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Life and Death at the Pe ş tera cu Oase

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HUMAN SERIES

Series Editors Russell L. Ciochon, The University of Iowa Bernard A. Wood, George Washington University

Editorial Advisory Board Leslie C. Aiello, Wenner-Gren Foundation Susan Ant ó n, New York University Anna K. Behrensmeyer, Smithsonian Institution Alison Brooks, George Washington University Steven Churchill, Duke University Fred Grine, State University of New York, Stony Brook Katerina Harvati, Univertit ä t T ü bingen Jean-Jacques Hublin, Max Planck Institute Thomas Plummer, Queens College, City University of New York Yoel Rak, Tel-Aviv University Kaye Reed, Arizona State University Christopher Ruff, John Hopkins School of Medicine , Washington University in St. Louis Carol Ward, University of Missouri

African Biogeography, Climate Change, and Evolution Edited by Timothy G. Bromage and Friedemann Schrenk Meat-Eating and Edited by Craig B. Stanford and Henry T. Bunn The Skull of afarensis William H. Kimbel, Yoel Rak, and Donald C. Johanson Evolution in Central Europe: The People of Doln í V ĕ stonice and Pavlov Edited by Erik Trinkaus and Ji ří Svoboda Evolution of the Hominin Diet: The Known, the Unknown, and the Unknowable Edited by Peter S. Ungar Genes, Language, & Culture History in the Southwest Pacifi c Edited by Jonathan S. Friedlaender The Lithic Assemblages of Qafzeh Erella Hovers Life and Death at the Pe ş tera cu Oase: A Setting for Modern Human Emergence in Europe Edited by Erik Trinkaus, Silviu Constantin, and Jo ã o Zilh ã o

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Life and Death at the Pe ş tera cu Oase A Setting for Modern Human Emergence in Europe

Edited by Erik Trinkaus , Silviu Constantin, Jo ã o Zilh ã o

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Library of Congress Cataloging-in-Publication Data

Life and death at the Pestera cu Oase : a setting for modern human emergence in Europe / edited by Erik Trinkaus, Silviu Constantin, and João Zilhão. p. cm. — (Human evolution series) Includes bibliographical references and index. ISBN 978–0–19–539822–9 (hardcover : alk. paper) 1. Prehistoric peoples—. 2. Cave dwellers—Romania. 3. Human evolution—Romania. 4. Human remains ()—Romania. 5. remains (Archaeology)—Romania. 6. Romania—Antiquities. I. Trinkaus, Erik. II. Constantin, Silviu. III. Zilhão, João. GN776.R8L54 2012 599.93’8—dc23 2012005790

ISBN: 978–0–19–539822–9

9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper

00_Trinkaus_Prelims.indd iv 8/31/2012 10:06:30 PM Contents

Contributors xi

I. INTRODUCTION AND BACKGROUND 1 1. Introduction (Erik Trinkaus, Silviu Constantin, Jo ã o Zilh ã o) 3

2. Exploration and Documentation of the Peş tera cu Oase (Ş tefan Milota, Mircea Gherase, Laurenţ iu Sarcină , Ricardo Rodrigo, Oana Moldovan, Erik Trinkaus, Silviu Constantin, Jo ã o Zilh ã o) 5

3. Problems, Approaches and Fieldwork: 2004–2005 (Jo ã o Zilh ã o, Ş tefan Milota, Ricardo Rodrigo, Silviu Constantin, Erik Trinkaus ) 16

II. THE CAVE AND ITS CONTENTS 31 4. The of the Ponor-Plopa System in Its Regional Context (Silviu Constantin) 33

5. The Ponor-Plopa Cave System: Description, Sediments, and Genesis (Silviu Constantin, Cristian-Mihai Munteanu, Ş tefan Milota, Lauren ţ iu Sarcina, Mircea Gherase, Ricardo Rodrigo, Jo ã o Zilh ã o) 41

6. Uranium-Series Dating on from the Ponor-Plopa Cave System (Silviu Constantin, Stein-Erik Lauritzen) 56

7. Electron Spin Resonance Dating of Vertebrate Remains from the Peş tera cu Oase (Rainer Gr ü n, Tegan Kelly, Maxime Aubert) 73

8. of the Pe ş tera cu Oase Faunal Remains (Thomas Higham, Eva Maria Wild) 77

9. Rock Magnetic Data of Late Sediments from the Pe ş tera cu Oase and Their Paleoclimatic Signifi cance (Cristian Panaiotu, Silviu Constantin, Că t ă lin Petrea, Viorel Horoi, Cristina-Emilia Panaiotu) 86

