Review of Palaeobotany and Palynology 120 (2002) 73^90

www.elsevier.com/locate/revpalbo

Holocene climate change and hydrarch succession in

lowland Amazonian Ecuador
Chengyu Weng a; , Mark B. Bush a , J. Stephen Athens b
a
Department of Biological Sciences, Florida Institute of Technology, College of Science and Liberal Arts,
150 W. University Blvd., Melbourne, FL 32901, USA
b
International Archaeological Research Institute Inc., 2081 Young St., Honolulu, HI 96826, USA
Received 1 February 2001; received in revised form 30 October 2001; accepted 12 November 2001

Abstract

Two sediment cores (Maxus 4 and Maxus 1) were obtained from lowland wetlands situated in the highly diverse
rain forests of the western Amazon basin, Ecuador. The cores, separated by ca. 57 km, were analyzed for fossil pollen,
charcoal and loss-on-ignition. The Maxus 4 core encompasses the last 9500 years, while the Maxus 1 core provides a
record for approximately the last 2000 years. As neither core registered evidence of human impacts or land use, an
unobscured image of Holocene vegetation and climate change was obtained. Alternating wet and drier periods on a
millennial scale characterized much of the Holocene. The unusually high values of Cecropia in the pollen sum between
8700 and 5800 cal yr BP suggest frequent disturbance or the presence of early successional habitat. After 5800 cal yr
BP a series of taxa rising and falling in prominence reveals a hydrarch succession that culminated in a relatively dry
period between ca. 4900 and 3700 cal yr BP. Seasonal flooding of the site is evident as local water tables rose between
3700 and ca. 1000 cal yr BP. After 1000 cal yr BP the modern wooded wetland was established. The second site
Maxus 1 reveals a detailed history of approximately last 2000 years. The principal feature of the record is a
disturbance event at about 1000 cal yr BP, followed by a period of hydrarch succession. 8 2002 Elsevier Science
B.V. All rights reserved.

Keywords: Amazonia; rain forest; fossil pollen; charcoal; climate change; hydrarch succession; Holocene

1. Introduction logical gradients and taxa with narrowly de¢ned

Amazonia supports the largest tropical rain for-
est system in the world. Within this vast area
there exists on all spatial scales an extraordinary
diversity of plants and animals. Fine-grained eco-
* Corresponding author.
E-mail address: cweng@¢t.edu (C. Weng).
niches combine to produce very high K- and
L-diversity (Gentry, 1988). The region of highest
K-diversity yet described is the Yasuni rain forests
of eastern Ecuador (Pitman et al., 1999). Consid-
eration of the origin of this diversity led to the
refugial hypothesis (Ha¡er, 1969) and, explicitly,
the identi¢cation of the Napo refugium. However,
Colinvaux et al. (2000) and Colinvaux (1996) re-
ject the refugial mechanism as an explanation of
the richness of species and endemics in the Napo
and other Amazonian regions. The immense bio-

0034-6667 / 02 / $ ^ see front matter 8 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 4 - 6 6 6 7 ( 0 1 ) 0 0 1 4 8 - 8

