MQ H2O. The
powders were then dissolved by stepwise addition of 7M HNO3 and
subsequently spiked using a mixed 229The236U spike. The solutions
were evaporated to dryness and re-dissolved in a mixture of H2O2
and concentrated HCl and HNO3 to remove organic matter. The
solution was boiled at 130  C, and evaporated to dryness again,
before the separation and purification procedures. Separation and
purification of U and Th fractions was performed using 500 ml ion
exchange columns and BioRad AG 1X8 resin. U and Th fractions
were dissolved in 1.8 ml 2% HCl and analysed on a Neptune ICP-MS
equipped with a Cetac Aridus nebulizer at CENIEH (Burgos, Spain)
following the procedures outlined in Hoffmann et al. (2007) and
Hoffmann (2008).
4.3. OSL dating
Sediment samples for OSL dating were collected by hammering
25 cm-long, opaque, plastic tubes (10 cm) into the sediment sec-
tions under no light conditions. Sample preparation for lumines-
cence measurements was performed under low intensity red light
conditions. Only the central portion of the tubes was removed for
dating. A three-day treatment with 10% HCl solution was used for
carbonate removal, followed by another three-day H2O2 (30%)
treatment for organic matter removal. The coarse grain fractions
(>63e90 mm) were separated through wet sieving. For the
extraction of the fine-grained quartz fraction (4e11 mm), the par-
ticles smaller than 11 mm were isolated from the fraction less than
63 mm by settling in Atterberg cylinders, according to the Stokes
law. This fraction was treated with hexafluorosilicic acid (H2SiF6)
for 10 days. Subsequently, the removal of the grains <4 mm was
carried out by centrifugation.
For the annual dose determination, radionuclide specific activ-
ities (U-238/Ra-226, Th-232, K-40) were measured using high
resolution gamma-ray spectrometry, and the dose rates were
calculated using the conversion factors tabulated by Adamiec and
Aitken (1998). Equivalent dose (De) measurements were under-
taken on the standard Risø TL/OSL-DA-20 reader, equipped with
blue LEDs (470  30 nm). IR (875  80 nm) LEDs were used for
infrared stimulation. Blue light stimulated OSL signal was detected
through a 7.5 mm thick Hoya U-340 UV filter. Irradiations were
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
7
carried out using the incorporated 90Sre90Y radioactive source,
calibrated against gamma dosed calibration quartz supplied by the
Risø National Laboratory, Denmark. A dose rate of 0.123 Gy/s for the
fine grains mounted on aluminium disks was obtained at the time
of measurement.
The measurement protocol was the Single Aliquot Regeneration
(SAR) applied on quartz (Murray and Wintle, 2000, 2003; Wintle
and Murray, 2006) (see Supplemental Information for a complete
description of the protocol). The OSL dating was done at the
Dating and Luminescence Dosimetry Laboratory, Babeș-Bolyai
University, Cluj-Napoca, Romania (see also supplemental material
for details).
4.4. Analysis of the sediment profiles
The sediments from the Excavation Chamber and the subterra-
nean terraces were collected from vertical trenches created by
removing w5 cm of the outer sediment layers. Duplicate sediment
samples were taken for: (1) granulometry, geochemistry, and rocke
magnetic properties measurements, and (2) screening of inverte-
brate fossil fauna, respectively.
Grain size measurements on fine samples were performed by
treating w5 g of the bulk sample, for 14 days, in a plastic box, with
w0.4 ml of a 1% solution of Na(PO3)n, n z 25 e Graham’s salt
(Merck). A quantity of w2.5 g sample was later extracted from the
box and treated again with w0.2 ml of 2% solution of Graham’s
salt. Each sample was analyzed on a HORIBA Partica LA-950V2
laser scattering particle size distribution analyzer to reveal grain
size fractions. The coarser samples were analyzed by vibrating
dry sieving of w100 g of the bulk sample and weighing the
sediment quantity retained on each sieve, on an OHAUS Scout
digital balance, down to the 500 mm fraction, which was subjected
to the same procedure as the fine samples. Calculations and
plots were done using the GRADISTAT 8 software (Blott, 2010). We
applied the Folk and Ward (1957) method, and logarithmic sta-
tistics. The magnetic properties, elemental geochemistry and
invertebrate fauna finds are still under investigation: they are not
significant for the scope of this paper and will be presented
elsewhere.
5. Results and discussion
5.1. U-series dating results
The results of the four flowstone samples dated from the
Excavation Chamber are given in Table 1. The samples have a low U-
content, ranging between 0.06 and 0.15 ppm and three of them
show low 230Th/232Th activity ratios ranging between10 and 35,
thus requiring age corrections for detrital Th.
The PU10-1 sample was a small, corroded, stalagmite that was
growing on top of the flowstone deposit directly above the shaft.
Its corrected age of c. 74  0.8 ka and its internal structure in-
dicates only that at that time there was already no water flow in
the upper levels of the cave. However, this has little significance
when considering the age measured for PU10-2, which was taken
from the outer flowstone crust layer that covers one of the walls of
the Shaft (Fig. 3). The sample yielded a reliable age of w291  9 ka,
which indicates that at that time the Shaft was already formed (i.e.
there was a relief of at least 15 m between the subterranean
stream and the upper passages).
Down in the Excavation Chamber, the age of the PU10-3
sample is important because it was obtained from the basal
part of a flowstone slab that is now suspended at w0.5 m above
the current excavation surface (Fig. S5). The sample is an ancient
flowstone that grew over a sediment stack that was
8 S. Constantin et al. / Quaternary International xxx (2013) 1e16
Table 1
Results of the U-series dating of speleothem samples. Isotopic ratios are given as activity ratios, errors are at 2s. Ages are corrected using
230
Th/238U ¼ 1  0.02 and 234U/238U ¼ 1  0.02.
238
Sample Lab number Description U [ng/g]
number
PU10-1 GEB-UTO-6 Small stalagmite; base 133.16  0.73 1.414
PU10-2 GEB-UTO-7 Flowstone crust on the 149.74  0.87 1.044
step; basal part
PU10-3 GEB-UTO-8 Flowstone at the excavation; 62.44  0.54 1.276
basal part
PU10-4 GEB-UTO-9 Stalagmite outer layer topping 81.63  0.47 1.514
sediments
subsequently removed. Its minimal age of c. 92  1.8 ka attests
for flowstone deposition process being active towards the end of
MIS 5 with flowstone crusts covering more ancient detrital
deposits.
Finally, in the opposite end of the Excavation Chamber, the
lateral flowstone was deposited aside a massive column and on top
of old silt deposits (sample PU10-4, Fig. S6) has a minimal age of
w146  6.8 ka. During sediment sampling, the old base of the
stalagmite, including a second such crust, laterally deposited, was
discovered buried in sediments at w0.9 m below (Fig. 5 and Fig. S6).
The complex structure of the speleothem indicates that there were
at least two ancient episodes of sediment accumulation in the
Excavation Chamber followed by periods when water level was low
enough to allow growth of stalagmites and flowstone. The second
such event took place some time before the end of MIS 6. Consid-
ering: (1) the suspended position of the PU10-3 flowstone and (2)
the current morphology of the outcrop, with the PU10-4 stalagmite
base partially exposed, we speculate that these accumulation
events were followed by periods of fluvial erosion and sediment
removal.
5.2. Cave sediments and OSL ages
Cave sediments successions studied from the three profiles in
the Scientific Reserve are presented in Fig. 5. Measured grain size
and statistical parameters are presented in Table 2 (see also Fig. S7).
In all profiles, the sediments predominantly include silt-size sedi-
ments which are finning towards the upper parts. The PU1 and PU2
upper parts consist of sub-horizontal plateaus showing ichnofa-
brics (cave bear prints, scratches, and nests) and, in the case of PU2,
two fully articulated cave bear skeletons.
For the PU1 profile, the statistical parameters calculated for
the grain size distribution of sediments show two very strong
positive correlations between median and mean (Md/Mzi) (r:
0.9692; P < 0.0001), and between median and sorting (Mzi/si) (r:
0.9275; P < 0.0001), the latter showing a lower degree of sorting
for the finest fraction. Positive correlations were calculated be-
tween the median and the skewness (Mzi/Ski) (r: 0.7933;
P < 0.0001) and the sorting and skewness (si/Ski) (r: 0.7119;
P < 0.0001), respectively. These indicate an excess of fine detritus
with a low degree of sorting, owing to an increased input of lutite
fraction (% lutite/Ski: r: 0.7464; P < 0.0001). The C/Md plot
Table 2
Grain size statistical parameters for the reference sediment sections. Sampling interval: 5 cm.
Profile Number of Clay (%) Silt (%) Sand (%) Clasticity index Mode 1 Mo1 (ø) Median Md (ø) Mean Mz (ø) Sorting si (ø) Skewness (Ski) Kurtosis (KG)
samples (N) C (ø)
PU1 31 0.12e43.87 56.13e89.1 0e9.6 2.13e6.04
PU2 50 0e68.79 29.83e96.93 0e59.01 0.56e6.43
PU4 46 0e40.09 20.02e92.15 1.08e77.04 0.76e4.08
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
232
Th/238U ¼ 1.250  0.625,
234
U/238U 230
Th/238U 232
Th/238U 230
Th/232Th Calculated age Corrected age
(ka) (ka)
 0.003 0.725  0.005 0.0202  0.0001 35.73  0.26 75.153  0.692 73.975  0.812
 0.002 0.985  0.006 0.0051  0 192.79  1.31 291.496  9.143 291.090  9.146
 0.003 0.771  0.004 0.0649  0.0002 11.88  0.06 96.694  0.921 92.450  1.814
 0.004 1.212  0.006 0.1202  0.0003 10.08  0.04 152.323  1.781 146.2696.822
(Passega diagram) (Fig. 6) displays a scattered pattern for Profile
PU1, suggesting a variable environment energy. However, most
values are parallel to the C ¼ Md line indicating that “graded
suspension” have been the main transport mechanism. The
Estimated Settling Velocity (ESV) of the particles varies between
0.58 and 81.77  103 cm/s. All parameters suggest a back-
swamp facies (sensu Bosch and White, 2004), with quasi-
stagnant water and short transitions to a channel facies,
marked by the higher sorting degree of several samples and
indicating a more dynamic water.
In the PU2 Profile, the values of Mo1, Md and Mzi are strongly
influenced by the percentage of lutite-sized clasts (% lutite/Mo1: r:
0.8614; P < 0.0001; % lutite/Md: r: 0.9105; P < 0.0001; % lutite/Mzi:
r: 0.9437; P < 0.0001). The sorting degree of the deposits increases
with the content of silt fractions (% silt/si: r: 0.8431; P < 0.0001)
and decreases with the content of lutite fractions (% lutite/si: r:
0.7594; P < 0.0001). A correlation between the main grain size
statistical parameters could not be established, possibly owing to a
more variable depositional environment. This variability is also
indicated by the ESV range of 0.01e5348.58  104 cm/s, as well as
by the very scattered projections of the PU2 Profile on the Passega
diagram. The sediment transport regimes correspond to either
“suspension or rolling” or “graded suspension” mechanisms.
All the above suggests a backswamp facies, with transitions to a
channel facies, the latter being probably caused by short increases
of the water dynamics. The grain size of the sediment supplied to
the cave has varied, possibly owing to changes within the source
area (one such event can be noted w1.6 m from the top and it is
marked by opposite trends for silt and arenite, and by higher values
of Mo1 and ESV).
Profile PU4 shows a higher dynamic depositional environment,
with no clear correlations between the main grain size statistical
parameters. The values of the clasticity index increase with the silt
fractions content (% silt/C: r: 0.7530; P < 0.0001), while those of the
first mode and mean increase with the lutite fractions content (%
lutite/Mo1: r: 0.8028; P < 0.0001; % lutite/Mzi: r: 0.8111;
P < 0.0001); the values of the clasticity index, median and mean
decrease with the arenite-sized clasts percentage (% arenite/C:
r: 0.8041; P < 0.0001; % arenite/Md: r: 0.9105; P < 0.0001; %
arenite/Mzi: r: 0.7882; P < 0.0001). The sorting degree of the
deposits decreases with the lutite fractions content (% lutite/si: r:
0.9240; P < 0.0001). The fluctuating environment energy is also
4.77e12.21 5.06e8.31 5.18e9.03 0.91e3.21 0.03e0.61 0.49e2.24
3.2e13.39 3.61e12.65 4.21e11.19 0.8e4.49 0.73e0.64 0.51e2.68
2.03e13.39 2.44e7.24 3.13e8.45 0.93e4.13 0.11e0.62 0.54e1.96
 S. Constantin et al. / Quaternary International xxx (2013) 1e16 9

