Progress in Physical Geography

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Physical Geography
Fire in the cerrado (savannas) of Brazil: an ecological review
Jayalaxshmi Mistry
Progress in Physical Geography 1998 22: 425
DOI: 10.1177/030913339802200401
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Progress in Physical Geography 22,4 (1998) pp. 425±448
Fire in the cerrado (savannas) of
Brazil: an ecological review
Jayalaxshmi Mistry
Department of Geography, Royal Holloway, University of London, Egham,
Surrey TW20 0EX, UK
Abstract: Fire is a major determinant of cerrado (savanna) vegetation in Brazil, and is used as a
management tool during the dry season. This has consequently induced a long history of fire
studies in the region. This article reviews past and present fire ecology studies in the cerrado, and
emphasizes the need for a more applied approach to future work.
Key words: fire; cerrado; Brazil; savanna.
I
Introduction
The Brazilian savanna, commonly called cerrado (meaning `closed' in Portugese) is a
complex vegetation form, characterized by a mosaic of physiognomies ranging from pure
grasslands through open scrubland to dense woodlands (Eiten, 1972; 1978) (Figure 1). It
occupies over 1.8 million km2, 22% of the Brazilian territory (Goodland, 1971a; Coutinho,
1990), and in terms of area, is second only to the Amazonian rain forest (Furley and
Ratter, 1988). The cerrado is centred on the Brazilian Planalto, characterized by a realm of
plateaux and high tablelands (termed chapadas) ranging in altitude from approximately
300 m to 1000 m above sea level (Ab'Saber, 1971).
Fire is a major determinant of the cerrado. Wildfires have been significant in the cerrado
since at least the middle Holocene, some 6000 years BP (Vernet et al., 1994), and indigenous people have been using fire for more than 32 000 years in central Brazil (Guidon
and Delibrias, 1986; Vicentini, 1993). Vegetation was burned in hunting and tribal wars
(Lukesch, 1969; Villas-Boas and Villas-Boas, 1976), to limit the growth of certain undesirable plant species, and to stimulate the production of certain native fruit-bearing
trees (Anderson and Posey, 1985; 1987). Today, the principal cause of fire in the cerrado is
through agricultural activities (Alho and Martins, 1995).
Vast areas of the cerrado are cleared and burned at the end of the dry season (August±
September) to plant crops such as soyabean, corn, rice, beans and cassava (Klink et al.,
1993). The more open forms of the cerrado (utilized as natural pastures) and planted
c Arnold 1998
*
0309±1333(98)PP199RA
426
Figure 1
cerradaÄo
The structural gradient of cerrado vegetation, from campo limpo to
pastures are frequently burned by cattle ranchers to promote fresh grass growth during
the dry season (Coutinho, 1990). Subsistence farmers burn arable areas to clear them of
weeds and other harvest residue during the dry season, in preparation for planting at the
beginning of the wet season (Mocelin, 1996).
Other burns arise from various causes, such as the control of shrubs in pastures, pest
control, negligence in fire management of intentionally burned areas (such as during the
cutting and burning of vegetation while cleaning the edges of highways and railroads),
and the falling of balloons with the wicks still alight during the June religious festivals
(Coutinho, 1990). Although carelessness with cigarettes does not seem to be relevant in
the cerrado, arson is common, mostly by local people who burn the vegetation for the
aesthetic value fire has (Coutinho, 1990).
As a major force in the cerrado, fire has been discussed ever since the classic works of
Saint-Hilaire (1824; 1827), Warming (1892) and LoÈfgren (1898). Today, the study of fire in
the cerrado is seen as increasingly important: both the population and agricultural
activities have grown substantially in the last 35 years; delimited areas of conservation,
such as national parks and reserves, are constantly invaded by criminal/accidental
wildfires; biodiversity is being threatened; there is global concern of greenhouse
warming, and the contribution of vegetation burning in the release of carbon dioxide;
and lack of research has obstructed fire management plans/policies being developed for
natural areas (Alho and Martins, 1995). This article will review the various aspects of fire
research in the cerrado, summarized in Figure 2. Generally, these can be divided into
abiotic, biotic, social studies and management studies.
II
Abiotic studies
Abiotic studies of fire in the cerrado have been divided into four main areas: the physical
nature of fire behaviour; the effects of fire on soil physical properties; the effects of
fire on nutrient cycling; and the types and quantities of gaseous particles released during
fires.
