AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 133:1060–1066 (2007)

Human Colonization of the Central Territory of Argentina: Design Matrix Models and Craniometric Evidence Mariana Fabra, Andre´s G. Laguens, and Darı´o A. Demarchi* Museo de Antropologı´a, Facultad de Filosofı´a y Humanidades, Universidad Nacional de Co´rdoba, Co´rdoba 5000, Argentina KEY WORDS

South America; Southern cone; peopling; craniometrics; correlation matrix analysis

ABSTRACT In this study we test several hypotheses about the peopling of the central territory of Argentina, archaeologically known as Sierras Centrales, by testing the association between four alternative models of the peopling of the area and cranial morphological variation through distance and matrix correlation analyses. Our results show that the ancient inhabitants of Sierras Centrales demonstrated close morphological

similarities with the Patagonian and Tierra del Fuego populations. Moreover, the correlation and partial correlation analyses suggest that the peopling of the Sierras Centrales most likely took place as a migratory wave proceeding from the present area of Northeastern Argentina, and continued southward to Patagonia and Tierra del Fuego. Am J Phys Anthropol 133:1060–1066, 2007. V 2007 Wiley-Liss, Inc.

The biological variation of the native populations of Argentina and the southernmost region of South America has been extensively studied, through the study of morphological (Cocilovo and Di Rienzo, 1984–1985; Cocilovo and Neves, 1988–1989; Rothammer et al., 1984, 1988– 1989; Lahr, 1995; Lalueza et al., 1996; Herna´ndez et al., 1997; Marcellino and Colantonio, 2000; Gonza´lez-Jose´ et al., 2001a,b, 2003; Fabra et al., 2005), and molecular data (Dipierri et al., 1998, 2000; Demarchi, 2000; Demarchi et al., 2001; Goicoechea et al., 2001; Dejean et al., 2004; Cabana et al., 2006). However, questions about the origins and strategies of dispersion of the colonizer populations remain unresolved and controversial, in part because the extinct populations that once inhabited the central mountains of Argentina, archaeologically known as Sierras Centrales, have received little attention (Ameghino, 1885, 1889; Gonza´lez, 1944; Cocilovo, 1984; Marcellino, 1992; Marcellino and Colantonio, 1993). The first reference to the inhabitants of the Sierras Centrales is found in the writings of Pedro Gonza´lez de Prado (during the expedition of Diego de Rojas in 1545) who named them Comechingones. This author noted that these native people were clearly different from the inhabitants of the surrounding plains. The Comechingones lived in pit-houses agglomerated in small settlements. They were excellent farmers, as well as llama breeders, wore clothes made from llama wool, and adorned their clothes with metal objects. The most striking feature that surprised the Spaniards was the appearance of the Comechingones: they were bearded people, with unusually dark skin and tall stature. Despite these distinct traits, most scholars who studied this group in the first half of the 20th century proposed that this population originated in the Central Andean region (Outes, 1911; Gonza´lez, 1944; Serrano, 1945). On the other hand, other researchers related the Comechingones to the extinct Huarpes, who also lived close to the Andean region, but more to the South. The Huarpes shared with the Comechingones

the high stature, dark skin, and dolichocephalic heads, in contrast to the typical Andean ‘‘morphotype,’’ i.e., shorter in stature and brachycephalic heads (Canals Frau, 1944, 1953). The oldest site dated in the Sierras Centrales is ‘‘Intihuasi,’’ which has been dated at *8,000 years BP, although it is thought that several other sites are even older. The associated culture (similar in certain aspects to Clovis of North America) has been named ‘‘Ayampitin,’’ and can be found in sites across the Sierras Centrales in San Luis and Co´rdoba provinces. The Sierras Centrales have a crucial importance to understanding the population processes of the southernmost region of South America, given its crossroad geographic location, as well as a possible common origin of their ancient inhabitants and the groups that colonized the southern extreme of the Americas, as suggested from both archeological and morphological evidences (Laguens et al., 2003; Fabra et al., 2005). The aim of this study is to test several hypotheses about the peopling of the central territory of Argentina by testing the association between four alternative models of the peopling of the area and cranial morphological variation through distance and matrix correlation analyses.

