Neogene and Quaternary Development of the Neotropical Rain
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Transcript of Neogene and Quaternary Development of the Neotropical Rain
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.Earth-Science Reviews 44 1998 147183
Neogene and Quaternary development of the neotropical rainforest: the forest refugia hypothesis, and a literature overview
Henry Hooghiemstra a,), Thomas van der Hammen a,b
a (Hugo de Vries Laboratory, Department of Palynology and PaleorActuo-ecology, Uniersity of Amsterdam The Netherlands Centre for
)Geo-ecological Research, ICG Kruislaan 318, 1098 SM Amsterdam, Netherlandsb
Tropenbos-Colombia, Apartado Aereo 036062, Bogota, Colombia
Received 13 March 1998; accepted 26 June 1998
Abstract
The upheaval of the northern Andes in Miocene and Pliocene time changed the drainage system in northern South
America significantly and caused the present-day rain forest areas of Choco and the Lower Magdalena Valley becameseparated from Amazonas. Plant diversity may have reached the highest level in the Miocene or Pliocene, and excessive
present-day phytodiversity may be regarded as a legacy of the Tertiary, rather than an evolutionary product of the
Quaternary. In the Quaternary strong temperature oscillations, related to the series of ice-ages, were superposed on the LateTertiary forest dynamics, which included river displacement and latitudinal migrations of the equatorial rain belt caloric
.equator with the rhythm of the precession cycle of orbital climate forcing. The hypothesis that claims a permanent rain
forest cover all over the Amazon basin during the last glacial is in contrast with the forest refugia hypothesis, which
accepts replacement of rain forest by savanna, or savanna forest, during dry climatic intervals. Both scenarios have beenevidenced by pollen records. In this paper, it is suggested that both hypotheses are not necessarily conflicting and apparently
did occur in different parts of the Amazon basin, and in different periods, depending on the climatological constraints. A
compilation of the most important literature concerning the vegetational, climatic, and environmental history of the rain
forest areas of Amazonas and Choco, and surrounding dry ecosystems has been included. q 1998 Elsevier Science B.V. Allrights reserved.
Keywords:Neogene; Quaternary; rain forest; forest refugia; Amazonas
1. Introduction
Apart from paleoclimatological studies using ma-
rine sediments, palynological and paleoecologicalstudies from the continents contributed immensely
during the last decades to the understanding of the
history of our environment and climate. Most atten-
)
Corresponding author. Tel.: q31-20-5257857; Fax:q31-20-
5257662; E-mail: [email protected]
tion has been dedicated to the temperate zones,
whereas the history of climate, environment, and
ecosystems of the tropical and subtropical areas is
less understood. Spatial studies covering the globe ofthe past decade, highlight the unacquaintedness with
the tropical part of the earth system.
Tropical rain forests are well known for their high .biodiversity Groombridge, 1992; Davis et al., 1997 .
But it is unclear which conditions in the past have
permitted the evolution of such high degree of diver-
0012-8252r98r$19.00 q 1998 Elsevier Science B.V. All rights reserved. .P I I : S 0 0 1 2 - 8 2 5 2 9 8 0 0 0 2 7 - 0
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( )H. Hooghiemstra, T. an der HammenrEarth-Science Reiews 44 1998 147183148
sity and, apparently, also such an effective conserva-
tion of species which developed during geological
time. Originally, the high biodiversity was ascribed
to the stability of the rain forest ecosystem during
the Quaternary. But gradually, it became clear that at
least parts of the tropical rain forest also had experi-
enced a dynamic history as a result of precipitation .change explaining the forest refugia , temperaturechange e.g., Van der Hammen, 1974; Gates, 1976;
.Colinvaux et al., 1989c, 1996; Bush et al., 1990 andriver dynamics Salo et al., 1986; Salo, 1987; Rasanen
.et al., 1987, 1992; Puhakka et al., 1992 . In fact, it is
unclear if environmental stability, or dynamic condi-
tions, have contributed to the present high biodiver-
sity of the tropical rain forests.
The objective if this paper is to provide a concise
history of the development of the neotropical rain
forest during the Neogene, to address the two hy-
potheses concerning the ice-age Amazon, and toprovide a bibliography concerning these topics.
2. Development of the South American tropical
rain forest since the Miocene
2.1. Upheaal of the Andes and deelopment of the
Amazon rier
The upheaval of the different Cordilleras of the
Andes during the Tertiary significantly changed the
river systems, atmospheric circulation and rainfallpatterns in northern South America. Locally, partial
upheaval occurred during certain intervals of theTertiary Van der Hammen, 1961; Case et al., 1971;
Van der Hammen et al., 1973; Kroonenberg et al.,
1990; Helmens and Van der Hammen, 1994; Cooper.et al., 1995 . There was relative subsidence of the
area immediately east of the Andes, giving rise to ahuge accumulation of sediments Rasanen et al.,
.1995; Paxton and Crampton, 1996 . Apart from un-
published studies of deep bore holes by oil compa-
nies, these sediment sequences are still little ex-
plored.
