Chapter 10

Who Were the Nasca? Population Dynamics in Pre-Columbian Southern Peru Revealed by Ancient DNA Analyses
Lars Fehren-Schmitz, Susanne Hummel and Bernd Herrmann

Abstract Through the analysis of ancient DNA from human mortal remains it is possible to gain access to a biohistoric archive containing relevant information about the structure of prehistoric populations. The data obtained help to answer questions related to migration processes and population relationships that could not be answered by the methods of cultural science alone. The aim of this study was to show to what extent the cultural evolution of the southern Peruvian Palpa area was accompanied by processes of population exchange. Bone and tooth samples of over 200 individuals from prehistoric burial grounds from southern Peru were collected and examined with the methods of ancient DNA analysis. The study focuses on the matrilineal population dynamics by the analysis of mitochondrial genetic markers. Mitochondrial haplogroups and types could be successfully determined for over 100 individuals from different archaeological periods. The obtained data were compared with mitochondrial data from recent Native American populations. The results allow us to describe to what extent cultural changes were influenced by allochthonous contributions to the gene pool and how changes in the socioecological complexity of the cultures affected the genetic composition of the Palpa valley population. Also, a significant differentiation of ancient coastal and highland populations in southern Peru is detectable as are changes in the mitochondrial haplogroup distribution patterns as a result of the emergence of the extensive highland empires in later South American prehistory.

L. Fehren-Schmitz (*) Johann Friedrich Blumenbach Institute of Zoology and Anthropology, Historical Anthropology and Humanecology, Georg-August-University Goettingen, Burgerstrae50, 37073 Gottingen, Germany e-mail: lfehren@gwdg.de

M. Reindel, G.A. Wagner (eds.), New Technologies for Archaeology, Natural Science in Archaeology, DOI 10.1007/978-3-540-87438-6_10, Springer-Verlag Berlin Heidelberg 2009

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10.1 Introduction
One of the recurrent questions of archaeological science is how far cultural change in a geographic region is connected to changes in the biological population structure, for example, by migration. The progression of archaeological theories from cultural history, processual archaeology, and postprocessual archaeology to the present developments were always accompanied by changes in the perception of migration as a potential factor in cultural evolution (Anthony 1990). There are many possible scenarios for the cause of cultural changes reaching from pure autochthonous developments or cultural diffusion with a stable population to culture change through population replacements by processes of migration or invasion (Burmeister 2000). Although changes in the material culture can be detected in the archaeological record by typological comparison, methods of cultural science allow no definite decision if new craft styles, ritual behavior, and so on were introduced by immigrating people or other mechanisms of diffusion. The only archive that can be used to resolve this question is humans themselves. Through the use of molecular genetic methods it is possible to gain access to a nonbiographic archive containing all relevant information about the biological composition of a population. Uniparental inherited genetic markers such as mitochondrial DNA (mtDNA) and the nonrecombining proportion of the Y-chromosome (nryDNA) have proven valuable for the reconstruction of the global spread of Homo sapiens and therewith the understanding of longer term global patterns of human diversification (Underhill and Kivisild 2007). Analyses of the maternal inherited mtDNA and paternal inherited nryDNA from recent populations were successfully used to throw light on the processes source populations, number of migrants, migration dates, routes, and so on accompanying the initial colonisation of the Americas (e.g., Torroni et al. 1993), Europe (e.g., Richards et al. 1998), and Australia (e.g., Hudjashov et al. 2007). Comparisons of the data from both genetic markers also allowed the analysis of sex-specific mobility patterns and therewith human migrational behavior (e.g., Wilder et al. 2004). All those studies use DNA from recent populations to reconstruct historic migration processes by utilising methods and calculations of population genetics. Through the development of methods and techniques to analyse ancient DNA (aDNA) from pre-/historic specimen physical anthropology gained an analytic tool that also allows the access to such genetic data from populations and cultures that vanished long ago. Instead of deducing historic processes from a recent genetic as-is state, these methods allow a diachronic comparison of the genetic relationship of ancient populations. Furthermore the analysis of ancient DNA allows, for example, the reconstruction of complex genealogies from individuals buried in prehistoric burial grounds as shown for the Bronze Age Lichtenstein Cave near Osterode, Germany (Schilz 2006) or species determination from prehistoric animal remains (Renneberg et al. this volume).

