ArticlePDF Available

The Genetic Structure of Populations from Haiti and Jamaica Reflect Divergent Demographic Histories

Authors:

Abstract and Figures

The West Indies represent an amalgamation of African, European and in some cases, East Asian sources, but the contributions from each ethnic group remain relatively unexplored from a genetic perspective. In the present study, we report, for the first time, allelic frequency data across the complete set of 15 autosomal STR loci for general collections from Haiti and Jamaica, which were subsequently used to examine the genetic diversity present in each island population. Our results indicate that although both Haiti and Jamaica display genetic affinities with the continental African collections, a stronger African signal is detected in Haiti than in Jamaica. Although only minimal contributions from non-African sources were observed in Haiti, Jamaica displays genetic input from both European and East Asian sources, an admixture profile similar to other New World collections of African descent analyzed in this report. The divergent genetic signatures present in these populations allude to the different migratory events of Africans, Europeans, and East Asians into the New World.
Content may be subject to copyright.
The Genetic Structure of Populations from Haiti
and Jamaica Reflect Divergent Demographic Histories
Tanya M. Simms,
1,2
Carol E. Rodriguez,
1
Rosa Rodriguez,
1
and Rene J. Herrera
1
*
1
Department of Molecular and Human Genetics, College of Medicine, Florida International University,
Miami, FL 33199
2
Department of Biological Sciences, Florida International University, Miami, FL 33199
KEY WORDS Greater Antilles; Afro-Caribbean; New World African; STRs; phylogenetic analyses
ABSTRACT The West Indies represent an amalga-
mation of African, European and in some cases, East
Asian sources, but the contributions from each ethnic
group remain relatively unexplored from a genetic per-
spective. In the present study, we report, for the first
time, allelic frequency data across the complete set of 15
autosomal STR loci for general collections from Haiti
and Jamaica, which were subsequently used to examine
the genetic diversity present in each island population.
Our results indicate that although both Haiti and
Jamaica display genetic affinities with the continental
African collections, a stronger African signal is detected
in Haiti than in Jamaica. Although only minimal contri-
butions from non-African sources were observed in Haiti,
Jamaica displays genetic input from both European and
East Asian sources, an admixture profile similar to other
New World collections of African descent analyzed in
this report. The divergent genetic signatures present in
these populations allude to the different migratory
events of Africans, Europeans, and East Asians into
the New World. Am J Phys Anthropol 142:49–66,
2010. V
V
C2009 Wiley-Liss, Inc.
The West Indian archipelago is bordered on the north
by the southeastern United States, on the west by Cen-
tral America, and on the south by the northern coast of
South America. This island chain is classified into two
groups: the Greater (northern) and Lesser (southeastern)
Antilles, the two encompassing a broad arc 4,000 km
long (Watts, 1987). The Antilles was first settled by
Amerindians about 2,500–3,000 years before the present
(YBP) (Keegan, 1995) who then served as slave laborers
for the Spanish Crown. Overwork, malnourishment, and
exposure to European diseases, concomitantly, led to the
drastic reduction of the native population and conse-
quently, a major deficit in the workforce (Rogozin
´ski,
1999). As a result, enslaved migrants from the West Afri-
can coast were imported into the New World beginning
from the early 1500s until the late 1800s (Knight and
Crahan, 1979). At the same time, struggles ensued among
the European powerhouses, namely Britain, Spain,
France, and the Netherlands, for control of the West In-
dian islands. Due to the different European influences,
the islands have experienced diverging cultural, economi-
cal, political, and demographic histories. In this study, we
attempt to characterize the genetic input from European,
African, and East Asian colonizations into the Greater
Antillean islands of Haiti and Jamaica.
Haiti was the largest and most profitable of the
French colonies in the New World during the 18th cen-
tury, monopolizing the world’s sugar (40%) and coffee
([50%) production (Knight, 2000). This rise in prosperity
was accompanied by the importation of an estimated
860,000 African slaves into the island (Curtin, 1975;
Coupeau, 2008) and, as a direct consequence, the num-
ber of Affranchis or gens de couleor (free coloreds and
mulattoes) increased (Coupeau, 2008). Slave rebellions
culminated in the Haitian Revolution (1789–1804) and
ultimately resulted in the emigration of large numbers
of French colonists and their slaves (Pamphile, 2001;
Treco, 2002), which reduced the population to half of its
original size (Ott, 1973). In 1804, Haiti was declared a
republic (Knight, 2000; Coupeau, 2008), and under the
newly established constitutions of the ‘‘world’s first Black
Republic’ (Heinl and Heinl, 2005), whites were prohibited
from purchasing land in the island until the American
occupation (1915–1934) (Coupeau, 2008). Consequently, a
schism was generated between the European and African
populations, most likely resulting in endogamy.
During the early 19th century, Jamaica surpassed
Haiti as the principal sugar-producing island in the
Antilles (Rogozin
´ski, 1999). Along with this affluence
came increased demands for plantation laborers that
were satisfied originally by European (Irish, Scottish,
and Welsh) white bond servants and later on, by the
introduction of approximately 914,902 African slaves
(Pepin, 2005) who, by the early 1800s, outnumbered the
Caucasians 20:1 (Mordecai and Mordecai, 2001). Accord-
ing to Pepin (2005), 90.4% of the total slave population
in Jamaica disembarked from West African ports while
Additional Supporting Information may be found in the online
version of this article.
*Correspondence to: Dr. Rene J. Herrera, Department of Molecu-
lar and Human Genetics, College of Medicine, Florida International
University, University Park, OE 304, Miami, FL 33199.
E-mail: herrerar@fiu.edu
Received 16 April 2009; accepted 24 August 2009
DOI 10.1002/ajpa.21194
Published online 16 November 2009 in Wiley InterScience
(www.interscience.wiley.com).
V
V
C2009 WILEY-LISS, INC.
AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 142:49–66 (2010)
the remaining 9.6% were transported from Southeast
Africa (Mozambique, Madagascar, and Tanzania)
although a smaller proportion [7.7% (Thomas, 1998; Pe-
reira et al., 2001)] from the same region has been
reported for the New World. Specifically, the Madagass,
those native to Madagascar, were known to be present in
Jamaica (Henriques, 1964). With the abolition of slavery
and the culmination of the Apprenticeship period
(1834–1838) throughout the British West Indies, massive
labor shortages arose again, motivating the Jamaican
government to contract more than 4,000 West European
[British and German] and approximately 1,000 Chinese
indentured servants to work on plantations between
1838 and 1918 (Lai, 1990, 1998). In addition to these
two groups, a limited number of African laborers from
Sierra Leone and Central Africa was also recruited
(Roberts, 1954; Mordecai and Mordecai, 2001). The
extent to which the aforementioned groups contributed
to the current population’s gene pool is uncertain since
many returned to their native land upon expiration of
their contracts (Roberts, 1954; Lai, 1990, 1998).
West African influences have become ingrained into
both Haitian and Jamaican societies due to the large pro-
portion of enslaved Africans transported from continental
Africa to these New World destinations. The Yoruba
speakers (natives of Nigeria and Benin), although not
the most prominent group, seem to have had the greatest
impact on Haitian society. The most practiced Yoruba
customs include Haitian Vodou, an amalgamation of the
religious beliefs of the Dahomey (present-day Benin)
people and Roman Catholics (de Heusch, 1995; Brown,
1997), and its associated festival, Rara (Tselos, 1996).
Yoruba imprints have also been detected in Haitian art
(Okediji, 2003; Roberts, 2004), music, dance (Tselos,
1996) and folklore (Roberts, 2004). Jamaica, on the other
hand, has not been impacted to such an extent by one
particular ethnic group, but instead displays influences
from various West African sources including the Akan
(Mordecai and Mordecai, 2001), Coromantee, Ibo, Fula,
Mandingo, and Yoruba peoples (Mason, 2000). Important
cultural traditions derived from West Africa include
obeah [magic] (Curtin, 1955), myalism [religion] (Curtin,
1955; Schuler, 1979), Jonkonnu [music and dance]
(Curtin, 1955; Bettelheim, 1979), ‘‘jerk,’’ [a method of
food preparation] and elements of the Jamaican dialect
(Mordecai and Mordecai, 2001).
Studies delineating the genetic ancestry of the Haitian
populace are limited. On the basis of the D1S80 polymor-
phic locus, Haiti was found to cluster with the continental
African groups in both principal component (PC) and
maximum likelihood (ML) analyses of worldwide popula-
tions (Herrera et al., 2004). In addition, using the poly-
marker system (PM) [lipoprotein receptor (LDLR), glyco-
phorin A (GYPA), hemoglobin G gamma globin (HBGG),
group specific component (GC), and D7S8], genetic affin-
ities between Haiti and other African derived New World
populations (French Antilles, Costa Rica, and Brazilian
Mulattoes) were observed in a neighbor-joining (NJ) tree
(Hidalgo et al., 2005). Haitian women were also subjects
in a clinical study examining the frequency of Duffy
antigen receptor for chemokines (DARC) single-nucleotide
polymorphisms (SNPs) in populations from the Carib-
bean, Europe, and the US. Seventy-five percent of the
Haitian samples examined exhibited the CC genotype
(absent among women of European ancestry) at levels
comparable to females of African descent from the US
(73%) (Grann et al., 2008).
In contrast to Haiti, the genetic structure of the
Jamaican population has been better characterized. In
the aforementioned study by Grann et al. (2008), the fre-
quency of the CC genotype among Jamaican females
(63%) is less than the levels present in either Haitian
(75%) or US African American (73%) women. Close
genetic ties between Jamaicans and continental Africans
have been detected through analysis of b-globin gene
polymorphisms (Antonarakis et al., 1984), autosomal
SNP and insertion/deletion (INDEL) frequency data
(Parra et al., 1998), mtDNA (Parra et al., 1998; McLean
et al., 2003, 2005), Y-chromosome markers (Bergen et
al., 1999; Benn Torres et al., 2007), as well as 28 ances-
try informative markers (AIMs) generated from autoso-
mal SNP and INDEL loci (Benn-Torres et al., 2008).
African genetic influences detected in Jamaica include
the insertion allele in the Y-chromosome Alu polymor-
phism (YAP), the marker defining Y-haplogroups D-
M174 and E-M96 (sub-Saharan African-specific), and the
presence of mtDNA haplogroup L, prevalent throughout
sub-Saharan Africa (Parra et al., 1998). European
genetic contributions, on the other hand, are generally
much lower, ranging from virtually no genetic influence
(McLean et al., 2003) up to 18.1% (Gibbs et al., 1972)
and vary depending on the marker system used (Parra
et al., 1998; McLean et al., 2005; Benn-Torres et al.,
2008). Furthermore, in addition to the presence of DNA
from these two sources, Benn-Torres and colleagues
(2008) have also reported limited Native American con-
tributions (3.2%) in Jamaica.
Reports surveying the genetic diversity present
throughout the remainder of the West Indian archipel-
ago also indicate strong genetic binds with continental
Africans, although varying degrees of genetic input from
other sources have been observed. In the Greater Antil-
les, for example, the Cuban population was found to
share the majority (45.3%) of its maternal component,
but only a limited portion (19.7%) of its Y-chromosome
lineages, with African sources (Mendizabal et al., 2008).
Also, in an earlier study, in which HLA class I polymor-
phisms were examined, genetic contributions from Africa
were detected in Cuban-Mulattoes at a comparable level
(45.32%) while the frequency in Cuban-Whites (21.46%)
was generally much lower (Ferrer et al., 2007). In addi-
tion, admixture estimates for the Puerto Rican popula-
tion reveal signals of gene flow from mainland Africa
[26% when JC virus strains were examined (Ferna
´ndez-
Cobo et al., 2001) and 27.2% based on mtDNA hap-
logroup data (Martı
´nez-Cruzado et al., 2005)], but in con-
trast to Cuba, Native Americans serve as the major con-
tributor to this island’s maternal gene pool.
Admixture profiles published for the Lesser Antillean
populations also indicate significant gene flow from the
African continent into the region. When Y-chromosome
markers are used, collections from Grenada, St. Kitts,
St. Vincent, St. Thomas, and Trinidad were all found to
receive greater than 75% of their paternal contributions
from African sources, whereas lower proportions were
exhibited by the Dominican (34.1%) and St. Lucian
(45.8%) samples. In the same group of populations, with
the exception of St. Thomas, the mtDNA was also pri-
marily African-specific (haplogroups L0, L1, L2, and L3),
although Native American (A, C, and D, ranging from
0.63% to 2.87%) and Eurasian (H, K, M, and N, ranging
from 0.32% to 1.91%) haplogroups were detected at low
frequencies (Benn Torres et al., 2007). Furthermore, in
two additional studies, commensurate levels of African
50 T.M. SIMMS ET AL.
American Journal of Physical Anthropology
influences (94%, 89.6%, and 86.8%) were reported for
Tobago (Miljkovic-Gacic et al., 2005), Barbados, and St.
Thomas (Benn-Torres et al., 2008), respectively, when
AIMs were employed. The authors also identify Euro-
pean and Native American influences at frequencies
ranging from 4.6% to 10.6% and 0.2% to 2.6%, respec-
tively (Miljkovic-Gacic et al., 2005; Benn-Torres et al.,
2008).
In the present study, allelic frequency data for
15 hypervariable autosomal STR loci are reported for the
first time for the current populations of Haiti and
Jamaica. Although the Jamaican populace has been
examined previously across the 13 Combined DNA Index
System (CODIS) STR loci (Budowle et al., 1999), the
increased number of markers analyzed in this report
allows for added resolution, thereby strengthening the
statistical analyses. Phylogenetic comparisons to world-
wide geographically targeted reference populations
reveal strong genetic affinities between Haitians, Jamai-
cans, the other New World groups of African descent
[New Providence (Bahamas) and African Americans
from the US], and the continental African collections.
Even though both Haiti and Jamaica exhibit their most
substantial genetic input from Africa, Jamaica, in con-
trast to Haiti, also displays greater genetic contributions
from Europe and East Asia.
MATERIALS AND METHODS
Sample collection and DNA extraction
Buccal swabs, along with genealogical information for
at least two generations, were collected at random from
unrelated individuals belonging to the general popula-
tions of Haiti (n5111) and Jamaica (n5119). All ethi-
cal guidelines were followed as set forth by the Florida
International University Institutional Review Board
(IRB). DNA extractions were performed using the
Gentra Buccal Cell Kit according to the manufacturer’s
specifications (Puregene, Gentra Systems, Minneapolis,
MN) and stored at 2808C.