J. Mistry 427
Figure 2 The different aspects of cerrado fire research
1
Fire behaviour
Fires in the cerrado are characteristically surface level and fast-moving, reaching speeds
of 30 m min71 (Kauffman et al., 1994) and consuming the herbaceous layer, but rarely
igniting the taller woody plants (Miranda and Miranda, 1993) (see Figure 3). Variables
describing fire behaviour include fire intensity, fire temperature, its duration (residence
time) and its spatial distribution, rate of fire spread, the pattern of fire spread, and flame
height (Whelan, 1995). Factors affecting these variables are fuels and their properties,
Figure 3 Fires in the cerrado
428
climate and topography (Whelan, 1995). The cerrado landscape is characterized by gently
sloping hills (Ab'Saber, 1971), and in view of this fact, topography probably plays an
important role in fire spread, which increases with degree of upslope due to fuel
preheating (McArthur, 1971). Unfortunately, research has yet to be carried out on this
aspect of fire behaviour in the cerrado, although modelling of fire spread in cerrado
landscapes is beginning (Roque et al., 1996). Fire behaviour in the cerrado has predominantly been studied in relation to climate, and fuels, which are intrinsically related
to physiognomy.
a Climate and fire: The prevailing climatic conditions at the time of the burn will
greatly affect the behaviour of the fire. This varies during the dry season. Generally, fires
during the early season (May±June) are patchy and of a low intensity, due to the high
moisture still present within the vegetation from the rainy season (Miranda and Miranda,
1993). The mid-season fires (July±August) are of a higher intensity and more homogeneous, since most of the combustible fuel may be dry, and they attain a maximum peak
in the late season of August±September (Coutinho, 1990). This period is particularly
favourable for the propagation of fire, as relative air humidity during the hottest
hours of the day (25±30 8C) can reach below 20%, and the days are very windy
(Coutinho, 1982) (see Table 1). In years of frosts, a great part of the epigeous phytomass
in the herbaceous/undershrub stratum dies. This, and the accompanying fall of leaves
from many trees and shrubs, accumulate on the soil as an easily dehydratable and highly
combustible material, greatly increasing the risk of fire (Coutinho, 1990). As the wet
season starts (September±October), the occurrence of fires drops markedly, and although
prescribed burning does not take place, the vegetation is still susceptible to burning,
particularly in areas where there have been no burnoffs for several years, and/or after a
sequence of hot days in the absence of rain (veranico) (Cochrane et al., 1988). Fire during
this period, however, is normally patchy, and of a low intensity, and frequently self
extinguishes rapidly (Miranda and Miranda, 1993) (see Table 1, September 1992).
b Fuel and fire: Combustible fuel in the cerrado consists of grass, leaves, twigs and
branches, but due to their high degree of flammability, grasses and other ground-layer
vegetation are considered the major source of combustible material in different cerrado
physiognomies (Kauffman et al., 1994). Although total fuel biomass is significantly
Table 1 Fire behaviour of prescribed burns in campo sujo during August and September of
1992 (vegetation subjected to 18 years without fire) and 1994 in the IBGE Reserve, Brasilia, DF
August
September
Fire behaviour parameters
1992
1994
1992
1994
Days without rain
Air temperature (8C)
Relative humidity (%)
Wind speed (m s71)
Speed of ®re front (m s71)
Fire intensity (kJ ms71)
96
26.5
29.2
1.0±1.9
0.13
1390
53
25.0
30.0
2.7
0.42
5889
2
26.6
38.4
1.5
0.15
1256
96
30.4
23.2
2.7±2.9
0.64
8134
Source: Modi®ed after Miranda et al. (1996).
J. Mistry 429
greater in cerrado sensu stricto and cerradaÄo, the biomass of grasses is considerably higher
in campo limpo and campo sujo (Coutinho, 1982; Pivello and Coutinho, 1992; Ward et al.,
1992; Miranda and Miranda, 1993; Kauffman et al., 1994), and this is reflected in the fire
behaviour of the forms.
For example, in a campo sujo, where grasses can represent up to 91% of the combustible
fuel, Ward et al. (1992), Cesar (1980) and Berardi (1994) measured temperatures of over
800 8C, whereas in cerrado sensu stricto, Miranda et al. (1993) recorded significantly lower
temperatures. Flame height and fire intensity have also found to be greatest in campo
limpo and campo sujo (Kauffman et al., 1994). Virtually no smouldering combustion
following flaming combustion was detected in either campo limpo or campo sujo (due to
the low woody biomass), compared to campo cerrado and cerrado sensu stricto, where
smouldering combustion was prevalent, suggesting that fire has a greater influence in
the open cerrado forms, and that more woody plants will be killed, rather than scorched,
in the closed physiognomies (Kauffman et al., 1994).
Differences in fire behaviour along the gradient of cerrado physiognomies can also be
explained according to fuel moisture content. Kauffman et al. (1994) found that fuel
moisture content was very low in dry, nongreen grass (22±29% dry weight basis),
comprising 78% of the fuel mass in campo limpo and 40% in campo cerrado. In contrast,
fuel moisture content of the woody component ranged from 118 to 140% (dry weight
basis) ± 28% of the fuel mass in campo cerrado yet less than 6% in campo limpo. Total mass
of water in fuels was calculated to be 5 3100 kg ha71 in campo limpo and campo sujo, and
5 4600 kg ha71 in campo cerrado and cerrado sensu stricto. The differences in moisture
content affect the ignitability of fuels, thus possibly contributing to the lower impact of
fire on the tree-dominated communities, in comparison to the grasslands (Kauffman et al.,
1994).
