Review of Palaeobotany and Palynology 133 (2005) 235 – 248

www.elsevier.com/locate/revpalbo

Late Quaternary grassland (Campos), gallery forest, fire and

climate dynamics, studied by pollen, charcoal and multivariate
analysis of the São Francisco de Assis core in western Rio Grande
do Sul (southern Brazil)
Hermann Behlinga,*, Valerio DePatta Pillarb, Soraia Girardi Bauermannc
a
Department of Geosciences, University of Bremen, B.O. Box 330440, 28334 Bremen, Germany
b
Departamento de Ecologia, Universidade Federal do Rio Grande do Sul, 91540-000 Porto Alegre, RS, Brazil
c
Laboratório de Palinologia, Universidade Luterana do Brazil (ULBRA), Rua Miguel Tostes 101, 90.420-280 Canoas, RS, Brazil
Received 22 July 2003; accepted 15 October 2004

Abstract

We present a detailed pollen and charcoal record of a 368-cm-long sediment core from the lowland Campos (grassland)
region near the city of São Francisco de Assis in the western Rio Grande do Sul State in southern Brazil. Based on four AMS
radiocarbon dates, the record represents the last about 22,000 cal yr BP. The region was naturally covered by Campos
throughout the recorded glacial and Holocene period under cold and relatively dry and warm and dry condition, respectively.
Initial expansion of gallery forest after 5170 cal yr BP indicates a change to wetter climatic conditions. Maximum extent of
gallery forest after 1550 cal yr BP reflects the wettest recorded period. There is some evidence of plant migration from the
eastern Atlantic coastal lowland reaching the western lowland, first after mid Holocene times by Cecropia and, possibly, species
of Myrsine and Moraceae and later, after about 1000 cal yr BP, by species of Alchornea and Acalypha. Multivariate analysis
revealed that the long-term pollen composition dynamics is a two-phase process, with random, chaotic changes characterizing
some periods when climate conditions were likely more stable, and directional compositional changes (phase transitions) in
periods coincide with major climate and/or anthropogenic changes. Natural fires were rare during full- and lateglacial periods,
but became frequent at the beginning of the Holocene, suggesting the beginning of human occupation of the western lowland at
that time. Highest fire frequency is found during the wet late Holocene period, suggesting an increase of indigenous populations
in the São Francisco de Assis region.
D 2004 Elsevier B.V. All rights reserved.

Keywords: southern Brazil; Campos; gallery forest; palaeoecology; palaeovegetation; palaeofire; palaeoclimate; Amerindians; pollen analysis;
multivariate analysis; last glacial maximum; Holocene

* Corresponding author.
E-mail addresses: Hermann.Behling@uni-bremen.de (H. Behling)8 vpillar@ecologia.ufrgs.br (V.D. Pillar).

