Origin of the R1b_KN subclade: a celtic migration from Armorica

Origin of the R1b_KN subclade: a celtic migration from Armorica?

John McEwan

12 July 2005

Background

The phase 1 and phase 2 analyses both identified a subclade tentatively labeled R1bi. Ken Nordtvelt had previously identified this cluster independently. To make the subclade more memorable and simpler to quote in future I will refer to it as R1b_KN. The majority of the subclade individuals appeared to be clustered in the Strathclyde region, at least from the results of my phase 1 analysis, which only used selected surnames. Ken observed the same phenomenon in his analysis. The subclade from the phase 1 analysis also appeared to be under-represented in Ireland, relative to other putative R1b clusters. However, a visual examination of this subclade in the 37 STR analysis of R1b identified a number of unusual properties in this subclade’s shape and surname distribution that deserved a closer examination. The relevant section of the 37 STR phylogram is shown below.

Figure 1. Overall phylogram of the 37 STR cluster analysis of R1a and R1b haplogroups (left) and an enlarged view of R1b_KN (shaded pink), individuals are shown with their haplogroup and sample identifier appended.

Visual pattern

The first observation is the shape of the subclade, it emerges from a single “neck” and extends upwards, with those near the top more closely related, while the “neighbours” at the bottom are more distantly related. In fact the cluster could be extended (and originally was) to include individuals down to Andriesen. Subsequent comments will include these individuals as well. How could such a subclade shape emerge? Well if it is not a statistical artifact, it could be a result of a migration. People closer to the bottom are remnants left behind as the group or tribe moved to a new location. At the end of the migration people would have had less time to diverge and branches would be shorter. In fact the movement could be dated from when they first started to migrate, the intervening branches and when they finally settled down. Deep branches, like the group from Kane to Andrieson could represent the founder population or a bifurcation i.e. split or branching in the migration pattern. Note a similar, but less marked pattern is also evident in the R1ba subclade that David Wilson has previously described. I will in future call this subclade R1b_DW.

Is this interpretation probable and what independent evidence can be assembled? There are three separate lines of evidence, the first is to calculate the TMRCA for the nodes along the branch and try to assign this to a known historical or “folk” migration. The second is to examine the likely origin of the surnames (or the location of the paternal ancestors given by the submitter). The first option, surnames, provides some indication of where their ancestors were around 1000-1200 AD, while the second is typically later than 1800AD. If ages of TMRCA are plotted against geographical location and some form of trend or correlation is observed then this is independent evidence that the migration was real. It also provides a separate way to identify and date the probable migration path. If the calculated correlation is significant then regardless of errors in assigning surname the pattern is likely to be real, albeit the subsequent interpretation may not be. The final source of evidence is the proportion of the cluster relative to the database size and how that relates to known historical migrations and population sizes.

Ancestor origin

A visual examination identified that names on the bottom of the cluster, Parker-Young are generally English, and those above lowland Scots with many from Strathclyde. The two out groups Kane-Donohue are primarily pre-Viking Celtic Irish while the final group is English with one European. Below in table 1 are listed the origin of ancestors as given in Ysearch in cluster order. The given origins have been color coded to ease interpretation again it can be seen that there is a distinct gradation from the bottom of the cluster from Europe or England to Scotland at the top. The Irish primarily appear as a branch within England.

The divergence times for individuals were also calculated using http://www.mymcgee.com/tools/yutility.html . The figures below are approximate and are those for the intersection of the individual at the “tip of the cluster” i.e. the path traverses along the “backbone” of the subclade cluster. Campbell_R1b_8CBUR was chosen as this person. Divergence times for each individual are also given for the R1b modal value which acts as a common outgroup from which all within the group were descended. Perhaps not surprisingly, TMCRA trend from 1200 years to in excess of 3000 years. The most distant branches are at a similar distance to the modal value of R1b.

Table 1. List of names in the extended R1b_KN cluster and origin of paternal ancestor as given in Ysearch (color coded for Scotland, England, Ireland and Europe). The order listed is the cluster order, but may differ slightly from the tree graph above because of plotting conventions. Time to most recent common ancestor versus individual 8CBUR is listed and summarized from table 4.

