Quaternary Research 61 (2004) 289 – 300

www.elsevier.com/locate/yqres

Reconstructing habitats in central Amazonia using megafauna,

sedimentology, radiocarbon, and isotope analyses
Dilce de Fátima Rossetti, a,* Peter Mann de Toledo, a
Heloı́sa Maria Moraes-Santos, a and Antônio Emı́dio de Araújo Santos, Jr. b
a
Museu Paraense Emı́lio Goeldi, Av. Perimetral, 1901, CP 399, CEP 66710-530 Belém, PA, Brazil
b
Universidade Federal do Pará, Centro de Geociências, Campus do Guamá S/N, Belém, PA, Brazil
Received 14 November 2002

Available online 30 April 2004

Abstract

A paleomegafauna site from central Amazonia with exceptional preservation of mastodons and ground sloths allows for the first time a
precise age control based on 14C analysis, which, together with sedimentological and y13C isotope data, provided the basis to discuss habitat
evolution within the context of climate change during the past 15,000 yr. The fossil-bearing deposits, trapped within a depression in the
Paleozoic basement, record three episodes of sedimentation formed on floodplains, with an intermediate unit recording a catastrophic
deposition through debris flows, probably favored during fast floodings. The integrated approach presented herein supports a change in
humidity in central Amazonia through the past 15,000 yr, with a shift from drier to arboreal savanna at 11,340 (F50) 14C yr B.P. and then to a
dense forest like we see today at 4620 (F60) 14C yr B.P.
D 2004 University of Washington. All rights reserved.

Keywords: Amazonia; Pleistocene; Paleontology; Mammals; Sedimentology; Radiocarbon dating; Landscape evolution

Introduction such a large and complex natural system. Although the

Deciphering the origin of the Amazon biodiversity has
been a challenge to the scientific community with special
interest in issues related to natural history and conservation
of communities and ecosystems. An important aspect of this
multidisciplinary field is the understanding of the main
historical factors relating physical and biological phenome-
na that acted upon the shaping of the modern biome as we
see today. In order to reconstruct the origin and the historical
events of the main ecological processes that took place to
form the rain forest, an analysis and organization of a series
of multidisciplinary data related to geology and climate and
a reasonable control of the fossil history are needed. So far,
geological and paleontological data are relatively scarce,
considering the continental dimensions of the Amazon
region, and the information available furnishes only a broad
view on the evolutionary patterns. The building of historical
datasets is an important contribution to the understanding of
* Corresponding author. Fax: (091) 249-0466.
E-mail address: rossetti@museu-goeldi.br (D. de Fátima Rossetti).
entire history back to at least the Early Tertiary is relevant to
these studies, the Pleistocene should be particularly
addressed, as it bears the closest relationship with the
modern ecosystem. Only a broad picture of what happened
in the Amazon region during the major ecological shifts
between ice-age aridity and more humid interglacial periods
has been provided so far (reviewed by Latrubesse, 2000).
This is mainly due to the following reasons: (1) geological,
palynological, and vertebrate paleontological data are still
scarce and spotty; (2) information refers only to some areas
located in southeastern and southwestern Amazonia; and (3)
the best information is related to times before 24,000 yr ago
(Latrubesse, 2000). The incomplete information has moti-
vated many debates, with the arid Amazonia refugia model
(Haffer, 1969; Prance, 1982) on one side against stability of
the forest throughout the Cenozoic on the other side
(Colinvaux et al., 2000; Colinvaux and Oliveira, 2001).
This paper reports a new fossil quarry bearing two
megafauna elements consisting of Haplomastodon waringi
and Eremotherium laurillardi from the locality of Itaituba,
State of Pará, northern Brazil (Fig. 1), in Central Amazonia,

0033-5894/$ - see front matter D 2004 University of Washington. All rights reserved.
doi:10.1016/j.yqres.2004.02.010
290 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300

Fig. 1. Location and geologic map of the Itaituba area in the state of Pará, northern Brazil, with the location of the fossil quarry bearing the megafauna of
mastodon (Haplomastodon waringi) and giant ground sloth (Eremotherium laurillardi).

