ARTICLE

BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN

MOCHE LORDS: DENTAL PHENOTYPES AND POLITICAL
ORGANIZATION IN ANCIENT PERU

Haagen D. Klaus, Walter Alva, Steve Bourget and Luis Chero

Between AD 100 and 800, the Moche culture emerged on the north coast of Peru. Diverse debates surround the nature of
Moche territorial and political centralization, sociopolitical identities, and the internal social diversity of Moche society.
Here we address some of these issues in a biodistance study based on phenotypic variation of inherited dental traits within
and between 36 individuals in the royal tombs of Sipán (Lambayeque valley), Úcupe (Zaña valley), and Dos Cabezas (Jeque-
tepeque valley). Metric and nonmetric dental trait data were analyzed using hierarchical cluster and R-matrix analyses. The
results independently indicate that the highest-level Sipán and Dos Cabezas lords likely represented different endogamous kin
groups, while limited gene flow occurred between groups of Moche lower nobility between the Lambayeque and Jequetepeque
regions. Although biology and material cultural link the Lord of Úcupe to Dos Cabezas, many objects in his tomb demonstrate
his participation the world of the Sipán elites. These Moche lords were, on some levels, bioculturally interconnected. None-
theless, the data broadly lend support to a “many Moches” model of sociopolitical structure, further casting doubt on earlier
one-dimensional visions of a centralized hegemonic Moche polity.

La cultura Moche surgió en la costa norte del Perú entre 100 y 800 dC. Diversos debates giran en torno a la naturaleza de la
centralización territorial y política Moche, las identidades sociopolíticas y la diversidad social interna de esta sociedad. En
este trabajo abordamos algunos de estos temas a partir de un estudio de biodistancia (o parentesco) basado en la variación
fenotípica de rasgos dentales hereditarios dentro y entre 36 individuos de las tumbas reales de Sipán (Valle de Lambayeque),
Úcupe (Valle de Zaña) y Dos Cabezas (Valle de Jequetepeque). Los datos de rasgos dentales métricos y no métricos se ana-
lizaron mediante análisis de agrupamiento jerárquico y de matriz R. Los resultados indican de manera independiente que los
señores del nivel más alto de Sipán y de Dos Cabezas probablemente representaron diferentes grupos de parentesco endogá-
micos, mientras que ocurrió un limitado flujo genético entre grupos de la nobleza baja de las regiones de Lambayeque y Jeque-
tepeque. Aunque la biología y la cultura material conectan al Señor de Úcupe con Dos Cabezas, muchos objetos en su tumba
demuestran su participación en el mundo de las elites de Sipán. De cierta manera, estos señores Moche estuvieron biocultural-
mente interconectados. Sin embargo, los datos apoyan un modelo de estructura sociopolítica de “múltiples Moches” que pone en
duda las visiones unidimensionales de una unidad política hegemónica Moche centralizada.

T
he Moche culture emerged approximately
1,900 years ago on the arid north coast of
Peru and flourished for centuries, produ-
cing artistic masterworks, monumental architec-
ture, and a narrative art style, perhaps second
only to that of the ancient Greeks. For most of
the twentieth century, Moche studies drew from
an art history paradigm and they were long envi-
sioned as the first centralized territorial Andean
state. Since the 1980s, the growth of Moche
field archaeology has elucidated the intricacies
of Moche political organization (Quilter and
Castillo 2010), forcing a fundamental rethinking
of the Moche phenomenon.

Haagen D. Klaus ▪ Department of Sociology and Anthropology, George Mason University, 4400 University Dr., Fairfax, VA
22030, USA (hklaus@gmu.edu, corresponding author)
Walter Alva ▪ Director, Museo Tumbas Reales de Sipán, Juan Pablo Vizcardo y Guzman 895, Lambayeque, Peru
Steve Bourget ▪ Collections Manager, Department of the Americas, Musée du quai Branly - Jacques Chirac 37 Quai Branly,
75007 Paris, France
Luis Chero ▪ Director, Museo de Sitio Huaca Rajada-Sipán, Avenida Augusto B. Leguía, CPM Sipán, Lambayeque, Peru
Latin American Antiquity, pp. 1–22
Copyright © 2018 by the Society for American Archaeology
doi:10.1017/laq.2018.46

1
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 2 LATIN AMERICAN ANTIQUITY
To date, 22 Moche elite tombs have been
excavated at the three centers of Moche political
power on the north coast of Peru during the
mid-first millennium AD: Sipán (Lambayeque
valley), Úcupe (Zaña valley), and Dos Cabezas
(Jequetepeque valley). What were the social
identities and relationships between these indivi-
duals? How did these groups of Moche elites in
neighboring valleys interact with each other?
Were they a single dynasty distributed across
the coast as part of centralized territorial Moche
state? Alternatively, do they represent more insu-
lar groups reflecting a decentralized, plural, and
contested political reality?
Bioarchaeological data provide often other-
wise unobtainable insights on social structure,
ideology, politics, and identity. Human remains
from these sites provide an initial opportunity
to reconstruct and interpret biological interaction
patterns within and between the Moche lords at
Sipán, Úcupe, and Dos Cabezas. We provide a
bioarchaeological analysis on interindividual
variation and preliminary population-structure-
related patterns of inherited dental traits among
36 individuals to provide insights into the nature
of Moche politics and organization and society.
The Moche Culture
Chronology and Characteristics
The desert coast of Peru has been home to a
sequence of complex cultures since 2600 BC.
Following the emergence and decline of several
early complex societies, the Moche were the
dominant cultural development in the north
coast during the first millennium AD (Figure 1;
Benson 2012; Bourget and Jones 2008; Shimada
1994a; Uceda and Mujica 1994, 2003). Their
iconography communicated a belief system insti-
tutionalizing themes surrounding death, sacri-
fice, and fertility (Bourget 2006; Donnan 1995;
Millaire 2002). Moche economy was fueled by
large-scale systems of tribute and labor organiza-
tion, irrigation agriculture, fishing, camelid herd-
ing, and trade (Bawden 1996; Benson 2012).
Moche politics appears unexpectedly complex,
sparking various intense debates.
Moche origins are poorly understood but
probably spanned multiple locations between
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AD 100 and 300, involving growth of the polit-
ical hierarchy of the preexisting Gallinazo cul-
ture, accompanied by a new religious formation
(Bourget 2016). The two most prominent loci
of these early developments occurred in a pre-
vailing northern polity centered around Sipán
in the Lambayeque region and in a southern pol-
ity located in the Moche-Chicama valley region
(Castillo and Donnan 1994; Shimada 1994b).
Separating these two regions was the Pampa de
Paiján, a 90 km intervalley desert expanse serv-
ing as a natural geopolitical and cultural barrier.
Southern territories were under the influence of
the Huacas de Moche site, while to the north of
the pampa, multiple autonomous polities devel-
oped in the Jequetepeque, Zaña, Lambayeque,
and La Leche valleys. There are numerous differ-
ences between northern and southern Moche
material culture, art styles, architecture, and his-
torical trajectories (Shimada 2010). The five
classical temporal phases developed by Larco
Hoyle are applicable only to the southern
Moche. In the north, an Early, Middle, and
Late Moche sequence is recognized that can be
further subdivided into Middle Moche A, B,
and C phases (Bourget 2014; Supplemental
Tables 1, 2).
An extended and abnormally intense El Niño
Southern Oscillation (ENSO) event was asso-
ciated with a series of social and political trans-
formations beginning around AD 600. The
southern Moche system mostly imploded, and a
complex transition to the Moche V/Late Moche
transpired (Koons and Alex 2014). At this time,
reconfigurations of Moche ideology and econ-
omy surfaced at Galindo (Moche valley), San
José de Moro (Jequetepeque valley), and
Pampa Grande (Lambayeque valley; Bawden
2005; Castillo 2001; Shimada 1994a). The final
political disintegration of the Moche unfolded
around AD 800, although a few strongholds per-
sisted for several more decades (Uceda 2010).
Moche Political Organization: Previous
Thinking and Unresolved Questions
Over the years, Moche studies featured sever-
al prominent themes, from descriptive and
chronology-building foci to art history (e.g.,
Donnan 1978; Kubler 1967). Thoughts on
Moche politics by Larco Hoyle (1938) and
 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS 3