Published data
A total of 35 geographically targeted reference popula-
tions reporting 15 loci and 36 collections genotyped at 13
loci were included for comparison to the Haitian and Ja-
maican groups presented in this study. The African
(East, West, and South) populations were chosen to
assess their impact on Haiti and Jamaica, since both
islands are known to have received genetic contributions
from this region during slave trade and post–slave trade
events. Inclusion of the Latin American collections is
warranted because of the substantial African component
detected by Simms et al. (2008). In addition, the East
Asian collections were included to survey the extent to
which this group influenced the Jamaican populace dur-
ing the indentured labor system. All collections, their
abbreviations, biogeographical origin, and average heter-
ozygosity values are listed in Table 1.
DNA amplification and STR genotyping
The complete set of 15 autosomal STR loci (D8S1179,
D21S11, D7S820, CSF1PO, D3S1358, TH01, D13S317,
D16S539, D2S1338, D19S433, vWA, TPOX, D18S51,
D5S818, and FGA) was amplified simultaneously using
the AmpFlSTR Identifiler PCR Amplification Kit
(Applied Biosystems, Foster City, CA) according to the
manufacturer’s instructions (Applied Biosystems, 2001).
Amplified products were separated and detected by high-
performance multicapillary electrophoresis in an ABI
3130xl Genetic Analyzer with POP 7
TM
separation
medium. ABI GeneScan
TM
500 LIZ
TM
was used as the
internal size standard. The fragment analysis files gen-
erated were examined using the GeneMapper
1
software
v3.2 which allows for sample comparisons to the com-
mercially available allelic ladder and subsequent geno-
type assignment (Butler et al., 2004).
Statistical and phylogenetic analyses
Allelic frequencies for the general populations of Haiti
and Jamaica were estimated using the GenePop v 3.4
program (Raymond and Rousset, 1995). Using this data,
histograms comparing the distribution of alleles in
both collections per STR locus were constructed with
Microsoft Excel. The Arlequin version 2.000 software
(Schneider et al., 2000) was used to generate observed
and expected heterozygosities (H
o
and H
e
, respectively)
along with Hardy-Weinberg equilibrium (HWE) Pvalues
and gene diversity indices (GDI). The Bonferroni adjust-
ment (a50.05/15 50.00333 for 15 loci) was applied to
minimize possible type I errors. In addition, to assess
the effects of topogeographical barriers (e.g., the Carib-
bean Sea and the Atlantic Ocean) on gene flow and
possible inbreeding in the three Afro-Caribbean collec-
tions, heterozygote deficiencies (F
IS
and corresponding
Pvalues) were computed for each of the 15 loci with
GenePop v 3.4 (Raymond and Rousset, 1995).
Intra-, inter-, and total population (Hs, Gst, and Ht,
respectively) genetic variance values were calculated
with the DISPAN program (Ota, 1993). The groups
examined were assembled according to biogeographic ori-
gin as shown in Table 1. Delta (d) values for each of the
15 STR loci were also computed (Listman et al., 2007)
for the grouped East versus West African collections to
ascertain marker information content, i.e., the ability of
the Identifiler loci to differentiate between the two
groups of populations. Any locus yielding a dvalue
0.30 (Vergara et al., 2009) is considered informative for
discriminating between the two sources.
The Carmody’s G-test program (Carmody, 1990) was
used to assess the statistical significance of pair-wise
population comparisons through 100,000 simulations,
both before and after application of the Bonferroni cor-
rection (a50.05/666 50.000075). Comparisons yielding
Pvalues greater than aindicate genetic homogeneity
among the populations.
Multidimensional scaling (MDS) plots (with associated
stress values) were constructed with the statistical pack-
age SPSS version 14.0 (SPSS, 2001) using Nei’s genetic
distances (Fst) generated with the PHYLIP 3.52c soft-
ware (Felsenstein, 2002). Phylogenetic comparisons of
the Haitian and Jamaican collections with pertinent
available reference populations were performed across
the complete set of 15 loci and were repeated using 13 of
the loci included in the set of 15 to allow inclusion of the
Dominican Republic in the analysis. In addition, the
New World (Haiti, Jamaica, New Providence, and two
US African American populations) and continental
African groups [Madagascar was not included given its
known Austronesian ancestry (Hurles et al., 2005;
Regueiro et al., 2008)] were analyzed by themselves to
better assess phylogenetic relationships among them.
51PHYLOGENETIC RELATIONSHIPS OF NEW WORLD AFRICANS
American Journal of Physical Anthropology
The aforementioned US African American collections,
which represent a Minnesotan (US1) and a general pop-
ulation from the United States (US2), were included as
additional representatives of New World Africans.
Admixture analyses were conducted to determine the
genetic contributions made to the gene pools of the
Haitian and Jamaican populations by three parental
groups encompassing the geographical regions of Africa,
East Asia and Europe [the Latin American collections
were not included as a parental source because of the
substantial proportion of their autosomal component
shared with continental Africans and European (Simms
et al., 2008)]. The Bahamian population of New Provi-
dence and the two US African American collections (US1
and US2) previously investigated by Simms et al. (2008)
were also reexamined for their source populations in
light of additional African data sets now available and
employed in the present study.
To calculate admixture proportions (m), three different
approaches were taken. Using the SPSS version 14.0
software (SPSS, 2001), genetic contributions from the pa-
rental sources were generated using the weighted least
squares (WLS) method proposed by Long et al. (1991).
Admixture was also quantified with the ADMIX 2.0 pro-
gram, which employs the coalescent-based approach
described by Bertorelle and Excoffier (1998). Standard
deviations were computed with bootstrapping imple-
menting 1,000 reiterations. Additionally, with the
LEADMIX program (Wang, 2003), admixture propor-
tions were computed using the least square regression
method of Roberts and Hiorns (1965). Each of the afore-
mentioned admixture estimators are referred to
throughout the remainder of the article as m
L
,m
Y
,and
m
RH
, respectively.
The admixture tests (Long et al., 1991) as well average
heterozygosity calculations were repeated for the five
New World African groups following the segregation
of the 15 STR markers into slow (0.001 mutations/
meioses) vs. fast (0.01 mutations/meioses) mutating
loci. The analyses were performed to examine the effects
of mutation rate on admixture proportions and genetic
diversity indices. Mutation rates for the Identifiler loci
TABLE 1. Populations analyzed
Biogeographical origin Population Abbreviation nAvg He Reference
Afro-Caribbean
a
Haiti HAT 111 0.80300 Present study
Jamaica JAM 119 0.77647 Present study
New Providence (Bahamas) NWP 221 0.78312 Simms et al., 2008
Afro-American
a
US African American 1
(Minnesota)
US1 157 0.80380 Gross and Budowle, 2006
US African American 2
(General)
US2 258 0.79573 Butler et al., 2003
West African Cabinda (Angola) ANG 110 0.79698 Beleza et al., 2004
Guinea-Bissau GUB 92 0.80249 Pereira et al., 2005
Equatorial Guinea EGU 134 0.89193 Alves et al., 2005
Namibia NAM 195 0.78293 Muro et al., 2008
East African Hutu (Rwanda) HUT 95 0.81193 Shepard and Herrera, 2006
Kenya KEN 65 0.81436 Shepard and Herrera, 2006
Madagascar MAD 67 0.80350 Regueiro et al., 2008
Maputo (Mozambique) MOZ 144 0.79580 Alves et al., 2004
Tanzania TAN 272 0.81062 Forward et al., 2008
Tutsi (Rwanda) TUT 126 0.77818 Regueiro et al., 2004
Uganda UGA 90 Gusma
˜o et al., 2006
South African Cape Town (South Africa) SAF 98 Kido et al., 2007
East Asian Chao Shan (South China) CCS 144 0.77793 Hu et al., 2005
Japan JAP 526 0.77127 Hashiyada et al., 2003
Korea KOR 231 0.77087 Kim et al., 2003
Malaysia MAL 210 0.78127 Seah et al., 2003
Philippines PHI 106 0.77793 De Ungria et al., 2005
Shaanxi Han (NW China) CSH 203 0.78232 Wang et al., 2005
Latin American Caracas (Central Venezuela) CAR 255 0.79267 Chiurillo et al., 2003
Central Mexico MEX 211 0.82827 Herna
´ndez-Gutie
´rrez
et al., 2005
Costa Rica CSR 500 0.85353 Rodrı
´guez et al., 2007
Dominican Republic
b
DMR 2565 0.89677
´az et al., 2008
Hispanic (US - Minnesota) HIS 151 0.80120 Gross and Budowle, 2006
Maracaibo (Venezuela) MAR 203 0.77896 Bernal et al., 2006
Metztitla
´n (Mexico) MET 180 0.75067 Gorostiza et al., 2007
Puerto Rican American
(Massachusetts)
PRA 205 0.79113 Zu
´n
˜iga et al., 2006
Rio de Janeiro (Brazil) BRA 300 0.85533 de Souza Go
´es et al., 2004
European Andalusia (South Spain) AND 114 0.77600 Coudray et al., 2007
Belaruse BLR 176 0.77160 Rebala et al., 2007
Caucasian (US - General) CAU 302 0.78173 Butler et al., 2003
Flemish (North Belgium) FLE 231 0.79533 Mertens et al.,2006
North and Central Poland POL 412 0.79447 Czarny et al., 2005
Spain SPN 342 0.76666 Camacho et al., 2007
a
These groups are combined in some analyses as New World Africans. For specific analyses see Materials & Methods.
b
The Average Heterozygosity (Avg He) for this group was calculated at the level of 13 loci.
52 T.M. SIMMS ET AL.
American Journal of Physical Anthropology
were provided by the NIST STR database at http://
www.cstl.nist.gov/div831/strbase/mutation.htm.
The Structure software v. 2.3.1, available at http://
pritch.bsd.uchicago.edu/structure/html, was used to infer
population substructure for both the Haitian and Jamai-
can collections (Pritchard et al., 2000). After uploading
the genotypic data into the program, a length of burn-in
period of 20,000 was selected and the number of Markov
chain Monte Carlo (MCMC) repetitions after burn-in
was also set to 20,000. When updating the ancestry
model specifications, the admixture model that performs
clustering without using population of origin information
and that assumes the allele frequencies in the popula-
tion to be independent, was chosen. In the current study,
the Structure analyses were performed with the number
of ancestral population fixed at K53and K54.
RESULTS
Intrapopulation diversity
Allelic frequencies for the 15 autosomal STR loci along
with H
o
,H
e
, HWE pvalues and GDI are listed in
Tables 2 and 3 for Haiti and Jamaica, respectively. His-
tograms generated from the allele-frequency data gener-
ally reveal no differences in the distribution of alleles
between the two collections but do illustrate variation in
the frequencies of alleles across loci (Supporting Infor-
mation Figure 1). In Jamaica, two loci (D16S539 and
D2S1338) deviate from HWE expectations (a50.05) but
were found to be statistically insignificant after applica-
tion of the Bonferroni adjustment. Interestingly, in the
Haitian population, 10 loci (D8S1179, D21S11, CSF1PO,
D16S539, D2S1338, D19S433, vWA, TPOX, D5S818, and
FGA) possess H
o
values exceeding H
e
, whereas only
three loci (D8S1179, D18S51, and FGA) in the Jamaican
collection have higher than expected heterozygozities.
Haiti and Jamaica both exhibit the highest genetic vari-
ability at locus D2S1338, with a GDI of 0.8887 and
0.8877, respectively, while D13S317, with values of
0.6717 for Haiti and 0.7132 for Jamaica, was found to be
least variable in both populations. The combined power
of exclusion (CPE) was [99.9% for both collections; how-
ever, noticeable differences between the two populations
were detected when comparing the combined power of
discrimination (CPD), 0.733211 for Haiti versus 0.69732
for Jamaica, and average heterozygosities, 0.80300 ver-
sus 0.77647, with Jamaica exhibiting the lower of the
two values.
The number of alleles present in the Haitian popula-
tion (160) and the Jamaican collection (158) outnumber
the average allele count (149) for the continental African
populations and are comparable in number to the other
New World collections of African descent [New Provi-
dence (173), US African American 1 (162) and US Afri-
can American 2 (168) (Simms et al., 2008)]. The Jamai-
can populace shares alleles D7S820-6 and FGA-45.2 with
Tanzania, whereas the Haitian collection possesses allele
D16S539-6 characteristic of Angola. Both Haiti and
Jamaica, along with New Providence, exhibit FGA-16.1
present in the East African nations of Madagascar and
Tanzania. In addition, only two of the five New World
African groups, Haiti and US African American 1 (US1),
carry allele FGA-24.3, which is not present in any conti-
nental African collection examined in this report. Other
alleles shared between these island populations and the
mainland African groups include FGA-19.2 and 30.2.
The highest intrapopulation variances (Hs) are
observed in the New World African populations (0.79889)
and the Afro-Caribbean collections (0.79712), both of
which exceed the values seen in the continental Africans
including (0.79411) and excluding Madagascar (0.79356).
The lowest Hs levels are observed among the European
(0.78461) and East Asian (0.78131) groups of populations
(Table 4). Despite the high-diversity levels observed for
the Afro-Caribbean group, F
IS
calculations and associ-
ated Pvalues (Table 5) reveal two loci in Haiti (D7S820,
TH01), two loci in Jamaica (TH01, D2S1338), and five
loci in New Providence (D21S11, CSF1PO, D19S433,
TPOX, and FGA) that exhibit lower than the statistically
expected number of heterozygotes.
Interpopulation diversity
Phylogenetic relationships among all populations
examined in this study were investigated using MDS
projections, pair-wise population comparisons (G-test)
and admixture analyses. In addition, Gst and Ht values
were computed to assess inter- and total population var-
iance, respectively, whereas dvalues were calculated to
evaluate the ability of the 15 Identifiler loci to statisti-
cally differentiate between the grouped East and West
African populations.
In the MDS plot of all populations (Fig. 1A), three
groupings according to biogeographical origin are evi-
dent: an East Asian cluster in the lower left quadrant, a
Latin American/European assemblage in the upper left
section (although a clear partitioning between the two is
apparent), and an African group occupying the right half
of the graph. At the periphery of the African cluster,
Jamaica, along with the two US African-American collec-
tions (US1 and US2), New Providence and Angola, segre-
gate away from the mainland African populations in the
direction of the Latin American/European conglomerate.
