Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

Speleothem record attests to stable environmental conditions during Neanderthal–modern human turnover in southern Italy

An Author Correction to this article was published on 13 July 2020

This article has been updated


The causes of Neanderthal–modern human (MH) turnover are ambiguous. While potential biocultural interactions between the two groups are still little known, it is clear that Neanderthals in southern Europe disappeared about 42 thousand years ago (ka) after cohabitation for ~3,000 years with MH. Among a plethora of hypotheses on Neanderthal extinction, rapid climate changes during the Middle to Upper Palaeolithic transition (MUPT) are regarded as a primary factor. Here we show evidence for stable climatic and environmental conditions during the MUPT in a region (Apulia) where Neanderthals and MH coexisted. We base our findings on a rare glacial stalagmite deposited between ~106 and ~27 ka, providing the first continuous western Mediterranean speleothem palaeoclimate archive for this period. The uninterrupted growth of the stalagmite attests to the constant availability of rainfall and vegetated soils, while its δ13C–δ18O palaeoclimate proxies demonstrate that Apulia was not affected by dramatic climate oscillations during the MUPT. Our results imply that, because climate did not play a key role in the disappearance of Neanderthals in this area, Neanderthal–MH turnover must be approached from a perspective that takes into account climatic and environmental conditions favourable for both species.

This is a preview of subscription content, access via your institution

Relevant articles

Open Access articles citing this article.

Access options

Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Fig. 1: Cave samples.
Fig. 2: PC time series.
Fig. 3: PC versus Mediterranean records.

Data availability

Data supporting this study are available in Supplementary Table 1 and Supplementary Table 2.

Change history


  1. Benazzi, S. et al. Early dispersal of modern humans in Europe and implications for Neanderthal behaviour. Nature 479, 525–529 (2011).

    CAS  PubMed  Google Scholar 

  2. Wolf, D. et al. Climate deteriorations and Neanderthal demise in interior Iberia. Sci. Rep. 8, 7048 (2018).

    CAS  PubMed  PubMed Central  Google Scholar 

  3. Mellars, P. A new radiocarbon revolution and the dispersal of modern humans in Eurasia. Nature 439, 931–935 (2006).

    CAS  PubMed  Google Scholar 

  4. Müller, U. C. et al. The role of climate in the spread of modern humans into Europe. Quat. Sci. Rev. 30, 273–279 (2011).

    Google Scholar 

  5. Staubwasser, M. et al. Impact of climate change on the transition of Neanderthals to modern humans in Europe. Proc. Natl Acad. Sci. USA 115, 9116–9121 (2018).

    CAS  PubMed  Google Scholar 

  6. Melchionna, M. et al. Fragmentation of Neanderthals’ pre-extinction distribution by climate change. Palaeogeogr. Palaeoclimatol. Palaeoecol. 496, 146–154 (2018).

    Google Scholar 

  7. Finlayson, C. & Carrion, J. S. Rapid ecological turnover and its impact on Neanderthal and other human populations. Trends Ecol. Evol. 22, 213–222 (2007).

    PubMed  Google Scholar 

  8. Stewart, J. R. The ecology and adaptation of Neanderthals during the non-analogue environment of Oxygen Isotope Stage 3. Quat. Int. 137, 35–46 (2005).

    Google Scholar 

  9. Genty, D. et al. Precise dating of Dansgaard–Oeschger climate oscillations in western Europe from stalagmite data. Nature 42, 833–837 (2003).

    Google Scholar 

  10. Pérez-Mejías, C. et al. Orbital-to-millennial scale climate variability during Marine Isotope Stages 5 to 3 in northeast Iberia. Quat. Sci. Rev. 224, 105946 (2019).

  11. Badino, F. et al. An overview of Alpine and Mediterranean palaeogeography, terrestrial ecosystems and climate history during MIS 3 with focus on the Middle to Upper Palaeolithic transition. Quat. Int. (2019).

  12. Higham, T. et al. The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512, 306–309 (2014).

    Google Scholar 

  13. Rey-Rodríguez, I. et al. Last Neanderthals and first anatomically modern humans in the NW Iberian Peninsula: climatic and environmental conditions inferred from the Cova Eirós small-vertebrate assemblage during MIS 3. Quat. Sci. Rev. 151, 185–197 (2016).

    Google Scholar 

  14. Benazzi, S. et al. The makers of the Protoaurignacian and implications for Neandertal extinction. Science 348, 793–796 (2015).