Name	Given origin in Ysearch	TMRCA yrs vs 8CBUR
Campbell_R1b_8CBUR	Scotland	0
McCutchan_R1b_4C39R	West Meath, Ireland	1650
Campbell_R1b_K3XKQ	Kilberry, Scotland	1650
Campbell_R1b_9MJ45	Inveraray, Scotland	1650
Urquhart_R1b_JWBVE	Knockbain, Scotland	1230
MacEachern_R1b_8K5XS	Scotland	2100
MacDonald_R1b_JZQQX	Idrigil, Isle of Skye, Scotland	1860
Forbes_R1b_4ASGK	Scotland	1650
Anderson_R1b_H5535	Northumberland, England	1860
Hamilton_R1_AM847	Scotland	2340
MacDonald_R1b_F4XG9	Knockintorran, Scotland	2340
Livingston_R1b_Q5BR3	Scotland	2100
McGregor_R1b_4WAZU	Monzie Parish, Scotland	1860
Stirling_R1b_KNES8	Perthshire, Scotland	1860
Stirling_R1b_RS2P7	Perthshire, Scotland	2100
McGregor_R1b_VBSFV	Scotland	2340
Stirling_R1b_GDW3X	Kinnettles, Angus, Scotland	1650
McMillan_R1b_V5URN	Scotland	2340
McMillan_R1b_YA5PE	Scotland	2340
McMillan_R1b_6UWW4	Midlothian, Scotland	2100
Anderson_R1b_YNXGB	Isle of Skye, Inverness-shire, Scotland	2100
McKinlay_R1b_7X6HZ	Row, United Kingdom	1860
Stein_R1b_E48PN	Scotland	1860
White_R1b_E6W6F	United Kingdom	2580
Jamieson_R1b_YRADM	Moray, Scotland	1650
Tagert_R1b1c_MABKF	Stewart county, Tennessee, USA	2340
Anderson_R1b_77P34	Lincolnshire, England	2340
Anderson_R1b_A7P6F	Lincolnshire, England	2340
Kincaid_R1b_Q2YWJ	Chester County, Pennsylania, Scotland	2580
McMillan_R1b_WBDKF	Scotland	2580
Young_R1b_CSRF9	Scotland	2340
Alexander_R1b_SQV7S	Raphoe, Donegal, Ireland	2100
Gray_R1b_VBP44	Scotland	3390
MacCorkle_R1b_Q777T	Argyllshire, Scotland	2580
Wilkinson_R1b_KFM4M	England	2850
Wilkinson_R1b_T8VPT	Ireland	2850
Hall_R1b_B4WFM	Ashbourne, England	2580
Wilson_R1b_3UEUC	Shetland Isles, Scotland	3390
Walker_R1b_MWE53	England	3120
Simmons_R1b_CV74C	England	3390
Parker_R1b_Z5PNU	England	3690
Clark_R1b_S286F	Hertfordshire area, England	3990
ODonoghue_R1b_2FMC4	Cavan, Ireland	3390
Donohoe_R1b_M7V3H	Ballyconnell, Cavan, Ireland	3390
Kane_R1b_NE5HD	Alverton, Ireland	1860
Barrett_R1b_W9TVC	Andover, Hampshire/Hantshaving, England	2850
Turk_R1b_2BFE5	ANTRIM, Ireland	2580
Blakeston_R1b_QRKMM	Beverley, East Riding, England	3390
Andriesen_R1b_UA5J3	Rotterdam, Netherlands	2850
Ming_R1b_Z8YYY	United Kingdom	2340
R1b		3120

In order to find the probable times of settlement in Scotland, England and Ireland the mean and SEM was calculated for each country of origin in table 3 below. The estimates suggest the group reached Scotland around 150BC, and England around 410BC, a branch reached Ireland about the same time as Scotland either via England or via Scotland, most likely England. The single sample from Europe suggests an even earlier divergence. When tested using a 2 tailed t-test, and assuming heterogeneous variances, the difference in TMCRA between Scotland and England was significant (p=0.004) i.e. you would expect to see this pattern 4 times in 1000 simply by chance.

Table 2. Summary of times to most recent common ancestor of 8CBUR for the individuals from various locations from table 1

origin	n	mean TMRCA vs 8CBUR	SEM
Scottish	26	2153	126
Irish	8	2228	250
English	15	2612	168
Europe	1	2850

Size of the migration

There are 50 individuals in the expanded cluster and 37 in the R1b_KN sub clade out of 713 individuals in total. It is well known that the Ysearch database is biased, but we can still obtain conservative estimates. It has been estimated that Roman Britain had around 2-3 million people (eg http://www.dartfordarchive.org.uk/early_history/politics.shtml) and they probably had expanded rapidly in the previous 500 years from perhaps 150,000 at the start of the iron age. Assuming all pre-Roman people in England were predominantly R1b, correcting for the proportion 713 haplotypes for those of R1a, Scandinavian and Western Europe origin provides an estimate of 74% of the total sampled were British or Irish origin i.e. 0.74 x 713 = 528 R1b haplotypes. If we also assume no preferential reproductive success for the haplotypes since that time, the cluster represents at least 37/528 x 2,000,000 = 140,000 and at most 50/528 x 3,000,000 = 284,000 people at the time of the Roman invasion. This population size and R1b haplotype is inconsistent with this group being part of a Roman associated immigration from Europe, given known numbers of Romans that came to Britain. It is recorded that the garrison itself was only originally 100, 000 soldiers reducing to 20,000 in later years. It is also extremely unlikely that the Roman soldiers made a beeline to settle in Strathclyde. In contrast if this group were part of an earlier iron age migration then the numbers become 37/528 x 150,000 = 10,500 to 50/528 x 150,000 = 14,200. These latter numbers are probably far more realistic. This would lend support to the migration being in pre-Roman times, but almost certainly one of the later Celtic migrations (otherwise it would not have the narrow necked feature consistent with a later immigration). The most consistent with the evidence are the Belgae, Laginian, or Goidelic invasions. Of these oral history suggests that the Laginian invasion is the best given the sparse evidence. It has been recorded that one tribe of the Laginian invasion (dated as ~300BC), the Dumnonii, settled in Scotland where they founded Dumbarton and established the kingdom later known as Strathclyde (http://en.wikipedia.org/wiki/Early_history_of_Ireland#The_Laginian_invasion ). This is the centre of origin for surnames associated with the later most recent twigs at the top of the R1b_KN cluster. The date also spans the period of MRCA for the English and Scottish individuals. However, these estimates are dependent on many assumptions so some care is required in their interpretation.