However, when only the African and African derived col-
lections are employed (Fig. 1B), Jamaica plots on the
outskirts of the central cluster at a considerable distance
from Angola and the New World African populations. In
contrast, Haiti maps among the continental Africans in
both dendrograms (Fig. 1A,B) but lies closest to two of
the West African collections (Guinea-Bissau and Equato-
rial Guinea) when all the populations are examined (Fig.
1A) and displays greater genetic affinities with Angola
(Dimension 1), US2, and Equatorial Guinea (Dimension
2) when only the African and African derived groups are
considered (Fig. 1B). At 13 loci, Haiti plots distantly
from the general populace of the Dominican Republic
(Supporting Information Figure 2), thereby suggesting a
differential contribution of source populations as well as
limited gene flow between the two portions of the island.
Although the Latin American collections plot close to the
European groups (Fig. 1A), they clearly segregate in two
directions: the Hispanics from Minnesota in the direction
of the East Asian cluster while the Brazilians and
Puerto Rican Americans head toward the African collec-
tions. The tendency of some Latin American groups to
segregate toward the East Asian collections is possibly
the result of the Amerindian component in those New
World populations. The stress value for the all popula-
tion projection was 0.10202, whereas estimates for the
African and African derived groups alone led to an out-
put of 0.19149.
53PHYLOGENETIC RELATIONSHIPS OF NEW WORLD AFRICANS
American Journal of Physical Anthropology
TABLE 2. Haitian allelic frequencies (n5111)
Allele D8S1179 D21S11 D7S820 CSF1PO D3S1358 TH01 D13S317 D16S539 D2S1338 D19S433 vWA TPOX D18S51 D5S818 FGA
5 0.0090 0.0090
6 0.1036 0.0045 0.0946
7 0.0045 0.0360 0.4775 0.0135
8 0.0045 0.2297 0.0676 0.2207 0.0045 0.0495 0.2928 0.0631
9 0.1532 0.0225 0.1486 0.0090 0.1937 0.2117 0.0045 0.0180
9.3 0.0315
10 0.0090 0.3018 0.2613 0.0090 0.0270 0.0856 0.0180 0.0811 0.0045 0.0721
10.1 0.0045
10.2 0.0045
11 0.0495 0.1667 0.2523 0.2928 0.2793 0.0450 0.0045 0.2748 0.1937
12 0.0586 0.1036 0.2883 0.4730 0.2027 0.1081 0.0045 0.0270 0.0586 0.3784
12.2 0.0495
13 0.1847 0.0315 0.0586 0.1306 0.1351 0.2838 0.0045 0.0045 0.0405 0.2387
13.2 0.0495 0.0045
14 0.3874 0.0045 0.0090 0.1261 0.0631 0.0360 0.1937 0.0991 0.0586 0.0270
14.2 0.0676 0.0090
15 0.2432 0.0045 0.3063 0.0045 0.0856 0.1937 0.1261
15.2 0.0586 0.0045
16 0.0541 0.3108 0.0676 0.0135 0.3108 0.2162 0.0045
16.1 0.0045
16.2 0.0180
17 0.0090 0.2117 0.1081 0.0045 0.1757 0.1532 0.0045
17.2 0.0045
18 0.0360 0.0405 0.1261 0.1036 0.0045
18.2 0.0045
19 0.0090 0.1712 0.0721 0.0991 0.0450
20 0.0676 0.0856 0.0586
20.2 0.0045
21 0.1396 0.0045 0.0180 0.0991
21.2 0.0045
22 0.1396 0.0045 0.0045 0.1847
23 0.1261 0.1802
24 0.0676 0.1667
24.3 0.0045
25 0.0631 0.1486
26 0.0045 0.0090 0.0495
27 0.0315 0.0315
28 0.3018
29 0.1532
30 0.2072
30.2 0.0180 0.0045
31 0.0721
31.2 0.0631 0.0090
32 0.0225
32.2 0.0495
33 0.0045
33.2 0.0090
34 0.0135
54 T.M. SIMMS ET AL.
American Journal of Physical Anthropology
Inter- and total population diversity indices (Gst and
Ht, respectively) are presented in Table 4. The highest
Gst values are recorded for the All populations group
(0.02400) and are lowest among the New World African
collections (0.00330), likely attributed to the genetic con-
tributions made to the populations in the group by simi-
lar sources. Total population variance (Ht) is also highest
in the All populations assemblage (0.81014) and is
followed in decreasing order by the Africans including
(0.80320) and excluding (0.80169) Madagascar, New
World Africans (0.80154), Afro-Caribbeans (0.79984),
Latin Americans (0.79840), East Asians (0.78807), and
Europeans (0.78789).
dvalues calculated for the grouped East versus West
African collections are provided in Table 6. The average
dacross the entire set of 15 loci was 0.088, and values at
each locus for the pair ranged from 0.045 to 0.117. Since
no locus attained a value 0.30, these markers are not
considered informative for statistically differentiating
between the two groups of populations.
In the G-test pair-wise analyses (Table 7), statistically
insignificant genetic differences are observed between
several of the New World and continental African collec-
tions, indicating a close genetic relationship between the
two groups of populations. Specifically, all five New
World African populations were not found to differ sig-
nificantly from Kenya or Equatorial Guinea. G-test results
also indicate genetic similarities between the Afro-Carib-
bean collections and Guinea-Bissau as well as between
Haiti, Jamaica, US1, and Angola. Given the genetic affin-
ities displayed by the New World and continental African
collections, in combination with the large proportion of
enslaved Africans transported to the New World, it is not
surprising that results from the pair-wise population
comparisons reflect genetic homogeneity between the Afro-
Caribbean and Afro-American collections.
Admixture estimates for the New World African collec-
tions employing grouped African, East Asian, and Euro-
pean populations as parentals are provided in Table 8.
The results generated from each of the three methods
used, as well as for the admixture tests involving the
segregated loci (Supporting Information Table 1), collec-
tively indicate that the most significant genetic contribu-
tions made to the gene pools of all five hybrid popula-
tions originate from Africans. In particular, m
L
and m
RH
estimates demonstrate that of all the New World collec-
tions, the Haitian populace receives the largest African
genetic component, sharing greater than 95% of its DNA
with this group, while comparable levels of genetic
input (ranging from 69.7% to 83.7%, depending on the
approach taken to quantify admixture) from African
sources are detected in the other hybrid populations.
The remaining genetic influences observed in New World
Africans, with the exception of New Providence, arise
from both the European and East Asian collections. In
Jamaica genetic contributions from the European (m
L
5
16%, m
Y
517.9% and m
RH
514.2%) and East Asian
(m
L
55.7%, m
Y
55.65% and m
RH
53.73%) sources are
commensurate among the methods employed, whereas in
Haiti and the two Afro-American populations, the degree
of European and East Asian admixture varies according
to the method employed. Interestingly, of the New World
African groups, Jamaica is the only collection to exhibit
significant variation in admixture proportions between
slow and fast mutating loci.
The divergent genetic signals observed in the admix-
ture profiles for Haiti and Jamaica are not reflected in
TABLE 2. (Continued)
Allele D8S1179 D21S11 D7S820 CSF1PO D3S1358 TH01 D13S317 D16S539 D2S1338 D19S433 vWA TPOX D18S51 D5S818 FGA
34.2 0.0045
35 0.0405
36 0.0045
Ho 0.8198 0.8559 0.7387 0.8378 0.7207 0.6757 0.6306 0.8198 0.8919 0.8829 0.8469 0.8108 0.8649 0.7568 0.8919
He 0.7511 0.8308 0.7967 0.7786 0.7509 0.6924 0.6717 0.8176 0.8887 0.8510 0.8076 0.7810 0.8809 0.7555 0.8685
HWE 0.3208 0.3520 0.3609 0.4647 0.6197 0.0740 0.7018 0.9226 0.4855 0.2813 0.9874 0.3768 0.9021 0.8624 0.6072
GDI 0.7511 0.8308 0.7967 0.7786 0.7508 0.6924 0.6717 0.8176 0.8887 0.8510 0.8076 0.7810 0.8809 0.7555 0.8685
Ho: observed heterozygosity; He: expexted heterozygosity; HWE: Hardy-Weinberg equilibrium pvalues; GDI: gene diversity index.
55PHYLOGENETIC RELATIONSHIPS OF NEW WORLD AFRICANS
American Journal of Physical Anthropology
TABLE 3. Jamaican allelic frequencies (n5119 )
Allele D8S1179 D21S11 D7S820 CSF1PO D3S1358 TH01 D13S317 D16S539 D2S1338 D19S433 vWA TPOX D18S51 D5S818 FGA
6 0.0084 0.1429 0.0336
7 0.0084 0.0546 0.3655 0.0210
8 0.1639 0.0672 0.2311 0.0336 0.0168 0.4286 0.0588
9 0.0126 0.0798 0.0420 0.1555 0.0294 0.2521 0.0042 0.1807 0.0210
9.3 0.0840
10 0.0378 0.3992 0.2941 0.0210 0.0336 0.1134 0.0084 0.0714 0.0420
10.2 0.0042
11 0.0714 0.2101 0.2017 0.2731 0.2983 0.0546 0.0084 0.2437 0.0084 0.2689
11.2 0.0084
12 0.1134 0.1176 0.2647 0.0042 0.4244 0.2101 0.1218 0.0210 0.0714 0.3445
12.2 0.0546
13 0.1513 0.0084 0.0630 0.1765 0.0924 0.2563 0.0084 0.0630 0.2479
13.2 0.0462 0.0042
14 0.2857 0.0042 0.0126 0.0630 0.0294 0.0168 0.2185 0.0630 0.0798 0.0084
14.2 0.0588 0.0084
15 0.2227 0.2983 0.0798 0.2437 0.1429 0.0042
15.2 0.0042 0.0252
16 0.0882 0.3109 0.0378 0.0168 0.2479 0.1975 0.0042
16.1 0.0042
16.2 0.0378
17 0.0168 0.2521 0.0966 0.0042 0.1681 0.1513
17.2 0.0042
18 0.0588 0.0630 0.1345 0.1092
18.2 0.0252
19 0.0084 0.2017 0.0882 0.1050 0.0588
19.2 0.0042
20 0.1176 0.0252 0.0336 0.0336
20.2 0.0042
21 0.1218 0.0042 0.0084 0.1050
21.2 0.0084
22 0.1092 0.0084 0.0084 0.1597
23 0.0966 0.0042 0.1891
24 0.0840 0.2017
25 0.0588 0.1008
26 0.0042 0.0084 0.0420
27 0.0336 0.0042 0.0336
28 0.2521 0.0168
29 0.2101 0.0042
30 0.1849 0.0042
30.2 0.0042
31 0.0630
31.2 0.0546
32 0.0294
32.2 0.0630
33 0.0084
33.2 0.0378
56 T.M. SIMMS ET AL.
American Journal of Physical Anthropology
the output of the Structure analyses. The bar plots gen-
erated for both groups (Supporting Information Figure
3), assuming either three (K53) or four (K54) ances-
tral populations, reveal no population substructure in ei-
ther collection. The graphs also indicate that both groups
receive equivalent contributions from the same ancestral
source populations. The general lack of population sub-
structure observed in Haiti and Jamaica is likely attrib-
uted to the limited number of loci (15) employed in the
analyses (Listman et al., 2007), which according to
Ardlie and colleagues (2002) should be much larger
([100) to reliably differentiate between highly admixed
populations sharing close genetic ties.
DISCUSSION
The Antilles is recognized as a ‘‘great melting pot,’’
(Knight and Crahan, 1979) with influences that stem
from Africa, Europe, and East Asia; however, the extent
to which these ancestral source populations have con-
tributed genetically to the island chain remains rela-
tivelyunexplored.Thus,inthisstudy,15autosomal
STR loci were typed in the general populations of Haiti
and Jamaica to assess the genetic implications of their
known divergent colonial histories and associated mi-
gratory events. This investigation represents the first
time that Haitian STR allelic frequency data are pre-
sented, whereas the Jamaican population is examined
across the full set of 15 STR loci since D2S1338 and
D19S433 were not previously reported (Budowle et al.,
1999). Inclusion of additional loci in the data set not
only increases the phylogenetic resolution between
closely related populations, but also generates more ro-
bust probability calculations important in forensic and/
or paternity testing.
Genetic diversity in the New World is expected to be
high due to genetic contributions from various founder
populations and subsequent admixture events over the
last 500 years. Historical accounts for the West Indies
indicate extensive gene flow from continental Africa dur-
ing the period of the Transatlantic slave trade in addi-
tion to varying degrees of genetic input from European,
Asian, and/or other sources (Parra et al., 1998;
Saunders, 2003; Herrera et al., 2004; Pepin, 2005; Benn
Torres et al., 2007; Benn-Torres et al., 2008). For
instance, Jamaica, whose motto is ‘‘Out of many one peo-
ple,’’ has been settled by a wide array of settlers includ-
ing enslaved Africans, Portuguese Jews, Scottish and
Irish bond servants, German and Chinese indentured
laborers, and later on Syrian (Henriques, 1964), Cuban
and Haitian (Thomas-Hope, 2003) nationals, whereas
Haiti, the ‘‘world’s first Black Republic,’’ has received
gene flow, for the most part, exclusively from Africa.
Given the high level of diversity anticipated for the
region, it is not surprising that average heterozygosity
values for all five New World collections (Table 1) are
comparable to the combined average heterozygosity of
the mainland African populations (0.79778) included in
the present study. With respect to Haiti (0.80300) and
Jamaica (0.77647), noteable differences in average hetero-
zygosities are observed for each collection upon segrega-
tion of the STR markers into slow versus fast mutating
loci [0.74325 versus 0.66387 (slow) and 0.81200 versus
0.79380 (fast) for Haiti and Jamaica, respectively], but
this disparity should be tempered considering the
unequal distribution of loci within the two categories.
