Population Structure during the Demise of the Moche (550–850 AD): Comparative Phenetic

Analyses of Tooth Trait Data from San José de Moro, Perú

Author(s): Richard C. Sutter and Luis Jaime Castillo
Source: Current Anthropology, Vol. 56, No. 5 (October 2015), pp. 762-771
Published by: The University of Chicago Press on behalf of Wenner-Gren Foundation for
Anthropological Research
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 762
Population Structure during the Demise of
the Moche (550–850 AD)
Comparative Phenetic Analyses of Tooth Trait Data
from San José de Moro, Perú
Richard C. Sutter and Luis Jaime Castillo
Department of Anthropology, Indiana University–Purdue
University, Kettler Hall G11A, Fort Wayne, Indiana 46805,
USA (sutterr@ipfw.edu)/Departamento de Humanidades–
Sección Arqueología, Pontificia Universidad Católica del
Perú, Avenida Universitaria 1801, San Miguel, Lima, Peru.
This paper was submitted 22 VIII 14, accepted 26 III 15, and
electronically published 1 X 15.
CA1 Online-Only Material: Supplement A
Population structure analyses and biodistance comparisons
for Middle Moche (550–700 AD), Late Moche (700–850 AD),
and Transitional (850–950 AD) period skeletal samples from
San José de Moro with a previously reported Middle Moche
period sample from Pacatnamú, a nearby Jequetepeque Val-
ley site, and eight samples from Huaca de la Luna and Cerro
Oreja, two sites in the Moche Valley, suggest that substan-
tial levels of extralocal gene flow into the Jequetepeque Valley
from the adjacent highlands to the east occurred during the
demise of the Moche polity there. This process strongly cor-
responds to evidence for a temporal increase in exotic grave
offerings at San José de Moro during the Late Moche and
subsequent Transitional periods. The broader implications
of these results for our understanding of the demise of the
Moche are discussed.
Societal collapse has long been a preoccupation of anthro-
pologists as a natural outgrowth of an interest in complexity
(Railey and Reycraft 2008; Schwartz and Nichols 2006; Tainter
1988; Yoffee 2006). Likewise, an interest in past migrations and
their impacts remains a critical area of anthropological study.
When forced political resettlement is not the driving force, both
push and pull factors—including a people’s familiarity with the
region, the presence of kin, trade interests, and information—
may influence peoples’ decision to migrate (Anthony 1990; Bret-
tell 2008). Migrations also can provoke a number of economic,
cultural, and political changes in both the source and recipient
populations (Stanish et al. 2010)—including strains on local re-
sources and infrastructure—and existing ethnic identities may
either be intensified or lose their salience (Barth 1969). Migra-
tory frontiers are typically active locations for social interac-
tions that result in culture change (Schortman and Urban 1994:
q 2015 by The Wenner-Gren Foundation for Anthropological Research.
All rights reserved. 0011-3204/2015/5605-0008$10.00. DOI: 10.1086
/683269
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Current Anthropology Volume 56, Number 5, October 2015
44), especially for understanding the impact of population dis-
persals’ influence(s) on the local indigenous culture of recipi-
ent populations (Baker and Kealhofer 1996; Newson 1976; Pels
1997).
How, then, can we best identify prehistoric migrations and
their consequences? Concomitant with archaeological infor-
mation, bioarchaeological data are critical to our understand-
ing of how and why people may have chosen to migrate as
well as what the social, cultural, and political implications
of migration were. For this paper, we focus on population
structure analyses and highlight both survey and excavation
results for the Jequetepeque Valley, Perú, in order to un-
derstand the population dynamics and varied sociopolitical
processes during and immediately following the demise of
the Moche within the region. We suggest that the varied
local responses observed during the Late Moche and subse-
quent Transitional periods resulted from the unique nature
of the preceding developments during the Middle Moche
Period as well as external migratory influences during a pe-
riod of environmental degradation. In addition to providing
a brief overview of the preceding Moche era and its subse-
quent decline, we highlight survey results and bioarchaeo-
logical data for the Jequetepeque Valley.
Archaeological Background
The Moche (200–850 AD) were a number of complex socie-
ties that likely represented nascent states or shifting confed-
erations of competing complex polities (Castillo 2001, 2003;
Castillo and Quilter 2010; Castillo and Uceda 2008) within
the coastal valleys of the north coast of Perú (fig. 1; see also
fig. A1; figs. A1–A12 in CA1 online supplement A). While
the nature of these polities and their degree of integration
remains the source of continued investigations, general con-
sensus suggests that there were two independent spheres of
influence: one in the north (Piura to Jequetepeque valleys) and
one in the south (Chicama to Huarmey valleys). Sociopolitical
developments in the southern region likely resulted from the
expansion of a multivalley state with its seat of power in the
Moche Valley. At the same time, the Jequetepeque Valley was
characterized by sociopolitical fragmentation of competing
Moche polities, each with their own independent irrigation
canals and fortified settlements (Castillo 2001; Castillo and
Quilter 2010; Castillo and Uceda 2008; although see Quilter
and Koons 2012 for a dissenting view).
For the Jequetepeque Valley, the cultural sequence is sim-
ilar to that reported for the southern Moche, although some
important differences exist. Castillo (2003) has developed a
comparative three-phase Moche ceramic chronology for the
Jequetepeque Valley that broadly corresponds to the tradi-
tional five-phase ceramic chronology developed for the south-
ern Moche (fig. 2; see also fig. A2). Moche settlement in lower
Jequetepeque during the Early Moche period (200–550 AD) was
primarily focused along the banks of the alluvial plain until
Sutter and Castillo Population Structure during the Demise of the Moche (550–850 AD) 763

