assemblage 12 (2013): 28-42

South American Extinctions, a case study: the Rodrigo Botet Collection of the Museum of Natural Science in Valencia, Spain

by KARINA VANESA CHICHKOYAN

This paper aims to discuss the - South American megafauna extinctions, taking into account the evidence of bones exhibiting cut marks, identified in the Rodrigo Botet Collection of Valencia, Spain, donated by Rodrigo Botet at the end of the 19th century. Due to the amount of material present and the completeness of some specimens, this Collection is often considered the most important assemblage of South American megafauna fossils housed in Europe. Pleistocene- Holocene extinction is quite a controversial topic and a variety of explanatory arguments have been put forward over the last 30 years. In South America, the loss of such a considerable amount of native fauna, in particular the megafauna, coincided with two important events: Homo sapiens’ expansion onto the continent and the rise of the temperatures, both of which affected the transition. In order to contribute to current understanding of this event, a taphonomical analysis was performed on the Rodrigo Botet Collection and these data were then contextualised with palaeoecological information. The taphonomic appraisal of this archived material was useful in order to identify bones with anthropic cut marks and, thus to provide new data for the debate on the South American megafauna extinction during the Pleistocene-Holocene transition.

Keywords: extinction, megafauna, South America, taphonomical analysis, palaeoecological relationships, zooarchaeology

Introduction Taphonomical analyses were conducted in order to differentiate between anthropic traces A taphonomical analysis was undertaken on or cut marks, and non-anthropic agencies (e.g., the Rodrigo Botet fossil megafauna collection, carnivores, wind or water erosion, plant which comes from the northeast of the growth, etc.). The analysis was performed Pampean region in Argentina (Figure 1). This using magnifying glasses and stereoscopic research was conducted in order to add new microscopes. This methodological approach is data to the debate on the megafauna extinction critical for analysing non-stratigraphic of the Pleistocene-Holocene transition museum material where context data has been (Chichkoyan 2011). This event is widely lost. The examination of a bone surface is discussed in archaeological literature (cf. useful to distinguish any kind of anthropic Barnosky et al. 2004; Cione et al. 2003, 2009; intervention and to obtain evidence of human Gutiérrez and Martínez 2008: 62, 63; Koch exploitation of different fauna. Consequently, and Barnosky 2006; Martin 1975, 1984; Sodhi these data can be used in the discussion of et al. 2009) as it coincided with the rise of the megafauna extinction in order to understand temperatures and the expansion of Homo humans’ impact upon the native sapiens into the continent. The southern palaeoenvironment during the Pleistocene- portion of South America provides an excellent Holocene transition in South America. Prior to case study to evaluate the influence of humans discussing this argument in full, it is important on these , as this is the last region to provide a brief history of the Collection as where humans migrated and where an well as the background of the material. autochthonous fauna developed. The last factor was the result of the previous isolation of this continent, until 3 MA when South America connected with North America. These History of the Collection native fauna, unique in the world, were characterised by their size, i.e., >1000 kg for The collection of material was gifted to megamammals or 44-999 kg for macro- Valencia City in 1889 by the engineer Jose , and survived until Pleistocene- Rodrigo Botet who obtained it from the Holocene transition, when humans appeared collector, Enrique de Carles (Belinchón et al. in the region (Cione et al. 2003, 2009). 2009: 31 to 41). De Carles was a naturalist from the Museum of Natural Science of

Karina Vanesa Chichkoyan – [email protected] © Chichkoyan 2013 University of Buenos Aires © assemblage 2013 CHICHKOYAN

Buenos Aires and undertook excavations in the analysis of the materials. At times it can including those in De La Plata, Salado and be difficult to understand the utility of Samborombon Rivers and Samborombon Bay analysing bones housed in museums that lack in the northern part of the Pampean region, context. For this research, a different point of Argentina (Figure 1). perspective was necessary, in which museums are considered the first essential part of an investigation. Although partial, the infor- mation recollected here is not only useful because it provides additional data contrib- uting to current discussions, but also aids in creating new research lines for the future. Lastly, in the case of bones with cut marks, these materials can be subjected to radio- carbon dating, allowing for a chronological framework of human intervention to be obtained.