Summary

What pattern is consistent with the observations and information available for the R1b_KN subclade? Well there are several, given the inaccuracy and uncertainty implicit in these calculations. However, the Pretanic invasion is an unlikely source, because of the observed R1b_KN subclade pattern and age, as are the Belgae and Goidelic by the distribution of surnames within the cluster. Finally, the pattern is inconsistent with a post Roman invasion migration based on recorded history, and the numbers involved. This leaves the Laginian Dumnonii who are recorded to have settled in the Strathclyde region some time after ~300BC and which also is the region with the highest concentration of surname origins within the subclade. The geographical grading of the surnames in the cluster with TMCRA divergence is supporting evidence. The Irish Kane-Donohoe “branch” also suggests that the invasion of Ireland was probably via England not Scotland based, but the evidence is weak. It is recorded that a related tribe, the Lagin, invaded and settled in Leinster.

Further material

The raw data for the extended cluster, and the results from the TMCRA and assumptions used are shown below.

Table 3. Raw data from the subclade in cluster order plus external modal haplotypes.

Ysearch Database Configuration - DNA Results Comparison

D
Y
S
3
9
3

D
Y
S
3
9
0

D
Y
S
1
9
/
3
9
4

D
Y
S
3
9
1

D
Y
S
3
8
5
a

D
Y
S
3
8
5
b

D
Y
S
4
2
6

D
Y
S
3
8
8

D
Y
S
4
3
9

D
Y
S
3
8
9
-
1

D
Y
S
3
9
2

D
Y
S
3
8
9
-
2

D
Y
S
4
5
8

D
Y
S
4
5
9
a

D
Y
S
4
5
9
b

D
Y
S
4
5
5

D
Y
S
4
5
4

D
Y
S
4
4
7

D
Y
S
4
3
7

D
Y
S
4
4
8

D
Y
S
4
4
9

D
Y
S
4
6
4
a

D
Y
S
4
6
4
b

D
Y
S
4
6
4
c

D
Y
S
4
6
4
d

D
Y
S
4
6
0

G
A
T
A
-
H
4

Y
C
A
-
I
I
a

Y
C
A
-
I
I
b

D
Y
S
4
5
6

D
Y
S
6
0
7

D
Y
S
5
7
6

D
Y
S
5
7
0

C
D
Y
a

C
D
Y
b

D
Y
S
4
4
2

D
Y
S
4
3
8

modal

R1b_given

R1b_dw_given

R1b_KN_given

R1a_given

I1_given

Campbell_R1b_8CBUR

McCutchan_R1b_4C39R

Campbell_R1b_K3XKQ

Campbell_R1b_9MJ45

Urquhart_R1b_JWBVE

MacEachern_R1b_8K5XS

MacDonald_R1b_JZQQX

Forbes_R1b_4ASGK

Anderson_R1b_H5535

Hamilton_R1_AM847

MacDonald_R1b_F4XG9

Livingston_R1b_Q5BR3

McGregor_R1b_4WAZU

Stirling_R1b_KNES8

Stirling_R1b_RS2P7

McGregor_R1b_VBSFV

Stirling_R1b_GDW3X

McMillan_R1b_V5URN

McMillan_R1b_YA5PE

McMillan_R1b_6UWW4

Anderson_R1b_YNXGB

McKinlay_R1b_7X6HZ

Stein_R1b_E48PN

White_R1b_E6W6F

Jamieson_R1b_YRADM

Tagert_R1b1c_MABKF

Anderson_R1b_77P34

Anderson_R1b_A7P6F

Kincaid_R1b_Q2YWJ

McMillan_R1b_WBDKF

Young_R1b_CSRF9

Alexander_R1b_SQV7S

Gray_R1b_VBP44

MacCorkle_R1b_Q777T

Wilkinson_R1b_KFM4M

Wilkinson_R1b_T8VPT

Hall_R1b_B4WFM

Wilson_R1b_3UEUC

Walker_R1b_MWE53

Simmons_R1b_CV74C

Parker_R1b_Z5PNU

Clark_R1b_S286F

ODonoghue_R1b_2FMC4

Donohoe_R1b_M7V3H

Kane_R1b_NE5HD

Barrett_R1b_W9TVC

Turk_R1b_2BFE5

Blakeston_R1b_QRKMM

Andriesen_R1b_UA5J3

Ming_R1b_Z8YYY

Distance from reference:

Zero

One

Two

Three+

Table 4. TCMRA using assumptions at the bottom of the table and data from the above table

