Applied Vegetation Science && (2014)

Does fire induce flowering in Brazilian subtropical

grasslands?
Alessandra Fidelis & Carolina Blanco

Keywords
Flowering; Forbs; Functional groups;
Graminoids; Mowing; Subtropical grasslands
Nomenclature
APG III
Received 11 July 2013
Accepted 20 December 2013
Co-ordinating Editor: John Morgan
Fidelis, A. (Corresponding author,afidelis@
^ncias, UNESP –
rc.unesp.br): Instituto de Biocie
Univ. Estadual Paulista, Departamento de
Bot^anica, Av 24A, 1515, CEP 13506-900, Rio
Claro, Brazil
Blanco, C. (carolynablanco@gmail.com):
^ncias, USP – Universidade de
Instituto de Biocie
S~ao Paulo, Departamento de Ecologia, Rua do
Mat~ao, Trav 14, 321, CEP 05508-900, S~ao
Paulo, Brazil
Abstract
Questions: We aimed to analyse the effect of fire on flowering in subtropical
grasslands, by addressing the following questions: will fire history affect flower-
ing? If yes, do fire feedbacks influence flowering or is it just the removal of
above-ground biomass? Are there differences in burned and mowed plots?
Location: Subtropical grasslands in Southern Brazil (30°03′S, 51°07′W).
Methods: We established plots in areas with different fire histories: 30 d (30
plots: five replicates), 1 yr (14 replicates), 3 yr (30 plots: five replicates) since
the last fire, in experimentally burned and mowed plots (14 replicates each). We
counted the number of flowering species, as well as the number of flowering
stalks.
Results: Graminoid species flowered in highest numbers 1 yr after fire, whilst
forbs had more species flowering just after fire, indicating different reproductive
strategies in post-fire environments. Mowing was not as efficient as fire in stim-
ulating flowering. Finally, the different functional groups showed different flow-
ering responses to time since last fire and to the different types of management.
Conclusions: Our results show fire stimulated flowering. Although mowing
can be a good alternative for maintaining plant diversity, our study showed that
this practice is not as efficient as fire in stimulating flowering. However, fire sea-
son should be noted as a limiting factor to the recovery of C3 grasses in these sub-
tropical grasslands, and annual burns may be harmful to C4 grasses, since they
delay their flowering to the next post-fire growing season.

Wrobleski & Kauffman 2003). Other factors found in post-

Introduction
In several ecosystems worldwide, fire can be responsible
for maintaining vegetation physiognomy, structure and
diversity (Bond & Keeley 2005). Flowering phenology in
plant communities can also be affected by fire, since it can
be a trigger for flowering in several perennial plants (La-
mont & Downes 2011). Some geophytes, for example, can
flower within days after a fire in fynbos and chaparral veg-
etation (Keeley et al. 1981; Keeley 1993; Tyler & Borchert
2002), and then disperse seeds before the bare soil is cov-
ered again by other species.
It is known that several abiotic factors are related to
flowering (Ratchke & Lacey 1985). The three major factors
are light (Hulbert & Society 1988; Brewer 1995; Tyler &
Borchert 2002; Paritsis et al. 2006; Borchert & Tyler 2009),
temperature (Keeley 1993; Elzinga et al. 2007; Borchert &
Tyler 2009) and water (Smith-Ramirez & Armesto 1994;
fire sites can also be related to the increase of flowering,
such as smoke (Keeley 1993; Tyler & Borchert 2002),
increase in the availability of nutrients (Lamont & Runci-
man 1993; Saulnier & Reekie 1995; Wrobleski & Kauffman
2003; Sardans et al. 2005; Coates et al. 2006; Elzinga et al.
2007) and decrease in competition (Howe 1994; Lunt
1994; Coates et al. 2006; Coates & Duncan 2009).
Peak flowering can be synchronized by fire season and
frequency (Platt et al. 1988). For example, summer burns
in prairies increased flowering diversity, favouring early-
flowering species such as Andropogon gerardii, Sorghastrum
nutans and Schizachyrium scoparium (all C4 grasses; Howe
1994). Additionally, more plants with flowers and more
flowering stalks were found after summer and autumn
than after spring fires (Bowen & Pate 2004). Shorter inter-
vals of fire could lead to a high proportion of species flow-
ering (Coates et al. 2006), whilst other studies showed that