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74 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
diversity of the upper Napo drainage has made
this area a top priority for permanent protection
(Dinerstein et al., 1995).
The Amazon basin has long been home to
Amerindian peoples (Roosevelt et al., 1996), and
scattered paleoecological data provide some local
insights into land use. However, at the larger
scale, evidence is lacking regarding their in£uence
on the forests (e.g. Bush and Colinvaux, 1988;
Behling, 1996; Behling and Da Costa, 2000). If
other areas of lowland Ecuador are a guide, ripar-
ian forests could have been in£uenced by human
use and disturbance for much of the Holocene.
Human occupation of lakeside sites and the culti-
vation of maize have a long (6000 years) history
in eastern Ecuador and other regions of Amazo-
nia (Athens and Ward, 1999; Bush and Colin-
vaux, 1988; Bush et al., 1989; Piperno and Holst,
1998; Piperno, 1990; Piperno and Becker, 1996).
Human use of this region, however, abruptly de-
clined 400^450 years ago with depopulation (Por-
ro, 1994). Similar abandonment is also inferred
from records from Panama between 400 and 350
yr BP (Bush and Colinvaux, 1994; Bush et al.,
1992; Piperno et al., 1991). The coincidence of
these changes with the arrival of Conquistadors
leaves little doubt that the introduction of Old
World diseases was the primary factor (Denevan,
1976; Denevan, 1992). Given this historical rec-
ord, it is obvious that the common perception of
the Amazon rain forest as primeval and largely
untouched by humans is misleading (Bale¤e,
1988). However, the extent to which anthropogen-
ic in£uence has been a factor shaping Amazon
forests, including those of the Yasuni area, is an
important issue in Amazonian ecology and con-
servation.
Similarly, it is important to establish whether
there have been signi¢cant climatic or environ-
mental changes during the Holocene. Servant et
al. (1981) suggested a profound dry phase in Bo-
livia that purportedly caused an expansion of sa-
vanna into rainforest areas in the mid-Holocene.
To date there has been no independent evidence
to support this contention, although there have
been many reports of more modest cycles of low-
ered lake levels from Holocene records across
Amazonia (e.g. Liu and Colinvaux, 1988; Piperno
PALBO 2448 23-7-02
and Becker, 1996; Behling and Hooghiemstra,
1999).
In this paper we present two paleoecological
records that provide the ¢rst complete Holocene
fossil pollen and charcoal evidence from the Ya-
suni region of Ecuador. We also use these data to
evaluate the impact of human activity and climate
change on Yasuni ecosystems over the last 9000
years.
1.1. Study area
The study area of the present project is roughly
the northwestern arm of Yasuni National Park
(Fig. 1). The Rio Napo and the Rio Yasuni ap-
proximate the northern and southern boundaries,
respectively, of this park subregion (Yasuni Na-
tional Park extends east to the present Peruvian
border and south to the Rio Curaray; the western
border is irregular ^ see Fig. 1). The elevations of
Yasuni National Park are between 175 and 400 m
above sea level.
Our principal study site, Maxus 4 (0‡27PS,
76‡37PW), lies near the Rio Yasuni, while Maxus
1 is near the Rio Napo, the latter being one of the
major rivers of the eastern Ecuadorian lowlands
(Fig. 1). The Maxus 4 core was collected from a
marsh in the £oodplain of the Yasuni River ap-
proximately 60 km southeast of the community of
Pompeya on the Napo River (Fig. 1). The Maxus
1 core was raised from what seems to be an in-
¢lled abandoned meander (perhaps of the nearby
Rio Indillana), and is located about 3 km south-
east of the Napo River. This site is covered with
evergreen tropical rain forest. The two sites are
separated by about 57 km, and the elevation of
both sites is about 220 m above sea level.
1.2. Climate
The climate of the Yasuni area is generally wet
and warm. Average monthly temperatures are be-
tween 24‡C and 27‡C for all months and average
annual precipitation is over 3000 mm. Rainfall
occurs in all months, but April^May and Octo-
ber^November are the wettest, and December^
February and August are the driest. As in other
areas, the largest current inter-annual variation in
 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90 75

Fig. 1. Sketch map showing the location of the coring in relation to rivers and the Yasuni National Park boundary. Maxus 4
and Maxus 1 are study sites in this paper. Maxus 5 is the study site for Athens and Ward, 1999, and mentioned in this paper.

precipitation is attributable to the El Nin‹o South-
ern Oscillation. Strong El Nin‹o events cause the
dry seasons to be more marked, which in turn
causes river levels to drop. Isolated climatic events
such as windstorms sometimes cause blowdowns
that open relatively large ( s 2 ha) canopy gaps
(Foster and Terborgh, 1998; Nelson et al., 1994).
Over the last several decades natural ¢res are not
known from the Yasuni area (Pitman, unpub-
lished data).
1.3. Vegetation
The Yasuni area is covered by a diverse, tall,
closed-canopy, moist evergreen forest; it is one of
the most species-rich forests yet described. Soil
moisture plays a determining role in vegetation
distribution and local forest types are broadly
classi¢ed into terra ¢rme forests, seasonally
£ooded forests, and swamp forests according to
their hydroperiod.
Terra ¢rme forests lie above the mean high riv-
er level and seldom, if ever, £ood. The most im-
portant woody families of the terra ¢rme are
Annonaceae, Arecaceae, Fabaceae, Lauraceae,
Melastomataceae, Meliaceae, Moraceae, Rubia-
ceae and Sapotaceae. Romoleroux et al. (1997)
estimated a total of 3114 tree and shrub species
for the Yasuni National Park and the adjacent
ethnic reserve. In investigations of 58 1-ha tree
plots, between 114 and 307 species were identi¢ed
per plot (Gentry, 1988 ; Pitman, unpublished
data).
The seasonally inundated forests may be
£ooded for up to 90 days each year. The com-
monest tree in both the seasonally inundated

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76 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90

Table 1
Sediment description of the study cores
Layer Depth Sediment Average accumulation rate
(cm)
Core 4 I 0^18 dark gray silty clay, ¢ne rootlet and plant remains :
>
>
II 18^64 dark brown peat with silt, plant remains become ¢ner with 0.41 mm/yr
depth <
>
>
III 64^146 brown clay with a small amount ¢brous material 8
IV 146^252 black peat, fair coarse with abundant macro-plant remains 0.81 mm/yr
V 252^386 dark gray clay
1.1 mm/yr
VI 386^417 silty clay, abundant woody pieces 
VII 417^505 dark gray clay, grayish brown in lower part 0.98 mm/yr
VIII 505^641 dark gray clay 0.65 mm/yr
IX 641^800 silty clay with sand N/A
Core 1 I 0^100 dark brown humic silt :
>
>
>
II 100^224 dark gray humic silt with a small amount of ¢ne sand >
III 224^266 peat, no sand 4.7 mm/yr
<
>
IV 266^330 same as layer II with gradual increase in clay near base >
>
>
V 330^469 grayish brown silty clay with ¢ne sand and plant remains 8
VI 469^680 brownish gray silty clay with ¢ne sand :
>
>
VII 680^704 gray silty clay
N/A
VII 704^730 gray silt with ¢ne sand <
>
>
VIII 730^736 brown silty clay with some ¢ne sand 8