Figure 1. Moche regions of influence (after Castillo and Quilter 2010:3).

the construction of new irrigation canals during the Middle
Moche period (550–700 AD), when settlement expanded first
to the south and later into the northern sections of the valley
(Castillo 2010; Castillo et al. 2008; Eling 1987; see also fig. A3).
The Late Moche period (LMP; 700–850 AD) is associated with
the environmentally influenced demise of the Moche’s presence
throughout the north coast region and concurrent settlement
shifts inland near the uptakes of irrigation canals. Paleocli-
matic data point to an extended period of drought for the
region that is dated between 650 and 730 AD (Shimada et al.
1991; Thompson et al. 1985). Within the Jequetepeque Valley,
the LMP was characterized by political decentralization and
shifting alliances among competing polities that emerged
along divisions in the extant irrigation systems (Castillo 2010;
Castillo et al. 2008; Swenson 2004). During the LMP, the
southern Jequetepeque Valley settlements might have been
abandoned or only intermittently occupied as a result of dune
encroachment (Dillehay, Kolata, and Moseley 2004; Dillehay,
Kolata, and Swenson 2009; Eling 1987; Hecker and Hecker
1982), while there was a simultaneous proliferation of hin-
terland settlements, each with their own highly varied, local-
ized ritual platforms and feasting wares (Castillo 2000, 2003,
2007, 2009, 2010; Dillehay, Kolata, and Moseley 2004; Dille-
hay, Kolata, and Swenson 2009; Hecker and Hecker 1995:78;
Swenson 2004:726, 732–737; 2007). In some instances, these
settlements—typified by multiple fortifications and piles of
sling stones—were located at distances of up to 3 km from
associated unfortified farming hamlets and water sources,
leading Swenson (2004, 2007) to suggest that the fortified sites
represent hinterland refuges from internecine warfare, where

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 764 Current Anthropology Volume 56, Number 5, October 2015

Figure 2. Archaeological sites of the Jequetepeque Valley (modified from Rosas 2010:927–930, fig. 5.2).

local feasting, ancestor worship, and other Moche rituals oc-
curred during the LMP (see fig. A4). Simultaneous with these
changes, adjacent highland Cajamarca influence expands
eastward, with recent excavations at Cerro Chepén revealing a
clear highland Cajamarca presence in the Jequetepeque Valley
(Cusicanqui and Caramanica 2011; Rosas 2007, 2010; see
figs. A5, A6). Radiocarbon dates and the presence of LMP
Moche fineline vessels recovered from structures at the site
indicate that it was contemporaneous with the LMP Moche
settlements (Rosas 2010). The exact nature of the relation-
ship between LMP Moche and Cajamarca peoples at present
is, however, unclear.
Throughout this period of local variation, influence from
the ceremonial center at San José de Moro appears to have
increased. During the LMP, San José de Moro became the
preeminent ritual center of the valley focused on a cult of
priestess, as well as serving as the valley’s primary funerary
locus, and the sole producer of fineline ceramics (Castillo
2001, 2009; McClelland, McClelland, and Donnan 2007; see
fig. A7). Fineline depictions of earlier (i.e., Early and Middle
Moche) male entities dramatically decreased in frequency
during the LMP, while depictions of elite females associated
with the recurring sacrifice theme predominated. Funerary
offerings associated with elite female burials in the LMP at
San José de Moro indicate that a lineage of priestesses played
an important role in Moche mortuary feasting and sacrificial
traditions (Castillo 2001, 2006; see fig. A8). Through his
ongoing excavations at San José de Moro, Castillo has un-
covered more than 1,000 burials dating to the terminal Mid-
dle Moche period (∼550–700 AD), Late Moche (∼700–850

