ARTICLE IN PRESS

Journal of Archaeological Science xxx (2009) 1–10

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Journal of Archaeological Science

journal homepage: http://www.elsevier.com/locate/jas

Guanaco (Lama guanicoe) isotopic ecology in southern South America:

spatial and temporal tendencies, and archaeological implications
Ramiro Barberena a, *, A. Zangrando b, Adolfo F. Gil c, Gustavo A. Martı́nez d, Gustavo G. Politis e,
Luis A. Borrero f, Gustavo A. Neme c
a
CONICET – IMHICIHU, Saavedra 15, 5 (1083), Buenos Aires, Argentina
b
CONICET – CADIC/Universidad de Buenos Aires. B. Houssay 200, Ushuaia, Tierra del Fuego, Argentina
c
CONICET – Museo de Historia Natural de San Rafael. Parque Mariano Moreno (5600), San Rafael, Mendoza, Argentina
d
CONICET – INCUAPA. Avenida del Valle 5737 (7400), Facultad de Ciencias Sociales, Universidad Nacional del Centro de la Provincia de Buenos Aires, Olavarrı́a, Argentina
e
CONICET – INCUAPA -Universidad Nacional del Centro de la Provincia de Buenos Aires, Universidad Nacional de La Plata, La Plata, Argentina
f
CONICET – IMHICIHU/Universidad de Buenos Aires. Saavedra 15, 5 (1083), Buenos Aires, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: We synthesize and discuss available isotopic data on Holocene guanaco samples from southern South
Received 20 June 2009 America, extending from Tierra del Fuego Island to northern Patagonia and the Pampean region. We
Received in revised form evaluate temporal and spatial tendencies on the basis of 91 samples (mainly based on d13Ccollagen values),
6 August 2009
presenting implications for paleodietary research in archaeology. We conclude that there are no strong
Accepted 11 August 2009
correlations with latitude in the macro-spatial scale, while there is a set of interesting patterns at smaller
regional scales. These patterns lead us to evaluate the role of ecologic and topographic variables
Keywords:
(e.g. canopy effect, variations in altitude, ecotones) in structuring isotopic variability. Finally, on the basis
Guanaco
Stable isotopes of the present analysis we suggest a number of hypotheses and perspectives for the use of stable isotopes
d13Ccollagen as geographic tracers of guanaco distribution in the past, and specific implications for the paleodietary
Spatial tendencies study of human samples.
Geographic tracers Ó 2009 Elsevier Ltd. All rights reserved.
South America

1. Introduction supra-regional unit of analysis covers a latitudinal range of 20 and

The zooarchaeological record from the South American Pampas
and Patagonia indicates that guanaco (Lama guanicoe) was the
main terrestrial staple for Holocene hunter-gatherer populations
(Borrero, 1990; Mengoni Goñalons, 1999; Politis and Pedrotta,
2006; Gutiérrez and Martı́nez, 2008; Neme and Gil, 2008; Morales
et al., 2009). Stable isotopes on human remains provide an inde-
pendent line of paleodietary evidence that offers quantitative
information. Nevertheless, stable isotopes data does not have an
intrinsic meaning and requires a contextual approach taking into
account local climatic and biogeographical conditions. Isotopic
ecology provides that context and offers a frame of reference for
the interpretation of isotopic data on human remains (Burton
et al., 2001). In this paper we synthesize the available isotopic data
for guanaco from Tierra del Fuego Island to northern Patagonia
and the Pampean regions in Argentina (the northern limit being
set at central Mendoza and central Buenos Aires provinces). This
* Corresponding author.
E-mail address: ramidus28@gmail.com (R. Barberena).
ca. 2400 lineal km, being located between 54 and 34 S (Fig. 1).
We begin presenting and discussing temporal and spatial
patterns by using stable isotopes as a dietary tracer for guanaco
samples. Then, we explore the potential of this information as
a geographic tracer of guanaco distribution in the past. Recent
research shows the important role that this data may have in the
context of species conservation decisions and management of
protected spaces (Etnier, 2004). Although it is not the main goal of
this paper, this supra-regional analysis will contribute to
a comparative evaluation of -among other issues- the human
consumption of marine and domesticated resources, such as maize.
1.1. Fitogeographic regions
Five main fitogeographic regions are currently represented in
the study area depicted in Fig. 1, all of which are -or were in recent
times- inhabited by guanacos: Pampean, Espinal, Monte, Patagonia,
and Sub-Antarctic (Cabrera, 1976; González et al., 2006; Abraham
et al., 2009). The Pampean fitogeographic Province occupies the
eastern plains between 31 and 39 south. Weather is temperate
with mean annual precipitations between 600 and 1100 mm.

