TMP B664

Quaternary International xxx (2013) 1e16
Contents lists available at ScienceDirect
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

of fossil remains preserved in caves, especially those of micro-
Cave deposits are largely recognized as holding the potential of mammals, have been long considered as a paleoclimate proxy.
valuable palaeoenvironmental and paleoclimatic archives (see, for Moldovan et al. (2011) have shown that even invertebrate fossil
example, Ford and Williams, 2007; White, 2007). Speleothems are associations from cave sediments may be used as another paleo-
currently considered as one of the most reliable cave archives for environmental proxy.
reconstructing paleoclimate evolution at regional scale (Fairchild The advance of direct and indirect dating techniques (e.g.
and Baker, 2012). Fluviatile or lacustrine sediments from caves Couchoud, 2006; Grün, 2006; Berger et al., 2008) has allowed for a
are also physical and chemical recorders of past climate and 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-
* Corresponding author. logical deposits is the correct interpretation of the evolution of the
E-mail address: [email protected] (S. Constantin). 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 (Fig. S1) that leads to a small chamber (the Excavation Chamber)
wide range of cave archives and/or the analysis of coeval proxies. and the Scientific Reserve.
The paleontological excavation of the sediment deposit from the The Scientific Reserve is the lower level of Urşilor Cave (w500 m in
Scientific Reserve in Urşilor (¼Bears) Cave (Robu et al., 2011) has length) and includes a generally large horizontal passage (10 15 m),
revealed the presence of an extensive bone bed including mostly with several small side-passages and fossil meanders (Fig. S2). A small
cave bear remains but also two cave lions and at least one hyaena stream (1e10 l/s) flows through the lower level and sinks into an
and one mustelid. The taphonomic analysis has revealed that the impenetrable sump in the Excavation Chamber. Along the lower
vertebrate remains may have been accumulated via multiple passage, the stream has deposited alluvial sediments consisting of
mechanisms such as pitfall entrapment, natural death (including fine sands, silts, and clays which often support stalagmite formations
drowning) or fluvial transport. As direct radiocarbon dating has on their top. Currently the stream meanders through w0.5e3 m-high
yielded results that are close to or exceed the limits of the method, alluvial terraces. In the Excavation Chamber, it sinks and flows
the dating of complementary cave deposits and the understanding through an accumulation of sediments that exceeds 200 m3 of a mix
of the speleogenetic scenario are crucial for taphonomical of sandy-clayish alluvia and fossil bone beds.
interpretations. Both cave levels host important cave bear bone assemblages
With only a few exceptions (Constantin, 2003; Petculescu and (both on the surface and within the sediment, scattered or fully
Murariu, 2009; Häuselmann et al., 2010; Trinkaus et al., 2013) articulated specimens) and cave bear ichnofabrics (nests, scratch
most studies on the evolution of karst systems from the Romanian marks, footprints and fur imprints). However, most articulated
Carpathians rely only on stratigraphical and geomorphological skeletons and ichnofabrics are found in the Lower Level whereas in
evidence. Urşilor Cave is no exception, with the earlier attempts to the Upper Level the fossil remains are more scattered and/or
reconstruct the evolution of the cave and the sedimentation covered by massive speleothem formations.
mechanisms and chronology (Hadnagy, 1977; Terzea, 1978, 1989; The genesis of the cave system has been largely interpreted by
Valenaş, 1979; Jurcsák et al., 1981) relying mostly on the analysis corroborating observations on the overall cave morphology and
of the mineralogy of sediments, cave morphology, and a partial correlations of the different cave levels with the local river terraces
paleontological record. (Valenaş, 1979; Rusu, 1981; Rusu and Racoviţa , 1981). In broad
This study focuses on the establishment of a chronological and terms, the evolution of the cave was thought to include the
evolutionary framework for the Urşilor Cave in the last w300,000 following stages:
years. It integrates the study of relevant cave deposits (cave sedi-
ments, speleothems, and fossil bones) which were directly or (1) an initial karst drainage towards the Cr aiasa Valley led to
indirectly dated by means of radiocarbon, U-series, and optically the formation of the Upper Level passages. This stage in-
stimulated luminescence (OSL). It also integrates new, detailed, cludes successive abandonments of older passages, in pace
geomorphological and topographical surveys of the cave that are with the deepening of the base level. The Twisted Passage
relevant to its speleogenesis. The aim of the study was to provide a was abandoned in favour of a new resurgence via the
solid chronological and paleoevolutive basis for the taphonomical Candlesticks Passage which was, in turn, abandoned for a
interpretation of the fossil accumulation from the Scientific Reserve new drainage via the Access Passage.
of Urşilor Cave. (2) the capture of the subterranean stream towards the end of the
“Emil Racoviţa” Passage. A new drainage was established to-
2. Background wards the Peştera de la Chişcau, a small (148 m) resurgence
cave located w40 m below the current entrance of Urşilor
Urşilor Cave is located on the western slope of the Apuseni Cave (Figs. 1 and 2). The greatest part of the passages between
Mountains (Western Carpathians, Romania), on the left side of the the sink and Peştera de la Chişc au are impenetrable and
Craiasa Valley, at 491 m a.s.l (Fig. 1). The cave was discovered in thought to be generally small in size. This may have caused
1975 by blasting in a local marble quarry. Owing to both its rich and periodic accumulations of water upstream the sink and the
complex Upper Pleistocene thanatocenosis, and the abundance of deposition of alluvial material throughout the Lower Level.
speleothems, the cave was immediately protected, studied and (3) during the Holocene, the massive speleothem deposition
surveyed over two years. Since 1980, the upper part of the cave was throughout the Upper Level has sealed the former cave en-
set up for tourism, becoming the most important show cave of trances and largely covered the old sediments, and bone
