Cited by 141Forensic and Genetic ResearchArchaeology and ancient environmental studiesArchaeological and Geological Studies

Ancient DNA from Chalcolithic Israel reveals the role of population mixture in cultural transformation

Open access
Harney, Éadaoin, May, Hila, Shalem, Dina, Rohland, Nadin, Mallick, Swapan, Lazaridis, Iosif, Sarig, Rachel, Stewardson, Kristin +4 more·2018·Nature Communications

Abstract

The material culture of the Late Chalcolithic period in the southern Levant (4500–3900/3800 BCE) is qualitatively distinct from previous and subsequent periods. Here, to test the hypothesis that the advent and decline of this culture was influenced by movements of people, we generated genome-wide ancient DNA from 22 individuals from Peqi’in Cave, Israel. These individuals were part of a homogeneous population that can be modeled as deriving ~57% of its ancestry from groups related to those of the local Levant Neolithic, ~17% from groups related to those of the Iran Chalcolithic, and ~26% from groups related to those of the Anatolian Neolithic. The Peqi’in population also appears to have contributed differently to later Bronze Age groups, one of which we show cannot plausibly have descended from the same population as that of Peqi’in Cave. These results provide an example of how population movements propelled cultural changes in the deep past.

AI snapshot

Written from the paper’s full text.

TL;DR

Genome-wide ancient DNA from 22 individuals buried in Peqi’in Cave, Israel, shows a homogeneous Late Chalcolithic population modeled as ~57% Levant Neolithic, ~17% Iran Chalcolithic, and ~26% Anatolian Neolithic ancestry. The results suggest population movement and mixture contributed to Chalcolithic cultural transformation, with later Bronze Age Levantine groups showing different ancestry patterns.

Key findings

  • The 20 unrelated Peqi’in individuals formed a tight PCA cluster and were used as the analysis population; median coverage was 358,313 targeted SNPs covered at least once (range 25,171–1,002,682), with average 0.97× coverage on the 1240k SNP targets. Nine of ten males belonged to Y-chromosome haplogroup T.
  • Levant_ChL was best modeled by qpAdm as a three-way admixture of Levant_N (57%), Iran_ChL (17%), and Anatolia_N (26%); two-way models failed at p > 0.05.
  • Genetic differentiation had already collapsed toward present-day levels by the Chalcolithic: median pairwise FST between Peqi’in and other West Eurasian Neolithic/Chalcolithic populations was 0.023 (range 0.009–0.061), versus 0.098 pre-Neolithic, 0.015 Bronze Age, and 0.011 present-day.
  • Bronze Age Levantine populations differed: Levant_BA_South could not plausibly descend directly from Levant_ChL and was best modeled as Levant_N + Iran_ChL, while Levant_BA_North fit better with Levant_ChL-related ancestry; qpWave required at least three ancestry sources for Levant_N, Levant_ChL, and Levant_BA_South. Allele findings included rs12913832 near OCA2 at 49% G, rs1426654 in SLC24A5 fixed for derived A, and rs6903823 at 20% in Levant_ChL.

Methods

  • The authors screened 48 skeletal remains from Peqi’in Cave, 37 of them petrous bones, extracted ancient DNA, built UDG-treated libraries, enriched for mitochondrial DNA and then ~1.2 million SNPs, and retained 22 individuals passing authenticity and quality criteria.
  • Authenticity filters included ≥3% cytosine-to-thymine errors at the final nucleotide, X-to-Y chromosome ratio consistent with sex, >95% mitochondrial consensus match, male X-chromosome contamination <2%, and ≥5,000 targeted SNPs covered at least once.
  • Analyses included PCA, ADMIXTURE (K = 11), FST and conditional heterozygosity, f3/f4 statistics, qpAdm, qpWave, and allele-frequency comparisons, using newly generated data merged with published datasets (HO+ and HOIll+).

Limitations

  • The paper states that these are currently the only ancient DNA samples available from the Levant in this time period and that future work will plausibly reveal genetic substructure in Chalcolithic samples across the broader region; it also cautions that skin-pigmentation inference from a single site should be viewed with caution and that Y-haplogroup calls for samples with fewer than 100,000 SNPs are uncertain.
  • The paper notes concern that three-way admixture findings might be an artifact of lacking samples closely related to true ancestral populations, and it used qpWave to test this; it also notes that without Levant_ChL data there may be insufficient leverage to detect Anatolian-related ancestry in Levant_BA_North.
  • Evident from the text: the analysis rests on a single cave site and 20 unrelated individuals, so it may not represent all Late Chalcolithic Levantine populations; admixture models depend on the available ancient reference populations and chosen outgroup sets.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1038/s41467-018-05649-9

Cites

59 of 59 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

33 of 63 cited sources checked · 9 supported · 0 flagged · accuracy not yet measured

  • Findingcitation

    ADMIXTURE model-based clustering analyses produced results consistent with PCA in suggesting that individuals from the Levant_ChL population had a greater affinity on average to Iranian agriculturalist-related populations than was the case for earlier Levantine individuals.

