View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by MPG.PuRe AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 155:600–609 (2014) Between Andes and Amazon: the Genetic Profile of the Arawak-Speaking Yanesha Chiara Barbieri,1 Paul Heggarty,2 Daniele Yang Yao,1 Gianmarco Ferri,3 Sara De Fanti,1 Stefania Sarno,1 Graziella Ciani,1 Alessio Boattini,1 Donata Luiselli,1* and Davide Pettener1 1Department of Biological, Geological and Environmental Sciences, University of Bologna, 40126 Bologna, Italy 2Department of Linguistics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany 3Dipartimento di Medicina Diagnostica, Clinica e di Sanita Pubblica, Universita degli Studi di Modena e Reggio Emilia, 41124 Modena, Italy KEY WORDS mtDNA; Y chromosome; STR; South America; language ABSTRACT The Yanesha are a Peruvian population and one INDEL diagnostic for assigning haplogroups). who inhabit an environment transitional between the We uncover sex-biased genetic trends that probably Andes and Amazonia. They present cultural traits char- arose in different stages: first, a male-biased gene flow acteristic of both regions, including in the language they from Andean regions, genetically consistent with high- speak: Yanesha belongs to the Arawak language family land Quechua-speakers and probably dating back to (which very likely originated in the Amazon/Orinoco Inca expansion; and second, traces of European contact lowlands), but has been strongly influenced by Quechua, consistent with Y chromosome lineages from Italy and the most widespread language family of the Andes. Tyrol, in line with historically documented migrations. Given their location and cultural make-up, the Yanesha Most research in the history, archaeology and linguistics make for an ideal case study for investigating language of South America has long been characterized by percep- and population dynamics across the Andes-Amazonia tions of a sharp divide between the Andes and Amazo- divide. In this study, we analyze data from high and nia; our results serve as a clear case-study confirming mid-altitude Yanesha villages, both Y chromosome (17 demographic flows across that ‘divide’. Am J Phys STRs and 16 SNPs diagnostic for assigning haplogroups) Anthropol 155:600–609, 2014. VC 2014 The Authors. American and mtDNA data (control region sequences and 3 SNPs journal of physical Anthropology published by Wiley Periodocals, Inc. INTRODUCTION On a local scale, meanwhile, the complex demographic histories of precolonial times are difficult to reconstruct, Particularly since the 1990s, population genetic stud- and different genetic data often hint at contradictory ies have taken a keen interest in the indigenous popula- explanations (Wang et al., 2007; Bisso-Machado et al., tions of the Americas, given their unique demographic 2012). Recent studies have focused on the dynamics background (Schurr et al., 1990; Rothhammer and Silva, behind the current distribution of language families, or 1992; Torroni et al., 1992). The Americas were the last on the role of the environment in shaping demographic major continents to be settled by humans, probably by a small founding population around 15–17 kya (Tamm et al., 2007; Fagundes et al., 2008; Kitchen et al., 2008), Additional Supporting Information may be found in the online yet they are home to surprisingly wide cultural and lin- version of this article. guistic diversity (Nettle, 1999). On a broad scale, the available genetic data clearly Grant sponsor: The European Research Council; Grant number: point towards a strong founder effect and a rapid spread ERC-2011-AdG 295733 grant (Langelin); Grant sponsor: The Fac- through the continent (Wallace et al., 1985; Lewis et al., ulty of Mathematical, Physical and Natural Sciences of the Univer- 2007), from the Bering Strait to Patagonia; this scenario sity of Bologna. is in agreement with archaeological indications of human presence in southern Chile from c. 14,500 ya This is an open access article under the terms of the Creative (Dillehay et al., 2008), although no actual human Commons Attribution-NonCommercial-NoDerivs License, which remains of this antiquity have yet been found there. permits use and distribution in any medium, provided the original Uniparental genetic markers in native American popula- work is properly cited, the use is non-commercial and no modifica- tions or adaptations are made. tions are affected by this bottleneck (Torroni et al., 1993; Bortolini et al., 2003; Schurr and Sherry, 2004; Bisso- *Correspondence to: Donata Luiselli; Department of Biological, Machado et al., 2012; Battaglia et al., 2013): for mito- Geological and Environmental Sciences, Univeristy of Bologna, via chondrial DNA (mtDNA) only four macro haplogroups Selmi 3, 