ARTICLE IN PRESS
Flora 202 (2007) 27–49
www.elsevier.de/flora
Adaptive strategies in burned subtropical grassland in southern Brazil
Gerhard Ernst Overbeck, Jörg Pfadenhauer
Technische Universität München, An Hochanger 6, 85350 Freising-Weihenstephan, Germany
Received 17 May 2005; accepted 2 November 2005
Abstract
Extensive parts of subtropical South America are covered by grassland vegetation, despite climatic conditions that
allow for forest development, and fire may have been an important factor in the evolutionary history of these
grasslands. In a regularly burned grassland area, situated in a forest–grassland-mosaic near Porto Alegre, RS, Brazil,
life form spectrum and plant species’ reaction to fire were examined, allowing for (1) a physiognomic description of the
grassland, and (2) a functional classification of grassland species in relation to fire. Grassland sites with different time
since the last fire occurred were compared between each other as well as to sites at the forest–grassland border. South
Brazilian grassland is dominated by hemicryptophytic caespitose graminoids that resist fires, but contains a large
number of geophytic or hemicryptophytic forbs, in general sprouting after fire. Shrubs, mostly sprouting species of the
grassland community, were present with high cover values even in recently burned areas. In contrast to Central
Brazilian Cerrado, trees were of little importance: most species found were forest pioneer species without the capacity
to survive fires unless growing on sites protected from fire or at the forest border where burns stop. Non-sprouting
species were of little importance in the community, and only two species found were therophytes. Lack of therophytes
in South Brazilian grassland vegetation deserves further attention.
r 2006 Elsevier GmbH. All rights reserved.
Keywords: Campos; Fire; Life form; Plant functional type; Raunkiaer
Introduction
While savanna ecosystems cover large parts of
tropical South America (e.g., Sarmiento 1990 for an
overview), wide parts of northern Argentina and
Uruguay are characterized by grasslands, extending
around the Rı́o de la Plata and continuing in southern
Brazil (Bilenca and Miñarro 2004; Soriano et al. 1992;
Fig. 1). In southern Brazil, natural grasslands (‘‘Campos’’) can be found in the southermost state Rio Grande
do Sul (RS; Fig. 2) and in the adjacent states of Santa
Catarina (SC) and Paraná (PR). The total extension of
Corresponding author. Fax: +09 8161 714143.
E-mail address: overbeck@wzw.tum.de (G.E. Overbeck).
0367-2530/$ - see front matter r 2006 Elsevier GmbH. All rights reserved.
doi:10.1016/j.flora.2005.11.004
the grasslands in those three states has been reduced
from originally 218.700 km2 (Longhi-Wagner 2003) to
today appoximately 136.800 km2 (Nabinger et al. 2000),
principally due to expansion of agri- and silvicultural
production. South Brazilian Campos grasslands, presenting a mixture of C4 and C3 grasses with C4
dominance, are under subtropical humid climate (Köppen’s Cfa) in southern RS state, and under warmtemperate humid climate on the highlands in northern
RS and adjacent SC (Köppen’s Cfb; Moreno 1961), in
the latter climatic region in mosaics with Araucaria
forests (see Fig. 1). Precipitation in southern Brazil
ranges from 1200 (southern RS) to over 2200 mm year1
(highest part of the planalto in northern RS), with no
dry season. Hydric deficits may occasionally occur in the
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G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
BRAZIL
PARAGUAY
25° S
30° S
URUGUAY
ARGENTINA
Porto
Alegre
35° S
Río de la
Plata
40° S
0
500
1.000 km
Fig. 1. Location of temperate and subtropical grasslands in
southern South America, between 25 and 381S (both shades of
gray). Grasslands south of the Rı́o de la Plata commonly are
considered to be Pampas grassland; those north of the Rı́o de
la Plata; Campos. Campos grasslands in Brazil depicted in
darker shade gray (sources: Soriano et al. 1992 for Argentina
and Uruguay; Leite 2002 for distribution of grasslands in
Brazil, modified).
Location of the
study site in
Porto Alegre
Atlantic forest
Semi-deciduous forest
Araucaria forest
Grassland
Coastal vegetation
Transitional vegetation on the
southeastern shield (mosaics of
grassland, shrublands and forests)
0
100
200
Fig. 2. Vegetation types in Brazil’s southernmost state, Rio
Grande do Sul (from Pillar and Quadros 1997, modified).
summer months, but current climatic conditions allow
for forest development (Lindman 1906; Rambo 1956).
Already Lindman (1906) had noted the contradiction
between the presence of grassland vegetation in southern
Brazil and humid climatic conditions that allow for
forest development. Similarly, the presence of grasslands
in a climate apparently supporting forest vegetation has
led to intense debate of the so-called ‘‘Pampas problem’’
in the Rı́o de la Plata region (e.g., Box 1986; Eriksen
1978; Walter 1967). Discussion, however, focused on
relations between climate and vegetation in a rather
static view, neglecting vegetation history and the
possibility of rather recent climatic changes. Phytogeographical and paleoecological data suggest that grasslands in southern Brazil are relicts of drier and cooler
conditions during the last glacial and post-glacial
periods, stabilized by herbivory and fire and subject to
forest invasion only relatively recently (Behling 2002,
2004; Bigarella 1971; Klein 1975; Pillar and Quadros
1997; Rambo 1953, 1956). In contrast to South African
savannas and grasslands, large native herbivores are
largely missing: they became extinct as the result of
climatic changes between the end of the last glaciation
and 8000 years BP (Kern 1994), coninciding with the
arrival of human populations in the region. The fire
history of southern Brazil is poorly known. Palynological studies indicate that fire was rare during early
post-glacial times, but became frequent about 7400
years ago (Behling et al. 2004), possibly as it may have
been used as a tool for hunting by indigeneous people
(Kern 1994). In Cerrado (Central Brazilian savannas),
fire is considered to have been present long before
human occupation, but frequency most probably has
increased with anthropogenic use (Hoffmann and
Moreira 2002; Miranda et al. 2002); the effect has been
a stabilization of open vegetation formations by
impeding establishment of forest species, and thus a
change in physignomy (Hoffmann 1996, 2000; Mirelles
et al. 1997). Recent modelling of the distribution of
vegetation types in southern Africa in relation to
climatic parameters and fire has shown that in regions
above a certain limit of precipitation (approx. 650 mm
for southern Africa), not climatic conditions, but
presence of fire determines vegetation physiognomy,
independent of historic human action (Bond et al. 2003).
While less data are available for southern Brazil, the
situation likely is similar.
Few studies have been conducted on the effect of fire
on grassland communities in southern Brazil. Single fire
events do not invoke major overall compositional
changes (Eggers and Porto 1994), and fire has been
shown to increase small-scale and short-term diversity
(Overbeck et al. 2005). Adaptations of South Brazilian
Campos species have not been studied explicitly so far. A
number of classification systems for plant adapations
exist, including some specifically related to fire. Noble
and Slatyer (1980) have presented a general model of
plant strategies in respect to fire, focusing on regeneration or colonization attributes; application of this model
to a community, however, requires detailed information
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G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
on life cycle, persistance and dispersal abilities of the
species present. Plants from fire-prone communities
commonly can be classified according to the two criteria
survival of fire and reproductive response to fire (Bond
and Van Wilgen 1996). For consideration primarily of
survival of fire, plants in fire-prone habitats have been
grouped into plants resisting fire with part of their
aboveground biomass (‘‘resisters’’, Rowe 1983), plants
dying back completely aboveground, but surviving due
to insulation of buds by the soil (‘‘endurers’’, Rowe
1983; ‘‘sprouters’’, Bond and Van Wilgen 1996) and into
plants having to germinate either from the seed bank
(‘‘evaders’’, Rowe 1983) or from newly dispersed seeds
(‘‘invaders’’, Rowe 1983). If recruitment occurs only
right after a fire event, non-sprouters have been termed
‘‘fire-recruiters’’ (as opposed to ‘‘fire-persisters’’; Keeley
1992). Similarily, chaparral species have been categorized into ‘‘obligate seeders’’, ‘‘obligate resprouters’’ and
‘‘facultative seeders/facultative resprouters’’ (e.g.,
Franklin et al. 2004; Keeley 1986, 1992). In poorly
known florae information on germination requirements
usually is unavailable for the majority of species, while
position and protection of regenerative buds can be
observed more readily. This is the principal criterion in
one of the classical functional type systems: Raunkiaer’s
life form spectrum (Raunkiaer 1934), applied for
prediction of climate in many ecosystems of the world
(see Cain 1950 for a compilation of life form spectra for
various climatic regions) and considered to be a
good predictor of disturbance (McIntyre et al. 1999).
The growth form system, an essentially synonymous
(Pillar and Orlóci 2004), but somewhat less strict
classification as not based on a clear criterion but on
general plant architecture, has been identified to be a
useful framework for classification into plant functional
types in various systems (e.g., Chapin et al. 1996;
Diaz and Cabido 1997; McIntyre et al. 1995). Raunkiaer’s life form system has been applied to fire-prone
systems (Batalha and Martins 2002; Chapman and
Crow 1981), but applicability of the system specifically
to fire has never been evaluated. In this paper,
we present the physiognomic composition of a grassland
community subject to frequent burns in southern Brazil,
and discuss functional attributes in relation to fire,
evaluating applicability of the classical life form
system and linking it to the main post-fire regeneration
mechanisms in an alternative classification. This
allows to describe the structure of the grasslands
via the physiognomic information given by Raunkiaer’s
system, while being able to discuss regeneration
responses in relation to fire. Further, we characterize
the principal plant types present in a brief, reviewlike way and compare the in general still poorly
studied South Brazilian Campos grassland with
other burned grassland systems concerning life form
composition.
29
Methods
Study site
Field work was carried out on Morro Santana, Porto
Alegre, RS, Brazil. Morro Santana (301030 S, 511070 W,
max. alt. 311 m a.s.l.), situated at the northern limit of
the Crystalline shield in RS, is covered by a mosaic of
forests (Atlantic forest) and grasslands. Floristic composition of the grassland area has been presented in
Overbeck et al. (2006a). The grasslands are species rich,
with a total of 430 species identified so far in an area of
about 220 ha, from a species pool of about 450–500
species. Borders between forest and grasslands have
been remarkably stable in the past decades, due to
frequent fires which prevent expansion of forest into
grassland. Isolated patches of shrubs and trees, considered nuclei of forest establishment, can be found
principally in areas with rock outcrops. Presence of fire,
today anthropogenic, throughout the past 1200 years
has been confirmed by a palynological study (Behling et
al., in press).
Data collection
Six pairs of transects (distance between transects in
each pair approximately 5 m) were installed from the
forest border into the grassland, in areas differing in
slope, aspect, degree of shrub encroachment and time
since the last fire. In October 2002, one transect in each
of the pairs was subjected to an experimental burn;
however, in two of the transects, situated in areas that
had burned almost 1 year before, the fire did not spread
due to lack of continuous biomass. This left us with four
transects burned in our experiments (group 1), four
transects burned a year before (group 2) and four
transects burned 3 or more years before (group 3). To be
able to sample both the herbaceous stratum and
characterize woody species distribution, we used a
stratified plot design. Each transect consisted of seven
contiguous large plots (LPs) of 4.5 m 4.5 m. The first
plot, situated directly at the forest–grassland border, did
not burn in any of the transects. Within LPs, three
contiguous medium-sized plots (MPs) of 1.5 m 1.5 m
were marked, totalling 21 contiguous plots from the
border into the grassland for each transect. In LPs, all
woody vegetation components (trees and shrubs) with
height above 80 cm were recorded (number of individuals per species), whereas in MPs, cover of all woody
individuals above 10 cm was recorded, using the Londo
(1976) decimal scale. Floristic composition of the
herbaceous layer was sampled in three small contiguous
plots (SPs) of 50 cm 50 cm, marked in the center
of LPs 1, 3, 5 and 7 (1 being closest, 7 farthest to the
forest border), using the Londo (1976) decimal scale.
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G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Additionally, cover of litter, rocks, mosses, open soil
and total cover of standing dead biomass were recorded
in SPs, while in MPs and LPs, total cover of herbs,
grasses (divided into living and dead), lianas, rocks,
mosses, litter and open soil and medium height of the
herbaceous stratum recorded. All surveys were conducted 2–3 months after the experimental fire. Additional structural surveys in MPs and LPs had also been
conducted directly before, and, in burned plots, 1 week
after the experimental burns.
Data analysis
Data from the three contiguous SPs was pooled to
one plot of 0.75 m2 by taking the mean value across
plots for each species or structural category. Species
were grouped into life forms according to Raunkiaer’s
(1934) system, with a modified classification into
subgroups (Table 1). Plants were grouped a posteriori,
i.e. according to potential life form of the species.
Instead of grouping woody species into phanerophytes
and chamaephytes as in Raunkiaer’s system, they were
categorized into the growth form categories shrubs and
trees, considered to reflect differences in plant carbon
allocation strategy (Bond and Midgley 2000; see also
Sarmiento and Monasterio 1983), thus avoiding the
necessity to fix a height limit between the two groups,
which would be arbitrary in subtropical vegetation.