0034-6667/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2004.10.004
236 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
1. Introduction
Several palaeoecological studies from the Campos
and Araucaria forest regions of the southern Brazilian
highlands have been carried out in the last decade.
Data from the states of Paraná (Serra Campos Gerais:
Behling, 1997), Santa Catarina (Serra do Rio Rastro,
Morro da Igreja, Serra da Boa Vista: Behling, 1993,
1995) and Rio Grande do Sul (Aparados da Serra:
Roth and Lorscheitter, 1993; São Francisco de Paula:
Behling et al., 2001; Cambará do Sul: Behling et al.,
2004) have proven that extensive areas of Campos
vegetation existed on the highlands through glacial,
early and mid Holocene times. The dominance of
Campos vegetation was attributed to cold and dry
glacial, and warm and dry early Holocene climates. A
dry season lasting probably about 3 months per year
was characteristic for the early and mid Holocene
period (Behling, 1997, 2002). Initial expansion of
Araucaria forests started by migration from the
gallery forests along the rivers about 3210 cal yr BP,
which indicates a turn to somewhat wetter climates. A
marked expansion of Araucaria forests started on the
highlands, replacing Campos vegetation in Santa
Catarina State about 930 cal yr BP ago, and in Paraná
State (Serra Campos Gerais) about 1400 cal yr BP
ago, reflecting a very humid climate without a marked
seasonal dry period. New data from the 42,000 14C
years BP old Cambará do Sul record indicate that the
initial expansion of Araucaria forests started by 4320
cal yr BP, and the marked expansion by 1100 cal yr
BP (Behling et al., 2004).
So far, nothing was known about the vegetation
and climate history of the Campos region in the
lowlands of Rio Grande do Sul State west and south
of the highland plateau. Our main questions are: Is the
Campos formation the natural vegetation of the
southern lowland? Do we have similar climate
changes in the southern lowlands, compared to the
southern highlands? What would be the effect of
climate change on the southern lowland vegetation?
What was the fire history in the lowland? How and
when began the pre-Columbian occupation in the
lowland?
To investigate the late Quaternary vegetation,
climate and fire dynamics, we cored a peat bog in
the São Francisco de Assis region of western Rio
Grande do Sul state. The site is interesting for at
present it is surrounded by grassland characteristic for
the western part of Rio Grande do Sul State, close to
gallery forest and not far from the edge of the forest
on the westernmost escarpments of the southern
Brazilian highlands.
2. Environmental setting
2.1. Geographical setting
The lowlands of Rio Grande do Sul (Fig. 1) lie at
elevations between 40 and 200 m south and west of
the South Brazilian highlands and north of the
crystalline shield. The lowland is a soft rolling
landscape, and includes some higher elevated hills.
Major rivers are the Ibibuı́, draining westward to the
Uruguay River, and the Jacuı́ River, draining eastward
to the Patos Lagoon and the Atlantic Ocean. More
information on the geography of the region can be
found in Rambo (1956) and in satellite images at
http://cdbrasil.cnpm.embrapa.br/rs.
The studied peat bog (29835V12WS, 55813V02WW,
100 m elevation) is situated on the Itajuru farm, about
5 km north from the Ibicuı́ River, 8 km southwest of
the city of São Francisco de Assis, in the western
region of Rio Grande do Sul state (Fig. 1). The site is
about 15 km SW from the westernmost escarpments
of the southern Brazilian plateau, which raises here
only to elevations of ca. 300 m. The Atlantic Ocean is
ca. 480 km to the east. The ca. 50
150 m large peat
bog is found in a shallow basin near the southern
margin of the Inhacundá River valley, ca. 600 m
distant and about 10 m above the river, to which it is
not connected.
2.2. Modern vegetation
General information on the flora and ecology of the
lowland Campos can be found in Boldrini (1997)
among others. The grassland Campos in the study
region develops on sandy substrate is highly diverse
and dominated by species of the genera Andropogon,
Aristida, Paspalum, Axonopus, Elyonurus in the
Poaceae family, of the genera Baccharis, Vernonia
and Senecio in the Asteraceae family, several Myrta-
ceae shrubs of the genera Eugenia, Psidium and
Campomanesia, and the palm Butia paraguayensis
 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248 237

Fig. 1. Map showing the location of Itajuru bog (cross in the image at the right) in the São Francisco de Assis region in Rio Grande do Sul State
in southern Brazil. Areas covered with forest are darker in the image. Escarpments covered with forest are at the right bottom.

