Ancient phylogeography: spread of haplogroups R1b, R1a and N

The previous post showed the potential use of TreeToM to visualize ancient DNA samples in maps together with their Y-DNA phylogenetic trees. I have written Newick trees for Y-chromosome haplogroups R1b-L388 (encompassing R-V1636 and R-P297, which in turn split into R-M73 and R-M269), R1a, and N.

I have reviewed some of the BAM files from my previous bulk analyses with YLeaf v.2, to add information that I had not previously included in the All Ancient DNA Dataset, and which might be relevant to the proper depiction of phylogenetic trees; in particular, positive and negative SNPs potentially distinguishing archaic subclades (“pre-” and “basal” branches) within the R1a, R1b, and N trunks:

  1. Haplogroup K split in Northern Asia.
  2. Haplogroup R1b Eurasiatic expansion.
  3. Haplogroup R1a East European expansion.
  4. Haplogroup N Siberian expansion.

NOTE. I don’t have currently time for much more, and SNP inference is a kind of “art” which requires experience assessing (available ancestral and derivative reads and their implications in) hundreds of ancient and modern samples which very few people have, and few of them dedicate time to ancient DNA, so if you see errors just contact me.

1. Haplogroup K split in Northern Asia

It seems quite likely that the expansions of early K subclades accompanied the different waves of ancestry described for AMH, Upper Palaeolithic and Epipalaeolithic expansions, although it is still unclear which precise route(s) R1a and R1b carriers took and when, because the few available ancient samples (and modern DNA distributions) are of little help regarding potential northern vs. southern Caspian paths for ca. 20,000-10,000 years ago.

1.2. Haplogroup K Newick tree

This is a simplified Newick tree for the available samples within the K trunk:

('K*':0.1,
 ('K2*':0.1,
 '(K2)':0.1,
  ('K2b*':0.1,
  '(K2b)':0.1,
   ('K2b2/P':0.1,
    ('P1':0.1,
     ('P337*':0.1,
      ('P284*':0.1
      )P284:0.1
     )P337:0.1
    )P1:0.1,
    ('(P2)':0.1,
     ('Q':0.1,
      ('(Q1)':0.1,
       ('(Q1a)':0.1,
        ('F746*':0.1,
         ('(YP1500)':0.1
         )YP1500:0.1
        )F746:0.1,
        ('(M25)':0.1
        )M25:0.1
       )Q1a:0.1,
       ('(Q1b)':0.1,
        ('(Q1b1)':0.1,
         ('(Q1b1a)':0.1,
          ('(M930)':0.1,
           ('(M3)':0.1
           )M3:0.1
          )M930:0.1,
          ('(Z780)':0.1,
           ('(M944)':0.1
           )M944:0.1
          )Z780:0.1
         )Q1b1a:0.1
        )Q1b1:0.1
       )Q1b:0.1
      )Q1:0.1
     )Q:0.1,
     ('R*':0.1,
      ('R1':0.1,  
       ('R1a':0.1,
        ('(YP4141)':0.1,
         ('YP5018*':0.1
         )YP5018:0.1
        )YP4141:0.5,
        ('M459*':0.1,
        '(M459)':0.1,
         ('(YP1272)':0.1
         )YP1272:0.7,
         ('(M198)':0.1
         )M198:0.6
        )M459:0.3
       )R1a:0.5,
       ('R1b':0.1,
        ('(L754)':0.01,
         ('(L389)':0.1,
          ('(V1636)':0.1,
           ('(CTS3187)':0.1
           )CTS3187:0.9
          )V1636:0.9,
          ('P297*':0.1,
          '(P297)':0.1,
           ('M73*':0.1,
           '(M73)':0.1
           )M73:0.6,
           ('M269*':0.1,
           '(M269)':0.1
           )M269:0.7
          )P297:0.2
         )L389:0.2,
         ('V2219*':0.01,
         '(V2219)':0.01,
          ('PF6340*':0.01,
          '(PF6340)':0.01,
           ('Y8457*':0.1,
           '(Y8457)':0.1
           )Y8457:0.1
          )PF6340:0.1
         )V2219:0.1
        )L754:0.1
       )R1b:0.1
      )R1:0.1,
      ('R2':0.1,
       ('R2a*':0.1,
       '(R2a)':0.1
       )R2a:0.1,
       ('R2b':0.1
       )R2b:0.1
      )R2:0.5
     )R:0.1
    )P2:0.1
   )K2b2:0.1
  )K2b:0.1
 )K2:0.1
)K:0.1

1.2. Haplogroup K ancient samples

These are the available ancient DNA samples until roughly 6000 BC, with the CSV code for TreeToM:

ancient-dna-haplogroups-k-p-q-r

(K2),57.7,71.1,black
(K2b),39.39,115.52,black
P284*,70.7,135.4,purple
P337*,70.7,135.4,purple
R*,52.9,103.5,red
F746*,56.05,92.87,deeppink
V2219*,46.15,12.21,blue
(M944),45.993056,-110.661389,deeppink
V2219*,44.29,4.46,blue
V2219*,44.5154854,23.52087725,blue
M459*,48.3,35.139,green
R2a*,34.45,48.116,brown
(R2a),34.45,48.116,brown
(R2a),34.45,48.116,brown
(R2a),34.45,48.116,brown
(R2a),34.45,48.116,brown
(YP1500),68.6,159.1,deeppink
(M930),-19.47718333,-44.03805556,deeppink
V2219,44.517,22.722,blue
(Z780),-19.47718333,-44.03805556,deeppink
P297,56.2833,25.1333,blue
(Y8457),48.3,35.139,blue
YP5018*,48.91422,33.76493,green
V2219*,44.640262,22.30333,blue
(Z780),-10.3222,-76.6666667,deeppink
V2219*,44.629711,22.6125561,blue
(Z780),-38.3605556,-60.2441667,deeppink
(Y8451),44.53,22.05,blue
(Q1),-10.3222,-76.6666667,deeppink
(M3),46.223056,-119.143333,deeppink
(M25),56.2833,25.1333,deeppink
PF6340*,44.53,22.05,blue
(Q1b),-38.3605556,-60.2441667,deeppink
M459*,61.65,35.65,green
(M459),43.98,26.4,green
P297*,53.4,50.4,blue
(P297),56.2833,25.1333,blue

1.3. Haplogroup K TreeToM map

This is the map of Eurasia (see also the map including Q samples in the Americas):

haplogroup-k2-r-q-expansion-north-eurasia

2. Haplogroup R1b Eurasiatic expansion

2.1. Haplogroup R1b Newick tree

This is the Newick tree for R1b-L388 (see here for the V2219 tree):

NOTE. I have used a simplified adjustment of node distances to reflect the TMRCA of early branches (where 0.1 is roughly a thousand years), but then I have kept 0.1 for L23 and subsequent branches to avoid overcrowding.