10. The Distributions of Finds and Features (Jo ã o Zilh ã o, Ricardo Rodrigo, H é l è ne Rougier, Ş tefan Milota) 100

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III. THE CAVE BEARS 125 11. and Paleobiology at the Pe ş tera cu Oase: Population Structure and Size Variability (Martina Pacher, J é r ô me Quil è s) 127

12. Taphonomic Analysis of the Cave Bear Remains from the Peş tera cu Oase (Martina Pacher, J é r ô me Quil è s) 147

13. An Ancient DNA Perspective on the Cave Bears from the Pe ş tera cu Oase (Mathias Stiller, Michael Hofreiter) 159

IV. THE OTHER VERTEBRATES 165 14. , Wolves, and from the Pe ş tera cu Oase (Emil Ş tiuc ă , Alexandru Petculescu) 167

15. Small Remains from the Pe ş tera cu Oase (Alexandru Petculescu) 185

16. The Ungulates from the Pe ş tera cu Oase (Simon A. Parfi tt, Adrian M. Lister) 189

17. Taphonomic Considerations of the Non-Ursid Vertebrate Remains (Erik Trinkaus, Martina Pacher) 204

18. Stable Isotopes and Dietary Patterns of the Faunal Species from the Peş tera cu Oase (Erik Trinkaus, Michael P. Richards) 211

V. THE HUMAN REMAINS 227 19. Radiocarbon Dating of the Pe ş tera cu Oase Human Remains (Erik Trinkaus) 229

20. The Human Mandible from the Pe ştera cu Oase, Oase 1 (Erik Trinkaus, Hé l è ne Rougier) 234

21. The Human Cranium from the Peş tera cu Oase, Oase 2 (Hé l è ne Rougier, Erik Trinkaus) 257

22. Contours of the Oase 2 Cranium (Sheela Athreya, H é l è ne Rougier) 321

23. The Internal Cranial Morphology of Oase 2 (Marcia Ponce de Le ó n, Christoph P. E. Zollikofer) 332

24. The Dental and Alveolar Remains of Oase 1 and 2 (Erik Trinkaus, Shara E. Bailey, H é l è ne Rougier) 348

25. The Oase 1 Human Stable Isotopes (Michael P. Richards, Erik Trinkaus) 375

VI. DISCUSSION 383 26. The Paleoenvironmental Context of the Pe ş tera cu Oase (Silviu Constantin, Jo ã o Zilh ã o, Erik Trinkaus) 385

27. Paleoanthropological Implications of the Pe ş tera cu Oase and Its Contents (Erik Trinkaus, Jo ã o Zilh ã o) 389

VII. REFERENCES 401 28. References 403

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Sheela Athreya Viorel Horoi Department of Anthropology, 234 Anthropology D é partement des Sciences de la Terre et de l’Atmosph è re, Building, 4352 Texas A&M University, College Station, Universit é de Qu é bec à Montr é al, Montr é al, Qu é bec TX 77843, USA H3C 3P8, ; and Institutul de Speologie “Emil Racovi ţă ,” str. Frumoasă 31, 010986 Bucharest, Maxime Aubert Romania Research School of Earth Sciences, Australian National University, Canberra ACT 0200, Tegan Kelly Research School of Earth Sciences, Australian National Shara E. Bailey University, Canberra ACT 0200, Australia Department Anthropology, New York University, 25 Waverly Place, New York NY 10003, USA Stein-Erik Lauritzen Department of Earth Sciences, Bergen University, Silviu Constantin Allegaten 41, N-5007, Bergen, Institutul de Speologie “Emil Racovi ţă ,” Department of Geospeleology and Paleontology, str. Frumoasă 31, Adrian M. Lister 010986 Bucharest, Romania Department of Palaeontology, Natural History Museum, Cromwell Road, London SW7 5BD, UK Mircea Gherase Pro Acva Grup, Str. Surduc, Nr. 1 (Sc. A, Ap. 4), 300552 Ş tefan Milota Timi ş oara, Romania Pro Acva Grup, Str. Surduc, Nr. 1 (Sc. A, Ap. 4), 300552 Timi ş oara, Romania Rainer Gr ü n Research School of Earth Sciences, Australian National Oana Moldovan University, Canberra ACT 0200, Australia Institutul de Speologie “Emil Racovi ţă,” Cluj Branch, Clinicilor 5, P.O. Box 58, 400006 Cluj-Napoca, Thomas Higham Romania Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History of Art, Dyson Perrins Cristian-Mihai Munteanu Building, South Parks Road, Oxford OX1 3QY, UK Institutul de Speologie “Emil Racovi ţă ,” Calea 13 Septembrie, nr. 13, 050711 Bucharest, Romania Michael Hofreiter Evolutionary Biology and Ecology, Department Martina Pacher of Biology, University of York, Wentworth Way, Institut f ü r Pal ä ontologie, Universit ä t Wien, Heslington, York YO10 5DD, UK Althanstrasse 14, 1090 Wien, Austria

