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. Martınez d, Gustavo G. Politis e, Luis A. Borrero f, Gustavo A. Neme c
a

CONICET – IMHICIHU, Saavedra 15, 5 (1083), Buenos Aires, Argentina 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, Olavarrıa, 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
b

a r t i c l e i n f o
Article history: Received 20 June 2009 Received in revised form 6 August 2009 Accepted 11 August 2009 Keywords: Guanaco Stable isotopes d13Ccollagen Spatial tendencies Geographic tracers South America

a b s t r a c t
We synthesize and discuss available isotopic data on Holocene guanaco samples from southern South America, extending from Tierra del Fuego Island to northern Patagonia and the Pampean region. We evaluate temporal and spatial tendencies on the basis of 91 samples (mainly based on d13Ccollagen values), presenting implications for paleodietary research in archaeology. We conclude that there are no strong 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 (e.g. canopy effect, variations in altitude, ecotones) in structuring isotopic variability. Finally, on the basis of the present analysis we suggest a number of hypotheses and perspectives for the use of stable isotopes as geographic tracers of guanaco distribution in the past, and specific implications for the paleodietary study of human samples. Ó 2009 Elsevier Ltd. All rights reserved.

1. Introduction 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 Gonalons, 1999; Politis and Pedrotta, ´ ´ 2006; Gutierrez and Martınez, 2008; Neme and Gil, 2008; Morales et al., 2009). Stable isotopes on human remains provide an independent 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

supra-regional unit of analysis covers a latitudinal range of 20 and 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; Gonzalez 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.

* Corresponding author. E-mail address: ramidus28@gmail.com (R. Barberena). 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

Guanaco troop in Tierra del Fuego Island (53 S). 2006. Study area and fitogeographic regions. 1978).. The Monte Province coincides with the so called South American Arid Diagonal (Bruniard.. Currently guanacos are more abundant in the Patagonian steppe and in the foothills of the Andean Mountains (Cajal. and Schinus mostly represented. Stable isotopes from archaeological samples can contribute to develop this endeavor in Holocene times. (2009).003 . but also has the ability to alternate seasonally between grazing and browsing. that flank the western side of South America from 37 to 54 south (at Tierra del Fuego Island. Martınez et al. Tierra del Fuego. This species typically feeds on the herbaceous stratum (grasses. et al. Gonzalez et al. The Espinal Province constitutes a belt surrounding the Pampa to the west (Fig. Guanaco (Lama guanicoe) isotopic ecology in southern South America:.08. Southern Patagonia. importantly. 1980). Archaeol. / Journal of Archaeological Science xxx (2009) 1–10 are shrub and herbaceous steppes that may have varying frequencies of C3 and C4 plants (although the former are usually dominant). 1996. Humid Pampas. steppe. 2007.. 2009). Bachelor males form a separate herd (Raedeke. flat terrain and low hiding cover (Cajal. The main community is a xerophytic shrub steppe represented by Larrea and Prosopis.. 5. The main communities represented in the areas considered here are shrub and herbaceous steppes. 1988. 1996). grass-likes [Juncaceae and Ciperaceae]. 1. occupying the different fitogeographic regions described. from open habitats and scrubland to open forests (Cajal. J. Fitogeographic regions: green. Guanaco populations from steppe. C4 and CAM taxa may be present in varying densities and have been isotopically identi´ ´ fied (Gomez Otero. The main communities Fig. R.. Central-northern Patagonia. black.2. The climate is dry and cold with mean annual precipitations between 150 and 500 mm. The main vegetal community represented is the herbaceous steppe with the genera Stipa. and forbs). and xerophytic woodlands with the genera Prosopis. This camelid lives in herds composed of females. it is bounded by the Monte to the east and by the Sub-Antarctic forests to the west (see below). young individuals (chulengos) and a dominant male. 4. 2).. Methodology This paper provides a comprehensive synthesis of guanaco stable isotope values in southern South America. The Sub-Antarctic Province is closely associated to the Andes mountains. Sub-Antarctic Province (based on Cabrera. 1). 1).jas. 1976). with mean annual precipitations between 80 and 200 mm. Barberena et al. Sci. The climate is temperate-cold and humid with precipitations between 800 and 2000 mm. 1978. Guanacos inhabit preferably in open areas with grasslands.. 2. Puig et al. There are records from the Monte Province indicating that guanacos do not strongly select against C4 grasses due to its capacity to digest high fibercontent taxa (Puig et al.2009. 1978).. grey. 2. 1980) and extending from the sea level up to 4500 masl.ARTICLE IN PRESS 2 R. Espinal Province. guanicoe) is the larger terrestrial wild mammal in southern South America.. straight lines. Ortega and Franklin. The dominant communities are deciduous and ever-green forests where Nothofagus is the main genera (Boelcke et al. with mean precipitations between 500 and 1100 mm associated with continental conditions that produce higher evapo-transpiration rates than in the Pampa. 6. 2006). which weighs between 85 and 120 kg ´ in average for different regions (recent review in Gonzalez et al. population dynamics and habitat use of guanacos in those regions are poorly known. Bonino and Pelliza Sbriller. Monte Province. according to forage availability. However. References: Study areas: 1. 1. Acacia. Please cite this article in press as: Barberena. Fig. 3. 2. Southern Patagonia. Llano. forest. Guanaco ecology The guanaco (L.. pre-cordillera.. doi:10. 1996. Poa. cordillera and transition zone between cordillera and steppe zones have been well studied with regard to food habits (Raedeke. this behavior allows to define the guanaco as an ‘adaptable mixed feeder’ that has the ability to digest low quality ´ forage (Puig et al. Until the Nineteenth century. see Fig. 2009). although they are known to inhabit forested environments in Tierra del Fuego Island (Raedeke. 1991.. Patagonia Province. 1985). 1980). The main criterion for the inclusion of the samples is the existence of accurate Fig.1016/j. and Briza among the most represented. yellow. The Patagonia Province extends from southern Mendoza (36 south) up to Tierra del Fuego Island (53 south). 1982). Pampean Province. the guanaco was present in almost all regions of Argentina. Centralsouthern Mendoza.