Time to Most Recent Common Ancestor (Years)

m
o
d
a
l

R
1
b
_
g
i
v
e
n

R
1
b
_
d
w
_
g
i
v
e
n

R
1
b
_
K
N
_
g
i
v
e
n

R
1
a
_
g
i
v
e
n

I
1
_
g
i
v
e
n

C
a
m
p
b
e
l
l
_
R
1
b
_
8
C
B
U
R

M
c
C
u
t
c
h
a
n
_
R
1
b
_
4
C
3
9
R

C
a
m
p
b
e
l
l
_
R
1
b
_
K
3
X
K
Q

C
a
m
p
b
e
l
l
_
R
1
b
_
9
M
J
4
5

U
r
q
u
h
a
r
t
_
R
1
b
_
J
W
B
V
E

M
a
c
E
a
c
h
e
r
n
_
R
1
b
_
8
K
5
X
S

M
a
c
D
o
n
a
l
d
_
R
1
b
_
J
Z
Q
Q
X

F
o
r
b
e
s
_
R
1
b
_
4
A
S
G
K

A
n
d
e
r
s
o
n
_
R
1
b
_
H
5
5
3
5

H
a
m
i
l
t
o
n
_
R
1
_
A
M
8
4
7

M
a
c
D
o
n
a
l
d
_
R
1
b
_
F
4
X
G
9

L
i
v
i
n
g
s
t
o
n
_
R
1
b
_
Q
5
B
R
3

M
c
G
r
e
g
o
r
_
R
1
b
_
4
W
A
Z
U

S
t
i
r
l
i
n
g
_
R
1
b
_
K
N
E
S
8

S
t
i
r
l
i
n
g
_
R
1
b
_
R
S
2
P
7

M
c
G
r
e
g
o
r
_
R
1
b
_
V
B
S
F
V

S
t
i
r
l
i
n
g
_
R
1
b
_
G
D
W
3
X

M
c
M
i
l
l
a
n
_
R
1
b
_
V
5
U
R
N

M
c
M
i
l
l
a
n
_
R
1
b
_
Y
A
5
P
E

M
c
M
i
l
l
a
n
_
R
1
b
_
6
U
W
W
4

A
n
d
e
r
s
o
n
_
R
1
b
_
Y
N
X
G
B

M
c
K
i
n
l
a
y
_
R
1
b
_
7
X
6
H
Z

S
t
e
i
n
_
R
1
b
_
E
4
8
P
N

W
h
i
t
e
_
R
1
b
_
E
6
W
6
F

J
a
m
i
e
s
o
n
_
R
1
b
_
Y
R
A
D
M

T
a
g
e
r
t
_
R
1
b
1
c
_
M
A
B
K
F

A
n
d
e
r
s
o
n
_
R
1
b
_
7
7
P
3
4

A
n
d
e
r
s
o
n
_
R
1
b
_
A
7
P
6
F

K
i
n
c
a
i
d
_
R
1
b
_
Q
2
Y
W
J

M
c
M
i
l
l
a
n
_
R
1
b
_
W
B
D
K
F

Y
o
u
n
g
_
R
1
b
_
C
S
R
F
9

A
l
e
x
a
n
d
e
r
_
R
1
b
_
S
Q
V
7
S

G
r
a
y
_
R
1
b
_
V
B
P
4
4

M
a
c
C
o
r
k
l
e
_
R
1
b
_
Q
7
7
7
T

W
i
l
k
i
n
s
o
n
_
R
1
b
_
K
F
M
4
M

W
i
l
k
i
n
s
o
n
_
R
1
b
_
T
8
V
P
T

H
a
l
l
_
R
1
b
_
B
4
W
F
M

W
i
l
s
o
n
_
R
1
b
_
3
U
E
U
C

W
a
l
k
e
r
_
R
1
b
_
M
W
E
5
3

S
i
m
m
o
n
s
_
R
1
b
_
C
V
7
4
C

P
a
r
k
e
r
_
R
1
b
_
Z
5
P
N
U

C
l
a
r
k
_
R
1
b
_
S
2
8
6
F

O
D
o
n
o
g
h
u
e
_
R
1
b
_
2
F
M
C
4

D
o
n
o
h
o
e
_
R
1
b
_
M
7
V
3
H

K
a
n
e
_
R
1
b
_
N
E
5
H
D

B
a
r
r
e
t
t
_
R
1
b
_
W
9
T
V
C

T
u
r
k
_
R
1
b
_
2
B
F
E
5

B
l
a
k
e
s
t
o
n
_
R
1
b
_
Q
R
K
M
M

A
n
d
r
i
e
s
e
n
_
R
1
b
_
U
A
5
J
3

M
i
n
g
_
R
1
b
_
Z
8
Y
Y
Y

modal

1020

2580

840

3840

9330

2100

1860

2100

1860

1230

1440

1230

840

1230

2100

1860

1650

1230

1020

840

1020

1230

1440

840

1020

1650

840

660

1230

840

1440

1020

1650

1440

2100

1440

2100

1230

2340

1650

3120

2100

1440

1020

2340

1860

1230

R1b_given

1020

1650

1440

4830

8310

3120

2850

2580

1860

2580

1860

1650

2340

2580

2850

1860

2100

1860

2100

2340

1440

1230

2580

1650

1440

1650

2100

1230

1860

1230

2340

1230

1860

1440

2100

1860

2850

2580

2100

1650

1440

1650

2100

1440

R1b_dw_given

2580

1650

3120

4830

9330

3990

3390

3120

4320

3120

4320

3690

3120

3990

4320

3390

3120

2850

3690

3120

2580

3690

2340

2850

3690

2580

3390

3690

3120

2580

2340

3120

2850

3120

3990

3120

2580

2850

2580

R1b_KN_given

840

1440

3120

3840

9330

1650

1440

1650

1230

660

1020

660

300

1020

1440

840

1230

1020

1440

1230

1860

1650

840

450

660

1650

660

1440

1230

1440

2100

1020

660

1650

1230

2340

1860

1650

2850

2100

4320

3990

2850

1650

1860

2850

2340

1860

R1a_given

3840

4830

3840

4320

3840

4320

3420

4320

3840

3420

3840

4320

4830

4320

3840

4320

3840

4320

3420

3840

3420

3840

4830

4320

4830

4320

5400

4830

4320

3840

5400

4830

3420

5400

3840

4320

3000

5400

I1_given

9330

8310

9330

4320

10530

9330

8310

9330

8310

9330

8310

9330

8310

10530

9330

8310

9330

8310

9330

8310

9330

8310

7440

8310

9330

7440

8310

9330

8310

7440

8310

7440

6690

8310

12000

8310

9330

Campbell_R1b_8CBUR

2100

3120

3990

1650

4320

10530

1650

1230

2100

1860

1650

1860

2340

2100

1860

2100

2340

1650

2340