Fig. 5. Stratigraphic logs of the three sediment profiles from the Scientific Reserve and tentative correlations between the different sediment levels. The dots indicate the position of
OSL ages, the diamonds that of the 14C ages, and the square indicates the UeTh age. OSL ages given in square brackets are considered less reliable but indicative for an older stage.
Radiocarbon ages for the top of the sediments are conservative estimates calibrated at 2s confidence level.

illustrated by the projections’ scattering on the Passega diagram
and by the ESV, which has values varying across a very large range:
3.19e1961.06  103 cm/s. The high variability of the ESV indicates
the occurrence of some water pulses. The projections on the Pas-
sega diagram (Fig. 6) indicate that sediments were transported as
bedload, mainly by either “rolling and suspension” or “suspension
and rolling”, and only secondarily by “graded suspension”. Similarly
to Profile PU2, in PU4 the sedimentological parameters point to an
overall backswamp facies, with short transitions to a channel facies,
caused by a higher dynamics of the depositional environment.
Overall, the sedimentology of the three investigated sediment
profiles shows a consistent pattern, with a typical backswamp
facies that includes transitions to channel facies during high-
energy hydraulic events. Evidence for such events increase from
PU1 through PU4 suggesting that the upstream sections were
mostly deposited at high water levels with low hydraulic energy
(reservoir lake) while the downstream sections experienced
alternating episodes of high water levels which deposited fine
sediments and torrential events that deposited coarser fractions
and might have eroded through or even wash-out the older
sediments.
The results of the OSL dating carried out on seven sediment
samples are presented in Table 3. The obtained ages clearly cluster
in two ranges. The first group shows theoretical ages of 210e
230 ka, with typical uncertainties of >10% (1s). These ages seem
to be unreliable. Legitimate doubts concerning a potential
incomplete bleaching of the quartz grains can be raised according
to the more general belief that quartz OSL dating for old (>70 ka)

Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
10 S. Constantin et al. / Quaternary International xxx (2013) 1e16

Fig. 6. The Passega diagram of the sediments from the underground terraces of the Lower Level in Urşilor Cave. The projections’ scattering degree increases with the depositional
environment energy, from Profile PU1 to profiles PU2 and PU4, as the sediment transport mechanisms diversify.

sediments is unreliable (Chapot et al., 2012; Timar-Gabor et al.,
2012). However, it is interesting to note that these ages are in
agreement with both the U/Th age of the PU10-4 flowstone (Fig. 5
and S6), as well as with the U/Th age of the PU10-2 flowstone that
grew onto the Shaft’s wall at w290 ka. We can only speculate that
the bottom sediments were deposited at some early stage, whose
age is about 210e230 ka.
The second group of ages belongs to the top of the sediment
sections and to the sediments in the excavation trench. They show
calculated OSL ages that cluster between w67 and 90 ka. These ages
appear to be more reliable, and they generally follow the strati-
graphical sequence. The ages measured for the Levels 4 and 5 of the
Excavation pit seem to be: (i) out of stratigraphical sequence (L5
yielded a younger age than L4) and (ii) in disagreement with the
radiocarbon ages of the incorporated fossil remains which are
younger than w47 ka (see Section 5.3). However, as L4 is a debris
cone originating from the sediments in the upper levels of the Shaft
(Fig. 4), its older OSL age is explicable.
We therefore consider that the top deposits within the profiles
indicate a secondary sediment inflow phase with sediments being
transported into the cave as early as w90 ka. The contradiction
between the OSL ages of sediments and the radiocarbon ages of the
incorporated fossil bones may be due to either: (i) an incomplete
reset of the OSL signal at the time of transport, or (ii) a reworking of
older sediments from the remote parts of the cave towards their
current position. Overall, although the OSL measurements did not
yield a precise chronology of the sediment sections, they suggest at
least two separate episodes of rapid sediment inflow into the cave.
5.3. Radiocarbon dating results
The results of radiocarbon dating are given in Table 4. From a
total number of 25 submitted samples, we discarded the results
that either show abnormally low %N (<0.5%) or collagen yields
lower than 0.8%. All ages were calibrated using the OxCal
4.2.2 program (Bronk Ramsey, 2009) and the IntCal09 calibration
curve (Reimer et al., 2009). The results are graphically shown in
Fig. 7.
Radiocarbon ages of the 15 fossil bone samples collected from
the Scientific Reserve range between c. 42 and 45 ka (median
values) for both surface and excavated finds. We have discarded
two samples taken from the lower part of the excavation (100 to

Table 3
OSL age dating results of the sediment samples. L4 and L5 are samples taken from sediments in the excavation’s Level 4 and 5, respectively. See Fig. 5 for the positions of the PU1
through PU4 samples. Alpha efficiency factor considered was 0.04  0.02. All uncertainties are 1s.

Sample number Location (cm from top) Equivalent Anual dose Water content (%) Age and total Random Systematic
dose (Gy) (Gy/Ka) error (ka) error (%) error (%)

L4 Excavation trench: 100 223  6 2.48  0.03 49  4.9 89.6  9.4 2.9 10.1
L5 Excavation trench 150 194  3 2.74  0.03 30  3 70.9  6.7 2.9 9.25
PU2-1 PU2: 270 563  28 2.47  0.03 68  6.8 228  27 5.2 10.8
PU2-2 PU2: 155 535  34 2.53  0.04 58  5.8 211  25 6.4 10.0
PU2-3 PU2: 70 208  3 3.11  0.04 36  3.6 66.9  6.3 1.8 9.2
PU1 PU1: 75 231  3 3.08  0.04 35  3.5 75.0  7.2 1.67 9.5
PU4 PU4: 115 515  19 2.35  0.03 33  3.3 219  21 3.9 8.9

Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
 S. Constantin et al. / Quaternary International xxx (2013) 1e16
Fig. 7. Radiocarbon ages of fossil bones from the Urşilor Cave, calibrated using OxCal
4.2.2 (Bronk Ramsey, 2009) and IntCal09 (Reimer et al., 2009). The NGRIP curve is
shown for reference with the Heinrich (H) events marked.
170 cm) that showed abnormally young ages. These samples also
show abnormally low %C with respect to %N measured for whole
bone (%Cb and %Nb). Normally, the %Cb would also include the “non
collagenous” carbon, which can be calculated using the equation %
Cncoll ¼ %Cb  (2.7  %Nb) (Bocherens et al., 2005, 2008). However,
many samples for which %Cb and %Nb was available show a
negative %Cncoll and the two rejected samples show %Cncoll
ranging between 1.07 and 1.87. Such a depletion in non-collagen
carbon was associated with an almost total loss of collagen
(Bocherens et al., 2008) but this does not seem to be the case here,
when considering the measured %Nb > 2. Considering: (i) the
possibility of preferential carbon leaching; (ii) the old age of the
samples, and (iii) the apparent stratigraphical inversion, we
decided to discard the samples which show a %C depletion higher
than 1%. Nevertheless, the issue of fossil samples that show such an
unexpected negative departure of %Cb when compared to %Nb re-
quires further attention and study.
Collagen reported yields (not available for the CEDAD labora-
tory) were generally low (0.8e7.5%), as typical for many bone ac-
cumulations from deep cave settings (e.g. Zilhao et al., 2007; for the
Peştera cu Oase). Taking into account that the ages are close to the
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
11
limit of radiocarbon, the calibrated values show relatively high
analytical uncertainties. However, for the Scientific Reserve all of
the samples match the w40e47 ka age range, produced by the
different laboratories involved in the dating. We therefore consider
that these bones have accumulated at c. 40 ka at the latest, as a
consequence of one or more flooding episodes.
On the other hand, the ages of the several fossil remains dated
from the Passage of the Pits and Access Passage are much younger.
Apart from a cave bear metacarpal dated at c. 41 ka, the other dated
remains (including one of an articulated cave bear skeleton) have
ages ranging between w31 and 36 ka. The ages seem to indicate
that the cave bears were using the Upper Passages longer than the
Lower ones.
5.4. Fossil accumulation and hydrological regimes
The excavation of Urşilor Cave yielded a large number of artic-
ulated skeletons of both cave bears and cave lions (at least 6 in-
dividuals). These finds provide evidence that the deposition of
sediments was caused by rapid flooding events and exclude the
hypothesis of accumulation formed solely by bone reworking. On
the other hand, the presence of interbedded flowstone within the
sediments and even of small stalagmites precipitated on top of cave
bear bones indicate that the sediment accumulation in the Exca-
vation Chamber formed during several successive stages.
The sediment terraces and clay-covered walls in the Scientific
Reserve of Urşilor Cave are unique repositories of both articulated
skeletons of cave bears and lions and their vital traces (scratches,
footprints, fur imprints, etc.) (Robu et al., 2011) that attest for ani-
mal activity following the sediment deposition. In the Scientific
Reserve terraces and the Excavation Chamber the ages of the arti-
culated skeletons range between 41 and 45 ka, thus establishing a
terminus ante quem for the deposition of sediments. In the Upper
Passages, the partially articulated skeletons attest for cave bear
presence as recently as w31 ka, but it seems that, for some reason,
the bear population may have been prevented from entering the
Lower Passages after c. 40 ka.
At the same time, the large number of cave bear vital traces
preserved in the soft clay sediments from the Scientific Reserve
indicates that no significant flooding occurred after c. 40 ka, as such
an event would have been destroyed them. As well, the walls of the
vertical shafts from the Passage of the Pits, which descend to almost
the same level as the river bed from the Scientific Reserve, are
covered with the same thin clay layer as in the Scientific Reserve,
and both present numerous bioglyphs, indicating post-flooding
cave bear activity.
Finally, the presence of coarser sediments in the Upper Passage,
which includes cave bear bones as old as 35e38 ka, is somehow
surprising. They indicate a hydrologically dynamic environment
and/or a reworking of older sediments due to a distinct water
source, (e.g. from a temporary stream originating in the upper part
of the Shaft). However, the precise location of the water inflow
remains uncertain.
5.5. The general evolution of Urşilor Cave
When putting together the chronological information obtained
so far on speleothems, sediments, and fossil remains from Urşilor
Cave, a broad scenario emerges for the cave evolution during the
last w300 ka. This scenario is inevitably incomplete, given the
uncertainties associated with the different dating results. Therefore
only some key-episodes or key-moments in the evolution of the
karst system can be securely dated. We present these episodes and
their associated chronology in Fig. 8 and describe them below.
12 S. Constantin et al. / Quaternary International xxx (2013) 1e16