which allowed precise age control. Integration of paleontol-
ogy and sedimentology, as well as radiocarbon and isotope
data, provides the basis for discussing the possible changes
in this landscape through the past 15,000 yr.
Geological framework and physiography
The Itaituba area is located in the southern margin of the
Amazonas Basin, which is a large (i.e., nearly 500,000 km2)
depression located in the middle and low Amazonas. This
basin is bounded by the Guianas Shield to the north, Brazil-
ian Shield to the south, Purus Arch to the west, and Gurupá
Arch to the east. The origin of the Amazonas Basin is related
to rifting associated with intraplate stretching during the
Paleozoic, which took place in three stages and gave rise to a
6500-m-thick sedimentary package represented by three
megasequences formed in the Ordovician – early Devonian,
middle/late Devonian –early Carboniferous and middle Car-
boniferous – Permian. The opening of the South Atlantic
Ocean and rise of the Andean Cordillera resulted in the
tectonic reactivation of the area during the Cretaceous –
Cenozoic and deposition of a fourth megasequence up to
500 m thick. This later phase of tectonism continued even in
more recent times in the late Quaternary, having a strong
effect in the development of the modern drainage system
(e.g., Costa and Hasui, 1997; Bemerguy, 1997).
The fossil-bearing sedimentary deposits emphasized in
this study rest in the left margin of the middle Tapajós River
and occur overlying limestone of the upper Carboniferous
 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300
Fig. 2. A view of the Itaituba fossil quarry, illustrating the sharp, erosional
contact between the Pleistocene deposits and the underlying Paleozoic
Itaituba Formation. Note the karstic structures at the bounding surface
(arrows) and the ragged, straight vertical segments of the basement,
probably resulting from fault displacement.
Itaituba Formation (Fig. 1). The Tapajós River runs through
a NE/SW-oriented normal fault zone, which is part of a
triple junction formed by strike– slip reactivation of Meso-
zoic structures during the Quaternary (Costa and Hasui,
1996). This river is in general straight but it becomes highly
meandering in the Itaituba region, with large coarse-grained,
sandy point bars. The fossil quarry is located in the flood-
plains, which at this locality reach up to 7 km wide and
extend through an area with relief <20 m. A hilly area with
Paleozoic rocks displaying altitudes up to 300 m occurs
surrounding the floodplains.
Fig. 3. Stratigraphy of the Itaituba fossil site, displaying the main characteristic megafauna-bearing sedimentary units formed from late Pleistocene upon a
basement with Paleozoic rocks.
291
Today, the central Amazonia is characterized by a humid
to subhumid, equatorial climate, with well-defined dry (June
to November) and rainy (December to May) seasons. The
registered mean annual precipitation is 2000 mm and the
mean temperature is 35j. Vegetation pattern is complex,
ranging from dense to open tropical forests, as well as areas
of cerrado.
Sedimentology of the fossil beds
Facies description
The Itaituba megafauna occurs within a thin (<2 m thick)
sedimentary package that directly overlies Carboniferous
limestone of the Itaituba Formation. A sharp erosional
contact occurs between these deposits, forming an uncon-
formity locally marked by a surface with erosional relief of a
few meters and with microkarstic structures that form
dissolution holes up to 15 cm deep (Fig. 2). The succession
with the megafauna also occurs laterally at the same
horizontal level as the limestone. In this case, the edge of
the Itaituba limestone is defined by a ragged, sharp surface
displaying straight, vertical segments (Fig. 2), which sug-
gest fault displacement. Although detailed tectonic studies
in the Itaituba area are still unavailable, regional works attest
that the Amazon area from Manaus to Belém displays a
variety of faults formed by a relatively young strike –slip
tectonism, which took place during the Miocene/Pliocene
and even more recently during the late Pleistocene/Holocene
(Bemerguy, 1997; Costa and Hasui, 1996).
Three sedimentary successions were distinguished in the
Itaituba fossil quarry (Fig. 3). The lowermost unit (unit I) is
up to 30 cm thick and rests directly on the basal unconfor-
292 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300

