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 Journal of South American Earth Sciences 14 (2001) 631±639
www.elsevier.com/locate/jsames

Holocene environmental changes in the SaÄo Francisco de Paula region,

southern Brazil
Hermann Behling a,*, Soraia Girardi Bauermann b, Paulo CeÂsar Pereira Neves b
a
Center for Tropical Marine Ecology, University of Bremen, Fahrenheitstraûe 6, Bremen D-28359 Germany
b
Universidade Luterana do Brazil (ULBRA), Departamento de Biologia,Rua Miguel Tostes 101, Canoas RS 90.420-280 Brazil
Received 1 February 2000; revised 1 January 2001; accepted 1 March 2001

Abstract
Holocene paleoenvironments have been interpreted from a radiocarbon dated pollen and charcoal record of the SaÄo Francisco de Paula
region on the southern Brazilian highland. Today the region is characterized by pastureland and small patches of disturbed Araucaria forest.
The region receives 2450 mm per annum. This is the highest precipitation rate in southern Brazil. Studied surface samples represent the
modern pollen analog of the anthropogenic in¯uenced vegetation. Pollen analytical studies of the 84 cm long core, collected from a small
basin with clay, organic matter and peat deposits, show that pollen and spores of the lower core section (84 and 46 cm depth) have been
almost destroyed. This period between ca. 7500 and 4000 14C yr bp was too dry for conservation of pollen and spores in the basin. Climate
must have been markedly drier than today during early and mid Holocene times. After 4000 14C yr bp, preserved pollen grains re¯ect wetter
conditions than before and indicate the predominance of campos (grassland) vegetation with small areas of Araucaria forest in the study
region. Forest expansion is documented since 1060 14C yr bp and expansion of Araucaria angustifolia trees itself since 850 14C yr bp. During
the last 1000 yr, rainfall must have been much more intensive with no or only short dry periods such as the modern climate. The results
con®rm the vegetational and climatic changes documented from the Araucaria forest region of Santa Catarina and Parana State. Concentra-
tion and accumulation rates of carbonized particles are somewhat higher during the last 850 14C yr bp than before, indicating an increased ®re
frequency. q 2001 Elsevier Science Ltd. All rights reserved.
Keywords: Holocene; Paleovegetation; Araucaria forest; Grassland (campos); Paleoclimate; Paleo®res; Southern Brazil

1. Introduction marked dry seasons (Hueck, 1966), is an excellent climate

Several paleoenvironmental studies from the Araucaria
forest region of the southern and southeastern Brazilian
highland, which have been carried out in the last few
years, have shed more light into Late Quaternary vegetation
and climate history of this region (e.g. Behling, 1995,
1997a,b, 1998; Behling and Lichte, 1997; Behling et al.,
1998; De Oliveira, 1992; Ledru, 1993). Compared to other
regions the southern Brazilian highland is poorly studied,
despite the importance of the region for climate change.
Paleoecological data from the Araucaria forest region are
an important contribution for understanding present-day
ecosystems, its stability and response to climate change.
Especially the conifer Araucaria angustifolia, which
requires a minimum of 1400 m annual rainfall and no
indicator.
SaÄo Francisco de Paula is an interesting locality to study
the vegetation and climate history and to compare with the
data of other studied sites from the Araucaria forest region.
The southernmost highland region receives the highest
precipitation rate of southern Brazil and is very important
for climate change studies. If past precipitation had been
only slightly reduced, this region would have been covered
by Araucaria forest since the beginning of the Holocene
warming. If past rainfall rates had been strongly reduced
and a marked dry season occurred, vegetational changes
similar to those documented for the other southern Brazilian
highland regions, would be expected.
2. Study region

The study area is found on the southernmost highland

* Corresponding author. Tel.: 149-421-238-0046; fax: 149-421-238-
region of southern Brazil at 900 m elevation in Rio Grando
0030. do Sul State. The core site is situated on the farm named
E-mail address: hbehling@zmt.uni-bremen.de (H. Behling). Fazenda do Pinto (29824 0 S, 50834 0 W), which is 8 km from
0895-9811/01/$ - see front matter q 2001 Elsevier Science Ltd. All rights reserved.
PII: S 0895-981 1(01)00040-2
632 H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639