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To conduct such investigations, methods and techniques developed for the analysis of DNA from living organisms have to be adapted to the specific characteristics of aDNA, DNA degradation patterns have to be examined, and methods of contamination control and prevention have to be improved. So the constant development of new methods and techniques is an essential need for palaeogenetic science. Detailed information on possible applications, methods, specifics, and possible problems faced with the analysis of aDNA are compiled in some review articles (e.g., Paabo et al. 2004; Willerslev and Cooper 2005) and two monographs published by the Palaeogenetics Group of the Department of Historic Anthropology, Gottingen (Herrmann and Hummel 1995; Hummel 2003).

10.1.1 The Peopling of South America from a Genetic Perspective

Important insights into the initial colonisation of the New World have been gained through molecular genetic studies particularly through the analyses of mtDNA of recent Asian and Native American populations. Today it is generally agreed that the first humans entered the Americas through or along the Beringian land bridge. Controversy arises when it comes to the timing, the precise route, and the number of major migrations. The most parsimonious model by todays state of knowledge from genetic data is a single initial migration with relatively few individuals about 20,00014,000 years ago along a Pacific coastal route (Merriwether et al. 1995; Silva et al. 2002; Schurr 2004; Lewis et al. 2007). The assumption of rapid spread of the people along the Pacific coast from Beringia to southernmost South America and therewith a pre-Clovis colonisation of the Americas is consistent with archaeological data (Dixon 2001) and the assured early 14C dates of South American archaeological contexts such as Monte Verde, Chile (Dillehay 1999). Demographic simulations also prove that even for a small founding population by this route it would be possible to cover such a distance Alaska to Tierra del Fuego in relatively short time (e.g., 1000 years) under retention of an effective population size (Fix 2002). Despite the richness of their cultures and the richness of environments that they inhabit, the Native Americans harbor a low level of genetic diversity. This is probably a consequence of the previously described terms of the initial peopling. Nearly all Native American mtDNA-haplotypes belong to four ancestral lineages, the mt-haplogroups labled A, B, C, and D (Torroni et al. 1993). These lineages are widely found throughout the Americas, but there is a lot of variation in frequencies among populations and geographic regions. A fifth founding mitochondrial haplogroup, designated X, is only found in indigenous populations of northern North America (Dornelles et al. 2005). All of those five major matrilineages (mt-haplogroups) were represented by only one (Schurr 2004) or a few (Tamm et al. 2007) sublineages (mt-haplotypes) in the initial founding population. The mt-haplogroups have a definite Asian

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ancestry and the genetic data indicate that the ancestral source population probably originated in south-central Siberia from where it migrated to Beringia and then into the New World (Schurr 2004). In the case of the Y-chromosomal DNA most male Native Americans belong to the two principal founding lineages C and Q (nomenclature: Y-Chromosome Consortium 2002). Haplogroup Q dominates with about 75% and for South America especially the subhaplogroup Q-M3 can be found, a group not existent outside the Americas (Underhill et al. 2001). Although understanding of the further peopling of North America has significantly progressed in the last years this is not the case for South America. There is compelling archaeological and genetic evidence to suggest that the continent was peopled by 1413,000 years ago (Dillehay 1999; Fuselli et al. 2003) but there is no agreement regarding the number of initial migrations and the migration routes (Lalueza et al. 1997; Rothhammer et al. 2001; Keyeux et al. 2002; Lewis et al. 2007). South America has a unique pattern of genetic diversity. Western (Andean) populations show a higher level of within-population diversity but short genetic distances to each other when compared to eastern (Amazonian) populations (Fuselli et al. 2003; Lewis et al. 2005). There is a very specific regional distribution of mitochondrial haplogroup frequencies (Fig. 10.1; Fehren-Schmitz 2008) and a high frequency of mtHaplotypes that are unique and not shared between different regions. Recent genetic comparison of modern Native American populations from all over South America based on mitochondrial Hyper Variable Region I (HVR I) sequences show that although there are regional differences in the patterns of genetic variation the low overall variance among these regions gives no evidence for several migrational waves (Lewis et al. 2007). Even if this makes it most parsimonious that the continent was peopled by one founding population, the exact routes, and if this wave split in different groups when passing the Isthmus of Panama remains uncertain. This lack of knowledge partly can be attributed to the circumstance that all available hypotheses regarding these questions are based on the analysis of recent Native American populations. There are only a few aDNA studies from South America with reliable results (e.g., Shimada et al. 2004; Moraga et al. 2005; Shinoda et al. 2006). Unfortunately most of these studies have neither a sample size that allows relevant population genetic calculations nor diachronic developments of settlement areas. The palaeogentic investigations conducted within the NascaPalpa project are the first largescale, diachronic aDNA studies for South America.