Romania. remains. The Holocene age of the typical “candlestick” sta-
The cave is carved in the Upper Jurassic (Tithonian) recrystallized lagmites from the Upper Level was also indicated by subse-
limestones of the Bihor Autochthonous unit, which is partially quent studies (Onac et al., 2002; Constantin S. and Onac B.P.,
overlain by the Arieşeni and Codru Nappes, respectively (V alenaş, unpublished data). No other age controls existed for the
1979) (Fig. 1). The cave comprises two distinct levels, separated by previous evolutionary stages.
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, 3. Studied profiles
The Twisted Passage, The Passage of the Pits, and the “Emil Racoviţa”
Passage, see Fig. 2) and a large chamber (The Intersection Chamber, 3.1. The Excavation Chamber
w10 15 25 m), totaling c. 1000 m of galleries. The two artificial
entrances correspond broadly with two former natural entrances The new paleontological excavation at Urşilor Cave is located at
that were sealed by sediments and flowstone precipitation. the northern (downstream) end of the Excavation Chamber (Fig. 3).
In the final part of the “Emil Racoviţa ” Passage, a smaller, The Excavation Chamber (w20 m 6e10 m) reaches a maximum
ascending passage leads to a breakdown cone formed by blocks of height of w20 m directly below the Shaft. At its southern end, the
greywacke and flowstone fragments embedded in a clay matrix. underground stream sinks laterally at the base of the left wall. The
The overall morphology of this cave sector suggests the presence of small passage is impenetrable for explorers but it is evident that the
a local fault with the debris originating from the surrounding non- stream flows at the limit between the limestone wall and the
karst deposits. This passage ends with a 17 m-drop (the Shaft) sediments accumulated in the chamber. The stream may be
j.quaint.2013.10.012
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 The floor of the chamber is covered by brown clay with both
chamber but becomes again impenetrable after w50 m. The walls scattered cave bear remains and partially articulated skeletons.
of the chamber are largely covered by massive flowstone forma- Toward the walls, some of these remains are encrusted in calcite.
tions; a thin film of clay covers the walls up to w4 m in height, The thickness of the sediment deposit can only be assessed in the
indicating a former stagnant water level. southern end, where it forms a 1.8 m-high step (Fig. 3).
j.quaint.2013.10.012
4
j.quaint.2013.10.012
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
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.
Fig. 3. Sketch of the Excavation Chamber with the position of the excavation trench The PU2 Profile (Fig. 2c; Fig. S10) is located w65 m from the
and the U/Th dated calcite samples.
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
The excavation is still in progress and the detailed paleonto- profile consists of a flat surface that preserves ichnofabrics (floor
logical and taphonomical results will be presented elsewhere. Here scratch marks, cave bear nests, partially preserved footprints), and
we will only summarize the stratigraphy of the excavation and the its elevation corresponds to that of the neighboring terrace surface
surrounding deposits that were sampled for dating purposes as well where the fully articulated cave bear skeletons can be found. As in
as for sedimentological, geochemical, and rock-magnetic studies. the case of PU4, sediment samples were collected at 5 cm intervals.
The excavation trench is “L” shaped, has a total area of 7 m2 and In addition, three sediment samples were taken for OSL measure-
a current depth of w1.6 m without reaching the bedrock (Fig. S3). ments: at 70 cm, 155 cm, and 270 cm from the terrace top,
Eight distinct sediment layers (labeled L1-8) were found, with L1, respectively.
L3, and L5 being very rich in cave bear remains while the remaining The PU1 Profile (Fig. 2c; Fig. S11) is located w135 m upstream
layers are sterile. The bone beds extend across the whole section from the Excavation, at the point bar of an underground meander,
while the lower sterile ones were thicker along the A1-D1 transect in the right-side wall of the passage. The thickness of the deposits
and become thinner towards the D4 square, where they lie over a is 1.55 m, slightly lower than that of the opposite cut-terrace. The
flowstone terrace. The transects reveal a channel facies that cor- deposits consist of brownish, laminated, poorly sorted muds. The
responds to the general direction of flow of the subterranean top of the sequence consists of the same flat surface with cave
stream (A1eD1) and a cave terrace facies towards the sides (D1e bear nests, bones and scratch marks. The profile’s position is
D4) consisting of interbedded cave sediments and flowstone above opposite to the terrace where the cave bears and lion articulated
L5 (Fig. 4). skeletons are found. The top levels can be considered coeval
The excavation yielded a large number of cave bear bone re- although (owing to local cave morphology) the sediment stack on
mains (both articulated and scattered), as well as one articulated the left bank is w0.5 m higher. Sediment samples were collected
cave lion skeleton (Fig. S4). Twelve cave bear bone fragments were along the profile at 5 cm intervals. One sediment sample for OSL
sampled for radiocarbon dating. In addition, sediments from two dating was collected at 75 cm below the top. In addition, four bone
layers (L4; 100 cm and L5; 150 cm) were sampled for OSL dating. samples for radiocarbon dating were collected from the articu-
In the northern side of the Excavation Chamber, a flowstone lated skeletons of cave bears and the lion resting on top of the
pavement suspended w0.5 m above the floor (Fig. S5) indicates opposite terrace.
deposition on top of a former alluvial terrace, subsequently
removed by erosion. A flowstone sample was collected from this 3.3. Additional samples (the Upper Passage)
speleothem for U-series dating. In addition, we collected two other
samples from the flowstone that covers both the wall and the top of Additional bone samples were collected for radiocarbon dating
the Shaft (Fig. 3). from two significant locations in the Upper Passage:
j.quaint.2013.10.012
1) The Passage of the Pits is an intricate network of horizontal they consists of limestone pebbles and clay indicating a highly
passages and shafts starting in the “Emil Racoviţ
a” Passage and dynamic depositional environment.
descending to w3 m above the current level of the underground