Fig. 1. Map showing site locations in the SaÄo Francisco de Paula area, State of Santa Catarina, southern Brazil. Shaded areas indicate distribution of the
Araucaria forest. Key: X ˆ Fazenda do Pinto core near the city of SaÄo Francisco de Paula; 1 ˆ Serra do Rio Rastro; 2 ˆ Morro da Igreja; 3 ˆ Serra da Boa
Vista; 4 ˆ Serra Campos Gerais (in Parana State).

the city of SaÄo Francisco de Paula, near the road (Estrada do
Meio) that runs northeast to the village Bom Jesus. The
studied deposits are found in a small rocky and shallow
basin that is 25 m in diameter.
The vegetation prior to post-Columbian settlement can be
described as a mosaic of campos (grassland) and small
patches of Araucaria forest. Dates on the ¯oristic composi-
tion of the southern Brazilian highland have been published
Ð e.g. Hueck (1953, 1966), Klein (1978, 1979, 1984),
Negrelle and Da Silva (1992), Rambo (1953, 1956) and
Por (1992). The campos vegetation is highly diverse and
characterized by species that belong to the main families
such as Poaceae, Cyperaceae, Asteraceae, Apiaceae and
Eriocaulaceae. Other important herbs belong to the genera
Polygala, Xyris, and Plantago. Continuously subtropical
Araucaria forest occurs in southern Brazil between latitudes
248 and 308S, primarily at elevations between 1000 and
1400 m, and in southeastern Brazil in isolated `islands'
between 18 and 248S at elevations between 1400 and
1800 m (Hueck, 1953) (Fig. 1). Important trees include,
for example, Araucaria angustifolia, Ocotea sp., Ilex para-
guariensis, Mimosa scabrella, and Podocarpus lambertii.
Atlantic rain forest is found in the lowland and on the slopes
to the east, while the `Rio Uruguai rain forest' is found in the
lowland and on the slopes to the west. The modern vegeta-
tion of the study region is strongly affected by pasture activ-
ities and some agriculture. The local vegetation of the basin
is a mosaic of grass and Sphagnum moss patches. The
surrounding vegetation is pastureland. Small areas of
disturbed Araucaria forest rich in Araucaria angustifolia
 H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639
Table 1
Radiocarbon dates for the Fazenda do Pinto core
Lab. no. Depth (cm) 14
C yr bp a 13
C/ 12C Ratio
Hv-20837 25±30 1185 ^ 190 219.6
Hv-20836 57±59 5775 ^ 190 220.8
Hv-20835 71±73 6750 ^ 215 221.6
a 13
C adjusted.
and Podocarpus are found some distance from the studied
basin. Pine (Pinus) plantations also occur in the study
region.
The climate of the study region is characterized as warm
and humid without marked dry periods (Nimer, 1989). The
study area is located in a zone with an average annual rain-
fall of over 2250 mm, which is the highest in southern
Brazil. Precipitation is very high because the southern high-
land forms an orographic barrier for southeasterly moisture.
As documented from the SaÄo Francisco de Paula station, the
climate records a mean annual precipitation of 2456 mm.
The mean annual temperature is 14.58C and the lowest
measured temperature is 26.58C (Nimer, 1989). The atmo-
spheric circulation of southern Brazil is dominated by the
South Atlantic anticyclone, a semi-permanent high pressure
system that transports moist tropical air masses over the
continent in easterly and northeasterly directions during
the entire year. Disturbances to this pattern are related to
perturbations of polar cold fronts that, upon meeting tropical
air masses, produce strong rainfall in southern Brazil
(Nimer, 1989; Hastenrath, 1991). Higher rainfall in southern
