981689

research-article2020
HOL0010.1177/0959683620981689The HoloceneRindel et al.

Research Paper

The Holocene

The distribution of the guanaco (Lama

2021, Vol. 31(4) 644­–657
© The Author(s) 2020
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guanicoe) in Patagonia during Late sagepub.com/journals-permissions

DOI: 10.1177/0959683620981689
https://doi.org/10.1177/0959683620981689

Pleistocene–Holocene and its importance

journals.sagepub.com/home/hol

for prehistoric human diet

Diego D Rindel,1,2 Bruno F Moscardi3

and S Ivan Perez2,3

Abstract
The great importance of the guanaco (Lama guanicoe) for the subsistence of the human populations of Patagonia during the Holocene has been demonstrated
in numerous studies. This species is considered as the main prey of the hunter-gatherer groups of the Patagonian interior, indicating the existence of a
strong ecological relationship of predator-prey type. Despite the importance of this ungulate for prehistoric human populations, its spatial distribution
throughout the Holocene has not been systematically explored, assuming that it was ubiquitously distributed in almost all of Patagonia. In this work, the
spatial distribution of the guanaco in Patagonia during different periods of the final Pleistocene and Holocene is explored using zooarchaeological and
paleontological evidence together with species distribution models and paleoclimatic reconstructions. Likewise, the spatial variation in the importance of
this species for the diet of human populations is assessed. The results show that there were large differences in the distribution of guanaco in Patagonia
over time. In general terms, this camelid suffered a constant contraction in range since the Late Pleistocene period to the present. There is an Andean
range and peri-cordilleran core of population distribution that remains observable during all this time. Human subsistence was strongly influenced by this
pattern of distribution. The differences in the guanaco distribution described in this work are important not only for archaeological and paleontological
problems, but also for the current conservation of guanacos.

Keywords
human subsistence, Lama guanicoe guanicoe, paleoclimatic data, paleontological information, species distribution models, zooarchaeological evidence

