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Chapter 10

Who Were the Nasca? Population Dynamicsin Pre-Columbian Southern Peru Revealedby 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 relevantinformation about the structure of prehistoric populations. The data

obtained help to answer questions related to migration processes and popu-

lation 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 matri-

lineal 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 differentia-

tion of ancient coastal and highland populations in southern Peru is detect-

able as are changes in the mitochondrial haplogroup distribution patterns asa 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, HistoricalAnthropology and Humanecology, Georg-August-University Goettingen,

Bu rgerstrae50, 37073 Gottingen, Germany

e-mail: [email protected]

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

159

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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 popula-

tion structure, for example, by migration. The progression of archaeologicaltheories 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 craftstyles, 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 biologi-

cal 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 longerterm 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 recentpopulations 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 genealogiesfrom individuals buried in prehistoric burial grounds as shown for the Bronze

Age Lichtenstein Cave near Osterode, Germany (Schilz 2006) or species deter-

mination from prehistoric animal remains (Renneberg et al. this volume).

160 L. Fehren-Schmitz et al.

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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 essentialneed 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 gen-

erally 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 withrelatively 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

10 Who Were the Nasca? 161

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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-ChromosomeConsortium 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 diver-

sity. 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 mt-

Haplotypes 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 large-

scale, 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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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 archae-

ological 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 anddisappearance 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

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)


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raised new questions regarding population relationships and population dis-

continuities 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 elite-

dominance 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 bychromosomal 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 high-

throughput 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 cir-

cular 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 chromoso-

mal 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 (Merriwetheret al. 1995). This twofold determination system for mt-haplogroups was devel-

oped to authenticate the results because possible fast-evolving mutational hot-

spots 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 real-

time PCR was developed. The mitochondrial HybProbe multiplex PCR on the

LightCycler 2.0TM (Roche) allows the simultaneous amplification and determi-

nation (by melting curves analysis) of all four polymorphisms characterising the

Fig. 10.2 Schematic representation of the human mitochondrial DNA genome with thelocation 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 real-

time 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 calcu-

lated (Tajima 1993; Nei 1987). Genetic distances between the populations werecalculated 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 popu-

lations 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 macro-

scopic 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 chronologi-cal 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 bone-

powder 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 compar-

ison (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 para-

meters, 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

Period / Culture Phase PernilAlto

Mollake

Chico

Jauranga

LosMolinos

LaMuna

Hanaq

Pacha

Monte

Grande

Paracas

Peninsula

Pacapa-

ccari

LIP 8

16

MH

1

5 2

LATE

MIDDLE

7

19

821

NASCA

EARLY15

4

27

LATE

MIDDLE12

48

PARACAS

EARLY 510

INITIALPERIOD

1ARCHAIC PERIOD


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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 reproduciblydetermine 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 poorerDNA preservation in the Palpa area may be explained by higher temperature

and soil pH-value, but also the chronologically changing environmental condi-

tions (Eitel and Ma chtle 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 asso-

ciated. Individuals from Los Molinos and La Mun a 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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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 ofthem all distinct haplotypes different from the founder types appear syn-

chronous in populations from different archaeological sites, cf. one in the

Jauranga (ParacasPalpa) and the ParacasPeninsula population. Ten haplo-

types persist over more than one time period and appear in different popula-

tions from the Palpa sites (cf. Fehren-Schmitz 2008). All analysed populations

have a high degree of genetic diversity as shown, for example, in the mitochon-

drial 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 statisticanalyses (not shown here) verify this conclusion and also that the high frequen-

cies 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 dis-

tances 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 therecent 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).

Table 10.2 Mitochondrial haplogroup frequencies and haplotype diversity (HD) for the

Analysed pre-columbian populations

Haplogroup Frequency

Group / Population na Hb A B C D Hdc

Paracas (Peninsula) 10 (6) 5 0,00 0,00 0,30 0,70 0,93Paracas (Palpa) 28 (25) 15 0,07 0,00 0,14 0,79 0,95

NascaRural (Palpa) 37 (30) 26 0,02 0,11 0,22 0,65 0,98

NascaUrban (Palpa) 28 (25) 17 0,00 0,18 0,43 0,39 0,97

Middle Horizon (Palpa) 11 (6) 6 0,00 0,27 0,36 0,36 1,00

Pacapaccari (Highlands) 16 (12) 6 0,00 0,69 0,31 0,00 0,86

a 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.


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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 processesof 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 con-

stant population in the Palpa area persisting into the Nasca period. Thesefindings 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 assump-

tion is supported by the fact that individuals from the elite burials of La Mun a

share mt-haplotypes with the rural populations. The emergence of haplogroup

B and the differences between the rural and urban populations suggest thatthere 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

Fig. 10.4 NJ-Tree based on the pairwise FST-distances of the analysed pre-Columbian popu-lations (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


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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 popula-

tion structure changes slightly in the Nasca period, and the urban Nasca

population differs from the rural can be best explained by the immigration offoreign 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 valleysas a consequence of climatic aggravations (Eitel and Ma chtle 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 fre-

quency of haplogroup B in recent indigenous Peruvian populations suggests

that the observed genetic changes are best explained by the expansions of thevast-reaching highland empires, especially the Inca. This assumption is sup-

ported 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 popu-

lations 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 pre-

Columbian 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 pat-

terns of the populations that settled in the Andean highlands were also affected

by terms of selective forces. It is imaginable that the harsh environmentalconditions 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, knowl-

edge 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.


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