10.1.2 Aims and Goals of the Palaeogenetic Investigations of Human Remains from the Palpa Area
One of the aims of the study presented here was to prove that large-scale ancient DNA investigations can be used to reveal complex population dynamics in

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Fig. 10.1 Recent regional distribution of mitochondrial haplogroup frequencies in native South American populations. For this illustration a reduced dataset is used. Refer to Fehren-Schmitz (2008) for a more detailed distribution map. (Data used: Torroni et al. 1993; Merriwether et al. 1995; Ward et al. 1996; Rickards et al. 1999; Moraga et al. 2000; Goicoechea et al. 2001; Keyeux et al. 2002; Fuselli et al. 2003; Lewis et al. 2005; Torres et al. 2006; Garcia et al. 2006; Lewis et al. 2007)

prehistoric settlement areas, and prove the existence of migration processes and their influences on culture as well as cultural influence on migration patterns. The Palpa area in southern Peru and the scientific conditions of the Nasca Palpa project offered the perfect conditions for such investigations. The archaeological evidence shows settlement continuity for this area until now, beginning with the Archaic period (about 3800 BC). In this long period of time the region faced many more or less dramatic cultural changes and the emergence and disappearance of archaeological cultures such as the Paracas and Nasca (Reindel this volume). The new insights into the cultural and ecological history of the southern Peruvian coast that arose from the project at the same time

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raised new questions regarding population relationships and population discontinuities or continuities, for instance:

 Was the Paracas tradition in southern Peru represented by a biological

uniform population or by solitary local groups sharing cultural traits (Silverman and Proulx 2002)?  Did the Nasca culture evolve from the Paracas tradition with a continuous population? Or was it introduced to the region by foreign culture-bearers?  Are there any biological traces of Wari influence in the Palpa region at the end of the Nasca period that justify the assumption of invasion or elitedominance scenarios as formulated by Allison (1979)? In addition to those questions that focus on the regional population dynamics in southern Peru the investigations should help to shed light on the peopling of western South America. As mentioned above there is a constant need for fundamental research, for example, to understand the degradation processes of DNA and to enhance the methods of authentication and contamination prevention. To answer the content-related questions new molecular assays had to be developed. For instance, real-time PCR technology was adapted to the specific requirements of aDNA research in the course of this project. The adoption of this technology and thereby development of new methods for low-copy DNA quantification and inhibition control (Westenthanner et al. in prep) allowed novel insights to differential DNA preservation in human bones (Fehren-Schmitz 2008) and by chromosomal topography. Additionally real-time PCR technology was used to develop new, faster SNP (single nucleotide polymorphism) detection assays for mitochondrial and Y-chromosomal haplogroup determination, allowing highthroughput analysis for aDNA.