Table 1 mixed with the carbonaceous matrix. Ground sloth bones,

List of bone elements of Haplomaston waringi and Eremotherium
including several parts of four incomplete E. laurillardi (one
laurillardi collected from the Itaituba site
newborn and three adults; Fig. 7; Table 1), occur within this
Cranial elements Post cranial elements
facies, being particularly concentrated at its base (Fig. 8).
H. waringi Isolated molars Not preserved The fossiliferous bed of unit II contains a large amount of
and fragments of
wood fragments, with individual logs reaching up to 15 cm
mandible skull
E. laurillardi Partial skull and Incomplete acropodial, long and 3 to 4 cm in diameter. The amounts of both fossils
jaw fragments of stylopodial, pectoral, and pebbles decrease upward, which is followed by an
three adult specimens and pelvic girdle elements increase in the amount of matrix, thus characterizing an
and one newborn plus several isolated vertebrae overall normal grading. The upper facies in unit II consists
individual and ribs belonging to four
of massive, carbonaceous mudstone bearing disperse quartz
individuals (3 adults and
1 newborn). Complete and pebbles and shells, the later sourced from the basement,
broken bones are identified as represented by Paleozoic rocks of the Itaituba Formation.
the following: femur, humerus, The mudstone displays a lighter color relative to the lower
tibia, fibula, ulna, radius, facies of this unit, due to a lower content of carbonaceous
scapula, pelvis, calcaneum,
debris. Fine-grained quartz sands are dispersed in the
astragalus, and other bones of
the manus and pes. mudstone. The contact between the conglomerate and the
mudstone in unit II is gradational, and the clay minerals
consist of kaolinite and illite, with only minor amounts of
mity. It consists of a sandy and a light-colored clay facies smectite (Figs. 5B and 6).
with a very low content in plant debris. The sandy facies is The uppermost unit III averages 20– 25 cm thick and
white to yellowish and comprises well-sorted, fine-grained, consists of black, highly organic mudstone. This unit con-
massive sands. The overlying clay facies is light gray to trasts with the underlying ones, as it does not contain neither
greenish and has no observable sedimentary structures. vertebrate fossils nor quartz pebbles, and it is characterized
Disarticulated bone fragments of the mastodon H. waringi by a high content of organic matter. The clay content is the
(Table 1; Fig. 4) were found concentrated at the base of this highest of the whole succession, being represented by up to
clay facies. The top of unit I is sharp and defined by vertical, 95% of the grain sizes. Similar to the underlying units, this
wedge-shaped holes that average 10 cm deep. These cavities succession also shows smectite, illite, and kaolinite, but it is
were filled with sediments derived from the overlying bed. interesting to notice that the relative proportions among
Grain size analysis shows that the contents of clay and silt these minerals are lower compared to unit I, where smectite
fractions in unit I reach up to 86%. Analysis of X-ray clearly dominates (Figs. 5C and 6).
diffraction of the clay minerals revealed the presence of
smectite, illite, and kaolinite, with the first being by far the Depositional history
dominant one (Figs. 5A and 6).
The intermediate unit II averages 1 m thick and consists The Itaituba fossil quarry is located in the central intra-
also of two facies. The lowermost one is represented by cratonic Amazon Basin (Fig. 1). It records an area of great
poorly sorted, massive conglomerate characterized by quartz stability, without significant sediment preservation since the
and, subordinately, limestone pebbles. Pebbles vary in size, Paleozoic, when the Itaituba Formation was formed in a
but are usually <3 cm in diameter, and they are sub- to well- shallow marine setting. It is possible that after this time,
rounded. The matrix is composed of mud and a high volume sedimentation returned to this area only at the end of the
of plant debris, which gives a black color to this facies. Pleistocene, when the sedimentary succession described
Granules and fine- to medium-grained quartz sands are here was formed. The sharp boundary between these depos-

Fig. 4. Fossils of H. waringi found at the base of unit I in the Itaituba fossil quarry, illustrating (A) a lateral view of a mandible and (B) an occlusal view of a
tooth. Scale bar, 5 cm.
 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300 293

Fig. 5. Results of X-ray analysis of clays from sedimentary units (A) I, (B) II, and (C) III in the Itaituba fossil quarry (I, illite; S, smectite; K, kaolinite).

its and the underlying Paleozoic rocks, forming vertical
straight segments, as well as the record of numerous fault
traces in the area as indicated by regional studies, suggest
that sedimentation might have been renewed due to fault
reactivation associated with the late Pleistocene/Holocene
phase of strike –slip deformation (Costa and Hasui, 1997).
Fault processes would have created traps in the limestone,
where the deposits and the megafauna of mammals accu-
mulated. Initially, the rate of fault displacement might have
been reduced, creating a shallow depression, where low-
energy sediment deposition took place. During this time
(Figs. 9A and 9B), a thin sedimentary succession, repre-
sented by unit I, was formed most likely through streams
(sandy facies) and then a small lake or pond (clay facies).
The low-energy conditions prevailing in the latter would
have been ideal for preservation of the H. waringi remains.
The abundance of smectite and illite relative to kaolinite in
these beds is suggestive of deposition under climates rela-
tively drier than for the upper units.
After lake formation, there was a period of nondeposi-
tion, when the lake surface was exposed to a period of
subaerial exposure (Fig. 9C), as interpreted from the vertical
wedge-shaped holes at the top of unit I, attributed to root
development. A renewed period of sediment accumulation
took place, forming a thicker succession, represented by unit
II. This unit formed under high-flow energy conditions, as
indicated by its high volume in quartz pebbles. The lower
facies records maximum flow energy when, together with
294 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300

Fig. 6. Relative proportions in clay minerals comparing the three

sedimentary successions of the study area, based on peak intensity as
indicated by the X-ray diffractometers.

the pebbles, a huge amount of vertebrate bones and plant

fragments, as well as quartz sands and muds, were all
brought together and deposited into the basin as debris
flows (Fig. 9D). As flow energy decreased, normal grading
was developed. Debris flows occur in most climatic regimes
but they are usually initiated in slope areas after heavy
rainfall (Leeder, 2001). The four incomplete skeletons of
ground sloths, including three adults and a juvenile, mixed Fig. 8. A detail of the base of unit II, with remains of E. laurillardi. Note
with a high volume of other disarticulated bones, suggest also abundant quartz pebbles, which decrease in size upward, forming
accidental death as the most likely, which together with the normal grading.