('(L389?)':0.1,
  ('(V1636)':0.1,
   ('(CTS3187)':0.1
   )CTS3187:0.9
  )V1636:0.9,
  ('P297*':0.1,
   '(P297)':0.1,
   ('M73*':0.1,
   '(M73)':0.1,
    ('(M478)':0.1,
     ('(L1433)':0.1
     )L1433:0.1
    )M478:0.2
   )M73:0.6,
   ('M269*':0.1,
   '(M269)':0.1,
    ('(PF7562)':0.1
    )PF7562:0.2,
    ('L23*':0.1,
    '(L23)':0.1,
     ('Z2103*':0.1,
     '(Z2103)':0.1,
      ('(Y13369)':0.1,
       ('(L584)':0.1,
        ('(FGC14590)':0.1,
         ('(Y19434)':0.1,
          ('(Y16852)':0.1,
           ('Y23838*':0.1,
           '(Y23838)':0.1
           )Y23838:0.1
          )Y16852:0.1
         )Y19434:0.1,
         ('(L945)':0.1
         )L945:0.1
        )FGC14590:0.1
       )L584:0.1
      )Y13369:0.1,
      ('(Y4364)':0.1,
       ('(Y4366)':0.1,
        ('(BY13830)':0.1,
         ('(BY3293)':0.1
         )BY3293:0.1
        )BY13830:0.1
       )Y4366:0.1
      )Y4364:0.1,
      ('(Z2106)':0.1,
       ('(Z2109)':0.1,
        ('(KMS67)':0.1,
         ('(KMS75)':0.1,
          ('(KMS75)':0.1,
           ('(KMS86)':0.1
           )KMS86:0.1
          )KMS75:0.1
         )KMS75:0.1
        )KMS67:0.1
       )Z2109:0.1
      )Z2106:0.1
     )Z2103:0.1,
     ('L51*':0.1,
     '(L51)':0.1,
      ('Z2118*':0.1,
      '(Z2118)':0.1
      )Z2118:0.1,
      ('P310*':0.1,
      '(P310)':0.1,
       ('L151*':0.1,
       '(L151)':0.1,
        ('(S1194)':0.1,
         ('(A8039)':0.1,
          ('(A8040)':0.1,
           ('(A8041)':0.1
           )A8041:0.1
          )A8040:0.1
         )A8039:0.1
        )S1194:0.1,
        ('(U106)':0.1,
         ('(FGC3861)':0.1
         )FGC3861:0.1,
         ('(Z18)':0.1
         )Z18:0.1,
         ('(Z2265)':0.1,
          ('(BY30097)':0.1,
           ('(Z381)':0.1
           )Z381:0.1
          )BY30097:0.1,
          ('(Z2265)':0.1,
           ('(BY30097)':0.1,
            ('(Z381)':0.1,
             ('(Z156)':0.1,
              ('(Z306)':0.1
              )Z306:0.1
             )Z156:0.1
            )Z381:0.1
           )BY30097:0.1
          )Z2265:0.1
         )Z2265:0.1
        )U106:0.1,
        ('P312*':0.1,
        '(P312)':0.1,
         ('(DF19)':0.1,
          ('(DF19)':0.1
          )DF19:0.1
         )DF19:0.1,
         ('(Z290)':0.1,
          ('(L21)':0.1,
           ('DF13)':0.1,
            ('L513)':0.1,
             ('S6365)':0.1,
              ('BY17)':0.1,
               ('S7106)':0.1,
                ('Z16386)':0.1
                )Z16386:0.1
               )S7106:0.1
              )BY17:0.1
             )S6365:0.1
            )L513:0.1,
            ('DF21)':0.1,
             ('Z30233)':0.1
             )Z30233:0.1
            )DF21:0.1
           )DF13:0.1
          )L21:0.1
         )Z290:0.1,
         ('(ZZ11)':0.1,
          ('(U152)':0.1,
           ('(PF6658)':0.1
           )PF6658:0.1,
           ('(Z56)':0.1,
            ('(Z43)':0.1,
             ('(Z145)':0.1,
              ('(PH2982)':0.1,
               ('(CTS6389)':0.1
               )CTS6389:0.1
              )PH2982:0.1,
              ('(BY1823)':0.1,
               ('(BY3951)':0.1
               )BY3951:0.1
              )BY1823:0.1
             )Z145:0.1
            )Z43:0.1
           )Z56:0.1,
           ('(L2)':0.1,
            ('(PR5365)':0.1
            )PR5365:0.1,
            ('(DF110)':0.1
            )DF110:0.1,
            ('(FGC4183)':0.1
            )FGC4183:0.1,
            ('(DF90)':0.1
            )DF90:0.1,
            ('(Z258)':0.1,
             ('(Z367)':0.1,
              ('(L20)':0.1
              )L20:0.1
             )Z367:0.1
            )Z258:0.1,
            ('(Z49)':0.1,
             ('(Z142)':0.1
             )Z142:0.1
            )Z49:0.1
           )L2:0.1
          )U152:0.1,
          ('(DF27)':0.1,
           ('(ZZ12)':0.1,
            ('(Z225)':0.1
            )Z225:0.1,
            ('(BY15963)':0.1
            )BY15963:0.1,
            ('(BY3332)':0.1,
             ('(PH4023)':0.1
             )PH4023:0.1
            )BY3332:0.1,
            ('(FT9510)':0.1,
             ('(FT64916)':0.1,
              ('(Y30814)':0.1
              )Y30814:0.1
             )FT64916:0.1
            )FT9510:0.1,
            ('(FGC78762)':0.1,
             ('(BY21074)':0.1,
              ('(PH2781)':0.1
              )PH2781:0.1
             )BY21074:0.1,
             ('(BY3235)':0.1,
              ('(BY61519)':0.1
              )BY61519:0.1
             )BY3235:0.1
            )FGC78762:0.1
           )ZZ12:0.1,
           ('(Z195)':0.1,
            ('(Z274)':0.1,
             ('(Z209)':0.1,
              ('(Z295)':0.1,
               ('(Z270)':0.1,
                ('(Z219)':0.1
                )Z219:0.1
               )Z270:0.1
              )Z295:0.1
             )Z209:0.1
            )Z274:0.1,
            ('(Z198)':0.1,
             ('(L165)':0.1
             )L165:0.1,
             ('(ZS312)':0.1,
              ('(BY3214)':0.1
              )BY3214:0.1
             )ZS312:0.1
            )Z198:0.1
           )Z195:0.1
          )DF27:0.1
         )ZZ11:0.1
        )P312:0.1
       )L151:0.1
      )P310:0.1
     )L51:0.1
    )L23:0.1
   )M269:0.7
  )P297:0.2
)L389:0.2

2.2. Haplogroup R1b ancient samples

This is the list of available ancient samples within the R1b-L388 tree ordered by date:

NOTE. I have tried to remove all samples with undefined subclades whenever they are not informative for the specific period and region (viz. undefined L23 in the Bell Beaker period, undefined L151 or P312 in the Bronze Age, etc.).