Received 21 August 2020; revised manuscript accepted 22 November 2020

Introduction 2008, 2013). The northern subspecies L. guanicoe cacsilensis,

After the megafaunal extinctions during the Pleistocene–Holo-
cene transition, which included the prehistoric horse (Hippidion
saldiasi) and the ground sloth (Mylodon darwini; Metcalf et al.,
2016; Villavicencio et al., 2016), the main surviving species in the
Patagonian plateau was the guanaco (Lama guanicoe; Borrero,
2009; Metcalf et al., 2016; Villavicencio et al., 2016). Phyloge-
netically, the guanaco is related to the Old World camels (order
Artiodactyla, family Camelidae, tribes Lamini and Camelini
respectively), whose ancestors probably diversified in central
North America approximately 16 million years ago (Heintzman
et al., 2015; Wheeler, 2012). Posteriorly, three million years ago,
the extinct Hemiauchenia crossed the Isthmus of Panama, colo-
nizing South America and giving origin to the genera Lama and
Vicugna. Within these genera, later diversification gave rise to the
guanaco, the vicuña (Vicugna vicugna), the llama (Lama glama)
and the alpaca (Lama pacos), which form the group of extant
South American camelids (Wheeler, 1995, 2012). The guanaco
and the vicuña are wild species, while the llama and the alpaca are
domestic species, products of the artificial selection of certain
traits carried out by diverse human populations between 6000 and
3500 years ago in different regions of the Andean area (Wheeler,
2012; Yacobaccio and Vilá, 2013). Among these species, the gua-
naco, with a body size ranging from 90 to 140 kg (Franklin, 2011),
is the most widely distributed of the South American camelids
and is currently differentiated into two subspecies (Marín et al.,
ranging from central Peru to central Chile, and L. guanicoe gua-
nicoe, distributed mainly in Patagonia, with small populations in
Cuyo and Pampa (Marín et al., 2008, 2013). Currently, the distri-
bution of this last subspecies is not homogenous within Patago-
nia, being more abundant in the Santa Cruz province of Argentina
due to bioclimatic conditions and anthropogenic factors (e.g.,
human settlements; Gavuzzo et al., 2015; Oliva et al., 2019; Tra-
vaini et al., 2015).
1
Instituto Nacional de Antropología y Pensamiento Latinoamericano
(INAPL), Ciudad Autónoma de Buenos Aires, Argentina
2
Consejo Nacional de Investigaciones Científicas y Técnicas
(CONICET), Argentina
3
División Antropología, Facultad de Ciencias Naturales y Museo
(FCNyM), Universidad Nacional de La Plata (UNLP), Argentina
Corresponding authors:
S Ivan Perez, División Antropología, Facultad de Ciencias Naturales y
Museo (FCNyM), Universidad Nacional de La Plata (UNLP), 122 y 60,
La Plata 1900, Argentina.
Email: ivanperezmorea@gmail.com
Diego D Rindel, Instituto Nacional de Antropología y Pensamiento
Latinoamericano (INAPL), 3 de febrero 1370, Ciudad Autónoma de
Buenos Aires, Buenos Aires C1426BJN, Argentina.
Email: drindelarqueo@yahoo.com
Rindel et al. 645
The archaeological studies show that L. guanicoe guanicoe
was the most important prey for prehistoric human populations
of the Patagonian inland during the Holocene (Borrero, 2009;
Mengoni Goñalons, 1999; Miotti and Salemme, 1999; Moscardi
et al., 2020). Particularly, the archaeological assemblages from
this region are mainly composed of guanaco bones that display
abundant evidences of human consumption (e.g. cut marks,
burned bones; Borrero, 2009; Mengoni Goñalons, 1999; Mos-
cardi et al., 2020). Moreover, archaeological and molecular data
for Patagonia indicate that after the Early Holocene both prehis-
toric human and guanaco populations grow rapidly, suggesting
the existence of a strong predator-prey ecological relationship
(Moscardi et al., 2020; Perez et al., 2016, 2017). The zooarchae-
ological evidence also shows that hunter-gatherers increased
their guanaco consumption after Middle Holocene (Bernal et al.,
2018; Moscardi et al., 2020; Perez et al., 2016). Despite the
importance of this camelid for prehistoric human populations
(Borrero, 2009; Moscardi et al., 2020), its spatial distribution
during the Holocene has not been systematically explored. Sev-
eral archaeological works assume that the guanaco was ubiqui-
tously distributed during the Holocene in almost all of Patagonia,
postulating higher abundance values than in the present (Men-
goni Goñalons, 1999; Miotti and Salemme, 1999; Perez et al.,
2016; Rindel, 2009), but probably displaying lower density val-
ues in North Patagonia (Borrero, 1981, 2008; Prates, 2008). The
greater abundance of this species in prehistoric times finds
empirical support in mitochondrial DNA data that indicate an
exponential growth of guanaco populations throughout the Holo-
cene, with an important decline towards historical times (Mos-
cardi et al., 2020; see also Perrig et al., 2019). Likewise, historical
records suggest a considerable reduction in guanaco populations
in recent centuries due to human hunting and other anthropo-
genic factors (Baldi et al., 2016; de Lamo, 2011). Overall, since
the guanaco suffered changes in its abundance and because its
presence is biased by human settlements (Gavuzzo et al., 2015),
the current spatial distribution is not representative of the ancient
dispersion.
The spatial distribution of the guanaco during the Holocene
can be explored using archaeological and paleontological locali-
ties. Although this spatial distribution can be directly studied
recording the localities where it was observed, the obtained maps
will probably be incomplete due to incomplete sampling and
taphonomic effects. In this context, species distribution models
(SDMs) have become a powerful modeling approach to estimate
maps of the potential distribution of a species (Elith and Leath-
wick, 2009). These models relate the species occurrence (or pres-
ence/absence) at specific locations to the main environmental or
bioclimatic conditions at those locations, using classical statisti-
cal methods (e.g. GLM) or machine-learning algorithms (Elith
and Leathwick, 2009; Hao et al., 2019). With the resulting model,
we can predict the spatial and temporal changes in species distri-
bution using the widely available bioclimatic layers (e.g., World-
Clim2; Fick et al., 2017). Although both data types, bioclimatic
layers and occurrence records, are important to estimate the
model, the final predicted distribution is made by projecting the
model onto the bioclimatic layers (Elith and Leathwick, 2009).
This approach has been extensively applied in ecology, biogeog-
raphy and conservation biology for studying the distribution of
extant species (Elith and Leathwick, 2009; Hao et al., 2019), and
more recently in paleontology (e.g., Varela and Fariña, 2016; Vil-
lavicencio et al., 2019). However, beyond its potential utility for
the study of the distribution of species in the past and the growing
availability of detailed paleo-bioclimatic reconstruction (Brown
et al., 2018; Fordham et al., 2017), species distribution models
have not been widely employed in archaeology to estimate the