    CAS  PubMed  Google Scholar 

  15. NGRIP Members. High-resolution record of Northern Hemisphere climate extending into the last interglacial period. Nature 431, 147–151 (2004).

    Google Scholar 

  16. Kudielka, G. et al. in 250 Million Years of Earth History in Central Italy: Celebrating 25 Years of the Geological Observatory of Coldigioco Vol. 542 (eds Koeberl, C. & Bice, D. M.) 429–445 (Geological Society of America, 2019);

  17. Budsky, A. et al. Western Mediterranean climate response to Dansgaard/Oeschger events: new insights from speleothem secords. Geophys. Res. Lett. 46, 9042–9053 (2019).

  18. Denniston, R. F. et al. A stalagmite test of North Atlantic SST and Iberian hydroclimate linkages over the last two glacial cycles. Clim. Past 14, 1893–1913 (2018).

    Google Scholar 

  19. Badertscher, S. et al. Pleistocene water intrusions from the Mediterranean and Caspian seas into the Black Sea. Nature Geosci. 4, 236–239 (2011).

    CAS  Google Scholar 

  20. Bar-Matthews, M., Ayalon, A., Gilmour, M., Matthews, A. & Hawkesworth, C. J. Sea–land oxygen isotopic relationships from planktonic foraminifera and speleothems in the Eastern Mediterranean region and their implication for paleorainfall during interglacial intervals. Geochim. Cosmochim. Acta 67, 3181–3199 (2003).

    CAS  Google Scholar 

  21. Yasur, G. et al. Climatic and environmental conditions in the Western Galilee, during Late Middle and Upper Paleolithic periods, based on speleothems from Manot Cave, Israel. J. Hum. Evol. (2019).

    Article  PubMed  Google Scholar 

  22. McDermott, F. Palaeo-climate reconstruction from stable isotope variations in speleothems: a review. Quat. Sci. Rev. 23, 901–918 (2004).

    Google Scholar 

  23. Drysdale, R. N. et al. Evidence for obliquity forcing of glacial Termination II. Science 325, 1527–1531 (2009).

    CAS  PubMed  Google Scholar 

  24. Luetscher, M. et al. North Atlantic storm track changes during the Last Glacial Maximum recorded by Alpine speleothems. Nat. Commun. 6, 6344 (2015).

    CAS  PubMed  PubMed Central  Google Scholar 

  25. Rasmussen, S. O. et al. A stratigraphic framework for abrupt climatic changes during the Last Glacial period based on three synchronized Greenland ice-core records: refining and extending the INTIMATE event stratigraphy. Quat. Sci. Rev. 106, 14–28 (2014).

    Google Scholar 

  26. Cheng, H. et al. The climatic cyclicity in semiarid-arid central Asia over the past 500,000 years. Geophys. Res. Lett. 39, L01705 (2012).

    Google Scholar 

  27. Columbu, A. et al. A long record of MIS 7 and MIS 5 climate and environment from a western Mediterranean speleothem (SW Sardinia, Italy). Quat. Sci. Rev. 220, 230–243 (2019).

    Google Scholar 

  28. Allen, J. R. M. et al. Rapid environmental changes in southern Europe during the last glacial period. Science 400, 740–743 (1999).

    CAS  Google Scholar 

  29. Tzedakis, P. C., Hooghiemstra, H. & Pälike, H. The last 1.35 million years at Tenaghi Philippon: revised chronostratigraphy and long-term vegetation trends. Quat. Sci. Rev. 25, 3416–3430 (2006).

    Google Scholar 

  30. Toucanne, S. et al. Tracking rainfall in the northern Mediterranean borderlands during sapropel deposition. Quat. Sci. Rev. 129, 178–195 (2015).

    Google Scholar 

  31. Hodell, D. A., Channell, J. E. T., Curtis, J. H., Romero, O. E. & Röhl, U. Onset of “Hudson strait” Heinrich events in the eastern North Atlantic at the end of the middle Pleistocene transition (~640 ka)?: Pleistocene Heinrich events. Paleoceanography 23, PA4218 (2008).

    Google Scholar 

  32. Kaufmann, G. & Dreybrodt, W. Stalagmite growth and palaeo-climate: an inverse approach. Earth Planet. Sci. Lett. 224, 529–545 (2004).

    CAS  Google Scholar 

  33. Columbu, A. et al. A long continuous palaeoclimate-palaeoenvironmental record of the last glacial period from southern Italy and implications for the coexistence of Anatomically Modern Humans and Neanderthals. Proc. European Geosciences Union (EGU) Conference (2020).

