Revista Argentina de Antropología Biológica ISSN: 1514-7991 [email protected] Asociación de Antropología Biológica Argentina Argentina

Aguirre-Samudio, Ana J.; González-Sobrino, Blanca Z.; Álvarez-Sandoval, Brenda A.; Montiel, Rafael; Serrano-Sánchez, Carlos; Meza-Peñaloza, Abigail GENETIC HISTORY OF CLASSIC PERIOD TEOTIHUACAN BURIALS IN CENTRAL Revista Argentina de Antropología Biológica, vol. 19, núm. 1, enero-junio, 2017, pp. 1-14 Asociación de Antropología Biológica Argentina Buenos Aires, Argentina

Available in: http://www.redalyc.org/articulo.oa?id=382249205002

How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative REVISTA ARGENTINA DE ANTROPOLOGÍA BIOLÓGICA Volumen 19, Número 1, Enero-Junio 2017 GENETIC HISTORY OF CLASSIC PERIOD TEOTIHUACAN BURIALS IN CENTRAL MEXICO

Ana J. Aguirre-Samudio1*, Blanca Z. González-Sobrino1, Brenda A. Álvarez-Sandoval2, Rafael Montiel2, Carlos Serrano-Sánchez3 y Abigail Meza-Peñaloza3

1Laboratorio de Antropología Genética. Instituto de Investigaciones Antropológicas. Universidad Nacional Autónoma de México. D. F. México 2Laboratorio Nacional de Genómica para la Biodiversidad, Unidad de Genómica Avanzada, CINVESTAV-IPN. Irapuato. México 3Laboratorio de Osteología. Instituto de Investigaciones Antropológicas. Universidad Nacional Autónoma de México. D. F. México

KEY WORDS ancient DNA; genetic analysis; mitochondrial haplogroups; prehispanic populations; barrios

ABSTRACT The ancient city of Teotihuacan was a great urban agreement with the number of migrants estimated. Tlailotlacan, and ceremonial center, whose population grew exceptionally another Teotihuacan household, was different following a small during the Classic Period (300–700 AC). Settlement patterns, interaction with Mazapa, Xolalpan, and La Ventilla. Through culture and burials have indicated an occupation that consisted the estimation of immigrants, the three households studied seem of groups of neighboring apartment compounds or barrios. We to have come into contact with Mayans from Xcaret in Yucatan, investigated the genetics of three apartment compounds in the which coincides with archaeological data reported. Genetic data Teotihuacan Valley through ancient DNA analysis to prove mul- could indicate that migration, along with reduced genetic drift, tiethnicity during the Classic Period. Amerindian mitochondrial may possibly have a more effective role among Teotihuacan haplogroups were identified in 10 subjects from San Francisco groups. This suggests that interchange with other groups did not Mazapa, 7 from San Sebastián Xolalpan, and 19 human bone restrict to commercial, service or governmental purposes, which tools from La Ventilla. These samples had a wide genetic diver- implies demographic integration and genetic fusion culminat- sity. Differences in genetic structures between the three house- ing in multiethnicity during the Classic Period in Teotihuacan. holds and seven ancient populations from central and southern Further studies can be directed to examine other households and Mexico were slight but significant p( <0.001) by FST analysis be- with future sequencing analysis. Rev Arg Antrop Biol 19(1), tween the three barrios studied. Xaltocan (post-conquest) was in 2017. doi:10.17139/raab.2017.0019.01.02

PALABRAS CLAVE análisis genético; ADN antiguo; haplogrupos mitocondriales; poblaciones prehispánicas; barrios RESUMEN La ciudad de Teotihuacan tuvo un gran crecimiento con 7 poblaciones antiguas del centro y sur de México. En los poblacional durante el Período Clásico (300-700 AC, del inglés análisis, Xaltocan fue congruente con el número de migrantes after Christ), cuando alcanzó el desarrollo urbano y llegó a ser estimado. Tlailotlacan, otro barrio de Teotihuacan, tuvo una un centro ceremonial de gran importancia. Los patrones de relación pequeña con los barrios estudiados. La estimación de asentamiento, cultura y los entierros excavados muestran una migrantes mostró que pudieron tener contacto con mayas de ocupación organizada en barrios. En este estudio se realiza el Xcaret en Yucatán, en coincidencia con los datos arqueológicos análisis genético, por medio del ADN antiguo, de tres barrios reportados. Los datos genéticos podrían señalar que la migra- ubicados en el Valle de Teotihuacan con el objetivo de identi- ción y poca deriva genética jugaron un papel importante entre ficar patrones de multietnicidad durante el Período Clásico. Se los grupos teotihuacanos, lo que sugiere intercambio con otros identificaron los haplogrupos mitocondriales amerindios en grupos por propósitos de comercio, servicios o gubernamenta- 10 individuos de San Francisco Mazapa, 7 de San Sebastián les, lo cual implica integración y fusión genética que determina Xolalpan y 19 residuos de herramientas óseas de La Ventilla. multietnicidad en Teotihuacan durante el período Clásico. Estos Estos barrios mostraron diversidad genética. El análisis de FST resultados pueden ser corroborados por estudios en otros barrios reveló poca estructura genética, pero estadísticamente signifi- de Teotihuacan y con futuros análisis de secuenciación. Rev Arg cativa (p<0.001), entre los barrios estudiados, en comparación Antrop Biol 19(1), 2017. doi:10.17139/raab.2017.0019.01.02

