Received: 11 November 2021 Revised: 23 July 2022 Accepted: 13 September 2022

DOI: 10.1002/ajpa.24626

RESEARCH ARTICLE

Diet composition and prey choice in prehistoric human

individuals from Northwest Patagonia: An application
of species distribution and isotope mixing models

Bruno F. Moscardi 1 | Valeria Bernal 1,2 | Marcio Silva Araújo 3 |

n 1,2 | Virginia A. Cobos 1,2 | Natalia Brachetta-Aporta 2,4
Florencia Gordo |
Raymond Lee 5 | Diego D. Rindel 1,2 | Paula N. Gonzalez 1,2 | Claudia Della Negra 6 |
S. Ivan Perez 1,2

1

n Antropología, Facultad de Ciencias
Divisio
Naturales y Museo, Universidad Nacional de Abstract
La Plata, La Plata, Argentina
Objectives: Ancient hunter-gatherer diets were heterogeneous, varying substantially
2
Consejo Nacional de Investigaciones
Científicas y Técnicas (CONICET), Buenos
across time and space, and frequently showing considerable intrapopulation varia-
Aires, Argentina tion. The diet composition of these human groups depended primarily on resource
3
Departamento de Biodiversidade,
availability, but also on the active selection of certain prey due to different bio-
Universidade Estadual Paulista “Júlio de
Mesquita Filho”, Rio Claro, SP, Brazil cultural factors. In this context, we explore resource availability, diet composition,
4

n en Paleobiología y
Instituto de Investigacio and prey choice in the human populations of the Middle-Late Holocene from North-
Geología, Universidad Nacional de Río Negro,
Viedma, Argentina
west Patagonia.
5
School of Biological Sciences, Washington Material and Methods: We employ species distribution models using current and
State University, Pullman, Washington, USA zooarchaeological data to estimate species availability throughout Northwest Patago-
6

n Provincial de Patrimonio Cultural,
Direccio
nia, and we use Bayesian stable isotope mixing models on a large number of samples
Ministerio de las Culturas, Provincia del
Neuquén, Neuquén, Argentina to analyze human diet composition at the individual level during the Middle-Late
Holocene. Finally, we calculate a prey selectivity index to address the different die-
Correspondence

n
S. Ivan Perez and Valeria Bernal, Divisio tary choices of human individuals in the region.
Antropología, Facultad de Ciencias Naturales y
Results: Our results show large differences in species available for consumption
Museo, Universidad Nacional de La Plata, La
Plata, Argentina. throughout the region, as well as a high dietary variation between human individuals,
Email: ivanperezmorea@gmail.com and
which is mainly related to their spatial location. Some species, such as guanaco, were
bernal.valeria@gmail.com
widely distributed and consumed in the region. Notably, species of small mammals
Funding information
were actively selected in several areas, indicating greater importance in human diets
Consejo Nacional de Investigaciones
Científicas y Técnicas, Grant/Award Numbers: than previously appreciated.
PIP-0729, PIP-2974; Universidad Nacional de
Discussion: Species availability does not appear as the only factor driving human
La Plata, Grant/Award Numbers: 11/N742,
11/N932 diets in the region, since prey choice seems to have been a recurring phenomenon
among these populations. The novel approach used in this study overcomes several
limitations of previous studies employing isotopic analysis in prehistoric human diets,
allowing new insights into the bioarchaeology of the region.

KEYWORDS
Bayesian mixing models, diet, hunter-gatherers, species distribution models, stable isotopes

Am J Biol Anthropol. 2022;1–17. wileyonlinelibrary.com/journal/ajpa © 2022 Wiley Periodicals LLC. 1

2 MOSCARDI ET AL.
1 | I N T RO DU CT I O N
Diet composition and subsistence strategies of prehistoric hunter-
gatherers play a predominant role in archaeological studies, being
essential for understanding human evolution and adaptation (Binford,
2001; Kelly, 2013; Lee & DeVore, 1968; Leonard, 2000; Ungar, 2006).
Contrary to the static and reductionist perspective of anthropological
studies up to the middle of the 20th century that tended to assume
high homogeneity among hunter-gatherers, diet composition and prey
choices in these groups are dynamic and complex phenomena, which
may vary substantially across time and space (Binford, 2001; Cordain
et al., 2002; Kelly, 2013; Lee & DeVore, 1968). Resource availability is
a key factor in determining diet composition and prey choice, along
with other cofactors such as nutritional requirements, consumer
knowledge, and technology, among others (Binford, 2001; Ellis et al.,
1976; Winterhalder & Smith, 2000). Consequently, studies of diet
composition and prey choice in hunter-gatherers would benefit from
the recognition of the spatiotemporal variation in resource availability
and potential diets, and the unraveling of their intrinsic and extrinsic
underlying factors.
Northwest Patagonia is a vast region that presents high environ-
mental heterogeneity, ranging from the temperate-cold Patagonian
forest in the southwest to the Monte desert in the northeast, and
showing high spatial variation in the occurrence of animal and plant
species (Arana et al., 2017; Oyarzabal et al., 2018), and therefore in
resource availability for hunter-gatherer groups. Previous archaeologi-
cal studies identified the main species consumed by prehistoric human
groups in the region, among which large herbivores and small mam-
mals (e.g., guanaco, lesser rhea, pichi, mountain viscacia), and some
wild C3 plants (e.g., alpataco) stand out (Barberena, Prates, & de
Porras, 2015; Cordero, 2010, 2011; Llano, 2015; Neme et al., 2011;
Otaola et al., 2012; Rindel, 2017). Recent studies suggest that human
groups with different diets coexisted throughout the Middle-Late

n et al.,
Holocene in Northwest Patagonia (Bernal et al., 2021; Gordo

n, et al., 2021). It has been proposed that the pre-
2018; Rindel, Gordo
historic human groups from North Neuquén relied almost exclusively
on terrestrial animals (mainly guanaco) while in neighboring areas
(South Mendoza and South Neuquén), other resources such as C3
wild plants and small mammals were also frequently incorporated

n
(Bernal et al., 2016; Cordero, 2011; Gil, Giardina, et al., 2014; Gordo
et al., 2018; Lema et al., 2012; Llano, 2015; Rindel, 2017; Rindel,

n, et al., 2021). Additionally, it has been proposed that these
Gordo
diet variations could have been related with the substantial differ-
ences in human density that existed in different areas of the region
during the Middle-Late Holocene, with high densities in the northern
(Central Mendoza in the Atuel and Diamante river basins) and south-
ern (South Neuquén and North Río Negro) areas, and with low human
densities in the central areas (South Mendoza and North Neuquén;
Barberena, Borrazzo, et al., 2015; Cobos et al., 2022; Gil, Villalba,

n et al., 2019; Rindel, Gordo
et al., 2014; Gordo
n, et al., 2021).
Considerable temporal changes in dietary patterns have been
described for these areas of Northwest Patagonia during the Late
Holocene (Bernal et al., 2016, 2021; Gil, Villalba, et al., 2014;
Lagiglia, 1982; Novellino et al., 2004). Particularly for Central-South
Mendoza, a broader diet has been proposed for the last 2000 years,
characterized by increased consumption of species with low energy
return. However, it is not clear if this was concomitant with a
decrease in consumption of high-ranking resources (i.e., prey which
optimizes the obtaining cost/caloric return ratio, such as guanaco;
Abbona et al., 2021; Bernal et al., 2016; Neme, 2007; Neme & Gil,
2008; Otaola, 2013; Wolverton et al., 2015). For North Neuquén, the
emergence of grinding tools at ca. 5000 BP and ceramics around

