Finucane 2006

Human and animal diet at Conchopata, Peru: stable isotope
evidence for maize agriculture and animal management practices

during the Middle Horizon
Brian Finucane
a,
*
, Patricia Maita Agurto
b
, William H. Isbell
c
a
Research Laboratory for Archaeology and the History of Art, University of Oxford, Dyson Perrins Building, South Parks Road, Oxford, OX1 3QJ, UK
b
Facultad de Ciencias Biologicas, Labortorio de Fauna, Universidad Nacional Mayor de San Marcos, Lima, Peru
c
Department of Anthropology, State University of New York, Binghamton, NY 13901, USA
Received 20 December 2005; received in revised form 20 March 2006; accepted 23 March 2006
Abstract
This paper reports d
13
C and d
15
N values for human and animal skeletal remains from the Middle Horizon (AD 550e1000) site of Conchopata
in the Peruvian highlands. The data indicate that maize was the dietary staple for both humans and the majority of animals at this urban site.
Camelids at the site segregated into two groups according to d
13
C values, reecting two distinct animal management strategies. Human infants
were found to have elevated d
15
N values, reecting the trophic level effect of breastfeeding. No sex based differences in diet were observed.
2006 Elsevier Ltd. All rights reserved.
Keywords: Conchopata; Stable isotopes; Camelids; Wari; Middle horizon; Maize; Ayacucho; Andes
1. Introduction
1.1. Wari
The Wari polity (AD550e1000) was the rst urban and state
level society to emerge in the Andean sierra [23e25,53]. As
such, this society represents a valuable case study in the devel-
opment of prehistoric states. This investigation focuses upon
elucidating the economy of the Wari state through the stable iso-
topic analysis of the human and animal remains from the most
intensively studied Wari site, Conchopata. Specically this
investigation seeks to determine both the role maize played in
sustaining urban life in Ayacucho as well as the character of
the animal management practices of the Wari state.
1.2. Maize in the Andes
Throughout the Americas there is a strong positive correla-
tion between intensive maize cultivation and sociopolitical
complexity. Every urban and state level society in Mesoamer-
ica, from Monte Alban and the Classic Maya to Teotihuacan
and Aztec civilizations of Central Mexico, had economies
based on maize. In North America the highest level polities,
the Pueblo communities of the Southwest, the complex chief-
doms in the Southeast, and the Mississippi and Ohio River
Valleys all relied upon maize as the staple crop [51].
Archaeobotantical evidence from coastal Ecuador indicates
that maize had reached the Andes from Mexico by w4200 BP,
long after indigenous cultigens such as quinoa, beans and tubers
had come into use [62]. The exotic grain was adopted by peoples
who were already sedentary agriculturalists. Isotopic evidence
indicates that for over a millennium, the contribution of maize
to human diet in Ecuador was insignicant [62].
The role of maize in the economies of ancient Andean
states is contested. The dominant reconstruction of Andean
economies, as expounded by John Murra [37,40], emphasizes
the contributions of indigenous crops and relegates maize to
the position of ceremonial crop. Other archaeologists consider
maize to be signicant primarily within the context of political
economies of Inka and pre-Inka societies [20,21,24,26,53].
States may have nanced their activities through chicha
* Corresponding author. Tel.: 44 1865 285213; fax: 44 1865 285220.
E-mail address: brian.nucane@merton.ox.ac.uk (B. Finucane).
0305-4403/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2006.03.012
Journal of Archaeological Science 33 (2006) 1766e1776
http://www.elsevier.com/locate/jas
(maize beer) fueled fests, but maize did not constitute a dietary
staple. The argument that maize cultivation played a decisive
role in the development of Andean states has not been widely
accepted [68].
1.3. Camelid pastoralism
Animal husbandry has been central to reconstructions of
prehistoriceconomyinthe Andes. Thedominant model, variously
termed verticality and ecological complementarity emphasizes
the movement of bulk resources between attitudinally dened
ecological zones and the direct control of communities over
these resources [36e40,69]. In the schema generated from eth-
nographic and ethnohistoric data, mixed llama and alpaca herd-
ing is coupled to tuber cultivation on the high altitude grassland
known as puna (>4100 m) and is spatially segregated from
maize cultivation in the quechua (2600e3400 m) and yunga
(2200e2600 m) zones [15,16,22,34,38,43]. As with lower ele-
vations, the wild ora of the puna is a C
3
plant community, typ-