The density of phorophyte individuals, and their effects on microclimate in the
different physiognomies, affects the behaviour of fire in the cerrado. Miranda et al. (1993)
recorded a maximum fire temperature of 260 8C in an area of cerrado sensu stricto which
had been protected for 15 years and burned three days after rain. Many patches of the
vegetation remained unburned. In contrast, campo sujo burned on the same day under the
same conditions, left only the woody vegetation unburned and attained temperatures in
the region of 650 8C. Shading of the fine fuel on the ground by trees and shrubs affects
the rate of fuel moisture loss, causing mosaic-like burns in the more closed cerrado types
(Miranda et al., 1993). In the open campo sujo, most of the dry matter is not in close
contact with the wet soil surface, and is well exposed to wind and solar radiation, so
moisture is quickly lost to the environment (Luke and McArthur, 1978; Miranda et al.,
1993).
`Time-since-last-fire' also determines the characteristics of cerrado fires. Areas protected
from fire for long periods of time will burn at higher temperatures than those areas
burned regularly, regardless of their physiognomic type, due to the build up of
combustible fuels (Miranda and Miranda, 1993; Miranda et al., 1993).
The behaviour of fire may be related to the height of the combustible fuel within the
herbaceous layer. In an analysis of the spatial distribution of temperatures, Miranda et al.
(1993) found that regardless of cerrado physiognomy, at 1 cm above the ground, 60 cm
height and 160 cm height, maximum temperatures ranged from 85 to 326 8C, 180 to 84 8C
and 107 to 650 8C, respectively. Also, the residence time above 60 8C varied from 90 to
270 seconds at 1 cm above the ground, 90 to 200 seconds at a 60 cm height, and 20 to
70 seconds at a 160 cm height. Other data confirm these results, which suggest that the
430
highest temperatures occur between 1 to 60 cm above the ground, and that the residence
time above 60 8C also peaks at this height range (Miranda and Miranda, 1993). Above
60 cm, fire temperatures decrease, with temperatures reaching peaks of up to 700 8C for
short periods of time.
2
Eects of soil physical properties
The degree of heating of the soil during a fire is a function of various factors such as
combustible phytomass per unit area, phytomass humidity and soil humidity (Coutinho,
1990). Most soil temperature measurements during cerrado fires have found relatively
small increases, e.g., around 50 8C, decreasing exponentially with depth, and becoming
more or less negligible at and below 5 cm depth (Coutinho, 1976; 1978; Cesar, 1980;
Miranda et al., 1993). These insignificant soil temperature changes are irrespective of the
physiognomic form being burned, and the maxima observed are unlikely to have any
direct effect on soil organic matter, microbial population or buried seeds (Miranda et al.,
1993).
The indirect effects on soil temperature of fires, however, may be important. Castro
Neves and Miranda (1996) found that between 10:00 a.m. and 2:30 p.m., the albedo in an
area of campo sujo reached 0.11. This value was reduced to 0.03 after the area was burned,
during which 94% of the vegetation was consumed. This decrease in albedo represented
an increase of about 10% in the energy absorbed. One month later, the albedo had only
reached 54% of the prefire value. This increase in the quantity of absorbed solar energy
could alter the soil microclimate, affecting soil micro-organisms and processes such as
seed germination (Frost and Robertson, 1987).
Soil exposure after vegetation burnoff could also increase the amplitude of the daily
thermal variation in the postburning period, and so influence soil processes. In the
same experimental fire described above, Castro Neves and Miranda (1996) measured the
soil heat flux, and found that it changed from 55.3+ 1.7 W m2 before the fire to
74.7+ 2.4 W m2 after the fire.
3
Eects on nutrient cycling
Generally, the soils underlying the cerrado are rather poor in mineral nutrients, are
distinctly acidic and have high levels of aluminium (Goodland, 1971b; Lopes, 1975;
Lopes and Cox, 1977). Fire is intimately related to this nutritional status since it is
involved with the cycling of mineral nutrients (Coutinho, 1990). Through the action of
fire, most of the epigeous biomass is rapidly mineralized, with nitrogen, carbon, sulphur
and to a lesser extent, phosphorous and potassium, volatilized and lost to the atmosphere, and the remaining material either deposited on the soil surface as ash, or removed
as particulate matter in smoke (Frost and Robertson, 1987).
In the cerrado, the immediate effects of burning result in a temporary increment at the
soil surface (0±5 cm) in concentrations of calcium, magnesium, phosphorus and
potassium, and the complete disappearance of aluminium, which can remain at zero
levels for up to 40 days (Cavalcanti, 1978; Coutinho, 1982). Deeper down in the soil there
is no change in these nutrients or in aluminium levels, suggesting that the ash deposited
on the topsoil is highly beneficial to the growth of herbaceous/undershrub plants
with superficial root systems, since they are provided with a large quantity of mineral
nutrients and a significant reduction in aluminium toxicity (Coutinho, 1990).