and terra ¢rme forests is the palm, Iriartea deltoi-
dea. The diversity of these £ooded forests is
slightly less than that of the terra ¢rme forest
(Pitman, unpublished data).
The swamp forests are the wettest and least
diverse of the forest types (Pitman, unpublished
data). Often they are dominated by Mauritia, a
large palm. The presence of Mauritia indicates a
consistently high water table, with standing water
often present 6 months of the year.
2. Methods
We present data from sediment cores, Maxus 4
and Maxus 1, two of a series of ¢ve cores raised
with a Colinvaux^Vonhout coring device from
wetlands alongside a petroleum access road in
eastern Ecuador in September, 1994 (Athens and
Ward, 1999). Upon opening the cores in the lab-
oratory, the sediments were described and sub-
sampled for pollen, loss-on-ignition (LOI) and ac-
celerator mass spectrometry (AMS) radiocarbon
dating. Samples of 0.5 cm3 were taken at 10^30-
cm depth intervals. Pollen samples were treated
with standard palynological procedures (Faegri
and Iversen, 1989) and a spike of exotic Lycopo-
dium clavatum spores was added to facilitate
quantitative analyses (Stockmarr, 1971). Identi¢-
cations were based on published atlases and keys
(Absy, 1979; Bartlett and Barghoorn, 1973; Co-
linvaux et al., 1999; Hooghiemstra, 1984; Roubik
and Moreno, 1991) and the pollen reference col-
lection at the Florida Institute of Technology
( s 3000 species). At least 300 terrestrial pollen
grains were counted for each sample. Pollen per-
centages are calculated based on the total of ter-
restrial pollen grains.
Organic content was determined by LOI at
105‡C, 550‡C and 1000‡C (Dean, 1974). Samples
selected for charcoal quanti¢cation and identi¢ca-
tion of 0.5 cm3 were sieved through a 180-Wm-
mesh screen after treatment with hot 10% KOH
and 0.1 M sodium pyrophosphate solution. Char-
coal fragments were scanned under a dissecting
microscope at a magni¢cation of U20.
AMS radiocarbon dates were obtained from
the Institute of Arctic and Alpine Research at
the University of Colorado and Beta Analytic.
The corresponding calendar ages were obtained
using the CALIB 4.2 computer program (Stuiver
and Reimer, 1993).

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 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90 77

Fig. 2. Age^depth curves for (top) Maxus 4 and (bottom) Maxus 1. Also shown are stratigraphical lithology and organic content
curves. The bottom date for Maxus 1 is rejected because it is apparently too young. Note the scale di¡erence of the axes of age
(horizontal axes) for the two cores.

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78 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90

Table 2
AMS 14 C dates from Maxus cores
14
Depth Laboratory No. Materials C age Calendar years
(cm) (yr BP) mean (range)
Core 4 60^64 NSRL-11814 leaves 1120 Q 30 1051, 1030, 992 (970^1059)
149^150 NSRL-11185 wood 3430 Q 35 3686, 3663, 3650 (3630^3696)
244^246 NSRL-11186 wood 4350 Q 35 4868 (4861^4969)
405^406 NSRL-11187 wood 5450 Q 40 6280 (6201^6292)
509^510 NSRL-11188 sediment 6480 Q 45 7339, 7339, 7381 (7286^7388)
626^627 NSRL-11189 wood 8180 Q 45 9042, 9071, 9088, 9132, 9147, 9176, 9183 (8992^9212)
Core 1 147^148 NSRL-11684 wood 140 Q 30 265, 218, 142, 24, 2 (1^274)
225^226 NSRL-11685 sediment 250 Q 30 298 (288^308)
465^469 Beta-80900 plant macroremains 1000 Q 60 929 (798^960)
702^704 NSRL-11486 wood 160 Q 35 271, 209, 198, 193, 146, 14, 3 (1^281)