Teotihuacan has been described as a great *Correspondence to: A. J. Aguirre-Samudio. Laboratorio de ancestral multiethnic center of the Basin of Antropología Genética. Instituto de Investigaciones Antro- Mexico during the Classic Period; it was an pológicas. Universidad Nacional Autónoma de México. Av. Universidad 3000. Coyoacan 04510. D.F. México. E-mail: economic and political center, and an important [email protected] point of interchange extending throughout Mex- ico and Central America (Parsons, 1987). The Financiamiento: Program to Support Research Projects and Technological Innovation, UNAM (PAPIIT N° IN402015, Teotihuacan period lasted for approximately IN400913); National Council of Science and Technology eight centuries, before and during the powerful (CONACYT 100936). pre-Hispanic Period, and went from a flourish- Recibido 18 Enero 2016; aceptado 4 Julio 2016 ing Teotihuacan to its decay. There were three principal settlement pat- doi:10.17139/raab.2017.0019.01.02 1 A. J. AGUIRRE-SAMUDIO ET AL./REV ARG ANTROP BIOL 19(1), 2017 terns in Teotihuacan: the Pre-Classic period front of the Ciudadela (López Austin and López (Patlachique phase), from 400 to 100 BC ap- Luján, 2010). proximately; the Middle Horizon (Tlamimi- Burials in the Teotihuacan Valley have been lolpa, Xolalpan, and Metepec phases), from 300 excavated since 1905, with numerous explora- to 650 AC; and the Coyotlatelco phase, from tions carried out in the last century, and continu- 700 to 900 AC (Parsons, 1987; Manzanilla, ing to date (Rattray, 1997; Rodríguez Manzo, 2014). Towards the late Metepec and Coyot- 2003). Excavations were also done in apart- latelco phases, changes in ceramics represented ment compounds and multi-family residences a break in the ceramic tradition compared with with a number of rooms, platforms, corridors, early Teotihuacan (1-200 AC), which Rattray and patios, such as Tetitla, Atetelco, Las Ca- describes as catastrophic (Rattray, 1987a). laveras, Cuanalan, Barrio de los Comerciantes The first human occupation of the Teotihua- (Rattray and Civera Cerecedo, 2003), La Ventilla can Valley began in the late Pre-Classic period (Gómez Chavez, 2000), Tlamimilolpa, Tlailot- (400-100 BC), when inhabitants of the valley lacan or Oaxaca Barrio (Spence and Gamboa lived from the natural resources, such as corn Cabezas, 2003; Ortega Cabrera, 2014), and and other plants and their derivatives, found Tlajinga (Storey and Widmer, 2003). These ru- in the surrounding areas of the San Juan river ral settlements or barrios, also known as ethnic (Manzanilla, 2014). Archaeological analysis enclaves, were a part of the greater Teotihuacan has shown cultural growth, with compounds society. Presumably, these barrios were defined of monumental architecture in the center, and by a form of organization into categories that groups of residential structures placed around contained different sectors, such as political, them. Urban development was established in economic, corporate, ethnic, and domestic areas the first century (Tzacualli phase, 1-150 AC), on for families; these may or may not have shared a territory covering an area of 4km2, which has the same cultural traditions as other inhabitants been identified as the biggest Pre-Classic site in in the city (Millon, 1973). These sites were used central Mexico (Sanders, 1965). At some point in the archaeological interpretation of materials during this period, the entire city covered a sur- located in domestic places for family, political, face of over 20km2 and was home to 20000 to religious, and socioeconomic integration activi- 30000 inhabitants (Millon, 1973). It is possible ties. Nevertheless, archaeological data do not that an increase in the number of residents in the support the current knowledge on this complex settlements may have been one of the causes for barrio structure, nor has genetic homogeneity residential re-organization. Another possibility been proven (Ortega Cabrera, 2014). is that it occurred owing to greater production Thus far, ancient DNA (aDNA) analysis has requirements (Spence, 1987). In the following allowed reconstructing a part of the genetic his- century, the Miccaotli phase (100-200 AC) was tory of ancient populations. Because of its ma- characterized by the construction of the Avenue ternal heritage, mitochondrial DNA (mtDNA) of the Dead and the Temple of the Feathered Ser- extracted from osseous samples provides a po- pent, an important ritual center in Teotihuacan tential source of genetic evidence. The use of (Manzanilla, 1995). Population growth reached mtDNA has an advantage in aDNA studies since an estimated of 125000 inhabitants (Kurtz et al., the molecule itself is contained in numerous mi- 1987) in the Tlamimilolpa phase (250-400 AC). tochondria within the cell, which increases the It was the most important demographic growth likelihood of obtaining DNA from preserved taking place in Teotihuacan, occurring in the bone material despite taphonomic action. The Middle Horizon or Classic Period, and repre- aDNA is always scarce and degraded, and to be senting a large urban model (Manzanilla, 1995) extracted under strict conditions to avoid contam- designed to provide enormous supplies of food, ination from modern DNA. In Mexico, genetic water, and dwellings, all of which implied gov- studies of pre-Hispanic populations are scarce ernmental organization and open commercial (Merriwether, 1994; González-Oliver, 2001; interchange. Teotihuacan culture was magnified Kemp et al., 2005; De la Cruz et al., 2008; by the Pyramid of the Moon, the Temple of the Solórzano Navarro et al., 2009; Álvarez-Sandoval Feathered Serpent and the Great Compound in et al., 2014, 2015). All samples showed the vari-

