Joumul of South Amuicon Eurth Sciences, Vol. 10, No. I, pp.

ZY-38, IY97
Pergamon Q 1997 Published by Elsevier Science Ltd
All nghu reserved. Pnnti in Great Bruin
PII: SO8959811(97)00003-S 08YS-YXlW7 SI7.LX) +o.Ca

Hypothesis on The Cause of Extinction of The South American

Mastodonts
‘G. FICCARELLI, ‘A. AZZAROLI, ‘A. BERTlNI, *M. COLTORTl, 3P MAZZA, ‘C. MEZZABOT-lA,
4M. MORENO ESPlNOSA, ‘L. ROOK and ‘D. TORRE

‘Department of Earth
*Department of Earth
sMuseum of Geology and
4Museo Ecuatoriano de
Sciences, University of Florence Via La Pira 4, 50121 Florence, Italy
Sciences, University of Siena Via della Cerchia 3,53 100 Siena, Italy
Paleontology, University of Florence Via La Pira 4,50121 Florence, Italy
Ciencias Naturales Cane Rumipamba y Avda. de 10s Shirys, Parque La
Carolina POBox 8976, Sue. 7, Quito, Ecuador

Abstract - Paleontological, geomorphological and sedimentological investigations on the Cangahua Formation in the Interandean
depression of Northern and Central Ecuador have provided information on the evolution of the Andean paleoenvironment during
the Late Pleistocene. Pyroclastic and windblown sediments were deposited during cold and dry phases of the last glaciation, inter-
rupted many times by the development of forest-steppe and steppe paleosoils during interstadials. An erosional phase which closed
the Cangahua sedimentation was followed by the deposition of colluvial sediments, characterized by a high number of minor
pedogenetic episodes. The colluviums are confidently referable to the Holocene. The upper part of the Cangahua Formation is rich
in mammal fossils and is probably referable to the Last Glacial Maximum.
The fossiliferous sequences suggest that mastodonts disappeared before mylodonts and equids. We hypothesize that the increased
cold and aridity of the Last Glacial Maximum. which deeply affected the Cordillera. caused the extinction of most of the mega-
fauna and the mastodonts seem to have been the most sensitive to the environmental degradation.
The final history of South American mastodonts, represented by Hupfomustodon and Sregomastodon, spans the latest Pleistocene
and probably the earliest Holocene. Huplomusrodon was dispersed in the highlands within the tropical belt and Sregomustodon in
plains of the southernmost part of Brazil, in Paraguay, Uraguay, Argentine, central and northern Chile.
Both Huplomusrodon and Stegomusrodon suffered the same negative effects of the Last Glacial Maximum when their habitats
underwent intense desertifications under dry and cold conditions. They disappeared in a mosaic way in the course of the latest
Pleistocene, the last representatives probably surviving in favorable restricted areas where however the considerably increased
selective pressure was in the long run devastating. In our opinion the human impact was not a determinant in causing mastodont
extinction. 0 1997 Published by Elsevier Science Ltd. All rights reserved
Resumen - Estudios paleontol6gicos geomorfol6gicos y sedimentol6gicos de la Formaci6n Cangahua, depresi6n Interandina de
la parte Septentrional y Central de Ecuador, han dado interesantes informaciones sobre la evoluci6n de1paleoambiente Andino en
el tardo Pleistocene. Depdsitos pirockticos y e6licos se han depositado durante las fases frlas y aridas de la ultima glaciacibn.
Estos depdsitos se interrumpen varias veces con paleosuolos caraterfsticos de la sfases interglaciales. Un period0 erosivo cierra la
sedimentaci6n de la Formacidn Cangahua a la cual siguen dep6sitos “colluviales” con vatias fases pedogen6tica.s. Estos sedimen-
tos se pueden datar coma holocenicos.
En la parte superior de la Formaci6n Canghaua abundan 10sWsiles de mamiferos referidos al dltimo mbimo glacial.
La sucesi6n fosilifera sujiere la desaparicion de 10s mastodontes antes que 10s milodontes y 10skquidos. Se puede hipotizar que un
aumento del frfo y la desettificacibn durante el dltimo maxim0 glacial pueda haber causado la desaparicidn de la megafauna. Los
mastodontes parecen ser 10s mh sensibles al degrado de1 ambiente.
La historia final de 10s mastodontes suram&icanos, Huplomustodon y Stegomusfodon. se desarrolla en el Pleistocene m&stardio y
el Holocene mh temprano. Huplomustodon ocupa las alturas de1 area tropical y Stegomustr~don las llanuras del sur Brazil, Para-
guay, Uruguay, Argentina y la parte Norte y Central de Chile. Huplomartodon y Stegomusrodon sufrien 10s mismos efkctos en el
tlltimo mbimo glacial quando las condiciones ambientales varian y se instaura una intensa desertiticacidn con un ambiente rni4s
iirido y frfo.
La desaparici6n sigui6 un modelo “a mosaico” durante el Pleistocene mas tardio. Los iiltimos ejemplares sobrevivien en algunas
areas restrictas. En estas areas. sin embargo, el grande acrecimiento de la presidn selectiva ha sido destructive.
Es nuestra opinidn que el impact0 human0 no ha sido determinante en la desaparicidn de 10s mastodontes.

INTRODUCTION Ledru (1993); Iriondo & Garcia (1993); Hansen et al.