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Sutter and Castillo Population Structure during the Demise of the Moche (550–850 AD) 765

AD), and the Transitional (∼850–950 AD) periods. While clus-
ters of boot tombs associated with Middle Moche ceramics
typify burials of the Middle Moche period, the subsequent
LMP is characterized by tombs that often contain a new, lo-
cally produced style of Moche fineline wares (see fig. A9) and
imported styles from the central coastal Nievería, adjacent high-
land Cajamarca (see fig. A10), and southern highland Wari tra-
ditions (see fig. A11), among others (Castillo 2000, 2001, 2007,
2009). The Transitional period is associated with the demise
of the Moche polity at San José de Moro and the subsequent
emergence of the north coast Lambayeque cultural tradition.
During the Transitional period, Castillo (2000) hypothesizes
that an absence of centralized power at San José de Moro per-
mitted a diversification of cultural expression. The early Tran-
sitional period is marked by eclecticism, with local ceramics
blending with foreign styles from the central coast, adjacent
highlands, and distant Ayacucho region (see fig. A12). In terms
of both quantity and quality, the exotic wares and objects in-
cluded as funerary offerings at San José de Moro are unparal-
leled in the north coast region (Castillo and Uceda 2008).
We focus on biodistance analyses of San José de Moro re-
mains from the Middle, Late, and Transitional periods. Using
burials from the Middle Moche Period as a baseline, does
the increased evidence of extraregional wares in the burials
of LMP and Transitional periods at San José de Moro reflect
an increase in extraregional gene flow? We hypothesize that if
these contacts do reflect increases in extraregional gene flow
at San José de Moro, then this will be reflected by bioindica-
tors of gene flow examined by this study (i.e., R matrix
biodistances, FST values, and positive Relethford-Blangero re-
siduals, as described below).
Material and Methods
Human nonmetric dental trait data are used to derive esti-
mates of genetic distance and variation among skeletal sam-
ples from the Middle, Late, and Transitional periods at San
José de Moro and data for nine other previously reported
north coast skeletal samples (Lambert et al. 2012; Sutter 2009;
Sutter and Cortez 2005; Sutter and Verano 2007; table 1). Data
for 31 nonmetric tooth traits were collected using standard-
ized casts and descriptions (Turner et al. 1991).
In order to make dental trait scores reported here com-
parable to those reported in other studies, teeth were scored
according to the individual count method (Turner and Scott
1977). In cases where an individual exhibited asymmetry in
the expression of a given trait, the greatest level of expression
is used. Before conducting biodistance analyses, traits with
fewer than 10 observations per skeletal sample were elimi-
nated. This resulted in the retention of 16 tooth cusp and
root traits. Additionally, those individuals who were missing
observations for more than 25% of the remaining traits were
eliminated from analyses.
This study estimates levels of genetic diversity (FST; Releth-
ford 1996, 2003; Relethford and Blangero 1990; Relethford,
Crawford, and Blangero 1997; Williams-Blangero 1989a, 1989b;
Williams-Blangero and Blangero 1989) and genetic distances
between the samples being studied using the R (relational) ma-
trix method (Relethford 2003; Relethford and Blangero 1990;
Relethford, Crawford, and Blangero 1997), using a code written
by Lyle Konigsberg for the statistical program R 2.15.2. The R
matrix provides an estimate of sample variability and the de-
gree of similarity to other samples being compared (Relethford