Brazil is also related to El NinÄo events (Martin et al., 1993;
McGlone et al., 1992; Ratisbona, 1976).
3. Methods
Deposits were cored in the central part of the basin using
a Russian corer. The 84 cm-long core was wrapped in plas-
tic and aluminum ®lm and stored under refrigeration (148C)
after return from the ®eld and before sampling. For radio-
carbon dating, three samples were taken from the core.
Moss and soil surface samples for modern pollen analogs
were collected in the surroundings of the studied basin and
in the disturbed Araucaria forest.
For pollen analysis, the samples (0.3 cm 3) were generally
taken at 4 cm intervals along the pro®le. All samples were
treated using standard methods with hydro¯uoric acid (47±
52%) and acetolysis. Pollen residues were mounted on
slides in a glycerin gelatin medium. Pollen preparation
included the addition of exotic Lycopodium spores to deter-
mine pollen concentration (grains/cm 3) and pollen in¯ux
(grains/cm 2/yr). Pollen samples were counted to a minimum
of 300 grains for surface samples and 500 grains for core
samples. The total pollen sum includes herbs, shrubs, and
trees, but not aquatic taxa, fern and moss spores, and algae
colonies of Botryococcus. Carbonized particles were
633
counted on pollen slides and expressed as percentage of
the total pollen sum, as concentration and in¯ux. Pollen
and spores from samples of the lower core section (depth
between 84 and 46 cm) were almost destroyed and not
included in the analysis. Pollen identi®cation relied on the
H. Behling's reference collection (containing about 1600
Brazilian species) and pollen morphological descriptions
in Behling (1993). Identi®ed taxa were classi®ed into
following ecological groups: campos, Araucaria forest,
other shrubs and trees, aquatics, and ferns. The software
programs tilia, tiliagraph, and coniss were used for
data plotting in pollen diagrams, calculations, and cluster
analysis (Grimm, 1987).
4. Results
4.1. Stratigraphy
The 84 cm-long core starts below the ®ne grass layer. The
stratigraphy from the Fazenda do Pinto core can be
described as follows:
0±8 cm Brown, weakly decomposed and
unconsolidated peat, with many plant
roots
8±14 cm Dark brown, compact, almost
completely decomposed peat, with
roots
14±64 cm Black, very compact, totally
decomposed organic material, with
some rootlets
64±84 cm Brown±gray clay, with a few rootlets
84 cm Gray clay on rocky subsurface
4.2. Radiocarbon data
Three conventional radiocarbon dates (Table 1) from the
84 cm long core provide chronological control. The base of
the core is extrapolated at 7590 14C yr bp. The pollen record
itself starts at 46 cm core depth at 3970 14C yr bp. Radio-
carbon ages for the two pollen zone (extrapolated or inter-
polated) are based on uncalibrated 13C-adjusted radiocarbon
ages.
4.3. Modern pollen data from surface samples
Pollen rain data from collected surface samples of the
pastureland near the studied basin and from the disturbed
Araucaria forest, which is found about 100 m from the core
site, are shown in Fig. 2. Pollen samples (Nos. 1±4) of the
campos area (pastureland) contain high percentages of herb
pollen (60±80%) mainly Poaceae, Cyperaceae, and Aster-
aceae. The Araucaria forest pollen sums are low (5±10%).
The sums of other shrub and tree pollen (5±30%) are higher.
Percentages of fern spores are low, except in one sample.
 634
H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639