South America underwent significant modifications in the
dynamics of landscape moulding due to the widescale cli-
matic deterioration which occurred during the last glacial
phase. Van Gee1 & Van der Hammen (1973); Shackleton et
al. (1983); Hooghiemstra (1984); Rind & Peetet (1985);
Van der Hammen (1978, 1983); Colinvaux er al. (1988);
Van der Hammen & Absy (1994); Clapperton (1993a,b);
(1984); Hooghiemstra & Melice (1994); Heusser & Shack-
leton (1994); Iriondo (1994) proposed reconstructions of
the possible climatic conditions and relevant geomorpho-
logic processes affecting South America during the Last
Glacial Maximum. The models proposed are a compen-
dium of the most important environmental modifications
and of the distribution of the main vegetation belts.

29
 G. FICCARELLI
In the northern and central Andes the glaciers seem to
have reached their greatest extension between 34.000 and
27.000 yr BP, when precipitation increased considerably.
This seems consistent with pollen evidence which suggests
a moderate development of highland pluvial forests as a
consequence of a slight climatic amelioration. All the
authors agree in recognizing a retreat of glaciers, a fall in
lake levels and a reduction of the flux of moisture from
25.000 yr B.P., due to a general cooling. Temperatures
were about 5”-8°C lower than today and this caused an
expansion of grasslands (Van der Hammen, 1983).
According to Van der Hammen & Absy (1994), during
the last glacial period (ca.22.000-14.000 yr B.P.) the tem-
perature in Amazonia turned about 2”-6” C colder than
today and precipitations declined by about 500-l BOO mm.
Both had important consequences on the vegetation. The
savanna stretched towards the equator and the pluvial for-
ests contracted. At the same time, wooded savannas were
replaced by open grassland savannas and other vegetation
boundaries also shifted. Opposite shifts occurred when
temperature and precipitation increased. Alternating open
and wooded savannas thus carefully reflect climatic oscil-
lations. During the last part of the Late Glacial (13.000-
10.000 yr BP) or at the beginning of the Holocene (10.000
yr B.P.), precipitation increased and water levels rose. The
pluvial forest in the tropical areas and the savannas in the
near desert areas reached their greatest extension.
At about 30.000 yr B.P., in Central Brazil the climatic
conditions were equable and more humid than today, while
between 17.000 and 12.000 yr B.P. pollen evidence is sug-
gestive of drier and colder landscapes (Ledru, 1993).
Between 13.000 and 11 .OOO yr B.P. the climate turned
cooler and more humid and, again according to Ledru, a
new cold and dry episode, which is correlated with the
so-called Younger Dryas, is documented between 11.OOO
and 10.000 yr B.P.
While in the northern and central Andes the glaciers
retreated, those in the southern Andes, fed by Pacific
western winds, reached their maximum extension
(20.000-18.000 yr B.P.) when the global climate became
colder (Clapperton, 1993a). However at different times,
during the later part of the last glacial cycle, it seems that
the snow-line lowered about 1.OOOm both in the tropical
and northern part of the southern Andes (Clapperton,
1987; Porter, 1981). Since about 12,500-12,000 yr B.P.
warmer temperatures and increased humidity gave rise to
conditions which favoured the development of the
Andean forest (Hansen et al., 1984; Heusser & Shackle-
ton, 1994).
According to Iriondo & Garcia (1993), between 18.000
and 8.500 yrs BP the Argentine plains were characterized
by an arid and cool climate which shifted the isoclimatic
lines about 750 km northeast of their present location.
Later on, between 8.500 and 3.500 yr B.P. a humid, sub-
tropical and tropical climate was re-established and shifted
the climatic limits about 800-900 km southwest of their
glacial age position.
er al.
In Southern Patagonia and Tierra de1 Fuego Clapperton
(1993b) hypothesizes, presumably at the Last Glacial
Maximum, a thermal decline of at least 10” C than the
present day.
Eolian morphologies suggest that climate in South
America was very cold and windy during the Last Glacial
Maximum, when almost 25% of the continent was covered
by deserts and much of the continent experienced much
drier conditions than now (Clapperton, 1993b).
THE NORTHERN AND CENTRAL ECUADORIAN
ANDES DURING THE LAST GLACIATION
The highest mountain peaks are covered by glaciers
whose tongues stretch down to the altitude of almost
5.000 m a.s.1. (Hasternath, 1981; Clapperton, 1987;
1990). During the last cold phase of the Pleistocene, the
glaciers had their end moraines at about 3.500 m of alti-
tude. Areas beyond the glaciers were possibly affected
by periglacial processes; however, because of Ecuador’s
latitude, these processes were probably restricted, since
no fossil periglacial features have ever been reported
from this region.
The study of a widespread eolian deposit, the Canga-
hua Formation, permitted the reconstruction of the envi-
ronmental conditions in the central valley of Ecuador
during the last glaciation (Sauer, 1965; Baldock, 1982;
Clapperton, 1990; 1991; Iriondo, 1994; Ficcarelli et al.,
1992). The formation is made of fine-grained sands and
silts derived by the intense eolian reworking of pyroclas-
tic deposits, containing variable amounts of plagioclases,
pyroxenes, amphiboles, quarz, biotite, rock fragments,
pumice and volcanic glasses. However, tephra intercala-
tions occur in almost all the deposits, sometimes contain-
ing fall-out products of the cordilleran volcanoes, such
as pumice, lapilli and rock fragments, with no evidence
of eolian reworking. Sauer (1965) first described these
deposits and recognized several facies, in particular, an
eolian and a lacustrine facies. He correlated these sedi-
ments with interglacial phases and identified three peri-
ods of deposition including a postglacial period, which
would correspond, according to Iriondo (1994), to an
extensive pyroclastic cover observed in the Amazonian
