Research ArticleANTHROPOLOGY

Pre-Clovis occupation of the Americas identified by human fecal biomarkers in coprolites from Paisley Caves, Oregon

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Science Advances  15 Jul 2020:
Vol. 6, no. 29, eaba6404
DOI: 10.1126/sciadv.aba6404
  • Fig. 1 The dominant fecal sterol and bile acids in fecal material of humans, canids, pigs, and bovids.

  • Fig. 2 Site location map.

    (A) Location of Paisley Caves in the western Great basin. (B) Spatial location of coprolites in cave 5. (C) Stratigraphic profiles in cave 5.

  • Fig. 3 Summary of the fecal lipid biomarker and bile acid profiles of the coprolites analyzed, their species identified by DNA (light gray), their radiocarbon age, and determination of species (dark gray).

    Samples with multiple radiocarbon dates are combined using Oxcal 4.3 “R_combine” command.

  • Fig. 4 Partial gas chromatograms illustrating the distribution of steroid compounds in coprolites identified as human or carnivore.

    (A) human (S226) and (B) carnivore (S249), where ● denotes n-alcohols of carbon chain length C20 to C32, and their corresponding bile acids (C) human (S226) and (D) carnivore (S249), where ▲ denotes diacids (methylated), ■ denotes hydroxy fatty acid methyl esters (TMS derivatives), and ⧫ denotes ω-hydroxy fatty acid methyl esters (TMS derivatives). IS denotes added internal standards: Preg-5en-3β-ol is the sterol standard, and hyocholic acid is the standard for bile acids.

  • Table 1 Classifications of coprolites and sediment sample based on the relative distributions of fecal biomarkers.

    LCA, lithocholic acid; DCA, deoxycholic acid; CDCA, chenodeoxycholic acid; CA, cholic acid; UDCA, ursodeoxycholic acid; 7-keto-LCA, 7-keto-lithocholic acid; 12-keto-LCA, 12-keto-lithocholic acid; n/a, not applicable; nd, not determined. Bile acids are presented in the order of abundance.