Table 1. Skeletal samples analyzed by this study

Sample Abbreviation Period Sample size

Moche Valley:
Cerro Oreja:
Salinar COSAL 329–205 BC (pre-Moche)a 51
Gallinazo 1 COG1 200–115 BC (pre-Moche)a 91
Gallinazo 2 COG2 115 BC–AD 127 (Early Moche)a 54
Gallinazo 3 COG3 ∼127–200 AD (Early Moche) 47
Huaca de la Luna:
Urban Sector HLLUS ∼300–600 AD (Middle Moche) 27
Platforms PLATS ∼400–600 AD (Middle Moche) 42
Jequetepeque Valley:
Pacatnamú:
H45CM1 PACAT ∼400–600 AD (Middle Moche) 27
San José de Moro:
Middle Moche SJMMM ∼400–600 AD (Middle Moche) 34
Late Moche SJMMT ∼600–850 AD (Late Moche) 16
Transitional SJMTRANS ∼850–950 AD (Transitional) 16
Moche Valley:b
Huaca de la Luna:
Plaza 3C P3C ∼300–500 AD (Middle Moche) 41
Plaza 3A P3A ∼600–650 AD (Late Moche) 22
a
From Lambert et al. (2012).
b
Executed male prisoners.

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 766
and Blangero 1990). The R matrix is a matrix of average kin-
ship coefficients between populations (off the diagonal, or rij)
and within populations (on the diagonal, or rii). Therefore,
samples with positive rij values are genetically more similar,
while those with negative values are less similar than average
(Irish 2010:9).
R matrix analysis is also used to derive both phenetic dis-
tances (Mahalanobis’s distance)—which reveal the patterns of
genetic relatedness primarily due to gene flow and ancestral-
descent relationships among the samples—and estimates of
the effects of genetic drift and gene flow, or FST. Theoreti-
cally, those populations that frequently shared mates will be
characterized by smaller distances between them than those
populations that rarely shared mates, while populations that
frequently shared mates should have small FST values (pop-
ulations exhibiting no differences between them will have an
FST p 0) and comparisons between those that rarely shared
mates will have large FST values. The Mahalanobis’s gener-
alized distance (d 2) for binary epigenetic traits (Konigsberg
1990) is calculated as a biodistance measure between two skel-
etal samples i and j, which represents the minimum genetic
distance between two groups being compared (Williams-
Blangero 1989b). This distance is determined using the fol-
lowing equation:
dij2 p (zik 2 zjk )0 T 21 (zik 2 zjk ),
where, as Konigsberg (1990:60) explains, “zik represents the
threshold value corresponding to a trait frequency of pik for
trait k in site i, zjk is the threshold value for trait k in site j, and T
is a pooled tetrachoric correlation matrix between the k traits.”
Recent work (Nystrom 2006; Stojanowski 2003, 2005, 2009)
has also used intrasite phenotypic variability to provide esti-
mates of extralocal levels of gene flow, as indicated by the sign
of Relethford-Blangero residuals, whereby positive residuals
indicate greater extralocal gene flow and negative residuals
suggest decreased extralocal gene flow. Importantly, both trait
heritabilities and effective population sizes for the samples
must be estimated for R matrix analyses.
Once differences in effective population size are accounted
for, any changes in heterozygosity will reflect effects of migra-
tion and population history (Relethford 1996). Proportional
estimates of effective population sizes are necessary for bias
correction of the R matrix and are either estimated from historic
records (Relethford 2003; Relethford et al. 1997; Stojanowski
2003, 2005, 2009) or archaeologically derived estimates of pop-
ulation densities or site habitation areas (Nystrom 2006). Un-
biased estimates of both the R matrix and FST were then calcu-
lated using formula from Relethford et al. (1997). Effective
population sizes used by this study are based on relative differ-
ences from estimates of irrigable land for both the Moche and
Jequetepeque valleys (Wilson 1999) and settlement pattern
studies for the Moche (Billman 1996) and Jequetepeque valleys
(Castillo 2010).
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Current Anthropology Volume 56, Number 5, October 2015
R matrix analyses for this preliminary study were conducted
using a conservative heritability estimate of 1, which produces
the minimum FST (Relethford and Blangero 1990). While future
analyses will explore FST using different estimates of heritabil-
ity, previous studies have demonstrated that patterns of R
matrix analyses are largely unaffected by different heritability
estimates and bias corrections (Nystrom 2006; Stojanowski
2005). Significance tests for differences in FST were determined
from standard error calculations (Relethford et al. 1997).
Results
The results of both the uncorrected and unbiased (corrected)
Mahalanobis’s d 2 values are presented in table 2. Examination
of the matrix of tetrachoric correlations among the 16 re-
maining traits indicated that all intertrait correlations were
low (1 0.18) and nonsignificant. The matrix of unbiased in-
tersite distances was analyzed using multidimensional scal-
ing procedures (fig. 3). In both cases, the two-dimensional
solutions captured more than 99% of the total variation, and
the final stress of the solutions was low. Inspection of the
resulting multidimensional scaling solution reveals that the
Moche Valley samples from Cerro Oreja and the local Huacas
at Moche site (i.e., HLLUS and PLATS) cluster near the center
of the solution along with the contemporaneous Pacatnamú
and Middle Moche San José de Moro samples of the Jeque-
tepeque Valley, while nonlocal human sacrificial victims from
Huacas at Moche (P3C and P3A) and the Late and Transi-
tional period samples from San José de Moro are more dis-
tantly related. These results are similar to those for the nine
previously reported samples examined by this study (Sutter
and Cortez 2005; Sutter and Verano 2007).
Focusing on the skeletal samples from the Jequetepeque
Valley, the biodistance results indicate that the Middle Moche
samples from Pacatnamú and San José de Moro are fairly
similar to both each other and contemporaneous samples
from the Moche Valley (Huaca de la Luna–Urban Sector and
Huaca de la Luna–Platform samples). Further, the broad
similarities reflected by the biodistance results are also indi-
cated by the inspection of both the biased (not presented here)
and unbiased R matrix results (table 2). Both the unbiased R
matrix and the biodistance results indicate that genetic dif-
ferentiation increased in the Jequetepeque Valley during both
the LMP and the subsequent Transitional period.
The biased phenotypic FST for the 12 prehistoric skeletal
samples is 0.018 (SE p 0.003), while the unbiased pheno-
typic FST is 0.03 (SE p 0.004). Both the biased and unbiased
FST values differ significantly from 0.0 (P ! .05) and are sim-
ilar to FST values reported for other prehistoric skeletal pop-
ulations (see Nystrom 2006; Stojanowski 2005). Inspection
of the Relethford-Blangero residuals (table 3) indicates that
all skeletal samples predating the LMP are characterized by
negative Relethford-Blangero residuals, while samples from
the LMP and Transitional periods at San José de Moro exhibit
positive residuals. These results suggest that the LMP and
 Table 2. Unbiased distances (below diagonal) and unbiased R matrix (diagonal and above) for 12 north coast skeletal samples analyzed by this study