side. Sauer and Iriondo both suggested a humid grass-
land depositional environment for the Cangahua Forma-
tion with temperatures slightly warmer than the present.
On the other hand, Clapperton & Vera (1986) and Clap-
perton (1990) referred this formation to the last cold
stage of the Pleistocene.
In the northern sector near Bolivar and S.Gabriel
(Carchi Province) (Figs. 1, 2), a composite sequence has
been described formed by various outcrops, correlated
using a number of reference tephra layers. The basal part
lies conformably over fluvial sediments, many hundreds
of meters thick, containing rare pyroclastites, debris
flows, breccias, agglomerates and lava flows. These
deposits form an extensive plateau whose elevation var-
ies according to the activity of recent normal faults, from
 Hypothesis on The Cause of Extinction of The South American Mastodonts
Fig. 1. Location map of the studied stratigraphic sections. 1)
Casario ‘IIquer, 2) Panamerican Highway, 3) Quebrada Cuesaca,
4) Rio Chiche, 5) Quebrada Chak.
2,800 to over 3,000 m a.s.1.. Breccias, agglomerates and
lava flows increase progressively toward the cordillieras
and interfinger with the products of the Boliche and
Mangus volcanoes. The transition to the Cangahua For-
mation is realized through the progressive reduction of
the elastic coarse sediments and the increase of pyroclas-
tic and eolian deposits. In the Casario Tuquer sequence
(Fig.2, section a), lahars, mud flow, ash fall and eolian
sediments are present at different levels. These sedi-
ments are in many cases weathered by pedogenesis and
form paleosoils which constitute an important source of
information on the past climatic environment. They are
represented by evolved and non evolved soil profiles.
The former, which can be more than 2 m thick, are
leached, rubified and contain argillic horizons, and were
developed under a forest cover during long lasting Inter-
glacial conditions. The less evolved soils are represented
by chernozems, vertisoils and brunizems which testify a
forest-steppe or a praire environment. The occurrence of
a Ca horizon made of pseudomicium and/or of thin nod-
ules or crusts is common. The brunizems may include a
thin argillic horizon frequently mixed with calcium car-
bonate (Bca). These paleosoils have been connected to
Interstadial conditions. Sometimes, a horizon made up of
calcium carbonate platy veins may occur at the base or at
the top of the profile revealing changes to open steppe or
even to desert environment (Gile et al., 1966).
The Cangahua sequence in the Carchi Province is com-
pleted by some sections along the Panamerican Highway
(Fig.2, section b). A rubified argillic horizon, located at the
top of the sequence and truncated by an unconformity, rep-
resents the most developed paleosoils below the present
day surface and could correspond to the last Interglacia-
WIULO-L-C
31
tion. A second unconformity, not associated with pale-
osoils, is present in a lower position. Minor unconformities
have also been observed inside the secondary valleys on
the flanks of the Boliche volcano, where stream sediments
are intercalated with eolian deposits.
At the top of the sequence, on the ridges (Fig.2, section
c), a black andosoil developed under Holocene climatic
conditions. The upper part of this paleosoil is marked by
increased conditions of profile aridity which led to the for-
mation of carbonate crusts a few decimeters thick. It seems
likely that this profile aridity has to be connected to the
strong deforestation which occurred in the area since Pre-
Colombian times. The profile is also locally truncated and
divided into two subunits by the presence of colluvial sedi-
ments which contain archaeological remains (Negative
Carchi Culture, 500-1500 AD). Contemporaneously, the
small valleys of the plateau were filled with colluviated
sediments at places containing stone artifacts suggesting
the presence of widespread preceramic occupation. Before
human intervention, a general downcutting of the Canga-
hua was under way.
Mammal remains come from three fossiliferous layers
in the upper part of the Cangahua Formation. The lower-
most level is characterized by a high amount of masto-
donts (Haplomastodon chimborazi) and a slightly lesser
abundance of mylodonts (Glossotherium). In the second
level, which lies directly above, Glossotherium is well
represented, cervids (Matama rujina) are rare, while
mastodonts apparently disappeared. A rich micromam-
ma1 assemblage, mostly represented by pastoral cri-
cetids, such as Phyllotis and Akodon, was recovered from
the upper level, where a Smilodon canine was also found.
Few isolated equid cheek teeth were recovered in the
uppermost part of the outcropping sequence. The present
record is suggestive of a succession of extinction waves,
which first affected the mastodonts, then the mylodonts
and, finally, the equids. On the contrary, cervids and
micromammals survived these extinction events without
apparent modifications, since the same species are
present in Holocene deposits (Fejfar et al., 1993). On the
basis of geomorphological, sedimentological and pale-
ontological evidence, the fossiliferous levels are confi-
dently referred to the latest Pleistocene.
Close to Quito, the Rio Chiche Sequence, already
described by Sauer (1965) and Clapperton & Vera (1986),
represents the most complete sequence of Central Ecuador.
It was investigated in this study with respect to the pale-
osoils present in it (Fig.3, section a). The transition from
underlying fluvial and laharic sediments is gradual, as in
the Car&i area. The Cangahua Formation contains many
pyroclastic layers and paleosoils formed under forest-
steppe and steppe vegetation. A rubified horizon, which
could be connected to an interglacial paleosoil, is present in
the lower part of the formation. No unconformities have
been observed. However, like in the Carchi Province, they
can be locally masked by paraconformities. Close to Rio
 60