We then modified this system according to survival
capacity to fire, obtaining the classification into fire life
forms presented in Table 3. In this new classification,
position of the buds (Raunkiaer’s criterion) is joined by
information on whether the plant dies (non-sprouter),
looses its aboveground biomass but can recover from
belowground organs (sprouter) or manages to keep at
least some aboveground biomass (resister) during a fire.
No evidence of stimulation of heat germination by fire
(as known for many hardseeded shrubs in mediterranean climate regions, e.g. Keeley and Fotheringham
Table 1.
2000) exists for the studied region (Overbeck et al.,
2006b); thus, it was not necessary (nor possible due to
information limited to a few species only) to include an
attribute for this in the classification. For both
Raunkiaer’s system and the fire life form classification,
individual species cover values were summed up to the
corresponding life form category and then the relative
cover of each category in relation to total vegetation
cover was calculated, allowing for comparison of areas
contrasting greatly in total vegetation cover in relation
to time since the last burn. Species that had not been
identified to the species level and could not be safely
assigned to any life form group were considered as part
of total vegetation cover in calculation of relative cover
values of each life form group, but otherwise left out
from the analyses (0.58% of sum of all frequencies). The
three plot groups representing different time since the
last fire were compared for differences between performance of each life form and fire life form, using
univariate analysis of variance with randomization
testing (MULTIV software, Pillar 2004; Euclidean
distance as resemblance measure, a ¼ 0:05 as probability limit for rejection of null-hypothesis). Cover of
standing dead biomass was treated the same way.
Further, border plots were compared to grassland plots
taken as a group. In addition to the vegetation life form
spectrum based on relative cover values, a floristic life
form spectrum based on species number per life form
group was elaborated.
For MP and LP data, species were grouped into
shrubs and trees (growth forms), into species commonly
known as grassland species, forest border species, or as
forest species (successional groups) and into species with
or without sprouting capacity after fire (regenerative
types). Total number of individuals (LPs) and total
cover (MPs) per plot for each of these groups was
compared between grassland plot treatment groups by
univariate analysis of variance with randomization
testing, and between border plots and grassland plots
Raunkiaer’s life forms and sub-types used in this study
Main type
Sub-type
Criteria
Therophytes
—
Geophytes
Bulbous geophytes
Rhizomateous geophytes
Forbs
Caespitose graminoids
No perennating buds; plants complete life cycle in 1 year; only seeds as
perennating structures
Bulbs or corms as perennating organs, situated below the soil surface
Rhizomes or tubers as perennating organs, situated below the soil surface
Forbs with meristems at the soil surface
Caespitose graminoids with meristems more or less at the soil surface,
usually in densely packed tussocks
In general multi-stemmed growth; according to common local denomination
of species
In general single-stemmed growth; according to common local
denomination of species
Herbaceous or woody plants without self-supporting structures
Hemicryptophytes
Shrubs
—
Trees
—
Lianas
—
Modified from Raunkiaer (1934).
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G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
taken as a group (Euclidean distance; a ¼ 0:05). Spearman rank correlations were calculated between total
number (LPs and MPs) or total cover (MPs only) of
woody individuals and the variables describing vegetation structure for all grassland plots to test if the woody
species component in the grassland was related to
structural factors of the community, such as density of
the herbaceous layer. For this, data from the survey
right before (unburned plots) and right after the burns
(burned plots) and from the survey conducted 2–3
months after the burns were used.
Results
Raunkiaer’s life form and fire life form spectra for
the community: data from SPs
When comparing relative cover of the different life
forms in the treatment groups (grassland plots with
different time since fire), few significant differences
could be found for any of the life form categories. Over
all treatment groups, hemicryptophytes amounted to
67.3% of relative cover (caespitose grasses alone to
50.1%), geophytes to 16.3% and shrubs to 15.1% (mean
values). Therophytes barely contributed to total cover
(Table 2A). Trees and lianas showed low cover values.
In border plots, relative cover of caespitose grasses was
significantly lower compared to grassland plots as a
group, in contrast to trees and lianas for which the
opposite was true. Cover of trees, but not shrubs, was
significantly higher at the border.
Concerning number of species in each life form group,
hemicryptophytic forbs were the most important,
followed by caespitose grasses, shrubs and rhizomateous
geophytes, with little differences between treatment
Table 2.
31
groups (Table 2B). Species numbers for the different
life-form groups did not differ much at the forest
border, except for liana and tree species. Therophytes
were almost completely absent.
Standing dead biomass differed significantly between
the three treatment groups (po0:001), with mean values
of 2.5% in recently burned SPs (group1), 9.8% in SPs of
group 2 and 23.2% in SPs of group 3. Standing dead
biomass almost completely could be contributed to
species from the Poaceae or Cyperaceae. In the border
plots (all unburned since 3 years or more), standing dead
biomass had a mean cover of 10.4%.
In the fire life form classification, sprouting species
summed up to 55.5% of relative cover in recently
burned plots (Table 3). Their contribution, however,
was markedly lower in plots of group 2, and relative
cover of hemicryptophytic sprouters differed significantly between groups 1 and 3. While cover of sprouting
forbs and shrubs thus decreased with time since fire,
relative cover of caespitose grasses, exclusively making
up the group of resisters in grassland plots, showed an
increase with time since fire. Non-sprouting species
contributed less than 10% of relative cover in all groups
of grassland plots. Border plots had significantly higher
relative cover of non-sprouting trees, sprouting trees and
lianas, and significantly less of caespitose grasses, which
contributed to less than 1/3 of relative cover at the forest
border.
Characterization of the woody stratum: data from
MPs and LPs
Cover values for shrub and tree species was higher in
MPs than in SPs, most likely due to the very small total
area size sampled by SPs. This under-representation of
shrubs and trees in the SPs has to be kept in mind when
Vegetation (A) and floristic (B) spectrum of Raunkiaer’s life forms (n ¼ 12 for grassland groups and border plots)
Therophytes
Geophytes*
Bulbous
Rhizomateous
Hemicryptophytes
Caespitose graminoids
Forbs
Shrubs
Trees
Lianas
A: Relative cover values (vegetation spectrum)
B: Percentage of species (floristic spectrum)
Group 1
Group 2
Group 3
Border
Group 1
Group 2
Group 3
Border
0.2%
19.6% a
0.7%
18.9%
62.1%
42.0%
20.1%
15.7%
2.1%
0.2%
0.0%
12.2% b
0.1%
12.1%
70.2%
51.0%
19.2%
17.2%
0.0%
0.0%
0.1%
17.0% ab
0.3%
16.7%
69.5%
57.2%
12.4%
12.3%
0.2%
0.7%
0.0%
11.9%
0.0%
11.8%
44.3%
31.2%
13.1%
9.8%
7.9%
25.8%
0.9%
19.3%
4.4%
14.9%
58.8%
25.4%
33.3%
17.5%
1.8%
1.8%
0.0%
17.8%
2.5%
15.3%
62.7%
28.0%
34.7%
19.5%
0.0%
0.0%
1.6%
14.8%
3.1%
11.7%
55.5%
23.4%
32.0%
22.7%
2.3%
3.1%
0.0%
14.0%
0.8%
13.1%
50.8%
20.5%
30.3%
16.4%
12.3%
6.6%
For the vegetation composition (based on relative cover), plots from group 1 and 2 differed significantly for geophytes (indicated by an asterisk in the
table; po0:05; different letters behind numbers indicate significant differences between groups for geophytes). Relative cover of all hemicryptophytes,
caespitose grasses, trees and lianas differed between border plots and all grassland plots taken as a group (indiated by italics in the table; po0:001).
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Table 3.
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Fire life form distribution in frequently burned grassland on Morro Santana, Porto Alegre, RS, Brazil
Relative cover values
Non-sproutersa
Therophytes
Hemicryptophytes
Shrubs
Trees
Sprouters (from belowground organs)
Geophytes
bulbous
rhizomateous
Hemicryptophytic forbs*
Shrubs
Trees
Lianas
Resisters (aboveground survival)b
Caespitose graminoids (hemicryptophytic)
Shrubs
Trees
Border
Gr. 1
Gr. 2
Gr. 3
2.4%
0.2%
1.3%
0.9%
0.0%
55.5%
19.6%
0.6%
18.9%
18.8% a
14.8%
2.0%
0.2%
42.0%
42.0%
0.0%
0.0%
9.0%
0.0%
6.1%
2.9%
0.0%
39.7%
12.2%
0.1%
12.1%
13.2% ab
14.3%
0.0%
0.0%
51.0%
51.0%
0.0%
0.0%
6.9%
0.8%
1.6%
4.3%
0.2%
36.5%
17.0%
0.3%
16.7%
10.9% b
8.0%
0.0%
0.7%
57.1%
57.1%
0.0%
0.0%
6.5%
0.0%
1.7%
0.7%
4.1%
63.0%
11.9%
0.0%
11.8%
12.6%
9.1%
3.7%
25.7%
31.3%
31.2%
0.0%
0.1%
Percentage of species
All Campos plots
Border plots
7.7%
1.1%
2.2%
2.8%
1.7%
67.9%
17.7%
3.3%
14.4%
30.9%
15.5%
1.1%
2.8%
24.3%
24.3%
0.0%
0.0%
14.5%
0.0%
4.1%
2.5%
8.2%
63.9%
13.9%
0.8%
13.1%
26.2%
13.9%
3.3%
6.6%
21.3%
20.5%
0.0%
0.8%
Sprouting hemicryptophytic forbs differed in relative cover value between groups of grassland plots (indicated by an asterisk; different letters behind
numbers indicate significant differences between groups for hemicryptophytic forbs). Groups printed in italics differed significantly between
grassland plots as a group and border plots concering relative cover value (po0:05).
a
Per definition, no non-sprouting geophytic species exist. See discussion for classification of caespitose grasses.
b
No resisting geophytes, hemicryptic forbs or lianas could be found in the study.
interpreting the data on the community level and
underlines the importance of a nested design when
working in communities with distinct herbaceous and
woody strata, where high richness of the herbaceaous
layer impedes working with larger plot size. Using the
MP data, in average, 5.4 individuals of shrubs and 0.1 of
trees could be found per m2 in the grassland. In all, 38 of
the species (25 shrubs, 13 trees) were sprouters and 21
non-sprouters (9 shrubs and 12 trees). Sprouters showed
substantially higher cover values than non-sprouters
both in grassland and at the border, however, more
pronounced in the grassland and relatively more in
recently burned areas. In general, cover of shrubs and
trees rose with time since fire (Table 4A). Tree species,
border species or forest species were of little overall
importance in the grassland’s woody species community, but present throughout the area, whereas grassland
shrubs clearly were the most important component of
the grassland. The only resisters were Leucothoe
eucalyptoides (Ericaceae), Butia capitata (Araceae), both
with thick bark, and Opuntia macrocantha (Cactaceae),
where at least part of the plant survives due to its high
moisture content.
In grassland LPs, a mean of 0.23 woody plants higher
than 80 cm could be found per m2, 0.21 of these being
shrubs and 0.02 trees (Table 4B). Number of woody
species was significantly higher (po0:001) in plots of
group 3, where 0.48 woody individuals above 80 cm
were present per m2, in comparsion to 0.05 and 0.14 in
treatment groups 1 and 2, respectively. The largest part
of woody species in the grassland were grassland species,
in contrast to the border, where species from the forest
or from the border were of importance as well, with in
general higher numbers of trees and shrubs from all
successional groups (Table 4B).
No strong correlations (with a correlation coefficient
above 0.4 or below 0.4) between number or cover of
woody species in LPs and MPs and structure of the
herbaceous stratum (cover of dead and live grasses,
herbs, lianas, litter, open soil, rocks, height of grass
layer) was detected, for either sampling data of
vegetation structure attributes.
Discussion
Applicability of Raunkiaer’s life form system to fire
Application of Raunkiaer’s system to tropical ecosystems has been criticized (Sarmiento and Monasterio
1983; Solbrig 1993) because many species cannot readily
be classified into the categories, as growth patterns may
undergo changes from one class to another, depending
on events such as heavy drought and fire. Trees, for
example, may resprout like shrubs, recovering their
ARTICLE IN PRESS
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
33
Table 4. Cover and number of shrubs and trees per m2, using data from (A) 232 plots of 1.5 m 1.5 m (cover values; all woody
species 410 cm) and (B) 84 plots of 4.5 m 4.5 m (number of individuals per m2, all woody species480 cm) on Morro Santana,
Porto Alegre, RS, Brazil
Total
Growth forms
Trees
Shrubs
Successional groups
Grassland species
Border/pioneer
species
Forest species
Regenerative types
Non-sprouters
Resprouters
Resisters
(A) MPs—cover (all individuals410 cm)
(B) LPs—number per m2 (all individuals480 cm)
(A1) Grassland with different
time since fire (Groups 1, 2, 3)
(A2) Open grassland
(Camp.) and border
(Bord.)
(B1) Grassland with
different time since fire
(Groups 1, 2, 3)
(B2) Open
grassland and
border
Gr. 1
Gr. 2
Gr. 3
Camp.