Figure 1. The north coast of Peru, highlighting major Moche sites including those discussed in the text. Map by
Haagen Klaus.

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 4 LATIN AMERICAN ANTIQUITY
participants of the 1940s Virú Valley Project
envisioned a single, centralized, hegemonic,
and expansionist Moche state with its capital at
the Huacas de Moche. This notion persisted for
decades. More recently, field excavations point
to a “many worlds of the Moche” model that
sees evidence of intricate and distinct constella-
tions of Moche peoples, politics, and social
identities (Benson 2012; Quilter and Koons
2012). These include a hybrid Moche-Vicús
polity/art style in the far northern coast
(Makowski 1994). Descendants of the earlier
Gallinazo peoples persisted as one or more social
or ethnic groups coexisting with, and even
outliving, the Moche in places (Bourget 2010;
Donnan 2009; Shimada and Maguiña 1994).
Simultaneously, it has become increasingly
apparent that northern Moche centers were
closely linked to each other and with their south-
ern counterparts (for a fuller exposition of this
argument, see Bourget 2014). Both northern
and southern Moche elites participated in a com-
mon cultural and religious framework. Yet, these
ideas were expressed differently at the three nor-
thern Middle Moche centers: Sipán, Úcupe, and
Dos Cabezas.
Research Questions
Archaeogenetic studies of the northern Moche
lords of Sipán, Úcupe, and Dos Cabezas have
been frustrated by poor ancient DNA preserva-
tion. Shimada and colleagues (2005, 2008) re-
covered authentic mitochondrial DNA (mtDNA)
from only nine individuals at Sipán. The results
point to the Sipán funerary platform being a mau-
soleum for a closely related Moche royal lineage.
Here, we analyze the dental morphology of these
and other Moche lords, whose teeth were far
more intact, to provide strong indications of gen-
etic relationships. Multivariate analysis of den-
tally inherited traits reflects both maternal and
paternal genetic signal traits, providing a basis
for reconstructing aspects of relationships
among northern Moche elites, including for
those with poor aDNA preservation. How were
lords at different centers across different valleys
related to one another? How did they interact?
Did they intermarry? What kinds of organiza-
tional corollaries might be reflected in biological
interaction patterns?
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In light of growing evidence for multiple
Moche “worlds,” we also recognize integrative
and harmonizing interregional mechanisms that
interconnected Moche rulers and their courts
into a single, shared elite tradition (Bourget
2016; Donnan 2010). Accordingly, we address
the traditional vision and hypothesize that pat-
terns of elite kinship and intermarriage (gene
flow) will demonstrate a biologically homogen-
ous group of Moche elites between Sipán,
Úcupe, and Dos Cabezas, reflecting an integrated
and centralized leadership ruling over the nor-
thern Moche peoples and territories during the
late Early and Middle Moche periods.
Materials and Methods
Sipán
Since 1987, 16 elite late Early Moche and Mid-
dle Moche tombs have been excavated at the
high-status monumental funerary platform of
Sipán Mound in the mid-Reque drainage in the
Lambayeque river Valley Complex (Supplemen-
tal Figure 1). These adobe brick chamber tombs
and their contents have been described in exten-
sive detail (see Alva 1994, 2001, 2015; Alva and
Donnan 1993), and include the iconic Lord of
Sipán (Tomb 1), the Old Lord of Sipán (Tomb
3), a Warrior-Priest (Tomb 14), and a highly
ranked Priest (Tomb 2). It contained an astonish-
ing range of gold and silver diadems, backflap
ornaments, exquisite drilled shell bead pectoral
necklaces, ritual paraphernalia including knife-
handled scepters, and the skeletal remains of
six tomb retainers.
The tombs at Sipán appear to encompass
four distinct categories: ruling male lords
(Figure 2; Tombs 1, 3, 14, 15), high-status
male priests (Tomb 2), a second-tier nobility
(Figure 3) consisting of male individuals span-
ning inferred military leaders, and assistants to
the lords and priests (Tombs 4–6, 7, 8, 9, 10,
11, 12, 16). The principal personages of inferred
political or military roles were buried in the nor-
thern sector of the platform. Religious personnel
were interred in the southern sector. Tomb retai-
ners (the females and the few children placed
adjacent to the principal personages in symbolic-
ally subordinate locations) have been varyingly
 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS 5

Figure 2. Examples of first-order (high elite) tombs at Sipán: the iconic Tomb 1, or “Lord of Sipán” (a), the
“Warrior-Priest” Tomb 14 (b), and the “Priest” of Tomb 3 (c). Images by Museo Tumbas Reales de Sipán. (Color
online)

Figure 3. Examples of second-order (lower nobility) tombs at Sipán: Tomb 8 (a), Tomb 10 (b), and Tomb 12 (c). Images
by Museo Tumbas Reales de Sipán. (Color online)

