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Perspectives in Plant Ecology, Evolution and Systematics 9 (2007) 101116
www.elsevier.de/ppees
Abstract
The South Brazilian grasslands occupy some 13.7 million ha and support very high levels of biodiversity. This paper
reviews the current state of ecological knowledge on South Brazilian Campos and of threats and challenges associated
with their conservation. The principal factors shaping grassland physiognomy and diversity are discussed, and
information is presented on diversity of plant species; best estimates suggest that 30004000 phanerophytes occur in
the South Brazilian grasslands.
It is argued that, despite their high species richness, Campos vegetation is not adequately protected under current
conservation policies. In the past three decades, approximately 25% of the grassland area has been lost due to land use
changes, and this trend continues. However, representation of Campos grasslands in conservation units is extremely
low (less than 0.5%), and the management in most of these is inadequate to preserve the grasslands, as grazing and re
are important factors for their persistence. In conclusion, the following urgent needs are identied: (1) to create more
conservation units in different regions, including different grassland types throughout southern Brazil, (2) to develop
proper management strategies where grasslands are subject to shrub encroachment and forest expansion, (3) to
conduct research on biodiversity and ecological processes in the Campos region and (4) to raise public awareness of the
value and vulnerability of this vegetation type.
r 2007 Rubel Foundation, ETH Zurich. Published by Elsevier GmbH. All rights reserved.
Keywords: Biodiversity; Conservation; Grassland; Land use change; Plant species; Preservation
Corresponding author. Academy of Spatial Research and Planning, Scientic Section III: Natural Resources, Environment, Ecology,
Hohenzollernstrae 11, 30161 Hannover, Germany.
Tel.: +49 511 3484 222.
E-mail address: gerhard_overbeck@yahoo.com (G.E. Overbeck).
Introduction
Brazil is one of the worlds megadiverse countries (e.g.
Barthlott et al., 1996; Lewinsohn and Prado, 2005), but
1433-8319/$ - see front matter r 2007 Rubel Foundation, ETH Zurich. Published by Elsevier GmbH. All rights reserved.
doi:10.1016/j.ppees.2007.07.005
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Fig. 1. Location of South Brazilian grasslands: (a) overview, (b) ofcial Brazilian classication of biomes (IBGE, 2004) and (c)
distribution of grasslands in Brazils southern region (abbreviation of states: RS: Rio Grande do Sul; SC: Santa Catarina, PR:
Parana).
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known as campo limpo or campo sujo (e.g. OliveiraFilho and Ratter, 2002). Describing the South Brazilian
grasslands as savannas and steppes is thus in disagreement with the accepted international use of these terms
(also see Marchiori, 2002).
Classical botanical and phytogeographical studies
(e.g. Lindman, 1906; Rambo, 1956a) and more recent
works on grassland vegetation in southern Brazil
(e.g. Boldrini, 1997; Pillar and Quadros, 1997; Overbeck
and Pfadenhauer, 2007), although without classicatory
objectives, prefer to refer to grassland formations in
southern Brazil simply as Campos. However, terms as
Campo limpo (clean grassland, i.e. without a woody
component) and Campo sujo (dirty, i.e. shrubby grassland) have become commonly used. In trying to
differentiate different types of grasslands within the
Campos region, most studies reect the two different
biogeographical domains (see above; see Tables 1 and 2
for a compilation of characteristic species) and regional
differences in Campos ora, with notably higher
contribution of C3 grasses (e.g. from the genera Briza,
Piptochaetium, Poa, Stipa) in the southern half of Rio
Grande do Sul (Burkart, 1975; Valls, 1975). Boldrini
(1997) describes six physiognomic regions of Campos
vegetation in Rio Grande do Sul state, considering local
oristic variations associated with climate, topographic
variation and soil heterogeneity. However, a good
portion of the variation in grassland physiognomy
(e.g. distinction between campo limpo and campo sujo)
and in the composition of the dominant species,
irrespective of region, seems to be determined by grazing
and re regimes (Pillar and Quadros, 1997). Altogether,
internal classication of the Campos grasslands is still in
need of further research concerning oristic and
structural differentiation and relative impacts of climate,
substrate and management. Henceforth, when we refer
to Campos or South Brazilian Grasslands, or Campos/
grassland region, without any further qualication, we
mean both grasslands associated with Araucaria forest
and the grasslands collectively considered as Pampa in
the current classication of biomes by IBGE (2004) (see
Fig. 4 for some impressions of Campos landscapes).
