Caribbean Journal of Science, Vol. 40, No.

3, 291-298, 2004 Copyright 2004 College of Arts and Sciences University of Puerto Rico, Mayaguez

Cranial Evidence of Pre-Contact Multiple Population Expansions in the Caribbean

ANN H. ROSS
North Carolina State University, Department of Sociology and Anthropology, Campus Box 8107, Raleigh, NC 27695-8107 Email: ann_ross@ncsu.edu ABSTRACT.The most recognized Caribbean population dispersal hypothesis is a direct jump from South America followed by dispersal into the Lesser Antilles and westward. This evidence primarily comes from the archaeological record, as skeletal material is scarce in the Caribbean due to generally poor preservation. This study evaluated the direct jump hypothesis along with other possible migration routes using cranial landmark data. A study of three-dimensional facial shape variation among pre-Contact Taino groups from Cuba, Puerto Rico, Jamaica, and Hispaniola, and pre-Contact groups from Mexico, Venezuela, Colombia, and Florida was conducted. Cuban Tainos differed from other Caribbean Taino groups, suggesting a dissimilar ancestry. No significant difference between the Caribbean Taino (excluding Cuba) groups and the South American groups was observed, a result that was consistent with the archaeological record for dispersal from South America into the Lesser Antilles. Cuba was also very distinct from the Florida series, a finding that contradicts hypotheses of possible migrations across the Straits of Florida. The differentiation of the Cuban Tainos from the rest of the American and Caribbean series suggests another source of population influx. KEYWORDS.Caribbean, population migrations, geometric morphometrics

INTRODUCTION The peopling of the New World and expansion into the rest of the Americas, including population movements into the Caribbean, is a topic of fundamental interest to physical anthropologists, geneticists, linguists, and archaeologists. Discussions and studies designed to uncover the mystery of Native American origins have not been without controversy, and have presented various dispersal models such as single or multiple waves of migrations. There has been a renewed interest in the peopling of the Caribbean with recent DNA, dental, and craniometric studies, an area that has traditionally been dominated by archaeology (Coppa et al. 2002; LaluezaFox et al. 2001; Lalueza-Fox et al. 2003; Martinez-Cruzado et al. 2001; Toro-Labrador et al. 2003; Ross et al. 2002). The Caribbean was settled by several waves of migrations of pre-ceramic and ceramic age settlers (Keegan 1995). Several migration routes were proposed for the settlement of the Caribbean, including preceramic colonists crossing the Yucatan passage or dispersing across the Straits of
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Florida into Cuba and then eastward. However, the most recognized Antillean dispersal hypotheses is a direct jump by agriculturalists from South America, followed by dispersal into the Lesser Antilles and westward (Keegan 1995; Moreira de Lima 1999). At the time of contact, two main indigenous groups inhabited the Caribbean islands. The Tainos, inhabited Hispaniola, Puerto Rico, the eastern part of Cuba, Jamaica, the Bahamas, and Turks and Caicos, while the Caribs inhabited the Windward Islands and Guadeloupe (Lalueza-Fox et al. 2003; Rouse 1986, 1992; Petersen 1994). There is evidence that a third group, the aceramic Ciboneys or Guanahatabeys, inhabited the western Greater Antilles including western Cuba, and who are believed to have originated from a preceramic Paleolithic people from Central America (Petersen 1994; Highfield 1994). However, the information available for this group is scarce. The origins of the Taino have been traced to the Orinoco River in Venezuela, and their expansion into the Lesser Antilles to Puerto Rico has been well documented on the basis of a characteristic

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type of pottery known as Saladoid (Rouse 1992; Keegan 1992). The early Saladoid people then gave rise to the Ostionoid around A.D. 600, with these groups eventually giving rise to the Tainos encountered by Columbus (Rouse 1992; Keegan 1992; Callaghan 2001). To evaluate these dispersal hypotheses, a preliminary study of facial shape variation was conducted among pre-contact Caribbean Taino groups and pre-contact groups from North and South America using three-dimensional landmark data. MATERIALS AND METHODS Eight groups totaling 103 individuals were used in this study. Group names, sample sizes, source of data, and proveniences are presented in Table 1. Specific information regarding the exact provenience of some of the samples is limited. Generally, all that is known is the location and date of acquisition of the remains. Males and females were pooled in order to incorporate all of the biological variation within a population. Ten homologous facial landmarks were utilized in the analyses (1. Alare left, 2. Alare right, 3. Dacryon left, 4. Dacryon right, 5. Ectoconchion left, 6. Ectoconchion

right, 7. Nasion, 8. Inferior orbital border left, 9. Superior orbital border left, and 10. Subspinale). Because the Tainos practiced intentional cranial vault reshaping, only facial landmarks were analyzed. The landmarks used in this study are standard craniometric landmarks, and detailed landmark descriptions are found in Howells (1973). A Microscribe 3-DX digitizer was used to obtain the x, y, and z coordinates for each landmark using the software ThreeSkull, written by Stephen D. Ousley. After digitizing the sets of landmark coordinates, it was necessary to scale, translate and rotate each configuration of points so that all skulls would be of comparable size, location and orientation. A Generalized Procrustes Analysis (or GPA) was used to perform these transformations to minimize the sum of squared distances between homologous landmarks on all skulls and scale specimens to a common size (Bookstein 1996; Rohlf and Slice 1990; Slice 1993; Ross et al. 2004). The GPA superimposition was performed using Morpheus et al., a cross-platform program written by Dennis E. Slice and available for downloading from the SUNY-Stony Brook Morphometrics homepage (Slice 1998). A Principal component analysis (PCA) of the covariance matrix was conducted on