Antecedents of the region

Argentina is located in southern South America, one of the biggest land masses in the Southern Hemisphere. The country has two distinct geographical areas: the lowlands to the east, with different vegetation mosaics, and the Andean Mountain range to the west, with marked altitudes (Figure 1). Due to Argentina’s Figure 1 Map of Argentina in the southern unusual shape, it has a great diversity of part of South America; 1: Andean Mountain biomes and climates that allows for different Range to the west; 2: lowlands to the east; A: adaptations to the environment (Argollo the provenance of the Rodrigo Botet 2006). During most of the Cenozoic, this Collection (the Pampean region); and B: the continent was an island, until 3 MA when it location of some archaeological sites joined with North America through an discussed in the text (image courtesy of intercontinental bridge, the Parana Isthmus. Instituto Geográfico Nacional, Argentina). This occurrence was one of the largest biogeographical events in the Americas. This All of the macromammals Darwin recognised event led to what was called the Great in the Beagle’s historical journey appear in the American Faunal Interchange, when various Rodrigo Botet Collection (Belinchón et al. , from both continents, migrated north 2009). The Collection held in Valencia, and south (Patterson and Pascual 1968; Potts considered the first European palaeontological and Behrensmayer 1992: 462, 463; Webb museum (Belinchón et al. 2009), and, 1978). although most of the material requires taxonomic re-identification, at least six orders The previous isolation of South America of mammals and megamammals are conclu- allowed the development of an indigenous sively present: Carnivora, Notoungulata, fauna of great variety, quite different in shape Litopterna, Perissodactyla, Proboscidea and and size to what had emerged on other Xenarthra (Martel San Gil and Aguirre continents. Among these fauna were the Enríquez 1964). When this material was Xenarthra, one of the earliest American orders originally excavated, the methodology of fossil of placental mammals that during the Great extraction was markedly different from that of American Faunal Interchange, extended to the today. There was no stratigraphical control, no north, while many carnivores, such as ursids systematic recording materials found in each and felids, entered into the south (Potts and stratum, or even a detailed description of the Behrensmayer 1992: 462, 463). Some site and the surrounding area. At the present skeletons of these native species can be seen in time, these kinds of procedures are de rigueur, Figure 2, while Figures 3, 4 and 5 depict ideal and it is impossible to do an excavation reconstructions of these native fauna. without a strict protocol. Further, every detail is taken into account, registered and processed

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Figure 2 Skeletons on display in the Museum of Natural Science of Valencia: A. Macrauchenia patachonica, B. Megatherium americanum, C. A type of Mylodontidae, D. A type of Glyptodontidae (photographs by Chichkoyan).

Within this complex migratory event, The Pampean region is crucial not only Argentina was located in a critical sector, because of its location, but also because it has amongst the Guayano-Brazilian and Andean- the best known South American faunal Patagonian sub-regions, which were home to a records, which have defined the stratigraphy variety of fauna (Morrone 2004). The key area for the whole continent (Patterson and Pascual within this system was the Pampean region 1968). This material, accumulated under arid (see Figure 1), a transitional area between or semi-arid environmental conditions, these two great biomes. Thus, this region has accompanied the deposition of sediments always been particularly susceptible to during much of the Quaternary (Prado and changes, in both flora and fauna, which tend to Alberdi 2010). Further, this region is expand and contract in relation to the characterised by having some of the earliest fluctuating climate (Prado and Alberdi 2010). evidence of human-megafauna interaction.

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Existing evidence of Human- Megafauna interaction

Megafauna bones, in association with human activities or with evidence of anthropic intervention, have been found in various sites in southern Province of Buenos Aires, located to the southeast of the Pampean region (see Figure 1). For the purposes of this article a selection of these sites are described below:

La Moderna: Located on the banks of Azul Creek and dated between 7500 to 7000 BP (Gutiérrez and Martínez 2008). Hunting and processing of clavicaudatus took place here, as inferred by the association

Figure 3 Reconstruction of Macrauchenia between lithic instruments and the remains of patachonica (image by Mauricio Anton, from the (Politis and Gutiérrez 1998). The Belinchón et al. 2009; reproduced with animal would have been quartered here and permission of publisher). the front and rear quarters would have been transported elsewhere. Some axial remains, i.e., the skull and the shell, were left in the site. This evidence corresponds to a ‘Kill Butchery Stage’ (Lyman 1994: 300) in which there is a brief human occupation and restricted butchery activities take place on site.