Log-transformed percentile data for 30 most
abundant pollen taxa in the Maxus 4 core were
ordinated using the PC-Ord 4.0 version of de-
trended correspondence analysis (McCune and
Me¡ord, 1999).
3. Results
3.1. Stratigraphy and LOI
Maxus 4 and Maxus 1 are 8.00 m and 7.36 m
long, respectively. Sediments of these cores consist
primarily of clay and peat, with silt and sand
becoming more important constituents in some
layers (Table 1, Fig. 2). A band of ¢ne white
volcanic ash was noted in Maxus 4 at 494^496
cm, and ash was also present in more dispersed
form between 641 and 798 cm.
In both cores, sediments fall into three basic
categories: those with V5% organic content, a
value consistent with local riverine sediment,
those with V15% organic content, consistent
with other closed basin lakes in the Amazonia
£oodplain, and sediments with s 75% organic
material, consistent with shallow, permanently sa-
turated swamps (Kalliola et al., 1993 ; Bush, un-
published data).
3.2. Chronology and sediment accumulation rate
Six AMS radiocarbon dates were obtained (Ta-
ble 2, Fig. 2) for Maxus 4. The dates are inter-
nally consistent with no inversions. An extrapo-
lated age for the bottom of the core is about 9850
cal yr BP, suggesting that it covers almost the
entire Holocene (Fig. 2). Average accumulation
rates were between 0.4 and 1.1 mm/yr. The high-
est rate of deposition was in the middle of the
core (252^505 cm, around 1 mm/yr; Table 1).
The sediments and their accumulation rates sug-
gest that the accumulation was continuous.
Four AMS dates were obtained for the Maxus
1 (Table 2, Fig. 2). The oldest date was 1000 Q 60
yr BP. The basal date was apparently too young
(probably from tree root materials), and is re-
jected (Fig. 2). The top two dates are consistent
with a relatively rapidly depositional environ-
ment. The extrapolated age of the bottom of the
core is 6 2000 years old (Fig. 2). We consider it
likely that the core represents only the late Holo-
cene period. The average sediment accumulation
rate for the top 470 cm is about 4.7 mm/yr, which
is much higher than Maxus 4.
3.3. Pollen analysis
3.3.1. Maxus 4
The sandy sediments at the base of the core
lack pollen, but all other layers are rich in pollen.
Based on a CONISS analysis (Grimm, 1987) and
major changes of abundant pollen taxa, the pollen
record is divided into ¢ve zones (Fig. 3).
Zone 1 ( s 570 cm; s 8300 cal yr BP): Pollen

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 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
PALBO 2448 23-7-02

Fig. 3. Pollen percentage diagram of selected taxa from Maxus 4. AMS radiocarbon dates, corresponding calendar year ages, and lithology are also shown on the
left of the diagram. Organic content in the sediment is shown on the right of the diagram for comparison.

79
80 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90

Fig. 4. Pollen concentration diagram of selected taxa from Maxus 4 (U1000 grains/cm3 ).

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 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
concentrations are 6 100 000 grains/cm3 (Fig. 4),
organic content of sediments in this zone is low
(V10%). The pollen assemblage is diverse and is
not dominated by any single type. Medium high
abundance of Malpighiaceae, Fabaceae, Myristi-
caceae, Combretaceae/Melastomataceae and Ficus
characterizes the pollen assemblages. Cecropia
pollen is scarce ( 6 20%). Polypodium spores are
very common (25^100% total pollen) (Fig. 3).
Zone 2 (570^350 cm, V8300^5700 cal yr BP):
Pollen concentrations are very high ( s 100 000
grains/cm3 , with the highest reaching 400 000
grains/cm3 ), and organic content is still low
(V10%) (Figs. 2 and 4). Extremely high Cecropia
pollen percentages (40^80%) (except at depth of
410 cm) characterize this zone. Concentrations
of Cecropia pollen are generally between 50 000
and 350 000 grains/cm3 . Triporate Urticaceae/
Moraceae and Myristicaceae pollen are also abun-
dant.
Zone 3 (350^150 cm, 5700^3500 cal yr BP):
Pollen concentrations drop to between 50 000
and 100 000 grains/cm3 . Organic content is very
high (up to 70%). Cecropia pollen percentages de-
crease to very low levels ( 6 5%). Fabaceae, Mal-
pighiaceae, Myristicaceae, Zanthoxylum, Lamia-
ceae, Ficus and Sloanea sequentially dominated
the pollen assemblages.
Zone 4 (150^70 cm, 3500-V1000 years ago):
Total pollen concentrations are very high again
due to high concentrations of Mauritia and Sloa-
nea (with as many as 250 000 grains/cm3 ), but
dropped near the top of the zone to 6 50 000
grains/cm3 . Organic content is persistently low
( 6 20%). Mauritia and Sloanea dominate the pol-
len percentages ( s 20% to nearly 90%, and V10^
75%, respectively). Iriartea is also an important
component (V5%). Selaginella and other trilete
spores (mostly from Cyathaceae) reach maximal
representation in this zone.
Zone 5 (70^0 cm, 6 1000 years ago): Pollen
concentration is low at the beginning of the
zone ( 6 100 000 grains/cm3 ), but increases to
very high levels in the upper portion of the zone
( s 200 000 grains/cm3 ). Organic contents are very
high (60^70%, matching the levels in zone 3).
Myristicaceae pollen is an abundant component
of this uppermost zone. Combretaceae/Melasto-
PALBO 2448 23-7-02
81
mataceae, Acalypha, Alchornea, Poaceae and As-
teraceae are important.
3.3.2. Maxus 1
Pollen concentration in this core is generally
lower than in Maxus 4 (all less than 65 000
grains/cm3 , except for levels of 243 cm and 461
cm, which are around 400 000 grains/cm3 ). The
low pollen concentrations are consistent with rel-
atively high sediment accumulation rate. The rec-
ord is divided into four zones based on a CONISS
analysis (Grimm, 1987) of the percentile data
(Fig. 5).
Zone 1 (736^475 cm, s V1000 cal yr BP):
Pollen concentration is 6 25 000 grains/cm3 . Or-
ganic content is very low ( 6 5%) (Figs. 2 and 6).
Pollen of Urticaceae/Moraceae, Combretaceae/
Melastomataceae, Iriartea and Piper is common
but none rises to dominate the assemblage. Cecro-
pia is generally between 15% and 20%, but stead-
ily increases its abundance in the upper portion of
the zone (to V35%). Fern spores, especially tri-
lete spores, are abundant at the beginning of the
zone, reaching their maximum representation in
the core (V10%), but decline markedly in abun-
dance near the top (Fig. 5).
Zone 2 (475^400 cm, 800^1000 cal yr BP): Pol-
len concentration is extremely high at 450 cm due
to high Cecropia concentration (nearly 400 000
grains/cm3 ), but then decreases to 6 50 000
grains/cm3 (Figs. 5 and 6). Organic content is be-
tween 15 and 20%. High Cecropia pollen percent-
age characterizes this zone ( s 30%, and reaching
V90% at 450 cm). Ficus and Inga pollen are
abundant at the top of this zone. Other pollen
types have low percentage representation due to
the abundance of Cecropia.
Zone 3 (400^225 cm, V800^300 cal yr BP):
Pollen concentration is generally between 40 000
and 80 000 grains/cm3 , but at level of 243 cm, it
reaches 480 000 grains/cm3 . Organic content is
low at the base of the zone (10^15%), but very
high at the top (230^270 cm, as high as 80%).
Iriartea pollen is the most abundant pollen type
in this zone (V30%). Zanthoxylum, Tapirira,
Acalypha and Combretaceae/Melastomataceae
are also important components of the pollen spec-
trum. Polypodium spores are consistently present
 82
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PALBO 2448 23-7-02

