Hublin, J.

English

Hublin, Jean-Jacques

HAL Descartes

The last Neanderthal

MPG.PuRe

Hal-Diderot

Hublin, J. 

COMMENTARYThe last NeanderthalJean-Jacques Hublina,1The mechanism of the Neanderthal extinction and theirreplacement by modern humans of African origin is oneof the most discussed issues in paleoanthropology.Central to this discussion are the questions of thechronological overlap between Neanderthal popula-tions and modern humans in Western Eurasia and theprecise geographical circumstances of this overlap.For a long time, the Vindija (Croatia) site was consid-ered to provide solid evidence for a long survival ofNeanderthals in Central/Southern Europe. Not onlydid directly dated Neanderthal remains from layerG1 of the site provide radiocarbon ages postdatingthe most widely accepted transition time of 40–35,000radiocarbon years ago (1), but the same layer alsoyielded a type of split-based bone points commonlyassigned to the Aurignacian (2), a stone artefact industryof the early Upper Paleolithic that, to date, only yieldedhuman remains of a modern nature (3). For some, thissituation implied the possibility of a long and complexinteraction between the two groups of hominins in thisregion and also falsified the notion of a systematic asso-ciation between defined archaeological assemblagesand specific biological populations at the time of thereplacement. In PNAS, Devièse et al. (4) provide newradiocarbon dates for the same Vindija Neanderthalsamples, dating them to before 40,000 14C B.P., sig-nificantly older than previous efforts dating this materialto 29–28,000 and 33–32,000 radiocarbon years (1). Thebone points of layer G1 could not be dated, but therange of ages obtained from faunal and human samplesin this layer suggests taphonomic mixing as a likelymechanism to explain their stratigraphic association inthis part of the Vindija stratigraphic sequence. The situ-ation in Vindija is therefore not at all exceptional, andprevious results can be explained by the effect of samplecontamination and layer admixture.The first message delivered by the new study is oneof prudence. Radiocarbon is the most precise methodof directly dating human fossil remains. However, atthe limit of its application range around 45–40,000 cal-endar years ago, which unfortunately corresponds to theperiod of replacement of the last Neanderthals, it ishighly sensitive to contamination. For a radiocarbondate of 25,000 y, 1% contamination by modern carbonwill produce an age that is 1,628 y too young, but for aradiocarbon date of 42,500 y, the apparent age shifts8,455 y toward the present (Fig. 1). This length of time isin the higher range of estimates for possible overlapbetween Neanderthals andmodern humans at the scaleof Western Eurasia (3, 5). It is therefore critical whendating organic material extracted from fossil bonesto authenticate it as composed of degraded proteins,essentially collagen, of the bone itself. From this per-spective, the development of extraction techniquesimplementing “ultrafiltration” in order to eliminate smallcontaminants represented a major step forward (6). Eversince, sample pretreatments have witnessed several im-portant improvements. In practical terms, these succes-sive advances mean that, for the transition period, thelarge number of radiocarbon dates on bone samplesproduced before 2004 should be used with great cau-tion, if not simply forgotten. To overcome contaminationissues in an even more thorough way, Devièse et al. (4)used a method based on the extraction of hydroxypro-line, an amino acid specific to collagen. Hydroxyprolinedating relies on significant bone sample sizes, as thedated carbon atoms are restricted to those deriving froma single amino acid making up roughly 10% of all aminoacid positions of mature collagen type I. Hence, bonesample sizes for hydroxyproline dating are larger thanFig. 1. Contamination effect on radiocarbon dates. Theblack dots represent three examples of radiocarbondates without contamination, and the red dots representthe effect of only 1% contamination by modern carbonon the same samples.aDepartment of Human Evolution, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, GermanyAuthor contributions: J.