    “ADMIXTURE model-based clustering analyses34 produced results consistent with PCA in suggesting that individuals from the Levant_ChL population had a greater affinity on average to Iranian agriculturalist-related populations than was the case for earlier Levantine individuals.”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites [34] Alexander, D. H., Novembre, J. & Lange, K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 19, 1655–1664 (2009). source
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • Premisecitation

    Previously published Bronze Age Levantine samples from 'Ain Ghazal (Levant_BA_South) and Sidon (Levant_BA_North) can be modeled as two-way admixtures, without the Anatolia_N contribution that is required to model the Levant_ChL population.

    “It was striking to us that previously published Bronze Age Levantine samples from the sites of 'Ain Ghazal in present-day Jordan (Levant_BA_South) and Sidon in present-day Lebanon (Levant_BA_North) can be modeled as two-way admixtures, without the Anatolia_N contribution that is required to model the Levant_ChL population24,26.”
    Quote in the paperSubstantial claim2 of 2 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source

      The section on how early farmers mixed reports admixture in southern Levantine Bronze Age populations, and Extended Data Table 2 identifies Levant_BA admixture using Iran_N and Levant_N references. These passages support a two-source model for the southern population, but they do not establish it for Bronze Age samples from Sidon or address the claimed contrast with Levant_ChL and its required Anatolia_N contribution. (LLM, conf=0.85)

      Experimental — accuracy not yet measured
    • cites [26] Haber, M. et al. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am. J. Hum. Genet. 101, 274-282 (2017). source

      The Main Text (Figure 2; page not specified) models Sidon_BA as a two-way mixture of Levant_N and Iran_ChL and reports that the Bronze Age Jordan samples from near ‘Ain Ghazal were modeled the same way. The cited paper does not establish that Anatolia_N is required to model Levant_ChL, so that comparison is unsupported by this source. (LLM, conf=0.86)

      Experimental — accuracy not yet measured
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • Findingown data

    Measured in terms of average coverage at SNPs, the increase in data quality is even more pronounced than in previous studies of ancient Near Easterners.

    “Measured in terms of the average coverage at SNPs, the increase is even more pronounced due to the higher quality of the data reported here than in previous studies of ancient Near Easterners24,26.”
    Quote in the paperSubstantial claim2 of 2 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source
    • cites [26] Haber, M. et al. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am. J. Hum. Genet. 101, 274-282 (2017). source
  • Interpretationcitation

    The placement of the Levant_ChL cluster is consistent with a previously observed pattern whereby chronologically later Levantine populations are shifted towards the Iran Chalcolithic population compared to earlier Levantine populations.

    “The placement of the Levant_ChL cluster is consistent with a previously observed pattern whereby chronologically later Levantine populations are shifted towards the Iran Chalcolithic (Iran_ChL) population compared to earlier Levantine populations, Levant_N (Pre-Pottery and Pottery Neolithic agriculturalists from present-day Israel and Jordan) and Natufians (Epipaleolithic hunter-gatherers from”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • Interpretationown data

    This may explain why Haber et al. did not detect the Anatolian Neolithic-related admixture in Levant_BA_North.

    “This may explain why Haber et al.26 did not detect the Anatolian Neolithic-related admixture in Levant_BA_North.”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Haber et al.26 Haber, M. et al. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am. J. Hum. Genet. 101, 274-282 (2017). source

      The Main Text (Figure 2 and its accompanying discussion; page unavailable) models Sidon_BA as a mixture of Neolithic Levant and Chalcolithic Iran ancestry, rather than reporting Anatolian Neolithic-related admixture. But our copy does not identify Sidon_BA as Levant_BA_North or explicitly report a test that failed to detect that specific admixture in that named group. (LLM, conf=0.76)

      Experimental — accuracy not yet measured
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • Classified as probably not a substantial claim. 20
  • May combine two claims. 20
  • The quote only partly states this claim. 4
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Lazaridis et al. showed that the Levant Bronze Age population from the site of 'Ain Ghazal, Jordan (2490-2300 BCE) could be fit statistically as a mixture of around 56% ancestry from a group related to Levantine Pre-Pottery Neolithic agriculturalists and 44% related to populations of the Iranian Chalcolithic.

    “Lazaridis et al.24 showed that the Levant Bronze Age population from the site of 'Ain Ghazal, Jordan (2490–2300 BCE) could be fit statistically as a mixture of around 56% ancestry from a group related to Levantine Pre-Pottery Neolithic agriculturalists (represented by ancient DNA from Motza, Israel and 'Ain Ghazal, Jordan; 8300–6700 BCE) and 44% related to populations of the Iranian Chalcolithic (”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Lazaridis et al.24 Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source

      Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Previous genome-wide ancient DNA studies from the Near East have revealed that at the time when agriculture developed, populations from Anatolia, Iran, and the Levant were approximately as genetically differentiated from each other as present-day Europeans and East Asians are today.