2 GENETIC HISTORY OF TEOTIHUACAN ation expected for Amerindian mtDNA, analyzed fragments were dated to the late Xolalpan phase. in most of the cases using restriction enzymes In addition, the remains of bone tool manufac- over amplified DNA to generate restriction frag- turing from 20 different right femurs and one ment length polymorphisms (RFLP) characteris- humerus were taken out from La Ventilla’s tic of each of the mtDNA haplogroups (Wallace et pits and dumps and were also sampled. These al., 1985). Historically, such polymorphism stud- fragments from La Ventilla, another neighbor- ies carried out around the world have been fo- hood belonging to the Tlamimilolpa to Metepec cused on research of the peopling of the Americas phases, were dated between 300 to 700 AC. The (Wallace et al., 1985; Shurr et al., 1990; samples corresponded to excavations in sec- Torroni et al., 1993; Stone and Stoneking, 1993; tors N1W2, S1W1, and S1W2 in Teotihuacan Merriwether et al., 1994), population expansion (Gomez Chavez, 2000; Meza-Peñaloza, 2015). by language family (Monroe et al., 2013), and hu- A total of 111 pre-Hispanic individuals were man dispersal out of Africa (Behar et al., 2008). analyzed; the origins of 75 of them are referenced In this research, aDNA analysis helped to in Merriwether et al. (1994), González-Oliver characterize the three households at the Teoti- et al. (2001), Juárez (2002), López-Armenta huacan Valley through mtDNA recovered from (2007), Herrera-Salazar (2007), Mata-Míguez et burials in San Francisco Mazapa, San Sebastián al. (2012). In the present study, 36 Teotihuacan Xolalpan. Remains of human bone material individuals from the Classic period (300-600 used to manufacture tools in La Ventilla were AC) and located in the Basin of Mexico were also utilized to characterize this barrio. We in- analyzed. Dating and geographical distribution vestigated multiethnicity among households are detailed in Table 1. during the Middle Horizon in Teotihuacan, based on the 2000 architectonic corporative Contamination precaution and control structures established in the city (Millon, 1973), Extraction and amplification setup were and subsequent demographic changes during its conducted under standard authentication cri- hegemony. Archaeological data provided evi- teria for ancient DNA analysis (Pääbo et al., dence of population growth and integration with 2004). The spaces were physically separated other regions in Mexico, thus making it possible in each procedure, cleaned with 10% bleach to predict genetic variation within the city. solution, 70% alcohol, and ultraviolet (UV)-ir- radiated for 45min. Autoclaved disposable ma- MATERIALS AND METHODS terials, gloves, mouth masks, hair covers, dis- posable cloths and equipment confined to the Samples workspace were used. The solutions used were also autoclaved and UV-irradiated. Prior to The skeletons studied belong to a collection eliminating any minor surface contamination, of pre-Hispanic individuals, which form part of bone samples were UV-irradiated and cleaned the national archaeological heritage. The collec- with gentle soap, 10% bleach solution, and tion is kept under the custody of the Instituto 70% alcohol. The clean fragments were placed Nacional de Antropología e Historia, which has in tubes with zircon beads and were homog- authorized their use for study purposes. enized in FastPrep®-24 instruments (MP Bio- Phalanxes and femur fragments of 11 indi- medicals). PCR pipettors were only used for viduals from San Francisco Mazapa’s burials ancient DNA analyses. Aerosol-resistant bar- were examined. These burials belonged to a rier tips to prevent crossover were utilized and household located in sector N2E2 (Serrano et the mix for PCR preparation was performed in al., 1991) of Teotihuacan’s planimetric map in a cabinet with continuous airflow (or positive), central Mexico (Millon, 1973). These burials filtered and UV-irradiated. date back to the Tlamimilolpa-Xolalpan phas- Molecular data obtained at the Anthro- es (250-400 AC). Fragments of radius, femur, pological Genetics Laboratory (Universidad tibia, and fibula of 8 subjects from sector N4E2 Nacional Autónoma de México, Mexico City) (Millon, 1973) at the San Sebastián Xolalpan were reproduced at the aDNA laboratory of the household were recovered and sampled. These National Laboratory of Genomics for Biodiver-