C 2007 V

WILEY-LISS, INC.

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Grant sponsor: FONCyT; Grant number: PICT 2003–15187; Grant sponsor: SECyT (Universidad Nacional de Co´rdoba). *Correspondence to: Darı´o A. Demarchi, Museo de Antropologı´a, Facultad de Filosofı´a y Humanidades, Universidad Nacional de Co´rdoba, Av. Hipo´lito Yrigoyen 174, Co´rdoba CP5000, Argentina. E-mail: [email protected] Received 10 August 2006; accepted 3 April 2007 DOI 10.1002/ajpa.20634 Published online 25 May 2007 in Wiley InterScience (www.interscience.wiley.com).

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COLONIZATION OF THE CENTRAL TERRITORY OF ARGENTINA TABLE 1. Craniometric series included in this study Population

Code

Region

Sample size

Description in Reference

Sierras Centrales Patagones Patagones Yamana Kaweskar Selknam Valles Calchaquı´es Bele´n San Juan Delta del Parana´

CBA RNE CHU YAM KAW SEL VCA BEL SJU DPA

Central Patagonia Patagonia Tierra del Fuego Tierra del Fuego Tierra del Fuego Northwest Northwest Northwest Northeast

38 62 99 82 25 80 141 22 16 50

Fabra (2005) Lalueza Fox et al. (1996) Lalueza Fox et al. (1996) Lalueza Fox et al. (1996) Lalueza Fox et al. (1996) Lalueza Fox et al. (1996) Constanzo´ (1942a) Paulotti et al. (1949) Constanzo´ (1942b) Torres (1911)

MATERIALS AND METHODS The area of study

population with a mixed economy based on agriculture, hunting, gathering, and fishing.

The Sierras Centrales are located between 30 and 338 (S) latitude and 62–658 (W) longitude, occupying the current territory of Co´rdoba and part of the San Luis provinces. The area is characterized by three mountain chains, the Sierras Grandes, Sierras Chicas, and Sierras Occidentales, that are separated by longitudinal valleys and high-altitude plains, or pampas (Capitanelli, 1979).

Analytical methods

The sample The sample from Sierras Centrales (CBA) includes 38 adult male skulls, preserved in public and private museums of Co´rdoba (Fabra, 2005). As mentioned earlier, the native inhabitants of the Sierras Centrales have been traditionally known as Comechingones, a generic denomination given by the first Spaniards of the 16th century, and popularized after Serrano (1945). Unfortunately, this homogenizing denomination hid the temporal and cultural variation of the region, inhabited at least from 8,000 BP (Gonza´lez, 1960). Archeologically, it is possible to recognize at least two different ways of life in the region: a hunter–gathered subsistence system, from the 8,000 BP until the adoption of agriculture, llama breeding, and sedentarism at pit-houses at around 1,500 BP. Although unfortunately there is no absolute chronology for the skulls analyzed here, most of the samples were recovered from agricultural contexts, therefore they probably correspond to the Late Holocene. In order to carry out interpopulation comparisons, we analyzed data published by several authors, including 577 adult males from nine pre-Hispanic human groups who inhabited at different times several geographic regions of the Southern Cone of South America (Table 1). The Patagonian samples from Rı´o Negro and Chubut (RNE, CHU) correspond to continental terrestrial hunter–gatherers, ethnographically known as Tehuelches. The Selk-Nam (SEL), or Ona, also represent terrestrial hunter–gatherer that inhabited the grasslands of the Isla Grande (Tierra del Fuego). The Kaweskar (KAW) and Yamana (YAM), with a subsistence strategy based on a marine hunter and gathering, inhabited the southern shores of the Tierra del Fuego archipelago. Samples from Salta (VCA), Catamarca (BEL), and San Juan (SJU) provinces belong to sedentary agricultural groups that lived in the northwest portion of the current territory of Argentina, in the Andean region. Finally, the sample from the Parana´ river delta (DPA), in eastern Argentina, represents a