    “Previous genome-wide ancient DNA studies from the Near East have revealed that at the time when agriculture developed, populations from Anatolia, Iran, and the Levant were approximately as genetically differentiated from each other as present-day Europeans and East Asians are today24,25.”
    Quote in the paperNot a substantial claim2 of 2 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source

      The ‘Extreme regional differentiation in the ancient Near East’ section (page not provided) reports that ancient populations at the four corners of its genetic quadrangle had FST values of 0.08–0.15, comparable to 0.09–0.13 between present-day West Eurasians and Han East Asians. But those four corners include the Levant and Iran alongside two European hunter-gatherer groups—not Anatolia. The passage therefore does not establish the stated comparison for populations from all three named regions relative to each other. (LLM, conf=0.85)

      Experimental — accuracy not yet measured
    • cites [25] Broushaki, F. et al. Early Neolithic genomes from the eastern Fertile Crescent. Science 353, 499–503 (2016). source

      The supplied Abstract passages say Early Neolithic populations from the Zagros and NW Anatolia descended from distinct ancestral populations and differed conspicuously. They do not establish the claim for Levantine populations or compare the degree of differentiation with that between present-day Europeans and East Asians; searches for those comparisons returned no passage. (LLM, conf=0.85)

      Experimental — accuracy not yet measured
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Haber et al. suggested that the Canaanite Levant Bronze Age population from the site of Sidon, Lebanon (~1700 BCE) could be modeled as a mixture of the same two groups albeit in different proportions (48% Levant Neolithic-related and 52% Iran Chalcolithic-related).

    “Haber et al.26 suggested that the Canaanite Levant Bronze Age population from the site of Sidon, Lebanon (~1700 BCE) could be modeled as a mixture of the same two groups albeit in different proportions (48% Levant Neolithic-related and 52% Iran Chalcolithic-related).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Haber et al.26 Haber, M. et al. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am. J. Hum. Genet. 101, 274-282 (2017). source
  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    By the Bronze Age, expansion of different Near Eastern agriculturalist populations-Anatolian, Iranian, and Levantine-in all directions and admixture with each other substantially homogenized populations across the region, thereby contributing to the relatively low genetic differentiation that prevails today.

    “By the Bronze Age, however, expansion of different Near Eastern agriculturalist populations—Anatolian, Iranian, and Levantine—in all directions and admixture with each other substantially homogenized populations across the region, thereby contributing to the relatively low genetic differentiation that prevails today24.”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductcitation

    The study treated libraries with Uracil-DNA glycosylase (UDG) to reduce characteristic ancient DNA damage at all but the first and last nucleotides.

    “We treated the libraries with Uracil–DNA glycosylase (UDG) to reduce characteristic ancient DNA damage at all but the first and last nucleotides29”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [29] Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil–DNA–glycosylase treatment for screening of ancient DNA. Philos. Trans. R. Soc. B 370, 20130624 (2015). source

      The Effectiveness of partial uracil-DNA-glycosylase treatment passage says partial UDG removes uracils within ancient DNA molecules while preserving substantial damage at the terminal bases. But it also says terminal-base damage is reduced compared with untreated libraries; the Library construction passage quantifies that reduction at about threefold. Thus the first and last nucleotides are not wholly exceptions to the reduction described in the claim. The indexed copy provides no page number for these passages. (LLM, conf=0.84)

      Experimental — accuracy not yet measured
  • May combine two claims.

    Study design and conductown data

    Levant_BA_South and Levant_BA_North are treated as separate populations for analysis, since the symmetry statistic f4(Levant_BA_North, Levant_BA_South; A, Chimp) is significant for a number of test populations A.

    “In what follows, we treat Levant_BA_South and Levant_BA_North as separate populations for analysis, since the symmetry statistic f4(Levant_BA_North, Levant_BA_South; A, Chimp) is significant for a number test populations A (|Z| ≥ 3) (Supplementary Data 5), consistent with the different estimated proportions of Levant_N and Iran_ChL ancestry reported in24,26.”
    Quote in the paperSubstantial claim2 of 2 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source

      The section ‘How diverse first farmers of the Near East mixed to form the region's later populations’ estimates that Iran Chalcolithic–related ancestry contributed about 44% to southern Levantine Bronze Age populations. Our copy does not give a corresponding northern estimate or report the stated north–south f4 test. The citing authors present that test and their decision to analyze the populations separately as their own analysis, not as findings of this source. (LLM, conf=0.82)

      Experimental — accuracy not yet measured
    • cites [26] Haber, M. et al. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am. J. Hum. Genet. 101, 274-282 (2017). source

      The Main Text (page not provided) estimates Levant_N and Iran_ChL ancestry proportions for Sidon_BA and says Jordan_BA can also be modeled as a mixture of those ancestries. It does not report a north–south comparison of the proportions or the cited f4 symmetry test. The decision to analyze the populations separately and that test are presented as the citing authors’ own analysis, not as findings of this source. (LLM, conf=0.78)

      Experimental — accuracy not yet measured
  • May combine two claims.

    Premisecitation

    Direct radiocarbon dating suggests that the cave was in use throughout the Late Chalcolithic (4500-3900 BCE), functioning as a central burial location for the region.

    “Direct radiocarbon dating suggests that the cave was in use throughout the Late Chalcolithic (4500–3900 BCE), functioning as a central burial location for the region21,23.”
    Quote in the paperNot a substantial claim1 of 2 sources checked
    • cites [21] Shalem, D. et al. Peqiʻin: A Late Chalcolithic Burial Site Upper Galilee, Israel (Kinneret Academic College Institute for Galilean Archaeology, 2013).