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Table 1. Characteristic families and species of grasslands in the Atlantic Forest biome (grasslands in northern Rio Grande do Sul,
Santa Catarina and Parana)
Amaryllidaceae
Hippeastrum breviflorum Herb.
Apiaceae
Eryngium horridum Malme
Eryngium pandanifolium Cham. & Schltdl.
Eryngium urbanianum H. Wolffa
Eryngium zozterifolium H. Wolffa
Asteraceae
Baccharis milleflora (Less.) DC.
Baccharis sagittalis (Less.) DC.
Baccharis uncinella DC.a
Calea phyllolepis Baker
Hypochaeris catharinensis Cabrera a
Noticastrum decumbens (Baker) Cuatrec.
Senecio juergensii Mattf .
Senecio oleosus Vell.a
Trichocline catharinensis Cabreraa
Campanulaceae
Lobelia camporum Pohl
Cyperaceae
Ascolepis brasiliensis (Kunth) Benth. ex C.B.Clarke
Bulbostylis sphaerocephala (Boeck.) C.B. Clarke
Carex brasiliensis A.St.-Hil.
Carex longii Mack. var. meridionalis (Kuk.) G.A. Wheeler
Eleocharis bonariensis Nees
Lipocarpha humboldtiana Nees
Pycreus niger (Ruiz & Pav.) Cufod.
Rhynchospora barrosiana Guagl.
Rhynchospora globosa (Kunth) Roem. & Schult.
a
Fabaceae
Adesmia ciliata Vogel
Adesmia tristis Vogel
Eriosema longifolium Benth.
Galactia neesii DC.
Lathyrus paranensis Burkart
Lupinus reitzii M. Pinheiro & Miottoa
Lupinus rubriflorus Planchueloa
Lupinus uleanus C. P. Sm.a
Macroptilium prostratum (Benth.) Urb.
Rhynchosia corylifolia Mart. ex Benth.
Trifolium riograndense Burkart a
Poaceae
Andropogon lateralis Nees
Andropogon macrothrix Trin.
Axonopus siccus (Nees) Kuhlm.
Axonopus suffultus (Mikan ex Trin.) Parodi
Bromus auleticus Trin. ex Nees
Paspalum maculosum Trin.
Paspalum pumilum Nees
Schizachyrium tenerum Nees
Stipa melanosperma J. Presl
Stipa planaltina A. Zanin & Longhi-Wagnera
Solanaceae
Petunia altiplana Ando & Hashimoto
Verbenaceae
Glandularia megapotamica (Spreng.) Cabrera & Dawson
Verbena strigosa Cham.
Endemic species.
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Table 2.
G.E. Overbeck et al. / Perspectives in Plant Ecology, Evolution and Systematics 9 (2007) 101116
Characteristic families and species of grasslands in the Pampa biome (grasslands in southern Rio Grande do Sul)
Apiaceae
Eryngium elegans Cham. & Schltdl.
Eryngium horridum Malme
Eryngium sanguisorba Cham. & Schltdl.
Asteraceae
Acmella bellidioides (Sm.) R. K. Jansen
Aspilia montevidensis (Spreng.) Kuntze
Aster squamatus (Spreng.) Hieron.
Baccharis coridifolia Spreng.
Baccharis dracunculifolia DC.
Baccharis trimera (Less.) DC.
Chaptalia runcinata Kunth
Eupatorium buniifolium Hook. et Arn.
Gamochaeta spicata (Lam.) Cabrera
Senecio brasiliensis (Spreng) Less. var. brasiliensis
Senecio cisplatinus Cabrera
Senecio oxyphyllus DC.
Stenachenium campestre Baker
Vernonia flexuosa Sims.
Vernonia nudiflora Less.
Cyperaceae
Carex bonariensis Desf. ex Poir.
Carex phalaroides Kunth
Carex sororia Kunth
Cyperus luzulae (L.) Retz
Eleocharis bonariensis Nees
Eleocharis dunensis Kuk.a
Eleocharis sellowiana Kunth
Kyllinga brevifolia Rottb.
Pycreus polystachyos (Rottb.) P. Beauv.
Rhynchospora holoschoenoides (Rich.) Herter
Rhynchospora megapotamica (A. Spreng.) H. Pfeiff.