Arroyo Seco 2: Located to the south of La Moderna and dated between 12150 to 11200 BP, based upon multiple determinations from megafauna bones (Steele and Politis 2009). In the lower layers of this archaeological site, there was evidence of human exploitation of Megatheium americanum as well as two types Figure 4 Glyptodon, Neosclerocalyptus and of horses: Equus neogeus and Hippidion sp. Eutatus (image by Mauricio Anton, from The front and rear quarters from these animals Belinchón et al. 2009; reproduced with as well as bones with possible traces of human permission of publisher). intervention were found at the site (Politis 1989).

Campo Laborde: Located along the banks of Talapaque River and dated between 8700 to 7750 BP (Gutiérrez and Martínez 2008). This site was interpreted as being the location of hunting and primary processing (i.e., evidence of rib bones with cut marks) of Megatherium americanum (Gutiérrez and Martínez 2008). Following processing, only meat and viscera were taken away, leaving the majority of animals’ bones on site (Politis and Messineo 2008).

Paso Otero 5: Located at the right margin of the Quequen River and dated from 10450 to Figure 5 Ideal reconstruction of a Pampean 10200 BP (Gutiérrez and Martínez 2008). At landscape with Megatherium americanum, this site burned bones of Megatherium Equus, Prosbocioidea and Glyptodon (image americanum and Hemiauchenia sp. were by Mauricio Anton, from Belinchón et al. found and interpreted as fuel material. 2009; reproduced with permission of publisher). Cueva Tixi: Located in the Tandilia Mountain System, 40 km from the Atlantic coast and

-32- assemblage dated between 10400 and 10000 BP. Bone material larger than 2cm of size was (Gutiérrez and Martínez 2008). Three selected for observation with magnifying glass, phalanxes and six osteoderms of Eutatus and binocular microscopy, utilising an seguini were found in association with lithic Olympus SZ-PT. Full taxonomic and tools and a hearth. This association of anatomical identification was not always material was interpreted as evidence of an provided for the material and the degree of anthropic intervention over this species identification available has been categorised as (Mazzanti and Quintana 1997; Politis and follows: Gutiérrez 1998). Identified: material was assigned to a definite The date range for these sites is indicative of a taxa and anatomical level; 3000-4000 year period of coexistence for ID: material identified at anatomical level but Homo sapiens and the native South American taxonomical identification below class not fauna (Politis and Messineo 2008: 111). possible; or Although there is some evidence of II: material neither identified at anatomical exploitation of megafauna in the south of the nor taxonomic level. Pampean region, the small guanaco (Lama guanicoe) is still considered to have been the most important animal resource (Cione et al. 2009; Politis et al. 2004). This scenario is General Results different to what happens in Patagonia for example. In this region an opportunistic In total, 11466 bone elements were analysed: behaviour was defined in relation to the 4474 elements were identified at species, exploitation of the megafauna (Borrero and genus, family or order level, 693 elements were Franco 1997). The analysis of bones housed in identified only by anatomical level (i.e. ID) and museums can contribute new evidence to 6299 fragments were unidentifiable (i.e. II) evaluate, clarify, and specify the sort of (Figure 6). impacts that human populations would have introduced. Homo sapiens must be seen as a new invader species (Lanata et al. 2008), that dispersed into the American palaeo- environments, which were not used to this novel predator (Cione et al. 2009).

Methodology

Taphonomical analysis of the Rodrigo Botet Collection was undertaken to examine the Figure 6 Percentage of the material analysed. fossil bones. These analyses allow for the identification of cut marks in animal bones At least 19 bone elements, 0.17% of the total, (i.e., from human and non-human agents), as exhibited evidence of human intervention well as the detection of weathering by multiple elements (see Table 1). Of the 52.6% of bone factors, which can be indicative of the palaeo- elements that were taxonomically identified, and post-depositional environment five taxa are represented: Eutatus, (Bonnichsen 1989; Lyman 2008). Glyptodontidae (Figure 7), Macrauchenia patachonica (Figure 14), Megatherium (Figure Much has been written on how to identify cuts 13), and Mylodontidae (Figures 8, 9, 10, 11, & of human origin and to what part of the 12). The rest of the material is distributed butchery process they correspond (Binford among three ID elements: one rib fragment, 1981; Lyman 1994; Shipman 1981; Walker and one shaft, and one fragment of mandible from Long 1977). For the purposes of this study a non-identified species, and six II elements. human traces (full methodology details in Three observations can be highlighted: the Chichkoyan 2011) were classified as cut marks relative age of cut marks (i.e., in relation to for simple cuts, percussion when the trace was other evidence of weathering on the more profound, and tools for objects fragments), the taxonomical identification of potentially fashioned by humans. the bone upon which cut marks were detected, and the anatomical identification of the bone upon which cut marks were noted.