Temporal tendencies There is a subtotal of 74 samples with available information on d13Ccollagen and radiocarbon dates (in all cases these two determinations were obtained from the same specimen).. Fig. In Fig. van Klinken et al. perhaps favoring C3 species.jas. Species with the C4 photosynthetic pathway display a global d13C average value between À13 and À12& (Ehleringer and Cerling. r ¼ À0. 1). / Journal of Archaeological Science xxx (2009) 1–10 3 taxonomic determinations. from the southern end of Tierra del Fuego Island. The samples from Central-northern Patagonia. As already suggested. although it neither allows ruling this possibility out. This wide geographical Please cite this article in press as: Barberena. 2001). 2000). We present a review of information on d13Ccollagen. The results presented below operate on two spatial scales of analysis: first. 4 we show the ranges of variation for the regions considered.2009. which provide information that is adequate for our goals (Table 2). The absence of marked temporal trends is not unexpected if we take into account the large spatial scale considered. This information includes d13Ccollagen results for 90 samples and 14C dates for 74 samples. Therefore. Of the 90 d13Ccollagen samples. suggesting a number of explanations for them. 3. we focus on the d13Ccollagen information since it provides a relatively large sample size (n ¼ 90). in the case of southern Santa Cruz we treat the samples that come from the Andean forests and the eastern steppes separately. we present isotopic values for 91 guanaco bone samples from six different regions: central-southern Mendoza province (n ¼ 12). Forested ecosystems are composed of C3 species and may present a distinctive isotopic signature. a regional scale. area 4) in southern Patagonia does not show any correlations between chronology and d13Ccollagen values (Borrero et al.003 . accompanied with an increment in precipitations and lake expan¨ sions (Schabitz. 2000). Palynological. 5. at 34 S (Fig. 1999). p 0. Therefore. During these times. 11&) for this species on the macro-regional scale. see Fig. and pedological information indicates the occurrence of important climatic changes ¨ during the Late Holocene (Schabitz.7& for a southern Mendoza sample. Different projects integrated by the authors have produced isotopic data for 67 guanaco samples.18. The descriptive statistics shown on Table 2 indicate a very wide isotopic range (of ca. This value does not indicate an important incidence of sample sizes on the isotope diversity recorded for each region. forest (n ¼ 11). located between 40 and 39 of southern latitude. which will be evaluated on the basis of future oriented sampling. R. geomorphological. there are 25 samples that were generated in the context of other projects. second. The database discussed here was not generated in relation to the specific goal addressed in this paper. in order to exclude specimens belonging to other Lama species. 2009.. On the other hand. This large amount of variation for this herbivore species constitutes an interesting pattern in itself.. 1. which may have affected the composition of vegetal communities modifying the relative abundance of C4 and C3 species. On the other hand. although the discussion is based only on the d13Ccollagen values.2% of this subsample..10. Results The information presented is already published in a large number of contributions (see references in Table 1). 2001). 34 S. Barberena et al. there is a low positive correlation between sample size and variance for these regions -which is clearly non-significant-: r ¼ 0. L’Heureux. 1991.41.. central-northern Patagonia (n ¼ 13).g. Heaton. Nevertheless. see contextual information in Martınez et al.4&.3& for a Pampean sample to À14. This preliminary inference needs further data in order to be tested.. Sci. 1). southern Patagonia. ´ r ¼ 0. being the product of diverse interests usually operating in regional or local spatial scales. our research contributes to identify interesting regional and supra-regional patterns. we can proceed to an evaluation of preliminary temporal and spatial tendencies on different scales of analysis. show a positive and significant correlation between chronology and d13Ccollagen values (n ¼ 11. therefore. 3) between these variables that suggests the absence of temporal tendencies in guanaco d13Ccollagen values at the macro-regional scale. 