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 6 LATIN AMERICAN ANTIQUITY
inferred as sacrifice victims, wives, or deputies
(Bentley and Klaus 2016). Newly published cali-
brated Bayesian radiocarbon and thermolumin-
escence dates (Aimi et al. 2017) define their
span from the mid-fourth century (Tomb 3) to
the mid-seventh century (Tomb 1; Supplemental
Tables 3, 4).
Úcupe
Another monumental Moche center was estab-
lished south of the Reque region on the south
bank of the Zaña river during the late Early
Moche era. The architectural complex of El
Pueblo de Úcupe covered 970 ha, the focal
point of which was a truncated adobe platform
mound some 81 m long, 55 m wide, and 18 m
high. A chamber tomb contained an elite in-
dividual dating to the Middle Moche A period
and was locally named the Lord of Úcupe
(Figure 4; Bourget 2014). Tomb contents and
internal organization closely mirrored practices
at Sipán. This adult male principal personage
was interred wearing full regalia (metal funerary
masks, multiple nose clips and diadems, and full-
sized metal war clubs fashioned from copper
sheeting) and was accompanied by three tomb
retainers (Supplemental Figure 2) and sumptuary
ceramics.
Dos Cabezas
Between 1997 and 2000, Donnan (2007)
excavated five virtually contemporaneous elite
tombs at Huaca Dos Cabezas in the Jequetepeque
valley (Figure 5). The earliest contexts were
Tomb A and Tomb B. Tomb A held the disarti-
culated remains of an adult male, but at some
point in antiquity, the tomb was opened, and
the skeletal remains were removed, manipulated,
and replaced (sensu Shimada et al. 2015). Tomb
B contained remains of four inferred retainers
and isolated skeletal elements, and was probably
a kind of an open sepulcher.
Tombs 1, 2, and 3 occurred slightly later in
this sequence. Each featured a centrally placed
male principal personage accompanied by
resplendent headdresses and diadems, gold, sil-
ver, copper, and gilded copper nose ornaments,
and sumptuary ceramics. A single adolescent
female accompanied the male in Tomb
1. Tomb 2, which was centrally located and the
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largest, featured a male principal personage
accompanied by two retainers. A similar context
was seen with the male in Tomb 3 buried with a
woman and an older child. Radiocarbon dates
place Dos Cabezas in Middle Moche A (aver-
aging between AD 410 and 620) and are contem-
poraneous with the Lord of Úcupe. The skeletal
remains of these Moche lords demonstrated
unusually tall stature and anomalous growth pat-
terns (Cordy-Collins and Merbs 2008). Biers and
colleagues (2017) considered these characteris-
tics consistent with endocrine disruptions
produced by castration.
Biodistance Analysis
Biological distance (biodistance) analysis is one
of about a dozen areas of bioarchaeological
research. It involves mathematical assessment
of similarity of phenotypes that reflect microevo-
lution or culturally structured patterns of genetic
variation and inheritance (Larsen 2015). Biodis-
tance analysis provides a record of kinship, gene
flow, ethnic group boundaries, postmarital resi-
dence patterns, migration, and other forms of
biological interactions, from intracemetery to
global scales (Buikstra et al. 1990; Larsen
2015; Pilloud and Hefner 2016; Stojanowski
and Schillaci 2006).
The basis of biodistance research is well
established by evolutionary theory. As popula-
tions exchange mates (genes), they become
phenotypically and genetically more alike;
those that do not exchange genes become
increasingly dissimilar at a rate determined by
genetic drift and their effective population sizes
(Stojanowski and Schillaci 2006). Although there
is never a 1:1 correspondence between pheno-
type and genotype, some dental and skeletal
phenotypes do serve as useful proxies of under-
lying polygenic variation. While cranial size and
shape provide more of a mirror on environmental
factors (Katz et al. 2017), tooth development is
strongly canalized and developmentally invari-
ant. Data include metric traits (linear measure-
ments such as length and width of a tooth
crown) and nonmetric traits (discrete or quasi-
continuous features involving the presence,
absence, or graded expressions of a trait; Hemp-
hill 2016a, 2016b; Irish 2016; Pilloud and
 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS 7

Figure 4. The tomb of the “Lord of Úcupe” at Huaca El Pueblo, Zaña valley, with a view of the exposed funerary bundle
(a); diadems, necklaces, and burial masks defined during excavation (b); and, the exposed principal personage of the
tomb (c). Photos by Steve Bourget. (Color online)

Kenyhercz 2016; Pilloud et al. 2016; Scott and
Turner 1997).
The heritability (h2) of various dental traits
remains incompletely studied and understand-
ings vary trait-by-trait. Mesiodistal incisor
crown dimensions demonstrate an extremely
high degree of genetic expression (h2 = 0.81 to
0.91; Dempsey et al. 1995). Most other dental
metric and nonmetric h2 trait values fall between
0.70 and 0.90 to indicate varying but consistently
dominant genetic (and not environmental) bases
of expression. At least 300 genes, including key
homeobox genes, play roles in tooth morphogen-
esis (Edgar and Ousley 2016). Some nonmetric
traits appear selectively neutral (Scott and Turner
1997), while others are varyingly shaped by
selection (Mizoguchi 2013). Still, on the time-
scale considered here, dental variation is most
directly shaped by differences in population
history and biological affinity.
Data Collection
Dental metric and nonmetric data were collected
from 20 individuals from Sipán, four individuals
in the tomb of the Lord of Úcupe, and nine indi-
viduals from the five tombs at Dos Cabezas, who
had more than 75% of their dentitions preserved
(Table 1). Metric data gathering was modeled
after Jacobi’s (2000) techniques based on classic
protocols established by Moorres and Reed
(1964) and Kieser (1990).
Metric data were gathered using a Mitutoyo
digital sliding caliper with sharpened tips cali-
brated to 0.01 mm with a direct USB input into
a Microsoft Excel spreadsheet. Measurements
were made during two separate passes through

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 8 LATIN AMERICAN ANTIQUITY

Figure 5. Tomb A (a), Tomb B (b), Tomb 1 (c), Tomb 2 (d), and Tomb 3 (e) at Huaca Dos Cabezas, Jequetepeque valley.
Images by and courtesy of Christopher B. Donnan. (Color online)

the material separated by several hours. For each
tooth, standard maximum mesiodistal and buc-
colingual tooth dimensions were documented
(detailed measurement protocols are presented
in the online Supplemental Text 1). Twenty-
seven nonmetric traits were scored using Turner
and others’ (1991) Arizona State University
Dental Anthropology Scoring System (Scott
and Irish 2017; Scott et al. 2016).
Data Preparation and Analysis
The raw dataset consisted of 56 metric variables
and 48 nonmetric variables per individual
(Table 2; for complete descriptions of the

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 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS 9

Table 1. Individuals from Sipán, Úcupe and Dos Cabezas Examined in This Work.