66

50

I
46

40

30
5
-=-r^ _--_r
- -:+:

2.5

20
10
:

,
15
, I

,
,
10
a b
C
* .,.
5 . . . ; :',
i L6

0
Fig. 2. Stratigraphic sections of the Casario Tuquer (a), Panamerican
Legend: 1 andosoils; 2 colluvial soils; 3 tephra; 4 tephra, scoria falls;
10 coarse lahars; 11 brunizem with Bt-Horizons (forest steppe soils);
conglomerates; 18 silts and sands; 19 Cretaceous marls; 20 weathered
Y
Highway (b), and Quebrada Cuesaca (c) sequences and their correlations.
5 chemozem; 6 chernozems containing “bolas de Canghaua”; 7 K-Horizon - Desert open steppe soil; 8 vertisoils; 9 fine lahars;
12 fragipan horizon; 13 Ieaches reddish paleosoils (tropical forest); I4 sands; 15 sands and gravels; 16 boulders; 17 gravels and
granite; 21 unconformity. Solid triangles indicate Carchi culture level.
 Hypothesis on The Cause of Extinction of The South American Mastodonts 33

Fig. 3. Stratigraphic sections of the Rio Chiche (a) and Quebrada ChaMn (b) sequences. For the legend, see Fig.2

Chiche, at Tumbaco, fauna1 remains were observed in col-
luvial deposits within the Cangahua, a few meters below a
black andosoil similar to the one observed in the Carchi
Province.
On a wide-scale view, the megafauna of the northern
part of the Ecuadorian Cordillera (Carchi, Imbabura and
Pichincha Provinces) is characterized by a predominance
of mastodonts (Haplomasbdon chimborazi) and a large
abundance of mylodonts (Glossotherium), while cervids
and equids are poorly represented.
A preliminary study of the sedimentary sequence of
Quebrada Chal& (Fig.3, section b), in the area of Punin,
south of Riobamba (Chimborazo Province) celebrated for
its fossil mammal content (Hoffstetter, 1952), has also
been carried out by the writers. The 14 m thick sequence is
subdivided into two cycles by an unconformity. The fauna
34 G. FICCARELLI
is preserved about 7 m from the top of the outcropping
sequence. Eolian deposits weathered by forest-steppe and
steppe soils characterize the upper part of the sequence.
Soils with a similar degree of evolution have not been
observed in the upper part of the Carchi sequences.
The paleontological investigations revealed a remark-
able abundance of equids and cervids and a modest occur-
rence of camelids. Extremely scanty element of mastodont
and mylodonts were found during these field surveys. In
literature mastodont remains are reported from the Punin
area, including the type specimen (presently lost) of the
Late Pleistocene mastodont, Haplomastodon chimborazi.
In any case, the occurrence of mastodonts in the Punin area
seems very rare. Hoffstetter (1952) reported the recovery
of a partial left femur of Cuvieronius tarijensis (= Cuviero-
nius hyodon) (Ficcarelli et al., 1995) during the excavation
of a trench in the Quebrada Colorada. The trench cuts
through the sequence and therefore the stratigraphical posi-
tion of the find is unknown. Unfortunately, most of the his-
torical data from Ecuador are doubtful. The mastodont
femur from Punfn belongs to a juvenile individual; for this
reason a possible attribution to H. chimborazi should thus
not be ruled out. The only true C. tarijensis material (two
adult fragmentary tusks) presently known from Ecuador
was donated by anonymous collectors to the Colegio
Benign0 Malo, at Cuenca, where it is still preserved today.
This material is reported from Bafios de Cuenca but with-
out reliable stratigraphical information. Some of the
present authors, revising the mastodont material of Ecua-
dor (Ficcarelli et al., 1995), have already stressed the erro-
neous stratigraphical and systematical attributions reported
in literature, and the uncertain provenance of part of the
fossil material known until now.
On the basis of paleontological evidence, it is difficult
to ascertain how the Carchi and Punin fauna1 successions
are correlated. The contrasting amounts of the faunas may
reflect different environmental conditions existing in the
two areas, the Punfn territory being more arid than Carchi,
as it is today, and/or a somewhat difference in age.
From the stratigraphical and sedimentological point of
view, it is possible to conclude that: 1) the Cangahua is
mainly composed of volcanic deposits, largely reworked
by eolian and colluvial activity, laid down during the cold
phases of the Pleistocene; 2) its deposition was interrupted
by the evolution of paleosoils sometimes associated with
unconformities; 3) it contains leached and rubified hori-
zons which represent interglacial soils testifying that the
deposition occurred during more than a single cold phase;
4) it is thicker in the northern part of the country and tapers
southwards.
The most significant fossil assemblages from the coastal
area are those from La Carolina (Santa Elena Peninsula,
Guayas Province), where, according to literature, masto-
donts, megatherids, equids and cervids are well repre-
sented. These finds are also referable to the latest
Pleistocene. At present, inferences can be only drawn from
literature, in the absence of direct observations by the writ-
ers. Spillmann (1942) and Hoffstetter (1952) contrast in the
et al.
interpretation of the La Carolina sequence. Spillmann
(1942) quoted the occurrence of three successive fossilifer-
ous layers, the lowest characterized by large-sized mam-
mals, the upper ones by smaller-sized forms. On the other
hand, Hoffstetter (1952) believed that all the specimens
came from a single geological horizon. Spillmann’s view is
consistent with the succession observed in the Carchi area.
However further geological investigations and a deeper
knowledge of the stratigraphic characters of the sequence
are needed.
In the light of the evidence from Carchi and Punin, the
writers believe that the successive disappearance of mas-
todonts, mylodonts, and, finally, equids was widespread
in the Interandean Cordillera of Ecuador. In the Cordill-
era no human remains or artefacts were found in associa-
tion with mastodonts. This suggests that these
megafauna elements may have died out before the arrival
of Man. Even the famous human skull from Punin was
found as an erratic and its stratigraphical position is
therefore unreliable.
Palynological analyses have been performed on sam-
ples collected in Carchi and Punin. Unfortunately, the
results have been disappointing, at the moment.
Although a high density of samples was taken and heavy
liquid enrichment techniques have been used because of
the peculiar nature of the sediments, all the samples
proved sterile.
DISTRIBUTION OF SOUTH AMERICAN
MASTODONTS DURING THE LATEST
PLEISTOCENE
Outside Ecuador, the geographical distribution of mas-
todonts in other parts of South America seems to outline
two different fauna1 provinces: a northern one, extending
along the tropical belt, from the Pacific to the Atlantic
coasts, and a southern one including Southern Brazil, Para-
guay, Uruguay, the Argentine plains and part of Chile (Fig.
4). The first is characterized by the occurrence of Haplo-
mastodon, the second by Stegomastodon. This distribution
may reflect the different ecological preferences of the two
genera. Elephants are our only living reference, although
they differ considerably from mastodonts, especially in the
dentition. Loxodonta and Elephas show broad habitat toler-
ance, since they live from near desert areas to savannas, to
woodlands, to tropical forests (Haynes, 1991). Unfortu-
nately, a major problem in comparing with extant species is
that present ranges are controlled by the severe pressure of
anthropic activity, and therefore some habits and some
relations with the environments are probably unnatural.
The dentition, skull and mandible structure and the
arrangement of the masticatory apparatus of South Ameri-
can mastodonts, as well as the fauna1 assemblages in which
their remains are usually found, suggest that these animals
may have been prevalently woodland to open woodland
dwellers; however it cannot be excluded that, like
elephants, their range of habitats may have been broader,
including grassland savanna. In particular, functional mor-
phological analyses of the dental structures suggest that
 Hypothesis on The Cause of Extinction of The South American Mastodonts 35