(Marchiori, 1995). More detailed descriptions of the
grassland on sandy areas in this region are found in
Trindade (2003).
Small patches or long strips of gallery forests
occur along rivers in-between extensive areas of
grassy vegetation. Patches of forest are also found on
escarpments of low elevated sandstone hills scattered
throughout the region. The species composition of
the forests contains taxa of the seasonal deciduous
forest (Teixeira et al., 1986) occupying the escarp-
ments of the southern Brazilian plateau, whose
westernmost representation is not far from the study
site. The composition of the forests is characterized
by species of Myrtaceae (Eugenia, Psidium),
Euphorbiaceae (Sebastiana, Alchornea, Sapium),
Mimosaceae (Acacia, Enterolobium, Parapiptade-
nia), Tabebuia (Bignoniaceae), Schinus and Lithraea
(Anacardiaceae), Celtis (Ulmaceae), Scutia (Rham-
naceae), Myrsine (Myrsinaceae), Erythrina (Faba-
ceae), Vitex (Verbenaceae) and Syagrus (Arecaceae).
The flora contains elements of the forests of the
Atlantic coast, the Paraná River valley and the Chaco
(Waechter, 2002).
The peat bog today is covered by grasses and
sedges and is used for cattle grazing. It has been partly
drained by shallow channels.
2.3. Modern climate
Southern Brazil is characterized by a warm
temperate and humid climate; precipitation is well
distributed throughout the year along a gradient
from south to north, with maximum precipitation on
the eastern highland plateau (Nimer, 1989; Ratis-
bona, 1976). In the study region, mean temperatures
range from 14 8C in July to 25 8C in January
(INPE/CPTEC, http://www.cptec.inpe.br/clima/monit/
monitor_brasil.shtml). Precipitation varies from year
to year ranging from 1200 to 2000 mm (Suertegaray et
al., 2001), and in some summers, water deficit may
occur (Buriol et al., 1979).
2.4. Human occupation
Perhaps the earliest evidence of human occupa-
tion in the western lowlands of the Rio Grande do
238 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
Sul State is dated about 15,400 cal yr BP at a site
in the Ibicuı́ River valley where lithic artifacts were
found with fossils of extinct megafauna; other early
evidence elsewhere with different lithic artifacts is
dated at 13,470 and 9550 cal yr BP (Kern, 1997a).
Horticulturists, who used forestland, arrived in the
Uruguay River around 2000 years ago (Schmitz,
1997). In the 17th Century, domestic cattle and
horses were introduced in the Jesuitical missions
east from the Uruguay River. After the abandon-
ment of the first missions, cattle spread naturally in
the Campos region until the establishment of farms
by Portuguese settlers after mid 18th Century
(Porto, 1954).
3. Methods
The peat deposit was cored in its deepest part using
a Russian corer. From bedrock, the total length of the
core was 380 cm. Sections of 50 cm length were
extruded on-site, wrapped in plastic and aluminum
foils and stored under cool (ca. +4 8C) and dark
conditions after return from the field and before
sampling. For radiocarbon dating, 1-cm slices of bulk
subsamples were taken. Organic matter was dated
through AMS at the laboratory of the University
Nqrnberg-Erlangen (Germany).
For pollen analysis, subsamples of 0.5 cm3 volume
were taken at 4- and 8-cm intervals along the 380-cm-
long core. All subsamples were processed with stand-
ard pollen analytical methods, using hydrofluoric acid
(HF) and acetolysis (Faegri and Iversen, 1989). To
determine the pollen concentration (grains/cm3) and
pollen accumulation rate (grains/cm2/year), one tablet
of exotic Lycopodium clavatum spores was added to
each subsample. Pollen and spores were in general
well preserved. Only the subsamples below 368 cm
core depth and subsamples between 220 and 195 cm
were poorly preserved and were not counted. A
minimum of 300 pollen grains was counted for each
subsample. The pollen sum includes herbs, shrubs and
trees, and excludes aquatic taxa, fern, moss and fungal
spores and the alga Botryococcus. Pollen identification
relied on the first author’s own reference collection
(containing about 2000 Brazilian species) and pollen
morphological descriptions in Behling (1993). Pollen
and spore data are presented in pollen diagrams as
percentages of the total pollen sum. Carbonized
particles (5–200 Am) were counted on pollen slides
to calculate concentration (particles/cm3) and accumu-
lation rate (particles/cm2/year).
The software TILIA, TILIAGRAPH and CON-
ISS were used for illustration of the pollen and
spore data, calculations and cluster analysis, respec-
tively (Grimm, 1987). The pollen diagrams include
individual records of the most abundant pollen and
spore taxa (Fig. 2), and records of the groups:
Campos, gallery forest, aquatics, and ferns (exclud-
ing Isoetes), mosses, Botryococcus, and concentra-
tion and accumulation rates of pollen and charcoal
particles, and a cluster analysis dendrogram (Fig. 3).
The zonation of the pollen record is based on
changes in the pollen assemblages and on the
cluster analysis.
Ordination by principal coordinates analysis
(PCoA) was used to reveal a synthetic view of
compositional transitions in the palaeocommunities
(Orlóci, 2000; Orlóci et al., 2002a), similarly as
done in Behling et al. (2004). Multivariate methods,
including ordination, are reviewed by Legendre and
Legendre (1998) and Podani (2000), among others.
Ordination methods allow the projection into a
lower number of dimensions, usually two, of
objects described in a multidimensional abstract
space defined by the variables. In this case, the
objects were subsamples and the variables pollen
taxa. PCoA was based on pairwise Euclidean
distances between subsamples. Distances were
computed from square root transformed pollen
percentages in order to reduce the excessive weight
of dominant pollen taxa. Indeterminate and spore-
bearing taxa were not included in the ordination. In
one analysis the complete trajectory of 61 records
for 83 taxa was examined and interpreted in terms
of past vegetation dynamics. In other ordination
analyses, segments of the vegetation trajectory were
examined in more detail. Rates of composition
change between time steps were computed by
dividing the Euclidean distance of adjacent sub-
samples by the time interval in calibrated years BP;
the analysis used the same distances, computed
from square root transformed pollen percentages,
that were used as input for the ordinations (see
references above). The analyses were performed by
SYNCSA software (Pillar, 2001).
 D

50

400
300
250
200
150
100

350
ep
th
Po (c
m
ac )

20
ea
e

40
60
80
C
yp
er

20
ac
ea
e

40
As
te
Ba rac
cc ea
Er ha e s
yn ris ub
Pl giu -typ f. A
an m e s
Bo tag -typ te
rio
rr o e id
Al eria aus ea
te tra e
rn
C a lis
ar nt -ty
y h pe
Sp op era
e hy
Am rma llac
a co ea
G ran ce e
om th
Se ph ace
n re a
Ap eci na- e/C
ia o-t Pf he
Bo cea ype affia nop
rre e -ty od
Er ria I pe ia
Herbs