Sipán Úcupe Dos Cabezas

Tomb 1 Principle Personage “Lord of Sipán” Burial 1 (Retainer 1) Tomb A Burial 1 (Inferred Principal
Personage)
Tomb 1 Retainer 1 “Primary Woman” Burial 2 (Principal Personage Tomb B Burial 1 (retainer)
“Lord of Úcupe)”
Tomb 1 Retainer 2 “Secondary Woman” Burial 3 (Retainer 2) Tomb B Burial 2 (retainer)*
Tomb 1 Retainer 3* Burial 4 (Retainer 3) Tomb B Burial 2 (retainer)
Tomb 1 Retainer 4 * Tomb B Burial 4 (retainer)*
Tomb 1 Retainer 5 * Tomb 1 Burial 1 (Principal Personage)
Tomb 1 Retainer 6 * Tomb 1 Burial 2 (retainer)
Tomb 1 Retainer 7 “Guardian”* Tomb 2 Burial 1(retainer)
Tomb 1 Retainer 8* Tomb 2 Burial 2 (Principal Personage)
Tomb 2 Principle Personage “Priest” Tomb 2 Burial 3 (retainer)
Tomb 2 Principle Personage Tomb 3 Burial 1 (retainer)
Tomb 2 Retainer 1 Tomb 3 Burial 2 (Principal Personage)
Tomb 2 Retainer 2 Tomb 3 Burial 3 (retainer)
Tomb 2 Retainer 3*
Tomb 2 Retainer 4*
Tomb 2 Retainer 5*
Tomb 3 Principal Personage “Old Lord of Sipán”*
Tomb 3 Retainer 1*
Tomb 5 Principal Personage “Musical Warrior”
Tomb 6 Principal Personage “Priest”
Tomb 7 Principal Personage
Tomb 7 “Guardian”
Tomb 8 Principle Personage A
Tomb 8 Principle Personage B
Tomb 10 Principal Personage
Tomb 11 Principle Personage*
Tomb 11 Retainer 1*
Tomb 12 Principal Personage
Tomb 12 Retainer 1
Tomb 14 Principal Personage “Warrior-Priest”
Tomb 14 Retainer 1
Tomb 15 Principal Personage “Young Noble”
Tomb 16 Principal Personage
Tomb 2 Retainer 1*
Tomb 2 Retainer 2*
* Not included in analysis due to more than 25% missing data.

statistical procedures conducted here, please see
the online Supplemental Text 1). In summary,
work began with removal of variables with
zero variance and data categories consisting of
more than 25% missing data. Interobserver
error for metric data was evaluated using one-
way ANOVA tests in SAS 9.4 (Supplemental
Table 5), and for nonmetric traits, Cohen’s
kappa tests were used. Each test revealed
strong nonsignificant error. Any missing metric
data were imputed using an expectation-
maximization algorithm. For nonmetric ordinal
values, the multiple imputation technique was
used. For each, missing data points were calcu-
lated via 10,000 iterations performed 25 times.
To maximize phenotypic coverage, metric
and nonmetric dental traits were combined into
a single dataset following Corruccini and Shi-
mada (2002), by converting them into z-scores
in SPSS 24. Then, conversion of z-scores
into c-scores minimized any effects of sex-
ual dimorphism. A Kolmogrov-Smirnov test
demonstrated a normal distribution of the dataset.
Distances were calculated using Malhalanobis’s
(1936) generalized distance (D2) that employs a
tetrachoric correlation in the calculation of a

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 10 LATIN AMERICAN ANTIQUITY

Table 2. Traits Used in Biodistance Analysis of Sipán, Úcupe, and Dos Cabezas.

M-D and B-L measurements (mm) I1, LC, P3, P4, M1, M2 M-D and B-L measurements (mm) I1, LC, P3, P4, M1, M2*
1 2
Labial Convexity, I , I Shoveling, I1, I2, LC
Shoveling, I1, I2, LC** Canine Distal Accessory Ridge**
Cusp number, M1, M2
Double Shoveling, I1, I2, LC Anterior Fovea, M1, M2
Interruption Groove, I1, I2, Groove Pattern, M1, M2
Tuberculum Dentale, I1, I2, LC
Canine Mesial Ridge** Distal Trigonid Crest, M1, M2
Canine Distal Accessory Ridge** Deflecting Wrinkle, M1, M2**
Metacone (Cusp 3), M1, M2 Protostylid, M1, M2
Hypocone (Cusp 4), M1, M2 Hypoconulid (Cusp 5), M1, M2
Metaconule (Cusp 5), M1, M2 Entoconulid (Cusp 6), M1, M2
Carabelli’s Trait, M1, M2*** Metaconulid (Cusp 7), M1, M2
Parastyle, M1, M2*** Enamel extensions/pearling P3, M1, M2
Enamel extensions/pearling P3, M1, M2 Peg-shaped incisor or molar ***
Peg-shaped incisor or molar, M1***, M2 Odontome, all premolars
Odontome, all premolars***
* Buccolingual measurements of the LC, LI1, and LI2 , removed from analysis due to more than 25% missing data.
** Nonmetric trait removed from analysis due to more than 25% missing data.
*** Nonmetric trait removed from analysis due to zero variance (universal absence).

covariance matrix to eliminate distortional
effects of trait intercorrelation. Hierarchical
cluster analysis using the average linkage tech-
nique was used to represent distances. This
method is simple, robust, and proven (Everitt
et al. 2001). To validate the clusters, a silhouette
measure of cohesion and separation was
employed.
A window on population structure was pro-
vided by an R-matrix analysis of the metric
data. Based on wholly different mathematics,
an R-matrix can also serve as an independent
test of cluster analysis results and interpretations.
The R-matrix is a standard method that estimates
population genetic heterogeneity and variance
via metric variation in RMET 5.0 (see Relethford
2016) available at: http://employees.oneonta.
edu/relethjh/programs/. It independently calcu-
lates a bias-corrected measurement of FST (an
expression of population genetic diversity), the
relative degree of external gene flow, and a prin-
cipal component analysis plot of Malhalanobis
distances calculated from the R-matrix.
Results
2
Based on the D proximity matrix, the average
linkage algorithm produced a dendrogram with
distinctive structure (Figure 6). The silhouette
measure of cohesion and separation value was
0.50—the breakpoint between “fair” and “good”
quality. Thus, we have confidence that the clusters
represent nonrandom patterns and mathematically
authentic relationships.
The individuals from Sipán, Úcupe, and Dos
Cabezas grouped into two primary clusters—one
dominated by the highest-ranking peoples from
Sipán and the other by the lords of Dos Cabezas.
The notional “Sipán Cluster” contains 18 indivi-
duals, 14 of whom were from Sipán. This cluster
featured two primary constituent groupings.
Cluster 1 Group 1 was composed of the principal
personage of Sipán Tomb 2, the “guardians” of
Tombs 2 and 7, and retainers from Dos Cabezas
Tombs B, 2, and 3. A particularly close distance
was observed between Sipán Tomb 1 Retainer 1
and the guardian of Sipán Tomb 2. Group 2 con-
tains the Lord of Sipán and the principal perso-
nages of the Sipán Tombs 6, 12, 14, and 16,
along with retainers from Tomb 1, 2, and 14.
Also in this group was the principal personage
of Dos Cabezas Tomb 1.
The “Dos Cabezas Cluster” contained 18
individuals. Three of the four principal perso-
nages of the Dos Cabezas tombs (Tombs A, 2,
and 3) are found along with all members of the

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 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS 11

Figure 6. Dendrogram representing estimated phenetic distances between the individuals from Sipán, Huaca el
Pueblo-Úcupe, and Dos Cabezas. Illustration by Haagen Klaus.