  34. Stewart, J. & Stringer, B. Human evolution out of Africa: the role of refugia and climate change. Science 335, 1317–1321 (2012).

    CAS  PubMed  Google Scholar 

  35. Ait Brahim, Y. et al. North Atlantic ice-rafting, ocean and atmospheric circulation during the Holocene: insights from Western Mediterranean speleothems. Geophys. Res. Lett. 46, 7614–7623 (2019).

  36. Sano, K. et al. The earliest evidence for mechanically delivered projectile weapons in Europe. Nat. Ecol. Evol. 3, 1409–1414 (2019).

    PubMed  PubMed Central  Google Scholar 

  37. Arrighi, S. et al. Backdating systematic shell ornament making in Europe to 45,000 years ago. Archaeol. Anthropol. Sci. 12, 59 (2020).

    Google Scholar 

  38. Arrighi, S. et al. Bone tools, ornaments and other unusual objects during the Middle to Upper Palaeolithic transition in Italy. Quat. Int. (2019).

  39. Marciani, G. et al. Lithic techno-complexes in Italy from 50 to 39 thousand years BP: an overview of lithic technological changes across the Middle–Upper Palaeolithic boundary. Quat. Int. (2019).

  40. Drysdale, R. N. et al. Precise microsampling of poorly laminated speleothems for U-series dating. Quat. Geochronol. 14, 38–47 (2012).

    Google Scholar 

  41. Hellstrom, J. U–Th dating of speleothems with high initial 230Th using stratigraphical constraint. Quat. Geochronol. 1, 289–295 (2006).

    Google Scholar 

  42. Cheng, H. et al. Improvements in 230Th dating, 230Th and 234U half-life values, and U–Th isotopic measurements by multi-collector inductively coupled plasma mass spectrometry. Earth Planet. Sci. Lett. 371–372, 82–91 (2013).

    Google Scholar 

  43. Scholz, D. & Hoffmann, D. L. StalAge – an algorithm designed for construction of speleothem age models. Quat. Geochronol. 6, 369–382 (2011).

    Google Scholar 

  44. Breitenbach, S. F. M. et al. COnstructing Proxy-Record Age models (COPRA). Clim. Past 8, 1765–1779 (2012).

    Google Scholar 

  45. Hendy, C. H. The isotopic geochemistry of speleothems-I. The calculation of the effects of different modes of formation on the isotopic composition of speleothems and their applicability as palaeoclimatic indicators. Geochim. Cosmochim. Acta 35, 801–824 (1971).

    CAS  Google Scholar 

  46. Mickler, P. J., Stern, L. A. & Banner, J. L. Large kinetic isotope effects in modern speleothems. Geol. Soc. Am. Bull. 118, 65–81 (2006).

    CAS  Google Scholar 

  47. Dreybrodt, W. & Scholz, D. Climatic dependence of stable carbon and oxygen isotope signals recorded in speleothems: from soil water to speleothem calcite. Geochim. Cosmochim. Acta 75, 734–752 (2011).

    CAS  Google Scholar 

  48. Columbu, A., Sauro, F., Lundberg, J., Drysdale, R. & De Waele, J. Palaeoenvironmental changes recorded by speleothems of the southern Alps (Piani Eterni, Belluno, Italy) during four interglacial to glacial climate transitions. Quat. Sci. Rev. 197, 319–335 (2018).

    Google Scholar 

  49. Columbu, A. et al. Early last glacial intra-interstadial climate variability recorded in a Sardinian speleothem. Quat. Sci. Rev. 169, 391–397 (2017).

    Google Scholar 

  50. Bard, E. et al. Hydrological conditions over the western Mediterranean basin during the deposition of the cold Sapropel 6 (ca. 175 kyr BP). Earth Planet. Sci. Lett. 202, 481–494 (2002).

    CAS  Google Scholar 

  51. Kim, S.-T. & O’Neil, J. R. Equilibrium and nonequilibrium oxygen isotope effects in synthetic carbonates. Geochim. Cosmochim. Acta 61, 3461–3475 (1997).

    CAS  Google Scholar 

  52. Tremaine, D. M., Froelich, P. N. & Wang, Y. Speleothem calcite farmed in situ: modern calibration of δ18O and δ13C paleoclimate proxies in a continuously-monitored natural cave system. Geochim. Cosmochim. Acta 75, 4929–4950 (2011).