ied by the perennial grasses Festuca, Calamogrostis, Stipa, and
Poa [15,61]. Within the puna there are wetlands or bofedales,
characterized by Distichia muscoides [67]. Despite the punas
thin air, frigid temperatures and intense solar radiation, the
region is extremely productive as rangeland for camelids (a
generic term applied to the species Lama glama, L. pacos,
L. guanaco and Vicugna vicugna) and for the cultivation of the
bitter potato (Solanum curtilobaum) which is freeze dried for
storage and transported as chun o. Animals are managed through
a rangestock strategy, with little if any foddering and their
manure is used to fertilize plots of tubers.
Yet, puna pastoralism was not and is not the only form of
camelid husbandry in the Andes. Alternative strategies of cam-
elid husbandry are documented ethnographically and archaeo-
logically. McCorkle [34] describes herding strategies at the
village of Usi in southern Peru. At the upper margin of maize
agriculture at 3700 m, the community utilizes a form of com-
pressed verticality by controlling lands ranging from 3200e
5000 m. Although some camelid herds are kept permanently
at puna estancias, most herds are instead kept close to the vil-
lage, being driven to pasture during the day and corralled at
night. During the dry season, the animals are grazed on the stub-
ble of elds. In all the animals spend nine months grazing inten-
sively between 3500e4000 m. Only when all available fodder is
exhausted are the animals driven to the puna to graze.
1.4. Stable isotope analysis
The study of paleodiet through stable isotope analysis
proceeds from the experimental observation that the isotopic
composition of animal tissues generally reects that of the
diet they consume (for a review see ref. [52]). Such analyses
utilize the variation in the ratios of the stable isotopes of car-
bon and nitrogen within ecosystems to measure the relative
contribution of different resources to animals diets [11,12].
The stable isotopes of carbon (
13
C and
12
C) and those of
nitrogen (
15
N and
14
N) are expressed in per mil (&) as d values:
d R sample=R standard 1 1000
where R
13
C/
12
C for the measurement of carbon and
15
N/
14
N for the measurement of nitrogen. The standards to
which samples are compared are the limestone Vienna PeeDee
Belemnite (VPDB) and atmospheric nitrogen (AIR) for carbon
and nitrogen respectively. As most terrestrial materials, includ-
ing plant and animal tissues contain less
13
C than VPDB, d
13
C
values are typically negative.
The fundamental variation in the ratios of stable carbon iso-
topes in terrestrial food webs stems from differences in the
photosynthetic pathways of plants. The overwhelming major-
ity of plants utilize the Calvin Cycle (C
3
) and have tissues with
an average d
13
C value of 26.5& [63]. The d
13
C values of
plants relying upon the Hatch-Slack pathway (C
4
), mainly
tropical grasses including such domesticates as maize, millet,
and sugarcane, are much higher, averaging 12.5& [63].
Plants utilizing a third pathway, Crassulacean acid metabolism
(CAM) have d
13
C values ranging from 27& to 12&. C3
ora are dominant in the Central Andean region [57].
The carbon isotope signature of animal proteins, such as
collagen in bone and dentin and keratin in hair and nails, pre-
dominantly reect the protein component of diet, as essential
amino acids are directly routed from diet to incorporation in
body tissue [30]. However, the relationship between dietary
protein and the isotopic composition of tissues is not one to
one. The d
13
C values of the collagen of large herbivores are
more positive by w5& relative to their dietary average
[30,63]. The collagen of carnivores in turn, reects the isoto-
pic signature of the animal protein in their diets, ultimately
derived from the plants eaten by primary consumers. The iso-
topic signature of omnivore collagen represents the contribu-
tion of both plant and animal protein to diet.
Further variation in d
13
C values is introduced by marine
and freshwater ecosystems. Marine sh tend be enriched in
13
C, with isotopic values resembling those of C
4
plants
whereas freshwater sh have variable isotopic signatures, pri-
marily reecting the differences in the concentration of dis-
solved inorganic carbonates which are the carbon source of
many aquatic plants [7,41,49].
Analysis of nitrogen stable isotopes provides additional res-
olution in the reconstruction of paleodiet by discriminating
between plant and animal protein sources, and distinguishing
marine from terrestrial protein [48,65]. Apart from nitrogen
xing legumes, terrestrial typically plants have d
15
N values
between 2 to 6&. The nitrogen isotope ratios of consumers,
