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QUATERNARY RESEARCH 34, 330-345 (1990)

Late Pleistocene Temperature Depression and Vegetation Change in


Ecuadorian Amazonia
MARK B. BUSH,* PAUL A. COLINVAUX,* MICHAEL C. WrEMANN,t
DOLORES R. PIPERNO,$ AND KAM-BIU L~ut
*Department of Zoology, The Ohio State University, 1735 Neil Ave., Columbus, Ohio, 43210; fDepartment
of Botany, Louisiana State University, Baton Rouge. Louisiana 70803; #Smithsonian Tropical Research
Institute, P.O. Box 2072, Balboa, Panama: and Department of Geography and Anthropology, Louisiana
State University. Baton Rouge, Louisiana 70803
Received September 14, 1989
Paleoecological (pollen, phytolith, and wood) analyses of sediments, radiocarbon dated 33,000 to
26,000 yr B.P.. from two sites in Ecuadorian Amazonia provide data that suggest a cooling of ca.
7.5C below present in equatorial lowlands from 33,000 to 30,000 yr B.P. A period of warming
followed in which novel species assemblages, a blend of montane and lowland floral components,
persisted for at least 4000 years. These data of forest community change, from sites lying within the
postulated glacial rain forest Napo refugium. provide the strongest paleoecological refutation of the
refugial hypothesis yet obtained. The large temperature depression at ca. 30,000 yr B.P. allows the
possibility that if maximum cooling at the equator was synchronous with the last glacial maximum
(LGM) of the northern hemisphere, freezing temperatures would have been experienced in parts of
lowland Amazonia between 25,000 and 18,000 B.P. 0 1990 University of Washington.

INTRODUCTION Although deglaciation was approximately


synchronous at low and middle latitudes,
Rind and Peteet (1985) identified prob- glacial maxima may have been asynchro-
lems in reconciling conservative last glacial nous. Recent models seeking to predict
maximum (LGM) sea-surface temperature conditions at the LGM consider only the
models postulating a ca. 2C cooling period 18,000 to 0 yr B.P., and, therefore,
(CLIMAP, 1981) with available empirical do not cover the period of maximum tem-
data that suggest tropical land temperatures perature depression in the tropics (e.g.,
fell by 9-10C (Flenley, 1979; Walker and CLIMAP, 1976, 1981; Kutzbach and
Flenley, 1979, Hooghiemstra, 1984). By Street-Per-r-Ott, 1985; Rind and Peteet, 1985;
doubling the cooling of the sea surface from Kutzbach and Guetter, 1986).
the predictions of CLIMAP (1981), Rind Although distinct changes in the montane
and Peteet (1985) calculated a ca. 6C trop- vegetation of the Andes have been docu-
ical land temperature depression and some mented since initial reports appeared 30
precipitation decrease at 18,000 to 15,000 yr years ago (van der Hammen, 1961), there
B.P. This temperature depression is still has been little consensus as to the impact of
conservative in view of the 10C depression this change on the lowland vegetation. Van
advocated by Flenley (1979). The few long der Hammen (1974) suggested that the mid-
paleobotanical records that exist from the elevational forests had contracted vertical
tropics are consistent in suggesting that the ranges during glacial maxima, to some ex-
maximum temperature depression occurred tent cushioning the downslope extension of
between 33,000 and 25,000 yr B.P. (e.g., montane floras into the lowlands, and that
Powell, 1970; Walker and Flenley, 1979; the upper limit of the lowland forest was not
van Gee1 and van der Hammen, 1973; van reduced by 1000 m but by only about 200 m
der Hammen, 1974; Hooghiemstra, 1984). (van der Hammen, 1974). This argument is
330
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Copyright 0 1990 by the University of Washington.
All tights of reproductmn in any form reserved.
AMAZONIAN PALEOTEMPERATURES 331