(P297),56.2833,25.1333,chartreuse
(P297),56.2833,25.1333,chartreuse
(L754),56.2833,25.1333,black
(P297),56.2833,25.1333,chartreuse
P297*,53.4,50.4,darkgreen
(P297),56.2833,25.1333,chartreuse
(M73),55.807759,22.422246,aqua
P297*,56.2833,25.1333,darkgreen
(V1636),52.22,48.1,#2F4F4F
(V1636),43.822691,43.350278,#2F4F4F
(L389?),51.63847621,46.61499023,black
(L389?),51.63847621,46.61499023,black
(L389?),51.63847621,46.61499023,black
(L389?),51.63847621,46.61499023,black
(L389?),51.63847621,46.61499023,black
P297*,58.272319,26.051586,darkgreen
(P297),56.2833,25.1333,chartreuse
(V1636),43.822691,43.350278,#2F4F4F
(M269),53.10453726,49.42989349,purple
(M269),53.10453726,49.42989349,purple
(M269),53.10453726,49.42989349,purple
P297*,56.2833,25.1333,darkgreen
(P297),49.88193151,87.87135903,chartreuse
(P297),57.504548,26.584483,chartreuse
(M269),56.78451992,40.45303345,purple
(M269),56.78451992,40.45303345,purple
(M73),53.305983,67.648166,aqua
(Z2103),50.7581128,85.56024972,blue
(Z2103),53.38,50.39,blue
(L23),53.38,50.39,darkblue
(Z2103),53.31,51.15,blue
(Z2109),50.75648626,85.56087884,indigo
(L23),49.88193151,87.87135903,darkblue
(Z2103),54.01169323,90.66188638,blue
(Z2103),51.49763611,85.97078889,blue
L23*,53.38,50.39,red
(Z2103),51.27,58.18,blue
(L945),50.75364627,85.55666955,skyblue
(L23),46.556,43.676,darkblue
(L945),50.7581128,85.56024972,skyblue
(Z2103),50.75364627,85.55666955,blue
(Z2109),49.134217,75.851817,indigo
(Z2109),50.75364627,85.55666955,indigo
(Z2109),53.87388889,91.46972222,indigo
(KMS75),52.71085,49.46685556,indigo
(Z2103),53.38,50.38,blue
(L23),54.35661,90.919186,darkblue
(BY3293),48.91,33.76,steelblue
(KMS86),53.16638889,50.67361111,indigo
(V1636),45.725651,43.986933,#2F4F4F
(Z2103),53.38,50.39,blue
(Z2103),45.537249,41.116544,blue
(Z2103),44.031853,43.129956,blue
(Z2103),45.76,18.57,blue
(Z2103),54.35661,90.919186,blue
(Z2109),53.58975,50.57129167,indigo
(KMS75),43.905354,43.521883,indigo
(Z2106),46.539,43.699,indigo
(L23),44.031853,43.129956,darkblue
(Z2103),54.01169323,90.66188638,blue
(Z2103),54.01169323,90.66188638,blue
(Z2103),53.34,50.34,blue
(KMS75),44.023591,42.818456,indigo
(Z2103),48.832683,83.909744,blue
(Z2103),44.05336,43.21034,blue
(L23),46.539,43.699,darkblue
(Z2103),46.539,43.699,blue
(Z2109),54.35661,90.919186,indigo
(Z2106),48.716,44.5,indigo
(Z2103),53.31,51.15,blue
Z2103*,45.535978,41.059517,crimson
(Z2103),53.708561,90.359808,blue
(L151),49.3628,3.4628,brown
(CTS3187),40.65,45.1167,gray
(P310),49.468332,9.744072,firebrick
(Z2103),49.96666667,44.66666667,blue
(U152),48.691,13.016,darkorange
(L2),42.419214,-3.484745,orange
(P312),52.733563,5.096183,sienna
(P297),69.47,18,chartreuse
(Z290),50.919204,-2.015441,burlywood
(P312),47.65,7.45,sienna
(L23),43.41022,-5.98496,darkblue
(Z2103),54.01169323,90.66188638,blue
(L51),47.383224,19.020252,maroon
(L151),50.410265,14.074597,brown
L151*,47.429954,19.051966,red
L151*,47.429954,19.051966,red
(L2),50.410265,14.074597,orange
(L2),50.410265,14.074597,orange
(L2),50.410265,14.074597,orange
(L2),50.410265,14.074597,orange
(L2),50.410265,14.074597,orange
(L2),50.410265,14.074597,orange
(L2),50.410265,14.074597,orange
(Z2109),49.96666667,44.66666667,indigo
(Z2109),47.429954,19.051966,indigo
(P312),47.65,7.45,sienna
(M269),43.9655,5.8126,purple
(Z2103),45.725651,43.986933,blue
(L2),47.59916,19.05456,orange
P310*,48.33,10.9,red
(Z2103),50.649,21.656,blue
(Z2103),47.383224,19.020252,blue
(Z2109),45.535978,41.059517,indigo
(PF6658),51.79,11.14,darkorange
(L151),51.16,-1.77,brown
(L2),50.272105,15.818985,orange
(L2),50.09,18.1,orange
(L21),51.22,-1.77,burlywood
(P312),45.46,6.213,Sienna
(DF27),51.79,11.14,goldenrod
(P312),49.153,6.102,sienna
(L2),48.31966,10.89153,orange
(DF27),37.95,14.083333,goldenrod
(L2),48.32,10.89,orange
(L51),48.691,13.016,maroon
L51*,50.1217475,14.4569508,red
(L151),40.22,-3.76,brown
(P312),40.22,-3.76,sienna
(ZZ11),48.33,10.9,peru
(PF6658),40.49033,-3.373888,darkorange
(L2),50.19,14.158,orange
(L2),50.19,14.158,orange
(L2),50.19,14.158,orange
(L2),50.19,14.158,orange
(L2),50.19,14.158,orange
(L2),50.19,14.158,orange
(L2),50.19,14.158,orange
(L2),48.33,10.895,orange
(L2),48.691,13.016,orange
(L2),48.841,12.75,orange
(L2),48.841,12.75,orange
(L2),48.841,12.75,orange
(Z290),51.16,-1.77,burlywood
(L2),48.25283,10.80879,orange
(L151),46.231707,7.351237,brown
(P312),48.8833,12.5332,sienna
(L2),48.8833,12.5332,orange
(L2),48.8833,12.5332,orange
(L2),50.87,16.86,orange
(L2),50.0504153,14.3667267,orange
(L151),50.649,21.656,brown
(L151),50.8316798,-0.1342324,brown
(L2),50.166667,14.316667,orange
(Z290),51.2377356,-2.998695,burlywood
(L21),51.1576798,-1.8103,burlywood
(P312),37.908,13.478,sienna
(L2),50.1217475,14.4569508,orange
(L2),50.1217475,14.4569508,orange
(DF27),37.908,13.478,goldenrod
(L51),42.419214,-3.484745,maroon
L51*,50.11,18.21,red
(P312),37.908,13.478,sienna
(P310),50.118,14.258,firebrick
(U106),55.381,13.445,fuchsia
(Z290),51.16,-1.77,burlywood
(Z290),51.16,-1.77,burlywood
(P312),49.153,6.102,Sienna
(FGC4183),42.72503,-3.844537,orange
(P312),54.522812,-1.308134,sienna
(L2),50.1217475,14.4569508,orange
(L2),50.1217475,14.4569508,orange
(L2),48.29573,10.89103,orange
(L2),50.1217475,14.4569508,orange
(P312),52.733563,5.096183,sienna
(P312),52.733563,5.096183,sienna
(Z2103),43.826337,42.716199,blue
(P312),55.97,-2.444,sienna
(P312),44.8,10.333333,sienna
(L2),48.29573,10.89103,orange
(P312),52.733563,5.096183,sienna
(L2),48.26656,10.87857,orange
(L2),48.29573,10.89103,orange
(P312),52.733563,5.096183,sienna
(L2),44.476,6.369,orange
(P312),52.733563,5.096183,sienna
L23*,53.58975,50.57129167,red
(P312),55.99,-2.537,sienna
(L2),48.29573,10.89103,orange
(L21),57.721,-3.3881,burlywood
(L2),48.26656,10.87857,orange
(L2),48.29573,10.89103,orange
(P312),48.26656,10.87857,sienna
(L2),48.25283,10.80879,orange
(Z30233),55.292132,-6.191685,burlywood
(P312),48.22281,10.84499,sienna
(Z305),50.0504153,14.3667267,deeppink
(P312),50.0504153,14.3667267,sienna
(L2),48.29573,10.89103,orange
(Z290),56.062,-3.419178,burlywood
(BY61519),38.5781,-2.8417,goldenrod
(Z2109),52.81666667,60.46666667,indigo
(DF21),55.292132,-6.191685,burlywood
(P312),37.908,13.478,sienna
(BY3332),38.8494,-0.87138,goldenrod
(L2),48.22281,10.84499,orange
(P297),40.14,45.26,chartreuse
(P312),51.33,1.36,sienna
(L2),48.22281,10.84499,orange
(Z290),51.471197,-0.266408,burlywood
(L2),48.22281,10.84499,orange
(L2),48.22281,10.84499,orange
(L2),48.22281,10.84499,orange
(L2),48.22281,10.84499,orange
L1433,52.81666667,60.46666667,#8FBC8F
(U106),52.733563,5.096183,fuchsia
(L2),48.22281,10.84499,orange
(L21),55.309983,-1.553328,burlywood
(Z225),42.44,-2.8867,goldenrod
(DF27),39.526799,-0.509749,goldenrod
(Z195),41.5373,2.1329,goldenrod
(L51),51.15,-1.75,maroon
(P312),48.22281,10.84499,sienna
(L151),38.6314,-0.8615,brown
(Y30814),42.7422292,-3.8377766,goldenrod
(ZZ12),36.132004,-5.343773,sienna
(ZZ12),36.132004,-5.343773,sienna
(L151),42.44,-2.8867,brown
(L21),55.292132,-6.191685,burlywood
(P312),52.696301,-0.34,sienna
(L2),48.17,10.814,orange
(Z198),41.2916,1.0638,goldenrod
(ZZ11),41.223,1.77416,peru
(P312),38.0393077,-7.6005786,sienna
(L151),42.57,-2.62,brown
(M269),40.8586,-5.3518,purple
L151*,50.5589705,13.802021,red
(BY15963),41.963046,-1.992604,goldenrod
(M269),53.043,57.066,purple
(Z290),50.919204,-2.015441,burlywood
(U106),56.97,9.552,fuchsia
(Z290),52.34,0.03,burlywood
(L23),37.730792,22.756382,blue
(Z16386),55.98,-2.896,burlywood
(Z2103),37.730792,22.756382,blue
(Z16386),52.17839,0.122544,burlywood
(P312),52.34,0.03,sienna
(ZZ12),42.57,-2.62,sienna
(DF27),41.4584,-5.2872,goldenrod
(Z2103),31.666667,34.566667,blue
(L21),52.122402,-0.511365,burlywood
(M269),32.6641744,35.2249937,purple
(Z290),52.122402,-0.511365,burlywood
(Z2103),36.9944,45.4744,blue
(Z290),52.122402,-0.511365,burlywood
(Z290),52.122402,-0.511365,burlywood
(M478),50.2249,81.83678333,aqua
(Z2103),39.2,46.4,blue
Y23838*,37.004831,45.458892,skyblue
(P312),57.721,-3.3881,sienna
(P312),39.999,3.891,sienna
(Z290),57.72,-3.388,burlywood
(FGC4183),50.5099972,14.05,orange
(Z2103),41.7516182,12.4286064,blue
(DF90),41.6026782,12.508636,orange
Z2118*,41.6433289,12.9126915,red
(PH4023),37.3244444,-4.005833333,goldenrod
(BY3951),41.7952421,12.7789564,orange