0305-4403/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2009.08.003

Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
 ARTICLE IN PRESS
2 R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10
Fig. 1. Study area and fitogeographic regions. References: Study areas: 1. Central-
southern Mendoza, 2. Humid Pampas, 3. Central-northern Patagonia, 4. Southern Pata-
gonia, steppe, 5. Southern Patagonia, forest, 6. Tierra del Fuego. Fitogeographic regions:
green, Pampean Province; black, Espinal Province; yellow, Monte Province; grey,
Patagonia Province; straight lines, Sub-Antarctic Province (based on Cabrera, 1976).
The main vegetal community represented is the herbaceous steppe
with the genera Stipa, Poa, and Briza among the most represented.
The Espinal Province constitutes a belt surrounding the Pampa to
the west (Fig. 1), with mean precipitations between 500 and
1100 mm associated with continental conditions that produce
higher evapo-transpiration rates than in the Pampa. The main
communities represented in the areas considered here are shrub
and herbaceous steppes, and xerophytic woodlands with the
genera Prosopis, Acacia, and Schinus mostly represented. The Monte
Province coincides with the so called South American Arid Diagonal
(Bruniard, 1982), with mean annual precipitations between 80 and
200 mm. The main community is a xerophytic shrub steppe rep-
resented by Larrea and Prosopis; importantly, C4 and CAM taxa may
be present in varying densities and have been isotopically identi-
fied (Gómez Otero, 2007; Martı́nez et al., 2009).
The Patagonia Province extends from southern Mendoza
(36 south) up to Tierra del Fuego Island (53 south); it is bounded
by the Monte to the east and by the Sub-Antarctic forests to the
west (see below). The climate is dry and cold with mean annual
precipitations between 150 and 500 mm. The main communities
Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
are shrub and herbaceous steppes that may have varying frequencies
of C3 and C4 plants (although the former are usually dominant). The
Sub-Antarctic Province is closely associated to the Andes mountains,
that flank the western side of South America from 37 to 54 south
(at Tierra del Fuego Island; Fig. 1). The climate is temperate-cold and
humid with precipitations between 800 and 2000 mm. The domi-
nant communities are deciduous and ever-green forests where
Nothofagus is the main genera (Boelcke et al., 1985).
1.2. Guanaco ecology
The guanaco (L. guanicoe) is the larger terrestrial wild mammal
in southern South America, which weighs between 85 and 120 kg
in average for different regions (recent review in González et al.,
2006; see Fig. 2). This camelid lives in herds composed of females,
young individuals (chulengos) and a dominant male. Bachelor males
form a separate herd (Raedeke, 1978). Guanacos inhabit preferably
in open areas with grasslands, flat terrain and low hiding cover
(Cajal, 1980), although they are known to inhabit forested envi-
ronments in Tierra del Fuego Island (Raedeke, 1978). This species
typically feeds on the herbaceous stratum (grasses, grass-likes
[Juncaceae and Ciperaceae], and forbs), but also has the ability to
alternate seasonally between grazing and browsing, according to
forage availability; this behavior allows to define the guanaco as an
‘adaptable mixed feeder’ that has the ability to digest low quality
forage (Puig et al., 1996; González et al., 2006). There are records
from the Monte Province indicating that guanacos do not strongly
select against C4 grasses due to its capacity to digest high fiber-
content taxa (Puig et al., 1996; Llano, 2009).
Until the Nineteenth century, the guanaco was present in almost
all regions of Argentina, occupying the different fitogeographic
regions described, from open habitats and scrubland to open
forests (Cajal, 1980) and extending from the sea level up to
4500 masl. Currently guanacos are more abundant in the Patago-
nian steppe and in the foothills of the Andean Mountains (Cajal,
1980). Guanaco populations from steppe, pre-cordillera, cordillera
and transition zone between cordillera and steppe zones have been
well studied with regard to food habits (Raedeke, 1978; Ortega and
Franklin, 1988; Bonino and Pelliza Sbriller, 1991; Puig et al., 1996).
However, population dynamics and habitat use of guanacos in
those regions are poorly known. Stable isotopes from archaeolog-
ical samples can contribute to develop this endeavor in Holocene
times.
2. Methodology
This paper provides a comprehensive synthesis of guanaco
stable isotope values in southern South America. The main criterion
for the inclusion of the samples is the existence of accurate
Fig. 2. Guanaco troop in Tierra del Fuego Island (53 S).
 ARTICLE IN PRESS
R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10
taxonomic determinations, in order to exclude specimens
belonging to other Lama species. This is a key issue north of ca.
34 S, given the presence of wild and domesticated camelid species
whose morphological identification is debated (Izeta, 2007;
L’Heureux, 2008).
The database discussed here was not generated in relation to the
specific goal addressed in this paper, being the product of diverse
interests usually operating in regional or local spatial scales. This
introduces sampling limitations in our analysis, given that there are
large non-sampled areas. Despite these limitations, our research
contributes to identify interesting regional and supra-regional
patterns, suggesting a number of explanations for them, which will
be evaluated on the basis of future oriented sampling. We present
a review of information on d13Ccollagen, d13Capatite, and d15N,
although the discussion is based only on the d13Ccollagen values,
which provide information that is adequate for our goals (Table 2).
Of the 90 d13Ccollagen samples, 72 are the product of 14C AMS
datings. These kinds of values have already been used for conti-
nent-wide isotopic discussions (e.g. van Klinken et al., 2000).
d13Ccollagen on herbivores is conditioned by the isotopic values of
the vegetal species regularly consumed. The existence of two main
photosynthetic pathways that predominate on contrasting climatic
and ecologic conditions provides the opportunity of evaluating
latitudinal and altitudinal isotopic tendencies (Iacumin et al.,
2000). Species with the C4 photosynthetic pathway display a global
d13C average value between 13 and 12& (Ehleringer and Cerling,
2001), being adapted to arid and hot climatic conditions. On the
other hand, species showing the C3 pathway have average d13C
values between 27 and 26&, and display lower adaptive effi-
ciency in settings characterized by high temperatures and water
restrictions (Tieszen, 1991; Ehleringer and Cerling, 2001). Forested
ecosystems are composed of C3 species and may present a distinc-
tive isotopic signature, given that they may be characterized by the
recycling of inert carbon that produces impoverished isotopic
values, which are transmitted through the successive steps in the
trophic chains (van der Merwe and Medina, 1991; Heaton, 1999).
3. Results
The information presented is already published in a large
number of contributions (see references in Table 1); therefore, our
main goal is to integrate these results within a specific frame of
research. Different projects integrated by the authors have
produced isotopic data for 67 guanaco samples; besides, there are
25 samples that were generated in the context of other projects.
Globally, we present isotopic values for 91 guanaco bone samples
from six different regions: central-southern Mendoza province
(n ¼ 12), the Humid Pampas (n ¼ 26), central-northern Patagonia
(n ¼ 13), southern Patagonia, steppe (n ¼ 25), southern Patagonia,
forest (n ¼ 11), and Tierra del Fuego Island (n ¼ 4) (Fig. 1, Table 1).
This information includes d13Ccollagen results for 90 samples and 14C
dates for 74 samples. Therefore, we can proceed to an evaluation of
preliminary temporal and spatial tendencies on different scales of
analysis. We present the complete stable isotope and radiocarbon
database in Table 1 and a descriptive statistics synthesis of this
information in Table 2.
As already suggested, we focus on the d13Ccollagen information
since it provides a relatively large sample size (n ¼ 90). The
descriptive statistics shown on Table 2 indicate a very wide isotopic
range (of ca. 11&) for this species on the macro-regional scale. This
range extends from 25.3& for a Pampean sample to 14.7& for
a southern Mendoza sample, presenting an average value of
20.4&. This large amount of variation for this herbivore species
constitutes an interesting pattern in itself, inviting to evaluate
which are the temporal and spatial factors that determine it.
Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
3
4. Temporal tendencies
There is a subtotal of 74 samples with available information on
d13Ccollagen and radiocarbon dates (in all cases these two determi-
nations were obtained from the same specimen). There is a very
weak negative correlation (n ¼ 74, r ¼ 0.18, p 0.10; Fig. 3) between
these variables that suggests the absence of temporal tendencies in
guanaco d13Ccollagen values at the macro-regional scale. If we limit
this analysis to the last 4500 14C years, which provide an 89.2% of
this subsample, we get a total lack of correlation (n ¼ 66, r ¼ 0.07,
p 0.53). The absence of marked temporal trends is not unexpected if
we take into account the large spatial scale considered, which
averages quite different climatic and ecological settings. Therefore,
we proceed to analyze values on smaller regional scales, sacrificing
sample size but gaining better control on the ecological homoge-
neity of the spatial units of analysis.
The samples from Central-northern Patagonia, located between
40 and 39 of southern latitude, show a positive and significant
correlation between chronology and d13Ccollagen values (n ¼ 11,
r ¼ 0.61, p 0.04; see contextual information in Martı́nez et al., 2009).
This region is currently located at a faunal and fitogeographical
ecotone between the Monte and Espinal Provinces (Abraham et al.,
2009; see Fig. 1). Palynological, geomorphological, and pedological
information indicates the occurrence of important climatic changes
during the Late Holocene (Schäbitz, 1994), which may have affected
the composition of vegetal communities modifying the relative
abundance of C4 and C3 species. During these times, this region
underwent climatic shifts from arid to semiarid conditions,
accompanied with an increment in precipitations and lake expan-
sions (Schäbitz, 2003), perhaps favoring C3 species. This hypothetic
situation may explain the tendency towards more depleted
guanaco values recorded for the last 2000 years. This preliminary
inference needs further data in order to be tested, but provides
a basis for introducing a temporal dimension in northern Patago-
nian isotopic ecology studies. On the other hand, information
available for different localities of Santa Cruz province (Fig. 1, area
4) in southern Patagonia does not show any correlations between
chronology and d13Ccollagen values (Borrero et al., 2009; Tessone and
Belardi, 2009).
5. Spatial tendencies
The regions considered present different ranges of isotopic
variation that may be partly explained by sample sizes (Table 3).
Nevertheless, there is a low positive correlation between sample
size and variance for these regions -which is clearly non-signifi-
cant-: r ¼ 0.41, p 0.412. This value does not indicate an important
incidence of sample sizes on the isotope diversity recorded for each
region, although it neither allows ruling this possibility out.
In Fig. 4 we show the ranges of variation for the regions
considered. For reasons that are discussed and justified below, in
the case of southern Santa Cruz we treat the samples that come
from the Andean forests and the eastern steppes separately. The
results presented below operate on two spatial scales of analysis:
first, a macro-regional scale, where the main issue to be considered
is the role of latitude in structuring guanaco isotopic values;
second, a regional scale, which allows us to enquire on the influence
of different ecologic and topographic contexts in the stable isotope
results.
5.1. Macro-regional scale
The macro-regional scale of analysis covers a latitudinal range of
20 , from the southern end of Tierra del Fuego Island, at 54 S, to
central Mendoza Province, at 34 S (Fig. 1). This wide geographical
 ARTICLE IN PRESS