internally inconsistent, as a modern lapse-


rate was used by van der Hammen to es- COLOMBIA

tablish the descent of montane forests but


was ignored when considering the vegeta-
tional changes downslope. The modern
lapse-rate across the Ecuadorean Andes is ECUADOR
calculated from recent weather station data
to be 5C cooling per 1000-m rise in altitude
(Catiadas, 1983). As lapse-rates are a phys-
ically determined property of air, they are
unlikely to have changed through time
(Webster and Streten, 1978; Stone and
Carlson, 1979).
Data describing the climatic conditions
prevalent in lowland Amazonia are critical
to the debate surrounding the refugial hy-
FIG. 1. Sketch map of Ecuador showing the loca-
pothesis (Haffer, 1969; Whitmore and tion of San Juan Bosco (*SJB), Mera (*M). sites of the
Prance, 1987; Colinvaux, 1987; Salo, 1987; fossil deposits, and the location of lakes from which
Gentry, 1989), which contends that disjunct surface samples were taken in relation to the 2000-m
centers of lowland species endemism ap- contour of the Andes, shown as the shaded area. Mod-
pear to be correlated with mid-elevational ern pollen sites are (1) Yaguar cocha; (2) Cunro; (3)
San Marcos; (4) Yambo; (5) Rum Turn; (6) Indanza; (7)
(500-2000 m) hilltops in Central and South Llaviucu; (8) Ayauchi; (9) Kumpak; (IO) Puyo Bog;
America. It has been proposed that these (11) Anangucocha; (12) Cuyabeno; (13) Lago Agrio;
hilltops captured orographic rainfall and (14) Santa Cecilia. All of Ecuadorian Amazonia is con-
supported lowland rain forests in isolated sidered to lie within the postulated Napo rain forest
refugia throughout a dry glacial period refugium.
(Haffer, 1987). The refugial hypothesis sug-
gests that thus isolated by arid lowlands, etation depression, a 4.5C minimum tem-
gene flow between refugia was restricted perature depression, and a change in the
and allopatric speciation took place. No un- local floral assemblages during the LGM.
equivocal paleoecological evidence for low- Their suggestion that Amazonian floral as-
land aridity in the Amazon has been forth- semblages are ephemeral is in accord with
coming and no sites radiocarbon dated to recent studies on temperate floras (Davis,
glacial times have yet been published from 1986; Huntley and Webb, 1989).
the entire Amazon basin except the site at In June 1988, a second exposure, similar
Mera (B-ion, 1984; Liu and Colinvaux, and contemporaneous to the one docu-
1985; Colinvaux, 1987; Salo, 1987) (Fig. 1). mented from Mera, but ca. 160 km south,
The concept of glacial aridity in the Ama- was discovered near the village of San Juan
zon, therefore, remains unsubstantiated Bosco, Ecuador. Analysis of fossil wood
(Colinvaux, 1987). and pollen from the San Juan Bosco depos-
The first empirical data to relate directly its, together with extended analyses of the
to mid-glacial Amazonian floral assemblages Mera deposits, is presented to test the hy-
and paleotemperature were from road-cut pothesis that temperature depression,
and stream-bank exposures at Mera, Ecua- rather than aridity, was the dominant prop-
dor (1100 m altitude; Liu and Colinvaux, erty of lowland Amazonia in glacial times.
1985). Softwood samples (since confirmed The data invite the further hypothesis that
to be Podocarpus) were dated to 33,000 and times of maximal cooling at the equator
26,000 yr B.P. and were used by Liu and were earlier than times of maximal glacia-
Colinvaux (1985) to argue for a 700-m veg- tion at high latitudes.
332 BUSH ET AL.

THE SITES AND SAMPLING branch and wood samples were collected
for identification and radiocarbon dating.
Both sites are at the western edge of the The second Mera site is exposed where the
Amazon basin in the Andean foothills, at Rio Pastaza has cut deeply into the hillside,
970 and 1100 m. Dense rain forest covers leaving a vertical cliff from which tree
the region, obscuring surficial geomorphic stumps protrude about 10 m above the
features, and extending upslope to about modern stream. Wood samples were col-
1200 m elevation before gradually merging lected from the debris of the cliff fall but the
into forests characteristic of higher eleva- stratum in the cliff itself was not reached.
tions. Both sites are at the sides of valleys
holding under-fit, rapid, clearwater streams. San Juan Bosco (3 3 45, 78 27 20)
As discussed below, the deposits include A tributary stream of the Rio El Triunfo
highly polleniferous peat or clay that re- has carved a deeply incised valley about 10
quired still-water conditions for its accumu- km south of San Juan Bosco (Fig. 1). A
lation, as well as what appear to be flood river-cut exposure on the steep side of this
deposits and assorted wood fragments tributary valley, at about 970 m altitude,
ranging from the small to whole tree revealed 5 m of stratified silts, sands, peb-
stumps. bles, and small boulders overlying a schis-
tose metamorphic rock. The stream, seen
Mera (I 29, 77 06)
during the dry season, was low, did not fill
The Mera site was described and illus- its present course, and appeared to carry
trated by Liu and Colinvaux (1985). The little sediment.
principal site is exposed in a road-cut in the The exposure was cleaned off and eight
valley of the Rio Pastaza, about 200 m from basic stratigraphic units were identified
the river and 10 m above the present flood- (Fig. 2). Large pieces of wood were evident
plain. Amorphous compacted, silty peat, in six of the eight depositional segments.
1.3 m thick overlies a deposit of silty sand Wood samples were collected for 14C dating
into which the road bed has been cut (Fig. and identification from each unit. The wood
2). Seven samples were collected at 30-cm appeared to be waterlogged, even during
intervals for palynological analysis, and the dry season, and laterally compressed.