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Types of Human Intervention Species Bone Type Instrument Percussion Cut Mylodontidae Rib 1 Mylodontidae Scapula 1 Megatherium sp. Rib 1 Undetermined Mandible 1 Undetermined Rib 1 Undetermined Shaft 1 Undetermined Undetermined 1 Undetermined Undetermined 1 Mylodontidae Atlas 1 Mylodontidae Metatarsal 1 Macrauchenia patachonica Rib 1 Macrauchenia patachonica Rib 1 Undetermined Undetermined 1 Undetermined Undetermined 1 Undetermined Undetermined 1 Undetermined Undetermined 1 Glyptodontidae Osteoderm 1 Eutatus sp. Osteoderm 1 Eutatus punctatus Osteoderm 1 TOTALS 3 8 8

Table 1 Bones with evidence of anthropic intervention.

First, this analysis aids in considering a broad manipulation, or archival of the material post- ‘taphonomic history’ of these fossils (i.e., a excavation, very unlikely (Chichkoyan 2011; chronology of the agents that affected the Chichkoyan et al. 2013). bones). Examination revealed that cut marks detected were inflicted before the intervention of other non-human agents (e.g., roots or trampling). The Mylodontidae atlas (see Figures 9 & 10) displays evidence of latter intervention by non-human agents. The same can be said regarding the Glyptodontidae osteoderm (Figure 7), which exhibits manganese spots that are evidence of post- depositional chemical processes which altered two continual walls that were already affected by human intervention. The Macrauchenia patachonica ribs (see Figure 14), the Mylodontidae rib (see Figure 8) and meta- tarsal (see Figure 12), as well as the ID and II fragments, both the bone and cut marks were observed to have the same superficial colour- ation. This colouration can be interpreted as evidence of a post-depositional environment that uniformly affected the material. This information gives greater weight to the interpretation of the marks being the result of Figure 7 Glyptodontidae (left) and pre-depositional human intervention upon the Eutatus (right) osteoderms (photograph bone. Further, this makes the possibility that by Chichkoyan). these marks occurred during transfer,

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Figure 8 Mylodontidae rib with three percussions (photographs by Chichkoyan).

Figure 9 Mylodontidae scapula with percussion (photographs by Chichkoyan).

Figure 10 Mylodontidae atlas (dorsal face). The circled areas are where cut marks were found (photographs by Chichkoyan).

Figure 11 Detail of Figure 10: Mylodontidae atlas (photographs by Chichkoyan).

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Figure 12 Mylodontidae metatarsal with cut marks in the distal epiphysis (photographs by Chichkoyan).

Figure 13 Megatherium sp. rib with the detail of the distal part where there are percussion marks (photographs by Chichkoyan).

Figure 14 Two Macrauchenia patachonica ribs with cut marks (photographs by Chichkoyan).

