Quaternary Research 80 (2013) 207217

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Quaternary Research
journal homepage: www.elsevier.com/locate/yqres

Pre-Columbian landscape impact and agriculture in the Monumental Mound region

of the Llanos de Moxos, lowland Bolivia
Bronwen S. Whitney a,, Ruth Dickau b, Francis E. Mayle a,c, J. Daniel Soto d, Jos Iriarte b
a

School of Geosciences, The University of Edinburgh, Drummond St., Edinburgh EH8 9XP, UK
Department of Archaeology, Laver Bldg., North Park Rd., University of Exeter, Exeter EX4 4QE, UK
Department of Geography and Environmental Science, University of Reading, Whiteknights, Reading RG6 6AB, UK
d
Museo de Historia Natural Noel Kempff Mercado, Universidad Autnomia Gabriel Ren Moreno, Av. Irala 565, Casilla 2489, Santa Cruz, Bolivia
b
c

a r t i c l e

i n f o

Article history:
Received 26 April 2013
Available online 13 July 2013
Keywords:
Pre-Columbian
Archaeology
Bolivia
Amazon
Pollen
Phytoliths
Maize
Savanna
Anthropogenic res
Charcoal

a b s t r a c t
We present a multiproxy study of land use by a pre-Columbian earth mounds culture in the Bolivian Amazon. The
Monumental Mounds Region (MMR) is an archaeological sub-region characterized by hundreds of pre-Columbian
habitation mounds associated with a complex network of canals and causeways, and situated in the forestsavanna
mosaic of the Llanos de Moxos. Pollen, phytolith, and charcoal analyses were performed on a sediment core from a
large lake (14 km2), Laguna San Jos (1456.97S, 6429.70W). We found evidence of high levels of anthropogenic
burning from AD 400 to AD 1280, corroborating dated occupation layers in two nearby excavated habitation
mounds. The charcoal decline pre-dates the arrival of Europeans by at least 100 yr, and challenges the notion
that the mounds culture declined because of European colonization. We show that the surrounding savanna soils
were sufciently fertile to support crops, and the presence of maize throughout the record shows that the area
was continuously cultivated despite land-use change at the end of the earth mounds culture. We suggest that burning was largely conned to the savannas, rather than forests, and that pre-Columbian deforestation was localized to
the vicinity of individual habitation mounds, whereas the inter-mound areas remained largely forested.
2013 University of Washington. Published by Elsevier Inc. All rights reserved.

Introduction
New archaeological work and the use of satellite imagery (Erickson,
2000a; McEwan et al., 2001; Heckenberger et al., 2008; Prssinen et al.,
2009; Lombardo, 2010; Schaan, 2012) have revealed an ever-increasing
wealth of complex archaeological sites in Amazonia in recent years,
countering the long-held view (e.g. Meggers, 1954) that Amazonia
was a largely untouched wilderness prior to European colonization
(Denevan, 1992, 2011). The Llanos de Moxos, a vast seasonally ooded
forestsavanna mosaic of lowland Bolivia, is one such region rich in
archaeological features, which is recognized as one of the most heavily
impacted regions of Amazonia in pre-Columbian times (Denevan, 2001;
Mayle et al., 2007; Erickson, 2008; Mann, 2008; Heckenberger and
Neves, 2009; Schaan, 2012). The numerous earthworks identied in
the Llanos de Moxos are regionally distinct, and feature raised elds
(Erickson, 1995; Walker, 2004, 2008; Lombardo et al., 2011a), habitation mounds (Erickson, 2000b; Erickson and Bale, 2006; Prmers,
2009; Lombardo and Prmers, 2010; Lombardo et al., in press),
ring-ditches (Walker, 2008; Prmers, 2009; Erickson, 2010), canals
and causeways (Erickson and Bale, 2006; Lombardo and Prmers,
2010; Lombardo et al., in press), and sh weirs (Erickson, 2000a). Despite
the effort invested in cataloguing and identifying the abundance of
Corresponding author.
E-mail address: b.whitney@ed.ac.uk (B.S. Whitney).

archaeological features in the Llanos de Moxos, the extent to which

its patchwork landscape is largely natural (Langstroth, 2011) or instead
domesticated by pre-Columbian human agency (Erickson, 2006), is
still under debate.
GIS analysis of the spatial distribution of earthwork features
(Lombardo, 2010; Lombardo and Prmers, 2010; Lombardo et al.,
in press), archaeological excavations of mounds and ring-ditches
(Dougherty and Calandra, 1984; Walker, 2004; Erickson and Bale,
2006; Walker, 2008; Prmers, 2009), and botanical surveys of modern
vegetation associated with articial earth mounds (Erickson and Bale,
2006; Erickson, 2010) have attempted to determine the extent of past
human agency in shaping the landscape and vegetation of the region.
These studies, however, do not offer a temporal dimension to the question of past human impact and legacy on the current vegetation mosaic.
In this study, we used palaeoecological methods to examine past vegetation change, agriculture, and land use associated with the monumental
mound culture in the southeastern Llanos de Moxos. The temporal perspective offered through this palaeoecological investigation provides
insight into pre-Columbian impact and examines the legacy of
past human activity within the modern landscape.
Study area and archaeological context
The Llanos de Moxos is an extensive hydrological savanna and forest mosaic (130,000 km2) (Fig. 1A) that experiences seasonal

0033-5894/$ see front matter 2013 University of Washington. Published by Elsevier Inc. All rights reserved.
http://dx.doi.org/10.1016/j.yqres.2013.06.005

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B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

ooding with the passage of the South American summer monsoon.