-J.H. wrote the paper.The author declares no conflict of interest.See companion article on page 10606.1Email: hublin@eva.mpg.de.10520–10522 | PNAS | October 3, 2017 | vol. 114 | no. 40 www.pnas.org/cgi/doi/10.1073/pnas.1714533114COMMENTARYcommonly used for ultrafiltration pretreatment and result in greaterSEs than with the ultrafiltration technique (table 2 of ref. 4). In thepresent state of the art, this approach can only be envisioned whenlarge fragments bearing little anatomical information are available.New methods, such as collagen peptide mass fingerprinting(ZooMS), might provide access to such human bone specimensby large-scale screening projects (7).Since other cases of Neanderthal late survival, such as RipparoMezzena in Northern Italy (8), have now been falsified, the latestoccurrences of Neanderthal remains in the European fossil recordcan now be documented by specimens discovered in associationwith so-called “transitional industries.” These industries display fea-tures inherited from local Middle Paleolithic assemblages producedby Neanderthals, combined with Upper Paleolithic innovations sim-ilar to those encountered in the Aurignacian. At Saint-Ce´saire and atthe Grotte du Renne, France, Neanderthal remains associated witha transitional industry called Chaˆtelperronian, which already displaysmany Upper Paleolithic features, were directly dated to 36,200 ±750 and 36,840 ± 660 14C B.P., respectively (7, 9). Another seriesof Neanderthal remains from the cave of Spy, Belgium, has alsobeen directly dated to ca. 36,000 14C B.P. (10). This age falls in thetime range of the Lincombian-Ranisian-Jerzmanowician (LRJ), a Eu-ropean transitional industry identified in this site. If one assumesthat the entire Chaˆtelperronian and LRJ were produced by lateNeanderthals, this would push the last occurrence date for thesepopulations to ca. 35,000 14C B.P. Converting this radiocarbon dateinto a calendar age corresponds to ∼40,000 y ago. By that time,modern humans producing Aurignacian industries had already oc-cupied the neighboring regions of Austria (11), Germany (12), andNorthern Italy (13) for some time. Interactions with modern groupsmay explain the cultural evolution of the last Neanderthals, but apuzzling issue remains regarding possible biological interactionsbetween the two groups. Introgression of Neanderthal DNA intothe modern genome has been widely demonstrated and analyzedthrough the endless availability of extant genetic data. It is quite likelythat gene flow also occurred in the opposite direction. However, todate not a single detailed nuclear DNA sequence has been recon-structed from one of these postcontact Neanderthal populations andit is therefore impossible to assess the magnitude and possible effectof modern DNA introgression into late Neanderthal populations.As far as archaeological assemblages of Middle Paleolithictype can be considered a good proxy to identify the occurrence ofNeanderthal populations in different parts of Western Europe, thesouth of the Iberian Peninsula has sometimes been proposed asan area of their late survival. Specifically, south of the Ebro river,Neanderthals may have survived several millennia after theirextinction in the rest of Western Europe (14). This idea is heavilydisputed, as the dating of some of the sites on which this modelwas based have been revised to older ages (15). Still, sites in theMula basin of Murcia, Spain, are claimed to document a modernreplacement of Neanderthals taking place as late as around37,000 y ago in calibrated chronology (16). The explanations pro-vided for this delay are primarily geographical and environmental,and partly relate to the expansion of forested environments dur-ing this time period in Iberian regions south of 40°N.If one takes a broader view away from the European continent,a series of recent studies have supported early modern humanexpansion in tropical Asia, as far as China (17), Laos (18), and Indo-nesia (19), and ultimately in Australia (20) more than 60,000 y ago.If substantiated by further discoveries, this situation would set thefar-west Eurasia as a region of delayed replacement for archaiclocal populations. If modern humans were then able to settle inAsian tropical forests and cross large bodies of water, it is difficultto conceive that their expansion in the Iberian Peninsula wouldThe last chapter of the Vindija saga reminds usonce more that the interpretation of materialfrom older excavations is quite challenging, asthe precise archaeological context of discoveriesthat took place several decades ago generallyremains a topic of continuous debate.have been stalled by a river and deciduous temperate forests.Regardless of the “Ebro frontier” issue, it is striking to considerthat the places where the latest survival of Neanderthals is bestdocumented by the direct dating of diagnostic human remainsactually correspond to areas where Neanderthal populationsdisplayed the highest