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Fire and flowering in subtropical grasslands
the longer the time since the last fire, the lower the propor-
tion of species flowering, as well as the number of flowers
per individual (Lamont & Runciman 1993; Lunt 1994).
Most studies on the effects of fire on flowering usually
analyse the responses of a single plant species or a group of
species. According to Jarrad et al. (2008), species can show
individual responses to fire, and thus studies that consider
multi-species responses should be carried out in order to
detect more generic trends (see Hinman & Brewer 2007
and references therein). Moreover, vegetation responses to
fire can also be evaluated by comparing the flowering time
of different functional groups: species flowering just after
fire have an advantage in replenishing their seed bank (La-
mont & Downes 2011), and hence assuring population
persistence. On the other hand, species that delay their
flowering in response to fire, by flowering only in the next
growing season, would be disadvantaged under annual
burnings. Thus, information about the effects of fire on
flowering of different species is important in order to
understand plant community dynamics in flammable eco-
systems.
Brazilian subtropical grasslands (known as Campos) are
ecosystems under the constant influence of disturbance,
mostly fire and grazing. They show an interesting mixture
of C3 and C4 grasses (Fidelis 2010) and a high diversity of
plant species (Boldrini et al. 2009). Besides the humid cli-
mate in southern Brazil, with low variation in precipitation
between months, grasslands are still present in these areas.
Therefore, disturbance is likely the major factor affecting
Campos grassland physiognomy and diversity (Overbeck
et al. 2007). Fire has been a frequent disturbance since
7400 cal
yr BP (Behling et al. 2004; Behling & Pillar
2007). Several plant species show adaptations to this type
of disturbance (e.g. presence of below-ground organs;
Fidelis et al. 2009) and they can recover rapidly after fire
(Fidelis et al. 2012) by resprouting mostly from the below-
ground bud bank (Fidelis et al., in press). Nowadays fires
are mostly anthropogenic and used as a management tool
by farmers to stimulate resprouting of graminoids, mostly
for cattle raising, in late winter, due to the high accumula-
tion of standing dead biomass (Heringer & Jacques 2002).
In Brazil, the use of fire as a management tool is a contro-
versial issue, even for grassland and savanna ecosystems
(Fidelis 2010; Fidelis & Pivello 2011). Although several
studies in southern Brazil have already shown the positive
effects of fire on plant populations (Fidelis et al. 2008, 2010)
and grassland community diversity and dynamics (Over-
beck et al. 2005; Fidelis et al. 2012), its use is very restricted
by law. Fire can efficiently open spaces within the dense
grass layer, allowing the establishment of other species, thus
maintaining the biodiversity of these subtropical grasslands
(Fidelis et al. 2012). Additionally, it can maintain dynamics
of some plant populations of different functional groups
2 Doi: 10.1111/avsc.12098 © 2014 International Association for Vegetation Science
A. Fidelis and C. Blanco
(forbs and shrubs), mostly by stimulating resprouting from
the bud bank (Fidelis et al. 2010; Fidelis et al., in press).
Mowing could be an alternative to fire since it removes
above-ground biomass and thus could also allow new colo-
nization and stimulate plants to resprout. However, Fidelis
et al. (2012) showed that fire removes litter and competi-
tors more efficiently than mowing, having a larger effect on
vegetation dynamics. However, less is known about the
effects of biomass removal (both by fire and mowing) on the
flowering phenology of subtropical grassland plant commu-
nities, which could later affect seedling recruitment and
plant establishment in these ecosystems.
Therefore, the present study is one of few to analyse the
effects of fire and mowing on flowering considering the
whole plant community. Thus, we aimed to: (1) evaluate
the effects of fire history on flowering of plant species and
functional groups in subtropical grasslands, as well as on
the number of flowering stalks produced; and (2) compare
the number of species flowering in burned and mowed
sites. We hypothesize that more flowering species will be
found in burned plots than in mowed ones, since fire is
more efficient in removing the above-ground biomass and
opening spaces within the vegetation (Fidelis et al. 2012).
Methods
Study area
This study was carried out in natural subtropical grasslands
located on a granitic hill in Southern Brazil (Morro San-
tana, 220 ha, 30°03′S, 51°07′W, 311 m a.s.l.). The climate
is subtropical humid, with annual rainfall of 1350 mm and
mean temperature of 22 °C (Livi 1999). The soils in the
study area are dystrophic red-yellow argisols (Garcıa-Mar-
tinez 2005).
Grasslands are located atop hills, composing a mosaic
with semi-deciduous seasonal forest and Atlantic Rain For-
est. Fire has been present in the study area throughout the
past 1200 yrs (Behling et al. 2007). Since cattle have not
been present for several years in the area, fire is the major
disturbance. Fire frequency is estimated at between
2–5 yrs, and it is set illegally mostly by local residents dur-
ing summer to maintain open areas for local community
practices, such as religious rituals and exploitation of local
plant diversity for medicines and handcrafts.
These grasslands are very rich in plant species (ca. 450–
500 plant species; Aguiar 1986) with a high diversity of