2100

1860

2580

1650

2340

2580

2340

2100

3390

2580

2850

2580

3390

3120

3390

3690

3990

3390

1860

2850

2580

3390

2850

2340

McCutchan_R1b_4C39R

1860

2850

3390

1440

3840

9330

1650

840

1230

1020

2100

1860

1230

1860

2100

1860

2100

2340

1860

2340

2580

2100

1650

2100

1440

2100

2340

2850

2340

1860

2580

2340

2850

2580

3120

4680

4320

1860

2100

2340

2850

3390

2580

Campbell_R1b_K3XKQ

2100

2580

3120

1650

3840

9330

1650

840

1440

1230

2340

2100

1650

1860

2100

2340

2100

1860

2100

2340

1860

2580

2340

2100

1650

1860

2340

1440

2340

2580

3120

2580

1650

2850

2100

2850

2100

2340

2580

2850

5040

3690

4320

1860

2340

2580

2850

3120

2580

Campbell_R1b_9MJ45

1860

2580

4320

1230

4320

8310

1650

1230

1440

1230

2100

1650

1440

1860

2100

1650

1860

2100

2580

1860

1440

1650

2340

1860

2340

1860

2580

2850

1650

3120

2100

3390

2850

3390

5040

3990

2100

2580

3390

2580

Urquhart_R1b_JWBVE

1230

1860

3120

660

3420

8310

1230

1020

1230

1440

840

1230

1650

1230

1650

1440

1860

1230

1860

1230

840

1860

840

1440

1230

1650

1860

1440

1020

2340

1440

2340

1860

2100

2850

2580

4320

3990

3390

1650

2340

2100

3390

2850

2340

MacEachern_R1b_8K5XS

1440

2580

4320

1020

4320

8310

2100

2340

2100

1440

1230

1020

1230

2100

1440

1650

1860

1650

1440

2100

1650

2580

2100

1440

1650

1440

1860

1650

2340

2580

2100

1650

2580

2100

2850

2340

3690

2850

3990

4320

2850

2340

2580

2340

3690

3120

2580

MacDonald_R1b_JZQQX

1230

1860

3690

660

4320

8310

1860

2100

1650

840

1230

840

1650

2100

1440

1650

1440

1860

1650

2340

2100

1230

840

1020

2340

1230

1860

1440

1860

2100

1650

1230

2340

1860

2580

2100

2340

3120

2850

4320

4680

2850

2100

2340

3390

2850

2340

Forbes_R1b_4ASGK

840

1650

3120

300

3840

9330

1650

1230

1650

1440

840

1020

840

1020

1650

1440

1020

1230

1020

1440

1230

1860

1650

1020

660

840

1650

660

1440

2100

1230

840

1440

2340

1860

1650

2850

2100

4320

3990

2850

1650

1860

2850

2340

1860

Anderson_R1b_H5535

1230

2340

3990

1020

3420

8310

1860

1230

1650

1020

1860

2100

1440

1650

1440

1650

1440

1860

1650

1230

1650

1230

1650

1860

2100

1860

1440

2340

1650

2850

2580

2100

3120

2580

3990

3390

1860

2340

1860

3120

2580

2340

Hamilton_R1_AM847

2100

2580

3990

1440

3840

9330

2340

2100

1860

1650

2100

1650

1860

1440

1650

2100

2340

2100

2340

1650

2100

1860

2100

1650

2100

1860

2100

2340

2580

3390

2100

1860

3120

2340

2850

2340

2850

2580

3120

2850

5040

4680

3690

2340

3390

2580

3990

3120

MacDonald_R1b_F4XG9

1860

2850

4320

1440

3840

8310

2340

2100

2340

1860

1650

1440

2100

1440

2100

1440

1860

2100

2340

2100

1650

2340

2100

2340

1860

1650

2100

1650

1860

2100

3120

1860

2850

2340

2580

2100

2580

2100

3120

2340

5430

5040

3990

2580

3120

2100

3690

3390

3120

Livingston_R1b_Q5BR3

1650

1860

3390

840

3840

9330

2100

1230

1650

1440

1020

1440

1650

1860

2100

1860

1650

1860

1230

1860

1230

1440

1230

1020

2100

1230

1860

2100

3120

1860

1230

2340

1860

2100

1650

2100

2850

2340

4680

4320

3390

2100

2580

1650

2850

2340

2100

McGregor_R1b_4WAZU

1230

2100

3120

1230

4320

8310

1860

1650

1230

1860

1650

1230

1650

2100

1020

840

1020

1860

2100

1650

1230

1440

2580

1440

1650

1440

1860

2340

1860

1440

2580

1860

2340

3120

2580

3690

3120

3990

3390

2850

1650

2340

3390

2850