40126 Bologna, Italy. E-mail: [email protected] are found at appreciable frequencies (A, B, C, and D), while for the Y chromosome just a single lineage is pre- Received 3 July 2014; accepted 4 September 2014 dominant (macro-haplogroup Q). This limited genetic variability contrasts with high differentiation between- DOI: 10.1002/ajpa.22616 populations, probably arisen through drift (Yang et al., Published online 17 September 2014 in Wiley Online Library 2010; Bisso-Machado et al., 2012; Roewer et al., 2013). (wileyonlinelibrary.com). Ó 2014 THE AUTHORS. AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY PUBLISHED BY WILEY PERIODOCALS, INC. THE GENETIC PROFILE OF ARAWAK-SPEAKING YANESHA 601 trajectories (Rothhammer and Dillehay, 2009; Callegari- although a few were borrowed from other forms of Que- Jacques et al., 2011; Amorim et al., 2013; Ramallo et al., chua, from nearby Amazonian languages, and from other 2013; Roewer et al., 2013; Sandoval et al., 2013). unidentified source languages (Adelaar, 2006). Quechua Both linguistic and environmental factors are unargu- influence seems to have largely ceased a few centuries ably important in understanding cultural diversity ago, but instead Spanish has brought another layer of across South America. Much of the continent falls under loanwords, since the Selva Central was first colonized by one of two extreme environments: the Amazon rainfor- Spanish Franciscan friars from 1635. Nonetheless, the est; or the Andes, the world’s second-highest mountain main sources of European migrations, starting in the range running north-south through the tropics, parallel 1850s, were Prussia and Italy (Contreras, 2007). to the Pacific coast. The Central Andes saw the emer- The latest census (INEI) puts the overall Yanesha popu- gence of one of the world’s few independent civilizations, lation at 9213, in villages scattered across three main alti- as complex societies arose here up to five millennia ago. tude levels: High Selva (HS) up to 1,800 m, Intermediate However, the main indigenous language family of the Selva (IS) up to 800 m and Low Selva (LS) up to 100 m. Andes, Quechua, expanded only in the last 1500 years As a rule, the higher the altitude the greater the degree of or so. Its spread to the highlands of Ecuador and Bolivia urbanization and westernization, the lower communities dates to the Inca and early Spanish colonial periods, but being the most isolated. What historical sources fail to Quechua’s core range in the highlands of central and reveal is the extent to which the HS and IS settlements southern Peru dates back to earlier expansive processes, reflect recent incoming Quechua (and even European) pop- with an obvious candidate being the ‘Middle Horizon’ ulations from the Andes, who switched language to Ara- expansion, c. 550–1000 CE, originating from the site of wak here, or original settlements by Arawak-speaking Wari, near modern Ayacucho (Beresford-Jones and Heg- populations arriving from Amazonia. garty, 2013). Both Wari and the Incas are widely pre- This study provides novel genetic data for the Yanesha sumed to have fostered significant population people, and discusses their genetic variability against movements through the Central Andes highlands, lead- the South American background. To this end, we ana- ing to considerable genetic homogenization. lyzed uniparental markers in 214 HS and IS samples, In the lowlands east of the Andes, the archaeological providing Y chromosome data (17 STRs and 16 SNPs record is much less detailed, frustrated also by poorer diagnostic for assigning haplogroups) and mtDNA data visibility and preservation in the Amazonian rainforest. (control region sequences and 3 SNPs and one INDEL Certainly, the linguistic panorama is unusually frag- diagnostic for assigning haplogroups). The aims of our mented and diverse. Four major language families are research are to: (1) characterize the genetic diversity of spoken—Arawak, Carib, Tupı, and Je— but in scattered the Yanesha population, and uncover potential differen- distributions and interspersed with dozens of other ces between the HS and IS communities; (2) estimate minor families and isolate language lineages not demon- the degree of any sex-biased patterns of contact; (3) strably relatable to any other (Campbell, 1997). Arawak investigate the origin of the Yanesha population, by dis- nonetheless ranks as the most widely dispersed family tinguishing Amazonian, Andean, and European gene in the Americas, scattered from the Antilles to pockets flows; and (4) explore a putative genetic signature for even in northernmost Argentina. Various competing Quechua and Arawak speakers in South America. hypotheses disagree on where the family originated, when it began to expand,