Annual precipitation ranges from 2000 mm (north) to 1200 mm
(south). Around the city of Trinidad, capital of the Department of Beni,
annual precipitation averages around 1800 mm (Pouilly and Beck,
2004). The majority (6080%) of annual rainfall occurs between December and March in the austral summer (Pouilly and Beck, 2004),
with a dry season lasting approximately ve months from May to September. The low relief of the basin, combined with thick hard-packed
Quaternary clays (Clapperton, 1993) gives rise to seasonally inundated
savannas that dominate much of the landscape (Hanagarth, 1993).
Tropical forest occupies areas of higher relief, specically forest islands
and abandoned river leves, that rise above the level of inundation
(Langstroth, 1996, 2011). The Llanos de Moxos is characterized by distinctive geo-ecological regions, and this landscape heterogeneity is shown to
have inuenced different land-use patterns by pre-Columbian people
(Lombardo et al., 2011b).
This study focuses on the Monumental Mound Region (MMR, after
Lombardo et al., in press), which is situated ca. 50 km east of Trinidad
(Fig. 1A), where over 100 large habitation mounds, locally called lomas,
have been identied (Lombardo and Prmers, 2010). More than 40 radiocarbon dates from the excavations of two lomas, Salvatierra and Mendoza,
constrain the MMR occupation period to between AD 500 and AD 1400
(Lombardo and Prmers, 2010). Many of these sites are situated on
abandoned river leves, where pre-Columbian people not only used,
but modied and built upon, naturally-occurring higher elevation
terrain for habitation, as evidenced by dense domestic refuse and construction layers. Of the various earthworks found in the MMR, habitation
mounds required the most labour for construction and are thought to
have featured prominently in political and ritual life (Lombardo and
Prmers, 2010).
Habitation mounds are typically constructed of an elevated platform, 35 m high, but can reach up to 21 m in height and cover areas as
large as 20 ha (Erickson, 2006; Lombardo and Prmers, 2010). They
form part of a more complex, well-planned architectural design that includes several associated structures, such as pyramidal mounds, plazas,

canals, causeways, and water reservoirs. For example, at Loma Salvatierra,

the pyramidal mounds, which reach to a height of 7 m, are arranged in a
U-shaped format enclosing a plaza area on top of the main habitation
mound. These large complexes also contain hundreds of burials belonging to all occupation periods, including smaller burial mounds possibly
built for high-ranking individuals, as shown by a male burial unearthed
in Salvatierra which was accompanied by prestigious grave goods including a metal plate, ear pieces, bracelets and a jaguar teeth collar
(Prmers, 2009). In addition to these large-scale habitation mounds,
smaller forest islands feature prominently in the area, the majority of
which are hypothesized to have been once occupied (Langstroth, 1996)
or articially constructed (Erickson, 2006; Lombardo and Prmers,
2010). GIS-based site-distribution analysis of the MMR suggests that
the high number and density of habitation mounds and forest islands,
together with their linking canals and causeways, formed a complex
infrastructure that, to date, comprises the only region in the Llanos de
Moxos in which a hierarchical, regionally organized, pre-Columbian
society is clearly documented (Lombardo and Prmers, 2010; Lombardo
et al., in press).
In contrast to the raised-elds region of the northwestern Llanos de
Moxos, there are no clear landscape features in the mounds region that
could represent ancient raised-eld remains (Lombardo and Prmers,
2010). An examination of soil properties in distinct geoecological
regions of the Llanos de Moxos has revealed that, although the MMR is
seasonally ooded, its soils exhibit higher fertility compared to the
northwest of the Llanos de Moxos where the raised elds are located
(Lombardo et al., in press). The relatively base-rich Andean-derived
soils of the mound region are hypothesized to be sufciently fertile to
have supported crops (Langstroth, 2011; Lombardo et al., in press). Furthermore, Lombardo et al. (2012, in press) suggest that the numerous
identied canals and ditches (totalling 957 km) represent drainage
and irrigation channels built to control water levels among crops
grown on savannas, but this has not been veried with eld excavation
data. Macrobotanical investigations of mound sediments, and starch
grain and phytolith analyses of ceramic plant-processing tools and

Figure 1. (A) Ecosystem map of northern Bolivia showing the location and extent of the Llanos de Moxos; (B) Detailed map showing the location of Laguna San Jos amid the patchwork vegetation, as well as the location of canals, causeways, mounds, and forest islands, after Lombardo and Prmers (2010). Also shown are the locations of Lomas Salvatierra
(1) and Mendoza (2), habitation mounds from which dated excavations constrain the earth mounds culture occupation period from AD 500 to AD 1400 (Prmers, 2009).

B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

containers excavated from Lomas Salvatierra and Mendoza, revealed

that the earth mounds culture relied on a diversity of plants, including
maize (Zea mays), manioc (Manihot esculenta), yam (Dioscorea spp.),
squash (Cucurbita spp.), peanut (Arachis hypogaea), cotton (Gossypium
sp.), and palm fruits (Arecaceae) (Bruno, 2010; Dickau et al., 2012).
However, the agricultural strategy of pre-Columbian people in this
seasonally ooded landscape remains unresolved (Lombardo and
Prmers, 2010).
The abundance of higher elevation terrain once occupied by
pre-Columbian people, on which forest is currently growing, has
led to speculation that the current spatial extent of forest cover
may be articially high due to the human-made alterations in relief
in the Llanos de Moxos mosaic landscape (Erickson, 2006). Currently,
approximately 25% of the patchwork landscape in the MMR region is
situated on higher, terra rme (non-ooded) ground that supports
forest (Lombardo et al., in press). If the majority of mounds, forest
islands, canals and ditches were created for habitation and drained-eld
agriculture, pre-Columbian people may have profoundly and irreversibly
changed the relative abundance of seasonally ooded versus terra rme
terrain. Following the decline of the pre-Columbian mounds culture,
possibly related to European colonization (Denevan, 2001; Erickson,
2010), it has been argued that forest recovery on abandoned mounds
and islands may have resulted in articially high levels of arboreal cover
in the MMR (Erickson, 2006).
Aims
This multiproxy palaeoecological study, combining fossil pollen,
phytolith and charcoal analyses, reconstructs vegetation and land-use
history from prior to the mound occupation period until present to elucidate the extent of vegetation changes associated with landscape alteration for habitation and agriculture. The aims of this study are threefold:
(i) to determine the agricultural strategy of the monumental mounds
culture; (ii) to determine the extent to which the pre-Columbian
mounds people cleared naturally occurring forest during the mounds
occupation period; and (iii) to ascertain whether they augmented the
modern level of forest cover in the MMR through the past creation,
and subsequent abandonment, of habitation mounds and drained
savannas.
Methods
Study site
Laguna San Jos (referred to henceforth as LSJ, 1456.97S,
6429.70W) is located close to the excavated and well-dated habitation mounds of Salvatierra and Mendoza (Fig. 1B) (Prmers, 2009).
Archaeological investigations revealed an intense domestic occupation
of these platform habitation mounds, as indicated by the large number
of pottery shards and animal bones recovered in distinct refuse layers
(Prmers, 2009; Jaimes Bentancourt, 2012). Radiocarbon dating of excavated layers in the mounds shows that the main occupation period
occurred from AD 500 to AD 1400 (Prmers, 2009; Dickau et al., 2012).
LSJ is a large, at-bottomed lake (14 km2) with a maximum depth
of ca. 1.0 m in the dry season, with no inowing rivers or streams. It is
situated among open seasonally inundated savannas (Fig. 1B), with a
fringing strip of forest b 20 m in width, around its shore. Lomas Salvatierra
and Mendoza are located approximately 5 km and 6 km, respectively,
from the nearest lake shore. LSJ is also situated among a number of canals,
ditches and reservoirs (Lombardo and Prmers, 2010), and the nearest
canal is located b100 m from the lake (Fig. 1B).
Sediment collection and eld surveys
A 30-cm sediment core was extracted ca. 500 m from the northeast
shore of LSJ (Fig. 1B) in July 2010 using a Perspex tube and piston from