COSAL COG1 COG2 COG3 HLLUS PLATS PACAT SJMMM SJMMT SJMTRANS P3C P3A
COSAL .026 .006 2.002 .003 2.009 2.002 2.005 2.001 2.005 2.007 2.004 2.001
COG1 .032 .018 .001 .003 2.004 2.001 2.004 2.005 2.007 2.005 2.003 .001
COG2 .048 .034 .018 2.002 .002 .001 2.003 2.004 2.008 2.005 .003 2.001
COG3 .046 .036 .047 .026 2.002 .000 2.007 2.008 .001 2.005 2.012 .005
HLLUS .065 .047 .036 .052 .022 .000 .000 .002 2.006 2.007 .001 .002
PLATS .041 .030 .028 .037 .033 .011 2.001 2.003 2.003 .004 2.002 2.003
PACAT .055 .045 .042 .059 .041 .031 .018 .070 .001 .006 .001 2.012
SJMMM .046 .046 .042 .059 .035 .033 .022 .017 .001 2.004 .005 2.006
SJMMT .069 .065 .067 .058 .066 .049 .051 .049 .033 .002 2.009 2.001
SJMTRANS .085 .073 .074 .081 .082 .049 .052 .071 .074 .046 2.010 2.015
P3C .067 .057 .045 .083 .053 .047 .049 .040 .084 .098 .033 2.003
P3A .062 .051 .055 .050 .052 .051 .077 .064 .069 .111 .072 .034

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Note. rii values are in bold. Unbiased FST p 0.03 (SE p 0.004).

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 768 Current Anthropology Volume 56, Number 5, October 2015

Figure 3. Southern versus northern Moche ceramic chronologies (after Castillo and Quilter 2010:3).