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Second, the bones with anthropic intervention cutting these bones in the process of found in the Rodrigo Botet Collection are a butchering are noticeably greater. Addition- representative sample of different megafauna ally, the ribs and atlas are related to the first species which became extinct at the time of the steps in butchering, in which the separation of Pleistocene-Holocene transition; excepting the skull from the rest of the body generally Eutatus, which became extinct during the leaves marks over the atlas, and the middle Holocene (Krmpotic and Scillato-Yané evisceration process is related to the marks on 2007; Krmpotic et al. 2009). Not surprisingly, the ribs (Binford 1981). At the site of La most of the material exhibiting signs of human Moderna, the ribs and skull were left after the intervention belongs to Mylodontidae, because initial carcass processing was complete (Politis this family was one of the best represented in and Gutiérrez 1998). Additionally, the site of the Collection (i.e., 25.12% or NISP of 1124), as Campo Laborde revealed a rib of Megatherium well as the Pampean region on the whole americanum with what has been interpreted (Miño-Boilini and Carlini 2009). Mega- as an anthropic cut mark (Gutiérrez and therium sp. (see Figure 13, rib), Macrauchenia Martínez 2008). When butchering large patachonica (see Figure 14, ribs), and animals, often the initial goal is to cut and Glyptodontidae (see Figure 7, osteoderms) transport ‘clean muscles’ to another locale, represent other taxa abundant during the leaving as many bones as possible where the Pleistocene that did not survive beyond the animal was slaughtered. Though a great deal of initial Holocene and which displayed indi- cutting can be involved in dismembering, the cations of human intervention. Megatherium long bones of the appendicular skeleton are americanum is considered to be the most covered by a significantly larger amount exploited species south of the Pampean region, muscle mass which offers up fewer possi- as is evidenced by finds at sites such as Arroyo bilities for the lithics to come in contact with Seco (Steele and Politis 2009), Campo Laborde the bones (Borrero and Martin 2012). The (Gutiérrez and Martínez 2008), and Paso presence of cut marks on the metatarsal of Otero 5 (Gutiérrez and Martínez 2008). There Mylodontidae (see Figure 12) and on one is markedly less evidence of prehistoric indeterminate shaft can be related to the size consumption of Macrauchenia patachonica. of the animal and to the near impossibility of Thus the identification of cut marks on fossils moving the front and the rear quarters without of this species within the Rodrigo Botet further butchering. The percussive marks on Collection provides compelling evidence of the scapula from Mylodontidae (see Figure 9) Macrauchenia patachonica interaction with can be also related to the detachment of limbs Homo sapiens. Examination of the from the rest of the body. Glyptodontidae osteoderms revealed fine striation marks on the walls. These marks were It is worth noting something about the interpreted as evidence of human intervention probability of scavenging of animals hunted by as this kind of work must be done on fresh other carnivores. The faunal population of the material, as post-mortem the bones rapidly Pampean region was characterised as lose collagen becoming brittle. Despite these unbalanced, with greater variation in and observations, more information is necessary in quantities of herbivores than carnivores order to interpret and understand how (Fariña 1996). Due to the size of the native osteoderms were manipulated by humans herbivorous fauna, only the carnivorous (Chichkoyan 2011; Chichkoyan et al. 2013). Smilodon populator, and later Panthera onca, Puma concolor, and Canis nehringi, were Finally, we can focus on the anatomical considered capable of preying upon them elements in which these anthropic traces were (Prevosti and Vizcaíno 2006). The size of these found. The axial skeleton is the most abundant herbivorous megafauna made them group within the sample, and included five ribs particularly unattractive for hunting by and one atlas. The appendicular skeleton is carnivores, and consequently there were fewer represented by one indeterminate shaft and a chances for them to be attacked. Thus, the Mylodontidae metatarsal. A fragment of possibility for humans to have scavenged over Mylodontidae scapula and a mandible frag- previously hunted animals is reduced. Lastly, ment of unknown taxon were also found. The there are notably no identifiable bones of rest of the material corresponds to the six II carnivores within the Rodrigo Botet Collection. fragments mentioned before and to the three osteoderms previously described (see Figure 7). It can be supposed that bones from the axial skeleton more commonly display cut marks as they represent those parts closest to the skin. Consequently, the chances of a lithic