2.3. Haplogroup R1b TreeToM maps

General R1b-L388 TreeToM map:

r1b-samples-ancient-dna

2.3.1. Haplogroup R1b Mesolithic-Neolithic expansions

This is a TreeToM map of Mesolithic and Neolithic samples, which trace a potential picture of the Eurasiatic-related expansions including Indo-Uralic and the paradigmatic Proto-Indo-European-speaking Khvalynsk culture:

NOTE. Not included is the potential R1b-M269 sample with Steppe ancestry from the Balkan Chalcolithic Smyadovo site, due to conflicting negative SNPs (for P) potentially indicating damage. If it is within the R trunk, as officially reported, then it’s likely R1b-M269, too, as it likely expanded with Suvorovo chiefs.

r1b-samples-mesolithic-khvalynsk

2.3.2. Haplogroup R1b in Yamnaya- and Afanasievo-related peoples

These are the samples and corresponding TreeToM map of Yamnaya, Afanasievo, and other Early Bronze Age groups. Particularly interesting are the basal L23* and Z2103* subclades found in the Don-Volga region, supporting Anthony’s (2019) description of it as cradle of Proto-Indo-Europeans.

I have selected the most conservative assignment, L23*, for the sample from Lopatino II, reported as xZ2103, potentially pre-L51 (Y410,xL51), but this map and the following period with East Bell Beakers suggest where the first full L51 samples are going to be found among the Yamnaya

NOTE. The Corded Ware sample from Obłaczkowo, RISE1, is not included due to lack of subclade beyond R1b-L754, and especially because of the conflicting archaeological context (hence origin and radiocarbon date), given that the tested individual should be the same as poz44, which has a different Y-DNA and mtDNA…

r1b-samples-yamnaya-afanasievo

ancient-r1b-haplogroup-yamnaya-afanasievo

2.3.3. Haplogroup R1b in Bell Beaker, Balkan, and Poltavka expansions

These are the samples and corresponding TreeToM map of European Late Neolithic – Early Bronze Age groups, such as Bell Beakers, Balkan BA, Catacomb, Poltavka. Particularly interesting are:

  • High variability proper of the expected Bell Beakers source – includinig basal subclades L51*, L151*, P310*, as well as Z2103 (and dubious U106, which I haven’t included despite the confirmed slightly later sample from Únětice) – among the East BBC Group (see another map with basal subclades).
  • Basal L23* from Samara within the Potapovka culture and Z2103 lineages in Sintashta, both a mixture of Abashevo chiefs and Poltavka herders in the development of the Proto-Indo-Iranian community.

r1b-samples-bell-beaker-potapovka

ancient-r1b-haplogroup-bell-beaker-catacomb

The following map is a close-up of the Bell Beaker and EEBA area and associated R1b lineages. Even without including the dubious U106 samples from Bavarian, Czech, and Hungarian BBC groups, it is evident from Bell Beaker and European Early Bronze Age data combined that the common origin of expansion of Yamnaya lineages with Bell Beakers was located in Central Europe closest to the Middle Danube, with the periphery showing bottlenecks under specific subclades.

ancient-r1b-haplogroup-bell-beakers

2.3.4. Haplogroup R1b and Bronze Age expansions

These are the samples and corresponding TreeToM map of Bronze Age groups from Eurasia. Particularly relevant in this period are the emergence of Yamnaya R1b-L23 lineages among Mycenaeans (Z2103 and potentially also xZ2103), and among early Italic-speaking peoples (basal Z2118* and Z2103, apart from L151), which – together with Pre-Tocharian Afanasievo and Pre-Indo-Iranian Poltavka – confirms the association of the expansion of R1b-L23-rich Yamnaya with the spread of Late Proto-Indo-European.

r1b-samples-bronze-age

ancient-r1b-haplogroup-bronze-age

And this is a close-up of Europe:

ancient-r1b-haplogroup-bronze-age-europe

3. Haplogroup R1a East European expansion

3.1. Haplogroup R1a Newick tree

I have used a simplified adjustment of node distances to reflect the TMRCA of early branches (where 0.1 is roughly a thousand years), but then I have kept 0.1 for M417 and subsequent branches to avoid overcrowding.