io la ce
Eu u ifoc a t ae
ph lac lia
Fa or ea -ty
ba bia e pe
Fa ca -ty
ba e I pe
Iri ce
da ae
Xy cea II
ri e
Ze s
a
M ma
el ys
a
M sto
yr m
M tace ata
yr a ce
s ae
Ac ine e
al
Al yph
ch a
M orn
or ea
C ace
el a
t e
Ar is /Ur
ec tic
La ce a ac
ea
th ae
Po are e
do a/S
M ca ch
yr rp in
i
H oph us us-
yd y
l ty
pe
U roc lum
tri ot
c y
M ula le
on ri
a
M ole
Gallery Forest

on te
M le psi o
on te la
t
Se ole psi e <
la te lat 50
g
Is in er e > um v
oe el ru 5
l 0
Ph tes a e cat um
xc e
ae u <
20

oc rre 50
er ns um
ou
s
40

la
Sp ev
ha is
gn
um
Zone

SFA-I
SFA-II
SFA-IV

SFA-III

Fig. 2. Pollen percentage diagram of the frequent and most important taxa of the São Francisco de Assis core (100 m altitude), grouped into Campos, gallery forest, aquatics and ferns.

239 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
 240
H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
Fig. 3. Summary pollen diagram of the São Francisco de Assis core, showing the AMS radiocarbon dates, stratigraphy, ecological groups, pollen sum numbers, pollen and charcoal
concentration and accumulation records, pollen zones and the cluster analysis dendrogram.
 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248 241

4. Results Table 2
Pollen zones from core São Francisco de Assis, showing core depth,
the calculated radiocarbon ages and the number of pollen
4.1. Stratigraphy subsamples of each pollen zone
Zone Depth Age range Age range No. of
The core is underlain by compact clayish sandy. (cm) (14C yr BP) (cal yr BP) subsamples
From the base from 380 to 370 cm, sandy gray clay
SFA-I 368–266 20,140 –10,490 22,150–12,360 26
is found (Fig. 3). Between 370 and 265 cm, brown- SFA-II 266–178 10,490 – 4620 12,360–5170 12
gray clay with organic matter accumulated. From SFA-III 178–106 4620 –1650 5170–1550 9
265 to 175 cm compact, completely decomposed SFA-IV 106–0 1650 to 51 1550 to 51 14
black organic matter accumulated. From 175 to 110
cm, the sediment is composed of dark brown to
black, very compact, completely decomposed by the SYNCSA software, the ages were not
organic material, containing rootlets. Between 110 rounded up.
cm and the grassy surface, the deposit is brown,
weakly decomposed and unconsolidated peat, with 4.3. Description of the São Francisco de Assis pollen
many plant roots. diagram

4.2. Radiocarbon dates The pollen assemblages of the 61 subsamples

counted are diverse. The pollen diagram displays the
Four AMS radiocarbon dates (Table 1) provide most frequent pollen and spore taxa of the 102
chronological control for the São Francisco de Assis different types identified (Figs. 2 and 3). About 12
pollen record indicating that the deposit is of full- pollen and spore types remain unknown. According to
glacial, lateglacial and Holocene age. The calibration major changes in pollen assemblages and the cluster
of the radiocarbon dates have been carried out after analysis, four local pollen zones (SFA-I to SFA-IV)
CALPAL (Weninger et al., 2004). The base of the were differentiated (Table 2). Pollen concentration and
core (377 cm depth) has an age of 23,011F441 cal pollen accumulation rates are relatively stable
yr BP (20,814F211 14C years BP), but good pollen throughout the record, but in general higher during
preservation starts only at an interpolated age of glacial times and at the beginning of the Holocene
about 22,150 cal yr BP. The radiocarbon dates and than during mid and late Holocene times.
sediment stratigraphy indicate that the sedimentation Zone SFA-I (22,150–12,360 cal yr BP, 20,140–
was probably continuous, but a sedimentary gap in 10,490 14C years BP, 368–266 cm) is marked by
the poorly pollen preserved sediments between 220 abundant Campos pollen (95%), primarily Poaceae
and 195 cm core depth is possible. Based on the and lower percentages of Cyperaceae, Asteraceae
radiocarbon dates, age ranges were calculated for subf. Asterioideae, including Baccharis-type and
each pollen zone (Table 2). Uncalibrated dates for Eryngium-type, the Plantago australis-type, Borre-
each pollen zone are shown in parenthesis. For the ria, Alternanthera, Caryophyllaceae and several other
multivariate analysis calibrated radiocarbon dates herb taxa. Gallery forest taxa (1–2%) are very rare and
have been used for each sample. For the analysis only represented by a few single grains of Melasto-
mataceae, Celtis, Arecaceae, Lithraea/Schinus-type
and Podocarpus. Pollen grains from Melastomataceae
Table 1 might have originated from Campos shrubs. Aquatic
Radiocarbon dates from core São Francisco de Assis taxa (1–4%) are represented by pollen of Myriophyl-
Lab. number Depth 14
C yr BP 13
C /12C r. Calendar age lum, Hydrocotyle and one single grain of Utricularia.
(cm) (cal yr BP) Fern spores (0–2%) are rare in this and the following
Erl-5158 106 1625F39 17.34 1501F55 zones. Spores of Isoetes were found exclusively in
Erl-5159 157 3231F42 15.14 3455F51 this zone. Moss spores are only represented by
Erl-5160 266 10,462F61 17.93 12,344F208 Phaeocerous laevis (1–3%). Carbonized particles are
Erl- 4052 377 20,814F211 23.55 23,011F441
relatively rare in zone SFA-I.
242 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248

