and South Asia (Fig. 3A and fig. S14). The detectionof an hpAsia2 strain in the Iceman’s stomach israther surprising because despite intensive sampling,only three hpAsia2 strains have ever been detectedin modern Europeans. Stomachs of modern Eu-ropeans are predominantly colonized by recom-binant hpEurope strains. Further analysis withthe STRUCTURE linkagemodel (23), used to detectancestral structure from admixture linkage dis-equilibrium, revealed that the ancient H. pyloristrain contained only 6.5% [95% probability in-tervals (PI) 1.5 to 13.5%] of the northeast African(AE2) ancestral component of hpEurope (Fig. 3B).Among European strains, this low proportion ofAE2 is distinct andhas thus far only been observedin hpAsia2 strains from India and Southeast Asia.In contrast, the three European hpAsia2 strains(Fig. 3B, black arrows) contained considerablyhigher AE2 ancestries than that of the H. pyloristrain of the Iceman (Finland 13.0%, PI 5.9 to 21.7;Estonia 13.2%, PI 6.2 to 22.3; and the Netherlands20.8%, PI 11.5 to 31.7), although 95% probabilityintervals did overlap. A principal componentanalysis (PCA) of the MLST sequences of thehpAsia2, hpEurope, and hpNEAfrica populationsrevealeda continuumalongPC1 that correlateswiththe proportion of AE2 ancestry versus AE1 ancestryof the isolates (Fig. 3C). The Iceman’s ancient H.pylori was separated from modern hpEuropestrains, and its position along PC1 was close tomodern hpAsia2 strains from India, reflecting itsalmost pure AE1 and very low AE2 ancestry.Comparative whole-genome analyses (neighbor

joining, STRUCTURE, and principle componentanalyses) with publicly available genomes (n = 45)confirmed the MLST result by showing that theIceman’s ancient H. pylori genome has highestsimilarity to three hpAsia2 genomes from India(figs. S15 to S17). Although the Iceman’sH. pyloristrain appears genetically similar to the extantstrains from northern India, slight differenceswere observed along PC2 in bothMLST (Fig. 3C)and genome PCAs (fig. S17) and in the neighborjoining tree (fig. S15). To further studygenomic-scaleintrogression, we performed a high-resolutionanalysis of ancestralmotifs using fineSTRUCTURE(24). The resulting linked co-ancestrymatrix (Fig. 4)showed that the ancient H. pylori genome shareshigh levels of ancestry with Indian hpAsia2 strains(Fig. 4, green boxes), but even higher co-ancestrywith most European hpEurope strains (Fig. 4, blueboxes). In contrast, the Iceman’s H. pylori shareslow ancestrywith the hpNEAfrica strain, amodernrepresentative of AE2 (Fig. 4, black box), and withEuropean strains originating from the IberianPeninsula, where the proportion of AE2 ancestryis relatively high (Fig. 4, white box) (4). Our samplesize (n = 1) does not allow further conclusionsabout the prevalence of AE1 in ancient Europe andthe course or rate of AE2 introgression. However,the ancient H. pylori strain provides the first evi-dence that AE2 was already present in CentralEurope during theCopperAge, albeit at a low level.If the Iceman H. pylori strain is representative ofits time, the low level of AE2 admixture suggeststhat most of the AE2 ancestry observed in hpEu-rope today is a result of AE2 introgression into

Europe after the Copper Age, which is later thanpreviously proposed (4, 6). Furthermore, ourco-ancestry results indicate that the Iceman’sstrain belonged to a prehistoric European branchof hpAsia2 that is different from the modernhpAsia2 population from northern India. Thehigh genetic similarity of the ancient strain tobacteria from Europe implies that much of thediversity present in Copper Age Europe is stillretained within the extant hpEurope population,despitemillennia of subsequent AE2 introgression.