distribution of main human prey (for an exception see Politis
et al., 2011).
The aim of this work is to estimate the distribution of the
main prey of Patagonian hunter-gatherers, the guanaco, for dif-
ferent periods between the end of the Late Pleistocene and the
present time. Because the hunter-gatherers groups in Patagonia
during the Holocene were small and mobile (Borrero, 1994–
1995; Miotti and Salemme, 1999), we assume that the ancient
guanaco distribution was mainly related to bioclimatic variables,
independently of human settlement. We expect that the guanaco
was ubiquitously distributed in almost all Patagonia during the
Holocene, with two moments of significant contraction: (a) after
the Late Pleistocene, associated to the large geographical and cli-
mate changes of Pleistocene–Holocene transition, and (b)
between Late Holocene and present times, mainly post 1800 AD,
related with the introduction of domestic livestock and the estab-
lishment of modern cities and towns, among other anthropogenic
factors. To explore these expectations, we use species distribu-
tion models together with archaeological and paleontological
occurrence records and paleo-bioclimatic models for the Late
Pleistocene and the main time periods or stages for the Holocene
(Cohen et al., 2020; Walker et al., 2018). Species distribution is
modeled using a maximum entropy approach (maxent; Phillips
et al., 2006, 2017), which uses environmental layers or biocli-
matic raster files and geo-referenced occurrence records to esti-
mate a model that predicts the suitability conditions for the
species presence in each raster pixel (Hijmans et al., 2017). We
employ this approach because is widely used in paleoecology
(e.g., Politis et al., 2011; Varela and Fariña, 2016; Villavicencio
et al., 2019) and is considered one of the most effective methods
for species distribution modeling with presence-only data (Elith
et al., 2010; Espinosa et al., 2018; Phillips et al., 2006; Politis
et al., 2011). Additionally, we explore the spatial variation in the
importance of guanaco for human diet in Patagonia during the
end of the Late Pleistocene and the main time periods of the
Holocene. The obtained results are used for discussing the impact
of paleo-distribution of guanaco in prehistoric hunter-gatherers
diet, as well as their implications for the sustainable management
and conservation of the species.
Methods
Guanaco occurrence data and bioclimatic layers
The guanaco distribution in the Late Pleistocene and three Holo-
cene periods was explored using an initial dataset of geographic
coordinates for 694 occurrence records compiled from archaeo-
logical and paleontological literature (Supplemental Table S1).
For all occurrence records included, we searched for radiocarbon
dates and calibrated them using SHCal13 curve and rcarbon
package for R software 3.4.3 (R-Development Core Team, 2019)
When radiocarbon dates were not available, reliable relative dates
given by the authors, based on contextual and/or stratigraphic
information (e.g., ceramic, European artifacts, extinct mega-
fauna), were taken into account. When an occurrence record did
meet these requirements it was included in the time periodifica-
tion (see below), otherwise, we excluded it from the dataset. We
also checked for very close sites in the same period, occupying
almost the same spatial location (~1 km), and deleted them to
avoid redundant records. Likewise, some archaeological and
paleontological sites with unreliable dates and stratigraphic and/
or taphonomic problems (e.g., Pali Aike) were not included in the
initial dataset. After sifting the database using these criteria, we
obtained a total of 327 presence and 60 absence data (Supplemen-
tal Table S2a and S2b). The dataset of the prehistoric guanaco
occurrence was divided in four time periods, according with
available paleoclimatic reconstructions (www.paleoclim.org;
Brown et al., 2018): Late Pleistocene (14.7–11.7 ka; N = 23),
Early Holocene (11.7–8.3 ka; N = 26), Middle Holocene (8.3–
4.2 ka; N = 51) and Late Holocene (4.2–0.3 ka; N = 227). The
646 The Holocene 31(4)
dataset of current occurrence of the Patagonian guanaco (N = 250)
was generated based on data available in the Global Biodiversity
Information Facility dataset (www.gbif.org; gbif code: 0052951-
200221144449610; Supplemental Table S3). All replicated data
points in the on-line database were deleted. The final sample size
for each period is within the range of other studies exploring the
paleo-distribution of species (e.g., Varela and Fariña, 2016; Varela
et al., 2010; Villavicencio et al., 2019).
For the species distribution modeling we used high-resolution
spatial climate data obtained from paleoclimate simulations
(Brown et al., 2018; Fordham et al., 2017). Specifically, we con-
sider as predictor the 19 standard bioclimatic variables available
in PaleoClim for the intervals: 14.7–12.9 ka (Bolling-Allerod),
11.7–8.3 ka (Greenlandian), 8.3–4.2 ka (Northgrippian) and 4.2–
0.3 ka (Meghalayan; www.paleoclim.org; Brown et al., 2018;
Fordham et al., 2017). These paleoclimate simulations were gen-
erated using general circulations models in atmosphere and
oceans, as well as considering shape of terrestrial landmasses,
CO2/N2O/CH4 levels and orbital parameters for a given time
period (Brown et al., 2018; Fordham et al., 2017). Specifically,
the paleoclimate simulations for the time period explored in this
work come from the climate model available in PaleoView
(Fordham et al., 2017) downscaled to a spatial resolution of
2.5 arc-minutes (~5 km; www.paleoclim.org; Brown et al., 2018).
The current species distribution was estimated using the same 19
standard bioclimatic variables available in WorldClim2, which
result from averaging the climate conditions for the years 1970–
2000 (www.worldclim.org; Fick et al., 2017). All current cli-
matic variables were obtained with a spatial resolution of
30 arc-seconds (~1 km).
Spatial distribution modeling of guanaco
Species distribution models have become the most widely used
modeling approach in quantitative ecology for projecting in the
space the potential for species distribution (Elith and Leathwick,
2009). These methods relate the species occurrence at specific
locations to the prevailing bioclimatic conditions – described in
spatial layers – at each location, and the resulting model is pro-
jected in the entire geographical space covered by the bioclimatic
variables. In this work, the spatial distribution of guanaco was
explored by using a maximum entropy distribution modeling
approach based on presence-only data (Phillips et al., 2006,
2017), as implemented in maxent function of the package Dismo
for the R software 3.4.3 (Hijmans et al., 2017; R-Development
Core Team, 2019). Maxent is a method proposed for making esti-
mations and predictions from incomplete data (Phillips et al.,
2006). In this sense, it is noteworthy that maximum entropy
method is considered the best option for species distribution mod-