('R1a':0.1,
 ('(YP4141)':0.1,
  ('YP5018*':0.1
  )YP5018:0.1
 )YP4141:0.5,
 ('M459*':0.1,
 '(M459)':0.1,
  ('(YP1272)':0.1
  )YP1272:0.7,
  ('(M198)':0.1,
   ('(YP1051)':0.1
   )YP1051:0.1,
   ('M417*':0.1,
   '(M417)':0.1,
    ('CTS4385*':0.1,
    '(CTS4385)':0.1
    )CTS4385:0.1,
    ('(Z645)':0.1,
     ('(Z283)':0.1,
      ('(Z282)':0.1,
       ('(PF6155)':0.1,
        ('(M458)':0.1
        )M458:0.1
       )PF6155:0.1,
       ('Y2395*':0.1,
       '(Y2395)':0.1,
        ('(Z284)':0.1,
         ('(Z287)':0.1,
          ('(CTS8401)':0.1
          )CTS8401:0.1
         )Z287:0.1
        )Z284:0.1
       )Y2395:0.1,
       ('(Z280)':0.1,
        ('(Z92)':0.1,
         ('(S24902)':0.1
         )S24902:0.1,
         ('(Y4459)':0.1,
          ('(YP617)':0.1
          )YP617:0.1
         )Y4459:0.1,
         ('(CTS1211)':0.1,
          ('(YP343)':0.1,
           ('(YP340)':0.1
           )YP340:0.1
          )YP343:0.1
         )CTS1211:0.1
        )Z92:0.1
       )Z280:0.1
      )Z282:0.1
     )Z283:0.1,
     ('(Z93)':0.1,
      ('Y39884*':0.1,
      '(Y39884)':0.1,
       ('(Y41571)':0.1
        )Y41571:0.1
     )Y39884:0.1,
      ('(YP1506)':0.1
      )YP1506:0.1,
      ('(YP5585)':0.1
      )YP5585:0.1,
      ('(Z94)':0.1,
       ('Y3*':0.1,
       '(Y3)':0.1
        )Y3:0.1,
       ('(Z2124)':0.1,
        ('(Z2122)':0.1
        )Z2122:0.1,
        ('(Z2125)':0.1,
         ('(YP413)':0.1
         )YP413:0.1,
         ('(S23592)':0.1,
          ('(YP1558)':0.1,
           ('(S23201)':0.1,
            ('(S21872)':0.1
            )S21872:0.1
           )S23201:0.1,
           ('(YP1456)':0.1
           )YP1456:0.1
          )YP1558:0.1
         )S23592:0.1,
         ('(Z2123)':0.1,
          ('(Y20746)':0.1
          )Y20746:0.1,
          ('(Y934)':0.1,
           ('(Y874)':0.1
           )Y874:0.1,
           ('(BY30762)':0.1
           )BY30762:0.1,
           ('(YP523)':0.1,
            ('(YP520)':0.1
            )YP520:0.1
           )YP523:0.1
          )Y934:0.1
         )Z2123:0.1
        )Z2125:0.01
        )Z2124:0.01
      )Z94:0.01
      )Z93:0.1
    )Z645:0.1
   )M417:0.3
  )M198:0.6
 )M459:0.3
)R1a:0.5

3.2. Haplogroup R1a ancient samples

This is the list of R1a cases used for the maps in the required CSV format:

NOTE. I have removed all samples with undefined subclades not informative for later dates (viz. undefined M459 in EMBA, undefined M417 in LBA, etc.). I have also removed the unlikely R1a cases from Cis-Baikal Neolithic samples, as well as the unlikely R-M198 (xZ93) from the Xiaohe cemetery, since it is more straightforward to assume that any Andronovo-related R1a case from Xinjiang should be, in fact, Z93+.