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Cristian Panaiotu Ricardo Rodrigo Bucharest University, Paleomagnetism Laboratory, Sociedade Torrejana de Espeleologia e Arqueologia, B ă lcescu 1, 010041 Bucharest, Romania Quinta da Lez í ria, 2350 Torres Novas, Portugal

Cristina-Emilia Panaiotu H é l è ne Rougier Bucharest University, Paleomagnetism Laboratory, Department of Anthropology, California State University B ă lcescu 1, 010041 Bucharest, Romania Northridge, 18111 Nordhoff St., Northridge, CA 91330, USA Simon A. Parfi tt Department of Palaeontology, Natural History Museum, Lauren ţ iu Sarcin ă Cromwell Road, London SW7 5BD, UK; and Institute Pro Acva Grup, Str. Surduc, Nr. 1 (Sc. A, Ap. 4), 300552 of Archaeology, University College London, 31–34 Timi ş oara, Romania Gordon Square, London WC1H 0PY, UK Mathias Stiller Alexandru Petculescu Department of Ecology and Evolutionary Biology, Institutul de Speologie “Emil Racovi ţă,” str. Frumoas ă 31, University of California Santa Cruz, 1156 High Street, 010986 Bucharest, Romania Santa Cruz, CA 95064, USA

C ă t ă lin Petrea Emil Ş tiuc ă Institutul de Speologie “Emil Racovi ţă ,” str. Frumoas ă 31, Institutul de Speologie “Emil Racovi ţă ,” str. Frumoas ă 31, 010986 Bucharest, Romania 010986 Bucharest, Romania

Marcia Ponce de Le ó n Erik Trinkaus Anthropologie, Y42 K 22, Winterthurerstrasse 190, 8057 Department of Anthropology, Washington University, Z ü rich, Switzerland Saint Louis MO 63130, USA

J é r ô me Quil è s Eva Wild Unit é Toulousaine d’Arch é ologie et d’Histoire (UMR Institut f ü r Isotopenforschung und Kernphysik der 5608), Maison de la Recherche, Universit é Toulouse Universit ä t Wien, VERA-Laboratorium, Waehringer Le Mirail, 5 all é es Antonio Machado, 31058 Toulouse Strasse 17, 1090 Wien, Austria c é dex 9, France Jo ã o Zilh ã o Universitat de Barcelona/ICREA, Departament de Michael P. Richards Prehistò ria, Historia Antiga i Arqueologia, Facultat de Department of Anthropology, University of British Geografi a i Hist ò ria, Montalegre 6, 08001 Barcelona, Columbia, Vancouver, B.C. V6T 1Z1, Canada; and Department of Human Evolution, Max-Planck Christoph P.E. Zollikofer Institute for Evolutionary Anthropology, Deutscher Anthropologie, Y42 K 22, Winterthurerstrasse 190, 8057 Platz 6, 04103, , Z ü rich, Switzerland

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Part Six Discussion

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2 6 The Paleoenvironmental Context of the Pe ş tera cu Oase

Silviu Constantin , Jo ã o Zilh ã o, and Erik Trinkaus

Introduction This chapter summarizes the conclusions on the pale- oenvironmental context of the Peş tera cu Oase that arose The complex research undertaken at the Peş tera cu Oase from different lines of evidence. Such evidence came from was triggered by the discovery of the oldest directly dated, the dating of a relatively large number of speleothems substantial modern human remains and by the legitimate (Chapter 6), the analysis of sedimentological and mag- hopes that the deposits might yield an archeological con- netic properties of two cave-sediment profi les (Chapters 5 text or at least archeological elements. Given the richness and 9), the large amount of data gathered from the study of the paleontological deposit and the inevitable com- of the cave bear remains (Chapters 11 and 12), as as plications associated with the evolution of an extensive of the herbivore (Chapter 16), micromammals cave system, a large array of investigation methods was (Chapter 15), and detailed of the bone accu- deployed, some of them targeting cave sectors beyond the mulation on the Panta Str ă mo ş ilor (Chapter 10). Oase sector sensu strictu. Although the initial goal of fi nd- ing more archeological elements was not achieved, the Pe ştera cu Oase team managed to gather a considerable The Timescale Issue amount of informative data concerning both the genesis of the cave system and its paleoenvironmental and pale- The largest diffi culty when trying to interpret past envi- oclimatic context. ronments from multiple archives from a given site is that In recent , paleoclimate/paleoenvironment different time scales are being used. When not coupled research of the Quaternary continental realm increas- with stable isotope records, datings alone ingly uses cave deposits as valuable archives that may would typically offer a statistical, broadscale view on contain relevant proxies. Cave speleothems may yield, what in the past may have been the periods more or less when accurately dated, high-resolution isotopic records, favorable for soil formation and underground sometimes annually resolved, that are relevant for deposition. understanding past climate changes. Even the simple The geological context and the inferred evolution of absence or presence of speleothems at a given moment the Ponor-Plopa cave system (Chapters 4 and 5) show in time is relevant to qualify surface climate conditions that the history of the system extends well beyond ~210 such as absence or presence of permafrost. Fossil remains ka cal BP, which is the oldest age measured for a speleo- preserved in cave sediments may offer insights into them from the Galeria Culcu ş urilor. This Marine Isotope the paleodiets or paleobiogeography of past , Stage (MIS) 7 age and the subsequent MIS 5 ages meas- although these are likely to apply to discrete moments ured for other speleothems formed on the rock terraces only. In a complex site, such as the Ponor-Plopa system, in Galeria Lung ă or fragments found within the Panta the research inevitably yields disparate elements of a Str ă mo şilor deposit show that at least during those inter- complicated puzzle that, for the most part, needs addi- glacials the upper passages of the system were already tional pieces of evidence to be resolved. formed and they functioned as dry-cave galleries with

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only occasional (if any) fl uviatile input. Speleothem for- Where cave records are concerned, it is interesting mation during interglacials in southwestern Romania to note that, in many sites, speleothem growth seems is well documented (Constantin, 2003) and attests for to be less affected by the warm–cold alternations. For a temperate climate with suffi cient rainfall to allow example, in Kleegruben Cave (Sp ö tl et al., 2006), located rapid calcite deposition. The large number of dated spe- at an elevation of more than 2000 meters in the Austrian leothems that started to grow during the Holocene or Alps, the speleothems actively grew during the stadials, the Tardiglacial, sometimes showing very high growth despite their high elevation. In the (south- rates (see Chapter 6), suggests that broadly similar cli- western France) record (Genty et al., 2010), speleothems mate conditions as today may be inferred for the MIS 5 were actively growing and have recorded the H5 and H4 and 7. events, while one shows a hiatus during the warm GI 14 The largest body of data obtained from the study of episode. These are only two examples of apparently par- the Pe ştera cu Oase corresponds to a shorter time range, adoxical speleothem growth during “cold” rather than within the middle MIS 3, between ~50 and ~30 ka cal BP, “warm” MIS 3 oscillations. Coming closer to the Peş tera a period that is known for its rapid climatic oscillations. cu Oase, the Poleva Cave record (Constantin et al., 2007), We can defi ne this as an intermediate timescale, where a although less well dated, shows a marked decrease in wider range of proxy data are available within the same δ 18 O values corresponding to the H5 event and a hiatus period and with similar uncertainties, and they may be marked by a clay intercalation broadly corresponding to corroborated or validated against each other. It is for GI 12. This is followed by a new decrease in δ 18 O values this time period that we may be able to “zoom in” in an centered at ~38 ka cal BP and considered to represent the attempt to describe the past environmental conditions of H4 event. Finally, in the Sofular Cave record from north- southwestern Romania. ern Turkey (Fleitmann et al., 2009), a shows a It has to be noted, however, that the average duration continuous growth (and isotope record) during the 50–35 of the climatic oscillations known in MIS 3 is only of a ka cal BP time range. couple of thousands years or so. When considering (and The previous examples suggest that MIS 3 climate combining) the typical uncertainties of different absolute oscillations may not represent suffi ciently drastic changes age dates, this means that one is not always able to estab- in temperature to prevent speleothem formation as a lish a satisfactorily explained climatic model and that any result of the absence of percolating water caused by either inferred environmental conditions should be approached aridity or permafrost conditions. Moreover, speleothem with caution. growth cessation associated with climatic oscillations Finally, there is always the issue of short timescale may represent a side effect of local changes in hydraulic records, such as the isotope records from single individu- regime of the karst aquifer or karst streams, including the als or even populations of mammals or high-resolution sudden fl ooding of passages. isotope records from speleothems. No matter how allur- In the present settings of southwestern Romania, ing such interpretations may be, they are always associ- with low elevation hills and mountains and within a mild, ated with dating uncertainties that are at least one order sub-Mediterranean climate, calcite speleothems precipi- of magnitude higher than the theoretical resolution of the tate abundantly in all seasons with higher growth rates record. These uncertainties should always be accounted during the winter (S. Constantin, experimental unpub- for, and it is with them in mind that we approach these lished data). In the Pe ştera cu Oase, had been concluding comments. actively growing during the last glacial maximum (LGM) in the Galeria Lungă (Chapter 6), so we may infer that even the LGM conditions were mild enough to prevent What Meant “Warm” and What Meant “Cold” the formation of a permafrost layer, at least above the during MIS 3 in Southwestern Romania? Plopa Plateau. Going further back in time, speleothems actively grew in the Pe ş tera cu Oase during MIS 3. In the The MIS 3 climate history broadly includes a typical alter- case of the PPL6b stalagmite, the theoretical growth rate nation of short periods (usually up to 2000 years long) was rather high, of ~6 cm/ka during the GI 8–GI 7 time of abrupt warming and cooling of the climate. Sudden period, similar to that calculated for some Holocene sta- warmings of the climate (Greenland Interstadials—GI) lagmites from Poleva Cave (Constantin, 2003; Constantin were followed by gradual coolings; the major cooling epi- et al., 2007). sodes (Greenland Stadials—GS) coincide with Heinrich The combined results of the speleothem dating from events. The timing of these events is currently well the Panta Stră mo ş ilor, together with the sedimentological resolved from both ice core and other continental (includ- and taphonomic study of the deposit, show that the tor- ing speleothem) proxies (Svensson et al., 2008; Sanchez rential event that generated Level 2 occurred prior to ~47 Go ñ i and Harrison, 2010). ka cal BP (Chapters 6 and 10). On the other hand, Level 2