Fig. 5. Pollen percentage diagram of selected taxa from Maxus 1. AMS radiocarbon dates and lithology are also shown on the left of the diagram. Organic content
in the sediment is shown on the right of the diagram for comparison.
C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90 83

Fig. 6. Pollen concentration diagram of selected taxa from Maxus 1 (U1000 grains/cm3 ).

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84 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90

Fig. 7. DCA ordination result of the ¢rst two axes for log-transformed pollen samples from Maxus 4. Rare species are down-
weighted. Eigenvalues for axis 1 and axis 2 are 0.275 and 0.069, respectively. Samples from each of the ¢ve pollen zones are dis-
tributed in a relatively close area (circled in the diagram), and samples in di¡erent zones are generally distributed in separate
areas, except for samples of zone 1 and zone 3, indicating the similarity of these two zones. Wet-adapted taxa (Mauritia, Ilex,
Iriartea and Sloanea) have the highest scores on axis 1. Thus, axis 1 expresses a dry to wet gradient. Zones are ordered from old
to young (from bottom to top), and the label of each sample means its level of depth (e.g. L284).

at s 10% of total pollen. The sediment for the
top part of the zone is peat, di¡erent from its
neighbors, which are silt with ¢ne sands. A peak
of Poaceae pollen (V30%) is found in this layer.
Zone 4 (225^0 cm, 6 300 cal yr BP): Pollen
concentration is low (30 000^60 000 grains/cm3 ).
Organic content increases from 6 10% at the
base to high ( s 60%) on the top. Cecropia and
Iriartea are moderately abundant. Alchornea, Cel-
tis, Trema and Urticaceae/Moraceae are also im-
portant components.
3.4. Charcoal
Almost no charcoal fragments were found in
either core. At a few levels, one or two fragments
of charcoal ( s 65 Wm) were seen, but they are too
rare to be meaningful.
3.5. Detrended correspondence analysis (DCA)
result
The DCA ordination of the Maxus 4 data (Fig.
7) produces a ¢rst axis that polarizes Mauritia-
dominated samples from other samples. This
axis is of increasing £ood duration. The second
axis polarizes lakeside £oras into those that are
dominated by Cecropia (negative extreme) from
those that are not. Although zones 1 and 3 are
very similar in their composition, there is little
overlap of samples between contiguous zones.
From this result, we know that samples from
the same pollen zones are relatively similar to
each other, and that changes between zones are
relatively great. This suggests that the environ-
mental conditions and vegetation components
were stable during the period represented by