species belonging to Poaceae, Asteraceae, Fabaceae and
Rubiaceae (Overbeck et al. 2006), and a high occurrence
of below-ground organs (Fidelis et al. 2009; Fidelis et al.,
in press). There is a tendency toward the loss of grassland
species where fire has been excluded for many years, and
an increase in shrub cover and forest expansion (M€ uller
et al. 2007). Frequently burned areas are characterized by
Applied Vegetation Science
A. Fidelis and C. Blanco
a continuous grass matrix, with both C3 (e.g. Briza subaris-
tata, Stipa filiculmis) and C4 (e.g. Andropogon lateralis, Aristida
flaccida, Axonopus sulfuttus) species. These areas are also very
rich in forb species (e.g. Chaptalia runcinata, Richardia gran-
diflora, Evolvulus sericeus), and the presence of several small
grassland shrubs (e.g. Baccharis cognata, Porophyllum lanceol-
atum, Vernonia nudiflora). Areas excluded from fire for at
least 5 yrs show a higher percentage of shrub cover,
mainly Heterothalamus psiadioides, Eupatorium ligulaefolium
and Baccharis leucopappa (all Asteraceae). Despite the high
cover of shrub species, grasses still occur, with a higher
amount of standing dead biomass. A loss of forb species
can also be observed. Most graminoids (C4) and forbs
flower at the end of the spring season (November) and
throughout the whole of summer (December–March).
Fewer species can be observed flowering during winter. C3
grasses, on the other hand, flower earlier (end of winter,
during spring season).
Methods
In order to test our hypothesis about differences between
fire and mowing, we used experimentally burned and
mowed plots. To test if fire history affected flowering of
plant community and functional groups, we used plots in
sites with different fire histories burned at different times
(1 mo, 1 and 3 yrs after fire). Therefore, we had the fol-
lowing treatments: F30D – 1 mo after fire (five and 14 rep-
licates, for more details see description of the methods
below), F1Y – burned 1 yr ago (14 replicates), F3Y –
burned 3 yrs ago (five replicates), M30D – mowed 1 mo
ago (14 replicates), and M1Y – mowed 1 yr before observa-
tions (four replicates). We summarize the number of plots
per treatment in Table 1.
Abiotic data
TinyTag data loggers (Gemini data loggers, TGP-4500,
measurements every 5 s) were used to measure PAR (pho-
tosynthetically active radiation, lmol
m 2
s 1), tempera-
ture at soil level (°C) and relative air moisture (%). We
measured the abiotic data in two pairs of plots: F30D and
F3Y. Measurements were carried out over 7 d. Due to the
lack of replication (only two loggers per site), only abiotic
data will be described.
Table 1. Description of treatments, experimental design and number of plots.
Treatment Description
Fire 9 mowing Plots established in areas mowed (M30D, M1Y) and burned (F30D, F1Y)
during summer; observations were carried out 1 mo and 1 yr after experiments
Fire history Plots established in areas burned after 1 mo (F30D), 1 yr (F1Y) and 3 yrs (F3Y)
Applied Vegetation Science
Doi: 10.1111/avsc.12098 © 2014 International Association for Vegetation Science
Fire and flowering in subtropical grasslands
Dates of flowering observations
Fire and mowing experiments were conducted in natural
subtropical grasslands during the summer of 2006 (Decem-
ber). Fourteen plots were burned (F) and 14 plots were
mowed (M, total of 28 plots of 5 9 5 m) in patches of
grasslands with the same fire history. All phenological
observations were conducted in January 2007 (30D) and
2008 (1Y) (14 plots/treatment). Due to an accidental fire,
we lost ten of the 14 mowed plots and thus, 1 yr after
treatments, we had data for only four replicates (M1Y). In
addition to the experimental plots, we monitored flower-
ing in several patches of grassland burned during summer
2007 (late Nov, in a total period of 2 wks, F30D) and oth-
ers that had not been burned for 3 yrs (F3Y). These fires
were man-caused and are very common in these grass-
lands, mostly during summer. Thus, these plots were
established at sites burned by non-experimental fires
within the same 2-wk period of the summer, as were the
experimental fires. The non-experimental plots, however,
were burned in 2007 rather than in 2006. We included
vegetation sampling on the non-experimental fires in
2007 to determine if the trends seen in the experimental
plots in the summer of 2006 were comparable to responses
to fire in the summer of 2007.
Since our aim was to analyse flowering differences
caused by fire history and disturbance type, we did not
repeatedly visit all plots throughout the entire year, but
quantified flowering of all species flowering during the
peak flowering season for the majority of species, which is
the summer in these grasslands.
Fire 9 flowering species
Plots of 0.25 m2 (0.5 9 0.5 m) were established randomly
within the different patches burned during summer 2007
and inside the experimental plots, in order to evaluate the
effects of fire since time of last fire (30 d, 1 and 3 yrs). We
used 30 plots in sites F30D (six plots/site, total of five sites)
and 30 plots in sites F3Y (six plots/site, total of five sites),
and 14 plots for F1Y (one in each experimental plot: 14
replicates; Table 1). All species that were flowering at the
time of the observations (during summer) were identified
and counted. Later, species were grouped into graminoids
(C3 and C4 species: Poaceae, Cyperaceae and Juncaceae),
Number of plots
14 plots/area
30 plots (6 plots/replicate = total of 5 replicates)
for F30D and F3Y, and 14 plots for F1Y
3
Fire and flowering in subtropical grasslands A. Fidelis and C. Blanco

