Brief Communication: Restricted Geographic
Distribution for Y-Q* Paragroup in South America

Graciela Bailliet,1* Virginia Ramallo,1 Marina Muzzio,1 Angelina Garcı́a,2 Marı́a R. Santos,1

Emma L. Alfaro,3 José E. Dipierri,3 Susana Salceda,4 Francisco R. Carnese,5 Claudio M. Bravi,1

Néstor O. Bianchi,1 and Darı́o A. Demarchi2

1Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biologı́a Celular (IMBICE),
CCT- CONICET-La Plata, Argentina
2Museo de Antropologı́a, Facultad de Filosofı́a y Humanidades, Universidad Nacional de Córdoba,
Córdoba, Argentina
3Instituto de Biologı́a de la Altura, Facultad de Humanidades y Ciencias Sociales,
Universidad Nacional de Jujuy, San Salvador de Jujuy, Argentina
4Departamento de Antropologı́a Biológica, Facultad de Ciencias Naturales y Museo,
Universidad Nacional de La Plata, La Plata, Argentina
5Sección Antropologı́a Biológica, Instituto de Ciencias Antropológicas, Facultad de Filosofı́a y Letras,
Universidad de Buenos Aires, Buenos Aires, Argentina

KEY WORDS Y chromosome; SNP; microsatellites; south America

ABSTRACT We analyzed 21 paragroup Q* Y chro-
mosomes from South American aboriginal and urban
populations. Our aims were to evaluate the phyloge-
netic status, geographic distribution, and genetic diver-
sity in these groups of chromosomes and compare the
degree of genetic variation in relation to Q1a3a haplo-
types. All Q* chromosomes from our series and five
samples from North American Q* presented the deri-
vate state for M346, that is present upstream to M3,
and determined Q1a3* paragroup. We found a restric-
tive geographic distribution and low frequency of
Q1a3* in South America. We assumed that this low
frequency could be reflecting extreme drift effects.

However, several estimates of gene diversity do not
support the existence of a severe bottleneck. The mean
haplotype diversity expected was similar to that for
South American Q1a3* and Q1a3a (0.478 and 0.501,
respectively). The analysis of previous reports from
other research groups and this study shows the high-
est frequencies of Q* for the West Corner and the
Grand Chaco regions of South America. At present,
there is no information on whether the phylogenetic
status of Q* paragoup described in previous reports is
similar to that of Q1a3* paragroup though our results
support this possibility. Am J Phys Anthropol 140:578–
582, 2009. VVC 2009 Wiley-Liss, Inc.

Seielstad et al. (2003) described the M242 Y-chromo-
some polymorphism defining the Q haplogroup (YCC,
2002), and proposed it as a founder for the Americas.
In relation to their geographic distribution, Q, simi-

larly to C, is currently present in Eurasia, suggesting
that they originated in Asia and then spread over
Europe and America. Frequencies for Q were lower than
17% (5% average) for Asian populations (Seielstad et al.,
2003), 18.8% for Siberia (Karafet et al., 2002), while for
North America they were up to 47% (13.9% average)
(Zegura et al., 2004; Bolnick et al., 2006). However, in
most South American populations Q* is poorly repre-
sented (\6%) (Bortolini et al., 2003; this study). Findings
from other authors make us presume that the divergence
between Q and Q1a3a (YCC, 2002; Karafet et al., 2008)
took place shortly before or during the Bering Strait
crossing (Lell et al., 2002), while the origin of Q might have
preceded the population expansion from Asia to the rest of
the world (Bortolini et al., 2003; Seielstad et al., 2003).
Karafet et al. (2008) described an extensively revised

Y-chromosome tree containing 311 distinct haplogroups
and incorporating approximately 600 binary markers.
They showed that Q haplogroup accumulated 17 muta-
tions defining 14 haplogroups; they also described that
all Q chromosomes from their series presented P36.2
downstream to M242, expecting that MEH2 could be a

subset of Q-P36.2. In this scenario, we assumed that the
presence of M346, downstream to MEH2 and upstream
to M3, evidences the phylogenetic status of our Q* para-
group chromosomes (Karafet et al., 2008). Anyway, we
still ignore the real status of Q* in America and Siberia
(Bortolini et al., 2003; Bolnick et al., 2006; Karafet et al.,
2002), though we could not discard that at least some
haplotypes from that Q* paragroup also presented
M346*G. Every sample from our series of Q* chromo-
somes presented M346, indicating that they belong to
the Q1a3* paragroup.