1994).07. et al. where the main issue to be considered is the role of latitude in structuring guanaco isotopic values. Globally. besides. which are transmitted through the successive steps in the trophic chains (van der Merwe and Medina. 2008). 2007. Archaeol. and d15N. We present the complete stable isotope and radiocarbon database in Table 1 and a descriptive statistics synthesis of this information in Table 2. d13Ccollagen on herbivores is conditioned by the isotopic values of the vegetal species regularly consumed.. 5. we get a total lack of correlation (n ¼ 66. This region is currently located at a faunal and fitogeographical ecotone between the Monte and Espinal Provinces (Abraham et al. species showing the C3 pathway have average d13C values between À27 and À26&.1. the Humid Pampas (n ¼ 26).. to central Mendoza Province.412. 2009.1016/j. steppe (n ¼ 25). This introduces sampling limitations in our analysis. 2003). p 0. and display lower adaptive efficiency in settings characterized by high temperatures and water restrictions (Tieszen. 72 are the product of 14C AMS datings. information available for different localities of Santa Cruz province (Fig. Ehleringer and Cerling. 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. This hypothetic situation may explain the tendency towards more depleted guanaco values recorded for the last 2000 years.61.04. 2009). For reasons that are discussed and justified below. at 54 S. which averages quite different climatic and ecological settings. p 0... If we limit this analysis to the last 4500 14C years. given that there are large non-sampled areas. we proceed to analyze values on smaller regional scales. which provide an 89. Spatial tendencies The regions considered present different ranges of isotopic variation that may be partly explained by sample sizes (Table 3). This is a key issue north of ca. Table 1). given that they may be characterized by the recycling of inert carbon that produces impoverished isotopic values. given the presence of wild and domesticated camelid species whose morphological identification is debated (Izeta. inviting to evaluate which are the temporal and spatial factors that determine it. 2009).ARTICLE IN PRESS R.53). This range extends from À25. and Tierra del Fuego Island (n ¼ 4) (Fig. presenting an average value of À20. sacrificing sample size but gaining better control on the ecological homogeneity of the spatial units of analysis. J. p 0. Tessone and Belardi. 1. which allows us to enquire on the influence of different ecologic and topographic contexts in the stable isotope results. r ¼ À0. southern Patagonia. this region underwent climatic shifts from arid to semiarid conditions. (2009). Guanaco (Lama guanicoe) isotopic ecology in southern South America:. These kinds of values have already been used for continent-wide isotopic discussions (e. 4. 1991. Macro-regional scale The macro-regional scale of analysis covers a latitudinal range of 20 . There is a very weak negative correlation (n ¼ 74.08. being adapted to arid and hot climatic conditions. a macro-regional scale. our main goal is to integrate these results within a specific frame of research. Despite these limitations. d13Capatite. doi:10. but provides a basis for introducing a temporal dimension in northern Patagonian isotopic ecology studies.

9 À19..LR1/2 44 San Antonio 1-FCS 45 San Antonio 2 S 1.. 2005 Bonomo.. Barberena et al. pers.9 Gil Gil Gil Gil Gil Gil Gil Gil Gil Gil Gil Gil et et et et et et et et et et et et al.4 À19.3 7.6 À19.ARTICLE IN PRESS 4 R.6 6. al..2 ´ Martınez.5 À18.E 40 El Tigre-FCS.6 6.9 5. 2008 ´ Martınez et al.6 À10. comm.1 À23. 2001 Politis et al.6 6..4 À19. Central-southern Mendoza (N ¼ 12) (*) 1 Arroyo El Desecho 10 2000 2 Agua de los Caballos 1025 3 Cueva de Luna 1300 ´ 4 El Indıgeno 3600 5 La Gotera 1500 6 Arroyo Malo 3 2000 7 Ojo de Agua 1600 8 El Sosneado 3 2050 9 Arroyo El Desecho 10 2000 10 La Corredera 1150 11 Agua de los Caballos 1 1025 12 Gruta El Manzano 1300 2.. 2009 Madrid et al.1 À18.6 À19.8 À23.4 À18... 2001 Politis et al...3 À18. al..7 À19. ´ Martınez. (2009).4 À19. 1999 Please cite this article in press as: Barberena..2009.ET.7 À23.3 À9. comm..3 À20.. 2005 Tessone et al.. Steele and Politis.8 À22. 2006 ´ Politis and Bayon. 2007 ´ Gomez Otero. al. J. steppe 52 La Siberia 2 53 La Siberia 2 ´ 54 Alero del Leon 55 Parapeto 4 ´ 56 Alero Gerasın II 57 Cerro Pampa 2 ´ 58 Solıs 59 Cerro Pampa 2 60 El Sosiego 2 61 A.0 À19.5 À18.7 À18.6 À25. 