REFERENCES AND NOTES

1. S. Suerbaum, C. Josenhans, Nat. Rev. Microbiol. 5, 441–452 (2007).2. P. Malfertheiner, F. K. Chan, K. E. McColl, Lancet 374,

1449–1461 (2009).3. R. M. Peek Jr., M. J. Blaser, Nat. Rev. Cancer 2, 28–37 (2002).4. D. Falush et al., Science 299, 1582–1585 (2003).5. Y. Moodley, B. Linz, Genome Dyn. 6, 62–74 (2009).6. B. Linz et al., Nature 445, 915–918 (2007).7. Y. Moodley et al., PLOS Pathog. 8, e1002693 (2012).8. Y. Moodley et al., Science 323, 527–530 (2009).9. T. Wirth et al., Proc. Natl. Acad. Sci. U.S.A. 101, 4746–4751 (2004).10. W. Müller, H. Fricke, A. N. Halliday, M. T. McCulloch,

J. A. Wartho, Science 302, 862–866 (2003).11. W. Haak et al., Nature 522, 207–211 (2015).12. A. Keller et al., Nat. Commun. 3, 698 (2012).13. I. Lazaridis et al., Nature 513, 409–413 (2014).14. P. Gostner, P. Pernter, G. Bonatti, A. Graefen, A. R. Zink,

Arch. Sci. 38, 3425–3431 (2011).15. A. F. Andersson et al., PLOS ONE 3, e2836 (2008).16. N. Segata et al., Genome Biol. 13, R42 (2012).17. S. Sawyer, J. Krause, K. Guschanski, V. Savolainen, S. Pääbo,

PLOS ONE 7, e34131 (2012).18. K. R. Jones, J. M. Whitmire, D. S. Merrell, Front. Microbiol. 1,

115 (2010).19. M. Uhlén et al., Science 347, 1260419 (2015).20. J. A. Gaddy et al., PLOS Pathog. 10, e1004450 (2014).21. S. T. Leach, H. M. Mitchell, C. L. Geczy, P. M. Sherman,

A. S. Day, Can. J. Gastroenterol. 22, 461–464 (2008).

22. J. K. Pritchard, M. Stephens, P. Donnelly, Genetics 155,945–959 (2000).

23. D. Falush, M. Stephens, J. K. Pritchard, Genetics 164,1567–1587 (2003).

24. D. J. Lawson, G. Hellenthal, S. Myers, D. Falush, PLOS Genet. 8,e1002453 (2012).

ACKNOWLEDGMENTS

We acknowledge the following funding sources: the South Tyroleangrant legge 14 (F.M., N.O.S., G.C., V.C., M.S., and A.Z.), theErnst Ludwig Ehrlich Studienwerk, dissertation completionfellowship of the University of Vienna (D.T.), the Graduate SchoolHuman Development in Landscapes and the Excellence ClusterInflammation at Interfaces (B.K. and A.N.), the European ResearchCouncil (ERC) starting grant APGREID (J.K. and A.H.), the NationalInstitutes of Health from the National Institute of GeneralMedical Sciences under grants R01 GM087221 (R.M.), S10RR027584 (R.M.), and 2P50 GM076547/Center for SystemsBiology (R.M). E. Leproust and O. Hardy are highly acknowledgedfor their help in the RNA bait design. We thank the sequencingteam of the Institute of Clinical Molecular Biology at Kiel University forsupport and expertise. We are grateful to E. Hütten for proofreading ofthe main text. We are grateful to Olympus, Italy, for providing us withequipment for endoscopy. F.M. and A.Z. conceived the investigation.F.M., B.K., D.T., R.G., J.K., A.N., Y.M., T.R., and A.Z. designedexperiments. P.M., L.E., E.E.V., M.S., F.M., and A.Z. were involved in thesampling campaign. F.M., B.K., M.R.H., J.H., U.K., and G.C. performedlaboratory work. F.M., B.K., D.T., A.H., M.R.H., N.O.S., B.L., R.L.M., R.G.,J.K., Y.M., and T.R. performed analyses. F.M. wrote the manuscriptwith contributions from B.K., D.T., A.H., M.R.H., U.K., N.O.S., V.C., B.L.,R.L.M., R.G., J.K., Y.M., A.N., T.R., and A.Z. Data are available from theEuropean Nucleotide Archive under accession no. ERP012908. Theauthors declare no competing interests.