10.2 Material and Methods 10.2.1 Molecular Genetic Markers and Analysis Systems
The studies presented here where mainly based on the analysis of mtDNA, and therewith the matrilinear population dynamics. Mitochondrial DNA is a circular double-stranded molecule present in the mitochondria of eukaryotic cells. Each cell contains hundreds to thousands of copies of mtDNA making it much more likely that even after the DNA has undergone degradation processes following death there are more mitochondrial DNA fragments preserved in human skeletal remains than chromosomal DNA. MtDNA is exclusively maternally inherited, lacks recombination, and evolves faster than chromosomal DNA (Pakendorf and Stoneking 2005). These characteristics enable us to trace related maternal lineages nearly unchanged back through time and make it the molecule of choice for phylogenetic and population genetic studies, most notably when analysing aDNA.

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To determine the mitochondrial haplotypes we analysed a 388 bp fragment of the mitochondrial HVR I (np1602116408). A modular analysis system consisting of eight primers generating overlapping PCR products was designed. The modular character of this system allows the amplification of fragments with different sizes: long (434 bp), medium (236261 bp), and short (157180 bp). Therewith it is possible to adapt the analysis to the specific DNA degradation grades encountered in different samples. The generated PCR products were further analysed by direct sequencing. In addition to the determination of the mitochondrial haplogroups by the specific HVR I polymorphisms we analysed four specific polymorphisms of the mitochondrial coding region determining the groups A, B, C, and D (Fig. 10.2). Three of the groups A, C, and D are characterised by SNPs (single base transversions or transitions) and group B is characterised through a 9 bp deletion at nucleotide position (np) 82728280 of the mitochondrial genome (Merriwether et al. 1995). This twofold determination system for mt-haplogroups was developed to authenticate the results because possible fast-evolving mutational hotspots of the HVR I and miscoding lesions caused by postmortem DNA damage exhibit a high risk of false determinations (Meyer et al. 1999; Willerslev and Cooper 2005; Gilbert et al. 2007). For the analysis of the four coding region polymorphisms a hybridisation probe-based multiplex PCR assay for the realtime PCR was developed. The mitochondrial HybProbe multiplex PCR on the LightCycler 2.0TM (Roche) allows the simultaneous amplification and determination (by melting curves analysis) of all four polymorphisms characterising the

Fig. 10.2 Schematic representation of the human mitochondrial DNA genome with the location of the three base substitutions and the 9 bp deletion determining the Native American haplogroups. For each haplogroup the nucleotide position (e.g., np 663) and realised base substitution (e.g., A) is mentioned. The labels inside the circle specify the respective functional region (genes) of the mitochondrial genome

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four Native American mt-haplogroups. Conventional methods used to determine such polymorphisms consist of several analysis steps and therewith are more time-consuming whereas the developed assay is a one-step procedure. In addition to the mt-haplogroups/-types also investigations on Y-chromosomal haplogroups were made. The Y-haplogroups are also characterised by SNPs (cf. Y-Chromosome Consortium 2002). A HybProbe multiplex assay for the realtime PCR was developed to determine the four most frequently occurring Y-chromosomal haplogroups in Native American populations: C, P, Q, and Q3 (as mentioned above). SNPs analysed were M130 (C>T) for C, M45 (G>A) for P, M242 (C>T) for Q, and M3 (C>T) for haplogroup Q3. All analysis systems were tested on modern DNA from European and South American peoples first before being applied to the aDNA analysis. For all analysed populations standard genetic diversity indices were calculated (Tajima 1993; Nei 1987). Genetic distances between the populations were calculated on haplogroup and haplotype level. Calculations for the measures were performed using the Arlequin software package (version 3.1, Excoffier and Schneider 2005). The obtained HVR I sequence data were also used to calculate distance-based phylogenetic trees and mean nucleotide distances between populations by the Mega 4.0 software (Tamura et al. 2007).