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 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
each pollen zone, and environmental changes
were substantial and relatively rapid between
zones.
4. Interpretation and discussion
4.1. Environmental history
The base of Maxus 4 is composed of clays with
sands. No pollen was preserved in this sediment,
indicating a prolonged oxidation prior to the for-
mation of the modern wetland. This sediment is
the common matrix found beneath wetlands in
western Amazonia and probably represents an an-
cient soil surface. About 9300 cal yr BP, clays
were deposited that are rich in pollen indicating
that the site had £ooded. The origin of the wet-
land is not known, but it could have been caused
by river migration or by tectonic activity. As no
coarse sands underlie this zone it is unlikely that
this site was a main river channel, but it could
have been a white water (varzea) or blackwater
(igapo) £ood system. The spectrum and concen-
tration of pollen deposition at this time is consis-
tent both with that of modern lowland rain forest
and with that of modern riverine muds.
Between 8700 and 5700 yr BP, Cecropia be-
comes the most abundant pollen type, dominating
the pollen spectrum. At lakes in western Amazo-
nia, Cecropia typically forms 15^20% of the pol-
len count (Bush and Colinvaux, 1988; Liu and
Colinvaux, 1988; Lyons-Weiler, 1992), but at
Maxus 4 Cecropia accounts for 40^80% of the
pollen count for 3000 years.
As Cecropia are short-lived pioneer trees that
thrive in disturbed settings, their abundance in
the early to mid-Holocene requires explanation.
In the absence of human disturbance, Cecropia
are fugitive species that invade tree-fall gaps and
primary successional habitats on the accreting
beaches associated with meandering rivers (Fos-
ter, 1990). The sudden £are in abundance of Ce-
cropia in Maxus 1 may be associated with a single
natural disturbance event (Fig. 5). Such a signa-
ture is consistent with an event that allows Cecro-
pia to establish a population for one generation
prior to being snu¡ed out by competition. How-
PALBO 2448 23-7-02
85
ever, at Maxus 4, some factor, or combination of
factors, maintained a large population of Cecro-
pia close to the coring site for three millennia.
One explanation of a prolonged dominance of
Cecropia in the pollen rain is that humans were
disturbing the landscape. At Ayauchi , Ecuador,
human disturbance was inferred from the pres-
ence of charcoal, high abundances of Poaceae
(15^20% of non-Cecropia pollen), Cecropia (40^
90%) and Zea mays (Bush and Colinvaux,
1988). A charcoal peak is also documented in
Maxus 5 that seems to be roughly contemporane-
ous with the Cecropia spike in Maxus 4, though it
was not accompanied by high Poaceae values or
Zea mays (Athens and Ward, 1999). Although the
increase in Cecropia in Maxus 4 might be re-
garded as a signal indicative of major human dis-
turbance, the near or complete absence of char-
coal, Poaceae pollen, or Zea pollen in this core
renders such an interpretation uncertain.
In the modern ecosystems, the principal place
in which large populations of Cecropia occur in
natural settings is where river migration results in
the accretion of a sandy beach. On such beaches
there is a predictable succession of Tessaria (As-
teraceae) and Salix (Salicaceae) followed by Ce-
cropia. In these settings, Cecropia can form large,
almost monotypic stands that are gradually in-
vaded by larger taxa such as Ficus and Guarea
(Foster, 1990; Foster et al., 1986). The Cecropia
phase probably lasts less than 40 years (Foster,
1990) at any one place and its population e¡ec-
tively undergoes local migration at the pace of
river migration, occupying new sandbars and
beaches as they form and stabilize.
At Cocha Cashu, Peru, a 160-year history of
successional changes was tracked in the pollen
rain following abandonment of an oxbow. Cecro-
pia pollen was typically between 15 and 35% of
the pollen count. The highest representation of
Cecropia pollen, ca. 50%, came about 100 years
after abandonment. After 160 years of isolation,
Cecropia representation had fallen to about 25%
(Bush, unpublished data). The Cocha Cashu rec-
ord, unfortunately, does not provide evidence of
an analog mechanism that could maintain unusu-
ally high Cecropia representation for thousands of
years.
86 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
At Maxus 4, if bankside populations are the
origin of the mid-Holocene peak of Cecropia, it
is evident that there was a higher energy £uvial
system causing a greater proportion of the £ood-
plain to remain in an early successional state.
Otherwise the river must have been very close to
the coring site (the Rio Yasuni is presently about
1 km distant from Maxus 4). Ra«sa«nen (personal
communication), however, has not seen evidence
of increased erosive activity in Ecuadorian rivers
at this time.
For Cecropia pollen to be found at 40^80%
suggests a very close source area, i.e. less than
0.5 km distant. Yet, Maxus 4 has a slow deposi-
tional rate (compared with Maxus 1), suggesting
that £ooding from nearby rivers was rare. Evi-
dence for a lack of regular £ooding also comes
from the pollen spectrum. River muds do not
contain such high percentages of Cecropia pollen
and contain a lower concentration of pollen than
is found at this site (Haberle, 1997 ; our unpub-