Úcupe tomb. In Cluster 2 Group 1, a strong phe-
netic similarity was observed between four Dos
Cabezas retainers, two of the three retainers at
Úcupe, and the principal personage of Dos Cabe-
zas Tomb 3. Cluster 2 Group 2 contained the
principal personages of Sipán Tombs 7 and 15,
along with retainers from Sipán Tombs 8 and
12 and the final retainer from Úcupe. Cluster 2
Group 3 consisted of the principal personages
of Sipán Tombs 5, 7, and 10, the principal

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 12 LATIN AMERICAN ANTIQUITY

Table 3. Diagonals of the R-matrix and Calculation of Unbiased FST and Standard Error (A), R-matrix Estimation of
Differential Gene Flow (Residual Values; B), and Malhalanobis (D2) Distance Matrix Derived from the Principal Components
Analysis of the R-matrix (C).

3A. FST ANALYSIS

Sample Biased R(ii) Unbiased R(ii) Standard Error
Sipán High Elites 0.758742 0.658742 0.054053
Sipán Lower Nobility 0.184183 0.084843 0.026679
Sipán Retainers 0.647363 0.601910 0.033662
Úcupe/Dos Cabezas Lords 0.336706 0.253373 0.032871
Úcupe Retainers 2.449121 2.28454 0.125373
Dos Cabezas Retainers 0.179194 9.095861 0.023980
FST = 0.759328
Unbiased FST = 0.662864
Standard Error = 0.005969

3B. R-MATRIX ESTIMATION OF

DIFFERENTIAL GENE FLOW
Observed Within- Expected Within-
Sample R(ii) Group Variance Group Variance Residual Value
Sipán High Elites 0.758742 0.977 0.986 − 0.009
Sipán Lower Nobility 0.184183 1.045 2.644 − 1.599
Sipán Retainers 0.647363 0.704 1.150 − 0.446
Úcupe/Dos Cabezas Lords 0.336706 1.98 2.157 − 0.169
Úcupe Retainers 2.449121 0.511 −3.705 4.217
Dos Cabezas Retainers 0.179194 0.619 2.612 − 1.994

3C. PCA- DERIVED D2 DISTANCE MATRIX

Subsample 1 Subsample 2 D2 Distance Value Standard Error
Sipán High Elites Sipán Lower Nobility 1.114493 0.100616
Sipán High Elites Sipán Retainers 0.132880 0.039484
Sipán High Elites Úcupe/ Dos Cabezas Lords 0.283828 0.057428
Sipán High Elites Úcupe Retainers 5.374515 0.240681
Sipán High Elites Dos Cabezas Retainers 1.223915 0.099673
Sipán Lower Nobility Sipán Retainers 0.930455 0.077629
Sipán Lower Nobility Úcupe/ Dos Cabezas Lords 0.475249 0.068186
Sipán Lower Nobility Úcupe Retainers 2.045585 0.154090
Sipán Lower Nobility Dos Cabezas Retainers 0.120455 0.046310
Sipán Retainers Úcupe/ Dos Cabezas Lords 0.058375 0.030466
Sipán Retainers Úcupe Retainers 5.257130 0.211363
Sipán Retainers Dos Cabezas Retainers 1.300709 0.084198
Úcupe Retainers Úcupe/ Dos Cabezas Lords 4.050351 0.203467
Úcupe Retainers Dos Cabezas Retainers 1.277657 0.121271
Úcupe/ Dos Cabezas Lords Dos Cabezas Retainers 0.796280 0.078614

personages of the Úcupe tomb, Dos Cabezas
Tomb A, and Dos Cabezas Tomb 2.
R-matrix analysis of population structure
results in a bias-corrected FST of 0.66 with a
low standard error (0.0059), meaning that the
sample possesses an extremely strong degree of
heterogeneous structure; put another way, the
constituent subsamples are very different from
one another (Table 3A). Except for the Úcupe
retainers1, all of the subsamples appear to
possess varying degrees of relatively negative
extralocal gene flow as compared to the model-
derived expectation of the broader population
as a whole (Table 3B). The PCA plot (Figure 7)
derived from the R-matrix (Table 3C) illustrates
closest similarity between the high Sipán elites
and the Sipán retainers. Increasing distance
emerges between them and the Lords of Dos
Cabezas, lower Sipán nobility, the Dos Cabezas
Retainers, and the Úcupe retainers.

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 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS 13

Figure 7. Principal coordinates analysis plot of scaled Malhalanobis distances obtained from the R-matrix. Illustration
by Haagen Klaus.

Discussion and very low R-matrix standard error all inde-

Phenotypic data should be considered a reflec-
tion of underlying genotypes rather than a pre-
cise representation of actual genes. That said,
the cluster analysis (a model-free, exploratory
interindividual analysis) and the R-matrix (a
model-bound, population genetic analysis) pro-
duced comparable results and indicate that it is
appropriate to reject the hypothesis. Biodistance
patterns between the lords of Sipán, Úcupe, and
Dos Cabezas cannot be reasonably interpreted as
a single, unified, or uniform elite lineage.
Any inferences drawn from these results must
be conservative because the samples are small
and we may never know how they may be
skewed or otherwise nonrepresentational due to
the devastating looting at these sites. Still, the
correlation of biodistance patterns with multiple
lines of archaeological contexts, the strong sil-
houette measure of cohesion and separation,
pendently speak to the likelihood that these
observations capture reflections of ancient bio-
cultural patterns that are mathematically reliable.
Of significant interest, the bias-corrected FST
value of 0.66 is extremely high, as the modern
human average is approximately 0.16 (Temple-
ton 2002). We can be virtually certain this
value does not represent the Moche population
as a whole. More likely, it is the signature of con-
siderably intense endogamy among those repre-
sented in a small sample drawn from a
proportionally very small breeding population
(see below).
Intrasite Biodistance Structures
High Elites and the Lower Nobility at Sipán.
Interindividual cluster analysis indicates the
Lord of Sipán appears most similar to the princi-
pal personages of Sipán Tombs 6, 12, 14, and 16,