MERA SAN JUAN BOX0

FIG. 2. Site stratigraphy of Mera and San Juan Bosco, Ecuador. Lettering on the right of each
column indicates stratigraphic units sampled for paleoecological analyses; numbering on left indicates
location of samples taken from layers where more than one sample was collected.
AMAZONIAN PALEOTEMPERATURES 333

There was no evidence to indicate slippage posefully designed program for the Macin-
or post-depositional perturbation, although tosh microcomputer (Eisner and Sprague,
the basal sediments had been subject to 1988).
some tilting (Fig. 2). A total of 15 samples
was collected. ANALOG POLLEN SAMPLES FROM
VARYING ALTITUDES
METHODS Fourteen modern and recent fossil analog
Longitudinal, transverse, and tangential sites (Fig. 1) are used to display the changes
sections of the wood samples were pre- in pollen representation with increasing al-
pared. The identification of all samples of titude. A marked discontinuity between the
Mera and San Juan Bosco wood were car- lowland and the upland pollen spectra is ap-
ried out using the General Unknown Entry parent (Fig. 3). The lowlands are character-
and Search System (Wheeler et al., 1986), a ized by high values of Cecropia (with the
computer database which includes more exception of Cuyabeno and Mera), Ficus,
than 2500 genera. The state of preservation Tremu, Aculyphu, and Palmae, while the
of the samples and the incomplete database upland sites are rich in Gramineae, Alnus,
prevented identification to species, but a Compositae, Cyperaceae, and Chenopodi-
strong indication of genus or family was uceuelAmurunthus. A survey of the flora
usually attainable. and pollen deposition in the Colombian
The sediments were analyzed for water Andes revealed similar patterns, with low
content, and samples of known mass and pollen percentages of Alnus and Weinmun-
volume were wet-sieved (124 pm) and then niu below 2500 m altitude (Grabandt, 1980).
oven-dried to determine proportion by When these two genera represented more
mass of coarse and tine particulate matter. than 2% of the total pollen sum, a signifi-
Phytoliths were isolated by density gra- cant relationship was found between local
dients, concentrations were determined by floral abundance and pollen representation.
the aliquot method, and counts were car- Close correlations are not sought with Gra-
ried out on a line and a coarse silt fraction bandts (1980) work, as the Andean and
for each sample (Piperno, 1988). Single sub-Andean forests of Colombia are rich in
samples from the silty sands and the peat of Quercus which can account for ~60% of
Mera were analyzed and samples from the pollen rain (Grabandt 1980), whereas
stratigraphic units A, C, H, and I (Fig. 2) Quercus is absent from Ecuador, prevent-
were analyzed from San Juan Bosco. ing meaningful comparison of the percent-
Quantitative pollen slides were prepared age data.
by the addition of exotic spores (Lycopo- At Mera the modern pollen sample from
dium clavatum; Stockmarr, 1971) and by a moss polster may be providing a more-
following standard palynological prepara- local pollen assemblage than the surface
tion procedures (10% HCl, 10% NaOH, ac- sediment samples from the lowland lakes.
etolysis, 1.9 s.g. bromoform separation; Mera provided a high representation of
Faegri and Iversen, 1975). The samples Gramineae for the lowlands, which may be
were mounted in glycerol and pollen was the result of the local pollen input to moss
counted to sums of 200 grains per sample at polsters and presumably represents a forest
500-1000x. Identifications are based on grass. The strong (>20%) representation of
our modern pollen reference collection of Urticaceae/Moraceae at Mera suggests a
about 3000 tropical American taxa and pub- later successional stage than would Cecro-
lished pollen descriptions (Absy, 1979; piu and this is in keeping with the forest
Bartlett and Barghoorn, 1973; Hooghiem- from which the sample was collected. Ce-
stra, 1984; Roubik and Moreno, in press). cropiu is comparatively scarce in the sea-
Pollen diagrams were drawn using a pur- sonally inundated, black-water Zgupo site
334 BUSH ET AL.
AMAZONIAN PALEOTEMPERATURES 335

of Macurococha, Cuyabeno, apparently an well as an equally large representation of


