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Quaternary Science Reviews 30 (2011) 481e487

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Quaternary Science Reviews

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How cold was it for Neanderthals moving to Central Europe duringwarm phases of the last glaciation?

Grzegorz Skrzypek a,*, Andrzej Wi�sniewski b, Pauline F. Grierson a

aWest Australian Biogeochemistry Centre, School of Plant Biology M090, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australiab Institute of Archaeology, The University of Wrocław, Szewska Str. 48, 50e139 Wrocław, Poland

a r t i c l e i n f o

Article history:Received 13 August 2010Received in revised form19 December 2010Accepted 20 December 2010Available online 26 January 2011

Keywords:PleistoceneNeanderthalsStable isotopeOxygenTeethPhosphateTemperature

* Corresponding author. Tel.: þ61 8 6488 4584; faxE-mail addresses: gskrzypek@yahoo.com, gr

(G. Skrzypek).

0277-3791/$ e see front matter � 2010 Elsevier Ltd.doi:10.1016/j.quascirev.2010.12.018

a b s t r a c t

Precise estimates of mean annual temperature (MAT) for when Neanderthals occupied Central Europeare critical for understanding the role that climatic and associated environmental factors played inNeanderthal migrations and in their ultimate extinction. Neanderthals were continuously present in therelatively warm regions of southern and Western Europe in the Pleistocene but only temporarily settledCentral Europe (CE), presumably because of its colder and less hospitable climate. Here, we presenta new approach for more spatially and temporally accurate estimation of palaeotemperatures based onthe stable oxygen isotope composition of phosphates extracted from animal teeth found at sites linkeddirectly to concurrent Neanderthal occupation. We provide evidence that Neanderthals migrated alongthe Odra Valley of CE during warmer periods throughout the Upper Pleistocene. The MATs during thesemigrations were about 6.8 �C for the warm phase of Oxygen Isotope Stage OIS 5aed (prior to the OIS4cold event) at w115e74,000 yr BP and about 6.3 �C during the early OIS 3 warm phase w59e41,000 yrBP. Our results show that temperatures during these phases peaked 2e4 �C above longer term estimatesfrom ice cores and pollen records. We argue that our approach can provide valuable insights intoevaluating the role of climate in human migration patterns in the Pleistocene.

� 2010 Elsevier Ltd. All rights reserved.

1. Introduction

While there is evidence from southern and western Europe thatconfirms a widespread distribution of the Neanderthal populationduring theearlyandmiddleWeichselianglaciations (w115e30,000yrBP), the presence of Neanderthals in Central Europe (CE)was sporadicand concentrated within relatively small areas (Harvati, 2007;Jiménez-Espejo et al., 2007). In all probability, the Alps and Carpa-thian Massifs were crossed relatively rarely and only during thewarmest phases of theWeichselian (van Andel et al., 2003). The OdraValley is considered to have been the major migration pathway forhominids from the South to the Central European highlands andlowlands through the Sudetes and CarpathianMountains, and severalarchaeological sites providing strong evidence of hunting andpresumable scavenging occur all along the Odra Valley (Wi�sniewskiet al., 2009, Wi�sniewski 2005). While the Neanderthals are consid-ered a robust species, accurate estimates of MAT for areas suchas Lower Silesia, hitherto unknown, are critical for assessing the

: þ61 8 64887925.zegorz.skrzypek@uwa.edu.au

All rights reserved.

expansion range of Neanderthals into CE, and thus how warm theclimate had to be to attract their migration (Tzedakis et al., 2007).