10.2.2 Sample Material and Sample Preparation

In different field campaigns (20042006) bone and teeth samples from 216 skeletonised or mummified human individuals were collected (Fig. 10.3). 172 of these were from different archaeological sites in the Palpa area and 44 individuals were from pre-Columbian burial grounds outside this area: Monte Grande (near the coast to the west of Palpa); Paracas Peninsula (Paracas

Fig. 10.3 Skulls of three human individuals from different archaeological sites of the Palpa area (left: Los Molinos; right: Pernil Alto) showing the encountered average state of macroscopic preservation. Individuals are largely skeletonised with some soft tissue conservation through mummification

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Cavernas 6), and Pacapaccari (Andean highlands to the east of Palpa). The Palpa samples were collected from chronologically varying sites so that it was possible to investigate a timeframe from the late Archaic period (approx. 3800 BC) to the Middle Horizon (6501000 AD). Sites bearing burials from more than one archaeological phase were preferred. Sampled sites, their chronological classifications, and the number of individuals sampled from them for each chronological period are listed in Table 10.1. Sample preparation for DNA extraction followed a standardised protocol (Hummel 2003). Bone and teeth fragments were mechanically pulverised and afterwards chemical procedures for decalcification and cellysation of the bonepowder followed. Automated DNA extraction and purification were performed with an EZ1 Biorobot (Qiagen). For each sample two or more extracts were made. This precaution allows an authentication of analysis results by comparison (Hummel 2003). In addition to the aDNA investigations, mt-haplogroup (cf. Fig. 10.1) and HVR I sequence data of over 4000 Native American individuals from different South American populations were collected from the literature to compare the pre-Columbian Palpa populations with the whole continent. Only a part of the obtained data and comparisons is presented here. For further information concerning the used extraction methods, primer sequences, analysis parameters, and statistic parameters, for example, refer to Fehren-Schmitz (in prep.). In the context of this text it is only possible to present a fractional

Table 10.1 Distribution of individuals sampled (number) tabulated by site and dating
Los Molinos Pernil Alto La Muna Peninsula Jauranga

Mollake

Paracas

Grande

Monte

Pacha

Pacapa-

Hanaq

Chico

Period / Culture

Phase

LIP

15

NASCA

LATE

MIDDLE

21

19

MH

15

EARLY

EARLY

INITIAL PERIOD

ARCHAIC PERIOD

10

48

MIDDLE

12

PARACAS

LATE

27

16

ccari

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amount of all generated genetic data, used datasets, population statistics, and other details of the conducted studies.

10.3 Results and Interpretation 10.3.1 Molecular Analysis Results and Data Evaluation
Despite recently formulated hypotheses that the ecological conditions of desert environments allow a maximum preservation time for DNA of approximately 700 years (e.g., Marota et al. 2002) the methods used and developed in the context of this study proved successful in the recovery of DNA from much older bone material stored in desert soils. Overall it was possible to reproducibly determine the mt-haplogoups of 131 individuals and mt-haplotypes (complete 388 bp HVR I sequence) of 105 individuals. Amplification of Y-chromosomal DNA was only successful for six individuals (all Y-haplogroup Q-M3), five from the site Pacapaccari (Andean highlands) and one from Jauranga (Palpa). Generally it can be stated that the grade of DNA preservation found in the individuals of the highland Chullpas proved to be better than in the individuals from desert burials. The stable level of humidity and temperature encountered in the Chullpas can be best compared to storage conditions in caves, which proved to be very good for DNA preservation (Burger et al. 1999). The poorer DNA preservation in the Palpa area may be explained by higher temperature and soil pH-value, but also the chronologically changing environmental conditions (Eitel and Machtle 2006) and therewith storage conditions of the region. For population statistics the data obtained from the different archaeological sites were grouped by time period and region (e.g., ParacasPalpa and Paracas Paracas Peninsula). For the Nasca period two extra divisions were made based on the socioeconomic character of the sites with which the individuals were associated. Individuals from Los Molinos and La Muna belong to the group Nasca urban whereas all other individuals from sites such as Jauranga or Hanaq Pacha are grouped as Nasca-rural (Table 10.1). All successfully analysed individuals belong to one of the four Native American haplogroups A, B, C, and D (see above). The distribution of the mt-haplogroups for the analysed populations is found in Table 10.2. For both Paracas populations there are very high frequencies of haplogroup D followed by haplogroup C with a much lower fequency. This dominancy of D persists into the Nasca time as seen with the Nasca (Palpa-rural) population (Table 10.2). For the urban Nasca population, however, D and C were determined with nearly equal frequencies. Parallel to the decrease of D there is an increase of haplogroup B which emerges first in the Palpa area with the Nasca period. This trend continues with the transition to the Middle Horizon (cf. Table 10.2). The frequency distributions of the Palpa populations differ significantly from the highland population of Pacapaccari. Here B is definitely the dominating group and C the only other