density and a continuous occupation for thetens of millennia preceding their final demise. This suggests thatalthough environmental conditions might have influenced thetempo of modern human penetration in Europe, the decisivefactor that slowed their settlement could have been the very pres-ence of relatively dense Neanderthal populations well adapted tothe Pleistocene environments of the middle latitudes.The last chapter of the Vindija saga reminds us oncemore that theinterpretation of material from older excavations is quite challenging,as the precise archaeological context of discoveries that took placeseveral decades ago generally remains a topic of continuous debate.The resolution of pending issues regarding the European expansionof modern humans will come from the study of new sites or thereassessment of earlier discovered oneswith entirely new approaches.Without relying too much on the hope of discoveries, such asspectacular human remains, major progress can be accomplishedin the near future through the implementation of new methodo-logical approaches ranging from the screening of anatomicallynondiagnostic human bone remains out of faunal assemblages bypeptide mass fingerprinting of collagen to improvements to comein the reduction of sample sizes for direct radiocarbon dating.1 Higham T, Ramsey CB, Karavanic´ I, Smith FH, Trinkaus E (2006) Revised direct radiocarbon dating of the Vindija G1 Upper Paleolithic Neandertals. Proc Natl AcadSci USA 103:553–557.2 Karavanic´ I, Smith FH (1998) The Middle/Upper Paleolithic interface and the relationship of Neanderthals and early modern humans in the Hrvatsko Zagorje,Croatia. J Hum Evol 34:223–248.3 Hublin J-J (2015) The modern human colonization of western Eurasia: When and where? Quat Sci Rev 118:194–210.4 Devièse T, et al. (2017) Direct dating of Neanderthal remains from the site of Vindija Cave and implications for the Middle to Upper Paleolithic transition. Proc NatlAcad Sci USA 114:10606–10611.5 Higham T, et al. (2014) The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512:306–309.6 Brown TA, Nelson DE, Vogel JS, Southon JR (1988) Improved collagen extraction by modified Longin method. Radiocarbon 30:171–177.7 Welker F, et al. (2016) Palaeoproteomic evidence identifies archaic hominins associated with the Chaˆtelperronian at the Grotte du Renne. Proc Natl Acad Sci USA113:11162–11167.Hublin PNAS | October 3, 2017 | vol. 114 | no. 40 | 105218 Talamo S, et al. (2016) Direct radiocarbon dating and genetic analyses on the purported Neanderthal mandible from the Monti Lessini (Italy). Sci Rep 6:29144.9 Hublin J-J, et al. (2012) Radiocarbon dates from the Grotte du Renne and Saint-Ce´saire support a Neandertal origin for the Chaˆtelperronian. Proc Natl Acad SciUSA 109:18743–18748.10 Semal P, et al. (2009) New data on the late Neandertals: Direct dating of the Belgian Spy fossils. Am J Phys Anthropol 138:421–428.11 Nigst PR, et al. (2014) Early modern human settlement of Europe north of the Alps occurred 43,500 years ago in a cold steppe-type environment. Proc Natl AcadSci USA 111:14394–14399.12 Higham T, et al. (2012) Τesting models for the beginnings of the Aurignacian and the advent of figurative art and music: The radiocarbon chronology ofGeißenklösterle. J Hum Evol 62:664–676.13 Benazzi S, et al. (2015) Archaeology. The makers of the Protoaurignacian and implications for Neandertal extinction. Science 348:793–796.14 Zilh~ao J (2000) The Ebro Frontier: A model for the late extinction of Iberian neanderthals.Neanderthals on the Edge: Papers from a Conference Marking the 150thAnniversary of the Forbes’ Quarry Discovery, Gibraltar, eds Stringer CB, Barton RN, Finlayson JC (Oxbow Books, Oxford), pp 111–122.15 Wood RE, et al. (2013) Radiocarbon dating casts doubt on the late chronology of the Middle to Upper Palaeolithic transition in southern Iberia. Proc Natl Acad SciUSA 110:2781–2786.16 Zilh~ao J, et al. (2016) Cueva Anto´n: A multi-proxy MIS 3 to MIS 5a paleoenvironmental record for SE Iberia. Quat Sci Rev 146:251–273.17 Liu W, et al. (2015) The earliest unequivocally modern humans in southern China. Nature 526:696–699.18 Demeter F, et al. (2015) Early modern humans and morphological variation in Southeast Asia: Fossil evidence from Tam Pa Ling, Laos. PLoS One 10:e0121193.19 Westaway KE, et al. (2017) An early modern human presence in Sumatra 73,000-63,000 years ago. Nature 548:322–325.20 Clarkson C, et al. (2017) Human occupation of northern Australia by 65,000 years ago. Nature 547:306–310.10522 | www.pnas.org/cgi/doi/10.1073/pnas.1714533114 Hublin

