ARTICLE IN PRESS

Quaternary International 185 (2008) 69–74

Extinction of Pleistocene megamammals in South America:

The lost evidence
Luis Alberto Borrero
CONICET-IMHICIHU-DIPA, Saavedra 15, Piso 5, 1083 Buenos Aires, Argentina
Available online 5 November 2007

Abstract

The extinction of the Pleistocene megamammals is one of the oldest unsolved questions of South American natural history. Data from
this continent are rarely considered in synthesis papers and books produced in the Northern Hemisphere, and sometimes there is a
claim that no adequate data can be found. The main point of this presentation is that there is a wealth of both paleontological
and archaeological South American data which may help in the evaluation of the different theories and can be used to interpret the
extinction event.
r 2007 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction of the dispute reaches the review papers produced in the

South American data concerning the relationship
between humans and megamammals are important, since
this continent witnessed one of the major extinction events
that occurred at the end of the Pleistocene (Martin and
Steadman, 1999; Cione et al., 2003). Dozens of species of
megamammals went extinct in South America ca.
10,000 BP, at least 1000 radiocarbon years after the human
colonization of the continent started (Lavallée, 1995;
Martinez, 2001). This period immediately followed a pulse
of very cold climate, partially coincident with the Antarctic
Cold Reversal in the southern part of the continent
(Sugden, 2005). Under these conditions, the classic dispute
between the competing hypotheses used to explain those
extinctions in other continents also takes place in South
America. Some researchers support human hunting as a
crucial cause (Cione et al., 2003), others favor climatic
change (Ficcarelli et al., 1997) and there are even some
defending the action of pathogens (Ferigolo, 1999). A
negative test of the hypothesis that maintains that there
was a differential extinction of southern stock faunas as
compared with northern stock faunas that migrated to
South America was published (Lessa et al., 1997).
Unfortunately, with few exceptions (i.e., Fiedel and
Haynes, 2004), very little of this South American aspect
E-mail address: dipa.imhicihu@conicet.gov.ar
Northern Hemisphere (Hubbe et al., 2007). Sites, analysis
and ideas produced in South America deserve to be taken
into account (Fig. 1).
2. Published record
This paper discusses data that have been published but
not completely used. When Martin and Wright (1967)
published their landmark volume Pleistocene Extinctions:
the Search for a Cause, it was probably difficult to find
somebody to cover the evidence from South America,
because at that time very few relevant sites were studied.
Several years later, when Quaternary Extinctions: a
Prehistoric Revolution was edited by Martin and Klein
(1984), a chapter by Gruhn and Bryan (1984) was the only
one focused in South America, and it was not a review
paper but a case study focused on the Taima Taima site in
Venezuela. More recently, synthetic papers were published
in Science (Barnosky et al., 2004), Proceedings of the
National Academy of Sciences (Steadman et al., 2005) and
Annual Review of Ecology and Evolutionary Systematics
(Koch and Barnosky, 2006), arguably among the most
important scientific publications in the world. These papers
are important contributions to the extinctions debate, but
still their treatment of the South American data is
incomplete. This situation signals a panorama that needs
to be rectified.