209

an anchored coring platform. The core was cut into 0.5-cm consecutive
increments, which were then stored in air tight plastic vials. Rapid vegetation surveys were conducted on the northeastern lake shore to aid
the interpretation and identication of pollen and phytolith data. Commonly encountered plant species were identied by DS and recorded
in Table 1. Voucher specimens were also collected and housed at the
Herbario del Oriente Boliviano (USZ) of the Noel Kempff Mercado
Natural History Museum in Santa Cruz, Bolivia.
Chronology
The core chronology is based on three AMS 14C dates, all of which
were obtained from non-calcareous bulk sediment. Due to the absence of terrestrial plant macrofossils and insufcient concentrations
of large (N 250 m) charcoal particles in the core sediments, bulk sediments were chosen for radiocarbon dating. The horizons chosen for
dating were based on key changes in the charcoal curve (presented
below). The three radiocarbon dates (Table 2) were calibrated using
the IntCal09 calibration curve in OxCal version 4.1 (Bronk Ramsey,
2009) and date ranges are reported with 95% condence intervals.
The age model was created through linear interpolation between
calibrated 14C dates and an assumed modern age for the core top
(sedimentwater interface) (Fig. 2). Age estimations were rounded
to the nearest 10 yr.
Multiproxy analyses
Pollen analysis was performed at 1-cm resolution, and samples measuring 1 cm3 were prepared following the standard chemical-digestion
protocol (Fgri and Iversen, 1989; Bennett and Willis, 2002), including
a sieving stage to concentrate large cultigen pollen types, such as
Z. mays (Whitney et al., 2012). Standard terrestrial pollen counts

Table 1
List of species recorded at LSJ and the vegetation types in which they were located. Also
listed is the collection number (DS) of voucher specimens for the herbarium of the
Noel Kempff Mercado Natural History Museum in Santa Cruz, Bolivia.
Collection
number

Species

Family

Vegetation type

1404
1405
1406

Inga ingoides (Rich.) Willd.

Rynchosia sp.
Ludwigia grandiora (Michx.)
Greuter & Burdet
Cecropia cf. concolor Willd.
Selenicereus cf. grandiorus (L.)
Britton & Rose
Hymenachne donacifolia
(Raddi) Chase
Ficus eximia Schott
Hydrocotyle cf. ranunculoides L. f.
Urera caracasana (Jacq.)
Gaudich. ex Griseb.
Panicum sp.
Digitaria ciliaris (Retz.) Koeler
Aeschynomene foliolosa Rudd
Pavonia corymbosa (Sw.) Willd.
Erythrina fusca Lour.
Ludwigia helminthorrhiza
(Mart.) H. Hara
Eclipta sp.
Momordica charantia L.
Hura crepitans L.
Canna glauca L.
Polygonum sp.
Marsilea sp.
Sterculia apetala (Jacq.) H. Karst.
Thalia geniculata L.

Fabaceae
Fabaceae
Onagraceae

Gallery forest
Gallery forest
Open forest

Urticaceae
Cactaceae

Gallery forest
Gallery forest

Poaceae

Gallery forest

Moraceae
Araliaceae
Urticaceae

Gallery forest
Open forest
Gallery forest

Poaceae
Poaceae
Fabaceae
Malvaceae
Fabaceae
Onagraceae

Open forest
Open forest
Open forest
Gallery forest
Gallery forest
Open forest

Asteraceae
Cucurbitaceae
Euphorbiaceae
Cannaceae
Polygonaceae
Marsileaceae
Sterculiaceae
Marantaceae

Gallery forest
Gallery forest
Gallery forest
Open forest
Open forest
Open forest
Gallery forest
Open forest

1407
1408
1409
1410
1411
1412
1413
1414
1415
1416
1417
1418

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B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

