Caribbean Journal of Science, Vol. 39, No. 1, 11-22, 2003 Copyright 2003 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez Mitochondrial DNA Analysis in Aruba: Strong Maternal Ancestry of Closely Related Amerindians and Implications for the Peopling of Northwestern Venezuela GLADYS TORO-LABRADOR1,OSWALD R. WEVER2, AND JUAN C. MARTI´NEZ-CRUZADO1,3 1Department of Biology, University of Puerto Rico, P.O. Box 9012, Mayagu¨ez, Puerto Rico 00681-9012 2Department of Internal Medicine, Dr. Horacio Oduber Hospital, Oranjestad, Aruba 3Corresponding author: [email protected] ABSTRACT.—The continental origin of human mtDNA can be identified by its haplogroup determination through restriction fragment length polymorphism (RFLP) analysis. Hair root samples from 16 Aruban informed volunteers were analyzed by this method. Thirteen samples had mtDNAs of Amerindian origin and the remainder had their origin in sub-Saharan Africa; thus suggesting a substantial Amerindian maternal ancestry in Arubans, and helping explain the high incidence of health-related conditions common in Am- erindian populations. Nine of the thirteen Amerindian mtDNAs belonged to haplogroup D, suggesting that despite intense Indian population movements through historical times, most of the mtDNAs shared a recent common origin. Our results, combined with the pre- and post-Columbian history of Aruba and northwestern Venezuela, lead us to hypothesize that the frequency of haplogroup D mtDNAs should be high in the Caquetı´o tribe of coastal Falco´n in Venezuela and at least moderate in the Guajiro Indians of eastern Colombia. In addition, if the native Aruban Indians contributed substantially to the current mtDNA pool of Arubans despite the repeated historical deportation events that presumably removed the complete popula- tions from the island, they must also have possessed haplogroup D mtDNAs in high frequency. Our results do not rule out an origin for the Amerinds of northwestern Venezuela in ancient Amazonian ceramic cultures. Because of its strict maternal inheritance, INTRODUCTION mtDNA studies only measure female in- High-resolution restriction endonuclease put. Thus, the results of such studies in analysis of the mitochondrial DNAs (mtDNAs) populations that have emanated from a of people from various continents, fol- colonization process are frequently not in lowed by phylogenetic reconstruction concordance with those obtained from analysis, has identified many groups of studies based on the Y chromosome, which evolutionarily related haplotypes (hap- displays strict paternal inheritance pat- logroups) that are mostly continent- terns. The results of combining data from specific. Thus, haplogroups can be defined mtDNA and Y chromosome studies typi- by sets of evolutionarily stable restriction cally reveal a predominance of asymmetric motifs that are common to all of their hap- matings between men from colonizing lotypes. Furthermore, the continental ori- populations and women from colonized gin of mtDNA obtained from a mixed ones (Pinto et al., 1996; Batista Dos Santos population can be determined by its hap- et al., 1999; Carvajal-Carmona et al., 2000). logroup identity. Hence, restriction frag- Aruban preceramic archaeological sites ment length polymorphism (RFLP) analy- belong to the Meso-Indian I period (Rouse ses have been used, sometimes in and Cruxent, 1963). The oldest dated site is combination with control region sequences, no more than 4500 years old. The Archaic to estimate the contribution of peoples settlers were apparently fisher-hunter- from different continents to the mtDNA gatherers organized in bands of 10 to 15 pool of mixed populations (Rando et al., people who sustained themselves prima- 1998; Alves-Silva et al., 2000; Green et al., rily on seafood (Versteeg, 1991). Polished 2000). axe-heads made of greenish-gray volcanic 11 12 G. TORO LABRADOR ET AL. material associated with the Meso-Indian large region that included the San Cristo´bal industry of Venezuela, and a sherd of a pot- area of the Andes, south of Lake Mara- tery disc found at the Canashitu preceramic caibo, and coastal areas from Lake Mara- site (500 BC-500 AD), suggest that the Ar- caibo to Cumana´as well as the islands of chaics traded with the ceramic cultures of Aruba, Curac¸ao, and Bonaire. Four subtra- Venezuela (Dahn, 1970). Because of their ditions developed from this expansion (Ol- proximity, we believe that the most likely iver, 1989), of which the Dabajuran subtra- candidates for such trade lived in the Para- dition took over coastal Falco´n, including guana´peninsula, located approximately 30 the Paraguana´peninsula and the islands of km south of Aruba. Aruba, Curac¸ao, and Bonaire (Fig. 1). The The most recent skeletal remains of Aru- Indians occupying this region were known ban Archaics have been found in Malmok as Caquetı´os by the Spaniards, and their (600 AD-900 AD). These remains bear clear language (Caquetı´o) belongs to the Arawa- distinctions when compared with those of kan family of languages (Oliver, 1989). the ceramic