stream (Fig. 2b; Fig. S12). Based on the phreatic morphology of 4. Analytical methods
the passages, the sediment fill and the overall topography of the
14
cave, this passage is considered to have functioned as an over- 4.1. AMS C dating
flow of the waters that sank in the Excavation Chamber.
Numerous cave bear remains, including one partially articulated Most of the AMS 14C dating was done at the Poznan Radio-
skeleton, have been found in this passage and three samples carbon Laboratory (Poland) (8 samples) and Centro di Datazione
were taken for dating. e Diagnostica (CEDAD) (University of Salento, Italy) (11 samples).
2) An old test-pit excavated in the Access Passage during the initial One sample was dated at the Vienna Environmental Research
research in the late ‘970s (Fig. 2a; Fig. S13) was reopened and Accelerator (VERA) and two samples were dated at the Oxford
two cave bear bone samples were taken from the freshly cleaned Radiocarbon Accelerator Unit (ORAU) and reported by Stuart and
wall of the pit at depths of 0.12 and 1.1 m, respectively. The Lister (2011). The analytical methods for collagen extraction,
sediments here are much coarser that in the lower level and 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.
j.quaint.2013.10.012
described in detail on the websites of the respective laboratories. carried out using the incorporated 90Sre90Y radioactive source,
Basically all laboratories use the collagen extraction procedure calibrated against gamma dosed calibration quartz supplied by the
originally described by Longin (1971), although with some Risø National Laboratory, Denmark. A dose rate of 0.123 Gy/s for the
modifications. At the Poznan Laboratory, collagen extraction was fine grains mounted on aluminium disks was obtained at the time
performed using the method described by Pietrowska and Goslar of measurement.
(2002) and the extracted collagen is then purified on Elkay 8 mm The measurement protocol was the Single Aliquot Regeneration
filters, and ultrafiltered on Vivaspin 15 MWCO 30 kD filters (SAR) applied on quartz (Murray and Wintle, 2000, 2003; Wintle
(Bronk Ramsey et al., 2004). Content of 14C in carbon samples and Murray, 2006) (see Supplemental Information for a complete
was measured following the protocol described by Goslar et al. description of the protocol). The OSL dating was done at the
(2004). A preliminary step includes a check for collagen degra- Dating and Luminescence Dosimetry Laboratory, Babeș-Bolyai
dation by measuring %C and %N in whole bone using a Flash University, Cluj-Napoca, Romania (see also supplemental material
EA1112 Series analyser (Thermo Scientific). The samples are for details).
regarded suitable for collagen dating if nitrogen content in bone
is not lower than 1%. The CEDAD laboratory does not apply an 4.4. Analysis of the sediment profiles
ultrafiltration step (Calcagnile et al., 2004, 2005) and, at the time
of analysis, it did not measure either %C or %N. The Oxford and The sediments from the Excavation Chamber and the subterra-
VERA collagen extraction procedures are similar, although nean terraces were collected from vertical trenches created by
treatment procedures are slightly different (Higham and Wild, removing w5 cm of the outer sediment layers. Duplicate sediment
2013). None of these laboratories measures %C and %N in samples were taken for: (1) granulometry, geochemistry, and rocke
extracted collagen as a routine, and hence the atomic C:N ratio magnetic properties measurements, and (2) screening of inverte-
cannot be calculated. brate fossil fauna, respectively.
Grain size measurements on fine samples were performed by
4.2. U-Th dating 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
Speleothem calcite subsamples were hand drilled, weighed, (Merck). A quantity of w2.5 g sample was later extracted from the
placed in clean Teflon beakers and covered with MQ H2O. The box and treated again with w0.2 ml of 2% solution of Graham’s
powders were then dissolved by stepwise addition of 7M HNO3 and salt. Each sample was analyzed on a HORIBA Partica LA-950V2
subsequently spiked using a mixed 229The236U spike. The solutions laser scattering particle size distribution analyzer to reveal grain
were evaporated to dryness and re-dissolved in a mixture of H2O2 size fractions. The coarser samples were analyzed by vibrating
and concentrated HCl and HNO3 to remove organic matter. The dry sieving of w100 g of the bulk sample and weighing the
solution was boiled at 130 C, and evaporated to dryness again, sediment quantity retained on each sieve, on an OHAUS Scout
before the separation and purification procedures. Separation and digital balance, down to the 500 mm fraction, which was subjected
purification of U and Th fractions was performed using 500 ml ion to the same procedure as the fine samples. Calculations and
exchange columns and BioRad AG 1X8 resin. U and Th fractions plots were done using the GRADISTAT 8 software (Blott, 2010). We
were dissolved in 1.8 ml 2% HCl and analysed on a Neptune ICP-MS applied the Folk and Ward (1957) method, and logarithmic sta-
equipped with a Cetac Aridus nebulizer at CENIEH (Burgos, Spain) tistics. The magnetic properties, elemental geochemistry and
following the procedures outlined in Hoffmann et al. (2007) and invertebrate fauna finds are still under investigation: they are not
Hoffmann (2008). significant for the scope of this paper and will be presented
elsewhere.
4.3. OSL dating
5. Results and discussion
Sediment samples for OSL dating were collected by hammering
25 cm-long, opaque, plastic tubes (10 cm) into the sediment sec- 5.1. U-series dating results
tions under no light conditions. Sample preparation for lumines-
cence measurements was performed under low intensity red light The results of the four flowstone samples dated from the
conditions. Only the central portion of the tubes was removed for Excavation Chamber are given in Table 1. The samples have a low U-
dating. A three-day treatment with 10% HCl solution was used for content, ranging between 0.06 and 0.15 ppm and three of them
carbonate removal, followed by another three-day H2O2 (30%) show low 230Th/232Th activity ratios ranging between10 and 35,
treatment for organic matter removal. The coarse grain fractions thus requiring age corrections for detrital Th.
(>63e90 mm) were separated through wet sieving. For the The PU10-1 sample was a small, corroded, stalagmite that was
extraction of the fine-grained quartz fraction (4e11 mm), the par- growing on top of the flowstone deposit directly above the shaft.