1040-6182/$ - see front matter r 2007 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2007.10.021
 ARTICLE IN PRESS
70 L.A. Borrero / Quaternary International 185 (2008) 69–74
Fig. 1. South America. Archaeological sites mentioned in the text.
3. Discussion
3.1. Sloths, mastodons and extinction dates
The work by Steadman et al. (2005) is a good example of
the discussion of the extinctions of megamammals based
on Last Appearance Dates (LADs), which are used as a
proxy for ‘‘extinction dates’’ of the different taxa (Stead-
man et al., 2005, p. 11763). These authors have to be
careful in selecting their data, and so they choose to reject
standard radiocarbon dates unless they are replicated by
AMS dates. This is a sound and necessary procedure in
order to obtain robust chronologies. One problem is that
when it comes to South America they have not analyzed all
the existent evidence before reaching their conclusions. For
example, they claim that, ‘‘The youngest reliable dates on
dung or tissue of extinct sloths have means of ca.
10,600–10,200 BP’’ (Steadman et al., 2005, p. 11765).
However, in their summary of available radiocarbon
dates for sloths in South America, several sites are missing,
and not all of them lack AMS dates for sloths. For
example, the open air site Arroyo Seco 2 which produced
AMS and standard Late Pleistocene and Early Holocene
radiocarbon dates for sloths (Politis and Madrid, 2001).
The Early Holocene dates for Arroyo Seco 2 are rejected by
the excavators (G. Politis, personal communication), but
they accept Early Holocene dates for sloths at Campo
Laborde (Messineo et al., 2004; Politis et al., 2004) and for
glyptodonts at La Moderna (Politis and Gutiérrez, 1998).
Three samples on sloth dung and ossicles from Gruta del
Indio also have Early Holocene dates (Garcı́a and Lagiglia,
2000). The Holocene survival of sloths—or glyptodonts—
should not be taken as a fact. However, a thorough
consideration of the stratigraphy, materials and dates of La
Moderna, Campo Laborde or Gruta del Indio is needed to
argue that the case of Holocene survival of megamammals
is not robust. Steadman et al. (2005, p. 11765) are ready to
accept Holocene dates for sloths in Cuba, in spite of
‘‘inadequate field notes and subsequent damage to the
sites’’. They even accept ‘‘a single whole-bone dateyon an
assortment of sloth bones’’ (p. 11766). Without comment-
ing on the evidence from Cuba, it is apparent that these
authors are using a different standard here.
In addition to the evidence concerning extinction dates,
one of the goals of Steadman et al. is to maintain that the
timing of extinction is synchronic with that of human
peopling, thus implicating humans as causal agents of
extinction. In terms of this goal, they made some poor site
choices. It might be suggested that this is derived from their
selection of sites characterized by sloth dung accumula-
tions, but the authors are not restricted by this criteria
(Steadman et al., 2005, p. 11765). An example is the
uncritical use of the evidence from Gruta del Indio,
Mendoza, Argentina, even when the quality of archae-
ological research associated with that site is not very high.
Accumulations of dung attributed to Megatherium sp. and
Mylodon sp. were found physically associated with flakes.
Examination of the papers by Semper and Lagiglia (1968)
and Lagiglia (2001), the original excavators of the site,
reveals the poor quality of the evidence, the difficulties in
understanding the provenance of the lithic samples and
other problems. The use of dynamite during the process of
excavation does not constitute grounds for direct dismissal
of the site, but it certainly indicates the need for careful
scrutiny of the evidence before using it. Due to these
interpretative difficulties, a single pollen column from
Gruta del Indio has produced completely different inter-
pretations (Lagiglia, 1975; D’Antoni, 1983; Mancini et al.,
2005). Only the most recent research by Garcı́a (2003) can
be used, and Garcı́a himself—whose goal was to make
some sense of the information from this site—is at pains to
understand the sequence. In an effort to suggest that the
evidence from Gruta del Indio is useful, Steadman et al.
(2005, p. 11764) claim that, ‘‘No charcoal was found with
or beneath the sloth dung’’. But it is also true that lithic
artifacts were found in the lower layers, dated ca.
25,000–435,000 BP (Garcı́a, 2003). Vertical migration of
flakes is accepted by the excavator (Lagiglia, 2001) and it is
indeed a plausible explanation for their presence within a
long sequence of sloth dung. Then, why choose this site?
Perhaps it was because the site is important and well
known, thanks to the fact that the preservation of bones
and soft tissues is excellent, and also because North
American researchers were involved in its study. It must
be clearly stated that Paul Martin and Austin Long were
not involved in the excavation problems of Gruta del
 ARTICLE IN PRESS
L.A. Borrero / Quaternary International 185 (2008) 69–74 71