    Sample
    ID
    Paisley
    catalog
    number
    Published aDNA
    results (3, 5)
    Ratio 2
    (12)
    Ratio 3
    (12)
    Coprostanol/
    cholesterol
    Bile acidsLipid
    biomarker
    results
    Age 14C yr B.P.
    1651294-PC-5/7D-
    4-3a
    H. sapiens mtDNA
    haplogroup B2,
    Canis latrans
    0.92.53.0DCA, 12-keto-LCA,
    7-keto-LCA, LCA, CA
    Human1308 ± 28 (OxA-16377)
    2231704-PC-
    5/12C-13-6a
    H. sapiens mtDNA
    haplogroup A
    0.12.70DCA, 12-keto-LCA
    LCA, CA
    Carnivore4950 ± 15 (UCIAMS-79710)
    2161704-PC-
    5/12C-13-4a
    L. rufus0.11.50DCA, CA, 12-keto-LCA
    CDCA, LCA
    Carnivore5380 ± 15 (UCIAMS-79715)
    2281704-PC-
    5/12C-12-6a
    H. sapiens mtDNA
    haplogroup A
    0.551.1DCA, 12-keto-LCA, 7-keto-LCA,
    LCA, CA.
    Human5595 ± 15 (UCIAMS-79702);
    5655 ± 15 (UCIAMS-79703)
    2261704-PC-
    5/12C-14-6a
    H. sapiens mtDNA
    haplogroup A
    0.713.64.0DCA, 12-keto-LCA,
    7-keto-LCA, LCA, CA
    Human5715 ± 15 (UCIAMS-76186)
    2241704-PC-
    5/12C-15-5a
    H. sapiens mtDNA
    haplogroup B
    0.100DCA, CA, 12-keto-LCA,
    LCA, CDCA
    Carnivore6115 ± 15 (UCIAMS-76185)
    2271704-PC-
    5/12C-14-9a
    H. sapiens mtDNA
    haplogroup A
    0.71.81.5DCA, 12-keto-LCA,
    LCA, CA, UDCA
    Human6970 ± 15 (UCIAMS-76180)
    2511830-PC-
    2/4D-33-2
    H. sapiens mtDNA
    haplogroup A
    0.200.2DCA, 12-keto-LCA, CA,
    7-keto-LCA, LCA
    Carnivore7025 ± 15 (UCIAMS-79713)
    2491830-PC-
    2/4C-34-101
    H. sapiens mtDNA
    haplogroup A
    0.23.80.2DCA, CA, 12-keto-LCA,
    CDCA, LCA,
    7-keto-LCA,
    Carnivore7490 ± 20 (UCIAMS-79704);
    7605 ± 20 (UCIAMS-79705)
    2501830-PC-
    2/4-D-33
    H. sapiens mtDNA
    haplogroup A
    0.91.412.4DCA, CA, LCA, 12-keto-
    LCA, 7-keto-LCA
    Human7645 ± 20 (UCIAMS-79712)
    2171704-PC-
    5/12A-16-9a
    H. sapiens mtDNA
    haplogroup A
    0.41.4DCA, 12-keto-LCA,
    LCA, CA, 7-keto-LCA,
    CDCA
    Human9170 ± 20 (UCIAMS-76183)
    2151704-PC-
    5/12A-16-10a
    H. sapiens mtDNA
    haplogroup A
    0.713.411.2CA, DCA, 12-keto-LCA
    LCA
    Human9585 ± 20 (UCIAMS-76181)
    1951294-PC-5/6B-
    40-6a
    H. sapiens mtDNA
    haplogroup B2,
    C. lupus/familiaris
    0.51.51.0CDCA, DCA, CA, LCAHuman/
    carnivore*
    10,050 ± 50 (β-213423);
    10,965 ± 50 (OxA-16376)
    2821896-PC-
    5/16A-24-7
    H. sapiens mtDNA
    haplogroup A
    0.71.06.4DCA, LCAHuman11,205 ± 25 (UCIAMS-90583);
    11,250 ± 25 (UCIAMS-90584)
    2831896-PC-
    5/16A-25-12a
    Camelidae0.22.4DCAn/a11,315 ± 25 (UCIAMS-90586);
    12,125 ± 30 (UCIAMS-90589)
    2801830-PC-
    5/11B-33
    H. sapiens mtDNA
    haplogroup A
    0.6nd6.9DCA, CA, LCA, 12-keto-
    LCA, 7-keto-LCA,
    CDCA
    Human12,050 ± 25 (UCIAMS-79707);
    12,165 ± 25 (UCIAMS-79706)
    1941294-PC-5/6B-
    50-5a
    H. sapiens mtDNA
    haplogroup A2,
    V. vulpes
    0.5nd32.2DCA, LCA, 12-keto-
    LCA, CA
    Human12,140 ± 70 (OxA-16495);
    12,260 ± 60 (β-216474)
    2711830-PC-
    5/11B-31-12
    P. leo0.31.40.2DCA, CDCACarnivoreFrom same context as
    coprolite 281.
    2811830-PC-
    5/11B-31-2
    H. sapiens mtDNA
    haplogroup B
    0.45.70.5DCA, CA, CDCA,
    12-keto-LCA, LCA
    Carnivore12,260 ± 30 (UCIAMS-76190);
    12,265 ± 25 (UCIAMS-76190)
    1851294-PC-5/7C-
    31-9a
    H. sapiens mtDNA
    haplogroup B2
    0.10.10.1DCAn/a12,290 ± 60 (β-213426);
    12,345 ± 55 (OxA-16497)
    1821374-PC-5/5D-
    31-2a5
    H. sapiens
    mtDNA
    haplogroup B?
    0.2ndndCarnivore12,275 ± 55 (OxA-16498);
    12,400 ± 60 (β-213424)

    *Possible mixing of fecal matter due to high proportion of CDCA in bile acids.

    †Several coprolites are associated with the same catalog number, and we cannot be 100% certain that we analyzed the same coprolite that was analyzed for mtDNA.

    ‡Potentially carnivore (unable to confirm due to lack of bile acids).

    Supplementary Materials

    • Supplementary Materials

      Pre-Clovis occupation of the Americas identified by human fecal biomarkers in coprolites from Paisley Caves, Oregon

      Lisa-Marie Shillito, Helen L. Whelton, John C. Blong, Dennis L. Jenkins, Thomas J. Connolly, Ian D. Bull

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      The PDF file includes:

      • Figs. S1 to S5
      • Tables S1 and S2
      • Legend for data S1

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