exhibit trophic level enrichment, as the d
15
N values increase
3 to 4&with each step up in the food chain [12,49,50]. Marine
plants are w4& enriched in
15
N relative to terrestrial plants.
This oral enrichment, in conjunction with the complexity
of aquatic food webs, results in the
15
N enrichment of marine
sh and mammals.
Most of the mass of osseous remains, 70% in bone and
tooth dentin and 98% in tooth enamel, is comprise of the min-
eral hydroxylapatitite (Ca
10
(PO
4
)
6
(OH)
2
). Structural carbonate
occurs when -CO
3
substitutes for phosphate and hydroxyl in
the apatite crystals [28,31]. This carbonate is derived from
1767 B. Finucane et al. / Journal of Archaeological Science 33 (2006) 1766e1776
blood bicarbonate generated by cellular metabolism and is on
average enriched in
13
C by w9& relative to diet [3,4].
Whereas, the isotopic composition of collagen chiey reects
the growth substrate, (i.e. protein), carbonate represents the
contributions of all dietary components [3,58].
The difference between the isotope values of bone carbonate
and collagen (D
13
C
carb-coll
) will vary depending on whether an
individual consumes a monoisotopic diet or polyisotopic diet.
Organisms consuming protein and carbohydrates with approxi-
mately equal d
13
C values will exhibit D
13
C
carb-coll
values of
w4.4&. Organisms consuming protein with a d
13
C value less
negative than bulk diet, as in the case of marine protein and C
3
carbohydrates, will exhibit D
13
C
carb-coll
values of <4.4&.
D
13
C
carb-coll
values >4.4& result from diets incorporating
protein with a d
13
C value more negative than whole diet, the
scenario expected in a population consuming maize as a die-
tary staple and protein of terrestrial origin [3,4].
Whereas, the carbonate in bone is subject to diagenetic alter-
ation in the postmorteminterval, enamel with its greater density
and crystal size is less prone to modication [28,29]. In contrast
to bone collagen which is remodeled throughout life, enamel
and primary dentin are deposited during discrete periods early
in life and do not undergo subsequent remodeling [46].
1.5. Site description
Conchopata is located at 74
12 west, 13
9 south at an
elevation of 2700 m in the highlands of central Peru (Fig. 1).
It is situated atop a ridge w2 km northeast of the city center
of modern Ayacucho. The site lies within the ecological region
Fig. 1. Map of the Ayacucho Valley showing ecological zones, archaeological sites and modern towns. Adapted from Anders [5, Fig. 1] .
known as quechua, a vegetational zone of thorn scrub forest
[32,61]. As the label implies, the wild ora of this zone is
comprised mainly of shrubs and cacti (CAM plants), such as
Opuntia sp., Agave Americana, Agracia macaranta, Cesalpina
spinosa, and Fourcroya anima.
Although the urban core of Conchopata is thought to have
covered over 20 ha, modern construction has reduced the area
of architectural remains to 3 ha [44]. Radiocarbon dates indi-
cate the earliest burials at Conchopata predate the Wari state,
though the majority of the sites extant architecture pertains to
the Middle Horizon [23]. Planned architecture at Conchopata
attributed to the Wari state includes a perimeter wall, two patio
compounds, several D-shaped temples, and two plazas [23].
Amongst these planned architectural complexes are the re-
mains of a dense network of domestic and mortuary structures.
2. Methods
Sample preparation and analysis were conducted at the
Research Laboratory for Archaeology and the History of Art
at Oxford University. Collagen was extracted from both faunal
and human samples following procedures described in Richards
and Hedges [45]. Isotopic analysis was conducted using a Carlo
Erba 1108 carbon and nitrogen elemental analyzer coupled to
a Europa Geo 20/20 mass spectrometer in continuous ow
mode. The isotopic values of all samples were measured relative
to laboratory standards of nylon and alanine whose isotopic
values are calibrated with respect to IAEA standards. All sam-
ples were analyzed in triplicates in separate batches. Analytical
errors are of the order of 0.2&for d
13
C and d
15
N.
Samples of tooth enamel were prepared for carbonate
analysis by rst separating a chip of enamel from the underly-
ing dentin using a Dremel tool and then shot blasting the frag-
ment with aluminum oxide powder to remove the exterior
surface (possibly subject to diagenetic alteration) and any re-
maining adhering dentin. The enamel samples were then
ground into powder using an agate mortar and pestle in the
presence of methanol to prevent the ejection of fragments
from the mortar. The methanol was subsequently evaporated
in a drying oven at 70