Fabaceae
Adesmia araujoi Burkarta
Adesmia bicolor (Poir.) DC.a
Adesmia latifolia (Spreng.) Vogel
Arachis burkartii Handroa
Clitoria nana Benth.
Desmodium incanum DC.
Lathyrus pubescens Hook.& Arn.
Macroptilium prostratum (Benth.) Urb.
Rhynchosia diversifolia M. Micheli
Stylosanthes leiocarpa Vog.
Trifolium polymorphum Poir.a
Hypoxidaceae
Hypoxis decumbens L.
Iridaceae
Herbertia pulchella Sweet
Sisyrinchium micranthum Cav.
a
Juncaceae
Juncus capillaceus Lam.
Juncus microcephalus Kunth
Oxalidaceae
Oxalis articulata Savigny
Oxalis eriocarpa DC.
Oxalis perdicaria (Molina) Bertero
Poaceae
Andropogon lateralis Nees
Andropogon selloanus (Hack.) Hack.
Andropogon ternatus (Spreng.) Nees
Aristida filifolia (Arechav.) Herter
Aristida jubata Arech.
Aristida laevis (Nees) Kunth
Aristida spegazzinii Arech.
Axonopus affinis Chase
Bothriochloa laguroides (DC.) Herter
Bouteloua megapotamica (Spreng.) O. Kuntzea
Briza subaristata Lam.
Coelorachis selloana (Hack.) Camus
Danthonia secundiflora Presl
Dichanthelium sabulorum (Lam.) Gould & C.A. Clark
Elyonurus candidus (Trin.) Hack.
Ischaemum minus J. Presl
Melica eremophila M.A. Torres
Melica rigida Cav.a
Panicum aquaticum Poir.
Paspalum dilatatum Poir.
Paspalum nicorae Parodi
Paspalum notatum Fl.
Paspalum pauciciliatum (Parodi) Herter
Paspalum pumilum Nees
Piptochaetium lasianthum Griseb.
Piptochaetium ruprechtianum Desv.
Piptochaetium stipoides (Trin. & Rupr.) Hack.
Saccharum trinii (Hack.) Renvoize
Stipa filifolia Neesa
Stipa megapotamia Spreng. ex Trin.
Stipa nutans Hack.
Stipa philippii Steud.a
Stipa setigera C. Presl
Rubiaceae
Borreria verticillata (L.) G.F.W. Meyer
Richardia humistrata (Cham. et Schlecht.) Steud.
Verbenaceae
Glandularia subincana Tronc.
Lippia asperrima Cham.
Phylla canescens (H.B.K.) Greene
Endemic species.
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Table 3. Diversity and vegetation structure in relation to time since last burn in 0.25 m2 grassland plots in Porto Alegre, RS, Brazil
(data from Overbeck et al., 2005)
Time since last re
Diversity (Shannon)
Number of species
Open soil (% cover)
Litter (% cover)
Standing dead biomass (% cover)
3 months
1 year
2 years
3 years
2.72 a
28 a
46.6 a
3.2 a
6.7 a
2.4 b
22.50 b
31.2 b
7.8 b
8.2 a
2.43 b
21.75 b
5.1 c
13.1 c
18 b
1.84 c
15.07 c
1d
37.2 d
28 c
Columns indicate different time since the last burn (3 months, 1 year, 2 years, 3 years or more). For each variable (row), different letters behind data
indicate signicant differences between plots with different time since the last re (po0.05), tested by randomization testing with analysis of variance.
Forestgrassland dynamics
In the absence of re and grazing, grasslands are
subject to shrub encroachment and, when in the vicinity
of forest vegetation, to forest expansion (Machado,
2004; Oliveira and Pillar, 2004; Muller et al., 2006); this
has been found for the South Brazilian plateau and for
the Central Depression, but there have been no studies
from the southern part of Rio Grande do Sul. Similar
results were found by Safford (2001) for highland
grasslands of southeastern Brazil. Increased density of
shrubs and trees in grassland and savanna vegetation
has been observed throughout the world in the last three
decades (e.g. Archer, 1990; van Auken, 2000; Roques
et al., 2001; Cabral et al., 2003), with different
hypotheses being proposed to explain these patterns
(such as climatic shifts or elevated global CO2 levels; e.g.
