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A Mainly Circum-Mediterranean Origin for West Eurasian and North African mtDNAs in Puerto Rico with Strong Contributions from the Canary Islands and West Africa - Héctor J. Díaz-Zabala 2016
has been attributed to Western European sources (Vilar et al. 2014). However, using Fisher’s Exact tests, median-joining networks, posterior probability analyses, and
principal component analyses,
we found that the origins of the WE-NA female ancestry of modern Puerto Ricans may be varied, with strong contributions from
Spain, the Canary Islands and populations related to contemporary Fulani, Serer and Wolof populations of West Africa. Substantial contributions were also detected from elsewhere in the circum-Mediterranean
Puerto Rico and the Caribbean prior to the Bight of Biafra (Moreno-Estrada et al. 2013). It is noteworthy that
despite its West African origin, U5b1b1b has not been
found among African-Americans (Just et al. 2008), nor in the English-speaking Caribbean (Benn-Torres et al. 2007), Cuba (Mendizabal et al. 2008), ***or Dominican
Republic (unpublished results)***. Thus, it is apparent that the arrival of the U5b1b1b haplotype to Puerto Rico was a rareMaternal lineages of West Eurasian
and North African origin account for 11.5% of total mitochondrial ancestry in Puerto Rico. Historical sources suggest that this
ancestry arrived mostly from European migrations that took place during the four centuries of the Spanish colonization of Puerto Rico. This study analyzed 101
mitochondrial control region sequences and diagnostic coding region variants from a sample set randomly and systematically selected using a census-based sampling
frame to be representative of the Puerto Rican population, with the goal of defining West Eurasian
Puerto Rican mtDNA set within the variation present amongst all reference populations.
Our study shows that up to 38% of West
Eurasian and North African mitochondrial ancestry in Puerto Rico most likely migrated from the Canary Islands. However
Mediterranean region and from West African populations related to the modern Wolof and Serer peoples from Senegal and the nomad Fulani who extend up to Cameroon. In conclusion, the West
Eurasian mitochondrial ancestry in Puerto Ricans is geographically diverse. However, haplotype diversity seems to be low and frequencies have been shaped
by population bottlenecks, migration waves, and random genetic drift. Consequently, approximately 47% of mtDNAs of West Eurasian and North African ancestry in Puerto Rico probably
arrived early in its colonial history.Puerto Ricans are an admixed population composed of three main ancestral subcontinental groups: Sub-Saharan African, European and Native American (Bryc
et al. 2010; Moreno-Estrada et al. 2013). Documentary sources supported by the archaeological record indicate that the
Spanish occupation of Puerto Rico began in
1506, and that by 1513 each of these ancestral populations coexisted on the island. Subsequent migration waves of mostly European and Sub-Saharan African
individuals occurred for at least four centuries after the beginning of colonization (Emmer 1999; Picó 2004). A previous study based on a sample set representative
of the population of Puerto Rico showed that
61.3% of Puerto Rican mitochondrial DNAs (mtDNAs) are of Native American origin, whereas 27.2% are Sub-Saharan
African and 11.5% West Eurasian (Martínez-Cruzado et al. 2005). Subsequent studies have produced consistent results (Vilar et al. 2014). However,
a study on
the autosomal ancestry of the same sample set identified average genome ancestry proportions of around 63.7% European, 21.2% Sub-Saharan African, and 15.2%
Native American (Via et al. 2011). Other studies conducted with Puerto Ricans living in the continental USA have produced similar results (Bryc et al. 2010;
Moreno-Estrada et al. 2013; Gravel et al. 2013). In Puerto Rico, European migrations began with the Spanish colonization
of the early 16th century. Historical sources indicate that the first migrations were fostered by the Spanish Crown’s selective migration policy,
which excluded “Jew”,
“Moor” and “Gypsy” men from settling on the island (Cifre De Loubriel 1964). Later during the 16th century, the Spanish Crown abandoned this policy and agreed
to welcome all migrants (Cifre De Loubriel 1964). Documentary sources indicate that the majority of
emigrants from Spain originated in Aragon, Basque Country,
Valencia, Galicia, Extremadura, Catalonia, Andalusia, Canary Islands, and Castile (Fernández Méndez 1970).The vast majority of Spaniards arriving to Puerto Rico were single men who, by 1506 had begun to intermarry with Native American women. The Governor
reported to the Crown in 1530 that, of a total of 369 “white” men in Puerto Rico, only 57 were married to “white” women (Brau 1904). The Crown took measures to
increase the number of “white” people on the island.