Grant sponsors: CONICET, CICPBA, ANPCyT, UNJu, Antorchas
Foundation of Argentina. G Bailliet, S Salceda, NO Bianchi, CM
Bravi, and DA Demarchi are members of the Consejo Nacional de
Investigaciones Cientı́ficas y Técnicas de la República Argentina
(CONICET).

*Correspondence to: Graciela Bailliet, IMBICE, 526 e/ 10 y 11. CC
403, La Plata 1900, Argentina. E-mail: gbailliet@imbice.org.ar

Received 5 December 2008; accepted 29 May 2009

DOI 10.1002/ajpa.21133
Published online 9 July 2009 in Wiley InterScience

(www.interscience.wiley.com).

VVC 2009 WILEY-LISS, INC.

AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 140:578–582 (2009)



MATERIALS AND METHODS

We analyzed 759 samples from nonrelated males of 11
South American native populations (Lengua and Ayoreo
from Paraguay; Huilliche and Pehuenche from Chile;
Wichi, Toba, Chorote, Mocovı́, Mapuche, Tehuelche, and
Humahuaca from Argentina), and seven admixed urban
populations (Jujuy, Catamarca, Tucumán, Salta, and
Córdoba from Argentina; and La Paz and Tarija from
Bolivia) (Table 1, Fig. 1). Because of Q* low frequency in
native populations, we also analyzed urban populations
considering that all Q* Y-chromosomes belong to Native
American male ancestors. Bolivian samples increased
the number of Andean samples in the series; Native
American ancestry in all urban samples can be inferred
from Q1a3a frequency (Table 1). These results contribute
to certify the high proportion of Native American
inheritance in urban populations.
All biological samples were collected with the informed

consent of the donors, encoded, and sent to our labora-
tory in anonymity for DNA testing.
For this study, we included samples corresponding to:

Q* paragroup (their phylogenetic status was confirmed by
the analysis of M9*G, M25*G, M120*T) (Underhill et al.,
2001), P27*C (Karafet et al., 1999), M242*T (Seielstad
et al., 2003), M3*C (Underhill et al., 1996)]. We sequenced
M346 (Sengupta et al., 2006) in all Q* samples of our se-
ries and in two Sioux, one Navajo, and two Zuni from
North American Q* (Bianchi et al., 1998), and identified
haplotypes using seven Y chromosome STRs [DYS19,
DYS389a, DYS389b, DYS390, DYS391, DYS392, and
DYS393 (de Knijff et al., 1997; Kayser et al., 1997)]. From
21 samples characterized as Q*, only 17 could be analyzed
for the seven specific microsatellites.
Lineage frequencies were calculated by direct count-

ing. Microsatellite haplotype affinities were assessed
through the Median Joining Network analysis (Bandelt
et al., 1999) after the Star Contraction procedure (For-
ster et al., 2001), using the Network software (v 4.2.1.0,
www.fluxusengineering.com). Data on Q* (Bortolini
et al., 2003; Bolnick et al., 2006) and 129 Q1a3a haplo-
types from the same populations included in this study
were analyzed (Table 1) (Bianchi et al., 1998; present
results).
To detect any possible differences in haplotype diver-

sity between South and North American samples related
to any bottleneck event, we computed several genetic di-
versity estimators (such as expected diversity, number of
alleles, allelic size range, and Garza–Williamson index),
using the Arlequin 3.1 software (Schneider et al., 2000).
Figure 2 and Table 2 show a total of 35 Q* paragroup

haplotypes corresponding to 17 haplotypes from our
South American series (identified with bold italic num-
bers in Table 2), and 19 North-American haplotypes
reported by Bolnick et al. (2006) (identified with one
asterisk preceding the population acronym in Table 2).