J. Mistry 431
However, for the tree/shrub layer which generally possesses deep root systems
(Rawitscher, 1942a; 1942b; Rawitscher et al., 1943), fire is detrimental, since nutrients
made available from burned leaves, flowers, fruits and branches are mostly transferred to
the herbaceous layer (Cavalcanti, 1978; Batmanian, 1983). Coutinho (1984) suggests that
compensation for this loss of nutrients from the tree/shrub vegetation is brought about
through the action of leaf-cutter ants (Atta sp.), which are frequently encountered in the
cerrado at a density of two to three nests/ha or more. Through their foraging activity,
nutrients are transported to chambers at depths of 6±7 m in the soil, and although this
process is limited to where the ants have their nests, in the long term, nutrients will be
absorbed by the deep roots of the tree/shrub stratum, and may determine cerrado
vegetation at the patch scale (Coutinho, 1984).
With the great disparity in cerrado forms, a variation in the relationship between fire
and nutrient dynamics along the physiognomic gradient would be expected. Kauffman
et al. (1994) looked at this association, and found that though the pool size of nitrogen,
carbon and sulphur (within the fuel load) increased along the gradient from campo limpo
to cerrado sensu stricto, the percentage of those nutrients lost by fire decreased. For
example, along the gradient, total mass of nitrogen increased from 24 to 55 kg ha71, but
nitrogen lost by fire was greater or equal to 90% of the pool in the grasslands, in
comparison to less than 56% in the tree-dominated communities. This is probably due to
the significantly greater amounts of readily accessible combustible fuel in the grasslands,
while nutrients remain locked up in the woody plants, which rarely burn (Pivello and
Coutinho, 1992; Kauffman et al., 1994).
Other data showed that in cerrado sensu stricto, greater quantities of nutrients were lost
as particulates, whereas in campo limpo, most of the nutrients were volatilized during fire
(Kauffman et al., 1994). These levels of losses through volatilization are especially
important because they are likely to be ecosystem losses (Raison et al., 1985) in contrast to
particulate losses, which may be redistributed within the ecosystem. However, the
amount of nutrients lost in the cerrado due to fire represents a minor proportion of the
total pool (in roots, soils and above-ground woody vegetation), and is likely to be
replaced through natural inputs, particularly through rainfall (Coutinho, 1979; Schiavini,
1984), in one to three years (Pivello-PompeÃia, 1985; Kauffman et al., 1994).
4
Eects on atmospheric emissions
Fires in tropical savannas are a major source of particulate matter and gaseous emissions
to the atmosphere (Crutzen and Andreae, 1990; Hao et al., 1990). Some emitted gases
such as CO2 , CH4 and CH3CL cause atmospheric warming, contributing to the greenhouse effect (Ramanathan et al., 1985), whilst others such as CO and NOx are involved in
complex chemical reactions in the troposphere, causing elevated levels of ozone and acid
precipitation (Crutzen, 1988; Crutzen and Andreae, 1990). Using techniques such as
satellite remote sensing, various workers have measured atmospheric emissions from
cerrado burning (e.g., Ward et al., 1992; Poth et al., 1995). In comparison with typical
tropical forest, and North American forest burns, results indicate that the cerrado has
lower emission factors, suggesting the contribution of savanna fires to global emissions
to be smaller than expected (Kaufman et al., 1992). However, correlations show a direct
link with burning and increased levels of ozone (Kirchoff and Alvala, 1996).
The chemical precursors necessary for ozone formation are believed to be provided by
biomass burning, and layers of ozone concentration enhancements have been observed
432
over cerrado regions during the dry season (Kirchoff and Marinho, 1994). Further evidence comes from studies of the large ozone bulge in the South Atlantic Ocean, near the
African coast. Preliminary studies at Natal, Brazil (Kirchoff and Nobre, 1986) and later in
the South Atlantic (Watson et al., 1990; Fishman et al., 1990; 1991) indicate the bulge to be
the result of transport and chemistry inducement by biomass burning in nearby South
America and African regions. Trajectory analyses also show that it is the Brazilian
sources which contribute to the South Atlantic tropical portion of the ozone bulge at
higher levels (above 500 hPa), whereas lower portions of troposphere in the South
Atlantic from coast to coast receive burning products from Africa (Fuelberg et al., 1996).
III Biotic studies
1 Plant community studies
The origins and the principal determinants of the cerrado vegetation have been discussed
since the last century (Saint-Hilaire, 1824; Warming, 1892). Fire has played an important
role in these deliberations. Several authors, such as Rizzini and Heringer (1962) and
Rizzini (1963), have considered the cerradaÄo as the forest type in the whole cerrado area,
the other structural types having been derived by human activities, particularly burning.
Though this hypothesis may be applicable to some restricted areas, it is improbable that
the whole cerrado region is determined by ancient anthropogenic burning (Sarmiento,
1983).