forbs (all other herbaceous species, except graminoids) and 1800

shrubs (all woody species <1.5-m high) for comparison 1600

among functional groups. 1400

PAR ( mol·m–2s–1)
1200
1000
Fire and mowing 9 flowering species
800

In order to analyse the effects of fire and mowing on the 600

flowering of species, we used only the experimental plots 400

(5 9 5 m, described above). We experimentally burned 200

14 plots and mowed the other 14 (at the soil level, at least 0

1 cm above soil, with litter removal after cutting the bio- 45

mass). Experiments were conducted during summer 40
(December–January; for more details see Fidelis et al. 35
2012). Thirty days after treatments (F30D and M30D), we 30
randomly established one plot (0.5 9 0.5 m, avoiding the 25
borders of the plots where fire burns irregularly) inside 20
each experimental plot, and identified and counted all spe- 15
cies that had flowers at the time of observation. The same 10
procedure was carried out 1 yr after experiments (F1Y and 5
M1Y). Species were also grouped in functional groups. 0
Since no shrub species had flowers at the time of observa- 120 F30D
tions, they were not further analysed to compare burned F3Y
100
Relative air moisture (%)

and mowed plots.

80

Statistical analysis 60

In order to evaluate the differences in the number of spe- 40

cies flowering, as well as the number of flowering stems
between treatments, we performed ANOVA using ran- 20

domization tests (10000 iterations), applied to Euclidean 0

distance. Randomization tests were chosen since there is 1 3 5 7 9 11 13 15 17 19 21 23

no restriction on normal distribution of data, as well as on Hours

the different number of sampling units used in this study

Fig. 1. (a) PAR (lmol
m 2
s 1), (b) temperature (°C), (c) relative air
(for more details, see Manly 2007). We used the software humidity (%) in sites with different fire histories: F30D (30 d after fire) and
MULTIV 2.5 (Pillar 2005) for all statistical analyses. F3Y (3 yrs after fire) in Brazilian Campos grasslands.