Fig. 8. Cartoon showing a possible scenario of the evolution of the Lower Level in Urşilor Cave between w290 ka and w40 ka. For simplicity, the last two evolutionary stages of the
cave, which are only documented by finds from the Upper Levels, are not shown in this figure (see text for details).

5.5.1. Stage 1: prior to w290 ka
The age of the flowstone covering the wall of the Shaft
(w291.1  9.15 ka) shows that more than w290 ka, the 17 m step in
the Excavation Chamber was already formed and the subterranean
stream was sinking at a lower level, at the base of the Shaft. The
overall morphology of the cave suggests that the small shafts in the
Passage of the Pits may have acted as overflow channels for the
water temporarily accumulated in the Excavation Chamber. The age
of the flowstone corresponds to MIS 8. Although it may be sur-
prising that flowstone deposition was active during this cold stage,
MIS8 was comparable to the presumably warmer MIS3 and displays
a very similar Dansgaard-Oeschger (D-O) variability (Sidall et al.,
2007). On the other hand, at the low altitude of Urşilor Cave, the
primary control for speleothem deposition would not normally be
the annual mean temperature but rather percolation water
availability.
5.5.2. Stage 2: between w210 and w145 ka
The age of w146.3  6.8 ka measured for the PU10-4 speleo-
them in the Excavation Chamber marks another chronological
constraint in the evolution of Urşilor Cave. The part of the stalag-
mite that grew atop the sediment terrace of PU4 clearly postdates
it. The OSL estimates for the sediment sample taken from w100 cm
below PU10-4, as well as the two other samples from the lower part
of the PU2 profile suggest a maximum age of c. 210 ka. We therefore
infer that at some time between w210 ka and w145 ka, water
flooded the Lower Passages having had a relatively high hydraulic
energy to deposit fine levels of sands. These episodes alternated
with periods when a backswamp lake formed and deposited the
finer silt or clay layers. The stratigraphy of the deposits, and espe-
cially the fact that the PU10-4 stalagmite is partially engulfed in
sediments (Fig. S6), suggest the existence of several flooding
events, separated by at least one interval of low waters allowing the
growth of the basal w70 cm of the PU10-4 stalagmite. All other
sedimentological parameters point to alternating episodes of low-
energy hydraulic regimes depositing fine sediments within a
backswamp facies, and short transitions to high-energy waters
which were able to deposit coarser sediments (channel facies) and
to partially erode older terraces.
No bone remains have been found in the sediments of this stage,
and there is no indication that the cave might have been inhabited
by any animals at that time. On the other hand, it is interesting to
note that the age measured for the PU10-4 stalagmite also fells
within a cold interval, MIS 6. Although calcite precipitation rates in

Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
j.quaint.2013.10.012
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of

Table 4
Radiocarbon measurements of the samples from Urşilor Cave.

Field number Lab. noa Specimen Location Depth Coll. yield (%) %Cb whole %Nb whole %Cncoll 14
C age Cal BP age Age range cal BP Comments
[cm] bone bone (median) (2s)

The Scientific Reserve (Lower Level)

PU/SR-012 OxA-22122 P. spelaea, upper I1 Terrace Surface 7.5 39000  1000 43363 44922e41971 Source: Stuart and
Scientific Lister, 2011
Reserve
PU/TA-001 OxA-22123 P. spelaea, upper I1 Terrace Surface 1.6 38600  1000 43078 44672e41721
Scientific
Reserve

S. Constantin et al. / Quaternary International xxx (2013) 1e16

PU/SR-025 VERA-5565 U. spelaeus canine Terrace Surface >1% 39000  1000 41209 44922e41971 Articulated skeleton
Scientific
RESERVE
PU/SR-024 LTL5510A P. spelaea, lower I3 Terrace Surface 36027  350 43363 41832e40486 Articulated skeleton
Scientific
Reserve
PU/A1-0-001 LTL5511A U. spelaeus lower M1 Excavation 10 35490  400 40697 41514e39524
PU/D4-0-076 LTL5513A U. spelaeus upper I3 Excavation 10 36625  500 41634 42431e40872
PU/D3-0-055 LTL5515A U. spelaeus upper I3 Excavation 12 39483  500 43629 44445e42823
PU/A1-0-002 LTL5512A U. spelaeus lower I2 Excavation 13 36529  370 41573 42156e41001
PU/D4-II-004 LTL5517A U. spelaeus upper I3 Excavation 14 40916  620 44688 45673e43719 Articulated skeleton
PU/D4-0-025 LTL5516A U. spelaeus lower M1 Excavation 15 36752  400 41718 42322e41114
PU/D3-0-057 LTL5514A U. spelaeus upper I3 Excavation 17 40252  600 44187 45145e43189
PU/A1-VIII-001 LTL5518A U. spelaeus lower M3 Excavation 70 36807  400 41754 42359e41148
PU/C1-VIII-062 Poz-48880 U. spelaeus mandible Excavation 80 1.1 5.3 2.1 0.37 37200  800 42051 43353e40892
PU/D3-IX-027 Poz-46548 U. spelaeus metacarpal Excavation 100 0.84 3.6 1.2 0.36 37700  600 42362 43270e41520 Articulated skeleton
PU/A1-XII-004 LTL5519A U. spelaeus metacarpal Excavation 110 41733  900 45276 47045e43936 Articulated skeleton
PU/C1-XV-039 Poz-53617 Panthera spelaea femora Excavation 157 0.93 2.8 1.5 L1.25 32850  350 37497 38544e36674 Articulated skeleton;
discarded age
PU/A1-XII-4g Poz-53618 U. spelaeus metacarpal Excavation 110 2.04 6.5 3.1 L1.87 34500  400 39563 40630e38670 Same as LTL5519A;
discarded age
The Access Passage
PU/GO-001 Poz-48737 U. spelaeus metacarpal Access Passage 12 3.15 8.2 3.2 0.44 32200  350 36751 37928e35565
The Passage of the Pits
PU/GP-003 Poz-48739 U. spelaeus metacarpal Passage of the Surface 3.57 7.5 3.1 0.87 36450  600 41504 42449e40455
Pits
PU/GP-002 Poz-48741 U. spelaeus metacarpal Passage of the Surface 0.8 1.5 0.5 0.15 24600  180 29441 30159e28845
Pits
PU/GP-001 Poz-48742 U. spelaeus femora Passage of the Surface 2.17 3.5 1.3 0.01 26750  230 31203 31467e30957 Articulated skeleton
Pits