SUPPLEMENTARY MATERIALS

www.sciencemag.org/content/351/6269/162/suppl/DC1Materials and MethodsFigs. S1 to S17Tables S1 to S13References (25–93)

15 August 2015; accepted 20 November 201510.1126/science.aad2545

PALEOCLIMATE

Reconciliation of the DevilsHole climate record withorbital forcingGina E. Moseley,1* R. Lawrence Edwards,2 Kathleen A. Wendt,1,2 Hai Cheng,2,3

Yuri Dublyansky,1 Yanbin Lu,2 Ronny Boch,1† Christoph Spötl1

The driving force behind Quaternary glacial-interglacial cycles and much associated climatechange iswidely considered to be orbital forcing. However, previous versions of the iconic DevilsHole (Nevada) subaqueous calcite record exhibit shifts to interglacial values ~10,000 yearsbefore orbitally forced ice age terminations, and interglacial durations ~10,000 years longerthan other estimates. Our measurements from Devils Hole 2 replicate virtually all aspectsof the past 204,000 years of earlier records, except for the timing during terminations,and they lower the age of the record near Termination II by ~8000 years, removing both~10,000-year anomalies.The shift to interglacial values now broadly coincides with the rise inboreal summer insolation, the marine termination, and the rise in atmospheric CO2, which isconsistent with mechanisms ultimately tied to orbital forcing.

Changes to Earth’s orbital configuration rel-ative to the Sun, known as the Milanko-vitch hypothesis, astronomical theory, ororbital forcing, have long been consideredthe leading theory for the primary mech-

anism driving Quaternary glacial-interglacialcycles (1–3) and associated climate change.The hypothesis is supported by a huge arrayof evidence from paleoclimate records acrossthe globe, which show that major shifts in

SCIENCE sciencemag.org 8 JANUARY 2016 • VOL 351 ISSUE 6269 165

RESEARCH | REPORTSon S

eptember 15, 2020

http://science.sciencem

ag.org/D

ownloaded from

climate took place throughout the Quater-nary on orbital time scales (2, 4, 5). One par-ticular seminal paleoclimate record, however,the ~500-thousand-year (ky) Devils Hole recordfrom Nevada (fig. S1) (36°25’N, 116°17’W; 719 mabove sea level), has challenged this hypoth-esis for nearly three decades (6–8). Of note isthe controversy (4, 9–11) surrounding the tim-ing of events at the end of the penultimateglacial period [Termination II (TII)]. At issueis whether TII preceded the rise in boreal in-solation, in which case TII could not result di-rectly from orbital forcing. The current DevilsHole chronology places the TII shift to inter-glacial values ~10 ky before the rise in borealsummer insolation (Fig. 1) and the duration ofthe last interglacial period to almost double itsequivalent in other records, both of which areinconsistent with a straightforward explanationin terms of orbital forcing.The controversial findings generated dis-

cussion on (i) dating accuracy (12–16) and re-solution (17, 18); (ii) the possibility that therecord represents a regional hydrological sig-nal (19–21); and (iii) phase leads and lagsbetween proxy records (7, 18, 22). Deviationfrom an orbital pacing was highlighted in theoriginal Devils Hole record (6), with a chronol-ogy based on alpha-counting 230Th and 234U/238U dating of sample DH-2. This record waslater replicated with higher-precision thermalionization mass spectrometric ages on sampleDH-11 and extended deeper in time (7, 8). Amechanism was immediately suggested where-by water-sourced 230Th incorporated duringcalcite growth could lead to artificially old ages(12, 14). However, measurements on the outersurfaces of DH-11 and DH-2, both of whichhad been submerged for an extended periodof time, showed that this process operatedonly at low efficiency, under the presumptionthat the adsorption of 230Th was irreversible(15, 16). Several years later, concordant 231Pa and230Th ages were obtained on an ~80-ky-old sub-sample of DH-11 (13), consistent with an accu-rate chronology for this portion of the record.In the following two decades, the accuracy ofthe DH-11 chronology remained unchallenged.More recently, marine cores off the west coast