Transitional populations of the Jequetepeque Valley received
substantial external gene flow, while—before the LMP—north
coast populations of the Jequetepeque and Moche valleys re-
ceived less external gene flow.
Discussion and Conclusions
The consensus is that extended periods of drought were
ultimately responsible for the social unrest and political de-
centralization that occurred throughout the north coast in
the LMP (Castillo and Uceda 2008; Dillehay 2001; Dillehay,
Kolata, and Moseley 2004; Dillehay, Kolata, and Pino 2004;
Shimada et al. 1991). Specifically, within the Jequetepeque
Valley, independent surveys and analyses of Late Moche set-
tlements undertaken by Swenson (2004) and the San José de
Moro Archaeological Project (Castillo et al. 2008; Cusicanqui
and Barrazuetea 2010; Ruiz 2004) have identified covariation
in hinterland fortified communities with both feasting wares
and ritual platforms that are associated with divisions in the
Late Moche irrigation systems. Rosas (2010:841) suggests that
the extended drought during the LMP likely increased the
demands for irrigation water in the lower Jequetepeque and
directly contributed to the Moche’s demise and apparent po-
litical balkanization along preexisting canal systems.
Likewise, environmental degradation may have also driven
highland peoples into the valley and resulted in the estab-
lishment of an LMP Cajamarca settlement at nearby Cerro
Chepén. Rosas (2010:847–848) suggests that the extended pe-
riod of drought between 650 and 730 AD was the primary mo-
tivating factor for Cajamarca peoples of either the Guzmango
or the Tantarica highland communities to seek fertile lands and
water in the lower Jequetepeque Valley. Although demands for
irrigation water would have already been strained, the river still
would have had water year round, and the region would have
been more hospitable than the adjacent highlands to the east.
Indeed, a politically fragmented landscape may have made the
Jequetepeque Valley a more inviting location for highlanders
to resettle, while local LMP Moche elites at San José de Moro
may have seen the new highland occupants of Cerro Chepén as
potential allies. Indeed, Rosas (2010:874) contends that Caja-
marca mourners from Cerro Chepén were likely the source
of exotic finewares encountered as funerary offerings at the fu-

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Sutter and Castillo Population Structure during the Demise of the Moche (550–850 AD) 769

Table 3. Relethford-Blangero residuals for 12 north coast skeletal samples

Sample rii Observed (vi ) Expected (E(vi )) Residual (vi 2 E(vi ))

COSAL .026 1.023 1.088 2.065
COG1 .018 .949 1.097 2.148
COG2 .018 .767 1.097 2.330
COG3 .026 .936 1.088 2.152
HLLUS .022 .821 1.093 2.272
PLATS .011 1.156 1.105 .051
PACAT .018 .944 1.096 2.152
SJMMM .017 .899 1.098 2.199
SJMMT .033 1.484 1.080 .404
SJMTRANS .046 1.383 1.066 .317
P3C .033 1.260 1.080 .179
P3A .034 1.328 1.079 .249
Note. Positive residuals indicate larger than average external gene flow. Significant values are in bold.

nerary site San José de Moro. Accordingly, the Cajamarca mi-
grants brought trade items with them and influenced the cul-
tural changes that are most apparent in exotic grave goods
during both the LMP and the Transitional periods at San José
de Moro. Indeed, we posit that the decrease in the local fineline
styles and the increase in eclecticism and blending of local and
exotic forms and styles that typifies locally produced terminal
LMP and Transitional period vessels may be related to the ar-
rival of Cajamarca migrants.
In support of this model, population structure results of
this study indicate that there was substantial external gene
flow into the Jequetepeque Valley following the Middle Moche
period. Before the LMP, unbiased biodistance and FST results
suggest that extensive gene flow existed among Middle Moche/

Figure 4. Multidimensional scaling of the unbiased distance values, as determined from R matrix analysis for 12 prehistoric north
coast skeletal samples. The biased phenotypic FST for the 12 prehistoric skeletal samples is 0.018 (SE p 0.003), while the unbiased
phenotypic FST is 0.03 (SE p 0.004). Both the biased and unbiased FST values differ significantly from 0 (P ! .05). Relative sample
heterozygosity (derived vi values from the R matrix) is represented by each sample’s point size. The unbiased biodistance and FST
results indicate that before the Late Moche Period, extensive gene flow existed among north coast populations up through the Middle
Moche period, suggesting that they formed a relatively coherent breeding population. Following the Middle Moche period, the
unbiased R matrix and biodistance results indicate that genetic differentiation increased as a result of significant levels of extralocal
gene flow into the Jequetepeque Valley during the Late Moche and Transitional periods.