Bord.
Gr. 1
Gr. 2
Gr. 3
Camp.
Bord.
20.3% b
29.3% a
35.8% a
28.4% b
45.2% a
0.05 b
0.14 b
0.48 a
0.23 b
0.65 a
1.8%
18.5% b
0.8%
28.5% a
2.9%
32.9% a
1.8% b
26.6%
21.9% a
23.3%
0.02
0.03 b
0.02
0.12 b
0.02
0.46 a
0.02 b
0.21
0.32 a
0.34
18.5% b
0.4%
28.0% a
0.5%
29.6% a
3.9%
25.3% a
1.6% b
14.3% b
10.3% a
0.03 b
0.00
0.12 b
0.00
0.43 a
0.03
0.19
0.01 b
0.21
0.14 a
1.4%
0.8%
2.3%
1.5% b
20.6% a
0.02
0.02
0.02
0.02 b
0.31 a
1.1% c
18.8% b
3.7% b
25.6% a
5.6% b
22.5%
15.1% a
27.4%
0.02
0.03 b
0.00
0.14 a
0.20 a
0.28 a
0.07 b
0.15 b
0.33 a
0.30 a
0.4%
0.0%
12.2% a
23.0%
ab
0.6%
0.3%
2.7%
0.00
0.00
0.00
0.00 b
0.03 a
Values presented comparing groups of grassland plots with different time since fire (A1, B1) and grassland plots as a group with border plots (A2,
B2). Different letters indicate significant differences between groups (po0:05), to be read per line separately for A1, A2, B1, B2.
biomass by ‘‘crowns on the ground level’’ (Bellingham
and Sparrow 2000). Further, the height limit of
25–50 cm between chamaephytes and phanerophytes
(Ellenberg and Mueller-Dombois 1966) is a consequence
of the height of temperate clima dwarf shrubs and
cannot be applied in tropical or subtropical systems,
where chamaephytes may grow higher: here, division
into growth forms (shrubs and trees), i.e. a categorization based on plant architecture, seems to be more useful
(Sarmiento and Monasterio 1983). The life form system
had originally been elaborated to characterize plant
strategies in face of an infavorable season, and not as a
reaction to disturbances, which shape, for example,
tropical savannas or subtropical grasslands. In systems
with recurrent disturbances, position of buds is not
solely a consequence of climatic conditions, but
influenced by other environmental factors, e.g. the
action of herbivory or fire (Cain 1950). The life form
system therefore is useful for description of vegetation
physiognomy, but does not necessarily serve for prediction of climatic conditions, as originally indended by
Raunkiaer.
Fire life forms
In vegetation subject to frequent burns, survival and
sprouting ability depend on the degree of bud protection: a chamaephyte or phanerophyte may have to
sprout from belowground buds after the fire, or a
hemicryptophyte may not be able to sprout at all.
Between growth or life forms, traits allowing for
survival of fire vary. Resistance in woody plants is
principally achieved by protective tissues, e.g. thick bark
with low heat conductivity (Bond and Van Wilgen 1996;
Gill 1981), often joined by elevation of buds above the
flammable vegetation layer (e.g., Givnish et al. 1986). In
tussock grasses, densely packed basal leaf sheats allow
for protection of aboveground meristems from fire (e.g.,
Gill 1981; Sarmiento 1992). Sprouting ability from
belowground organs is usually a consequence of buds
protected by the soil, where temperature effects on plant
tissue stay below values causing damage (451–55 1C for
most plants; Levitt 1980): higher temperatures are only
reached in the uppermost centimeter of soil (e.g., Auld
and Bradstock 1996; Bradstock and Auld 1995; Miranda et al. 1993; Silva et al. 1991).
In the studied grassland, very few species manage to
keep aboveground live biomass through a fire. However,
meristems of tussock grasses, making up more than 50%
of plant cover 1 year after a burn, remained alive slightly
above the soil surface; species from this group thus have
to be considered as resisters, even though their behavior
does not differ much from hemicryptophytes sprouting
from buds at the soil surface. Individuals of Eryngium
horridum (Apiaceae), a large rhizomateous rosette
species with fibrous leaves and high water content,
sometimes did not burn completely and managed to
ARTICLE IN PRESS
34
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
keep some live biomass at the center of the leaf rosettes,
however, regaining aboveground biomass after the fire
through sprouting of new rosettes (Fidelis et al.
submitted), i.e. essentially behaving like a sprouting
species. The cactus species O. macrocantha and Pavonia
ottonis, sparsely present in the studied grassland, will
not burn due to their high water content, but, if fire is
intense enough, aboveground biomass will die due to the
effect of fire temperature on plant tissue. It is difficult to
classify them into the fire life form categories, as
resprouting can occur from below- or aboveground
organs, depending on fire severity and amount of
lignified structures. In a study in subtropical grasslands
in Texas, Bunting et al. (1980) showed that 20–65% of
plants from three Opuntia species did not sprout after
fire, but that mortality rate rose to 60–80% in a period
of 4 years after fire, as a consequence of increased
herbivory and fungal or bacterial infections on firedamaged plants. Resisting capacity for these species thus
is clearly lower than that of the ‘‘true’’ resisters, in our
study only the tree L. eucalyptoides (Ericaceae) and the
palm B. capitata (Arecaceae).
A small percentage of species in the studied grassland
was not able to resprout, but depended on regeneration
from seeds after the disturbance (obligate seeders; Bond
and Van Wilgen 1996). This group of non-sprouters
comprised plants from all life forms, except for
geophytes. This latter group should be the most
successful of the fire life forms considering survival of
burns (Taylor 1978), explaining why frequent fires often
favour sprouting forbs, often geophytes, over grasses,
mostly hemicryptophytes (Daubenmire 1968; Rowe
1983). In hemicryptophytes, the large majority of the
species in our grassland plots either were resprouters
(53.9% of species) or resisters (only the caespitose
grasses, 42.3%). In the group of shrubs, however, 84.9%
of species in grassland plots showed the ability to
resprout, indicating that position of aboveground buds
alone is a poor predictor of fire survival for the studied
grassland. Both the application of Raunkiaer’s system
and of the fire life form classification showed little
overall changes between different treatment groups,
reflecting slight shifts in relative importance in different
post-fire stages due to differences in competitive ranking
(Suding 2001), but no prounced changes in plant types.
When classifying plant species into fire life form
categories, two things have to be kept in mind: fire
intensity may differ between fires and according to local
site characteristics, and resistance capacity in respect to
fire may not be the same for all life stages of plants (e.g.,
Bond and Van Wilgen 1996; Gill 1981). These two
factors, alone or in combination, in some cases may
allow for survival of aboveground biomass in individuals even without protected buds on protected sites (in
the study area, for example, between rock outcrops,
where no continuous grass layer exists to support a fire)
and for damage of even belowground plant parts in
others when in an early life-cycle phase or when
subjected to extremely severe fire event, e.g. on sites
with higher fuel availability (see example of Cactus
species above). Sensitivity of sprouting species to fire
will further depend on season of fire and thus on specific
phenology of plant active phases and carbonhydrate
storage (Bond 1997; Chidumayo 2006), requiring to
study the phenological cycles of the species in question
for a more detailled prediction of the effect of fire on a
grassland community (Loucks et al. 1985; Sarmiento
and Monasterio 1983; Towne and Owensby 1984).
Brief functional characterization of main species
groups in relation to fire
Non-sprouting species
No significant differences could be found for cover
values of non-sprouting species, i.e. obligate seeders,
between the treatment groups, suggesting that, at least
in their majority, they cannot be considered to be fire
followers as found in some shrubland or grassland
communities (e.g., Ghermandi et al. 2004; Keeley et al.
1981), but will need some time to regain their pre-fire
cover values through new seedling recruitment from the
seed bank or through seed dispersal (see also Overbeck
et al. 2005, 2006b). Fire annuals sensu Keeley et al.
(1981) were absent in the studied grassland. Within the
group of non-sprouters, a small number of hemicryptophytic species with lack of protected buds was present in
all treatment groups. Especially common were Dichondrea sericea and Evolvulus sericeus, two small creeping
herbs of similar habitus (Convolvulaceae) that also
appeared in great quantities in a study of the seed banks
in the grasslands on Morro Santana (Müller and
Overbeck, unpubl.). Of great importance considering
cover values, especially in treatment groups 2 and 3 with
more time since fire, were the shrubs Heterothalamus
psiadioides and Baccharis dracunculifolia, which do not
resprout, in contrast to a number of other Asteraceae
shrub species with very similar habitus. Especially H.
alienus showed very strong recruitment in some plots of
group 2, where it had been present with large cover
values before the last burn (field observation). As this
species also was very abundant in the seed bank in the
study area (unpubl. data), most probably the creation of
open soil by the fire event will have been responsible for
its regeneration. In contrast to chaparral (Christensen
1985; Keeley 1986), frequent fires apparently do not
eliminate seeders in subtropical grassland.
Graminoids
Caespitose grasses, in their vast majority tussock
grasses with the C4 photosynthetic pathway, together
with Cyperaceae species accounted for over 50% of
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G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
vegetation cover in the studied grassland. We did not
found any annual grasses, and only a small percentage
of grasses was stoloniferous or rhizomateous, in contrast
to grazed grasslands in the region (Boldrini and Eggers
1997; Rodrı́guez et al. 2003). The majority of the
caespitose grasses found in the study on Morro Santana
also occur in the Brazilian Cerrado, including most of
the dominant species (e.g., Andropogon lateralis, Leptochoryphium lanatum, Elyonurus muticus). Tussock
grasses present densely packed basal shoots that allow
for survival of fire despite aboveground location of
meristems (Gill 1981); young plants, however, may be
seriously affected by fire in their survival (e.g., Silva and
Castro 1989). Some tussock grasses depend on regular
burns on the long run: without periodic removal of
biomass through fire, shading by dead biomass inhibits
survival and tillering and higher humidity under the
litter may cause death and decay of underground plant
parts within a few years (e.g., Bond et al. 2003; Morgan
and Lunt 1999; Silva et al. 1990, 1991; Vogl 1974).
Increased photosynthetic activity after fire has been
documented for Andropogon gerardii in tallgrass prairie
(Knapp 1985), and increased growth rates after burning
are known for many caespitose grass species (Silva et al.
1991), e.g. for A. lateralis, one of the most dominant
grasses in southern Brazil (Trindade and Da Rocha
2001). Sexual reproduction is stimulated by burning in
many savanna grass species (Gill 1981; Sarmiento 1992).
Especially grasses from the Andropogoneae tribe, of
high importance in the studied grassland, are extremely
productive under suitable, i.e. sufficiently humid,
climate conditions, producing large amounts of badly
decomposing biomass (Bond et al. 2003), thus allowing
for yearly burns where dominant (see also Overbeck et
al. 2005).
Woody plants
Woody species in South Brazilian Campos differ
strinkingly from woody species both in African and
neotropical savanna. In these systems, trees, commonly
resisting fire because of protective bark, characterize
vegetation physiognomy (Sarmiento and Monasterio
1983 for an overview). Tree species found in grassland
communities in southern Brazil, in contrast, are pioneer
trees that usually do not tolerate fire; frequent fires thus
should impede recruitment of forest pioneer trees in
grassland and thus forest expansion (Müller et al.
unpubl.). The woody stratum in Campos grassland is
comprised principally of grassland shrubs, in their
majority sprouters, with species from the Asteraceae
having the greatest importance both in terms of species
number and cover values. Usually, these shrubs do not
exceed 2 m in height, and some of them remain within
the height of the grass layer. In the absence of
disturbance, the shrubs may form dense stands, often
as belts along the forest–grassland border (Oliveira and
35
Pillar 2004). However, it has not been studied so far if
and in which ways they facilitate establishment of forest
species and to what extent they suppress flammability,
thus possibly accelerating vegetation change to forest
when fire intervals become longer. In southern Brazil,
fires reduce the shrub component more or less to the
ground level, with rapid recovery of biomass after the
fire. Flowering and fructification for the majority of
grassland shrubs (e.g., sprouting Asteraceae) may take
place within 3 months after the fire (field obs.), for some
species even less (e.g., the resprouter Schinus weinmanniaefolius, Anacardiaceae: fruits two months after fire).
Distribution of shrubs in the system is rather patchy;
this may be due to differences in recruitment behavior
(populations of sprouting species should be spatially
more stable), differences in edaphic conditions or
different fire history.
From field observations and considering the lack of
significant correlations between the shrub/tree layer and
the herbaceous layer, we conclude that shrub density does
not become high enough as to impede growth of
caespitose grasses and forbs, at least in fire intervals of
less than 5 years. If recruitment of shrub species were
facilitated by fire (presence of open soil), an interaction
between fire regime (colonization sites) and presence of
shrubs in nearby unburned areas (seed sources) could
exist. For South Brazilian grasslands, studies adressing
functional characteristics, life history strategies and
population dynamics of woody species under different
disturbance regimes are necessary, especially to be able to
compare Campos grassland to Brazilian Cerrado or other
savanna systems. Under present climatic conditions and
without recurrent disturbances, South Brazilian grasslands will turn into shrublands, or into a mosaic of
grassland, shrublands, and forest islands, subsequently
leading to forest development (Müller and Forneck 2004;
Oliveira and Pillar 2004; Müller et al. 2006).