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [23] Segal, D., Carmi, I., Gal, Z., Smithline, H. & Shalem, D. Dating a Chalcolithic burial cave in Peqi’in, upper Galilee, Israel. Radiocarbon 40, 707–712 (1998). source

      The Discussion says radiocarbon dates confirm cave use over several Chalcolithic centuries and that the cave served as a burial center. It does not establish continuous burial use throughout the specified 4500–3900 BCE span, and its account distinguishes earlier, pre-burial use from the burial phase. Page numbers are unavailable in our copy. (LLM, conf=0.84)

      Experimental — accuracy not yet measured
  • May combine two claims. The quote only partly states this claim.

    Study design and conductcitation

    The study evaluated each individual for evidence of authentic ancient DNA by limiting to libraries with a minimum of 3% cytosine-to-thymine errors at the final nucleotide, requiring X-to-Y-chromosome ratio characteristic of sex, >95% matching to mtDNA consensus, and <2% contamination for males.

    “We evaluated each individual for evidence of authentic ancient DNA by limiting to libraries with a minimum of 3% cytosine-to-thymine errors at the final nucleotide29, by requiring that the ratio of X-to-Y-chromosome sequences was characteristic of either a male or a female, by requiring >95% matching to the consensus sequence of mitochondrial DNA30, and by requiring (for males) a lack of variation”
    Quote in the paperSubstantial claim3 of 3 sources checked
    • cites [29] Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil–DNA–glycosylase treatment for screening of ancient DNA. Philos. Trans. R. Soc. B 370, 20130624 (2015). source
    • cites [30] Fu, Q. et al. DNA analysis of an early modern human from Tianyuan Cave, China. Proc. Natl Acad. Sci. 110, 2223-2227 (2013). source

      The citation is attached to the cytosine-to-thymine error criterion. In Results and Discussion (page unavailable), the paper uses end-of-fragment substitutions as evidence of ancient DNA and reports frequencies of 25–30% close to fragment ends. It does not specify a minimum of 3% at the final nucleotide as a library-selection rule. (LLM, conf=0.84)

      Experimental — accuracy not yet measured
    • cites [32] Korneliussen, T. S., Albrechtsen, A. & Nielsen, R. ANGSD: analysis of next generation sequencing data. BMC Bioinforma. 15, 356 (2014). source

      The citation is attached to the minimum 3% terminal cytosine-to-thymine error criterion. The ABBA-BABA section says transitions have high error rates in ancient genomes, but our copy does not give a 3% threshold or describe terminal-base errors as an authenticity test. Searches for those criteria found no supporting passage. (LLM, conf=0.74)

      Experimental — accuracy not yet measured
  • May combine two claims.

    Premisecitation

    Previously reported median pairwise FST between populations in pre-Neolithic periods is 0.098 (range: 0.023-0.153), in Bronze Age periods 0.015 (range: 0.002-0.045), and in present-day West Eurasian populations 0.011 (range: 0-0.046).

    “relative to a previously reported median pairwise FST of 0.098 (range: 0.023–0.153) observed between populations in pre-Neolithic periods, 0.015 (range: 0.002–0.045) in the Bronze Age periods, and 0.011 (range: 0–0.046) in present-day West Eurasian populations24.”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source

      The section “Extreme regional differentiation in the ancient Near East” reports high FST among early West Eurasian populations and says differentiation had fallen to present-day low levels by the Bronze Age. It also gives 0.046 as the upper bound for present-day West Eurasian pairs. Our copy does not report the three claimed medians or the full ranges for the pre-Neolithic and Bronze Age periods. (LLM, conf=0.86)

      Experimental — accuracy not yet measured
  • The quote only partly states this claim.

    Premisecitation

    Later Bronze Age individuals all belonged to haplogroup J.

    “and later Bronze Age individuals, all of whom belonged to haplogroup J24,26.”
    Quote in the paperNot a substantial claim2 of 2 sources checked
    • cites [24] Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419 (2016). source

      The Abstract says the sample includes Bronze Age individuals, but does not give their haplogroups. The section on Basal Eurasian ancestry identifies haplogroup E in earlier Natufian and Levantine Neolithic populations; searches for Bronze Age haplogroup J and Y-chromosome lineages found no passage establishing that all later Bronze Age individuals belonged to J. (LLM, conf=0.72)

      Experimental — accuracy not yet measured
    • cites [26] Haber, M. et al. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am. J. Hum. Genet. 101, 274-282 (2017). source

      The Main Text and Table 1 (page not specified) report that both male Sidon Bronze Age individuals had Y-chromosome haplogroup J. The claim's 'all' is broader than this evidence: three other Sidon individuals were female and had no Y-haplogroup assignment, and the paper does not establish that all later Bronze Age individuals belonged to J. (LLM, conf=0.88)

      Experimental — accuracy not yet measured
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The allele (G) at rs6903823 has been argued to have been under positive selection by Mathieson et al.