n & Novellino, 2017; Lema et al.,
2000 BP (Della Negra, 2008; Gordo
2012) were not related to greater incorporation of plants, but to an
increase in the consumption of large herbivores during the later Late

n et al., 2018; Gordo
Holocene (Bernal et al., 2021; Gordo
n &
Novellino, 2017). Similarly, in Central and South Neuquén and North
Río Negro the trend in the zooarchaeological record was toward
higher consumption of guanaco despite the increase in the number of
exploited taxa throughout the Holocene (Rindel, 2017).
Regardless of the advances in our knowledge about the main
resources consumed in Northwest Patagonia, their importance or rela-
tive contribution to the diet of human populations (i.e., diet composi-
tion) from each area is still debated (Bernal et al., 2016, 2021; Gil

n et al., 2018). Moreover,
et al., 2020; Gil, Villalba, et al., 2014; Gordo
studies directly investigating prey choice are absent in the region. Pre-
vious studies are generally based on presence-absence or frequency
data of animal remains and, to a lesser extent, of vegetables
(i.e., zooarchaeological and archaeobotanical data; Cordero, 2011; Cri-
velli Montero et al., 1996; Fernández, 1988–1990; Lema et al., 2012;
Neme et al., 2011; Otaola et al., 2012; Rindel, 2017), providing evi-
dence of their relative contribution on a regional scale (Cordero,
2007, 2010; Gil et al., 2020; Neme & Gil, 2008; Otaola et al., 2012;

n, et al., 2021;
Pérez & Batres, 2008; Rindel, 2017; Rindel, Gordo
among others). During the last decades the employment of stable iso-
topes, along with the use of mixing models, has been a breakthrough
in the study of the diet composition of prehistoric human populations
in the region (Bernal et al., 2016, 2021; Fernández & Panarello, 2001;

n et al., 2018;
Gil et al., 2011; Gil, Villalba, et al., 2014; Gordo
Novellino et al., 2004). These studies have also focused on a regional
scale, both in resource availability and human mean diet composition,
with significant analytical and conceptual limitations (Bernal et al.,
2021). An alternative and promising approach is to use isotopic data
to analyze diet composition at an individual scale and explore human
individual prey choice according to spatial variation in resource avail-
ability for human consumption (Broughton & O'Connell, 1999; Kelly,
2013; Winterhalder & Smith, 2000).
Here, we study variation in diet composition and prey choice
within Northwest Patagonia, estimating the spatial variation in the
availability of species for human consumption by employing species
distribution models (SDMs; Guisan et al., 2017) and using isotope-
based mixing models to estimate individual human diets (Adams et al.,
2017). Specifically, the aim of this work is to study the spatiotemporal
variation in diet composition and prey choice of individuals from
prehistoric human groups from Northwest Patagonia during the

n et al., 2019; Figure 1), considering
Middle-Late Holocene (Gordo
MOSCARDI ET AL. 3

systematically the geographic differences in the availability of species
for consumption. Because Northwest Patagonia presents spatial varia-
tion in environmental characteristics, we expect to find significant dif-
ferences in resource availability and past human populations diets. To
evaluate these expectations, we used occurrence records of different
species, zooarchaeological information, and isotopic data together
with species distribution modeling (SDM; Elith & Leathwick, 2009;
Guisan et al., 2017; Schmitt et al., 2017), Bayesian mixing models
(BMM; Moore & Semmens, 2008; Stock et al., 2018), a prey choice
index (Zandonà et al., 2011) and Stable Isotope Bayesian Ellipses
approach (Jackson et al., 2011). Whereas Bayesian diet estimation is
not new in bioarchaeology, the estimation of species availability using
SDM has been scarcely explored. By relating the occurrence of a
given species with the local bioclimatic conditions, this method pro-
jects an estimated potential distribution for the species in space
(Guisan et al., 2017). The approach used in this work allows a replica-
ble analysis of stable isotope data, while considering the most appro-
priate resources to incorporate into the mixing models and avoiding
issues related to grouping the human consumers by external criteria in
populations that differ in mobility and home ranges (Adams et al., 2017;
Araújo et al., 2011; Bernal et al., 2021; Gil et al., 2020; Layman et al.,
2012; Moore & Semmens, 2008; Phillips et al., 2014).
2 | MATERIAL AND METHODS
We developed a robust approach to evaluate the variation in the diet
composition and prey choice of human populations from Northwest
Patagonia (Figure 1). First, we determined the list of species poten-
tially consumed by human populations in the region based on previous
studies (e.g., Barberena, Prates, & de Porras, 2015; Cordero, 2010,
2011; Fernández, 1988–1990; Llano, 2015; Otaola et al., 2012;

n, et al., 2021). Then, we modeled the past
Rindel, 2017; Rindel, Gordo
potential spatial distributions of these species throughout the region
using SSDM (Elith & Leathwick, 2009; Guisan et al., 2017; Schmitt
et al., 2017). Since the occurrence data in the paleontological and
archaeological record is scarce for many species, and because neither
the climatic conditions nor species composition has changed signifi-
cantly in the region in the last 6000 years, we modeled species distri-
butions using both current and zooarchaeological occurrence data

F I G U R E 1 Map of Northwest Patagonia showing (a) the human burials and the zooarchaeological sites used in this study, and (b) the species
diversity index together with the human geographical clusters (i.e., human individuals that share the same set of available species, according to
the SSDMs, in their burial locations). The map was generated using the software QGIS v. 3.16.6 “Hannover” (http://www.qgis.org/es/site/).
4 MOSCARDI ET AL.
(Marcott et al., 2013; Pires et al., 2020; Rindel, 2017). The diet com-
position of each individual was estimated using isotopic data and
Bayesian Mixing Models, considering the local species availability esti-
mated based on SSDMs (Adams et al., 2017; Moore & Semmens,
2008; Schmitt et al., 2017; Stock et al., 2018). We calculated an index
of prey choice or selectivity (henceforth “prey selectivity” as it is used
in ecological literature; Zandonà et al., 2011) that compares the con-
tribution of resources to human diets with resource local availability
(i.e., the species local relative abundance) in order to gain insight into
how human populations actively selected or rejected resources.
Finally, we applied the Stable Isotope Bayesian Ellipses approach to
measure the variation in individual diets (Jackson et al., 2011).
2.1 | Species distribution models
In order to estimate species potential distributions (or probabilities of
occurrence) in Northwest Patagonia, we used occurrence records
from the Global Biodiversity Information Facility data set (www.gbif.
org). The selected species, based on archaeological information from
previous studies, include five large herbivores (guanaco [Lama guani-
coe], Patagonian huemul [Hippocamelus bisulcus], southern pudu [Pudu
puda], lesser rhea [Rhea pennata], and greater rhea [Rhea americana]);
two small armadillos (large hairy armadillo [Chaetophractus villosus]
and pichi [Zaedyus pichiy]); four medium size rodents (mountain visca-
cia [Lagidium viscacia], plains viscacia [Lagostomus maximus], Patago-
nian mara [Dolichotis patagonum], and coypu [Myocastor coypus]); and
four C3 plants (algarrobo [Prosopis flexuosa], alpataco [Prosopis alpa-
taco], chañar [Geoffroea decorticans], and araucaria [Araucaria arau-
cana]) (Table S1). All considered species present some direct or
indirect evidence of probable consumption by humans in the studied
region (Barberena, Borrazzo, et al., 2015; Bernal et al., 2021; Cordero,