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 14
whereas retainers from Tombs 1, 6, and 14 were
assigned membership in other clusters. These
tombs span more than a century at Sipán and
indicate continuity of this inferred hereditary lin-
eage between Middle Moche A and B periods.
The largest distances appear between the group-
ing of the Lord of Sipán/Tomb 12 principal per-
sonage/Tomb 14 principal personage and the
priest of Tomb 2. This reinforces previously
defined distinctions between the tombs with
more secular overtones (northern side of the plat-
form) and those associated with religious leader-
ship (southern side of the platform). They
perhaps represented differentially ranked and
spatially segregated lineages within the funerary
platform.
Concerning the retainers, women buried with
the Lord of Sipán appear to have been varyingly
linked to him but at somewhat greater distances.
At least one of these women (Tomb 1 Retainer B)
was associated with the principal personage of
the second-order Tomb 16. Perhaps female kin
of lower-ranked lords became retainers to more
highly ranked lords. Still, the identity of these
women remains a vexing question (Bentley and
Klaus 2016). They are commonly inferred as
either wives or kin. These results are consistent
with either possibility, and the R-matrix analysis
shows the Sipán high elite and Sipán retainers
possessed very similar FST values and shared
very close relationships on the PCA plot.
Kin and wife are not necessarily mutually
exclusive concepts among Andean royalty. Rost-
worowski (2000) noted that the prevalent rule of
succession among yunga (north coast region)
nobility involved the brother of the deceased
(rather than his son) who assumed rulership,
and this pattern of succession continued to the
last brother of that generation before passing to
the subsequent generation. Given Shimada and
others’ (2005) evidence of maternal linkages
between several Sipán lords, the Moche may
have practiced an avunculate system of descent
where the son of the ruler’s sister had priority
over the ruler’s son, potentially promoting
“cases of incestuous marriage between ruling
brother and sister” (Rostworowski 2000:178).
Kinship Heterogeneity at Dos Cabezas. At
Dos Cabezas, most lords also appear to share
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LATIN AMERICAN ANTIQUITY
close mutual relationships, particularly between
the lords of Tomb A and Tomb 2. Because
Tomb A preceded Tomb 2, temporal biological
continuity of a local royal lineage is again likely.
This linkage also indicates the disturbed remains
in Tomb A was of a person biologically part of
the local aristocracy, possibly even the original
principal personage of the tomb. Other strong
phenetic connections existed between the princi-
pal personage of Dos Cabezas Tomb 3, the retain-
ers of Dos Cabezas Tombs B, 1, 2, and 3, and
Retainers 1 and 4 at Úcupe. While they were
spread across geographic space and overlapped
during Middle Moche A, these people were evi-
dently linked by kinship. And although the lord
of Dos Cabezas Tomb 1 shared the same abnor-
mal growth anomalies with the other lords of Dos
Cabezas, it did not involve immediate biological
ties. The larger Dos Cabezas cluster possessed a
more heterogeneous FST value and contained a
diverse cross-section of retainers and personages
from other tombs and sites. Perhaps Sipán was
somehow more “exclusive” or bounded.
Northern Moche Biodistance in Regional
Perspective
Gene Flow and Intermarriage. The data indi-
cate potential gene flow and intermarriage
between the Moche leadership at Sipán and
Dos Cabezas over the span of perhaps three to
five generations (Middle Moche A [ca. AD
390–500] to Middle Moche B [ca. AD 550–
700]). Cluster analysis demonstrates an intri-
guing pattern of principal personages grouping
at the bottoms of Clusters 1 and 2. Eight Sipán
individuals (principal personages of Sipán
Tombs 5, 7, 10, and 15, along with Sipán
Tomb 8 Retainers C and D, and the retainer of
Tomb 12) show affinity with the Dos Cabezas
Moche. The PCA plot of the R-matrix shows
lower Sipán nobility (as a group buried mostly
in the southern [lower] sector of the Sipán funer-
ary platform) falling on the other side of the
population centroid from the high Sipán elite.
These dissimilarities could reflect a few
possibilities.
One is that the lower nobility did not
exchange significant quantities of genes with
the lords of the Tombs 1, 2, 6, 12, and 14.
 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS
Relative to the high Sipán elite, these lower-
status lords were more homogenous (inbred)
with a smaller FST and greater degree of negative
gene flow. Cluster analysis suggests that the lord
of Tomb 14 is in fact the most different (isolated)
of all. Because it is highly probable that the
Moche lordship featured a very small effective
population size, intermarriage would rapidly
make these people more similar.
Cluster analysis and R-matrix data indicate
these distinctions may reflect a social status bar-
rier to widespread intermarriage and that the
lower Sipán elites represented a different lineage.
Most second-order Sipán tombs held a biodis-
tance affinity to the Jequetepeque lords, distinct
from the principal Sipán dynastic core group.
Knowledge and memory of this ancestry may
have created a perception of “otherness,” and
perhaps sometimes encoded in idiosyncrasies
of funerary ritual. For example, Sipán Tomb 10
was atypical in several respects, containing the
body of an adolescent male accompanied by
only a few ornaments. The body was oriented
north-south (looking to the south), unlike the
standard south-north orientation. This young
male’s closest biological relationships included
the principal personages of Dos Cabezas Tomb
A and Tomb 2. Was the Sipán Tomb 10 person-
age positioned to face south looking towards the
region of geographical ancestry?
Similarly, cluster analysis associated the prin-
cipal personage of Sipán Tomb 15 with Dos
Cabezas. This tomb represents one of the first
burials at Sipán, and the corpse was also posi-
tioned to face south. Some of the grave goods
are extremely reminiscent of Jequetepeque cer-
amic styles (in particular, two ceramic owl bot-
tles; Chero 2015:Figure 177). Although this
person is unlikely to be a lineage founder at
Sipán, a Dos Cabezas connection suggests bio-
cultural linkages between Lambayeque and
Jequetepeque valleys from the early days of
Sipán, if not before.
Paralleling possible “otherness” at Sipán,
three retainers in Dos Cabezas Tombs 1, 2, and
3 were evidently most closely related to the retai-
ners from Sipán Tombs 1, 2, and 7. They were
also clearly of a lower status. Similarly, a close
affinity linked the principal personage of Dos
Cabezas Tomb 1 (containing the least wealth
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15
and complexity of the Dos Cabezas tombs) to
the principal personages of Sipán Tombs 1, 6
and 162.
Any gene flow between Sipán and Dos Cabe-
zas might be understood in terms of strategies
involving the different ways regional royal
lineages maintained ties with one another.
Nevertheless, these data indicate this interchange
may not have been equal: more people buried at
Sipán bore a biological correlation to the Dos
Cabezas lineage. R-matrix results lend additional
credence to this idea. High elites at Sipán demon-
strate both higher (more diverse) FST values and
a residual value closer to zero, indicating greater
relative extralocal gene flow among the Dos
Cabezas lords. Thus, directional gene flow is
suggested with Sipán high elites characterized
by a greater intake of genes (mates) from differ-
ent sources other than themselves. One possibil-
ity is Dos Cabezas occasionally sending brides
or grooms to Sipán to establish and maintain pol-
itical alliances. We hypothesize that these people
and their descendants formed part of this lower
elite stratum.
Identity and Territory at Úcupe. The affin-
ities between the Lord of Úcupe and his retainers
with the Dos Cabezas Moche are surprising. At
some point in Middle Moche times, the Collique
intervalley canals were constructed. Sipán would
have almost certainly controlled these canals to
link themselves and the Lambayeque Valley
Complex with the Zaña valley to the south (Shi-
mada 1994a). This was not simply about increas-
ing agricultural productivity. Considering the
political significance of water access in north
coast politics (Netherley 1984), canals embodied
deep statements regarding political power and
resource control. Sipán is geographically closer
to Huaca el Pueblo-Úcupe than Dos Cabezas
(Supplemental Figure 3), and it is reasonable to
hypothesize that Huaca el Pueblo was a satellite
of Sipán, extending their influence into the
Zaña valley. Many prominent items including
one of the Úcupe principal personage’s funerary
masks, V-shaped diadems, and other items
resemble those at Sipán (Supplemental Fig-
ure 4A, B, C). We also think it likely that the
Lord of Úcupe and the men in Sipán Tombs 2,
3, and 14 all shared the role of Figure B/D in
 16 LATIN AMERICAN ANTIQUITY