In the successional paradigm, many authors, such as Rawitscher (1942a) and Ferri
(1944), thought that the physiognomic gradient was determined by fire, with cerradaÄo
representing the `fire climax'. However, in the new conceptual framework of savanna
modelling (Frost et al., 1986; Medina, 1987; Goldstein et al., 1988), there is no one single
cause that governs cerrado formation. An approach employing `hierarchy theory'
(Solbrig, 1991) stresses the relative importance of all savanna determinants, at different
spatial and temporal scales. Fire is a key determinant of the cerrado, and is significant at a
range of scales (see Figure 4). It affects vegetation composition and structure and individual species, many of which have adapted to fire over time.
a Effects of fire on the woody layer: The death of established woody plants by fire is a
rare phenomenon in the cerrado (Ramos and Rosa, 1992). Many species possess pyrophytic characteristics, the most notable being the strong suberization of trunks and
branches, permitting thermal isolation of living internal tissues (Eiten, 1994). Some trees
protect their dormant apical buds using dense, hairy cataphylls, e.g., Anemia anthriscifolia
(Rachid-Edwards, 1956). In contrast, others have exposed dormant apical buds, and
these frequently die during the fires. A few days later, adventitious buds may sprout
from the branches, resulting in sympodial growth of the stems (Eiten, 1982). This imparts
the most characteristic feature of the taller shrubs and trees, which enables a cerrado to be
recognized on sight ± its tortuosity (Eiten, 1994). Other fire adaptations include the
capacity to produce vigorous sprouts from subterranean roots following the total
carbonization of the aerial branches (Rachid-Edwards, 1956). Even the seedlings of
certain tree species may present this type of adaptation (Dionello, 1978).
Still, for the majority of less established plants, such as seedlings and young
individuals, fire is detrimental, causing the total destruction of aerial parts (Hoffman,
1996). The subsequent reduction in tree recruitment favours more open forms of cerrado,
J. Mistry 433
Figure 4 A hierarchy model of cerrado key determinants. (Note: Determinants
at one level in the hierarchy are constrained by those above. Determinants at
the same level of the hierarchy interact frequently and strongly)
Source: Mistry, 1996, Modified after Solbrig, 1991
whereas protection from fire allows the woody vegetation over the herbaceous to establish, and succession continues to the closed cerradaÄo when other factors are not limiting
(Ferri, 1973; Ratter et al., 1973; 1978; Ratter, 1991; Henriques, 1993). This hypothesis was
confirmed by Moreira (1992; 1996) in central Brazil, using line intercept transects to
sample frequently burned and fire-protected areas of different cerrado physiognomies.
Table 2A shows the average number of individuals per transect for the different physiognomies. It indicates that fire protection does lead to an increase in woody elements in all
physiognomies. Ramos (1990) likewise found that woody plants doubled in density in
areas protected from fire.
Table 2 Average number of woody individuals and species per transect. Standard errors are
shown in parentheses and the significant difference ( p 5 0.05) between protected and
unprotected for each physiognomy are indicated with letters
A) No. of individuals
B) No. of species
Physiognomy
Protected
Protected
CerradaÄo
Cerrado sensu stricto
Campo cerrado
Campo sujo
349
323
246
257
(+ 43.9)b
(+ 7.0)b
(+ 12.9)b
(+ 17.9)b
Unprotected
268
246
182
178
(+ 22.1)a
(+ 11.4)a
(+ 6.4)a
(+ 11.1)a
58
54
49
48
(+ 3.2)b
(+ 0.7)a
(+ 3.2)b
(+ 0.3)b
Source: Modi®ed after Moreira (1996).
Unprotected
46
52
43
44
(+ 2.6)a
(+ 3.5)a
(+ 0)a
(+ 0.5)a
434
In addition, fire has been shown to affect the structure of cerrado vegetation. Moreira
(1992; 1996) found that the number of trees taller than 2 m was greater in fire-protected
areas, regardless of physiognomy. Ramos (1990) discovered that the basal area of trees
was two times greater in areas protected from fire for 13 years compared to those burned
every two years. Mistry (1996), studying cerrado denso vegetation in central Brazil, also
detected similar trends. Fire protection led to a greater tree height, as well as an increase
in individuals with thicker trunks and low-level branches.
With the occurrence of fire, these structural characteristics of the vegetation are
modified. The number of small individuals (0±50 cm in height) increases in all physiognomies except the campo sujo, subshrubs growth forms become more common (Moreira,
1992), lower branches are eliminated reducing structural complexity, and there is a
decrease in individuals with large girths (Mistry, 1996). The difference registered for the
campo sujo above confirms observations made for other cerrado areas (Coutinho, 1982;
1990): it seems that fire is important for the physiognomic equilibrium of some open
forms like the campo sujo. Fire protection thus increases the presence of thick-stemmed
phorophytes that are multibranching near the base (Eiten, 1994), and the overall
structural complexity of the vegetation (Mistry, 1996), but as fire frequency increases, the
height of the vegetation is lowered, large-girthed trees are eliminated and structural
complexity is diminished (Moreira, 1992; Mistry, 1996).