M459*,48.3,35.139,darkgreen
YP5018*,48.91422,33.76493,black
M459*,61.65,35.65,darkgreen
(M459),43.98,26.4,green
M459*,52.22,48.1,darkgreen
Y3*,49.71,37.58,deeppink
(M459),55.6837241,31.4954019,green
(YP1272),45.725651,43.986933,chartreuse
(M459),56.78451992,40.45303345,green
(Z93),46.536389,30.656667,pink
(Z93),43.7,28.516667,pink
(YP1272),59.4266389,28.055402,chartreuse
(M417),48.932,12.259,blue
CTS4385*,52.2966783,17.5455648,indigo
(Z283),59.0967586,25.3486914,lightcyan
(M417),48.932,12.259,blue
(Z94),53.66,50.67,pink
(CTS8401),55.703,11.855,gold
M417*,50.009,10.18,red
(Z645),56.025,14.233,lightblue
(Y2395),55.917796,24.913053,goldenrod
Y2395*,58.4098135,15.6245252,brown
(Y2395),58.325533,26.152339,goldenrod
(CTS4385),50.5447396,13.8351164,darkblue
(Z645),59.0967586,25.3486914,lightblue
Y2395*,58.6018994,26.3499016,brown
(M417),51.42,11.68,blue
CTS4385*,50.5390744,13.8288292,indigo
(Z645),61.658497,17.003742,lightblue
(CTS4385),50.54412,13.834065,darkblue
(CTS4385),51.42,11.68,aqua
(M417),51.42,11.68,blue
(CTS4385),51.42,11.68,darkblue
(CTS4385),51.42,11.68,darkblue
(CTS4385),51.42,11.68,darkblue
(CTS4385),51.42,11.68,darkblue
(CTS4385),51.42,11.68,darkblue
(CTS4385),51.42,11.68,darkblue
(YP1456),52.453,55.16,hotpink
(Z645),50.23,18.22,lightblue
(Z2123),52.91277778,50.99055556,#C71585
(Z2123),50.2249,81.83678333,#C71586
(Z2123),53.876,59.076,#C71587
(CTS1211),55.767926,22.417098,aqua
(Z2123),52.81666667,60.46666667,#C71588
(S24902),52.66,18.88,#20B2AA
(Y874),52.81666667,60.46666667,#C71611
(Z2123),52.81666667,60.46666667,#C71589
(Z93),52.81666667,60.46666667,pink
(Z93),52.81666667,60.46666667,pink
(Z94),52.81666667,60.46666667,purple
(Z2124),52.81666667,60.46666667,fuchsia
(Z2124),52.81666667,60.46666667,fuchsia
(Z2124),52.81666667,60.46666667,fuchsia
(Z2125),52.81666667,60.46666667,fuchsia
(Z2123),52.81666667,60.46666667,#C71590
(Z2123),52.81666667,60.46666667,#C71591
(Z2123),52.81666667,60.46666667,#C71592
(Z93),54.1469927,50.8544665,pink
(Z2123),52.81666667,60.46666667,#C71593
(Z93),52.81666667,60.46666667,pink
(Z2123),48.78176667,76.45303333,#C71594
(Z2123),52.81666667,60.46666667,#C71595
(Z280),54.0702,55.5442,aqua
(Z280),54.0702,55.5442,aqua
(S23592),50.09,74.44,hotpink
(Z93),52.5834,56.8465,pink
(Z2124),50.09,74.44,fuchsia
(Z2123),49.12,75.81,#C71596
(YP5585),54.0702,55.5442,orchid
(Z2124),50.4501,57.04005556,fuchsia
(S23592),47.77798333,72.00968333,hotpink
(Z2123),45.13237778,79.36439444,#C71597
(Z2123),46.7881094,17.8398853,#C71598
(Z2123),48.78176667,76.45303333,#C71599
(Z93),53.12,48.37,pink
(Z2123),53.08,50.36,#C71600
(Z94),54.0702,55.5442,purple
(BY30762),54.0702,55.5442,#C71610
(S23592),50.09,74.44,hotpink
(Z93),42.132249,25.492723,pink
(Z93),48.74175,76.99573333,pink
(Z2123),42.68943333,73.11748333,#C71601
(S23592),55.1924,89.3053,hotpink
(S23592),55.1924,89.3053,hotpink
(S23592),55.1924,89.3053,hotpink
(S23592),55.1924,89.3053,hotpink
(S23592),55.1924,89.3053,hotpink
(YP1456),55.1924,89.3053,hotpink
(Z2123),48.78176667,76.45303333,#C71602
(Z2125),53.87313333,91.46046667,fuchsia
(Z2123),42.68943333,73.11748333,#C71603
(Z93),46.96586667,80.03473333,pink
(S23592),46.96586667,80.03473333,hotpink
(S23592),55.1924,89.3053,hotpink
(Z2124),46.94785,79.98808333,fuchsia
(S23592),54.17931667,91.57886667,hotpink
(S23592),46.94785,79.98808333,hotpink
(Z2124),42.68943333,73.11748333,fuchsia
(Z2123),50.2249,81.83678333,#C71604
(Z93),52.54,50.5,pink
(Z2124),53.03,50.39,fuchsia
(YP1506),48.1,54.44,maroon
(Z2122),50.82,75.6,magenta
(Z280),53.043,57.066,aqua
(Z2123),42.68943333,73.11748333,#C71605
(Z2124),40.54,64.99,fuchsia
(S23592),52.954,90.187,hotpink
(Z2124),53.456,85.447,fuchsia
(Z93),48.5055,88.9621,pink
(YP617),54.362324,23.333318,aqua
(YP520),53.152,91.05,#C71612
(Z93),46.96586667,80.03473333,pink
(Z93),55.51,77.05,pink
(Z2123),48.2144,67.0205,#C71606
(Z93),48.12247971,89.91481606,pink
(Z93),48.12247971,89.91481606,pink
(Z93),48.12247971,89.91481606,pink
(Z93),48.5055,88.9621,pink
(Z93),48.5055,88.9621,pink
(Z2123),50.82,75.6,#C71607
(Z93),50.2249,81.83678333,pink
(Z2123),50.2249,81.83678333,#C71608
(Z93),48.99500819,88.22292153,pink
(Z93),40.37722222,71.855,pink
(Z280),51.89583333,11.04666667,aqua
(Z2123),49.691667,99.696111,#C71609
(Z94),34.75146111,72.40193611,purple
(Z283),59.45,25.133333,lightcyan
(YP340),59.5051279,25.52381115,aqua
(Z2125),54.67583333,90.85361111,fuchsia
(Z93),46.4107,29.4901,pink
(Z93),38.718548,61.692287,pink
(Y3),34.75938611,72.37083333,deeppink
(YP1548),49.060646,75.5032,hotpink
(YP1456),48.48834,74.42547,hotpink

3.3. Haplogroup R1a TreeToM maps

This is the TreeToM map with all samples until the Early Iron Age:

ancient-r1a-subclades-map

3.3.1. Haplogroup R1a Mesolithic – Neolithic expansion

These are the Mesolithic and Neolithic samples up until just before the Corded Ware expansion, and the map they form (with slightly modified colors for easier visualization):

NOTE. The R1a-Y3* sample from Alexandria (I6561) has a radiocarbon date ca. 4000 BC that should be questioned based on its fully formed Corded Ware-like ancestry (different from coeval or later samples from the area), Y3 subclade, and LP alleles appearing 1,000 years earlier than they should, but I include it nevertheless in case its real date corresponds to a Pre-Yamnaya or Yamnaya phase (say, ca. 3500-2900 BC) from the Middle Dnieper.

r1a-samples-mesolithic-neolithic

ancient-r1a-subclades-mesolithic-neolithic-map

3.3.2. Haplogroup R1a in Corded Ware and MLBA Sintashta-Potapovka

It is hard not to notice that the division of R1a-M417 subclades before and during the expansion of Corded Ware groups runs ostensibly through the Volhynian-Podolian Upland, exactly where archaeologists have traditionally estimated the origin of the Corded Ware culture. That is, until the fiasco of conventionally selecting a poorly understood (and catastrophically named) “Yamnaya ancestry” component as defining a single population, which has thrown the field into disarray.

r1a-samples-corded-ware-mlba

ancient-r1a-subclades-corded-ware-map

3.3.3. Haplogroup R1a in the Bronze Age

These are the samples from the Bronze Age until just before the Scythian and later expansions from Asia. The most interesting aspect in Europe is the apparent replacement of most R1a-CTS4385 lineages prevalent in CWC groups of Central Europe, which can be attributed to the massive migration of Bell Beakers.

The later gradual expansion of Z645 subclades in the Eastern EEBA province – seen in the Bronze Age samples from Iwno-Trzciniec and Chłopice-Veselé (in the previous map), and later in the Halberstadt LBA sample – suggest resurgence events and bottlenecks among expanding Balto-Slavs.

r1a-samples-mlba-bronze-age

ancient-r1a-subclades-bronze-age

Here is a close-up of the R1a-Z93 tree and samples in Eurasia, which point (together with later samples, such as those from modern Finno-Ugric peoples and Hungarian Conquerors) to the presence of basal subclades in the Andronovo-like Horizon, stemming from Abashevo migrants through the Seima-Turbino expansion.

ancient-r1a-z93-subclades-bronze-age

4. Haplogroup N Siberian expansion

The assessment of haplogroup N is complicated by the current lack of a proper regional and temporal transect of Siberia. Nevertheless, the different available patterns in selected periods shed some light on the most likely causes of its bottlenecks.