of the Americas recorded changes in temperaturebefore marine d18O terminations (21). These ob-servations were interpreted in terms of changingsurface currents resulting in changing temper-ature. If Devils Hole recorded these early changesin regional temperature, the difference betweentheDevilsHole record and themarine d18O recordwould be explained.The record was then extended to the Holocene,

using samples DHC2-3 and DHC2-8 (23), both

collected from Devils Hole 2 (24), a cave similarto Devils Hole and 200 m away from it. Theportion near TI could also be explained in termsof changing surface currents in the easternPacific (21).Despite the apparent resolution, the con-

troversy has subsequently been revived. GreatBasin dripstone (fig. S1) records exhibit shifts to

interglacial values around the time of marineTII (25, 26) and TI (27) and therefore replicateneither of the early shifts documented at DevilsHole (7, 8, 23). Thus, the controversy currentlycenters onmajor chronological discrepancies be-tween the subaerially formed Great Basin drip-stone and subaqueously formed Devils Holerecords.

166 8 JANUARY 2016 • VOL 351 ISSUE 6269 sciencemag.org SCIENCE

1Institute of Geology, University of Innsbruck, Innrain 52, 6020Innsbruck, Austria. 2Department of Earth Sciences, University ofMinnesota, 310 Pillsbury Drive SE, Minneapolis, MN 55455,USA. 3Institute of Global Environmental Change, Xi'an JiaotongUniversity, Xi'an 710049, China.*Corresponding author. E-mail: gina.moseley@uibk.ac.at†Present address: Institute of Applied Geosciences, Graz Universityof Technology, Rechbauerstraße 12, 8010 Graz, Austria.

Fig. 1. Comparison of Devils Hole records with other paleoclimate archives. (A) 230Th ageswith 2s uncertainties for DHC2-8 (light blue) (23), DHC2-3 (light-mid blue) (23), DH-11 (lightgreen) (7, 8, 23), DH-2 (dark green) (this study), DH2-E (mid-dark blue) (this study), and DH2-D(dark blue) (this study). VPBD, Vienna Pee Dee belemnite standard. (B) d18O records for samples in(A) [same color codes and references as in (A)] and 65°N July insolation (dark gray) (32). (C) d13Crecords for DH-2, DH2-E, and DH2-D [same color codes as in (A)]. (D) Growth rate of samples fromthis study with [same color codes as in (A)]. (E) Stacked global benthic d18O records (5). (F) Ant-arctica CO2 composite curve (38–40). (G) 230Th ages on mammillary calcite with 2s uncertainty(24). Pre-folia, red; post-folia, black; pre-growth hiatus, blue. Arrows indicate the direction of water-table change.

RESEARCH | REPORTSon S

eptember 15, 2020

http://science.sciencem

ag.org/D

ownloaded from

To address this controversy, we tested the re-producibility of the Devils Hole record by re-analyzingDH-2 (fig. S2) (6) fromDevilsHole properand by analysis of four newly drilled cores (fig. S2)from Devils Hole 2. The previously studied sam-ples were all collected at substantial depth belowthe current water table (DH-2 = –21 m; DH-11 =–30 m; DHC2-3 = –25 m; DHC2-8 = –60 m),whereas the new cores from Devils Hole 2 werecollected from above the water table (DH2-D =+2.1m;DH2-E = +0.8m;DH2-G =+4.5m;DH2-J =+5.5 m) (fig. S3).Considering our data, the results confirm

the overwhelming majority of the features ofthe original studies (6–8, 23), verifying the orig-inal analytical results and the reliability withwhich the groundwater system and the calciteprecipitated from it record climate. Portions ofthe record have now been replicated four timeswith similar results. The general character andrange of d18O variations, and the U isotopiccompositions and concentration, are all sim-ilar to those in the original reports (6, 8, 23)(Fig. 1 and table S1). Further, large portions ofour chronology replicate the original chronol-ogies (8, 23). Between 28 to 112 thousand yearsago (ka), DH2-D replicates at significantly higherresolution the timing of the previous chronol-ogy (8, 23) (Fig. 1), while further adding supportfor the timing of the isotopic maximum thatoccurs at about the time of marine isotope stage5a, with an age of 82.5 ± 0.7 ka, replicating theoriginal 230Th age of 80.6 ± 2.5 ka (7, 8) and the231Pa age 82.5 ± 2.8 ka (13).

