Harvard University
Ancient DNA Analysis of St. Mary’s City Lead Coffin Burials October 23 2016 working paperDavid Reich1,2,3, Kristin Stewardson1,2, Iosif Lazaridis1, Swapan Mallick1,2,3, Nadin Rohland1 and Douglas Owsley41 Department of Genetics, Harvard Medical School, Boston, MA 02115, USA2 Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA3 Broad Institute of MIT and Harvard, Cambridge, MA 02142, USA4 Department of Anthropology, National Museum of Natural History, Smithsonian Institution, Washington DC 20560, USATo whom correspondence and request for data should be addressed: David Reich (reich@genetics.med.harvard.edu) and Douglas Owsley (owsleyd@si.edu)OverviewThe goal of this report is to document the screening for ancient DNA of bone samples from each of three Lead Coffin Burials from Historic St. Mary’s City.MethodsThe screening of these samples was based on a modified version of protocols previously used in several published papers from the Reich laboratory at Harvard Medical School.PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5IYWFrPC9BdXRob3I+PFllYXI+MjAxNTwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA 1-4 Table 1 shows the results. Briefly:A Dremel tool with a sterile disposable drill bit was used to take 43-82 mg of powder from each sample after cleaning the surface with a disposable sanding disk.DNA from the powder was extracted for each sample.PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5EYWJuZXk8L0F1dGhvcj48WWVhcj4yMDEzPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 530uL or 3uL of DNA extract was converted into barcoded non-UDG-treated Illumina sequencing libraries, a total of five libraries for the three samples.PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Sb2hsYW5kPC9BdXRob3I+PFllYXI+MjAxNTwvWWVhcj48
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ADDIN EN.CITE.DATA 6 Two of the libraries passed standard wet laboratory quality control (the others had evidence of inhibition).The two libraries passing quality control were hybridized to oligonucleotide probes targeting the mitochondrial genome.PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NZXllcjwvQXV0aG9yPjxZZWFyPjIwMTQ8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 1,7 The enriched libraries were sequenced on an Illumina NextSeq 500 instrument using 2×76?bp reads. Identifying barcodes and adapters were trimmed, read pairs with at least 15 base pairs of overlapping sequences were merged, and the merged sequences were mapped to the RSRS mitochondrial DNA reference genome, ADDIN EN.CITE <EndNote><Cite><Author>Behar</Author><Year>2012</Year><RecNum>10064</RecNum><DisplayText><style face="superscript">8</style></DisplayText><record><rec-number>10064</rec-number><foreign-keys><key app="EN" db-id="zaad5fxd72psvqet9a95xd2qadrapppw5ap0" timestamp="1464732526">10064</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Behar, D. M.</author><author>van Oven, M.</author><author>Rosset, S.</author><author>Metspalu, M.</author><author>Loogvali, E. L.</author><author>Silva, N. M.</author><author>Kivisild, T.</author><author>Torroni, A.</author><author>Villems, R.</author></authors></contributors><auth-address>Estonian Biocentre and Department of Evolutionary Biology, University of Tartu, Tartu, Estonia. behardm@</auth-address><titles><title>A "Copernican" reassessment of the human mitochondrial DNA tree from its root</title><secondary-title>Am J Hum Genet</secondary-title></titles><periodical><full-title>Am J Hum Genet</full-title><abbr-1>American journal of human genetics</abbr-1></periodical><pages>675-84</pages><volume>90</volume><number>4</number><keywords><keyword>Animals</keyword><keyword>Base Sequence</keyword><keyword>Biological Evolution</keyword><keyword>DNA, Mitochondrial/*classification/*genetics</keyword><keyword>Databases, Genetic</keyword><keyword>Genetic Variation</keyword><keyword>Haplotypes</keyword><keyword>Humans</keyword><keyword>Molecular Sequence Data</keyword><keyword>Mutation</keyword><keyword>Neanderthals/genetics</keyword><keyword>*Phylogeny</keyword></keywords><dates><year>2012</year><pub-dates><date>Apr 6</date></pub-dates></dates><isbn>1537-6605 (Electronic)