Patterns of spatial distribution among woody plants also seem to alter as a
consequence of fire. Sambuichi (1991) demonstrated that trees and shrubs change from
a random distribution in periodically burned areas to an aggregated distribution in fireprotected areas. The greatest increase in terms of frequency of individuals is in the
16±63 cm height class and the 3±11 cm circumference class (Ramos, 1990). Seed dispersal
and plant establishment in many species may be local in relation to the mother plant,
confirming the idea that it is the younger individuals that are most affected by fire
(Hoffmann, 1996).
This effect of fire on seedlings and their establishment has been investigated through a
small number of studies of particular woody species. Oliveira and Silva (1993), working
with two species of Kielmeyera, showed that seedlings had a high survivorship despite
fire occurrence in the first year. In a study of Dalbergia miscolobium, fire was attributed a
major mortality factor for seedlings less than one year old, but not for older seedlings,
which could resprout from basal buds (Franco et al., 1996). Hoffmann (1996) surveyed
12 cerrado species, and found that burning had an overall negative effect on seedling
establishment in the first year following fire, which may be attributed to higher
temperature and desiccation exposure following woody cover removal. But by the
second year following fire, establishment returned to control levels.
The cerrado has the richest floral diversity of all the world's savannas (Sarmiento, 1983;
Eiten, 1994), and according to Heringer et al. (1977), 774 woody plant species are known.
Frequent fires, however, have shown to diminish the species diversity of cerrado woody
vegetation. Table 2B shows the number of woody species in protected and frequently
burned cerrado physiognomies, and indicates the decrease in diversity with fire (Moreira,
1992). Similar findings were recorded by Sambuichi (1991).
In addition, fire can induce changes in species composition. In the cerrado, the
differences in woody species composition between protected and unprotected are most
striking in the cerradaÄo. Moreira (1992) observed the absence in a burned area of five of
the ten most abundant species in the protected area, indicating the fire sensitivity of this
physiognomy. Emmotum nitens, Ocotea spixiana and Alibertia edulis are typical cerradaÄo
J. Mistry 435
Table 3
Example of species belonging to different `fire-sensitivity' groups
`Fire-sensitivity'
categories
Species
Group description
Species Ì'
Rapanea guianensis, Miconia albicans, Miconia
fallax, Miconia burchellii, Salacia crassifolia,
Protium ovatum, Byrsonima crassa,
Blepharocalyx suaveolens
Very sensitive to ®re
Species È'
Roupala montana, Miconia ferruginata,
Miconia pohliana, Eremanthus glomerulatus,
Didymopanax macrocarpum, Qualea multi¯ora,
Aspidosperma macrocarpum, Couepia
grandi¯ora, Dalbergia violacea, Styrax
ferruginea, Connarus fulvus, Davilla elliptica,
Neea theifera, Syagrus comosa
Sensitive to ®re but can sustain
a rare ®re
Species `P'
Syagrus petraea, Piptocarpha rotundifolia
Pro®t from ®re
Species `T'
Byrsonima verbascifolia, Byrsonima
coccolobifolia, Aspidosperma tomentosum,
Erythroxylum tortuosum, Palicourea rigida,
Vellozia ¯avicans, Acosmium dasycarpum,
Tabebuia ochracea, Machaerium opacum,
Tocoyena formosa
Fire tolerant
Source: Modi®ed after Sambuichi (1991).
species, and were conspicuously absent in the burned site. Moreira also identified two
species of fire-sensitive shrubs: Miconia albicans and Miconia cf pohliana. Shrubs were
always considered `typically pyrophytic' (Coutinho, 1990), yet these results propose that
it may not only be tree species that are most susceptible to fire (Moreira, 1992).
Sambuichi (1991) assigns `fire-sensitivity' labels to cerrado woody species to explain the
compositional differences between fire-protected and burned areas in central Brazil
(Table 3). Fire-protected sites comprise species extremely sensitive to fire (designated Ì'),
and species which are sensitive to fire but can tolerate a rare fire (designated È'). The
majority of these species are zoocorica, and they are frequently encountered in the
cerradaÄo formations of the region (Sambuichi, 1991).
Species characteristic of burned areas are categorized as pyrophytic (designated `P')
when they benefit from fire, and tolerant (designated `T') if they can sustain fire.
Comparing these groups of species, Sambuichi (1991) found that in a periodically burned
area, 60% were of groups `P' and `T', and 40% of Ì' and È', whereas in a protected
area, 30% were of groups `P' and `T' and 70% of Ì' and È'. Evidently, long-term fire
suppression alters the composition of the cerrado and augments its susceptibility to fire
(Sambuichi, 1991).
b Effects of fire on the herbaceous layer: The herbaceous flora is dominant of the
open cerrado forms, and is considered to be `typically pyrophytic' (Coutinho, 1990).