4.1. Haplogroup N Newick tree

This is the Newick tree for the reported N subclades (no distance between nodes for clarity):

('(N)':0.01,
('(N1)':0.01,
 ('N1a*':0.01,
 '(N1a)':0.01,
  ('N1a1':0.1,
   ('M178*':0.1,
   '(M178)':0.1,
    ('Tat*':0.1,
    '(Tat)':0.1,
     ('(L708)':0.1,
      ('(M2126)':0.1,
       ('(M2019)':0.1,
        ('(M2058)':0.1,
         ('(A9408)':0.1,
          ('(PH1612)':0.1,
           ('(PH1896)':0.1
           )PH1896:0.1
          )PH1612:0.1
         )A9408:0.1,
         ('(M2016)':0.1,
          ('(M1993)':0.1
          )M1993:0.1
         )M2016:0.1
        )M2058:0.1
       )M2019:0.1,
       ('L1026*':0.1,
       '(L1026)':0.1,
        ('(Y6058)':0.1,
         ('(Y16323)':0.1,
          ('(F4205)':0.1,
           ('(Y16220)':0.1,
            ('(F22331)':0.1
            )F22331:0.1
           )Y16220:0.1
          )F4205:0.1
         )Y16323:0.1,
         ('(CTS10760)':0.1,
          ('(VL29)':0.1,
           ('(L550)':0.1,
            ('(F22331)':0.1
            )F22331:0.1,
            ('(L1025)':0.1,
             ('(Y4706)':0.1
             )Y4706:0.1
            )L1025:0.1,
            ('(Y4341)':0.1,
             ('(Y4339)':0.1
             )Y4339:0.1
            )Y4341:0.1
           )L550:0.1,
           ('(CTS9976)':0.1,
            ('(N1022)':0.1,
             ('(Y5005)':0.1,
              ('(Y5003)':0.1
              )Y5003:0.1
             )Y5005:0.1
            )N1022:0.1
           )CTS9976:0.1
          )VL29:0.1
         )CTS10760:0.1
        )Y6058:0.1,
        ('(Z1936)':0.1,
         ('(Z1934)':0.1
         )Z1934:0.1,
         ('(Y13850)':0.1
         )Y13850:0.1
        )Z1936:0.1
       )L1026:0.1
      )M2126:0.1
     )L708:0.1
    )Tat:0.1
   )M178:0.1
  )N1a1:0.1,
  ('(L666)':0.01
  )L666:0.1
 )N1a:0.1,
 ('(N1b)':0.01,
  ('(Z4781)':0.01
  )Z4781:0.1
 )N1b:0.1
)N1:0.1,
 ('N2*':0.01,
 '(N2)':0.01,
  ('N2a*':0.01,
  '(N2a)':0.01
  )N2a:0.1
 )N2:0.1
)N;

4.2. Haplogroup N ancient samples

These are the available samples in chronological order:

(Z4781),42.9995032,125.9816054,green
(L666),52.286944,104.249167,salmon
(L666),51.698333,103.703056,salmon
(L666),51.698333,103.703056,salmon
(L666),51.698333,103.703056,salmon
(N1b),40.96330795,119.0258789,green
(N1b),40.96330795,119.0258789,green
(N1b),40.96330795,119.0258789,green
(N1b),40.96330795,119.0258789,green
(N1b),40.949724,113.2540836,green
(N1b),40.949724,113.2540836,green
(N1b),40.949724,113.2540836,green
Tat*,53.188889,103.368056,deeppink
N2a*,53.305983,67.648166,purple
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
(N1b),42.18025,119.1854168,green
N2a*,50.46,49.2,purple
N2a*,50.46,49.2,purple
(N2),51.698333,103.703056,blue
(N1b),42.35675615,120.1673876,green
(N1b),42.35675615,120.1673876,green
(N1b),42.35675615,120.1673876,green
L1026*,69.0618033,36.381134,red
L1026*,69.0618033,36.381134,red
(L1026),49.691667,99.693333,violet
(N2),56.016,92.866,blue
N2*,47.7300788,20.0910629,red
(N1b),41.9202562,118.7020825,green
(L1026),59.5,26.533333,violet
(L550),59.55,26.15,goldenrod
(L550),58.406944,21.993611,goldenrod
(M178),42.159444,77.406389,pink
(F22331),52.61805556,76.74944444,firebrick
(L1026),56.5046381,16.4293858,violet
(L1025),55.743786,24.228036,goldenrod
(Y16323),46.2219,20.4809,pink
(F4205),47.4244,19.0661,firebrick
(F4205),45.377,21.7188,firebrick
(L1026),47.0244,19.1236,pink
(Y16323),47.0612,19.2754,brown
(Y16323),47.0612,19.2754,brown
(Y16323),46.5816,20.2266,brown
(Y16323),46.5064,20.5377,brown
(F4205),46.8964,19.6897,firebrick
(F4205),47.0612,19.2754,firebrick
(F4205),47.0612,19.2754,firebrick
(F4205),46.6242,19.8585,firebrick
(F4205),47.0612,19.2754,firebrick
(Y16323),46.6225,19.2875,brown
(F4205),46.9755,19.0171,firebrick
(F4205),47.0612,19.2754,firebrick
(F4205),47.0612,19.2754,firebrick
(F4205),46.7628,19.7533,firebrick
(F4205),46.7628,19.7533,firebrick
(F4205),46.7628,19.7533,firebrick
(F4205),46.7628,19.7533,firebrick
(F4205),46.7628,19.7533,firebrick
(F4205),46.7628,19.7533,firebrick
(F4205),46.7628,19.7533,firebrick
(VL29),58.17,22.25,orange
(L550),58.17,22.25,goldenrod
(L550),58.17,22.25,goldenrod
(L550),58.17,22.25,goldenrod
(L550),58.17,22.25,goldenrod
(L550),58.17,22.25,goldenrod
(L550),67.29,14.42,goldenrod
(Y4706),58.17,22.25,goldenrod
(PH1896),48.3402,22.1211,magenta
(Z1936),48.2004,21.5311,cyan
(Z1936),48.2004,21.5311,cyan
(Z1936),48.4242,21.6109,cyan
(Z1936),48.3794,21.758,cyan
(Z1936),48.4242,21.6109,cyan
(M2019),48.327874,21.719835,magenta
(M2019),48.327874,21.719835,magenta
(L550),56.8057977,16.7301606,goldenrod
(Y13850),47.2749,20.4469,cyan
(L1034),46.9359,20.0969,cyan
(PH1612),46.8306,20.7344,magenta
(VL29),57.34,18.19,orange
(L550),57.63,18.28,goldenrod
(Y5005),57.34,18.19,goldenrod
(VL29),54.78,31.87,orange
(L550),58.38,13.65,goldenrod
(L550),58.38,13.65,goldenrod
(Y4339),59.616667,17.716667,goldenrod
(Y5003),58.38,13.65,goldenrod
(M1993),70.7,135.4,magenta
(L550),59.4275,26.709444,goldenrod
(L550),58.516667,22.7,goldenrod
(L550),58.057222,26.498333,goldenrod

4.3. Haplogroup N TreeToM maps

ancient-dna-n1a-haplogroup

4.3.1. Haplogroup N Mesolithic and Neolithic expansions

These are the early N subclades available and the corresponding map, which shows the most likely origin of the split of N lineages west of Lake Baikal:

NOTE. I did not include the reported N-Tat of a Comb Ware group by Chekunova because of the dubious combination of subclade and radiocarbon date in that location, although admittedly R1a-Tat is old enough to appear anywhere during the Neolithic…

n1c-samples-mesolithic-neolithic

ancient-n1a-haplogroup-neolithic

4.3.2. Haplogroup N Eurasian Bronze Age

These are the available Bronze Age samples until just before the (Pre-Scythian) nomadic expansions. Notice the L1026* from Lovozero related to the expansion of Palaeo-Arctic peoples, and the L1026 from Khövsgöl in the Eastern Steppes, a region under the impact of Karasuk-related ancestry, suggesting its presence among (and expansion with) the Scythian communities of Siberia. Both cases help explain the later emergence of the so-called “Siberian ancestry” in Eastern and Northern Europe.

n1c-samples-bronze-age

ancient-n1a-haplogroup-bronze-age

4.3.3. Haplogroup N and Iron Age nomads

The westward expansion of nomads throughout Northern Eurasia brought different haplogroups from Siberia across the Urals, with hg. N probably already prevalent among Iranian-, East Uralic-, and Altaic-speaking populations of Asia and possibly already on both sides of the Urals due to the integration of forest hunter-gatherers among expanding Corded Ware-related groups.