unfit habitat for Cecropia. the high Andean taxa transported, on this
Podocarpus never attains values greater hypothesis, from above. That a rich flow of
than 1% in any of these samples, even in suspended peat from a stream should be so
modern Andean forests in which the genus nicely mingled with local peat and pollen
is an important element. This genus, de- seems vanishingly unlikely. Accordingly,
spite having pollen looking very much like we conclude that the peaty part of the Mera
that of Pinus, apparently liberates much section, with its included fossils, represents
less pollen; therefore, we attach more sig- organic matter contributed by plants grow-
nificance to the presence of relatively low ing in the immediate vicinity of the site.
percentages of Podocarpus than one would The silts and sands underlying the Mera
for Pinus. peat and making up most of the section at
San Juan Bosco, however, suggest deposi-
ORIGIN OF THE FOSSILS: LOCAL tion in low energy streams or in diverticulae
OR TRANSPORTED or temporary impoundments of such
streams. But these deposits are also as rich
Pollen, phytolith, and macrofossil analy- with pollen as Ecuadorian lake samples,
ses reveal the presence in both Mera and from both modern Andean and lowland
San Juan Bosco deposits of taxa now con- taxa. Because the Andean genera Alnus
fined to elevations 1000-2000 m higher in and Weinmannia alone represent up to 50%
the Andes. Therefore, either these taxa of total pollen, the transport hypothesis
once grew at low elevations or their pollen would require that influx of these taxa
and wood were transported downslope, should exceed the influx of locally growing
presumably by the streams in whose val- lowland taxa, even after the long stream
leys the deposits lie. The postulate of long- journey. We know of no precedent for the
distance transport requires that objects as required overwhelming of high local pollen
varied as large tree stumps, phytoliths, and concentrations in an impoundment by the
pollen grains should be moved together for contributions of stream flows from a distant
many kilometers horizontally and up to 2 source.
km vertically before being deposited to- The pollen deposition required by the
gether more than 600 m above the Amazon transport hypothesis is from water with
bottom land. This seems highly unlikely. enough energy to move large tree trunks
The postulate can, however, be examined from the high Andes also. In these streams
with evidence from within the sediments pollen would have been carried in a very
themselves. dilute suspension and could not have been
Pollen concentrations in the Mera peat (6 deposited simultaneously with the larger
x 105-1 .2 x lo6 pollen grains cm3- ) are as debris. Furthermore, no correlation is
high as any known from Ecuadorian lakes, found between the presence of montane
both Amazonian and Andean, demonstrat- wood or pollen types and sediment type:
ing that the deposit collected in still water Drimys wood is found in clays through to
with very little input of allochthonous min- coarse sands (Fig. 4); Alnus and Weinman-
eral matter, as should be present if the im- nia pollen are found in clays, amorphous,
poundment had been fed by streams from greasy, peats, and fine sands; and montane
above. The only way this deposit could be wood and pollen types are found in the
the result of long-distance transport would same depositional bands, despite requiring
be mass transfer of the peat, either as a different transportational and depositional
floating mass or as a turbid flow. However, energies.
this explanation can be excluded because This analysis, therefore, permits the re-
pollen in the Mera peat includes a large rep- jection of the hypothesis of long-distance
resentation of lowland taxa throughout, as transport and we conclude that all pollen
336 BUSH ET AL.

z
2m E San Juan Bosco Mera
z 22 % Dry mass Bloslratlgraphy
P
z 0 0 20 40 60 80

lDrrmy.9

Podocarpus
Magnoliaceae
ZOW
1 37
H 2 40
3 42 IDnmys
1 43
2 44 Ahus /
345 Grammae
4 47 ZOW
5 49
0 20 40 60
% Water (wet mass)

FIG. 4. Silt, sand and water fractions for sediments from Mera and San Juan Bosco, Ecuador with
biostratigraphic annotation. n , % dry mass silt and clay fraction; i3, % dry mass sand fraction, +, %
wet mass water.

and wood samples from both sites repre- resin canals but did contain longitudinal pa-
sent plants that grew locally. renchyma. Podocarpus is, in Ecuador, a
Both the Mera and the San Juan Bosco predominantly Andean genus, but a single
sites are presently too steep to accommo- lowland species, P. rospigliossii Pilger is
date either of the old depositional systems documented at ~600 m elevation from Ec-
indicated by the stratigraphy. Between uadorian Amazonia (Renner et al., 1989). A
33,000 and 26,000 yr B.P. the two localities nongymnospermous wood sample tenta-
must have had a much-less-incised topog- tively identified as cf. Tovomira (Liu and
raphy that allowed lower energy streams Colinvaux, 1985) has since been identified
than are seen today. So drastic a change in as a member of a less-advanced genus. This
topography suggests local tectonic activity. sample, which has scalariform perforations
Local tectonism would also explain the ap- and poorly developed longitudinal paren-
parent contemporaneity of the Mera and chyma, is probably a species of Weinman-
San Juan Bosco sites. That the streams ad- nia, Symplocos, or Laplacea (Table 1).
jacent to both sites lie in underfit valleys is San Juan Bosco wood samples from the
consistent with stream down-cutting fol- uppermost sedimentary stratum (A, Fig. 2)
lowing tectonic movement. The low-energy and a lower layer (H3, Fig. 2) were 14C
depositional environment in the old im- dated to 26,020 + 300 (B-27144) and 30,990
poundments may also have been empha- k 350 yr B.P. (B-27145), respectively.
sized by reduced stream flow correspond- Wood samples were found to be poorly pre-
ing to relative aridity during the Andean served but identifiable to family or genus in
glacial maximum (Hastenrath and Kutz- many instances (Table 1). A wood type rep-
bath, 1985). resented in samples (A, C, and H3) had a
distinctive structure attributable to Drimys
PALEOECOLOGICAL RESULTS (Winteraceae), identifiable by the absence
of vessels and the presence of heteroge-
Wood samples from the basal material of neous rays containing large upright cells.
the Mera sites were dated to 26,530 + 270 Both South American Drimys species are
(B-10170) (Mera 1) and 33,520 + 1010 yr montane, and the only member of the ge-
B.P. (B-9618) (Mera 2). Most of the wood nus, found in Ecuador, is D. granadensis
from both sites was gymnospermous, al- L.F. (C. Ulloa, personal communication,
most certainly Podocarpus, as it had no 1989).
AMAZONIAN PALEOTEMPERATURES 337