3 A. J. AGUIRRE-SAMUDIO ET AL./REV ARG ANTROP BIOL 19(1), 2017

sity (CINVESTAV, Instituto Politécnico Na- cional) using high-resolution melting (HRM) analysis (Álvarez-Sandoval et al., 2015) in or- der to authenticate aDNA results. Collagen was observed as indirect evidence of DNA survival prior to ascertaining the biochemical preserva- This study This study This study This study References Juárez, 2002 tion in the bone material utilized (see Aguirre López Armenta, 2007 López Herrera Salazar, 2007 Herrera Salazar, et al., 2011). Merriwether et al., 1994 Mata-Míguez et al., 2012 Mata-Míguez et al., 2012 González-Oliver et al., 2001 DNA extraction, amplification, and statisti- cal analysis Powdered samples, between 300-500mg, were digested with extraction buffer (50µl 0.5M EDTA pH 8, 750µl of 10% SDS, 200µl Chiapas 20mg Proteinase K) overnight at 37°C in slow Basin of Mexico Basin of Mexico Basin of Mexico Basin of Mexico Basin of Mexico Basin of Mexico Basin of Mexico agitation. Phase separation was done by cen- Xcaret, Geographic distribution

Puebla, Basin of Mexico trifugation for 5min at 4°C with shaking at 13 500 rpm, followed by the GENECLEAN® procedure based on silica matrix (www.mpbio. - - - com). The resulting pellet was dried at room 0.619 0.733 0.236 0.374 0.571 0.467 0.714 0.593 temperature for 10min and re-suspended in 30-

Diversity (H) 50µl deionizer and distilled water conditions previously reported in our laboratory (Aguirre 0 4 0 0 0 5 0 0 3 13 50 et al., 2011). % D Duplicated extracts were used for analysis of the markers that define Native American 0 7 0 8 0 0 0 21 14 14 100 % C mtDNA lineage clusters (Stone and Stonek- ing, 1993). 5-8µl of DNA extract was used 0 4 0 0 0 in PCR reaction mixture, which contained 1x 20 30 16 30 43 25 % B buffer, 1mg/ml BSA, 2.0mM MgCl2, 200µM mix dNTP, 0.6µM each primers, 1U AmpliTaq 0 60 30 88 58 70 43 58 50 100 100

% A % Gold® polymerase (Applied Biosystems) in a total volume of 25µl. Cycling had an initial de- naturation step which was performed at 95°C 9 9 4 8 7 15 10 24 19 10 36 for 5min; 60 cycles at 95°C for 20sec, 55°C (or 60°C in the case of Hae III site primers) Haplogroup frequencies, geography distribution and dating frequencies, 1. Haplogroup TABLE Sample size for 30sec, 72°C for 30sec, and an extension of 72°C for 10min. PCR products were visualized using 3% Methaphord® agarose-gel electro- phoresis. RFLP analysis by enzyme digestion Date was done at 37°C for HinC II (for haplogroup 600-800 AC 600-800 500-700 AC 500-700 600-800 AC 600-800 300-700 AC 300-700 300-500 AC 300-500 250-400 AC 250-400 250-400 AC 250-400 300-700 AC 300-700 1100-1500 AC 1100-1500 1300-1350 AC 1300-1350 1400-1500 AC 1400-1500 C), Hae III (for haplogroup A), and Alu I (for haplogroup D). Genotypes were observed in 15% polyacrylamide gel. For statistical analysis the sum of sample haplogroups from the San Sebastián Xolalpan, La Ventilla, and San Francisco Mazapa barrios Copan Xcaret Cholula was considered as Teotihuacan representative. Population La Ventilla La Tlailotlacan Teotihuacan Tlailotlacan was not included in the unit, since

San Francisco Mazapa this barrio has been archaeologically described San Sebastian Xolalpan Xaltocan (pre-conquest) Xaltocan (post- conquest) as either Oaxacan or external in origin.