ticles smaller than 11 mm were isolated from the fraction less than Its corrected age of c. 74 0.8 ka and its internal structure in-
63 mm by settling in Atterberg cylinders, according to the Stokes dicates only that at that time there was already no water flow in
law. This fraction was treated with hexafluorosilicic acid (H2SiF6) the upper levels of the cave. However, this has little significance
for 10 days. Subsequently, the removal of the grains <4 mm was when considering the age measured for PU10-2, which was taken
carried out by centrifugation. from the outer flowstone crust layer that covers one of the walls of
For the annual dose determination, radionuclide specific activ- the Shaft (Fig. 3). The sample yielded a reliable age of w291 9 ka,
ities (U-238/Ra-226, Th-232, K-40) were measured using high which indicates that at that time the Shaft was already formed (i.e.
resolution gamma-ray spectrometry, and the dose rates were there was a relief of at least 15 m between the subterranean
calculated using the conversion factors tabulated by Adamiec and stream and the upper passages).
Aitken (1998). Equivalent dose (De) measurements were under- Down in the Excavation Chamber, the age of the PU10-3
taken on the standard Risø TL/OSL-DA-20 reader, equipped with sample is important because it was obtained from the basal
blue LEDs (470 30 nm). IR (875 80 nm) LEDs were used for part of a flowstone slab that is now suspended at w0.5 m above
infrared stimulation. Blue light stimulated OSL signal was detected the current excavation surface (Fig. S5). The sample is an ancient
through a 7.5 mm thick Hoya U-340 UV filter. Irradiations were flowstone that grew over a sediment stack that was
j.quaint.2013.10.012
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 232
Th/238U ¼ 1.250 0.625,
230
Th/238U ¼ 1 0.02 and 234U/238U ¼ 1 0.02.
238 234
Sample Lab number Description U [ng/g] U/238U 230
Th/238U 232
Th/238U 230
Th/232Th Calculated age Corrected age
number (ka) (ka)
PU10-1 GEB-UTO-6 Small stalagmite; base 133.16 0.73 1.414 0.003 0.725 0.005 0.0202 0.0001 35.73 0.26 75.153 0.692 73.975 0.812
PU10-2 GEB-UTO-7 Flowstone crust on the 149.74 0.87 1.044 0.002 0.985 0.006 0.0051 0 192.79 1.31 291.496 9.143 291.090 9.146
step; basal part
PU10-3 GEB-UTO-8 Flowstone at the excavation; 62.44 0.54 1.276 0.003 0.771 0.004 0.0649 0.0002 11.88 0.06 96.694 0.921 92.450 1.814
basal part
PU10-4 GEB-UTO-9 Stalagmite outer layer topping 81.63 0.47 1.514 0.004 1.212 0.006 0.1202 0.0003 10.08 0.04 152.323 1.781 146.2696.822
sediments
subsequently removed. Its minimal age of c. 92 1.8 ka attests (Passega diagram) (Fig. 6) displays a scattered pattern for Profile
for flowstone deposition process being active towards the end of PU1, suggesting a variable environment energy. However, most
MIS 5 with flowstone crusts covering more ancient detrital values are parallel to the C ¼ Md line indicating that “graded
deposits. suspension” have been the main transport mechanism. The
Finally, in the opposite end of the Excavation Chamber, the Estimated Settling Velocity (ESV) of the particles varies between
lateral flowstone was deposited aside a massive column and on top 0.58 and 81.77 103 cm/s. All parameters suggest a back-
of old silt deposits (sample PU10-4, Fig. S6) has a minimal age of swamp facies (sensu Bosch and White, 2004), with quasi-
w146 6.8 ka. During sediment sampling, the old base of the stagnant water and short transitions to a channel facies,
stalagmite, including a second such crust, laterally deposited, was marked by the higher sorting degree of several samples and
discovered buried in sediments at w0.9 m below (Fig. 5 and Fig. S6). indicating a more dynamic water.
The complex structure of the speleothem indicates that there were In the PU2 Profile, the values of Mo1, Md and Mzi are strongly
at least two ancient episodes of sediment accumulation in the influenced by the percentage of lutite-sized clasts (% lutite/Mo1: r:
Excavation Chamber followed by periods when water level was low 0.8614; P < 0.0001; % lutite/Md: r: 0.9105; P < 0.0001; % lutite/Mzi:
enough to allow growth of stalagmites and flowstone. The second r: 0.9437; P < 0.0001). The sorting degree of the deposits increases
such event took place some time before the end of MIS 6. Consid- with the content of silt fractions (% silt/si: r: 0.8431; P < 0.0001)
ering: (1) the suspended position of the PU10-3 flowstone and (2) and decreases with the content of lutite fractions (% lutite/si: r:
the current morphology of the outcrop, with the PU10-4 stalagmite 0.7594; P < 0.0001). A correlation between the main grain size
base partially exposed, we speculate that these accumulation statistical parameters could not be established, possibly owing to a
events were followed by periods of fluvial erosion and sediment more variable depositional environment. This variability is also
removal. 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
5.2. Cave sediments and OSL ages diagram. The sediment transport regimes correspond to either
“suspension or rolling” or “graded suspension” mechanisms.
Cave sediments successions studied from the three profiles in All the above suggests a backswamp facies, with transitions to a
the Scientific Reserve are presented in Fig. 5. Measured grain size channel facies, the latter being probably caused by short increases
and statistical parameters are presented in Table 2 (see also Fig. S7). of the water dynamics. The grain size of the sediment supplied to
In all profiles, the sediments predominantly include silt-size sedi- the cave has varied, possibly owing to changes within the source
ments which are finning towards the upper parts. The PU1 and PU2 area (one such event can be noted w1.6 m from the top and it is
upper parts consist of sub-horizontal plateaus showing ichnofa- marked by opposite trends for silt and arenite, and by higher values
brics (cave bear prints, scratches, and nests) and, in the case of PU2, of Mo1 and ESV).
two fully articulated cave bear skeletons. Profile PU4 shows a higher dynamic depositional environment,
For the PU1 profile, the statistical parameters calculated for with no clear correlations between the main grain size statistical
the grain size distribution of sediments show two very strong parameters. The values of the clasticity index increase with the silt
positive correlations between median and mean (Md/Mzi) (r: fractions content (% silt/C: r: 0.7530; P < 0.0001), while those of the