The locations and character of early and middle WeichselianNeanderthal sites north of the Sudetes and Carpathian Mountainsindicate clearly that these premodern humans likely slowlymigrated over generations to the north, when climate conditionsbecome more favorable (Soffer, 2009). Subsequently, the Neander-thals’ northern reach gradually contracted when the climate againbecame colder, partially due tomigration back to the south butmostlikely primarily due to local extinction (Hublin and Roebroeks,2009). This scenario is confirmed by archeological evidence froma few sites within the Carpathian and Czech Massifs that containNeanderthal remains (Rink et al., 1996; Valde-Nowak, 2009). Northof these massifs, Neanderthal sites preceding the First Pleniglacial(OIS 4) in Central Europe are represented by several locations withevidence of hunting and other activities in rock shelters, cavesand open-air sites: 1) Zwole�n site, Central Poland, OIS 5a-OIS 4(Schild, 2005); 2) Königsaue site, AeC, SaxonyeAnhalt, OIS 5a(Mania, 1999); 3) Salzgitter-Lebenstedt site, Lower Saxony, OIS 5a(Gaudzinski and Roebroeks, 2000); 4) Raj Cave, layer 6, CentralPoland, OIS 5a (Patou-Mathis, 2004) and 5) Hallera Avenue,Wroc1aw, SW Poland, OIS 5a-d (Wi�sniewski et al., 2009). Neander-thal exploitationof this part of CE is also confirmed for thebeginning

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of OIS 3, but at a few locations only: 1) Lichtenberg, Lower Saxony(Veil et al., 1994); 2) Ksiecia Józefa and Piekary IIa in Kraków (Ziebaet al., 2008) and again 3) Hallera Avenue, Wroc1aw, SW Poland(Wi�sniewski et al., 2009) (Fig. 1).

The Hallera Avenue site is one of the largest archaeologicalexcavation sites in CE. Moreover, the Hallera Avenue site providesunique evidence of Neanderthal occupation during both warmclimatic episodes, OIS 3 and OIS 5aed (Wi�sniewski et al., 2009). Theremains in this locality were gathered by Palaeolithic huntersduring their occupation and thus present an excellent geochemicalarchive for estimation of MAT. In this study we have analyzed thestable oxygen isotope composition of phosphates (d18OP) fromwell-preserved samples of animal teeth, including bovids, horses,mammoths and rhinoceros. By analysing these “dining scraps” ofPalaeolithic hunters, we can estimate temperatures directly asso-ciated with the time of their occupation during warm phases of theearly and middle Pleistocene.

2. Materials and methods

2.1. Location and geology

Bone and teeth samples were collected from Hallera Avenue(Wroc1aw/Poland), one of the largest open-air archaeologicalexcavation sites in Central Europe, with a total area >800 m2 anddepth of 2e4 m (Wi�sniewski et al., 2009; Wi�sniewski 2005). The

Fig. 1. Location of the studied site at Hallera Av. in Wroc1aw (SW Poland) and other Neandertin Czech Massif (Valde-Nowak, 2009). 3) Zwole�n site, Central Poland (Schild, 2005); 4) KöSaxony (Gaudzinski and Roebroeks, 2000); 6) Raj Cave, layer 6, Central Poland (Patou-Mathisin Kraków (Zieba et al., 2008). The map based on ASTER GDEM, open access product of ME

Hallera Avenue site is a palimpsest (a multifunctional site used forseveral seasons) situated on the outskirts of the Wroc1aw IceMarginal Valley and the Wroc1aw Plain, on a small elevation raisedca. 12e15 m above the present water level of the Odra River (51�N17�E, altitude 124 m a.s.l.) (Fig. 1). The riverine deposits buildingthis elevation consist of four major lithological complexes: (A)a lower horizon of stone boulder pavement derived during erosionof moraine till; (B) a middle horizon of alluvial gravelesand sedi-ments deposited by the braided river; and (C) an upper horizon ofsands and gravels accumulated by the ephemeral, low-energybraided river; (D) fine grained slope sediments belonging to theinfillings of thermo-carst palaeolakes (Wi�sniewski et al., 2009). Theartefacts and bones from the older phase of the settlement occurredon the stone boulder pavement surface (complex A) and partially inthe alluvial sediments of complex B, while the artefacts and bonesof the younger phase of settlement were found within the lowerpart of sediments of complex C (Figs. 2 and 3).