0002-9297 (Linking)</isbn><accession-num>22482806</accession-num><urls><related-urls><url> using the Burrows Wheeler Aligner ADDIN EN.CITE <EndNote><Cite><Author>Li</Author><Year>2009</Year><RecNum>38</RecNum><DisplayText><style face="superscript">9</style></DisplayText><record><rec-number>38</rec-number><foreign-keys><key app="EN" db-id="zaad5fxd72psvqet9a95xd2qadrapppw5ap0" timestamp="0">38</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Li, H.</author><author>Durbin, R.</author></authors></contributors><auth-address>Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, CB10 1SA, UK.</auth-address><titles><title>Fast and accurate short read alignment with Burrows-Wheeler transform</title><secondary-title>Bioinformatics</secondary-title><alt-title>Bioinformatics</alt-title></titles><periodical><full-title>Bioinformatics</full-title><abbr-1>Bioinformatics</abbr-1></periodical><alt-periodical><full-title>Bioinformatics</full-title><abbr-1>Bioinformatics</abbr-1></alt-periodical><pages>1754-60</pages><volume>25</volume><number>14</number><edition>2009/05/20</edition><keywords><keyword>*Algorithms</keyword><keyword>Genomics/*methods</keyword><keyword>Sequence Alignment/*methods</keyword><keyword>Sequence Analysis, DNA/methods</keyword><keyword>*Software</keyword></keywords><dates><year>2009</year><pub-dates><date>Jul 15</date></pub-dates></dates><isbn>1367-4811 (Electronic)
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ADDIN EN.CITE.DATA 3 Sequenced was performed and data processed in the same way as for the mitochondrial analysis with the exception that we mapped to the whole genome, using the reference sequence hg19. Table 1: Results on the three St. Mary’s City individuals Note: Libraries S2098.E1.L2, S2098.E1.L3, S2097.E1.L2, and S2097.E1.L3 are not listed because of reagent failures.Results (1) Summary of screening for each of the three individuals? A single library was made for HSMC-99-100/HSMC#1 (the adult male). This library passed quality control in the sense that at least 24% of cytosines in the last nucleotide of sequences were misread as thymines (authentic ancient DNA is expected to have at least 10% ADDIN EN.CITE <EndNote><Cite><Author>Sawyer</Author><Year>2012</Year><RecNum>1905</RecNum><DisplayText><style face="superscript">11</style></DisplayText><record><rec-number>1905</rec-number><foreign-keys><key app="EN" db-id="zaad5fxd72psvqet9a95xd2qadrapppw5ap0" timestamp="0">1905</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Sawyer, S.</author><author>Krause, J.</author><author>Guschanski, K.</author><author>Savolainen, V.</author><author>Paabo, S.</author></authors></contributors><auth-address>Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany. susanna_sawyer@eva.mpg.de</auth-address><titles><title>Temporal patterns of nucleotide misincorporations and DNA fragmentation in ancient DNA</title><secondary-title>PLoS One</secondary-title><alt-title>PloS one</alt-title></titles><periodical><full-title>PLoS One</full-title><abbr-1>PloS one</abbr-1></periodical><alt-periodical><full-title>PLoS One</full-title><abbr-1>PloS one</abbr-1></alt-periodical><pages>e34131</pages><volume>7</volume><number>3</number><keywords><keyword>Animals</keyword><keyword>Base Composition</keyword><keyword>DNA/*genetics</keyword><keyword>DNA Damage</keyword><keyword>*DNA Fragmentation</keyword><keyword>DNA, Mitochondrial/*genetics</keyword><keyword>Gene Library</keyword><keyword>Gorilla gorilla</keyword><keyword>Humans</keyword><keyword>Paleontology/methods</keyword><keyword>Polymerase Chain Reaction/methods</keyword><keyword>Sequence Analysis, DNA</keyword><keyword>Time Factors</keyword></keywords><dates><year>2012</year></dates><isbn>1932-6203 (Electronic)
1932-6203 (Linking)</isbn><accession-num>22479540</accession-num><urls><related-urls><url>), the match rate to the mitochondrial consensus sequence is 99.6% indicating minimal contamination, and the coverage on autosomal targets is 326,801 SNPs.? Two libraries were successfully made for HSMC-99-200/HSMC#2 (the adult female), which were both inhibited or failed amplification. We stopped after two attempts.? Two libraries were successfully made for HSMC-99-300/HSMC#3, the second of which gave results consistent with authentic ancient DNA. A fraction 19% of cytosines in the last nucleotide of sequences were misread as thymines, the match rate to the mitochondrial consensus sequence is 99.9% indicating minimal contamination, and the coverage on autosomal targets is 618,843 SNPs.(2) The adult male and infant are genetically male and have minimal contaminationThe observed ratio of X chromosome to Y chromosome SNP coverage for the two samples that gave working ancient DNA (the adult male and the infant) were both consistent with male sex. Because males have a single X chromosome it is possible to use heterozygosity on chromosome X to quantify contamination (using the ANGSD tool). ADDIN EN.CITE <EndNote><Cite><Author>Korneliussen</Author><Year>2014</Year><RecNum>8672</RecNum><DisplayText><style face="superscript">12</style></DisplayText><record><rec-number>8672</rec-number><foreign-keys><key app="EN" db-id="zaad5fxd72psvqet9a95xd2qadrapppw5ap0" timestamp="0">8672</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Korneliussen, T. S.</author><author>Albrechtsen, A.</author><author>Nielsen, R.</author></authors></contributors><auth-address>Centre for GeoGenetics, Natural History Museum of Denmark, Copenhagen, Denmark. thorfinn@binf.ku.dk.