Fig. 2. Percentage diagram of surface samples from the surroundings of the studied basin (samples 1±4) and from the disturbed Araucaria forest of the study region (samples 5±8).
 H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639

Fig. 3. (a) Pollen percentage diagram from the Fazenda do Pinto core. (b) Summary pollen percentage diagram from the Fazenda do Pinto core, including the records of pollen concentration, pollen in¯ux, and
carbonized particles, and the cluster analysis dendrogram.
635
636 H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639

Fig. 3. (continued)
 H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639 637

Table 2 The pollen assemblages (Nos. 5±8) of the secondary

List of identi®ed taxa from the Fazenda do Pinto core Araucaria forest contain low percentages of campos pollen
Campos Araucaria Forest (7±16%), while sums of the Araucaria forest group are high
Abutilon/Malvastrum-type Araucaria angustifolia (70±90%), especially the high percentages of Araucaria and
Amaranthaceae/ Ilex Podocarpus pollen. Percentages of other shrub and tree
Chenopodiaceae I pollen are low, and percentages of fern spores are higher
Amaranthaceae/ Mimosa scabrella-type
than in the samples from the campos area.
Chenopodiaceae II
Ambrosia-type Podocarpus
Baccharis-type Sebastiania commersoniana 4.4. Fossil pollen data
Borreria I Symplocos lanceolata-type
Borreria II The pollen record from the Fazenda do Pinto core starts at
cf. Apium Other Shrubs and Trees a core depth of 46 cm (Fig. 3a and b). Samples below 46 cm
Convolvulaceae Alchornea contain almost destroyed pollen grains and were not
Cuphea Allophylus included in the diagram. The percentage diagram illustrates
Cyperaceae Alnus(long distance transport)
Eriocaulon/Paepalanthus Arecaceae
the most abundant pollen and spore taxa (Fig. 3a) of the 104
Eryngium-type Celtis different types, including 7 unknown types. The list of iden-
Gomphrena/Pfaf®a-type cf. Juniperus ti®ed taxa is given in Table 2. The pollen concentration and
Iridaceae I Chrysopyllum viride-type pollen in¯ux in the sediments are somewhat lower in the
Iridaceae II Croton lower part of the core than in the upper part. Based on major
Lamiaceae Daphnopsis
Other Asteraceae subf. Ephedra(long distance transport)
changes in the pollen assemblages, primarily by the increase
Asteroideae of Araucaria angustifolia pollen grains, two local pollen
Other Asteraceae subf. Ephedra tweediana-type (long zones of the diagram can be estimated: Zone SFP-I (46±
Cichorioideae distance transport) 20 cm, 3970±850 14C yr bp, 6 samples) and Zone SFP-II
Pamphalea Ericaceae (20±0 cm, 850 14C yr bp Ð i.e. modern, 5 samples).
Papilionaceae I Lamanonia speciosa-type
Papilionaceae II Matayba
The pollen diagram is characterized by abundant herb
Papilionaceae III Melastomataceae/Combretaceae pollen (80±85%), primarily Poaceae, with lower percen-
Petunia-type Meliaceae tages of Cyperaceae, Baccharis-type, other Asteraceae,
Plantago Moraceae/Urticaceae (P2) Eryngium-type, and several other herb taxa. The highest
Poaceae Moraceae/Urticaceae (P3) values of Poaceae pollen (50±60%) are found in Zone
Polygala Myrsine
Ranunculus bonariensis Myrtaceae
SFP-I. The Poaceae pollen decrease in Zone SFP-II to
Senecio Nothofagus dombeyi-type (long 29% at the core top, while Cyperaceae pollen increase
distance transport) from 10 to 40%. Pollen grains from the Araucaria forest
Spermacoce Phrygilanthus acutifolius group (Araucaria, Podocarpus, Ilex, Mimosa scabrella-
Verbena isabellii-type Piper type, Symplocos lanceolata-type) are rare in Zone SFP-I
Vernonia-type Prockia crucis-type
Vicea/Lathyrus Roupala
(about 2%) and higher in Zone SFP-II (about 5%). The
Xyris Sapium Araucaria angustifolia pollen, especially, increase from
Zea mays Sapotaceae about 1% in Zone SFP-I to 3% in Zone SFP-II. The group
Sebastiania schottiana-type of other shrubs and trees (mainly Moraceae/Urticaceae,
Aquatics Sloanea Myrsine, Melastomataceae, and Myrtaceae) shows low
Hydrocotyle Solanum I
Ludwigia Solanum II
sums of about 5% in Zone SFP-I. Sums increase up to
Typha Struthanthus 10±13% at the end of Zone SFP-II. Fern spores are rela-
Utricularia Styrax tively low in percentages, while moss spores of Sphagnum
Tetrorchidium rubrivenium and Phaeoceros laevis are relatively high. Algae colonies of
Ferns Trema-type Botryococcus are frequent in the lower part of Zone SFP-I
Cyathea-type Weinmannia
Dicksonia sellowiana Zanthoxylum-type
and in the uppermost sample. Concentration and in¯ux of
Hymenophyllum-type charcoal particles are somewhat lower in Zone SFP-I than in
Isoetes Mosses Zone SFP-II.
Lycopodium alopecuroides-type Phaeoceros laevis
Lycopodium clavatum-type Sphagnum
Monolete psilate ,50 mm 5. Discussion and conclusion
Monolete psilate .50 mm Algae
Monolete verrucate ,50 mm Botryococcus The use of modern pollen analogs from surface
Monolete verrucate .50 mm
Pteris-type Unknowns
samples of the study region for reconstruction of vegeta-
Trilete psilate Type 1±7 tional changes is somewhat limited, because of the strong
Trilete verrucate human in¯uence on the vegetation. However, surface
samples indicate that the local pollen rain of the pastureland
and Araucaria forest species is quite high, while pollen