During the Middle Miocene, there were still con-
nections between the present-day Amazon basin and
the Caribbean through the Maracaibo area, and prob- . ably also with the Pacific Hoorn et al., 1995 Fig.
.1 . The rivers of northwestern Amazonia were run-
ning westward, to the Pacific, and northward to the
Caribbean, forming what might be called a paleo-
Orinoco river system reaching the Caribbean in theMaracaibo area. During the Middle Miocene 1610
.million years BP marine incursions occurred in the
Amazon basin during periods of high sea level stands,
leading to extensive mangrove vegetation in the pre-sent-day rain forest area Hoorn, 1993a, 1994a,b,c;
.Hoorn et al., 1995; Rasanen et al., 1995 . Estuarinelacustrine phases occurred when sea level
was lower, giving way to many types of swamp
vegetation and to seasonally inundated forest. Thesetemporal and spatial alternations between salt and
fresh water ecosystems caused a dynamic and di-
verse history for different geographical areas, possi-
bly stimulating floral evolution and biodiversity in
some areas and extinction in others. At places where
lower montane forest was near, altitudinal and latitu-
dinal shifts of vegetation belts may have stimulated
exchange of flora elements and subsequent ecologi-
cal adaptation. Therefore, it may be expected that
vegetation communities have changed continuously
through time. The present-day communities shouldbe regarded as the result of a very long and diverse
history, and a reflection of the present time-slice. As
an effect of the upheaval of the Eastern Cordillera of
the northern Andes, the mouth of the paleo-Orinoco
River is closed during the Late Miocene and the
present Orinoco River was formed. The rivers in the
northwestern part of the present-day Amazon basin
have changed their course toward the east, forming
part of the Amazon river system as we know it
today.
. . .Fig. 1. Paleogeographic maps of the early Middle Miocene a , late Middle Miocene b , and Late Miocene to Present c of northern South
America. The development of the northern Andes, the separation of Choco and Lower Magdalena rain forest areas from the main area ofAmazonian rain forest, and the Middel Miocene fluviolacustrine system in the present-day central Amazon basin is shown. At various
places in the present Amazon basin mangrove vegetation developed under brackish water conditions during periods of high sea-level stands.
Between the Late Miocene and the Quaternary the modern Orinoco River system developed from the paleo-Orinoco River, and the Amazon .River developed as a transcontinental drainage system towards the Atlantic. Modified after Hoorn et al. 1995 .
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The rain forest area of the Amazon basin became
separated from the rain forests along the PacificOcean, extending from Ecuador to Panama Choco
.biogeographic area , and in the Magdalena Valley .Hoorn et al., 1995 . The neotropical flora of north-
ern South America developed during the entire Ter-
tiary and got its final touch during the Pliocene andQuaternary Van der Hammen, 1974; Gentry, 1982a;
Van der Hammen and Cleef, 1986; Duque-Caro,.1990b; Hooghiemstra and Cleef, 1995 . Phytogeo-
graphic studies e.g., Cleef, 1979; Van der Hammen.and Cleef, 1986 show the source areas of that part
of the neotropical flora that reached northern South
America by migration.
2.2. Impoerishment of Amazonian flora since the
Miocene?
As far as the pollen flora in sediment cores from
the Amazon basin represent the species richness of .the vegetation phytodiversity correctly, we may
.draw some tentative conclusions. Hoorn 1994c
found ca. 280 pollen types in river valley sediments .of Miocene age, whereas Urrego-Giraldo 1994
found 140 pollen types in Holocene sediments of a
comparable environmental setting. It seems that
Miocene plant diversity may have been considerably
higher than today, and that we have to take into
account the possibility of later periods of significant .extinction. Also Wijninga 1996 speculates on the
basis of his palynological and paleobotanical studiesof sediment sequences of Miocene, Pliocene and
Quaternary age that an impoverishment of the
Neotropical flora could have occurred. More in gen-
eral, aspects of the history and ecology related to
biodiversity in the northern Andes and in Amazonas
were summarized and discussed in Van der Hammen .in press, a .
3. Impact of Quaternary climatic change
The final uplift of the Eastern Cordillera of
Colombia to its present-day elevation took place
mainly between 6 and 3 Ma. Upheaval was docu-
mented on the basis of palynological, paleobotanical,and geological studies Van der Hammen et al.,
1973; Helmens, 1990; Helmens and Van der Ham-.men, 1994; Wijninga, 1996 .
The climate and vegetation history of the Pliocene
and Quaternary of the northern Andes is relatively
well known and numerous papers deal with thisaspect e.g., Van der Hammen and Gonzalez, 1960,
1964; Van der Hammen et al., 1973, 1980r81;
Hooghiemstra, 1984, 1989; Kuhry, 1991; Kutzbach
et al., 1993, 1998; Hoorn, 1994c; Hooghiemstra and
Cleef, 1995; Hoorn et al., 1995; Van der Hammen
and Hooghiemstra, 1995, 1997; Wijninga, 1996;.Mommersteeg, 1998 .
At the end of the Pliocene, between 3.2 to ca. 2.5
Ma, there is a significant decrease of temperature on
a global scale. The Pleistocene starts with a major
cold period, considered as the first glacial of theQuaternary stage 100 of the marine oxygen isotope
.record . This cooling is also clearly registered in the
deep bore hole of Funza-II between ca. 470 m and400 m core depth Hooghiemstra and Ran, 1994;
Van der Hammen and Hooghiemstra, 1995, 1997;
.Hooghiemstra and Cleef, in prep. . Pleistocene tem-perature fluctuations at the high plain of Bogota .2550 m , were in the order of 88C.
XThe Funza-I pollen record from Bogota 4850 N;X . 74812 W shows that the 20,000-years rhythm pre-
.cession cycle of the Milankovitch climate forcing
was continuously present, whereas the 100,000-years .rhythm eccentricity cycle occurred only during the
.last ca. 0.8 Ma Hooghiemstra et al., 1993 . Climate
forcing in the precession band is of great importance
as it relates to the latitudinally shifting Intertropical
.Convergence Zone ITCZ , the major system thatdetermines the geographical distribution of precipita-
tion near the equator. As tropical rain forest requires
minimally 1500 1800 mm annual precipitation with-
out dry season, oscillations of the ITCZ in the past,
and the history of the related monsoon system, are of
crucial importance for the history of tropical rain
forest.
3.1. Quaternary climatic conditions in Choco
Air masses from the Pacific Ocean were continu-
ously forced to ascend above the narrow area be-tween the Pacific coast and the Western Cordillera of
Colombia causing heavy cloud formation and con-
vective rains. There is little reason to suppose that
this area with rain forest experienced significant
changes during the Quaternary. The impact of the
precession-forced migrations of the ITCZ-related rain
belt, as discussed for the Amazon basin, are most
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( )H. Hooghiemstra, T. an der HammenrEarth-Science Reiews 44 1998 147183 151
possibly masked by the heavy convective rains at the
western side of the Andes. For that reason it is
hypothesised that the continuous high precipitation
rates during the Quaternary facilitated a continuous
rain forest cover in the Choco biogeographic area.We assume that during the Quaternary Choco did notexperience significant perturbations of a magnitude
that could have impact on the geographical distribu- .tion of rain forest. Gentry 1986a,b mention the fact
that montane forest taxa, such as Podocarpus,
Hedyosmumand Ilex, are present at low elevation in
the very wet Pacific forest. These observations warn
us not to use the presence of montane forest trees at
low elevation as firm evidence of climatic cooling at .sea level Colinvaux, 1996 . Pollen records from
Choco that cover pre-Holocene time are not avail-able.
3.2. Quaternary climatic conditions in AmazonasDuring the Neogene, the neotropical montane and
Amazonian flora was enriched by migration of taxafrom southern source areas austral-antarctic ele-
.ments . After the connection between Central and
South America was established around 54 million .years ago Keigwin, 1982 , the flora was also en-
riched by elements from the holarctic phytogeo-graphic area Van der Hammen, 1974; Cleef, 1979;
Gentry, 1982a; Van der Hammen and Cleef, 1983,.1983r1984, 1986 .
The rich neotropical lowland flora, which devel-oped during the Miocene and Pliocene under alter-
nating conditions fresh water swamps and periods.with marine incursions suffered during the Quater-
nary the series of ice-ages. During ice-ages tempera-
tures at sea-level were some 58C lower and such
conditions lasted several times longer than the
warmer interglacial periods. In fact glacial conditionsare more normal than the present-day ones Colin-
.vaux, 1996, 1997 . These interglacialrglacial tem-
perature changes caused altitudinal shifts of vegeta-
tion belts in areas near mountains, e.g., along the
foot of the Andes and in centralreastern Brazil. This
may have caused exchange of elements between
floras of different altitudinal belts which may have
stimulated biodiversity in the periphery of the rain
forest area.
Northern South America was also influenced by
precession driven climatic change, which was evi-
denced in the long continental pollen record of Funza .for the Quaternary Hooghiemstra et al., 1993 . A
considerable part of the Amazon basin experienced
probably the above mentioned 20 kyr rhythmic
change in precipitation. During relatively dry inter-
vals of the precession cycle only the wettest parts ofthe Amazon basin ) 15001800 mm annual pre-
.cipitation without dry season kept its rain forest
cover, leading to the occurrence of forest refugia .sensu Haffer 1969 . Effects of orbital forcing on the
vegetation were also evidenced on the Yucatan .Peninsula Leyden et al., 1994 . Pollen diagrams
from the Colombian Andes and southern Amazonas
show that precipitation maxima are reflected as
forest phases in the Amazon basin, and as phases
with wet vegetation in the Colombian Andes and
occur at intervals of ca. 20,000 years distance, i.e.,
ca. 55,000 BP, ca. 35,000 BP, and ca. 6000 BP
.Hooghiemstra, 1995; Fig. 2 . Also the pollen record .of e.g., Salitre, south of Amazonia Ledru, 1993 ,
and the pollen record from Lake Valencia in .Venezuela Leyden, 1985 show similar precipitation
fluctuations.
4. Conflicting scenarios of the ice-age Amazon?
Paleoecological data from pollen records on the
Pleistocene history of the Amazonian rain forest is
still scarce, whereas first evidence from ColombianChoco rain forest is now available Behling et al., in
.press . Amazonian evidence is based on geomorpho-logical and geological studies e.g., Irion, 1976a,b,
1984, 1989; AbSaber, 1982; Bigarella and Ferreira,. 1985 , on palynological studies e.g., Van der Ham-
men, 1972, 1974; Absy, 1985; Colinvaux, 1987c,
1989c, 1996; Bush and Colinvaux, 1988; Liu and
Colinvaux, 1988; Absy et al., 1991; Van der Ham-
men and Cleef, 1992; Ledru, 1993; Van der Ham-
men and Absy, 1994; Urrego-Giraldo, 1997; Ledru.
et al., 1998a,b and biogeographical patterns Haffer,1969; Prance, 1973, 1978, 1982b; Andersson, 1979;.Brown, 1982; Gentry, 1982b; Beven et al., 1984 .
During the past years, there has been considerable
discussion about these data and hypotheses, and
several new theories have been put forward, to ex-
plain the very high biodiversity of the Amazonianrain forest and the diversity patterns Colinvaux,
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. . Fig. 2. Pollen records of Carajas Brazilian lowland forest , Lake Fuquene Colombian montane forest belt , Lake La Primavera .Colombian paramo . Vegetation indicative of high precipitation is present around 27,000 BP and 6000 BP: forest in Carajas and a
and TPN 21B. Vegetation indicative of low precipitation is present around 18,000 BP: savanna in Carajas and dry paramo in Fuqu . . . .Geel and Van der Hammen 1973 , Melief 1985 and Salomons 1986 , respectively. Taken from Hooghiemstra 1995, 1997 .
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( )H. Hooghiemstra, T. an der HammenrEarth-Science Reiews 44 1998 147183 153
.1996, 1997 . Here we will discuss briefly the palyno-
logical evidence.
4.1. Late Quaternary pollen records of Amazonas
and the forest refugia hypothesis
.When Haffer 1969 put forward his forest refugia
hypothesis on the basis of zoological evidence, no
pollen records existed from the Amazonian rain for-
est. Based on distribution patterns of Amazonian
forest birds, Haffer recognized centres of dispersal,
which were assumed to reflect refugia where the
tropical fain forest ecosystem survived climatologi-
cal dry conditions. Areas outside such refugia as-
sumedly experienced so much reduction in precipita-
tion that rain forest was replaced by savanna and
savanna forest vegetation.The first pollen record of Amazonas Katira; 98S;
.638W was published in 1972 by Van der Hammen.Subsequently, more records became available. Paly-
nologists working in areas where during the last
ice-age rain forest was replaced by savanna, e.g., inRondonia Van der Hammen, 1972, 1974; Van der
.Hammen and Absy, 1994 supported the forest refu-
gia hypothesis. But palynologists working in other
areas of the Amazon basin, e.g., in the Ecuadorian XAmazonas and the area of Lake Pata 0816 N;
X . 66841 W Colinvaux, 1987a,b, 1996; Bush et al.,.1990 did not find evidence for a replacement of rain
forest by savanna. The controversy about Haffersforest refugia hypothesis was born. Papers discussing
and supporting the forest refugia hypothesis are e.g., . .Simpson-Vuilleumier 1971 ; Prance 1973, 1982a ;
. .Brown et al. 1974 ; Van der Hammen 1974 ; An- . .dersson 1979 ; Brown and AbSaber 1979 ; Haffer
. .1979, 1982, 1987a,b ; Steyermark 1979, 1982 ; . .Andrade-Lima 1982 ; Gentry 1982c, 1992 ; Simp-
. .son 1982 ; Mayr and OHara 1986 ; Aguilar-Del- . .gado 1987 ; Van der Hammen and Absy 1994 ,
.and Hooghiemstra 1997 . Papers claiming the forest
refugia hypothesis cannot be supported by data are .e.g., Colinvaux 1979, 1987c, 1996, 1997 ; Endler
. . .1982 ; Livingstone 1982 ; Nelson et al. 1990 , and .Haberle 1997 .
4.2. Two scenarios
According to our view, both of the above-men-
tioned scenarios did occur. In fact, four aspects are
important when the forest history of the Amazon
basin is considered. .1 The annual migration of the caloric equator
.ITCZ between about 88N in July and 38S in Jan-
uary causes an annual latitudinal shift of the equato-
rial rain belt, leading to seasonal variations in pre-
cipitation. Most of the area has two dry and two
humid periods, whereas the distal areas experience
only one dry and one humid season. . 2 The precession cycle of orbital forcing e.g.,
.Hays et al., 1976; Berger, 1989 causes an oscillation
of the equatorial rain belt as described under the first
point with a period of about 20,000 years. Under the
present-day orbital configuration the southern hemi-
sphere is tilted towards the sun and the caloric .equator ITCZ lies south of the geographical equa-
tor. This configuration brings most precipitation to
the central and southernmost part of the Amazon
basin. Half a precessional cycle in the past 11,000.years ago , the northern hemisphere was tilted to-
wards the sun and the caloric equator was north of
the geographical equator. This configuration brings
most precipitation to the northernmost part of the
Amazon basin and adjacent Caribbean. .3 The temperature record of the Quaternary was
modulated by the glacialrinterglacial cycles. Long
pollen records from the Colombian Andes at 2550 melevation, such as Funza-I and Funza-II e.g.,
.Hooghiemstra and Ran, 1994 , Funza IIArFuquene-
. X
VII Mommersteeg, 1998 , and Fuquene-II 5827 N;X . .73846 W Van Geel and Van der Hammen, 1973
show with high resolution the temperature record of
northern South America with changes of some 88C.
But it is widely accepted now that the tropical
lowlands experienced also temperature oscillations
of some 58C during the last glacialrinterglacial cyclee.g., Colinvaux, 1987c, 1996; Bush et al., 1990;
.Van der Hammen and Absy, 1994 . It seems plausi-
ble that the composition of the Amazonian lowland
forest changed during the Quaternary. Montane arbo-
real taxa could reach lower elevations under condi-tions of lower temperature andror high precipitation .Gentry, 1986a,b . The modern forest composition
should be seen as the situation of the present time-
slice rather than a constant characteristic of the
Quaternary. .4 The concave shape of the eastern slopes of the
Andes between 58N and 158S act as a trap for humid
Atlantic air masses causing continuous convective
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( )H. Hooghiemstra, T. an der HammenrEarth-Science Reiews 44 1998 147183154
rains in northwestern Amazonas, irrespective of pre-
cession forcing. Therefore, the northwestern part of
the Amazon basin most possibly received continu-
ously high precipitation and explains in this area a
continuous cover of rain forest during the Quater-
nary. Also other minor areas may have experienced a
continuous rain forest cover related to a complex,
and yet poorly understood, pattern of precipitation
related to Walker circulation and an Amazonian
Convergence Zone.
Considering the four aspects mentioned above, it
is very plausible that the paleo-ecological history
was markedly different depending on the geographi-
cal location in the Amazon basin. A continuous
forest cover, expansion of savannas and dunes, and
lowering of temperatures occurred. The hypothesis
claiming a continuous rain forest cover in the Ama- .zon basin put forward by the school of Colinvaux ,
and the forest refugia hypothesis put forward by theschool of Haffer, Prance, Gentry, and Van der Ham-
.men do not exclude each other but reflect two
extremes out of a spectrum of different regional
paleo-ecological histories which are summarized in
the following section.
4.3. Regional egetation histories of the Amazon
basin
( )4.3.1. Site Katira 98S; 638W
Among the first palynological papers on the vege-
tation history of Amazonas was the pollen record of . .Katira Rondonia by Van der Hammen 1972, 1974
.and Absy and Van der Hammen 1976 . This site is
located in the rain forest area and the sequence
represents the sediment infill of a minor valley. The
interpretation of this sequence was improved when
some years ago AMS radiocarbon dates became
available. The pollen record shows rain forest vege- .tation around 50,000 BP Middle Pleniglacial age
and vegetation dominated by grass savanna around .18,000 BP Pleniglacial age Van der Hammen and
.Absy, 1994; Van der Hammen, in press, a . Todaythe site is in the rain forest area and recent sediments
from the area show that pollen grains of rain forest
elements dominate.
( X X )4.3.2. Site Carajas 15832 S; 47840 W
Another important pollen record from lake sedi-
ments comes from the top of table mountains at ca.
700 m elevation in the rain forests of the Carajas
area in eastern Amazonia. The vegetation on the
table mountains consists of edaphically determined
low forest and shrub and open savanna like vegeta-
tion. Rain forest covers the area around the tablemountains and their slopes Absy et al., 1991; Van
der Hammen, 1992; Van der Hammen and Absy,. .1994 . The radiocarbon dated sequence Fig. 2 rep-
resents the last approximately 60,000 years and the
pollen record shows three intervals during which
savanna vegetation is dominant: around ca. 65,000
BP, ca. 40,000 BP and the period of ca. 25,000 to
10,000 BP. These periods of savanna vegetation
alternate with three periods in which forest vegeta-tion dominate around 55,000 BP, 35,000 BP, and
.6000 BP . Three dry savanna periods and three wet
forest periods are evidenced and do show a rhythm
of ca. 20,000 years, related to the precession cycle
.Hooghiemstra, 1995 .
4.3.3. Sites Pantano de Monica and Araracuara
Another relevant pollen diagram is from sedi-
ments of a forest-marsh on top of the lower terrace
of the Caqueta River, south of Mariname Island X X .0844 S; 72804 W in the Araracuara region of
Colombia Amazonas Urrego-Giraldo, 1994, 1997;.Van der Hammen, in press, a . Radiocarbon dates of
this approximately 4-m sequence indicate Upper
Pleniglacial, Late Glacial and Holocene age. The
Late Pleistocene part is dominated by forest vegeta-tion, but the record shows a specific composition
which is considerably different from the Holocene
forest composition. During the Upper Pleniglacial
part Myrtaceae dominate the forest, followed by
Ilex; the presence of some Gramineae pollen grains
and especially of some grains of Podocarpus is
notorious. In this area, there was apparently no Late
Pleniglacial replacement of rain forest by dry sa-
vanna-like vegetation. The local forest composition
was partly different and there might have been lo-
cally more open vegetation, or caatinga type ofvegetation, on sands of the higher terraces. So, it
seems that under climatic marginal conditions,
edaphic factors play an important role in changes in
the composition of forest. This was also observed in X X .pollen record Araracuara-1 0840 N; 72830 W , lo-
cated in the Bonnetia-dominated shrub savannas near
the air strip of Araracuara: a sharp transition from
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( )H. Hooghiemstra, T. an der HammenrEarth-Science Reiews 44 1998 147183 155
Rapateaceae and Xyridaceae dominated open sa-
vanna into Bonnetia-dominated shrub savanna and
dwarf forest was registered. Radiocarbon dates are
lacking in these sandy sediments extremely poor in
organic carbon, but most probably this record repre-sents Late Holocene time Hooghiemstra, unpub-
.lished data .
( X X )4.3.4. Site Lake Pata 0816 N; 66841 W
Based on 12 radiocarbon dates, the sediments of .the upper 160 cm of core Lake Pata Fig. 3 repre-
.sent the last 42,000 years Colinvaux et al., 1996b .
The pollen diagram includes some 45 pollen spectra
and represent an ice-age record like the Carajas
diagram. Reduced precipitation is evidenced from
roughly 30,000 to 14,000 BP. The pollen record
shows, however, that sufficient precipitation re-
mained to support rain forest. Increased representa-
.tion during the last glacial maximum LGM ofcool montane trees, such as Podocarpus, Hedyos-
mum and Ilex, may indicate that temperatures had
lowered. These results are in many respects in agree-
ment with other information. Lake Pata is located in .a predicted forest refugium Fig. 3 and the pollen
record supports the concept published by Van der .Hammen and Absy 1994 . Also the more dominant
role of Ilex in the last glacial Amazonian forests isin agreement with the records from Araracuara Van
.der Hammen, in press, a; Behling et al., in press .
The dry period from roughly 30,000 to 14,000 BP
corresponds to the period of dry conditions regis- .tered in Carajas Fig. 2 , but also in the Colombian
Andes e.g., Lake Fuquene; Van Geel and Van der.Hammen, 1973 and is most possibly related to the
precession cycle.
( )5. The last glacial maximum LGM in Amazonas
First CLIMAP estimates indicated for the LGM
sea surface temperatures only a small temperaturedecrease of ca. 128C CLIMAP Project Members,
.1976, 1981 . However, palynological data from alti-
tudes between 2500 and 4000 m in the Colombian
and Venezuelan Andes showed during the LGM a
temperature decrease of 798C as compared to mod-
ern conditions. The glacial lapse rate may have been
slightly higher than today because of the drier air: .0.7 instead of 0.68C per 100 m Bakker, 1990 .
. .Colinvaux 1989c and Bush et al. 1990 inferred
from Amazonian pollen records a LGM temperature
decrease of maximally 68C. This estimate has been
based on increased percentages of pollen grains of
montane forest taxa, mainly Podocarpus and
Hedyosmum. The question how much temperature
lowered at sea level during the LGM is still in .debate. The observations by Gentry 1986a, 1986b
that montane trees do occur at low elevation under
very high precipitation regimes, such as in Choco,seems of crucial importance. But at this moment we
do not have a calibration tool to use this information
for quantitative temperature estimates. The lowering
of temperature in the tropical lowlands during glacial
time is still in debate. There might be an uncertainty
of "28C, but for the time being a value of 48" 28C
for the LGM temperature depression in the Amazonbasin seems to be a safe figure and interval CLIMAP
Project Members, 1976, 1981; Rind and Peteet, 1985;Colinvaux, 1989c, 1996; Bush et al., 1990; Anderson
and Webb, 1994; Guilderson et al., 1994; Van der
Hammen and Absy, 1994; Broecker, 1995; Stute et
al., 1995; Colinvaux et al., 1996a,b; De Oliveira and.Colinvaux, 1996; Van der Hammen, in press, a .
5.1. Reduced LGM precipitation and the geographi-
cal location of forest refugia
In view of the fact that the climate was much
. drier north Lake Valencia, Venezuela , west Co-. .lombian Andes and south Brazil of Amazonia and
inside parts of Amazonia Rondonia, Carajas, Lake.Pata during the LGM, we may as a first approxima-
tion, and on the basis of the available data, estimate
the effect of the reduction of precipitation in the
entire area of Amazonia. For that aim we may use
the modern precipitation map of Amazonia and the .sites of Katira Van der Hammen and Absy, 1994 ,
Georgetown Wijmstra and Van der Hammen, 1966;. Van der Hammen, 1974 and Carajas Absy et al.,
.1991 , where the forest disappeared, and Araracuara .Van der Hammen et al., 1992a and Lake Pata .Colinvaux et al., 1996b , where the forest did not
.disappear Fig. 3 . Moreover there is the fact that
today we find savannas, and dry forest or cerrado,
where the annual rainfall is less than 1500 mm. If the
rainfall in the area was reduced by 500 mm, Carajas
would be in the savanna zone, and if it were reduced
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1000 mm, Georgetown and Katira could also be
under savanna, whereas Araracuara and Lake Pata .would remain under forest Fig. 3 . Notorious is the
permanent existence of rain forest in a large area in
western to northwestern Amazonia, besides other
relatively large areas in the centre and northwest of
Amazonia. This reconstruction is, of course, only a
first approximation that should be corrected on the
basis of new and more detailed precipitation maps
and new palynological data.
This hypothesis of a reduced modern precipita-
tion pattern to explain the geographical position of
the LGM forest refugia only makes use of a globally
lower precipitation level during the LGM and the
geography of the Andes. The earlier mentioned as-
pects of precessional climate forcing and the
glacialrinterglacial temperature oscillations will
make this assumption more complex. We assume
that, depending on the geographical area, the vegeta-tion in the Amazon basin during the Late Quaternary
was either permanently rain forest, possibly partly
with a different forest composition compared to pre-
sent-day situation, or experienced periods with
semi-deciduous tropical forest, or even was replaced
by different types of savanna-like vegetation, all
depending changes in annual precipitation, seasonal-
ity in precipitation, temperature changes, and geo-
graphical setting. But in fact the hypothesis of a
reduced modern precipitation pattern is elegant in the
sense that it provides good possibilities to be testedwhen evidence from new sites comes available. In
this respect the results of the newly published42,000-years-long pollen record of Lake Pata Colin-
.vaux, 1996 are interesting: there is evidence of a
drier climate but forest was not replaced by
savanna-like vegetation during the LGM. This site is
located in the area indicated by Van der Hammen .and Absy 1994 as a potential forest refugium.
Therefore, the record of Lake Pata is unable to
invalidate the forest refugium hypothesis and is in
agreement with the pattern predicted.
5.2. Eidence of high precipitation in the Andes and
Colombian Amazonas
During the Middle Pleniglacial cold and wet pe-
riod, in the Andes considerable quantities of gravels
and sands, partly of fluvioglacial origin, were de-posited in the valleys Van der Hammen et al.,
.1980r81 . In the same period terrace gravels, sands
and silts were deposited in the Middle Caqueta area,and probably also by other west Amazonian rivers .Van der Hammen et al., 1992b . After ca. 30,000
BP during the cold and dry Late Pleniglacial, rivers
incised in these sediments and only started sedimen-
tation again during the Late Glacial and Holocene,viz. after approximately 12,500 BP Van der Ham-
.men et al., 1992b . In lower Amazonia, rivers incised
also in earlier terrace sediments, in this case because
of the low sea-level stand during glacial age. During
the following Late Glacial and Holocene sea-levelrise, the lower Amazon valley became a large
estuary, until it was filled up with sediments, and
the present-day broad zone with varzea vegetation,
characterized by seasonal inundation, was formed. It
is clear, therefore, that not only the vegetation suf-
fered the effect of the glacial climate change, but
also the river systems.
6. The ice-age Amazon as reflected in the offshore
marine sediments
Pollen analysis of offshore marine sediment cores
have at several locations proven to be a valuable
source of information concerning environmental and
climatic change on the adjacent continent. Therefore,
a palynological study of cores from the Amazon fan
was expected to be a source of information of the
environmental conditions of the ice-age Amazon.
However, there are several reasons to doubt the
value of the pollen signal from Amazon fan sedi-
ments.
.Fig. 3. Maps showing the modern distribution of rain forest in relation to the present-day 2500, 2000, and 1500 mm precipitation isoline a . . . .After a reduction of precipitation of 25% b and 40% c new 1500 mm iso-contours show the potential area with rain forest. Scenario c
meets the available palynological data for the glacial Amazon: presence of rain forest in Mera, Araracuara and Pata, and replacement of rain .forest by savanna-like vegetation in Katira Rondonia , Carajas, and Georgetown. The situation might also have been intermediate between
. . . .scenarios b and c , forest areas being larger than indicated in c . Modified after Van der Hammen and Absy 1994 .
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First, many different environments supply pollen
to the Amazonian drainage system, of which the
interpretation may be unclear: graminaceous pollen
grains may reflect open savanna-like vegetation, but
also floating grass-rich meadows on the Amazonian .rivers and in the varzeas e.g., Hoorn, 1994c , which
makes the interpretation of such records unsure.
Secondly, glacial periods with sea-level stands
some 120 m below the present-day level caused
massive erosion in the lower parts of the Amazon
drainage system. These sediments have been trans-
ported to the offshore Amazon fan and have been
re-deposited there again. This process must have
caused a mixing of pollen grains from different
source areas and of different ages, possibly including
sediments from earlier glacial-interglacial cycles.
Both aspects should be taken into account when the
pollen signal of sediments from the Amazon fan are
used to infer glacial environmental conditions of theinterior of the Amazon basin.
A third aspect relates to the gallery forests that
often accompany the river valleys in savanna areas.