0.9692; P < 0.0001), and between median and sorting (Mzi/si) (r: first mode and mean increase with the lutite fractions content (%
0.9275; P < 0.0001), the latter showing a lower degree of sorting lutite/Mo1: r: 0.8028; P < 0.0001; % lutite/Mzi: r: 0.8111;
for the finest fraction. Positive correlations were calculated be- P < 0.0001); the values of the clasticity index, median and mean
tween the median and the skewness (Mzi/Ski) (r: 0.7933; decrease with the arenite-sized clasts percentage (% arenite/C:
P < 0.0001) and the sorting and skewness (si/Ski) (r: 0.7119; r: 0.8041; P < 0.0001; % arenite/Md: r: 0.9105; P < 0.0001; %
P < 0.0001), respectively. These indicate an excess of fine detritus arenite/Mzi: r: 0.7882; P < 0.0001). The sorting degree of the
with a low degree of sorting, owing to an increased input of lutite deposits decreases with the lutite fractions content (% lutite/si: r:
fraction (% lutite/Ski: r: 0.7464; P < 0.0001). The C/Md plot 0.9240; P < 0.0001). The fluctuating environment energy is also
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 4.77e12.21 5.06e8.31 5.18e9.03 0.91e3.21 0.03e0.61 0.49e2.24
PU2 50 0e68.79 29.83e96.93 0e59.01 0.56e6.43 3.2e13.39 3.61e12.65 4.21e11.19 0.8e4.49 0.73e0.64 0.51e2.68
PU4 46 0e40.09 20.02e92.15 1.08e77.04 0.76e4.08 2.03e13.39 2.44e7.24 3.13e8.45 0.93e4.13 0.11e0.62 0.54e1.96
j.quaint.2013.10.012
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 PU1 through PU4 suggesting that the upstream sections were
and by the ESV, which has values varying across a very large range: mostly deposited at high water levels with low hydraulic energy
3.19e1961.06 103 cm/s. The high variability of the ESV indicates (reservoir lake) while the downstream sections experienced
the occurrence of some water pulses. The projections on the Pas- alternating episodes of high water levels which deposited fine
sega diagram (Fig. 6) indicate that sediments were transported as sediments and torrential events that deposited coarser fractions
bedload, mainly by either “rolling and suspension” or “suspension and might have eroded through or even wash-out the older
and rolling”, and only secondarily by “graded suspension”. Similarly sediments.
to Profile PU2, in PU4 the sedimentological parameters point to an The results of the OSL dating carried out on seven sediment
overall backswamp facies, with short transitions to a channel facies, samples are presented in Table 3. The obtained ages clearly cluster
caused by a higher dynamics of the depositional environment. in two ranges. The first group shows theoretical ages of 210e
Overall, the sedimentology of the three investigated sediment 230 ka, with typical uncertainties of >10% (1s). These ages seem
profiles shows a consistent pattern, with a typical backswamp to be unreliable. Legitimate doubts concerning a potential
facies that includes transitions to channel facies during high- incomplete bleaching of the quartz grains can be raised according
energy hydraulic events. Evidence for such events increase from to the more general belief that quartz OSL dating for old (>70 ka)
j.quaint.2013.10.012
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., between the OSL ages of sediments and the radiocarbon ages of the
2012). However, it is interesting to note that these ages are in incorporated fossil bones may be due to either: (i) an incomplete
agreement with both the U/Th age of the PU10-4 flowstone (Fig. 5 reset of the OSL signal at the time of transport, or (ii) a reworking of
and S6), as well as with the U/Th age of the PU10-2 flowstone that older sediments from the remote parts of the cave towards their
grew onto the Shaft’s wall at w290 ka. We can only speculate that current position. Overall, although the OSL measurements did not
the bottom sediments were deposited at some early stage, whose yield a precise chronology of the sediment sections, they suggest at
age is about 210e230 ka. least two separate episodes of rapid sediment inflow into the cave.
The second group of ages belongs to the top of the sediment
sections and to the sediments in the excavation trench. They show 5.3. Radiocarbon dating results
calculated OSL ages that cluster between w67 and 90 ka. These ages
appear to be more reliable, and they generally follow the strati- The results of radiocarbon dating are given in Table 4. From a
graphical sequence. The ages measured for the Levels 4 and 5 of the total number of 25 submitted samples, we discarded the results
Excavation pit seem to be: (i) out of stratigraphical sequence (L5 that either show abnormally low %N (<0.5%) or collagen yields
yielded a younger age than L4) and (ii) in disagreement with the lower than 0.8%. All ages were calibrated using the OxCal
radiocarbon ages of the incorporated fossil remains which are 4.2.2 program (Bronk Ramsey, 2009) and the IntCal09 calibration
younger than w47 ka (see Section 5.3). However, as L4 is a debris curve (Reimer et al., 2009). The results are graphically shown in
cone originating from the sediments in the upper levels of the Shaft Fig. 7.
(Fig. 4), its older OSL age is explicable. Radiocarbon ages of the 15 fossil bone samples collected from
We therefore consider that the top deposits within the profiles the Scientific Reserve range between c. 42 and 45 ka (median
indicate a secondary sediment inflow phase with sediments being values) for both surface and excavated finds. We have discarded
transported into the cave as early as w90 ka. The contradiction 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
j.quaint.2013.10.012
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
Fig. 7. Radiocarbon ages of fossil bones from the Urşilor Cave, calibrated using OxCal indicates that no significant flooding occurred after c. 40 ka, as such
4.2.2 (Bronk Ramsey, 2009) and IntCal09 (Reimer et al., 2009). The NGRIP curve is an event would have been destroyed them. As well, the walls of the
shown for reference with the Heinrich (H) events marked. 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,
170 cm) that showed abnormally young ages. These samples also
and both present numerous bioglyphs, indicating post-flooding
show abnormally low %C with respect to %N measured for whole
cave bear activity.
bone (%Cb and %Nb). Normally, the %Cb would also include the “non
Finally, the presence of coarser sediments in the Upper Passage,
collagenous” carbon, which can be calculated using the equation %
which includes cave bear bones as old as 35e38 ka, is somehow
Cncoll ¼ %Cb (2.7 %Nb) (Bocherens et al., 2005, 2008). However,
