A Holocene vegetational and climatic record from the Atlantic rainforest belt of coastal State of...

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Review of Palaeobotany and Palynology 131 (2004) 181–199

A Holocene vegetational and climatic record from the Atlantic

rainforest belt of coastal State of Sao Paulo, SE Brazil

Maria Judite Garcia*, Paulo Eduardo De Oliveira,Eliane de Siqueira, Rosana Saraiva Fernandes

Laboratorio de Geociencias, Universidade Guarulhos, Prac�a Thereza Cristina 1- Centro, Guarulhos, Sao Paulo CEP 07023-070, Brazil

Received 6 February 2003; accepted 26 March 2004

Abstract

Holocene vegetation and climate have been reconstructed by means of pollen analysis in a 643-cm-long core from the

Jacareı peat deposits (23j17VS, 45j58VW, 550 m a.s.l.) within the Atlantic tropical rainforest belt in the State of Sao Paulo, SE

Brazil.

Three conventional 14C dates indicate that the onset of peat formation started at 9720 years BP. Due to sampling restrictions,

the record encompasses the period between 9720 and ca. 1950 years BP. The palynological content of the samples permitted the

recognition of five distinct climatic periods between 9700 and ca. 1950 years BP: humid and cool climate from 9720 to ca. 8240

years BP, humid and warm from ca. 8240 to ca. 3500 years BP, cooler and moister than today from 3500 to 1950 years BP. The

return of a cool climate at the late Holocene is suggested by the reappearance of montane and humid forest taxa such as

Araucaria, Drimys, Daphnopsis, Ericaceae, Podocarpus and Myrsine in the upper sections of the pollen diagram.

Throughout its formation, the Jacarei peatbog has had a very different botanical composition. Gleichenia was the most

important taxon in the peat bog from 9720 to 8240 years BP, followed by Selaginella, Polypodium and Asplenium until 5400

years BP. A Sphagnum/Lycopodium dominated peat was established from 5400 to 3500 years BP, followed by Gleichenia/

Sphagnum from 3500 to ca. 1950 years BP.

The interpretation of the pollen and spore diagrams permitted a correlation between the vegetational and climatic signal

contained in the Jacareı peatbog with other locations in southeastern and southern Brazil.

D 2004 Elsevier B.V. All rights reserved.

Keywords: Holocene; pollen; Sao Paulo; Atlantic forest; Araucaria; Brazil

1. Introduction

The Brazilian Atlantic Forest domain, together

with Amazonia, represents one of the most floristical-

ly diverse regions of the world (Silva and Filho, 1982;

0034-6667/$ - see front matter D 2004 Elsevier B.V. All rights reserved.

doi:10.1016/j.revpalbo.2004.03.007

* Corresponding author.

E-mail address: [email protected] (M.J. Garcia).

Oliveira-Filho and Fontes, 2000). One of the most

important attributes of its high bioversity can be

illustrated by the impressive levels of endemism.

According to Morellato and Haddad (2000), nearly

50% of all animal and plant species belonging to the

Atlantic Forest region biota are endemic.

Despite its biological importance, very little is

known about the vegetational and climatic history

of this ecosystem because the low number of pollen

M.J. Garcia et al. / Review of Palaeobotany and Palynology 131 (2004) 181–199182

records available in the literature. One important

source of information on the environmental history

of the Brazilian Atlantic Forest are the pollen rich

peatbog deposits of the Paraıba do Sul River

Valley, which cover a 2400-km2 area (Garcia, 1994).

These peats are characterized by lithologies of 6–

20 m in thickness and overlie basal strata varying

from clays to sand with dates ranging from the late

Pleistocene (>30,000 years BP) to the late Holocene

(Turcq et al., 1992). These lithological character-

istics and the highly sinuous aspect of a long

meandering riverine system are suggestive of former

climatic change that has occurred regionally during

the late Quaternary.

Presently, the organic nature of these sediments

allows extensive cultivation of rice and vegetables

along the valley, which are exported to urban areas of

southeastern Brazil. The use of the peatbog sediments

as a source of fertilizers for crops, fuel and of a

variety of industrial products (Shimada and Carvalho,

1980; Villwock et al., 1980; Mendes and Costa, 1981;

Kumoto et al., 1985), indicates its importance to the

local economy. These economical activities have

dramatically intensified during the last decade and

resulted in a widespread destruction of the organic

sediments of the valley, which potentially bear pale-

ocological records. We have sampled three peat

locations in the valley prior to the establishment of

destructive practices caused by agriculture and peat

exploitation. We present here only the results for the

Jacareı site. The other two pollen records will be

published elsewhere.

Because there is no information from late Quater-

nary pollen studies in the valley, paleoenvironmental

information has been indirectly derived from radio-

carbon dates of selected peat cores studied by Mello

and Moura (1991), Suguio et al. (1992), Turcq et al.

(1992) and Moura and Mello (1991).

Based on neoctectonical data, Riccomini (1989)

proposed that, from an extrapolated age of 52,000 to

27,480 years BP, erosional events produced extensive

alluvial deposits with the formation of peatbogs

occurring only in the last 2000 years. On the other

hand, a much older date of 20,160 years BP for the

beginning of peat accumulation in the valley was

revealed by a 6-m peatbog core raised by Turcq et

al. (1992) in Jacareı, Sao Paulo. This allowed the

authors to conclude that the formation of these peat-

bogs was controlled primarily by a rise of the water

table at 20,000 years BP and after 8000 years BP.

These investigations suggest that the peat deposits

of the Paraıba River Valley have been formed at least

since the late Pleistocene, possibly under different

climatic regimes.

The primary objective of this study was to recon-

struct the late Quaternary paleovegetation and paleo-

climatic history from the Jacarei peat deposits and to

correlate it with other palynological records from

different vegetation types of southeastern, southern

and southwestern Brazil.

2. Study site

2.1. Geographical setting of the Jacareı peat deposits

The Jacareı peat site (23j17VS, 45j58VW, 550 m

elevation) is located in the mid section of the Paraıba

do Sul River valley, geologically interpreted as a

graben of 20 km width and 173 km length, extending

in a SW–NE direction. The extensive peat deposits,

located at 550 m elevation, are delimited by the Serra

da Mantiqueira and the mountain complex of the Serra

do Mar which border the valley on its western and

eastern flanks, respectively (Fig. 1).

2.2. Climate

Precipitation patterns within the valley are directly

related to the influence of the west/east shifts of the

South Atlantic High Pressure System, especially dur-

ing the southern hemisphere summer months (Decem-

ber, January and February) and the seasonal shifts of

the Polar Air Mass System, during the winter. During

the summer, the influence of equatorial air masses

becomes important as the ICTZ (Intertropical Conver-

gence Zone) reaches its southernmost limit in March

(Nimer, 1989a). Precipitation in the valley ranges

from 1200–1500 mm in the lowlands to 2500 and

3000 mm in the Serra da Mantiqueira and Serra do

Mar, respectively (Radambrasil, 1983). The number

of dry months is generally not higher than 2.

Mean annual temperature ranges from 19 to 21 jCin the lowlands to < 15 jC at elevations higher than

2000 m (Radambrasil, 1983). At Jacareı, the mean

temperature for July, the coldest month, is 15 jC. The

Fig. 1. Study site location in relation to the Serra da Mantiqueira and the Serra do Mar complex. The Paraıba do Sul River, which initially flows in a northeastern–southwestern

direction, is fed by streams born in both mountainous regions. The peatbogs of the valley occur between 46j00 and 45j15VW longitude.