L _ Savono 7

Arnozon Cane

Arid 8 Flarh
Floods a MIII Savono, Coatlngor
a Campor Cbrrodos A-
ICE UPS a . . A.
VALLET GLACI

Aroucorla ford

Limit of S~a~natly/P~rrnniolly
PotaQonian Ice Cap Froxrn Ground

A Haplomastodon chimborazi
W Stegomastodon ex gr. piatensis-superbus

Fig. 4. Distribution of mastodonts in South America during the Last Glacial Maximum (modified from Clapperton, 1994).

Haplomastodon was primarily a woodland dweller and that
Stegomustodon probably preferred environments ranging
from woodland to a more open grassland savanna with
trees and shrubs. The mastodonts of the tropical belt were
probably confined by desert areas, like the ones now occtr-
ring in Peru and northern Chile, and pluvial forests. The
mastodonts farther south were likely blocked by pluvial
forests, to the north, and by the hostile conditions of the
southernmost parts of Patagonia and Chile.
As in Ecuador, mastodonts seem to have disappeared
from Argentina before equids (Tonni, pers. comm.), since
their remains have never been found in latest Pleistocene
deposits. This seems to reflect the progressive climatic
36 G. FICCARELLI
deterioration, with increasing aridity, which developed
during the Last Glacial Maximum (Van der Hammen,
1983; Ledru, 1993; Heusser & Shackleton, 1994) and
which is the most probable cause of these successive waves
of extinctions. The occurrence of Stegomastodon in the
Tagua-Tagua deposits of central Chile and at Monteverde,
in the northern part of southern Chile, dated to some
13.000-11.000 yr BP. (Nuiiez Atencio, 1994; Politis etaf.,
1995), may attest to the fact that this particular area at the
end of the Pleistocene was more humid than today and may
therefore have been characterized by a richer vegetation
cover. Also today, Patagonia and the westernmost part of
northern Argentina are more arid than central-southern
Chile as a consequence of the fact that the prevailing air
masses from the Pacific Ocean rise on the windward side of
the Cordillera, precipitate their humidity and cause the for-
mation of a dry rainshadow desert on the lee side of the
Cordillera, in Argentina. This might have been even more
intense during the Last Glacial Maximum because of a
northern shift of the Pacific anticyclone.
The reconstruction of the paleoenvironment of Brazil
and of the Amazon Basin is even more complex, due to the
transition from pluvial forests to steppes. Haplomastodon
dwelt at higher altitudes suggesting that woodland condi-
tions existed in these areas; Stegomastodon lived more to
the south (Rio Grande do Sul), in lands where the vegeta-
tion was probably more open. The only well dated masto-
dont finds are those collected in Late Pleistocene deposits
at Barra do Antonilo and Toca de Garrincho in the Sao
Raimundo Nonato region, State of Piaui (NE Brazil), and
referred to Haplomastodon (Guerin et al., 1993; Guidon et
al., 1994). According to these authors, Haplomastodon dis-
appeared from these localities at the end of the Late Pleis-
tocene. The poor information prevents a reliable
reconstruction of the stratigraphical distribution of the two
genera and also any conclusive remarks relative to their
contemporaneity, to their possible ecological limits and to
the timing of their extinction.
CONCLUSIONS
In the present state of knowledge, finding the causes and
precise time of extinction of South American mastodonts
certainly is not an easy task. There is no doubt that Haplo-
mastodon disappeared from the Northern Ecuadorian Cor-
dillera in the course of the Last Glacial Maximum and that
its extinction preceeded that of mylodonts and equids. This
seems to have occurred, in the same order, also in the Cen-
tral Cordillera (Punin, Chimborazo Province). In the light
of our present knowledge it is not possible to ascertain if
the extinctions in the megafauna have been successive also
in the coastal area (La Carolina, Santa Elena peninsula,
Guayas Province) of Ecuador.
At the moment it is unknown if the megafauna distrib-
uted throughout the tropical belt underwent differential
extinctions because of the progressive increase in aridity of
the Latest Pleistocene.
et al.
In the Argentine plains, Stegomastodon is not reported
from the youngest deposits of the Last Glacial Maximum