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Table 10.2 Mitochondrial haplogroup frequencies and haplotype diversity (HD) for the Analysed pre-columbian populations Group / Population Paracas (Peninsula) Paracas (Palpa) NascaRural (Palpa) NascaUrban (Palpa) Middle Horizon (Palpa) Pacapaccari (Highlands)
a

na 10 (6) 28 (25) 37 (30) 28 (25) 11 (6) 16 (12)

Hb 5 15 26 17 6 6

Haplogroup Frequency A B C 0,00 0,07 0,02 0,00 0,00 0,00 0,00 0,00 0,11 0,18 0,27 0,69 0,30 0,14 0,22 0,43 0,36 0,31

D 0,70 0,79 0,65 0,39 0,36 0,00

Hdc 0,93 0,95 0,98 0,97 1,00 0,86

n = Number of individuals with successful haplogroup determination; the number of individuals with sucessfully determined mt-haplotypes follows in brackets. b H = Number of determined different haplotypes in the population. c Hd = Haplotype diversity (Nei 1987). The value relates to the HVR I sequence data not to the haplogroup data.

detected (Table 10.2). The frequencies of haplogoup D of all pre-Colombian coast populations analysed in the context of this study are much higher than in the recent indigenous Peruvian populations and those of the other northern and southern central Andean regions (Fig. 10.1). The HVR I sequence data from the 105 successfully typed individuals could be assigned to 57 mitochondrial haplotypes of which 32 were single-haplotypes. Twenty-five haplotypes could be detected in more than one individual. Four of them all distinct haplotypes different from the founder types appear synchronous in populations from different archaeological sites, cf. one in the Jauranga (ParacasPalpa) and the ParacasPeninsula population. Ten haplotypes persist over more than one time period and appear in different populations from the Palpa sites (cf. Fehren-Schmitz 2008). All analysed populations have a high degree of genetic diversity as shown, for example, in the mitochondrial Haplotype-Diversity (Hd, Table 10.2). Even though there is a diachronic and group specific change in the haplogroup frequencies, there is no increase of overall mitochondrial genetic variability. Other conducted population statistic analyses (not shown here) verify this conclusion and also that the high frequencies of group D cannot be explained in terms of genetic isolation. Conducted genetic distance calculations based on the HVR I sequence data support the considerations derived from the haplogroup data. Very low distances can be determined between both Paracas period populations and the rural Nasca group. These three populations cluster in a genetic distance-based neighbour joining (NJ) tree (Fig. 10.4). The urban Nasca and Middle Horizon populations exhibit a higher distance to the other three pre-Columbian coast populations but still cluster within the same branch of the tree whereas the recent indigenous populations of Peru (data: Fuselli et al. 2003; Lewis et al. 2005) and the pre-Columbian Andean highland site of Pacapaccari form a distinct branch of the tree (Fig. 10.4).

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Fig. 10.4 NJ-Tree based on the pairwise FST-distances of the analysed pre-Columbian populations (circles) and four recent Native American populations (triangles) from Peru. The calculations are based on HVR I data. The tree is drawn to scale, with branch lengths in the same units as those of the evolutionary distances used to infer the phylogenetic tree