    “that has been argued to have been under positive selection by Mathieson et al.31”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [31] Mathieson, I. et al. Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528, 499–503 (2015). source

      The ‘Twelve signals of selection’ section discusses selection at several named SNPs and loci, but the passages in our copy do not identify rs6903823 or its G allele as a target. Searches for rs6903823 and for ‘rs6903823 G positive selection’ found no passage addressing that specific allele. (LLM, conf=0.72)

      Experimental — accuracy not yet measured
  • May combine two claims.

    Premisecitation

    The archaeological record is marked by dramatic changes in settlement patterns, large-scale abandonment of sites, many fewer items with symbolic meaning, and shifts in burial practices, including the disappearance of secondary burial in ossuaries.

    “aspects of the archeological record marked by dramatic changes in settlement patterns43, large-scale abandonment of sites52–55, many fewer items with symbolic meaning, and shifts in burial practices, including the disappearance of secondary burial in ossuaries56–59.”
    Quote in the paperSubstantial claim1 of 9 sources checked
    • cites [43] Rowan, Y. M. & Golden, J. The Chalcolithic period of the Southern Levant: a synthetic review. J. World Prehistory 22, 1–92 (2009). source
    • cites [52] Gilead, I. The history of the Chalcolithic settlement in the Nahal Beer Sheva area: the radiocarbon aspect. Bull. Am. Sch. Orient. Res. 296, 1–13 (1994). source
    • cites [53] Bourke, S. et al. The chronology of the Ghassulian Chalcolithic period in the southern Levant: new 14 C determinations from Teleilat Ghassul, Jordan. Radiocarbon 43, 1217–1222 (2001). source

      Split into 2 atoms — 0 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 2 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured
    • cites [54] Bourke, S. J. & Lovell, J. L. Ghassul, chronology and cultural sequencing. Paléorient 30, 179–182 (2004). source
    • cites [55] Gilead, I. in Culture, Chronology and the Chalcolithic: Theory and Transition (eds Lovell, J. L. & Rowan, Y. M.) 12–24 (Oxbow Books, 2011).

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [56] van den Brink Edwin, C. in Culture, Chronology and the Chalcolithic: Theory and Transition(eds Lovell, J. L. & Rowan, Y. M.) 61–70 (Oxbow Books, 2011).

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites [57] Vardi, J. & Gilead, I. Chalcolithic early bronze age I transition in the Southern Levant: the Lithic perspective. Paléorient 39, 111–123 (2013). source
    • cites [58] Milevski, I. The transition from the Chalcolithic to the early bronze age in the Southern Levant in socio-economic context. Paléorient 39, 193–208 (2013). source
    • cites [59] Braun, E. & Roux, V. The late Chalcolithic to early bronze age I transition in the Southern Levant: determining continuity and discontinuity or “Mind the Gap”. Paléorient 29, 15–22 (2013). source
  • Classified as probably not a substantial claim. The quote only partly states this claim.

    Study design and conductcitation

    We assessed the quality of each library at the screening stage using three standard methods for determining ancient DNA authenticity: mitochondrial genome matching rate, C-to-T substitution rate, and X-chromosome contamination.

    “We assessed the quality of each library at the screening stage using three standard methods for determining ancient DNA authenticity.”
    Quote in the paperSubstantial claim3 of 3 sources checked
    • cites [30] Fu, Q. et al. DNA analysis of an early modern human from Tianyuan Cave, China. Proc. Natl Acad. Sci. 110, 2223-2227 (2013). source

      The Results and Discussion section (“DNA Sequence Authenticity”; page unavailable) uses mitochondrial sequence agreement and C→T substitutions to assess whether DNA in the Tianyuan libraries is ancient and largely from one individual. Our copy does not describe X-chromosome contamination as a third method or say that these three methods were used to assess each library at screening. (LLM, conf=0.87)

      Experimental — accuracy not yet measured
    • cites [29] Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil–DNA–glycosylase treatment for screening of ancient DNA. Philos. Trans. R. Soc. B 370, 20130624 (2015). source

      The Introduction describes C-to-T mismatches and mitochondrial consensus mismatches as evidence of ancient DNA authenticity. The sequencing and analysis section also reports mitochondrial contamination estimates, but our copy does not describe X-chromosome contamination as a screening method. It therefore supports some, but not all, of the three methods attributed to it. (LLM, conf=0.86)

      Experimental — accuracy not yet measured
    • cites [32] Korneliussen, T. S., Albrechtsen, A. & Nielsen, R. ANGSD: analysis of next generation sequencing data. BMC Bioinforma. 15, 356 (2014). source

      The Results and discussion > ABBA-BABA section (page unavailable) describes filtering error-prone transitions when analyzing an ancient Denisovan genome, not screening libraries for authenticity. Searches for mitochondrial matching rates, C-to-T substitution rates, and X-chromosome contamination found no passage describing the three claimed methods. (LLM, conf=0.77)

      Experimental — accuracy not yet measured
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductcitation

    We excluded two individuals from further analysis because genetic patterns showed they were first-degree relatives of higher coverage samples in the dataset.