Consequently, many studies of this strata have concentrated on the adaptations various
species have to fire. Some have annual life cycles, growing and developing during the
436
rainy months, thus escaping the dangers of dry-season fires as seeds (Coutinho, 1990).
Many perennial species possess subterranean organs such as bulbs, underground shoots,
rhizomes and xylopodia, which avoid damage from fire (Coutinho, 1982). The densely
imbricated sheaths of some grasses provide protection by limiting combustion due to
inadequate aeration, e.g., Aristida pallens (Rachid-Edwards, 1956). Some woody species
present in the herbaceous stratum develop their entire system of trunks and branches
subterraneously, with only the small vegetative branches or yearly reproductive sprouts
protruding above the soil. This example of cryptophytism can be found in trees such as
Anacardium pumilum and Andira humilis and among palms such as Acanthococos emensis
and Attalea exigua (Rawitscher et al., 1943; Rawitscher and Rachid, 1946; Lopez-Neranjo,
1975).
Burning induces flowering in many species, such as in the orchid family (Oliveira et al.,
1996), and fruit dehiscence in many others (Coutinho, 1982). Some seeds of the Mimosa
genus require a thermal shock in order to germinate (Coutinho and Jurkewics, 1978;
Coutinho, 1982; Almeida and Silva, 1989). Coutinho (1977) postulates that fire has a
beneficial effect on these species by cleaning out obstructing vegetation, thus facilitating
pollination and seed dispersion. Several species possess special woody underground
organs termed `lignotubers' or `xylopodia', which are thick and lignified, and have
dormant buds in their upper region at ground level or some centimetres below
(Coutinho, 1982). These include species such as `Lantana montevidensis, Isostigma
peucedanifolium, Macrosiphonia martii and Stylosanthes capitata. Coutinho et al. (1978)
measured the nutrient content of the first two species before and after fire, and demonstrated that these species use their xylopodia for storing nutrients after fire. Although
they have a poor root system, they absorb nutrients from the ash with great efficiency,
possibly explained by the presence in many roots of vesicular-arbuscular mycorhyzae
(Teixeira, 1986).
Apart from fire adaptation studies, the herbaceous layer of the cerrado has not been as
intensively studied as the woody layer. Moreira (1992) investigated the effects of burning
on grass cover between different cerrado physiognomies. She found a significant difference between protected and burned areas only in the cerradaÄo. Here, protected cerradaÄo
had less grass cover than the burned area, suggesting that fires support and encourage
growth in grassy vegetation, prevalent in the more open forms of cerrado (Moreira, 1992).
Other studies have focused on the effects of fire on individual grass species. Echinolaena
inflexa, a common cerrado C3 grass, has shown to be consistent in frequency between
burned and unburned areas (Klink and Solbrig, 1996), although Parron (1992) found that
seed production is halved in burned areas, and Murakami and Klink (1996) showed that
in burned areas, E. inflexa does not attain the architectural complexity seen in protected
sites. Neto et al. (1995), comparing tiller dynamics and leaf production among native
grasses, found that the tiller survival rate after fire was greatest for E. inflexa and Axonopus barbigerus, with A. barbigerus producing leaves continuously during the subsequent
dry period. It has been suggested that fire affects life strategies of cerrado grass species,
with some such as E. inflexa being an `r' strategist (opportunistic with rapid population
growth, typical of variable environments) in burned areas and `K' strategist (having slow
growth, favouring stable environments) in protected areas (Miranda and Klink, 1996).
Although the herbaceous layer of cerrado vegetation is extremely rich (Heringer, 1971)
and may comprise over 1550 species (Heringer et al., 1977), the effects of fire on this high
floral diversity is poorly understood. Scattered data suggest that contrary to the woody
layer, periodic fires may increase and/or maintain the species diversity of the herbaceous
J. Mistry 437
layer. For example, fewer orchid species were found in unburned areas compared to
burned areas (Oliveira et al., 1996).
c Plant invasions: African grasses are the most common exotics invading cerrado areas
today, and the most studied. Some, such as Hyparrhenia rufa, were accidentally introduced into Brazil, brought in slave ships during colonial times, whereas others, including
Melinis minutiflora, were purposely used in the conversion of natural to artificial pastures
(Alho and Martins, 1995). The implications of these biological invaders are wide,
including reduced productivity in agricultural areas, economic costs related to control
methods, as well as ecological adverse effects such as reduced biodiversity in natural
areas due to interspecies competition (Alho and Martins, 1995). It has also been
hypothesized that exotic plants may alter the fire regime in the cerrado, as they do in
other ecosystems (D'Antonio and Vitousek, 1992; Smith and Tunison, 1992).