10.3.2 Ancient DNA Data Translated to the Population History of South Peru
Combined with knowledge of the cultural and ecological history the population genetic data can be translated to reconstruct the population dynamic processes of the Palpa region for the investigated timeframe. The genetic similarity of the Paracas populations from the Palpa area and the peninsula, and also the low genetic distance between both allows the conclusion that the Paracas culture of the southern Peruvian coast formed a uniform population. The results also show that there is a high gene flow, thus migration frequency, within the distribution area of the Paracas culture but no significant genetic exchange with the surrounding populations, for example, from the Andean highlands. Distance calculations and distribution patterns also show that there is a constant population in the Palpa area persisting into the Nasca period. These findings combined with archaeological evidence such as parallels in ornaments and the existence of early geoglyphs (Reindel and Gruen 2006) suggest that the Nasca culture of this area evolved from the bearers of the Paracas culture. Even though the urban and rural Nasca populations exhibit a degree of genetic distance there is no evidence that this results from foreign influences, as it would be expected with an elite dominance migration scenario. This assumption is supported by the fact that individuals from the elite burials of La Muna share mt-haplotypes with the rural populations. The emergence of haplogroup B and the differences between the rural and urban populations suggest that there is a higher amount of genetic exchange with populations from outside the investigated coast area, maybe the highlands. This might be a result of the increase of socioeconomic complexity in the Nasca period which concurrently

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would have caused an increase of the migrational pull-factor of this area. A grade of social complexity as is reached in the Nasca culture requires craft and administration specialists (Service 1962). The fact that the genetic population structure changes slightly in the Nasca period, and the urban Nasca population differs from the rural can be best explained by the immigration of foreign specialists into the urban settlements. At the same time these results verify the urban, respectively, central character of this settlements. There are no significant genetic changes from the Nasca period to the Middle Horizon and therewith no evidence for allochthonous contributions to the Palpa gene pool. Invasion or colonisation scenarios can be rejected as a cause for the demise of the Nasca culture in this area. The decrease of population density in the Late Nasca period and the Middle Horizon contrasted with the constantly high mt-haplotype diversity fits best with scenarios suggesting that the main settlement area of the people shifts eastwards into the Andean valleys as a consequence of climatic aggravations (Eitel and Machtle 2006). The pre-Columbian populations of the southern Peruvian coast significantly differ from the recent Peruvian and ancient highland populations although they are very similar to the recent Native American populations of Middle and South Chile (cf. Fig. 10.1; Fehren-Schmitz 2008). Mitochondrial distribution patterns of todays coastal Peru therefore have to be a result of cultural and political processes succeeding the time span investigated in this study. The high frequency of haplogroup B in recent indigenous Peruvian populations suggests that the observed genetic changes are best explained by the expansions of the vast-reaching highland empires, especially the Inca. This assumption is supported by the affinity of the recent Chilean populations to the pre-Columbian Peruvian coastal populations. The maximum southward geographic extension of the Inca empire is congruent with the recent border between populations exhibiting mitochondrial distribution patterns similar to recent Peruvian populations and Chilean populations that exhibit the ancient coastal patterns (Fig. 10.1).

10.4 Future Prospects

The conducted investigations and conclusions show that the use of the archive ancient DNA allows insights into prehistoric population dynamics that could not be achieved by the analysis of archaeological sources or the use of modern DNA alone. Although it was possible to gain novel information about preColumbian colonisation and migration processes in western Southern America, especially the southern Peruvian coastal regions, there is still no information about equal processes in the Andean highlands. The only available highland population datasets (this study; Shinoda et al. 2006) date into the Middle Horizon or the Inca period. To reveal the colonisation and migration processes that took place in this second major cultural area of western South America,

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studies have to be conducted involving populations dating into the earlier time periods especially those analysed in the present studies. Furthermore it should be investigated how far the genetic distribution patterns of the populations that settled in the Andean highlands were also affected by terms of selective forces. It is imaginable that the harsh environmental conditions physiological stressors such as hypoxia (Moran 2000) of the highlands and therewith the necessary adaptive reactions also had influence on the genetic structures of the populations (Fehren-Schmitz 2008). To answer these questions new genetic markers have to be explored and analysis systems suitable for aDNA analysis have to be developed. As mentioned above, knowledge about the colonisation history of South America is still not satisfying. The presented studies show that one method of resolution for this could be the genetic analysis of more pre-Columbian populations. The analysis of recent native South American populations alone will not solve the problem.