Considering our data sample by sample, start-ing with our deepest sample (DH-2: –21 m), wefind that within quoted uncertainties, the newlyanalyzed portion of DH-2 replicates both theoriginal DH-2 (6) and DH-11 records (7, 8), thusverifying the original analytical results, includingmeasurement of the U and Th isotopes used incalculating ages. The first hint of an age dis-crepancy occurs at about the time of TII, becausethe shift in d18O values in our DH-2 record isnominally later than the shift in DH-11 (7, 8)(Figs. 1 and 2).Moreover, comparison of our two highest-

elevation d18O records (DH2-D: +2.1; DH2-E:+0.8 m) to those from deeper cores yields largediscrepancies in chronology, well outside ofanalytical error (Figs. 1 and 3 and fig. S4). Thesediscrepancies are largely confined to and clearlyobserved at times that correspond to each ofthe last two terminations, but are best resolvedduring TII because calcite deposition rates arehigher than during TI (Fig. 1 and fig. S5). The230Th chronology of the shift to interglacial val-ues is systematically younger for cores col-lected at higher elevation (Figs. 1 and 2), withthe shallowest core recording a time of 132.2 ±1.5 ka (24) for the midpoint of the shift to in-terglacial values, as compared with a value of142 ± 3 ka (18) for the deepest sample forwhich TII data have been reported. The shifttoward glacial values at the end of the lastinterglacial is similar in all records (Figs. 1 and2). Therefore, the duration of the last intergla-cial d18O peak as recorded in the samples also

shifts systematically with sample elevation,with a last interglacial duration of 16.1 ky (mea-sured from the midpoint of rise to the mid-point of fall) recorded in DH2-D as comparedto a DH-11 duration of ~22 ky (table S2) (7, 8, 18).Considering just the portion of the peakthat records the highest d18O values, DH2-Drecords a 6-ky duration (from 127 to 121 ka),which is close to half the duration recorded inDH-11 (7, 8, 18). Thus, the shallowest core recordsthe latest shift to interglacial conditions andthe shortest duration for the last interglaciald18O peak (Figs. 1 and 2).The explanation for the relationship between

apparent age and sample elevation probably liesin 230Th incorporated in the growing calcite fromthewater, as raised previously (12, 14) and arguedto be negligible (15, 16). The discussion at thattime assumed irreversible adsorption of 230Thonto the walls (12, 14–16). In light of our data andconsidering advanced understanding in the dis-tribution of 230Th in seawater (28, 29), we con-sider irreversible exchange unlikely. Indeed,vertical profiles through the ocean water columnshow that both particulate and dissolved 230Thincrease in concentration with depth, indicatingreversible exchange of 230Th between water andparticulate matter (28). Furthermore, sharp in-creases in dissolved 230Th in the upper 200 mof the ocean have been demonstrated (29). The

SCIENCE sciencemag.org 8 JANUARY 2016 • VOL 351 ISSUE 6269 167

Fig. 2.Timing of TII and the last interglacial in speleothem records. (A) DH-2 (dark green) (this study)and DH-11 (light green) (7, 8, 23). (B) DH2-D (dark blue) (this study) and DH2-E (mid-blue) (this study).(C) Great Basin composite dripstone record (“Leviathan chronology,” pink) (30); Lehman Cave (red) (26)(dark red) (25); and 65°N July insolation (gray) (32).Themidpoints of transitions are indicatedwith x’s andvertical dashed lines.

Fig. 3. TI in Devils Hole and Great Basin paleo-climate records. (A) 230Th ages with 2s un-certainties: DHC2-8 (light blue) (23), DHC2-3(light-mid blue) (23), DH2-E (mid blue) (thisstudy), and DH2-D (dark blue) (this study). (B)d18O records for samples in (A) with same colorcodes and 65°N July insolation (dark gray) (32).(C) d18O Great Basin composite dripstone record(“Leviathan chronology”) (30) and 65°N Julyinsolation (dark gray) (32).