RESULTS

Of the 759 samples analyzed, we found 21 Q* chromo-
somes. Lengua and Ayoreo populations showed the high-
est Q* frequency in our series (7/24, 29.2%, and 2/9,
22.2%, respectively). Lower values were observed for
Mocovı́ (2/40, 5%), Mapuche, Huilliche (1/26, 3.8%), and
Wichi (1/120, 0.8%) (Table 1, Fig. 1). Q* Y-chromosome
frequency for urban populations ranged from 0 for Cata-
marca, Jujuy, and Tucumán to 5.5% for Tarija; Salta, La

Paz, and Córdoba showed 4.8, 3.4, and 0.6% intermedi-
ate frequencies, respectively (Table 1, Fig. 1).
To determine the phylogenetic status of this para-

group, we analyzed the M346 mutation that is positioned
upstream to M3, accordingly with the phylogeny recently
published by Karafet et al. (2008). We confirmed the
presence of M346 G in all samples of our series, and in
the Q* paragroup samples from North America: two
Sioux, one Navajo, and two Zuni (data not shown, Bian-
chi et al., 1998). This evidences that probably most of
the Q* chromosomes from North America (Bolnick et al.,
2006) also present M346 G, this being the reason why
we consider that all these samples belong to the Q1a3*
paragroup, closely related with Q1a3a (Table 2, Fig. 2).
We succeeded in characterizing the full set of seven

microsatellites (DYS 19, 389 I, 389 II, 390, 391, 392, and
393) in 17 of the 21 Q* chromosomes; each one of the 17
fully tested chromosomes exhibited a private single hap-
lotype (Table 2, Fig. 2). Haplotypes 1 and 2 were found
in two Lengua, and haplotype 3 was exhibited by one
Creek individual. Similar haplotypes were described in
one Guaranı́, three Ingano, and one Wayuu (Haplotype
2; Bortolini et al., 2003) (Table 2, Fig. 2). Haplotype
1 was also found in the Q1a3a haplogroup proposed as
the haplotype from which M3 originated (Bortolini et al.,
2003).
Figure 2 shows the network of South and North

American Q* haplotypes, in which the South American
haplotypes occupy a central position. Because it was not
possible to compare them with Siberian Q* haplotypes,
we cannot assert further conclusions about their
phylogenetic status.
The mean number of alleles were 3.00, 3.43, and 5.00

for North American Q*, South American Q1a3* and
Q1a3a, respectively. The range of allele size was 2.14,
2.71, and 4.00, respectively. Expected diversity was
0.482 for North American and 0.478 for South American

TABLE 1. Q1a3* and Q1a3a frequencies in
South American populations

Populations
Linguistic
lineagesa N Q1a3* Q1a3a

Paraguay
Lengua (tribal) Mascoian 24 0.292 0.667
Ayoreo (tribal) Zamucoan 9 0.222 0.556

Argentina
Wichi (tribal) Mataco 120 0.008 0.375
Toba (tribal) Guaicuruan 12 0 0.833
Chorote (tribal) Mataco 9 0 0.889
Mocovı́ (tribal) Guicuruan 40 0.050 0.550
Mapuche (tribal) Mapudungun 26 0.038 0.538
Tehuelche (tribal) Chon 20 0 0.650
Humahuaca (urban) Spanish 31 0 0.839
Jujuy (urban) Spanish 46 0 0.360
Salta (urban) Spanish 72 0.048 0.444
Catamarca (urban) Spanish 98 0 0.112
Tucumán (urban) Spanish 17 0 0.118
Córdoba (urban) Spanish 156 0.006 0.147

Chile
Huilliche (tribal) Mupudungun 26 0.038 0.461
Pehuenche (tribal) Mupudungun 18 0 0.833

Bolivia
La Paz (urban) Spanish 29 0.034 0.655
Tarija (urban) Spanish 72 0.055 0.500

Total 759 0.029 0.427

Q1a3*, Q1a3a (YCC, 2002; Karafet et al., 2008).
a Ethnologue language database: http://www.ethnologue.com.