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effects of altitude on temperature, in this portion of

southeastern Brazil, can be documented by the num-

ber of days with frosts per year: Campos do Jordao, at

1600 m elevation, has an average of 46 days with frost

whereas the average for Itatiaia, at 2199 m, is 56 days

(Nimer, 1989b). By comparison, the number of days

with frost at Jacareı is 3–4 (Nimer, 1989a).

According to the Koppen classification system, the

climate of the lowlands is Cwa—representing a hot

and humid climate with dry winters. The high-eleva-

tion area of the surrounding mountains has a climate

identified as Cwb, a temperate climate, with dry

winters (Garcia, 1994). The climate of the Paraıba

do Sul River valley, as for most of southeastern Brazil,

is controlled by the semi-permanent high-pressure

system of the South Atlantic Anticyclone, which

generates easterly–northeasterly tropical air flow. Ad-

ditional moisture reaches the region during the sum-

mer (December – January –February) and winter

(July–August–September) months related to the an-

nual southward displacement of the ITCZ and inva-

sions of cold Antarctic polar fronts, respectively

(Nimer, 1989a).

2.3. Vegetation

Historical documents indicate that prior to the

arrival of European settlers in the 1500s, the vegetation

of the middle Paraıba do Sul River Valley was com-

posed primarily of tropical forests. According to

Radambrasil (1983), the original vegetation of the

Paraıba do Sul River Valley was made up of evergreen

Atlantic rainforest on both humid mountain ranges of

the Serra do Mar and Serra da Mantiqueira and by a

seasonal semi-deciduous forest within the valley (Figs.

2 and 3). Large tracts of this seasonal forest can still be

found in some locations. According to Nimer (1989a),

the distribution of the different vegetation types in the

valley appears to be in response to modern climatic

patterns. For example, the high-altitude Araucaria

forests occur within the 13 jC isotherm of mean annual

temperature. Mean annual precipitation is >1500 mm,

with 1–2 dry months. The montane Atlantic forest on

the eastern and western facing slopes of the Serra da

Mantiqueira and Serra do Mar complex is under 2000

mm mean annual precipitation, and 1–2 dry months

(Nimer, 1989a). According to Radambrasil (1983),

mean annual precipitation in the Paraıba do Sul valley

is only 1200 mm, a direct consequence of the rain

shadow effect produced by the eastern mountain ranges

of the Serra do Mar complex. The semi-deciduous

tropical forests of the studied site area are characterized

by arboreal taxa such as Aspidosperma, Bombax,

Centrolobium, Copaifera, Cedrela, Chorisia, Dalber-

gia, Esenbeckia, Lecythis, Melanoxylon, Ocotea,

Schizolobium and many others.

With increasing elevation, different forest compo-

sitions are observed within the domain of the Atlantic

rainforest. For example, at elevations above 1500 m

cold and moist adapted taxa prevail. These are Arau-

caria angustifolia, Podocarpus lambertii, Drimys

brasiliensis, Clethra brasiliensis, Clusia, Ericaceae

(Gaylussacia and Leucothoe) Hedyosmum brasi-

liense, Melastomataceae, Myrtaceae, Vochysia lauri-

folia, Talauma organensis, Cariniana excelsa,

Ocotea, Myrsine, Roupala, Ilex, Weinmannia and

others. These lush montane forests, characterized by

Araucaria predominating in the canopy as an emer-

gent tree, have similar botanical composition and

physiognomy to the Araucaria forests found in south-

ern Brazil (Wanderely et al., 2002; Joly, 1976).

Arboreal taxa found in elevations < 1500 m are

Alchornea, Arecastrum, Callophyllum, Cecropia,

Cordia, Croton, Geonoma, Melastomataceae, Myrta-

ceae, Piptadenia, Rubiaceae, Tabebuia, Tapirira,

Virola and Xylopia among others. Small patches of

species-poor Cerrado vegetation can still be found on

sandy soils of the valley. But open savanna-like

vegetation such as Cerrado is present mostly beyond

the Serra da Mantiqueira, in southern State of Minas

Gerais (Radambrasil, 1983).

Presently, representative areas of all these vegeta-

tion types can still be found in the valley in the form

of fragments in various stages of plant succession.

The original vegetation of the peatbogs has been

seriously impacted by agricultural practices but its

overall plant composition can be inferred from few

remnants of undisturbed wetland, located within the

Jacarei municipal area, believed to be transitional

stages to peatbogs. The most important taxa in these

undisturbed bogs are Sphagnum, Lycopodium, Blech-

num, Polypodium sp., together with other swamp

vegetation elements such as Poaceae, Asteraceae,

Cyperaceae, Eriocaulon, Typha, Ludwigia and Polyg-

onum hydropiperoides. In certain portions of these

undisturbed flooded areas, one notices the presence of

Fig. 2. Vegetation map of southeastern and southern Brazil showing potential distribution of different plant formation prior to pre-colonial settling at ca. 500 years BP (modified from

Radambrasil, 1983) and the location (numbered circles) of pollen records discussed in the text.

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Fig. 3. Schematic representation of the vegetation types found along a topographical profile of the Serra da Mantiqueira, the Paraıba do Sul River valley and the Serra do Mar.

Selected taxa illustrate a small portion of the total plant biodiversity of the local vegetation types (after Radambrasil, 1983).

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M.J. Garcia et al. / Review of Palaeobotany and Palynology 131 (2004) 181–199 187

abundant small trees of Cecropia, Melastomataceae

and shrubs of Mimosa and Baccharis.

2.4. Drainage and pollen source

The Paraıba do Sul River drains the western

portion of the Serra dos Monos, Serra dos Patis and

the Serra do Mar, referred to as the Serra do Mar

complex, and the eastern slopes of the Serra da

Mantiqueira in the State of Sao Paulo, between

latitudes 23j30VS and 22j45VS. These mountain

regions, forested prior to the European settlement,

are likely to have left an allochthonous pollen signal

in the peatbog sediments. Incoming pollen from the

valley forests and peatbog vegetation are likely to be

responsible for an autochthonous pollen signature.

In order to verify the pollen contribution from the

Serra da Mantiqueira and Serra do Mar, surface sedi-

ment pollen samples were collected in different modern

vegetation types along an altitudinal gradient ranging

from 550 to 1587 m, shown on Tables 1 and 2. In

general, the majority of pollen taxa present in these

surface samples are represented by percentage values

very similar those obtained by Behling et al. (1997) and

Behling (1997b) in the native Atlantic rainforest of

State of Parana, southern Brazil. However, the arboreal

pollen signature at this altitudinal range in the Serra da

Mantiqueira (Table 1) appears to reflect different suc-

cessional stages of the remaining native vegetation. For

instance, above 1000 m elevation, there is a higher

representation of arboreal pollen, when compared to

sites on the valley. The high altitudinal forests mainly

consist of old successional and some primary forests

that have been preserved in state and municipal parks,

Table 1

Modern pollen rain along an altitudinal transect within the Paraıba do Su

Location Elevation

(m)

Vegetation type

Campos do Jordao, SP 1575 Araucaria forest



Santo Antonio do Pinhal, SP 1087 Transitional Araucaria/

semi-deciduous forest

Jacareı, SP 578 Semi-deciduous forest



Jacareı, SP 550 Preserved peatbog veget

Results are given as a percentage of the pollen sum.

whereas in Jacareı there is a predominance of second-

ary and more open forests. This could account for

higher terrestrial herb pollen percentages at lower

elevations. The modern pollen signal from the pre-

served peatbog vegetation is primarily coming from

aquatic and terrestrial herbs and does not include a

contribution from the highland forests.