RESEARCH | REPORTSon S

eptember 15, 2020

http://science.sciencem

ag.org/D

ownloaded from

systematic increase of 230Th ages with depth(Figs. 1, 2, and 4) in Devils Hole may thus be theresult of increasing concentrations of 230Thdown the water column. The strong correla-tion (coefficient of determination, R2 = 0.97)between the age of the TII midpoint in eachsample versus depth (Fig. 4) (24) is consistentwith a reversible exchange–generated increaseof 230Th with depth in the TII Devils Hole watercolumn.The appearance of age anomalies during ter-

minations may result from several factors. First,the water table was higher than at presentduring both TI and TII, before it progressivelydeclined during the respective following inter-glacials (Fig. 1), in agreement with progressivelydrier conditions during interglacials (30, 31). Onewould therefore expect that for a given elevation,the water 230Th concentration would have in-creased during the pluvial periods associatedwithterminations and then decreased over the courseof the proceeding interglacial. Second, lower dep-ositional rates during times of the anomalies(Fig. 1 and fig. S5) may play a role in the in-corporation of excess 230Th onto the calcite (12).Finally, wetter conditions associated with higherwater tables could be associated with increasedgroundwater flow rates and 230Th fluxes into theopen fractures.Regardless of the details of the mechanism, if

progressively higher levels of 230Th at depth areresponsible for the anomalies, then the shal-lowest core, which records the youngest ages, hasthe most accurate chronology. Thus, we con-sider the chronology of the DH2-D d18O recordto be the closest to the true age. Because theDH2-D record is in close agreement with thetiming of d18O shifts in Great Basin dripstonerecords [Figs. 2 and 3; particularly consideringan up to 2-ky groundwater transit time in theDevils Hole system (23)], we conclude that DH2-Dhas negligible initial 230Th and provides an ac-curate chronology.

In contrast to DH-11 (7, 8), the timing of TIIand the duration of the last interglacial d18Opeak (either midpoint to midpoint or peak valueduration) in the DH2-D record are all consistentwith orbital forcing. The DH2-D d18O shift tointerglacial values is broadly coincident with therise in boreal summer insolation (32), the rise inatmospheric CO2 (33), the marine termination(34), and the TII Weak Monsoon Interval (4).The mechanism for 18O enrichment in precipita-tion could be increasing temperature (6–8, 21, 23),in which case the temperature rise could becaused by the rise in global atmospheric CO2,which has been closely tied to orbital forcing(4). However, it is also likely that changes inthe proportion of summer (high-d18O today) towinter (low-d18O today) meteoric precipitation(i.e., changes in seasonality) played a role. If so,the low d18O values before the glacial-interglacialtransition could be partly explained by a largerproportion of low-d18O cool season rainfall as-sociated with pluvial conditions. The pluvial con-ditions in turn, could be caused by the splittingof the jet stream (35) around the LaurentideIce Sheet to the north. In this case, the propor-tion of (low-d18O) winter season moisture woulddiminish as the ice sheet melted, as a result ofincreased boreal summer insolation. The risein insolation could also trigger (high-d18O sum-mer) North American monsoon-like rainfallin the region, increasing the mean annual d18Oof precipitation. We conclude that some com-bination of these orbitally based processes con-tributed to the observed shift in d18O. Temperaturechanges of Devils Hole water have been small(36), so that the temperature-dependent calcite-water fractionation would have had little effecton calcite d18O. Also, the rainfall seasonalitymechanisms do not directly relate to regionaltemperature and could plausibly explain theabsence of a Devils Hole “lead” that is analogousto the lead in temperature observed in the ma-rine cores (21).Finally, in our (DH2-D) chronology, the prom-

inent low in d13C at the end of TII is shifted toyounger values by about 7 ky (Fig. 1) relative tothe DH-2 and DH-11 (37) records, thus coincid-ing with the boreal summer insolation peak.Because this could be a time of regional warmthand relatively high warm-season rainfall, ourtiming supports the idea (37) that such d13Clows are caused by more extensive vegetationcover and productivity in the source region forthe aquifer.Our chronologies from Devils Hole have

demonstrated that there is a systematic offsetin the age of calcite deposited at increasingdepths in these open fractures across glacialterminations, thus helping to solve one of thegreat paleoclimate enigmas of the past threedecades.