n et al., 2018;
2010, 2011; Fernández, 2008; Gil et al., 2020; Gordo
Otaola et al., 2012; Rindel, 2017). We also estimated the distributions
of three large carnivores (gray fox [Lycalopex griseus], red fox [Lycalo-
pex culpaeus], and puma [Puma concolor]) that could have been occa-
sionally consumed (Rindel, 2017) and which were direct competitors
of humans for the animal resources. Complementing the species
occurrence records from the Global Biodiversity Information Facility,
we also employed a data set of zooarchaeological assemblages from
97 archaeological sites spread across Northwest Patagonia (Figure 1;
Table S2). Only assemblages with radiocarbon date less than
6000 years before present and geographic coordinates were included
in the analyses. For all species, we checked for very close records
(ca. 1 km) and deleted them to avoid redundancy, as well as duplicate
records. After sifting the database using these criteria, we obtained a
total of 2399 occurrence records (Table S1).
We also obtained the 19 current bioclimatic variables available in
WorldClim2 (www.worldclim.org; Fick & Hijmans, 2017). We com-
pared the values of these variables with those corresponding to the
Late Holocene (from http://www.paleoclim.org/) and both are highly
correlated (r = 0.999 for Annual Mean Temperature and Annual Pre-
cipitation), as expected given the similarities in the region's climate
during the Middle-Late Holocene and the present (Llano et al., 2020).
The bioclimatic variables are computed from monthly minimum, maxi-
mum and mean temperature, and mean precipitation in the period
1970–2000 (Fick & Hijmans, 2017). The obtained variables represent
annual climatic trends that are important to species distribution,
including spatial variation in temperature and precipitation, seasonal-
ity and extreme values of the variables for the coldest, warmest, dri-
est, and wettest months and quarters (Fick & Hijmans, 2017; Guisan
et al., 2017). The bioclimatic variables were downloaded in GeoTiff
format, using a spatial resolution of 30 arc-seconds (
1 km), and
clipped in QGIS v. 3.16.6 “Hannover” (QGIS-Development Core
Team, 2020) using South American contour as a mask layer.
We used a stacked species distribution modeling approach
(SSDM; Guisan et al., 2017; Schmitt et al., 2017) to model the current
and the current plus ancient species distributions. This approach
relates georeferenced species occurrence records to the bioclimatic
conditions at the same location, projecting the result to the entire
geographical space covered by the bioclimatic variables. The spatial
distribution of each species was estimated using a maximum entropy
(Maxent) distribution model (Phillips et al., 2006; Phillips et al., 2014),
which is considered the best method when presence-only data are
used (Elith et al., 2010; Espinosa et al., 2018; Phillips et al., 2006;
Politis et al., 2011). The species distribution models were estimated
using the algorithms implemented in the packages SSDM for the R
software 4.0.3 (Guisan et al., 2017; R-Development Core Team, 2021;
Schmitt et al., 2017). The performance of each distribution model was
evaluated using the area under the receiver operating characteristic
(ROC) curve (AUC statistic; Elith & Leathwick, 2009). For a detailed
review of these methods see Guisan et al. (2017), and for an applica-
tion in archaeology see Rindel, Moscardi, and Perez (2021).
2.2 | Mixing model isotopic analyses for individual
diet composition
In order to estimate the diet composition of ancient humans in the
region, we analyzed isotopic values of δ13C and δ15N from bone colla-
gen of 181 adult individuals of both sexes. These individuals belong to
archaeological sites excavated in the current Argentinean provinces of
Neuquén (n = 76), Mendoza (n = 83), La Pampa (n = 20), Río Negro (n
= 1), and Chubut (n = 1; Table S3). For Neuquén we provide isotopic
data for 40 individuals not previously published. All these individuals
were determined to be adults based on two criteria: the obliteration
of the spheno-basilar suture and eruption of the third molar
(Buikstra & Ubelaker, 1994). The samples come from archaeological
sites dated between ca. 6000 and 200 Calibrated Years Before Pre-

n
sent (Cal BP; Bernal et al., 2020; Gil et al., 2011, 2016, 2020; Gordo
et al., 2018, 2019).
With the aim of establishing the contribution of different sources
to the diets of human individuals, we generated a database of δ13C
and δ15N values of 418 samples of animals (bone collagen) and plants
(edible fruit or seed) from the region (Table S4). The isotopic values
for 355 samples were obtained from previous studies, whereas the
MOSCARDI ET AL. 5

T A B L E 1 Isotopic values of
Species N Class δ13C δ15N
considered resources introduced in the
mixing models Araucaria araucana 9 Plants 22.60 ± 1.01 0.88 ± 2.22
Dolichotis patagonum 10 Small mammals 19.47 ± 1.76 3.62 ± 0.97
Hippocamelus bisulcus 8 Large herbivores 21.03 ± 1.04 1.73 ± 1.05
Lagostomus maximus 50 Small mammals 16.67 ± 2.47 4.56 ± 2.46
Lama guanicoe 108 + 57a Large herbivores 19.25 ± 0.99 5.24 ± 1.17
Myocastor coypus 4 + 1a Small mammals 22.12 ± 0.31 4.14 ± 0.16
Prosopis alpataco 10 Plants 24.82 ± 1.93 1.14 ± 2.22
Prosopis flexuosa 24 Plants 23.76 ± 1.18 4.50 ± 3.71
Pudu puda 5 Large herbivores 21.82 ± 0.60 9.12 ± 2.48
Rhea americana 8 Large herbivores 20.04 ± 0.92 5.80 ± 0.96
Rhea pennata 8 + 6a Large herbivores 20.85 ± 1.06 6.00 ± 0.92
Zaedyus pichiy 30 + 4a Small mammals 17.98 ± 1.99 6.51 ± 1.46
a
Weighted mean of different areas.