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Discussion possibility of a high reproductive rate, could have influenced the fact that this animal The evidence found in the analysis of the survived well into the Holocene like the Rodrigo Botet Collection is significant as it previously mentioned guanaco. indicates human populations might have had substantial access to megafauna in the The South American megafauna were adapted Pampean region. However, in order to to live in open areas with cold environments, understand the importance of these findings, it as were characteristic of the Pampean region is necessary to consider them within the during the Pleistocene. A series of interglacial context of human-megafauna interaction. It is and glacial periods demonstrated the ability of essential to take into account the general these animals to adapt to changing environ- palaeoecological conditions and relationships mental conditions as the habitats of these that developed during the Pleistocene- animals contracted and expanded along with Holocene transition, and how Homo sapiens’ the fluctuating climatic conditions. Yet, the incursion into this region played out. arrival of Homo sapiens during the last interglacial could have unbalanced the natural The South American megafauna were adaptive strategies of megafauna. Upon predominantly K-selection strategy species arrival, humans found different megafauna (MacArthur and Wilson 1967), characterised species isolated in shrinking biomes, serving as by low population density, few offspring per ecological shelters, due to rising temperatures. individual, late sexual maturity and long The area from where the Collection comes gestation periods. K-strategists tend to have could have acted as an ecological shelter. The few newborns with low survival rates, making Pampean region was an open space which their populations especially vulnerable during maintained the arid climatic conditions of the times of palaeoclimatic fluctuation (i.e., the Pleistocene that facilitated the survival of these Pleistocene-Holocene transition) (Cione et al. megamammals. When the human population 2009). In light of these characteristics, human arrived, they most likely found this native predation upon these animals might have fauna already reduced to their minimal viable noticeably affected the density of native number for survival. Humans might have megafauna populations. Effects need not be incorporated them as a novel resource and immediate however; as there is also evidence, though exploitation might have been sporadic, as noted previously, that Homo sapiens and this new predator probably had a negative native fauna coexisted for at least 3000-4000 impact on the natural survival adaptations of years. Nevertheless, the palaeoenvironmental the megafauna. changes for the region combined with some utilisation of megafauna by humans might The invasive character (sensu Lanata et al. have eventually negatively impacted the native 2008) of the Homo sapiens populations in the fauna. different palaeoenvironments implies a differential usage of the diverse resources in It is important to note that these animals were the process of dispersion through the isolated not subjected to human predation or biomes. In this way, the Rodrigo Botet technology when they evolved on the South Collection can be compared to evidence from American continent. As is commonly witness- south of the Pampean region. In general, ed in alien-native interactions (Kondoh 2006), similarities with the exploitation of Mega- the megafauna probably were slow to develop therium and Eutatus can be detected; the defensive or evasive behaviours in order to former was consumed in Arroyo Seco (Politis adapt to a new threat. The megafauna, which 1989) and Campo Laborde (Gutierrez and lumbered across the landscape, were an easy Martinez 2008), and utilised for fuel in Paso target for the new human population, Otero 5 (Gutierrez and Martinez 2008); while equipped with lithic technology (Cione et al. Eutatus was recorded at Cueva Tixi (Gutierrez 2009; Gutiérrez and Martínez 2008; Patterson and Martinez 2008). Only Campo Laborde and Pascual 1968: 447). The majority of taxa presents clear evidence of cut marks on a identified in the Rodrigo Botet Collection fit Megatherium rib. Though megafauna were the K-strategist paradigm (i.e., Mylodontidae, utilised in both regions, in the south, Macrauchenia patachonica, Megatheium sp., Megatherium was the most exploited animal; and Glyptodontidae). In contrast, Eutatus, however, to the north, Mylodontidae was the although a Pampean species exploited by most important taxon as evidenced by the humans, perhaps could have survived for a Rodrigo Botet Collection. In contrast to what greater period of time as it was not a K- happened in the Pampean region, in Patagonia strategist. Eutatus’ reduced size, weighing less a lower encounter rate for humans and than 50 kg (Krmpotic et al. 2009), and the megafauna was postulated. The archaeological