4 R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10

Table 1
Isotopic and radiocarbon results for guanaco samples from southern South America.
14
# Site Altitude Latitude Longitude Lab code C date d13Ccol d13Cap d15N References
(masl) (South) (West)
1. Central-southern Mendoza (N ¼ 12) (*)
1 Arroyo El Desecho 10 2000 35 110 70 30 USF 5905 – 19.1 10.7 4.3 Gil et al., 2006
2 Agua de los Caballos 1025 35 220 68 180 USF 5906 – 14.7 5.0 Gil et al., 2006
3 Cueva de Luna 1300 36 050 60 430 USF 5907 – 19.4 11.1 4.6 Gil et al., 2006
4 El Indı́geno 3600 34 300 69 590 USF 6173 – 9.1 Gil et al., 2006
5 La Gotera 1500 35 520 69 570 USF 8354 – 18.7 6.2 Gil et al., 2006
6 Arroyo Malo 3 2000 34 520 69 540 USF 8355 – 18.8 4.8 Gil et al., 2006
7 Ojo de Agua 1600 35 90 69 380 USF 8356 – 18.7 6.6 Gil et al., 2006
8 El Sosneado 3 2050 34 500 69 540 USF 8357 – 18.9 6.1 Gil et al., 2006
9 Arroyo El Desecho 10 2000 35 110 70 30 USF 5913 – 18.8 -8.9 4.3 Gil et al., 2006
10 La Corredera 1150 36 310 68 320 USF 8864 – 19.3 6.3 Gil et al., 2006
11 Agua de los Caballos 1 1025 35 220 68 180 USF 8865 – 18.5 7.6 Gil et al., 2006
12 Gruta El Manzano 1300 36 040 69 430 USF 8866 – 17.6 7.6 Gil et al., 2006

2. Humid Pampas (N ¼ 26)

13 Zanjón Seco 2 100 38 100 59 100 CAMS 4993 3070  40 19.8 5.6 Politis et al., 2001
14 Zanjón Seco 2 100 38 100 59 100 CAMS 4994 3080  40 19.5 6.3 Politis et al., 2001
15 Calera 200 36 580 60 140 AA 67732 3008  46 18.5 Politis et al., 2005
16 Calera 200 36 580 60 140 AA 67733 2075  44 18.8 Politis et al., 2005
17 Calera 200 36 580 60 140 AA 67735 1748  42 19.2 Politis et al., 2005
18 Calera 200 36 580 60 140 AA 64617 2232  55 20.8 Politis et al., 2005
19 Calera 200 36 580 60 140 AA 71669 3390  170 22.6 Politis et al., 2005
20 Calera 200 36 580 60 140 AA 71671 3005  66 19.1 Politis et al., 2005
21 Cortaderas 90 38 200 59 390 AA 67736 2270  190 23.6 Massigoge, 2007
22 Nutria Mansa 1 <20 38 240 58 150 AA 55114 2705  66 25.3 Bonomo, 2005
23 Nutria Mansa 1 <20 38 240 58 150 AA 55115 3080  110 25.0 9.0 Bonomo, 2005
24 Nutria Mansa 1 <20 38 240 58 150 AA 55116 2920  110 24.6 Bonomo, 2005
25 Nutria Mansa 1 <20 38 240 58 150 USF 8751 – 25.0 12.2 Bonomo, pers. comm.
26 Arroyo Seco 2 110 38 210 60 140 AA 24052 7540  80 19.8 Steele and Politis, 2009
27 Arroyo Seco 2 110 38 210 60 140 AA 52613 8390  410 23.3 Steele and Politis, 2009
28 Quequén Salado 1 <20 38 490 60 320 Beta 360  40 18.6 Madrid et al., 2002
169820
29 Quequén Salado 1 <20 38 490 60 320 Beta 790  40 19.0 Madrid et al., 2002
157398
30 Quequén Salado 1 <20 38 490 60 320 Beta 940  40 19.1 Madrid et al., 2002
157397
31 Quequén Salado 1 <20 38 490 60 320 Beta 960  40 18.9 Madrid et al., 2002
169821
 0  0
32 Quequén Salado 2 <20 38 49 60 33 Beta 1720  40 19.3 Madrid et al., 2002
169822
 0  0
33 Tres Reyes 1 180 37 56 60 34 AA 7070 1845  50 20.4 Madrid and Barrientos, 2000
34 Tres Reyes 1 180 37 560 60 340 AA 7971 2235  50 19.9 Madrid and Barrientos, 2000
35 La Barrancosa 180 37 560 60 080 AA 59507 1676  46 20.2 Messineo, 2008
36 San Clemente VI <20 35 140 57 160 AA 29412 935  55 16.4 Paleo and Pérez Meroni, pers. comm.
37 Paso Otero 1 100 38 120 59 70 AA 72844 3056  42 19.0 Martı́nez, 2006
38 La Olla 4 <20 38 590 61 21́ AA-80664 6960  71 19.9 Politis and Bayón, pers. comm.

3. Central-northern Patagonia (N ¼ 13)

39 El Tigre- FCS.ET1.E <20 39 460 62 220 Ua 22561 455  45 19.9 Martı́nez, 2008
40 El Tigre-FCS.ET.C20 <20 39 460 62 220 AA 81830 473  43 20.1 Martı́nez et al., 2009
41 El Tigre-FCS.ET.C19 <20 39 460 62 220 AA 81834 536  43 20.2 Martı́nez et al., 2009
42 Loma Ruiz 1- LR1/1 <20 39 130 62 380 AA 53331 1615  50 17.8 Martı́nez, 2008
43 Loma Ruiz 1- LR1/2 <20 39 130 62 380 AA 53332 1935  44 16.2 Martı́nez, 2008
44 San Antonio 1-FCS <20 39 390 62 090 AA 81832 773  44 19.9 Martı́nez et al., 2009
45 San Antonio 2 S 1.1 <20 39 390 62 090 AA 77966 764  45 20.9 Martı́nez, 2008
46 San Antonio 2-FCS <20 39 390 62 090 AA 81831 988  44 19.7 Martı́nez et al., 2009
47 Angostura 1 70 40 100 64 110 AA 2551 938  45 23.9 Prates, 2008
48 Negro Muerto 60 39 500 65 170 AA 62794 398  46 19.5 Prates, 2008
49 Negro Muerto 60 39 500 65 170 AA 62795 483  43 22.8 Prates, 2008
50 Costa de Chubut <20 42 USF – 21.3 8.2 Gómez Otero, 2007
51 Chubut near coast <20 42 USF – 21.4 5.2 Gómez Otero, 2007

4. Southern Patagonia, steppe (N ¼ 25)

52 La Siberia 2 300 48 540 71 150 UGA 10014 1710  40 18.6 Tessone et al., 2005
53 La Siberia 2 300 48 540 71 150 UGA 10013 1100  40 19.0 Tessone et al., 2005
54 Alero del León 300 48 540 71 150 UGA 10009 2190  50 19.4 Tessone et al., 2005
55 Parapeto 4 300 48 540 71 150 UGA 08705 2010  50 19.4 Tessone et al., 2005
56 Alero Gerası́n II 300 48 540 71 150 UGA 10017 1580  40 19.4 Tessone et al., 2005
57 Cerro Pampa 2 300 48 540 71 150 UGA 10019 310  40 19.7 Tessone et al., 2005
58 Solı́s 300 48 540 71 150 UGA 10018 1140  40 19.8 Tessone et al., 2005
59 Cerro Pampa 2 300 48 540 71 150 UGA 10020 170  40 20.6 Tessone et al., 2005
60 El Sosiego 2 200 50 90 72 350 GX 25278 1920  40 20.5 Carballo Marina et al., 1999
61 A. Piedra Quemada 500 50 30 72 170 GX 26196 650  40 20.8 Carballo Marina et al., 1999

Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
 ARTICLE IN PRESS

R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10 5

Table 1 (continued )
14
# Site Altitude Latitude Longitude Lab code C date d13Ccol d13Cap d15N References
(masl) (South) (West)
62 V. Piedra Quemada 2 500 50 30 72 170 GX 25775 520  40 20.3 Carballo Marina et al., 1999
63 Cabo Vı́rgenes 7 40 52 190 68 210 GX 25773 160  40 20.1 Borrero et al., 2006
64 Cabo Vı́rgenes 8 < 20 52 200 68 230 GX 27868 240  40 21.0 Borrero et al., 2006
65 Cabo Vı́rgenes 4 < 20 52 200 68 230 GX 27864 2000  40 21.0 Borrero et al., 2006
66 Cabo Vı́rgenes peat < 20 52 200 68 230 GX 27865 1510  30 22.3 Borrero et al., 2009
67 Cerro León 1, 1 400 50 510 72 120 GX 27863 4340  40 20.8 Borrero et al., 2009
68 Cerro León 1, 2 400 50 510 72 120 GX 27866 2850  40 19.8 Borrero et al., 2009
69 Orejas de Burro 1, II 145 52 070 69 330 Ua 21902 3490  50 19.8 Borrero et al., 2009
70 Cóndor 1, 4E, II 150 51 890 69 360 Ua 24658 965  40 20.3 Borrero et al., 2009
71 Cóndor 1, 4E, III 150 51 890 69 360 Ua 24658 1360  65 19.5 Borrero et al., 2009
72 La Carlota 70 50 510 72 140 Beta 1070  40 20.1 Campan et al., 2007
215184
73 Cabo Vı́rgenes 22 40 52 190 68 220 GX 32586 660  50 21.3 Borrero et al., 2009
74 Cerro León 3 400 50 360 72 160 GX 32583 4370  50 19.7 Borrero et al., 2009
75 Cabo Vı́rgenes <20 52 200 68 220 USF 582 – 21.0 2.2 Barberena, 2002
76 Cerro Verlika 1 1100 50 360 72 160 GX 25277 1685  70 21.3 Franco et al., 1999

5. Southern Patagonia, forest (N ¼ 11)

77 Alero del Bosque 225 50 200 72 310 Beta 91301 3110  50 22.0 Franco et al., 1999
78 Lago Roca 3 250 50 200 72 310 Beta 91302 170  30 24.0 Franco et al., 1999
79 Alice 1, 1 200 50 200 72 310 Beta 1420  70 22.0 Borrero et al., 19981999
112231
80 Alice 1, 2 200 50 300 72 400 Beta 1480  70 24.9 Borrero et al., 19981999
112232
81 Alice 2 200 50 300 72 400 GX 27174 740  60 20.4 Borrero et al., 19981999
82 Chorrillo Malo 2, 1 200 50 90 72 360 Beta 3790  80 19.0 Franco and Borrero, 2003
148743
83 Chorrillo Malo 2, 4 200 50 90 72 360 GX 25279 9740  50 22.1 Franco and Borrero, 2003
84 Rincón Amigo 300 50 30 72 170 Beta 1840  40 24.6 Carballo Marina et al., 1999
138991
85 CCP 7, Capa 5 N8A 900 47 570 72 050 UGA 866 5400  64 20.7 Aschero et al., 2007
86 CCP7, 17 (2) C11B 900 47 570 72 050 UGA 870 8362  68 20.8 Aschero et al., 2007
87 CCP7, 19 D9C 900 47 570 72 050 UGA 873 10690  72 23.0 Aschero et al., 2007

6. Tierra del Fuego Island (N ¼ 4)

88 Túnel I <20 54 490 68 090 AC 703 5280  100 20.5 Alberó et al., 1986
89 Túnel I <20 54 490 68 090 AC 704 3190  100 21.8 Orquera and Piana, 1996
90 San Genaro 2 <20 53 190 68 170 USF386/243 – 21.4 14.2 4.9 Guichón et al., 2001
91 Las Mandı́bulas 3 <20 53 120 68 380 Beta 1190  50 20.9 Favier Dubois and Borrero, 2005
122882

Note: (*) contextual information indicates that the Central-southern Mendoza samples are Holocene in age.

range provides a great window to evaluate the incidence of latitude
in relation with isotopic variation. Since global climate presents
a marked latitudinal configuration (Strahler, 1982), and considering
that climate is one of the main conditioners of the abundance of C3
and C4 vegetal species (Ehleringer and Cerling, 2001), this isotopic
record is adequate to evaluate macro-regional variations in
guanaco isotopic ecology.
An analysis of correlation between latitude and d13Ccollagen
values produces a result of r ¼ 0.28 (p < 0.01), indicative of
a negative and very weak correlation between these variables at the
large macro-regional scale. Although it is expected that this
tendency will experiment modifications with increasing amounts
of isotopic data, we suggest that this result is not the product of
biases due to small sample sizes. In order to support this statement,
we explore climatic and ecologic variables acting on smaller scales,
which are averaged at the macro-regional analysis producing the
emergent lack of correlation observed between latitude and
isotopic values.
Isotopic results for guanaco are distributed in two large lat-
itudinal subsets: the southern set corresponds to the regions of
Tierra del Fuego and southern Patagonia, located between 54 and

Table 2
Synthesis of isotopic data for guanaco samples from southern South America.

Region Samples d13Ccollagen d13Capatite d15N 14

C
Dates
Central-southern 12 11 4 11 –
Mendoza
Humid Pampas 26 26 2 2 25
Central-northern 13 13 – 2 11
Patagonia
Southern Patagonia, 25 25 – 1 24
steppe
Southern Patagonia, 11 11 – – 11
forest
Tierra del Fuego 4 4 1 1 3
Total 91 (100%) 90 (98.9%) 7 (7.7%) 17 (18.7%) 74 (81.3%)
Fig. 3. Temporal tendencies in d13Ccollagen values for Holocene guanaco samples.

Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
 ARTICLE IN PRESS

6 R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10

Fig. 4. Ranges of variation in d13Ccollagen values on guanaco samples.

48 S; the northern set includes Central-northern Patagonia,
Central-southern Mendoza, and the Humid Pampas, being
emplaced between 42 and 34 S. There is an intermediate eight
degrees wide area that is void of information, between 47 and
42 S, clearly limiting our ability to evaluate latitudinal variations in
a continuous manner. From now we treat these two sets of isotopic
data separately.
When evaluating the correlation between latitude and d13Ccollagen
values within each of the two spatial sets we obtain the following
results: the southern set produces an r ¼ 0.13 (p 0.41), whereas the
northern set produces an r ¼ 0.37 (p < 0.01). Both of these values
are negative and very low indicating, as already seen at the macro-
regional scale, the absence of a strong correlation between these
variables. Nevertheless, given that both sets of samples average quite
different ecological contexts, we consider it is necessary to proceed
one step further by subdividing these sets in terms of general
ecological or climatic information. Therefore, we isolate assemblages
from areas that are more ecologically homogeneous (fitogeographic
Provinces in Fig. 1; Cabrera, 1976).
In the case of the southern group we segregate the samples
coming from the western forest environments and from the eastern
steppes, given that these ecosystems may show isotopic variation
that is to a large extent independent of latitude (van der Merwe and
Medina, 1991). In the northern group we separate the samples from
the Humid Pampas from those from Central-northern Patagonia
and Central-southern Mendoza, which are located within the
climatic-ecologic zone defined as the ‘South American Arid Diag-
onal’ (Bruniard, 1982). Although these decisions are based on the
present configuration of these ecosystems, there is important
paleoecological data that indicates their temporal depth, lending
support to their use for organizing Late Holocene samples. In the
case of the southern area, paleoecological data indicates that
changes in forest extension do not affect the suggested segregation
of the samples (Huber et al., 2004; Mancini et al., 2008). In the case
of the northern group, on the other hand, diverse lines of paleo-
climatic evidence indicate that the Arid Diagonal experimented
changes in its specific configuration (Mancini et al., 2005), but these
changes do not affect the separation suggested for Late Holocene
samples (Fig. 1; Zárate, 2002; Labraga and Villalba, 2009).
We suggest that this segmentation of the southern and northern
sets of samples allows isolating the incidence of latitude in the
isotopic data, shown as unimportant so far, by removing some local
ecologic and climatic variables from the analysis. Next we explore
the correlation between latitude and d13Ccollagen values for both sets

Fig. 5. d13Ccollagen values and latitude. Note: the r values do not include the samples from the southern Patagonian forest and the Humid Pampas.

Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
 ARTICLE IN PRESS
R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10
Fig. 6. Correlations between altitude and d13Ccollagen and d15N values in Central-
southern Mendoza.
considering only the samples from the steppes, in the southern set,
and those located in the Arid Diagonal, in the northern set. There-
fore, we are respectively excluding the cases from the southern
Patagonian forests and the northern Humid Pampas (Fig. 5).
The correlation results are different to those obtained previ-
ously, since the value for the southern set is of r ¼ 0.68 (p < 0.005)
and the value for the northern set is of r ¼ 0.57 (p < 0.005).
Comparing these values with those obtained for the global
southern (r ¼ 0.13, p 0.41) and northern (r ¼ 0.37, p < 0.01) sets,
we can see that there is a marked change, with correlations being
more negative and statistically significant in the present analysis.
This change appears more strongly in the southern set (Fig. 5).
Therefore, we suggest that at this regional scale of analysis, and
discriminating the samples in terms of their ecological context, we
identify an important incidence of latitude on the guanaco d13Ccollagen
values. This indicates a situation different to those recorded at both
the macro-regional scale and at the ecologically undifferentiated
regional analysis.
There is another interesting spatial pattern in Fig. 5, consisting
in the gradual enrichment of isotopic values recorded indepen-
dently in the southern and northern geographical sets. A t test of
these two groups of samples produces a result of t ¼ 1.731 (p 0.08),
suggesting that they are not statistically different despite the large
geographical gap that separates them. As already suggested, we
consider that this global similitude emphasizes that latitude is not
the only -neither the main-factor operating at the large macro-
regional scale.
5.2. Regional scale
The results presented above indicate that there is not a latitudinal
tendency in the guanaco isotopic data at the macro-regional scale. On
the other hand, there is a negative and relatively important
Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
7
correlation between latitude and d13Ccollagen values identifiable at
smaller regional scales, if a consideration of ecological context is
introduced in the analysis. Next we develop regional cases that
allow exploring the role played by selected ecological and topo-
graphical variables. These cases will be used to illustrate the
potential of isotopic information as a geographic tracer of guanaco
ranges in the past.
In southern Patagonia we have identified an isotopic signature of
guanacos inhabiting forest ecosystems (for the regions of lago
Argentino and Parque Nacional Perito Moreno, in Santa Cruz
province; it must be mentioned that although the two samples
from Túnel site in Tierra del Fuego also come from a closed forested
setting, they do not show a ‘canopy signature’). These samples
present depleted d13Ccollagen values, which may be accounted for by
the canopy effect characteristic of closed forested systems with
a slow recycling of available carbon (van der Merwe and Medina,
1991). Ecologic information attributes the behavior of inhabiting
forested settings only to guanacos from Tierra del Fuego Island
(Raedeke, 1978), characterized by a largely forested southern
region, and therefore this information extends this pattern to the
southern end of the continent. This isotopic signature provides
interesting geographic information on guanaco movements in the
past, providing a new proxy of its persistent use of forest ecosys-
tems in the continent. On the other hand, this pattern also provides
an important factor to be considered in human paleodietary
reconstructions, most importantly for the identification of the
consumption of marine resources, which is a key topic in Patago-
nian archaeology (Barberena, 2002; Gómez Otero, 2007; Borrero
et al., 2009). The existence of a forest signature in some guanacos
may suggest the need to adjust the terrestrial end-line used for the
interpretation of human samples on smaller spatial scales.
This situation, which is suggested here for southern regions
located at the eastern flank of the Andes, may be even more
important for the Pacific coastlines where marine foods were
usually the main staple, and for northern Patagonia, with wider
forest ecosystems and an apparent more fluid connectivity between
eastern and western flanks of the Andes (facilitated by general
biogeographical conditions). This record will have to be reevaluated
in the future on the basis of a larger dataset, although it can be stated
that its implications for human paleodietary research will remain.
The southern Mendoza region provides information to evaluate
the relationships between isotopic values and topographic varia-
tion. The data available on the vegetal isotopic ecology shows that
there is an important increase in the frequency of C4 grasses in the
lower altitudinal eco-zones (at 1000/1500 masl), while they are
absent at higher altitudes of the Andes mountain range (Cavagnaro,
1988; Llano, 2009). There is a set of 11 samples from this region
(Table 1) that allows an initial assessment of the influence of alti-
tude on guanaco d13Ccollagen and d15N values. Of this subset, 10
samples come from settings between 1000 and 2000 masl, which
are separated by a minimum distance of ca. 50–100 km. In Fig. 6 we
present analyses of correlation for d13Ccollagen and d15N values with
altitude, indicating negative and weak correlations (Gil et al., 2006,
2009). Sample size is small and spatially heterogeneous for solving
this issue, although a number of expectations that will be tested in
the future can be developed on the basis of the available data. The
altitudinal range between 1000 and 2000 masl represented in
these samples corresponds to a transitional area between
C4-dominated herbaceous communities, in the lowlands, to
C3-dominated communities, in the highlands (Cavagnaro, 1988;
Llano, 2009). Despite this situation, there are no spatial patterns in
the distribution of isotopic values with the exception of one
enriched sample from a low altitude setting that fits with the
expected pattern. Interestingly, if we exclude this outlier from
 ARTICLE IN PRESS
8 R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10
Table 3
Descriptive statistics for d13Ccollagen on guanaco samples.
Region Samples Average
Central-southern Mendoza 11 18.7
Humid Pampas 26 20.6
Central-northern Patagonia 13 20.3
Southern Patagonia, steppe 25 20.2
Southern Patagonia, forest 11 22.1
Tierra del Fuego 4 21.2
Total 90 20.4
the analysis, the r value drops to 0.16 (p 0.64), suggesting that
altitude is not structuring the isotopic values in this sample.
There are two main factors that need to be considered in order
to develop this issue: the size and spatial configuration of guanaco
home ranges -that may integrate close isotopically distinct altitu-
dinal floors-, and guanaco foraging patterns and grass selectivity.
Available information already mentioned indicates that guanacos