****This included ordering “white”, enslaved Christian women to be sent to Puerto Rico in 1512, ****and instituting a policy
of subsidizing the migration of Spanish families by offering free tools and seeds (Thomas 1997; Díaz-Soler 2000). From this initiative, 50 families consisting of 207
people arrived to Puerto Rico in a single ship in 1520. However, constant conflict with local Native Americans, frequent hurricanes, the little gold found in Puerto
Rico, and the gold rush that developed in Perú at the time stymied the development of a stable settler population (Fernández Méndez 1970).
In 1532, the Spanish crown enacted an order that limited the number of enslaved Sub-Saharan Africans on island farms to five per each “white” peasant.
Hence, peasants were sent to Puerto Rico mostly from the poorest parts of Spain, especially the southern provinces of Andalusia and Extremadura, but also from
Castile and the Canary Islands (Fernández Méndez 1970). In 1695, 20 families numbering 100 members were sent to Puerto Rico from the Canary Islands by
request of the Governor (Brau 1904).
This event marked the beginning of subsequent migration waves from the Canary Islands that lasted for the next two
centuries (Cifre De Loubriel 1964). Between 1720 and 1730, Puerto Rico
s. Northwest Africa was defined including Mauritania (n = 47), Morocco (n = 61), West Sahara (n = 14), Tunisia (n = 46), Moroccan Berbers
(n = 53) and Tunisian Berbers (n = 26).
Because of their high representation of haplogroup U, a group of West African populations was made from Fulani, Wolofs
and Serers (n = 25). Middle East is composed of Dubaians (n = 193), Palestinians (n = 198), Bedouin (n = 73), Druze (n = 120), Nubians (n = 34), Egyptians (n =
Development Core Team 2013). Results WE-NA mtDNA variability in Puerto Rico. We found a total of 34 WE-NA
mtDNA clades in our population sample of 101 unrelated individuals. Samples belonging to haplogroup U, initially identified by the HinfI site at nucleotide
position (np) 12308 when using a mismatched primer (Martínez-Cruzado et al. 2005),
were the most frequent, accounting for 33% of our total sample. Samples
belonging to haplogroup H were the second most frequent contributing around 28%of the total sample, and samples belonging to haplogroup J accounted for 23%.
Thus, whereas haplogroups J and U combined account only for approximately 26.5% of the mtDNAs in the region encompassing Europe and the Caucasus, in Puerto
Rico they account for 56% of all mtDNAs of WE-NA ancestry. More specifically
The most frequent haplotype (n = 10), corresponding to the U5b1b1b clade, is one mutational step away (np 16320) from a node with equal amount of samples from
Crete, Austria and Germany (Figure S2).
Clade U5b1b1b is regarded as specific to West Africa, associated to the Serer, Wolof and Fulani populations from Senegal
and northern Cameroon (Rando et al. 1998; Achilli et al. 2005; Coia et al. 2005; Cerny et al. 2006). The scarcity of HVR-II sequences
Posterior probability analysis.
Our posterior probability (Pos) analysis of all subhaplogroups combined
**does not fit the historical expectation of a major
contribution to the Puerto Rican WE-NA mtDNAs from the Iberian Peninsula** (Table 4, Figure S6). Instead, WE-NA mtDNA sequences in Puerto Rico are in
general
more likely to originate in the Canary Islands with a Pos value of 14.23% (SD=3.48%),
or from the Wolof and Serer from Senegal and the nomad Fulani of
West Africa (Pos = 10.38%, SD = 3.03%). However, the contribution of these West African populations seems to be largely restricted to haplogroup U (Table 4, Figure
S6), suggesting there exists no population in the database which carries a similarly diverse distribution of haplogroups as found in the Puerto Rican WE-NA mtDNAs,
with the possible exception of the Canary Islands. In consistency with this pattern, the most common Puerto Rican WE-NA haplogroups present origin probabilities widely different
7.62%, SD = 5.53%), and North Western Europe (Pos = 7.11%, SD = 5.36%). Haplogroup H, on the other hand shows no region or population with a particularly
high probability of origin (Table 4). Hence, our posterior probability analysis
suggests that WE-NA mtDNAs in Puerto Rico are likely derived from a wide geographical range,
with the Canary Islands as the largest contributor.Table 4. Posterior Probability of Origin (Pos) for Puerto Rican WE-NA
mtDNA) ancestries in the admixed populations (Carvalho-Silva et al. 2001; Bryc et al. 2010). This has been confirmed for Puerto Rico (Vilar et al. 2014), and also
the Canary Islands (Pinto et al. 1994, 1996), where a large proportion of mtDNAs, but not nuclear markers, were found to be of Guanche origin, in addition to those
belonging to other
subjugated populations such as Berbers or Guineans. Thus, the bimodality of the obtained haplotype frequency distribution
interesting.