totalled 300 grains, and large pollen grains (N53 m), concentrated
through the ne-sieving methodology, were scanned for Z. mays and
other crop taxa producing large pollen, such as M. esculenta (manioc)
and Ipomoea batatas (sweet potato). Pollen identication was made
with reference to tropical pollen oras (Roubik and Moreno, 1991;
Colinvaux et al., 1999), a digital pollen catalogue (Bush and Weng,
2007), and a Neotropical pollen reference collection consisting of
N1000 specimens, which predominantly originate from lowland Bolivia,
housed at the School of Geosciences, The University of Edinburgh. Z.
mays pollen grains were identied using the criteria outlined in Holst
et al. (2007), including an examination of the distribution of intertectile
columellae using phase contrast at 1000 magnication. Equivalent
volumes of residue were scanned for large grains, as determined by
counting to approximately the same number of exotic Lycopodium
spores in each sample (Stockmarr, 1971; Whitney et al., 2012). Calculation of 95% condence intervals was achieved using a modication of
Maher's lognormal distributions in Psimpoll (http://chrono.qub.ac.uk/
psimpoll/psimpoll.html, accessed 24/04/2013).
Phytoliths were analyzed at 2- to 3-cm resolution to complement the
pollen analysis. Preparation of samples for phytolith analysis followed the
protocol outlined in Piperno (2006). Samples measuring approximately
5 cm3 were pre-treated to remove clays through deocculation, agitation,
and gravity sedimentation. Samples were divided into silt (A/B-fraction,
b50 m) and sand (C-fraction, N 50 m) fractions. A 3-ml subsample of
each fraction was digested for carbonates (36% HCl) and organics (70%
HNO3), and phytoliths were concentrated by heavy liquid otation
using ZnBr2 prepared to a density of 2.3 g/cm3. Approximately 10 mg
(not standardized) of extracted phytolith residue was mounted from
each sample. If processed samples yielded less than 10 mg, all of the residue was mounted and scanned. Entellan mounting medium was used
to permit three-dimensional viewing of phytoliths. For the A-fraction,
phytoliths were examined and described at 500 magnication. A minimum of 200 phytoliths was counted, and the rest of the microscope slide
was scanned to identify any other diagnostic types. For the
C-fraction, the entire slide was scanned at 200 magnication and
all diagnostic phytoliths counted. Phytoliths were identied using
a comparative collection of over 750 tropical and subtropical species from South America, housed at the University of Exeter
Archaeobotany Laboratory. The collection includes taxa from lowland Bolivia, which were obtained from the Herbario del Oriente
Boliviano of the Noel Kempff Mercado Natural History Museum
in Santa Cruz, Bolivia. Reference works were also consulted for the
identication of phytoliths (e.g., Piperno, 2006; Iriarte and Paz,
2009; Dickau et al., 2013; Watling and Iriarte, 2013).
Macroscopic charcoal was analyzed at 1-cm resolution to determine
past levels of anthropogenic burning. Samples measuring 1 cm3 were
rst treated with 5% (w:v) sodium pyrophosphate to disaggregate the
clayey sediments and then rinsed through 250- and 125-m sieves, as
is standard methodology (Whitlock and Larsen, 2001). The two separate size fractions (N250 m, 250125 m) were retained and counted
for charcoal fragments using a stereomicroscope. Charcoal concentrations were plotted with both pollen (Fig. 3) and phytolith (Fig. 4) stratigraphic plots so that levels of anthropogenic burning and vegetation
change can be directly compared. All stratigraphic plots were created
in C2 (Juggins, 2003).

Results
Modern vegetation
Results of the rapid vegetation survey are presented in Table 1.
Much of the survey was conducted in the thin strip of gallery forest
that surrounds LSJ, where the majority of the 23 recorded species
were found. Gallery forest trees next to the lake are of low stature, measuring ca. 1012 m, and dominant trees species include Inga ingoides
and Hura crepitans. The grass, Hymenachne donacifolia, is also abundant
in this gallery forest. The fringing marshy habitat along the lake shore is
dominated by Thalia geniculata, and Ludwigia grandiora is frequently
present. LSJ, and the gallery forest that surrounds it, are located within
open seasonally inundated savannas interspersed with forest islands
and patches of degraded forests (Fig. 1B).
Stratigraphy and chronology
The 30-cm core is composed of uniformly brown-grey clay with very
low organic content (b2% by loss-on-ignition) that grades gradually to
brown-grey clay mixed with sand near the base of the core (Fig. 3).
The absence of any abrupt lithological changes suggests that sedimentation was continuous throughout the 30-cm core, representing approximately the past 3000 yr. The sedimentation rates at LSJ are comparable
to those of other lowland Bolivian lakes of similar size, whether they are
based upon bulk-sediment dates (Mayle et al., 2000; Burbridge et al.,
2004) or terrestrial plant macrofossils (Whitney et al., 2011). There
are no reversals in the dating. The agedepth relationship for the
bottom 6 cm of core, however, was determined by extrapolation from
the lowest date, thus these oldest age estimates should be viewed
with caution.
Pollen
Pollen preservation was very poor in the lowermost 5 cm of the
core, so these samples were not analyzed for pollen (Fig. 3). Pollen assemblages in all samples are dominated by herb taxa. Poaceae (2546%)
and Cyperaceae (1428%) are most abundant throughout the core,
but Mimosa (06%), Gomphrena (04%), Chenopodiaceae/Amaranthus
(02%), and Asteraceae (27%), of which Ambrosia (01%) was counted
separately, are all consistently present throughout. Common tree pollen
types that occur throughout the core include Cecropia (920%),
Moraceae/Urticaceae (813%), Anadenanthera (04%), Celtis (04%),
Gallesia (26%), and Acalypha (03%). Palm (Arecaceae) pollen is
present throughout (13%), but often poorly preserved and in low
abundance. Calculated 95% condence intervals (not shown) demonstrate that there are no signicant changes in the percent abundance
of key pollen types throughout the record, with the exception of the
surface sample (01 cm), which contains signicantly lower abundance
of Cecropia (10%), and the lowest analyzed sample (25 cm), which contains higher Poaceae values compared to the rest of the analyzed assemblages. The poor preservation of pollen grains in this lowermost sample,
compared with the rest of the core, may explain the relatively high
abundance of robust Poaceae pollen grains in this sample.

Table 2
Details of AMS radiocarbon dating results and calibration using the IntCal09 curve.
Publication
code

Sample identier

Conventional 14C age

(yr BP 1)

Carbon content
(% by wt.)

13CVPDB 0.1

Calibrated date AD
(2)

SUERC-43149
SUERC-34151
SUERC-43150

San Jose 1213

San Jose 2222.5
San Jose 24.525

743 38
1061 37
1739 35

0.6
0.43
0.3

19.9
18.4
17.8

1297 82
960 65
311 89

B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

211

Figure 2. Agedepth model for the 30-cm sediment core from Laguna San Jos. The model is built using three AMS 14C dates calibrated using IntCal09 (Bronk Ramsey, 2009); the 2
error is presented for the calibrated dates.