The New World Africans also exhibit the highest intra-
TABLE 3. (Continued)
Allele D8S1179 D21S11 D7S820 CSF1PO D3S1358 TH01 D13S317 D16S539 D2S1338 D19S433 vWA TPOX D18S51 D5S818 FGA
34.2 0.0084
35 0.0336
36 0.0126
45.2 0.0042
Ho 0.8740 0.7983 0.7143 0.7899 0.7227 0.6555 0.6807 0.7563 0.8235 0.8319 0.8151 0.6723 0.8908 0.7311 0.8908
He 0.8217 0.8459 0.7524 0.7927 0.7464 0.7641 0.7132 0.7846 0.8877 0.8548 0.8237 0.7202 0.8815 0.7449 0.8720
HWE 0.5772 0.3424 0.9637 0.9435 0.7683 0.1598 0.1949 0.0178 0.0040 0.3589 0.4727 0.6787 0.6876 0.9691 0.4080
GDI 0.8217 0.8459 0.7524 0.7927 0.7464 0.7641 0.7132 0.7846 0.8877 0.8548 0.8237 0.7202 0.8815 0.7449 0.8720
Ho: observed heterozygosity; He: expexted heterozygosity; HWE: Hardy-Weinberg equilibrium pvalues; GDI: gene diversity index.
57PHYLOGENETIC RELATIONSHIPS OF NEW WORLD AFRICANS
American Journal of Physical Anthropology
population variance level of any group examined
(Table 4).
Although dvalues generated for the 15 Identifiler loci
(Table 6) do not support the segregation of the African
collections into East and West, noticeable differences
between the two groups of populations are observed in
both the MDS analysis (Fig. 1B) and the G-test (Table
7). In the MDS plot of African and African derived popu-
lations, noteworthy is the partitioning of the East Afri-
can collections. Although the East African collections
from Mozambique, Tanzania, and Kenya were found to
group with the West African (Angola, Equatorial Guinea,
and Guinea-Bissau) and New World populations in the
central cluster, the Uganda, Hutu, and Tutsi collections
lie on the fridges of this assemblage. A plausible expla-
nation for the lack of discrimination observed between
the East and West African populations is the Bantu
expansion, which is known to have been a venue for the
demic diffusion of a number of West African markers,
including Y-haplogroups E1b1a, E2, and B2a (Berniell-
Lee et al., 2009), as well as mtDNA haplogroup L sub-
clades (specifically L1a and L2a), into East Africa (Salas
et al., 2002). Moreover, the Bantu diaspora is also likely
the cause for genetic similarities observed in the G-test
between Kenya and the West African groups (with the
exception of Namibia).
Admixture proportions generated for the five New
World African populations (Table 8), regardless of the
method employed, indicate that their strongest genetic
signal derives from the African continent. Given the
large number of African laborers [estimates ranging
from about 6,000 (Dominican Republic) to greater than
1 million (Brazil) slaves (Pepin, 2005)] transported to the
region during the period of the Transatlantic slave trade,
these findings are not surprising. In Jamaica, the rela-
tively high rate of the YAP insertion [ranging from
17.89% (Parra et al., 1998) to 59.09% (Benn Torres et al.,
2007)] in combination with elevated levels of Y-hap-
logroup E1b1a-M2 (about 43%) (Athey, 2006; Benn
Torres et al., 2007) and mtDNA haplogroup L (25.86%)
Fig. 1. A. Multidimensional scaling plot (stress 50.10202) of all populations at 15 loci. B. Multidimensional scaling plot
(stress 50.19149) of the African and African derived collections.
58 T.M. SIMMS ET AL.
American Journal of Physical Anthropology
TABLE 4. Inter-, intra-, and total population genetic variance
Locus
Afro-Caribbean New World African
a
African w/o
Madagascar
African with
Madagascar East Asian Latin American European All populations
Gst Hs Ht Gst Hs Ht Gst Hs Ht Gst Hs Ht Gst Hs Ht Gst Hs Ht Gst Hs Ht Gst Hs Ht
D8S1179 0.00476 0.78859 0.79236 0.00493 0.79008 0.79399 0.01146 0.77362 0.78259 0.01253 0.77533 0.78517 0.00762 0.84279 0.84926 0.00533 0.79687 0.80114 0.00469 0.80477 0.80856 0.02144 0.79769 0.81517
D21S11 0.00262 0.83717 0.83937 0.00249 0.84052 0.84262 0.00937 0.85160 0.85966 0.01000 0.85301 0.86162 0.00871 0.81690 0.82408 0.00407 0.83425 0.83765 0.00263 0.84413 0.84636 0.01917 0.83997 0.85639
D7S820 0.00549 0.77387 0.77814 0.00461 0.77871 0.78231 0.00811 0.76163 0.76786 0.00910 0.76344 0.77045 0.00944 0.75886 0.76608 0.00753 0.79102 0.79702 0.00428 0.80119 0.80463 0.02157 0.77684 0.79397
CSF1PO 0.00214 0.78204 0.78372 0.00187 0.78084 0.78230 0.01527 0.77695 0.78899 0.01457 0.77571 0.78718 0.00874 0.72283 0.72920 0.00330 0.71758 0.71996 0.00529 0.72430 0.72815 0.01486 0.74692 0.75819
D3S1358 0.00213 0.74892 0.75052 0.00349 0.75207 0.75470 0.01227 0.73200 0.74109 0.01231 0.72975 0.73884 0.01191 0.71936 0.72803 0.01200 0.75931 0.76853 0.00319 0.79479 0.79733 0.01623 0.74802 0.76036
TH01 0.00588 0.73376 0.73810 0.00472 0.73893 0.74243 0.01389 0.72753 0.73778 0.01299 0.72930 0.73889 0.01452 0.69914 0.70944 0.01458 0.77066 0.78206 0.00847 0.77510 0.78172 0.05882 0.74208 0.78845
D13S317 0.00331 0.70077 0.70309 0.00247 0.70627 0.70802 0.00704 0.70777 0.71278 0.01095 0.71409 0.72199 0.00608 0.80024 0.80513 0.01072 0.80819 0.81695 0.00354 0.77766 0.78042 0.03782 0.75766 0.78744
D16S539 0.00363 0.79869 0.80159 0.00348 0.79425 0.79702 0.00870 0.78878 0.79570 0.00967 0.78850 0.79619 0.00957 0.77487 0.78235 0.00764 0.77957 0.78557 0.00244 0.75587 0.75772 0.01811 0.77984 0.79423
D2S1338 0.00216 0.88510 0.88702 0.00247 0.88726 0.88945 0.01061 0.88152 0.89097 0.01043 0.88107 0.89036 0.00706 0.86761 0.87377 0.00894 0.86449 0.87229 0.00414 0.86760 0.87121 0.02051 0.87396 0.89225
D19S433 0.00135 0.84174 0.84288 0.00214 0.84083 0.84263 0.01017 0.82757 0.83607 0.01120 0.82926 0.83865 0.01380 0.79961 0.81079 0.00778 0.82661 0.83310 0.00290 0.76447 0.76670 0.02022 0.81494 0.83175
vWA 0.00342 0.81135 0.81414 0.00317 0.81307 0.81565 0.00947 0.81916 0.82700 0.00980 0.81737 0.82546 0.00681 0.79667 0.80213 0.00751 0.79039 0.79637 0.00403 0.80725 0.81051 0.01900 0.80596 0.82156
TPOX 0.00655 0.75879 0.76379 0.00491 0.76191 0.76567 0.01585 0.76994 0.78234 0.02124 0.76262 0.77917 0.00567 0.62619 0.62976 0.00926 0.67459 0.68089 0.00481 0.61521 0.61818 0.03776 0.69746 0.72483
D18S51 0.00310 0.87691 0.87964 0.00262 0.87786 0.88017 0.00658 0.86494 0.87067 0.00708 0.86545 0.87162 0.00742 0.85343 0.85981 0.00356 0.87685 0.87998 0.00457 0.87159 0.87559 0.01820 0.86864 0.88474
D5S818 0.00273 0.74781 0.74986 0.00259 0.75047 0.75242 0.00627 0.74582 0.75053 0.01009 0.75214 0.75980 0.00761 0.78432 0.79033 0.01213 0.71660 0.72540 0.00400 0.71021 0.71306 0.02917 0.74265 0.76496
FGA 0.00229 0.87133 0.87333 0.00378 0.87035 0.87366 0.00776 0.87454 0.88138 0.00897 0.87468 0.88260 0.00468 0.85688 0.86091 0.00614 0.87367 0.87907 0.00358 0.85508 0.85814 0.01136 0.86781 0.87778
All loci 0.00339 0.79712 0.79984 0.00330 0.79889 0.80154 0.01015 0.79356 0.80169 0.01131 0.79411 0.80320 0.00858 0.78131 0.78807 0.00796 0.79204 0.79840 0.00415 0.78461 0.78789 0.02400 0.79070 0.81014
a
The New World African assemblage includes the Afro-Caribbean populations along with the two US African-American collections.
TABLE 5. Heterozygote deficiencies for the Afro-Caribbean populations
Locus
Haiti Jamaica New Providence
Ho He F
IS
PHo He F
IS
PHo He F
IS
P
D8S1179 0.8198 0.7511 20.0920 0.9934 0.8740 0.8217 20.0640 0.9383 0.8326 0.8017 20.0390 0.8563
D21S11 0.8559 0.8308 20.0300 0.1716 0.7983 0.8459 0.0560 0.1832 0.7783 0.8441 0.0780 0.0019
D7S820 0.7387 0.7967 0.0730 0.0270 0.7143 0.7524 0.0510 0.3209 0.7376 0.8287 0.0560 0.0792
CSF1PO 0.8378 0.7786 20.0760 0.7320 0.7899 0.7927 0.0040 0.5900 0.7195 0.8057 0.0820 0.0016
D3S1358 0.7207 0.7509 0.0400 0.4083 0.7227 0.7464 0.0320 0.5445 0.7557 0.7577 0.0030 0.4488
TH01 0.6757 0.6924 0.0240 0.0162 0.6555 0.7641 0.1430 0.0163 0.7557 0.7619 20.0040 0.4157
D13S317 0.6306 0.6717 0.0610 0.4027 0.6807 0.7132 0.0460 0.4918 0.7104 0.7767 0.0200 0.0597
D16S539 0.8198 0.8176 20.0030 0.5326 0.7563 0.7846 0.0360 0.1389 0.7919 0.8076 0.0130 0.0946
D2S1338 0.8919 0.8887 20.0040 0.6875 0.8235 0.8877 0.0730 0.0147 0.8552 0.8925 0.0380 0.0580
D19S433 0.8829 0.8510 20.0380 0.8002 0.8319 0.8548 0.0270 0.3725 0.7873 0.8292 0.0500 0.0491
vWA 0.8469 0.8076 20.0490 0.9187 0.8151 0.8237 0.0100 0.3964 0.7964 0.8117 0.0190 0.5672
TPOX 0.8108 0.7810 20.0380 0.7439 0.6723 0.7202 0.0670 0.3706 0.6833 0.7835 0.1280 0.0150
D18S51 0.8649 0.8809 0.0180 0.2617 0.8908 0.8815 20.0110 0.6247 0.8959 0.8815 20.0200 0.7320
D5S818 0.7568 0.7555 20.0020 0.2589 0.7311 0.7449 0.0190 0.5483 0.7330 0.7513 0.0240 0.0949
FGA 0.8919 0.8685 20.0270 0.8145 0.8908 0.8720 20.0220 0.9037 0.9140 0.8856 20.0350 0.0000
p\0.05 are in bold.
(Parra et al., 1998), corroborate the large African compo-
nent detected in this population in the present study.
Africa’s prominent contribution to the Haitian gene pool
is supported by an earlier D1S80 study in which Haiti
was found to cluster with continental Africans in PC and
ML analyses of worldwide populations (Herrera et al.,
2004). In addition, phylogenetic assessments performed
by Simms and colleagues (2008) reveal strong genetic
affinities between New Providence in the Bahamas, the
two US African American groups and the continental
Africans.
Of the five New World African groups, the percent con-
tribution from continental Africa is greatest in Haiti for
two of three admixture tests performed. The larger per-
centage of African DNA present in the Haitian gene pool
is supported by Haiti’s position amidst the African collec-
tions in the MDS plot of all populations (Fig. 1A). Within
this cluster, Haiti maps in closest proximity to Guinea-
Bissau and Equatorial Guinea in both Dimensions.
Likewise, when only the African and African derived pop-
ulations are considered, the Haitian populace still exhib-
its genetic affinities with the West Africans. In addition,
pair-wise population comparisons (Table 7) performed also
indicate genetic homogeneity between Haiti/Angola, Haiti/
Equatorial Guinea, and Haiti/Guinea-Bissau. These find-
ings are supported by a previous study, based on the
D1S80 polymorphic locus, in which Haiti was found to
map equidistant from Benin and a general collection from
the Bahamas in the PC plot of African and New World
African populations (Herrera et al., 2004).
Admixture data for the Jamaican collection reveals
commensurate contributions from Africa (ranging from
76.5% to 82.1%), regardless of the method employed to
quantify admixture. Genetic ties with the African conti-
nent are not only observed in the clustering of Jamaica,
along with the other New World African groups, on the
fringes of the African assemblage in Figure 1A, but are
also detected in the results of the pair-wise comparisons
(Table 7), which indicate genetic similarities between
Jamaica, the West African collections (Angola, Equato-
rial Guinea, and Guinea-Bissau), and Kenya. Within the
assembly of New World populations (Fig 1A), noteworthy
is the position of Angola, which maps distantly from the
remaining continental African groups. The close proxim-
ity of Jamaica and Angola in Figures 1A (Dimensions
1 and 2) and B (Dimension 2) most likely derives from
the large proportion of enslaved Africans [approximately
142,725 persons according to Pepin (2005)] transported
to the island from West-Central Africa, the region
encompassing Angola and the Congos. Genetic influences
in Jamaica from West Africa have also been detected in
earlier studies in which elevated levels (approximately
70%) of the Benin b
S
-haplotype, prevalent throughout
West-Central Africa [Cameroon, Nigeria, and Benin),
have been reported (Wainscoat et al., 1983; Antonarakis
et al., 1984; Nagel et al., 1984; Pante-de-Sousa, 1998).
Also, in a more recent study examining mtDNA haplo-
types, the Jamaican population was found to branch to-
gether with samples native to Sierra Leone in NJ trees
(McLean et al., 2005). This clustering is expected since
indentured servants from this region of West Africa
were transported to Jamaica during the postslavery pe-
riod.