2005 Tessone et al.C20 41 El Tigre-FCS. 2005 Politis et al. 1999 Carballo Marina et al.9 À19.3 6..8 À16.6 7.2 Politis et al. 2009 ´ Martınez et al.ET1.2 5.FCS. 2005 Politis et al. Humid Pampas (N ¼ 26) ´ 13 Zanjon Seco 2 ´ 14 Zanjon Seco 2 15 Calera 16 Calera 17 Calera 18 Calera 19 Calera 20 Calera 21 Cortaderas 22 Nutria Mansa 1 23 Nutria Mansa 1 24 Nutria Mansa 1 25 Nutria Mansa 1 26 Arroyo Seco 2 27 Arroyo Seco 2 ´ 28 Quequen Salado 1 29 30 31 32 33 34 35 36 37 38 ´ Quequen Salado 1 ´ Quequen Salado 1 ´ Quequen Salado 1 ´ Quequen Salado 2 Tres Reyes 1 Tres Reyes 1 La Barrancosa San Clemente VI Paso Otero 1 La Olla 4 USF USF USF USF USF USF USF USF USF USF USF USF 5905 5906 5907 6173 8354 8355 8356 8357 5913 8864 8865 8866 – – – – – – – – – – – – À19. 2005 Tessone et al.. 2005 Politis et al....8 À19.. 2005 Tessone et al.3 À18. 2008 ´ Martınez.. 2008 ´ Gomez Otero. comm.1 4.. Guanaco (Lama guanicoe) isotopic ecology in southern South America:. 2005 Bonomo.5 À17. 2005 Tessone et al.7 À18. doi:10.8 À19.4 À19.9 À18.6 À25. 2007 48 48 48 48 48 48 48 48 50 50 540 540 540 540 540 540 540 540 90 30 71 71 71 71 71 71 71 71 72 72 150 150 150 150 150 150 150 150 350 170 UGA 10014 UGA 10013 UGA 10009 UGA 08705 UGA 10017 UGA 10019 UGA 10018 UGA 10020 GX 25278 GX 26196 1710 Æ 40 1100 Æ 40 2190 Æ 50 2010 Æ 50 1580 Æ 40 310 Æ 40 1140 Æ 40 170 Æ 40 1920 Æ 40 650 Æ 40 À18..9 À20. 2008 Prates.8 À18.8 6. 2008 ´ Martınez et al. al. Archaeol. 2008 Prates...C19 42 Loma Ruiz 1.1 À9.2 À19.3 5.8 Tessone et al. et al.. 2009 ´ Martınez.7 À11.2 À16... 2008 ´ Paleo and Perez Meroni.3 À21. al. 2002 Madrid et al.9 À19. 2005 Politis et al..9 À20.003 .8 À20. 2000 Madrid and Barrientos. al.ET.08.. Piedra Quemada 39 39 39 39 39 39 39 39 40 39 39 42 42 460 460 460 130 130 390 390 390 100 500 500 62 62 62 62 62 62 62 62 64 65 65 220 220 220 380 380 090 090 090 110 170 170 Ua 22561 AA 81830 AA 81834 AA 53331 AA 53332 AA 81832 AA 77966 AA 81831 AA 2551 AA 62794 AA 62795 USF USF 455 Æ 45 473 Æ 43 536 Æ 43 1615 Æ 50 1935 Æ 44 773 Æ 44 764 Æ 45 988 Æ 44 938 Æ 45 398 Æ 46 483 Æ 43 – – À19.jas.1 46 San Antonio 2-FCS 47 Angostura 1 48 Negro Muerto 49 Negro Muerto 50 Costa de Chubut 51 Chubut near coast 4.6 -8. 2009 Steele and Politis.4 À19.4 8.0 À24.5 À22.2 À20.5 À20. 2002 Madrid and Barrientos. 2009 ´ Martınez. 2005 Politis et al.. 2005 Carballo Marina et al.1 À20. 2007 Bonomo.2 4..0 À12. 3. al...6 À19. 2005 Tessone et al. pers. # Site Altitude (masl) Latitude (South) 35 35 36 34 35 34 35 34 35 36 35 36 110 220 050 300 520 520 90 500 110 310 220 040 Longitude (West) 70 68 60 69 69 69 69 69 70 68 68 69 30 180 430 590 570 540 380 540 30 320 180 430 Lab code 14 C date d13Ccol d13Cap d15N References 1... Southern Patagonia. Sci..2 À17.7 À19. Central-northern Patagonia 39 El Tigre. al.8 À19..8 À21. 2005 Massigoge. 2002 Madrid et al.1016/j. 2002 Madrid et al.9 À20.. al.1 4. 2002 Madrid et al..1 À14. R.0 À19. 2005 Bonomo. al.0 4. 2006 2006 2006 2006 2006 2006 2006 2006 2006 2006 2006 2006 100 100 200 200 200 200 200 200 90 <20 <20 <20 <20 110 110 <20 <20 <20 <20 <20 180 180 180 <20 100 <20 (N ¼ 13) <20 <20 <20 <20 <20 <20 <20 <20 70 60 60 <20 <20 (N ¼ 25) 300 300 300 300 300 300 300 300 200 500 38 38 36 36 36 36 36 36 38 38 38 38 38 38 38 38 100 100 580 580 580 580 580 580 200 240 240 240 240 210 210 490 59 59 60 60 60 60 60 60 59 58 58 58 58 60 60 60 100 100 140 140 140 140 140 140 390 150 150 150 150 140 140 320 38 490 38 490 38 490 38 49 37 37 37 35 38 38   0 60 320 60 320 60 320 60 33 60 60 60 57 59 61   0 56 560 560 140 120 590 0 34 340 080 160 70 ´ 21 0 CAMS 4993 CAMS 4994 AA 67732 AA 67733 AA 67735 AA 64617 AA 71669 AA 71671 AA 67736 AA 55114 AA 55115 AA 55116 USF 8751 AA 24052 AA 52613 Beta 169820 Beta 157398 Beta 157397 Beta 169821 Beta 169822 AA 7070 AA 7971 AA 59507 AA 29412 AA 72844 AA-80664 3070 Æ 40 3080 Æ 40 3008 Æ 46 2075 Æ 44 1748 Æ 42 2232 Æ 55 3390 Æ 170 3005 Æ 66 2270 Æ 190 2705 Æ 66 3080 Æ 110 2920 Æ 110 – 7540 Æ 80 8390 Æ 410 360 Æ 40 790 Æ 40 940 Æ 40 960 Æ 40 1720 Æ 40 1845 Æ 50 2235 Æ 50 1676 Æ 46 935 Æ 55 3056 Æ 42 6960 Æ 71 À19.LR1/1 43 Loma Ruiz 1. 2000 Messineo..3 À25. pers. al.0 À19.6 À20. 2008 ´ Martınez et al. / Journal of Archaeological Science xxx (2009) 1–10 Table 1 Isotopic and radiocarbon results for guanaco samples from southern South America. 2005 Tessone et al...0 À19. 2009 Prates.