Results
Abiotic data
There was an increase of ca. 228% in PAR in sites burned
after 1 mo in comparison to sites excluded from fire for
3 yrs (Fig. 1a). Temperature at soil level also increased
7 °C in these sites (Fig. 1b), whilst relative air humidity
decreased 21% at 11:00 h. In general, relative air moisture
was 7% higher in sites excluded from fire for 3 yrs (Fig. 1c).
Fire history
Thirty days after fire, an average of three to four plant
species per plot had flowers (ca. 20% of species present in
the plots), but most plant species flowered 1 yr after fire
(P ≤ 0.05; Fig. 2a). The number of flowering species
decreased 3 yrs after fire in comparison to F30D
(P = 0.0002). The same tendency was observed for grami-
noid species: more species showed flowers 1 yr after fire
(34% of all graminoid species, P ≤ 0.05); the number of
species flowering after 3 yrs decreased (P = 0.0001;
Fig. 2b). Forb species, on the other hand, had a different
response to fire: more species were observed flowering
30 d after fire (F30D) in relation to F1Y (P = 0.007) and
F3Y (P = 0.0001; Fig. 2c), and the number of species
blooming decreased in the absence of fire (P ≤ 0.05).
Finally, shrub species showed no significant differences
between time since last fire (Fig. 2d). One year after fire,
no shrub species had flowers and only a few flowered 30 d
and 3 yrs after fire at the time of observations.
The C3 and C4 grasses showed the same tendency of
graminoid species from the analysis considering the
comparison between functional groups: more species

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4 Doi: 10.1111/avsc.12098 © 2014 International Association for Vegetation Science
A. Fidelis and C. Blanco Fire and flowering in subtropical grasslands

(a) (b)

(c) (d)

Fig. 2. Number of species flowering: total (a), graminoid (b), forb (c) and shrub (d) in sites under different fire histories: F30D – 30 d after fire, F1Y – 1 yr
after fire and F3Y – 3 yrs after fire in Brazilian Campos grasslands. Euclidean distance between sampling units was the resemblance measure used and
10,000 iterations were calculated for the randomization test. Different letters denote significant differences (P ≤ 0.05). The squares between boxes
represent the median, boxes represent 25% interquartiles and lines represent maximum and minimum values. Y-axis in the figures has different scales.

flowered 1 yr after fire than in plots with other fire histo- tively), Desmanthus tatuhyensis, Macroptilium prostratum
ries (P < 0.05; Table 2). More C4 grasses had flowers 1 yr (both Fabaceae, presence of xylopodium), Chaptalia runci-
after fire in comparison to C3 species (P = 0.003), but no nata (Asteraceae, absence of below-ground organs) and
significant differences could be found at F30D and F3Y Oxalis conorrhiza (Oxalidaceae, presence of bulbs). Other
(P > 0.05). species followed the general tendency and were found
As shown in Table 3, species from different functional flowering at higher frequency 1 yr after fire, such as Arist-
groups showed different responses to fire. Some species ida flaccida, Axonopus suffultus (C4 grasses) and Aspilia monte-
flowered only 30 d after fire, such as Calamagrostis viridifl- vidensis (Asteraceae, absence of below-ground organs).
avescens, Leptocoryphium lanatum (C3 and C4 grasses, respec- Several species were not found flowering in sites F3Y (e.g.
Stipa filiculmis, a C3 grass and Evolvulus sericeus, Convolvula-
ceae with no below-ground organs).
Table 2. Number of C3 and C4 grass species flowering in sites under three
different fire histories: F30D – 30 d after fire, F1Y – 1 yr after fire and F3Y –
3 yrs after fire in Brazilian Campos grasslands. Euclidean distance between Fire 9 Mowing
sampling units was the resemblance measure used and 10,000 iterations
were calculated for the randomization test. Different small letters denote As shown in Fig. 3a, more flowering species were found in
significant differences (P ≤ 0.05) between grass species (C3 and C4) and dif- burned than in mowed plots (P = 0.02). The number of
ferent capital letters denote significant differences between sites (F30D, species increased after 1 yr, and burned plots had a higher
F1Y and F3Y).
number of flowering species than mowed plots (P = 0.02).
Grass species Number of species flowering The same tendency was observed for graminoid species
(Fig. 3b): more species flowered in burned than in mowed
F30D F1Y F3Y
plots (at F30D – P = 0.04; and at F1Y – P = 0.0006). Never-
C3 0.24  0.09 aA 1.73  0.38 aB 0.31  0.11 aA theless, there were no significant differences in the
C4 0.48  0.11 aA 3.64  0.43 bB 0.55  0.18 aA
number of forb flowering species between treatments (fire