Fig. 7. Examples of fossils of E. laurillardi found in unit II of the Itaituba fossil quarry, illustrating (A) a calcaneum, (B) a dorsal view of a skull from an adult,
and frontal views of skulls from (C) an adult and (D) a newborn. Scale bar, 5 cm.
 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300
Fig. 9. Proposed reconstruction of landscapes for the Itaituba area, as
indicated by the depositional units described in this paper. (See text for
explanations).
occurrence in the debris flow deposits, is consistent with a
catastrophic event. An event such as a flash flooding might
have promoted slope instability, death of the ground sloths,
and their transportation into the basin together with the other
debris. Considering the proposed fault displacement as the
origin for this Pleistocene basin, an alternative explanation
295
would be that these debris flows were favored by a combi-
nation of fast flooding and fault reactivation, which created
space to accommodate the three described sedimentary
units. Holocene clay – pebbly conglomerate beds rich in
reworked Tertiary fossil vertebrates extensively exposed in
western Amazonia have been attributed to a catastrophic
flooding resulting from the sudden draining of glacial Lake
Titicaca (Campbell and Frayley, 1984; Campbell et al.,
1985). Although we do not have evidence to support their
model, it is interesting to note that coarse-grained deposits
would be expected in such a flooding event and the high
amount of quartz pebbles associated with the debris flow
deposits from the base of unit II could be a record of such a
large-scale phenomenon. However, the data presented in
this study are local and allow only speculation about the
origin of the flooding event.
The upper facies in unit II is also attributed to debris
flows, but this deposit differs from the underlying one as it
bears a higher amount of matrix, has only dispersed verte-
brate fossils, and shows a high content of shell fragments
eroded from the Paleozoic basement. These characteristics
point to a genesis related to waning flows as the most likely.
The high content in plant debris supports a vegetated area,
and the prevalence of kaolinite relative to the other clay
minerals suggests deposition during a period of higher
humidity compared to the underlying stratigraphic unit.
The sharp, undulating surface at the top of unit II
suggests another period of nondeposition. This was fol-
lowed by the accumulation of a thin succession, represented
by the black clay in unit III. This deposit records the return
to basin stability and sediment accumulation in low energy
areas of the floodplain, which was then densely vegetated
(Fig. 9E). The high amount of organic matter and clay
minerals (up to 96%) is consistent with this interpretation.
The low proportional difference between smectite and illite
relative to kaolinite and the high content in plant debris
point to an environment developed under high humidity
(e.g., Tucker, 1981; Chanley, 1989).
Radiocarbon dating
Four samples were dated at the Beta Analytic Radiocar-
bon Dating Laboratory (Table 2). Carbon from samples 1
and 2, corresponding to the Haplomastodon and Eremothe-
rium derived from depositional units I and II, respectively,
was extracted from collagen using alkali (NaOH) washes,
reduced to graphite (100%C), and dated by accelerator mass
spectrometer (AMS). The first sample might include some
exogenous carbon within the collected organics due to
degradation of bone protein, but sample 2 provided accurate
measurement. Samples 2 and 3, corresponding to wood and
organic sediments derived from depositional units II and III,
respectively, were dated by scintillation spectrometer. The
first sample was pretreated with acid to remove carbonates
and weaken organic bonds, washed with alkali to remove
296 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300

Table 2 28x and from 12 to 28x, respectively (Merwe, 1982;