Indio, and in fact their chronological work (Long et al., Table 1

1998), together with the recent efforts of Garcı́a (2003) Rating system (Mead and Meltzer, 1984, p. 442)
constitutes the only hope for the understanding of that Score
complex site. That is one reason why the Early Holocene
dates from this site need to be considered, since they are Material dated
bona fide taxon dates. However, it is clear that the quality Derived from extinct fauna
Collagen 5
of the contextual evidence is very poor, and that not much
Body perishables (dung, hide, hair) 5
light on human association can come from Gruta del Indio Apatite 3
without careful scrutiny of its stratigraphy and research Whole bone 1
history.
Derived from other organic material
Another bad choice is Monte Verde. Barnosky et al. Charcoal (elemental) 6
(2004, supporting text, p. 5) wrote that, ‘‘In South Wood 5
America, evidence for utilization of extinct megafauna by Peat 3
humans exists at a few sites’’, and then quote the paper by Organic mud 3
Soil 3
Meltzer et al. (1997). However, the presence of mastodon
Shell 2
(Cuvieronius hyodon) at this site is only indicative of the Terrestrial carbonate 1
existence of that taxon in the area, and its relationship with
Strength of association
humans is only secondary. Those mastodon bones were
Strong 3
collected by prehistoric people from defleshed carcasses, Unknown or medium 2
since the dirt found embedded in the bones was not local Weak 1
(Karathanasis, 1997). Although this interaction between
humans and mastodons is exciting, Monte Verde is a bad
choice of a site to study the role of humans in the extinction Table 2
of megamammals. Radiocarbon dates, Tres Arroyos 1 (Borrero, 2003; Massone and Prieto,
In discussing how the LADs for ground sloths track the 2004)
first arrival of people, a claim for the extinction of an Lab Date (BP) Material
undescribed species of ground sloth identified through
DNA at Cuchillo Curá, Neuquén, is quoted (Steadman et Dic-2732 10,2807110 Camelide bone
Dic-2733 10,4207100 Terrestrial mammal bone
al., 2005). Two radiocarbon dates of 14,6657150 BP (Ua-
Beta-20219 11,8807250 Terrestrial mammal bone
13871) and 13,7507230 BP (GX-21149) (Hofreiter et al., Beta-101023 10,600790 (AMS) Charcoal, hearth 2
2003) are available. This is a fine piece of research that Beta-113171 10,580750 (AMS) Charcoal, hearth 3
suggests the extinction of one species of ground sloth OXA-9245 10,575765 (AMS) Dusicyon avus
before human arrival. The potential significance of this OXA-9246 10,630770 (AMS) Vicugna sp.
OXA-9247 10,685770 (AMS) Hippidion saldiasi
finding for the relationship between humans and the
OXA-9248a 11,085770 (AMS) Panthera onca mesembrina
extinction of ground sloths is not discussed. OXA-9666 10,1307210 (AMS) Charcoal, hearth 4

3.2. The rating system of radiocarbon dates Uncalibrated radiocarbon years.

a
Below human occupation layer.

Barnosky et al. (2004, supporting text, p. 13), in a paper

that supports the view that humans were a critical cause of
the Late Pleistocene extinctions, sustain that, ‘‘The South
America datesyneed detailed evaluation before they can
be considered robust’’, and that this evidence ‘‘awaits
critical analysis’’ (p. 74). However, in their review they
basically quote secondary sources, instead of the original
papers. It can be sustained that the use of the original
sources constitutes the best way to conduct ‘‘critical
analysis’’.
Using Mead and Meltzer’s (1984, p. 442) criteria of
association—a rating system that considers useful only
those cases with grades 8 or 9 (Table 1)—they analyze the
radiocarbon dates and the strength of their association
with megamammals in several regions. This system
evaluates radiocarbon dates on the basis of the material
being dated, with charcoal, bone collagen, body perish-
ables, such as dung or hair, and wood providing reliable
results. The system also considers the strength of the
association on the basis of stratigraphic and contextual
information (Mead and Meltzer, 1984, pp. 441–442). From
the application of this rating system, Barnosky et al. (2004,
supporting text, p. 13) conclude that, ‘‘Very few, if any, of
the South American dates reported in the literature would
receive a grade of 8 or 9’’.
Thus, the conclusion is that no reliable information is
contributed by South American sites. However, at several
sites, for example, Tres Arroyos 1 and Tierra del Fuego,
there are several concordant AMS dates made on charcoal
(Massone and Prieto, 2004) (Table 2). These qualify as a 6
in Mead and Meltzer’s rating system.
The association is as good as they come—small circular
discrete hearths with taxonomically determined megamam-
mal bones which are broken, cut-marked and burnt. Dates
on the bone collagen of different species are also
concordant with those from the hearths. In addition, these
 ARTICLE IN PRESS
72 L.A. Borrero / Quaternary International 185 (2008) 69–74