Figure 8. Comparison of features from Dos Cabezas (left column) and Huaca el Pueblo (right column) regarding overall
form and layout (a, b), zigzag walls (c), X-shaped low relief wall decoration (d), and adobe brick makers’ marks (e). All
are absent at Sipán. Photos by Steve Bourget. (Color online)

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 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS
the Moche Presentation Theme. Similar associa-
tions are absent at Dos Cabezas.
Beyond this, an Úcupe-Dos Cabezas connection
is far more obvious. The iconography and design of
some grave goods tie Úcupe to Dos Cabezas, such
as ceramic representations of the so-called Moche
“engineer,” diadems with the “Crested Animal/
Seahorse,” and the similarities between the style
and design of other masks worn by the Lord of
Úcupe and the personage of Dos Cabezas Tomb
2 (Supplemental Figure 4D, E).
Architecturally, Huaca el Pueblo is a direct
analog of Dos Cabezas (Figure 8). Although
Dos Cabezas is twice the size of Huaca el Pueblo,
their layouts and cardinal orientations are identi-
cal. They share identical superimposed stepped
platform construction, a zigzag platform mound
wall design, and identical adobe brick molds
with shared “makers’ marks.” A long wall from
the northwest corner of the platform turns north
to delimit the border of the main plaza at
Huaca el Pueblo and Dos Cabezas. The same
wall at both sites is decorated with low-relief
X-shaped geometric murals with a white dia-
mond on a black background (Bourget 2014).
Dos Cabezas and Úcupe lords were buried in
adobe brick chamber tombs adjacent to the pri-
mary platform mound. Placement of a retainer
beneath a principal personage is only seen at
Dos Cabezas and Úcupe. Not one of these shared
features is present at Sipán.
Huaca El Pueblo seems to have been a satel-
lite of Dos Cabezas. Did it represent a northern
expansion of the Jequetepeque Moche to secure
water sources, subjects, tribute labor, and culti-
vatable land via implantation secondary center
at Úcupe? This may be, but objects in the tomb
of Lord of Úcupe demonstrate that something
even more complicated unfolded here. At least,
this lord appears literally and figuratively
between two worlds, and we may be glimpsing
some kind of hybrid sociopolitical identity
bridging the Moche to his north and south.
Further, it is important to recognize that
Andean concepts of territory differed starkly
from Western notions. Ethnohistorical sources
describe how communities of one lord lived in
the “territory” of another (Rostworowski 1985).
Whatever was going on, it may represent some-
thing far less predatory. A better understanding
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17
of who built the Collique intervalley canals,
and exactly when this happened will be import-
ant to explore this issue.
Moche Macropolitics. In terms of the larger
political landscape, three considerations emerge.
First, negative residual values lend support to a
hypothesis of elite endogamy and minimal verti-
cal gene flow coming from other (lower) social
groups or strata—in other words, socially allo-
patric relationships driving observed heterogen-
eity (see Klaus et al. [2017] for a comparative
Middle Sicán study). Perhaps echoing this,
elite/nonelite differences were expressed in early
postcontact north coast mythology that justified
social inequality via cosmology (Rowe 1948).
Second, cluster analysis produced distinctive
Sipán and Dos Cabezas-dominated groups
along with independent demonstration of large
(heterogeneous) FST for the sample. Further cor-
responding subsample FST variation, residual
variation, and principal components analyses
results are equally consistent with expectation
of a “many Moches” model that anticipates fairly
explicit internal political differentiation and
social and structural diversity (Benson 2010,
2012; Quilter and Koons 2012). This also reso-
nates with Donnan’s (2011) vision of one or
more Moche royal courts in the various north
coast valleys and who marked distinctive group
identities with divergent artistic substyles.
Shimada (2010:72) further considered how
Moche monuments were the seats of semi-
autonomous elite lineages in each valley, loosely
tied to each other through marriage alliances, gift
exchange, and reciprocal obligations focused by
a shared elite material culture and politico-
religious system. Here, a footprint of between-
lineage intermarriage among lower nobility
seems apparent. Diachronic coherence of these
lineages probably also reflects lasting and stable
forms of kinship-based rule. Such would favor
dispersed authority and power across a social
landscape and not unstable charismatic kingship
(Shimada 2010).
Third, the growing consensus regarding rela-
tive ideological and cultural unity among
Moche lords in these diverse valleys does not
necessarily imply biological unity. Structurally
speaking, each northern valley likely possessed
 18
at least one elite administrational component and
they were “Moche” in their own ways. Con-
structed on principles of kinship, they produced
degrees of solidarity among those who managed
the social, economic, and ceremonial functions
of local society/societies in that valley—particu-
larly among the lower nobility. We think our data
resonate with Benson’s (2010) model involving
multiple lordships that involved elite competition
and cooperation. Suzerainty (where a dominant
polity permits subordinates extensive autonomy
and sovereignty) and reciprocity-driven cooper-
ation must also be investigated further.
Golte (1994) considered the Moche repre-
sented relatively independent political entities
participating in overarching economic spheres,
trade networks, and production of sumptuary
elite goods most consistent with decentralized,
competitive polities further preconditioned by
geography (Benson 2012). Each north coast
river valley is circumscribed as mostly individual
river drainages. This physiogeography likely
encouraged development of distinct spatial sys-
tems resulting in self-generating and variable
expressions of sociopolitical self-sufficiency and
autonomy (Benson 2012:27). In the north, there
are V-shaped headdresses as Sipán, the “hat
with bangles” in the looted Vicús metalwork,
and associations with Figure B/D of the Presenta-
tion Theme among Sipán and Úcupe lords. In the
south, there are multipointed crowns atop the
heads of figures in the Huaca de la Luna and
Huaca Cao Viejo Complex Theme murals, and
at Huaca de la Luna, priests are associated with