In order to determine the long distance transport,

especially of well-known anemophilous pollen, we

present in Table 2 only the results for Araucaria,

Podocarpus and Myrtaceae, three important ane-

mophilous pollen types in the regional vegetation.

The complete pollen spectra of all samples will be

published elsewhere. According to Behling et al.

(1997), Araucaria angustifolia pollen from the high-

lands of Parana was deposited in pollen traps ca. 100

km away, at the coast of Santa Catarina. In the same

study, the authors found for Podocarpus a maximum

value of 0.6%, in a plot containing four trees/hectare,

which yielded only 0.8% of the pollen sum. As shown

in Table 2, the modern low percentage and concen-

tration values of Araucaria pollen, in the low eleva-

tion vegetation of the Paraıba do Sul Valley, can be

explained by wind dispersal from regional planted

ornamental trees. The significant value of 2.2% for

Podocarpus at 570 m elevation suggests that this

taxon can be found in the semi-deciduous forests in

the valley, whereas values < 1%, according to Behling

et al. (1997), must indicate long distance transport.

In summary, the results, presented in Tables 1 and 2,

indicate that there is no significant contribution from

the highlands to the pollen spectra found in modern

surface sediments of the Paraıba do Sul River valley.

The low percentage and concentration values of wind-

l River Valley, Sao Paulo

% Tree

pollen

% Shrub

pollen

% Terrestrial

herbs

% Aquatic

herbs

94.3 4.8

90.4 9

79.6 14.8

85.9 11.5

48.6 44.1 0.5

57.0 1.1 35.3

48.9 0.3 40.6

ation 12.2 0.5 34.8 50.7

Table 2

Number of grains counted, percentage and concentration values in the modern pollen rain for Araucaria, Podocarpus and Myrtaceae along an

altitudinal gradient

Elevation Araucaria angustifolia Podocarpus Myrtaceae

(m)Grains

counted

% Conc.

(103)

Grains

counted

% Conc.

(103)

Grains

counted

% Conc.

(103)

1575 44 10.5 49 339 81.1 378 4 1 4

1563 25 4.9 15 360 70.3 219 34 6.6 20

1568 36 12.7 40 164 57.7 183 11 3.9 12

1087 8 1.1 4.7 3 0.4 1.8 45 6.2 26

578 0 0 0 1 0.3 0.4 26 6.5 11

570 2 0.4 1.1 10 2.2 5 51 11.3 28

550 0 0 0 1 0.3 0.3 0 0 0

550 2 0.3 744 1 0.3 0.7 0 0 0


dispersed taxa such as Araucaria, Podocarpus and

Myrtaceae in the modern surface sediments of the

valley suggest a minor contribution from the highland

forests. Therefore, we expect the pollen spectra found

in the peatbog sediments to be representative of the

vegetation that covered the Paraıba do Sul River valley

before the period of human impact on the vegetation.

3. Materials and methods

3.1. Sediment coring, 14C dating and pollen analysis

A 643-cm-long peat core was drilled at the south-

ern edge of the Jacareı peatbog, at 23j15VS–45j55VW(Fig. 1), using a 4-cm diameter piston corer designed

by the Geological Survey of Finland. Compacted clay

sediments found below the 643-cm depth prevented

further drilling. The undisturbed sediments were re-

trieved in sections of 90 cm each. Each section was

contained in PVC tubing, sealed and transported to the

laboratory. The sediments were described and sam-

pled for pollen analysis and radiocarbon dating.

Because of possible reworking of the top sediments

by modern agricultural practices, the top 50 cm of the

sequence was discarded and sampling was conducted

along the core at 20–30-cm intervals.

Three bulk samples collected at 106, 432 and 642

cm depths were dried at 40 jC to prevent fungal

growth and sent to Beta Analytic (USA) where they

were cleaned from roots and other plant materials and

radiocarbon dated.

The palynological methodology used is described

in Faegri and Iversen (1989). For silicate removal, the

chemical pre-treatment consisted of concentrated HF,

for 48 h at room temperature. Humic acids were

removed by 10% KOH followed by acetolysis (nine

parts of acetic anhydride and one part sulfuric acid).

Palynomorphs found in the residues were stained with

an alcoholic safranin solution. Permanent slides were

made by mounting the residues on the glass coverslips

with cellosize gum and glasslides containing Entellan

medium. An average of 353 pollen grains were

counted per level and pollen sums varied from 178

to a maximum of 1475 grains per level in poor and

rich samples. Pollen sum included trees, shrubs and

aquatic taxa but excluded spores. The objective of

extensive counting in the latter samples was to obtain

as much information as possible on the former re-

gional vegetation.

The palynological profiles were graphed by using

the Tilia/Tiliagraph software and pollen zones were

established after running cluster analysis for strati-

graphically constrained samples using the CONISS

software (Grimm, 1987).

Due to the high frequency of Lycopodium and

myrtaceous taxa in the original local vegetation, the

use of either Lycopodium and Eucalyptus exotic

markers as means of assessing pollen concentrations

(Stockmarr, 1971) was avoided.

4. Results and interpretation

4.1. Stratigraphy, 14C dating and sedimentation rates

Lithology of the sediments, shown in Fig. 4,

consisted of peaty soil (0–50 cm, not sampled),

Fig. 4. Arboreal pollen percentage diagram, with pollen zones and lithology of the Jacareı peat core.

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ligneous peat (50–460 cm), clayey peat (460–579

cm) and clay (579–643 cm).

Radiocarbon dates indicate that sedimentation at

the Jacareı section started at the onset of the Holocene

as shown by the 14C chronology shown on Table 3.

The record encompasses the interval from ca. 1950 to

9700 years BP. Conventional and interpolated dates

indicate that peat sedimentation rates varied from 1.3

mm/year, between 9720 and ca. 8200 years BP, to

0.82 mm/year, between 8200 to ca. 5400 years BP.

4.2. Palynological results

Well preserved pollen and spore grains prevailed in

most samples, with the exception of samples at 460

and 490 cm, and the basal samples at 519, 611 and

643 cm depths.

The percentage pollen diagrams are given in Figs.

4 and 5. The profiles of pteridophytic and algal spores

are given in Fig. 6. The complete list of all taxa found

in the analysis as well as their occurrence in the

respective neotropical vegetation types is given in

Table 4.

The results of the Coniss cluster analysis, shown

on Fig. 4, suggest five pollen zones, which are

displayed in all pollen diagrams. Because the material

was used up for geochemical analyses, further 14C

dating was not possible and ages for some pollen

zones were based on linear interpolation.

4.2.1. Zone JA1 (9720–ca. 8850 years BP)

This zone is subdivided into subzones JA1a and

JA1b, which correspond to clay and the peat sedi-

ments, respectively. It is characterized by >60% of the

aquatic taxon Polygonum aff. hydropiperoides. Her-

baceous taxa found are mainly Poaceae and Astera-

ceae. Gleichenia spores account for almost 100% of

all spores. Other pteridophytic components of this

zone are Selaginella and Polypodium. Arboreal pollen

Table 3

Radiocarbon dates of the Jacareı peat sediments

Laboratory no. Sediment type Depth

(cm)

Age

(14C years BP)

Beta-68157 ligneous peat 106–107 4130F 70

Beta-68156 ligneous peat 432–433 8100F 90

Beta-68155 clay 642–643 9720F 100

become important in the final stage of subzone JA1b,

when Pisonia accounts for over >60%.