and anyhow, even in these areas, it became extinct before
the other megafauna representatives.
Paleontological evidence from Chile suggests the
occurrence of Stegomastodon until about 11.000 yr B.P.
Some researchers believe that the anthropic impact of
the so-called hunter-gatherers may have been the main or
an additional cause of extinction of the megafauna in South
America and in general on a world-wide scale during the
latest Pleistocene and the onset of the Holocene. In the
opinion of the present authors there is enough evidence to
exclude Man as a possible cause of extinction of masto-
donts. In the Ecuadorian Andes, and in the Ecuadorian and
northern Peruvian Coasts, mastodonts became extinct
before Man arrived. The same occurred in the Argentine
plains. Man apparently colonized the Argentine plains and
central-southern Chile, the so-called “Southern Cone”,
about 13.000-11 .OOOyr B.P.(Politis et al., 1995). However,
in a few localities, mastodonts actually coexisted with Man
for a certain lapse of time before their complete disappear-
ance. Haplomastodon is associated with Man in northeast-
ern Brazil (Guerin et al., 1993; Guidon et al., 1994) and at
Taima-Taima, in the Coro Peninsula, Venezuela (Bryan et
al., 1978; Bryan, 1986); Stegomastodon occurs with Man
at Tagua-Tagua and at Monteverde, Chile (Montane, 1968;
Dillehay, 1986; Dillehay & Collins, 1988; Nufiez Atencio,
1994). Correal Urrego (1981) reported an archaeological
site at Tibito, Bogota Sabana in Columbia, where artifacts
are associated with mastodont remains (cfr. Marshall et al,,
1984). This last fauna1 assemblage is however unreliable
because of the occurrence of taxa which seem to be sugges-
tive of different stratigraphical provenance.
The writers believe that the extinction of mastodonts in
South America is connected with significant climatic
changes which caused intense and abrupt environmental
modifications, too extreme for their adaptive capacity.
These marked climatic modifications, certainly triggered
by the Last Glacial Maximum cooling, deeply affected the
distribution and amount of precipitation. This in turn had
important consequences on the distribution of foraging and
arid areas and, finally, on the survival of mastodonts, which
disappeared in a mosaic way. Mastodonts were probably
forced to leave the regions characterized by a dry and cold
climate and to concentrate in favorable restricted areas,
where however the resulting selective pressure consider-
ably increased. As these large-sized animals require wide
foraging areas, most suffered the negative consequences of
this drastic environmental choking, which may have given
rise to delays in sexual maturity and decrease in fertility, as
has been observed in extant elephants confined within
restricted areas (Haynes, 1991). The imbalance between
feeding requirements and territory productivity may have
been as devasting. In these critical circumstances the
human impact may have hastened an anyway unavoidable
fa@ for these large-sized mammals.
In this scenario Haplomastodon probably disappeared
from the highlands during the Last Glacial Maximum and
 Hypothesis on The Cause of Extinction of The South American Mastodonts
probably survived slightly longer in some restricted areas
of the Amazon Basin andfor the lowlands of Brazil. If the
age of the sequence is younger than that of Carchi, La
Carolina could have been another possible refuge area.
However a post-Pleistocene persistence of Huplunmtodun
in the above mentioned areas is only speculative as at
present no data justify such a conclusion. Stegomastodon
certainly disappeared from the Argentine plains before the
end of the Last Glacial Maximum but survived in limited
areas of central Chile and perhaps in southern Brazil if it
would be proved that the finds from Rio Grande do Sul are
more recent.
Acknowledgments- The authors express their deep gratitude to Dr. R.
Pascual and E.P Tonni of the Departamento Cientifico de Paleontoiogia
des Vertebrados. Museo de La Plata, Argentina; Dr. A.Ramos, Museo
Argentino de Ciencias Naturales “Bernardino Rivadavia” de Buenos
Aires; Dr. D. Frassinetti C., Museo National de Historia Natural de San-
tiago del Chile; Dr. S.A. Azevedo, Museo National de Rio de Janeiro, for