Notes: All depths are in cm below surface. Collagen yields are given in percent of extracted collagen with respect to the initial weight of the sample. %Cncoll is the %C from “non collagen” (%Cncoll ¼ %Cb  (2.7  %Nb).
a
Lab acronyms: OxA e Oxford Radiocarbon Accelerator Unit; VERA e Vienna Environmental Research Accelerator; LTL e CEDAD Laboratory, Salento; POZ e Poznan Radiocarbon Laboratory.

13
14 S. Constantin et al. / Quaternary International xxx (2013) 1e16
Romanian karst are known to be generally lower during glacials
than during interglacials, speleothem formation is documented
during MIS 6 in many cave sites located at middle latitudes (e.g. Bar-
Matthews et al., 2003; Constantin et al., 2006; Wainer et al., 2011).
The stereotype according to which speleothem formation is more
likely to correspond to warm climate phases while sediment ac-
cumulations are more likely related to colder ones cannot be
ascertained. Hence, we do not have any evidence to link these first
episodes of sediment deposition to either a warm or a cold stage
and those sediments may have been transported during either
MIS7 or MIS6, or both.
5.5.3. Stage 3: w92.5 ka
At c. 92.5 ka, the PU10-3 flowstone from the downstream end of
the Excavation Chamber was growing on top of previously depos-
ited sediments. It is now suspended at w 0.5 m above younger
sediments and fossil bones, thus adding evidence for the scenario of
periodical high-energy hydraulic events able to wash out the pre-
viously deposited sediments. On the other hand, the precipitation
of this flowstone indicates an extended time period of low waters,
as flowstone is known for its low growth-rates and there are no
visible impurities in the cross-section to indicate some sort of pe-
riodical flooding. The obtained age corresponds to the beginning of
the cold MIS 5b. When correlated with the PU10-4 and PU10-2
ages, it suggests that colder periods may have been favorable for
uninterrupted speleothem deposition and that the sediment in-
flows might have been linked to warmer stages.
5.5.4. Stage 4: between w92.5 and w40 ka
Sediments from the top part of the PU2 profile, the PU1 profile,
as well as those from the excavated trench, have shown broadly
concordant OSL ages, which are interpreted as being younger than
w90 ka. In addition, they all show similar sedimentological fea-
tures: they almost exclusively include silts and laminated clays and
point towards a backswamp-type depositional facies.
The main ambiguity concerning the real age of these sediments
stems from the fact that the two samples taken from the excavation
trench correspond to stratigraphical layers that include fossil bones
which are as young as w42 ka. Hence, we can only assume that
either the OSL signal was not completely reset or that the sedi-
ments have been reworked from deeper parts of the cave. Either
way, the depositional event(s) took place at some time between
w47 and 45 ka the ages of the articulated skeletons lying on top of
the alluvial terraces and w42 ka (the youngest reliable radiocarbon
age of an articulated skeleton found in the excavation trench).
When considering the stratigraphy of the deposit in the
excavation trench, at least two flooding events are apparent.
Their overall sedimentologic and taphonomic features suggest
short but intense flooding, with the instant burial of numerous
cave bears (and at least one cave lion), as shown by the presence
of their articulated skeletons. Given the large uncertainties of the
radiocarbon dates, the timing of these events is hard to separate.
The presence of L8 flowstone crusts above the L5 suggests that
the floods were separated in time by at least several hundred
years.
However, considering that all excavated speleothems are posi-
tioned towards the side of the passage (i.e. the lateral “terrace”), we
cannot exclude that the first flooding event was followed by a rapid
decrease of water level during which the stream has flown only
along the central channel, thus allowing for subaerial deposition of
speleothems. A new flood would have add a new layer of fine
sediments and bones (both carcasses and reworked), covering the
entire floor of the chamber, including the small lateral terrace. If
this scenario repeated, it seems plausible that during the
“erosional” phases bones which were at that time positioned along
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
the central channel were scrambled and transported downstream.
Any new flooding would bring new, either younger or older
(reworked), fossil remains to fill these gaps. This may explain why
some discrepancies may exist between radiocarbon ages of remains
from what is now perceived as a “single” stratigraphic level.
We cannot speculate about when the bear occupation started in
the Scientific Reserve of Urşilor Cave. What is clear is that by
w45 ka this cave sector was hosting a large number of cave bears,
and possibly also cave lions. It was also about this time when the
first flooding event raised the PU2 river terrace by about 1 m and
created the PU1 terrace deposit. Subsequently, the cave bears
continue to live in the cave, and all terraces and sediment faces
have preserved their vital traces. The same intensive cave bear
usage of the cave is documented as well in the Upper Level. The
next flooding events led to the accumulation of new fossil remains,
which may originate from either the Lower Level (by fluvial
transport of bones or drowning) or the Upper Level (by fluvial
transport or individual animals falling down the Shaft).
At the present state of research, the interpretation of the cave
system evolution during this stage faces several uncertainties
associated with both the OSL and radiocarbon ages and also to our
understanding of the accumulation processes. However, virtually
all dated fossil remains show ages between w45 and w42 ka. While
it is very tempting to correlate this taphonomic event with a cli-
matic oscillation during MIS3, evidence is not solid enough to
ascertain one specific event. For instance, the H5 event (as many
other MIS3 climatic oscillations) was very short (between 47.74 and
46.68  0.8 ka according to Genty et al., 2010) and the errors
associated with the radiocarbon ages at this time are simply too
high to establish such a correlation. In the case of Peştera cu Oase
(SW Romania), where a similar thanatocenosis of a roughly similar
age was described, the favored interpretation was that at the onset
of GIS 12 the snow cover that accumulated during the w1000
years-long H5 event rapidly melted and generated a massive
flooding event (Constantin et al., 2013b).
5.5.5. Stage 4: between w40 ka and w30 ka
There is no reliable radiocarbon date or other evidence to
prove cave bear presence in the Lower Level of Urşilor Cave after c.
40 ka. There is also no hard evidence attesting to any notable