Conventional and AMS radiocarbon dates of the Itaituba quarry samples.
Tieszen, 1991). This allows making a distinction from
14 13
Sample Dep. Type of C yr B.P. C/12C Cal year B.P. forest- to grass-dominated vegetation, an approach that has
unit material (x)
been used to characterize modern and ancient vegetation
4 III Wood 4620 (F60) 29.6 5570 – 5540; patterns (e.g., Merwe and Medina, 1991; Bird et al., 1992;
5470 – 5270;
Magnusson et al., 2002; Kastner and Goñi, 2003). However,
5170 – 5070
3 II Wood 37,700 (F540) 31.3 Outside the empiric studies have shown a broader range of values, and
calibration range even overlaps, within plant groups (Medina et al., 1986).
2 II Bone 11,340 (F50) 26.9 13,760 – 13,700; For the particular case of Amazonia, depleted values as low
collagen 13,470 – 13,140 as 37x have been recorded for the undergrowth vegeta-
1 I Bone 15,290 (F70) 28.5 18,730 – 17,860
tion and 31.5x for the upper canopy (Merwe and Medina,
collagen
1991).
Stable carbon isotope was measured in materials derived
secondary organic acids, and then combined with acid again from the three stratigraphic units of the Itaituba site (Table
to provide more accurate dating. Sample 4 was repeatedly 1). Collagen from H. waringi collected in unit I indicates
washed with acid (HCl) to ensure the absence of carbonates. y13C value of 28.5x. Unit II shows values of 26.9 and
The 14C results confirm that, during the past 15,000 yr, 31.3x for collagen from E. laurillardi and the reworked
sedimentation in the Itaituba quarry did not take place as a wood fragment, respectively. Organic debris derived from
continuum, but through different episodes of deposition unit III gave a y13C value of 29.6x. Before interpreting
alternated with erosion. The bone fragment of H. waringi these data, the collagen – diet spacing must be considered. In
collected from unit I was preserved within mud deposits general, the ratio in bulk plants is transferred to higher
formed in a low-energy setting (i.e., lake or pond) and does trophic levels, but the absorption of carbon by the collagen
not show any evidence for reworking. The age obtained by may vary according to metabolic rates, food preferences,
AMS indicates 15,290 (F70) 14C yr B.P. Despite the body size, and, possibly, phylogenetic distances (e.g.,
possibility that some exogenous carbon might have been Merwe, 1982; Merwe and Medina, 1991; Tieszen, 1991).
added to this sample, given some degradation of the bone The collagen –diet relationship of extinct megafauna can be
protein attributed to subaerial exposure during formation of only estimated. Considering the enrichment of collagen
a discontinuity surface at the top of unit I, the indicated age relative to diet for large size browsers of +5.3x (Merwe
is consistent with the stratigraphic position of this sample. and Medina, 1991), the isotope data obtained from the
The two samples from unit II provided very distinctive Haplomastodon and Eremotherium of the Itaituba site
radiocarbon ages. Hence, sample 2, corresponding to bone indicate a C3 vegetation corresponding to 33.8 and
material of E. laurillardi, indicated 11,340 (F50) 14C yr 32.2x, respectively. Considering these corrections, the
B.P, while the wood fragment was dated at 37,700 (F540) y13C values of the Itaituba site, ranging from 28 to
14
C yr B.P. Because the analyzed ground sloth bone came 33.8x, indicate the prevalence of C3 plant types in this
from an articulated specimen, and bone protein was excep- central Amazonian area at least for some of the past 37,700
tionally well preserved, it is concluded that deposition of yr. That is not to say, however, that a dense forest vegetation
unit II took place at 11,340 (F50) 14C yr B.P. The wood would have dominated through this time, as arboreal sav-
sample indicating a much older age was clearly reworked annas with more than 40% of tree cover might display y13C
from older deposits underlying the studied sedimentary values that are undistinguishable from forest values (Merwe,
succession, or from nearby depositional sites, during mass 1982; Magnusson et al., 2002). Taking this into account, the
failure through debris flows. Unfortunately, the small pieces y13C isotope results must be used in integration with
of this material did not allow further studies concerning the geological and paleontological data in order to provide a
type of vegetation. full discussion of paleolandscapes in the study area, as
Unit III was deposited much more recently, as revealed presented below.
by radiocarbon analysis of its organic content, indicative of
4620 (F60) 14C yr B.P. These data are consistent with the
presence of a discontinuity surface at its base, which is The use of megafauna as a paleoecological indicator
attributed to another period of nondeposition and/or erosion.
Two major points must be taken into account regarding
the usefulness of megafauna as past environmental indica-
y13C isotope data tors. The first is related to definition of paleoecological
variables, such as of broad or specific preferences for open
y13C data have been increasingly used as an important and/or closed habitats. The second is related to the historical
tool for reconstructing ancient landscapes regarding C3- and control of events, which poses the major restriction in
C4-dominated plants. This is possible because, in general efforts of timing correlation between the different megafau-
the y13C values of C3 and C4 plants range from 26 to na sites.
 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300