Zone SFA-II (12,360–5170 cal yr BP, 10,490–4620 are present with slightly lower percentages while
14
C years BP, 266–178 cm) continues to be charac- Cyperaceae higher. Borreria, Alternanthera and
terized by high amounts of Campos pollen. Poaceae Caryophyllaceae pollen are less frequent. Sums of

Fig. 4. Ordination diagrams of pollen percentages from the 61 samples in the São Francisco de Assis record. The diagrams map the vegetation
trajectory for the last 22,146 cal yr BP in (a) and (b), and part of the trajectory (22,146 to 11,202 cal yr BP) in (c) and (d). Calibrated years BP
are indicated for some points. (a and c) Trajectory of 61 samples, (b and d) of 83 and 59 taxa, respectively. In (b) and (d), taxa are shown in
positions proportional to their correlation level with the ordination axis (only taxa with the highest correlations are indicated); the corresponding
diagrams overlapped would form biplots. The tick marks on the axes indicate the origin coordinates. Taxa abbreviations are in Table 3.
 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248 243

the gallery forest taxa continue low. For the first time, Table 3
a few single Myrtaceae pollen was found in this zone. List of abbreviations of pollen taxa in the ordination diagrams
(Fig. 4)
Aquatic taxa are rare, represented by a few single
Abbreviation Species name
grains. Moss spores represented by Phaeocerous
laevis increase markedly during this zone (5–35%). Acal Acalypha
Alch Alchornea
Sphagnum spores occur in trace amounts. The
Alte Alternanthera
concentration and accumulation rates of carbonized Ambr Ambrosia-type
particles are markedly higher, especially during the AmCh Amaranthaceae/Chenopodiaceae
second part of the zone. Api1 Apiaceae I
Zone SFA-III (5170–1550 cal yr BP, 4620–1650 Api2 Apiaceae II
14 AsLi Asteraceae subf. Cichorioideae
C years BP, 178–106 cm) is marked by very high
AsTu Asteraceae subf. Asterioideae
values of Campos pollen (90–95%). Percentages of Bacc Baccharis-type
Asteraceae subf. Asterioideae, the Baccharis-type and Bogr Borreria latifolia-type
the Plantago australis-type are lower. Gallery forest Cary Caryophyllaceae
taxa (3–5%) are still rare, but higher than in the Celt Celtis
Crot Croton
previous zone. First pollen grains of Myrsine and Cuph Cuphea
Moraceae/Urticaceae are found. Phaeocerous laevis Cype Cyperaceae
spores are lower. Values of carbonized particles Echi Echinodorus
remain high. Erio Eriocaulaceae
Zone SFA-IV (1550 to 51 cal yr BP, 1650 to 51 Euph Euphorbia-type
14 Fab1 Fabacae I
C years BP, 106–0 cm) continues to be dominated
Fab2 Fabaceae II
by high proportions of Campos pollen (80–90%), but GoPh Gomphrena–Pfaffia-type
sums are lower than in zone SFA-III. Pollen of gallery Hydr Hydrocotyle
forest taxa (5–16%) are now markedly higher Irid Iridaceae I
represented, especially by the increase of Melastoma- Irre Iridaceae II
LaSc Latharea–Schinus-type
taceae and Myrtaceae. Acalypha and Alchornea
Lami Lamiaceae
pollen are found only in this zone. There is a decrease Mela Melastomataceae
of the gallery forest sums in the uppermost two MoUr Moraceae/Urticaceae
subsamples. Moss spores from Phaeocerous laevis Myri Myriophyllum
decrease during this zone. One single pollen grain Myrs Myrsine
Myrt Myrtaceae
from Zea mays occurs at 118 cm core depth.
Pamp Pamphalea
Accumulation rates of carbonized particles are in Pipe Piper
general higher than in the previous zone, but decrease Plau Plantago australis-type
at the end of this zone. Poac Poaceae
Podo Podocarpus
Rann Ranunculus
4.4. Results of the multivariate analysis
Scut Scutellaria-type
Seco Sebastiana commersoniana
The ordination diagram in Fig. 4a–b maps the Sene Senecio-type
trajectory of pollen composition changes from the full Sper Spermacoce
glacial period to 51 cal yr BP (AD 2001). The two- Trem Trema-type
Trix Trixis
dimensional ordination diagram accounts for 42% of
Utri Utricularia
the total variation in the data set with 83 taxa in 61 ViLa Vicia–Lathyrus
subsamples (Table 3). The main ordination axis is Xyri Xyris
indicating pollen composition change from predom-
inance of Campos during the glacial period (on the
right, see taxa in Fig. 4b) to presence of subtropical Pollen composition dynamics depicted by the
gallery forest taxa from mid Holocene (4841 cal yr ordination analysis shows phases of random, chaotic
BP) to the present (on the left in Fig. 4b). changes and phases in which the changes were
244 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248