08.4 À20.9 À14. 2009 Borrero et al.. 2007 Aschero et al.8 À21...3%) Fig. 2001 Favier Dubois and Borrero. 1 200 80 81 82 83 84 85 86 87 Alice 1.0 Franco et al. 2003 Franco and Borrero.2 5. 1999 Borrero et al.9 À20.3 d13Cap d15N References Carballo Marina et al.2009. Guanaco (Lama guanicoe) isotopic ecology in southern South America:. 2006 Borrero et al.1016/j. Tierra del Fuego Island (N ¼ 4) ´ 88 Tunel I <20 ´ 89 Tunel I <20 90 San Genaro 2 <20 ´ 91 Las Mandıbulas 3 <20 54 54 53 53 490 490 190 120 68 68 68 68 090 090 170 380 AC 703 AC 704 USF386/243 Beta 122882 5280 Æ 100 3190 Æ 100 – 1190 Æ 50 À20.0 À21. / Journal of Archaeological Science xxx (2009) 1–10 Table 1 (continued ) # 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 Site V.7%) 74 (81. Although it is expected that this Table 2 Synthesis of isotopic data for guanaco samples from southern South America. range provides a great window to evaluate the incidence of latitude in relation with isotopic variation...jas.. 1998À1999 Franco and Borrero.. Please cite this article in press as: Barberena.. this isotopic record is adequate to evaluate macro-regional variations in guanaco isotopic ecology..8 À23. 2007 6. 2 Alice 2 Chorrillo Malo 2.. II ´ Condor 1. 4E.1 À21..8 À19. 2001)...1 À24.. 19 D9C 200 200 200 200 300 900 900 900 50 200 50 200 50 200 50 300 50 300 50 90 50 90 50 30 47 570 47 570 47 570 72 310 72 310 72 310 72 400 72 400 72 360 72 360 72 170 72 050 72 050 72 050 Beta 91301 Beta 91302 Beta 112231 Beta 112232 GX 27174 Beta 148743 GX 25279 Beta 138991 UGA 866 UGA 870 UGA 873 3110 Æ 50 170 Æ 30 1420 Æ 70 1480 Æ 70 740 Æ 60 3790 Æ 80 9740 Æ 50 1840 Æ 40 5400 Æ 64 8362 Æ 68 10690 Æ 72 À22.8 À19.0 À21. 1999 520 Æ 40 160 Æ 40 240 Æ 40 2000 Æ 40 1510 Æ 30 4340 Æ 40 2850 Æ 40 3490 Æ 50 965 Æ 40 1360 Æ 65 1070 Æ 40 660 Æ 50 4370 Æ 50 – 1685 Æ 70 2. et al. we suggest that this result is not the product of biases due to small sample sizes..ARTICLE IN PRESS R.9 ´ Albero et al. R. 1999 Aschero et al. 1 ´ Cerro Leon 1.5 À20. 1999 Borrero et al. 2009 Borrero et al. 2009 Borrero et al.7 À20..7 À21.003 . Region Central-southern Mendoza Humid Pampas Central-northern Patagonia Southern Patagonia. 2005 Note: (*) contextual information indicates that the Central-southern Mendoza samples are Holocene in age. 4E. An analysis of correlation between latitude and d13Ccollagen values produces a result of r ¼ À0.0 À22. II ´ Condor 1. III La Carlota ´ Cabo Vırgenes 22 ´ Cerro Leon 3 ´ Cabo Vırgenes Cerro Verlika 1 Altitude (masl) 500 40 < 20 < 20 < 20 400 400 145 150 150 70 40 400 <20 1100 Latitude (South) 50 52 52 52 52 50 50 52 51 51 50 52 50 52 50 30 190 200 200 200 510 510 070 890 890 510 190 360 200 360 Longitude (West) 72 68 68 68 68 72 72 69 69 69 72 68 72 68 72 170 210 230 230 230 120 120 330 360 360 140 220 160 220 160 Lab code GX 25775 GX 25773 GX 27868 GX 27864 GX 27865 GX 27863 GX 27866 Ua 21902 Ua 24658 Ua 24658 Beta 215184 GX 32586 GX 32583 USF 582 GX 25277 14 5 C date d13Ccol À20. 1986 Orquera and Piana. indicative of a negative and very weak correlation between these variables at the large macro-regional scale. 1998À1999 Borrero et al. 3. (2009). In order to support this statement.... 2009 Barberena. Since global climate presents a marked latitudinal configuration (Strahler. 2009 Borrero et al. steppe Southern Patagonia... 1998À1999 Borrero et al. Piedra Quemada 2 ´ Cabo Vırgenes 7 ´ Cabo Vırgenes 8 ´ Cabo Vırgenes 4 ´ Cabo Vırgenes peat ´ Cerro Leon 1. 4 ´ Rincon Amigo CCP 7.4 À19. and considering that climate is one of the main conditioners of the abundance of C3 and C4 vegetal species (Ehleringer and Cerling.0 À22..0 À22. Southern Patagonia.3 À19. 2002 Franco et al. which are averaged at the macro-regional analysis producing the emergent lack of correlation observed between latitude and isotopic values.9%) 7 (7. 2007 Aschero et al. 1 Chorrillo Malo 2.28 (p < 0. Isotopic results for guanaco are distributed in two large latitudinal subsets: the southern set corresponds to the regions of Tierra del Fuego and southern Patagonia. Archaeol.. forest (N ¼ 11) 77 Alero del Bosque 225 78 Lago Roca 3 250 79 Alice 1. 