The megafauna association, mostly including large mas-
todons (Haplomastodon) and ground sloths (Eremothe-
erium), is common in several Brazilian sites and has been
used to support changes from closed to more open habitats
during the late Cenozoic in northern South America (Rancy,
1991). Studies based on the analysis of body design led to
relate mastodon with a savanna-like environment and Ere-
motherium with a forest edge (Webb and Rancy, 1996;
Rancy, 2000). Based on this information, it has been
proposed that such mammals are reliable indicators for
reconstructing past Amazon landscapes. However, four
main problems must be considered: (1) most of the studies
have solely reported the occurrence of taxa, without precise
dating; (2) the data available have included only transported
fossils along riverbanks, which do not allow one to test the
hypotheses on the timing and mode of environmental
changes in northern South America; (3) the findings of
Pleistocene mammals in the Brazilian Amazon, especially
mastodons and ground sloths, are concentrated in the
western and southern marginal areas, leaving a void of
information about the advance and retreat of open-environ-
ment/savanna-like habitats in the central and northern por-
tions of the basin; and (4) findings regarding the combined
association of Haplomastodon and Eremotherium have been
regarded as a stratigraphical marker for the late Pleistocene/
Holocene (Rancy et al., 1984), which has led interpretations
of regional geographical correlation to be so far highly
speculative.
Furthermore, the use of megafauna elements as habitat
indicators (Owen-Smith, 1988) has been questioned on the
basis of the statement that large mammals, like tapirs,
may develop adaptive capability to different types of
landscapes (Colinvaux et al., 2000). In fact, some modern
African proboscideans, which are modern counterparts of
mastodons, occupy a variety of habitats, including open
savanna, wet marsh, thorn bush, semidesert scrub, and
even deep forest, and the Asiatic elephant, which also
shows large body size, varies in habitat from grassy plains
to thick jungles (Nowak, 1999). The wide distribution for
such large-size animals, and mostly their record in deep
forest habitats, leads us to review the overall assumption
that mastodons were actually restricted to savanna-like
habitats.
However, one must take into account the large differ-
ence in sizes among the megafauna associated with the
Pleistocene deposits throughout the Amazon, such as
ground sloths, mastodons, glyptodonts, pampatheres, tox-
odons, camelids, large-sized capybaras, and litopterns,
which is in great contrast to the small size range of
the modern mammals that inhabit the dense Amazon
forest, like tapirs and artiodactyls (Webb, 1991; Webb
and Rancy, 1996; Rancy, 2000; Croft, 2001). This
strongly supports the presence of relatively more arid
climate regimes and the existence of large areas with
more open, though not necessarily grass-dominated, Pleis-
tocene environments.
297
A similar approach is provided in the case of terrestrial
sloths. As opposed to mastodons, there is no modern
analog for these animals, thus any effort toward ecological
reconstructions must rely on its geographical distribution,
fauna associations, and inferences from biomechanical
information. Eremotherium displays, with the modern
puma (Felis concolor), one of the largest latitudinal
distributions of a mammalian species in America (Hoff-
stetter, 1982; Toledo, 1986; Cartelle and de Iullis, 1995).
Pleistocene occurrences of this terrestrial sloth are known
from southeastern North America to southern Brazil. The
other sister taxon, Megatherium, was restricted to southern
South America with northern limits at the central portions
of the Andean valleys. There is a consensus among most
of the authors (e.g., Toledo, 1986; Cartelle, 1999; Webb,
1999) that the large herbivores were mixed feeders and
that the pan-American megafauna taxa inhabited a mosaic
of savanna with forest patches. The broad latitudinal and
altitudinal biogeographic distribution suggests that Eremo-
therium occupied a wide range of ecological habitats and
might have fed on a variety of plant types sourced from
gallery forests, open woodlands, and shrub-covered areas.
This is particularly suggested on the basis of morpholog-
ical features of dental patterns and postcrania, which
resulted in a combination of a unique body design and
large size (up to 6 m in length). Such adaptations include
a doubled parallel chisel-like tooth wear pattern adapted
for cutting/crushing of foliage, twigs, and possibly fruits;
long anterior limbs and large-clawed manus, which were
very efficient for branch reaching while displaying a
bipedal stance and particularly for defense, avoiding
predators in open environments (a frequent incursion in
dense forests would increase the vulnerability against
ambush predators such as large cats); and long tongues
frequently used in the process of food gathering. The
hairy and thick skin was adapted for protection against
plant structures such as thorns and needles. Such charac-
teristics suggest that these ground sloths were better
adapted to open vegetation communities than to dense
forests.
In addition, paleontological data support that megatheres
displayed a gregarious social behavior. This is shown by the
Itaituba ground sloths and other findings, such as the one
from the Toca dos Ossos in the State of Bahia, central
Brazil, where a large number of complete skeletons were
recovered from a natural trap cave (Cartelle and Bohórquez,