    CAS  Google Scholar 

  53. Drysdale, R. N. et al. Stalagmite evidence for the precise timing of North Atlantic cold events during the early last glacial. Geology 35, 77–80 (2007).

    CAS  Google Scholar 

  54. Vanghi, V. et al. Climate variability on the Adriatic seaboard during the last glacial inception and MIS 5c from Frasassi Cave stalagmite record. Quat. Sci. Rev. 201, 349–361 (2018).

    Google Scholar 

  55. Regattieri, E. et al. A MIS 9/MIS 8 speleothem record of hydrological variability from Macedonia (F.Y.R.O.M.). Glob. Planet. Change 162, 39–52 (2018).

    Google Scholar 

  56. Ford, D. & Williams, P. Karst Geomorphology and Hydrology (John Wiley & Sons, 2007).

  57. Bajo, P. et al. Stalagmite carbon isotopes and dead carbon proportion (DCP) in a near-closed-system situation: an interplay between sulphuric and carbonic acid dissolution. Geochim. Cosmochim. Acta 210, 208–227 (2017).

    CAS  Google Scholar 

  58. Fairchild, I. J. & Treble, P. C. Trace elements in speleothems as recorders of environmental change. Quat. Sci. Rev. 28, 449–468 (2009).

    Google Scholar 

  59. Longinelli, A. & Selmo, E. Isotopic composition of precipitation in Italy: a first overall map. J. Hydrol. 270, 75–88 (2003).

    CAS  Google Scholar 

  60. Dansgaard, W. Stable isotopes in precipitation. Tellus 16, 436–468 (1964).

    Google Scholar 

  61. Lisiecki, L. E. & Raymo, M. E. A Pliocene–Pleistocene stack of 57 globally distributed benthic δ18O records. Paleoceanography 20, PA1003 (2005).

    Google Scholar 

  62. Frisia, S., Borsato, A., Preto, N. & McDermott, F. Late Holocene annual growth in three Alpine stalagmites records the influence of solar activity and the North Atlantic Oscillation on winter climate. Earth Planet. Sci. Lett. 216, 411–424 (2003).

    CAS  Google Scholar 

  63. Johnston, V. E. et al. Evidence of thermophilisation and elevation-dependent warming during the Last Interglacial in the Italian Alps. Sci. Rep. 8, 2680 (2018).

    CAS  PubMed  PubMed Central  Google Scholar 

  64. Belli, R. et al. Regional climate variability and ecosystem responses to the last deglaciation in the northern hemisphere from stable isotope data and calcite fabrics in two northern Adriatic stalagmites. Quat. Sci. Rev. 72, 146–158 (2013).

    Google Scholar 

  65. Pozzi, J. P. et al. U–Th dated speleothem recorded geomagnetic excursions in the Lower Brunhes. Sci. Rep. 9, 1114 (2019).

    PubMed  PubMed Central  Google Scholar 

  66. Regattieri, E. et al. Holocene Critical Zone dynamics in an Alpine catchment inferred from a speleothem multiproxy record: disentangling climate and human influences. Sci. Rep. 9, 17829 (2019).

  67. Regattieri, E. et al. A continuous stable isotope record from the penultimate glacial maximum to the Last Interglacial (159–121 ka) from Tana Che Urla Cave (Apuan Alps, central Italy). Quat. Res. 82, 450–461 (2014).

    CAS  Google Scholar 

  68. Isola, I. et al. Speleothem U/Th age constraints for the Last Glacial conditions in the Apuan Alps, northwestern Italy. Palaeogeogr. Palaeoclimatol. Palaeoecol. 518, 62–71 (2019).

    Google Scholar 

  69. Zhornyak, L.V. et al. Stratigraphic evidence for a “pluvial phase” between ca 8200–7100 ka from Renella cave (Central Italy). Quat. Sci. Rev. 30, 409–417 (2011).

    Google Scholar 

  70. Columbu, A. et al. Late quaternary speleogenesis and landscape evolution in the northern Apennine evaporite areas. Earth Surf. Process. Landf. 42, 1447–1459 (2017).

    Google Scholar 

  71. Frisia, S. et al. Holocene climate variability in Sicily from a discontinuous stalagmite record and the Mesolithic to Neolithic transition. Quat. Res. 66, 388–400 (2006).

    CAS  Google Scholar 

  72. Francke, A. et al. Sedimentological processes and environmental variability at Lake Ohrid (Macedonia, Albania) between 637 ka and the present. Biogeosciences 13, 1179–1196 (2016).