Bioinformatics Centre, Department of Biology, University of Copenhagen, Ole Maaloes Vej 5, Copenhagen, DK-2200, Denmark. albrecht@binf.ku.dk.
Centre for GeoGenetics, Natural History Museum of Denmark, Copenhagen, Denmark. rasmus_nielsen@berkeley.edu.
Department of Integrative Biology and Statistics, UC-Berkeley, 4098 VLSB, Berkeley, California, 94720, USA. rasmus_nielsen@berkeley.edu.</auth-address><titles><title>ANGSD: Analysis of Next Generation Sequencing Data</title><secondary-title>BMC Bioinformatics</secondary-title><alt-title>BMC bioinformatics</alt-title></titles><periodical><full-title>BMC Bioinformatics</full-title><abbr-1>BMC bioinformatics</abbr-1></periodical><alt-periodical><full-title>BMC Bioinformatics</full-title><abbr-1>BMC bioinformatics</abbr-1></alt-periodical><pages>356</pages><volume>15</volume><dates><year>2014</year></dates><isbn>1471-2105 (Electronic)
1471-2105 (Linking)</isbn><accession-num>25420514</accession-num><urls><related-urls><url> There was no evidence of contamination for either individual (both Z<2 standard errors from zero). Point estimates were 1.6% contamination for the adult male and 0.7% for the infant male.(2) Mitochondrial DNA haplogroups are different for the two individualsMitochondrial DNA is inherited from mother to child. The mitochondrial DNA haplogroups inferred based on the consensus sequences are T2b4 for the adult male, and V for the infant. These are both consistent with Western European ancestry. The fact that the two haplogroups are different excludes the individuals from being full siblings.(3) Y chromosome haplogroups are consistent with being the same for the two individuals? The Y chromosome haplogroup call for the adult male is R1b1a2a1a (supported by the P311:18248698A>G and P310:18907236A>C mutations).? The Y chromosome haplogroup call for the infant male is a more resolved version of that for the adult male, that is, R1b1a2a1a2 (supported by the P312:22157311C->A mutation). These two individuals belong to a haplogroup that is common in people of western European descent and so the results are consistent with but do not prove a patrilineal relationship. (4) Nuclear analysis implies a father-son pairPrincipal Component Analysis using smartpca in EIGENSOFT ADDIN EN.CITE <EndNote><Cite><Author>Patterson</Author><Year>2006</Year><RecNum>193</RecNum><DisplayText><style face="superscript">13</style></DisplayText><record><rec-number>193</rec-number><foreign-keys><key app="EN" db-id="zaad5fxd72psvqet9a95xd2qadrapppw5ap0" timestamp="0">193</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Patterson, N.</author><author>Price, A. L.</author><author>Reich, D.</author></authors></contributors><auth-address>Broad Institute of Harvard and MIT, Cambridge, Massachusetts, United States of America.</auth-address><titles><title>Population structure and eigenanalysis</title><secondary-title>PLoS genetics</secondary-title><alt-title>PLoS Genet</alt-title></titles><periodical><full-title>PLoS Genet</full-title><abbr-1>PLoS genetics</abbr-1></periodical><alt-periodical><full-title>PLoS Genet</full-title><abbr-1>PLoS genetics</abbr-1></alt-periodical><pages>e190</pages><volume>2</volume><number>12</number><edition>2006/12/30</edition><keywords><keyword>Computer Simulation/statistics & numerical data</keyword><keyword>Genetic Markers</keyword><keyword>*Genetic Variation</keyword><keyword>Genetics, Medical/*methods/statistics & numerical data</keyword><keyword>Genetics, Population/*methods/statistics & numerical data</keyword><keyword>Humans</keyword><keyword>*Models, Genetic</keyword><keyword>Models, Statistical</keyword><keyword>Principal Component Analysis/*methods</keyword></keywords><dates><year>2006</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1553-7404 (Electronic)
1553-7390 (Linking)</isbn><accession-num>17194218</accession-num><work-type>Research Support, N.I.H., Extramural
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ADDIN EN.CITE.DATA 1 indicate that both individuals are Western European in ancestry, clustering closely with present-day English individuals (Fig. 1). We used PLINK ADDIN EN.CITE <EndNote><Cite><Author>Purcell</Author><Year>2007</Year><RecNum>9371</RecNum><DisplayText><style face="superscript">14</style></DisplayText><record><rec-number>9371</rec-number><foreign-keys><key app="EN" db-id="zaad5fxd72psvqet9a95xd2qadrapppw5ap0" timestamp="1451680833">9371</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Purcell, S.</author><author>Neale, B.</author><author>Todd-Brown, K.</author><author>Thomas, L.</author><author>Ferreira, M. A.</author><author>Bender, D.</author><author>Maller, J.</author><author>Sklar, P.</author><author>de Bakker, P. I.</author><author>Daly, M. J.</author><author>Sham, P. C.