In subzone JA1b, there is a reduction in Gleiche-

nia, Polygonum and other herbs synchronous with an

increase of pollen grains belonging to Pisonia, an

arboreal taxon. Also found in this subzone, in low

percentages, are Erythroxylum, Ilex, Sebastiana and

Virola.

4.2.2. Zone JA2 (ca. 8850–ca. 8240 years BP)

This zone is markedly different from the previous

zone by the conspicuously continuous decline of

Gleichenia spores, which are replaced by Selaginella,

Polypodium, Cyathea, Asplenium and other ferns. The

disappearance of Pisonia from the pollen record is

off-set by the appearance of various arboreal taxa such

as Alchornea, Erythroxylum, Ilex, Machaerium, Coc-

coloba, Schizolobium, Myrsine, Sebastiana, Sorocea,

Symplocos and other Atlantic rainforest taxa. This

zone also shows a progressive increase in herb pollen,

especially Poaceae and Cyperaceae and marks a

substantial increase in freshwater algal spores belong-

ing to Mougeotia, Spirogyra and Zygnema.

4.2.3. Zone JA3 (ca. 8240–ca. 5400 years BP)

This zone is characterized by a progressive in-

crease in Poaceae grains, initially contributing to 30%

and increasing to 60% at the end of zone. This

increase in grasses is synchronous with the increase

in Lycopodium and various algal spores. Sphagnum

attains, initially, maximum percentage values but

disappears before the beginning of the following

pollen zone. The arboreal pollen found in this zone

is represented primarily by Coccoloba, a common tree

in the coastal Brazilian forests (Joly, 1976; Rizzini,

1997), and minor elements including Araucaria, Cel-

tis, Erythroxylum, Machaerium, Pisonia, Myrsine,

Sebastiana (aff. brasiliensis), Schizolobium and

others.

4.2.4. Zone JA4 (ca. 5400–3500 years BP)

This zone is characterized mainly by shifts in the

composition and abundance of spores such as Lyco-

podium, which declines progressively, contrasting

with the simultaneous increase of Asplenium, Cya-

thea, Gleichenia and Sphagnum spores. The highest

values of Poaceae reach 60% in this zone. Arboreal

polle percentages are similar to those observed in the

Fig. 5. Pollen percentage diagram of the Jacareı showing the downcore changes in the contribution of terrestrial and aquatic herbs.

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Fig. 6. Spore percentage diagram of the Jacareı peatbog showing bryophytes, pteridophytes and algae.

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Table 4

List of identified pollen and spore taxa and their grouping according to habitat

Abutilon—B, TF, CE Acanthaceae—H, some A, B, TF, CE, W

Alchornea—A, TF Althernanthera—H, CE, W

Alnus—A (long distance dispersal) -exotic Amaranthaceae/Chenopodiaceae—H, TF, W, CE

Araucaria—A, TF (montane) Ambrosia—H, TF, CE, W

Apocynaceae—A, B (here), TF, CE, W Apiaceae—H, TF, CE, W

Banara—A, TF Apocynaceae—A, B, H (here), TF, CE, W

Bauhinia—A, TF, CE, W Asphodelus—H, TF

Bignoniaceae—A (aff. Tabebuia), B, L, TF, CE, W Asteraceae, A, mainly herbs, TF, CE, W

Cariniana—A, TF Brassicaceae—H, TF, CE, W

Cassia—A, TF, CE, W Bredemeyera—H, L, TF, CE, W

Celtis—A, TF Caryophyllaceae—H, TF, CE, W

Coccoloba—A, TF (montane in Sao Paulo) Commelinaceae—H, TF, CE, W

Clusia—A, TF, W Drymaria—H, TF

Croton—A, TF, CE, W Gentianaceae—H, TF, W

Daphnopsis—A, TF Gomphrena—H, TF, CE, W

Drimys—A, TF (montane) Iridaceae—H, TF, CE, W

Ericaceae aff. Gaylussacia—B, TF, (montane in Sao Paulo) Justicia—H, TF, CE, W

Ericaceae aff. Leucothoe—B, TF (mainly montane) Lamiaceae—H, TF, CE, W

Fabaceae—A, TF, CE, W Liliaceae—H, TF, CE, W

Hedyosmum—A, TF (mainly montane in Sao Paulo) Paepalanthus—H, TF, CE, W

Ilex—A, TF, CE, W (mainly high elevation) Perezia—H, TF

Machaerium—A, TF, CE, W Pfaffia—H, TF, CE, W

Malpighiaceae—A (here), B, H, TF, CE, W Poaceae—H, TF, CE, W

Malpighiaceae—L (in diagram as Peixotoa) Primulaceae—H, TF, CE, W

Melastomataceae A, B, rarely H, TC, CE, W Pyrostegia—H, TF, CE

Mimosa A, B, TF, CE, W Saxifragaceae—H, TF, CE, W

Myrsine—A, TF (montane) Vernonia—H, TF, CE, W

Myrtaceae A, B, TF, CE, W (mainly high

elevation in southeastern Brazil)

Convolvulaceae—H, L (here), TF, CE, W

Ouratea—A, TF, CE Passiflora—L, TF, CE, W

Palmae (in diagram as Arecaceae) A, TF, CE, W

Petiveria A, TF Borreria—Q (here), terrestrial herb

in general but some aquatic

Pisonia A, TF, CE, W (montane Atlantic Forest in Sao Paulo) Drosera—Q

Podocarpus A, TF (mainly montane, although scattered

trees also found at elevations < 1000 m in Sao Paulo

Juncaceae—Q

Pseudobombax—A, TF Lentibulariaceae—Q

Rauwolfia A, TF Ludwigia—Q

Schizolobium A, TF Myriophyllum—Q

Sebastiana brasiliensis A, TF, CE Polygonum hydropiperoides—Q

Sessea A, TF

Sorocea A, TF Anemia—S

Symplocos A, TF ( mainly montane in Sao Paulo) Asplenium—S

Tiliaceae A, TF, CE, W Blechnum type—S

Ulmaceae A. TF Ctenitis—S

Urticaceae/Moraceae A, B, some herbs, TF Cyathea—S

Virola A, TF Dennstaedtiaceae—S

Histiopteris—S

Lycopodium clavatum—S

Meesea—S

Phaeoceros—S

Polypodium type—S

Pteridaceae—S

(continued on next page)


Table 4 (continued)

Virola A, TF Riccia—S

Schizaeaceae—S

Selaginella—S

Sphagnum—S

Vittaria—S

Cosmarium, Debarya, Micrasterias, Mougeotia, Spirogyra,

Trachiscia, Zygospore/Desmidiaceae, Zygnema—G

A=arboreal, B = shrub, H = terrestrial herb, L= liana, Q= aquatic herb, S = pteridophyte, G = algae, based on Croat (1978), TF = tropical forest,

CE = savanna, W= found in various vegetation types, based on De Oliveira (1992), De Oliveira et al. (1999), Flenley (1979), Gentry (1993),

Joly (1976), Marchant et al. (2002), Schultz (1985), Van der Hammen (1979).


previous zone, although there is a marked change in

floristic composition of the vegetation with the de-

crease in Coccoloba and the appearance of Cassia,

Gaylussacia, Leucothoe and Solanaceae.