have an important foraging flexibility, allowing them to forage C4
grasses with high fiber content (Puig et al., 1996; Llano, 2009). This
information suggests that guanacos would not strongly select
against C4 species, something that can be independently supported
on the basis of the latitudinal variations documented here. We use
this preliminary information to suggest a hypothesis to be tested in
the future: Late Holocene guanacos from southern Mendoza
occupied and foraged on different altitudinal floors located
between 1000 and 2000 masl. This would have the consequence of
producing an averaged isotopic signal that fails to show strong
altitudinal variations. When associated to variations in vegetal
isotopic ecology, isotopic information has a great potential to
provide information on the altitudinal distribution of South
American camelids (Fernández and Panarello, 1999–2001; Yaco-
baccio et al., 2009). Given that human consumption of isotopically
enriched guanaco meat may mimic a low-level consumption of C4
domesticated species, like Zea mays (see Tykot et al., 2009), this
issue is relevant for the isotopic evaluation of maize consumption
by humans, which is a key topic for the archaeology of the South
Central Andes (Gil et al., 2006).
Guanaco samples from the Humid Pampas and Central-
northern Patagonia present widest levels of isotopic variation as
indicated by their variance values (5.72& and 3.86& respectively;
see Table 3, Fig. 3). On the basis of the available information, we
do not have any convincing explanations for these patterns,
although it can be suggested that it is related with the ecological
complexity of these regions. In the Humid Pampas, guanacos may
have locally foraged on a number of different herbaceous
communities (Cabrera, 1976; Prieto, 1996), whose isotopic signa-
tures are not precisely known yet, which have the potential to
produce the isotopic divergence recorded. Central-northern
Patagonia, on the other hand, constitutes a faunistic and
fitogeographic ecotonal zone connecting the Monte and Espinal
provinces (Abraham et al., 2009). Besides, part of the isotopic
variation observed can be explained on the basis of the temporal
changes that we have recorded which can be associated with
climatic variations towards enhanced arid conditions. The four
regional cases briefly presented here highlight the importance of
differing levels of regional ecological complexity in structuring
guanaco isotopic variation, dictating the appropriate spatial scale
to conduct isotopic ecology studies.
6. Conclusions
The review of available information presented here allowed us
to identify a number of new and interesting tendencies on guanaco
d13C data. We have not recognized any temporal trend, the only
Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
Median Standard deviation Variance Maximum value Minimum value
18.8 1.4 2.1 14.7 19.8
19.8 2.4 5.8 16.4 25.3
20.1 2.0 3.9 16.2 23.9
20.1 0.8 0.71 18.6 22.3
22 1.8 3.44 19 24.9
21.2 0.6 0.3 20.5 21.8
19.9 2 3.8 14.7 25.3
exception being the regional case of Central-northern Patagonia. In
terms of spatial analysis, we have identified operating on different
spatial scales. At the largest level of analysis, corresponding to the
macro-regional scale, we did not identify strong correlations
between latitude and d13Ccollagen isotopic values; although there is
a gradual tendency towards enriched average values at lower lati-
tudes, this tendency is neither strong nor lineal (Fig. 5). On the
contrary, by reducing the scale of analysis and introducing
a discrimination of the samples in terms of their ecological context,
we do record a strong negative correlation between latitude and
d13Ccollagen values. At this smaller scale we are able to isolate the
incidence of latitude, as a direct conditioning of global climate, on
guanaco isotopic values. As already suggest, when we shift to larger
geographical scales we introduce another independent variables in
the analysis, such as ecological context, topography, and altitude,
producing a wider dispersion of isotopic values and reducing the
correlation with latitude per se.
The discussions developed on regional scales provide an example
of the role of stable isotopes as a geographic tracer of guanaco home
ranges, as exemplified with the case of a forest signal in southern
Patagonian samples. This information is potentially useful for
species conservation and protected areas management (Etnier,
2004) by providing a proxy of the past spatial distribution of
guanacos. As suggested by González et al. (2006: 172), the integra-
tion of data on past distributional ranges, population dynamics,
foraging behavior and genetic distinctiveness (Kadwell et al., 2001)
will help to define conservation units. Finally, by characterizing the
inter-regional isotopic variability on guanaco, which was a main
staple for Holocene hunter-gatherers, this information will have
important implications for comparative paleodietary research on
human samples by means of stable isotopes.
Acknowledgements
We would like to acknowledge Augusto Tessone for his thorough
reading of the paper, which helped to clarify our arguments and
provided valuable ideas on how to present them. To Luciano Prates
for the important discussions maintained and for providing infor-
mation, as well as to Cristina Bayón, Mariano Bonomo, Agustina
Massigoge, Pablo Messineo, M. Clara Paleo and Mercedes Pérez
Meroni for allowing us to use their unpublished results; to Teresa
Civalero, Mariana De Nigris, Hugo Yacobaccio and Juan Bautista
Belardi for their comments. The analyses developed in our projects
were funded by the following institutions: Consejo Nacional de
Investigaciones Cientı́ficas y Técnicas, Universidad de Buenos Aires,
Agencia Nacional de Promoción de la Ciencia y la Tecnologı́a,
National Geographic Society, Universidad Nacional del Centro de la
Provincia de Buenos Aires (INCUAPA), and Museo de Historia
Natural de San Rafael.
References
Abraham, E., del Valle, H., Roig, F., Torres, L., Ares, J., Coronato, F., Godagnone, R.,
2009. Overview of the geography of the Monte Desert biome (Argentina).
Journal of Arid Environments 73 (2), 144–153.
 ARTICLE IN PRESS
R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10
Alberó, M., Angiolini, F., Piana, E., 1986. Discordant ages related to reservoir effect of
associated archaeological remains from the Túnel Site, Beagle Channel,
Argentine Republic. Radiocarbon 28 (2A), 748–753.
Aschero, C., Bozzuto, D., Civalero, T., De Nigris, M., Di Vruno, A., Dolce, V., Fernández, N.,
González, L., Sacchi, M., 2007. Nuevas evidencias sobre las ocupaciones tempranas
en Cerro Casa de Piedra 7. In: Morello, F., Martinic, M., Prieto, A., Bahamonde, G.
(Eds.), Arqueologı́a de Fuego-Patagonia. Levantando piedras, desenterrando hue-
sos. y develando arcanos. CEQUA, Punta Arenas, pp. 569–576.
Barberena, R., 2002. Los lı́mites del mar. Isótopos estables en Patagonia meridional.
Sociedad Argentina de Antropologı́a, Buenos Aires.
Boelcke, O., Moore, D., Roig, F., 1985. Transecta Botánica de la Patagonia Austral.
CONICET-Instituto de la Patagonia-Royal Society, Buenos Aires.
