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Quaternary Research
journal homepage: www.elsevier.com/locate/yqres
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).
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
208
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).
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
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
210
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
Carbon content
(% by wt.)
13CVPDB 0.1
Calibrated date AD
(2)
SUERC-43149
SUERC-34151
SUERC-43150
743 38
1061 37
1739 35
0.6
0.43
0.3
19.9
18.4
17.8
1297 82
960 65
311 89
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.
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.
212
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).
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
214
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
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
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savanna.
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