-38- assemblage record of megafauna and human interaction is Thus, when human populations entered the scarce and some assemblages do have region, they found a native fauna under carnivore marks. It is proposed that the different types of stress, probably grouped in megafauna were opportunistically scavenged regions more favourable for their survival (e.g., by humans (Borrero and Franco 1997; Borrero the Pampean area). In the Pampean region, and Martin 2012). Consequently, it is cold and dry conditions allowed large animals important to take into consideration the to survive and be exploited by humans. diverse impacts that Homo sapiens could have produced over the populations of native fauna. The exact role Homo sapiens played in the Homo sapiens’ distribution and diversity extinction of South American megafauna might have been related to the characteristics should be investigated further. One of the key of the different palaeoecological patches, and issues is the dearth human traces on bones, this would have influenced human actions which is the most direct way to assess the within each of them. For example, it can be degree of anthropic exploitation. Most con- seen that there are some similar trends in both clusions have been made by cultural areas of the Pampean region, although the association in controlled excavations. The variability of the species consumed probably limited evidence of cut marks may be a result depends on the different abundance of the of the massive size of these animals or due to native population in each patch. In contrast, in the carcasses subsequent usage as fuel. Patagonia, fewer relations between humans and megafauna were registered overall. This It is evident that the climate of the Pleistocene- trend can be connected to the different Holocene transition played an important role palaeoenvironments of the regions as well as in the distribution and depletion of these the environmental niches of the particular animals. Megafauna populations would have taxa. been at minimal viable numbers when human groups entered the Pampean region. Further, The material found in the Rodrigo Botet the different regions of the landscape would Collection adds weight to the premise that have allowed a heterogenic rate of invasiveness Homo sapiens influenced the survival of the of humans onto the continent, according to the South American native megafauna in the diversity of resources present in each Pampean region in a negative way. This impact ecological patch. could have been facilitated by the shrinking ecological niches of these species as well as by For the future, regional studies are necessary the climatic fluctuations of the period. The in order to amass a body of comparative data exploitation of some animals in a population from palynological, sedimentological, and that was already in decline due to climatic archaeological studies. Further zooarchaeo- changes was potentially enough to lead them logical analyses must be undertaken to to extinction. The possibility of scavenging by understand the processing chain of these humans is temporarily rejected due to the lack megamammals. When the extinction of South of evidence of carnivores in the Rodrigo Botet American mammals is analysed, it is Collection. important to consider and account for the differences between genera and species involved and their distribution over the landscape. It can no longer be postulated that Conclusion the whole process did not have palaeoregions and species differentiation. The causes of megafauna extinction are far from clear. Positions as to whether extinction The discussion of megafauna extinction at the was caused by humans and/or climatic Pleistocene-Holocene transition is still not fluctuations are prevalent in the literature finished. Further research must be done in the (Borrero and Franco 1997; Brook and Bowman Pampean and in surrounding regions to delve 2004; Cannon and Meltzer 2004; Fiedel and into all aspects of the consumption of the Haynes 2004; Grayson and Meltzer 2003; megafauna. A more detailed comprehension of Haynes 2007; Koch and Barnosky 2006). landscape changes due to Homo sapiens According to the position taken in this intervention will be obtained in this way. investigation, the South American megafauna would have been declining when Homo sapiens arrived in the Pampean region. Climatic fluctuations already detrimentally affected the reproduction and development of megafauna species during the Pleistocene.

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Acknowledgements Borrero, L. A. and Martin F. M.. 2012 Ground sloths and humans in southern Fuego- This paper is part of my Masters Degree Thesis Patagonia: taphonomy and archaeology. that was completed thanks to the Erasmus World Archaeology 44 (1): 102-117. Mundus Scholarship between 2008 and 2010. I want to acknowledge my thesis Director and Brook, B.W. and Bowman, D.M.J.S. 2004. The Co-directors: Bienvenido Martinez-Navarro, uncertain blitzkrieg of Pleistocene megafauna. Margarita Belinchón and Jose Luis Lanata for Journal of Biogeography 31: 517-523. their advice and support. The material was analysed between April and May of 2010, Cannon, M.D and Meltzer, D.J. 2004. Early thanks to the Museum of Natural Science of Paleoindian foraging: examining the faunal Valencia. I want to thank Miquel de Renzi for evidence for large mammal specialization and his advices in the material’s analysis; the regional variability in prey choice. Quaternary different institutions that gave me the space Science Reviews 23: 1955–1987. and opportunity to develop this work: to the University of Ferrara, specially to Marta Chichkoyan, K. V. 2011. Grandes Mamíferos Arzarello, to Universitat Rovira i Virgili, in del Sur: Extinciones Sudamericanas y la particular Robert Sala and the different Colección Rodrigo Botet del Museo de investigators from the Institut Català de Ciencias Naturales de Valencia, España. Paleoecologia Humana i Evoluciò Social [Online] Colección Tesis. IIDyPCa-UNRN- (IPHES). A special acknowledgment to my CONICET. Bariloche. ISBN 978. friends in Spain: Graciela Fernandez and Jorge http://iidypca.homestead.com/PublicacionesI Szigety, for their support and company when I IDyPCa/Chichkoyan/Chichkoyan.html. was there. Chichkoyan, K.V., Belinchón, M., Lanata, J. L. and Martínez-Navarro, B. 2013. La Colección Rodrigo Botet y la extinción de la megafauna Bibliography en la Región Del Plata (Argentina). Paper presented in I International Congress of Archaeology of Del Plata Basin. Argollo, J. 2006. Aspectos Geológicos. In: Moraes, M.R., Øllgaard, B., Kvist, L.P., Cione, A.L., Tonni, E.P. and Soibelzon, L. Borchsenius, F. and Balslev, H. (eds.). 2003. The Broken Zig-Zag: Late Cenozoic large Botánica Económica de los Andes Centrales. mammal and tortoise extinction in South La Paz: Mayor University of San Andrés, pp1- America. Revista del Museo Argentino de 10. Ciencias Naturales 5 (1): 1-19.

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