Resprouting forbs
High diversity of South Brazilian Campos is largely a
consequence to the high number of forb species (see e.g.,
Overbeck et al. 2006a), in their vast majority sprouters.
In other burned grasslands, e.g. North American
tallgrass prairie (Benson et al. 2004) or Australian
temperate grassland (Morgan 1999), resprouting has
been identified to be the prevalent strategy for post-fire
biomass recovery in forbs. High productivity of
caespitose grasses and accumulation of litter leads to
the loss of forbs due to dominance effect of graminoids
in an apparently relatively random process of thinning
out of herbaceous species irrespective of their habitus
(Overbeck et al. 2005). Litter, which will accumulate in
the absence of fire, grazing or mowing, in general is
considered to reduce forb density and richness in
grasslands around the world (e.g., Facelli and
Pickett 1991; Knapp and Seastedt 1986). In the studied
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G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
grassland, richness of (sprouting and non-sprouting)
forb species thus can be directly linked to the fire cycle
(Leach and Givnish 1996), even though these should not
be important in terms of flammability. Studies of
population biology and life history of herbaceous
species in southern Brazil are missing almost entirely
(but see Fidelis et al. submitted).
Life-form composition of Brazilian subtropical
grassland in a global perspective
High importance of shrubs and increased recruitment
of forest pioneer species in sites close to the forestborder (seed sources) demonstrates the suitability of the
Campos biome’s climate for woody vegetation types; in
fact, longer unburned grasslands in southern Brazil
resemble secondary succession in areas formerly covered
by forests (Reitz and Klein 1964).
Similarily to Campos grassland (this study; Boldrini
1993; Garcia et al. 2002; Rodrı́guez et al. 2003) a lack of
annual species has also been noted for Brazilian
Cerrado, in contrast to many other grasslands and
savannas worldwide (Batalha and Martins 2002; Sarmiento and Monasterio 1983). Bond et al. (2003) suggest
for subtropical Southern Africa that perennial sprouters
are characteristic of fire-dependent grasslands, while
annuals prevail in climatically defined grassland, i.e. in
regions where development of woody vegetation types is
impeded by insufficient moisture supply. Similiarily, in
Australian temperate lowland grasslands the forb
component is dominated by perennials in all but the
driest regions (Lunt and Morgan 2001). Most of the
Cerrado biome today is considered to be under climate
suitable for forest establishment, but – like in southern
Africa – fire, interacting with climatic and edaphic
factors, determines actual vegetation physionomy (Henriques and Hey 2002; Oliveira-Filho and Ratter 2002).
The same seems to be true for South Brazilian grassland.
In the subtropical Campos region, with a humid climate
without any dry season, no climatic restriction favors an
annual life cycle. On the other hand, in many environments, annual species are known to profit from
disturbance events (Belsky 1992; Grime 1977) and, in
many fire-prone grasslands, opportunistic post-fire
colonizers appear in early post-fire vegetation development (e.g., Laterra et al. 2003; Lunt and Morgan 2001;
Ramsay and Oxley 1986). For tallgrass prairie, some
studies have shown that contribution of annuals
increases with disturbance (fire, grazing) intensity (e.g.,
Collins 1987; Engle et al. 2000; Gibson et al. 1993), while
in other studies, therophytes were almost missing
(Towne and Owensby 1984); however, we do not know
of any study adressing life form composition on the
gradient from tallgrass to shortgrass prairie. In some
Australian temperate grasslands, exotic annuals dom-
inate the seed bank and the post-fire seedling flora,
which has been attributed to the paucity of native
annuals (Morgan 1998, 2001) and to long-term grazing
that led to a loss of native species in the seed bank (Lunt
1990). Exotic annuals could not be found in the grassland
studied by us, despite the low contribution of native
annuals. Does the ‘‘niche for annuals’’ exist in southern
Brazil? The lack of therophytes in South Brazilian
grassland likely needs to be addressed in a comparative
study of grasslands systems around the world, under
various climatic conditions and under different disturbance regimes. The life form spectrum is a simple, yet
meaningful and readily usable classification; however, at
present surprisingly little reliable data on life-form
spectra for grassland communities can be found in the
literature. If joined by regeneration attributes, like in our
study, interpretational value of Raunkiaer’s system
should be enhanced considerately, allowing for comparative analysis of the influence of climatic factors and
disturbance on composition and vegetation dynamics in
grasslands in a global perspective.
Acknowledgments
We would like to thank Sandra Cristina Müller and
Valério DePatta Pillar, Departamento de Ecologia,
UFRGS, Porto Alegre, RS, Brazil, for the good
cooperation in the project and for discussion. Our thanks
go to staff at the Departamento de Botânica, UFRGS,
for help with identifying plant species. G.O. received a
Ph.D. grant by the German National Academic Foundation (Studienstiftung des deutschen Volkes). The project
was supported by the DFG and DAAD (Germany) and
CAPES (Brazil) in a ProBral-cooperation.
Appendix A. Species recorded in 144 plots of
0.25 m2, pooled to 48 plots of 0.75 m2, on Morro
Santana, Porto Alegre, RS, Brazil
Given are family, species name, life form (LF;
modified after Raunkiaer (1934); th: therophyte, gb:
bulbous geophyte, gr: rhizomateous geophyte, h: hemicryptophyte (excluding caespitose graminoids), hc:
hemicryptophytic caespitose graminoid; s: shrub, t: tree,
l: liana) and regenerative type after fire (RT; r: resister, s:
sprouter, n: non-sprouter), mean cover value (C) and
frequency (F) of species per group of plots, separated for
grassland plots at the forest-grassland border (border,
n ¼ 12), and plots in the open grassland, grouped into
plots burned three months before (Group 1; n ¼ 12),
burned a year before (Group 2; n ¼ 12) and unburned
for 3 years or more (Group 3, n ¼ 12). No information
is given for species not totally identified (Table A1).
Table A1
Family
Acanthaceae
Amaranthaceae
Amaryllidaceae
Anacardiaceae
Apiaceae
RT
n
s
s
s
s
s
s
s
s
s
n
s
s
s
s
s
s
s
s
s
s
s
s
s
s
l
gr
h
h
h
h
h
h
s
h
h
s
h
s
s
n
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
Group 1
Group 2
Group 3
C
F
C
F
C
F
C
F
0.009
0.005
0.021
0
0.028
0.005
0
0.289
0.005
0.009
0.046
0.009
0.141
0
0
0.037
0.023
0
0
0.023
0.002
0
0.002
0
0.002
0
0
0.882
0.014
0.014
0.009
0
0
0
0
0.005
0.014
0
0
0
1
1
4
0
3
1
0
6
1
3
2
1
2
0
0
3
1
0
0
1
1
0
1
0
1
0
0
8
1
2
3
0
0
0
0
1
1
0
0
0
0
0
0.100
0
0.012
0
0
0.037
0.519
0.069
0.009
0
0
0
0.037
0.102
0
0
0
0.171
0.002
0.007
0
0.579
0
0.019
0.019
0
0
0
0.072
0.063
0
0.005
0.005
0
0.056
0
0.014
0.028
0
0
8
0
2
0
0
4
3
7
3
0
0
0
2
7
0
0
0
5
1
1
0
1
0
2
2
0
0
0
7
6
0
1
1
0
3
0
4
2
0
0
0.058
0.007
0.148
0
0.023
0.468
0
0.104
0.049
0
0
0.046
0
0.185
0.005
0
0
0.347
0.002
0.046
0.069
0
0
0.056
0.007
0
0
0.032
0.109
0.005
0.003
0
0.005
0
0.028
0.005
0.183
0
0
0
7
3
2
0
1
8
0
4
5
0
0
5
0
7
1
0
0
9
1
7
1
0
0
2
2
0
0
5
10
1
1
0
1
0
4
1
11
0
0
0
0.049
0.005
0
0.016
0
0.252
0.634
0.039
0.069
0
0
0.016
0.053
0.035
0
0.002
0.009
0.127
0.046
0.005
0.005
0.076
0
0.053
0.037
0
0.002
0.009
0.023
0.009
0
0.005
0
0
0.032
0
0.030
0
0
0
7
1
0
3
0
6
3
5
10
0
0
4
4
5
0
1
1
5
1
2
1
4
0
5
4
0
1
2
3
2
0
1
0
0
5
0
3
0
37
s
h
gr
gb
s
gr
gr
gr
gr
gr
h
l
h
h
h
h
Border
ARTICLE IN PRESS
Justicia brasiliana Roth
Stenandrium Nees sp.
Pfaffia tuberosa (Moq. ex DC.) Hicken
Habranthus gracilifolius Herb.
Schinus weinmanniaefolius Engl.
Eryngium ciliatum Cham. & Schlecht.
Eryngium elegans Cham. & Schlecht.
Eryngium horridum Malme
Eryngium pristis Cham. & Schlecht.
Eryngium sanguisorba Cham. & Schlecht.
Hydrocotyle exigua (Urb.) Malme
Forsteronia glabrescens Müll.Arg.
Rumohra adiantiformis (G.Forst.) Ching
Achyrocline satureioides Gard.
Acmella bellidioides (Sm.) R.K.Jansen
Aspilia montevidensis (Spreng.) Kuntze
Asteraceae 1
Asteraceae 2
Baccharis articulata Pers.
Baccharis cognata DC.
Baccharis dracunculifolia DC.
Baccharis leucopappa DC.
Baccharis ochracea Spreng.
Baccharis patens Baker
Baccharis rufescens Spreng.
Baccharis sessiliflora Vahl
Baccharis trimera (Less.) DC.
Calea serrata Less.
Calea uniflora Krasch.
Chaptalia integerrima (Vell.) Burkart
Chaptalia runcinata Kunth
Chaptalia sinuata Baker
Conyza bonariensis (L.) Cronquist
Conyza chilensis Spreng.
Eupatorium ascendens Sch.Bip. ex Baker
Eupatorium intermedium DC.
Eupatorium ivaefolium L.
Eupatorium lanigerum Hook. & Arn.
Eupatorium ligulaefolium Hook. & Arn.
Eupatorium tweedianum Hook. & Arn.
LF
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Apocynaceae
Aspidiaceae
Asteraceae
Species
Family
Asteraceae
Cyperaceae
Heterothalamus psiadioides Less.
Hieracium commersonii Monnier
Hypochoeris sp.
Lucilia acutifolia Cass.
Lucilia nitens Less.
Noticastrum gnaphalioides (Baker) Cuatrec.
Orthopappus angustifolius Gleason
Porophyllum lanceolatum DC.
Pterocaulon alopecuroideum DC.
Pterocaulon rugosum Malme
Senecio heterotrichius DC.
Stenachaenium riedelii Baker
Stevia aristata D.Don ex Hook. & Arn.
Stevia cinerascens Sch.Bip. ex Baker
Verbesina subcordata DC.
Vernonia flexuosa Sims
Vernonia nudiflora Less.
Viguiera anchusaefolia (DC.) Bak.
Dolichandra cynanchoides Cham
Macfadyena unguis-cati (L.) A.H.Gentry
Moritzia ciliata DC.
Dyckia leptostachya Baker
Parodia ottonis (Lehm.) N.P.Taylor
Wahlenbergia linarioides DC.
Spergularia grandis Cambess.
Halimium brasiliense Gross.
Commelina erecta L.
Convolvulus crenatus Vahl
Dichondra sericea Sw.
Evolvulus sericeus Sw.
Ipomoea nitida Griseb.
Bulbostylis sp.
Bulbostylis closii Barros
Bulbostylis juncoides (Vahl) Kük. ex Osten
Bulbostylis sphaerocephalus C.B.Clarke
Carex phalaroides Kunth
Cyperaceae 1
Cyperus aggregatus Endl.
Cyperus incomtus Kunth
Cyperus lanceolatus Poir.