eling when we used presence-only data (Elith et al., 2010; Espi-
nosa et al., 2018; Phillips et al., 2006; Pliscoff and Fuentes-Castillo,
2011; Politis et al., 2011). So, it is especially well suited for pale-
ontological and archaeological studies, which typically depends
on few presence data. The result of the maxent function in Dismo
is a potential for distribution value ranging from 0 to 1, indicating
the minimum and maximum potential for the species distribution.
Particularly, maxent estimates a potential for distribution by
exploring a probability distribution close to the uniform, the max-
imum entropy distribution, considering constraints that represent
our incomplete knowledge about the potential distribution (Phil-
lips et al., 2006). This method uses information on the potential
distribution of a species based on bioclimatic variables, or other
environmental variables, and the constraints are the expected
value of each variable where the species occurred, that is, its
empirical average (Phillips et al., 2006). In our case, we use bio-
climatic variables from WorldClim2 and PaleoClim, represented
as raster files, and presence-only data (i.e., locations with recorded
occurrence; Supplemental Tables S2 and S3) to predict the poten-
tial distribution of the species. Comparatively, we also explore the
potential for distribution of all archaeological sites using presence
data (i.e., locations of sites) obtained from Perez et al. (2016). We
compared distribution models for guanaco and all archaeological
sites, as well as among periods, using the I similarity index for
niche overlapping (Warren et al., 2008). This statistic is based on
the Hellinger distance and varies between 0 (no overlap) and 1
(total overlap).
The performance of each distribution model was evaluated
using the AUC statistic (i.e., area under the receiver operating
characteristic [ROC] curve; Elith and Leathwick, 2009), which
measure the explicative or discrimination power of the model,
using 80% of sample to train and the remaining 20%, together
with 1,000 random points as background data, to test the model,
considering 5 random k-folds or replicates for cross-validation. In
other words, the AUC statistic allows for evaluating the capacity
of our species distribution models to separate the presences from
the absence of the species (Jiménez-Valverde, 2014). In a ROC
curve, sensitivity or true positive rate (TPR; the probability that
the species is present) is plotted against commission or false posi-
tive rate (FPR), which is equal to 1 - TNR (specificity or true
negative rate, the probability that the species is absent). The mid-
dle or diagonal line in this graph, where the AUC value is 0.5,
represent models with discrimination values no better than the
ones obtained by chance. The AUC range between 0 and 1, where
values close to 1 show high discrimination power and a value
below 0.5 indicates that the model is worse than chance (Jiménez-
Valverde, 2014). Because AUC has showed some limitations in
practical applications (Allouche et al., 2006), the model perfor-
mance was also evaluated using maximum TSS (TPR+TNR-1;
that is, the threshold at which the sum of the true positive rate
[TPR: sensitivity] and true negative rate [TNR: specificity] is
highest; Allouche et al., 2006; Somodi et al., 2017). For the spe-
cies distribution models, we used the default arguments available
in the maxent function of Dismo that are relevant to model fitting.
The obtained prediction values or potential distribution surfaces
were saved as raster file and plotted in QGIS 3.14 “Pi” (QGIS-
Development Core Team, 2020) using a discrete interpolation.
Spatial and temporal variation in archaeological
guanaco exploitation
Whereas presence data and maxent algorithm only inform on the
values of potential distribution of the guanaco in Patagonia for
different periods and, therefore, about its potential availability for
human populations, the spatial variation in the economic impor-
tance of this species can be explored with other proxies. In this
regard, we compiled a dataset of 276 sites/assemblages corre-
sponding to all the major areas of Patagonia and southeast Pampa.
All sites/assemblages included in the database have geographic
coordinates, radiocarbon dates or reliable contextual chronologi-
cal information (especially in the case of the last period consid-
ered, with numerous assemblages corresponding to historical
moments). All sites included in the archaeological sample show
evidence of processing (cut marks, percussion marks and/or
burned bones). From these, we collect the guanaco NISP (NISPg;
Lyman, 2008) for each assemblage/sites included in the dataset,
as well as the total NISP of potential human prey. For the total
NISP estimation we use specimens determined at least at genera
level, excluding species that clearly are of taphonomic origin
(e.g., small rodents, unless they show human processing marks),
as well as dasipodid plates (when the original authors make this
distinction) and fragmented rhea’s eggs. We use these measures
(Supplemental Table S4) to evaluate the differential frequency of
guanaco remains in archaeological sites in the different regions of
Patagonia. We used NISP because it is the most frequently
Rindel et al. 647
Table 1.  AUC and TSS statistics for the maxent model of the studied periods. The mean and deviation values for these statistics were
calculated on five replicates.
Periods AUC Mean-AUC Deviation-AUC
Late Pleistocene 0.9850 0.9629 0.0247
Early Holocene 0.9942 0.9637 0.0081
Middle Holocene 0.9767 0.9699 0.0088
Late Holocene 0.9819 0.9784 0.0019
Current 0.9877 0.9883 0.0038
published zooarchaeological data in Patagonia. However, it is
important to consider that because NISP considers each fragment
bone as a unit, this measurement is sensitive to fragmentation pat-
terns by human processing or taphonomic processes (Grayson,
1984; Lyman, 2008). Alternative measurements, such as the Min-
imal Number of Individuals (MNI), which is robust to the effect
of these factors, were not used because they are not widely avail-
able in the literature.
We describe the spatial and temporal variation in the con-
sumption of guanaco for prehistoric hunter-gatherers by
directly plotting the guanaco NISP divided by the total NISP of
potential human prey (prop-NISPg) values, as well as the abso-
lute values of NISPg, using a color gradient on the map of
current Patagonia. The location of each point represents the
geographical coordinates of the archaeological site from which
the NISP values were obtained. For sites including several
components or assemblages, in order to avoid biases in the
exploration of the spatial differences in the contribution of
guanaco to human diet along Late Pleistocene–Holocene,
we calculated the median for relative and absolute values. Due
to the large differences in the absolute NISPg values along
the region, we smoothed the variable using a log10-transfor-
mation. The spatial variation in the NISP of guanaco was