Longman and Jen k, 1992; Bond and Midgley, 2000;
Sternberg, 2001). As the climate in southern Brazil is
conductive to the development of forests, changes in the
disturbance regime, especially grazing and re regimes,
should be decisive factors for vegetation changes at
forestgrassland boundaries (Pillar and Quadros, 1997;
Scholes and Archer, 1997; Pillar, 2003; Langevelde et al.,
2003; Bond, 2005). Modelling of vegetation development in South Africa has shown that areas above a
certain precipitation limit (650 mm for South Africa)
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26
24
22
20
18
16
14
12
10
8
6
4
2
0
24
22
20
18
16
14
12
10
8
6
4
2
0
44
40
36
(a)
last fire in grassland: >3 years
> 80cm
> 30cm to < 80cm
< 30cm
(b)
last fire in grassland: 2 years ago
(c)
last fire in grassland: 1 year ago
32
28
24
20
16
12
presence
of rock
outcrops
8
4
0
31 28 25 22 19 16 13 10
grassland
7
-
4
border
2
-
11 14 17 20 23 26
forest (m)
Fig. 2. Number of tree individuals along the forestgrassland gradient according to size class intervals (height) in a natural
forestgrassland mosaic under the inuence of re in South Brazil (data from Muller, 2005). Grassland areas in (a) have not burned
for more than 3 years, in (b) were burned 2 years ago and in (c) 1 year ago. Please note different scales on the y-axes.
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Campos biodiversity
Information on plant biodiversity of the Campos is far
from complete. Boldrini (1997) estimated a total number
of 3000 grassland plant species in Rio Grande do Sul
state alone, and Klein (1975, 1984) estimated approximately 4000 species. While this is lower than the number
proposed for Brazils Cerrado region (6000 vascular
species; Furley, 1999), it should be noted that Cerrado
(total area 2 million km2) covers a much larger area than
South Brazilian Campos, and therefore also includes a
wider range of climatic and edaphic conditions (Furley,
1999) than the comparatively uniform Campos region
(Ministerio do Meio Ambiente (MMA), 2000); thus it
also includes a greater diversity of vegetation types from
grassland to forest physiognomies (e.g. Oliveira-Filho
and Ratter, 2002 for an overview).
The federal governments Project of Conservation
and Sustainable Use of Brazilian Diversity (PROBIO;
MMA, 1996) sponsored workshops that identied
approximately 900 priority areas to be conserved
throughout the country (MMA, 2002; Silva, 2005) and
led to oristic and faunistic inventories in areas
previously unstudied in southern Brazil as well. Grasslands on the South Brazilian plateau, i.e. grasslands
within the Atlantic forest biome (totalling 1,374,000 ha,
i.e. 1/10 of total Campos area), in Rio Grande do Sul
and Santa Catarina where included in this project: 1082
species, 95 of them endemics and 35 endangered by
extinction, could be listed in the course of these studies,
taking into account the available literature, herbarium
records and eld investigations (Boldrini, 2007). No
exhaustive compilations exists for grasslands in the
southern or more western parts of Rio Grande do Sul.
Despite these recent advances, the South Brazilian
grasslands remain one of the regions for which large
portions are still insufciently known (Giulietti et al.,
2005). A thorough analysis of the ora of the entire
grassland region is still not possible (as is probably the
case in other Brazilian biomes as well), but some broad
patterns are clear. The most species-rich plant families
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Cultivated pastures
The intensication of farming systems has led to
increases in the area of cultivated pastures. Despite the
high productivity and forage potential of many native
species, they are not commercially exploited, and
cultivated pastures are mainly formed using exotic
species (Nabinger et al., 2000). In 1996, 7.0 million ha
in the southern region of Brazil were covered by sown
pastures, mainly with non-native species. Important
cultivated grass species are e.g. Axonopus jesuiticus
(Araujo) Valls, P. notatum var. saurae Parodi, both
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111
26S
30
54
50W
Fig. 3. Levels of representativeness (gap status levels) in conservation units larger than 1,000 ha of the different phytoecological
regions in Rio Grande do Sul state. The levels range from 0% (1) to 7.1% (5). The ranges of each level of representativeness are not
equal neither are levels continuous. See text for exact values for the different vegetation types.
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Acknowledgements
The research leading to this paper was part of cooperation projects supported by CAPES (Brazil), the
German Academic Exchange Service (DAAD), the
German Research Foundation (DFG) and the State of
Bavaria (Germany). V.P. received support from CNPq
(Brazil). Our thanks go to Peter J. Edwards and to
Catherine Burns for checking the manuscript.
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