Table 4
Summary results of ordination analyses performed with segments of the pollen composition trajectory where phase transitions were evident
Phase transition Period considered Number of Main ordination axis Decreasing taxa Increasing taxa
(cal yr BP) in the analysis subsamples (% of total variation)
(cal yr BP)
9799 to 9268 11,022 to 8206 11 28.1 Sene, Api2, GoPh, Ambr, Cuph, Cype
Myri, Api1, Bogr, Plau
4889 to 4359 9268 to 2255 15 24.8 Cype, AsLi, Xyri Myrs, GoPh, Api1, Eryn, Mela
2255 to 1562 3011 to 1309 9 28.0 Bacc, GoPh, Eryn, Api1, Bogr Myrt, Xyri, MoUr, Sper, Mela
Taxa with the highest positive or negative correlations with the main ordination axis are indicated as decreasing or increasing during the
trajectory segment. Taxa abbreviations are in Table 3.

clearly directional. Directional jumps are most cene, which coincided with higher proportions of
evident from 11,528 to 10,876 cal yr BP, from gallery forest tree taxa.
5494 to 4841 cal yr BP, from 2267 to 1442 cal yr
BP; also, the change was clearly directional in the
last two centuries. Separate ordinations including 5. Interpretation and discussion
these periods revealed which taxa were the most
correlated with the directional changes (Table 4). The pollen analytical results from the São Fran-
From 22,146 to 13,689 14C years BP, the process cisco de Assis record document that the lowlands in
did not show, at the temporal resolution, a clear eastern Rio Grande do Sul State were naturally
directional trend. Fig. 4c–d displays the vegetation covered by Campos vegetation during full- and
dynamics during this period in a separate ordina- lateglacial periods (zone SFA-I). Gallery forests,
tion; grasses, other herbs, short and tall shrubs, forming today along the rivers did not exist at that
typical of the Campos vegetation, characterized this time. The evidence of only a few pollen grains
period; tree pollen taxa was present in small belonging to gallery forest taxa may suggest a few
quantities. After this period, a more directional isolated trees or shrubs along the rivers. The climate
change in pollen composition is evident towards
an increase of tree taxa (left side of the diagram in
Fig. 4a–b: Myrsine, Myrtaceae, Moraceae/Urtica-
ceae, Celtis, Alchornea, Trema, Schinus), most
likely in the gallery forest. The process, however,
was characterized by back and forth changes at
shorter time scales, an indication of short-term
random variation superimposed on a long-term
deterministic trend.
Rates of pollen composition change between
adjacent time points (Fig. 5) were lower and more
stable during the glacial period, increased from
11,202 to 9571 cal yr BP, decreased to a minimum
between 8919 and 6309 cal yr BP and increased
by 1442 cal yr BP to levels observed until the
present.
Concentration of carbonized particles (Fig. 6) was
lower during the glacial period and increased for the
Fig. 5. Rate of change (velocity) in pollen composition computed
first time at about 11,855 cal yr BP, reaching a for 61 time steps and 83 pollen taxa based on the sediment profile at
maximum around 8919 cal yr BP. Interestingly, São Francisco de Assis, Brazil. Years on the horizontal scale are in
carbonized particles decreased after the mid Holo- calibrated years BP.
 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
Fig. 6. Forest pollen taxa percentages (black line) and concentration of carbonized particles (gray line) from 22,146 cal yr BP to present.
was relatively dry and too cold to form gallery forests
along the rivers. Shallow water conditions occurred in
the studied basin during glacial times, indicated by the
occurrence of the submerged taxa Myriophyllum and
Isoetes. Shallow lake conditions during glacial times
have also been observed in records from the southern
highlands (e.g., Behling, 1995). The pollen record
suggests that there is no marked vegetation change
from the full-glacial to the lateglacial. Indeed, the
rates of compositional change, as indicated by multi-
variate analysis, were much slower during this period
than in the Holocene. However, a long-term direc-
tional trend was revealed, probably related to changes
in climate, with short-term random variation in shorter
time scales. Fire was rare on the Campos during
glacial times.
The transition from the lateglacial to the Holocene
(radiocarbon date at 266 cm is 10,460 14C yr BP)
shows a change from brown-gray clay with organic
matter below 265 cm core depth to completely
decomposed black organic matter above 265 cm,
reflecting a change from cold and dry to warm and dry
conditions. During the early Holocene (zone SFA-II),
Campos vegetation continued to dominate and sub-
tropical gallery forests were not apparent. The
composition of the Campos vegetation changed. For
example Borreria, Alternanthera and Caryophylla-
ceae were less frequent in the Campos than during the
glacial period. In the basin, the shallow lake environ-