To control variation due to sex differences, only males were included in the analysis. Sex attribution was carried out following standard methods outlined in Buikstra and Ubelaker (1994). Cranial measurements on the CBA series were based on the methods presented by Buikstra and Ubelaker (1994), defined previously by Martin and Saller (1957). The cranial variables used in the analysis were bizygomatic diameter (3.zy-zy), basion-bregma height (4.ba-b), basion-prostion length (6.ba-pr), upper facial height (10.n-pr), minimum frontal breadth (11.ftft), upper facial breadth (12.fmt-fmt), nasal height (13.nns), nasal breadth (14.al-al), orbital breadth (15.d-ec), orbital height (16), interorbital breadth (18.d-d), foramen magnum length (22), and breadth (23) Parenthetical numbers and letters indicate Buikstra and Ubelaker (1994) codification. The pooling of data from different sources and methods to calculate craniometric variables deserves some consideration. While several authors recommended against pooling data from different sources, others suggested low levels of interobserver error, and used series recorded by different sources to study craniometric variation at intrapopulational (Sjovold, 1978; Lalueza Fox et al., 1996), regional (Munford et al., 1995; Kozintsev et al., 1999; Jantz and Owsley, 2001), and global scales (Relethford, 1994; Herna´ndez et al., 1997; Powell and Neves, 1999; Gonza´lez-Jose´ et al., 2001a; Hemphill and Mallory, 2004). The population samples used in this study was recorded by different researchers using different recording methods. For example, Lalueza Fox et al. (1996) used Martin and Saller’s set of traits (Martin and Saller, 1957), while Torres (1911), Constanzo´ (1942a,b), and Paulotti et al. (1949) used early 20th century Monaco’s Convention (Papillaut, 1908). Considering that, we selected for the analysis 13 variables that can be found and homologue in all five sources, with the least number of missing values.

Biological and geographical distances To estimate biological affinities among 10 pre-Hispanic populations from the Southern Cone, we calculated square Euclidean distances based on the mean values for the 13 craniometric traits, listed above. We also computed a geographic distance matrix as linear distances

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in kilometers between pairs of localities, considering the epicenter of the sample distribution. This distance matrix was later used as a co variable to control for influence of geography in the correlation analysis. To further assess the comparative degree of interpopulation genetic variation, from the square Euclidean distance matrix, we constructed a bi-dimensional multidimensional scaling (MDS) plot (Kruskal, 1964). MDS plots permit the evaluation of between-population genetic distances in a visually tractable number of dimensions.

Design matrices and matrix correlation analysis Design matrices are similarity matrices constructed to evaluate hypothesis based on hypotheses of biological relationships between human populations and, as stated by Sokal et al. (1997), they can be used to measure agreement between two or more hypotheses or theories and the observed variation. They describe the relative distance expected between populations under a specific model (Waddle et al., 1998). In a design matrix, each element represents a hypothetical difference. The value given to each pair of localities is chosen taking into consideration the hypothesis suggested in each particular model (Table 2). These design matrices have been applied in numerous studies (Social et al., 1992, 1997; Waddle, 1994; Waddle et al., 1998). Association between design matrices, biological, and geographical distance matrices were analyzed through matrix correlation analysis, which is a powerful tool to evaluate the explanatory power of several hypotheses by measuring their correspondence with observed morphological variation (Gonza´lez-Jose´ et al., 2001a). To evaluate the significance of the correlations between biological and design matrices, we employed the Mantel test (Mantel, 1967). This test assumes that two matrices were obtained independently. It tests the pairwise association between elements in two matrices, computing the running total o the element-by-element or Hadamard product of the two matrices. Significance of the correlation was determined by a permutation test. First, we calculated the correlation coefficient between biological and design matrices. Values of r ranges from 1 (perfect correlation) to
1 (inverse correlation). Values near to 0 indicate no association between both matrices. If there is a positive and significant correlation it can be said that the model corresponds well to observed biological relationships. The Smouse–Long–Sokal test (Smouse et al., 1986) extends Mantel’s statistic to three or more matrices and tests whether an association between two matrices (in this case, biological and design matrices) is significant when one matrix (in this case, geographical matrix) is held constant. All computations were carried out using the program NTSYS 2.11S (Exeter Software).

Archaeological models for the peopling of the Sierras Centrales Four design model matrices were constructed representing hypothetical alternative routes for the human peopling of the current territory of Argentina, in a broader sense, and particularly of the Sierras Centrales region. These models are mostly based on the work of Gonza´lez-Jose´ et al. (2001a) and our own hypotheses about colonization processes (Fig. 1).

TABLE 2. Pairwise hypothetized distance under different peopling models Populations

M1

M2

M3

M4

CBA-RNE CBA-CHU CBA-YAM CBA-KAW CBA-SEL CBA-VCA CBA-BEL CBA-SJU CBA-DPA RNE-CHU RNE-YAM RNE-KAW RNE-SEL RNE-VCA RNE-BEL RNE-SJU RNE-DPA CHU-YAM CHU-KAW CHU-SEL CHU-VCA CHU-BEL CHU-DPA YAM-KAW YAM-SEL YAM-VCA YAM-BEL YAM-SJU YAM-DPA KAW-SEL KAW-VCA KAW-BEL KAW-SJU KAW-DPA SEL-VCA SEL-BEL SEL-SJU SEL-DPA VCA-BEL VCA-SJU VCA-DPA BEL-SJU BEL-DPA SJU-DPA

1 1 0 0 1 0 0 0 1 0 1 1 0 1 1 1 0 1 1 0 1 1 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 0 0 1 0 1 1

0 0 1 1 0 1 1 1 0 0 1 1 0 1 1 1 0 1 1 0 1 1 0 0 1 0 0 0 1 1 0 0 0 1 1 1 1 0 0 0 1 0 1 1

1 1 1 1 1 0 0 0 1 0 0 0 0 1 1 1 0 0 0 0 1 1 0 0 0 1 1 1 0 0 1 1 1 0 1 1 1 0 0 0 1 0 1 1

0 0 0 0 0 1 1 1 0 0 0 0 0 1 1 1 0 0 0 0 1 1 0 0 0 1 1 1 0 0 1 1 1 0 1 1 1 0 0 0 1 0 1 1

Models 1 and 2 propose two separate migration routes for the peopling of the Southern Cone of South America. One route followed the Andean chain, reaching the southernmost extreme of the continent. The populations of the Tierra del Fuego archipelagos (YAM, KAW) were representatives of this migration. The other route came from the Northeast, following the Parana´ and Uruguay riversides and/or the Atlantic Ocean coasts, peopling the plains of Patagonia and Tierra del Fuego. The Selk-Nam were the southernmost representatives of this migration route. Model 1 (M1). The populations that colonized the Sierras Centrales (CBA) derived from the Andean route. All the ‘‘Andean’’ populations should share similar morphological patterns, therefore a distance value of zero was set between CBA and these groups (VCA, BEL, SJU, YAM, KAW). Populations descendant from the second migratory group were assigned a value of 1, to reflect different morphological patterns (DPA, RNE, CHU, SEL). Model 2 (M2). The populations who colonized the Sierras Centrales came from the Northeast. Therefore, CBA

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COLONIZATION OF THE CENTRAL TERRITORY OF ARGENTINA

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Fig. 1. Map of the Southern Cone of South America, showing the location of 10 pre-Hispanic populations included in the analysis and hypothesized routes of peopling with emphasis on the colonization process of the Sierras Centrales.

shares a value of zero with the populations that derived from this route, and a value of 1 with those populations that derived from the northwestern route. Models 3 and 4 propose a different scenario. The Southern Cone was peopled by two different migratory waves. One wave came from the Andes but remained north, in the current territory of northwestern Argentina. The other wave came from the Northeast, following the Parana´ and Uruguay riversides and/or the Atlantic Ocean coasts. This migratory wave continued to the South, peopling Patagonia and Tierra del Fuego. All the Tierra del Fuego populations were descendants of this wave. Model 3 (M3). The population that colonized Sierras Centrales was part of the Northwestern wave and, therefore, CBA share with these populations (SJU, VCA, BEL) a distance value of zero, and a value of 1 with the

remaining southern populations (DPA, RNE, CHU, YAM, KAW, SEL). Model 4 (M4). The population that colonized CBA was part of the Northeastern wave and shares a value of zero with those populations that derived from this wave, and a value of 1 with those populations that derived from the Northwestern wave.

RESULTS The MDS plot based on the square Euclidean distance matrix, representing morphological relationships among the 10 pre-Hispanic series is presented in Figure 2. The stress value achieved was 0.13, which is considered fairly good (Kruskal, 1964), revealing a good correspondence between the plot and the original distance matrix. The three Northwestern populations (VCA, SJU, BEL)

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M. FABRA ET AL. TABLE 3. Correlation and partial correlations, holding geography (GEO) constant, between biological distances and four alternative models of peopling for the Cone South and Sierras Centrales Models

Biological distance

Biological distance (GEO)

M1 M2 M3 M4


0.139 (ns) 0.136 (ns) 0.404* 0.716**


0.143 (ns) 0.173 (ns) 0.195 (ns) 0.664**

ns ¼ not significant. *P < 0.051; **P < 0.01 (after 1000 random permutations).

Fig. 2. MDS plot for 10 Southern Cone populations based on the Square Euclidean distance matrix. Stress ¼ 0. 13. Populations’ abbreviated names are presented in Table 1.

cluster together on the left side of the plot, whereas the other populations fall on the right side of the figure. The three Fueguian samples (YAM, KAW, SEL) cluster together in the lower right quadrant of the plot, whereas the two Patagonian samples (CHU, RNE), fall together, close to the center of the plot. CBA is placed on the right extreme of axis I, almost equidistant to the other Southern samples. The correlation coefficient between biological and geographic (GEO) distance matrices is fairly high and statistically significant (r ¼ 0.469, P ¼ 0.005). This result suggests that the morphological variation observed is related to spatial separation. In order to eliminate the geographic influence, besides correlation analysis, we carried out the Smouse–Long–Sokal test with GEO held constant. Results of correlation and partial correlation analyses between the four design matrices and biological distances are given in Table 3. By far, model 4 presents the highest correlation value (r ¼ 0.716, P ¼ 0.009). In this model, the peopling of Sierras Centrales is proposed as part of a migratory wave proceeding from the Northeast and continued to the South peopling Patagonia and Tierra del Fuego. When the geographic variable is held constant (Smouse–Long–Sokal test), the partial correlation is still high and statistically significant (r ¼ 0.664, P ¼ 0.004). Model 3 also present an r value marginally significant, but when GEO is held constant, the correlation drops to a low, insignificant value.

DISCUSSION During the last century several hypotheses about time and origin of human entry into the Sierras Centrales region were elaborated. The first scholar who proposed an ancient peopling of the area was Florentino Ameghino (1885). His ideas about human antiquity in America were strongly criticized. Instead, it was widely believed that the peopling of Sierras Centrales was recent (Serrano, 1945). Nevertheless, Montes’s research about ‘‘Miramar fossil man’’ (Montes, 1960) and principally Rex Gonza´lez’s work at the Intihuasi cave (San Luis, Argentina) demonstrated that this region was already inhabited at least from 8,000 BP (Gonza´lez, 1960). In the first decades of the 20th century several authors proposed that the colonization of this region proceeded from the Central Andes region (Outes 1911; Gonza´lez, 1944; Serrano, 1945). Later, others authors