remaining 63 samples were processed by us and obtained from differ-
ent locations of the Neuquén province. We explored the spatial varia-
tion in isotopic values for each species and found no considerable
differences between North Neuquén and South Mendoza, which is in
line with previous findings (Bernal et al., 2021), taking into account
that they correspond to the same biogeographic transition zone
(Table S4b; Arana et al., 2017). Bernal et al. (2021) also showed that
there are no significant differences among mean resource values of
different phytogeographic areas for the species widely distributed in
Northwest Neuquén, such as guanaco, probably due to the wide
range of mobility of the large herbivores from the region. Table 1 pre-
sents the mean and standard deviation for the δ13C and δ15N values
of animals and plant resources from the region.
All samples not previously published were processed in the Labo-
ratory of Molecular Biology of the La Plata Museum (FCNyM, UNLP,
La Plata, Argentina). The bones were processed to separate the colla-
gen and mineral fractions, using a standard procedure to clean and
grind each sample that employs abrasive techniques and ultrasonic

n et al., 2018). After the cleaning of the samples, they
washing (Gordo
were demineralized by immersing them in a solution of 2% HCl for
72 h, changing the acid daily. The samples were treated with NaOH
0.1 M during 24 h before and after the osseous demineralization to
eliminate other post-depositional organic material introduced in the

n et al., 2018). The resulting
bones after the burial process (Gordo
osseous demineralized samples were dried at 60  C in a drying oven

n et al., 2018; Tykot, 2004). For plants, 19 samples from mod-
(Gordo
ern wild specimens (Araucaria araucana and Prosopis alpataco) were
collected and the edible tissues (seeds) were extracted. These tissues
were ultrasonically cleaned with distilled water, cut into smaller

pieces, and dried at 66 C, to be finally grounded into powder.
The estimation of stable isotope abundance ratios (13C/12C and
15
N/14N) was done in the UC Davis Stable Isotope Facility using a
continuous-flow gas-ratio mass spectrometer Finnigan Delta PlusXL,
and in the Washington State University (WSU) Stable Isotope Core
Laboratory using a Costech (Valencia) Elemental Analyzer interfaced
with a GV Instruments (Manchester, UK) Isoprime mass spectrometer.
The isotopic relative abundances were measured relative to the ratios
of those same isotopes in standard materials and expressed as the dif-
ference (δ value) per mil (‰) between the ratio of isotopes (13C/12C
or 15N/14N) in the sample and the standard, by using Vienna Pee Dee

n et al.,
Belemnite (VPDB) and Air as standards, respectively (Gordo
2018). Precision for isotope analyses was <0.2‰ for δ13C and <0.5‰
for δ15N based on the standard deviation of replicate working stan-
dards calibrated against NIST reference materials. The precision of the
measurements for human samples was checked for both isotopes, and
the ratio C/N (Table S3) was used to assess the quality of the collagen
(van Klinken, 1999). The resource samples analyzed in the WSU Sta-
ble Isotope Core Laboratory facility showing low nitrogen sample
peaks were considered less reliable and discarded from the analyses.
Finally, we applied the Bayesian mixing model implemented in the
MixSIAR package for R 4.0.3 (Stock et al., 2018) to analyze the propor-
tional contribution of different sources (animal and C3 plant species)
to the diets of each human individual (consumers). The Bayesian mix-
ing model uses Markov chain Monte Carlo (MCMC) algorithms to cal-
culate a posterior probability distribution, estimating the mean and
deviation of the diet proportions of each individual (Table S5), and
therefore considering error in diet estimation (Moore & Semmens,
2008). Particularly, this method incorporates uncertainty associated
with multiple resources, fractionation values, and isotope signatures
(Moore & Semmens, 2008; Stock et al., 2018). The resources used for
the human individual diet estimation were selected according to the
species availability estimated based on the SSDMs. It is important to
remark that even though some individuals display the same set of
available resources for consumption, such as it is shown in Figure 1b,
diet estimations were not constrained to an overall or mean diet esti-
mation. Conversely, human diet composition was estimated at the
individual level. For this purpose, we set each individual as a fixed fac-
tor and used the parameters recommended by the authors for MCMC
in the “very long” option, estimating the diet individually and calculat-
ing the “process error” (Moore & Semmens, 2008; Stock et al., 2018).
The isotope fractionation values used for different resources were
δ13C 1.0 ± 0.3/δ15N 4.0 ± 1.0 for all animals and δ13C 3.9 ± 1.4/δ15N
6 MOSCARDI ET AL.
2.2 ± 0.3 for C3 plants (Bernal et al., 2016; Bocherens & Drucker,
2003; Hare et al., 1991). Because the discriminatory power of the
Bayesian mixing model decreases with large numbers of resources
and when their isotopic signal is similar (Layman et al., 2012), we
include only the species with a potential distribution >0.1 in the
human burial location (Bernal et al., 2021; Phillips et al., 2014). Due to
the non-homogeneous spatial distribution of species in the region, the
maximum number of seven resources suggested by Phillips et al.
(2014) was not exceeded for any human individual diet estimation.
The Suess effect for modern samples was corrected by adding
+1.5‰ to δ C values, as it is frequently applied (Tessone, 2010;
13
Tessone et al., 2014). To corroborate this value, we built a database of
δ13C values from 556 faunal specimens (247 modern and 309 archaeo-
logical) and performed comparisons between the same species in the
same region (Table S6). Specifically, we calculated the difference of
the mean and median between archaeological and modern samples of
a given species from the same region, weighting the quality of the
comparisons according to the quantity and diversity of the samples.
The grand means and medians resulted in a difference of around
1.5‰ (Table S6).
2.3 | Prey choice analyses
For each human individual sample, we calculated the relative abun-
dances of resource species (pi) in the spatial location where the human
sample was collected, that is, the local availability of resources. For
each species i, we divided i's estimated local potential of occurrence
by the sum of the potentials of occurrence of all species present in
this location. We acknowledge that this approach is based on the
assumption that the actual abundance of a species is proportional to
its potential distribution estimated by the SSDM. To the extent that
the relationship between estimated potential distribution and actual
abundances are consistent among species, this approach should pro-
vide robust estimates of local relative abundances of species. To
investigate if humans were selectively choosing to feed on specific
species while avoiding other species, we calculated a prey selectivity
index (Zandonà et al., 2011) as follows: Li = ri  pi, where Li is the prey
selectivity index for the species i, ri is the relative contribution of spe-
cies i to an individual's diet estimated from the isotopic mixing model,
and pi is the relative abundance of species i in the human burial loca-
tion. The Li index varies between +1 and 1. When Li > 0, the species
proportional contribution to the diet is higher than its relative local
abundance, that is, the human consumer is actively selecting species i;
when Li < 0, the species contribution to the diet is lower than its local
abundance, that is, the human consumer is avoiding species i; and if Li
= 0, the species contribution to the diet is in direct proportion to its
local abundance, that is, the human consumer is consuming species
i in proportion to its local availability. The spatial variation in all vari-
ables was visualized using QGIS v. 3.16.6 “Hannover.” We also esti-
mated if the different species were significantly chosen in the positive
or negative direction by calculating the mean Li of human individuals
for each considered species, and applying a Bootstrap resampling
method with 10,000 replicates on the Li individual values to establish
confidence intervals, using the Boot package for R software 4.0.3 (R-
Development Core Team, 2021).
Additionally, in order to explore the temporal dietary changes in
the region, we divided human samples in three periods following
recent studies (Bernal et al., 2021; Gil et al., 2020): (i) older than
2000 years Cal BP (pre-2 k), (ii) dated between 1999 and 500 years
Cal BP (post-2 k) and (iii) assigned to the interval 499–200 years BP
(historical). Out of the 181 individuals analyzed, 168 were dated or
assigned to a chronological period by contextual information
(e.g., presence of pottery, remains of domesticated vegetables, post-
contact material evidence; Table S3), while the remaining samples of
unknown dating were excluded.
2.4 | Intra-group diet variation analyses
We applied the Stable Isotope Bayesian Ellipses approach to measure
the intra-group variation in individual diets, using δ13C and δ15N iso-
tope data of individual consumers, and considering the geographical
human clusters—reconstructed based on the local species availability
estimated with SSDMs—as group factor. The area of the standard
ellipse (SEA) is equivalent to the standard deviation for 2D data and
can be interpreted as a measure of variation in energy pathways
(δ13C) and trophic position (δ15N) among individuals. SEA = πab,
where a and b are obtained from the eigenvalues of the bivariate data.
We employed the Bayesian approach, where the SEA value is
obtained from the posterior distribution resulting from an MCMC sim-
ulation. We plotted the credible intervals for the vector of posterior
Bayesian Standard Ellipse Areas (SEA.B) for each studied human geo-
graphical cluster, displaying box edges that represent 50, 75, and 95%
of the Bayesian posterior distribution. The Stable Isotope Bayesian
Ellipses were calculated in the SIBER package for R (Jackson et al.,
2011). In addition, we plotted the δ13C and δ15N values for all individ-
uals grouped by geographical cluster, using convex hulls to display the
observed differences in intra-group variation.
3 | RE SU LT S
3.1 | Species distribution and diversity
Our species distribution models show considerable differences in spe-
cies diversity across Northwest Patagonia (Figure 1b; see Figures S1
to S4). The geographic distributions of large herbivores, small mammals
(i.e., rodents and armadillos), and plants obtained from the data sets
that combine current and zooarchaeological information are shown in
Figures S1, S2, and S3. With a few exceptions, the distributions esti-
mated from these data agree with those obtained using current data
only (Figures S1 to S3). Based on the potential distribution of the ana-
lyzed species, we found that some human individuals share the same
set of available resources (i.e., human individuals exhibiting potential
distribution values >0.1 of the same species in its burial location;
MOSCARDI ET AL.
F I G U R E 2 Human individual consumption and prey choice of each species with wide distribution in Northwest Patagonia. On top, human
individual consumption estimated by MixSir mixing models. On the bottom, prey selectivity index (Li), where positive values indicate an active
selection of a prey and negative values indicate a relative rejection
Figures S1 to S3). These human individuals sharing the same set of
potentially available resources are henceforth referred to as geo-
graphical clusters, and only represent 12 groups of individuals that
display the same potential diet (Figure 1b; Figures S5 to S7).
3.2 | Individual diet composition and prey choice
Because we have shown that some species display wide potential dis-
tribution throughout the region, while others occur particularly in the
north or the south, the following consumption and prey choice results
are presented according to this criterion. Moreover, since mountain
viscacia, large hairy armadillo and chañar display very similar distribu-
tions (Figures S1 to S3) and isotopic values (Figure S8) to guanaco,
pichi, and algarrobo, respectively, but exhibit weaker archaeological
evidence of consumption, only the latter were included in the MixSIR
analyses. Additionally, we excluded the three carnivorous species
because their isotopic values are also similar to armadillos and there is
less archaeological evidence of their consumption. The resource isoto-
pic values effectively used in MixSIR to estimate human individual diet
composition show low similarity between their means in each geo-
graphical cluster (Figures S9 and S10).
Figures 2 to 4 show the effective consumption of each species by
human individuals (estimated by MixSIR) and the Prey Selectivity Index
7
used to estimate prey choices (i.e., differences between available [rel-
ative potential distribution estimated using SSDM] and consumed
resources [estimated based on isotope data and MixSIR]) in the geo-
graphic location of the burial sites. The species with wider potential
distribution throughout Northwest Patagonia are guanaco, lesser rhea,
pichi, and coypu (Figures S1 and S2). Guanaco was highly consumed
by human individuals throughout most of the region, especially in the
middle portion (North Neuquén), but its consumption decreased
toward the north and was very low in the Southwest forest. Accord-
ingly, this camelid was selectively chosen by humans, particularly in
Northwest Neuquén (Figure 2). Lesser rhea, in turn, shows both high
consumption and prey choice values in East Neuquén. Pichi is the
other highly consumed species by prehistoric human groups in the
region, presenting the higher consumption and selectivity values in
the north, and declining gradually towards the south. Finally, coypu
was only frequently consumed in the southern portion of the region
(Figure 2).
Regarding the species consumed by humans in the north of the
region (Figure 3), greater rhea was consumed in low frequencies and
showed prey choice values according to its potential distribution.
Plains viscacia and Patagonian mara also show low values of con-
sumption in the north of the region, but with higher values in the
northeast area particularly for plains viscacia (Figure 3). Accordingly,
the selectivity index displays the highest values in the northeast area
8 MOSCARDI ET AL.