bones—Hippidion saldiasi, Vicugna sp.—were found in Table 5

physical association with projectile points and other Radiocarbon years, Piedra Museo (Miotti et al., 2003)
instruments—which qualifies as a 3. Only one standard Lab Date (BP) Material
date—Beta-20219—can be considered as an outlier. Thus,
this is a South American site with a grade of 9. Moreover, LP-949 92307105 Lama guanicoe bone
this is not an isolated case. Other examples of sites which LP-859 97107105 Lama guanicoe bone
AA-8428 10,400780 (AMS) Camelidae bone
provide abundant chronological evidence for Pleistocene
OXA-9249 10,470760 (AMS) Charcoal
mammals and human activities include Cueva del Medio GrA-9837 10,470765 Charcoal
(Nami and Nakamura, 1995) (Table 3), Cueva Lago Sofı́a OXA-8527 10,390770 (AMS) Lama guanicoe bone
1 (Prieto, 1991) (Table 4) and Piedra Museo (Miotti et al., OXA-8528 10,925765 (AMS) Hippidion saldiasi bone
2003) (Table 5). Thus, the evidence indicates that there are AA-27950 11,000765 (AMS) Charcoal
AA-20125 12,890790 (AMS) Charcoal
sites in South America with a grade 8 or 9. OXA-9509 9950775 (AMS) Charcoal
But, is this ranking of dates the best criterion? It is a OXA-9508 93507130 (AMS) Charcoal
good approximation, but one that needs to be updated (D. OXA-9507 10,1007110 (AMS) Lama sp. bone
Meltzer, personal communication). At Tres Arroyos, there
Uncalibrated radiocarbon years.
is a very strong evidence for a behavioral association
between humans and megamammals because there are
good charcoal and bone dates from hearths, all providing
contemporaneous ages. The existence of a taphonomic evaluation of the bones and deposits found at Tres Arroyos
(Borrero, 2003) is another reason to be optimistic about the
character of the association. In other words, contextual
Table 3 information is the basis of that interpretation.
Radiocarbon dates, Cueva del Medio (Nami and Nakamura, 1995) On a more general level, there are taphonomic reasons to
Lab Date (BP) Material
suggest that charcoal dates do not always apply to the
bones or other objects found in the same deposits. Mead
PITT-0344 95957115 Charcoal and Meltzer (1984) make it clear that, if a date was
Beta-40281 9770770 Bone obtained on a bone of any particular species, that date does
Gr.N-14913 10,310770 Charcoal
not apply to other species present in that faunal
Beta-58105 10,3507130 Bone
Beta-52522 10,430780 Charcoal assemblage: ‘‘Our position is predicated on the belief that
Gr.N-14911 10,5507120 Bone it is better to assume an association is weak until proven
NUTA-1811 10,7107100 (AMS) Hippidion saldiasi otherwise’’ (Mead and Meltzer, 1984, p. 442).
Beta-39081 10,9307230 Charcoal This is a position with which I strongly agree. Evidence
PITT-0343a 12,3907180 Burnt bone
for vertical migration of bones caused by a number of
NUTA-2331 10,8607160 (AMS) Hippidion saldiasi
NUTA-2197 11,0407250 (AMS) Lama cf. owenii cultural and non-cultural processes constitutes just one of
NUTA-1734 10,4307100 (AMS) Lama cf. owenii many reasons (Schiffer, 1987; Borrero, 1990). Then,
NUTA-1737 11,1207130 (AMS) Lama cf. owenii independent data are required, and in many cases it is
NUTA-2330 10,9607150 (AMS) Lama cf. owenii better to obtain radiocarbon dates on the bones themselves
NUTA-1735 10,4507100 (AMS) Lama guanicoe