LF
s
gr
h
gr
gr
h
h
s
h
h
s
gr
h
h
h
h
s
h
l
l
h
gr
s
th
h
h
h
l
h
h
gr
hc
hc
hc
hc
gr
gr
gr
gr
gr
RT
n
s
s
s
s
s
s
s
s
s
n
s
s
s
s
s
s
s
s
s
s
s
s
n
n
s
s
s
n
n
s
r
r
r
r
s
s
s
s
s
Border
Group 1
Group 2
Group 3
C
F
C
F
C
F
C
F
0
0
0
0
0
0
0.009
0.007
0.009
0
0
0
0.016
0.005
0.032
0.014
0.023
0
0.009
0.002
0.005
0
0
0
0.002
0
0.035
0.005
0.005
0.030
0.009
0
0
0.007
0.023
0
0
0
0.005
0
0
0
0
0
0
0
1
2
1
0
0
0
1
1
2
3
6
0
1
1
1
0
0
0
1
0
5
1
1
7
1
0
0
1
1
0
0
0
1
0
0
0
0
0.021
0.009
0
0.005
0.044
0.019
0.067
0
0.014
0.009
0.035
0.028
0.162
0.093
0
0
0
0
0.139
0.028
0
0
0.005
0.002
0
0.012
0.056
0
0.012
0.007
0.046
0.007
0
0
0.002
0
0
0
0
0
5
1
0
1
8
1
5
0
1
1
4
2
9
6
0
0
0
0
1
1
0
0
1
1
0
3
8
0
2
2
9
1
0
0
1
0
0
0.113
0.032
0
0.081
0.009
0.012
0.102
0.026
0
0.081
0.014
0.009
0.002
0.102
0
0.065
0.109
0.019
0
0
0
0
0
0
0
0.012
0.012
0
0.009
0.049
0
0.009
0.012
0.009
0.030
0
0.005
0.016
0
0.007
10
4
0
11
2
2
8
5
0
4
3
1
1
2
0
6
9
2
0
0
0
0
0
0
0
3
2
0
2
7
0
2
2
1
1
0
1
4
0
2
0.188
0
0.009
0.021
0.023
0.019
0.007
0.021
0
0.069
0.012
0
0.005
0.030
0.002
0.132
0.116
0
0
0
0
0
0
0.002
0
0.009
0.007
0.023
0.019
0.032
0
0
0
0.016
0.007
0.007
0
0
0
0
2
0
1
4
2
1
2
7
0
7
2
0
1
5
1
9
10
0
0
0
0
0
0
1
0
2
2
1
3
8
0
0
0
5
2
2
0
0
0
0
ARTICLE IN PRESS
Boragniaceae
Bromeliaceae
Cactaceae
Campanulaceae
Caryophylaceae
Cistaceae
Commelinaceae
Convolvulaceae
Species
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Bignoniaceae
38
Table A1. (continued )
Cyperaceae
Dioscoreaceae
Ebenaceae
Ericaceae
Erythroxylaceae
Euphorbiaceae
Euphorbiaceae
Lamiaceae
Liliaceae
Linaceae
Lythraceae
Malpighiaceae
Malvaceae
t
h
h
h
s
gr
s
h
s
h
h
gr
gr
h
s
h
gr
h
gb
gb
h
h
h
s
h
h
gb
h
h
l
s
s
s
n
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
n
s
s
s
s
s
s
s
0.023
0
0.009
0
0.120
0.002
0.005
0.009
0.037
0
0.012
0
0
0.002
0
0
0.009
0
0.007
0.002
0
0.021
0
0.007
0
0.014
0.009
0
0.005
0.009
0
0
0
0.002
0
0.002
0.005
0
0.012
0.002
0
0
0
0.009
0.014
0.009
4
0
3
0
4
1
1
1
3
0
2
0
0
1
0
0
1
0
1
1
0
4
0
1
0
3
2
0
1
2
0
0
0
1
0
1
1
0
1
1
0
0
0
1
2
1
0.192
0.016
0.025
0
0
0
0
0
0.049
0.005
0.019
0
0.002
0
0.005
0.002
0.049
0
0.028
0
0.009
0.039
0.014
0.039
0.005
0.009
0
0.007
0.009
0.016
0
0.005
0.012
0.005
0.002
0
0.012
0.009
0
0
0
0.005
0.005
0
0
0
12
4
5
0
0
0
0
0
6
1
4
0
1
0
1
1
3
0
4
0
1
5
1
2
2
4
0
1
2
3
0
2
2
2
1
0
1
1
0
0
0
1
1
0
0
0
0.079
0.007
0.030
0.009
0
0
0
0
0.139
0.005
0.042
0
0
0
0
0.028
0
0.042
0.009
0.079
0.005
0.053
0
0
0
0.039
0.021
0
0
0.012
0.009
0.005
0
0.028
0.007
0.046
0.060
0
0.630
0
0
0.002
0
0
0.023
0.010
11
3
6
1
0
0
0
0
7
1
7
0
0
0
0
4
0
2
1
1
1
8
0
0
0
4
4
0
0
2
2
1
0
3
2
5
4
0
4
0
0
1
0
0
3
1
0.116
0
0.009
0
0
0.002
0
0.005
0.074
0.032
0.014
0.005
0
0
0
0.012
0.023
0.005
0.039
0
0.005
0.021
0
0.009
0
0.007
0.039
0
0.014
0
0
0.002
0
0.009
0.002
0
0.002
0
0
0
0.005
0
0
0.009
0.002
0.005
10
0
3
0
0
1
0
1
7
2
5
1
0
0
0
3
3
1
7
0
2
5
0
1
0
1
5
0
2
0
0
1
0
1
1
0
1
0
0
0
1
0
0
1
1
1
39
r
r
s
s
s
n
r
n
n
n
s
ARTICLE IN PRESS
Iridaceae
hc
hc
gr
gr
l
t
t
t
s
s
h
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Fabaceae
Rhynchospora globularis Small
Rhynchospora setigera Boeck
Scleria sellowiana Kunth
Scleria sp.
Dioscorea multiflora Mart. ex Griseb.
Diospyros inconstans Jacq.
Leucothoe eucalyptoides DC.
Erythroxylum argentinum O.E. Schulz
Croton cf. nitrariaefolium Baill.
Croton thermarum Müll.Arg.
Euphorbia selloi (Klotzsch & Garcke) Boiss. in DC.
Euphorbiaceae 1
Euphrobiaceae 2
Sebastiania brasiliensis Spreng.
Tragia emrichii Herter
Aeschynomene elegans Cham. & Schlecht.
Centrosema virginianum Benth.
Chamaecrista nictitans Moench
Clitoria nana Benth.
Collaea stenophylla Benth.
Crotalaria tweediana Benth.
Desmanthus tatuhyensis Hoehne
Desmodium affine Schltdl.
Desmodium incanum DC.
Galactia gracillima Benth.
Galactia marginalis Benth.
Macroptilium prostratum Urb.
Mimosa parvipinna Benth. in Hook
Rhynchosia diversifolia Micheli
Stylosanthes montevidensis Vogel
Zornia sericea Moric.
Cypella coelestis (Lehm.) Diels
Herbertia pulchella Sweet
Sisyrinchium macrocephalum Graham
Sisyrinchium scariosum I.M.Johnst.
Sisyrinchium vaginatum Spreng.
Glechon squarrosa Benth.
Hyptis stricta Benth.
Salvia procurrens Benth.
Nothoscordum bonariense Beauverd
Cliococca selaginoides (Lam.) C.M.Rogers & Mild.
Cuphea glutinosa Cham. & Schlecht.
Janusia cf. guaranitica A.Juss.
Krapovickasia urticifolia (A. St.-Hil.) Fryxell
Pavonia hastata Cav.
Sida rhombifolia L.
Family
Malvaceae
Melastomataceae
Myrsinaceae
Myrtaceae
Myrtaceae
Orchidaceae
Oxalidaceae
Species
s
s
t
t
t
s
t
gb
gb
gb
h
hc
hc
hc
hc
hc
hc
hc
hc
hc
hc
gr
hc
gr
hc
hc
hc
hc
gr
hc
hc
gr
hc
hc
hc
gr
hc
gr
hc
hc
RT
s
s
n
n
n
s
s
s
s
s
s
r
r
r
r
r
r
r
r
r
r
s
r
s
r
r
r
r
s
r
r
s
r
r
r
s
r
s
r
r
Border
Group 1
Group 2
Group 3
C
F
C
F
C
F
C
F
0.021
0.032
0.046
0.009
0.046
0
0.093
0
0
0
0.095
0.093
0
0
0
0
0
0.002
0.039
0.046
0
0
0.037
0
0
0
0
0
0.083
0.019
0.009
0.150
0.588
0
0.171
0.000
0.051
0.049
0.005
0.014
3
2
2
1
1
0
1
0
0
0
3
6
0
0
0
0
0
1
4
4
0
0
2
0
0
0
0
0
5
2
1
8
8
0
3
0
4
3
1
1
0.019
0.037
0
0
0
0
0.056
0.005
0.009
0.016
0
0.218
0.044
0.146
0
0
0
0
0.590
0.199
0.005
0.127
0
0.056
0
0
0
0
0
0.014
0
0.044
0.287
0.032
0.296
0
0
0
0.000
0.000
2
6
0
0
0
0
1
1
1
3
0
4
4
7
0
0
0
0
11
7
2
6
0
1
0
0
0
0
0
2
0
6
5
4
4
0
0
0
0
0
0
0.155
0
0
0
0
0
0
0
0.002
0
0.123
0
0.113
0
0.407
0.014
0.350
0.257
0.676
0
0
0.468
0.102
0
0.010
0.019
0.014
0
0.026
0.010
0.174
0.454
0
0.528
0
0
0
0
0
0
5
0
0
0
0
0
0
0
1
0
3
0
4
0
5
2
7
5
9
0
0
6
3
0
3
2
3
0
2
1
8
8
0
5
0
0
0
0
0
0.058
0.012
0
0.009
0
0.009
0
0
0.007
0.009
0
0.551
0.326
0.000
0.053
0.069
0
0.016
0.556
0.188
0
0
0
0.359
0.005
0.005
0.014
0.030
0.000
0.171
0.000
0.025
1.116
0.019
0.044
0.005
0
0
0.005
0
4
4
0
2
0
1
0
0
3
3
0
5
5
0
2
1
0
2
5
5
0
0
0
8
1
1
4
3
0
6
0
6
9
2
3
1
0
0
2
0
ARTICLE IN PRESS
Wissadula glechomatifolia R.E.Fr.
Tibouchina gracilis (Bonpl.) Cogn.
Myrsine coriacea R.Br
Myrsine umbellata Mart.
Blepharocalyx salicifolius O.Berg
Campomanesia aurea O.Berg
Myrcia palustris DC.
Stenorrhynchus arechavaletsmii Barb.Rodr.
Oxalis brasiliensis Lodd.
Oxalis conorrhiza Jacq.
Peperomia pereskiaefolia H.B.& K.
Andropogon lateralis Nees
Andropogon leucostachyus H. B. & K.
Andropogon selloanus (Hack.) Hack.
Andropogon ternatus Nees
Aristida circinalis Lindm.
Aristida condylifolia Caro
Aristida filifolia (Arechav.) Herter
Aristida flaccida Trin. & Rupr.
Aristida laevis Kunth
Aristida venustula Arechav.
Axonopus argentinus Parodi
Axonopus suffultus (Mikan ex Trin.) Parodi
Axponopus sp.
Briza calotheca (Trin.) Hack
Briza sp.
Briza subaristata Lam.
Briza uniolae Nees ex Steud.
Calamagrostis viridiflavescens Steud.
Danthonia montevidensis Hackel & Arech.
Danthonia secundiflora J.Presl & C.Presl
Dichanthelium sabulorum (Lam.) Gould & C.A.Clark
Elionurus muticus (Spreng.) Kuntze
Eragrostis polytricha Nees
Leptocoryphium lanatum Nees
Melica brasiliana Ard.
Oplismenus hirtellus (L.) P.Beauv.
Panicum ovuliferum Trin.
Panicum peladoense Henrard
Paspalum mandiocanum Trin.
LF
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Piperaceae
Poaceae
40
Table A1. (continued )
Poaceae
Scrophulariaceae
hc
hc
hc
hc
s
h
h
h
l
h
h
h
t
s
h
h
h
h
t
t
s
th
h
l
s
r
r
r
r
s
s
s
s
s
s
s
s
n
s
s
s
s
s
n
n
s
n
s
s
s
0.118
0.037
0.016
0
0.046
0.030
0
0.086
0.007
0.102
0
0.019
0.093
0.037
0
0
0
0
0.002
0
0
0
0
0
0
0.241
0.009
0.044
0
0.009
0
0.023
0.007
0.023
0.007
0.009
0.005
0
0
0.002
0.028
0
0
0
0
0.069
4
2
3
0
1
2
0
4
1
1
0
2
1
2
0
0
0
0
1
0
0
0
0
0
0
4
1
3
0
1
0
2
2
1
2
3
2
0
0
1
3
0
0
0
0
1
0.014
0
0.023
0.005
0.093
0
0.005
0.044
0.023
0.012
0
0
0.039
0.002
0.067
0
0
0.005
0.009
0
0.005
0
0
0.005
0.005
0
0
0.093
0
0
0
0
0
0
0
0.002
0.102
0
0
0
0
0
0.012
0
0.002
0
2
0
4
1
1
0
1
3
4
2
0
0
3
1
3
0
0
1
1
0
1
0
0
1
1
0
0
6
0
0
0
0
0
0
0
1
6
0
0
0
0
0
3
0
1
0
0.060
0
0.035
0
0.046
0.044
0
0.713
0.104
0.331
0.009
0.012
0.116
0.009
0.009
0
0
0
0
0.012
0
0
0
0
0
0.030
0.039
0.012
0
0
0.032
0
0.019
0
0
0.076
0.039
0
0
0
0
0
0
0.005
0
0
3
0
3
0
2
4
0
5
8
6
1
2
1
1
1
0
0
0
0
1
0
0
0
0
0
4
4
2
0
0
5
0
5
0
0
10
5
0
0
0
0
0
0
1
0
0
0.012
0
0.227
0
0
0.234
0.005
0.030
0
0.005
0
0.093
0.106
0.007
0.593
0.002
0.002
0
0
0
0
0
0.002
0
0
0.025
0
0.063
0.014
0.016
0
0
0.009
0
0
0.007
0.063
0.005
0.002
0
0
0.019
0.002
0
0.009
0
4
0
8
0
0
6
1
5
0
1
0
5
2
2
5
1
1
0
0
0
0
0
1
0
0
2
0
7
2
3
0
0
3
0
0
3
7
1
1
0
0
1
1
0
1
0
41
Smilacaceae
Solanaceae
r
s
r
r
r
r
r
r
r
r
r
r
r
r
r
ARTICLE IN PRESS
Sapindaceae
hc
gr
hc
hc
hc
hc
hc
hc
hc
hc
hc
hc
hc
hc
hc
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Polygalaceea
Rubiaceae
Paspalum plicatulum Michx.