F I G U R E 3 Human individual consumption and prey choice of each species distributed in the north of Northwest Patagonia. On top, human
individual consumption estimated by MixSir mixing models. On the bottom, prey selectivity index (Li), where positive values indicate an active
selection of a prey and negative values indicate a relative rejection

for the last species (Figure 3). It is noteworthy that this great exploita-
tion of plains viscacia occurs in the area without guanaco distribution
and that adding the guanaco to the mixing model does not change this
result (results not shown). Despite its northern distribution, the algar-
robo was the plant consumed by the largest number of individuals
across the region, showing positive selectivity values in Southwest
Mendoza, although its contribution to their diets was low (Figure 3).
For the south portion of the region, southern pudu appears to
have been consumed in low proportions (Figure 4). Conversely, the
Patagonian huemul, the other deer species inhabiting the Southwest
forest, showed relatively high consumption and prey choice values
(Figure 4). The two C3 plant species of the South showed a reduced
consumption both in absolute terms and in relation to their potential
distribution. Araucaria was the only plant species with high selectivity
values, which were only found in individuals belonging to the Center-
West of the current province of Neuquén (Figure 4). The alpataco, in
turn, was only consumed in a few sites of Northwest Neuquén, where
it displays positive selectivity values since its potential distribution is
low for this area (Figure 4).
In Figures S11 to S13 we show the relative consumption of each
species by temporal periods. The spatial pattern of consumption remains
relatively constant over time and the apparent differences found in some
cases probably reflect the absence of samples for a given period. Particu-
larly, among the widely distributed species, the pattern of consumption
is similar among all periods for guanaco and pichi. Likewise, lesser rhea
displays similar patterns, although the consumption of this species could
not be evaluated for the pre2k period in the east of Neuquén due to the
lack of human samples (Figure S11). In the north of the region, a similar
pattern of consumption of all species (greater rhea, plains viscacia, Pata-
gonian mara, and algarrobo) was observed across the periods (Figure
S12). Finally, the lack of human samples from the south of the region for
some time intervals limited the evaluation of temporal changes in the
consumption of coypu and other species distributed in this area (Figures
S11 and S13).
Figure 5 displays the mean and bootstrap confidence interval of
the prey selectivity index (Li mean) of each human individual by the
consumed species. The species that showed high mean selectivity
values statistically different from zero, indicating positive prey choice
by humans, are guanaco and pichi among the species with wide distri-
bution, plains viscacia in the north, and Patagonian huemul and arau-
caria in the south of the region. Therefore, these species were
consumed in a higher proportion than expected given their spatial
availability. Two species of the south, southern pudu, and alpataco, pre-
sented mean selectivity values not statistically different from zero,
meaning they were consumed in direct proportion to their spatial avail-
ability. In contrast, the mean negative values obtained for lesser rhea,
coypu, greater rhea, Patagonian mara, and algarrobo indicate a lower
consumption than expected based on their potential distribution.
MOSCARDI ET AL. 9

F I G U R E 4 Human individual consumption and prey choice of each species distributed in the south of Northwest Patagonia. On top, human
individual consumption estimated by MixSir mixing models. On the bottom, prey selectivity index (Li), where positive values indicate an active
selection of a prey and negative values indicate a relative rejection

F I G U R E 5 Mean prey selectivity

index (Li) and bootstrap confidence
interval for each considered species. For
each species the mean was calculated
from the Li values of human individuals.
Species are ordered according to their
predominant geographical distribution
10 MOSCARDI ET AL.