in order to have reliable LADs. In an ideal situation, dates
NUTA-2332 10,7107190 (AMS) Lama guanicoe
NUTA-1812 10,8507130 (AMS) Lama guanicoe on both charcoal and bones should be obtained.
The chronology of human arrival to South America is
Uncalibrated radiocarbon years. also a critical issue. Barnosky et al. (2004, p. 71) state that,
a
Nami and Nakamura do not accept this date.
‘‘In South America, generally accepted dates place humans
in coastal Chile and Patagonia at 12.9–12.5 ka BP’’.
However, there is not really a consensus on the age of
Table 4 human settlement in South America, since most of the
Radiocarbon dates, Cueva Lago Sofı́a 1 (Prieto, 1991; Massone and
dates older than 12,000 BP are isolated outliers in strata
Prieto, 2004)
usually dated between 11,000 and 10,000 BP, such as at
Lab Date (BP) Material Guitarrero Cave (Lynch, 1980), Telarmachay (Lavallée et
al., 1985), Pachamachay (Rick, 1980), Cueva del Medio
PITT-0684 11,570760 Charcoal
OXA-8635 10,710770 (AMS) Lama guanicoe
(Nami and Nakamura, 1995), Los Toldos (Cardich et al.,
OXA-9319 10,780760 (AMS) Hippidion saldiasi 1973) or Piedra Museo (Miotti et al., 2003). Only at Monte
OXA-9504 10,3107160 (AMS) Hippidion saldiasi Verde are dates of more than 12,000 BP replicated
OXA-9505 10,1407120 (AMS) Pseudalopex culpaeus (Dillehay and Pino, 1997), and this is a site which is highly
A-7283 10,9107260 Charcoal controversial in the archaeological community.
PITT-0939a 12,9907490 Mylodon darwini
Finally, there is an observation that what is required
Uncalibrated radiocarbon years. are, ‘‘Improvements in the chronology of extinction
a
Below human occupation layer. and paleoclimatic reconstructionyfor South America’’
 ARTICLE IN PRESS
L.A. Borrero / Quaternary International 185 (2008) 69–74
(Barnosky et al., 2004, p. 74). Good chronology is indeed
necessary, and this discussion has already shown that some
improvements exist but were not taken into account. It also
remains true that, ‘‘association of artifacts and extinct
megafauna are generally accepted, although the criteria for
accepting association is not consistently specified’’ (Koch
and Barnosky, 2006, p. 236), which constitutes an excellent
reason to analyze each case on its own. Only careful
scrutiny of the available information can support or reject
a case of human association with megamammals. As for
the paleoclimatic evidence, the data base for the last 20,000
years is abundant (i.e., Clapperton, 1993; McCulloch et al.,
2000; Markgraf, 2001; Heusser, 2003; Sugden, 2005), with
most of these sources written in English and published in
Europe or USA. It is a truism that we always need more,
but even more important it is also true that we need to use
what we already have.
4. Conclusions
In the end, there is no basis to sustain the statement that,
‘‘until more comprehensive analyses are undertaken,
little else can be said with certainty’’ (Barnosky et al.,
2004, p. 71), or, in contrast, that we already know the
chronology of sloth extinctions and human dispersal in
South America (Steadman et al., 2005, p. 11765). These
and many other issues are still open to alternative
interpretations. The problem is not so much about the
presence of mistakes in the analyses as the lack of use of