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Fire and flowering in subtropical grasslands A. Fidelis and C. Blanco

Table 3. Most common species flowering (frequency of individuals cies were found. Graminoid species showed the highest
flowering, %) in sites under different fire histories: F30D – 30 d after fire, number of species flowering 1 yr after fire, whilst forb spe-
F1Y – 1 yr after fire and F3Y – 3 yrs after fire in Brazilian Campos
cies had more species with flowers just after fire (F30D).
grasslands.
Although shrubs are suggested to have more plastic
Species Growth Below-ground F30D F1Y F3Y responses than herbaceous species, and thus presumably
form structure should be able to respond better to changes in the environ-
Briza subaristata C3 grass No 27.3 57.2 60 ment (Paritsis et al. 2006), our study showed that shrub
Calamagrostis C3 grass No 100 0 0 flowering was not affected by fire.
viridiflavescens Grasses are known to be the most fire-resilient compo-
Stipa filiculmis C3 grass No 0 50 0 nent in the plant community, due to their high capacity to
Aristida flaccida C4 grass No 0 92 13
resprout after fire (Bond 2004; Vesk & Westoby 2004).
Axonopus suffultus C4 grass No 12.5 67 27.8
Leptocoryphium C4 grass No 50 0 0
Most graminoid species in the study area are caespitose
lanatum grasses, with no storage organs (Overbeck & Pfadenhauer
Chaptalia runcinata Forb No 63.2 0 0 2007). Even though caespitose grasses have no storage
Evolvulus sericeus Forb No 54.2 30 0 organs, they can accumulate nutrients in soil just beneath
Aspilia montevidensis Forb No 0 40 0 clones (‘nutrient islands’; Derner et al. 1997; Briske &
Desmanthus Forb Yes 58.5 0 0 Derner 1998; Derner & Briske 2001), assuring resources
tatuhyensis
for growth and reproduction. Therefore, graminoids might
Macroptilium Forb Yes 75 0 0
prostratum
have used the high input of nutrients after fire to first
Oxalis conorrhiza Forb Yes 55.6 0 0 regenerate their above-ground biomass, whilst resource
allocation for reproduction costs was used in the following
year. A delayed response to fire concerning flowering was
and mowing) for both F30D (P = 0.26) and F1Y (P = 0.79; also observed for Aristida mohrii, which was related to time
Fig. 3c). of rainfall (Shepherd et al. 2012). However, in these sub-
tropical grasslands, rainfall is more or less well distributed
Discussion throughout the whole year. Hence, we believe grasses
would have a competitive advantage in post-fire environ-
Does fire induce flowering in the Brazilian Campos
ments due to their rapid build-up of biomass (Fidelis et al.
grasslands?
2012). Flowering therefore would be delayed for the next
Fire affected the number of species flowering in Brazilian growing season. Another interesting result was the flower-
subtropical grasslands, when we compared the recently ing of several C3 species 30 d after fire. These species flow-
burned plots (1 mo and 1 yr after fire) and those surveyed ered in spring, just before fire. After fire, they resprouted
3 yrs after fire. However, when one considers the effects of vigorously and flowered again; therefore fire itself does not
fire on the different functional groups, interesting tenden- seem to be detrimental to C3 species, as affirmed by some

(a) (b) (c)

Fig. 3. Number of species flowering – total (a), graminoids (b) and forbs (c) – under four different treatments: 30 d after treatment and 1 yr after
treatments with fire or mowing in Brazilian Campos grasslands. Euclidean distance between sampling units was the resemblance measured used and
10,000 iterations were calculated for the randomization test. Asterisks denote significant differences (P ≤ 0.05) between treatments (fire and mowing) for
each fire history. Squares between boxes represent median, boxes represent 25% interquartiles and lines denote maximum and minimum values. Y-axis in
the figures has different scales.