3.3 | Diet variation
Finally, isotopic bivariate variation and SIBER results for each geo-
graphical cluster are displayed in Figure 6. The biplot (Figure 6a) and
credible intervals of the SEA.B (Figure 6b) show large differences in
intra-group variation among geographical clusters. The variation in
diet composition was higher in the north—clusters from South
Mendoza—and the south of the region—clusters from South Neuquén
and Rio Negro—. Conversely, the variation in the center of Northwest
Patagonia, particularly in North Neuquén, was considerably lower,
showing a posterior distribution that does not overlap with some clus-
ters from the northernmost clusters (Figure 6). These results suggest
that the degree of among-individual diet variation was higher in the
groups from South Mendoza and South Neuquén, which incorporated

F I G U R E 6 Inter-individual
isotopic variation of each human
geographical cluster. (a) Biplot of
δ13C and δ15N human values
showing different symbols,
colors, and convex hulls for each
geographical cluster. (b) Isotopic
variation of each human
geographical cluster estimated by
SIBER. The SIBER method uses
the standard ellipse area to
obtain a single estimate of the
variation in δ13C and δ15N human
isotopic values, and the Bayesian
method to generate credible
intervals. These intervals are
displayed in box edges that
represent 50%, 75%, and 95% of
the Bayesian posterior
distribution
MOSCARDI ET AL.
large diversity of resources, and lower in North Neuquén, where the
groups have a lower diversity of species available for consumption.
4 | DISCUSSION
Our results show that there was a significant spatial variation in the
species available for consumption by the human populations of North-
west Patagonia in the Middle-Late Holocene (Figures S1 to S3). Some
species have a widespread distribution, such as pichi and guanaco,
whereas others are spatially restricted. The spatial restriction of some
species such as the Patagonian huemul, southern pudu, and araucaria
only found in the Southwest forest had been documented in the
archeological record (Aguirre & Pérez, 2015; Fernández, 2008;
Fernández et al., 2015; Pérez & Batres, 2008), whereas for others the
distribution predicted by the model had not been previously reported.
In this sense, one remarkable finding is the absence of guanaco in the
northeast of the region and the presence of plains viscacia and Pata-
gonian mara only in this area (Figures S1 and S2). Additionally, lesser
rhea is predominantly distributed along peri-cordilleran areas, while
greater rhea is present only in the northeast portion of the region
(Figure S1). The three main carnivores of the region are distributed
primarily along the cordilleran and peri-cordilleran areas, where the
largest herbivores are also mainly distributed (Figure S4). These results
highlight the high variability present in the region: while the northern
and southern areas show a high potential distribution of species
(although different in each case), the central sector of Neuquén, espe-
cially toward the east, is comparatively poor in both animal and vege-
table species for human consumption (Figure 1b; Figures S1 to S3). In
this regard, the application of species distribution models to the study
area allowed us to obtain novel results. Species distribution models
are a powerful tool to evaluate resource availability for human popula-
tions, since they allow estimating the distribution of prey species
together with relevant paleoclimatic variables in a spatially explicit
way (Brown et al., 2018; Fordham et al., 2017). However, although
they are widely used in ecology, biogeography, conservation biology,
and more recently in paleontology (Elith & Leathwick, 2009; Hao
et al., 2019; Varela & Fariña, 2016; Villavicencio et al., 2019) their use
in archaeology is relatively restricted (for exceptions see Franco &
Camps, 2020; Franklin et al., 2015; Politis et al., 2011, 2018; Rindel,
Moscardi, & Perez, 2021).
In a scenario of species diversity and spatial variation in their
availability for human consumption, it is important to explore not only
diet composition, but also whether prey were exploited in close rela-
tion to their potential distribution or, alternatively, if some prey choice
or selection was made by prehistoric human groups (Broughton &
O'Connell, 1999; Kelly, 2013; Winterhalder & Smith, 2000). To evalu-
ate both aspects of human diet variation, in this work we estimate diet
composition at an individual scale using MixSir models (Moore &
Semmens, 2008) and prey choice employing a selectivity index
(Zandonà et al., 2011), in which the deviation from the null value
implies a choice for or against a certain species. Our results show that
the species distribution is not necessarily reflected in the patterns of
11
consumption of the human individuals, suggesting that prey choice
was a recurrent phenomenon in the region for some species. In this
sense, a marked trend towards a greater contribution of small mam-
mals in the diet was found north of the Colorado River (current Prov-
ince of Mendoza). This area is characterized by a relatively low
consumption of guanaco, especially considering its high potential dis-
tribution, and by an almost null consumption of both lesser and
greater rheas (Figures 2 and 3). Our results suggest that human popu-
lations of this area performed a high positive selection on pichi, while
plains viscacia was positively selected only in the northeast (Figures 2
and 3). Accordingly, zooarchaeological and ethnohistorical evidence
also suggest that a variety of species of small animals were frequently
consumed in the region, even becoming the main prey in some areas

n, et al., 2021). Like-
(Gil et al., 2020; Otaola et al., 2012; Rindel, Gordo
wise, some previous studies have also shown a higher consumption of
meat over vegetables (Gil et al., 2020), as observed in our results.
Conversely, plains viscacia have been found very rarely in the archae-
ological sites from the peri-cordilleran and northeastern extreme of
Mendoza (Table S2; Fernández et al., 2009; Giardina et al., 2017; Gil,
2006; Neme & Gil, 2009; Otaola et al., 2012), even though our results
support high consumption of this species.
In Northwestern Neuquén the trend is towards a high consump-
tion of guanaco, being this species consumed in higher frequencies
than expected by its potential distribution (Figure 2). The high contri-
bution of guanaco to human diet in this area had been previously
shown on the basis of isotopic and zooarchaeological data (Barberena,