graphed using QGIS 3.14 “Pi” (QGIS-Development Core
Team, 2020).
Results
Guanaco potential distribution
The predictive capacity of maxent is high for all models, with
values of AUC higher than 0.95, but with TSS showing lower
values for the most recent periods (Table 1). The potential distri-
butions for the four prehistoric periods and current time studied
here are presented in the Figures 1 and 2. In these maps, grey
represents areas close to 0 potential of finding guanacos and
strong red represents areas with high potential. Dramatic
changes in guanaco distribution over time are observed. These
distribution patterns are not identical to the distribution and den-
sity of all archaeological sites, such as it is observed in Supple-
mental Figures S1 and S2. The I similarity values between
guanaco and all sites distributions vary between 0.92 and 0.96
(Late Holocene = 0.952; Middle Holocene = 0.927; Early Holo-
cene = 0.962; Late Pleistocene = 0.938). The main differences
are observed for Central Chile and the Atlantic coast; where
there is a high density of archaeological sites for several periods,
but not guanaco records. The relative contribution of the biocli-
matic variables to the final maxent model is presented in Supple-
mental Table S5. The mean temperature of coldest quarter
(Bio_11) is a main bioclimatic variable for all models; with tem-
perature seasonality (Bio_4) being important for the two ancient
periods, the precipitation of coldest quarter (Bio_19) for the
Middle Holocene, isothermality (Bio_3) for Late Holocene and
the precipitation of wettest month (Bio_13) for the current
distribution. When we plotted the values of these variables
for the geographical coordinates of the sites for the different
periods (Supplemental Figure S3), we observed that the current
TSS Mean-TSS Deviation-TSS
0.5222 0.2981 0.2063
0.7565 0.2876 0.1754
0.4602 0.2339 0.0973
0.2517 0.1921 0.0414
0.0150 0.0889 0.0681
distribution is the period with the most different behavior. A
Kruskal-Wallis test confirms these differences, mainly for the
temperature seasonality and precipitation of the wettest month.
The maximum predicted potential distribution is observed in
the Late Pleistocene. The core of potential distribution for this
period occurs along the Andean range and its eastern slopes,
including large portions of inland Patagonia, especially the
Deseado Massif and neighboring areas. The predicted values of
potential distribution are intermediate for Pampa and Tierra del
Fuego, being low for the rest of Patagonia (Figure 1). In the Early
Holocene there is a notable retraction in the predicted distribution
of the guanaco, mainly in the Patagonian inland (Figure 1). From
north to south, there are continuous high values of potential distri-
bution along the Andean range and eastern slopes, from south
Mendoza to Tierra del Fuego. The high predicted potential in the
Deseado Massif and neighboring areas is spatially more restricted,
but continues to spread to the Atlantic coast. Areas with intermedi-
ate values of potential distribution are observed across northern
Patagonia and southeast Pampa. There is also high predicted
potential distribution in the Atlantic coast of the current province
of Chubut (Figure 1). In the Middle Holocene there is again a
retraction in the predicted distribution of the guanaco in the Pata-
gonian inland and a high potential distribution along the Andean
range and eastern slopes (Figure 1). An expansion of the high
potential distribution values is also observed in northeast Patago-
nia and southeast Pampa with respect to previous periods. In the
rest of Patagonia there are predominantly low potential distribu-
tion values, but with patches of intermediate potential values in the
Deseado Massif and some regions of the coast. Finally, in the Late
Holocene, the Andean range and eastern slopes remains as a sector
with high predicted potential distribution of guanacos, but the val-
ues are moderate in some places (Figure 1). Likewise, compared to
the previous period, the trend towards high potential values
increased in the Atlantic coast, including the southeast Pampa.
The current potential distribution of guanaco populations is
presented in Figure 2, indicating a further contraction compared
to the distribution of the species in the prehistoric periods. Sup-
plemental Figure S4 presents a UPGMA (Unweighted Pair Group
Method with Arithmetic mean) cluster analysis of I similarity,
clearly showing that the current distribution of guanaco is the
most different among all estimated distribution maps. High pre-
dicted values of potential distribution are observed in the south-
ernmost extreme of continental Patagonia, northern Tierra del
Fuego, Atlantic coastal sectors of Chubut province and isolated
patches in northern Neuquén and southern Mendoza provinces.
Intermediate values of potential distribution are observed in the
central portion of Tierra del Fuego and in some coastal and inte-
rior areas of the southern tip of Santa Cruz province. Low values
of potential distribution are found in the central portion of conti-
nental Patagonia and in the southeast of Pampa (Figure 2).
Spatial and temporal variation in guanaco
exploitation
The spatial variation in the relative consumption of guanaco along
the Late Pleistocene–Holocene is graphed in Figure 3 and the
648 The Holocene 31(4)
Figure 1.  Guanaco potential distribution for the studied periods of Late Pleistocene–Holocene.
absolute values in Supplemental Figure S5. The NISP values indi-
cate a high use of this species only in southern Patagonia (Deseado
Massif, Santa Cruz) at Late Pleistocene, with lower values in
Última Esperanza (Chile), the Andean range in North Patagonia
(Río Negro province) and southeast Pampa. Towards the Early
Holocene, a high consumption of guanaco is observed in southern
Patagonia (Deseado Massif) and the Andean range in North Pata-
gonia (Río Negro and Neuquén [south portion] provinces). Lower
frequencies of human exploitation are observed in other regions
of Patagonia, such as North Neuquén and South Mendoza, Pampa
and Tierra del Fuego. In the Middle Holocene, high evidence of
consumption is observed in the same regions as in the previous
period, but with an increase in guanaco consumption in southern
Tierra del Fuego and southeast Pampa. A continuous increase in
the quantity of sites with evidence of guanaco consumption is
observed globally for Patagonia from the Late Pleistocene to the
Late Holocene, being considerably higher in the last period (Fig-
ures 3 and 4). In the Late Holocene, with the highest relative
frequency of sites with evidence of human exploitation (Figure
4), the regions with relatively higher consumption are the same as
in the previous period (Figure 3 and Supplemental Figure S5). It
is noteworthy that the level of guanaco consumption in the Atlan-
tic coast is low or absent in all periods, which contrasts to the
evidence obtained in the interior of Patagonia, especially in its
southernmost portion. In this regard, low evidence of consump-