Paspalum sp.
Piptochaetium montevidense (Spreng.) Parodi
Poaceae/trib. Andropogoneae 5
Saccharum asperum Steud.
Schizachyrium microstachyum (Ham.) Roseng., B.R.Arill. & Izag.
Schizachyrium spicatum (Spreng.) Herter
Schizachyrium tenerum Nees
Setaria parviflora (Poiret) M.Kerguélen
Setaria vaginata Spreng.
Sporobolus multinodis Hackel
Stipa filiculmis Delile
Stipa filifolia Nees
Stipa tenuiculmis Hackel
Trachypogon montufari Nees
Poaceae 1
Poaceae 2
Poaceae 3
Poaceae 4
Poaceae 5
Poaceae 6
Poaceae/trib. Andropogoneae 1
Poaceae/trib. Andropogoneae 2
Poaceae/trib. Andropogoneae 3
Poaceae/trib. Andropogoneae 4
Monnina oblongifolia Arechav.
Borreria capitata DC.
Borreria fastigiata K.Schum.
Borreria verticillata G.Mey.
Chiococca alba Hitchc.
Diodia apiculata K.Schum.
Diodia cymosa Cham.
Galium uruguayense Bacigalupo
Guettarda uruguensis Cham. & Schlecht.
Psychotria carthagenensis Jacq.
Relbunium hirtum K.Schum.
Richardia grandiflora Steud.
Richardia humistrata Steud.
Rubiaceae sp.
Cupania vernalis Cambess.
Dodonaea viscosa Jacq.
Angelonia integerrima Spreng.
Gerardia communis Cham. & Schlecht.
Mecardonia herniarioides (Cham.) Pennell
Smilax campestris Griseb.
Cestrum strigillatum Ruiz & Pav.
Family
Solanaceae
Sterculariaceae
Styracacae
Symplocaceae
Turneraceae
Verbenaceae
Species
h
h
s
t
t
t
s
s
h
h
h
h
l
RT
s
s
s
n
s
s
s
s
s
s
s
s
s
Border
Group 1
Group 2
Group 3
C
F
C
F
C
F
C
F
0
0
0
0.009
0.116
0.002
0
0
0.002
0.007
0.002
0.012
0.236
0.002
0
0
0
0
0
0
0
0
0
1
1
1
0
0
1
1
1
2
4
1
0
0
0
0
0
0
0
0.002
0.009
0
0
0.009
0.009
0
0
0.030
0
0.007
0
0
0
0
0
0
0
0
0
1
1
0
0
1
2
0
0
3
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0.009
0
0.005
0
0.007
0.007
0
0
0.002
0
0
0
0
0
0
0
0
0
0
0
2
0
1
0
3
1
0
0
1
0
0
0
0
0
0.016
0
0.012
0
0
0
0.002
0.009
0
0.060
0.002
0
0.002
0
0
0.002
0.002
0.002
0.002
0.002
3
0
3
0
0
0
1
1
0
3
1
0
1
0
0
1
1
1
1
1
ARTICLE IN PRESS
Petunia integrifolia (Hook.) Schinz & Thell.
Petunia ovalifolia Miers
Waltheria douradinha A. St.Hil.
Styrax leprosum Hook. et Arn.
Symplocos tetandra Mart.
Symplocos uniflora Benth.
Turnera selloi Arechav.
Turnera sidoides L.
Glandularia megapotamica (Spreng.) Cabrera & Dawson
Lantana montevidensis (Spreng.) Briq.
Verbena ephedroides Cham.
Verbena pseudojuncea Gay
Cissus striata Ruı́z & Pav.
Sp. 1
Sp. 2
Sp. 3
Sp. 4
Sp. 5
Sp. 6
Sp. 7
LF
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Vitaceae
Unidentified
42
Table A1. (continued )
Table B1
Family
Anacardiaceae
Arecaceae
Asteraceae
Fabaceae
Lamiaceae
Lauraceae
Malvaceae
f
c
c
b
c
c
c
c
c
c
c
c
c
c
c
c
c
c
c
c
c
b
f
f
b
f
c
c
f
f
b
c
c
c
f
c
c
f
f
f
GF
t
s
t
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
t
t
t
t
s
s
t
t
s
s
s
s
t
s
s
t
s
t
Border
Group 1
Group 2
Group 3
C
F
C
F
C
F
C
F
0
0.67
0.83
0
0
1.94
0
0
0
0.28
0
0.72
0
0.94
0.78
0.28
0.28
0.56
0.92
0.28
1.44
0
0
0
0
0.11
2.44
0.39
0
0
0
0
0
0.22
0
0.5
0.22
0
0
0
0
6
2
0
0
4
0
0
0
1
0
5
0
3
3
1
1
1
6
1
13
0
0
0
0
1
11
2
0
0
0
0
0
2
0
3
2
0
0
0
0
0.33
0
0
0.33
5.69
0
0
0.24
0.69
0
1.43
0.25
0.25
2.21
0
0.47
0.19
0.85
0.25
3.71
0
0
0
0
0
0.43
0.42
0
0
0
0
0.47
0
0
0
0
0
0
0
0
6
0
0
3
36
0
0
5
1
0
17
6
3
26
0
3
2
19
3
54
0
0
0
0
0
10
9
0
0
0
0
3
0
0
0
0
0
0
0
0
2.53
0
0
0.53
8.07
0
0.19
0.69
0.06
0
0.51
0.29
0.06
3.19
0
0
0.63
2.68
0
4.56
0
0
0
0
0
2.75
0
0
0
0
0
0
0
0
1.01
0.06
0
0
0
0
18
0
0
4
53
0
2
4
1
0
7
6
1
32
0
0
9
37
0
63
0
0
0
0
0
35
0
0
0
0
0
0
0
0
13
1
0
0
0
0
0.07
0.06
0
1.44
3.85
0.97
0
0.88
2.72
0.06
1.85
0.64
0.97
2.6
0
0.19
3.97
0.74
0
3.26
0
0
0
0
0
3.1
0.72
0
0
0
0
1.25
0
0
0.11
0
0
0
0
0
2
1
0
12
30
7
0
7
13
1
28
10
4
30
0
2
16
14
0
48
0
0
0
0
0
31
7
0
0
0
0
7
0
0
2
0
0
0
0
43
Meliaceae
Monimiaceae
Myrsinaceae
s
s
r
s
s
s
n
s
s
s
s
s
s
s
s
n
s
n
s
s
s
r/s
s
n
r
n
n
n
n
s
s
s
s
s
n
s
s
n
n
n
SG
ARTICLE IN PRESS
Euphorbiaceae
Lithraea brasiliensis Marchand
Schinus weinmanniaefolius Engl.
Butia capitata Becc.
Baccharidastrum triplinervium (Less.) Cab.
Baccharis articulata Pers.
Baccharis cognata DC.
Baccharis dracunculifolia DC.
Baccharis leucopappa DC.
Baccharis ochracea Spreng.
Baccharis patens Baker
Baccharis rufescens Spreng
Baccharis sessiliflora Vahl
Baccharis trimera (Less.) DC.
Eupatorium intermedium DC.
Eupatorium ligulaefolium Hook. & Arn.
Eupatorium pedunculosum Hook. & Arn.
Eupatorium tweedianum Hook. & Arn.
Heterothalamus psiadioides Less.
Porophyllum lanceolatum DC.
Verbesina subcordata DC.
Vernonia nudiflora Less.
Opuntia monacantha Haw.
Maytenus cassineformis Reiss.
Diospyros inconstans Jacq.
Leucothoe eucalyptoides DC.
Erythroxylum argentinum O.E. Schulz
Croton cf. nitrariaefolium Baill.
Croton thermarum Müll.Arg.
Sebastiania brasiliensis Spreng.
Sebastiania serrata Müll.Arg.
Calliandra tweedii Benth.
Collaea stenophylla Benth.
Mimosa parvipinna Benth. in Hook
Hyptis mirabilis Briq.
Ocotea puberula (Rich.) Nees
Pavonia hastata Cav.
Sida rhombifolia L. –
Trichilia clausseni C.DC.
Mollinedia elegans Tul.
Myrsine coriacea (Sw.) R.Br.
RT
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Cactaceae
Celastraceae
Ebenaceae
Ericaceae
Erythroxlaceae
Euphorbiaceae
Species
44
Table B1. (continued )
Family
Species
RT
SG
GF
Border
C
Myrsinaceae
Myrtaceae
Myrtaceae
Solanaceae
Styracaceae
f
c
f
f
f
c
f
f
f
f
f
f
f
b
f
b
f
f
f
t
s
t
t
t
s
t
t
t
s
s
t
t
s
t
s
t
t
t
0
0.39
0
0
0
0.11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0.21
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
F
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0.22
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Group 3
F
0
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0.13
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
F
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
ARTICLE IN PRESS
Rutaceae
Sapindaceae
n
s
s
s
s
s
n
n
n
s
n
s
n
n
s
s
n
s
s
F
Group 2
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Nyctaginaceae
Rosaceae
Rubiaceae
Myrsine umbellata Mart.
Campomanesia aurea O.Berg
Eugenia hyemalis Camb.
Myrcia palustris DC.
Myrciaria cuspidata O.Berg
Psidium L.
Guapira opposita (Vell.) Reitz
Quillaja brasiliensis Mart.
Guettarda uruguensis Cham. & Schltdl.
Psychotria carthagenensis Jacq.
Psychotria leiocarpa Cham. & Schltdl.
Zanthoxylum rhoifolium Lam.
Cupania vernalis Cambess.
Dodonaea viscosa Jacq.
Matayba elaeagnoides Radlk.
Cestrum strigilatum Ruı́z & Pav.
Styrax leprosum Hook. et Arn.
Symplocos tetrandra Mart.
Symplocos uniflora Bedd.
Group 1
ARTICLE IN PRESS
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
45
Table C1
Familiy
Species
RT
SG
GF
Border
Group 1
Group 2
Group 3
Anacardiaceae
Lithraea brasiliensis Marchand
Schinus weinmanniaefolius Engl.
Butia capitata Becc.
Baccharidastrum triplinervium (Less.) Cab.
Baccharis articulata Pers.
Baccharis cognata DC.
Baccharis dracunculifolia DC.
Baccharis leucopappa DC.
Baccharis ochracea Spreng.
Baccharis patens Baker
Eupatorium intermedium DC.
Eupatorium ligulaefolium Hook. & Arn.
Eupatorium pedunculosum Hook. & Arn.
Eupatorium tweedianum Hook. & Arn.
Heterothalamus psiadioides Less.
Porophyllum lanceolatum DC.
Vernonia nudiflora Less.
Opuntia monacantha Haw.
Maytenus cassineformis Reiss.
Diospyros inconstans Jacq.
Leucothoe eucalyptoides DC.
Erythroxylum argentinum O.E. Schulz
Croton cf. nitrariaefolium Baill.
Croton thermarum Müll.Arg.
Sebastiania brasiliensis Spreng.
Sebastiania serrata Müll.Arg.
Calliandra tweedii Benth.
Collaea stenophylla Benth.
Mimosa parvipinna Benth. in Hook
Hyptis mirabilis Briq.
Pavonia hastata Cav.
Trichilia clausseni C.DC.
Myrsine coriacea (Sw.) R.Br.
Myrsine guianensis (Aubl.) Kuntze
Myrsine umbellata Mart.
Campomanesia aurea O.Berg
Eugenia dimorpha Berg
Eugenia uniflora L.
Myrcia palustris DC.
Myrciaria cuspidata O.Berg
Guapira opposita (Vell.) Reitz
Guettarda uruguensis Cham. & Schltdl.
Psychotria carthagenensis Jacq.
Zanthoxylum rhoifolium Lam.
Casearia decandra Jacq.
Cupania vernalis Cambess.
Dodonaea viscosa Jacq.
Matayba elaeagnoides Radlk.
Cestrum strigilatum Ruı́z & Pav.
Styrax leprosum Hook. et Arn.
Symplocos tetrandra Mart.
Symplocos uniflora Bedd.