n et al., 2018; Rindel, 2017), in line
Prates, & de Porras, 2015; Gordo
with the trend found for overall Patagonia (Moscardi et al., 2020; Rin-
del, Moscardi, & Perez, 2021). Likewise, pichi shows negative values
of selectivity in this area, despite being consumed in considerable fre-
quency, due to its extremely high potential distribution (mean = 0.99),
suggesting that this species was partly ignored. Conversely, the alpa-
taco shows relatively low levels of consumption, but appears as a
desired resource due to its relative scarcity in the area. On the other
hand, with a much lower diversity of available species (Figure 1b),
individuals from Eastern Neuquén relied mostly on lesser rhea and
pichi, actively selecting those prey over guanaco, which was also con-
sumed but with less frequency, and especially, over alpataco, which
appears as a highly negatively discriminated species (Figures 2 and 3).
Accordingly, the archaeological sites of this area are extremely domi-
nated by the remains of these three faunal species (Table S2; Della
Negra & Novellino, 2002; Menegaz, 1996; Rindel et al., 2018).
In the Southwest and Centerwest Neuquén forest, Patagonian
huemul, coypu and araucaria appear as the three main consumed spe-
cies (Figure 4). Being exclusive of this area, both Patagonian huemul
and araucaria show positive selectivity index means (Figures 4 and 5),
while coypu shows a negative value due to its much lower consump-
tion in the rest of the region. On other hand, southern pudu was con-
sumed according to its potential distribution, whereas guanaco and
pichi were consumed with much less frequency than would be
expected due to their estimated potential distribution, showing the
large amount of dietary choices that individuals had due to the sub-
stantial diversity of species available for consumption in the
12
temperate Southwest forest (Figures 2 and 4). These results are in
good agreement with the zooarchaeological evidence, which shows a
high prevalence of Patagonian huemul remains among the large faunal
diversity in the archeological sites from this area (Table S2; Fernández
et al., 2015; Pérez & Batres, 2008; Silveira et al., 2014). Likewise, both
ethnohistorical and ethnobotanical evidence inform the dietary rele-
vance of araucaria gathering during the summer season for the native
populations of the region (Aagesen, 2004; dos Reis et al., 2014;
Fernández, 1988–1990; Ladio et al., 2011). However, as with other
species of small mammals aforementioned, the coypu has been found
very rarely in the archaeological sites (Table S2), even though our
results show a high consumption of this species in the area.
From the aforementioned, it is evident that there is a good fit
between our results and other lines of evidence such as zooarchaeolo-
gical data. However, a number of discrepancies have also been
observed, especially in relation to the importance of certain small
mammal resources that have little or even null presence in the
zooarchaeological record. Although it is not possible to reach defini-
tive conclusions in this regard at the moment, we suggest that these
differences are due to a number of factors that should be evaluated in
the future. First, it is possible that these differences arise from difficul-
ties for taxonomic identification of small mammals and other small
prey in the zooarchaeological record. Furthermore, due to their small
size and more fragile bone structure, the remains of small prey tend to
present poorer preservation and be more fragmented making their
recovery and identification even more difficult (Behrensmeyer et al.,
1979; Shaffer & Sanchez, 1994; Von Endt & Ortner, 1984). In this line,
ethnoarchaeological information has repeatedly indicated the possibil-
ity of the complete consumption of this type of prey, and/or its con-
sumption without evidence of processing, making its evaluation
difficult in the archaeological record (Binford, 1981; Stahl, 1982,
1996). Finally, and more important for the purposes of this work,
these differences might arise from the nature of the data: while the
zooarchaeological record offers information about multiple consump-
tion events at the population level and on a broad temporal scale, the
diet modeling from isotopic data shows evidence of individuals' con-
sumption over approximately the last 10 years period prior to their
death (Ambrose, 1993; Davis & Pineda Munoz, 2016). In this sense,
the latter offers a robust means to evaluate the particular decisions of
each individual, adding another layer of complexity to the general
trends observed in the zooarchaeological record.
We also need to remark that the differences between isotopic
and zooarchaeological results can be related to a number of limita-
tions of isotopic analyses. Firstly, there are similarities in mean isoto-
pic values and species distribution for some species probably
consumed by prehistoric human groups, such as pichi and large hairy
armadillo, and guanaco and mountain viscacia (Figures S1, S3, and S8).
Because of these similarities, it is practically impossible to differenti-
ate the importance of these pairs of species for the human diet using
the approach employed in this work. Potentially, we could weigh the
contribution to the human diet of each of these pairs of species using
other archeological information. However, whereas there is much
more evidence of guanaco consumption than mountain viscacia, it
MOSCARDI ET AL.
should be noted that it is difficult to differentiate Dasypodidae spe-

n, et al., 2021). Addi-
cies in an archaeological context (Rindel, Gordo
tionally, the specimens of some species consumed by humans display
variable differences in isotopic values (Table S4; Figures S8 to S10),
which makes it difficult to characterize these species isotopically;
although Bayesian mixing models consider this intraspecific variation
in the resources. Prior work in the region has suggested that these dif-
ferences could be related to micro-regional patterns linked to vegeta-
tion units (Gil et al., 2020), but other work have questioned this
relationship (Bernal et al., 2021), and we have not found any clear pat-
tern (Table S4b). Future works need to explore the impact of the iso-
topic diet variation of prey on human diet composition estimations.
Finally, it could be thought that the zooarchaeological information
used and the potential availability of prey estimated for a given geo-
graphical area could generate some bias in the subsequent isotopic
results. In this sense, the selection of potential prey species to be
introduced in the mixing models is a general problem for isotopic
studies. In this work, we approached this issue in a systematic and
replicable way, reducing the chances of subjective bias. Moreover,
MixSir diet composition estimation for each human individual based
on isotopic data are relatively efficient for weighing between the spe-
cies that were included in the model which were consumed and which
were not. For example, we included the Patagonian mara and lesser
rhea in geographical areas where there is archaeological evidence of
their consumption, but the isotopic analysis showed that they were
just scarcely consumed (Figures 2 and 3).
There are different possible explanations for the observed spatial
differences in species consumption and prey choice in Northwest Pat-
agonia. One of the possible ways to explain these differences, and
particularly the high consumption of small prey in some areas, are the
changes in human density over time and space (Flannery, 1969; Stiner
et al., 2000; Winterhalder & Smith, 2000). Nutritional stress arising
from increased human density may push people to include suboptimal
resources, with lower cost/energy return ratios, in their diets
(Flannery, 1969; Kelly, 2013; MacArthur & Pianka, 1966; Stiner et al.,
2000; Winterhalder & Smith, 2000). In this sense, human populations
from Northwest Patagonia increased considerably during the Middle
and Late Holocene, with an early increase strongly supported by
molecular data (7500–6000 years Cal BP) and a later one clearly
described by absolute dates (3000–2500 years Cal BP; Barberena,
Prates, & de Porras, 2015; Cobos et al., 2022; Gil et al., 2016; Perez
et al., 2016, 2017). The demographic expansion during the Middle
Holocene has been associated with an economic intensification based
on the consumption of different ensembles of resources that occurred
following a population bottleneck in the region (Barberena et al.,
2017). On the other hand, the regional increase in human population
density suggested by absolute dates during the Late Holocene was
greater in South Mendoza and in the Southwest forest than in North
Neuquén (Cobos et al., 2022). Particularly, for South Mendoza, a pro-
cess of intensification during the last 2000 years related to these
demographic changes has been postulated and debated (Abbona
et al., 2021; Bernal et al., 2016; Neme, 2007; Neme & Gil, 2008;
Otaola, 2013; Wolverton et al., 2015). Although our results are
MOSCARDI ET AL.
compatible with a broad diet including species with low energy return
for South Mendoza, we do not find significant temporal dietary
changes in the area during this period (Figures S11 to S13), suggesting
that the process which generated these dietary and demographic dif-
ferences between South Mendoza and other areas may have occurred
before the study period. For the Southwest forest there is evidence of
incorporation of additional resources during the Late Holocene and
historical times, including a high proportion of vegetable resources
(e.g., araucaria; Fernández, 1988–1990; Ladio et al., 2011) as was
observed in our results, and with a relative increase in the diversity of
consumed animal species (Bernal et al., 2018; Rindel, 2017). It is
remarkable that in this area there is also a positive selection of Pata-
gonian huemul (Figure 5). Since small mammals are resources rich in a
variety of nutrients essential for human life (e.g., essential fatty acids),
their greater incorporation could benefit survival and reproduction
(Haws & Hockett, 2004; Hockett & Haws, 2003; Jenike, 2001; Rindel,