ment disappeared and organic matter accumulated.
The increased presence of Cyperaceae reflects the
245
development of a wetland. The marked increase of the
moss Phaeocerous laevis is a good indication for
warm conditions. This moss indicates also drier soils
and less vegetation cover in the areas surrounding the
peat bog (Behling, 1995; Behling et al., 2004). Pollen
preservation in the subsamples between 195 and 220
cm core depth is very poor and also suggest dry
conditions. A possible sedimentary hiatus, which
would also cause an error in the rate of change
analysis, cannot be excluded for this period. Fires
became frequent with the beginning of the Holocene.
In view of the results of the Cambará do Sul core from
the southernmost highland in northeastern Rio Grande
do Sul State (about 450 km east of the site), where
fires became frequent only after 7400 cal yr BP and
not at the beginning of the Holocene, it is suggested
that the increase of fire frequency in the São Francisco
de Assis might be anthropogenic. There is archeo-
logical evidence that suggested that Amerindians
already lived in the lowlands in western the Rio
Grande do Sul State by that time (Dillehay et al.,
1992; Kern, 1997a). Seasonally dry conditions during
the early Holocene may have favoured frequent fires,
especially after 12,300 cal yr BP. Indeed, a phase
transition between 11,528 and 10,876 cal yr BP was
detected by multivariate analysis, mainly character-
ized by the decline of Senecio and Apiaceae and
increase of Cuphea and Cyperaceae. Probably both,
warmer and drier conditions and the increase of fire
frequency may have changed the composition of the
Campos vegetation.
246 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
Since mid Holocene times (zone SFA-III) at 5170
cal yr BP subtropical gallery forests developed along
rivers. This was detected by multivariate analysis as a
phase transition between 5494 and 4841 cal yr BP
and as a sharp increase in the rate of compositional
change. This phase transition was mainly character-
ized by a decrease of Cyperaceae and Asteraceae
subf. Cichorioideae and increase of Myrsine. The
plant diversity of this forest was at that time relatively
low, represented only by a few tree species. The
formation of gallery forest reflects a change to wetter
conditions. This is also supported by the decline of
the moos Phaeocerous laevis. The lower proportions
of different Asteraceae species and Plantago aus-
tralis may also be related to wetter conditions.
Geomorphological evidence presented by others
(see review in Kern, 1997b) of a dry period after
mid Holocene is not supported by our findings. A
change to wetter conditions was also recorded after
4320 cal yr BP in the Cambará do Sul core (Behling
et al., 2004). At the São Francisco de Paula site, also
located on the southernmost highlands in northeastern
Rio Grande do Sul pollen was preserved only after
4500 cal yr BP (interpolated age) (Behling et al.,
2001). Fires, probably of anthropogenic origin, were
still frequent in the Campos during mid to late
Holocene times. A sign of agricultural activity is
evident at the end of this period, by a single Zea
mays pollen grain at 1960 cal yr BP (1950 14C yr BP,
118 cm core depth). The evidence of only one corn
pollen grain in the pollen record may suggest either a
small-size corn plantations near the site or plantations
distant to the site, which is consistent with archeo-
logical evidence that the Guarany, who were horti-
culturists, arrived in the Uruguay River around this
time (Schmitz, 1997).
Based on floristic analysis, Rambo (1961), Reitz
and Klein (1964) and Klein (1975, 1980) showed that
there were several plant migration routes in southern
Brazil during the Holocene. Species migrated into the
western lowlands in Rio Grande do Sul mostly by two
routes, from the eastern Atlantic coastal lowland and
from the western Rio Paraná valley. The evidence of
Myrsine at the beginning of pollen zone SFA-III, and
somewhat later the occurrence of Moraceae indicate
migration of species from both forest areas. The
evidence of Cecropia pollen at 2880 cal yr BP, though
in low numbers, indicates migration from the Atlantic
coastal lowland, where the genus is restricted at
present, to the western lowlands of Rio Grande do
Sul, reflecting wetter and warmer conditions at that
time. The migration route was likely along the
lowlands south of the South Brazilian highlands and
north of the crystalline shield, where at present
Cecropia and other typical Atlantic taxa, such as
Euterpe edulis, reach sites ca. 250 km eastward from
the study area.
During the late Holocene period (zone SFA-IV),
after 1550 cal yr BP, gallery forest expanded in the
study region, forming larger areas along the rivers and
some patches of forest in river valleys. This change
was detected by multivariate analysis as a phase