culture that arrived approxi- The dominant hypothesis is that the mately at 1000 AD. The Archaics possessed Arawakan family of languages originated high, narrow, and long dolichocephalic approximately 5500 years ago in the junc- skulls, while those from the ceramic culture tion between the Negro and Amazon rivers were low, wide, and short (brachyce- (Rouse, 1986). Proto-Arawakan speakers phalic). Furthermore, all Archaic skulls had migrated north, upstream along the Negro shovel-shaped incisors while such teeth are River, through the Casiquiare Canal, and rare in skulls from the ceramic culture (Ver- downstream along the Orinoco River. steeg, 1993; Tacoma, 1991). Splits occurred at the Meta-Orinoco and the The ceramic culture of Aruba belonged Apure-Orinoco junctions. Some of these an- to the Dabajuran subtradition of the cient people migrated further downstream Dabajuroid tradition. Because of similar el- along the Orinoco River, all the way to the ements in their pottery, the Dabajuroid and coast, and to the Lesser and Greater An- Tierroid traditions of western Venezuela tilles, while others migrated west, up- are believed to share a common ancestral stream along the Meta or the Apure River. culture that may have existed somewhere Those that migrated up the Meta River between the Turbio, Upper Cojedes, and reached the Andes. Some of those migrat- Upper Yaracuy rivers (Oliver, 1995). The ing up the Apure River diverged up the Dabajuroid tradition expanded in the Mar- Cojedes River and eventually reached the acaibo basin after 445 BC; it occupied a coast migrating downstream along the Ya- FIG. 1. Geographic distribution of the Dabajuran subtradition (shaded area). MITOCHONDRIAL DNA ANALYSIS IN ARUBA 13 racuy River. The latter may have given rise the presence of Indians on the islands (Ver- to the Macro-Dabajuroid tradition (Oliver, steeg and Ruiz, 1995). Because of the com- 1989). plexity of the Aruba cave labyrinths, it is The Dabajurans of Aruba developed an possible that the Indians were mostly na- agrarian lifestyle. Thus, whereas prece- tives who had escaped deportation, but ramic sites have large shell middens, sug- they could have been recent migrants from gesting that the Archaics consumed many the mainland. In addition, substantial bivalves and oysters (Versteeg and Ruiz, mainland to Aruba migrations of Indian es- 1995), ceramic sites have comparatively capees occured from 1529 to 1556 during fewer shells (Versteeg and Rostain, 1997) the development of the Venezuelan colony among which those of the queen conch, (Haviser, 1991). Strombus gigas, predominate (Boerstra, Aruba was neglected by the Spaniards 1983). from 1533 until the Dutch conquest of 1636, There is strong evidence that Aruban when Spanish and Amerindian languages Dabajurans maintained close contacts with (especially Caquetı´o) were widely spoken. mainland cultures. For example, close to Upon the Dutch conquest the Spaniards 700 chert flakes were found at the Tanki fled, and all the Indians were deported to Flip ceramic site (Heidecker and Siegel, the mainland because they were regarded 1969), and since no chert source is found in as sympathetic to the Spaniards (Hartog, Aruba, these cherts must have been im- 1961). However, in that same year the ported. In addition, the avemorphic motifs Dutch West Indian Company assigned painted in pottery found in Aruba seem to Aruba the duty of breeding horses and have originated in non-Dabajuroid cul- cattle, and Indians were chosen for these tures. These motifs likely came from the La endeavors because they had a good repu- Guajira peninsula of Colombia, where the tation as wild-horse hunters. Thus, Indian oldest site containing artifacts with these migration to Aruba was stimulated. Also, motifs occurs in the Rancherı´a Valley. Ap- some Indians in war with Spaniards west of parently, only later did the avemorphic Maracaibo fled to Aruba (Hartog, 1961). motifs made it to the Venezuelan The importance of Aruba diminished af- Dabajuran site of Los Me´danos (Oliver, ter the 1648 Dutch Peace Treaty with Spain 1989), suggesting that the Aruban Indians and the island was neglected again. In 1655, traded not only with their Dabajuran kins the Dutch West Indian Company recog- in Venezuela, but with other cultures as nized free Indians in Aruba as trade part- well. ners. These Indians were assigned a piece It is estimated that some 450 to 600 Indi- of land on which to maintain themselves ans lived in Aruba at the time of the Span- through cultivation; they also cut and sold ish discovery (Dijkhoff, 1997) in 1499. To- wood and exploited marine resources (Ver- gether with Curac¸ao and Bonaire, Aruba steeg and Ruiz, 1995). In his description of was declared an island without use in 1513, the Aruban way of life during the second and two years later some 2000 Caquetı´o In- half of the 17th century, A.O. Exquemelin dians from the three islands combined points out that the Indians spoke Spanish, were transported to Hispaniola to work in were Catholic, and were visited frequently mines (Hartog, 1961).