</author></authors></contributors><auth-address>Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA 02114, USA. shaun@pngu.mgh.harvard.edu</auth-address><titles><title>PLINK: a tool set for whole-genome association and population-based linkage analyses</title><secondary-title>Am J Hum Genet</secondary-title><alt-title>American journal of human genetics</alt-title></titles><periodical><full-title>Am J Hum Genet</full-title><abbr-1>American journal of human genetics</abbr-1></periodical><alt-periodical><full-title>Am J Hum Genet</full-title><abbr-1>American journal of human genetics</abbr-1></alt-periodical><pages>559-75</pages><volume>81</volume><number>3</number><keywords><keyword>Genetic Linkage/*genetics</keyword><keyword>Genome, Human/*genetics</keyword><keyword>Humans</keyword><keyword>Polymorphism, Single Nucleotide</keyword><keyword>Population/*genetics</keyword><keyword>*Software</keyword></keywords><dates><year>2007</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0002-9297 (Print)
0002-9297 (Linking)</isbn><accession-num>17701901</accession-num><urls><related-urls><url> to test the relatedness of the sample pair, and find that PI_HAT is 0.4274, consistent with being first-degree relatives.Figure 1: Principal components analysis. We projected the two ancient individuals onto the first two principal components of 392 present-day European individualsPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MYXphcmlkaXM8L0F1dGhvcj48WWVhcj4yMDE2PC9ZZWFy
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ADDIN EN.CITE.DATA 2,15,16. The two colonial individuals cluster closely to each other at the edge of the cluster of 10 present-day English individuals.ConclusionTaken together, these results suggest that HSMC-99-100/HSMC#1, the adult male identified as Philip Calvert, is a first degree relative of HSMC-99-300/HSMC#3, the unknown infant who is genetically identified as a male. The two individuals do not share the same mitochondrial DNA sequence which excludes them as full siblings. Thus, they are a father and son, consistent with their matching Y chromosome haplogroups. Data AvailabilityAutosomal and mtDNA BAMs of the two individuals are available from the European Nucleotide Archive under accession number PRJEB16735. Genotype data from the two individuals are available from the Reich lab datasets web page at genetics.med.harvard.edu/reich/Reich_Lab/Datasets.html.References ADDIN EN.REFLIST 1Haak, W. et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature, doi:10.1038/nature14317 (2015).2Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient Near East. Nature, doi:10.1038/nature19310 (2016).3Mathieson, I. et al. Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528, 499-503, doi:10.1038/nature16152 (2015).4Skoglund, P. et al. Genomic insights into the peopling of the Southwest Pacific. Nature, doi:10.1038/nature19844 (2016).5Dabney, J. et al. Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments. Proceedings of the National Academy of Sciences of the United States of America 110, 15758-15763, doi:10.1073/pnas.1314445110 (2013).6Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil-DNA-glycosylase treatment for screening of ancient DNA. Philosophical transactions of the Royal Society of London. Series B, Biological sciences 370, 20130624, doi:10.1098/rstb.2013.0624 (2015).7Meyer, M. et al. A mitochondrial genome sequence of a hominin from Sima de los Huesos. Nature 505, 403-406, doi:10.1038/nature12788 (2014).8Behar, D. M. et al. A "Copernican" reassessment of the human mitochondrial DNA tree from its root. American journal of human genetics 90, 675-684, doi:10.1016/j.ajhg.2012.03.002 (2012).9Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics 25, 1754-1760, doi:10.1093/bioinformatics/btp324 (2009).10Fu, Q. et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature, doi:10.1038/nature14558 (2015).11Sawyer, S., Krause, J., Guschanski, K., Savolainen, V. & Paabo, S. Temporal patterns of nucleotide misincorporations and DNA fragmentation in ancient DNA. PloS one 7, e34131, doi:10.1371/journal.pone.0034131 (2012).12Korneliussen, T. S., Albrechtsen, A. & Nielsen, R. ANGSD: Analysis of Next Generation Sequencing Data. BMC bioinformatics 15, 356, doi:10.1186/s12859-014-0356-4 (2014).13Patterson, N., Price, A. L. & Reich, D. Population structure and eigenanalysis. PLoS genetics 2, e190, doi:10.1371/journal.pgen.0020190 (2006).14Purcell, S. et al. PLINK: a tool set for whole-genome association and population-based linkage analyses. American journal of human genetics 81, 559-575, doi:10.1086/519795 (2007).15Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513, 409-413, doi:10.1038/nature13673 - supplementary-information (2014).16Patterson, N. et al. Ancient admixture in human history. Genetics 192, 1065-1093 (2012). ................
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