tion appears on Río Negro (Figures 3 and Supplemental Figure
S5) and Chubut coasts (Supplemental Figure S5) in the Late
Holocene, but it is comparatively low or moderate.
In Figure 5 the spatial distribution of zooarchaeological
assemblages without the presence of guanaco is displayed. This
figure shows the archaeological sites where there is no guanaco
and could represent the limits of the region with preponderance of
guanaco and how it changed over time. For the Late Pleistocene
there are zooarchaeological assemblages without guanaco in the
southern portion of continental Patagonia and in southeast Pampa.
In the Early Holocene, all recorded sites present guanaco. By
Rindel et al. 649
Figure 2.  Current guanaco potential distribution in the study area.
The outlier points observed in North Patagonia and South Pampa
correspond to isolated small guanaco groups inhabiting Protected
Natural Areas, which clearly are outside the current optimal
conditions for the distribution of the species.
Middle Holocene, zooarchaeological assemblages without gua-
naco are observed on the coast of southern Tierra del Fuego and
Chubut, as well as in some sites of the Andean range, mainly in
North Patagonia. This distribution is similar to that of the Late
Holocene, but in this last period the assemblages without guanaco
are registered in the Atlantic coast in Tierra del Fuego, Santa Cruz
and Buenos Aires provinces. The relative changes in the fre-
quency of sites without evidence of guanaco consumption
throughout the Late Pleistocene–Holocene are similar to those
observed in sites with presence of this species (Figure 4).
Discussion
Guanaco distribution during Late Pleistocene–
Holocene
Our results show important changes in the distribution of gua-
naco, and its consequent availability for human populations in
Patagonia over time, including the Pampa region as the northeast
limit. The widely accepted idea that the guanaco could be homo-
geneously distributed in Patagonia during the Holocene (Garrido
et al., 1981; Moscardi et al., 2020; Raedeke, 1979; Rindel, 2009;
among others), should be discarded. This species did not show a
continuous distribution in any period and suffered a continuous
range contraction since the Late Pleistocene period to the present
time (Figure 1 and Supplemental Figure S4). Additionally, at the
same time that the guanaco underwent a decrease in its potential
distribution, the Patagonian territory experienced a large surface
area reduction, from the largest surface area in Late Pleistocene
– which was around twice the current territory – to the current
surface area, established after ca. 8000 years BP (Cione et al.,
2009; Porter et al., 1984). Additionally, all of these changes in
available surface area and distribution during the Holocene
occurred in a context of exponential demographic growth of
guanaco populations, probable following a trajectory that was not
constrained by human predation (Moscardi et al., 2020).
Although the distribution of guanaco changed over time, there
is a core of population that remains from Late Pleistocene to the
present time in the Andean range and eastern slopes. All of these
areas have been characterized by presenting rich grasslands,
which varied in extension after Late Pleistocene (McCulloch
et al., 2000), and/or by being close to rich seasonal pastures (Oliva
et al., 2001). Because the potential distribution for Andes range
and nearby areas remains unchanged throughout the different
periods, this region could be of particular relevance for guanaco
dynamics in the past. In the present time, this area has high pri-
mary productivity (Oliva et al., 2001), and probably the produc-
tivity was even higher during the late Pleistocene and Early
Holocene (Cione et al. 2009; McCulloch et al., 2000; Tonni and
Carlini, 2008; Villavicencio et al., 2016), explaining the high val-
ues of potential distribution of guanacos. Although the guanaco
does not have an extreme dependence on water –since it is not an
obligate drinker (Rick and Moore, 2001; Yacobaccio et al., 2017)
and there are even anecdotal reports that it can drink salty water
(Darwin, 1845; Gerken et al., 2019)–, the high potential for distri-
bution in the Andean range also could be related to larger water
availability in this area or to indirect water supply on the pastures
that the animals consume. Outside this area, some steppe environ-
ments present variable but generally high potential for guanaco
distribution, such as the Deseado Massif. However, in the other
sectors of Patagonia, especially in its northern portion, there are
always low potential values. The southeast Pampa and Patagonian
Atlantic coast present a variable distribution, with periods with
high potential values, especially in the Late Holocene, and peri-
ods with a low potential distribution for guanaco.
The results of species distribution models can be influenced
by biases and uncertainties associated with problems in data sur-
vey, such as poor taxonomic identification, inaccurate recording
of the geographical coordinates of the locations and the existence
of a long-time from the original observation or record (Meyer
et al., 2016; Tessarolo et al., 2014). In our study, we avoid these
problems by using records coming from identifications realized
by professional archaeologists in well excavated and accurately
dated archaeological sites; almost all studied or re-studied in the
last two decades (Supplemental Table S2). However, the coverage
of our survey design could be biased by the distribution of archae-
ological sites, our main source for presence data of guanaco in the
past. Notwithstanding, it is remarkable that our results suggest
that the distribution pattern of guanaco for different periods are
not directly related to the distribution and density of archaeologi-
cal sites, such as it is observed in Supplemental Figures S1 and
S2. Although the I similarity indexes showed high values between
the distribution of both types of data, the distribution maps
inferred from all archaeological sites from Patagonia displayed a
wider distribution. The distribution of guanaco and all archaeo-
logical sites mainly differ in the Atlantic coast and Central Chile,
where all archaeological sites displayed high values of potential
distribution (Supplemental Figure S1) and higher density values
(Supplemental Figure S2). This pattern of distribution differs
from that estimated for guanaco in different periods (Figure 1),
suggesting that our paleo-distribution estimations are robust and
do not depend directly on the distribution of archaeological sites.
Andes range distribution and seasonality
The high potential distribution in the Andes range and nearby
areas throughout the different periods has several implications
for guanaco and human population dynamics. In the present
time, the areas closest to the Andes range are subject to intense
seasonality, and are practically uninhabitable during the winter
months (Pallo, 2012; Rindel, 2009). This has consequences on
650 The Holocene 31(4)