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A. Fidelis and C. Blanco
authors (Llorens & Frank 2004). Instead, fire season
should be noted as a limiting factor for the recovery of C3
grasses in these subtropical grasslands.
In subtropical grasslands, the number of forb species
decreases with longer fire intervals (Fidelis et al. 2010,
2012). A bud bank analysis showed that with 6 yrs of fire
exclusion, the bud bank of forb species significantly
decreased and thus they disappear from these grasslands
when fire is absent for longer periods (Fidelis et al., press).
Hence the availability of new microsites should be rapidly
colonized by forbs before grasses dominate and suppress
their growth. One main consequence of fire in these grass-
lands is the increase in bare soil and the removal of com-
petitors, in relation to areas where fire is excluded (Fidelis
et al. 2012). Since most of the forbs in these grasslands
have below-ground organs, such as bulbs, rhizomes and
xylopodia (Fidelis et al. 2009; Fidelis et al., in press), allo-
cation for both growth and reproduction assure their rapid
recovery and high proportion of species flowering.
Forb species with below-ground organs were able to
flower just after fire (Table 2). Studies on the effects of fire
on forb species populations in these grasslands showed sim-
ilar results: after fire, several individuals of Vernonia flexuosa
(presence of rhizophore) and Chaptalia runcinata (absence
of below-ground organ, both Asteraceae) were observed
(Fidelis et al. 2010). A fluctuation in fructan content in As-
teraceae was observed during the sprouting and flowering
phases, confirming this allocation of nutrients to growth
and flowering (Carvalho & Dietrich 1993). Finally, forb
species flowered earlier than other functional groups,
which can be an advantage in a post-fire environment
where competition is still low, there is an increase in bare
soil and thus seeds can reach the soil. Additionally, this
strategy can increase the accumulation of seeds stored in
the seed bank, assuring reproductive success for this func-
tional group if the next fire coincides with flowering or seed
maturation (Lamont & Downes 2011) or when the grass
matrix is already so thick that seeds cannot reach the soil.
Our results show differences between forbs and grasses
with respect to the timing of flowering following fire.
Regardless of the origin of taxonomic differences in flower-
ing phenology, however, an important ecological conse-
quence of burning in the summer could be the co-
existence of functional groups that show peak flowering at
different times since the last fire. Moreover, annual burns
in these grasslands may be harmful for C4 grasses, since
they delay their flowering until the next summer. The
results of the present study provide additional support for
the use of prescribed fire to maintain species diversity in
these grasslands and to support co-existence of different
functional and species groups by differences in their regen-
eration strategies and flowering. However, as suggested in
Hinman & Brewer (2007), we agree that very frequent
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Doi: 10.1111/avsc.12098 © 2014 International Association for Vegetation Science
Fire and flowering in subtropical grasslands
fires would have negative effects because some species
(such as graminoid species) showed delayed responses to
fire, flowering at higher frequency only 1 yr after fire.
Is mowing an alternative to fire?
As shown by our results, burned sites had a higher number
of species flowering than mowed plots. Our results differ
from those of Hulbert & Society (1988), where there was
no difference between the number of plants bolting after
fire and mowing. Besides, both fire and mowing open the
habitat and increase light interception on the soil surface.
Lamont & Downes (2011) suggested that the increase in
nutrient availability in burned patches is a relevant factor
for flowering.
The presence of below-ground organs in several forbs
species might have led to lack of differences in number of
species flowering between burned and mowed plots. Forb
species presented no differences in number of species flow-
ering between mowing and fire experiments. The removal
of above-ground biomass itself might have triggered
resource allocation to growth and reproduction. Grami-
noid species, on the other hand, did not show the same
pattern, and thus fire may have enhanced the flowering,
probably through the input of nutrients, since most of the
graminoids present in the area have no rhizomes (reserve
organs).
Several factors can trigger flowering after fire, such as