4 GENETIC HISTORY OF TEOTIHUACAN

We calculated haplogroup diversity (H) and RESULTS frequencies. Hierarchical clustering of spatial frequency of the groups was constructed in The population studied comprised 36 indi- the Statistica software, which was explored viduals from three households from the Teoti- by Principal Components Analysis. Also, the huacan Valley. centroid method calculated the distance be- Three of the samples were taken out of the tween two clusters for all variables and cases; database since it was not possible to obtain the the merged cluster is a weighted combination final genotype from one of them, and for the re- of the centroids of the two individual clusters. maining two, results could not be replicated. Hierarchical clustering was analyzed with the Mitochondrial ancestry by standard restric- algorithm K-Means Clustering (within Statis- tion fragment RFLP for haplogroups A, B, C, tica) to calculate distance or similarity matrix D were identified: 58% assigned to A, 25% to between all pairs of cases. Euclidean distances B, 14% to C, and 3% to D haplogroups (Table between clusters were obtained. 1). The number of haplogroups was smaller in Genetic differentiation levels between the Mazapa group with 2 variants, in contrast paired populations were estimated by Analy- to La Ventilla and Xolalpan, which had 3 and sis of Molecular Variance (AMOVA), pair- 4 variants respectively; Cholula (these groups wise FST distance, and Slatkin’s distances with were located geographically in Central Mexico) 10000 permutations using Arlequin version 3.0 and Copan (in Honduras) had 1, Tlailotlacan 2, (Excoffier et al., 2005). The resulting groups Xaltocan and Xcaret (from Southeast Mexico) from the structure analysis were employed to 3 haplogroups. The major diversity (H) was construct a dendrogram by the UPGMA (Un- between Teotihuacan and Xaltocan ranging be- weighted Pair-Group Method with Arithmetic tween 0.593 and 0.733, distributed in the Ba- Averaging) method using matrix distance of sin of Mexico, toward the central region of the linearized FST. country (Table 1). The estimated rates of gene flow for the 10 In the Principal Component Analysis, Teoti- population comparisons were computed, the huacan households of La Ventilla and post-con- matrix was based on of previous matrix of Slat- quest Xaltocan along with the total frequencies kin’s linearized FST t/M=FST/(1-FST), where two from La Ventilla/Mazapa/Xolalpan (all with 4 populations of size N exchange a fraction m of haplogroups) and Mazapa were closer to Xola- migrants of each generation, and mutation rate lpan and Tlailotlacan with 50% of haplogroups u is negligible compared to the migration rate A and D (Fig. 1). m, following equilibrium between migration The Centroid Method showed that the and drift. Additionally, the calculated gene flow groups from Bonampak (at the south of Mex- considers that only two populations exchanged ico), Cholula, Xcaret, Tlailotlacan, and Copan migrants, whose estimator was M=Nm for hap- were cases distant from the center, but classified loid populations, where M=1-FST/2FST (Excof- as a small cluster that included 3 cases. Xalto- fier, 2006). Linear regression was calculated can (post-conquest) and Teotihuacan, Tlailotla- between M and geographic distance (in km) can excluded, were fairly close to the centroid. in Sigma plot v.10.0. Infinite values of migra- Despite the dimensions registered at centroid, a tion rates estimated were collocated only for principal cluster was found with algorithm K- graphic representation. To test the hypothesis Mean, clustering in 10 cases (cluster no 1), and a that Teotihuacan was a multiethnic city dur- second cluster classified Copan (Table 2), whose ing the Mesoamerican Classic Period and that Euclidean distances were 1 vs 2 of 0.5806, and it extended toward other parts of present-day between cluster 2 vs 1 of 0.3371. Mexico or even Central America, we compared Altogether these analyses showed several the samples reported from Mayan sites such as distinctive populations, Copan standing out for Xcaret, Copan, and Bonampak; also Cholula its high incidence of haplogroup C. Also, Xal- culture; and Tlailotlacan or Oaxaca barrio and tocan (pre-conquest), far from the center, was Xaltocan (see Table 1 for references. For loca- different due to its proportion of haplogroups A tions see Fig. 4). and B. Cholula and Bonampak formed a cluster

5 A. J. AGUIRRE-SAMUDIO ET AL./REV ARG ANTROP BIOL 19(1), 2017

Fig. 1. Plot of principal components for the Amerindian haplogroups in 10 populations from ancient Mexico. because of their high proportion of haplogroup tral America. Such hypothesis cannot be con- A, followed by the Xcaret population. firmed with the current comparisons between Historical archaeological references tell us the genetic distances, where genetic variance of exchange between groups from Teotihua- among groups was undetectable, and inside can and central and southern Mexico to Cen- of proven groups there was a higher range of

TABLE 2. Cluster analysis with Centroid Methods and K-Means clustering between cases

Cases Principal Factor Distance from cluster center Distance from cluster center (Centroid) (k-mean) (k-mean) Cluster 1 Cluster 2

La Ventilla -0.180 0.088 - Mazapa 0.099 0.094 - Bonampak 0.862 0.201 - Tlailotlacan -0.692 0.228 - Xcaret 0.525 0.140 - Xolalpan -0.645 0.186 - Teotihuacan -0.197 0.079 - Cholula 0.862 0.201 - Xaltocan (post- 0.103 0.034 - conquest) Xaltocan (pre-conquest) -0.038 0.241 - Copan -2.082 - 0

6 GENETIC HISTORY OF TEOTIHUACAN

77.23 and 80.59 percent of variation (Table valley of Mexico. In contrast, Mayans from

3). The pairwise FST results showed structure Copan showed a higher genetic distance (Fig. population between Teotihuacan, central and 2) indicating a modest relationship with Teoti- southern Mexico groups (FST=0.228 versus huacan.