Figure 3.  Archaeological relative guanaco samples (prop-NISP) calculated as NISPg/total-NISP per period in the study area using quantiles for
distributing the values.

Figure 4.  Relative frequency of sites with and without guanaco throughout Late Pleistocene–Holocene. Consider that the total of sites with
guanaco is 327 and without guanaco 60.
Rindel et al. 651
Figure 5.  Sites without evidence of guanaco consumption
(Supplemental Table S2b).
several aspects of the socio-ecology of guanaco, such as diet,
patterns of use of space and social organization (Puig and Videla
1995, 2000; Raedeke, 1978). Specifically, some groups of guana-
cos are migratory and others sedentary, usually having the last
ones very small ranges of action and high territoriality (Burgi,
2017). The most documented current populations that carry out
migratory movements are placed in Torres del Paine (southern
Chile; Ortega and Franklin, 1995), Cuncumén and Chillepin
(northern Chile; Contreras et al., 2006) and La Payunia (Men-
doza-Neuquén, Argentina; Bolgeri, 2016). Some migratory
movements are large (like in La Payunia, 85 km on average)
while others are shorter (12 km in Torres del Paine), including
altitudinal mobility (as in Cuncumén and Chillepin, from 1000 to
4000 m.a.s.l). These movements probably were larger in the past
(Claraz, 1988; Moreno, 1969; Musters, 1964; discussed by Bol-
geri, 2016). The topographic complexity of a region seems to be
one of the intervening factors in the movements; when there are
important altitudinal differences in reduced areas, migrations are
less extensive, while movements are greater in more homoge-
neous areas. In this sense, the movements seem to be mainly gov-
erned by the seasonal availability of food (Baldi et al., 2016;
Contreras et al., 2006; Ortega and Franklin, 1995). The observed
mobility patterns in different populations of guanacos could have
several consequences for the ecology and evolution of the spe-
cies. In this sense, it has been observed in African ungulate spe-
cies such as wildebeest and in North American bison, pronghorn
and elk that, in the absence of effective controls by predators,
migrations are presented as a way of maintaining high population
densities that would not be possible in sedentary populations
(Milner-Gulland et al., 2011).
However, it is not clear if the migratory groups observed in
guanacos could be a relatively recent phenomenon, product of
density-dependent processes of the guanaco populations, or if it
was present in the prehistoric populations. Some archaeological
works have strongly supported this last conclusion for northwest
(Hajduk and Lezcano, 2007) and southern Patagonia (Goñi,
2010). This is the case, for example, of the establishment of logis-
tically organized guanaco hunting systems in Santa Cruz on the
Pampa del Asador, Guitarra, Strobel, Cardiel Chico, San Adolfo,
and Lake Buenos Aires plateaux, with occupations that begin in
the Middle Holocene and reach their maximum development
during the Late Holocene (Belardi et al., 2013; Dellepiane, 2019;
Goñi, 2000, 2010, Goñi et al., 2000–2002; Gradín, 1976, 1996;
Rindel, 2009). Although there are fewer systematic studies, the
evidence for the Tromen Massif in North Neuquén also suggests
logistically organized guanaco hunting systems (Rughini et al.,
2020). These practices were maintained until historical times in
Patagonia. In this sense, several ethnohistorical sources refer to
patterns of seasonal use of the Andean range and peri-cordilleran
high plateaux, particularly mentioning that practice in a hunting-
gathering context (Coan, 2006; Moreno, 1969; Musters, 1964;
Prichard, 2003; among others). From a discussion of historical
sources, Crivelli Montero (2017) points out a similar exploitation
in northwestern Patagonia, which remains to this date in the area
in a pastoral context. Conversely, certain historical sources
account for the inhospitable, waterless and resourceless character
of some areas of the Patagonian interior, especially in the northern
portion of the study area (Claraz, 1988; Musters, 1964).
Impact of guanaco on hunter-gatherer distribution
and diet
The geographical distribution and seasonal movement of guanaco
populations, together with previously described demographic
changes (Moscardi et al., 2020), have probably impacted in
human population distributions and subsistence during the Late
Pleistocene–Holocene (Miotti and Salemme, 1999; Moscardi
et al., 2020; Perez et al., 2016). Our results show that several areas
with low potential distribution of guanacos also present little evi-
dence of human presence in Patagonia, such as it was observed
from the spatial density of calibrated radiocarbon dates (Supple-
mental Figure S2; Perez et al., 2016) and suggested by the I simi-
larity results. This agreement is especially evident in several
sectors of the inland Patagonia, which has lower values of poten-
tial distribution of guanaco and lower frequencies of human occu-
pation than neighboring areas (Figure 1, Supplemental Figures S1
and S2; Perez et al., 2016).
Also, our results show a high agreement between the areas
with high potential distribution and high consumption of gua-
naco, as it was observed from the absolute and relative NISP for
guanaco, a proxy of human consumption. Some areas, such as the
Deseado Massif and the Andes range in South Patagonia and
neighboring zones, have high values of potential distribution and
high NISP of guanaco from the end of Late Pleistocene to Late
Holocene. The archaeological record in these regions is character-
ized by a heavy dependence on guanaco (Dellepiane, 2019; De
Nigris, 2004; De Nigris and Catá, 2005; Rindel, 2009; Rindel and
Bourlot, 2014). The same result is observed for northwest Patago-
nia (northern Neuquén and southern Mendoza) during the Late
Holocene (Cordero, 2007; Durán, 2000; Fernández, 2008; Neme
et al., 1999; Otaola et al., 2012). Towards the Middle and Late
Holocene the importance of guanaco is also registered in south-
east Pampa, where the archaeological sites show a diet strongly
orientated to the consumption of this species (Álvarez, 2014;
Álvarez and Salemme, 2015; Bonomo and Massigoge, 2004;
Martínez et al., 2005; Messineo et al., 2013; Scheifler, 2019). For
this region, it is interesting to note that the fluctuations in the
potential availability of guanaco are consistent with trends in the
zooarchaeological record, where a generalized strategy in
resource consumption is detected during the Late Pleistocene,
Early Holocene and the first part of the Middle Holocene, while
in the second part of the Middle Holocene and during Late Holo-
cene, the diet heavily focuses on guanaco consumption (Álvarez,
2014; Gutiérrez and Martínez, 2008; Scheifler, 2019). The south-
east Pampa could be a limit of the distribution of guanaco, as can
be seen in the advances and contractions in the guanaco potential
distribution throughout the Holocene (Politis et al., 2011). It is
important to note, however, that low values of potential for
652 The Holocene 31(4)
Figure 6.  Guanaco potential distribution for Late Holocene and the current time, together with the terrestrial ecoregions of the