1982). These characteristics are strong evidence to discard a
closed canopy forest as the preferential habitat of these
animals, since group behavior among large mammals in
modern habitats is more frequently observed in open
habitats.
Finally, it has been demonstrated that Eremotherium
had a high browser adaptation (Toledo, 1998), as sug-
gested by its giant size, the large claws of the upper
members, and the possibility of standing in the upright
position.
298 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300
Habitat evolution in central Amazonia: an integrated
approach
Evolutionary models relating to the Amazonian biodi-
versity are based on the assumption that global climate
during the Quaternary was the major driving force of
habitat modification, which affected and, at the same time,
enhanced the changes in precipitation, depositional/ero-
sional levels, and, ultimately, the geomorphologic frame-
work. Despite a recent proposal favoring a continuous
forest cover throughout most of the Amazon region since
the mid-Cenozoic (Colinvaux and Oliveira, 2001), a mul-
titude of scientific fields, coming mainly from geology,
palynology, genetics, and biogeography, has led to the
general acceptance that the Amazonian ecosystem experi-
enced several alternating wet and dry periods, which
resulted in forest and savanna expansion, respectively
(Mörner et al., 2001; van der Hammen, 2001; Haffer,
2001). Among these data, pollen analysis has been so far
the only reliable source from the fossil record, providing
information about Pleistocene paleoenvironmental changes
in the Amazonia with precise age control (Absy et al.,
1991; Sifeddine et al., 2001). However, the pollen record
is still scarce and spotty, providing only a broad picture of
what happened during the major ecological shifts of glacial
and interglacial periods.
There are few studies of the Amazonian megafauna with
precise stratigraphic control (e.g., Rancy, 1991; Latrubesse
and Rancy, 1998). These works have been crucial to support
dry periods, with savanna expansion in the western Ama-
zonia. This paper represents the first documentation based
on an integrated approach using paleontology, sedimentol-
ogy, and radiocarbon and isotope data that allow insights
into past habitats from the latest Pleistocene in a central
Amazonian area. This time was characterized by a world-
wide drop in sea surface temperature of 1 – 4j, with the
proposed impact in low-latitudinal areas, such as the Ama-
zonia, being represented by several periods of dry climate,
with changes in river discharge and sedimentation and
development of savanna vegetation (van der Hammen,
2001).
The data collected from the Itaituba site do not appear to
indicate any drastic landscape changes at least during the
past 15,000 yr, although slight variation in vegetation
density seems to have taken place through this time. The
y13C data, as discussed above, unequivocally discard any
significant contribution of tropical grassland savanna vege-
tation, recording instead a landscape dominated by C3
plants. However, the low content in plant debris observed
in the muddy unit I is more consistent with scarce vegetation
15,000 yr ago, suggesting a period with a tendency to
aridity. It is interesting to notice that the abundance of
smectite relative to the other clay minerals is consistent
with an area undergone to low humidity. In addition,
although probably not exclusively, Haplomastodon has been
preferentially found in association with open paleohabitats.
Based on the combination of these data, we envisage a
landscape with arboreal savannas for the study area 15,000
yr ago.
It is appropriate to include a brief discussion on the
potential paleoenvironmental significance of the wood as-
sociated with the fossil sloths. This is because the obtained
age of 37,700 14C yr B.P. is close to those from many other
Amazonian sites, which have been related to open habitats
(e.g., Rasänen et al., 1990; van der Hammen et al., 1992;
Latrubesse and Rancy, 1998). As previously mentioned, the
wood fragment dated here was reworked from older depos-
its, being derived either from underlying beds or from
nearby depositional sites. In either case, it records a period
of sedimentation taking place before deposition of the
studied deposits. The y13C value of 31x could suggest
a dense forest rather than open habitats in this central
Amazon area at 37,700 14C yr B.P., but this interpretation
is biased considering the reworking nature of this material
and the absence of any further information related to this
depositional time.
At 11,340 (F50) 14C yr B.P., the landscape seems to
have remained similar, though the humidity might have
been slightly enhanced. This is suggested by the relative
increase in kaolinite relative to other clay minerals in unit II.
The high content of logs and plant debris in this unit could
also be a further support for the proposed increased humid-
ity. However, the deposits with abundant quartz pebbles
attributed to debris flows conform to the presence of open
land areas. This information suggests an environment with
arboreal savannas similar to the one that occurred at 15,000
yr ago. The presence of E. laurillardi in unit II is consistent
with a landscape with abundant trees, as this ground sloth
genus had a high browser nature. The possibility of a diet
including upper canopy leaves could explain the slightly
heavier y13C values of the giant sloths, as in a same area the
undergrowth vegetation is usually depleted in y13C relative