FST=0.194, p=0.00, Table 3). Structure analy- To try to understand the genetic distance sis detected significant differences between 2 found, we estimated the number of migrants groups, which accounted for 1.64% of the ge- (Nm) between the populations studied. The es- netic variance; whilst subgroups within groups timate comparisons of gene flow for 10 popu- explained 17.7%. In this second analysis, lations are given in Table 4. Some infinite val- Bonampak was removed due to its inconven- ues were noted, indicating higher Nm, where iently small N, and because the variation be- gene flow exceeded any effects of genetic tween groups was negative (point 1 in Table 3). drift decreasing the differentiation, as was the A dendrogram constructed by UPGMA case of the comparisons of Mazapa vs. Xola- with FST values showed the arrangements of lpan, Mazapa vs. La Ventilla (p<0.01), Xola- populations (Fig. 2) with the same hierarchical lpan vs. La Ventilla (p=0.028), and Xaltocan clustering among distances with a difference (post-conquest) with La Ventilla/Xolalpan/Ma- between Copan and the rest of the cases. Ge- zapa. Another higher Nm was between Xcaret/ netic distance analysis grouped Mayans from Cholula (p<0.02). Also, higher values were the Xcaret and Cholula populations followed obtained between the populations from Xalto- by other branches that divided into Teotihua- can (post-conquest) and Xcaret /Tlailotlacan can and Xaltocan (post-conquest) groups, sug- (Nm>4, p=0.07), Xcaret vs. Mazapa (p=0.07), gesting migrations into Teotihuacan due to the Maya vs. La Ventilla (p<0.01), Xaltocan (pre- dominant role the city held during the Classic conquest) vs. La Ventilla (p<0.01), and Xalto- Period and because of population movements can (pre-conquest) vs. Xolalpan (Nm=19.49, occurring in the Post-Classic Period around the p<0.01). In Table 4, all Nm <1 describe a low

TABLE 3. Hierarchical analysis of molecular variance (AMOVA) using different structures from the Teotihuacan groups in Mexico. Describe percent of variation

Groups Among Among Within FST (p) FSC (p) FCT (p) groups populations populations within groups

1.Central 3 Maya -8.87 31.65 77.23 0.228 0.291 -0.089 Mexico, and populations, (0.000) (0.000) (0.746) South of Mexico 4 Teotihuacan to Honduras groups included with 3 populations from Central Mexico

2.Teotihuacan, 2 Maya 1.64 17.77 80.59 0.194 0.181 0.0164 Central Mexico populations, (0.000) (0.000) (0.241) and South of 4 Teotihuacan Mexico groups and 3 populations from Central Mexico

1.Includes groups from central Mexico: Cholula, Teotihuacan (Xolalpan, La Ventilla, Mazapa), Xaltocan; and Mayan – speaking people from southern Mexico: Xcaret, Bonampak; and Honduras: Copan. 2.Includes Teotihuacan, Tlailotlacan, Cholula, Xaltocan, and mayenses: Xcaret y Copan.

7 A. J. AGUIRRE-SAMUDIO ET AL./REV ARG ANTROP BIOL 19(1), 2017

Fig. 2. Dendogram based on UPGMA of mitochondrial haplogroups’ DNA for 10 populations from central and south Mexico. Xaltocan presents two data: post-conquest (postc) and pre-conquest (prec).

gene flow. Genetic drift may influence each of otihuacan groups (p=0.036, Table 4). Linear the populations, such as Copan and Teotihua- regression was calculated from 10 population can (included three studied groups), Cholula comparisons depending on their geographic and Teotihuacan groups, Xaltocan and Copan distance (Fig. 3). Infinite Nm values are rep- /Cholula (p<0.01), Tlailotlacan with other Te- resented by 13 Nm (r2=0.12994, p<0.0001) in

TABLE 4. Mitochondrial DNA pairwise comparisons by M=Nm-values (below diagonals) and p-values (above diagonals) between prehispanic populations from Mexico

1 2 3 4 5 6 7 8 9 10

1 0.142 0.020 0.070 1.000 0.002 0.103 0.000 0.073 0.000 2 0.111 0.429 0.801 0.210 0.020 0.359 0.073 0.717 0.000 3 ∞ 0.000 0.687 0.020 0.041 0.615 0.227 0.440 0.001 4 4.868 0.162 1.933 0.165 0.005 0.721 0.010 0.491 0.000 5 1.150 0.299 0.676 ∞ 0.028 0.151 0.003 0.194 0.000 6 4.977 0.473 2.429 ∞ ∞ 0.041 0.315 0.186 0.000 7 0.905 0.186 0.610 1.936 2.210 3.652 0.036 0.644 0.000 8 4.473 0.448 2.556 ∞ ∞ ∞ 2.662 0.234 0.000 9 4.910 0.323 2.589 ∞ ∞ ∞ 8.231 ∞ 0.000 10 0.735 0.309 0.590 3.000 19.496 4.084 ∞ 3.862 10.223

1: Xcaret, 2: Copan, 3: Cholula, 4: Mazapa, 5: Xolalpan, 6: La Ventilla, 7: Tlailotlacan, 8: Teotihuacan, 9: Xaltocan (post-con- quest), 10: Xaltocan (pre-conquest). ∞ are infinite values. Bold numbers indicate significant values since comparison between the study groups.