4.2.5. Zone JA5 (ca. 3500–1950 years BP)

This zone represents a departure from all previous

zones. It is marked by substantial increase in forest

taxa, represented mainly by Bauhinia, Ilex and

Alchornea. Rare pollen taxa found in this zone are

Araucaria, Drimys, Clusia, Daphnopsis, Ericaceae

(aff. Gaylussacia and aff. Leucothoe), Podocarpus

and Myrsine. There is a general decrease in peat

forming mosses such as Lycopodium concomitant

with an increase in Gleichenia. Sphagnum is repre-

sented by its highest percentage value in the overall

record before it declines rapidly towards the end of the

zone. Together with these trends, the algae decrease

sharply.

5. Interpretation of the pollen zones

At the onset of the Jacareı record at ca. 9700 years

BP (zones JA1a and JA1b), the Jacareı site was

dominated mostly by Gleichenia, an invasive fern

on humid soils in the more temperate tropics (Joly,

1976; Bold et al., 1980). Also present during this

phase of bog formation is Selaginella, a genus of

tropical club mosses that requires high moisture levels

for growth (Bold et al., 1980). Therefore, the local

climate, at the onset of sedimentation, was probably

cooler and moister than present. Because Gleichenia

is a taxon typically found in montane regions with

cool climates (Joly, 1976; Tryon and Tryon, 1982;

Stannard, 1995) its replacement in zone JA2 by

Polypodium, Pteridaceae, Selaginella and Asplenium

suggests warm and moist climates at ca. 8000 years

BP. It is likely that the decline of Pisonia is also a

consequence of warmer climates since this genus,

although found in various vegetation types in Brazil,

is very common in subtropical forests of southern

Brazil (Schultz, 1985).

At the beginning of zone JA3, starting at ca. 8200

years BP, Araucaria pollen appears for the first time

in the Jacarei peatbog pollen diagram with low

percentages, suggesting the presence of scattered trees

of this taxon in the valley. This conclusion is sup-

ported by the modern surface pollen spectra from

different vegetation types along an altitudinal gradient

in the valley (Table 2) showing that values greater

than 0.5% of Araucaria are unlikely to be attributed to

long distance pollen transport from high-altitude for-

ests of the Serra da Mantiqueira. The pollen spectra of

zone JA3, following the occurrence of Araucaria

pollen, suggest a decline in moisture levels as rates

of peat accumulation drop from 1.30 (zones JA1 and

JA2) to 0.82 mm/year (zones JA3 and JA4). This

decrease in moisture levels after 8100 years BP is

supported by the increase in herb pollen, which

prevails in the pollen diagram until the end of zone

JA4 at ca. 4100 years BP.

The pollen spectra of zone JA5 (ca. 3500–1950

years BP) indicate the return of humid conditions as

gallery forests expanded. This phase also suggests

climatic conditions favoring the appearance of cold-

adapted montane taxa (Ilex, Daphnopsis, Podocarpus,

Gaylussacia, Leucothoe) in the surrounding vegeta-

tion. The decrease in algae is possibly a consequence

of the increase in forest cover at the expense of the

peatbog area. These humid and somewhat cool con-

ditions that continue until ca. 1950 years BP correlates


well with the increase in Atlantic forest elements

observed by Scheel-Ybert (2000) from ca. 2300 to

2000 years BP on the coast of Rio de Janeiro. No

vegetational and climatic information can be obtained

for the last 2000 years in the Jacareı site due to the

sampling restriction caused by possible reworking of

sediments by modern agricultural practices.

6. The environmental history of the Jacareı peat

deposits in relation to Holocene climatic history of

southeastern Brazil

The Jacareı Holocene climatic history starts at

9700 years BP under a humid climate which was

colder than present. A similar climatic trend has been

observed at the onset of the Holocene at Serra Negra

(De Oliveira, 1992), Salitre (Ledru, 1993; Ledru et al.,

1996), in the Pantanal wetlands of southern Mato

Grosso (Bezerra, 1999) and in the Caatinga region

of northeastern Brazil (De Oliveira et al., 1999).

However, early Holocene dry climates have also been

reported in various locations of southern and south-

eastern Brazil by Behling (2002) at Lake Pires (Beh-

ling, 1995), Lake Silvana (Rodrigues-Filho et al.,

2002) and at Lagoa dos Olhos, in the Cerrado region

of central Minas Gerais (De Oliveira, 1992).

In the Serra da Mantiqueira Highlands (Morro de

Itapeva) of Campos do Jordao, Behling (1997a)

identified for the early Holocene the formation of a

cloud forest, reflecting a warm and moist climate on

the east facing slopes of the Serra do Mar and a drier

climate on the western highland plateau of the Man-

tiqueira mountain chain. Moisture levels in the Morro

de Itapeva did not reach modern values until the last

4000 years.

It is possible that the early Holocene humid con-

ditions reconstructed at Jacareı could be explained by

the differential moisture gradient proposed by Behling

(1997a) for the Paraıba do Sul Valley. Thus, greater

moisture along the eastern slopes of the coastal

mountain range would result in a higher water-table

in the valley.

Cold and humid climates lasted in Jacareı until ca.

8100 years BP when the dominant peatbog taxon

Gleichenia is replaced by Lycopodium, Polypodium

and Sphagnum. This transition is marked by a strati-

graphic change from clayey to ligneous peat sedi-

ments. Similar and synchronous cooler conditions,

although under dry climate, were observed at the

onset of the Holocene at the Mantiqueira highlands

by Behling (1997a).

In the Sao Paulo basin, this cooler phase is possibly

represented by the increase in Ilex, Myrtacae and

Hedyosmum pollen at 8120 years BP as reported by

Takiya (1997). In the Salitre (Ledru, 1993; Ledru et

al., 1996) pollen record Araucaria persists on the

landscape until 8000 years BP. In the sample dated

at 8100 years BP at Jacareı, Araucaria is represented

by 1.9% of the pollen sum, which is close to twice the

modern value for the Santo Antonio do Pinhal (tran-

sitional Araucaria forest) surface samples and signif-

icantly different from 0.3%, which can be accounted

solely by long distance transport (Behling et al.,

1997). These results support the conclusion that ca.

8100 years BP scattered trees of Araucaria and other

montane forest elements were actually growing in the

valley, thus suggesting a slightly cooler climate at that

time. It is possible that the moisture necessary for

Araucaria to grow can be explained by a more

shallow water-table fed by high precipitation on the

eastern slopes of the Serra do Mar. This hypothesis is

supported by the fact that this humid phase in Jacareı

(Zones JA1 and JA2) is synchronous with the one

observed in the Atlantic rainforest region of Lago

Pires (17jS) in eastern Minas Gerais reported by

Behling (1995).

From 8240 to ca. 5400 years BP dry climates

established in the region which is attested by the

substantial increase in grasses (Poaceae) and the drop

in sedimentation rate from 1.30 (Zones JA1 and JA2)

to 0.82 mm/year (zones JA3 and JA4). These dryer

conditions, which persisted until 3500 years BP, are

also well correlated with those observed by Behling

(1995) at Lago do Pires and on the western slopes of

the Serra de Itapeva (ca. 60 km in straight line from

the studied site) where they persisted until ca. 3000

years BP (Behling, 1997a).

A major climatic change appears in the record of

Jacareı at ca. 3500 years BP as indicated by the abrupt

increase of arboreal pollen grains from montane taxa,

e.g. Ilex, accompanied by low percentages of Arau-

caria, Ericaceae (aff. Gaylussacia and aff. Leuco-

thoe), Hedyosmum, Ilex, Myrtaceae, Podocarpus and

Myrsine. On the coast of the southern State of Sao

Paulo, Bissa (1998) shows a synchronous and similar


vegetational trend with the increase of montane At-

lantic forest elements (Ilex, Hedyosmum and Myrta-

ceae) at 4010 years BP and a substantial increase in

Ilex at 2820 years BP. The return of humid conditions

at the late Holocene appears to be a general climatic

trend in southeastern Brazil as documented at the

Morro de Itapeva at ca. 3000 years BP as well as in

the central area of the Brazilian Cerrados (Barberi,

2001; Barberi-Ribeiro, 1994; Ferraz-Vicentini, 1993;

Salgado-Labouriau, 1997; Salgado-Labouriau et al.,

1997, 1998; Parizzi et al., 1998).