Z. mays pollen is present in the majority of samples (Fig. 3), but

absent in the uppermost two samples, and is least abundant in the
oldest samples (1925 cm). Maize pollen was found in the lowest
sample analyzed for pollen, dating to ca. AD 180. Concentrations of

maize pollen are very low throughout, thus we reported only the
number of grains found in each sample (Fig. 3), which approximately equates to maize pollen concentration per 0.25 cm3 of
sediment.

Figure 3. Relative percent abundance of key pollen types, including raw count data for Zea mays grains that were retrieved using the methodology of Whitney et al. (2012). The
diagram is zoned according to charcoal data (presented on the right) to highlight the occupation period in the Monumental Mounds Region.

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B.S. Whitney et al. / Quaternary Research 80 (2013) 207217
Figure 4. Relative percent abundance of phytolith data. Rare types (b2%) are marked with (+). As shown in Figure 3, the data are zoned according to changes in the charcoal concentrations to highlight the occupation period in the Monumental Mounds Region (presented on the right of the diagram).

B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

Phytoliths
Phytolith assemblages are better preserved at the base of the core
compared to pollen. Therefore, one phytolith sample was analyzed at
29 cm (860 BC) where no pollen was preserved. Similar to the pollen
assemblages, phytolith assemblages are dominated by herb taxa.
However, the phytolith assemblages show more variability through
the core (Fig. 4), particularly with uctuations in the abundance of
Poaceae and Asteraceae. Asteraceae dominates the phytolith assemblages toward the base of the core (2765%), between 29 cm (860
BC) and 21 cm (AD 990), after which it declines to 512%, before
peaking again at 11 cm (AD 1380) (31%). Poaceae phytoliths are
less abundant at the base of the record (1329%), with a very low
value of 4% at 26 cm (80 BC). Poaceae phytoliths are highly abundant
(3249%), however, in the uppermost four samples dating from
around AD 1500 until present. The most common phytoliths of the
Poaceae family originate from Panicoid grasses, which are typical of
various terra rme and wetland habitats in tropical regions. Panicoid
phytoliths are highest in the uppermost three horizons analyzed at
6 cm, 3 cm, and 1 cm, dating from around AD 1670 until present.
Other abundant Poaceae phytoliths include Bambusoideae (17%)
and Oryzoideae (09%), the latter of which shows a small peak at
9 cm (AD 1500). Additional herb phytoliths that occur commonly
throughout the core include Marantaceae (618%), common to the
fringing wetland vegetation of the lake, and Heliconia (611%), observed throughout the wet savanna of the region.
Phytoliths of arboreal taxa show lowest abundance in the bottom
section of the core (1124%), from 29 to 21 cm, dating from approximately 860 BC to AD 1000 (Fig. 4). As is common practice, the phytolith
assemblages were not spiked with an exotic marker to determine absolute concentrations (Stockmarr, 1971), as is typical of pollen analysis.
Thus, it cannot be determined whether the low values for tree taxa at
the base of the core are caused by inated Asteraceae values within a
closed sum. The arboreal phytoliths are characterized by abundant
globular granulates, which peak at 16 cm (AD 1170), corresponding
to the middle of the mound occupation period. Arecaceae globular
echinate phytoliths are also common among assemblages (1028%),
with the exception of the lowest sample analyzed at 29 cm (860 BC)
where they do not exceed 5%. The number of burned phytoliths in each
analyzed horizon was not systematically recorded, although charred
phytoliths were found to be more abundant between 21 and 11 cm
depth (ca. AD 1000 to 1380).
Charcoal
Charcoal concentrations in both size fractions show a similar trend;
concentrations are lowest at the base of the core (Figs. 3 and 4), between 30 and 26 cm (1120 BC to 80 BC), and increase after 25 cm
(AD 180), the horizon at which the oldest maize pollen grain was
recorded. Charcoal values peak around 21 cm (AD 1000). After AD
1000, charcoal concentrations decline steadily until ca. AD 1280 after
which they remain low until present.
Discussion
Interpretation of pollen and phytolith signals
A comparison of modern phytolith and pollen assemblages from
soils and articial traps, respectively, from vegetation plots in a diversity
of tropical lowland ecosystems (Gosling et al., 2005, 2009; Burn et al.,
2010; Jones et al., 2011; Dickau et al., 2013) has shown that phytoliths
and pollen are complementary proxies for the reconstruction of tropical
vegetation. In particular, the taxonomic resolution of grass and herb
phytoliths is sufciently high to distinguish wetland and terra rme
savanna ecosystems (Dickau et al., 2013), which cannot be achieved
through pollen analysis (Jones et al., 2011). Furthermore, pollen