to the upper canopy (Merwe and Medina, 1991). Finally, as
shown in the foregoing discussion, the presence of Eremo-
therium in these deposits is in itself highly suggestive that a
dense forest was not present yet in the Itaituba area around
11,000 yr ago, when open woodlands seem to have domi-
nated the habitat.
At 4620 (F60) 14C yr B.P. even moister conditions seem
to have prevailed, with the establishment of a forest vege-
tation similar to that seen today in the Itaituba area. The
y13C value of 29.6x for organic soils indicates a C3 tree
cover (values obtained from modern Amazon soils are
usually less than 27x according to Magnusson et al.,
2002). However, the high amount of organic matter and
clay minerals, the latter with low proportional difference
between smectite and illite relative to kaolinite, is consistent
with an environment developed under higher humidity than
those recorded for the underlying units. Hence, it is pro-
posed that after sediment deposition through debris flows
(unit II), there was a return to low-energy deposition, with
development of ponds and/or marshes along floodplains in a
 D. de Fátima Rossetti et al. / Quaternary Research 61 (2004) 289–300
forested area undergoing higher humidity than in the previ-
ous time intervals recorded here.
While pollen data from deep-sea fan hemipelagic and
continental shelf sediments through the past 50,000 yr
support that the Amazon Basin forests were not extensively
replaced by savanna vegetation during the glacial period
(Heberle and Maslin, 1999; Kastner and Goñi, 2003), other
sources of information led to the proposal of drastic changes
in environmental conditions during this time interval. A
wide array of data from different fields of biology and
geology has pointed out successive periods of dry and wet
climate throughout the Pleistocene and on, which resulted in
changes from savanna to rain-forest vegetation. For in-
stance, in a seminal work revising the hypotheses to explain
the origin of species in Amazonia, Haffer (2001) has
eloquently put strong arguments for drastic changes in
climate pattern supported by a large body of hard evidence.
In addition, pollen data from the Carajás area shows a period
of dry climate with savanna vegetation between 25,000 –
10,000 yr ago (Absy et al., 1991) and 22,000 – 13,000 yr ago
(Sifeddine et al., 2001). According to these authors, heavy
rainfall and high sediment inflow with variable lake levels
and low organic carbon seem to have prevailed between
13,000 and 10,000 yr ago, as a result of climates transition-
ing from arid to humid. From 10,000 to 8000 yr ago there
was a relative increase in humidity, which was followed by
drier conditions up to 4000 yr ago, when humidity returned,
giving rise to development of rain forests (Sifeddine et al.,
2001). Studies along the Rio Negro record abundant sus-
pended load, with formation of white water between 14,000
and 4000 yr ago, attributed to increased erosion during
relatively more arid conditions (Latrubesse and Franzinelli,
1998). Only after this time there was a change to black
waters, characterized by low suspension load and high
organic content as we see today.
Our integrated analysis supports a progressively in-
creased humidity in the Itaituba area through the past
15,000 yr, which was directly reflected by a simultaneous
change in vegetation cover from arboreal savanna to dense
rain forest as we see today. These data add new insights to
the discussion of Amazonian climate and landscape during
the Quaternary, an issue still largely controversial.
Final remarks
The available historical data related to climate evolution
throughout the Pleistocene and Recent in the Amazon
region are still insufficient to provide a detailed character-
ization of all shifting episodes. There seems to be an overall
fairly good agreement on the major climatic patterns when
data from the study area are compared with those from other
places located thousands of kilometers apart throughout
northern South America, with a change from arid to rela-
tively more humid conditions in the past 15,000 yr. How-
ever, the data from the Itaituba area do not reveal any drastic
299
change in landscape in central Amazonia. Instead, integra-
tion of geological, paleontological, and isotope data
revealed only a slight change in humidity and, as a result,
vegetation density, with a shift from arboreal savanna to
forest. Such interpretation can be easily accommodated with
the data obtained from Amazon deep sea fan sediments
(e.g., Heberle and Maslin, 1999; Kastner and Goñi, 2003),
as arboreal savanna would show y13C values that might be
indistinguishable from those of forest vegetation. The in-
formation provided in this paper should be taken as an
additional source of data in the design of large-scale climatic
scenarios, but a word of caution must be considered in the
interpretations, as the modern landscape in the Central
Amazônia area is complex, with open and dense forest
and even areas with savannas.
Acknowledgments
We acknowledge the support given by Dr. Ima Célia
Vieira as the chair-in-charge of the Goeldi Museum during
the initial fieldwork. We thank Mr. Antônio Anildo Aguiar
and Mr. Joelson Aguiar, who first reported the fossil
occurrence on their farm, geologists Every Aquino (DNPM)
and Elias Leão Moraes (SEMMA) for field assistance, and
to Dr. Walter Neves (USP) for payment for one 14C analysis.
Finally, we thank two anonymous reviewers for comments
that improved this paper significantly.
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