579Q* IN SOUTH AMERICA

American Journal of Physical Anthropology



Q1a3* lineages, and 0.501 for South American Q1a3a
haplotypes. Distance assessed by the addition of square
size differences (RST) was 0.0196, not significant (P 5
0.144) between North American Q* and South Ameri-
can Q1a3* haplotypes. Meanwhile, distances between
Q1a3a haplotypes and North and South American Q*
were 0.0796 and 0.614, respectively, both statistically
significant (P \ 0.001). Fixation index for the three
groups was 0.069 (P 5 0.001). Finally, the Garza–Wil-
liamson index (the number of alleles divided by the
allelic range, expected to be low in bottlenecked popula-
tions and close to one in stationary populations) was
[0.96.

DISCUSSION

To analyze the phyletic origin of the Q* lineages in our
series, we confirmed the status of Q1a3* by the presence
of M346, upstream to M3. Even when there are no spe-
cific mutations that define Q1a3* by itself, this finding
confirms the close phylogenetic relationship between
Q1a3* and Q1a3a. In this case, Q1a3* would have
entered Siberia from Asia (Gonzales-José et al., 2008).
As complementary evidence of its Asian origin, M 346
has been previously described in India and Pakistan as
Q4 (Sengupta et al., 2006). Furthermore, Q is exclusively
distributed in North Western and North Eastern Siberia,

Fig. 1. Map of South American sampling localities were Q* paragroup is represented by circles, with an area proportional to
frequency. See Table 1 for names, size, and language of populations samples.

580 G. BAILLIET ET AL.

American Journal of Physical Anthropology



all these Q presented P36 derivated allele (Karafet
et al., 2002). The analysis of M346 in the Siberian Q
haplogroup would help to draw further conclusions about
the complexity of populations that were the source of
current American populations (Karafet et al., 2002;
Bortolini et al., 2003).
The genetic diversity of Q* (Bolnick et al., 2006) and

Q1a3* (this study) was similar, and they showed the
same genetic distance from Q1a3a. Even when Q* and
Q1a3* seem to be a little less diverse than Q1a3a, we
cannot confirm any bottleneck effect. These analyses,
and the presence of M346 G in Q* North American sam-
ples (data not shown), make us presume that most North
American Q* might also be Q1a3*.
When our data were pooled with those from Bortolini

et al. (2003), a particular geographic distribution of Q*
chromosomes was observed in South American popula-
tions: the highest concentration was shown by the
Northwest border (67% Ingano, 48% Zenu, and 21%
Wayuu) (Bortolini et al., 2003), followed by the Gran
Chaco region (29% Lengua, 22% Ayoreo) (this study),
whereas the remaining populations showed frequencies
below 6%.
Different sources of evidence support the hypothesis of

a coastal colonization route into South America (Fuselli
et al., 2003; Wang et al., 2007), thus explaining Q* high
frequency in the Northwest border. Furthermore, South
American inland population has been proposed to derive
from the subsampling of western towards eastern popu-
lations (Fuselli et al., 2003; Wang et al., 2007). This pat-
tern could explain the absence of Q* in most Eastern
populations due to genetic drift, while similarities among
different populations mismatching geographic distribu-
tion are attributed to founder effects. However, this can-
not explain the high frequency and the ancestral status
of lineages from the Gran Chaco area. Interestingly, this
population shows the highest diversity level in South
America (Demarchi et al., 2001; Cabana et al., 2006;
Crossetti et al., 2008).
The low frequency of the Q1a3* paragroup observed in

South America suggests that it is probably being lost by
an eventual bottleneck or genetic drift, especially when
we consider its small effective population size among
South American hunting-gathering groups. However,
several estimators show a similar degree of diversity for

North and South American samples. These values were
similar to those found for Q1a3a lineages from South
America and allow to rule out the possible existence of a
bottleneck event explaining the low representation of the
Q1a3* paragroup in South America.