8 GENETIC HISTORY OF TEOTIHUACAN

Fig. 3. Number of migrants estimated by geographic distance among the10 populations compared. A linear regression analysis can be observed; populations’names of pairwise comparisons are M:Maya (from Xcaret), Ch:Cholula, V:Ventilla, Mz:Mazapa, Tl:Tlailotlacan, X:Xolalpan, Xa:Xaltocan (pre-conquest), Xp:Xaltocan (post-conquest). the graph. There was no correlation between tween the rural residents of the three barrios geography and number of migrants. studied. Tlailotlacan did not group close in the cluster formed by La Ventilla-Mazapa- DISCUSSION Xolalpan-Xaltocan, but it was part of a larger cluster of 10 cases, which could be visualized Genetic variation was identified in La as a different residence compound with partial Ventilla, San Francisco Mazapa, and San Se- integration in Teotihuacan. bastián Xolalpan households, and it is evi- The analysis of population structure found denced by the distributions of haplogroup fre- a small molecular variance among groups. It quency. In the case of La Ventilla, the sample was possible to observe a slight genetic dif- comes from remains of bones used as tools, ferentiation within the three studied groups, which could represent the group, given that suggesting that migration and genetic drift previous studies were able to demonstrate that could lead to a more effective role between bones turned into artifacts belonged to inhab- the groups. itants from this barrio (Meza Peñaloza, 2015). Genetic distance analysis identified two Also, through stable isotope analysis, it was important branches: the first linked Mayans found that such materials presented a local di- from the Xcaret and Cholula populations, etary consumption (Arnaud, 2014). suggesting ancient relationships; the second Residential compounds showed to be com- branch grouped the three households studied posed by distinctive groups across the popula- with post-conquest Xaltocan; a third, small tions compared. Despite this, similar haplo- branch contained Tlailotlacan and pre-con- group frequencies as those observed in Teoti- quest Xaltocan (Fig. 2). The suits of cluster huacan (Table 1) have been reported through- analyses showed links formed between La out Mexico, and are maintained in present Ventilla-Mazapa-Xolalpan, suggesting a pro- day (A–50%, B–24%, C–18%, D–8%), even nounced population dynamic in Teotihuacan though regional differences may be variable during the Classic Period in the Basin of (Gonzalez-Sobrino et al., 2015). Principal Mexico and regions such as Xcaret in Quin- Component Analysis showed closeness be- tana Roo.

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Archaeological interpretations have Pairwise comparison highlights the rela- showed that the different households shared tion between Xaltocan (haplogroup frequen- religious, government, commercial, and cies after the conquest) and La Ventilla and burial activities. We added genetic relations Xolalpan. History provides an explanation for shown by the genetic distance between the Te- this connection: during the Middle Horizon, otihuacan groups, which could correspond to the settlement system extended over several the ethnic view of barrio described previously hundred hectares, which included monumen- by archaeologists (Millon, 1973). tal architecture, civic-ceremonial architec- Paleomigration markers or isotopic an- ture, sites of variable size, occupation in the thropology may perhaps also reflect patterns hills, and highly populated regions spread- of population movements within Teotihuacan. ing over the Valley of Mexico. Azcapotzal- In fact, it is possible to distinguish ethnic areas co, located to the west of the Texcoco Lake, in Teotihuacan by cluster analysis of craft pro- was an important site in the Middle Horizon duction, such as in Tlajinga, where elements (Parsons, 1987). In this region, Xaltocan was were found to be related to other regions in situated 35km north of modern Mexico City, Teotihuacan (Altschul, 1987). In addition, Te- whose demographic and genetic history was opancazco, another barrio from Teotihuacan, related to the Aztec conquest (Mata-Miguez displays foreign activity from different Mes- et al., 2012). Accordingly, the settlement pat- oamerican coastal regions as far as Honduras terns that extended to the Valley of Mexico and (Manzanilla, 2015). In the are consistent with the variability found in same way, sacrifices associated to the Pyra- this study, and it may be reflected by genetic mid of the Feathered Serpent have been iden- frequencies of maternal inheritance (view Ta- tified by isotopic analysis as foreign in origin, ble 4, p<0.01). maybe to demonstrate the hegemony of Teoti- La Ventilla and Mazapa households pre- huacan (White et al., 2002). A previous study sented evidence of Mayan integration, es- from other households such as Oaxaca, Cueva pecially from Xcaret, due to their mobility de las Varillas, Oztoyohualco, and Barrio de toward Teotihuacan (p<0.01, p=0.07 trend, los Comerciantes showed similar migratory respectively) by the estimated gene flow. movements in Teotihuacan, where residence Previous interpretations of Mayan ceramics change was continuous throughout people’s found in Teotihuacan considered the interac- lives (Price et al., 2000). In all the barrios tion of both societies. Clayton (2005) identi- described above, as well as in the sacrificial fied a very important relationship, over the site, evidence is directed toward identifying course of five centuries from the late Pre- multiethnic features in Teotihuacan, which is classic through late Classic, influencing po- consistent with lower genetic differentiation litical Mayan organization from Teotihuacan. and connection between the three households The interaction between Teotihuacan groups studied, thus indicating integration. and Mayans occurred in sub-regions such as Migration patterns were drawn from close , Petén, and , but there was to remote regions, estimated by M values, no direct interaction with Copan (Clayton, as an indirect method of gene flow. Statis- 2005). Lower numbers of migrants estimated tics significance displayed a strong relation between Teotihuacan and Copan were statis- between the Teotihuacan groups of Mazapa, tically tested (Table 4), confirming no con- Xolalpan, and La Ventilla, but not of Tlailot- nection between the two sites, as shown by lacan. The Oaxaca barrio has been described the origin of the ceramics. as a distinctive enclave within Teotihuacan, We found no correlation between Cholula showing clear mobility from the Oaxacan re- and Teotihuacan (p<0.01), but the gene flow gion (Ortega Cabrera, 2014). Indeed, oxygen between Mayans from Xcaret and Cholula isotope analysis revealed that the Tlailotlacan was high. Rattray, under ceramic analysis, barrio received immigrants from the Valley particularizes Cholula as a midpoint between of Oaxaca, over 100km away (White et al., Monte Albán (Oaxaca) and Teotihuacan 2004). (Rattray, 1987b). It is possible that the genetic