hydraulic events, and the fact that several w45 ka old skeletons
remained preserved on top of the terraces of the Scientific Reserve
actually rules out any major floods. Radiocarbon dates indicate the
presence of cave bears in the Upper Passages as late as w30 ka.
This raises the question of where might have been the original
entrance(s) used by cave bears to get into the cave. As the natural
entrances are now sealed by sediments and speleothems, we can
only assume that the entrance that allowed cave bears in the
Upper Passages continued to exist. On the contrary, a second
entrance which may have allowed animal access into the Lower
Level was already sealed.
The mechanism of deposition of the last sediments levels in the
Upper Passages is still uncertain. In the test trench of the Access
Passage the sediment consists of pebbles, coarse sand and only a
low amount of clay, accounting for a high hydraulic energy, while
also including cave bear bone fragments reliably dated to w35e
37 ka. The only reasonable explanation for this would be that a
second stream flowed through the Upper Passages at that time,
although its origins are unknown. This contradicts the common
views (e.g. Valenaş, 1979) according to which Urşilor Cave was
created by a single underground stream.
5.5.6. Stage 5: after w30 ka and through the Holocene
The latest dated cave bear remains from the Passage of the Pits
have ages close to c. 30 ka. In the absence of any evidence, we cannot
 S. Constantin et al. / Quaternary International xxx (2013) 1e16
speculate about the time when cave bears and other large mammals
stopped using the cave. The only chronological controls are: (1) the
abundant presence of the candlestick stalagmites in the Upper
Level, including some grown within hibernation nests or covering
fossil remains and which are known to be Holocene (Onac et al.,
2002; Constantin, 2003) and (2) the fact that the natural en-
trances were clogged by sediments and/or sealed by flowstone at
the time of discovery. Holocene speleothem formation is known to
be very fast and abundant in caves of Romania (Constantin, 2003;
Constantin et al., 2006, 2013a). We speculate that the closure of
the Urşilor Cave was caused by either small collapses at the end of
the Last Glacial Maximum (LGM) or clogging by speleothem depo-
sition or a combination of both. We have no evidence to document
the cave evolution during the MIS2 and the last deglaciation.
5.6. Summary: reconstruction of the sedimentary history of the
cave and the taphonomy of fossil deposits
The sedimentological research and dating of the various de-
posits in Urşilor Cave led to a better understanding of the evolution
of the cave during the last w300 ka and substantially changed some
of the hypotheses concerning the cave’s history. The cave bear fossil
assemblage in the Lower Level was formerly associated with a
massive collapse in the Shaft sector that prevented bears from
exiting the cave. This is now shown to be erroneous and, even
though a collapse cannot be totally ruled out, it did not influence
the fate of the large mammals from this sector since it happened
much earlier.
Uranium-series dating of several flowstone levels associated
with the OSL dating on sediments and the radiocarbon dating of
fossil bones indicate at least two broad periods of sediments input
into the Lower Level of the cave. These probably included alter-
nating depositional and erosional phases which complicated the
interpretation of the fossil deposit and scattered remains. Our data
indicate that only the last floods (sometime between w47 and
w40 ka) were responsible for the accumulation of fossil remains
and for the death of the cave bears and lions.
On the other hand, the fact that the Shaft was already formed at
that time indicates that the animals used another entrance, since it
is unlikely that so many individuals would enter so deep into the
cave while coping with a 17 m deep step. Such an entrance could
have been located in the middle part of the cave, close to the
present-day Excavation Chamber. This entrance must have been
opened even after the last flooding, as cave bears were still leaving
their vital traces throughout.
The fact that in the Upper Passages cave bears continued to live
for at least 10,000 years longer suggests that these passages were
actually a “separate” cave, with the Shaft acting as a barrier for the
bear population. The presence of different sediments in these
passages, indicative of a higher hydraulic gradient, can only be
explained by admitting that a second underground stream was still
flowing through the upper parts of the system concurrently with
the lower level stream. Why and when this stream has abandoned
the upper passages is unknown, but a possible explanation may be
the early reactivation of the European rivers during the last
deglaciation (Ménot et al., 2006).
When trying to reconcile all the data yielded by our research,
the former theories that explained the evolution of the cave as a
unitary karst system affected by underground stream piracy and
collapses seem to be too simplistic. The final picture that emerged,
although hindered by some chronological uncertainties, shows a
more complicated history, in which one karst system may develop
in such a way that different parts of it have separate sedimentary
histories for some time. Such an evolution inevitably affects the
distribution of cave animal populations and our conclusions on a
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
15
given thanatocenosis. This probably happens more often than
considered in many reported cave sites, as shown in the case of the
Peştera cu Oase (Constantin et al., 2013a) when the initial,
simplistic approach was soon challenged by field data and the final
history of the fossil accumulation proved to be more complicated
than initially thought.
6. Conclusions
This paper described the sedimentological, geomorphological,
and geochronological work undertaken at the Urşilor Cave with the
aim of understanding the taphonomy of the fossil accumulation in
the Excavation Chamber. The initial hypotheses, which assumed
that the fossil remains accumulation was caused by a combination
of collapses, river deepening, and underground flooding, were
largely erroneous. The “separation” of the cave system through the
Shaft step was much older than previously thought and the accu-
mulation of sediments in the lower level of the cave consisted of a
long history of depositional and erosional events. It was only the
last suite of such events (broadly dated between 47 and 40 ka) that
led to the accumulation of fossil remains. The upper levels of the
cave continued to be occupied by cave bears and seem to have
evolved separately with respect to the lower ones, even having
their own drainage system.