7. The origins of the Jacareı peat deposits

Thick peat deposits are usually found in temperate,

tropical and subtropical regions and are the result of

humid conditions in riverine valleys or depressions

(Damman, 1978; Garcia, 1994; McCabe, 1991). How-

ever, the global distribution of Sphagnum peats indi-

cates that its occurrence is restricted to cool and warm

temperate regions (Lottes and Ziegler, 1994; McCabe,

1991). These authors point out that peat formation is

contingent upon high precipitation levels, continuous-

ly distributed over the year, in association with

lowered temperatures and that peat accumulated in

tropical areas when the climate was cooler than

present. This latter conclusion was also reached by

Chateauneuf et al. (1991) who analyzed the properties

of African peats and verified that deposits rich in

organic matter accumulated in Africa in the late

Pleistocene or in the early Holocene under climates

colder than present. Additionally, more evidence for a

possible correlation between cool climates and peat

accumulation in tropical areas comes from the work of

Pezeril et al. (1986), who defined the last glacial

maximum, from 20,000 to 12,000 years BP as the

period with the greatest peat accumulation in Senegal.

Greatest peat accumulation in Jacareı is attained in

zone JA1 when climate was probably wetter and

cooler than present, with a mean sedimentation rate

of 1.3 mm/year. During this time, the prevalent peat

forming taxon was Gleichenia, which is presently

found at higher elevations of the Serra da Mantiqueira

and other montane areas of southeastern and north-

eastern Brazil (Joly, 1976; Tryon and Tryon, 1982;

Stannard, 1995). According to Tryon and Tryon

(1982), Gleichenia is adapted to cool and moist

climates, which explains its frequent distribution in

high southern latitudes as well as in montane and

alpine-like regions of the neotropics. Although this

observation supports partially the idea that peat accu-

mulation in tropical regions is fostered by colder

climates, a significant peat accumulation in Jacareı

was brought about by Sphagnum and Lycopodium,

from ca. 5400 to 3500 years BP, but under a climate,

possibly warm and relatively drier than present.

8. Paleoclimatic implications

The palynological analyses of the Jacareı peat core

permit some inferences on paleoclimatic changes such

as the behavior of the polar cold fronts, the ICTZ and

possibly of ENSO-like events during the Holocene of

the State of Sao Paulo.

The humid and cold early phase that lasted from

9720 to ca. 8240 years BP, suggested by the increase

of Gleichenia and the presence of Araucaria forests

taxa in the record, is in accordance with similar

climatic signals in the Sao Paulo Basin (Takiya,

1997) as well as in Salitre (MG) (Ledru, 1993; Ledru

et al., 1996). This humid and cold phase, attributed to

a northward displacement of the polar fronts into a

large area of southeastern Brazil during the early

Holocene allowed a faster accumulation of peat in

the Paraıba do Sul River valley.

Lower precipitation levels inferred for the period

between 5400 and 3500 years BP appear to be well

correlated with drier climates in theMantiqueira moun-

tain region and in the Lago do Pires and with increased

seasonality during the mid Holocene in the coast of Rio

de Janeiro (Scheel-Ybert, 2000). This drier phase has

been interpreted as a consequence of stronger influence

of the dry tropical continental air masses (Behling,

1997a), which can be tentatively explained by a sup-

posedly intensified Atlantic Anticyclone.

A significant climatic change is suggested by the

pollen spectra of pollen zone JA5 (ca. 3500–1950

years BP) representing the return of cooler and moist

climates in the valley, which is synchronous with high

lake levels in other areas of southeastern Brazil, such as

Lagoa Santa (Parizzi et al., 1998), Lagoa Olhos

D’Agua (De Oliveira, 1992) and Iguape (Bissa,

1998). Paleohydrological studies conducted in Lake

Infernao, in Sao Paulo, also suggest high precipitation


levels from 3500 to 3000 years BP (Lobo et al., 2001).

According to De Oliveira et al. (1999), this humid

phase between 4000 years BP and the present in

southern and southeastern Brazil is contrasted by

prevailing and synchronous semi-arid climates in

northeastern Brazil. These authors argue that there

might be a possible link between reported ENSO-like

variability in some areas of southeastern Brazil during

the late Holocene (Martin and Suguio, 1992; Martin et

al., 1993) and reported dry climates in the northeast.

During these climatic phases, moister and cooler con-

ditions prevailed in southern and some areas of south-

eastern Brazil possibly due to the blockage of polar

fronts, while in northeastern Brazil dry climates per-

sisted. According to De Oliveira et al. (1999), another

hypothesis for this apparent regional climatic asymme-

try after 4000 years BP between these two geographical

areas refers to a possible northward displacement of the

ICTZ, which became stationary over central and south-

ern North America. This secondary hypothesis is

supported by reported high humidity levels after 4000

years BP in Panama (Bush et al., 1992), in Haiti (Curtis

andHodell, 1993) and in theYucatan Peninsula (Hodell

et al., 1991, 1995). Further investigation and additional

records from coastal State of Sao Paulo are required for

the testing of these hypotheses. Nonetheless, the

Jacareı record brings some important contribution to

the understanding of the climatic changes in the im-

portant tropical forest belt of the Brazilian coast.

Acknowledgements

The authors thank Dr. Murilo Rodolfo de Lima (In

Memoriam) for support during the first stage of this

work. This research was supported in part by the

Universidade Guarulhos, CNPq (The National Re-

search Council of Brazil), CESP (The Energy

Company of the State of Sao Paulo) and Project

FAPESP nr. 2000/03960-5 (Historia da Exumac�ao da

Plataforma Sul-americana, a exemplo da regiao

sudeste brasileira: Termocronologia por trac�os de

fissao e sistematica Ar/Ar e Sm/Nd). Thanks are due

to Dr. J.P. Ybert (IRD-France), Dr. Kenitiro Suguio,

Dr. Setembrino Petri, Mary E.C. de Oliveira and

Claudio Riccomini (University of Sao Paulo) for

suggestions and guidance through the development of

this work. We are deeply thankful to Dr. Vera

Markgraf and Dr. Henry Hooghiemstra for their

careful revision and suggestions, which improved

the text significantly. Thanks are also due to Dr.

Herman Behling (University of Bremen-Germany)

and Dr. Maria Lea Salgado-Labouriau (University of

Brasilia, Brazil) for their suggestions and comments.

References

Barberi, M., 2001. Mudanc�as paleoambientais na regiao dos cerra-

dos do Planalto Central durante o Quaternario Tardio: O estudo

da Lagoa Bonita, DF. PhD thesis. Institute of Geosciences. Uni-

versidade de Sao Paulo, Sao Paulo, SP, Brazil, 210 pp.

Barberi-Ribeiro, M., 1994. Paleovegetac�ao e paleoclima no Quater-

nario Tardio da Vereda de Aguas Emendadas, DF, MS Thesis,

Brasılia, Universidade de Brasılia, 110 pp.

Behling, H., 1995. A high resolution Holocene pollen record from

Lago do Pires, SE Brazil: vegetation, climate and fire history.