213

assemblages in articial traps within savannas have been shown to capture the pollen of wind-blown arboreal taxa, such as Moraceae, from
neighbouring forest (Jones et al., 2011), but in contrast, arboreal
phytoliths are shown to reect a local signal in soils sampled from the
same savanna plots (Dickau et al., 2013). In general, however, arboreal
taxa are identiable to a higher taxonomic resolution palynologically
than is achievable through phytolith analysis. In the savannaforest
mosaic of the Llanos de Moxos, therefore, the complementary combination of pollen and phytolith analyses is an excellent approach (Mayle
and Iriarte, in press) for the examination of pre-Columbian land use.
Pollen and phytolith analyses are complementary, not only in terms
of bridging taxonomic gaps inherent to each proxy, but also in terms of
their differing spatial scales of source vegetation. Phytolith assemblages
generally reect in situ decay of plant matter from vegetation growing
locally (Piperno, 1988, 2006; Iriarte et al., 2010). Although alluvial and
colluvial processes might provide a mechanism for transportation and
redeposition of phytoliths (Fredlund and Tieszen, 1994; Dickau et al.,
2013), phytoliths do not undergo the same degree of transportation
as does wind-blown pollen, for example. Thus the catchment area of
phytolith assemblages derived from lake sediments is likely to represent
the vegetation adjacent to the shoreline, particularly in large, atbottomed lakes, such as LSJ, where there is no mechanism that concentrates deposition of plant remains towards the centre of the lake.
This is borne out by the surface-sediment phytolith assemblage,
which is consistent with the inventory of local vegetation around the
lake (Table 1).
By contrast, decades of pollen catchment modelling research in
temperate latitudes (Jacobson and Bradshaw, 1981; Sugita, 1994,
2007) have shown that: i) pollen assemblages from lakes with
large surface areas, similar to LSJ, represent averaged regional vegetation (N 50 km 50 km) (Sugita et al., 1999), and ii) large lakes are insensitive to detecting patch-size vegetation change in the surrounding
landscape (Sugita, 1994; Sugita et al., 1999). The applicability of these
pollen catchment studies for our study area might, understandably, be
questioned, given the much higher proportion of wind-pollinated tree
taxa in temperate forests compared with those of the tropics. However,
we are condent of the regional representativeness of the pollen signal
from LSJ because the key arboreal pollen types in the modern (core-top)
assemblage are wind-pollinated and well-dispersed, such as Moraceae
and Cecropia (Gosling et al., 2005, 2009; Burn et al., 2010; Jones et al.,
2011). Furthermore, the most common arboreal taxa in the modern pollen assemblage are rare or absent in the shore-line forest
(e.g. Moraceae, Anadenanthera, Gallesia) (Table 1), but common constituents of the terra rme forests across the mounds region (Langstroth,
1996; Erickson and Bale, 2006). Thus, the phytolith signal in LSJ most
likely represents the local shoreline and savanna vegetation immediately surrounding the lake (b 1 km), whereas the pollen record reects the
aggregate proportions of forest versus savanna vegetation of the MMR.
The pollen data indicate that in the ca. 4500-km2 region in which
mounds and forest islands were mapped (Lombardo and Prmers,
2010), the proportion of forested terrain, which currently covers 25%
of the landscape, was largely unaltered over the past 2000 yr. Given
the large size of LSJ (14 km2), uctuations in forest cover on the scale
of individual mounds (b20 ha) or clusters of mounds could have occurred without having been detected in the pollen record (Sugita et
al., 1999), especially given that the cumulative area of mapped mounds
and forest islands is b 800 ha (Lombardo and Prmers, 2010) or b1% of
the total forested landscape. However, if a high proportion of the forest
cover was cleared during the mound occupation period, a clear decline
in arboreal pollen would be expected alongside the peak in charcoal, especially with respect to wind-dispersed Moraceae pollen, which is the
most abundant pollen type in modern assemblages of lowland Bolivian
tropical forests (Gosling et al., 2005, 2009; Burn et al., 2010). Instead,
our 2000-yr pollen record shows no signicant change in the abundance of this pollen type. Any change in the levels of forest cover, therefore, must have occurred in relatively small patch sizes, perhaps

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B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

restricted to individual habitation mounds, with forest in the intermound areas kept largely intact. Fuel for anthropogenic res might
have originated from these smaller forest clearances, but given that the
regional-scale relative abundances of forest and savanna did not change
signicantly, it is more likely that the charcoal peak originated from
extensive burning of savannas, either to clear natural savanna for initial
cultivation or to clear and maintain agricultural elds after harvest.
By contrast, the phytolith data, which likely represent shoreline
vegetation and savannas proximal to the lake, show signicant uctuations throughout the record. In particular, Asteraceae dominates the
lowest assemblages (860 BC to AD 1000) and Poaceae values rise at
the top of the core (post AD 1500). Asteraceae is often associated
with disturbed or open vegetation (Piperno, 2006). However, in an
examination of soil phytolith assemblages in a diversity of ecosystem
types, Dickau et al. (2013) demonstrated that Asteraceae phytoliths
originate from a number of sources, such as semi-deciduous and
evergreen forests, and terra rme (cerrado) and wetland savannas.
Therefore, it is difcult to interpret the high Asteraceae values that
occur at the base of the core. Given that the diagnostic Asteraceae
phytoliths are broad and at platelets, their high abundance in the
lower core section may reect the ease by which they are transported
on re draughts, compared to other phytolith morphotypes, because
the peak in Asteraceae broadly coincides with higher levels of burning.
The ecological signicance of the high abundance of Asteraceae
phytoliths, however, is largely speculative. Variations in Poaceae
phytoliths show there was a greater expanse of grass adjacent to
the lake in the most recent samples (post AD 1670). Within the Poaceae
sum, however, the levels of terra rme taxa (Bambusoideae, Chusquea)
versus the wetland grass subfamily (Oryzoideae) show little variation
through the core, which implies that the area of seasonally inundated
savannas near the lake was not signicantly altered over the past
3000 yr. Phytoliths indicative of shoreline taxa and seasonally ooded
savanna, such as Heliconia spp. and Thalia spp. (Marantaceae), a common component of the shoreline vegetation (Table 1), show little variation throughout the record, which suggests that the fringing wetlands
of LSJ were not heavily impacted by past human activity.
Pre-Columbian occupation and agriculture
The charcoal data show that the greatest change throughout the
~ 3000-yr record occurred from AD 400 to AD 1280. Charcoal particles N100 m are generally considered to represent a local burning
signal (Whitlock and Larsen, 2001), but given that LSJ has a very
large surface area, which is known to increase charcoal catchment
size (Power et al., 2010), we conservatively interpret that only the
N250-m size fraction represents local burning. The similar pattern of
change shown in both charcoal size fractions, however, implies that
higher levels of local and extra-local burning, compared to present, occurred in the same time period. The origin of these past res was likely
anthropogenic because multiple studies demonstrate that climate during the late Holocene in the Bolivian Amazon became progressively
wetter over the past 2000 yr (Mayle et al., 2000; Burbridge et al.,
2004; May et al., 2008). Therefore, the pattern of increased re from
AD 400 to AD 1280 does not follow the expected trend if climate was
the key control on burning at this time, pointing to an anthropogenic
cause for the changes in re regime at our site. Furthermore, this period
of increased burning is coeval with the dates of human occupation determined through archaeological excavations (AD 500 to AD 1400) at
the neighbouring Mendoza and Salvatierra mounds (Prmers, 2009).
We therefore argue that the charcoal peak reects the height of anthropogenic burning in the mound region.
The decline of the mound culture occupation is well-constrained by
a bulk sediment date of AD 1297 82 from the end of the charcoal
peak (12.513 cm) (Table 2). Although this represents a mid-range
age estimate for the decline in charcoal at LSJ, the 95% condence
range of the date spans AD 1220 to AD 1380, which coincides with the