surprising. They indicate a hydrologically dynamic environment
many samples for which %Cb and %Nb was available show a
and/or a reworking of older sediments due to a distinct water
negative %Cncoll and the two rejected samples show %Cncoll
source, (e.g. from a temporary stream originating in the upper part
ranging between 1.07 and 1.87. Such a depletion in non-collagen
of the Shaft). However, the precise location of the water inflow
carbon was associated with an almost total loss of collagen
remains uncertain.
(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 5.5. The general evolution of Urşilor Cave
samples, and (iii) the apparent stratigraphical inversion, we
decided to discard the samples which show a %C depletion higher When putting together the chronological information obtained
than 1%. Nevertheless, the issue of fossil samples that show such an so far on speleothems, sediments, and fossil remains from Urşilor
unexpected negative departure of %Cb when compared to %Nb re- Cave, a broad scenario emerges for the cave evolution during the
quires further attention and study. last w300 ka. This scenario is inevitably incomplete, given the
Collagen reported yields (not available for the CEDAD labora- uncertainties associated with the different dating results. Therefore
tory) were generally low (0.8e7.5%), as typical for many bone ac- only some key-episodes or key-moments in the evolution of the
cumulations from deep cave settings (e.g. Zilhao et al., 2007; for the karst system can be securely dated. We present these episodes and
Peştera cu Oase). Taking into account that the ages are close to the their associated chronology in Fig. 8 and describe them below.
j.quaint.2013.10.012
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 it. The OSL estimates for the sediment sample taken from w100 cm
The age of the flowstone covering the wall of the Shaft below PU10-4, as well as the two other samples from the lower part
(w291.1 9.15 ka) shows that more than w290 ka, the 17 m step in of the PU2 profile suggest a maximum age of c. 210 ka. We therefore
the Excavation Chamber was already formed and the subterranean infer that at some time between w210 ka and w145 ka, water
stream was sinking at a lower level, at the base of the Shaft. The flooded the Lower Passages having had a relatively high hydraulic
overall morphology of the cave suggests that the small shafts in the energy to deposit fine levels of sands. These episodes alternated
Passage of the Pits may have acted as overflow channels for the with periods when a backswamp lake formed and deposited the
water temporarily accumulated in the Excavation Chamber. The age finer silt or clay layers. The stratigraphy of the deposits, and espe-
of the flowstone corresponds to MIS 8. Although it may be sur- cially the fact that the PU10-4 stalagmite is partially engulfed in
prising that flowstone deposition was active during this cold stage, sediments (Fig. S6), suggest the existence of several flooding
MIS8 was comparable to the presumably warmer MIS3 and displays events, separated by at least one interval of low waters allowing the
a very similar Dansgaard-Oeschger (D-O) variability (Sidall et al., growth of the basal w70 cm of the PU10-4 stalagmite. All other
2007). On the other hand, at the low altitude of Urşilor Cave, the sedimentological parameters point to alternating episodes of low-
primary control for speleothem deposition would not normally be energy hydraulic regimes depositing fine sediments within a
the annual mean temperature but rather percolation water backswamp facies, and short transitions to high-energy waters
availability. which were able to deposit coarser sediments (channel facies) and
to partially erode older terraces.
5.5.2. Stage 2: between w210 and w145 ka No bone remains have been found in the sediments of this stage,
The age of w146.3 6.8 ka measured for the PU10-4 speleo- and there is no indication that the cave might have been inhabited
them in the Excavation Chamber marks another chronological by any animals at that time. On the other hand, it is interesting to
constraint in the evolution of Urşilor Cave. The part of the stalag- note that the age measured for the PU10-4 stalagmite also fells
mite that grew atop the sediment terrace of PU4 clearly postdates within a cold interval, MIS 6. Although calcite precipitation rates in
j.quaint.2013.10.012
j.quaint.2013.10.012
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/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/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
Romanian karst are known to be generally lower during glacials the central channel were scrambled and transported downstream.
than during interglacials, speleothem formation is documented Any new flooding would bring new, either younger or older
during MIS 6 in many cave sites located at middle latitudes (e.g. Bar- (reworked), fossil remains to fill these gaps. This may explain why
Matthews et al., 2003; Constantin et al., 2006; Wainer et al., 2011). some discrepancies may exist between radiocarbon ages of remains
The stereotype according to which speleothem formation is more from what is now perceived as a “single” stratigraphic level.
likely to correspond to warm climate phases while sediment ac- We cannot speculate about when the bear occupation started in
cumulations are more likely related to colder ones cannot be the Scientific Reserve of Urşilor Cave. What is clear is that by
ascertained. Hence, we do not have any evidence to link these first w45 ka this cave sector was hosting a large number of cave bears,
episodes of sediment deposition to either a warm or a cold stage and possibly also cave lions. It was also about this time when the
and those sediments may have been transported during either first flooding event raised the PU2 river terrace by about 1 m and
MIS7 or MIS6, or both. created the PU1 terrace deposit. Subsequently, the cave bears
continue to live in the cave, and all terraces and sediment faces
5.5.3. Stage 3: w92.5 ka have preserved their vital traces. The same intensive cave bear
At c. 92.5 ka, the PU10-3 flowstone from the downstream end of usage of the cave is documented as well in the Upper Level. The
the Excavation Chamber was growing on top of previously depos- next flooding events led to the accumulation of new fossil remains,
ited sediments. It is now suspended at w 0.5 m above younger which may originate from either the Lower Level (by fluvial