Lantana camara L.
s
s
r
s
s
s
n
s
s
s
s
s
n
s
n
s
s
r/s
s
n
r
n
n
n
n
s
s
s
s
s
s
n
n
n
n
s
s
n
s
s
n
n
s
s
n
n
n
s
s
n
s
s
r
f
c
c
b
c
c
c
c
c
c
c
c
c
c
c
c
c
b
f
f
b
f
c
c
f
f
b
c
c
c
c
f
f
f
f
c
b
f
f
f
f
f
f
f
f
f
b
f
b
f
f
f
b
t
s
t
s
s
s
s
s
s
s
s
s
s
s
s
s
s
s
t
t
t
t
s
s
t
t
s
s
s
s
s
t
t
t
t
s
t
t
t
t
t
t
s
t
t
t
s
t
s
t
t
t
s
0.25
0.17
0.25
0
0
0.08
0.08
0
0
0.08
1.00
0.50
0.42
0
0.17
0.17
0
0.17
0
0.08
0.25
0.17
0
0
0.25
0.17
0.17
0
0.25
0.17
0.33
0.17
0.50
0.17
0.58
0.42
0
0.08
0.33
1.00
0.08
0.08
0.33
0.08
0.25
0.08
1.92
0.17
0.08
0.17
0.17
0.92
0.17
0.08
0
0
0
0
0
0
0
0
0.33
0
0
0
0
0.25
0
0
0
0.04
0
0
0
0
0
0
0
0
0
0.04
0
0
0
0
0
0
0
0
0
0.13
0.08
0
0
0
0
0
0
0.04
0
0
0
0
0.08
0
0.13
0.13
0
0
0
0.33
0
0
0.13
0
0
0.54
0
0
0
1.13
0.04
0
0
0
0
0
0
0
0
0
0
0.08
0
0
0.13
0.00
0
0
0
0
0
0
0
0.04
0
0.04
0
0
0
0
0
0.08
0.04
0
0
0.08
0
0.13
0.13
0
0
0.46
1.13
0.83
0.04
0.17
1.33
0.21
2.13
0
0.04
2.29
1.33
0.08
0
0
0
0
0
0.17
0.04
0
0
0
0.08
0.58
0
0.13
0
0
0.04
0
0
0.04
0
0.13
0.17
0
0.04
0
0
0
0
0.08
0.08
0.04
0
0
0.13
0
Arecaceae
Asteraceae
Asteraceae
Cactaceae
Celastraceae
Ebenaceae
Ericaceae
Erythroxlaceae
Euphorbiaceae
Euphorbiaceae
Fabaceae
Lamiaceae
Malvaceae
Meliaceae
Myrsinaceae
Myrtaceae
Nyctaginaceae
Rubiaceae
Rutaceae
Salicaceae
Sapindaceae
Solanaceae
Styracaceae
Verbenaceae
ARTICLE IN PRESS
46
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Appendix B. Woody species 410 cm, recorded
in 252 MPs (1.5 m 1.5 m) on Morro Santana,
Porto Alegre, RS, Brazil
Given are family, species, regenerative type (RT; r:
resister, s: sprouter, n: non-sprouter), successional group
(SG, c: grassland species, b: forest pioneer/border
species; f: forest species), growth form (GF; s: shrub, t:
tree) mean cover per plot (C) and frequency (F) per
group of plots, separated for grassland plots at the
forest–grassland border (Bord., n ¼ 36) and plots in
the open grassland, grouped into plots burned three
months before (Group 1; n ¼ 72), burned a year before
(Group 2; n ¼ 72) and unburned for 3 years or more
(Group 3, n ¼ 72) (Table B1).
Appendix C. Woody species 480 cm, recorded
in 84 LPs (4.5 m 4.5 m) on Morro Santana,
Porto Alegre, RS, Brazil
Given are family, species, regenerative type (RT; r:
resister, s: sprouter, n: non-sprouter), successional group
(SG, c: grassland species, b: forest pioneer/border species;
f: forest species), growth form (GF; s: shrub, t: tree) and
mean number of individuals per plot, separated for
grassland plots at the forest–grassland border (Bord.,
n ¼ 12) and plots in the open grassland, grouped into
plots burned three months before (Group 1; n ¼ 24),
burned a year before (Group 2; n ¼ 24) and unburned for
3 years or more (Group 3, n ¼ 24) (Table C1).
References
Auld, T.A., Bradstock, R.A., 1996. Soil temperatures after the
passage of a fire: do they influence the germination of
buried seeds? Austral. J. Ecol. 21, 106–109.
Batalha, M.A., Martins, F.R., 2002. Life form spectra of
Brazilian Cerrado sites. Flora 197, 452–460.
Behling, H., 2002. South and southeast Brazilian grasslands
during Late Quaternary times: a synthesis. Palaeogeogr.
Palaeoclimatol. 177, 19–27.
Behling, H., Pillar, V.D., Orlóci, L., Bauermann, S.G., 2004.
Late Quaternary Araucaria forest, grassland (Campos), fire
and climate dynamics, studied by high resolution pollen,
charcoal and multivariate analysis of the Cambará do Sul
core in southern Brazil. Palaeogeogr. Palaeoclimatol. 203,
277–297.
Behling, H., Pillar, V.D., Müller, S., Overbeck, G., in press.
Late Holocene vegetation and fire dynamics of Morro
Santana, in Porto Alegre, southern Brazil. Appl. Veg. Sci.
Bellingham, P.J., Sparrow, A.D., 2000. Resprouting as a life
history strategy in woody plant communites. Oikos 89,
409–416.
Belsky, A.J., 1992. Effects of grazing, competition, disturbance
and fire on species composition and diversity in grassland
communities. J. Veg. Sci. 3, 187–200.
Benson, E.J., Hartnett, D.C., Mann, K.H., 2004. Belowground
bud banks and meristem limitation in tallgrass prairie plant
populations. Am. J. Bot. 91, 416–421.
Bigarella, J.J., 1971. Variacoes climáticas no Quaternário
superior do Brasil e sua datacao radiométrica pelo método
do Carbono 14. Paleoclimas 1, 1–22.
Bilenca, D.N., Miñarro, F.O., 2004. Àreas valiosas de pastizal
em las pampas y campos de Argentina, Uruguay y sur de
Brasil. Fundación Vida Silvestre Argentina, Buenos
Aires.
Boldrini, I.B., 1993. Dinâmica da vegetação de uma pastagem
natural sob diferentes niveis de oferta de forragem e tipos
de solos, Depressão Central, RS. Ph.D. Thesis, Faculdade
de Agronomia, UFRGS, Porto Alegre.
Boldrini, I.B., Eggers, L., 1997. Directionality of succession
after grazing exclusion in grasslands in the South of Brazil.
Coenoses 12, 63–66.
Bond, W.J., 1997. Functional types for prediction changes in
biodiversity: a case study in Cape fynbos. In: Smith, T.M.,
Shugart, H.H., Woodward, F.I. (Eds.), Plant Functional
Types: their Relevance to Ecosystem Properties and
Global Change. Cambridge University Press, Cambridge,
pp. 174–194.
Bond, W.J., Midgley, G.F., 2000. A proposed CO2-controlled
mechanism of woody plant invasion in grasslands and
savannas. Global Change Biol. 6, 865–869.
Bond, W.J., Van Wilgen, B.W., 1996. Fire and Plants.
Chapman & Hall, London.
Bond, W.J., Midgley, G.F., Woodward, F.I., 2003. What
controls South African vegetation-climate or fire? S. Afr. J.
Bot. 69, 1–13.
Box, E.O., 1986. Some climatic relationships of the vegetation
of Argentina, in global perspective. Veröff. Geobot. Inst.
ETH, Stiftung Rübel 91, 181–216.
Bradstock, R.A., Auld, T.A., 1995. Soil temperatures during
experimental bushfires in relation to fire intensity: consequences for legume germination and fire management in
south-eastern Australia. J. Appl. Ecol. 31, 76–84.
Bunting, S.C., Wright, H.A., Neuenschwander, L.F., 1980.
Long-term effects of fire on Cactus in the southern mixed
prairie of Texas. J. Range Manage. 33, 85–88.
Cain, S.A., 1950. Life forms and phytoclimate. Bot. Rev. 16,
1–32.
Chapin, F.S., Bret-Harte, M.S., Hobbie, S.E., Zhong, H.,
1996. Plant functional types as predictors of transient
responses of arctic vegetation to global change. J. Veg. Sci.
7, 347–358.
Chapman, R.R., Crow, G.E., 1981. Application of Raunkiaer’s life form system to plant species survival after fire.
Bull. Torr. Bot. Club 108, 472–478.
Chidumayo, E.N., 2006. Fitness implications of late bud break
and time of burning in Lannea edulis (Sond.) Engl.
(Anacardiaceae). Flora 201 (7), 588–594.
Christensen, N.L., 1985. Shrubland fire regimes and their
evolutionary consequences. In: Pickett, S.T.A., White, P.S.
(Eds.), The Ecology of Natural Disturbance and Patch
Dynamics. Academic Press, Orlando, pp. 85–100.
ARTICLE IN PRESS
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Collins, S.L., 1987. Interaction of distrubances in tallgrass
prairie: a field experiment. Ecology 68, 1243–1250.
Daubenmire, R., 1968. Ecology of fire in grasslands. In:
Gragg, J.B. (Ed.), Advances in Ecological Research.
Academic Press, London, pp. 209–266.
Diaz, S., Cabido, M., 1997. Plant functional types and
ecosystem function in relation to global change. J. Veg.
Sci. 8, 463–474.
Eggers, L., Porto, M.L., 1994. Ação do fogo em uma
comunidade campestre secundaria, analisada em bases
fitossociologicas. Bol. Inst. Biociências UFRGS 53,
1–88.
Ellenberg, H., Mueller-Dombois, D., 1966. A key to Raunkiaer plant life forms with revised subdivisions.
Ber. Geobot. Inst. ETH, Stiftg. Ruebel, Zuerich 37,
56–73.
Engle, D.M., Palmer, M.W., Crockett, J.S., Mitchell, R.L.,
Stevens, R., 2000. Influence of late season fire on early
successional vegetation of an Oklahoma prairie. J. Veg. Sci.
11, 135–144.
Eriksen, W., 1978. Ist das Pampaproblem gelöst? Naturwissensch. Rundsch. 31, 142–148.
Facelli, J.M., Pickett, S.T.A., 1991. Plant litter: its dynamics
and effects on plant community structure. Bot. Rev. 57,
1–32.
Fidelis, A., Overbeck, G.E., Pfadenhauer, J., Pillar, V.D.,
submitted. Effects of disturbance on population biology of
a rosette species in grasslands in southern Brazil. Plant
Ecol.
Franklin, J., Coulter, C.L., Rey, S.J., 2004. Change over 80
years in a southern California chaparral community related
to fire history. J. Veg. Sci. 15, 701–710.
Garcia, E.N., Boldrini, I.B., Jacques, A.V.A., 2002. Dinâmica
de formas vitais de uma vegetação campestre sob diferentes
práticas de manejo e exclusão. Iheringia Ser. Bot. 57,
215–241.
Ghermandi, L., Guthmann, N., Bran, D., 2004. Early post-fire
succession in northwestern Patagonia grasslands. J. Veg.
Sci. 15, 67–76.
Gibson, D.J., Seastedt, T.R., Briggs, J.M., 1993. Management
practices in tallgrass prairie: large- and small-scale experimental effects on species composition. J. Appl. Ecol. 30,
247–255.
Gill, A.M., 1981. Fire adaptive traits of vascular plants. In:
Mooney, H.H., Bonnicksen, N.L., Christensen, N.L.,
Lotan, J.E., Reiners, W.A. (Eds.), Fire Regimes and
Ecosystem Properties, United States Forest Service. General Technical Report WO-26, pp. 208–230.
Givnish, T.J., McDiarmid, R.W., Buck, W.R., 1986. Fire
adaptation in Neblinaria celiae (Theaceae), a high-elevation
rosette shrub endemic to a wet equatorial tepui. Oecologia
70, 481–486.
Grime, J.P., 1977. Evidence for the existence of three primary
strategies in plants and its relevance to ecological and
evolutionary theory. Am. Nat. 111, 1169–1195.
Henriques, R.P.B., Hey, J.D., 2002. Patterns and dynamics of
plant populations. In: Oliveira, P.S., Marquis, R.J. (Eds.),
The Cerrados of Brazil: Ecology and Natural History of a
Neotropical Savanna. Columbia University Press, New
York, pp. 140–158.
47
Hoffmann, W.A., 1996. The effects of fire and cover on
seedling establishment in a neotropical savanna. J. Ecol. 84,
383–393.
Hoffmann, W.A., 2000. Post-establishment seedling success in
the Brazilian Cerrado: a comparison of savanna and forest
species. Biotropica 32, 62–69.
Hoffmann, W.A., Moreira, A.G., 2002. The role of fire in
population dynamics of woody plants. In: Oliveira, P.S.,
Marquis, R.J. (Eds.), The Cerrados of Brazil: Ecology and
Natural History of a Neotropical Savanna. Columbia
University Press, New York, pp. 159–177.