TABLE 1. Group names, sample sizes, source of data, and proveniences of the samples used in the present study. Sample name Cuban Taino (Arawak) Hispaniola Taino (Arawak) Puerto Rico Taino (Arawak) Jamaica Taino (Arawak) Colombia Venezuela Mexico Tarasco Florida

N 21 16 9 7 5 4 27 14

Provenience (ca. 800-1500 A.D.) Museo de Montane, Havana, Cuba (ca. 800-1500 A.D.) National Museum of Natural History, Smithsonian Institution, Washington D.C. (ca. 800-1542 A.D.) American Museum of Natural History, New York City (ca. 800-1500 A.D.) National Museum of Natural History, Smithsonian Institution, Washington D.C. Pre-Contact, American Museum of Natural History, New York City Pre-Contact, American Museum of Natural History, New York City Carl Lumholtz collection, American Museum of Natural History, New York City Pre-Contact (1300-1400 A.D.) 3-D coordinates provided by Steve Ousley, National Museum of Natural History, Smithsonian Institution, Washington D.C.

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the GPA transformed coordinates to reduce dimensionality and derive principal component scores for subsequent multivariate analyses. A multivariate analysis of variance (MANOVA) was performed on the first five principal component scores to test for mean shape differences between groups. The degree of differentiation among groups was assessed using Mahalanobis D2 or generalized squared distance of the principal component scores. In addition, an UPGMA Clustering analysis was performed from the generalized distance matrix to characterize relative shape similarities between the groups (Sneath and Sokal 1973). These analyses were performed using the SAS system for Windows Version 8 (SAS n.d.). RESULTS The MANOVA detected significant between group differences (Wilks = 0.257; F value = 4.19; df = 35; p < = 0.0001). The first five principal components accounted for 38%, 18%, 15%, 8%, and 7%, roughly explaining 86% of the total variation. D2 values based on the first five principal component scores are presented in Table 2. The furthest removed populations are Cuba and Florida. Hispaniola, Jamaica, Puerto Rico, Venezuela, and Colombia are not significantly different, and Mexico and Colombia do not differ significantly. Figure 1 shows an anterior view of the superimposition of the mean configuration for all populations. This superimposition or overlay illustrates that Cuba and Florida

are morphologically very different from the rest of the Americans. In Cubans, the alares were more superiorly placed, the inferior orbital border was more laterally and inferiorly placed, the superior orbital border was more superiorly placed, and nasion was more inferiorly placed. By contrast, in Floridians, the alares were oriented more inferiorly, subspinale was more laterally and inferiorly placed, inferior orbital border was more medially placed, and nasion was superiorly oriented. Figure 2 presents the difference between the Mexican and Colombian mean configurations as magnified (X2) difference vectors, which are very similar suggested by the short vectors. The difference vectors show the direction and magnitude of the difference between one mean configuration and another. The difference between the pooled mean forms of three Caribbean Taino groups (Hispaniola + Puerto Rico + Jamaica) and Cuba is illustrated in Figure 3. Here Cubans had more superiorly placed alares, a more antero- inferiorly placed nasion, more medially placed dacryons, a laterally placed inferior orbital border, and infero-medially oriented ectochonchions. The magnitude of the morphological differences between Cuba and Florida and between Mexico and Florida were also pronounced as observed by the length of the difference vectors (Fig. 4 and 5.). In the UPGMA clustering analysis (Fig. 6), two distinct groupings were observed. One cluster included Colombia, Venezuela, and Mexico, while the other included Hispaniola, Jamaica, and Puerto Rico. Notably,

TABLE 2. Generalized squared Mahalanobis distance using principal component scores (D2) showing the degree of differentiation among the groups. Group Colom Cuba FL Hisp Jam Mex PR Venz Colom 0 5.015* 3.620* 1.380 1.773 0.738 1.493 0.244 Cuba 0 11.822 4.024 6.217 5.339 3.775 4.938* FL Hisp Jam Mex PR Venz

0 4.329 2.629* 5.871 5.408 3.184

0 0.365 2.675 0.471 0.680

0 3.547* 1.152 0.911

0 1.620 0.898

0 0.667

All groups significant at p < 0.0001 except those marked *, which are significant at p < 0.05. Figures in boldface indicate no significance.