sediments and fossil bones, thus adding evidence for the scenario of transport of bones or drowning) or the Upper Level (by fluvial
periodical high-energy hydraulic events able to wash out the pre- transport or individual animals falling down the Shaft).
viously deposited sediments. On the other hand, the precipitation At the present state of research, the interpretation of the cave
of this flowstone indicates an extended time period of low waters, system evolution during this stage faces several uncertainties
as flowstone is known for its low growth-rates and there are no associated with both the OSL and radiocarbon ages and also to our
visible impurities in the cross-section to indicate some sort of pe- understanding of the accumulation processes. However, virtually
riodical flooding. The obtained age corresponds to the beginning of all dated fossil remains show ages between w45 and w42 ka. While
the cold MIS 5b. When correlated with the PU10-4 and PU10-2 it is very tempting to correlate this taphonomic event with a cli-
ages, it suggests that colder periods may have been favorable for matic oscillation during MIS3, evidence is not solid enough to
uninterrupted speleothem deposition and that the sediment in- ascertain one specific event. For instance, the H5 event (as many
flows might have been linked to warmer stages. 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
5.5.4. Stage 4: between w92.5 and w40 ka associated with the radiocarbon ages at this time are simply too
Sediments from the top part of the PU2 profile, the PU1 profile, high to establish such a correlation. In the case of Peştera cu Oase
as well as those from the excavated trench, have shown broadly (SW Romania), where a similar thanatocenosis of a roughly similar
concordant OSL ages, which are interpreted as being younger than age was described, the favored interpretation was that at the onset
w90 ka. In addition, they all show similar sedimentological fea- of GIS 12 the snow cover that accumulated during the w1000
tures: they almost exclusively include silts and laminated clays and years-long H5 event rapidly melted and generated a massive
point towards a backswamp-type depositional facies. flooding event (Constantin et al., 2013b).
The main ambiguity concerning the real age of these sediments
stems from the fact that the two samples taken from the excavation 5.5.5. Stage 4: between w40 ka and w30 ka
trench correspond to stratigraphical layers that include fossil bones There is no reliable radiocarbon date or other evidence to
which are as young as w42 ka. Hence, we can only assume that prove cave bear presence in the Lower Level of Urşilor Cave after c.
either the OSL signal was not completely reset or that the sedi- 40 ka. There is also no hard evidence attesting to any notable
ments have been reworked from deeper parts of the cave. Either hydraulic events, and the fact that several w45 ka old skeletons
way, the depositional event(s) took place at some time between remained preserved on top of the terraces of the Scientific Reserve
w47 and 45 ka the ages of the articulated skeletons lying on top of actually rules out any major floods. Radiocarbon dates indicate the
the alluvial terraces and w42 ka (the youngest reliable radiocarbon presence of cave bears in the Upper Passages as late as w30 ka.
age of an articulated skeleton found in the excavation trench). This raises the question of where might have been the original
When considering the stratigraphy of the deposit in the entrance(s) used by cave bears to get into the cave. As the natural
excavation trench, at least two flooding events are apparent. entrances are now sealed by sediments and speleothems, we can
Their overall sedimentologic and taphonomic features suggest only assume that the entrance that allowed cave bears in the
short but intense flooding, with the instant burial of numerous Upper Passages continued to exist. On the contrary, a second
cave bears (and at least one cave lion), as shown by the presence entrance which may have allowed animal access into the Lower
of their articulated skeletons. Given the large uncertainties of the Level was already sealed.
radiocarbon dates, the timing of these events is hard to separate. The mechanism of deposition of the last sediments levels in the
The presence of L8 flowstone crusts above the L5 suggests that Upper Passages is still uncertain. In the test trench of the Access
the floods were separated in time by at least several hundred Passage the sediment consists of pebbles, coarse sand and only a
years. low amount of clay, accounting for a high hydraulic energy, while
However, considering that all excavated speleothems are posi- also including cave bear bone fragments reliably dated to w35e
tioned towards the side of the passage (i.e. the lateral “terrace”), we 37 ka. The only reasonable explanation for this would be that a
cannot exclude that the first flooding event was followed by a rapid second stream flowed through the Upper Passages at that time,
decrease of water level during which the stream has flown only although its origins are unknown. This contradicts the common
along the central channel, thus allowing for subaerial deposition of views (e.g. Valenaş, 1979) according to which Urşilor Cave was
speleothems. A new flood would have add a new layer of fine created by a single underground stream.
sediments and bones (both carcasses and reworked), covering the
entire floor of the chamber, including the small lateral terrace. If 5.5.6. Stage 5: after w30 ka and through the Holocene
this scenario repeated, it seems plausible that during the The latest dated cave bear remains from the Passage of the Pits
“erosional” phases bones which were at that time positioned along have ages close to c. 30 ka. In the absence of any evidence, we cannot
j.quaint.2013.10.012
speculate about the time when cave bears and other large mammals given thanatocenosis. This probably happens more often than
stopped using the cave. The only chronological controls are: (1) the considered in many reported cave sites, as shown in the case of the
abundant presence of the candlestick stalagmites in the Upper Peştera cu Oase (Constantin et al., 2013a) when the initial,
Level, including some grown within hibernation nests or covering simplistic approach was soon challenged by field data and the final
fossil remains and which are known to be Holocene (Onac et al., history of the fossil accumulation proved to be more complicated
2002; Constantin, 2003) and (2) the fact that the natural en- than initially thought.