South American data.
The impression is that the review papers criticized here
are fine contributions to the understanding of Pleistocene
extinctions in North America or other continents, but that
their treatment of South America is incomplete. Statements
in favor or against the Holocene survival of megamammals
in South America, for example, should be the result of
careful analysis of the published evidence. It is true that
many sources are only available in Spanish or Portuguese.
However, the solution is not only that this information
becomes available in English, but also that English
speaking authors begin to use what is written in other
languages. Since the overkill position is a crucial compo-
nent of many popularized versions of the history of human
colonization of the continent, some of which picture
humans as destroyers of their resources and environments
(see Grayson and Meltzer, 2003, p. 590), the use of the
totality of the written output becomes a necessity.
Acknowledgments
To Joaquin Arroyo-Cabrales and Eileen Johnson for
their invitation to contribute to this volume in honor of our
mutual friend Oscar Polaco. To David Meltzer for his
comments to the oral presentation of this paper. Also, my
thanks to Ramiro Barberena and Marcia Bianchi for their
help with the map.
73
References
Barnosky, A.D., Koch, P.L., Feranec, R.S., Wing, S.L., Shabel, A.B.,
2004. Assessing the causes of Late Pleistocene extinctions on the
continents. Science 306, 70–75.
Borrero, L.A., 1990. Taphonomy of guanaco bones in Tierra del Fuego.
Quaternary Research 34, 361–371.
Borrero, L.A., 2003. Taphonomy of the Tres Arroyos 1 rockshelter, Tierra
del Fuego, Chile. Quaternary International 109–110, 87–93.
Cardich, A., Cardich, L., Hajduk, A., 1973. Secuencia arqueologica y
cronologia radiocarbonica de la Cueva 3 de Los Toldos (Santa Cruz,
Argentina). Relaciones de la Sociedad Argentina de Antropologı́a 7,
85–123.
Cione, A., Tonni, E.P., Soibelzon, L., 2003. The broken zig-zag: Late
Cenozoic large mammal and tortoise extinctions in South America.
Revista del Museo de Historia Natural Bernardino Rivadavia 5 (1),
1–19.
Clapperton, Ch., 1993. Quaternary Geology and Geomorphology of
South America. Elsevier, Amsterdam.
D’Antoni, H., 1983. Pollen analysis of Gruta del Indio. Quaternary of
South America and Antarctic Peninsula 1, 83–104.
Dillehay, T., Pino, M., 1997. Radiocarbon chronology. In: Dillehay, T.
(Ed.), Monte Verde. A Late Pleistocene Settlement in Chile. Vol. 2.
The Archaeological Context. Smithsonian Institution Press, Washing-
ton, DC, pp. 41–52.
Ferigolo, J., 1999. Late Pleistocene South-American land-mammal
extinctions: the infection hypothesis. Quaternary of South America
and Antarctic Peninsula 12, 279–310.
Ficcarelli, G., Azzaroli, A., Bertini, A., Coltorti, M., Mazza, P.,
Mezzabotta, C., Moreno Espinosa, M., Rook, L., Torre, D., 1997.
Hypothesis on the cause of extinction of the South American
mastodonts. Journal of South American Earth Sciences 10 (1), 29–38.
Fiedel, S., Haynes, G., 2004. A premature burial: comments on Grayson
and Meltzer’s ‘‘requiem for overkill’’. Journal of Archaeological
Science 31, 121–131.
Garcı́a, A., 2003. On the coexistence of man and extinct Pleistocene
megafauna at Gruta del Indio. Radiocarbon 45, 33–39.