lished data). In Maxus 4, a band of silty clay with
a markedly di¡erent pollen signature and low pol-
len concentration occurs at 4 m and it is probable
that this represents either a phase of £ooding or a
single £ood event. Thus, if a signi¢cant propor-
tion of the pollen was derived from £oodwater,
the pollen composition of Maxus 4 would be ex-
pected to resemble that of rivers, rather than this
Cecropia-rich spectrum. It seems improbable that
a river with enough energy to maintain extensive
areas of primary succession lies so close to Maxus
4 that its marginal vegetation dominates the pol-
len spectrum, but does not regularly £ood into
this adjacent wetland.
The pollen concentration data (Fig. 4) reveal
more detail of the Cecropia-dominated zone.
The initial rise of Cecropia provides a peak in
pollen concentration. The only other pollen taxon
to show a strong response at this time is Combre-
taceae/Melastomataceae. Both of these families
occur in almost all habitats in the modern forests,
and both could contribute to early successional
phases, although Miconia spp. (Melastomataceae)
are especially abundant in early successional envi-
ronments. Given the paucity of other forest ele-
ments, this Cecropia peak is consistent with the
formation of a riverine primary succession of Ce-
PALBO 2448 23-7-02
cropia near to the site. Overlying samples contain
higher concentrations of forest taxa suggesting
that there may be a separate explanation for the
occurrence of high concentrations of Cecropia in
these levels.
Studies in semi-deciduous forests on Barro Col-
orado Island, Panama, showed that Cecropia is
strongly over-represented in local pollen spectra
by a factor of about 17 relative to its basal area
in the adjacent forest (Bush and Rivera, 1998,
2001). In mature forests of the Yasuni, Cecropia
contributes ca. 6 1% basal area in £oodplain for-
ests and terra ¢rme forests (Pitman, personal
communication). Therefore, a background repre-
sentation of Cecropia in the pollen record of ca.
10^15% might be expected. Additional representa-
tion is likely as Cecropia take advantage of light
availability and gap formation at lake or river
margins. This additional source of Cecropia e¡ec-
tively doubles its representation in the pollen rec-
ord.
At Maxus 4, after the initial successional phase
at the base of the Cecropia zone, the proportion
of Cecropia pollen is around 60%, approximately
two to three times the amount normally found.
Given that the forest still contains many large
trees (as indicated by other components of the
pollen spectrum), it appears probable that canopy
turnover was more rapid. Increased gap forma-
tion would favor increased Cecropia abundance.
One possible explanation is that periodic drought,
such as that experienced in 1983 on Barro Colo-
rado Island could cause mortality and increased
canopy turnover. In that drought, the most
strongly a¡ected individuals were large canopy
trees ( s 20 cm dbh), and their mortality increased
from a ‘normal’ rate of 2% per annum to almost
7% per annum during the three years following
the drought (Condit et al., 1995, 1996). It will
be noted that this single drought induced a
three-fold increase of large tree mortality, and
by inference, a substantial increase in canopy
gap formation.
The amount of canopy gap formation needed
to explain the increase in Cecropia pollen in the
Maxus 4 record would be about a doubling of gap
area. Hence, the probable gap formation caused
by a severe drought is about the right scale of
 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
event to account for the Cecropia abundance. As
there is no evidence to suggest that the moist for-
est of this region was replaced by an overall drier
assemblage, we infer that the droughts were epi-
sodic with su⁄cient return periodicity to maintain
relatively high rates of canopy turnover. If this
hypothesis is correct, the cycle of droughts would
have started approximately 8700 cal yr BP and
ended approximately 5800 cal yr BP. Interest-
ingly, Athens and Ward (1999) suggest drier con-
ditions at this same time in Maxus 5, though they
infer a climate regime characterized by more in-
tense seasonality than at present. Distinguishing
between cyclical droughts, as we believe for
Maxus 4, and seasonally dry conditions as sug-
gested for Maxus 5, however, will be a consider-
able challenge for paleoecology.
The Yasuni inference of drought conditions co-
incides with the marked lowering of lake level at
Lake Titicaca, which receives most of its water
from the winds carrying Amazonian moisture
into the Andes (Hastenrath, 1985). Coinciden-
tally, this same period has been suggested to be
a time of reduced El Nin‹o activity (Keefer et al.,
1998; Sandweiss et al., 1996). However, the root
cause of these drought events remains enigmatic.
The onset of wetter conditions may have caused
the drop in Cecropia abundance at 5800 cal yr BP.
After 5800 cal yr BP, the pollen spectrum is more
typical of undisturbed lowland forest. A sequence
of species rising and falling in abundance suggests
a hydrological succession. Very similar patterns of
succession have been noted in late Holocene
Ecuadorian sequences from An‹angucocha (Frost,
1988) and Zancudococha (Lyons-Weiler, 1992). In
the Colombian savannas of the Llanos Orientales,
Behling and Hooghiemstra (1999) suggest the
presence of a wet phase from ca. 6350 cal yr
BP. This further corresponds remarkably closely