terminal occupation dates determined by Prmers (2009). Our dated

charcoal curve bolsters the archaeological evidence that the mound culture declined in the pre-contact period, challenging the widely-held assumption (Denevan, 2001; Erickson and Bale, 2006; Erickson, 2010)
that the earth-moving pre-Columbian cultures of the Llanos de Moxos
disappeared as a result of European colonization and the introduction
of Old World diseases, such as smallpox, after AD 1492.
Despite the reduced charcoal values relating to the decline in the
mound culture, however, the continuous presence of maize throughout
almost the entire core indicates that people continuously lived and cultivated crops in the mound region from at least AD 180 until present.
This agrees with the rst European accounts of the southeastern Llanos
de Moxos in AD 1617, in which they noted the impressive area of elds
under cultivation and the vast quantity of maize produced (Denevan,
1966). Thus, although we show that the pre-Columbian mound culture
managed the landscape more intensively through re in the period
dating from AD 400 to AD 1280, the presence of maize pollen clearly
indicates that the region was continuously occupied through both the
pre- and post-Columbian periods. We can reject the possibility that
these pollen grains originated from wild teosinte, the pollen of which is
similar in morphology and size to that of maize (Holst et al., 2007), because teosinte is native only to Mesoamerica and does not occur naturally
in southwestern Amazonia (Doebley, 1990; Fukunaga et al., 2005).
Maize pollen grains are large (55110 m diameter) and poorly dispersed from their source plants, which means they are grossly underrepresented in lake sediments (Jarosz et al., 2003; Lane et al., 2010).
At LSJ, the recovery of maize pollen should be further hampered because the core site is located 500 m offshore and the lake is both shallow and at-bottomed, which prevents sediment focusing towards
the middle of the lake. Maize pollen, however, was found in most horizons. The recovery of this poorly dispersed pollen type from this site
suggests that the crops were grown in savannas adjacent to the lake,
and possibly along the lake shore. Our data support the interpretation
of Lombardo et al. (in press), who suggest that the relatively fertile
soils east of Trinidad, created in the mid-Holocene through the migration of the Rio Grande and with it, the deposition of base-rich Andean
sediments (Plotzki et al., 2011), were capable of supporting agriculture.
The relatively higher soil fertility east of Trinidad, compared with other
areas in the Llanos de Moxos, has been offered as an explanation for the
absence of raised elds in the mound region (Lombardo et al., 2011b, in
press), which is supported by our palynological evidence for maize
agriculture close to LSJ.
Although Dickau et al. (2012) showed that the pre-Columbian earth
mounds culture relied on a diversity of crops for food, only maize pollen
was recovered from our record. The sieving methodology employed for
this study (Whitney et al., 2012) has proven effective for the isolation of
Cucurbita spp. (squash), M. esculenta (manioc), and I. batatas (sweet potato) at other archaeological sites in the tropical Americas (Iriarte et al.,
2012; Rushton et al., 2013). However, pollen of these other staple crops
is noticeably absent from the record at LSJ, which could be due to one of
two possibilities; either crops other than maize were grown too far from
the lake shore to be detected in the record, or maize dominated the diet
of the pre- and post-Columbian societies in the mound region. Given
that Dickau et al. (2012) reported greater abundance of starch grains
and phytoliths from maize, compared with other crops, in ceramics
from neighbouring excavated mounds (Mendoza and Salvatierra), it is
likely that maize was the staple crop in this region.
Pre-Columbian landscape impact in the Monumental Mounds Region
With the exception of higher abundance of Asteraceae phytoliths
at the base of the core, which, as discussed, might signify any number
of taphonomic or local vegetation changes, the pollen and phytolith
record at LSJ displays no signicant changes associated with the
peak period of mound occupation, as indicated by the peak in charcoal and higher concentrations of Z. mays pollen. Despite the elevated