REFERENCES AND NOTES

1. C. Emiliani, J. Geol. 63, 538–578 (1955).2. J. D. Hays, J. Imbrie, N. J. Shackleton, Science 194, 1121–1132

(1976).3. J. Imbrie, J. Geol. Soc. London 142, 417–432 (1985).4. H. Cheng et al., Science 326, 248–252 (2009).

5. I. E. Lisiecki, M. E. Raymo, Paleoceanography 20, 1–17(2005).

6. I. J. Winograd, T. B. Coplen, B. J. Szabo, A. C. Riggs, Science242, 1275–1280 (1988).

7. I. J. Winograd et al., Science 258, 255–260 (1992).8. K. R. Ludwig et al., Science 258, 284–287 (1992).9. R. N. Drysdale et al., Science 325, 1527–1531 (2009).10. A. L. Thomas et al., Science 324, 1186–1189 (2009).11. G. Marino et al., Nature 522, 197–201 (2015).12. R. L. Edwards, C. D. Gallup, Science 259, 1626–1627

(1993).13. R. L. Edwards, H. Cheng, M. T. Murrell, S. J. Goldstein, Science

276, 782–786 (1997).14. N. J. Shackleton, Nature 362, 596 (1993).15. K. R. Ludwig, K. R. Simmons, I. J. Winograd, B. J. Szabo,

A. C. Riggs, Science 259, 1626–1627 (1993).16. K. R. Ludwig et al., Nature 362, 596 (1993).17. C. D. Gallup, R. L. Edwards, R. G. Johnson, Science 263,

796–800 (1994).18. I. J. Winograd, J. M. Landwehr, K. R. Ludwig, T. B. Coplen,

A. C. Riggs, Quat. Res. 48, 141–154 (1997).19. W. S. Broecker, Nature 359, 779–780 (1992).20. P. M. Grootes, in Climatic Change in Continental Isotopic

Records, P. K. Swart et al., Eds. (Geophysical Monograph 78,American Geophysical Union, 1993), pp. 37–46.

21. T. D. Herbert et al., Science 293, 71–76 (2001).22. T. J. Crowley, K.-Y. Kim, Science 265, 1566–1568

(1994).23. I. J. Winograd et al., Quat. Res. 66, 202–212 (2006).24. Materials and methods are available as supplementary

materials on Science Online.25. J. D. Shakun, S. J. Burns, P. U. Clark, H. Cheng, R. L. Edwards,

Geophys. Res. Lett. 38, L18701 (2011).26. M. Cross et al., Quat. Sci. Rev. 127, 186–198 (2015).27. E. Steponaitis et al., Quat. Sci. Rev. 127, 174–185

(2015).28. M. P. Bacon, R. F. Anderson, J. Geophys. Res. 87, 2045–2056

(1982).29. S. B. Moran, M. A. Charette, J. A. Hoff, R. L. Edwards,

W. M. Landing, Earth Planet. Sci. Lett. 150, 151–160(1997).

30. M. S. Lachniet, R. F. Denniston, Y. Asmerom, V. Polyak,Nat. Commun. 5, 1–8 (2014).

31. J. S. Munroe, B. J. Laabs, Geol. Soc. Am. Bull. 125, 322–342(2013).

32. A. L. Berger, J. Atmos. Sci. 35, 2362–2367 (1978).33. J. R. Petit et al., Nature 399, 429–436 (1999).34. J. F. McManus, D. W. Oppo, J. L. Cullen, Science 283, 971–975

(1999).35. COHMAP Members, Science 241, 1043–1052 (1988).36. T. Kluge, H. P. Affek, Y. Dublyansky, C. Spötl, Earth Planet. Sci.