    “We excluded two individuals from further analysis, as the genetic patterns observed using the method described in Kuhn et al.73. Showed that they were first-degree relatives of higher coverage samples in the dataset.”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [73] Kuhn, J. M. M., Jakobsson, M. & Günther, T. Estimating genetic kin relationships in prehistoric populations. PLoS ONE 13, e0195491 (2018). source
  • Classified as probably not a substantial claim.

    Study design and conductcitation

    We excluded libraries with X-contamination estimates greater than 1.5%.

    “we excluded libraries with X-contamination estimates greater than 1.5%32.”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites [32] Korneliussen, T. S., Albrechtsen, A. & Nielsen, R. ANGSD: analysis of next generation sequencing data. BMC Bioinforma. 15, 356 (2014). source
  • May combine two claims.

    Findingreasoning

    The dataset is of exceptional quality given the typically poor preservation of DNA in the warm Near East, with a higher proportion of samples yielding appreciable coverage of ancient DNA than has previously been obtained from the region.

    “The dataset is of exceptional quality given the typically poor preservation of DNA in the warm Near East, with a higher proportion of samples yielding appreciable coverage of ancient DNA than has previously been obtained from the region, likely reflecting the optimal sampling techniques we used and the favorable preservation conditions at the cave.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    The samples from Peqi'in Cave form a tight cluster in PCA, supporting the grouping of these individuals into a single analysis population.

    “In a plot of the first and second principal components (Fig. 3a), the samples from Peqi’in Cave form a tight cluster, supporting the grouping of these individuals into a single analysis population”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • The quote only partly states this claim.

    Findingown data

    Only the model involving Iran_ChL remained plausible after adding Iran_N to the outgroup set.

    “Only the model involving Iran_ChL remained plausible.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • May combine two claims.

    Findingown data

    The Levant_ChL cluster overlaps in the PCA with a cluster containing Neolithic Levantine samples, although it is slightly shifted upward on the plot toward a cluster corresponding to samples from the Levant Bronze Age.

    “The Levant_ChL cluster overlaps in the PCA with a cluster containing Neolithic Levantine samples (Levant_N), although it is slightly shifted upward on the plot toward a cluster corresponding to samples from the Levant Bronze Age, including samples from 'Ain Ghazal, Jordan (Levant_BA_South) and Sidon, Lebanon (Levant_BA_North).”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingreasoning

    The quality of ancient DNA obtained from the Peqi'in Cave samples is excellent relative to other sites in the Near East.

    “The quality of ancient DNA obtained from the Peqi’in Cave samples is excellent relative to other sites in the Near East.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Findingown data

    The Levant_BA_South can be modeled as a mixture of Levant_N (58%) and Iran_ChL (42%), but has little if any additional Anatolia_N-related ancestry.

    “while the Levant_BA_South can be modeled as a mixture of Levant_N (58%) and Iran_ChL (42%), but has little if any additional Anatolia_N-related ancestry”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    The allele (G) at rs6903823 occurs with an estimated frequency of 20% in the Levant_ChL, 17% in the Levant_BA_South, and 15% in the Iran_ChL populations, while it is absent in all other populations.

    “occurs with an estimated frequency of 20% in the Levant_ChL, 17% in the Levant_BA_South, and 15% in the Iran_ChL populations, while it is absent in all other populations.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Findingown data

    The 22 samples that passed contamination and quality control tests have an average of 0.97× coverage on the 1240 k SNP targets and an average of 358,313 SNPs covered at least once.

    “The data from the 22 samples that passed contamination and quality control tests are reported in Supplementary Table 1, with an average of 0.97× coverage on the 1240 k SNP targets, and an average of 358,313 SNPs covered at least once.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationown data

    Both the increasing genetic diversity over time and the reduced differentiation between populations as measured via FST are consistent with a model in which gene flow reduced differentiation across groups while increasing diversity within groups.

    “Both the increasing genetic diversity over time, and the reduced differentiation between populations as measured via FST, are consistent with a model in which gene flow reduced differentiation across groups while increasing diversity within groups.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • May combine two claims.

    Interpretationreasoning

    A population that harbored ancestry more closely related to Levant_ChL than to Levant_N contributed to the Levant_BA_North population, even if it did not contribute detectably to the Levant_BA_South population.

    “These results imply that a population that harbored ancestry more closely related to Levant_ChL than to Levant_N contributed to the Levant_BA_North population, even if it did not contribute detectably to the Levant_BA_South population.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • May combine two claims.

    Interpretationreasoning

    The Levant_ChL population may not be directly ancestral to these later Bronze Age Levantine populations, because if it were, we would also expect to detect an Anatolia_N component of ancestry.

    “This suggests that the Levant_ChL population may not be directly ancestral to these later Bronze Age Levantine populations, because if it were, we would also expect to detect an Anatolia_N component of ancestry.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • May combine two claims.

    Interpretationown data

    The Levant_ChL population is descended from a population related to Levant_N, but also harbors ancestry from non-Levantine populations related to those of Iran or the Caucasus that Levant_N does not share.

    “This suggests that the Levant_ChL population is descended from a population related to Levant_N, but also harbors ancestry from non-Levantine populations related to those of Iran or the Caucasus that Levant_N does not share (or at least share to the same extent).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    The study attached unique barcodes to libraries to minimize the possibility of contamination.