2.2. Dating and stable isotope analyses

Nine samples were collected for Optically Stimulated Lumines-cence dating (OSL): five from horizon B, two from horizon C andtwo fromhorizon D (Fig. 2). Each samplewas sieved out and treatedaccording to the protocol described by Murray and Wintle (2000).The OSL analyses were performed using a Daybreak 2200 OSLreader equipped with combined blue (470�5 nm) and infrared

hal sites in the region: 1) Ob1azowa Cave in Carpathian (Rink et al., 1996); 2) K�ulna Cavenigsaue site, AeC, SaxonyeAnhalt (Mania, 1999); 5) Salzgitter-Lebenstedt site, Lower, 2004); 7) Lichtenberg, Lower Saxony (Veil et al., 1994); 8) Ksiecia Józefa and Piekary IIaTI and NASA.

Fig. 2. Sediment succession at the Hallera Ave. archaeological site in Wroc1aw (SW Poland): schematic sequence of sediments and results of OSL dating.

Fig. 3. Selected samples from complex AB: bovid skull with horns within the context of river sediments (A) scale 40 cm; on insert: the skeleton of a bovid with marked foundelements (gray and black (B)). Examples of retouched tools (side scrapers) made of erratic flint; scale 3 cm (C), photo A.W.

G. Skrzypek et al. / Quaternary Science Reviews 30 (2011) 481e487 483

G. Skrzypek et al. / Quaternary Science Reviews 30 (2011) 481e487484

(880 � 80 nm) LED OSL modules; luminescence emission wasdetected by an EMI 9235QA photomultiplier. The uncertainty of theOSL dating is based on ten replicates of each sample (Fig. 2). All OSLanalyses were undertaken in the Department of Radioisotopes,Silesian University of Technology (Gliwice, Poland).

In total, 27 samples of teeth and bones were selected for stableisotope analysis. Only well-preserved samples, clearly associatedwith either horizon AB or C, were chosen (the list of samples inAppendixS1/Tab.A). Phosphateswere extracted asAg3PO4 followingthe procedure of Stephan (2000)with a buffermodified according toWiedemann-Bidlack et al., (2008). The d18OP values of Ag3PO4 wereanalyzed using a Temperature Conversion Elemental Analyzer(TC/EA) coupled to a Thermo Finnigan DeltaPlus XLwith an externalreproducibly better than 0.3&. National Institute of Standards andTechnology (USA) standard, NBS 120c Florida phosphate rock,treated in the same way as the samples was used as the analysisquality controls, confirming high precision of stable isotope analyses(Appendix S1/Tab. B). Normalization to the VSMOW internationalscale was based on four international standards provided by Inter-national Atomic Energy Agency (IAEA) following multi-pointnormalization (Paul et al., 2007).

In all samples, the d18OC of carbonate was analyzed after pre-treatment, using a Thermo Finnigan Gasbench II coupled withThermo Finnigan DeltaPlus XL mass spectrometer, followinga procedure adapted from Paul and Skrzypek (2006). Analyzedcarbon and oxygen isotopic compositions were normalized to theinternational VPDB scale using multi-point normalization (Paulet al., 2007) and three IAEA standards. The external reproducibilitywas better than 0.1& for carbon, and 0.2& for oxygen isotopecompositions. The d18OC in VPDB was recalculated to the VSMOWscale using the equation: d18OVSMOW ¼ 1.03091 � d18OVPDB þ 30.91.All stable isotopes analyses were performed at the West AustralianBiogeochemistry Centre, The University of Western Australia.

2.3. Calculations

We first verified the preservation of the stable oxygen isotopecomposition in bioapatite based on the correlation of d18OC incarbonates to the d18OP of phosphates, following the procedure ofArppe and Karhu (2010) and a recent equilibration equation byLécuyer et al. (2010): d18OC ¼ 1.035(�0.017) � d18OP þ 8.33(�0.30).We found the d18O of bone samples, in contrast to those of teeth,were not well-preserved. The difference in d18OP between a pair ofsamples originating from the same individual was 0.8& for skulland tooth (sample no 20 and 24) or 1.4& for jaw and tooth (sampleno 25 and 26). Consequently, we used tooth samples only withconfirmed well-preserved bioapatite, and therefore samples no 1,17, 20, 24, 25, 26, 27 and 32were excluded from further calculationsas they were not well-preserved (for verification procedures seeAppendix S1/Tab. C).