In the case of hg. N1a-VL29 (from a branch Y6058 in common with Avar N1a-F4205 lineages), its expansion around the Baltic (including Scandinavia and the West Baltic) is preceded by a minimal infiltration of likely origin in marriage alliances of Akozino warrior-traders, who had in turn intense contacts with nomads to the south (also attested in Iranian loans among Uralic dialects), with no discernible recent impact over the Baltic linguistic landscape.

n1c-samples-iron-age

ancient-n1a-haplogroup-iron-age

4.3.4. Haplogroup N in the Middle Ages

The Siberian origin of N lineages pushed into the Cis-Urals by Altaic-speaking peoples – some of which eventually spread with the Hungarian Conquerors – can be inferred from previous Iron Age expansions, as well as from the modern wide distribution of N-Y13850 subclades among disparate Turkic-speaking populations, from Turan to Samara in the north, and to Anatolia in the south.

Similarly, many N1a-VL29 (especially N1a-L550) subclades found in the East Baltic today spread eastwards with Germanic-speaking peoples during the Viking migrations in the Middle Ages, adding to the known radical founder effects that happened roughly at the same time as Finnic spread to the north and east from its Estonian homeland.

n1c-samples-middle-ages

ancient-n1a-haplogroup-middle-ages

Conclusion

There is an obvious complementary distribution of R1b-M269 vs. R1a-M198, particularly R1b-L23 vs. R1a-M417, precisely during the relevant period of Indo-European vs. Uralic splits and expansions from Eastern Europe.

Only after the evolution of Bell Beakers into the European Early Bronze Age cultures, and of Poltavka herders into the Sintashta-Potapovka-Filatovka community, appeared different R1a subclades integrated into disparate (previously R1b-rich) Indo-European-speaking groups, expanding further under distinct regional bottlenecks corresponding to already developing dialects.

And only much later did N subclades, probably widely distributed among Palaeosiberian peoples during the Seima-Turbino phenomenon, start to have an impact within R1a-rich Uralic-speaking communities around the Urals, Arctic, and West Siberia, at the same time as other haplogroups (such as I1, I2, E1b, J2, or J1 subclades) either resurged or infiltrated and later expanded with Indo-European-speaking societies in Europe, too.

haplogroup-subclade-bottleneck
Simplified model of evolution of an ethnolinguistic community in terms of Y-chromosome haplogroups and admixture. I created it to explain the Balto-Slavic evolution in the South-Eastern Baltic, roughly depicting a (A1) Yamnaya replacement of (B) Corded Ware with (A2) Bell Beakers in the West Baltic / Northern European Plain, then (A3) resurgence events on their way to the East Baltic, and then the (A4) infiltration and expansion of haplogroup N. See full comment in its context.

It has already become evident (see e.g. the recent Mittnik et al. 2019) the relevance of a proper SNP inference to track fine-scale population movements together with ancestry estimations, in order to properly distinguish potential prehistoric language expansions. It is amazing that in 2020 many labs are still working with outdated software or simplistic automated inferences for haplogroup assignments, instead of carefully reviewing each sample’s available data by hand.

See also

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Syfyfan
Syfyfan

Hello 🙂

There will be a project called HistoGenes, which is a hungarian project with other countries’ collaboration. It’s about East europe in 400AD-900AD. I know it’s in hungarian, but I hope that it will be available in other languages. The archeologists, genetics, and lot of other science will work together to figure out what happened in this period of time. Can you check it and follow it?

Furioso
Furioso

Have you watched this?: https://youtu.be/Pe4jnBdVxjw

AK.
AK.

R1b-L23+ Sintashta and R1b+ L23 Yamnaya/Afansievo/Bell Beaker, derive from the same source.

A.K.
A.K.

Steppe ancestry evolution/ mutations/locations- within Yamnaya proper R1b L23+ & and proto Yamnaya R1b V1636+clusters.

R1b-L23&R1bV1636 dated samples + Yfull estimates[L23+ formed 6400 ybp,& V1636+formed 15600 ybp.]

I0443, Yamnaya/North,Samara, 3300-2700 BCE–L23+
[(L49+, L23+, PF6399+, L150+, L1353+, PF6509+, M269+, CTS12478+, L51-, Z2105-)]
Yfull-R-L23PF6404 * L478/PF6403 * L23/S141/PF6534formed 6400 ybp, TMRCA 6100 ybpinfo

1}I0122, ,Khvalynsk,Volga, 4700 BC-R1b-V1636+
2}PG2001, PG2004, Eneolithic steppe,Northern Caucasus, 4336-4047 BC-R1b-V1636+
3}I1635, Kura-Araxes,Armenia, 2600-2400 BC-R1b-V1636+
Yfull-R-V1636Y36360 * BY21443 * BY15342+96 SNPsformed 15600 ybp, TMRCA 6600 ybpinfo

A.K.
A.K.

Interesting geographic connections between different Yamnaya mtdna lines on the steppe. Two close proximity samples I0443 and I0439. Samara bend.

I0443, Yamnaya/North,Samara, 3300-2700 BCE–R1b L23+ Z2105-
W3a1a/W3a

1}poz222 YAM Bronze Age Ukraine F -3023
2}poz208 YAM Bronze Age Ukraine U -2882
3}I3607 BBC Copper Age Germany M -2350
3}I0116 UNC Bronze Age Germany M -2134
4}I4332 BABA Bronze Age Croatia F -1618
5}SCY196 SCU Iron Age Moldova U -400

I0439+I0438, Yamnaya/North, Samara Bronze Age Russia M -3321/3021-R1b-Z2109+ KMS-75+
U5a1a1

1}RISE508 BARu Bronze Age Russia F -3331
2}RISE507 BARu Bronze Age Russia F -3322
3}I4132 BBC Copper Age Germany M -2500
4}I6481 BBC Copper Age Germany M -2500
5}RISE560 BBC Neolithic Germany M -2500
6}HUGO_171 BBC Bronze Age Germany F -2387
7}I4252 BBC Copper Age Poland M -2285
8}I1767 BBC Bronze Age Great Britain M -2200
9}I4069 BBC Copper Age The Netherlands M -2188
10}POST_6 EBAGe Bronze Age Germany M -2009
11}I7577 BABI Bronze Age Great Britain M -1386
12}Kivutkalns153 BABL Bronze Age Latvia M -800
13}DA53 IAKa Iron Age Kazakhstan F -453
14}SCY197 SCU Iron Age Moldova M -400
15}PCA0052 MAPl Middle Ages Poland F 110

Carlos Quiles
Carlos Quiles

I think RISE508 and RISE507 are the same sample.

Pálfalvi László
Pálfalvi László

Hello
Have you read the article about Sardinia’s genome in history? It’s very interesting.

A.K.
A.K.

Nice job,[ regardless of interpretation geographic parsimonious spread] in visualizing/showing, distribution of relevant snp’s to the debate.

Hodo Scariti
Hodo Scariti

Very interesting blog entry.

Only a question: I saw in the first map that you have linked Villabruna R1b sample to V2219*… may I ask for the source of that association? I remained on Villabruna as R1b on its path to P297.

Thanks for attention.