C and the remaining powder collected.
Samples of enamel were analyzed at the Department of Earth
Sciences, Oxford University. Powdered samples of enamel were
reacted with 100% H
3
PO
4
at 90
in a common acid bath for

6 min. Isotopic measurements were made using a VG Prism
IRMS with an analytical precision of better than 0.1&.
Faunal samples from Conchopata were analyzed to provide
a dietary baseline. Forty samples were selected to represent
taxa known ethnographically and archaeologically to have
been major sources of animal protein in Ayacucho Valley
(see Table 1 and Fig. 2). These include camelids: both llamas
(Lama glama) and alpacas (Vicugna pacos), (difcult to distin-
guish osteologically), the domestic guinea pig (Cavia porcel-
lus) and the leaf mouse (Phyllotis). Both guina pigs and
mice are commensal species and should therefore act as indi-
cators of the resources available in the anthropogenic environ-
ment of Conchopata. 40 samples of human bone and tooth
were selected from 38 individuals (see Table 2 and Fig. 3).
In order to obtain human samples representing both sexes
and a range of age categories, osetological data collected by
Tung [70,71] were consulted. The lead author of this article
Table 1
Archaeological contexts and isotopic values of Conchopatas faunal samples
Sample EA Locus Taxon Bone d
13
C
(&)
d
15
N
(&)
C:N
CI1A 73T 1649 Camelid Femur 18.9 5.9 3.21
CI3A 142 2693 Camelid Humerus 9.7 7.5 3.19
CI7A 205 3512 Camelid Metapodial 9.0 6.3 3.33
CI14A 40B 3141 Camelid Rib 11.2 9.6 3.20
CI19A 44A 2592 Camelid Femur 11.2 6.4 3.22
CI22A 65 3034 Camelid Femur 11.9 4.6 3.16
CI23A 104T5 2092 Camelid Phalanx 16.8 5.0 3.21
CI24A 6 2806 Camelid Metapodial 9.8 6.6 3.19
CI26A 168 3039 Vicugna
pacos
Mandible 19.2 8.6 3.28
CI27A 168 3039 Camelid Mandible 9.2 6.4 3.22
CI37A 82 2651 Camelid Rib 9.3 7.6 3.21
CI38A 73N 1604 Camelid Femur 19.5 8.0 3.21
CI39A 77 1550 Camelid Femur 8.2 5.1 3.21
CI40A 73S 1550 Camelid Tibia 18.8 8.9 3.26
CI41A 73S 1550 Camelid Tibia 18.5 5.0 3.23
CI8A 40B 3115 Cavia
porcellus
Tibia 11.3 9.3 3.17
CI9A 40B 3115 C. porcellus Tibia 7.4 8.9 3.22
CI10A 40B 3118a C. porcellus Femur 9.9 8.4 3.21
CI12A 40B 3118a C.porcellus Femur 18.6 8.2 3.26
CI15A 40B 3094 C. porcellus Femur 11.8 9.1 3.18
CI16A 40A 3130 C. porcellus Femur 7.0 10.3 3.23
CI17A 31 3106 C. porcellus Tibia 12.6 7.8 3.24
CI34A 82 2651 C. porcellus Femur 8.2 8.1 3.27
CI36A 82 2651 C. porcellus Femur 8.5 8.1 3.32
CI20A 77 1956 Phyllotis sp. Femur 11.9 8.6 3.17
CI21A 77 1956 Phyllotis sp. Humerus 12.9 15.5 3.16
CI35A 82 2651 Phyllotis sp. Femur 11.4 10.2 3.4
Fig. 2. Butchered camelid remains, source of sample CI7A.
is responsible for the nal sex and age estimates provided in
Table 2.
Only samples of bone with collagen yields of >2% were
considered in this study. Collagen with C:N ratios outside the
2.9e3.6 range were also excluded, as such samples are likely to
have undergone diagenetic alteration [9]. Collagen samples
with low concentrations of carbon and nitrogen (below 30% and
10%by mass respectively) were also excluded, as lowconcentra-
tions of these element correlates with diagenetic alteration [1].
3. Results
The isotopic values of individual human samples are prese-
nted in Table 2, and the descriptive statistics for bones and teeth
are presented in Tables 3 and 4 respectively. The carbon stable
isotope values of the human collagen from Conchopata range
from 13.2& to 8.2& with an average value of 10.7&.
The carbonates in human tooth enamel have a mean d
13
C value
of 3.9&. The mean difference between paired samples of
enamel carbonate and dentin collagen (D
13
C
Enamel-dentin
) is
7.2&. This spacing between carbonates and collagen indicates
that Conchopatans consumed a diet with an energy component
more enriched in
13
C than the protein component.
Five samples of collagen, CI16H, CI44H, CI50H, CI69H
and CI72H, all from the bones and teeth of infants exhibit
anomalously high d
15
N values, in excess of 13& (Fig. 4). In
t test comparison of these infants with all other humans, the
infants are signicantly more enriched in
15
N (P < 0.005).
The isotopic signatures of males and females are not signi-
cantly different (t-test, P < 0.05).
The isotope values for individual faunal samples are pre-
sented in Table 1 and summarized by taxa in Table 5. The
Table 2
Archaeological contexts and stable isotope values of human samples from Conchopata, including bone and dentin collagen as well as enamel carbonate
Sample EA Locus Burial Sex Age (years) Bone/tooth d
13
C (&)
collagen
d
15
N (&)
collagen
d
13
C (&)
enamel
carbonate
C:N
collagen
CI4H* 104T5 2107 17 F 50 Femur 11.2 10.5 3.25
CI5H* 104T5 2107 17 F 50 LM
3
11.3 11.1 1.2 3.35
CI8H 205 3521 107 F 50 RPM
2
13.2 10.4 3.7 3.34
CI10H 208 3547 108 F 35e40 LM
3
10.8 11.9 4.0 3.23
CI19H 105 2095 98.2 F? 23e27 RM
2
4.4
CI20H 105 2095 F 20e24 LM
3
10.6 11.1 3.7 3.20
CI23H 178 3148 95.1 M 21e30 LM
2
4.0
CI24H 178 3148 95.4 F? 15e18 LM
2
11.2 8.3 3.8 3.22
CI27H 178 3148 95.3 M 21e25 LM
2
4.0
CI29H 179 3207 98.1 M 22e24 Pubis 10.4 12.3 3.44
CI32H 151 2858 50 F? 30e35 RM
2
3.8
CI33H 151 2858 50 F? 30e35 Skull 9.8 8.7 3.22
CI34H 100 1339 8 ? 6e7 Rm
2
3.9
CI36H 150 3067 42 ? 11e14 Rib 8.5 11.9 3.3
CI39H 147 2884 42 ? 6e7 RM
1
6.0
CI40H 147 2884 42 ? 6e7 Skull 10.7 10.5 3.26
CI43H 178 3148 95 ? 18e20 Femur 9.9 9.8 3.21
CI45H 187 3335 101 ? 3e4 Lm
2
11.7 12.4 3.9 3.27
CI46H 187 3335 101 ? 3e4 Femur 12.5 10.2 3.27
CI47H 187 3335 101 F 22e28 Os Coxa 11.6 10.5 3.17
CI52H 89A 2052 M 35e45 Os Coxa 8.2 10.2 3.19
CI56H
44A 950 M 18e22 RM