In the next step, we calculated the d18OW value of environmentalwater based on the d18OP of phosphates of unaltered teeth samples.We used published relationships between d18OP and d18OW forindividual studied animal species, which are typically presented asd18OP ¼ a � d18OW þ b (Appendix S1/Tab. D). Where necessary, werecalculated to the form d18OW ¼ a � d18OP þ b, based on theoriginal d18OW and d18Op data presented in D’Angela and Longinelli(1990), Ayliffe et al. (1992) and Chillón et al. (1994).

We then derived equations capturing varying spatial resolutionto describe the current relationship between d18OW and Tair forseveral locations across Europe. We used data from the GlobalNetwork of Isotope in Precipitation (GNIP/WISER IAEA database,2006) from: 1) the closest GNIP station to the study area, Kraków,which was about 300 km east (Appendix S1/Tab. F); 2) ten locationsat similar elevations with GNIP stations with the long observation

records that were within w500e1000 km to the east, west andsouth of study area, including Kraków (Appendix S2/Tab. D); and 3)all GNIP stations for continental Europe (Appendix S1/Tab. A,B).Finally, we estimated the mean annual temperature (MAT) forHallera Avenue using these derived equations and the d18OW thatwe calculated based on tooth d18OP.

3. Results and discussion

The evidence integrated from Optically Stimulated Lumines-cence dating (Fig. 2) and a variety of independent lithological,petrographical and sedimentological records, as well as correlationwith other profiles from excavation sites (Wi�sniewski et al., 2009),confirms that the horizons contained artefacts and animal remainsare linked with two warm phases of the Pleistocene. The lower ABhorizon fromtheHalleraAvenue site correlateswithOxygen IsotopeStage (OIS) 5aed (w115e74,000 yr BP) of the BrørupeOdderadeperiod while the upper C horizon with the beginning of OIS3(w59,000 yr BP) of the Oerel period (Fig. 2).

The d18OP values of tooth samples of bovids, horses, mammothsand rhinoceros collected from both horizons varied between 12.4and 15.0& (Appendix S1/Tab. D). This variation is attributed todifferent environmental conditions influencing the stable isotopecomposition ofwater consumed by these animals, but is also affectedbydifferences inmetabolic rates amongmammals (e.g. Bernard et al.,2009; Ayliffe et al.,1992; Chillón et al.,1994; D’Angela and Longinelli,1990). After accounting formetabolic differences among species, thed18OW was �10.7 � 0.8& during the older settlement phase duringOxygen Isotope Stage OIS5a (horizon AB), and �11.0 � 0.9& duringthe younger settlement beginning in OIS3 (horizon C). The observedrange of d18OW values estimated from the d18O of phosphates fromthe Hallera Avenue site is relatively narrow, despite being calculatedbased on several species from two horizons (AB or C). This findingsuggests that the water consumed by these animals had a similarisotope composition. The accumulation of each of these assemblageslikelyoccurredover a short time spanduring twoepisodes (AB andC)andat a scale of yearsedecades, rather thanhundredsof years. Today,d18O values around�11& are characteristic of precipitation falling inSweden and Norway, but not Central Europe. Further, the d18OWvalue of modern precipitation at the sampling site is �9.1 � 0.6&,which equates to a mean annual temperature of 9.1 �C (51�N 17�E,altitude 250 m a.s.l.; see red arrows on Fig. 4).