The comparable African admixture proportions exhib-
ited by the New Providence and US African American
collections (US1 and US2) echoes data published by Simms
et al. (2008). However, with the introduction of additional
African populations in the present study (Guinea-Bissau,
Namibia, Tanzania, and Uganda), the contributions from
Africa have decreased on average by 0.93%, 5.76%, and
10.8% when the Roberts and Hirons (1965), Long (1991),
and Bertorelle and Excoffier (1998) approaches were
employed, respectively. Despite this decrease, genetic affin-
ities between these collections and the continental African
groupsarestillthemostprominent.
The remaining influences present in each of the New
World African population’s gene pool are contributed by
the European groups and, except for the New Providence
collection, the East Asian populations as well. The
genetic input from Europeans is minimal in Haiti (with
the exception of the m
Y
estimator) when compared with
the other New World African populations that exhibit
much larger contributions (ranging from 14.2% to
36.3%). These findings are corroborated by the MDS plot
of all populations (Fig. 1A), which illustrates the latter
populations plotting closely together on the outskirts of
the African assemblage but in the direction toward the
European/Latin American group while the Haitian col-
lection maps amidst the mainland African populations
within the African cluster.
European gene flow into Jamaica is supported by previ-
ous studies which report admixture profiles portraying
from as little as no apparent contributions up to 2% when
mtDNA haplotypes were analyzed (McLean et al., 2003,
2005), 6.8% when autosomal SNP and INDEL frequency
distributions were employed (Parra et al., 1998), 12.4%
when a panel of 28 AIMs generated from SNP and INDEL
frequency data was utilized (Benn-Torres et al., 2008) and
18.1% when glucose-6-phosphate dehydrogenase (G6PD)
deficiency alleles were examined (Gibbs et al., 1972).
Genetic contributions from Europe most likely derive
from the influx of British soldiers, white bond servants
and, later, as a result of the immigration of Western Euro-
pean indentured laborers. In contrast to the other New
World African collections, the admixture proportions for
Haiti, with the exception of the coalescent based
approach, indicate only minor contributions from Euro-
pean sources. The limited presence of European DNA in
the current population is likely due to ramifications of the
Haitian Revolution in which the French colonists were ei-
ther killed or coerced out of the island.
TABLE 6. Delta (d) values for the grouped East versus
West African populations
Locus WA/EA
D8S1179 0.107
D21S11 0.095
D7S820 0.063
CSF1PO 0.117
D3S1358 0.045
TH01 0.104
D13S317 0.080
D16S539 0.110
D2S1338 0.063
D19S433 0.078
vWA 0.063
TPOX 0.105
D18S51 0.086
D5S818 0.097
FGA 0.105
Average 0.088
WA: West African; EA: East African.
60 T.M. SIMMS ET AL.
American Journal of Physical Anthropology
TABLE 7. G-test
Population US1 US2 ANG SAF EGU GUB HAT HUT JAM KEN MAD MOZ NAM NWP TAN TUT UGA
US1 231.3660 272.1790 396.6248 231.1540 284.6625 232.8402 290.1581 186.0889 202.7056 345.3930 369.3774 504.0229 211.5335 370.5138 487.7301 436.9200
US2 0.0506 292.7030 399.4874 244.9366 282.1639 227.0359 342.1582 216.0961 217.6445 382.9854 336.3618 544.4847 260.9949 400.2196 523.5327 483.2588
ANG 0.0001 0.0000 310.2818 221.7053 282.4472 239.2100 260.0201 262.8510 198.6299 371.3746 334.9184 290.9848 297.1790 327.6299 378.0364 468.1497
SAF 0.0000 0.0000 0.0000 333.2830 326.4199 279.4336 355.1657 348.8613 243.6538 385.7041 246.7927 419.0391 414.8787 330.5381 440.1677 510.8528
EGU 0.0309 0.0099 0.0238 0.0000 251.2966 169.9370 299.3431 235.1849 194.7626 364.9853 317.1875 352.9726 239.0506 334.4801 407.6942 456.9542
GUB 0.0000 0.0000 0.0000 0.0000 0.0005 238.6009 323.5085 251.3156 213.7887 354.1472 312.9706 394.3167 265.0812 321.6954 440.6943 416.2239
HAT 0.0440 0.0886 0.0120 0.0000 0.9480 0.0109 335.4467 230.7371 198.6350 332.5014 297.7143 337.0298 236.7576 308.1873 414.9631 416.9154
HUT 0.0000 0.0000 0.0001 0.0000 0.0000 0.0000 0.0000 301.1588 210.3549 376.5460 352.6128 462.6141 344.6294 333.5834 361.3040 328.6649
JAM 0.7290 0.1866 0.0002 0.0000 0.0122 0.0006 0.0462 0.0000 195.1255 333.7593 305.3622 419.8413 228.4719 299.7603 405.2145 367.2077
KEN 0.1050 0.0247 0.0916 0.0003 0.1682 0.0193 0.2982 0.0516 0.2052 306.9289 225.2373 299.9806 213.9525 227.2508 297.4595 241.7360
MAD 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 371.9299 506.4148 421.1929 437.2512 461.2226 501.2495
MOZ 0.0000 0.0000 0.0000 0.0005 0.0000 0.0000 0.0000 0.0000 0.0000 0.0035 0.0000 486.0583 408.3094 283.4961 523.5334 519.5912
NAM 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 548.6736 534.3686 634.0131 622.8959
NWP 0.3478 0.0142 0.0000 0.0000 0.0440 0.0009 0.0847 0.0000 0.1261 0.0754 0.0000 0.0000 0.0000 450.5210 484.4134 462.1212
TAN 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0365 0.0000 0.0001 0.0000 0.0000 481.9146 527.5989
TUT 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 537.8627
UGA 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0003 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
CCS 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
JAP 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
KOR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
MAL 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
PHI 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
CSH 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
CAR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
MEX 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
CSR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
HIS 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
MAR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
MET 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
PRA 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
BRA 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
AND 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
BLR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
CAU 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
FLE 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
POL 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
SPN 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000
TABLE 7. (Continued)
Population CCS JAP KOR MAL PHI CSH CAR MEX CSR HIS MAR MET PRA BRA AND BLR CAU FLE POL SPN
US1 1321.3734 1815.1772 1571.8273 1145.4186 956.6950 1366.2150 720.6636 1018.0546 1026.5992 681.2700 717.2574 1314.0172 618.4646 503.5044 681.6089 949.8528 1044.4484 1018.1482 1313.8867 1147.5273
US2 1612.5165 2444.9363 1978.6023 1474.4789 1150.3481 1668.1620 924.5727 1335.5060 1435.1771 863.0743 943.2885 1650.6353 830.7430 673.8070 873.9698 1193.3600 1413.5024 1345.8695 1775.0813 1548.8081
ANG 1319.5687 1743.0122 1547.0536 1230.0575 994.3469 1340.3058 893.2872 1170.4983 1226.6724 855.5585 906.1671 1417.8441 785.2439 656.7479 883.2368 1118.5775 1275.2731 1231.9263 1470.5598 1355.2976
SAF 1347.2368 1787.0132 1613.6172 1294.1763 1088.1547 1417.8231 951.1696 1255.0952 1228.6013 935.6351 1016.5126 1419.9254 881.2035 788.1117 946.0394 1259.1360 1347.4409 1312.3367 1584.3025 1425.7090
EGU 1410.5985 1908.0112 1703.4598 1264.7218 1045.0815 1449.3634 853.7404 1166.7423 1237.9307 842.0637 892.4207 1452.4832 754.5046 653.2888 831.1876 1117.4847 1275.5774 1177.8361 1488.0153 1311.7319
GUB 1101.0726 1417.4559 1327.3401 1044.8998 898.3178 1120.6151 669.1605 942.4453 929.3496 685.0397 746.6887 1180.8689 649.6805 563.4520 760.4780 960.4456 1036.7103 1000.3499 1215.0681 1098.9459
HAT 1258.7485 1702.6776 1540.2103 1153.0020 987.1977 1291.3613 781.3412 1056.8923 1114.7231 778.7883 821.7631 1295.0150 726.1169 589.4932 806.1568 1076.9105 1138.1791 1094.1213 1371.2069 1187.7039
HUT 1243.8484 1588.8331 1444.2677 1138.2894 1010.2482 1298.2859 803.4444 1042.8290 1083.9535 805.6285 805.2376 1316.9272 691.0500 647.6494 803.9218 982.7222 1098.8838 1081.3782 1249.1531 1185.1730
JAM 1105.7494 1451.3796 1325.4003 987.2772 861.8778 1130.8444 616.8490 880.0184 918.5863 637.9739 668.6646 1130.4785 577.1983 482.6524 666.4562 912.5247 978.4950 948.8259 1174.8984 1039.3077
KEN 965.3720 1132.9982 1112.2170 886.5229 784.0193 959.9223 577.6357 754.5463 736.7450 576.3684 587.2698 931.8091 535.9065 430.8232 619.4303 823.0012 835.1530 827.1734 983.0800 891.3480
MAD 708.5034 868.1801 857.8117 625.9515 527.1189 775.9848 553.3486 678.7587 725.1730 529.5678 579.8524 803.1285 558.0521 532.5978 656.8502 799.5281 863.2474 849.9046 943.9898 908.0583
MOZ 1516.8990 1978.8864 1779.2944 1456.1948 1166.9080 1575.8204 989.1948 1346.6300 1351.3932 956.1947 1024.2223 1573.4152 905.7759 824.1256 970.4476 1301.5133 1439.9630 1420.3245 1726.0402 1557.8480
NAM 1964.9546 2927.8337 2514.9470 1966.7240 1467.2649 2218.7803 1549.8578 1926.5803 2248.2058 1367.1688 1593.1064 2180.9568 1340.3010 1211.3700 1316.3779 1716.7960 2128.6575 1949.6779 2491.4854 2229.7070
NWP 1640.0826 2419.7532 2044.1027 1505.4727 1227.9337 1753.9027 916.2577 1304.4275 1401.6000 834.9598 918.8211 1625.9066 753.7961 658.0684 871.0754 1150.2979 1319.9277 1277.3502 1710.0082 1462.1938
TAN 1851.1737 2691.5598 2241.6169 1783.1946 1353.2587 1953.3273 1328.6235 1714.4022 1863.9269 1228.1897 1356.3922 2000.8979 1146.5701 1061.1766 1203.8531 1595.0778 1900.5359 1805.7930 2265.0481 2074.0374
TUT 1533.4893 1974.8872 1805.7122 1471.2946 1165.1117 1633.0477 1071.8092 1358.7449 1459.8461 1011.6371 1045.1583 1561.3545 976.0688 926.4795 976.5187 1242.6163 1480.4247 1403.4790 1682.7332 1564.5051
UGA 1159.6693 1464.1675 1357.8638 1083.6028 974.2226 1209.6449 863.1320 1019.6701 1109.8484 778.4283 827.5852 1233.6215 809.3176 715.9099 845.9526 1026.6661 1078.6869 1088.3894 1281.5101 1225.9478
CCS 431.2647 353.6910 432.0402 356.7864 279.7270 1125.1412 1056.0265 1340.8743 849.0118 1002.9550 1209.4363 1094.1124 1297.0906 889.4481 1176.4991 1356.3972 1272.7761 1577.0469 1384.5605
JAP 0.0000 293.7658 741.7219 497.2908 487.8763 1469.3901 1516.2504 1839.5123 972.6217 1249.6879 1583.5695 1436.7079 1865.7714 1030.0175 1547.7587 1952.4344 1806.0680 2485.0547 2096.0361
KOR 0.0000 0.0000 636.6891 459.4520 344.5786 1187.6215 1165.9557 1402.4498 844.1308 1037.3330 1330.2899 1180.2168 1468.2697 891.8792 1247.3016 1462.3920 1391.4364 1799.3508 1547.2969
MAL 0.0000 0.0000 0.0000 289.0813 472.2081 918.8443 877.5515 1212.1229 618.0319 803.8587 1130.5956 867.9807 1118.8350 695.2513 972.6280 1192.6473 1130.7333 1423.9149 1216.8807
PHI 0.0000 0.0000 0.0000 0.0000 377.1124 877.9362 854.0881 1016.2513 674.0782 786.1637 1106.5436 830.0125 957.3215 711.6191 914.1631 1044.2216 990.5612 1174.5063 1085.7383
CSH 0.0000 0.0000 0.0000 0.0000 0.0000 1131.8489 1023.6318 1295.5643 789.0430 979.0354 1224.3311 1062.3735 1312.8925 874.6797 1201.5341 1388.6058 1331.6224 1709.8162 1476.5314
CAR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 434.5750 291.5373 206.2046 194.9020 757.1349 323.2320 335.2340 351.6024 551.6054 503.3499 533.7566 753.9863 527.7271
MEX 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 545.4443 261.7632 378.3874 583.8787 574.1693 686.5186 493.6120 726.5411 783.9633 740.4211 1093.4510 840.4451
CSR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 248.0351 317.3796 779.3468 519.6517 604.9189 491.1570 716.8347 726.0587 752.0983 1151.3206 829.2176
HIS 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.1002 0.0023 0.0001 220.7598 442.0474 304.3576 352.9807 378.2934 507.0346 451.7975 484.1142 674.2834 510.1093
MAR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.4610 0.0000 0.0000 0.0103 611.7493 369.8294 340.7615 303.9688 479.3408 423.3698 457.7372 618.5443 461.2663
MET 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 941.7846 1130.1987 910.9257 1208.4436 1256.2998 1165.7612 1651.3641 1436.8680
PRA 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 353.1628 376.5916 593.3844 509.8419 543.5773 734.4340 528.5404
BRA 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 379.7395 570.6100 528.9551 564.9875 784.5393 570.0999
AND 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 259.4031 254.3532 240.6117 295.3663 191.7305
BLR 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 286.8939 272.8293 202.3072 336.5193
CAU 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 229.5211 276.5790 287.8519
FLE 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0030 330.2069 319.8255
POL 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0602 0.0000 0.0000 387.8344
SPN 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0.0329 0.0000 0.0000 0.0000 0.0000
Refer to Table 1 for key to abbreviations.
Pair-wise population comparisons yielding statistically insignificant differences both before (in bold) and after (in italics) applying the Bonferroni correction.
Signals of East Asian gene flow into the Jamaican pop-
ulation most likely stem from the influx of indentured
servants contracted from China between 1852 and 1884
to replace the dwindling African sources (Lai, 1990).