1996 ´ Guichon et al. Capa 5 N8A CCP7. J. 2006 Borrero et al. 2009 Campan et al. Barberena et al. Temporal tendencies in d13Ccollagen values for Holocene guanaco samples. 17 (2) C11B CCP7. 2003 Carballo Marina et al...0 À24.8 À20.6 À20.3 À20. 2009 Borrero et al. 2 Orejas de Burro 1. 2007 Borrero et al. we explore climatic and ecologic variables acting on smaller scales.3 À20. 1999 Franco et al. 2009 Borrero et al.. located between 54 and d13Ccollagen d13Capatite d15N 11 26 13 25 11 4 2 – – – 11 2 2 1 – 14 C Dates – 25 11 24 11 4 4 1 91 (100%) 90 (98.7%) 1 3 17 (18.2 4.01). Sci. doi:10..3 À19...0 À24..1 À21. forest Tierra del Fuego Total Samples 12 26 13 25 11 tendency will experiment modifications with increasing amounts of isotopic data.5 À21. 2006 Borrero et al. 1982).

d13Ccollagen values and latitude. 48 S... We suggest that this segmentation of the southern and northern sets of samples allows isolating the incidence of latitude in the isotopic data. doi:10. In the case of the southern group we segregate the samples coming from the western forest environments and from the eastern steppes. 1. we isolate assemblages from areas that are more ecologically homogeneous (fitogeographic Provinces in Fig. 4. In the northern group we separate the samples from the Humid Pampas from those from Central-northern Patagonia and Central-southern Mendoza. being emplaced between 42 and 34 S.13 (p 0. From now we treat these two sets of isotopic data separately. In the case of the southern area.41). the northern set includes Central-northern Patagonia. we consider it is necessary to proceed one step further by subdividing these sets in terms of general ecological or climatic information. clearly limiting our ability to evaluate latitudinal variations in a continuous manner. Mancini et al. 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 À0.01). Please cite this article in press as: Barberena. Zarate. Sci. by removing some local ecologic and climatic variables from the analysis. given that both sets of samples average quite different ecological contexts. R. Central-southern Mendoza. et al. 1. the absence of a strong correlation between these variables. 2009). In the case of the northern group. paleoecological data indicates that changes in forest extension do not affect the suggested segregation of the samples (Huber et al. (2009). on the other hand. as already seen at the macroregional scale. Labraga and Villalba. Note: the r values do not include the samples from the southern Patagonian forest and the Humid Pampas.. which are located within the climatic-ecologic zone defined as the ‘South American Arid Diagonal’ (Bruniard.. lending support to their use for organizing Late Holocene samples.08. there is important paleoecological data that indicates their temporal depth.003 . 2008). Cabrera. 1982). diverse lines of paleoclimatic evidence indicate that the Arid Diagonal experimented changes in its specific configuration (Mancini et al. 1991). 5. 2004.37 (p < 0. 2002.1016/j.2009. 1976). Next we explore the correlation between latitude and d13Ccollagen values for both sets Fig. between 47 and 42 S. shown as unimportant so far. given that these ecosystems may show isotopic variation that is to a large extent independent of latitude (van der Merwe and Medina. whereas the northern set produces an r ¼ À0. Therefore. Both of these values are negative and very low indicating. / Journal of Archaeological Science xxx (2009) 1–10 Fig.. and the Humid Pampas. Ranges of variation in d13Ccollagen values on guanaco samples.. Archaeol.jas. 2005). Barberena et al. Although these decisions are based on the present configuration of these ecosystems.. Guanaco (Lama guanicoe) isotopic ecology in southern South America:. but these changes do not affect the separation suggested for Late Holocene ´ samples (Fig. Nevertheless.. There is an intermediate eight degrees wide area that is void of information. J.ARTICLE IN PRESS 6 R.