n, et al., 2021), thus generating higher human density. This
Gordo
would be another possible and non-contradictory explanation for the
dietary differences in Northwest Patagonia, and particularly for the
higher consumption of small prey in some areas. Certainly, other cul-
tural variables, whose analysis goes beyond the scope of this study,
such as knowledge, hunting technology, social organization, and sub-
sistence strategies may also have played a pivotal role in the dietary
selections of these populations.
Finally, we found differences among individuals that share the
same set of available resources given their geographical location
(Figure 6), particularly for South Mendoza and Southwest forest
where there is evidence of higher human density during the Middle-
Late Holocene than in the North Neuquén (Cobos et al., 2022), as well
as more diversity of species available for human consumption (Figure
1b). Although inter-individual variation has been widely covered by
examples in ecology (e.g., Araújo et al., 2011; Bolnick et al., 2003) and
in the archaeology of complex, resource-producing societies
(e.g., Irvine & Erdal, 2020; Kinaston et al., 2013; Kusaka et al., 2010), it
remains less explored in hunter-gatherer bioarchaeology, and virtually
unexplored in Northwest Patagonia. In these last cases, the analysis of
diet variation in the literature implicitly assumes that individuals
within populations had equal access to all resources consumed by the
populations in these areas. However, there are several reasons to
believe that individuals did not consume resources homogeneously,
among which gender and age-based differences in diets (Kelly, 2013;
Tessone et al., 2015; Turner et al., 2010) and differences in social sta-
tus (Phaff et al., 2015; Testart, 1982; Turner et al., 2010) are worth
noting. Additionally, numerous examples among populations of animal
species indicate that the expansion of dietary niches, in which novel
food resources are included in the diet of a population, is achieved
through increased inter-individual variation (Araújo et al., 2011;
Bolnick et al., 2007; Pires et al., 2011). This means that not all individ-
uals expand their niches to include all dietary items in their diets, but
instead some individuals specialize on novel strategies to avoid com-
petition with conspecifics (Van Valen, 1965). The implications of these
results for an area inhabited by hunter-gatherer groups with access to
a high diversity of both animal and plant species, as well as with a
13
small contribution of domestic resources (Bernal et al., 2016; Gil,

n et al., 2018), cannot be ignored and needs to be dis-
2006; Gordo
cussed in greater depth in future studies.
5 | FI NA L R E M A R KS
In conclusion, our study shows that modeling individual diets allows
us to detect diverse spatial patterns that coexisted in different parts
of Northwest Patagonia, and that these patterns are not only deter-
mined by the availability of resources, since the selection of certain
prey was a recurring phenomenon. Previous studies had already indi-
cated a more important contribution of animal species over plants in

n et al., 2018; Rin-
the area (Bernal et al., 2016; Gil et al., 2020; Gordo

n, et al., 2021), as well as a relevant role of small mammals
del, Gordo

n, et al., 2021). Our results allow us
(Bernal et al., 2021; Rindel, Gordo
to expand these claims, indicating an even more significant role of cer-
tain species with little presence in the archaeological record. The rea-
son for these results probably resides in a high environmental
heterogeneity, displaying differences between the western and east-
ern sectors of the region and from north to south, and thus generating
localized distributions of plants and animals available for human con-
sumption, as well as different prey choice patterns. Finally, we want
to highlight a methodological aspect of our work: the use of isotopic
data allows the analysis of variability in individual diets, an aspect that
is difficult to resolve from other archaeological data, thus enriching
the discussions generated from other lines of evidence such as
zooarchaeology (Cordero, 2007, 2010; Gil et al., 2020; Neme & Gil,
2008; Otaola et al., 2012; Perez & Batres, 2008; Rindel, 2017; Rindel,

n, et al., 2021; among others). Moreover, since inter-individual
Gordo
variation in dietary resources acquisition and consumption due to
diverse factors is a widespread phenomenon in many species, model-
ing the diet individually appears as an appropriate approach which, in
turn, allows to avoid the clustering of human consumers in ad hoc
groups (Adams et al., 2017; Araújo et al., 2011; Bernal et al., 2021;
Bolnick et al., 2003). In this regard, we believe that beyond the limita-
tions of the isotopic analysis, we presented a promising approach in
bioarchaeology, especially for hunter-gatherer groups, where the
available data indicate significant differences in the importance of
diverse wild species of plants and animals through space and time,
whose relative importance to human diets is debated.
AUTHOR CONTRIBU TIONS
Bruno F. Moscardi: Conceptualization (equal); data curation (lead);
formal analysis (equal); investigation (equal); writing – original draft
(equal). Valeria Bernal: Conceptualization (equal); funding acquisition
(lead); investigation (equal); writing – original draft (equal). Marcio
Silva Araújo: Conceptualization (equal); methodology (lead); writing –

n: Resources (equal);
review and editing (equal). Florencia Gordo
writing – review and editing (equal). Virginia A. Cobos: Resources
(equal); writing – review and editing (equal). Natalia Brachetta-Aporta:
Writing – review and editing (equal). Raymond Lee: Resources (equal);
writing – review and editing (equal). Diego Rindel: Data curation
14
(equal); writing – review and editing (equal). Paula N. Gonzalez: Pro-
ject administration (lead); writing – original draft (equal). Claudia Della
Negra: Resources (equal); writing – review and editing (equal). S. Ivan
Perez: Conceptualization (equal); data curation (equal); formal analysis
(equal); supervision (lead); writing – original draft (equal).
ACKNOWLEDGMENTS
We want to thank Augusto Tessone for kindly sharing some of his
data. Bruno F. Moscardi is deeply grateful to Federico Moscardi for a
lifetime of guidance and support. We also thank two anonymous
reviewers whose comments helped improve this manuscript.
CONF LICT OF IN TE RE ST
The authors declare that they have no conflict of interest.
DATA AVAI LAB ILITY S TATEMENT
All data sets used in this study are provided as supplementary material
(Tables S1 to S6).
ORCID
S. Ivan Perez https://orcid.org/0000-0002-6543-5545
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n, F., Cobos, V. A., Brachetta-Aporta, N., Lee,
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(2022). Diet composition and prey choice in prehistoric human
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