Southernmost South America (WWF’s ecoregions, see http://maps.tnc.org/gis_data.html; Olson and Dinerstein, 2002).
distribution should not necessarily be understood as insufficient
for human needs. In fact, our results indicate that in areas with
low estimated potential for distribution of guanaco (for example
in northern Patagonia) this animal is present in some archaeologi-
cal sites. However, it is also evident from previous discussion that
in areas of low potential for distribution of guanaco, alternative
resources were exploited in a larger frequency (e.g., Borrero,
1981; Prates, 2008).
In this context, there is also a good fit between the zooarchaeo-
logical assemblages or sites that did not register the presence of
guanaco and our paleo-distribution results. These sites show a
peripheral distribution with respect to the areas with high poten-
tial for guanaco distribution. Particularly, in the forest areas of
both slopes of the Andes range of Santa Cruz, Chubut and Río
Negro, as well as those of the Chilean Patagonia, which do not
show high consumption of guanaco, the prehistoric groups sub-
sisted on the basis of the exploitation of huemul, pudu, foxes,
birds, and other small prey (Adán et al., 2001; Fernández et al.,
2015, 2019; Fuentes and Mena Larraín, 2010; among others). On
the other hand, in the northern sectors of Patagonia there is an
early use of lesser fauna: mollusks, fish, birds, deer, dasipodids,
and rodents have been part of the human diet since the early times
of occupation of the area (Berón, 2004; Mange, 2019; Martínez
et al., 2014; Prates, 2008; Rindel et al., in press; Stoessel and
Alcaráz, 2017). On the coast it can be observed that both the pres-
ence of guanaco and its frequency of consumption are discontinu-
ous. Here the relatively early use of marine resources such as
marine mammals, fish and mollusks made human occupation
possible. The exploration of coastal resources begins in different
parts of Pampa (7600 BP; Bonomo et al., 2013), continental Pata-
gonia (7200 BP; Gómez Otero, 2006; Favier Dubois and Scartas-
cini, 2012; Zubimendi et al., 2015) and Tierra del Fuego (6500
BP; Zangrando, 2009) during the Middle Holocene. This period is
characterized by a low potential distribution of guanaco along the
coast, but a relatively high density of archaeological sites (Sup-
plemental Figure S2). However, the frequency of marine resources
is variable along space and time in the Atlantic coast (Moreno
et al., 2011; Scartascini, 2014, 2017). Although the use of coastal
resources is well established in all sectors towards 2500 years BP,
the main frequency of use on the north Patagonian and Pampa
coast occurs between 4900 and 2200 years BP, on the Chubut
coast between 1000-400 years BP and an increase in the use of
marine resources on the Santa Cruz coast only occurs in the last
1000 years BP (Moreno et al., 2011; Scartascini, 2014, 2017; Zilio
et al., 2014). In this sense, this variability in the appearance and
development of coastal occupations was probably related to the
discontinuous potential distribution of the guanaco on the coast
from the Middle Holocene onwards (Figure 1 and Supplemental
Figure S1).
Implications for current guanaco distribution
Finally, it is important to highlight the relevance of our results not
only for archaeological and paleontological problems, but also for
the sustainable management and conservation of guanacos in the
present times. We must remark that the core of high potential dis-
tribution that remained almost invariant over time in the Andean
range and eastern slopes is no longer maintained in the current
distribution of guanaco populations. This situation generates iso-
lated patches of guanaco populations in the north (Figure 6), such
as has been discussed previously by Gavuzzo et al. (2015), Tra-
vaini et al. (2015) and Oliva et al. (2019). Several factors could
have generated the differences between the current and past gua-
naco distributions (Supplemental Figure S4), shaping the largest
restriction in the former. The main explored bioclimatic variables
have shown similar ranges of variation, but differences in median
values, between the current distribution and the past ones (Sup-
plemental Figure S3). In this context, for 10,000 years the gua-
naco was almost the only species occupying the large herbivore
niche of Patagonia (Hernández et al. 2019). This has probably
made the animal a poor competitor (Baldi et al., 2004; Raedeke,
1982; but see Iranzo et al., 2013), which is reflected in the current
inverse relationship between the guanaco and domestic fauna
density, especially ovine livestock (Baldi et al., 2004; Marino
Rindel et al. 653
et al., in press; Pedrana et al., 2010). Moreover, the current pres-
sure of human activities, which is partly related to the human
settlements (i.e. cities and towns), produces subsequent fragmen-
tation of habitats of these isolated northern populations, as cur-
rently occurs with the guanaco populations of Auca Mahuida and
La Payunia (Novaro et al., 2007; Rivas et al., 2015). In this sense,
the distribution and densities surveyed by current transects of
guanaco populations addresses not only the preference of certain
environments, but also the effect that human activities have on
this species (Gavuzzo et al., 2015; Marino et al., in press; Oliva
et al., 2019; Travaini et al., 2015). It is evident that a suitable
conservation strategy for this species must contemplate these
situations.
Conclusion
By using paleontological and archaeological data, together with
species distribution models, we have observed along the late
Pleistocene–Holocene a continuous reduction in guanaco distri-
bution. The geographic restriction of guanaco is more pronounced
in the present time. These differences in guanaco distribution dur-
ing the late Pleistocene and Holocene probably impacted human
demography and subsistence, generating a background upon
which human populations have diversified culturally. By address-
ing the distribution patterns of guanaco in the past, we also have
established an ecological reference to compare with the current
distribution, allowing us to measure the present and past human
impact on this species. Future studies combining paleontological
and archaeological information with ecological modelling will
allow a better understanding of the factors driving past and pres-
ent communities.
Acknowledgements
We are grateful to Fernando Oliva for his useful bibliographic sug-
gestions. We thank Florencia Gordón and Valeria Bernal for the
comments made to an earlier version of the manuscript. We also
thank three anonymous reviewers who help to improve this work.
Funding
The author(s) disclosed receipt of the following financial support
for the research, authorship, and/or publication of this article:
This work was supported by grants from Agencia Nacional de
Promoción Científica y Tecnológica (PICT 2014-2020/2134),
Universidad Nacional de La Plata (PI 2020-2023/N932), Con-
sejo Nacional de Investigaciones Científicas y Técnicas (PIP
2015-2020/729).
ORCID iD
S. Ivan Perez https://orcid.org/0000-0002-6543-5545
Supplemental material
Supplemental material for this article is available online.
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