TABLE 1. WOODANDPHYTOLITHSFROMSANJUAN Bosco AND MERA

Pollen zone Section Wood Phytoliths


San Juan Bosco
SJ-B 4 A-H2 Aibizia (Bl No phytoliths
Aspidosperma-type (B,Dl
Drimys (A,B,C)
Magnoliaceae (A)
Wienmannia-type (B,D,E2,H2)
SJB-3 H3 Drimys No phytoliths
SJB-2 II-4 No wood No phytoliths
SJB-1 I5 No wood No phytoliths
Mera
MER-2 Podocarpus Gramineae (<l%,O%)
Weinmannia-type Palmae (92%, 14%)
Marantaceae (2%,0%)
Chrysobalanaceae-type (4%,0%1
Undetermined (1%,86%)
MER-I Podocarpus Gramineae (74%,0%)
(composition of grasses:
Festucoid
14%, Panicoid 14%. Bambusoidi
Chloridoid 2% Other Gramineae (44%)
Chrysobalanaceae-type (26%,0%)

Note. Notations in parentheses indicate the levels in which the wood types were found and the percentage
composition of line fraction and coarse fraction phytoliths.

The Pollen Record sis; Hill, 1979b) modified to detrend axes


using second-order polynomials (ter Braak,
Pollen preservation in most samples was 1988).
good, as indicated by the presence of fragile The TWINSPAN classification of the
grains, e.g., Cyperaceae and two-pored Ur- matrix (Fig. 7) reveals an initial division
ticaceae/Moraceae; large numbers of small that separates the samples of pollen zone
grains, e.g., Melastomataceae and up to SJB-4 (1) from all other samples (0). The
30% Weinmannia-type (psilate to microre- group 0 was subdivided into those of the
ticulate, tricolporate 8-12 pm); and the Mera peats, MER-2 (OO), and the silty sands
presence of exosporia on many monolete of Mera and San Juan Bosco with the low-
spores. Pollen concentrations expressed as ermost clay sample SJB-3 (01). At the next
grains/cm3 ranged from 1.2 x lo6 to 2 x lo5 divisive level, SJB-1 and SJB-3 are isolated
grains/cm3 at Mera and from 3.5 x lo5 to (01 I) from the silty sand groups of MER-1
1.7 x IO4 grains/cm3 at San Juan Bosco. and SJB-2. This unconstrained classifica-
Pollen percentage diagrams are given in tion has shown a strong affinity between
Figures 5 and 6. Zoning and interpretation samples in the pollen zones SJB-2 and
are based in part on statistical analyses. MER-1 and also, perhaps surprisingly, an
Data matrices for statistical analysis and affinity between samples SJB-1 and SJB-3.
pollen diagramming were constructed using The polarization of the later San Juan Bos-
taxa attaining >3% total pollen and spores, co and Mera assemblages represents differ-
or those present in at least three samples. ent ecological successions taking place at
The pollen data matrices from Mera and the two sites, or the effect of the geograph-
San Juan Bosco were combined and ana- ical and altitudinal separation. This classi-
lyzed by means of TWINSPAN (Two-Way fication formed the basis of the construc-
Indicator Species Analysis; Hill, 1979a) and tion of local pollen zones.
DCA (Detrended Correspondence Analy- The DCA analysis of the same data set
0 51115 0 peat + = 1%

FIG. 5. Percentage pollen diagram of taxa attaining >5% from Mera. Ecuador. Details of the occurrence of rare taxa for Figures 5 and 6 are available
from the senior author.
AMAZONIAN PALEOTEMPERATURES 339
340 BUSH ET AL.