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 770
Moche IV populations of the north coast of Peru, indicating
a largely coherent breeding population. The unbiased esti-
mates of genetic distances (fig. 3) indicate a lack of biological
structure, with no evidence for isolation by distance. How-
ever, as the Moche’s influence began to wane throughout the
region during the LMP, evidence for external contacts in the
form of exotic ceramics from the central coast (i.e., Nievería)
and highlands (i.e., Cajamarca, Chachapoyas, Wari) becomes
evident from grave lots at San José de Moro (fig. 4). Both
the unbiased biodistance results (table 2) and the Relethford-
Blangero residuals indicate that the LMP population repre-
sented by the sample from San José de Moro received sub-
stantial external gene flow and began to differentiate from
the prior Jequetepeque Valley Middle Moche samples from
San José de Moro and Pacatnamú. The levels of external gene
flow and genetic differentiation continued to increase dur-
ing the subsequent Transitional period at San José de Moro.
Importantly, Castillo and Uceda (2008) emphasize that the
appearance of exotic wares is unique to San José de Moro and
that such wares are largely absent from other LMP and Tran-
sitional period sites from the north coast of Perú. Therefore,
the corresponding levels of external gene flow and culture
contact evident during the LMP and Transitional periods at
San José de Moro may not reflect changes at other contem-
poraneous north coast sites.
In the absence of a Cajamarca skeletal sample, we cannot
directly assess the ultimate source(s) of extralocal gene flow
into the Jequetepeque Valley. However, given the afore-
mentioned concomitant influx of both external cultural and
genetic influence detected at the mortuary center San José
de Moro, we tentatively propose that both the population
structure analyses presented here as well as grave good data
from the site and from recent excavations at Cerro Chepén
lend support to a model of an environmentally driven dis-
persal of Cajamarca peoples from the adjacent highlands as
the most likely source for the extra-local genetic and cultural
influences that are apparent during the demise of the LMP
Moche polity in the Jequetepeque Valley.
In conclusion, we suggest that the demise of the Moche
(200–850 AD) polity within the Jequetepeque Valley stemmed,
in part, from external influences that began during envi-
ronmental degradation in the LMP. The inferred arrival of
highlander Cajamarca migrants from the east implied by the
extralocal gene flow into the valley that we detect in our
population structure analyses likely exacerbated an already
politically fragmented landscape and put an additional strain
on the demands for irrigation water, which in turn was likely
to have contributed to the Moche polity’s demise. As the
Moche collapsed, there was both political and economic de-
centralization concurrent with the proliferation of fortified
hinterland sites and the reconfiguration of alliances that formed
along preexisting irrigation systems. We also suggest that the
arrival of Cajamarca environmental refugees contributed to the
presence of LMP exotic grave goods and the subsequent cultural
hybridization detected in San José de Moro grave offerings dur-
ing the subsequent Transitional Period.
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All use subject to JSTOR Terms and Conditions
Current Anthropology Volume 56, Number 5, October 2015
Acknowledgments
Excavations and surveys by L. J. Castillo in the Jequetepeque
Valley, Director of the Programa Arqueológica San José de
Moro, were conducted under Instituto Nacional de Cultura
(INC) Permit 151-2008-DN/INC. R. C. Sutter’s research on
the human dentitions from San José de Moro was funded, in
part, by a Post-PhD Research Grant from the Wenner-Gren
Foundation and an Indiana University Office of the Vice
Provost for Research Faculty Research Support Grant, while
the research conducted during 2001 on the comparative
skeletal samples from the Moche Valley and the sample from
Pacatnamú was funded by National Science Foundation grant
9816958. Key facilities and logistic support for this research
conducted on comparative materials during 2001 were pro-
vided by both the Museo de Arqueología, Antropología e His-
toria de la Universidad Nacional de Trujillo and the Instituto
Nacional de Cultura–La Libertad. We are grateful to our col-
leagues who granted R. C. Sutter permission to examine their
collections, including Christopher Donnan, José Carcelén,
and Santiago Uceda. We also thank Margaret Brown Vega
and two anonymous reviewers who provided constructive
comments on this manuscript. Any and all errors in inter-
pretation or fact are our own.
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