Bonino, N., Pelliza Sbriller, A., 1991. Composición botánica de la dieta del guanaco
(Lama guanicoe) en dos ambientes contrastantes de Tierra del Fuego, Argentina.
Ecologı́a Austral 1 (2), 97–102.
Bonomo, M., 2005. Costeando las llanuras. Arqueologı́a del litoral marı́timo pam-
peano. Sociedad Argentina de Antropologı́a, Buenos Aires.
Borrero, L., 1990. Fuego-Patagonian bone assemblages and the problem of
communal guanaco hunting. In: Davis, L.B., Reeves, B.O.K. (Eds.), Hunters of the
Recent Past. Unwin Hyman, London, pp. 373–399.
Borrero, L., Barberena, R., Franco, N., Charlin, J., Tykot, R., 2009. Isotopes and rocks:
geographic organization of Patagonian hunter-gatherers. International Journal
of Osteoarchaeology 19 (2), 309–327.
Borrero, L., Franco, N., Barberena, R., Borella, F., Campan, P., Carballo Marina, F.,
Cruz, I., Favier Dubois, C., Guichón, R., L’Heureux, L., Mancini, V., Manzi, L.,
Martin, F., 2006. Arqueologı́a de Cabo Vı́rgenes y Cañadón Gap. In: Cruz, I.,
Caracotche, M.S. (Eds.), Arqueologı́a de la Costa Patagónica. Perspectivas para su
Conservación. UNPA, Rı́o Gallegos, pp. 212–228.
Borrero, L., Franco, N., Carballo Marina, F., Martin, F., 1998–1999. Arqueologı́a de
Estancia Alice, Lago Argentino. Cuadernos del INAPL 18, 31–48.
Bruniard, E., 1982. La diagonal árida argentina. Un lı́mite climático real. Revista
Geográfica IPGH 95, 5–20.
Burton, R., Snodgrass, J., Gifford-González, D., Guilderson, T., Brown, T., Koch, P.,
2001. Holocene changes in the ecology of northern fur seals: insights from
stable isotopes and archaeofauna. Oecologia 128, 107–115.
Cabrera, A., 1976. Regiones Fitogeográficas Argentinas. Enciclopedia Argentina de
Agricultura y Jardinerı́a II (1). Acme, Buenos Aires.
Cajal, J., 1980. Situación del guanaco y estrategia de la conservación de los camélidos
en la República Argentina. Subsecretarı́a de Ciencia y Tecnologı́a, Buenos Aires.
Campan, P., Carballo Marina, F., Manzi, L., 2007. Arqueologı́a de Estancia La Carlota
(Campo Volcánico Pali Aike, Argentina). In: Morello, F., Prieto, A., Martinic,
Bahamonde, G. (Eds.), Arqueologı́a de Fuego-Patagonia. Levantando piedras,
desenterrando huesos . y develando arcanos. CEQUA, Punta Arenas, pp. 687–699.
Carballo Marina, F., Borrero, L., Franco, N., Belardi, J., Horwitz, V., Campan, P., Martin, F.,
Muñoz, A., Borella, F., Garcı́a, F., Lanata, J., 1999. Arqueologı́a de la costa del lago
Argentino, rı́o La Leona y pampas altas intermedias. Praehistoria 3, 13–33.
Cavagnaro, J., 1988. Distribution of C3 and C4 grasses at different altitudes in
a temperate arid region of Argentina. Oecologia 76, 273–277.
Ehleringer, J., Cerling, T., 2001. Photosynthetic pathways and climate. In:
Schulze, E.D., Heimann, M., Harrison, S., Holland, E., Lloyd, J., Prentice, I.C.,
Schimel, D.S. (Eds.), Global Biogeochemical Cycles in the Climate System.
Academic Press, New York, pp. 267–277.
Etnier, M., 2004. The potential of zooarchaeological data to guide pinniped
management decisions in the eastern North Pacific. In: Lyman, R.L., Cannon, K.P.
(Eds.), Zooarchaeology and Conservation Biology. University of Utah Press, Salt
Lake City, pp. 88–102.
Favier Dubois, C., Borrero, L., 2005. Playas de acreción: cronologı́a y procesos de
formación del registro arqueológico en la costa central de la bahı́a San Sebas-
tián, Tierra del Fuego (Argentina). Magallania 33 (2), 93–108.
Fernández, J., Panarello, H., 1999–2001. Isótopos del carbono en la dieta de herbı́-
voros y carnı́voros de los Andes jujeños. Xama 12–14, 71–85.
Franco, N., Borrero, L., 2003. Chorrillo Malo 2: initial peopling of the Upper Santa
Cruz Basin. In: Bonnichsen, R., Miotti, L., Salemme, M., Flegenheimer, N. (Eds.),
Where the South Winds Blow. Ancient Evidences of Paleo South Americans,
Center for the Studies of the First Americans. Texas A&M University Press, Texas,
pp. 149–152.
Franco, N., Borrero, L., Belardi, B., Carballo Marina, F., Martin, F., Campan, P., Favier
Dubois, C., Stadler, N., Hernández Llosas, I., Cepeda, H., Muñoz, S., Borella, F.,
Muñoz, F., Cruz, I., 1999. Arqueologı́a del cordón Baguales y sistema lacustre al sur
del Lago Argentino (Provincia de Santa Cruz, Argentina). Praehistoria 3, 65–86.
Gil, A., Neme, G., Tykot, R., Novellino, P., Cortegoso, V., Durán, V., 2009. Stable
isotopes and maize consumption in Central Western Argentina. International
Journal of Osteoarchaeology 19 (2), 215–236.
Gil, A., Tykot, R., Neme, G., Shelnut, N., 2006. Maize on the frontier. Isotopic and
macrobotanical data from Central-Western Argentina. In: Staller, J., Tykot, R.,
Benz, B. (Eds.), Histories of Maize. Multidisciplinary Approaches to the Prehis-
tory, Biogeography, Domestication, and Evolution of Maize. Academic Press,
New York, pp. 199–214.
Gómez Otero, J., 2007. Dieta, uso del espacio y evolución en poblaciones cazadoras-
recolectoras de la costa centro-septentrional de Patagonia durante el Holoceno
Medio y Tardı́o. Ph.D. Dissertation, Facultad de Filosofı́a y Letras, Universidad de
Buenos Aires, Argentina.
González, B., Palma, E., Zapata, B., Marı́n, J., 2006. Taxonomic and biogeographical
status of guanaco Lama guanicoe (Artyodactila, Camelidae). Mammal Review 36
(2), 157–178.
Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
9
Guichón, R., Borrero, L., Prieto, A., Cárdenas, P., Tykot, R., 2001. Nuevas determi-
naciones de isótopos estables para Tierra del Fuego. Revista Argentina de
Antropologı́a Biológica 3 (1), 113–126.
Gutiérrez, M., Martı́nez, G., 2008. Trends in the faunal human exploitation during
the Late Pleistocene and Early Holocene in the Pampean region (Argentina).
Quaternary International 191, 53–68.
Heaton, T., 1999. Spatial, species, and temporal variations in the 13C/12C ratios of c3
plants: implications for palaeodiet studies. Journal of Archaeological Science 26,
637–649.
Huber, U., Markgraf, V., Schäbitz, F., 2004. Geographical and temporal trends in late
quaternary fire histories of Fuego-Patagonia, South America. Quaternary
Science Reviews 23 (9–10), 1079–1097.
Iacumin, P., Nikolaev, V., Ramigni, M., 2000. C and N stable isotope measurements
on Eurasian fossil mammals, 40,000 to 10,000 years BP: herbivore physiologies
and palaeoenvironmental reconstruction. Palaeogeography, Palaeoclimatology,
Palaeoecology 163, 33–47.
Izeta, A.D., 2007. Zooarqueologı́a del sur de los valles Calchaquı́es (Provincias de
Catamarca y Tucumán, República Argentina). In: BAR International Series 1612.
Archaeopress, Oxford.
Kadwell, M., Fernández, M., Stanley, H., Baldi, R., Wheeler, J., Rosadio, R., Bruford, M.,
2001. Genetic analysis reveals the wild ancestors of the llama and the alpaca.
Proceedings of the Royal Society of London B 268, 2575–2584.
Labraga, J., Villalba, R., 2009. Climate in the Monte Desert: past trends, present
conditions, and future projections. Journal of Arid Environments 73 (2),
154–163.
L’Heureux, G., 2008. El estudio arqueológico del proceso coevolutivo entre las pobla-
ciones humanas y las poblaciones de guanaco en Patagonia meridional y norte de
Tierra del Fuego. In: Archaeopress, BAR International Series, vol. 1751 Oxford.
Llano, C., 2009. Photosynthetic pathways, spatial distribution, isotopic ecology, and