ACKNOWLEDGMENTS

We thank all DNA donors for making this work
possible. We are most grateful to Michael H. Crawford
for his critical review of the original version of this
manuscript.

Fig. 2. Microsatellite network for North and South Ameri-
can Q* paragroup haplotypes represented in Table 2. Black
circles, South American haplotypes (this study); gray circles,
South East North American haplotypes; white circles, North
East North American haplotypes (Boltnick et al., 2006). Circle
size is proportional to frequency.

TABLE 2. Q* haplotypes in North and South America

Haplotypesa 19 389a 389b 390 391 392 393
Populationsb

(casesc)

1 13 13 30 24 10 14 13 L (1)
2 13 12 32 24 10 14 13 L (1)
3 13 12 29 24 10 14 13 *CRK (1)
4 13 13 29 24 10 14 14 *SC (1),

*OC (1),
*SC (1)

5 13 13 30 23 10 13 13 Map (1)
6 13 13 31 23 10 13 13 Mo (1)
7 13 13 29 23 10 15 13 L (1),

*CHO (3)
8 13 12 28 23 10 15 13 *SC (1)
9 13 14 30 23 10 15 13 *CHO (1)

10 14 13 30 24 10 14 13 A (1)
11 14 13 29 24 10 14 13 B (1)
12 14 15 29 24 10 14 13 L (1)
13 14 13 27 24 10 14 13 L (1)
14 14 14 32 24 10 14 13 B (1)
15 14 13 30 24 10 14 12 *CA(1),

*SIO (1),
*WC (3)

16 14 13 29 24 10 14 12 *MC (1)
17 14 14 29 24 10 14 12 B (1)
18 14 13 30 23 10 14 13 *SIO (2),

*OC (1)
19 14 14 29 23 10 14 13 B (1)
20 14 13 30 23 10 14 14 *CA (2),

*MC (2)
21 14 12 28 23 10 14 14 *SIO (1)
22 14 12 28 25 10 14 14 *SIO (1)
23 14 13 30 23 10 14 14 *WC (1)
24 15 14 30 24 10 14 14 *SHA (1)
25 14 13 32 24 10 13 13 Mo (1)
26 14 14 29 24 10 15 13 A (1)
27 13 13 29 25 10 14 13 *CRK (1)
28 14 13 29 24 11 13 13 Cor (1)
29 15 14 29 23 10 14 13 L (1)
30 13 16 26 25 10 14 14 Sal (1)
31 13 13 30 24 9 14 12 *OC (1)
32 16 10 29 24 10 14 14 *OC (1)
33 13 14 32 23 10 14 13 *OC (1)
34 13 13 29 24 10 12 13 *SC (1)
35 14 13 29 24 11 14 13 *OC (1)

a South American populations (bold).
b South American populations: A, Ayoreo; L, Lengua; B, Bolivia;
Cor, Córdoba; Map, Mapuche; Mo, Mocovı́; Sal, Salta. (*) North
American haplotypes from Bolnick et al. (2006): CA, Cheyenne
Arapaho; CHIC, Chickasaw; CHO, Choctaw; CRK, Creek; KIC,
Kickapoo; MC, Minnesota Chippewa; MIC, Micmac; OC, Okla-
homa Red Cross Cherokee; SC, Stillwell Cherokee; SEM, Semi-
nole; SHA, Shawnee; SIO, Sisseton Wahpeton Sioux; TMC, Tur-
tle Mountain Chippewa; WC, Wisconsin Chippewa.
c Number of cases in parenthesis.

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