TWINSPAN

;--__ AXIS 2 DCA

SJB 2

1 SD

FIG. 7. TWINSPAN and DCA results of fossil pollen data from Mera and San Juan Bosco utilizing
taxa listed in Figures 5 and 6. Closed circles, Mera; open circles, San Juan Bosco. Bar scale indicates
I standard deviation of species turnover (Hill. 1979b). Dotted lines delimit TWINSPAN groupings, and
lettering within dotted line indicates samples enclosed. Lettering outside dotted line denotes local
pollen zones.

provided two readily interpretable axes San Juan Bosco record was dominated by
(Fig. 7). Stratigraphically adjacent samples swampland taxa there was a greater input
were joined by a straight line and at-rowed of forest taxa to the Mera deposit.
to indicate the progression from oldest to Axes 3 and 4 were not readily interpret-
youngest (after Clark, 1986). Taxa strongly able.
associated with the extremes of axes are The pollen diagrams from San Juan Bos-
shown in Figure 7. The older samples from co and Mera reveal very similar trends,
both sites lie at the positive extreme of Axis with the basal material of both sections be-
1 and the younger samples lie at the nega- ing rich in Alnus and Gramineae. In both
tive extreme of the axis. The length of the records a change in sediment type coin-
axis, 2.4 standard deviations of species cides with a change in pollen spectra (zones
turnover, suggests with >95% confidence MER-1 to MER-2 and SJB-1 to SJB-2; Figs.
that the two associations polarized by this 5 and 6, respectively); the loss of Alms and
axis were discrete populations. Gramineae; the first record of Apeiba,
The gradient represented by Axis 2 ap- Bombacaceae, and Alchornea; and marked
pears to be from open water to forest. That increases in two-pored Urticaceae/Mora-
the San Juan Bosco samples and the Mera ceae, Melastomataceae, and Weinmannia-
samples at the negative extreme of the first type. Changes in the local depositional en-
axis have been polarized by the second axis vironment across this boundary are also ev-
suggests that local differences in the two ident with the occurrence of still-water
sites are being described and that while the swampland indicators Cyperaceae, Sagit-
AMAZONIAN PALEOTEMPERATURES 341

taria, and Lycopodium (foveolate) above pollen samples that we have recorded in
the sedimentary change. In this upper sec- Ecuador (Fig. 3), the only site with a similar
tion of the Mera record Podocarpus pollen pollen spectrum is Llaviucu (3120 m alti-
attains values in excess of 5%, IO times the tude). As with both Llaviucu and the Alne-
representation found in any of the analog turn-meadow, Weinmannia, Podocarpus,
sites. The presence of Podocarpus wood in Proteaceae, Symphonia, and Tiliaceae
the same samples strongly suggests signifi- were scarce or absent from the lower Mera
cant Podocarpus populations growing and San Juan Bosco samples. The abun-
nearby. dance of festucoid phytoliths and the com-
parative scarcity of panicoid, chloridoid,
The Phytolith Record and bambusoid types (Table 1) is character-
istic of high Andean (sub-Paramo) grass-
The sediments from San Juan Bosco con- lands and would be consistent with an en-
tained no phytoliths, but the samples from vironment of Alnetum with meadows (van
Mera were rich in them. In the lower sandy
der Hammen, 1961; Grabandt, 1980). These
silts of Mera there were few large phyto- habitats lie somewhat higher than the lower
liths, but many small ones. The results of limit of the sub-Andean forest and would be
the phytolith analysis are given in Table 1.
consistent with an altitude of about 2500 m.
The higher percentage of Palmae phyto-
From this we postulate a minimum vegeta-
liths in MER-2 does not necessarily indi-
tion depression of ca. 1500 m, which, as-
cate a strongly palm-dominated forest, be-
suming a moist air lapse rate of 5C 1000
cause similar concentrations of palm phy-
m -, is equivalent to about 7.5C cooling if
toliths are present in soils beneath the
the descent of vegetation were controlled
mature lowland forests of Barro Colorado
by temperature alone. The descent might
Island where palms are scattered in the un-
also have been directly influenced by frosts
derstory (Piperno, 1988). In the coarse silt
at low altitudes because at the modern alti-
fraction from the same sample phytoliths
tude of 2500 m in central Ecuador occa-
are abundant and are represented by Pal-
sional frosts occur, as minimum tempera-
mae (14%) and an assemblage of largely un- tures can drop to - 0.6C (data for Ambato
identified phytoliths, which bear a strong
2540 m altitude; Centro Ecuatoriano de In-
similarity to assemblages found beneath
vestigacion Geografica, 1983).
modern, mature semi-evergreen and ever-
An alternative hypothesis would be that
green forests in Panama.
intense aridity brought about open condi-
tions in which pollen influx was extremely
PALEOCLIMATE INTERPRETATION:
low and certain taxa, e.g., Alnus and
33,000 TO 30,000 YR B.P.
Gramineae, would be over-represented, but
The presence of high values for Grami- the details of our pollen spectra make this
neae (up to 17%) and Alnus (up to 45%) in most unlikely. Our modern, seasonally
the Mera zone MER-1 and San Juan Bosco flooded, blackwater site (Cuyabeno, Fig. 3)
zones SJB-l-3 is strongly indicative of provides a spectrum of pollen from a low-
Andean forests (after van der Hammen, land subject to local seasonal aridity but
1974). Alnus is seldom found lower than reveals no similarity with the fossil data.
2400 m altitude and its combination with Aridity would not result in the downslope
high levels of Gramineae is suggestive of movement of montane taxa and, further-
the Alnetum-meadow (2560 m altitude) pol- more, we would not expect Alnus to be
len spectrum of van der Hammen (1961) in more over-represented from long-distance
which Alnus reaches ca. 50% total pollen transport than such mid-altitude, anemo-
and spores, and Gramineae attains ca. 35% philous taxa as Bursera, Byrsonima, Pal-
total pollen and spores. From the surface mae, Urticaceae/Moraceae, and Cyper-
342 BUSH ET AL.