3
9.4 11.2 4.5 3.17
CI57H
44A 950 M 18e22 Skull 10.4 10.3 3.23

CI58H 110 2014 ? 20e35 Femur 11.2 9.0 3.31
CI59H 110 2112 M? Adult Femur 11.3 11.0 3.39
CI61H 110 1993 ? 3e7 Femur 11.5 10.2 3.2
CI62H 39 1434 ? Adult Humerus 11.9 8.8 3.24
CI64H 40A 3097 83.4 M 35e50 Humerus 12.3 10.3 3.29
CI67H 39C 1818 ? Adult 1st MT 8.4 12.9 3.23
CI49H 179 3224 98.2 ? 15e21 months ri
1
3.6
CI16H 179 3224 98.2 ? 15e21 months lm
2
10.2 15.3 3.3 3.27
CI72H
!
205 3521 105 ? 15e18 months Rib 11.9 13.8 3.19
CI74H
!
205 3521 105 ? 15e18 months rm
2
5.9
CI50H 179 3224 98.2 ? 15e21 months ri
1
9.2 13.8 3.3 3.24
CI69H 205 3521 104 ? 10e12 months Rib 9.6 15.1 3.23
CI71H 205 3521 105 ? 10e12 months rm
2
4.1
CI44H 187 3335 101 ? 18e30 months lm
2
9.5 13.8 3.8 3.2
CI41H 147 2884 42 ? Neonate Skull 10.7 11.8 3.38
CI60H 110 2112 ? Neonate Ulna 9.8 11.5 3.2
CI11H 208 3547.1 108 ? Fetal Tibia 10.6 11.1 3.19
Three individuals (indicated by *,

, and
!
respectively) are represented by both bone and tooth samples.
average d
13
C values of the phyllotis samples, CI20A, CI21A,
and CI35A is 12.1& (s 0.8). CI21A has a high d
15
N
value of 15.5&, whereas CI20A and CI35A have d
15
N value
of 8.6& and 10.2& consistent with theoretical values for ter-
restrial herbivore collage.
The fteen cavia samples have d
15
N values that varied from
6.5&to 10.3&, all within the observed range for terrestrial om-
nivore collagen (mean 8.3&, s 0.9) [12,66]. The d
13
Cvalues
are more variable, ranging from 18.6&up to 7.0&, with an
average value of 11.0&(s 3.6). The variance of the guinea
pig carbon isotope values is signicantly greater than that of
humans (F-test, P < 0.005).
The seventeen camelid samples exhibit diversity in their
isotopic values similar to those of cavia (Fig. 4). The d
15
N
values of the camelid samples range from 4.6&up to a maxi-
mum of 9.6&with a mean value of 6.6&(s 1.5). Such var-
iable d
15
N values for camelid tissue are consistent with
experimental data from control feeding studies of camelids
and reect physiological adaptation to aridity [2,55].
Conchopatas camelids segregate into two discrete sub-
groups according to d
13
C values (Fig. 4). The rst group, com-
posed of samples CI3A, CI4A, CI5A, CI7A, CI14A, CI19A,
CI23A, CI24A, CI27A, CI37A, and CI39A has a mean d
13
C
value of 10.0&, whereas the second group represented by
samples CI1A, CI22A, CI26A, CI38A, CI40A, and CI41A
have mean a d
13
C value of 18.6&. The difference between
the carbon isotope values of these two groups is statistically
signicant (t-test, P < 0.005).
Both the cavia and the camelids are signicantly depleted
in
15
N relative to Conchopatas humans (t-test, P < 0.005).
4. Discussion
4.1. Humans
The nitrogen values of Conchopatas humans are consistent
with terrestrial omnivores indicating the sites occupants con-
sumed a substantial quantity of animal protein. This conclu-
sion is consonant with the isotopic spacing between human
and animal d
15
N values (w4&). Although the observed nitro-
gen isotope signatures could result from the consumption of
marine protein, this is extremely unlikely considering the sites
distance from the ocean and the high transportation costs asso-
ciated with transporting sh or shellsh to the sierra from the
Pacic. Furthermore all excavated deposits were passed
through 1 cm screens and the only objects of maritime origin
recovered were shell, almost exclusively Spondylus sp. a pres-
tige good imported from Ecuador and not a foodstuff.
The high d
15
N values of ve individuals at Conchopata
reect a trophic level shift between breast milk and the body tis-
sues of suckling infants (Fig. 4). The individuals with elevated
d
15
N values, CI16H, CI44H, CI50H, CI69H and CI72H were
all infants at the time of death and were isotopically predators
upon their mothers. Fetal and neonate remains (<1 year) from
the site, CI11H, CI60H, and CI41H, do not exhibit such elevated
15
Nlevels in their collagen, as their tissues formed in utero prior
to breast feeding and therefore directly reect maternal diet. The
enrichment of these infants is consistent with the ndings of
previous paleodietary research and studies of modern breastfed
infants [17,18,27].
The high d
13
C values of human collagen and carbonate
from Conchopata reect a maize based diet. The high
D
13
C
Enamel-dentin
values for the teeth analyzed indicate that
bulk diet was more enriched in
13
C than dietary protein, at
least during childhood when these teeth formed.
Even though highly improbable, the consumption of marine
protein could conceivably result in the observed carbon isotope
Fig. 3. Suboor cist tomb containing the remains of an adult female (CI8H)
and two infants (samples CI71H, CI72H, and CI74H).
Table 3
Descriptive statistics for the stable isotope of bone collagen
d
13
C (&)
bone collagen
all
d
15
N (&)
bone collagen
non-infant
d
15
N (&)
bone collagen
infant
N Mean 1s N Mean 1s N Mean 1s
21 10.6 1.2 16 10.7 1.2 5 14.4 .7
Due to the trophic level elevation resulting from breast feeding, the mean d
15
N