Journal of Paleolimnology 14, 253–268.

Behling, H., 1997a. Late Quaternary vegetation, climate and fire

history from the tropical mountain region of Morro de Itapeva,

SE Brazil. Palaeogeography Palaeoclimatology Palaeoecology

129, 407–422.

Behling, H., 1997b. Late Quaternary vegetation, climate and fire

history in the Araucaria forest and campos region from Serra

Campos Gerais (Parana), S. Brazil. Review of Palaeobotany and

Palynology 97, 109–121.

Behling, H., 2002. South and southeast Brazilian grasslands during

Late Quaternary times: a sythesis. Palaeogeography Palaeocli-

matology Palaeoecology 177, 19–27.

Behling, H., Negrelle, R.B., Colinvaux, P.A., 1997. Modern pollen

rain data from the tropical Atlantic rain forest, Reserva Volta

Velha, South Brazil. Review of Palaeobotany and Palynology

87, 287–299.

Bezerra, M.A.O., 1999. O uso de multi-trac�adores na reconstruc�aodo Holoceno no pantanal Mato-Grossense, Corumba, MS. Cen-

ter for Biological and Health Sciences. Universidade Federal de

Sao Carlos, Sao Carlos, SP, Brazil.

Bissa, W.M., 1998. Reconstituic�ao dos paleoambientes de uma

planıcie holocenica no curso inferior do Rio Ribeira (Muni-

cıpio de Iguape, SP). MS Thesis. Department of Geography.

FFLCH, Universidade of Sao Paulo, Sao Paulo, SP, Brazil,

82 pp.

Bold, H.C., Alexopoulos, C.J., Delevoryas, T., 1980. Morphology

of Plants and Fungi Harper and Row, New York, 819 pp.

Bush, M., Piperno, D.R., Colinvaux, P., De Oliveira, P.E., Krissek,

L., Miller, M.C., Rowe, W.E., 1992. A 14,300-yr paleocological

profile of a lowland lake in Panama. Ecological Monographs 62,

251–275.

Chateauneuf, J.-J., Farjanel, G., Laggoun-Defarge, F., Pezeril, G.,

Bikwemu, G., 1991. Petrological and physico-chemical prop-

erties of some African peats in relation to their suitability for

carbonization. Bulletin Societe Geologique de France 162,

423–435.


Croat, T.C., 1978. Flora of Barro Colorado Island. Stanford Uni-

versity Press, Palo Alto, CA, 943 pp.

Curtis, J., Hodell, D.A., 1993. An isotopic and trace element study

of ostracods from lake Miragoane, Haiti: a 10,500 year record of

paleosalinity and paleotemperature changes in the Caribbean.

Climate change in continental isotopic records. Geophysical

Monograph 78, 135–152.

Damman, A.W.H., 1978. Distribution and movement of elements in

ombrotrophic peat bogs. Oikos 30, 480–495.

De Oliveira, P.E., 1992. A palynological record of Late Quaternary

vegetational and climatic change in southeastern Brazil. PhD

thesis. The Ohio State University, Columbus, Ohio, 238 pp.

De Oliveira, P.E., Barreto, A.M.F., Suguio, K., 1999. Late Pleis-

tocene/Holocene climatic and vegetational history of the Bra-

zilian caatinga: the fossil dunes of the middle Sao Francisco

River. Palaeogeography Palaeoclimatology Palaeoecology 152,

319–337.

Faegri, K., Iversen, J., 1989. Textbook of Pollen Analysis, 4th ed.

Wiley, New York, 295 pp.

Ferraz-Vicentini, K.R., 1993. Analise palinologica de uma vereda em

Cromınia, GO. MS Thesis. Universidade de. Brasılia, 136 pp.

Flenley, J.R., 1979. The Equatorial Rain Forest: A Geological His-

tory. Butterworth, London, 162 pp.

Garcia, M.J., 1994. Palinologia de turfeiras Quaternarias do medio

Vale do Rio Paraıba do Sul, Estado de Sao Paulo. PhD thesis.

Institute of Geosciences. Universidade de Sao Paulo, Sao Paulo,

Brazil, 354 pp.

Gentry, A., 1993. A Field Guide to Families and Genera of Woody

Plants of Northwest South America (Colombia, Ecuador, Peru)

With Supplementary Notes on Herbaceous Taxa. Conservation

International, Washington, DC, 895 pp.

Grimm, E.C., 1987. CONISS: a Fortran 77 program for stratigraph-

ically constrained cluster analysis by the method of the incre-

mental sum of squares. Computers & Geosciences 13, 13–35.

Hodell, D.A., Curtis, J.H., Jones, G.A., Higuera-Gundy, A., Bren-

ner, M., Bindford, M.W., Dorsey, K.T., 1991. Reconstruction of

Caribbean climate change over the past 10,500 years. Nature

352, 790–793.

Hodell, D.A., Curtis, J.H., Brenner, M., 1995. Possible role of

climate in the collapse of Classic Maya civilization. Nature

375, 391–394.

Joly, A.B., 1976. Botanica. Introduc�ao a taxonomia vegetal Comp.

Editora Nacional, Sao Paulo, 777 pp.

Kumoto, E.T., Constanzo Jr., J., Monticeli, J.J. 1985. Minerais

energeticos: carvao, turfa e rochas oleıgenas. CESP-Compan-

hia Energetica de Sao Paulo-Serie Pesquisas e Desenvolvi-

mento, vol. 14. CESP, Sao Paulo, 58 pp.

Ledru, M.P., 1993. Late Quaternary environmental changes in Cen-

tral Brazil. Quaternary Research 39, 90–98.

Ledru, M.P., Braga, P.I.S., Soubies, F., Fournier, M., Martin, L.,

Suguio, K., Turcq, B., 1996. The last 50,000 years in the

Neotropics (Southern Brazil): evolution of vegetation and cli-

mate. Palaeogeography Palaeoclimatology Palaeoecology 123,

239–257.

Lobo, I., Mozeto, A.A., Aravena, R., 2001. Paleohydrological in-

vestigation of Infernao Lake, Moji-Guac�u River watershed, Sao

Paulo, Brazil. Journal of Paleolimnology 26, 119–129.

Lottes, A.L., Ziegler, A.M., 1994. World peat occurrence and the

seasonality of climate and vegetation. Palaeogeography Palae-

oclimatology Palaeoecology 106, 23–37.

Marchant, R., Almeida, L., Behling, H., Berrio, J.C., Bush, M.,

Cleef, A., Duivenvoorden, J., Kappelle, M., De Oliveira, P.,

Oliveira-Filho, A.T., Lozano-Garcia, S., Hooghiemstra, H.,

Ledru, M.P., Ludlow-Wiechers, B., Markgraf, V., Mancini, V.,

Paex, M., Prieto, A., Rangel, O., Salgado-Labouriau, M.L.,

2002. Distribution and ecology of parent taxa of pollen lodged

within the Latin American Pollen Database. Review of Palae-

obotany and Palynology 121, 1–75.

Martin, L., Suguio, K., 1992. Variation of coastal dynamics during

the last 7,000 years recorded in beach-ridge plains associatedwith

river mouths: example from the central Brazilian coast. Palae-

ogeography Palaeoclimatology Palaeoecology 99, 119–160.

Martin, L., Fournier, M., Mourguiart, P., Sifeddine, A., Turcq, B.,

Absy, M.L., Flexor, J.-M., 1993. Southern oscillation signal in

South American paleoclimatic data of the last 7000 years. Qua-

ternary Research 39, 338–346.