most importantly for the identification of the consumption of marine resources. Ecologic information attributes the behavior of inhabiting forested settings only to guanacos from Tierra del Fuego Island (Raedeke.57 (p < 0. 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). 10 samples come from settings between 1000 and 2000 masl. 2006..37. they do not show a ‘canopy signature’). et al.1016/j.jas. suggesting that they are not statistically different despite the large geographical gap that separates them. This change appears more strongly in the southern set (Fig. The correlation results are different to those obtained previously. Barberena et al. 1991).68 (p < 0. 5. These samples present depleted d13Ccollagen values. indicating negative and weak correlations (Gil et al. if we exclude this outlier from Please cite this article in press as: Barberena.. Gomez Otero. 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. in the lowlands. in the southern set. The altitudinal range between 1000 and 2000 masl represented in these samples corresponds to a transitional area between C4-dominated herbaceous communities. In Fig. Regional scale The results presented above indicate that there is not a latitudinal tendency in the guanaco isotopic data at the macro-regional scale. 5.731 (p 0. while they are absent at higher altitudes of the Andes mountain range (Cavagnaro. we consider that this global similitude emphasizes that latitude is not the only -neither the main-factor operating at the large macroregional scale.003 . it must be mentioned that although the two samples ´ from Tunel site in Tierra del Fuego also come from a closed forested setting. 5). Correlations between altitude and d13Ccollagen and d15N values in Centralsouthern Mendoza.. with wider forest ecosystems and an apparent more fluid connectivity between eastern and western flanks of the Andes (facilitated by general biogeographical conditions). with correlations being more negative and statistically significant in the present analysis.2009. Interestingly. we can see that there is a marked change. Therefore. considering only the samples from the steppes. we are respectively excluding the cases from the southern Patagonian forests and the northern Humid Pampas (Fig. in the highlands (Cavagnaro. Of this subset. Borrero et al. This isotopic signature provides interesting geographic information on guanaco movements in the past. providing a new proxy of its persistent use of forest ecosystems in the continent.. Llano. 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. which are separated by a minimum distance of ca. R. As already suggested. 2009). may be even more important for the Pacific coastlines where marine foods were usually the main staple. This indicates a situation different to those recorded at both the macro-regional scale and at the ecologically undifferentiated regional analysis. and for northern Patagonia. 2009). Sample size is small and spatially heterogeneous for solving this issue. and discriminating the samples in terms of their ecological context. 1988. J.2. if a consideration of ecological context is introduced in the analysis. characterized by a largely forested southern region. p < 0. consisting in the gradual enrichment of isotopic values recorded independently in the southern and northern geographical sets. Guanaco (Lama guanicoe) isotopic ecology in southern South America:. in Santa Cruz province. Archaeol. 6. Llano. On the other hand. A t test of these two groups of samples produces a result of t ¼ 1. to C3-dominated communities... there is a negative and relatively important correlation between latitude and d13Ccollagen values identifiable at smaller regional scales. which is suggested here for southern regions located at the eastern flank of the Andes. although it can be stated that its implications for human paleodietary research will remain.08.13.005). Next we develop regional cases that allow exploring the role played by selected ecological and topographical variables. / Journal of Archaeological Science xxx (2009) 1–10 7 Fig. and therefore this information extends this pattern to the southern end of the continent. There is another interesting spatial pattern in Fig.ARTICLE IN PRESS R. This record will have to be reevaluated in the future on the basis of a larger dataset. and those located in the Arid Diagonal. 2007. Sci. which is a key topic in Patago´ nian archaeology (Barberena.41) and northern (r ¼ À0. On the other hand. doi:10. This situation. p 0.005) and the value for the northern set is of r ¼ À0. 2009). The southern Mendoza region provides information to evaluate the relationships between isotopic values and topographic variation. 5). 1978). we identify an important incidence of latitude on the guanaco d13Ccollagen values. These cases will be used to illustrate the potential of isotopic information as a geographic tracer of guanaco ranges in the past. There is a set of 11 samples from this region (Table 1) that allows an initial assessment of the influence of altitude on guanaco d13Ccollagen and d15N values. since the value for the southern set is of r ¼ À0. in the northern set. this pattern also provides an important factor to be considered in human paleodietary reconstructions. 1988. Despite this situation. 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. although a number of expectations that will be tested in the future can be developed on the basis of the available data. Comparing these values with those obtained for the global southern (r ¼ À0. we suggest that at this regional scale of analysis. 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. 6 we present analyses of correlation for d13Ccollagen and d15N values with altitude..01) sets.08). Therefore. (2009). 50–100 km. 2002.