Demographic data reports that the Chinese comprise
0.2% of the total Jamaican populace (Mordecai and
Mordecai, 2001), supporting the minor East Asian com-
ponent detected in the present study. Although less
likely, genetic input from East Asia may have also arisen
from remnants of the indigenous Arawak Indian popula-
tion still present in Jamaica before the British occupa-
tion of the island (Henriques, 1964). Genetic imprints
(SNP and INDEL AIMs) of this group have been
reported by Benn-Torres and collaborators (2008) who
observed minor Native American genetic influences
(3.2%) in the island. In contrast, Haiti exhibits minimal
East Asian contributions (0.3%) only when the Long
(1991) method is employed. Genetic influences from this
region may be attributed to residual Native American
input or to gene flow from East Asian migrants ubiqui-
tous to the New World.
Overall, our results substantiate greater genetic
contributions to Haiti from Africa than to Jamaica,
which shares larger portions of its autosomal compo-
nent with the European and East Asian groups. The
prominent presence of African DNA in the Haitian
and Jamaican populations is likely the result of the
transportation of hundreds of thousands of enslaved
Africans to the colony from ports along Africa’s West
coast. Our data clearly demonstrates that among the
New World African collections, the contributions of
different ancestral groups of populations, namely Afri-
cans, Europeans, and East Asians, varies and corrobo-
rates information from historical accounts of migra-
tory events.
ACKNOWLEDGMENTS
The authors thank Mr. Carlos Velarde for his technical
assistance and Ms. Alicia Cadenas, Ms. Sheyla Mirabal,
Dr. Maria Reguerio, Dr. Diane Rowold, and Ms. Marisil
R. Wright for their scholarly contributions.
LITERATURE CITED
Alves C, Gusma
˜o L, Damasceno A, Soares B, Amorim A. 2004. Con-
tribution for an African autosomic STR database (AmpFlSTR
Identifiler and Powerplex 16 system) and a report on genotypic
variations. Forensic Sci Int 139:201–205.
Alves C, Gusma
˜oL,Lo
´pez-Parra AM, Soledad Mesa M, Amorim
A, Arroyo-Pardo E. 2005. STR allelic frequencies for an African
population sample (Equatorial Guinea) using AmpFlSTR Iden-
tifiler and Powerplex 16 kits. Forensic Sci Int 148:239–242.
Antonarakis SE, Boehm CD, Serjeant GR, Theisen CE, Dover
GJ, Kazazian HH. 1984. Origin of the b
S
-globin gene in
Blacks: the contribution of recurrent mutation or gene conver-
sion or both. Proc Natl Acad Sci 81:853–856.
Applied Biosystems. 2001. AmpFlSTR Identifiler PCR amplifica-
tion kit user’s manual. Foster City, CA: Applied Biosystems.
Ardlie KG, Lunetta KL, Seielstad M. 2002. Testing for popula-
tion subdivision and association in four case-control studies.
Am J Hum Genet 71:304–311.
Athey W. 2006. Haplogroup prediction from Y-STR values using
a Bayesian-Allele-Frequency approach. J Genet Geneal 2:
34–39.
Beleza S, Alves C, Reis F, Amorim A, Carracedo A, Gusma
˜oL.
2004. 17 STR data (AmpFlSTR Identifiler and Powerplex 16
system) from Cabinda (Angola). Forensic Sci Int 141:193–196.
Benn-Torres J, Bonilla C, Robbins CM, Waterman L, Moses TY,
Hernandez W, Santos ER, Bennett F, Aiken W, Tullock T,
Coard K, Hennis A, Wu S, Nemesure B, Leske MC, Freeman
V, Carpten J, Kittles RA. 2008. Admixture and population
stratification in African Caribbean populations. Ann Hum
Genet 72:90–98.
Benn Torres J, Kittles RA, Stone AC. 2007. Mitochondrial and
Y Chromosome diversity in the English-Speaking Caribbean.
Ann Hum Genet 71:1–9.
Bergen AW, Wang CY, Tsai J, Jefferson K, Dey C, Smith KD,
Park SC, Tsai SJ, Goldman D. 1999. An Asian-Native Ameri-
can paternal lineage identified by RPS4Y resequencing and
by microsatellite haplotyping. Ann Hum Genet 63:63–80.
Bernal LP, Borjas L, Zabala W, Portillo MG, Ferna
´ndez E,
Delgado W, Tovar F, Lander N, Chiurillo MA, Ramı
´rez JL,
TABLE 8. Admixture analyses for the New World African populations
Parental groups m
L
m
Y
m
RH
Haiti
African 0.955 60.022
a
0.778177 60.122908
b
1.0021 60.0359
b
East Asian 0.003 60.025
a
20.0758699 60.193892
b
20.0219 60.0347
b
European 0.043 60.027
a
0.297693 60.252631
b
0.0198 60.0527
b
Jamaica
African 0.783 60.020
a
0.764788 60.126145
b
0.8212 60.0657
b
East Asian 0.057 60.022
a
0.0564919 60.179226
b
0.0373 60.0423
b
European 0.160 60.024
a
0.17872 60.239307
b
0.1415 60.0657
b
New Providence (Bahamas)
African 0.780 60.017
a
0.783536 60.105421
b
0.8366 60.0243
b
East Asian 0.000 60.019
a
20.0707197 60.148
b
20.0547 60.0315
b
European 0.220 60.021
a
0.287184 60.186574
b
0.2181 60.0379
b
US African American 1 (Minnesota)
African 0.780 60.019
a
0.697312 60.0705579
b
0.8276 60.0440
b
East Asian 0.004 60.022
a
20.0609425 60.217413
b
20.0215 60.0443
b
European 0.216 60.024
a
0.36363 60.230357
b
0.1939 60.0550
b
US African American 2 (General)
African 0.782 60.016
a
0.711886 60.0651698
b
0.8228 60.0297
b
East Asian 0.030 60.018
a
20.0178861 60.158548
b
0.0101 60.0210
b
European 0.188 60.019
a
0.306 60.188193
b
0.1671 60.0360
b
a
Standard error.
b
Standard deviation.
American Journal of Physical Anthropology
63PHYLOGENETIC RELATIONSHIPS OF NEW WORLD AFRICANS
Garcı
´a O. 2006. Genetic variation of 15 STR autosomal loci in
the Maracaibo population from Venezuela. Forensic Sci Int
161:60–63.
Berniell-Lee G, Calafell F, Bosch E, Heyer E, Sica L,
Mouguiama-Daouda P, van der Veen L, Hombert J, Quintana-
Murci L, Comas D. 2009. Genetic and demographic implica-
tions of the Bantu expansion: insights from human paternal
lineages. Mol Biol Evol 26:1581–1589.
Bertorelle G, Excoffier L. 1998. Inferring admixture proportions
from molecular data. Mol Biol Evol 15:1298–1311.
Bettelheim J. 1979. Jamaican Jonkonnu and related Caribbean
festivals. In: Knight F, Crahan M, editors. Africa and the
Caribbean. Baltimore, MA: The Johns Hopkins University
Press. p 80–100.
Brown KM. 1997. Systematic remembering, systematic forgetting:
Ogou in Haiti. In Barnes ST, editor. Africa’s Ogun: Old world
and new. Bloomington, IN: Indiana University Press. p 65–89.
Budowle B, Moretti T, Baumstark AL, Defenbaugh DA, Keys
KM. 1999. Population data on the thirteen CODIS core short
tandem repeat loci in African Americans. U.S. Caucasians,
Hispanics, Bahamians, Jamaicans, and Trinidadians. J Foren-
sic Sci 44:1277–1286.
Butler JM, Buel E, Crivellente F, McCord BR. 2004. Forensic
DNA typing by capillary electrophoresis using the ABI Prism
310 and 3100 genetic analyzers for STR analysis. Electropho-
resis 25:1397–1412.
Butler JM, Schoske R, Vallone PM, Redman JW, Kline MC.
2003. Allele frequencies for 15 autosomal STR loci on U.S.
Caucasian, African American, and Hispanic populations.
J Forensic Sci 48:908–911.
Camacho MV, Benito C, Figueiras AM. 2007. Allelic frequencies
of the 15 STR loci included in the AmpFlSTR
1
Identifiler
TM
PCR Amplification kit in an autochthonous sample from
Spain. Forensic Sci Int 173:241–245.
Carmody G. 1990. G-test. Ottowa, Canada: Carleton University.
Chiurillo MA, Morales A, Mendes AM, Lander N, Tovar F,
Fuentes A, Ramı
´rez JL. 2003. Genetic profiling of a central
Venezuelan population using 15 STR markers that may be of
forensic importance. Forensic Sci Int 136:99–101.
Coudray C, Calderon R, Guitard E, Ambrosio B, Gonza
´lez-
Martı
´n A, Dugoujon J, Allele frequencies of 15 tetrameric
short tandem repeats (STRs) in Andalusians from Huelva
(Spain). Forensic Sci Int 168:e21–e24, 2007.
Coupeau S. 2008. The history of Haiti. Wesport, CT: Greenwood
Press.
Curtin PD. 1975. Economic change in pre-colonial Africa:
Sengambia in the era of the slave trade. Madison, WI: The
University of Wisconsin Press.
Curtin PD. 1955. Two Jamaicas: the role of ideas in a tropical
colony 1830–1865. Cambridge, MA: Harvard University Press.
Czarny J, Grzybowski T, Derenko MV, Malyarchuk BA, S
´liwka
DM. 2005. Genetic variation of 15 STR loci (D3S1358, vWA.
FGA, TH01, TPOX, CSF1PO, D5S818, D13S317, D75820,
D16S539, D2S1338, D8S1179, D21S11, D18S51, and
D19S433) in populations of north and central Poland. Foren-
sic Sci Int 147:97–100.
de Heusch L. 1995. Kongo in Haiti: a new approach to religious
syncretism. In: Davis DJ, editor. Slavery and beyond: the
African impact on Latin America and the Caribbean. Lanham,
MD: SR Books. p 103–120.
de Souza Go
´es AC, da Silva DA, Fonseca Gil EH, da Silva MTD,
Pereira RW, de Carvalho EF. 2004. Allele frequencies data
and statistic parameters for 16 STR loci - D19S433, D2S1338,
CSF1PO, D16S539, D7S820, D21S11, D18S51, D13S317,
D5S818, FGA, Penta E, TH01, vWA, D8S1179, TPOX,
D3S1358 in the Rio de Janeiro population, Brazil. Forensic
Sci Int 140:131–132.
De Ungria MCA, Roby RK, Tabbada KA, Rao-Coticone S, Tan
MMM, Hernandez KN. 2005. Allele frequencies of 19 STR loci
in a Philippine population generated using AmpFlSTR multi-
plex and ALF singleplex systems. Forensic Sci Int 152:281–284.
´az V, Rivas P, Olivero C, Carracedo A. 2008. The distribution
of allele frequencies of 15 STRs in Dominican population.
Forensic Sci Int: Genet Supplement Series 1:322–325.
Felsenstein J. 2002. Phylogeny inference package (PHYLIP),
Version 3.6a3. Seattle: Department of Genetics, University of
Washington.
Ferna
´ndez-Cobo M, Jobes DV, Yanagihara R, Nerurkar VR,
Yamamura Y, Ryschkewitsch CF, Stoner GL. 2001.
Reconstructing population history using JC virus: Amerinds,
Spanish, and Africans in the ancestry of modern Puerto
Ricans. Hum Biol 73:385–402.
Ferrer A, Naza
´bal M, Companioni O, Ferna
´ndez de Cossı
´o ME,
Camacho H, Cintado A, Benı
´tez J, Casalvilla R, Sautie
´M,
Villareal A,
´az T, Marrero A, Ferna
´ndez de Cossı
´oJ,
Hodelin A, Leal L, Ballester L, Novoa LI, Middleton D,
Duen
˜as M. 2007. HLA class I polymorphism in the Cuban
population. Hum Immunol 68:918–927.
Forward BW, Eastman MW, Nyambo TB, Ballard RE. 2008.
AmpFlSTR
1
Identifiler
TM
STR allele frequencies in Tanzania,
Africa. J Forensic Sci 53:245–247.
Gibbs WN, Ottey F, Dyer H. 1972. Distribution of Glucose-6-
Phosphate Dehydrogenase phenotypes in Jamaica. Am J Hum
Genet 24:18–23.
Gorostiza A, Gonza
´lez-Martı
´n A, Ramı
´rez CL, Sa
´nchez C,
Barrot C, Ortega M, Huguet E, Corbella J, Gene
´M. 2007. Al-
lele frequencies of the 15 AmpFlSTR Identifiler loci in the
population of Metztitla
´n (Estado de Hidalgo), Me
´xico. Foren-
sic Sci Int 166:230–232.
Grann VR, Ziv E, Joseph CK, Neugut AI, Wei Y, Jacobson JS,
Horwitz MS, Bowman N, Beckmann K, Hershman DL. 2008.
Duffy (Fy), DARC, and neutropenia among women from the
United States, Europe and the Caribbean. Brit J Haematol
143:288–293.
Gross AM, Budowle B. 2006. Minnesota population data on 15 STR
loci using the Identifiler
1
kit. J Forensic Sci 51:1410–1413.
Gusma
˜oL,Sa
´nchez-Diz P, Gomes I, Alves C, Carracedo A
´,
Prata MJ, Amorim A. 2006. Genetic analysis of autosomal
and Y-specific STRs in the Karimojong population from
Uganda. Int Congr Ser 1288:213–215.
Hashiyada M, Itakura Y, Nagashima T, Nata M, Funayama M.
2003. Polymorphism of 17 STRs by multiplex analysis in
Japanese population. Forensic Sci Int 133:250–253.
Heinl RD, Heinl NG. 2005. Written in blood: the story of the
Haitian people 1492–1995. Lanham, MD: University Press of
America.
Henriques F. 1964. Jamaica: land of wood and water. New York,
NY: London House & Maxwell.
Herna
´ndez-Gutie
´rrez S, Herna
´ndez-Franco P, Martı
´nez-Tripp S,
Ramos-Kuri M, Rangel-Villalobos H. 2005. STR data for 15
loci in a population sample from the central region of Mexico.
Forensic Sci Int 151:97–100.
Herrera RJ, Adrien LR, Ruiz LM, Sanabria NY, Duncan G.
2004. D1S80 single-locus discrimination among African popu-
lations. Hum Biol 76:87–108.