10 GENETIC HISTORY OF TEOTIHUACAN relationship between Teotihuacan and Cholu- the Zapotec enclave. Actually, the origin of la was not exhibited at this time in history, and Teotihuacan groups is not well known and any relationship could have been generated is still a topic for discussion. What we put due to commercial purposes, using Cholula as forward for consideration in this study is that a midpoint in the trade route toward Teotihua- people arrived in Teotihuacan from differ- can. An unrepresentative sample could also ent places as a consequence of the boom in explain the absence of a relationship. Mexico during the Classic Period. The Clas- Why did people migrate to Teotihuacan sic Period was of great importance across from different places in the Middle Horizon Mesoamerica in regions such as the Valley of or in the Classic Period? Answering is com- Oaxaca (Blanton, 1987), Mayan sites like Pe- plicated. Some current theories on why hu- ten, Tikal and Guatemala (Coe, 1972), Tlax- mans migrate point to a search for a more fa- cala (Snow, 1972), the Atemajac Valley in the vorable scenario or a more attractive climate; archaeological zone of Ixtepe, Zacatecas to other reasons include cultural or political op- the west of Mexico (Corona Núñez, 1972; pression or discrimination, natural disasters Hers, 2014); central and northern Veracruz or wars, and cultural motives such as similar (Brüggemann, 2014), and the northeastern religions and language (Norton, 2009). In this region (Michelet, 2014). Period, subsistence activities consisted in the Genetic studies through ancient mitochon- practice of agriculture, and the utilization drial DNA have allowed drawing converging of wild plants and local fauna within Teoti- lines, which may be subject to discussion or huacan’s surrounding areas in the Basin of exploration (Fig. 4). In general, these dynam- Mexico (McClung, 1987). Food sources were ic patterns appeared in ancient Mesoamerica stored and their organization and control of- according to archaeological data, and point to fered advantage over other places. Industry Teotihuacan being a multiethnic society. Our during the Patlachique, the Miccaotli and the results support this multiethnicity; however, Tlamimilolpa phases (in the Classic Period) it is necessary to consider other households, grew rapidly. Thus, obsidian technology was to increase the sample size, and to perform necessary in commerce, and artisan work- sequencing to examine the distribution of shops interchanged and exported products to frequencies. In the meantime, the households Mayan areas; under this argument the obsid- examined in this study showed the following: ian industry was under the control of Teoti- a) they seem to form a group; b) to exhibit a huacan’s state power (Spence, 1987). As the similar variability related to frequencies in ur- center of production and distribution of ob- ban groups in modern Mexico; c) and to show sidian, Teotihuacan was a commercially im- the same distribution observed in present-day portant place and culturally attractive to other haplogroups. regions. According to this development, it has been estimated that, at its peak during the CONCLUSIONS Classic Period, there were 125000 to 200000 inhabitants in Teotihuacan (Millon, 1981), The results obtained in this study suggest 51 000 in the Valley of Oaxaca during late probable gene flow due to population growth Monte Albán I (Kowalewski et al., 1989), and in Teotihuacan, where geographic distance was 35000 in Tikal during the late Classic Period not a barrier for interchange. The ancient people (Sanders, 1973). Indeed, agriculture was piv- of México were interrelated in economic, com- otal, which conditioned Mesoamerican life- mercial, political, and religious contexts, which style and survival, promoting the gene flow made genetic joins possible. We propose a rep- between central and southern areas in Mexico resentation of putative population movements, (López-Austin, 1989). represented in Figure 4, during the Classic Pe- A hypothesis suggests that people from riod, where genetic distance and routes con- Teotihuacan had arrived from other places, verge in Teotihuacan, suggesting multiethnicity. which is sustained by similarities in ceram- The contribution of the haplogroups obtained is ics; it was the case of Tlailotlacan, part of limited, and should be taken with caution since

11 A. J. AGUIRRE-SAMUDIO ET AL./REV ARG ANTROP BIOL 19(1), 2017

Fig. 4. Mathematical interpretation of population movements that putatively occurred during the Classic Period (300-700 AC) among indigenous groups from south and central Mexico. Routes are based upon the genetic dis- tances and migrants estimated with M-values. It is possible that there was a route originating in the south (Mayans from Xcaret) and possibly going through Cholula, until they arrived in Teotihuacan (thin black lines); but people from Cholula had less contact with Mazapa, Xolalpan, and Tlailotlacan (thick, light gray lines) except those from La Ventilla, who could have some relationship (thick, dark gray lines). In Teotihuacan there was an interchange between the three barrios studied, except Oaxaca barrio.

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