“flying feline” back ornaments. We strongly sus-
pect that these are indexical of distinct elite
groups, lineages, and identities within the broader
constellation of a shared elite Moche system (Ben-
son 2012). This harkens back to Berezkin’s
(1978) iconographic analysis where he argued
that prior to late Moche IV, the five sets of princi-
pal deities/supernatural actors were representative
of five principal elite lineages. Perhaps this work
has observed the biodistance signatures of two
such lineages.
Conclusions
Not long ago, Alva (2001:243) observed that the
nature, scope, and extent of the Sipán dynasty in
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LATIN AMERICAN ANTIQUITY
particular—and the rest of the elite Moche in
general, was unknown—as was the nature of
interactions of lords between valleys and the
extent of Moche political centralization. This
work represents a first step in addressing these
questions using the phenotypic record of inher-
ited dental traits. The data support the inference
that the Sipán and Dos Cabezas lords were mem-
bers of different dynastic kin groups generally
concordant with a “many Moches” model of pol-
itical organization, where these groups were linked
on some levels but followed their own distinct tra-
jectories, agendas, and approaches to alliance-
building—inconsistent with neo-evolutionary
visions of a single hegemonic Moche state.
In this and other work (Klaus et al. 2017; Sut-
ter and Castillo 2015), dental phenotypic vari-
ation is increasingly shown as a highly useful
approach in the archaeology of the north coast
of Peru. In the future, increased sample size
and expanded geographic scope can further
define the biological record of social interaction
patterns and politics throughout Moche realms.
We can further test the concept of a northern
north coast/southern north coast geopolitical,
ethnic, and linguistic bipartition (Rivet 1949;
Shimada 1994b) and if these factors were bar-
riers to gene flow. What might biodistance ana-
lysis data reveal about the maintenance of
non-Moche sociopolitical identities at places
such as the Virú and the Santa valleys (Bourget
2010; Chapdelaine 2008)? Was politically moti-
vated intermarriage with the Moche either
spurned or encouraged in their political calcu-
lus? Regional macropopulation structure and
evolutionary processes can be pursued further
(Klaus 2008), as well as defining the likely
coexistence of finer-scale regional identities
and interactions. That is, Sipán and Dos Cabezas
may not have ruled their entire respective
regions, and their interactions with other peers
and subordinates await a clearer understanding,
as does regional FST variation. Excavation of
contemporaneous commoner skeletal samples
will go far to meet these goals.
These findings represent a starting point and
provide new hypotheses for the study of Moche
biology, population structures, and politics.
Future work must consider the fluidity, symbo-
lisms, and situational dimensions of these
 [Klaus et al.] BIOLOGICAL DISTANCE PATTERNS AMONG THE NORTHERN MOCHE LORDS
relationships. Furthermore, the promise of next
generation ancient DNA and proteomic sequen-
cing technologies will no doubt expand our hor-
izons in ways we cannot yet anticipate in the
ongoing effort to reconstruct the history, struc-
ture, and legacy of one of the most remarkable
societies of the ancient South America.
Acknowledgements. We owe debts of gratitude to Christo-
pher Donnan, Ceyra Pasapera, Kimberly Jones, and the staffs
of the Museo Tumbas Reales and the Museo del Sitio de
Huaca Rajada for their numerous contributions. Joe Luce,
Marisa McKane, and JoEllen Perez assisted in data collection
between 2009-2011. Data collection was supported by grants
from Utah Valley University. Dan Temple provided expert
assistance with the RMET software. Chris Donnan, Clark
Larsen, Charles Merbs, Izumi Shimada, and Dan Temple pro-
vided valuable critical commentary and editorial input on
earlier drafts of this article.
Notes
1. The high positive extralocal gene flow among the
Úcupe retainers may signify very different origins for these
three people, but we are certain that their small subsample
size generated a spurious subsample FST value and cannot
be considered any further in the discussion of the R-matrix
results. In one alternate data analysis run, their removal chan-
ged the overall bias-corrected FST value to 0.40 producing a
slightly different but still very unusually high result.
2. This may imply that the shared growth abnormality
was not an inherited endocrine abnormality but indeed an
acquired disorder (Biers et al. 2017).
Data Availability Statement. The raw biodistance datasets
used in this study are digitally curated by HDK and can be
shared with interested scholars.
Supplementary Materials. For supplementary materials
accompanying this article, visit https://doi.org/10.1017/laq.
2018.46.
Supplemental Figure 1. View of the superimposed funer-
ary platforms at Sipán, demonstrating the complex building
history of the structure and tomb locations in the northern
(left) and southern (right) halves of the platform. Illustration
by the Museo Tumbas Reales de Sipán.
Supplemental Figure 2. Retainer burials in the tomb of the
Lord of Úcupe: Burials 1, 3, and 4 (A, B, and C, respectively).
Illustrations by Bruno Alva.
Supplemental Figure 3. Geographic locations and dis-
tances between Sipán, Úcupe, and Dos Cabezas. Map by
Haagen Klaus.
Supplemental Figure 4. Various items in the tomb of the
Lord of Úcupe bear strong stylistic and iconographic resem-
blance to items discovered at Sipán to the north, such various
V-shaped diadems with outstretched hands (A); pectoral
necklace manufactured from Spondlyus spp. and Conus sp.
shell (B), and spiral earspool repoussé decoration (C).
Other items, such as a a blackware vessel of the so-called
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19
“Engineer” (D) and a diadem possessing “Crested Animal-
Seahorse” elements (E) appear to be connections to Dos
Cabezas to the south. Photos by Steve Bourget.
Supplemental Table 1. Northern Moche stylistic
sequence and associated inferred chronology.
Supplemental Table 2. Correlation and overlap between
Phase IV and Middle Moche subphases.
Supplemental Table 3. Stratigraphic distribution of tombs
in the Sipán funerary platform.
Supplemental Table 4. Chronometric date estimations for
Sipán and Dos Cabezas tombs.
Supplemental Table 5. ANOVA tests, metric data.
Supplemental Text 1. Statistical Procedures.
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