    “We extracted DNA28 and built next-generation sequencing libraries to which we attached unique barcodes to minimize the possibility of contamination.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [28] Dabney, J. et al. Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments. Proc. Natl Acad. Sci. 110, 15758-15763 (2013). source
  • May combine two claims.

    Study design and conductown data

    A total of 20 Peqi'in samples appear to be unrelated to each other to the limits of our resolution, and we used these as our analysis set.

    “A total of 20 Peqi’in samples appear to be unrelated to each other to the limits of our resolution (that is, genetic analysis suggested that they were not first, second, or third degree relatives of each other), and we used these as our analysis set.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductnone given

    The study restricted to individuals with at least 5000 of the targeted SNPs covered at least once.

    “We also restricted to individuals with at least 5000 of the targeted SNPs covered at least once.”
    Quote in the paperNot a substantial claim
  • Classified as probably not a substantial claim.

    Premisecitation

    An allele at rs1426654 in the SLC24A5 gene is one of the most important determinants of light pigmentation in West Eurasians.

    “an allele at rs1426654 in the SLC24A5 gene which is one of the most important determinants of light pigmentation in West Eurasians41”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites [41] Soejima, M. & Koda, Y. Population differences of two coding SNPs in pigmentation-related genes SLC24A5 and SLC45A2. Int. J. Leg. Med. 121, 36–39 (2007). source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The Chalcolithic period in the Levant witnessed major cultural transformations in virtually all areas of culture, including craft production, mortuary and ritual practices, settlement patterns, and iconographic and symbolic expression.

    “The Chalcolithic period in the Levant witnessed major cultural transformations in virtually all areas of culture, including craft production, mortuary and ritual practices, settlement patterns, and iconographic and symbolic expression43.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [43] Rowan, Y. M. & Golden, J. The Chalcolithic period of the Southern Levant: a synthetic review. J. World Prehistory 22, 1–92 (2009). source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Y-chromosome haplogroup T is a lineage thought to have diversified in the Near East.

    “a lineage thought to have diversified in the Near East46”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites [46] Mendez, F. L. et al. Increased resolution of Y-chromosome haplogroup T defines relationships among populations of the Near East, Europe, and Africa. Hum. Biol. 83, 39–53 (2011). source
  • May combine two claims.

    Interpretationreasoning

    The Levant_BA_South population may represent a remnant of a population that formed after an initial spread of Iran_ChL-related ancestry into the Levant that was not affected by the spread of an Anatolia_N-related population, or perhaps a reintroduction of a population without Anatolia_N-related ancestry to the region.

    “The Levant_BA_South population may thus represent a remnant of a population that formed after an initial spread of Iran_ChL-related ancestry into the Levant that was not affected by the spread of an Anatolia_N-related population, or perhaps a reintroduction of a population without Anatolia_N-related ancestry to the region.”
    Quote in the paperSubstantial claim
  • Classified as probably not a substantial claim. May combine two claims.

    Study design and conductown data

    We used qpAdm with additional outgroups and repeated analysis until all but one admixture model was eliminated.

    “we repeated qpAdm with each additional outgroup added into the “Right” list until all but one admixture model was eliminated.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductnone given

    We performed ADMIXTURE analysis with 20 replicate analyses for each k between 2 and 14 and retained the highest likelihood replicate.

    “For each value of k between 2 and 14, we performed 20 replicate analyses, and we retained the highest likelihood replicate for each k.”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    We restricted data from sample I1183 to include only sequences with evidence of C-to-T substitution in order to minimize contamination.

    “We restricted data from sample I1183 to include only sequences with evidence of C-to-T substitution in order to minimize contamination which was evident in the full data from these samples.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationown data

    The movements of people within the region of the southern Levant were remarkably dynamic, with some populations, such as the one buried at Peqi'in, being formed in part by exogenous influences.

    “We show that the movements of people within the region of the southern Levant were remarkably dynamic, with some populations, such as the one buried at Peqi’in, being formed in part by exogenous influences.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductnone given

    Samples with fewer than 100,000 SNPs have too little data to confidently identify the correct Y-chromosome haplogroup.

    “we note that samples with fewer than 100,000 SNPs have too little data to confidently identify the correct haplogroup”
    Quote in the paperNot a substantial claim
39 of 39
  • The paper relies on this without support.

    The Peqi’in Cave individuals are representative of the Late Chalcolithic population of the southern Levant.

    “while we use the broad name “Levant_ChL” to refer to these samples, we recognize that they are currently the only ancient DNA available from the Levant in this time period and future work will plausibly reveal genetic substructure in Chalcolithic samples over the broad region”
  • The paper relies on this without support.

    The archaeological finds at Peqi’in Cave share distinctive characteristics with other Chalcolithic sites, both to the north and south, including secondary burial in ossuaries with iconographic and geometric designs.

    “The archaeological finds at Peqi’in Cave share distinctive characteristics with other Chalcolithic sites, both to the north and south, including secondary burial in ossuaries with iconographic and geometric designs.”
2 of 2

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

  • The ancestry of Chalcolithic individuals from Peqi'in cave included an additional component related to earlier Anatolian farmers, which was excluded as a substantial source for later Bronze Age groups from the Southern Levant, except for coastal groups from Sidon and Ashkelon.