Jean-Jacques Hublin

Crossref

Proceedings of the National Academy of Sciences

HAL: Hyper Article en Ligne

Hublin, J.J.

Leiden University Scholary Publications

COMMENTARYThe last NeanderthalJean-Jacques Hublina,1The mechanism of the Neanderthal extinction and theirreplacement by modern humans of African origin is oneof the most discussed issues in paleoanthropology.Central to this discussion are the questions of thechronological overlap between Neanderthal popula-tions and modern humans in Western Eurasia and theprecise geographical circumstances of this overlap.For a long time, the Vindija (Croatia) site was consid-ered to provide solid evidence for a long survival ofNeanderthals in Central/Southern Europe. Not onlydid directly dated Neanderthal remains from layerG1 of the site provide radiocarbon ages postdatingthe most widely accepted transition time of 40–35,000radiocarbon years ago (1), but the same layer alsoyielded a type of split-based bone points commonlyassigned to the Aurignacian (2), a stone artefact industryof the early Upper Paleolithic that, to date, only yieldedhuman remains of a modern nature (3). For some, thissituation implied the possibility of a long and complexinteraction between the two groups of hominins in thisregion and also falsified the notion of a systematic asso-ciation between defined archaeological assemblagesand specific biological populations at the time of thereplacement. In PNAS, Devièse et al. (4) provide newradiocarbon dates for the same Vindija Neanderthalsamples, dating them to before 40,000 14C B.P., sig-nificantly older than previous efforts dating this materialto 29–28,000 and 33–32,000 radiocarbon years (1). Thebone points of layer G1 could not be dated, but therange of ages obtained from faunal and human samplesin this layer suggests taphonomic mixing as a likelymechanism to explain their stratigraphic association inthis part of the Vindija stratigraphic sequence. The situ-ation in Vindija is therefore not at all exceptional, andprevious results can be explained by the effect of samplecontamination and layer admixture.The first message delivered by the new study is oneof prudence. Radiocarbon is the most precise methodof directly dating human fossil remains. However, atthe limit of its application range around 45–40,000 cal-endar years ago, which unfortunately corresponds to theperiod of replacement of the last Neanderthals, it ishighly sensitive to contamination. For a radiocarbondate of 25,000 y, 1% contamination by modern carbonwill produce an age that is 1,628 y too young, but for aradiocarbon date of 42,500 y, the apparent age shifts8,455 y toward the present (Fig. 1). This length of time isin the higher range of estimates for possible overlapbetween Neanderthals andmodern humans at the scaleof Western Eurasia (3, 5). It is therefore critical whendating organic material extracted from fossil bonesto authenticate it as composed of degraded proteins,essentially collagen, of the bone itself. From this per-spective, the development of extraction techniquesimplementing “ultrafiltration” in order to eliminate smallcontaminants represented a major step forward (6). Eversince, sample pretreatments have witnessed several im-portant improvements. In practical terms, these succes-sive advances mean that, for the transition period, thelarge number of radiocarbon dates on bone samplesproduced before 2004 should be used with great cau-tion, if not simply forgotten. To overcome contaminationissues in an even more thorough way, Devièse et al. (4)used a method based on the extraction of hydroxypro-line, an amino acid specific to collagen. Hydroxyprolinedating relies on significant bone sample sizes, as thedated carbon atoms are restricted to those deriving froma single amino acid making up roughly 10% of all aminoacid positions of mature collagen type I. Hence, bonesample sizes for hydroxyproline dating are larger thanFig. 1. Contamination effect on radiocarbon dates. Theblack dots represent three examples of radiocarbondates without contamination, and the red dots representthe effect of only 1% contamination by modern carbonon the same samples.aDepartment of Human Evolution, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, GermanyAuthor contributions: J.