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features a micromammal association that is indicative of To conclude, corroborating the various pieces of a cool climate with water that remain unfrozen information obtained from the cave deposits at Peş tera at least during the warm season (Chapter 15). Namely, the cu Oase for the middle part of the MIS 3, it appears rea- presence of Microtus nivalis , a species currently found at sonable to consider that, at least in this region, the cold elevations above 1600 m, indicates an episode of strong stages were, in fact, more like prolonged periods of hard, cooling that might be correlated to the H5 event. but still temperate, winters while the interstadials were The units found in the Pu ţ ul and considered brief phases of sudden warming causing episodes of tor- as being debris fl ows washed from the Oase sector all rential fl ooding of the underground galleries. Why this show low magnetic susceptibility values, again indica- cyclic scenario lasted only until about ~35 ka ago remains tive of a colder climate, but the chronology of this pro- an unanswered question that, as in many cases in karst fi le is uncertain (Chapter 9). While it is clear that the bone regions, may well have more to do with fast surface inci- accumulation in the uppermost, excavated part of Level sion and local drainage reorientations. 2 was caused by a torrential event (or series of events), it is hard to ascertain whether this happened during the H5 event or at the onset of GI 12. This is especially diffi cult The MIS 3 Environment at the Pe ş tera cu Oase considering the short duration of the H5 event, between 47.74 and 46.68 ± 0.8 ka cal BP (Genty et al., 2010) and the When compared with present-day , the fossil spe- typical dating uncertainties for both speleothem and radi- cies associations discovered at the Peş tera cu Oase would ocarbon dates connected with Level 2. Considering the rather be expected in regions located at northern lati- reduced catchment area of the Plopa Plateau, from which tudes and higher elevations than the ones of the Aninei the fl uvial waters that fl ooded the şPe tera cu Oase must Mountains. have originated, our preferred interpretation for the exca- As shown in Chapter 16, the large body size of the vated, upper reaches of Level 2 is that at the onset of GI 12 Cervus elaphus (the largest known in Pleistocene Europe the snow cover that accumulated during the ~1000 and similar to large North American wapiti) suggests long H5 event rapidly melted and generated a massive that vegetation conditions were highly favorable at Oase fl ooding event. ~50 ka cal BP, with a ready supply of high-quality for- Further evidence from the Panta Stră mo ş ilor spe- age and moderate amounts of winter snow. On the other leothem and stratigraphy study shows that a new hand, the association of described from Level 2 lower-energy fl ooding event occurred at some point in of the excavations indicates a rather colder climate and time between ~42.5 and ~40.2 ka cal BP and displaced to requires the presence of some water reservoirs. The gen- the Panta Stră mo şilor part of the bones accumulated in the erally large body size of the cave bears (Chapter 11) sug- upper galleries in the meantime, thus forming the Level 1 gests favorable ecological conditions, while the inferred massive bone bed. It is interesting to note that the more omnivorous diet than has been proposed for cave faunal association includes species typical for a temper- bears elsewhere (Chapter 18) also requires diverse food ate climate such as Cricetus cricetus but also M. nivalis , sources, likely including fi sh. Indeed, the large body size which is indicative of a colder climate. While this may be of the bears and the omnivorous diet indicated by their a refl ection of successive temperate and cold phases, the isotopic profi les may be covariates of a relatively rich lower hydraulic energy of the event that deposited Level environment for these animals, despite the lack of correla- 1 shows that less water was available (either as melted tion between the limb dimensions and δ15 N values within snow or torrential streams). the Oase bear sample. Moreover, the combination of these It is interesting to note that the deposition of the large cervids and ursids, together with the micromammal Oase 2 human remains took place on top of a surface remains from the Panta Stră mo ş ilor and the presence of fossilized by the basal calcite layers of the PPL6b sta- wolves throughout the sequence (Chapters 14 and 15), lagmite with an age of ~40.5 ka cal BP, which immedi- indicate a diversity of habitats available in the vicinity of ately precedes the H4 event (38.23–39.35 ± 0.62 ka cal the Ponor-Plopa system. It must also have been a reason- BP; Genty et al., 2010). The fact that the stalagmite grew ably favorable environment for the exploitation by early continuously until ~35 ka cal BP shows, again, that cli- modern human foraging populations, given their pres- matic conditions were not severe enough to prevent soil ence at least in the immediate vicinity of the Peş tera cu formation and water infi ltration even during cold H4 Oase. times. Unfortunately, the low uranium content and the The assemblage of fossil remains, together with the microstructure of the stalagmite PPL6b are not suitable hydraulic regime required for the formation of Level 2, for a high-resolution and well-dated isotope profi le, so a suggests that the Plopa Plateau must have been, during direct assessment of climatic conditions during this time the ~60–47 ka cal BP time interval, a rather fl at karst sur- could not be done. face where snowmelt or torrential water accumulated at