aceae growing on the actual site. We there- readvance burying the paleosols (Clapper-
fore find no evidence to support an ton, 1987b). Clappertons data provide a
hypothesis of intense aridity rather than maximal date for the onset of the cold pe-
temperature depression, or of aridity play- riod of ca. 33,000 yr B.P. The overlapping
ing a major role in lowering vegetation dates of Mera, 33,520 + 1010 yr B.P. for the
belts. cold period and 34,200 + 1080/950 (SRR-
Another hypothesis is that cooling was a 3034) for a paleosol at Laguna Pisayambo
local effect of descending glaciers, and (Clapperton, 1987b), suggest that the cold
driven by increased precipitation. Any period started abruptly.
downslope movement that did result from
this mechanism would be halted near the PALEOCLIMATE INTERPRETATION:
0C mean annual isotherm; given a constant 30,000 TO 26,000 YR B.P.
temperature, this would be some 3000 m The data from Mera and San Juan Bosco
higher than the sites of San Juan Bosco and suggest the period 30,000 to 26,000 yr B.P.
Mera. In addition, most empirical data sug- to have been one of warming, as Alnus and
gest that precipitation in the northern Gramineae are lost from the record and
Andes during the period 33,000 to 30,000 yr lowland elements, e.g., Marantaceae (cur-
B.P. was slightly less than, or roughly the rently seldom recorded above 1500 m in Ec-
same as, at present (van der Hammen, uador; Kennedy et al., 1988), and Palmae,
1974; Flenley, 1979; Bush and Colinvaux, Apeiba, Erythrina, Rhus, and Proteaceae
1990). Therefore, the hypothesis of a plu- become established. However, this was not
vial period feeding glaciers to yield local a pure lowland community, as montane el-
cooling can be rejected. ements (e.g., Drimys, Podocarpus, Hedy-
We conclude, therefore, that the ob- osmum, and Weinmannia-type) were abun-
served vegetation changes at our two study dant. The Drimys is suggestive of modern
sites were the product of a regional cooling, altitudes over 2500 m, while Podocarpus, in
suggesting a 75C temperature depression association with Hedyosmum and Wein-
at low altitudes near 0 latitude during the mannia, is characteristic of altitudes of
period 33,000 to 30,000 yr B.P. This esti- 2000 m and above. Alnus was still present at
mate is consistent with the estimate of full- Mera at higher values than any of our sur-
glacial cooling of 7.4C at high altitudes, face samples sites below 3000 m. This as-
based on geomorphic evidence (Clapper- semblage of mixed lowland and montane el-
ton, 1987a). ements is apparently without modern ana-
The basal sample of San Juan Bosco ap- log. The failure of lowland plants to
pears to exhibit a less-extreme montane dominate this forest during the period
flora than the overlying samples, and in the 30,000 to 26,000 yr B.P. may be due to cli-
TWINSPAN analysis was grouped with the matic fluctuations that allowed the incum-
upper zone SJB-3 which appears to repre- bent cold-tolerant species to hold their
sent a more moderate climatic period. ground. From such a community, where no
Thus, zone SJB-1 may document the onset modern analog even approximates the spe-
of the coldest period. The lowermost date cies assemblage, it is extremely difficult to
from Mera, taken from a large single piece quantify climatic change.
of wood, gives an effective minimum date
for the onset of this cold period of 33,520 ? THE AMAZON AT THE LAST
1010 yr B.P. (B-9618). GLACIAL MAXIMUM
Paleosols at three sites in central Ecua- The LGM is generally accepted to have
dor suggest a period of comparative been between 25,000 and 18,000 yr B.P.
warmth from >43,000 to ca. 33,000 yr B.P., throughout many temperate regions. How-
the end of which is manifested in a glacial ever, the dating of the coldest period in the
AMAZONIAN PALEOTEMPERATURES 343