implications for pre-hispanic human diets in central-western Argentina.
International Journal of Osteoarchaeology 19 (2), 130–143.
Madrid, P., Barrientos, G., 2000. La estructura del registro arqueológico del sitio
Laguna Tres Reyes 1 (Provincia de Buenos Aires): nuevos datos para la inter-
pretación del poblamiento humano del Sudeste de la Región Pampeana a los
inicios del Holoceno Tardı́o. Relaciones de la Sociedad Argentina de Antropo-
logı́a XXV, 179–206.
Madrid, P., Politis, G., March, R., Bonomo, M., 2002. Arqueologı́a microrregional en el
sudeste de la Región Pampeana Argentina: el curso del rı́o Quequén Salado.
Relaciones de la Sociedad Argentina de Antropologı́a XXVII, 327–355.
Mancini, M., Páez, M., Prieto, A., Stutz, S., Tonello, M., Vilanova, I., 2005. Mid-
Holocene climatic variability reconstruction from pollen records (32 –52 S,
Argentina). Quaternary International 132 (1), 47–59.
Mancini, V., Prieto, A., Páez, M., Schäbitz, F., 2008. Late Quaternary vegetation and
climate of Patagonia. In: Rabassa, J. (Ed.), Developments in Quaternary Sciences,
vol. 11. Elsevier, Amsterdam, pp. 351–367.
Martı́nez, G., 2006. Arqueologı́a del curso medio del rı́o Quequén Grande: estado
actual y aportes a la arqueologı́a de la región pampeana. Relaciones de la
Sociedad Argentina de Antropologı́a XXXI, 249–275.
Martı́nez, G., 2008. Arqueologı́a del curso inferior del rı́o colorado: estado actual del
conocimiento e implicaciones para la dinámica poblacional de cazadores-rec-
olectores pampeano-patagonicos. In: Cazadores Recolectores del Cono Sur, 3.
Martı́nez, G., Zangrando, A., Prates, L., 2009. Isotopic ecology and human paleodiets
in the lower basin of the Colorado River, Buenos Aires Province, Argentina.
International Journal of Osteoarchaeology 19 (2), 281–296.
Massigoge, A., 2007. Procesos de formación del registro arqueológico en el sitio
Cortaderas (partido de San Cayetano, provincia de Buenos Aires). Intersecciones
en Antropologı́a 8, 197–214.
Mengoni Goñalons, G., 1999. Cazadores de guanacos de la estepa patagónica.
Sociedad Argentina de Antropologı́a, Buenos Aires.
Messineo, P., 2008. NonEnInvestigaciones arqueológicas en la cuenca superior del
arroyo Tapalqué (Partidos de Olavarrı́a y Benito Juárez, Provincia de Buenos
Aires). Ph.D. Dissertation. Facultad de Ciencias Naturales y Museo, Universidad
Nacional de La Plata, Argentina.
Morales, M., Barberena, R., Belardi, J., Borrero, L., Cortegoso, V., Durán, V., Guerci, A.,
Goñi, R., Gil, A., Neme, G., Yacobaccio, H., Zárate, M., 2009. Reviewing human-
environment interactions in arid regions of Southern South America during the
past 3000 years. Palaeogeography, Palaeoclimatology, Palaeoecology 272.
Neme, G., Gil, A., 2008. Faunal exploitation and agricultural transitions in the South
American agricultural limit. International Journal of Osteoarchaeology 18,
293–306.
Orquera, L., Piana, E., 1996. El sitio Shamakush I (Tierra del Fuego, República
Argentina). Relaciones de la Sociedad Argentina de Antropologı́a XXI, 215–265.
Ortega, I., Franklin, W.,1988. Feeding habitat utilization and preference by guanaco male
group in the Chilean Patagonia. Revista Chilena de Historia Natural 61, 209–216.
Politis, G., Martı́nez, G., Bonomo, M., 2001. Alfarerı́a temprana en sitios de caza-
dores-recolectores de la Región Pampeana (Argentina). Latin American Antiq-
uity 12 (2), 167–181.
Politis, G., Messineo, P., Kaufmann, C., Barros, M., Alvarez, M.C., Di Prado, V.,
Scalise, R., 2005. Persistencia ritual entre cazadores-recolectores de la llanura
pampeana. Boletı́n de Arqueologı́a PUCP 9, 67–90.
Politis, G., Pedrotta, V., 2006. Recursos faunı́sticos y estrategias de subsistencia en el
este de la región pampeana durante el Holoceno tardı́o: el caso del guanaco (Lama
guanicoe). Relaciones de la Sociedad Argentina de Antropologı́a XXXI, 301–336.
Prates, L., 2008. Los indı́genas del rı́o Negro. Un enfoque arqueológico. Sociedad
Argentina de Antropologı́a, Buenos Aires.
 ARTICLE IN PRESS
10 R. Barberena et al. / Journal of Archaeological Science xxx (2009) 1–10
Prieto, A., 1996. Late quaternary vegetational and climatic changes in the Pampa
grassland of Argentina. Quaternary Research 45, 73–88.
Puig, S., Videla, F., Monge, S., Roig, V., 1996. Seasonal variation in guanaco diet (Lama
guanicoe) and food availability in northern Patagonia, Argentina. Journal of Arid
Environments 34, 215–224.
Raedeke, K., 1978. El guanaco de Magallanes. Su distribución y biologı́a. Publicación
Técnica 4. Corporación Nacional Forestal, Santiago de Chile.
Schäbitz, F., 1994. Holocene climatic variations in Northern Patagonia,
Argentina. Palaeogeography, Palaeoclimatology, Palaeoecology 109 (2–4),
287–294.
Schäbitz, F., 2003. Estudios polı́nicos del Cuaternario en las regiones áridas del sur
de Argentina. Revista del Museo Argentino de Ciencias Naturales 5 (2),
291–299.
Steele, J., Politis, G., 2009. AMS 14C dating of early human occupation of southern
South America. Journal of Archaeological Science 36 (2), 419–429.
Strahler, A., 1982. Geografı́a Fı́sica. Omega Editorial, Barcelona.
Tessone, A., Belardi, J., 2009. Evaluación de variaciones temporales del d13C y d15N
en el colágeno de herbı́voros de los lagos Tar y San Martı́n (provincia de Santa
Cruz, Patagonia). In: Proceedings of the 1st Congreso Nacional de Zooarqueo-
logı́a Argentina.
Tessone, A., Zangrando, A., Barrientos, G., Valencio, S., Panarello, H., Goñi, R., 2005.
Isótopos estables del carbono en Patagonia meridional: datos de la cuenca del
Please cite this article in press as: Barberena, R., et al., Guanaco (Lama guanicoe) isotopic ecology in southern South America:..., J. Archaeol. Sci.
(2009), doi:10.1016/j.jas.2009.08.003
lago Salitroso (Provincia de Santa Cruz, República Argentina). Magallania 33 (2),
21–28.
Tieszen, L., 1991. Natural variations in the carbon isotope values of plants: impli-
cations for archaeology, ecology and paleoecology. Journal of Archaeological
Science 18, 227–248.
Tykot, R., Falabella, F., Planella, T., Aspillaga, E., Sanhueza, L., Becker, C., 2009. Stable
isotopes and archaeology in central Chile: methodological insights and inter-
pretative problems for dietary reconstruction. International Journal of Osteo-
archaeology 19 (2), 156–170.
van der Merwe, N., Medina, E., 1991. The canopy effect, carbon isotope ratios and
foodwebs in Amazonia. Journal of Archaeological Science 18, 249–259.
van Klinken, G., Richards, M., Hedges, R., 2000. An overview of causes for stable
isotopic variations in past European human populations: environmental,
ecophysiological, and cultural effects. In: Ambrose, S.H., Katzenberg, M.A. (Eds.),
Biogeochemical Approaches to Paleodietary Analysis. Kluwer Academics/
Plenum Press, New York, pp. 39–63.
Yacobaccio, H., Morales, M., Samec, C., 2009. Towards an isotopic ecology of
herbivory in the Puna ecosystem: new results and patterns on Lama glama.
International Journal of Osteoarchaeology 19 (2), 144–155.
Zárate, M., 2002. Los ambientes del Tardiglacial y Holoceno en Mendoza. In: Gil, A.,
Neme, G. (Eds.), Entre montañas y desiertos: arqueologı́a del sur de Mendoza.
Sociedad Argentina de Antropologı́a, Buenos Aires, pp. 9–42.