Several factors have complex influences on the oxygen isotopecompositionofwater, including air temperature, amount effects anddistance from the sea (Ró _zanski et al.,1993). Therefore,most reliableestimates of Tair are calculated from the direct relationship betweenTair and d18OWat a particular study location and based on long-termobservations (Fricke and O’Neil, 1999). Based on the equationderived for the Kraków GNIP station (Tair ¼ 1.80�d18OWþ26.02,Appendix S1/Tab. F) the estimatedMATwas 6.8� 1.5 �C for the OIS5settlement phase (AB) and 6.3�1.6 �C for theOIS3 settlement phase(C) (Fig. 4 and Appendix S1/Tab. E). These estimates assume thatatmospheric circulation patterns during the Palaeolithic periodwere similar to those of today and that the positions of major bariccentres governing the distributions of rains in Europehave persistedin similar positions. However, variations in atmospheric circulationover time due to expansion or shrinkage of the Scandinavian ice capmay have influenced the total amount of precipitation at the studiedlocation aswell as the ratio between rains originating fromdifferentdirections (Hubberten et al., 2004).

In general, two major air masses govern climate in this part ofCE: the western and north-western Atlantic circulation and theeastern continental circulation (Przybylak et al., 2010). In order toaccount for any possible changes in atmospheric circulations, wedesigned a cross-verificationprocedure to assess the accuracy of our

Fig. 4. Calculation scheme for annual mean air temperature based on the stable oxygen isotope composition of animal teeth from the AB complex (OIS5). The example sample withamean d18OP value of tooth phosphates of 13.8& (orange line) and standard deviation 0.8& (gray shadow) is indicated on the left axis. The d18OWof environmental water�10.7� 0.8&(x-axis) was calculated based on published equations for horse, bison, elephant and deer (D’Angela and Longinelli,1990; Bernard et al., 2009; Ayliffe et al., 1992; Chillón et al.,1994). Airtemperature of 6.8� 1.5 �C (right y-axis) was calculated based on the equation describing the relationship between Tair and d18OW for Kraków (blue points and line), following Tütkenet al. (2007); graydots indicate data for the entire continent of Europe. The currentMAT for the sampling site (9.1 �C) and d18OW (�9.1&) are shown in red. See theOnline SupplementaryInformation for data set and calculations details. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article).

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estimates. We randomly selected a subset of additional nine GNIPstationswith long d18OWand Tair records, locatedw500e1000 km tothe east, west and south of our study site, which are also at similaraltitude to the Hallera Avenue site (Appendix S1/Tab. C, D). If, duringthe Palaeolithic period, the climate was more continental, the rela-tionship between Tair and d18OW in our archaeological site should bemore similar to that observed currently in, for example, Minsk(Belarus), or even farther to the east: Moscow, or St. Petersburg(Russia). On the other hand, if the fraction of rains coming from theNorthernAtlanticwas higher, this relationshipwould be expected tomore closely resemble that now observed in Central Germany (e.g.,Weil am Rhein). If the North Atlantic circulation influence was evenstronger, then the relationship would be closer to that of WesternGermany (e.g., Stuttgart, Karlsruhe). Consequently, we also usedGNIP stations fromZagreb/Croatia and Ljubljana/Slovenia in order tosimulate higher temperatures and more rain originating from theSouth. However, we found that temperatures for the upper Pleisto-cene at the Hallera Avenue site, estimated from equations derivedfrom different GNIP stations, did not vary from temperatures esti-mated based on for the data from the GNIP station at Kraków. Themean MAT based on equations for these nine locations variedaround 6.9 � 1.6 �C for horizon AB and 6.4 � 1.6 �C for horizon C(Appendix S1/Tab. E). As further confirmation of these results, wegenerated a general equation (Tair ¼ 1.41 � d18OW þ 21.63) for theentire continent of Europe (using >13 k records from GNIP/WISERdatabase 2006). This equation also gave similar results to equationsbased on GNIP form Kraków. However, the continental-scale esti-mates of temperature were slightly less variable; 6.6 � 1.1 �C forhorizon AB and 6.2� 1.3 �C for horizon C (Appendix S1/Tab. A). Thisconsistency in estimated temperatures confirms that the fraction ofprecipitation coming from different directions over a year likely had

minor influence on the stable isotope composition of precipitation,and Tair was the major factor governing the d18OW value. Conse-quently, our original approach based on the data from the neareststation at Kraków provided a reliable estimate of temperature forthe Hallera Avenue site.