    CAS  Google Scholar 

  73. Moseley, G. E. et al. NALPS19: sub-orbital scale climate variability recorded in Northern Alpine speleothems during the last glacial period. Clim. Past 16, 29–50 (2020).

Download references


We thank all local speleologists that helped with the 2014 and 2019 fieldwork at Pozzo Cucù, Sant’Angelo, Zaccaria and Messapi caves: G. Loperfido, S. Inguscio, G. Ragone, P. Lippolis, A. Lacirignola, D. Leserri, M. Marraffa, O. Lacarbonara, F. Semeraro, S. Calella, P. Calella, C. Pastore, C. Marchitelli, R. Romanazzi, R. Cupertino, G. Caló and F. Lorusso (Gruppo Speleologico Martinese, CARS Altamura, Gruppo Speleologico Neretino, Gruppo Ricerche Carsiche Putignano, Gruppo Puglia Grotte and Gruppo Escursionistico Speleologico Ostunense), as well as the Bellanova family for access to Messapi Cave. A.C., J.D.W. and V.C. are also grateful to all members of Gruppo Speleologico Martinese for their logistic help and warm hospitality in Martina Franca. Thanks also to M. Parise (University of Bari) for help during 2014 fieldwork; A. Reina (Polytechnic University of Bari) for his enthusiasm in supporting this research; V. Casulli and R. Laragione of Castellana Grotte for their interest in supporting this study; M. Wimmer and M. Luetscher (Innsbruck University) for their help during laboratory work; L. Pisani (Bologna University) for the DEM figure used in Extended Data Fig. 1; and L. Calabrò (Bologna University) for drilling of sample SA1. A.C. is supported by Leonardo Da Vinci Grant 2019 (DD MIUR, no. 787, 15/04/2019); S.B. is supported by ERC grant no. 724046—SUCCESS (, and H.C. by NSFC grant no. 41888101. This research received financial contributions from both Grotte di Castellana and Federazione Speleologica Pugliese.

Author information

Authors and Affiliations



A.C. and V.C. conceived and designed the experiments. A.C., V.C., C.S., J.H. and H.C. performed the experiments. A.C. and S.B. analysed the data. A.C., V.C., C.S., S.B. and J.D.W. contributed with materials and analysis tools. A.C. wrote the paper with input from all co-authors.

Corresponding author

Correspondence to Andrea Columbu.

Ethics declarations

Competing interests

The authors declare no competing interests

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data

Extended Data Fig. 1 Cave locations and sampling.

a) Apulia region with colour-coded topography; black dots indicate major caves in Puglia (from, while red circles report caves explored for this project. b) PC stalagmite found close to its original growth position. c) PC milling subsampling for δ18O and δ13C.

Extended Data Fig. 2 Intra-millennia events.

Intra-stadial and interstadial events recorded by PC δ18O compared to Greenland ice core δ18O (24). The events are reported, in both curves, with grey shading.

Extended Data Fig. 3 PC from DO 23 to 19.

Similarities between Chinese speleothem δ18O (orange curve26), PC δ18O (blue curve, this study) and Greenland ice core δ18O (green curve15) from ~110 to ~65 ka, during DO events 23 to 19.

Extended Data Fig. 4 Growth rate and [234/238U]i.

Comparison between PC δ13C, growth rate, δ18O and [234/238U]i.

Extended Data Fig. 5 Materials.

Speleothems (except PC, Fig. 1 main text) examined in this study. Blue rectangles indicate the location of sampling for U-Th dating.

Extended Data Fig. 6 Age model.

Top: comparison between StalAge and COPRA 2σ range and the resulting average age model used in this work. Bottom: Propagation of positive and negative 2σ uncertainty in the various age models. U-Th ages are shown by yellow dots and black 2σ error bars.

Extended Data Fig. 7 Hendy test.

Subsamples were extracted from individual growth lamina. δ13C-δ18O correlation (r) provided for each tested layer. The absence of a strong correlation between δ13C and δ18O and of a systematic increase from the centre to the flank indicate that calcite was deposited under quasi-equilibrium conditions (see Hendy test discussion for details).

Supplementary information

Reporting Summary

Supplementary Table 1

U–Th dataset.

Supplementary Table 2

Stable isotope time series.

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Columbu, A., Chiarini, V., Spötl, C. et al. Speleothem record attests to stable environmental conditions during Neanderthal–modern human turnover in southern Italy. Nat Ecol Evol 4, 1188–1195 (2020).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

This article is cited by


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing