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Tertiary 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 been
evidenced 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. q1998 Elsevier Science B.V. All
rights reserved.
Keywords: Neogene; Quaternary; rain forest; forest refugia; Amazonas
1. Introduction
Apart from paleoclimatological studies using ma-
rine sediments, palynological and paleoecological
studies 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: hooghiemstra@bio.uva.nl
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 of
the 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.
. PII: S0012- 8252 98 00027- 0
( ) H. Hooghiemstra, T. an der HammenrEarth-Science Reiews 44 1998 147183 148
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 , temperature
X
The 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 that
determines the geographical distribution of precipita-
tion near the equator. As tropical rain forest requires
minimally 15001800 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
( ) 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 not
experience 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,
Hedyosmum and 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 Amazonas
During the Neogene, the neotropical montane and
Amazonian flora was enriched by migration of taxa
a
n
d
e
r
H
a
m
m
e
n
r
E
a
r
t
h
-
S
c
i
e
n
c
e
R
e
i
e
w
s
4
4
1
9
9
8
1
4
7
1
8
3
1
5
2
. . . Fig. 2. Pollen records of Carajas Brazilian lowland forest , Lake Fuquene Colombian montane forest belt , Lake La Primavera Colombian paramo and profile TPN 21B
. Colombian paramo . Vegetation indicative of high precipitation is present around 27,000 BP and 6000 BP: forest in Carajas and a wide paramo belt in Fuquene, La Primavera
. and TPN 21B. Vegetation indicative of low precipitation is present around 18,000 BP: savanna in Carajas and dry paramo in Fuquene. Diagrams after Absy et al. 1991 , Van
. . . . Geel and Van der Hammen 1973 , Melief 1985 and Salomons 1986 , respectively. Taken from Hooghiemstra 1995, 1997 .
( ) 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.
X
Amazonas 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 Haffers
forest 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
X X
.
0844 S; 72804 W in the Araracuara region of
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
( ) 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-