I ACLI-Stetson Caribbean Law Program at the Cayman Island Law School THE IMPLICATIONS OF SELECT GLOBAL INITIATIVES ON ECONOMIC DEVELOPMENT IN THE CARl BBEAN Roger M. Groves Associ Professor Florida Coastal School of Law WINTER 2011 / —I Zm -Q m m fl’, . ‘Ii I a -‘‘9 ci. I. I ri. 4nn. Hum. Gene!. (2001), 65, 137—151 137 Printed iv Great Britain MtDNA from extinct Tainos and the peopling of the Caribbean C. LALUEZA-FOX’, F. LUNA CALDERON’, F. CALAFELL’, B MORERA3 AND J. BERTRANPETIT’ Secció Antropologia, Dept. Bioiogia Animal. Facuitat de Biologia, Universitat de Barcelona.. Barcelona. Spain ‘Depa4ta.mento de Antropoiogia Fi’sica, Museo del Hombre Dominicano, Santo Domingo, Replblica Doininicana; Universidad T\Ta,cjo?jai Pedro Henriguez Ureña, Repüblica Dominica.na Unita,t de Biologia. E’voiutiva, Faculta.t de Ciències de la Saint i de la Vida, Universita.t Pompeu Fabra, Barcelona, Spain (Received 10.7.00. Accepted 30.11.00) SUMMARY Tainos and Caribs were the inhabitants of the Caribbean when Columbus reached the Americas; both human groups became extinct soon after contact, decimated by the Spaniards and the diseases they brought. Samples belonging to pre-Columbiari Taino Indians from the La Caleta site (Dominican Republic) have been analyzed, in order to ascertain the genetic affinities of these groups in relation to present-day Amerinds, and to reconstruct, the genetic and demographic events that took place during the peopling of the Caribbean. Twenty-seven bone samples were extracted and analyzed for mtDNA variation. The four major Amerindian mtDNA lineages were screened through amplification of the specific marker regions and restriction enzymatic digestion, when needed. The HVRI of the control region was amplified with four sets of overlapping primers and sequenced in 19 of the samples. Both restriction enzyme and sequencing results suggest that only two (C and D) of the major mtDNA lineages were present in the sample: 18 individuals (75 %) belonged to the C haplogroup, and 6 (25%) to the D haplogroup. Sequences display specific substitutions that are known to correlate with each haplogroup, a fact that helped to reject the possibility of European DNA contamination. A low rate of Taq inisincorporations due to template damage was estimated from the cloning and sequencing of different PCR products of one of the samples. High frequencies of C and D haplogroups are more common in South American populations, a fact that points to that sub-continent as the homeland of the Taino ancestors, as previously suggested by linguistic and archaeological evidence. Sequence and haplogroup data show that the Tainos had a substantially reduced mt.DNA diversity, which is indicative of an important founder effect during the colonization of the Caribbean Islands, assumed to have been a linear migratory movement from mainland South America following the chain configuration of the Antilles. during his first discovery voyage, in 1492, he was INTRODUCTION greeted by indigenous people who called them- When Christopher Colombus reached two of selves Tairios. At that time, Columbus was the Greater Antilles (Bahamas and Hispaniola) convinced of having arrived in either Japan or China; later he changed his mind, and, believing Unitat Correspondence: Jaume Bertranpetit de he had reached India, called the abor Biologia Evolutiva, Facu]tat de Ciencies de Ia Sa]ut i de Ia Vida, Universitat Poinpeu Fabra, C. Dr. Aiguader 80, igines ‘Indians’, a misleading name for the 08003 Barcelona, Spain. Tel: (+3493) 542 28 40: Fax Native Americans that has remained in use to (+3493) 542 28 02. E-mail: jaume. bertranpetit@cexs. upf. es this day. Thus, the wrong and biased perceptions 138 C. LALuEzA-Fox AND OTHERS of Westerners about Caribbean aborigines date were established mainly in Venezuela. The back to the very first moment both cultures Tainos consisted of hierarchical societies collided. However, we don’t really know what organized into chiefdoms; they had advanced the Tainos thought about the Spaniards, since agricultural techniques that allowed them to they were extinguished iii just one or two establish some settlements of thousands of generations after this first contact, decimated by inhabitants, with ceremonial squares and ball the harsh treatment of the Spaniards and the game courts. In contrast, the Caribs were diseases they brought with them. It is difficult to ferocious nomadic. hunters that raided the Taino know how many people were killed during this villages, expanding from the South through the process of extinction; according to different Lesser Antilles. In addition to Tainos and Caribs, authors they could have numbered between 2 there were other groups at Columbus’ times: the and 7 million throughout the Caribbean so-called Arawaks, inhabitants of Trinidad and (Tjbelaker, 1992; Crawford. 