granting them access to the collections of their Institutions.
The work was financially supported by C.N.R. and M.U.R.S. grants.
Coltorti M. dealt with Geomorphology and Stratigraphy Ficcarelli G,
Azzaroli A., Bertini A., Mazza P., Mezzabotta C., Moreno Espinosa M.,
Rook L. and Terre D. dealt with Paleontology and Stratigraphy.
REFERENCES
Baldock, J.W., 1982. Geologiu de1 Ecuudar: Boletin de 11 Explicucion de1
Mapa Geol&ico de lu Reptiblicu de1 Ecuado,: escala 1:1.ooO.000.
Direci6n General de Geologia y Minas, Quito, Ecuador, 66 p.
Bryan, A.L., 1986. Paleoamerican Prehistory as seen from South
America. In: New Evidencefor the Pleistocene Peopling in the Ameri-
cus (edited by A. Bryan), pp. I-14. Center for the Study of Early Man,
University of Maine.
Bryan, A.L., Casamiquela, R.M., Cruxent, J.M., Gruhnn, R. & Ochsenius,
C., 1978. An El Jobo Mastodon Kill at Taima-Taima, Venezuela. Sci-
ence, 200, 1275-1277.
Clapperton, C.M., 1987. Glacial geomorphology. Quaternary glacial
sequence and paleoclimatic inferences in the Ecuadorian Andes. In:
Internutionul Geomorpholagy 19g6, Part II (edited by V. Gardiner).
Wiley, London, 843-870.
Clapperton, C.M., 1990. Glacial and volcanic geomorphology of the
Chimborazo-Carihuairazo massif, Ecuadorian Andes. Transactions
Royul Satiety, Edinburgh, 81.91-116.
Clapperton. C.M., 1991. Origin and characteristics of debris avalanches in
the Ecuadorian Andes. XIII Internutional INQUA Congress, Abstract
volume 61, Beijing.
Clapperton, C.M., 1993a. Quuternury Geolagy und Geomorpholagy of
South America. Elsevier Amsterdam.
Clapperton, CM.. 1993b. Nature of environmental changes in South
America at the last Glacial Maximum. Palaeugeugruphy, Pulueocli-
matology, Pulueoecology, 101, 189-208.
Clapperton, CM. & Vera, R., 1986. The quatemary glacial sequence in
Ecuador. A reinterpretation of the work of W. Sauer. Journul of Quu-
ternary Science, 1, 45-56.
Colinvaux, P.A., Olson, K. & Kam-Biu, L., 1988. Late-glacial and
Holocene pollen diagrams from two endorheic lakes of the Inter-
Andean Plateau of Ecuador. Review of Puleobotuny und Palynology,
55.83-99.
Correal Urrego, G., 198I. Evidencias Culturales y Megafauna Pleistoce-
nica in Colombia. Bogotd, Fondacicin de Investiguciones Arqueokgi-
cus Nucionales, 12. 1-148.
37
Dillehay, T.D., 1986. The Cultural Relationships of Monte Verde: Late
Pleistocene Settlement Site in the Sub-Antartic Forest of South-
Central Chile. In: New Evidence for the Pleistocene Peopling of the
Americus (edited by A. Bryan), Center for the study of Early Man.
University of Maine, 3 19-337.
Dillehay, T.D.& Collins, M., 1988. Early Cultural Evidence from Monte
Verde in Chile. Nurure, 332, 150-152.
Falgu&es, C., Fontugne, M., Chauchat, C.. Guadelli, J.L., 1994. Datations
radiom&riques de I’extinction des grandes faunes pl&stoc&nes au
Wrou. Camptes Rendus de 1’Acudemie des Sciences, Puris, 319, (III),
26 l-266.
Fejfar, 0.. Blasetti, A., Calderoni, G., Coltorti, M.. Ficcarelli, G., Masini,
E, Rook, L. & Terre, D., 1993. New fossil finds of Cricetids in North-
em Ecuador. Documents des Luborutaires de GPoiagie, Lyan, 125,
151-167.
Fejfar, 0.. Ficcarelli, G., Mezzabotta, C., Moreno Espinosa, M.. Rook, L.
& Terre, D., in press. First record of a Copemyne-Peromyscine form
in South America. Hypotheses on its ancestry in Palearctic. Actu Zoo-
lagica Cracoviensiu, 38.
Ficcarelli, G., Azzaroli, A., Borselli, V., Coltorti, M.. Dramis, F., Fejfar,
0.. Hirtz, A. & Terre, D. 1992. Stratigraphical and Paleontological
aspects of Late Quatemary deposits in the Interandean Depression of
Northern Ecuador. Journal of South American Eurth Sciences, 6 (3).
145-1.50.
Ficcarelli, G., Borselli, V., Moreno Espinosa. M. & Terre. D. 1993. New
Hapbmastudon finds from the Late Pleistocene of Northern Ecuador.
Geabibios,26 (2), 23 I-240.
Ficcarelli, G.. Borselli, V., Herrera, G., Moreno Espinosa. M. & Torre, D.,
1995.Taxonomic remarks on the South American mastodonts referred
to Huplomustadan and Cuvieronius. Geobios, 28 (6). 745-756.
Gile, L.H.. Peterson, F.F. & Grossman, R.B., 1966. Morphological and
genetic sequences of carbonate accumulation in desert soils. Sail Sci-
ence, lOl(5). 3 16-360.
Guerin, C., Curvello, M.A., Faure, M., Hugueney, M., Mourer-Chauvir&