Considering the ages of the dated speleothems and those of the
other cave deposits, it appears that: (1) speleothem precipitation
was not prevented during climatic cold periods; (2) sediment
depositional events were more likely linked to warmer and wetter
periods. We tentatively assign the fossil accumulation from the
Excavation Chamber to a series of floods that most probably took
place at the end of the H5 event and the onset of the GIS 12. The
speleogenesis of the Urşilor Cave, which appears to be more
complicated than initially thought, may bring to attention the need
for careful research of the sedimentary history of any paleonto-
logical site located in deep-cave settings.
Acknowledgements
This research was funded through the KARSTHIVES Project e
“Climate archives in karst”, funded by CNCS e UEFISCDI Grant
PCCE_ID 31/2010. We are grateful to Tomasz Goslar and Hervé
Bocherens for insightful discussions on the radiocarbon dating is-
sues, to Martina Pacher for her help with the VERA dating, and to
Oana Moldovan for her constant help during this research. We
thank Valerica Toma and Augustin Nae for their help in the field and
the Apuseni National Park for granting the excavation permit and
field assistance.
SC designed research and secured funding. MR, AP, MV, IM, MK,
SC, VD, and CP performed research. CMM, RDR, and CP have done
the sedimentological study, VD and DH have done the U-series
dating, VA and ATG carried out the OSL dating, and SC and MR wrote
the paper. All coauthors contributed to improve the manuscript.
Appendix A. Supplementary data
Supplementary data related to this article can be found at http://
dx.doi.org/10.1016/j.quaint.2013.10.012.
References
Adamiec, G., Aitken, M., 1998. Dose rate conversion factors: update. Ancient TL 16,
37e50.
Bar-Matthews, M., Ayalon, A., Gilmour, M., Matthews, A., Hawkesworth, C.J., 2003.
Seaeland oxygen isotopic relationships from planktonic foraminifera and spe-
leothems in the Eastern Mediterranean region and their implication for
16 S. Constantin et al. / Quaternary International xxx (2013) 1e16
paleorainfall during interglacial intervals. Geochimica Cosmochimica Acta 67
(17), 3181e3199.
Berger, G.W., Pérez-González, A., Carbonell, E., Arsuaga, J.L., Bermúdez de
Castro, J.M., Ku, T.L., 2008. Luminescence chronology of cave sediments at the
Atapuerca paleoanthropological site, Spain. Journal of Human Evolution 55 (2),
300e311.
Blott, S.J., 2010. GRADISTAT Version 8.0-A Grain Size Distribution and Statistics
Package for the Analysis of Unconsolidated Sediments by Sieving or Laser
Granulometer. http://www.kpal.co.uk/gradistat.html.
Bocherens, H., Drucker, D., Billiou, D., Moussa, I., 2005. Une nouvelle approche pour
évaluer létat de conservation de l’oset du collagène pour les mesures iso-
topiques (dation au radiocarbone, isotopes stables du carbone et de l’azote).
L’anthropologie 109, 557e567.
Bocherens, H., Drucker, D., Billiou, D., Geneste, J.-M., Kervzo, B., 2008. Grotte
Chauvet (Ardèche, France): a natural “experiment” for bone diagenesis in
karstic context. Palaeogeography, Palaeoclimatology, Palaeoecology 266,
220e226.
Bordea, S., Bleahu, M., Bordea, J., 1975. Date noi stratigrafice structurale asupra
Bihorului de vest. Unitatea de Urm at şi Unitatea de Vetre. D ari de seama ale
şedinţelor. Inst. Geol. Geof., Bucharest, pp. 61e83 (in Romanian).
Bosch, R.F., White, W.B., 2004. Lithofacies and transport of clastic sediments in
karstic aquifers. In: Sasowsky, I.D., Mylroie, J.E. (Eds.), Studies of Cave Sedi-
ments: Physical and Chemical Records of Paleoclimate. Kluwer Academic/
Plenum Publishers, New York, pp. 1e22.
Bronk Ramsey, C., 2009. Bayesian analysis of radiocarbon dates. Radiocarbon 51 (1),
337e360.
Bronk Ramsey, C., Higham, T., Bowles, A., Hedges, R., 2004. Improvements to the
pretreatment of bone at Oxford. Radiocarbon 46 (1), 155e163.
Calcagnile, L., Quarta, G., D’Elia, M., Rizzo, A., Gottdang, A., Klein, M., Mous, D.J.W.,
2004. A new accelerator mass spectrometry facility in Lecce, Italy. Nuclear In-
struments and Methods in Physics Research 223e224C, 16e20.
Calcagnile, L., Quarta, G., D’Elia, M., 2005. High resolution accelerator-based mass
spectrometry: precision, accuracy and background. Applied Radiation and Iso-
topes 62 (4), 623e629.
Chapot, M.S., Roberts, H.M., Duller, G.A.T., Lai, Z.P., 2012. A comparison of natural- and
laboratory-generated dose response curves for quartz optically stimulated lumi-
nescence signals from Chinese Loess. Radiation Measurements 47, 1045e1052.
Constantin, S., 2003. Evolutii paleoclimatice în Cuaternar pe baza speleotemelor din
carstul Munţilor Banatului şi Mehedinţi. Univ of Bucharest, Bucharest, p. 324
(PhD. thesis)(unpublished, in Romanian).
Constantin, S., Lauritzen, S.-E., Lundberg, J., 2006. New data on the chronology of
the Termination II and paleoclimate during MIS 5, based on a study of a sta-
lagmite from Closani Cave (Romania). In: Onac, B.P., T amas, T., Constantin, S.,
Persoiu, A. (Eds.), Archives of Climate Change in Karst. Karst Waters Institute
USA, pp. 98e100.
Constantin, S., Munteanu, C.M., Milota, Ş., Sarcina, L., Gherase, M., Zilhão, J., 2013a.
The Ponor-Plopa Cave system: description, sediments, and genesis. In:
Trinkaus, E., Constantin, S., Zilhão, J. (Eds.), Life and Death at the Peştera cu Oase
e a Setting for Modern Human Emergence in Europe. Oxford University Press
USA, New York, pp. 41e55.
Constantin, S., Zilhão, J., Trinkaus, E., 2013b. The paleoenvironmental context of the
Peştera cu Oase. In: Trinkaus, E., Constantin, S., Zilhão, J. (Eds.), Life and Death at
the Peştera cu Oase e a Setting for Modern Human Emergence in Europe.
Oxford University Press USA, New York, pp. 385e388.
Couchoud, I., 2006. Etude pétrographique et isotopique de spéléothèmes du sud-
ouest de la France formés en contexte archéologique, vol. 1. Université
Bordeaux, p. 346 (PhD thesis).
Fairchild, I.J., Baker, A., 2012. Speleothem Science. FRom Process to Past Environ-
ments. Willey-Blackwell, Chichester, p. 432.
Folk, R.L., Ward, W.C., 1957. Brazos River bar: a study in the significance of grain size
parameters. Journal of Sedimentary Petrology 27, 3e26.
Ford, D.C., Williams, P., 2007. Karst Hydrogeology and Geomorphology. Wiley, p. 562.
Genty, D., Comborieu-Nebout, N., Peyron, O., Blamart, D., Wainer, K., Mansuri, F.,
Ghaleb, B., Isabello, L., Dormoy, I., von Grafenstein, U., Bonelli, S., Landais, A.,
Brauer, A., 2010. Isotopic characterization of rapid climatic events during OIS3
and OIS4 in Villars Cave stalagmites (SW-France) and correlation with Atlantic
and Mediterranean pollen records. Quaternary Science Reviews 29, 2799e2820.
Goslar, T., Czernik, J., Goslar, E., 2004. Low-energy 14C AMS in Poznan  radiocarbon
laboratory, Poland. Nuclear Instruments and Methods B 223-224, 5e11.