tropics is more problematic, primarily be- Heusser, 1989) and provides some evidence
cause of a lack of data. Some evidence sug- that the westerlies were not necessarily
gests that the time of greatest cold might strengthened markedly during the LGM.
have been earlier in the tropics but it has
not been satisfactorily dated (Powell, 1970; THE STATUS OF THE NAP0 REFUGIUM
Walker and Flenley, 1979; Bush and Colin- If the refugial hypothesis (Haffer, 1969;
vaux, 1990). Other authors have chosen to Whitmore and Prance, 1987) is to explain
align their chronologies with those of north- tropical species diversity, the lowland to
ern hemisphere glaciations and either leave upland transition of Ecuador is a key re-
their cores entirely undated (Hooghiem- gion. About 50% of Ecuadors plant species
stra, 1984) or do not present radiometric occupy the 900-3000 m elevation range,
dates older than ca. 25,000 yr B.P. (van der which accounts for only 10% of the land
Hammen, 1974; van der Hammen et al., area (Balslev, 1988). High rates of ende-
1981). The period of intense cold from mism in the flora (39% of the mid-elevation
33,000 to 30,000 yr B.P. suggested by our flora is endemic to Ecuador) tempted refu-
data comes at a time of global glacier ex- gialists to postulate the Pleistocene Napo
pansion, as suggested by oxygen-isotope rain forest refugium, with 80-100% cer-
data from deep-sea cores with ocean- tainty (Brown, 1987). Both Mera and San
surface temperatures close to those of the Juan Bosco lie within the proposed Napo
LGM (Martinson et al., 1987). If the LGM refugium (Fig. l), yet no evidence was re-
of the northern hemisphere was associated corded to suggest a long, continuous his-
with a synchronous cooling event in the tory of unchanging rain forest habitats.
tropics, the temperature depression experi- The downslope movement of montane
enced in Amazonia during the LGM would vegetation (e.g., Alnus, Drimys, Thalic-
probably have been as cold as or even trum, Weinmannia, festucoid grasses, and
colder than the 7SC cooling we document Podocarpus) documented at Mera and San
from 33,000 to 30,000 yr B.P. Conse- Juan Bosco, and the probability of occa-
quently , freezing temperatures would have sional frosts, cannot be reconciled with per-
again penetrated into lowland Amazonia. If sistence of intact rain forest communities
the cooling synchronous with the LGM was above the basal lowlands as the refugial hy-
not as extreme, then the coldest period in pothesis requires. Just as we recorded a
the lowland tropics was not synchronous blend of montane and lowland taxa in the
with the LGM, as has been suggested for post 30,000 yr B.P. sections from these
some tropical locations (Powell, 1970; sites, it is possible, indeed likely, that some
Walker and Flenley, 1979; Flenley, 1979). of the lowland taxa were capable of surviv-
Further climatic reconstructions modeling ing even the coldest period from 33,000 to
the temperature of the lowland American 30,000 yr B.P. Thus, some species of low-
tropics should incorporate a probable cool- land rain forest plants may have had a con-
ing of ca. 7-8C. tinuous history of occupation at these sites,
Our data suggest that current climatic which is very different from suggesting that
models overestimate the temperature of the whole communities remained intact
equatorial American landmass during the throughout this period. We suggest that
late Pleistocene, one consequence of which Pleistocene communities were made up of
is to overestimate the temperature differen- novel assemblages of taxa without modern
tial between the low and middle southern analog and, like their Holocene counter-
latitudes (CLIMAP, 1986; Kutzbach and parts, were the product of rapid environ-
Guetter, 1986). This revised temperature mental change, often too rapid for compet-
differential is important because it drives itive exclusion to remove less adapted
the westerly wind belt (Markgraf, 1989; plants from local communities.
344 BUSH ET AL.

ACKNOWLEDGMENTS Davis, M. B. (1986). Vegetation-climate equilibrium.


Vegetatio 67 (2).
We are indebted to the government and people of
Eisner, W., and Sprague, A. P. (1988). MACPOL-
Ecuador, especially Hans and Trudi Steinitz for their
LEN: Pollen Data Entry and Diagramming Software
hospitality. Paulo De Oliveira, Dr Michael C. Miller,
for the Macintosh. Byrd Polar Research Center,
Dr. Miriam Steinitz-Kannan, Melanie Riedinger, and Ohio State University.
Faust0 Sarmiento are thanked for their assistance in
Faegri. K., and Iversen. J. (1975). Textbook of Pol-
the field and formative discussions. This work was
supported by Grants ATM-8789685 and BSR 8021539 len Analysis. Munksgaard, Copenhagen.
of the NSF. Flenley, J. R. (1979). A Geological History of Trop-
ical Rainforest. Butterworths, London.
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