1650

Stirling_R1b_KNES8

1020

2100

3120

1230

4830

10530

1860

2100

1650

1230

1650

2340

1860

1020

300

1020

1230

1860

1650

1440

1230

1440

2340

1650

1860

2580

2100

1650

2580

2340

2580

3120

2340

2850

4320

3690

3120

2340

1650

1860

2850

1440

Stirling_R1b_RS2P7

840

1860

3120

1020

4320

9330

2100

2340

1860

1440

1020

1440

2100

1650

840

300

840

1020

1650

1440

1230

1020

1230

2100

1440

1650

2340

1860

1440

2340

2100

2340

2850

2100

3120

2580

3990

2850

2100

1860

1650

3120

2580

1650

McGregor_R1b_VBSFV

1020

1860

2850

1440

4320

9330

2340

2100

1860

2100

1860

1440

1650

2340

2100

1860

1020

840

1020

2100

1860

1230

1440

1650

2100

1230

1650

2100

1650

2580

2340

1860

2580

2340

1650

2340

1860

3120

2340

4320

3690

2580

1860

2100

1650

2100

1650

Stirling_R1b_GDW3X

1230

2100

2850

1230

3840

9330

1650

1860

2100

1230

1650

1230

1440

1650

1230

1020

1230

1020

1650

1230

1440

1230

1020

2100

1230

1020

1650

2580

2100

1650

2580

2100

1860

2850

2340

3120

2580

4320

3690

2850

1860

2340

1440

2850

2340

1860

McMillan_R1b_V5URN

1440

2340

2850

1860

3840

8310

2340

2580

1860

2580

2340

1860

2100

2340

1860

1650

2100

1650

120

1230

2100

1860

1650

2850

1860

1650

2340

2100

2850

2100

1650

2100

1860

2340

2850

2340

3120

2580

3690

3990

3390

1650

2100

1440

3690

2340

2850

McMillan_R1b_YA5PE

1440

2340

2850

1860

3840

8310

2340

2580

1860

2580

2340

1860

2100

2340

1860

1650

2100

1650

120

1230

2100

1860

1650

2850

1860

1650

2340

2100

2850

2100

1650

2100

1860

2340

2850

2340

3120

2580

3690

3990

3390

1650

2100

1440

3690

2340

2850

McMillan_R1b_6UWW4

1440

2340

3690

1650

3840

9330

2100

2580

2340

2580

1860

2100

1650

1860

2100

1230

2100

1650

1440

1860

1230

1440

1230

1440

2580

1860

1650

2340

2100

3120

2340

1860

2850

2580

2340

3120

2340

2580

3990

3120

2100

2340

1230

3390

2340

2580

Anderson_R1b_YNXGB

840

1440

3120

840

3840

9330

2100

1860

1230

1440

1230

1020

1230

2100

2340

1440

1650

1440

1230

1440

2100

1440

450

1020

1650

1020

1440

1650

1440

2340

1860

1440

2340

1860

2100

2580

3990

3390

2340

1650

1860

1650

2850

2340

1650

McKinlay_R1b_7X6HZ

840

1440

3120

450

3840

9330

1860

1650

1440

840

1440

840

660

1230

1650

1860

1230

1020

1440

1230

1860

1230

450

660

2100

1020

1440

1230

1440

2340

1440

1020

2100

1650

2580

2100

2580

4320

3990

2850

1650

1860

3120

2580

1860

Stein_R1b_E48PN

1020

1230

2580

660

4320

8310

1860

1650

1860

1650

840

1650

1020

840

1650

1020

1440

1230

1650

1020

1650

1440

1020

660

1860

1020

1230

1440

2100

1650

1230

2100

1440

1860

1440

1650

2340

1860

4320

3120

1860

1650

2850

1650

White_R1b_E6W6F

1650

2580

3690

1650

4320

8310

2580

2100

2340

1860

1440

2340

1650

2100

2580

2340

2100

2850

2580

1650

2100

1860

1440

2340

2100

2340

2580

2340

2850

2340

1860

3120

1860

3990

3690

2580

2340

3120

2580

Jamieson_R1b_YRADM

840

1650

2340

660

3840

9330

1650

1440

1860

840

1440

1230

660

1230

1860

1650

1230

1440

1650

1440

1230

1860

1020

1440

1230

1860

1230

2100

1650

1230

1860

1650

1860

1440

2580

1860

4320

3690

2580

1440

1860

1650

2580

2100

1650

Tagert_R1b1c_MABKF

660

1440

2850

1440

4320

9330

2340

2100

2340

1440

1860

1440

1650

2100

1860

1650

1440

1650

1020

1650

1440

1020

1440

1230

1650

2100

1650

2100

1650

2100

1440

2340

1440