    “A study of Chalcolithic individuals from Peqi’in cave in the Galilee (present-day Israel) showed that the ancestry of this earlier group included an additional component related to earlier Anatolian farmers, which was excluded as a substantial source for later Bronze Age groups from the Southern Levant, with the exception of the coastal groups from Sidon and Ashkelon (Feldman et al., 2019; Harney”

    From The Genomic History of the Bronze Age Southern Levant (2020) · cites this paper as Harney et al., 2018

    Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.

    Experimental — accuracy not yet measured
  • The genetic distinctiveness of the Sidon individuals is compatible with previous findings that Chalcolithic Levantine individuals from Peqi'in Cave are consistent with contributing some ancestry to the Sidon individuals, but not to the 'Ain Ghazal ones.

    “The genetic distinctiveness of the Sidon individuals is also compatible with previous findings that Chalcolithic Levantine individuals from Peqi’in Cave are consistent with contributing some ancestry to the Sidon individuals, but not to the ‘Ain Ghazal ones (Harney et al.,”

    From The Genomic History of the Bronze Age Southern Levant (2020) · cites this paper as Harney et al., 2018

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

  • A Bronze Age South model does not remain plausible under all listed conditions

    Minor

    The text says the Levant_N and Iran_ChL model remains plausible under all conditions, but Table 2 reports p = 0.0139 for that model with the 09NWAZ outgroups, below the paper's p > 0.05 plausibility threshold.

1 of 1

Cites

Flagged first, then by citations

Research works printed but not linked: 7

  1. Shalem, D. Iconography on Ossuaries and Burial Jars from the Late Chalcolithic Period in Israel in the Context of the Ancient Near East.PhD dissertation, Haifa Univ. (2008).
  2. Tadmor, M. et al. The Nahal Mishmar hoard from the Judean desert: technology, composition, and provenance. Atiqot 27, 95–148 (1995).
  3. Levy, T. E. in The Archaeology of Society in the Holy Land (ed. Levy, T. E.) 226–244 (Leicester University Press, 1995).
  4. Merhav, R., Heltzer, M., Segal, A. & Kaufman, D. in Studies in the Archaeology and History of Ancient Israel in Honour of Moshe Dothan (eds. Heltzer, M., Segal, A. & Kaufman, D.) 21–42 (Haifa University Press, 1993).
  5. Bar-Yosef, O. & Ayalon, E. Chalcolithic ossuaries: what do they imitate and why? Qadmoniot 34, 34–43 (2001).
  6. Yadin, Y. The earliest record of Egypt’s military penetration into Asia? Some aspects of the Narmer Palette, the “Desert Kites” and mesopotamian seal cylinders. Isr. Explor. J. 5, 1–16 (1955).
  7. Yeivin, S. Early contacts between Canaan and Egypt. Isr. Explor. J. 10, 193–203 (1960).

Cited by

Sorted by citations

Authors
Harney, É., May, H., Shalem, D., Rohland, N., Mallick, S., Lazaridis, I., Sarig, R., Stewardson, K., Nordenfelt, S., Patterson, N., Hershkovitz, I., & Reich, D.
Affiliations
  • Harvard University (US)
  • Tel Aviv University (IL)
  • Kinneret College on the Sea of Galilee (IL)
  • Broad Institute (US)
  • Howard Hughes Medical Institute (US)
Corresponding author
  • Harney, Éadaoin
  • May, Hila
Venue
Nature Communications
Published
2018-08-20
Volume
9
Issue
1
Pages
3336-3336
ISSN
2041-1723 (ISSN-L)
OpenAlex
W2887731084
Cited by
141
References
59
Citations by year
2026: 7 · 2025: 52 · 2024: 18 · 2023: 11 · 2022: 16 · 2021: 11 · 2020: 14 · 2019: 11 · 2018: 1
FWCI
5.15
Type
journal article
Language
en
Open access
Locations
  • Nature Communications (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • Europe PMC (PubMed Central) (submittedVersion, cc-by)
  • PubMed Central (submittedVersion, cc-by)
  • RePEc: Research Papers in Economics (submittedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
Funders
  • National Science Foundation (1032255)
  • Howard Hughes Medical Institute
  • Dan David Prize
  • National Institutes of Health (GM100233)
  • National Institute of General Medical Sciences (R01 GM100233)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Forensic and Genetic Research
  • Physical Sciences › Earth and Planetary Sciences › Paleontology › Archaeology and ancient environmental studies
  • Social Sciences › Arts and Humanities › Archeology › Archaeological and Geological Studies
Keywords
Late Chalcolithic, ancient DNA, human mobility, cultural change, Southern Levant, Anatolian Neolithic, prehistoric population genetics
MeSH terms
DNA, Ancient, Culture, Genetics, Population, Humans, Israel, Mutation, Genetic Variation, Genetic Heterogeneity, Geologic Sediments, Caves

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-10.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-10.

Cited sources are checked by gpt-6-sol.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

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