Such forests prevent pollen grains from savanna
vegetation to reach the river system and we expect
savanna vegetation is poorly represented in the
river-transported pollen spectrum.
The pollen record from core Leg 155 shows a
very monotonous representation of pollen taxa from
all possible source areas: montane forest, lowland
rain forest, gallery forest along rivers, swamps, opensavanna vegetation, aquatic vegetation and fern
.spores Haberle, 1997; Hoorn, 1997 . Moreover, the
interpretation of the pollen record of Gramineae is
ambiguous, as grasses may represent open savanna-
like vegetation, as well as floating meadows rich in
grasses, that occur frequently on Amazonian riverssee also the discussion in Hoorn, 1994c and Hoorn
. . et al., 1995 . Haberle 1997 and Colinvaux 1996,.1997 interpret the monotonous records and low
representation in the Amazon fan sediments of pollen
grains from open vegetation types as evidence of low
representation of savanna-like vegetation and a sta-
ble rain forest cover since the last interglacial. For
the three reasons mentioned above, we think such
conclusions are not justified and Amazon fan pollen
records are very difficult to use as a document to
infer environmental change in the huge drainage
basin.
7. Conclusions
.1 The upheaval of the northern Andes since the
Middle Miocene led to a separation between the rain
forest of Amazonia at one hand, and those of the
Choco and the Magdalena Valley at the other hand. .2 In the intracontinental basin between the old
shields and the newly formed Andes, estuarine
lacustrine phases occurred when sea level was low,
and coastal environments with mangrove vegetation
occurred during high sea-level stands. These tempo-
ral and spatial alternations between salt and fresh
water ecosystems caused a dynamic and diverse
history for different geographical areas, possibly
stimulating floral evolution and biodiversity in some
areas and extinction in others. .3 Although data are scarce, it may be possible
that extinction of plant taxa was a more common
phenomenon in the Quaternary than previouslythought. MiocenerPliocene plant diversity may have
been larger that at present-day. Huge modern phyto-
diversity perhaps should be better regarded as a
legacy from the Tertiary rather than as a product of
the Quaternary. .4 Precession-related changes in the geographical
.position of the caloric equator equatorial rain belt ,
and river dynamics as a result of small tectonic
movements, have possibly been constant factors of
stress on the Amazonian rain forest ecosystem. In the
Quaternary significant changes in temperature, re-lated to the series of ice-ages, increased environmen-
tal stress. .5 The forest refugia hypothesis and the opin-
ion that Amazonian rain forest was not replaced by
savanna-like vegetation during the last ice-age under
climatic dry conditions are not necessarily conflict-
ing. According to our view both scenarios did occur
and represent the extreme scenarios under dry and
wet climatic conditions. .6 Four aspects are important when the forest
.history of the Amazon basin is considered: a the .annual migration of the caloric equator ITCZ be-
tween about 88N and 38S causing an annual latitudi-
nal shift of the equatorial rain belt, leading to sea- .sonal variations in precipitation; b the precession
cycle of orbital forcing causing an oscillation of the
equatorial rain belt with a period of about 20,000 .years; c the temperature oscillations at sea-level of
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some 58C during Quaternary glacial cycles causing
the composition of the Amazonian lowland forest .changed all over the Quaternary of composition; d
the concave shape of the eastern slopes of the Andes
between 58N and 158S act as a trap for humid
Atlantic air masses causing continuous convective
rains in northwestern Amazonas, irrespective of pre-
cession forcing. This explains continuous forest cover .forest refugium in northwestern Amazonas.
.7 The modern composition of Amazonian rain
forests should be seen as the situation of the present
time-slice, rather than a constant characteristic of the
Quaternary. .8 Using a simple reduced modern precipitation
pattern for the Amazon basin the geographical posi-
tion of the LGM forest refugia can be estimated. A
reduction of precipitation between 25 and 40% ex-
plains the available pollen evidence, including sites
with an uninterrupted forest cover Mera, Araracuara,.Pata and sites where rain forest was replaced by
.savanna Katira, Carajas, Georgetown . .9 The monotonous pollen records for all ecolog-
ical groups from sediment cores of the Amazon fan
evidence a thoroughly mixed pollen association orig-
inating from many different vegetation types in the
huge Amazonian drainage basin. The monotonous
grass record cannot be regarded as evidence for the
absence or existence in the Amazon basin of periods
with much savanna. We postulate savanna is poorly
represented in the river-borne pollen signal becauseof the gallery forests along the rivers. Pollen records
from Amazon fan sediments are unsuitable to infer
with any detail vegetational and environmental
changes in the remote hinterland.
Acknowledgements
The first author would like to thank the Nether- .lands Foundation for Scientific Research NWO for
the invitation to participate in the annual Huygens
Lecture, which was the immediate motive to prepare
this paper. Paul Colinvaux is thanked for his gra-
cious contribution in this lecture evening between
opponents. Hermann Behling is thanked for provid-
ing additional references. An anonymous reviewer is
thanked for critical comment and improvement of
the English. This paper is dedicated to the scientific
community of the PoleEquatorPole transect
.through the Americas PEP-I , which forms an exit-
ing forum for interdisciplinary cooperation.
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