Hidalgo PC, Bengochea M, Abilleira D, Cabrera A, Alvarez I.
2005. Genetic admixture estimate in the Uruguayan popula-
tion based on the loci LDLR. GYPA, HBGG, GC and D7S8.
Int J Hum Genet 5:217–222.
Hu SP, Yu XJ, Liu JW, Cai KL. 2005. Analysis of STR polymor-
phisms in the Choa Shan population in South China. Forensic
Sci Int 147:93–95.
Hurles ME, Sykes BC, Jobling MA, Forster P. 2005. The dual
origin of the Malagasy in island Southeast Asia and East
Africa: evidence from maternal and paternal lineages. Am J
Hum Genet 76:894–901.
Keegan WF. 1995. Modeling dispersal in the prehistoric West
Indies. World Archaeol 26:400–420.
Kido A, Dobashi Y, Fujitani N, Hara M, Susukida R, Kimura H,
Oya M. 2007. Population data on the AmpFlSTR Identifiler
loci in Africans and Europeans from South Africa. Forensic
Sci Int 168:232–235.
Kim Y, Hwang J, Kim Y, Lee S, Chung N, Goh H, Kim C, Kim
D. 2003. Allele frequencies of 15 STR loci using AmpFlSTR
Identifiler kit in a Korean population. Forensic Sci Int
136:92–95.
Knight F. 2000. The Haitian Revolution. Am Hist Rev 105:103–
115.
American Journal of Physical Anthropology
64 T.M. SIMMS ET AL.
Knight F, Crahan M. 1979. The African migration and the ori-
gins of an Afro-American society and culture. In: Knight F,
Crahan M, editors. Africa and the Caribbean. Baltimore, MA:
The Johns Hopkins University Press. p 1–19.
Lai L. 1990. Chinese indentured labor migrations to the British
West Indies in the nineteenth century. In Sanjek R, editor.
Caribbean Asians: Chinese, Indian and Japanese experiences
in Trinidad and the Dominican Republic. New York, NY:
Queens College. p 3–30.
Lai L. 1998. The Chinese in the West Indies 1806–1995: a docu-
mentary history. Kingston, Jamaica: The Press University of
the West Indies.
Listman JB, Malison RT, Sughondhabirom A, Yang B, Raaum
RL, Thavichachart N, Sanichwankul K, Kranzler HR, Tang-
wonchai S, Mutirangura A, Disotell TR, Gelernter J. 2007.
Demographic changes and marker properties affect detection
of human population differentiation. BMC Genet 8:21.
Long JC, Williams RC, McAuley JE, Meids R, Partel R,
Tregellas M, South SF, Rea AE, McCormick B, Iwaniec U.
1991. Genetic variation in Arizona Mexican Americans: esti-
mation and interpretation of admixture proportions. Am J
Phys Anthropol 84:141–157.
Martı
´nez-Cruzado JC, Toro-Labrador G, Viera-Vera J, Rivera-
Vega MY, Startek J, Latorre-Esteves M, Roma
´n-Colo
´nA,
Rivera-Torres R, Navarro-Milla
´nIY,Go
´mez-Sa
´nchez E,
Caro-Gonza
´lez HY, Valencia-Rivera P. 2005. Reconstructing
the population history of Puerto Rico by means of mtDNA
phylogeographic analysis. Am J Phys Anthropol 128:131–155.
Mason P. 2000. Jamaica: A guide to the people, politics and cul-
ture. Brooklyn, NY: Interlink Books.
McLean DC, Spruill I, Argyropoulos G, Page GP, Shriver MD,
Garvey WT. 2005. Mitochondrial DNA (mtDNA) haplotypes
reveal maternal population genetic affinities of Sea Island
Gullah-Speaking African Americans. Am J Phys Anthropol
127:427–438.
McLean DC, Spruill I, Gevao S, Morrison EYSA, Bernard OS,
Argyropoulos G, Garvey WT. 2003. Three novel mtDNA
restriction site polymorphisms allow exploration of population
affinities of African Americans. Hum Biol 75:147–161.
Mendizabal I, Sandoval K, Berniell-Lee G, Calafell F, Salas A,
Martı
´nez-Fuentes A, Comas D. 2008. Genetic origin, admix-
ture, and asymmetry in maternal and paternal human line-
ages in Cuba. BMC Genet 8:213.
Mertens G, Mommers N, Cardoen E, De Bruyn I, Jehaes E,
Rand S, Van Brussel K, Jacobs W. 2006. Flemish population
genetic analysis using 15 STRs of the Identifiler
1
kit. Int
Congr Ser 1288:328–330.
Miljkovic-Gacic I, Ferrell RE, Patrick AL, Kammerer CM,
Bunker CH. 2005. Estimates of African, European and Native
American Ancestry in Afro-Caribbean men on the island of
Tobago. Hum Hered 60:129–133.
Mordecai M, Mordecai P. 2001. Culture and customs of Jamaica.
Westport, CT: Greenwood Press.
Muro T, Fujihara J, Imamura S, Nakamura H, Yasuda T,
Takeshita H. 2008. Allele frequencies for 15 STR loci in
Ovambo population using AmpFlSTR
1
Identifiler kit. Leg
Med 10:157–159.
Nagel RL. 1984. The origin of the hemoglobin S gene: clinical,
genetic and anthropological consequences. Einstein Q J Biol
Med 2:53–62.
Okediji MB. 2003. The shattered gourd: Yoruba forms in twenti-
eth-century American art. Seattle, WA: The University of
Washington Press.
Ota T. 1993. DISPAN: genetic distance and phylogenetic analy-
sis. University Park, PA: Institute of Molecular Evolutionary
Genetics, Pennsylvania State University.
Ott T. 1973. The Haitian Revolution 1789–1804. Knoxville, TN:
The University of Tennessee Press.
Pamphile LD. 2001. Haitians and African Americans. Gainse-
ville, FL: University Press of Florida.
Pante-de-Sousa G, Mousinho-Ribeiro RC, Santos EJM, Zago
MA, Guerreiro JF. 1998. Origin of the Hemoglobin S gene in
a northern Brazilian population: the combined effects of slave
trade and internal migrations. Genet Mol Biol 21:427–430.
Parra EJ, Marcini A, Akey J, Martinson J, Batzer MA, Cooper
R, Forrester T, Allison DB, Deka R, Ferrell RE, Shriver MD.
1998. Estimating African American admixture proportions by
use of population-specific alleles. Am J Hum Genet 63:1839–
1851.
Pepin J. 2005. From the Old World to the New World: an eco-
logic study of population susceptibility to HIV infection. Trop
Med Int Health 10:627–639.
Pereira JV, Serra A, Anjos MJ, Vide MC, Corte-Real F, Vieira
DN. 2005. Allele frequencies for fifteen STR loci in Guinea-
Bissau population. J Forensic Sci 50:243–244.
Pereira L, Macaulay V, Torroni A, Scozzari R, Prata MJ,
Amorim A. 2001. Prehistoric and historic traces in the
mtDNA of Mozambique: insights into the Bantu expansions
and the slave trade. Am J Hum Genet 65:439–458.
Pritchard JK, Stephens M, Donnelly P. 2000. Inference of popu-
lation structure using multilocus genotype data. Genetics
155:945–959.
Raymond M, Rousset F. 1995. GENEPOP (version 1.2): a popu-
lation genetics software for exact test and ecumenicism.
J Hered 86:248–249.
Rebala K, Wysocka J, Kapin
´ska E, Cybulska L, Mikulich AI,
Tsybovsky IS, Szczerkowska Z. 2007. Belarusian population
genetic database for 15 autosomal STR loci. Forensic Sci Int
173:235–237.
Regueiro M, Carril JC, Pontes ML, Pinheiro MF, Luis JR,
Caeiro B. 2004. Allele distribution of 15 PCR-based loci in the
Rwanda Tutsi population by multiplex amplification and cap-
illary electrophoresis. Forensic Sci Int 143:61–63.
Regueiro M, Mirabal S, Lacau H, Caeiro JL, Garcia-Bertrand
RL, Herrera RJ. 2008. Austronesian genetic signature in
East African Madagascar and Polynesia. J Hum Genet
53:106–120.
Roberts DF, Hiorns RW. 1965. Methods of analysis of the genetic
composition of a hybrid population. Hum Biol 37:38–43.
Roberts GW. 1954. Immigration of Africans into the British
Caribbean. Population Studies 7:235–262.
Roberts K. 2004. The influential Yoruba past in Haiti. In:
Falolola T, Childs MD, editors. The Yoruba diaspora in the
Atlantic world. Bloomington, IN: Indiana University Press.
p 177–182.
Rodrı
´guez A, Arrieta G, Sano
´u I, Vargas MC, Garcı
´aO,
Yurrebaso I, Pe
´rez JA, Villalta M, Espinoza M. 2007. Popula-
tion genetic data for 18 STR loci in Costa Rica. Forensic Sci
Int 168:85–88.
Rogozin
´ski J. 1999. A brief history of the Caribbean. New York,
NY: Plume.
Salas A, Richards M, De la Fe T, Lareu M, Sobrino B,
Sa
´nchez-Diz P, Macaulay V, Carracedo A
´. 2002. The making
of the African mtDNA landscape. Am J Hum Genet
71:1082–1111.
Saunders G. 2003. Cultural Perspectives. In: Bahamas Depart-
ment of Archives Bahamas Heritage Festival. Nassau, Baha-
mas: Government Printing Department. p 1–13.
Schneider S, Kueffer JM, Roessli D, Excoffler L. 2000. Arlequin v.
2.000: a software for population genetics data analysis. Geneva:
Genetics and Biometry Laboratory, University of Geneva.
Schuler M. 1979. Myalism and the African religious tradition in
Jamaica. In: Knight F, Crahan M, editors. Africa and the
Caribbean. Baltimore, MA: The Johns Hopkins University
Press. p 65–79.
Seah LH, Jeevan NH, Othman MI, Jaya P, Ooi YS, Wong PC,
Kee SS. 2003. STR data for the AmpFlSTR Identifiler loci in
three ethnic groups (Malay. Chinese, Indian) of the Malaysian
population. Forensic Sci Int 138:134–137.
Shepard EM, Herrera RJ. 2006. Genetic encapsulation among
Near Eastern populations. J Hum Genet 51:467–476.
Simms TM, Garcia C, Mirabal S, McCartney Q, Herrera RJ.
2008. The genetic legacy of the Transatlantic Slave Trade in
the island of New Providence. Forensic Sci Int: Genet 2:310–
317.
SPSS for Windows, Rel. 11.0.1. 2006. Chicago: SPSS Inc.
Thomas H. 1998. The slave trade: the history of the Atlantic
slave trade 1440–1870. London: Macmillan Publishers Ltd.
American Journal of Physical Anthropology
65PHYLOGENETIC RELATIONSHIPS OF NEW WORLD AFRICANS
Thomas-Hope E. 2003. Irregular Migration and Asylum Seekers
in the Caribbean. United Nations University, WIDER. Discus-
sion Paper No. 2003/48.
Treco RN. 2002. The Haitian diaspora in the Bahamas. Miami,
FL: Florida International University.
Tselos SE. 1996. Threads of reflection: costumes of Haitian
Rara. Afr Arts Special Issue Arts of Vodou 29:58–102.
Vergara C, Caraballo L, Mercado D, Jimenez S, Rojas W,
Rafaels N, Hand T, Campbell M, Tsai YJ, Gao L, Duque C,
Lopez S, Bedoya G, Ruiz-Linares A, Barnes KC. 2009. African
ancestry is associated with risk of asthma and high total se-
rum IgE in a population from the Caribbean Coast of Colom-
bia. Hum Genet 125:565–579.
Wainscoat JS, Bell JI, Thein SL, Higgs DR, Serjeant GR, Peto
TEA, Weatherall DJ. 1983. Multiple origins of the sickle
mutation: evidence from bS gene cluster polymorphism. Mol
Biol Med 1:191–197.
Wang J. 2003. Maximum-likelihood estimation of admixture
proportions from genetic data. Genetics 164:747–765.
Wang Z, Yu R, Wang F, Li X, Jin T. 2005. Genetic polymor-
phisms of 15 STR loci in Han population from Shaanxi (NW
China). Forensic Sci Int 147:89–91.
Watts D. 1987. The West Indies patterns of development, cul-
tural and environmental change since 1492. New York, NY:
Columbia University Press.
Zu
´n
˜iga J, Ilzarbe M, Acunha-Alonzo V, Rosetti F, Herbet Z,
Romero V, Almeciga I, Clavijo O, Stern JNH, Granados J, Frid-
kis-Hareli M, Morrison P, Azocar J, Yunis EJ. 2006. Allele fre-
quencies for 15 autosomal STR loci and admixture estimates in
Puerto Rican Americans. Forensic Sci Int 164:266–270.
American Journal of Physical Anthropology
66 T.M. SIMMS ET AL.
... Initially relying upon classical genetic markers such as proteins found in blood and then eventually actual DNA sequences, researchers learned that genetic variation in contemporary populations reflected both distant and more recent events in human history (Jobling, Hurles, and Tyler-Smith 2004, 51). More recently, scholars are also increasingly using genetic research to interrogate biogeographic ancestry 4 as it relates to questions of ethnogenesis, ethno-national identity formation, and creolization of peoples in Africa and the African diaspora, including Latin America and the Caribbean (Benn Torres et al. 2008;Benn Torres, Stone, and Kittles 2013;Deason et al. 2012;Fendt 2012;Gaieski et al. 2011;Gomez, Hirbo, and Tishkoff 2014;Mendizabal et al. 2008;Price et al. 2017;Ruiz-Linares et al. 2014;Simms 2010;Stefflova et al. 2011;Tishkoff et al. 2009). ...
... We acknowledge, however, that given the small sample size of the Accompong Town community and corresponding standard errors of the admixture estimates, our assertions based on this particular analysis warrant additional sampling and testing. Nonetheless, the European admixture in the Accompong Town Maroon sample is comparable to that observed in the general Jamaican populace, in which estimates range from 10 to 16%, according to several previous studies (Benn Torres, Stone, and Kittles 2013;Benn-Torres et al. 2008;Simms et al. 2010). The similar ancestry between Accompong Town Maroons and the general Jamaican populace reflects the common West African ancestry between the two groups. ...