Garcı́a, A., Lagiglia, H., 2000. Avances en el estudio del registro
pleistocénico tardı́o de la Gruta del Indio (Mendoza). Cuadernos del
Instituto de Antropologı́a y Pensamiento Latinoamericano 18,
167–174.
Grayson, D.K., Meltzer, D.J., 2003. A requiem for North American
overkill. Journal of Archaeological Science 30, 585–593.
Gruhn, R., Bryan, A.L., 1984. The record of Pleistocene megafaunal
extinctions at Taima-Taima, northern Venezuela. In: Martin, P.S.,
Klein, R.G. (Eds.), Quaternary Extinctions. A Prehistoric Revolution.
University of Arizona Press, Tucson, USA, pp. 128–137.
Heusser, C.J., 2003. Ice Age Southern Andes. A Chronicle of Paleoeco-
logical Events. Elsevier, Amsterdam.
Hofreiter, M., Betancourt, J.L., Sbriller, A.P., Markgraf, V., MacDonald,
H.G., 2003. Phylogeny, diet, and habitat of an extinct ground sloth
from Cuchillo Curá, Neuquén Province, Southwest Argentina.
Quaternary Research 59, 364–378.
Hubbe, A., Hubbe, M., Neves, W., 2007. Early Holocene survival of
megafauna in South America. Journal of Biogeography 34 (9), 1642–1646.
Karathanasis, A.D., 1997. X-ray diffraction, X-ray fluorescence, and
differential scanning calorimetry analysis of sediments from selected
features and bones. In: Dillehay, T.D. (Ed.), Monte Verde. A Late
Pleistocene Settlement in Chile. Vol. 2. The Archaeological Context
and Interpretation. Smithsonian University Press, Washington, DC,
pp. 817–824.
Koch, P., Barnosky, A.D., 2006. Late Quaternary extinctions: state of the
debate. Annual Review of Ecology and Evolutionary Systematics 37,
215–250.
Lagiglia, H., 1975. Primer diagrama polı́nico de la estratigrafı́a
arqueológica argentina. In: Actas y Trabajos del Primer Congreso de
Arqueologı́a Argentina, Rosario, pp. 163–176.
 ARTICLE IN PRESS
74 L.A. Borrero / Quaternary International 185 (2008) 69–74
Lagiglia, H., 2001. El Paleoindio del Atuel en Sudamérica (Análisis de la
cronologı́a absoluta del Paleoindio del Atuel). Notas del Museo de
Historia Natural de San Rafael 48, 5–12.
Lavallée, D., 1995. Promesse D’Amérique. La Préhistoire de l’Amérique
du Sud. Hachette, Paris.
Lavallée, D., Julien, M., Wheeler, J.C., Karlin, C., 1985. Telarmachay.
Chasseurs et Pasteurs prehistoriques des Andes. ADPS-ERS, Paris.
Lessa, E.P., Van Valkenburgh, B., Fariña, R.A., 1997. Testing hypotheses
of differential mammalian extinctions subsequent to the Great
American Biotic Interchange. Palaeogeography, Palaeoclimatology,
Palaeoecology 135, 157–162.
Long, A., Martin, P.S., Lagiglia, H., 1998. Sloths and humans at Gruta
del Indio, Argentina. Radiocarbon 40, 693–700.
Lynch, T., 1980. Guitarrero Cave. Early Man in the Andes. Academic
Press, New York.
Mancini, M.V., Paez, M.M., Prieto, A.R., Stutz, S., Tonello, M.,
Vilanova, I., 2005. Mid-Holocene climatic variability reconstruction
from pollen records (321–521S, Argentina). Quaternary International
132, 47–59.
Markgraf, V. (Ed.), 2001. Interhemispheric Climate Linkages. Academic
Press, San Diego.
Martin, P.S., Klein, R.G. (Eds.), 1984. Quaternary Extinctions. A
Prehistoric Revolution. University of Arizona Press, Tucson, USA.
Martin, P.S., Steadman, D.W., 1999. Prehistoric extinctions on islands
and continents. In: MacPhee, R.D.E. (Ed.), Extinctions in Near Time.