C.. 1993. La faune pleistodne du Piaui (nordeste du Bn%il): implica-
tions paleoecologiques et biochronologiques. Quuternuria nova, 3,
303-341.
Guidon, N..Parenti. F., Da Luz, M., Gubrin, C.& Faure, M., 1994. Le plus
ancien peuplement de I’Amt%ique:le PalColitique du nordeste Br&il-
ien. Bulletin de la Societe Prkhistorique Frun~uise, 91 (4-5), 246
250.
Hansen, B., Wright, H.E. jr. & Bradbury, J.P., 1984. Pollen studies in the
Jumin area, Central Peruvian Andes. Geobgicul Society ufAmerica
Bulletin, 95, 1454-1465.
Hastemath, S., 198I. Glaciation oftheEcuudariun Andes. Balkema, Rot-
terdam, I59 p.
Haynes, G., 1991. Mammoth Mustadants & Elephrmts, Biology Behavior
und the Fossil Record. Cambridge University Press, 413 p.
Heusser, L.E. & Shackleton, N.J.. 1994. Tropical climatic variation on the
pacific slopes of the Ecuadorian Andes based on a 25.000-years pollen
record from Deep-sea sediment core tri163-31B. Quaternary
Research, 42,222-225.
Hoffstetter, R. 1952. Les Mammiferes Pltistocbnes de la Republique de
I’Equateur. Mimoires de lu Saci& G~ologique de Frunce, 31 (I-4).
66, l-391.
Hooghiemstra, H., 1984. Vegetutional und climufic history of the High
Plain of Bog& Columbia: u continuous record of the lust 3.5 million
years. J. Cramer, Vaduz.
Hooghiemstra, H., & Melice. J.L., 1994. Pleistocene evolution of orbital
periodicities in the high-resolution pollen record Funza I, Eastern Cor-
dillera, Colombia. Special Publicutions International Association uf
Sedimentolagy, 19, 117-126.
38 G. FICCARELLI et al.
hiondo. M. & Garcia N.O., 1993. Climatic variations in the Argentine
Plains during the last 18,000 Years. Palueogeogruphy, Pulueoclimu-
tology, Pulueoecology, 101.209-220.
hiondo. M., 1994. The Quatemary of Ecuador. Quuternury International,
21, 101-I 12.
Ledru, M.P., 1993. Late Quaternary environmental and climatic changes
in Central Brazil. Quaternury Reseurch, 39,90-98.
Marshall, LG.. Berta, A., Hoffstetter. R.. Pascual, R., Reig, O.A.,
Bombin, M. & Mones, A., 1984. Mammals and stratigraphy: geo-
chronology of the continental Mammal-bearing Quatemary of South
America. Pulueovertebrutu, Mimoire Exrruordinuire, l-76.
Montanb. J., 1968. Paleo-Indian Remains from Laguna de Tagua-Tagua.
Central Chile. Science, 161, 1137-I 138.
Nuhes Atencio. L. 1994. El sitio Paleoindio de Tagua-Tagua y sus rela-
ciones en el Cono Sur. Puper presenred at rhe XI Congreso Nucionul
de Aryueologiu Argentina. Sun Rufuel, Mendozu.
Politis, G.G., Prado, J.L. & Beukens. RI?, 1995. The human impact in
Pleistocene -Holocene extinctions in South America. In: Ancient Peo-
ples and Landwupes (edited by E. Johnson), Museum of Texas Tech-
nology University, 187-205.
Porter, SC., 1981. Pleistocene glaciation in the southern Lake District of
Chile. Quuternury Research, 24,269-292.
Rind, D. & Peteet, D., 1985.Terrestrial conditions at the last glacial maxi-
mum and CLIMAP sea-surface temperature estimates: are. they con-
sistent? Quutemury Reseurch, 24, l-22.
Sauer, W., 1965. Geobgiu de1 Ecuudor. Editorial de Ministerio de Edu-
cation. 383 p.
Shackleton, N.J., Imbrie, J. & Hall, M.A., 1983. Oxygen and carbon iso-
tope record of East Pacific core V 19-30: implications for the forma-
tion of deep water in the late Pleistocene North Atlantic. Eurth und
Plunerury Science L.etter.s, 65, 233-244.
Spillmann. F. 1942. Contribution al conocimiento de fosiles nuevos de la
avifauna Ecuadoriana en el Pleistocene de Santa Elena. Proceedings
l>f the 8th Americun Science Congress, 4,315-389.
Van der Hammen, T., 1978. Stratigraphy and environments of the Upper
Quatemary of the El Abra conoide and rock shelters (Columbia). Pul-
ueogeogruphy, Pulueoclimurology, Pulueoecobgy. 25, II l-162.
Van der Hammen, T., 1983. Palaeocology and palaeogeography of savan-
nas. In: Ecosystems of the world n. 13 (edited by F. Bourliere), Else-
vier Science Publication Company, 19-35.
Van der Hammen, T. & Ahsy, M.L., 1994. Amazonia during the last gla-
cial. Pulueogeogruphy. Pulueoclimutology Pulueoecology, 109,247-
261.
Van Geel, B. & Van der Hammen, T., 1973. Upper Quaternary vegetational
and climatic sequence of the Fuguere area (eastern Cordillera, Colum-
bia). Pulueogeogruphy, Puluer~limutology, Pulueoecology, 14.9-92.