Lett. 400, 251–260 (2014).37. T. B. Coplen, I. J. Winograd, J. M. Landwehr, A. C. Riggs,

Science 263, 361–365 (1994).38. D. Lüthi et al., Nature 453, 379–382 (2008).39. L. Bazin et al., Clim. Past 9, 1715–1731 (2013).40. D. Veres et al., Clim. Past 9, 1733–1748 (2013).

ACKNOWLEDGMENTS

This work was supported by the Austrian Science Fund (FWF)project no. FP263050 to C.S. and in part by NSF grants1103403 to R.L.E. and H.C., 1103320 to R.L.E., and NSFC grant41230524 to H.C. This research was conducted under researchpermit numbers DEVA-2010-SCI-0004 and DEVA-2015-SCI-0006issued by Death Valley National Park. We thank I. J. Winogradfor providing the study with sample DH-2, M. Wimmer forpreparation and measurement of the stable isotopes, K. Wilsonand R. Freeze for assistance in the field, and M. Cross fordiscussion of Great Basin paleoclimate. Data can bedownloaded from the National Oceanic and AtmosphericAdministration’s National Centers for Environmental Information(www.ncdc.noaa.gov/paleo/paleo.html).

SUPPLEMENTARY MATERIALS

www.sciencemag.org/content/351/6269/165/suppl/DC1Materials and MethodsSupplementary TextFigs. S1 to S6Tables S1 and S2References (41–59)

10 September 2015; accepted 2 December 201510.1126/science.aad4132

168 8 JANUARY 2016 • VOL 351 ISSUE 6269 sciencemag.org SCIENCE

Fig. 4. The age of the TII d18O midpoint of –15.7per mil versus depth below the water table[+5.5 m above present elevation (24)] for eachsample. DH-11 (light green) (7, 8, 23), DH-2 (darkgreen) (this study), DH2-E (mid blue) (this study),and DH2-D (dark blue) (this study). Uncertaintiesare 2s. R2, coefficient of determination.

RESEARCH | REPORTSon S

eptember 15, 2020

http://science.sciencem

ag.org/D

ownloaded from

Reconciliation of the Devils Hole climate record with orbital forcing

Christoph SpötlGina E. Moseley, R. Lawrence Edwards, Kathleen A. Wendt, Hai Cheng, Yuri Dublyansky, Yanbin Lu, Ronny Boch and

DOI: 10.1126/science.aad4132 (6269), 165-168.351Science 

, this issue p. 165Sciencesamples.interaction with groundwater, which preferentially affected the deeper original samples but not the new shallowerexpected on the basis of orbital forcing. The age offset displayed by the older samples apparently was caused by

present analyses of a new set of data from Devils Hole that show that the deglaciation indeed occurred at the timeal.etpresumption of so-called orbital forcing, however, suggesting that deglaciation began ~10,000 years too early. Moseley

orbit around the Sun. The classic record of the penultimate deglaciation from Devils Hole, Nevada, did not reconcile the Glacial cycles are in part controlled by the pattern of incident solar energy determined by the geometry of Earth's

The difference is all in the water

ARTICLE TOOLS http://science.sciencemag.org/content/351/6269/165

MATERIALSSUPPLEMENTARY http://science.sciencemag.org/content/suppl/2016/01/06/351.6269.165.DC1

CONTENTRELATED

http://science.sciencemag.org/content/sci/354/6310/296.5.fullhttp://science.sciencemag.org/content/sci/354/6310/296.4.fullhttp://science.sciencemag.org/content/sci/354/6310/296.3.full

REFERENCES

http://science.sciencemag.org/content/351/6269/165#BIBLThis article cites 54 articles, 21 of which you can access for free

PERMISSIONS http://www.sciencemag.org/help/reprints-and-permissions

Terms of ServiceUse of this article is subject to the

is a registered trademark of AAAS.ScienceScience, 1200 New York Avenue NW, Washington, DC 20005. The title (print ISSN 0036-8075; online ISSN 1095-9203) is published by the American Association for the Advancement ofScience

Copyright © 2016, American Association for the Advancement of Science

on Septem

ber 15, 2020

http://science.sciencemag.org/

Dow

nloaded from