1992). At the be the Guianas, and the Guanajuatabeys, inhabi ginning of the 16th century, to replace the tants of West Cuba. decreasing Tainos as agricultural and mining The names of the Caribbean groups and the labour, the Spaniards brought African slaves languages they spoke are a source of debate (Kiple, 1984), who came to constitute the major among scholars; it seems that both Tainos and present-day human substratum in the Caribbean. Island Caribs spoke Arawakan languages that Despite claims of Taino heritage survival in some belong to the Equatorial sub-family, in the rural communities in the east of Cuba, it must be Equatorial-Tucanoan family (Ruhlen, 1991). In concluded that, after 500 years of cultural and contrast, the Mainland Caribs spoke Caribbean genetic disruption, the original Caribbean people languages, which are classified into the Macro have disappeared forever as a distinct human Carib subfamily, within the Ge-Pano-Carib fain group. The study of the so-called Black Caribs ily (Greenberg, 1987; Ruhlen, 1991). The exist from Belize (Monsalve & Hagelberg, 1997), a ence of some words with clear Caribbean origin in population which is presumed to derive from the the language of the Island Caribs poinìts to a close admixture of Island Caribs with West African relationship with the Mainland Caribs. The slaves, illustrates the limitations of working with original homeland of the Taino groups in main the highly admixtured modern Caribbean popu land South America is more controversial. lations, since at least 16 of the 17 sequences Archaeological evidence shows that the found were clearly of African origin. Therefore, Caribbean area was already settled by 5000 B.C.; we need to rely on ancient DNA analysis if we however, it has been suggested that the direct want to know the genetic affinities of these ancestors of the Tainos might have come from groups in relation to the other peoples of the populations that migrated from the Lower Americas. Orinoco Valley, the Guianas or Trinidad and By the time of Columbus, and according to the Tobago, around 1000 B.C. Thereafter, they Spanish chroniclers, there were two main human undertook a long series of voyages, from one groups in the Caribbean, the Tainos, and the island to another, progressing from the mainland Caribs (whose name is the source of the region’s to the Lesser Antilles and from there to the name). The Tainos inhabited Ia Hispaniola, Greater Antilles, eventually mixing with or Puerto Rico, the east of Cuba, and probably pushing west the pre-existing populations, like Jamaica, the Bahamas, and the Turks and Caicos the Guanajuatabeys. The islands are so close to Islands, while the Caribs inhabited the Wind one another that, with three exceptions, it is ward Islands and Guadeloupe (Rouse, 1986, possible to see the next island in the migratory 1993). The latter group — sometimes called Island chain. If this hypothesis is correct. the peopling Caribs — was culturally related to some mainland of the Caribbean had to take place as a linear American groups (called Mainland Caribs), that migratory movement from South East to North mtDI’L4 from extinct Caribbean Indians 139 West, following the chain configuration of the tribe, suggesting a much more complex scenario Antilles Islands. than that expected from the three-migration Therefore, whether or not the Caribbean was model. Subsequent genetic studies (Horai et al. peopled from South-America is a hypothesis that 1993; Torroni et at. 1993a, 1993b) demonstrated can be reliably explored with ancient DNA that the Native American rntDNAs clustered in analysis. The vast majority of ancient DNA few, but relatively deep, lineages that were studies have been based on the analysis of widespread along the continent and not re mitochondrial DNA (mtDNA). This cytoplasmic stricted to any particular ethnic group or genorne has a better chance of recovery, since a linguistic family. The ubiquity of the Native cell with a single copy of the nuclear genome can American mtDNAs in Asia suggested that a contain several thousand copies of the mtDNA single initial migration into America, instead of genolne. MtDNA has been widely used as a successive migration waves, was a more plausible molecular tool for reconstructing the history of scenario (Merriwether et at. 1995; Merriwether & present-day human populations, by virtue of its Ferrell, 1996). From that common rnitochondrial special evolutionary properties, such as a rapid founding pooi, different demographic events mutation rate relative to nuclear DNA, lack of would have produced the differences observed recombination and maternal inheritance (Avise, among present-day Native American popu 1986; Stoneking, 1993). lations, thus complicating the interpretation of In the Americas, many studies have shown both genetic and ethnohistorical data (Forster et that most of the mtDNA of Amerindian PoPu at. 1996). lations falls into four major lineages (named ‘A’, The purpose of this study is to recover mtDNA ‘B’, ‘C’, ‘D’), primarily defined by specific from pre-Columbian Taino remains from rntDNA markers (Schurr et al. 1990; Torroni et Hispaniola (Dominican Republic) to ascertain at. 1992; 1993b, 1994; Horai et at.