Grün, R., 2006. Direct dating of human fossils. Yearbook of Physical Anthropology
49, 2e48.
Hadnagy, A., 1977. Studiul micromineralogic şi rolul mineralelor grele in corelarea
sedimentelor de peşter a. Nymphaea 107e128.
Häuselmann, A.D., Häuselmann, P., Onac, B.P., 2010. Speleogenesis and deposition of
sediments in Cioclovina Uscata  Cave, Sureanu Mountains, Romania. Environ-
mental Earth Sciences 61, 1561e1571.
Higham, T., Wild, E.M., 2013. Radiocarbon dating of the Peştera cu Oase faunal re-
mains. In: Trinkaus, E., Constantin, S., Zilhão, J. (Eds.), Life and Death at the
Peştera cu Oase e a Setting for Modern Human Emergence in Europe. Oxford
University Press USA, New York, pp. 77e85.
Hoffmann, D.L., 2008. 230Th isotope measurements of femtogram quantities for U-
series dating using multi ion counting MIC MC-ICPMS. International Journal of
Mass Spectrometry 275, 75e79.
Please cite this article in press as: Constantin, S., et al., Reconstructing the evolution of cave systems as a key to understanding the taphonomy of
fossil accumulations: The case of Urşilor Cave (Western Carpathians, Romania), Quaternary International (2013), http://dx.doi.org/10.1016/
j.quaint.2013.10.012
Hoffmann, D.L., Prytulak, J., Richards, D.A., Elliott, T., Coath, C.D., Smart, P.L.,
Scholz, D., 2007. Procedures for accurate U and Th isotope measurements by
high precision MC-ICPMS. International Journal of Mass Spectrometry 264,
97e109.
Jurcsák, T., Plopiş, R., Ignat, D., Şerban, M., Popa, E., 1981. Date privind fauna fosil
aa
Peşterii Urşilor (Munţii Bihor). Nymphaea VIII-IX, 161e257.
Longin, R., 1971. New method of collagen extraction for radiocarbon dating. Nature
230, 241e242.
Ménot, G., Bard, E., Rostek, F., Weijers, J.W.H., Hopmans, E.C., Schouten, S.,
Damsté, J.S.S., 2006. Early reactivation of European rivers during the last
deglaciation. Science 313, 1623e1625.
Moldovan, O.T., Mihevc, A., Miko, L., Constantin, S., Meleg, I.N., Petculescu, A.,
Bosak, P., 2011. Invertebrate fossils from cave sediments: a new proxy for pre-
Quaternary paleoenvironments. Biogeosciences 8, 1825e1837.
Murray, A.S., Wintle, A.G., 2000. Luminescence dating of quartz using an improved
single-aliquot regenerative-dose protocol. Radiation Measurements 32, 57e73.
Murray, A.S., Wintle, A.G., 2003. The single aliquot regenerative dose protocol:
potentials for improvement in reliability. Radiation Measurements 37, 377e381.
Onac, B.P., Constantin, S., Lundberg, J., Lauritzen, S.E., 2002. Isotopic climate record
in a Holocene stalagmite from Urşilor Cave (Romania). Journal of Quaternary
Science 17, 319e327.
Petculescu, A., Murariu, D., 2009. The First Ecological Reconstruction of Under-
ground Environment from Romania, Cioclovina Uscata  Cave. Editura Uni-
versitara , Bucharest, p. 138.
Pietrowska, N., Goslar, T., 2002. Preparation of bone samples in the Gliwice radio-
carbon laboratory for AMS radiocarbon dating. Isotopes in Environmental and
Health Studies 38 (4), 267e275.
Reimer, P.J., Baillie, M.G.L., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Bronk
Ramsey, C., Buck, C.E., Burr, G.S., Edwards, R.L., Friedrich, M., Grootes, P.M.,
Guilderson, T.P., Hajdas, I., Heaton, T.J., Hogg, A.G., Hughen, K.A., Kaiser, K.F.,
Kromer, B., McCormac, F.G., Manning, S.W., Reimer, R.W., Richards, D.A.,
Southon, J.R., Talamo, S., Turney, C.S.M., van der Plicht, J., Weyhenmeyer, C.E.,
2009. IntCal09 and Marine09 radiocarbon age calibration curves, 0-50,000
years cal BP. Radiocarbon 51 (4), 1111e1150.
Robu, M., Petculescu, A., Panaiotu, C., Doeppes, D., Vlaicu, M., Dragusin, V.,
Kenesz, M., Moldovan, O.T., Constantin, S., 2011. New insights on the cave bear
population from the Ursilor Cave, Romania. Quaternaire 4 (Suppl.), 107e116.
Rusu, T., 1981. La grotte des ours de Chiscau (Monts Apuseni). Revue Roumaine de
Géologie, Géophysique et Géographie. Géographie 25 (2), 193e204.
Rusu, T., Racoviţ a, G., 1981. Peştera Urşilor de la Chişca u. Ocrotirea Naturii şi a
Mediului Înconjura tor 25 (1), 57e71.
Sasowsky, I.D., Mylroie, J.E. (Eds.), 2004. Studies of Cave Sediments. Physical and
Chemical Records of Paleoclimate. Kluwer Academic/Plenum Publishers, New
York, p. 330.
Sidall, M., Stocker, T.F., Blunier, T., Spahni, R., Schwander, J., Barnola, J.-M.,
Chapellaz, J., 2007. Marine Isotope Stage (MIS) 8 millennial variability strati-
graphically identical to MIS 3. Paleoceanography 22, PA1208. http://dx.doi.org/
10.1029/2006PA001345.
Stuart, A.J., Lister, A.M., 2011. Extinction chronology of the cave lion Panthera spe-
laea. Quaternary Science Reviews 30, 2329e2340.
Terzea, E., 1978. Départament de Bihor (note préliminaire). Travaux de l’Institut de
Spéologie “Emile Racovitza”. Dépôts de remplisage et mammifères quaternaires
de “Peştera Urşilor” de Chişca u, vol. XVII, pp. 139e144.
Terzea, E., 1989. Les Arvicolidés (Rodentia, Mammalia) du Pléistocène Moyen de
Chişcau-1, vol. XXVIII, pp. 57e72 (Depart. de Bihor, Roumanie). Travaux de
l’Institut de Spéologie “Emile Racovitza”.
Timar-Gabor, A., Vasiliniuc, Ş., Vandenberghe, D.A.G., Cosma, C., Wintle, A.G., 2012.
Investigations on the reliability of SAR-OSL equivalent doses obtained for quartz
samples displaying dose response curves with more than one component.
Radiation Measurements 47, 740e745.
Trinkaus, E., Constantin, S., Zilhão, J. (Eds.), 2013. Life and Death at the Peştera cu
Oase -ea Setting for Modern Human Emergence in Europe. Oxford University
Press USA, New York, Human Evolution Series, p. 528.
Valenaş, L., 1979. Studiu complex al zonei Valea Cr aiasa-Valea Vârtoape, cu referire
special a la Peştera Urşilor (Munţii Bihor). Nymphaea VII, 139e176.
Wainer, K., Genty, D., Blamart, D., Daëron, M., Bar-Matthews, M., Vonhof, H.,
Dublyansky, Y., Pons-Branchu, E., Thomas, L., van Calsteren, P., Quinif, Y.,
Caillon, N., 2011. Speleothem record of the last 180 ka in Villars Cave (SW
France): investigation of a large?18O shift between MIS6 and MIS5. Quaternary
Science Reviews 30, 130e146.
White, W.B., 2007. Cave sediments and paleoclimate. Journal of Caves and Karst
Studies 69 (1), 76e93.
Wintle, A.G., Murray, A.S., 2006. A review of quartz optically stimulated lumines-
cence characteristics and their relevance in single-aliquot regeneration dating
protocols. Radiation Measurements 41, 369e391.
Zilhao, J., Trinkaus, E., Constantin, S., Milota, S., Gherase, M., Sarcina, L., Danciu, A.,
Rougier, H., Quilès, J., Rodrigo, R., 2007. The Pestera cu Oase people: Europe’s
earliest modern humans. In: Mellars, P., Boyle, K., Bar-Yosef, O., Stringer, C.
(Eds.), Rethinking the Human Revolution. McDonald Institute for Archaeolog-
ical Research, Cambridge, pp. 249e262.
Zupan Hajna, N., Pruner, P., Mihevc, A., Schnabl, P., Bosák, P., 2008. Cave sediments
from the PostojnskaePlaninska cave system (Slovenia): evidence of multi-
phase evolution in epiphreatic zone. Acta Carsologica 37 (1), 63e86.