continued postulating biological similarities with Andean groups (Mendonc¸a, 1983; Mendonc¸a et al., 1985) and Santiago del Estero (Cocilovo, 1984; Castagnino, 1985). For instance, Cocilovo (1984) proposed the existence of two or three stages in the ‘‘evolution’’ of the populations of the Central Mountains of Argentina. Cocilovo’s first stage is a migratory wave from the Andes of peoples possessing the Ayampitı´n technological complex at 8,000 BP. In the second stage, at 2,000 BP, the human inhabitants of Sierras Centrales would have experienced local evolution and gene flow with peoples from the Andean and Santiago del Estero province regions. This author also argued that people followed two migratory routes into Argentina, one proceeding from Andes—Co´rdoba and Santiago del Estero’s human inhabitants should be ascribed to this route—and one from the Northeast (Cocilovo and Di Rienzo, 1984–1985). According to climatic and archaeological evidences, Colantonio and Marcellino (2000) observed craniometric affinities between the inhabitants of Sierras Centrales and other groups from the Andean region, principally from Bolivia and North of Chile. On the other hand, other authors observed biological affinities between populations from the Central-South Atlantic shores of Brazil, Northeastern Argentina (Cocilovo and Neves, 1988–1989), and Sierras Centrales (Torres Mazzuchi, 1975). Other authors have also suggested a marked biological similarity between groups from the Northeast and Central regions of Argentina, but also extending these morphological affinities to some groups from the western region of Argentina, known as Cuyo (Marcellino and Colantonio, 1983). The correlation analysis indicates that M4 fits best with the observed biological pattern of variation (r ¼ 0.716, P ¼ 0.009), even when geography is held constant (r ¼ 0.664, P ¼ 0.004). It is therefore the most plausible scenario of peopling of the southernmost region of South America, and particularly, of the Sierras Centrales region. This model is similar to M2, in the sense that both propose a peopling process in a Northeast to South direction, but they differ in the populations involved. In M4 we propose that all the populations of Patagonia and Tierra del Fuego originated from the Northeastern wave. M2 proposed that only one of the Fueguian populations (SEL) who shares similar patterns of subsistence with the continental populations—terrestrial hunter–gatherers, derived from the Northeast route, whereas the rest of the Fueguian populations (YAM, KAW) derived from the Andean route. The results, as observed in Figure 2, show close morphological similarities between the three Fueguian populations, most likely related to a common ancestor and, as

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COLONIZATION OF THE CENTRAL TERRITORY OF ARGENTINA several authors suppose, to the retention of morphological characteristics of the first settlers of the Americas (Lahr, 1995, 1996; Lalueza Fox et al., 1997). Our results are in agreement with those presented by Me´ndez and Salceda (1995) and with our own findings based on non metric cranial data (Fabra et al., 2005), and support our hypothesis for the peopling process of the central territory of Argentina (Laguens et al., 2003). In this hypothesis we propose that the Sierras Centrales of Argentina began to be colonized by humans in the Pleistocene–Holocene transition. The first colonizers probably followed a Southeast to Northwest general direction, following least-cost routes (Anderson and Gillam, 2000) through plains and rivers, pursuing mega fauna, in an East–West general direction. This initial entrance should be seen as a deviation, migratory option, or group fission, from a Northeast–South principal migratory route that resulted in the peopling of the Pampas and Patagonia. This colonization process should not have necessarily occurred before the peopling of Patagonia and the Pampas. Rather, it could have been the result of fission of populations settled in riversides after the initial entrance. Generally, fission occurs when a population reaches a limit related with environment and social tension; so fission may occur when effective colonization is established. Conversely, the peopling of the Sierras Centrales may have occurred from the Pampas, in a South–North direction. Unfortunately, no clear riverine or paleochannel evidence exists to support either hypothesized route. Geomorphological studies at the high pampas of Sierras Centrales revealed that these areas had more benign climate conditions during the driest periods when compared with areas in equivalent latitude. These conditions made the high pampas the places chosen by megafauna for their humid and coldest conditions in comparison with oriental plains (Laguens et al., 2003). Although the archaeological record of the first inhabitants of Sierras Centrales is fragmentary and limited, there is a high probability that alluvial plains of the Parana´ tributaries were the first initial migration routes, due to their potential as least-cost routes with a variety of resources. Then, the peopling of the central territory of Argentina may have occurred following these routes, in a East–West direction, probably related to fauna migration to better climatic and geomorphological conditions. However, these conclusions must be taken with a note of caution. The study of a larger sample, representative of the different subregions of the Sierras Centrales, as well as the analysis of ancient DNA (which is currently carried out in our laboratory), is necessary to further evaluate the biological variation in Central Argentina in order to support (or not) the results obtained in this study.

ACKNOWLEDGMENTS M. Fabra is a doctoral fellow, and A.G. Laguens and D.A. Demarchi are Investigator Career Members of the Consejo Nacional de Investigaciones Cientı´ficas y Te´cnicas de la Repu´blica Argentina (CONICET). We are especially grateful to the museums that allowed access to their collections, particularly to the Museo de Antropologı´a, Facultad de Filosofı´a y Humanidades, Universidad Nacional de Co´rdoba. We are also most grateful to Graciela Cabana who kindly revised an earlier version of this article.

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American Journal of Physical Anthropology—DOI 10.1002/ajpa