-J.H. wrote the paper.The author declares no conflict of interest.See companion article on page 10606.1Email: hublin@eva.mpg.de.10520–10522 | PNAS | October 3, 2017 | vol. 114 | no. 40 www.pnas.org/cgi/doi/10.1073/pnas.1714533114COMMENTARYcommonly used for ultrafiltration pretreatment and result in greaterSEs than with the ultrafiltration technique (table 2 of ref. 4). In thepresent state of the art, this approach can only be envisioned whenlarge fragments bearing little anatomical information are available.New methods, such as collagen peptide mass fingerprinting(ZooMS), might provide access to such human bone specimensby large-scale screening projects (7).Since other cases of Neanderthal late survival, such as RipparoMezzena in Northern Italy (8), have now been falsified, the latestoccurrences of Neanderthal remains in the European fossil recordcan now be documented by specimens discovered in associationwith so-called “transitional industries.” These industries display fea-tures inherited from local Middle Paleolithic assemblages producedby Neanderthals, combined with Upper Paleolithic innovations sim-ilar to those encountered in the Aurignacian. At Saint-Césaire and atthe Grotte du Renne, France, Neanderthal remains associated witha transitional industry called Châtelperronian, which already displaysmany Upper Paleolithic features, were directly dated to 36,200 ±750 and 36,840 ± 660 14C B.P., respectively (7, 9). Another seriesof Neanderthal remains from the cave of Spy, Belgium, has alsobeen directly dated to ca. 36,000 14C B.P. (10). This age falls in thetime range of the Lincombian-Ranisian-Jerzmanowician (LRJ), a Eu-ropean transitional industry identified in this site. If one assumesthat the entire Châtelperronian and LRJ were produced by lateNeanderthals, this would push the last occurrence date for thesepopulations to ca. 35,000 14C B.P. Converting this radiocarbon dateinto a calendar age corresponds to ∼40,000 y ago. By that time,modern humans producing Aurignacian industries had already oc-cupied the neighboring regions of Austria (11), Germany (12), andNorthern Italy (13) for some time. Interactions with modern groupsmay explain the cultural evolution of the last Neanderthals, but apuzzling issue remains regarding possible biological interactionsbetween the two groups. Introgression of Neanderthal DNA intothe modern genome has been widely demonstrated and analyzedthrough the endless availability of extant genetic data. It is quite likelythat gene flow also occurred in the opposite direction. However, todate not a single detailed nuclear DNA sequence has been recon-structed from one of these postcontact Neanderthal populations andit is therefore impossible to assess the magnitude and possible effectof modern DNA introgression into late Neanderthal populations.As far as archaeological assemblages of Middle Paleolithictype can be considered a good proxy to identify the occurrence ofNeanderthal populations in different parts of Western Europe, thesouth of the Iberian Peninsula has sometimes been proposed asan area of their late survival. Specifically, south of the Ebro river,Neanderthals may have survived several millennia after theirextinction in the rest of Western Europe (14). This idea is heavilydisputed, as the dating of some of the sites on which this modelwas based have been revised to older ages (15). Still, sites in theMula basin of Murcia, Spain, are claimed to document a modernreplacement of Neanderthals taking place as late as around37,000 y ago in calibrated chronology (16). The explanations pro-vided for this delay are primarily geographical and environmental,and partly relate to the expansion of forested environments dur-ing this time period in Iberian regions south of 40°N.If one takes a broader view away from the European continent,a series of recent studies have supported early modern humanexpansion in tropical Asia, as far as China (17), Laos (18), and Indo-nesia (19), and ultimately in Australia (20) more than 60,000 y ago.If substantiated by further discoveries, this situation would set thefar-west Eurasia as a region of delayed replacement for archaiclocal populations. If modern humans were then able to settle inAsian tropical forests and cross large bodies of water, it is difficultto conceive that their expansion in the Iberian Peninsula wouldThe last chapter of the Vindija saga reminds usonce more that the interpretation of materialfrom older excavations is quite challenging, asthe precise archaeological context of discoveriesthat took place several decades ago generallyremains a topic of continuous debate.have been stalled by a river and deciduous temperate forests.Regardless of the “Ebro frontier” issue, it is striking to considerthat the places where the latest survival of Neanderthals is bestdocumented by the direct dating of diagnostic human remainsactually correspond to areas where Neanderthal populationsdisplayed