increase in light (Hulbert & Society 1988; Brewer 1995;
Tyler & Borchert 2002; Paritsis et al. 2006; Borchert &
Tyler 2009), temperature (Elzinga et al. 2007; Keeley
1993; Paritsis et al. 2006) and soil nutrients (Saulnier &
Reekie 1995; Wrobleski & Kauffman 2003; Sardans et al.
2005; Coates et al. 2006; Elzinga et al. 2007). The increase
of light (228%) and temperature (7 °C) could also be
observed in our plots 30 d after fire, and might have influ-
enced flowering in these plots. We did not measure light
and temperature in mowed plots, and both factors can
influence the differences found in our experiments.
Although most of the litter is removed after mowing, the
amount of bare soil in these plots was lower than in burned
plots 30 d after experiments (Fidelis et al. 2012), and
therefore burned plots probably received more light than
mowed ones. Another important factor would be smoke
(Lamont & Runciman 1993), which has a large effect on
geophytes, grasstrees and cycads (Lamont & Downes
2011). Therefore, smoke should be taken into account as a
possible cue related to flowering in these grasslands, since
some species are typical geophytes, such as Habranthus gra-
cilifoilius (Amaranthaceae), which flower only after fire
and were not observed in any other plot 1 yr after burning.
The input of nutrients in post-fire environments might
also explain the increase in flowering, as shown in some
7
Fire and flowering in subtropical grasslands
studies (Lamont & Runciman 1993). Nutrient addition in
combination with clipping and increased light can trigger
flowering (Brewer 1995). On the other hand, if no nutri-
ents were added and repeated clipping (annual) was per-
formed, a larger allocation to reproduction is observed at
the expense of vegetative propagation in some grass spe-
cies (Brewer et al. 2009). To flower, plants re-allocate
nitrogen and phosphorus stored in leaves, roots and stor-
age organs, thus reducing vegetative growth (Saulnier &
Reekie 1995; Wyka & Galen 2000), given that carbohy-
drate storage is not affected (Wyka & Galen 2000). Hence,
fire may be a better management tool for inducing flower-
ing in subtropical grasslands than mowing, not only
because it is more effective at stimulating flowering in
graminoids than is mowing, but also because repeated
flowering triggered by repeated mowing could lead to
increased nutrient limitation of clonal growth. Moreover,
after fire, there is probably an input of nutrients to the soil,
which could be beneficial for plants and positively affect
the build-up of flowering structures.
Conclusions
Our study was one of the few to demonstrate the effects of
fire on the flowering of an entire plant community, reveal-
ing interesting strategies of forbs and graminoids. Forbs
take advantage of the availability of new microsites to
establish and flower when the strong competitors (grami-
noids) are reduced. Graminoids, on the other hand, invest
more in vegetative growth just after fire, re-allocating
more resources to flowers 1 yr after fire. Summer burns
stimulated C3 grasses to flower again, as most of grami-
noids and forbs in subtropical grasslands flower at the end
of spring season and throughout the whole summer per-
iod. Few species flower during winter, and most C3 grasses
flower before the C4 species, at the end of winter and
beginning of spring. Although mowing might be an accept-
able alternative to fire management of these grasslands,
our study corroborates previous reports: mowing is not as
efficient as fire in stimulating flowering and plant growth,
probably because of the effects of smoke, the increase in
light, temperature and/or N and P availability in post-fire
environments. Therefore, in addition to faunal-saving
strategies and monitoring plans to assure the maintenance
of community diversity, prescribed and well-controlled
fires at the correct frequency and season could be consid-
ered an alternative management tool for these grasslands.
Acknowledgements
We are grateful to the security guards of the Universidade
Federal do Rio Grande do Sul (UFRGS) for giving us their
support during fieldwork. We also thank Ilsi Boldrini for
8 Doi: 10.1111/avsc.12098 © 2014 International Association for Vegetation Science
A. Fidelis and C. Blanco
helping with plant identification. We thank Alexandre
Igari and Letıcia Dadalt, and three anonymous reviewers
for their valuable suggestions on earlier drafts of this man-
uscript. Finally, we thank Andrea Skiba for the English
review of the manuscript. A.F. received financial support
from KAAD.
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