... The East Asian admixture among the Accompong sample, however, is higher than the 6% East Asian admixture observed in the general population (Simms et al. 2010). This is especially notable considering that the sample size in the study based on the general populace was over four times as large as the sample size from Accompong: 111 versus 26 participants, respectively. ...
Article
Full-text available
While scholars like Henry Louis Gates, Jr., have popularized the use of genetic data (DNA) as a source of African American history and ancestry, similar studies are lacking for other peoples of African descent. The current study is an interdisciplinary, collaborative research project undertaken by a historian of Africa and the African diaspora in the Americas and a genetic anthropologist. Methodologically, we incorporate traditional sources of historical inquiry, as well as oral history and genetic data (mitochondrial, Y chromosome, and autosomal DNA), to investigate and suggest answers to the following question in popular and scholarly debates about the ethnogenesis of the Jamaican Maroons: to what extent did the indigenous Taínos form part of their early history? Taking the Maroon community of Accompong Town as a case study, we use a novel approach that reconsiders and disrupts mainstream scholarly discourses on Maroon ethnogenesis in Jamaica and, by extension, the larger circum-Caribbean world.
... All countries encompassing LAC present a level of admixture from the three ancestral populations, except for Haiti and Barbados showing only European and sub-Saharan African admixture. Of note, ancestry feasibly reflects the demographic history of LAC: Caribbean islands present a higher proportion of sub-Saharan African ancestry, as demonstrated by studies from Bahamas, Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Dominica, St. Lucia, Barbados, St. Vincent, Grenada, and Trinidad and Tobago (Benn- Torres et al., 2008;Bryc et al., 2010;Diaz-Horta et al., 2010;Simms et al., 2010;Torres et al., 2013;Marcheco-Teruel et al., 2014;Mathias et al., 2016;Martin et al., 2017;Fortes-Lima et al., 2018); Central America, Andean regions, and Ecuador have populations with high Native American ancestry (Arias et al., 2002;Wang et al., 2008;Nuñez et al., 2010;Ruiz-Narváez et al., 2010;Moura et al., 2015;Castro-Pérez et al., 2016;Mathias et al., 2016;Martin et al., 2017); and Brazil, Paraguay, Uruguay, Argentina, and Chile present high proportions of European ancestry (Ruiz-Linares et al., Homburger et al., 2015;Moura et al., 2015). Although based on mtDNA and Y-chromosome length polymorphisms, Native American subpopulations from French Guiana also present a proportion of Sub-Saharan African ancestry, which derived from historical contact with a group of escaped slaves who lives in the region nowadays (Mazières et al., 2007). ...
Article
Full-text available
Genomics can reveal essential features about the demographic evolution of a population that may not be apparent from historical elements. In recent years, there has been a significant increase in the number of studies applying genomic epidemiological approaches to understand the genetic structure and diversity of human populations in the context of demographic history and for implementing precision medicine. These efforts have traditionally been applied predominantly to populations of European origin. More recently, initiatives in the United States and Africa are including more diverse populations, establishing new horizons for research in human populations with African and/or Native ancestries. Still, even in the most recent projects, the under-representation of genomic data from Latin America and the Caribbean (LAC) is remarkable. In addition, because the region presents the most recent global miscegenation, genomics data from LAC may add relevant information to understand population admixture better. Admixture in LAC started during the colonial period, in the 15th century, with intense miscegenation between European settlers, mainly from Portugal and Spain, with local indigenous and sub-Saharan Africans brought through the slave trade. Since, there are descendants of formerly enslaved and Native American populations in the LAC territory; they are considered vulnerable populations because of their history and current living conditions. In this context, studying LAC Native American and African descendant populations is important for several reasons. First, studying human populations from different origins makes it possible to understand the diversity of the human genome better. Second, it also has an immediate application to these populations, such as empowering communities with the knowledge of their ancestral origins. Furthermore, because knowledge of the population genomic structure is an essential requirement for implementing genomic medicine and precision health practices, population genomics studies may ensure that these communities have access to genomic information for risk assessment, prevention, and the delivery of optimized treatment; thus, helping to reduce inequalities in the Western Hemisphere. Hoping to set the stage for future studies, we review different aspects related to genetic and genomic research in vulnerable populations from LAC countries.
... Several mtDNA lineages from Ceramic Age populations did not persist into the present-day, and Caribbean islanders vary in the extent of autosomal ancestry they can trace to precontact populations. [6][7][8]11,42,43,74 ing on one or more island communities and often seeking to characterize the impact of local sociocultural and historical processes on biocultural diversity, 9,10,[38][39][40]43,45,49,50,60,65,75,77,[81][82][83][84][85][86][87][88] (2) Regional histories surveying broad patterns of genetic diversity across several islands which are usually grouped together due to some unifying geographic, historical or linguistic factor (e.g., the Lesser Antilles, the former French colonies, the Anglophone islands), 11,[46][47][48]74,76,89 (3) [10][11][12] and, more recently, whole sequencing of both historic and present-day human genomes. 5,42 Overall, this large body of work has found that most Caribbean islanders can trace their genetic ancestries to two or more continental populations. ...
Article
Full-text available
As the last American region settled by humans, yet the first to experience European colonization, the Caribbean islands have a complex history characterized by continuous migration, admixture, and demographic change. In the last 20 years, genetics research has transformed our understanding of Caribbean population history and revisited major debates in Caribbean anthropology, such as those surrounding the first peopling of the Antilles and the relationship between ancient Indigenous communities and present‐day islanders. Genetics studies have also contributed novel perspectives for understanding pivotal events in Caribbean post‐contact history such as European colonization, the Atlantic Slave Trade, and the Asian Indenture system. Here, I discuss the last 20 years of Caribbean genetics research and emphasize the importance of integrating genetics with interdisciplinary historic, archaeological, and anthropological approaches. Such interdisciplinary research is essential for investigating the dynamic history of the Caribbean and characterizing its impact on the biocultural diversity of present‐day Caribbean peoples.
... Furthermore, other studies examining the genetic structure of populations from Haiti observed that Haitians exhibited minimal contributions from non-African populations in their gene pool. This suggests that Haitians may reflect similar prevalences of hemoglobinopathies as do populations from Sub-Saharan Africa, which are highly prevalent [64]. Finally, a report from the Ouest department in Haiti observed a prevalence of approximately 20% of glucose 6 phosphate dehydrogenase deficiency in a population of school-aged children further suggesting that hemoglobinopathies may be prevalent in this population [65]. ...
Article
Full-text available
Background In Haiti, differences in the prevalence of anemia between urban and rural areas have been observed.Objective To identify moderating factors that may help explain the difference in the prevalence of anemia in children from poor urban vs. rural areas of Haiti.Methods This cross-sectional study used secondary data from urban and rural school-based trials that assessed the effectiveness of a nutrition intervention. The study was registered at ClinicalTrials.gov as NCT02747524. A total of 300 rural- and 981 urban- children between 2.5-13 years of age were included in this analysis. Effect modification in a binary logistic generalized linear mixed model was conducted using sample weights in SPSS® version 26. Models were adjusted for age and income. School cluster was included as random effect.ResultsIn rural areas, stunting was more prevalent in children with anemia vs. no anemia, (16.6%, and 6.3%, P = 0.008), respectively. Also, rural children with anemia lived with fewer adults vs. rural children with no anemia, ([Formula: see text] = 2.83±1.29, and 3.30±1.54, P = 0.005), respectively. In poor urban areas, helminth morbidities were more frequent in children with anemia vs. no anemia, (21.9% vs. 13.9, P = 0.011), respectively. In the combined sample, stunting, [AOR = 2.05; 95%CI (1.32-3.18)], age [AOR = 0.89; 95%CI (0.85-0.93)], and households with more adults [AOR = 0.77; 95%CI (0.67-0.87)] were associated with anemia. Effect modification by place of residence was observed in households with more adults (t = 3.83, P
... mates varies between island populations(Benn Torres et al., 2008;Deason et al., 2012;Moreno-Estrada et al., 2013;Simms, Rodriguez, Rodriguez, & Herrera, 2010;Vilar et al., 2014). These interisland distinc- tions in ancestry point to differences in historical, demographic, and sociopolitical factors present throughout each island's colonization period and postcolonial migrations ...
Article
Full-text available
Objectives From a genetic perspective, relatively little is known about how mass emigrations of African, European, and Asian peoples beginning in the 16th century affected Indigenous Caribbean populations. Therefore, we explored the impact of serial colonization on the genetic variation of the first Caribbean islanders. Materials and methods Sixty‐four members of St. Vincent's Garifuna Community and 36 members of Trinidad's Santa Rosa First People's Community (FPC) of Arima were characterized for mitochondrial DNA and Y‐chromosome diversity via direct sequencing and targeted SNP and STR genotyping. A subset of 32 Garifuna and 18 FPC participants were genotyped using the GenoChip 2.0 microarray. The resulting data were used to examine genetic diversity, admixture, and sex biased gene flow in the study communities. Results The Garifuna were most genetically comparable to African descendant populations, whereas the FPC were more similar to admixed American groups. Both communities also exhibited moderate frequencies of Indigenous American matrilines and patrilines. Autosomal SNP analysis indicated modest Indigenous American ancestry in these populations, while both showed varying degrees of African, European, South Asian, and East Asian ancestry, with patterns of sex‐biased gene flow differing between the island communities. Discussion These patterns of genetic variation are consistent with historical records of migration, forced, or voluntary, and suggest that different migration events shaped the genetic make‐up of each island community. This genomic study is the highest resolution analysis yet conducted with these communities, and provides a fuller understanding of the complex bio‐histories of Indigenous Caribbean peoples in the Lesser Antilles.
Book
Full-text available
L’objectif principal de cette boîte à outils est de faciliter le diagnostic des nombreuses facettes de l’inégalité sociale qui prévaut en Amérique latine et dans les Caraïbes et de fournir des informations pertinentes sur les politiques sociales mises en œuvre dans différents pays de la région qui se sont avérées efficaces dans la réduction de ces inégalités. Il s’agit également d’apporter des données sur les principales normes internationales propres à faciliter, dans le cadre d’une approche fondée sur les droits, la formulation et la mise en œuvre de politiques sociales susceptibles d’atténuer les inégalités en question. Le diagnostic ainsi posé permet de cerner les défis qui doivent être relevés par les politiques sociales, notamment pour pouvoir progresser dans l’amélioration des conditions de vie des groupes de population restés à la traîne. Cet ensemble d’expériences constitue un point de départ pour explorer et élargir le cadre des possibilités de réponse face aux inégalités.
Book
Full-text available
This toolkit aims to facilitate the assessment of the multiple dimensions of social inequality in Latin America and the Caribbean, and to provide relevant information on social policies implemented in different countries in the region that have effectively reduced inequality. It also supplies information on key international standards that, starting from a rights-based approach, will facilitate the formulation and implementation of social policies that will reduce inequality. The analysis draws attention to the challenges that need to be taken up by social policymakers in order to improve the living conditions of the population groups that have been left the furthest behind. The descriptions of these policies are intended to be a starting point for exploring and expanding upon the array of possible responses to inequality.
Book
Full-text available
El propósito de esta caja de herramientas es facilitar el diagnóstico de las múltiples dimensiones de la desigualdad social presentes en América Latina y el Caribe, y proporcionar información relevante acerca de políticas sociales implementadas en diversos países de la región que han logrado reducir dicha desigualdad. Asimismo, se da a conocer normativa internacional clave que, desde un enfoque de derechos, contribuye a la formulación e implementación de políticas sociales reductoras de las desigualdades. A partir del diagnóstico realizado, se desprenden desafíos que deben ser enfrentados por medio de las políticas sociales y que dicen relación con la posibilidad de avanzar en la mejora de las condiciones de vida de los grupos poblacionales más rezagados. El conjunto de experiencias identificadas representa una base para explorar y expandir el marco de posibilidades de respuesta a la desigualdad.
Preprint
Full-text available
Latin America in general, and in particular the Caribbean islands are a ideal material for population genetics studies, due to the complex factors that influenced the contributions of Amerindians, Europeans, and Africans to the gene pools of different populations. The admixture studies have greatly contributed to understand the historical process in the conformation of the present Antillean populations. The divergent genetic differences present in the Antillean islands indicate to the different migratory process of Europeans, Africans and Native Americans into Caribbean basin. Also in each of the Caribbean islands have been produced microevolutionary, demographic, historical and social processes that particularize each other. América Latina en general, y en particular las islas del Caribe son un material ideal para estudios de genética de poblaciones, debido a los complejos factores que influyeron en las contribuciones de los amerindios, europeos y africanos a los genofondos de diferentes poblaciones. Los estudios de mezcla han contribuido en gran medida a comprender el proceso histórico en la conformación de las actuales poblaciones antillanas. Las diferencias genéticas divergentes presentes en las islas antillanas indican diferentes procesos migratorios de los europeos, africanos y americanos nativos en la cuenca del Caribe. También en cada una de las islas del Caribe se han producido procesos microevolutivos, demográficos, históricos y sociales que particularizan a las mismas entre sí. RESUMO A América Latina em geral, e particularmente as ilhas caribenhas, são um material ideal para estudos de genética de populações devido aos fatores complexos que influenciaram as contribuições de ameríndios, europeus e africanos nos pools gênicos das diferentes populações. Os estudos de miscigenação contribuíram grandemente para o entendimento dos processos históricosna formação das populações atuais das Antilhas. As diferenças genéticas divergentes presentes nas Antilhas são um indicativo dos diferentes processos migratórios de europeus, africanos e nativos americanos no Caribe. Além disso, em cada uma das ilhas caribenhas ocorreram processos microevolutivos, demográficos, históricos e sociais que particularizam cada uma.
Article
We describe a model-based clustering method for using multilocus genotype data to infer population structure and assign individuals to populations. We assume a model in which there are K populations (where K may be unknown), each of which is characterized by a set of allele frequencies at each locus. Individuals in the sample are assigned (probabilistically) to populations, or jointly to two or more populations if their genotypes indicate that they are admixed. Our model does not assume a particular mutation process, and it can be applied to most of the commonly used genetic markers, provided that they are not closely linked. Applications of our method include demonstrating the presence of population structure, assigning individuals to populations, studying hybrid zones, and identifying migrants and admixed individuals. We show that the method can produce highly accurate assignments using modest numbers of loci—e.g., seven microsatellite loci in an example using genotype data from an endangered bird species. The software used for this article is available from http://www.stats.ox.ac.uk/~pritch/home.html.