the highest density and a continuous occupation for thetens of millennia preceding their final demise. This suggests thatalthough environmental conditions might have influenced thetempo of modern human penetration in Europe, the decisivefactor that slowed their settlement could have been the very pres-ence of relatively dense Neanderthal populations well adapted tothe Pleistocene environments of the middle latitudes.The last chapter of the Vindija saga reminds us oncemore that theinterpretation of material from older excavations is quite challenging,as the precise archaeological context of discoveries that took placeseveral decades ago generally remains a topic of continuous debate.The resolution of pending issues regarding the European expansionof modern humans will come from the study of new sites or thereassessment of earlier discovered oneswith entirely new approaches.Without relying too much on the hope of discoveries, such asspectacular human remains, major progress can be accomplishedin the near future through the implementation of new methodo-logical approaches ranging from the screening of anatomicallynondiagnostic human bone remains out of faunal assemblages bypeptide mass fingerprinting of collagen to improvements to comein the reduction of sample sizes for direct radiocarbon dating.1 Higham T, Ramsey CB, Karavanić I, Smith FH, Trinkaus E (2006) Revised direct radiocarbon dating of the Vindija G1 Upper Paleolithic Neandertals. Proc Natl AcadSci USA 103:553–557.2 Karavanić I, Smith FH (1998) The Middle/Upper Paleolithic interface and the relationship of Neanderthals and early modern humans in the Hrvatsko Zagorje,Croatia. J Hum Evol 34:223–248.3 Hublin J-J (2015) The modern human colonization of western Eurasia: When and where? Quat Sci Rev 118:194–210.4 Devièse T, et al. (2017) Direct dating of Neanderthal remains from the site of Vindija Cave and implications for the Middle to Upper Paleolithic transition. Proc NatlAcad Sci USA 114:10606–10611.5 Higham T, et al. (2014) The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512:306–309.6 Brown TA, Nelson DE, Vogel JS, Southon JR (1988) Improved collagen extraction by modified Longin method. Radiocarbon 30:171–177.7 Welker F, et al. (2016) Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne. Proc Natl Acad Sci USA113:11162–11167.Hublin PNAS | October 3, 2017 | vol. 114 | no. 40 | 105218 Talamo S, et al. (2016) Direct radiocarbon dating and genetic analyses on the purported Neanderthal mandible from the Monti Lessini (Italy). Sci Rep 6:29144.9 Hublin J-J, et al. (2012) Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian. Proc Natl Acad SciUSA 109:18743–18748.10 Semal P, et al. (2009) New data on the late Neandertals: Direct dating of the Belgian Spy fossils. Am J Phys Anthropol 138:421–428.11 Nigst PR, et al. (2014) Early modern human settlement of Europe north of the Alps occurred 43,500 years ago in a cold steppe-type environment. Proc Natl AcadSci USA 111:14394–14399.12 Higham T, et al. (2012) Τesting models for the beginnings of the Aurignacian and the advent of figurative art and music: The radiocarbon chronology ofGeißenklösterle. J Hum Evol 62:664–676.13 Benazzi S, et al. (2015) Archaeology. The makers of the Protoaurignacian and implications for Neandertal extinction. Science 348:793–796.14 Zilh~ao J (2000) The Ebro Frontier: A model for the late extinction of Iberian neanderthals.Neanderthals on the Edge: Papers from a Conference Marking the 150thAnniversary of the Forbes’ Quarry Discovery, Gibraltar, eds Stringer CB, Barton RN, Finlayson JC (Oxbow Books, Oxford), pp 111–122.15 Wood RE, et al. (2013) Radiocarbon dating casts doubt on the late chronology of the Middle to Upper Palaeolithic transition in southern Iberia. Proc Natl Acad SciUSA 110:2781–2786.16 Zilh~ao J, et al. (2016) Cueva Antón: A multi-proxy MIS 3 to MIS 5a paleoenvironmental record for SE Iberia. Quat Sci Rev 146:251–273.17 Liu W, et al. (2015) The earliest unequivocally modern humans in southern China. Nature 526:696–699.18 Demeter F, et al. (2015) Early modern humans and morphological variation in Southeast Asia: Fossil evidence from Tam Pa Ling, Laos. PLoS One 10:e0121193.19 Westaway KE, et al. (2017) An early modern human presence in Sumatra 73,000-63,000 years ago. Nature 548:322–325.20 Clarkson C, et al. (2017) Human occupation of northern Australia by 65,000 years ago. Nature 547:306–310.10522 | www.pnas.org/cgi/doi/10.1073/pnas.1714533114 Hublin

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The last Neanderthal

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