for infants is shown separately.
Table 4
Descriptive statistics for the isotopic composition of dentin collagen and enamel carbonates
d
13
C (&) dentin collagen all d
15
N (&) dentin collagen
non-infant
d
15
N (&) dentin collagen infant d
13
C (&) enamel carbonate D
13
C
Enamel-dentin
(&)
N Mean 1s N Mean 1s N Mean 1s N Mean 1s N Mean 1s
10 10.7 1.2 6 10.6 1.1 4 13.8 1.2 19 3.9 0.98 10 7.2 1.6
The nitrogen isotope values of primary dentin deposited during infancy is presented separately, due to the trophic level effect of breastfeeding.
values for collagen. But the d
13
C values of the enamel carbon-
ates indicate that bulk diet, not just protein was enriched in
13
C. A pure CAM diet could potentially produce the observed
carbon isotope values. However, the only indigenous CAM
food consumed in Ayacucho today is tuna, the fruit of the prickly
pear (Opuntia sp.). Tuna is not a staple now, nor is it likely to
have been one in the past, as cactus horticulture is not capable
of sustaining dense urban populations.
Therefore the most plausible explanation is that maize was
the mainstay of human diet at Conchopata. Indeed, if 17&
and 3& are taken to represent the d
13
C values for the bio-
genic carbonates of pure C
3
and pure C
4
consumers respec-
tively, then Conchopatans appear to have eaten diets
consisting almost entirely of maize and maize fed animals.
The conclusion that maize was a dietary staple at Conchopata
is strongly supported by the isotopic values of the more numer-
ous faunal samples (see below). The stable isotope composition
of human and animal collagen at the Wari town of Conchopata,
reopens a long standing debate amongst Andean archaeologist
regarding the role of maize in complex societies of prehispanic
Peru. Contrary to the assertions of Murra [39,42] maize was not
merely a ceremonial crop which gained prominence only in the
Inka era. Rather, maize was the single most important food for
this urban society in the Andean sierra.
4.2. Camelids
The divergence between the d
13
C enriched and the d
13
C de-
pleted Conchopata camelids, as well as between Conchopatas
enriched animals and depleted camelids from both archaeolog-
ical and modern contexts, reects the practice of two distinct
forms of animal husbandry in the ancient Andes (Fig. 5).
The rst strategy of animal management, typical of CI1A,
CI22A, CI26A, CI38A, and CI40A as well as most of the
non-Conchopatan animals is rangestock grazing on puna pas-
ture characterized by C
3
ora. The second system of animal
husbandry, the approach typical of the majority of Conchopa-
tan camelids analyzed, is maize foddering. The uniqueness of
this second approach, to settlements such as Conchopata and
the Middle Sican center of Pacatnamu, indicates that the man-
agement of camelids in an urban context within a state level
polity was different than that of modern Andean herders [64].
Animals managed under a rangestock strategy are grazed in
agriculturally marginal areas. The abundant ethnographic liter-
ature on Andean pastoralists emphasizes the division between
the herders of the puna and the farmers of the quechua zone
[15,16,22,34,43,59]. The two are distinct, yet complementary
elements of Andean society characterized by dual organization.
As a result of the physical separation of herds and elds, animal
forage consists of wild, predominantly C
3
ora. The d
13
C
values of the non-Conchopata camelids, as well as Conchopata
samples CI1A, CI22A, CI26A, CI38A, CI40A, and CI40A are
consistent with this style of rangestock animal management.
The rangestock diet, characteristic of modern puna animals,
is more typical of alpacas raised on high grasslands than of
llamas. Contemporary camelid husbandry, characterized by
composite llama-alpaca herds, is an anomaly [54,60]. As a
result of competition with Old World domesticates (swine,
cattle, and sheep) llamas have largely been relegated to the
high altitude refugia of the puna. Prior to the introduction of
these exotic species, the range of llamas extended from the
highlands to the coast. Puna pastoralism in this period was
instead focused upon the more environmentally restricted
alpaca. As these animals were raised primarily for their
wool, the cold puna which promotes coat growth was the pre-
ferred environment [19,60]. Although alpaca products, such as
wool and charki, were probably common at lower elevations,
the animals themselves were not. Thus, generalized camelid
herding arose only recently with the disruption of more spe-
cialized strategies of animal management.
When the ecological differences between these species are
considered, in conjunction with optimal animal management
strategies, the potential for dietary differences becomes ap-
parent. Although it is difcult to distinguish llamas from al-
pacas osteologically by any criteria apart from size, it is
signicant that the only camelid positively identied as an al-
paca, CI26A, had an almost entirely C
3
puna-type diet.
0
2
4
6
8
10
12
14
16
18
-25 -20 -15 -10 -5 0
1
5
N