2340

1650

3120

3690

2850

1860

1230

2850

2580

1650

Anderson_R1b_77P34

1230

1650

3690

1230

4320

8310

2340

2100

2340

1860

1230

1650

1440

1860

2340

2100

1440

1860

1650

2100

1650

2340

1650

1230

2340

1860

1230

120

1860

2100

1650

1230

2340

1440

3120

2340

2850

2340

3690

2850

3390

3990

3120

2100

1860

2340

3390

3120

2100

Anderson_R1b_A7P6F

1230

1650

3690

1230

4320

8310

2340

2100

2340

1860

1230

1650

1440

1860

2340

2100

1440

1860

1650

2100

1650

2340

1650

1230

2340

1860

1230

120

1860

2100

1650

1230

2340

1440

3120

2340

2850

2340

3690

2850

3390

3990

3120

2100

1860

2340

3390

3120

2100

Kincaid_R1b_Q2YWJ

840

1650

2580

1440

3420

9330

2580

2340

2580

1650

2340

1860

1440

2100

2580

2100

1860

1650

2100

1440

2100

1230

1860

1440

1650

2340

2100

2580

1860

2100

1650

2580

1650

3390

2580

2100

1860

1650

2580

2100

1860

McMillan_R1b_WBDKF

1440

2100

3390

2100

3840

8310

2580

2850

3120

2850

1860

2580

2100

3390

3120

2340

2580

2340

2580

2850

3120

2340

2100

2340

2100

1650

2100

1440

2100

2850

1860

2850

2100

2580

2100

2580

1860

2850

3990

3690

2580

2340

2580

2850

2340

Young_R1b_CSRF9

1440

2100

3690

1020

3420

7440

2340

2580

1650

1440

2100

1650

1230

1860

2100

1860

2100

1860

2340

2100

2340

1860

1440

1650

2580

1650

2100

1650

2100

840

1650

1440

3390

2580

2340

3990

2580

3990

3390

2340

2580

2100

3120

2100

2580

Alexander_R1b_SQV7S

1020

1230

3120

660

3840

8310

2100

1860

1650

1020

1650

1230

840

1440

1860

1230

1440

1650

1440

1860

1650

1860

1440

1020

1230

2340

1230

1650

1230

1650

2100

840

1230

660

2580

1650

1440

2850

1860

3690

3390

3120

1650

2580

2100

Gray_R1b_VBP44

1650

1860

3120

1650

4830

9330

3390

2580

2850

3120

2340

2580

2340

1440

2340

3120

2850

2340

2580

2340

2580

2100

2850

2340

2100

2850

1860

2100

2340

2850

1650

1230

2580

2100

2340

1860

3390

2340

4320

4680

3990

2340

1860

2100

2850

2100

2580

MacCorkle_R1b_Q777T

1440

1230

3120

1230

4320

7440

2580

2340

2100

1440

2100

1860

1440

1650

2340

1860

2340

2100

2340

2100

2580

1860

1650

1440

2340

1650

1440

2100

1860

1440

660

1230

2580

1650

1440

2850

1860

3690

1860

2100

2580

2100

Wilkinson_R1b_KFM4M

2100

2340

2580

2340

4830

7440

2850

3390

2340

2850

2580

2340

2850

2580

2100

2340

2580

2340

1650

1860

2340

2100

2580

1860

2340

1860

2100

3120

2580

2850

3390

2580

120

1230

2340

1860

2580

2100

4320

3990

3120

2580

2100

2340

2580

2100

Wilkinson_R1b_T8VPT

2100

2340

2580

2340

4830

7440

2850

3390

2340

2850

2580

2340

2850

2580

2100

2340

2580

2340

1650

1860

2340

2100

2580

1860

2340

1860

2100

3120

2580

2850

3390

2580

120

1230

2340

1860

2580

2100

4320

3990

3120

2580

2100

2340

2580

2100

Hall_R1b_B4WFM

1440

1230

2340

1860

4320

8310

2580

2100

2850

1860

2850

2100

1860

2580

2340

2100

1650

2340

2580

2340

1650

1860

2340

2100

1440

2340

1440

2340

1860

2100

2580

1650

2100

1650

1230

1650

1020

1650

1230

3990

3690

3120

1860

1650

2100

1440

Wilson_R1b_3UEUC

2100

1860

3120

1650

5400

9330

3390

2580

2340

2850

1860

2850

2340

1860

2580

2850

2580

2100

3120

2850

2340

2850

3120

2100

1650

2340

1440

2340

2850

2100

2580

1650

2340

1440

2340

1650

1440

2340

1440

5040

4320

2580

2850

3390

2850

Walker_R1b_MWE53

1230

1440

2850

1650

4830

8310

3120

2580

2340

2850

2100

2340

1650