trances were clogged by sediments and/or sealed by flowstone at
the time of discovery. Holocene speleothem formation is known to 6. Conclusions
be very fast and abundant in caves of Romania (Constantin, 2003;
Constantin et al., 2006, 2013a). We speculate that the closure of This paper described the sedimentological, geomorphological,
the Urşilor Cave was caused by either small collapses at the end of and geochronological work undertaken at the Urşilor Cave with the
the Last Glacial Maximum (LGM) or clogging by speleothem depo- aim of understanding the taphonomy of the fossil accumulation in
sition or a combination of both. We have no evidence to document the Excavation Chamber. The initial hypotheses, which assumed
the cave evolution during the MIS2 and the last deglaciation. that the fossil remains accumulation was caused by a combination
of collapses, river deepening, and underground flooding, were
5.6. Summary: reconstruction of the sedimentary history of the largely erroneous. The “separation” of the cave system through the
cave and the taphonomy of fossil deposits Shaft step was much older than previously thought and the accu-
mulation of sediments in the lower level of the cave consisted of a
The sedimentological research and dating of the various de- long history of depositional and erosional events. It was only the
posits in Urşilor Cave led to a better understanding of the evolution last suite of such events (broadly dated between 47 and 40 ka) that
of the cave during the last w300 ka and substantially changed some led to the accumulation of fossil remains. The upper levels of the
of the hypotheses concerning the cave’s history. The cave bear fossil cave continued to be occupied by cave bears and seem to have
assemblage in the Lower Level was formerly associated with a evolved separately with respect to the lower ones, even having
massive collapse in the Shaft sector that prevented bears from their own drainage system.
exiting the cave. This is now shown to be erroneous and, even Considering the ages of the dated speleothems and those of the
though a collapse cannot be totally ruled out, it did not influence other cave deposits, it appears that: (1) speleothem precipitation
the fate of the large mammals from this sector since it happened was not prevented during climatic cold periods; (2) sediment
much earlier. depositional events were more likely linked to warmer and wetter
Uranium-series dating of several flowstone levels associated periods. We tentatively assign the fossil accumulation from the
with the OSL dating on sediments and the radiocarbon dating of Excavation Chamber to a series of floods that most probably took
fossil bones indicate at least two broad periods of sediments input place at the end of the H5 event and the onset of the GIS 12. The
into the Lower Level of the cave. These probably included alter- speleogenesis of the Urşilor Cave, which appears to be more
nating depositional and erosional phases which complicated the complicated than initially thought, may bring to attention the need
interpretation of the fossil deposit and scattered remains. Our data for careful research of the sedimentary history of any paleonto-
indicate that only the last floods (sometime between w47 and logical site located in deep-cave settings.
w40 ka) were responsible for the accumulation of fossil remains
and for the death of the cave bears and lions.
Acknowledgements
On the other hand, the fact that the Shaft was already formed at
that time indicates that the animals used another entrance, since it
This research was funded through the KARSTHIVES Project e
is unlikely that so many individuals would enter so deep into the
“Climate archives in karst”, funded by CNCS e UEFISCDI Grant
cave while coping with a 17 m deep step. Such an entrance could
PCCE_ID 31/2010. We are grateful to Tomasz Goslar and Hervé
have been located in the middle part of the cave, close to the
Bocherens for insightful discussions on the radiocarbon dating is-
present-day Excavation Chamber. This entrance must have been
sues, to Martina Pacher for her help with the VERA dating, and to
opened even after the last flooding, as cave bears were still leaving
Oana Moldovan for her constant help during this research. We
their vital traces throughout.
thank Valerica Toma and Augustin Nae for their help in the field and
The fact that in the Upper Passages cave bears continued to live
the Apuseni National Park for granting the excavation permit and
for at least 10,000 years longer suggests that these passages were
field assistance.
actually a “separate” cave, with the Shaft acting as a barrier for the
SC designed research and secured funding. MR, AP, MV, IM, MK,
bear population. The presence of different sediments in these
SC, VD, and CP performed research. CMM, RDR, and CP have done
passages, indicative of a higher hydraulic gradient, can only be
the sedimentological study, VD and DH have done the U-series
explained by admitting that a second underground stream was still
dating, VA and ATG carried out the OSL dating, and SC and MR wrote
flowing through the upper parts of the system concurrently with
the paper. All coauthors contributed to improve the manuscript.
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 Appendix A. Supplementary data
deglaciation (Ménot et al., 2006).
When trying to reconcile all the data yielded by our research, Supplementary data related to this article can be found at http://
the former theories that explained the evolution of the cave as a dx.doi.org/10.1016/j.quaint.2013.10.012.
unitary karst system affected by underground stream piracy and
collapses seem to be too simplistic. The final picture that emerged, References
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Quaternary International xxx (2013) 1e16

Contents lists available at ScienceDirect

Reconstructing the evolution of cave systems as a key to

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

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

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

In the southern end of the chamber, a massive stalagmite has its

3.2. Subterranean terraces proﬁles

Along the subterranean stream, two terrace proﬁles were also

limit of radiocarbon, the calibrated values show relatively high

5.4. Fossil accumulation and hydrological regimes

The excavation of Urşilor Cave yielded a large number of artic-

The Scientiﬁc Reserve (Lower Level)

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

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