(
)

A
I
R

13
C () VPDB
Camelids Guinea Pigs Mice Infants
Neonates, Children, Juveniles and Adults
C
3
Plant
Consumers
C
4
Plant
Consumers
Fig. 4. Carbon and nitrogen stable isotopic values of human and animal bone
collagen from Conchopata. Infants (age 1e3 years) are shown separately from
all other human age categories.
Table 5
Descriptive statistics for the isotopic composition of animal collagen
Phyllotis sp. (n 3) Cavia porcellus (n 15) Camelids
d
13
C (&) d
15
N (&) d
13
C (&) d
15
N (&) d
13
C (&) enriched n 11 d
13
C (&) depleted n 6 d
15
N (&) all n 17
Mean 1s Mean 1s Mean 1s Mean 1s Mean 1s Mean 1s Mean 1s
12.1 0.8 11.44 3.6 8.3 0.9 11.0 3.6 10.4 1.1 17.9 1.5 6.6 1.5
Camelids are segregated into two groups according to carbon stable isotope values.
Moreover, isotopic evidence from camelid ber supports the
ecological distinction between llamas and alpacas. Alpaca -
ber is ner and longer than llama wool, and consequently is
the material of choice for textiles, especially polychrome tex-
tiles featuring political and religious iconography. By mea-
suring the stable isotope ratios of polychrome textiles, it is
possible to target the diets of alpacas. Camelid (alpaca) tex-
tiles from the Osmore Drainage in southern Peru yielded an
average d
13
C value of 18& (adjusted), consistent with
a rangestock diet [42]. As the d
13
C values of samples
CI1A, CI22A, CI38A, CI40A, and CI40A resemble those
of the Osmore textiles, it is possible that these ve animals
were alpacas whereas the more d
13
C enriched Conchopata
animals were llamas (Fig. 3).
The d
13
C values of the other 12 Conchopata camelids
(CI3A, CI4A, CI5A, CI7A, CI14A, CI19A, CI23A, CI24A,
CI25A, CI27A, CI37A, and CI39A) are among the highest
yet measured in the Andes and reect maize consumption,
rather than grazing upon C
3
plants. The reliance of these ani-
mals upon large quantities of maize is consistent with the C
4
diets of the humans, guinea pigs and mice at Conchopata
and reects the prevalence of maize in the Ayacucho Valley
during the Middle Horizon.
The dietary emphasis upon a single domestic C
4
plant to the
exclusion of wild and domestic C
3
plants also suggests that the
forage of these animals was articially constrained by humans.
Such severe dietary restriction signals the limited mobility of
these animals within a thoroughly anthropogenic environment.
Either these camelids were conned to corrals for much of their
lives, or their grazing was restricted almost exclusively to the
stubble of maize plots. Corralling seems the more likely expla-
nation as the continuous foraging of animals in agricultural
plots would have been detrimental to seasonal crop production.
Maize farmers in the Ayacucho Valley continue to carry chala
(maize stalks and hulks) from the eld to their herds, though
Old World species have largely replaced camelids in the region.
Counter to Andean models of ecological complementarity
and verticality, maize cultivation and camelid pastoralism
were not spatially segregated, but instead appear to have
comprised an agro-pastoral complex in the quechua zone.
It is energetically improbable that maize was transported
thousands of vertical meters from the quechua zone to ani-
mals on the puna. Rather, than carrying maize to the moun-
tains, the camelids were brought to the corn. Camelids in the
quechua were either fed chala or were allowed to graze the
stubble of maize plots. Camelid dung could in turn have been
used to fertilize adjacent maize plots. The existence of such
an agro-pastoral complex calls into question interpretations
of Andean society which envision a timeless economic di-
chotomy between farmers and herder, elds and ocks, un-
derpinning a general cultural dualism [6,34].
It is possible that the maize foddering of Conchopatas
camelids was an element of an animal management system
wedded to caravan trade. The remains of exceptionally large
juvenile camelids (macho camelidos) recovered at the site
may represent animals purpose bred to be beasts of burden.
Specialized animal husbandry practices appear to have been
coupled to a breeding regime that created and maintained
a breed of pack llamas. The Wari practice of multiple forms
of camelid husbandry tailored to the intensive exploitation of
specialized breeds is consistent with a growing body of re-
search on Andean camelid management in antiquity [13,35].
4.3. Phyllotis
The d
13
C enrichment of the mice (CI20A, CI21A, and
CI35A) indicates they consumed substantial quantities of C
4
plants (i.e. maize) or the tissue of animals foddered on C
4
plants, such as camelids. The d
15
N ratio of sample CI21A
may result from the consumption of marine resources by this
mouse, an unexpected nding for the reasons mentioned above,
i.e. distance from sea and absence of sh bones. If the mouse
represented by sample CI21Awas consuming substantial quan-
tities of marine protein, it was not in the form of sh.
Although it must be emphasized that dietary insights drawn
from a single specimen are speculative, there is a scenario
which could account for the d
15
N enrichment of this rodent.
A diet composed primarily of marine plants would produce
the observed isotopic value, yet leave little if any physical res-
idue in the archaeological record [10]. The d
15
N and d
13
C
-25
-20
-15
-10
1
3
C