to rising levels documented in two central Ama-
zonian lakes at ca. 6500 cal yr BP (Bush et al.,
2000), and suggests the widespread onset of wetter
climatic conditions in the Holocene at this time.
The short-lived peaks of pollen types evident in
the Maxus 4 record could re£ect the successional
phases as the basin ¢lls with sediment. However,
it is equally likely to be a relatively stochastic se-
ries of events in which a small pool is serially
PALBO 2448 23-7-02
87
invaded by swamp taxa. By 4900 cal yr BP,
when the pool had almost ¢lled with sediments,
peat started to form. A renewed £ooding event at
ca. 3700 cal yr BP is suggested by a resumption of
clay deposition and the dominance of Mauritia.
This transition has been found in several sites in
Amazonia (Behling and Hooghiemstra, 1999;
Bush et al., 2000; Frost, 1988; Lyons-Weiler,
1992; Mayle et al., 2000) and probably represents
a regional increase in precipitation.
At about 1000 cal yr BP, the hydrarch succes-
sion resumes and Mauritia is replaced by wetland
elements such as Myristicaceae (cf. Iryanthera)
and Combretaceae/Melastomataceae. For the
last 1000 years, the hydrological and vegetational
conditions have been relatively stable, supporting
an essentially modern vegetation.
This hydrarch succession in the last 1000 years
can be seen most clearly in Maxus 1. The basal
sediments of this core are similar to those at the
base of Maxus 4, except pollen is preserved in
Maxus 1, although the concentrations are very
low (Fig. 6). The initiation of the Maxus 1 wet-
land is probably similar to that of Maxus 4. This
lake was formed during the wet period docu-
mented in Maxus 4 (3700^1000 cal yr BP). Sands
in the basal sediments are consistent with a river-
ine origin and are overlain by sandy clays typical
of an abandoned oxbow lake that is still season-
ally £ooded. A Cecropia peak occurred at about
1000 cal yr BP when the sediment changed from
sandy clay to silty clay. This sedimentary transi-
tion may indicate the complete abandonment of
the lake and a cessation of periodic £ooding by
the river. The AMS date indicates that this peak
follows soon after the apparent onset of drier
conditions at Maxus 4. The Cecropia trees may
represent colonization of exposed shoreline, or a
severe drought that opened canopy gaps. Unlike
the early Holocene record of Maxus 4, this Cecro-
pia peak is quickly followed by peaks of Ficus,
Cedrela and other trees in a typical succession
(Foster et al., 1986; Foster, 1990). By about 300
cal yr BP, sediments almost ¢lled the lake. Rela-
tively high abundances of Poaceae and ferns sug-
gest the formation of a £oating mat, which has
been steadily invaded to provide the present
wooded swamp. Evidence for human land use is
88 C. Weng et al. / Review of Palaeobotany and Palynology 120 (2002) 73^90
surprisingly sparse in Maxus 1 despite the pres-
ence of nearby archeological sites (Netherly,
1997).
5. Conclusion
Our data from two cores in the eastern Ecua-
dorean lowlands document an almost complete
Holocene sequence that re£ects vegetational and
climatic change. Unusually, these cores seem to
lack evidence for human landscape disturbance,
which fortuitously allows a clearer vision of nat-
ural changes in the record than has been obtained
from other Ecuadorian sites. This ¢nding also
suggests a rather low or negligible impact on the
landscape by prehistoric Amerindians, presum-
ably indicative of quite low population densities
in the inter£uvial areas.
A long-lasting plateau of Cecropia pollen
(8700^5800 cal yr BP) is tentatively interpreted
to indicate a period in which increased tree mor-
tality led to more gap formation and a relatively
high population of Cecropia. One explanation to
account for this mortality is sequences of severe
droughts occurring on a decadal timescale. The
timing of this drought-prone period corresponds
well with other early Holocene records from
Amazonia, adjacent savannas, and the Andes.
After 5800 cal yr BP, more uniform conditions
allowed the development of mature forests, and a
hydrarch succession that gradually ¢lled the lake
with sediment. The last 5800 years have seen al-
ternating wet and dry millennial-scale events. The
e¡ects of these oscillations of water level are evi-
dent in the hydrology of the site and in the com-
munities of wetland plants, rather than in any
regional change in vegetation types. We infer
that climatic conditions between 5800 and 4900
cal yr BP were relatively wet, from 4900 to 3700
cal yr BP they were relatively dry, from 3700 to
1000 cal yr BP they were relatively wet, and from
1000 cal yr BP to present they were (and are)
relatively dry.
As might be expected in such a wet region, the
landscape appears to have supported moist trop-
ical rain forest throughout the Holocene. How-
ever, the climatic oscillations evident in this core
PALBO 2448 23-7-02
may have been important in helping to maintain
species diversity through continual low levels of
ecosystem disturbance (Connell, 1978) that pre-
vented competitive exclusion.
Acknowledgements
We thank the government and people of Ecua-
dor for allowing us to work in their country. This
study was supported by a grant from the National
Scienti¢c Foundation DEB-9732951. Drs. H. Beh-
ling and H. Hooghiemstra reviewed the manu-
script and provided valuable suggestions to im-
prove it.
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