McCabe, P.J., 1991. Tectonic controls on coal accumulation. Bul-

letin Societe Geologique de France 162, 227–282.

Mello, C.L., Moura, J.R.S., 1991. Significado paleoambiental da

aloformac�ao Manso: um marco estratigrafico no Holoceno do

medio Vale do Rio Paraıba do Sul. Resumos do 3j. Con-

gresso da Associac�ao Brasileira dos Estudos do Quaternario

(ABEQUA), Belo Horizonte, MG. Publicac�ao Especial, vol. 1.

ABEQUA, Sao Paulo, pp. 49–52.

Mendes, V.A., Costa, A.C., 1981. Potencialidades da turfa no Nor-

deste do Brasil e suas possibilidades de emprego. Actas do X

Simposio de Geologia do Nordeste, Recife, vol. 1, pp. 132–144.

Morellato, L.P.C., Haddad, C.F.B., 2000. The Brazilian Atlantic

Forest. Biotropica 32, 786–792.

Moura, J.R.S., Mello, C.L., 1991. Classificac�ao aloestratigrafica do

Quaternario superior na regiao de Bananal (SP/RJ). Revista

Brasileira de Geociencias 21, 254–263.

Nimer, E., 1989a. Climatologia do Brasil. IBGE, Rio de Janeiro,

421 pp.

Nimer, E., 1989b. A circulac� ao atmosferica e as condic� oes do

tempo como fundamento para a compreensao do clima. Geo-

grafia do Brasil. Regiao Centro-Oeste, vol. 1. IBGE, Rio de

Janeiro, pp. 23–24.

Oliveira-Filho, A.T., Fontes, M.A.L., 2000. Patterns of floristic

differentiation among Atlantic forests of southeastern Brazil

and the influence of climate. Biotropica 32, 793–810.

Parizzi, M.G., Salgado-Labouriau, M.L., Kohler, H.C., 1998. Gen-

esis and environmental history of Lagoa Santa, southeastern

Brazil. Holocene 8, 311–321.

Pezeril, G., Chateauneuf, J.-J., Diop, C.E.W., 1986. La tourbe des

Niayes au Senegal. INQUA/1986. Dakar Symposium ‘‘Change-

ments globaux en Afrique’’. INQUA, Dakar, pp. 385–392.

Radambrasil Project Members, 1983. Levantamento de recursos

naturais. Vol. 32. Folhas SF.23/24. Rio de Janeiro/Vitoria.

Ministerio das Minas e Energia. Rio de Janeiro, RJ, Brazil,

775 pp.

Riccomini, C., 1989. O rift continental do Sudeste do Brasil. PhD

thesis. Institute of Geosciences, Universidade de Sao Paulo, Sao

Paulo, Brazil, 304 pp.


Rizzini, C.T., 1997. Tratado de Fitogeografia do Brasil. Aspectos

Ecologicos, Sociologicos, e Florısticos Ambito Cult. Ed., Rio de

Janeiro, 747 pp.

Rodrigues-Filho, S., Behling, H., Irion, G., Muller, G., 2002. Evi-

dence for lake formation as a response to an inferred Holocene

climatic transition in Brazil. Quaternary Research 57, 131–137.

Salgado-Labouriau, M.L., 1997. Late Quaternary paleoclimate in

the savannas of South America. Journal of Quaternary Science

12, 371–379.

Salgado-Labouriau, L., Casseti, V., Ferraz-Vicentini, K.R., Martin,

L., Soubies, F., Suguio, K., Turcq, B., 1997. Late Quaternary

vegetational and climatic changes in cerrado and palm swamp

from Central Brazil. Palaeogeography Palaeoclimatology Pa-

laeoecology 128, 215–226.

Salgado-Labouriau, M.L., Barberi, M., Ferraz-Vicentini, K.R., Par-

izzi, M.G., 1998. A dry climatic event during the Late Quater-

nary of tropical Brazil. Review of Palaeobotany and Palynology

99, 115–129.

Scheel-Ybert, R., 2000. Vegetation stability in the Southeastern

Brazilian coast area from 5500 to 1400 14C yr BP deduced from

charcoal analysis. Review of Palaeobotany and Palynology 110,

111–138.

Schultz, A., 1985. Introduc�ao a botanica sistematica. Ed Universi-

dade Federal do Rio Grande do Sul, Porto Alegre, 414 pp.

Shimada, H., Carvalho, W.S., 1980. Perspectivas de utilizac�ao da

turfa para fins energeticos no Estado de Sao Paulo. Semi-

nario: Contribuic�ao da Geologia a Busca e Aproveitamento

de Fontes Energeticas (Convencionais Ou Nao). Sociedade

Brasileira Geologia Boletim, vol. 1. Comunicac� oes, Sao

Paulo, pp. 33–64.

Silva, A.F., Filho, H.F.L., 1982. Composic�ao florıstica de um trecho

da mata atlantica da encosta no municıpio de Ubatuba (Sao

Paulo, Brasil). Revista Brasileira de Botanica 5, 43–52.

Stannard, B.L., 1995. The flora of the Pico das Almas, Chapada

Diamantina, Bahia, Brazil. Royal Botanical Gardens, Kew,

853 pp.

Stockmarr, J., 1971. Tablets with spores used in absolute pollen

analysis. Pollen et Spores 13, 615–621.

Suguio, K., Turcq, B., Cervant, M., Pressinoti, M.N., Soubies, F.,

1992. Late Quaternary detritic deposits in Southeastern Brazil:

palaeohydrological interpretations. Palaeoclimatic Changes

and the Carbon Cycle. Serie Geoquımica Ambiental, vol. 1.

IGC-USP, Sao Paulo, pp. 47–53.

Takiya, H., 1997. Estudo da Sedimentac�ao Neogenico-Quaternaria

no municıpio de Sao Paulo: caracterizac�ao dos depositos e suas

implicac�oes na geologia urbana. PhD thesis. Institute of Geo-

sciences, Universidade de. Sao Paulo, Sao Paulo, Brazil.

Tryon, R.M., Tryon, A.F., 1982. Ferns andAllied PlantsWith Special

Reference to Tropical America. Springer, Berlin, 857 pp.

Turcq, B., Suguio, K., Martin, L., Flexor, J.M., 1992. Late Quater-

nary organic deposition in Southeastern Brazil. Paleoclimatic

Changes and the Carbon Cycle. Serie Geoquımica Ambiental,

vol. 1. Instituto Geociencias, Universidade de Sao Paulo, Sao

Paulo, Brazil, pp. 41–46.

Van der Hammen, T., 1979. History of flora, vegetation and climate

in the Colombian Cordillera Oriental during the last five million

years. In: Larsen, K., Holm-Nielsen, L.B. (Eds.), Tropical Bot-

any. Academic Press, London, pp. 25–32.

Villwock, J.A., Denhardt, E.A., Loss, E.L., Hofmeister, T., 1980.

Turfas da provıncia do Rio Grande do Sul-Geologia do Deposito

Aguas Claras. Anais do Congresso Brasileiro de Geologia-Cam-

boriu, 31 (1), 500–512.

Wanderely, M.G.L., Shepherd, G.J., Giulietti, A.M., Melhem, T.S.,

Bittrich, V., Kameyama, C. (Eds.), 2002. Flora fanerogamica do

Estado de Sao Paulo, vol. 2. FAPESP/Ed. Hucitec, Sao Paulo,

391 pp.

A Holocene vegetational and climatic record from the Atlantic rainforest belt of coastal State of...

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