2100

2580

2100

2580

2340

2100

1860

2340

2100

1650

1860

1440

2340

1650

2100

2340

1440

1860

1440

1860

1020

1440

2100

1020

3990

4320

3690

2340

1650

2100

2580

2100

Simmons_R1b_CV74C

2340

2100

3120

2850

4320

8310

3390

3120

2580

3390

2850

3690

3120

2850

3120

2850

3690

2850

3120

2580

2340

3120

2580

2340

3690

2580

3990

2850

3390

2850

2580

1650

2340

2100

1440

5040

4320

3120

2340

2580

2850

3690

2580

Parker_R1b_Z5PNU

1650

1860

3120

2100

3840

7440

3690

3120

2850

3390

2580

2850

2100

2580

2850

2340

3120

2850

2580

2340

2580

1860

1650

2850

1650

1860

2580

1860

2340

1860

2100

1230

1440

1020

1440

3990

2850

2100

1860

2340

2850

2580

Clark_R1b_S286F

3120

2850

3990

4320

5400

8310

3990

4680

5040

4320

3990

4320

3990

5040

5430

4680

3990

4320

3990

4320

3690

3990

4320

3990

4320

3120

3390

2850

3990

3690

4320

3690

4320

3990

5040

3990

5040

3990

2850

2580

2340

2850

3690

3120

3690

ODonoghue_R1b_2FMC4

3120

2580

3120

3990

5400

7440

3390

4320

3690

3990

4320

4680

3990

4680

5040

4320

3390

3690

3990

3690

3990

3390

3990

4320

3990

3690

3990

3390

3990

3390

4680

3690

3990

3690

5040

4320

3990

2850

1440

2580

3390

3120

2850

Donohoe_R1b_M7V3H

2100

3120

2850

4830

6690

3390

4320

3990

3390

2850

3390

3690

3990

3390

2850

3120

2850

2580

2850

3390

3120

2340

2850

3120

3690

2580

2850

3120

2580

3690

3390

3120

3990

3690

3120

4320

3690

4320

3990

2580

1440

2580

3390

2850

3690

2850

2340

Kane_R1b_NE5HD

1440

1650

2580

1650

3420

8310

1860

2100

1650

2340

2100

1650

1860

2340

2580

2100

1650

2340

2100

1860

1650

2100

1650

1860

2580

1440

1860

2100

2580

2340

1650

2340

1860

2580

1860

2580

2340

3120

2850

2340

2580

1650

1440

2340

1650

2100

Barrett_R1b_W9TVC

1020

1440

2580

1860

5400

12000

2850

2100

2340

2580

2340

2580

2340

1860

2340

3390

3120

2580

2340

1650

1860

2100

2340

2100

2340

1860

2340

1860

1230

1860

2340

2580

1650

1860

2580

1860

2580

1650

2340

2100

2850

3390

1650

1230

1860

2340

1650

Turk_R1b_2BFE5

1020

1650

2850

1860

3840

8310

2580

2340

2580

2100

2340

1860

2580

2100

1650

2340

1860

1650

1440

1230

1650

1860

1650

2340

1650

1230

2340

1650

2580

2100

1650

2100

1650

2580

1650

2580

1860

2850

3390

2850

1440

1230

2100

1650

2100

Blakeston_R1b_QRKMM

2340

2100

2580

2850

4320

9330

3390

2850

3390

3690

3390

2850

3120

3990

3690

2850

3390

2850

3120

2100

2850

3690

3390

2850

3120

2850

3120

2580

2850

3390

2580

2850

3120

2580

2850

2580

2340

1650

2850

2100

2850

2340

3690

3390

3690

2340

1860

2100

1650

1860

Andriesen_R1b_UA5J3

1860

2100

2580

2340

3000

9330

2850

3390

3120

3390

2850

3120

2850

2340

2580

3120

3390

2340

2850

2580

2100

2340

2580

2850

3120

2100

2580

3120

2100

2850

2100

2580

2100

3390

2580

3690

2850

3120

2850

1650

2340

1650

2340

Ming_R1b_Z8YYY

1230

1440

2580

1860

5400

9330

2340

2580

2340

2580

2340

1860

2340

3120

2100

1650

1440

1650

1860

2850

2580

1650

1860

1650

2580

1650

2100

1860

2340

2580

2100

2580

2100

1440

2850

2100

2580

3690

2850

2340

2100

1650

2100

1860

2340

0-270 Years

300-570 Years

600-870 Years

900-1170 Years

- Infinite allele mutation model is used
- Average mutation rate: 0.0024
- Values on the diagonal indicate number of markers tested
- Probability is 50% that the TMRCA is no longer than indicated
- Average generaton: 30 years