Keeley, J.E., 1986. Resilience of mediterranean shrub communities to fire. In: Dell, B., Hopkins, A.J.M., Lamont,
B.B. (Eds.), Resilience in Mediterranean-ype Ecosystems.
Dr. W. Junk Publishers, Dordrecht, pp. 95–112.
Keeley, J.E., 1992. Recruitment of seedlings and vegetative
sprouts in unburned Chaparral. Ecology 73, 1194–1208.
Keeley, J.E., Fotheringham, C.J., 2000. Role of fire in
regeneration from seed. In: Fenner, M. (Ed.), Seeds: the
Ecology of Regeneration in Plant Communities. CABI
International, Wallingford, pp. 311–330.
Keeley, S.C., Keeley, J.E., Hutchinson, S.M., 1981. Postfire
succession in the herbaceous flora in southern California
chaparral. Ecology 62, 1608–1621.
Kern, A.A., 1994. Antecedentes Indı́genas. Editora da Universidade, Porto Alegre.
Klein, R.M., 1975. Southern Brazilian phytogeographic
features and the probable influence of upper quaternary
climatic changes in the floristic distribution. Bol. Paranaense Geociências 33, 67–88.
Knapp, A.K., 1985. Effect of fire and drought on the
ecophysiology of Andropogon gerardii and Panicum virgatum in a tallgrass prairie. Ecology 6, 1309–1320.
Knapp, A.K., Seastedt, T.R., 1986. Detritus accumulation
limits productivity of tallgrass prairie. BioScience 36,
662–668.
Laterra, P., Vignolio, O.R., Linares, M.P., Giaquinta, A.,
Maceira, N., 2003. Cumulative effects of fire on a tussock
pampa grassland. J. Veg. Sci. 14, 43–54.
Leach, M.K., Givnish, T.J., 1996. Ecological determinants
of species loss in remnant prairies. Science 273,
1555–1558.
Leite, P.F., 2002. Contribuição ao conhecimento fitoecologico
do Sul do Brasil. Ciênc. Ambiente 24, 51–74.
Levitt, J., 1980. Responses of Plants to Environmental Stress.
I. Chilling, Freezing, and High Temperature Stress.
Academic Press, New York.
Lindman, C.A.M., 1906. A vegetação no Rio Grande do Sul.
Universal, Porto Alegre.
Londo, G., 1976. The decimal scale for releves of permanent
quadrats. Vegetatio 33, 61–64.
Longhi-Wagner, H., 2003. Diversidade florı́stica dos campos
sul-brasileiros: Poaceae. In: 5. Congresso Nacional de
Botânica 2003, Belém. Desafios da Botânica brasileira no
novo milênio: inventário, sistematização e conservação da
diversidade vegetal. Sociedade Botânica do Brasil 2003,
vol. 1, pp. 117–120.
Loucks, O., Plumb-Mentjes, M.L., Rogers, D., 1985. Gap
processes and large-scale disturbance in sand prairies. In:
Pickett, G., White, P.S. (Eds.), The Ecology of Natural
ARTICLE IN PRESS
48
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Disturbance and Patch Dynamics. Academic Press, San
Diego, pp. 71–83.
Lunt, I.D., 1990. Impact of an autumn fire on a long-grazed
Themeda triandra (Kangaroo Grass) grassland: implications for management of invaded, remnant vegetations.
Victorian Nat. 107, 45–51.
Lunt, I.D., Morgan, J.W., 2001. The role of fire regimes in
temperate lowland grasslands of southeastern Australia. In:
Bradstock, R.A., Williams, J.E., Gill, A.M. (Eds.), Flammable Australia: the Fire Regimes and Biodiversity
of a Continent. Cambridge University Press, Cambridge,
pp. 177–197.
McIntyre, S., Lavorel, S., Tremont, R.M., 1995. Plant lifehistory attributes: their relationship to disturbance response
in herbaceous vegetation. J. Ecol. 83, 31–44.
McIntyre, S., Lavorel, S., Landsberg, J., Forbes, T.D.A., 1999.
Disturbance response in vegetation-towards a global
perspective on functional traits. J. Veg. Sci. 10, 621–630.
Miranda, A.C., Miranda, H., Dias, O.d.F., Dias, B.F.d.S.,
1993. Soil and air temperatures during prescribed cerrado
fires in Central Brazil. J. Trop. Ecol. 9, 313–320.
Miranda, H.S., Bustamante, M.M.C., Miranda, A.C., 2002.
The fire factor. In: Oliveira, P.S., Marquis, R.J. (Eds.), The
Cerrados of Brazil: Ecology and Natural History of a
Neotropical Savanna. Columbia University Press, New
York, pp. 51–68.
Mirelles, M.L., Klink, C.A., Silva, J.C.S., 1997. Un modelo de
estados y transiciones para el cerrado brasileno. Ecotropicos 10, 45–50.
Moreno, J.A., 1961. O clima do Rio Grande do Sul. Secretaria
de Agricultura, Porto Alegre.
Morgan, J.W., 1998. Composition and seasonal flux of the soil
seed bank of specis-rich Themeda triandra grasslands in
relation to burning history. J. Veg. Sci. 9, 145–156.
Morgan, J.W., 1999. Defining grassland fire events and the
response of perennial plants to annual fire in temperate
grassland of southern Australia. Plant Ecol. 144, 127–144.
Morgan, J.W., 2001. Seedling recruitment patterns over 4
years in an Australian perennial grassland community with
different fire histories. J. Ecol. 89, 908–919.
Morgan, J.W., Lunt, I.D., 1999. Effects of time-since-fire on
the tussock dynamics of a dominant grass (Themeda
triandra) in a temperate Australian grassland. Biol. Conserv. 88, 379–386.
Müller, S.C., Forneck, E., 2004. Forest-grassland mosaics in
the hills of Porto Alegre: a study case of forest expansion
patterns on Santana hill, Rio Grande do Sul, Brazil. In:
Porto, M.L. (Ed.), Workshop ‘‘Proteção e manejo da
vegetação natural de Porto Alegre com base em pesquisa de
padrões e dinâmica da vegetação’’. PPG Ecologia,
UFRGS, Porto Alegre, pp. 29–37.
Müller, S.C., Overbeck, G.E., Pfadenhauer, J., Pillar, V.D.,
2006. Plant functional types of woody species related to fire
disturbance in forest-grassland ecotones. Plant Ecology
(online first).
Nabinger, C., De Moraes, A., Maraschin, G.E., 2000. Campos
in Southern Brazil. In: Lemaire, G., Hodgson, J.G., de
Moraes, A., Cavalho, P.C.F., Nabinger, C. (Eds.), Grassland Ecophysiology and Grazing Ecology. CABI Publishing, Wallingford, pp. 355–376.
Noble, I.R., Slatyer, R.O., 1980. The use of vital attributes to
predict successional changes in plant communites subject to
recurrent disturbanes. Vegetatio 43, 5–21.
Oliveira, J.M., Pillar, V.D., 2004. Vegetation dynamics on
mosaics of Campos and Araucaria forest between 1974
and 1999 in Southern Brazil. Commun. Ecol. 5,
197–201.
Oliveira-Filho, A.T., Ratter, J.A., 2002. Vegetation physiognomies and woody flora of the Cerrado biome. In: Oliveira,
P.S., Marquis, R.J. (Eds.), The Cerrados of Brazil: Ecology
and Natural History of a Neotropical Savanna. Columbia
University Press, New York, pp. 91–120.
Overbeck, G.E., Müller, S.C., Pillar, V.D., Pfadenhauer, J.,
2005. Small-scale dynamics after fire in South Brazilian
humid subtropical grassland. J. Veg. Sci. 16, 655–664.
Overbeck, G.E., Müller, S.C., Pillar, V.D., Pfadenhauer, J.,
2006a. Floristic composition, environmental variation and
species distribution patterns in a burned grassland in
southern Brazil. Braz. J. Biol. 66, 1073–1090.
Overbeck, G.E., Müller, S.C., Pillar, V.D., Pfadenhauer, J.,
2006b. No heat-stimulated germination found in herbaceous species from burned subtropical grassland. Plant Ecol
184, 237–243.
Pillar, V.D., 2004. MULTIV: Multivariate Exploratory
Analysis, Randomization Testing and Bootstrap Resampling. User’s Guide v. 2.3.10. Departamento de Ecologia,
UFRGS, Porto Alegre, RS, Brazil. Available at /http://
ecoqua.ecologia.ufrgs.brS.
Pillar, V.D., Orlóci, L., 2004. Character-based community
analysis: the theory and an application program. Electronic
version. Available at: /http://www.ecoqua.ecologia.
ufrgs.brS (accessed 4 April 2005).
Pillar, V.D., Quadros, F.L.F., 1997. Grassland-forest boundaries in southern Brazil. Coenoses 12, 119–126.
Rambo, B., 1953. Historia da flora do Planalto rio-grandense.
An. Bot. Herbárium Barbosa Rodrigues 5, 185–232.
Rambo, B., 1956. A flora fanerogamica dos aparados
riograndeses. Sellowia 7, 235–298.
Ramsay, P.M., Oxley, E.R.B., 1986. Fire temperatures and
postfire plant community dynamics in Ecuadorian grass
páramo. Vegetatio 124, 129–144.
Raunkiaer, C., 1934. The Life Forms of Plants and Statistical
Plant Geography. Claredon, Oxford.
Reitz, P.R., Klein, R.M., 1964. O reino vegetal de Rio do Sul.
Sellowia 16, 9–118.
Rodrı́guez, C., Leoni, E., Lezama, F., Altesor, A., 2003.
Temporal trends in species composition and plant traits
in natural grasslands of Uruguay. J. Veg. Sci. 14,
433–440.
Rowe, J.S., 1983. Concepts of fire effects on plant individuals
and species. In: Wein, R.W., MacLean, D.A. (Eds.), The
Role of Fire in Northern Circumpolar Ecosystems. Wiley,
Chicester, pp. 135–154.
Sarmiento, G., 1990. Ecologı́a comparada de ecosistemas de
sabanas en América del Sur. In: Sarmiento, G. (Ed.), Las
sabanas americanas. Aspecto de su biogeografia, ecologia y
utilizacion. CIELAT, Mérida, pp. 15–56.
Sarmiento, G., 1992. Adaptive strategies of perennial
grasses in South American savannas. J. Veg. Sci. 3,
325–336.
ARTICLE IN PRESS
G.E. Overbeck, J. Pfadenhauer / Flora 202 (2007) 27–49
Sarmiento, G., Monasterio, M., 1983. Life forms and
phenology. In: Bourlière, F. (Ed.), Tropical Savannas.
Elsevier, Amsterdam, Oxford, New York, pp. 79–108.
Silva, J.F., Castro, F., 1989. Fire, growth and survivorship in a
neotropical savanna grass Andropogon semiberbis in Venezuela. J. Trop. Ecol. 5, 587–600.
Silva, J.F., Raventos, J., Caswell, H., 1990. Fire and fire
exclusion effects on the growth and survival of two savanna
grasses. Acta Oecol. 22, 783–800.
Silva, J.F., Raventos, J., Caswell, H., Trevisan, M.C., 1991.
Population responses to fire in a tropical savanna grass,
Andropogon semiberbis: a matrix model approach. J. Ecol.
79, 345–356.
Solbrig, O.T., 1993. Plant traits and adaptive strategies: their
role in ecosystem function. In: Schulze, E.-D., Mooney,
J.A. (Eds.), Biodiversity and Ecosystem Function. Springer,
Berlin, pp. 97–116.
Soriano, A., León, RJ.C., Sala, O.E., Lavado, R.S., Deregibus,
V.A., Cahuepé, O., Scaglia, A., Velazquez, C.A., Lemcoff,
J.H., 1992. Rı́o de la Plata grasslands. In: Coupland, R.T.
49
(Ed.), Ecosystems of the World 8A. Natural Grasslands.
Introduction and Western Hemisphere. Elsevier, Amsterdam, pp. 367–407.
Suding, K.N., 2001. The effect of spring burning on competitive
ranking of prairie species. J. Veg. Sci. 12, 849–856.
Taylor, H.C., 1978. Capensis. In: Werger, M.J.A. (Ed.),
Biogeography and Ecology of Southern Africa. Dr. W.
Junk, The Hague, pp. 171–229.
Towne, E.G., Owensby, C.E., 1984. Long-term effects of
annual burning at different dates in ungrazed Kansas
Tallgrass Prairie. J. Range Manage. 37, 392–397.
Trindade, J.P.P., Da Rocha, M.G., 2001. Rebrotamento de
capim caninha (Andropogon lateralis Nees) sob o efeito do
fogo. Ciênc. Rural 31, 1057–1061.
Vogl, R.J., 1974. Effects of fire on grasslands. In: Kozlowski,
T.T., Ahlgren, C.E. (Eds.), Fire and Ecosystems. Academic
Press, New York, pp. 139–194.
Walter, H., 1967. Das Pampaproblem in vergleichend ökologischer Betrachtung und seine Lösung. Erdkunde 21,
181–202.