(
V
P
D
B
)
-5
0
Conchopata
Argentina
Modern Samples [14]
Argentina
Archaeological
Samples [14]
Peru
Archaeological
Samples [8]
Peru
Modern Samples [51]
Argentine
Modern Samples [44]
Peru
Archaeological
Samples [43]
(keratin +1.85)
Peru
Archaeological
Samples [66]
C
4
Plant
Consumers
C
3
Plant
Consumers
Fig. 5. Carbon stable isotope values of Conchopatas camelids compared to the isotopic signatures of other archaeological and modern camelids in the Andes.
values of this mouses collagen closely resemble those of
coastal camelids from Chilca and Paloma thought to have
been foddered on seaweed [10]. The consumption of seaweed
in ancient Ayacucho is consistent with modern practices in
the region. Cochoyuyo (Porphyra columbina) is a prized food-
stuff in the highlands of southern Peru [33]. As recently as
25 years ago, herders from Ayacucho descend to the coast
with their llamas and to gathered cochoyuyo. The seaweed
was formed into cakes and dried before being loaded onto
llamas for the return trip to the sierra, where this exotic vegeta-
ble is sold in the markets of the Ayacucho Valley [33]. Although
the seaweed consumed in Ayacucho is today brought from Ca-
mana by truck, cochoyuyo remains a delicacy and features in
the feasts of wedding and religious festivals. Cochoyuyo is
also an important source of iodine, an essential nutrient absent
in the glacially depleted soils of the Andean sierra.
Another possible explanation for the extreme d
15
N value of
this specimen, as well as the
15
N enrichment of the coastal
camelids, is the physiological effect of aridity upon both
rodent and plants it was consuming [2,55].
4.4. Cavia
As dependent scavengers, guinea pigs have diets reecting
available human table scraps [17]. The tissues of these ani-
mals would therefore be expected to approximate an isotopic
average of household food waste over the life of the rodent.
Although an individual animals diet might be broad, there
should be low variability of isotopic values between individ-
uals assuming they had access to similar foodstuff in similar
quantities. The greater diversity in carbon isotope values of
Conchopatas guinea pigs relative to the sites humans is
therefore surprising (Fig. 4). The variable consumption of
maize by guinea pigs may result from seasonal variation in
the time of the animals death. Differential consumption of
maize by guinea pigs may also reect differential distribution
of maize waste in Conchopatas architectural spaces. Some
households may have processed maize within the home, re-
sulting in large quantities of husks and cobs in the residence.
Other households may have obtained maize as kernels, the
ears having been shucked and kernels removed elsewhere.
Such residences would have provided less maize waste for
guinea pig fodder.
Spatial variation in maize availability may be reected in the
isotopic values of the guinea pigs deposited in Locus 3118a of
EA40B(Table 1). One guinea pig (CI12A) derived very little of
its dietary protein from maize or other C
4
plants, whereas an-
other animal recovered from this deposit (CI10A) obtained the
bulk its dietary protein in the form of maize. These cuy may
constitute a deposit to which multiple households contributed.
The Conchopata deposit in Locus 3118a resembles a cache of
mummied guinea pigs recovered at the Inka site of Lo Demas,
a deposited attributed by Sandweiss and Wing [47] to a
curandero. Whether the Conchopata animals were deposited
in a dedicatory, diagnostic, or propiatory situation remains ob-
scure, but it is clear that cuy served more than a culinary role
within Wari society.
5. Conclusion
Bone collagen d
13
C values for human and animals from the
Middle Horizon site of Conchopata reect reliance upon
maize as a dietary staple. These ndings are signicant
because:
(1) They indicate that a maize based subsistence economy
supported the Wari state.
(2) They suggest the existence of maize-camelid, agro-pastoral
complex in Ayacucho during the Middle Horizon.
These isotopic insights into the economy of Conchopata
challenge existing models of prehistoric economy in the
Central Andes that emphasize indigenous cultigens and which
project presentist interpretations based on contemporary eth-
nographic data into the distant past. It is now clear that the
animal management practices of this urban state level society
were different from the practices of modern Andean peasants.
Furthermore, as in the complex societies of Mesoamerica,
maize was the source of the surplus calories that supported
the dense populations and specialists associated with urbanism
in Ayacucho.
The d
15
N values of human bone collagen reveal that Con-
chopatas residents consumed substantial quantities of terres-
trial protein, such as the camelids and guinea pig analyzed
in this study. The elevated d
15
N values of human infants reect
breast feeding during the rst year of life.
The isotopic data presented here represent the rst for the
Wari Empire and this is one of the few studies of ancient hu-
man diet in the Andes where human isotope values are consid-
ered in conjunction with those of animals remains from the
same site. The data also represents the earliest evidence for
maize as a dietary staple in the Peruvian highlands. Further
isotopic analyses of human skeletal material from Ayacucho
and other regions of the Andean sierra are necessary to deter-
mine when maize became a staple crop in the region, how
widespread dependence on maize was during prehistory, and
what role maize agriculture played in the increase in sociopo-
litical complexity during the rst millennium AD. It is how-
ever, intriguing to note a correlation between the apparent
late adoption of maize as a dietary staple and the late develop-
ment of cities and states in the Central Andes relative to
Mesoamerica and other nuclear areas of the world [53,56].
Might maize have been the crop that civilized the Andes?
Acknowledgements
This research was supported in part by the Rhodes Trust and
grants from the University of Oxfords Institute of Archaeology
and Merton College, Oxford. Stable isotope measurements for
enamel samples were taken by Norman Charnley in the Depart-
ment of Earth Science, University of Oxford. The authors are
indebted to Anita Cook and Martha Cabrera for access to the
osteological material analyzed in this study. Tifny Tung pro-
vided useful suggestions on the selection of samples from the
Conchopata skeletal population. We gratefully acknowledge
Elsa Tomasto, Mellisa Lund, Irela Vallejo, Gonzalo Rodriguez,
and Alberto Carbajal for their patient assistance in the export of
the samples from Peru as well as Perus Instituto Nacional de
Cultura for permission to export the samples in Oxford. Thanks
are due to Tamsin OConnell, Robert Hedges, Peter Ditcheld
and Jessica Pearson for technical assistance and advice
during the preparation of samples for isotope analysis. We thank
the two anonymous reviewers who provided constructive
comments.
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Human and animal diet at Conchopata, Peru: stable isotope

evidence for maize agriculture and animal management practices

in a common acid bath for

44A 950 M 18e22 RM

44A 950 M 18e22 Skull 10.4 10.3 3.23

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