638 H. Behling et al. / Journal of South American Earth Sciences 14 (2001) 631±639
transport by wind from grassland taxa into the forest is low.
Wind-transported pollen from forest taxa to the pastureland
(surface samples were taken at a distance of ca. 100 m into
the pastureland) is even lower. In particular, Araucaria and
Podocarpus pollen have high percentages in forest surface
samples and very low values in surface samples of the grass-
land. The transport of these pollen grains over distance
seems to be minor, but it has to be considered that the
areal extent of the Araucaria forest is quite small compared
to the huge pastureland areas of the study region. Studies on
modern pollen rain in the Atlantic rain forest lowland in
Santa Catarina State indicate that Araucaria pollen grains
can be transported quite well over some distance (Behling et
al., 1997). Future results from modern pollen rain transect
studies from undisturbed Araucaria forest to campos vege-
tation will be very useful for detailed reconstruction of past
vegetation.
The pollen analytical results from the SaÄo Francisco de
Paula region indicate that the climate was apparently too dry
for conservation of pollen and spores in the studied shallow
basin during the period from ca. 7500 14C yr bp (extrapo-
lated) until ca. 4000 14C yr bp. Annual precipitation was
signi®cantly lower during the early and mid-Holocene
than it is today, probably in the range 30±40%. It is possible
that a 3-month dry season occurred at during that time
(Behling, 1997b). During the period from ca.
4000 14C yr bp until ca. 1000 14C yr bp, the pollen record
documents the study region to be covered primarily with
campos vegetation, with a few small patches of Araucaria
forest. Because of the preserved pollen in the deposits, this
period must have been somewhat wetter and the annual dry
period somewhat shorter than before. The initial forest
expansion is evident at 1060 14C yr bp. Araucaria angusti-
folia itself became more frequent at 850 14C yr bp. These
vegetational changes re¯ect the change to modern climatic
conditions with high precipitation rates (.2200 mm) and
with no or very short dry periods.
The new results con®rm the documented environmental
changes from the studied Araucaria forest and campos
regions (Fig. 1) of Santa Catarina State and Parana State
(Behling 1995, 1997b). Paleoenvironmental data from sites
in Serra do Rio Rastro, Morro da Igreja, Serra da Boa Vista,
and Serra Campos Gerais (in ParanaÂ) show that huge areas
of campos vegetation were still found on the highlands of
southern Brazil during early and mid-Holocene times,
re¯ecting warm and dry climates with an annual dry
season of probably about three months (Behling, 1997b).
Initial expansion of Araucaria forests started from gallery
forests along the rivers about 3000 yr bp and re¯ect a some-
what wetter climate than before. A pronounced expansion
of Araucaria forests on the highland, replacing the
campos vegetation, is found in Santa Catarina State since
1000 yr ago and in Parana State (Serra Campos Gerais)
since 1500 yr ago. This late Holocene period is character-
ized by a very humid climate without a marked dry period.
It has been reported that the Araucaria forest on the south-
ern Brazilian highland, under natural conditions, is in
expansion today (e.g. Klein, 1960, 1975; Rambo, 1956,
1960).
Wetter conditions on the southern Brazilian highland can
be explained by a stronger in¯uence of cold fronts during
the last 1000±1500 yr, and there is less in¯uence from dry
tropical continental air masses. Moist tropical Atlantic air
masses, which transport rain to southern Brazilian, might
have had a stronger in¯uence during this late Holocene
period.
The southernmost highland is still mainly covered by
campos vegetation, with only poor coverage by Araucaria
forest compared to other southern highland regions. Only
during the last millenium have patches of Araucaria forest
in the SaÄo Francisco de Paula region increased in area and in
numbers. In the record, Araucaria angustifolia reached its
maxima of only 3% of the pollen sum. The reason for this
lessening of forest expansion seems to be the short time,
about 800 yr, between the change to very humid climatic
conditions that occurred about 1000 yr ago and the forest-
inhibiting use of the campos as pastureland for the last 200±
250 yr.
The increase of carbonized particles suggests a higher ®re
frequency during the last 850 yr. This is probably related to
human activity, because documented wetter climatic condi-
tions would reduce ®re frequency. How much ®re may have
favored (Soares, 1979) or blocked the expansion of the
Araucaria forest is still unclear. The role of paleo®res in
the Araucaria and campos region should be studied in more
detail.
Acknowledgements
We thank the owner of Fazenda do Pinto for his kind
permission to do ®eldwork on the farm. We thank Jorge
Rabassa, Mirta Quattrocchio, and an unknown reviewer
for providing constructive reviews. The NiedersaÈchsisches
Landesamt fuÈr Bodenforschung (Hannover) is thanked for
radiocarbon dating. The ®rst author thanks the Deutsche
Forschungsgemeinschaft (DFG) for the scholarship.
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