Long-term d18OW records from Greenland ice cores and frag-mented local records from various sources including pollen, sedi-ments and fossilised insect provide other approaches for estimatingtemperatures during the interglacial periods. The d18O chronologiesfrom Greenland ice encompass >150,000 yr and are continuousand highly accurate at the place of ice accumulation (Barbante et al.,2006; Dansgaard et al., 1993). However, while reliable for detectingglobal episodes of significant warming and cooling (Barbante et al.,2006), both the resolution and sensitivity of ice-core methods forpast temperature estimates at particular ice likely decrease withincreasing distance from Greenland. While chronologies based onpollen, insect and plant microfossil records provide a more localrecord of overall conditions in places of potential human presence,they are not as precise as stable isotope ice palaeothermome-try. Their resolution is low due to several limitations, includingecosystem inertia, ranges in temperature tolerance of plants, orslow or differential rates of successional changes in vegetation (e.g.,Skrzypek et al., 2009). Moreover, the uncertainty over dating ofarchaeological sites make it difficult to accurately link ice or pollenbased temperature estimates with the exact time of human occu-pation at any particular site. In contrast, reconstructed absolutepalaeotemperatures based on tooth phosphate isotope compositionreflect only the period when a particular animal was living andhunted (D’Angela and Longinelli, 1990). Consequently, the methodpresented here is especially attractive for tracing temperatureduring humanmigration and provides temperatures when humans

G. Skrzypek et al. / Quaternary Science Reviews 30 (2011) 481e487486

were present, regardless of the accuracy of dating of archaeologicalsites. All that is needed are well-preserved animal remains that arelinked with confirmed human presence.

We estimated temperatures from teeth phosphates that are2e4 �C higher than previous estimate based on pollen analyses andvegetation patterns for both the OIS5 (Komar et al., 2009) and OIS3(Huntley and Allen, 2003). Compared to our study, van Andel et al.(2003) also estimated significantly lower temperatures for OIS3thatwerebasedon large scale-modelling fromGreenland ice. On theother hand, our temperature estimates are similar to recent esti-mates for Fennoscandia (Väliranta et al., 2009 (OIS 5c); Engels et al.,2008 (OSI3)). Together, our data strongly suggest that climate in thewarmphases of the Pleistocenewasmilder thanpreviously thought.

4. Conclusion

In summary, while many other factors may have played roles inaccelerating or restricting the northern migration of Neanderthalsin Europe, including snow cover, winter versus summer tempera-ture, and changes in vegetation composition after cold events, theMAT likely played one of the most important roles. Our studyconfirms that MAT were w6.8 �C when Neanderthals were movinginto Central Europe prior to the OIS4 cold event and that Nean-derthals probably returned to the Odra Valley on the beginning ofOIS3, when the MAT was w6.3 �C. Put in a modern context, thesepalaeotemperatures are higher than the current MAT in Stockholm/Sweden (5.8 �C) or Riga/Latvia (6.0 �C).

The most important limitation to accurate assessment of climateconditions during Neanderthal migration into Central Europe andelsewhere is not uncertainties in the developed climatic models, butinaccuracy in the chronological linking of a climate model toaparticulararchaeological site.Ourapproachovercomes thisproblemby providing direct estimation ofmean air temperature at the precisetime of occupation by Palaeolithic hunters, regardless of the uncer-tainty in dating.

Acknowledgments

We thank J.R. Dodson (ANSTO), M. McCulloch (UWA) andL. Simmons (UWA) for valuable comments on an earlier version ofthe manuscript. We are also indebted to Anna Miko1ajczyk for herhelp in themap preparation. Special thanks to Dr. AgnieszkaWójcikfor introducing G.S. to A.W. whichmade this study possible. Helpfulcomments from an anonymous reviewer are also acknowledged.G.S. is supported by a Research Fellowship from the John de LaeterCentre of Mass Spectrometry, Western Australia.

Appendix. Supplementary material

Supplementary material related to this article can be foundonline at doi:10.1016/j.quascirev.2010.12.018.

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