1999–2001. something that can be independently supported on the basis of the latitudinal variations documented here. The discussions developed on regional scales provide an example of the role of stable isotopes as a geographic tracer of guanaco home ranges.. and guanaco foraging patterns and grass selectivity. et al.003 ... this issue is relevant for the isotopic evaluation of maize consumption by humans.0 0. On the contrary. Acknowledgements We would like to acknowledge Augusto Tessone for his thorough reading of the paper. On the basis of the available information. Prieto. although it can be suggested that it is related with the ecological complexity of these regions.9 0. When associated to variations in vegetal isotopic ecology..8 À25. The four regional cases briefly presented here highlight the importance of differing levels of regional ecological complexity in structuring guanaco isotopic variation.6 2 Variance 2. Universidad de Buenos Aires.2009. steppe Southern Patagonia. and altitude. 3).8 1. Available information already mentioned indicates that guanacos have an important foraging flexibility. ´ ´ Agencia Nacional de Promocion de la Ciencia y la Tecnologıa. as well as to Cristina Bayon. This information suggests that guanacos would not strongly select against C4 species. Roig.71 3.8 0. which is a key topic for the archaeology of the South Central Andes (Gil et al. Archaeol.jas. Fig. 1996).08. Sci. Besides.3 À24. like Zea mays (see Tykot et al. constitutes a faunistic and fitogeographic ecotonal zone connecting the Monte and Espinal provinces (Abraham et al. F.4 2. Finally...8 À25. (2006: 172). Barberena et al. to Teresa Civalero.4 Median 18. by reducing the scale of analysis and introducing a discrimination of the samples in terms of their ecological context.9 Standard deviation 1. / Journal of Archaeological Science xxx (2009) 1–10 Table 3 Descriptive statistics for d13Ccollagen on guanaco samples. Conclusions The review of available information presented here allowed us to identify a number of new and interesting tendencies on guanaco d13C data. the integration of data on past distributional ranges. 2004) by providing a proxy of the past spatial distribution of ´ guanacos. Guanaco samples from the Humid Pampas and Centralnorthern Patagonia present widest levels of isotopic variation as indicated by their variance values (5.2 À19. by characterizing the inter-regional isotopic variability on guanaco. such as ecological context.. see Table 3. Yacobaccio et al...2 À20. As suggested by Gonzalez et al.3 the analysis.2 À22. isotopic information has a great potential to provide information on the altitudinal distribution of South ´ American camelids (Fernandez and Panarello. guanacos may have locally foraged on a number of different herbaceous communities (Cabrera. doi:10. The analyses developed in our projects were funded by the following institutions: Consejo Nacional de ´ ´ Investigaciones Cientıficas y Tecnicas. Universidad Nacional del Centro de la Provincia de Buenos Aires (INCUAPA).8 À20. on the other hand. 144–153.. on guanaco isotopic values. Mariano Bonomo. 2009. Mariana De Nigris. del Valle.. J. we have identified operating on different spatial scales.9 À21. this tendency is neither strong nor lineal (Fig..4 À16. References Abraham. although there is a gradual tendency towards enriched average values at lower latitudes. allowing them to forage C4 grasses with high fiber content (Puig et al. as a direct conditioning of global climate. we did not identify strong correlations between latitude and d13Ccollagen isotopic values. Clara Paleo and Mercedes Perez Meroni for allowing us to use their unpublished results.3 À23. J... topography. M.16 (p 0. which have the potential to produce the isotopic divergence recorded. This information is potentially useful for species conservation and protected areas management (Etnier.3 3. foraging behavior and genetic distinctiveness (Kadwell et al. corresponding to the macro-regional scale. 1996. this information will have important implications for comparative paleodietary research on human samples by means of stable isotopes. As already suggest. suggesting that altitude is not structuring the isotopic values in this sample.5 À14. which helped to clarify our arguments and provided valuable ideas on how to present them. 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. Llano.1 À21. In the Humid Pampas. Agustina ´ Massigoge. This would have the consequence of producing an averaged isotopic signal that fails to show strong altitudinal variations...8 Maximum value À14.4 2. population dynamics. Guanaco (Lama guanicoe) isotopic ecology in southern South America:. 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 altitudinal floors-. (2009). the only exception being the regional case of Central-northern Patagonia. Region Central-southern Mendoza Humid Pampas Central-northern Patagonia Southern Patagonia. Hugo Yacobaccio and Juan Bautista Belardi for their comments. forest Tierra del Fuego Total Samples 11 26 13 25 11 4 90 Average À18.ARTICLE IN PRESS 8 R. We have not recognized any temporal trend. we do not have any convincing explanations for these patterns. 2009). L.1 À22 À21.1016/j. whose isotopic signatures are not precisely known yet. which was a main staple for Holocene hunter-gatherers. Central-northern Patagonia. we do record a strong negative correlation between latitude and d13Ccollagen values.. 2009). Overview of the geography of the Monte Desert biome (Argentina). 2001) will help to define conservation units. In terms of spatial analysis.3 À20. At this smaller scale we are able to isolate the incidence of latitude.2 À18. Please cite this article in press as: Barberena.6 À19 À20. Pablo Messineo.44 0. National Geographic Society. 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. Torres. 2009). E.6 À20. when we shift to larger geographical scales we introduce another independent variables in the analysis.8 3.7 À16. 5).1 5. as exemplified with the case of a forest signal in southern Patagonian samples. 2006). dictating the appropriate spatial scale to conduct isotopic ecology studies.64). Coronato. 1976. F. the r value drops to À0. Godagnone. To Luciano Prates for the important discussions maintained and for providing infor´ mation. Ares. H. R.. and Museo de Historia Natural de San Rafael. producing a wider dispersion of isotopic values and reducing the correlation with latitude per se. 6.7 À20.7 Minimum value À19. At the largest level of analysis.9 À22. R.1 À20.72& and 3. Given that human consumption of isotopically enriched guanaco meat may mimic a low-level consumption of C4 domesticated species. 2009). Journal of Arid Environments 73 (2).8 À19.86& respectively.

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