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FIG. 1. Mean cranial landmark locations after Generalized Procrustes Analysis for Cuba (dark blue spheres), Hispaniola (light blue spheres), Puerto Rico (Aqua spheres), Jamaica (light green spheres), Colombia (dark green spheres), Venezuela (yellow-green spheres), Mexico (yellow spheres), Florida (orange spheres) and the grand mean, the mean configuration for all groups (shown in red with connecting links). 1. Alare left, 2. Alare right, 3. Dacryon left, 4. Dacryon right, 5. Ectoconchion left, 6. Ectoconchion right, 7. Nasion, 8. Inferior orbital border left, 9. Superior orbital border left, and 10. Subspinale.

FIG. 2. Mean cranial landmark differences shown as vectors from Mexico (light gray) to Colombian (dark gray). Vectors magnified X2.

Florida and Cuba branch away from the rest of the American and Caribbean series, suggesting they have a dissimilar origin. DISCUSSION Since craniofacial morphological similarities primarily reflect genetic relationships (Ross et al. 2002; Sparks and Jantz

2002; Ross 2004), the various population expansion hypotheses into the Caribbean can be tested using three-dimensional facial landmark data. To begin with, the morphological similarities between the Mexican and the South American series concurs with earlier craniometric studies by Ross et al. (2002), who found similarities between Mexico and coastal Ecuador that suggested

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FIG. 3. Mean cranial landmark difference shown as vectors from pooled Hispaniola + Jamaica + Puerto Rico (light gray) to Cuba (dark gray). Vectors magnified X2.

FIG. 4. Mean cranial landmark differences shown as vectors from Cuba (light gray) to Florida (dark gray). Vectors magnified X2.

early population expansion. The morphological similarity between the Caribbean Taino (Puerto Rico, Jamaica, and Hispaniola) samples, which reflects similar origins, is not surprising. On the other hand, the dissimilarity between Cuban Tainos and the rest of the Caribbean Tainos, was unexpected, and suggested these groups had distinct origins. As stated above, several migration routes have been proposed for the settlement of the Caribbean, including pre-ceramic colonists crossing the Yucatan passage or dis-

persing across the Straits of Florida from North America into Cuba and then dispersing eastward. However, the most widely recognized Antillean dispersal hypothesis is a direct jump from South America followed by dispersal into the Lesser Antilles and westward (Keegan 1995; Moreira de Lima 1999). The results of this study are partly consistent with the hypothesis of a population expansion originating in South America and dispersing westward demonstrated by the cranial evidence or close biological affinities and lack of significant dif-

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FIG. 5. Mean cranial landmark differences shown as vectors from Mexico (light gray) to Florida (dark gray). Vectors magnified X2.

ferences among Puerto Rico, Jamaica, and Hispaniola Tainos and the South American (Colombia and Venezuela) groups. These findings are supported by genetic data. In a recent mtDNA study, Lalueza-Fox and coworkers (2001) found that pre-Columbian Tainos from Hispaniola had reduced mtDNA diversity and high frequencies of C and D haplogroups, indicating a founder effect during the colonization of the Caribbean and a South American origin concurring with the results of this study. However, a more recent genetic study of preColumbian Ciboneys from Cuba yielded inconclusive results that could not rule out Central America as a possible source (Lalueza-Fox et al. 2003). Similar results were obtained in a craniometric study, which suggested a dissimilar ancestry for Cuban Ciboneys (Ross et al. 2002). Likewise, the present study strongly suggests a different origin for the later Cuban Taino population. The possibility of Florida as a potential source can be discounted as Cuban Tainos differ significantly from the Floridian series. In addition, computer simulations of maritime voyaging patterns indicate that contact between Florida and Cuba would have been sporadic at best (Callaghan 2003). This simulation showed that a success rate from southern Florida was only possible if high paddling speeds were

maintained as lower speeds would have resulted in being swept into the Atlantic (Callaghan 2003). The results of this study, although preliminary, suggest at least two separate migration routes and possible population sources for the peopling of the Caribbean. One likely route is a northwest movement from South America, evidenced by the close affinity of the Caribbean Taino (excluding Cuba) and South American groups. The second possible source is from Central America to account for the Cuban Taino differentiation, which is subject to future and ongoing testing. The results of this study, although based on some small sample sizes, emphasize the need for a systematic and comprehensive sampling of the Americas and Caribbean before conclusions or interpretations can be made with regards to early morphological variation or migration models in the Americas. Acknowledgements.I would like to thank Steve Ousley for making ThreeSkull and the Florida sample available, Dennis Slice for generating the illustrations and for his continued guidance, Antonio MartinezFuentes of the Montane Museum in Cuba for his assistance with the collections, and Lori Baker for assisting in collecting the Cuban data (funded by the Bass Endowment). This project was funded by a Smithsonian

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FIG. 6. Phenogram derived from the UPGMA clustering for all groups showing two distinct nodes. One node is comprised of Colombia, Venezuela, and Mexico, while the other cluster includes Hispaniola, Jamaica, and Puerto Rico. Florida and Cuba branch away notably from the rest of the American and Caribbean series, indicating a dissimilar origin.

Visitor Grant (through Doug Ubelaker) and an American Museum of Natural History Collections Study Grant. LITERATURE CITED
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