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the bottom of dolines and ponors, to possibly form tem- sedimentary within the Ponor-Plopa karst system and porary fl oodplains or even heather moors. the geological and paleontological mix of the Pe ş tera cu Between ca. 45 ka and ca. 23 ka cal BP, the δ 13 C iso- Oase. As is often the case, our abilities to correlate climatic tope record of a stalagmite from Poleva Cave (Constantin indicators with more global (or even continental) patterns et al., 2007) shows a steady increase in δ 13 C values that are compromised by the level of precision in the dating was interpreted as a reduction in soil biological activity techniques available for the relevant time period (in this toward the end of MIS 3 and toward the LGM. At the case mostly MIS 5 to 3) and the diffi culties in correlating Pe ş tera cu Oase, this gradual cooling and possible change depositional events or periods with specifi c time hori- in vegetation type corresponds with the presence of ibex, zons. Yet, despite these limitations, what emerges is an whose typical habitat include a more rocky terrain with, overall pattern of a relatively moderate at least, scrubs and grass patches. climate for this portion of the southwestern Carpathians, the Aninei Mountains, with harsher conditions settling in toward MIS 2 and during the LGM. Despite fl uctua- Conclusion tions across cooler and warmer phases, indicated by the sedimentology, the speleothem formations and the micro- The complex geological history of the Ponor-Plopa system mammals, the regional MIS 3 appears to have been a rel- and variable use of the Pe ştera cu Oase and its environs atively rich environment for larger mammals, especially by various mammalian species presents a multidimen- cave bears and , and probably those that subsisted in sional puzzle for sorting out the paleoenvironmental part on one or both of these large mammals, the wolves conditions during the accumulation of the deposits, both and the .

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Part Seven References

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