transition between 2267 and 1442 cal yr BP mainly
characterized by a decline of Baccharis, Gomphrena–
Pfaffia-type, and Eryngium and increase of Myrtaceae
and Xyris. The increase of forest reflects clearly wetter
conditions, related to higher rainfall and probably a
shorter dry season. This has also been shown by
results from the southern highlands (Behling, 2002).
The increase of Myrtaceae and the decrease of the
moos Phaeocerous laevis during this zone is a good
indication for this wettest recorded period. During this
period, Alchornea and Acalypha (at 1090 cal yr BP
and at 970 cal yr BP, respectively) migrated into the
study region. Changes to wetter conditions are
recorded in the Cambará do Sul core by the expansion
of Araucaria forests after 1100 cal yr BP. At the São
Francisco de Paula site, forest expanded after 960 cal
yr BP. Forest expansion apparently started somewhat
earlier in the western lowlands, but maximum gallery
forest cover was reached at 1090 cal yr BP (78 cm
core depth).
High accumulation rates of carbonized particles
indicate that fires were most frequent during this
period, despite of wetter conditions. Abundant fires
suggest that the human occupation of the study region
may have increased at that time. The slight decrease of
forests at the end of the zone may be related to
deforestation.
6. Conclusion
The pollen and charcoal records from São Fran-
cisco de Assis reveal for the first time evidence for
palaeoenvironmental changes in of the southern
 H. Behling et al. / Review of Palaeobotany and Palynology 133 (2005) 235–248
Brazilian lowland since the last glacial maximum.
Four AMS radiocarbon dates provide time control for
the about 22,000 cal yr BP old pollen record.
Pollen data document that the lowland of western
Rio Grande do Sul State was covered with natural
Campos vegetation throughout the recorded glacial and
postglacial period. Modern gallery forests did not exist
during glacial times and developed in form of small
strips along rivers only after 5170 cal yr BP. Gallery
forest expansion occurred after 1550 cal yr BP with a
maximum at 1090 cal yr BP. There is evidence of plant
species migration from the western Atlantic lowlands
to the São Francisco de Assis region by Myrsine and
Moraceae since at about 5200 cal yr BP and by
Alchornea and Acalypha after about 1000 cal yr BP.
Multivariate analysis of the pollen data reveals
palaeovegetation trajectories in ordination space and
measure the rate of compositional change. The
dynamics of pollen composition involved both
undirected, chaotic phases and phases with directed,
sometimes fast vegetation changes, supporting the
theory that vegetation dynamics is complex charac-
terized by a combination of deterministic and
chaotic behavior (Anand, 2000; Anand and Orlóci,
1997; Orlóci et al., 2002b). This has also been
observed in the analysis of the Cambará profile
(Behling et al., 2004).
The palaeovegetation record indicates that Campos
vegetation existed in the area under a relatively dry and
cold climate during glacial times, and warm and dry
conditions during the post glacial times. A change to
wetter conditions is reflected by the development of
gallery forest and gallery forest expansion beginning at
5170 cal yr BP and especially after 1550 cal yr BP,
respectively. The palaeoclimatic results from the São
Francisco de Assis core show that past climate changes
in the southernmost highlands in Rio Grande do Sul
occurred at similar times also in the southern lowlands.
The charcoal record documents that natural fires
were rare during glacial times. Fire became frequent
only at the beginning of the Holocene, suggesting
anthropogenic fires by the first occupation of the
western lowlands in Rio Grande do Sul. In the
southernmost highlands, fire became very frequent
only after 7400 cal yr BP, suggesting an earlier human
occupation in the lowlands. The change to warm and
dry climatic conditions and frequent fires may have
been an important factor for changing the floristic
247
composition of the Campos. The highest fire fre-
quency occurred only during the late Holocene under
the wettest climatic conditions. The population of
Amerindians probably increased during that time in
the region around São Francisco de Assis. There is
evidence of corn (Zea mays) plantations at 1960 cal
yr BP.
Acknowledgments
We are grateful to Carlos Alberto Silveira (owner
of Itajuru Farm), Fernando Quadros and José Pedro
Trindade for providing invaluable assistance in the
field, and László Orlóci for suggestions on the
manuscript. The authors thank Vera Markgraf,
Marie-Pierre Ledru and one anonymous reviewer
for providing constructive comments. The project
was supported by Conselho Nacional de Desenvol-
vimento Cientı́fico e Tecnológico (CNPq; Brazil)
with a research grant and fellowship given to Valério
De Patta Pillar.
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