Causes, Contexts, and Consequences. Kluwer Academic/Plenum
Publishers, New York, pp. 17–55.
Martin, P.S., Wright, H.E. (Eds.), 1967. Pleistocene Extinctions: the
Search for a Cause. Yale University Press, New Haven, CT.
Martinez, G., 2001. ‘Fish-tail’ projectiles point and megamammals:
new evidences from Paso Otero 5 (Argentina). Antiquity 75,
523–528.
Massone, M., Prieto, A., 2004. Evaluación de la modalidad cultural Fell 1
en Magallanes. Chungara 1, 303–315.
McCulloch, R.D., Bentley, M.J., Purves, R.S., Hulton, N.R.J., Sugden,
D.E., Clapperton, C.M., 2000. Climatic inferences from glacial and
palaeoecological evidence at the Last Glacial Termination, southern
South America. Journal of Quaternary Science 15, 409–417.
Mead, J., Meltzer, D., 1984. North American Late Quaternary extinctions
and the radiocarbon record. In: Martin, P.S., Klein, R.G. (Eds.),
Quaternary Extinctions. A Prehistoric Revolution. University of
Arizona Press, Tucson, USA, pp. 440–450.
Meltzer, D.J., Grayson, D.K., Ardila, G., Barker, A.W., Dincauze, D.F.,
et al., 1997. On the Pleistocene antiquity of Monte Verde, southern
Chile. American Antiquity 62, 659–663.
Messineo, P., Politis, G., Rivas, M.I., 2004. Cazadores tempranos y
megamamiferos tardios en la region pampeana: el sitio Campo
Laborde. Paper presented at the XX Congreso Nacional de
Arqueologı́a Argentina, Rio Cuarto.
Miotti, L., Salemme, M., Rabassa, J., 2003. Radiocarbon chronology at
Piedra Museo locality. In: Miotti, L., Salemme, M., Flegenheimer, N.
(Eds.), Where the South Wind Blows. Ancient Evidence of Paleo South
Americans. Center for the Study of the First Americans, College
Station, pp. 99–104.
Nami, H.G., Nakamura, T., 1995. Cronologı́a radiocarbónica con AMS
sobre muestras de hueso procedentes del sitio Cueva del Medio
(Ultima Esperanza, Chile). Anales del Instituto de la Patagonia 23,
125–133.
Politis, G., Gutiérrez, M., 1998. Gliptodontes y cazadores-recolectores de la
región pampeana (Argentina). Latin American Antiquity 9 (2), 111–134.
Politis, G., Madrid, P., 2001. Arqueologı́a pampeana: estado actual y
perspectivas. In: Berberian, E.E., Nielsen, A. (Eds.), Historia
Argentina Prehispánica, vol. 2. Editorial Brujas, Córdoba, pp.
737–814.
Politis, G., Messineo, P.G., Kaufmann, C.A., 2004. El poblamiento
temprano de las llanuras pampeanas de Argentina y Uruguay.
Complutum 15, 207–224.
Prieto, A., 1991. Cazadores tempranos y tardı́os en la Cueva 1 del lago
Sofı́a. Anales del Instituto de la Patagonia 20, 75–99.
Rick, J., 1980. Prehistoric Hunters of the High Andes. Academic Press,
New York.
Schiffer, M.B., 1987. Formation Processes of the Archaeological Record.
The University of New Mexico Press, Albuquerque, USA.
Semper, J., Lagiglia, H., 1968. Excavaciones arqueológicas en el Rincón
del Atuel (Gruta del Indio). Revista Cientı́fica de Investigaciones 1 (4),
89–158.
Steadman, D.W., Martin, P.S., MacPhee, R.D.E., Jull, A.J.T., McDonald,
H.G., Woods, C.A., Iturralde-Vinent, M., Hodgins, G.W.L., 2005.
Asynchronous extinction of Late Quaternary sloths on continents and
islands. Proceedings of the National Academy of Sciences 102,
11763–11768.
Sugden, D. (Ed.), 2005. Late Glacial events in southernmost South
America and their global significance, Geografiska Annaler 87A (2),
271–408.