B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

levels of anthropogenic burning centred at AD 1000, there is no discernible decline in arboreal taxa in pollen assemblages associated
with the charcoal peak (Figs. 3 and 4). The large surface area of LSJ
(14 km2), however, means that the LSJ pollen signal predominantly
represents the regional-scale vegetation (N50 km 50 km) in which
the mounds are situated (Sugita et al., 1999), and the fossil pollen signal
will not detect localized patch-scale changes comparable in area to individual mounds or mound complexes (Sugita, 1994, 2007). Thus, the
static arboreal pollen curves demonstrate that any forest clearances
must have occurred at patch-size scales which were cumulatively insufcient in area to markedly reduce the regional-scale extent of forest
cover. If there had been large-scale deforestation associated with this
pre-Columbian mound culture, then one would instead expect to nd
a clear expansion in arboreal pollen due to forest recovery after the
end of the mound culture at AD 1280. This pattern, however, is not
observed in our pollen data.
Lombardo et al. (in press) argue that the pre-Columbian mound
culture altered the hydrology in the savannas through the creation
of drainage ditches and canals to irrigate elds. Although our pollen
record indicates that maize crops were grown in the vicinity of LSJ,
near which canals and reservoirs can also be identied, the phytolith
assemblages do not show any clear changes in the relative abundance
of wetland versus terra rme herb taxa coincident with the charcoal
peak and occupation as identied through archaeological excavations
(Prmers, 2009). Our phytolith data therefore imply that, although
these canal and ditch features are common on the landscape, the
pre-Columbian landscape engineering did not greatly alter the relative abundance of wetland versus terra rme habitat in the creation
and maintenance of agricultural elds in close proximity to the lake
shore. Instead, one of the greatest changes in the phytolith record occurs towards the top of the core (post AD 1670) where the abundance
of Panicoid grasses is much higher compared to the pre-Columbian
period. The latter may reect changes associated with European colonization and the maintenance of forest clearings near the lake after
the introduction of cattle in AD 1682 (Langstroth, 2011). These phytolith assemblages suggest that post-contact local clearances adjacent
to the lake might have been greater than the level of impact that
occurred during the pre-Columbian period.
The number and diversity of earthwork structures in the Llanos de
Moxos, which augmented the extent of terra rme habitat in this naturally
mosaic environment, has led many to conclude that the Llanos de Moxos is
a largely anthropogenic landscape (Erickson, 2006, 2010; Lombardo and
Prmers, 2010). In the MMR east of Trinidad, hundreds of habitation
mounds and occupied forest islands have been identied, alongside numerous canals, ditches and reservoirs (Lombardo and Prmers, 2010),
and vegetation surveys of abundant economically-important plant
species growing on and around habitation mounds (Erickson and Bale,
2006), have all supported the hypothesis that the conguration of the
landscape and its associated vegetation was heavily manipulated by
pre-Columbian people. Given the vast number of these anthropogenic
earthworks across the landscape, one might reasonably speculate that
much of the forest was also cleared during the period of earth mounds occupation (Erickson, 2006). An unexpected nding from our pollen data
from LSJ, however, is that the majority of forest across the MMR was
left standing by the earth mounds culture. We also show that the extent
of forest habitat was not altered signicantly through savanna drainage
or the creation of articial relief, as this too would have been reected
in our pollen record. Anthropogenic impact on savannas was greater, as
pre-Columbian people intensely burned and managed the savanna landscape, and the cultivation of crops likely occurred in existing savanna
areas, without necessitating the clearance of additional land. Low
re activity after AD 1280, inferred from our charcoal data, supports the assertion of Erickson and Bale (2006) that the modern
landscape reects the product of less intense human management
in recent times and a reduction in the annual burning of the
savanna.

215

Instead of altering the regional abundance of forest habitat, we infer

that the pre-Columbian earth mounds culture built many of their earthworks by augmenting the existing topography on abandoned river leves (Lombardo and Prmers, 2010). Their modication of the existing
landscape did not cumulatively create enough new terra rme habitat
outside areas of naturally higher relief to demonstrably alter levels of arboreal taxa in the pollen record. We infer that the pre-Columbian earth
mounds culture inuenced vegetation on local scales, equivalent to individual articial mounds or mound complexes, but did not deforest the
inter-mound area, which comprises the vast majority of the regional forested landscape. Although our data show that forest was not extensively
cleared in the period occupied by the mounds culture, the regional-scale
pollen record from LSJ is insensitive to detecting the extent to which
people altered the species composition of forest growing on and around
individual habitation mounds, such as the selective management to encourage the growth of economically-useful species (Erickson and Bale,
2006). Further analysis of small lakes (b1 ha), which reect localized
vegetation signals, will be required to understand the temporal changes
in composition of forest on and around the habitation mounds and forest
islands.
Conclusions
Peak levels of anthropogenic burning, inferred from charcoal data,
demonstrate that the height of the monumental mounds culture occurred
from AD 400 to AD 1280 in the Llanos de Moxos. The end of the mound
occupation and associated land-use change is well-constrained by radiocarbon dating which precedes European arrival in the New World by at
least 100 yr. The pre-contact date of the charcoal decline contradicts the
notion that earth-mound cultures of the Llanos de Moxos declined as
a result of European colonization and associated introduction of Old
World diseases (Denevan, 2001; Erickson and Bale, 2006; Erickson,
2010). We demonstrate that maize was a staple crop, as supported by
previous archaeobotanical analyses of excavated mounds that showed
maize was abundantly processed. Our results support the hypothesis
that specialized earth structures, such as raised elds, were not required
to grow maize on the relatively fertile soils of the savannas in the mounds
region. Despite the reduction in anthropogenic burning after AD 1280,
associated with land-use change and the decline of the earth mounds
culture, the region was continuously cultivated with maize through
much of the colonial period.
Aerial photographs and satellite imagery have demonstrated that
some of the greatest evidence of landscape alteration in the Amazon to
date (Denevan, 2001; Mayle et al., 2007; Erickson, 2008) comes from
the vast forestsavanna mosaic landscape of the Llanos de Moxos. Despite
this strong evidence of human modication of the Monumental Mounds
Region, contrary to expectation, we found no evidence from our pollen
data for regional-scale deforestation in pre-Columbian times. Any deforestation associated with mound occupation must have occurred at scales
too small to be detectable by the regional-scale pollen signal of our study
site, Laguna San Jos (LSJ). Furthermore, the landscape engineering that
created additional terra rme habitat also occurred at scales too small to
be detectable by the pollen record from LSJ. Instead, the inuence of
landscape engineering on vegetation likely occurred at the scale of individual articial mounds or mound complexes. However, because the pollen signal from our lake site reects regional scale forest composition,
our record is insensitive to anthropogenic forest management at and
around individual habitation mounds. Consequently, we cannot rule
out the possibly that this pre-Columbian culture domesticated their
forests by actively selecting for economically-useful species over other
less useful species (Erickson and Bale, 2006).
Acknowledgments
Research for this manuscript was supported by a Leverhulme
Trust research project grant (F/00158/Ch) awarded to FEM and JI.

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B.S. Whitney et al. / Quaternary Research 80 (2013) 207217

Radiocarbon dates were granted by the NERC radiocarbon facility to

FEM (allocation numbers 1527.1010 and 1623.0312). Fieldwork logistical support was provided by the Noel Kempff Mercado Natural
History Museum, Santa Cruz, Bolivia, and Programa de Conservacin
de la Paraba Barba Azul, Trinidad, Beni Department, Bolivia. We
thank Oscar Saavedra for his guidance and eld assistance in the
mounds region, and also Jane Bunting, University of Hull, for her valuable advice on the representativeness of pollen records from large
lakes. We also thank two anonymous reviewers whose comments
improved this manuscript.
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