Evidence for this was recently found through a study investigating fire behaviour in
cerrado invaded by the African grass Melinis minutiflora (Berardi, 1994). Thermocouples
were used to measure the spatial pattern of fire temperatures, and residence times of
these temperatures in M. minitiflora dominated campo sujo. The results are shown in
Figure 5. Compared to native campo sujo fires, where the highest temperatures are usually
in the first 100 cm above the ground (most of the fuel is situated here), and rarely exceed
800 8C (Cesar, 1980; Miranda et al., 1993), fire temperatures in M. minutiflora were highest
at 60 and 160 cm heights, with peaks of 817 8C and 1006 8C respectively, and flames over
six metres. Also, native fires normally have very short residence times, whereas in the
M. minutiflora fire, average temperatures of 300 8C were found to last over three minutes.
These results have important consequences for native cerrado vegetation. Normally,
cerrado woody species have thick bark insulating inner living tissues from the effects of
fire, and higher parts of the trunk and leaves are rarely affected (Coutinho, 1990). This
could be reversed where fires are sustained by M. minutiflora. For example, Guedes (1993)
calculated that the critical bark thickness below which the cambium will be killed for a
42-second exposure to a temperature of 180 8C would be 3 mm. This suggests that many
woody plants could be killed rather than scorched by fire (Berardi, 1994). Also the higher
peak temperatures at higher heights could contribute to adverse effects in the canopy
layers. All these factors, could, in turn, reduce species diversity. Berardi (1994) also
points out the effects on nutrient cycling. Fires within M. minutiflora would result in a
greater loss of nutrients due to the higher temperatures attained, which would volatize a
greater proportion of nutrients and lead to larger ecosystem losses.
2
Fauna
There is abundant information about the death and survival of particular cerrado animals
from fire, their related adaptive strategies, the dependence of many animals on fire
`products', e.g., deer species licking ash, and the postfire flush of new grass, flowers,
fruits and seeds during a period of food scarcity in the dry season (Sick, 1965; Alho, 1981;
Coutinho, 1990; Rodrigues, 1996). Yet, scientists are still uncertain of the effects of fire on
animal populations, and this has put question marks on the use of fire as a management
tool in natural areas.
Few groups of animals have been studied. For lizard populations, results show that
periodic heterogeneous burns probably help to maintain populations and species
diversity, which will fall with an increase in fire frequency and homogeneity (Araujo
438
Figure 5 Graph of temperature against time during Melinis fire
Source: After Berardi, 1994
et al., 1996). Periodic fires have also shown to be beneficial for insect herbivores, such as
leaf-galling midges, which increase substantially in burned areas (Prado, 1989; Prada
et al., 1995; Vieira et al., 1996). In ant populations, fire reduced the arboreal species by
69%, whereas underground colonies were hardly affected (Naves, 1996).
Among other studies conducted, the effects of fire on guilds of spiders (Dall'Aglio,
1992), on plant±insect interactions (Righetti, 1992; Seyffarth et al., 1996) and on the
construction of ant nests (Dias, 1993) have been investigated. Of bird communities,
most studies have concentrated on the effects of fire in gallery forest, a formation
common along waterways in the cerrado region, rather than actual cerrado vegetation
(e.g., Figueiredo and Cavalcanti, 1992; Marini and Cavalcanti, 1996). These studies,
J. Mistry 439
however, stress the importance of habitat diversity in the cerrado for animals during and
after fires (Mares et al., 1986; Marini and Cavalcanti, 1996). Nevertheless, for managing
fire in natural areas, data on other wildlife, such as the tamandua, guara wolf and deer,
are necessary, and this, as yet, is still lacking.
3
Social studies
There have been no specifically designed studies for investigating the human use of fire
in the cerrado. To date, information has been a byproduct of various anthropological
studies, all of which have concentrated on indigenous people. Anderson and Posey
(1985; 1989) observed KayapoÂ people who burn the cerrado during the dry season. They
found that the timing of the burns was determined by tribal elders, who used both
astrological and ecological indicators. The time of fruiting of Caryocar brasiliense, a key
food source for the Indians, was probably the most important of these indicators. The
burns were effectively controlled: precautionary firebreaks were constructed by removing dry grass and shrubs, and during the fire, branches were employed to avoid fires
penetrating other areas.
Indigenous tribes also use fire for hunting, and as signals to other tribes (NimuendajuÂ,
1983; Maybury-Lewis, 1984). Linguistic studies have indicated a rich vocabulary of `fire'
words in cerrado tribes such as the Xavante (McLeod and Mitchell, 1980; Giaccaria and
Heide, 1984; Hall et al., 1987). This suggests that fire has an important role to play in
indigenous lives. Unfortunately, no other details of fire use, such as frequency and size of
areas burned, are available, which could contribute towards drawing up management
plans for other cerrado areas.
This type of information is particularly necessary of farmers, who today are the
primary cause of fires in the cerrado (Coutinho, 1990). Various fire regimes have been
documented (Table 4) but, as yet, little else is known. Mocelin (1996), interviewing
extension officers in the Distrito Federal about farmer fire activities, confirmed
hypotheses and local assumptions that farmers burn pastures to encourage fresh
regrowth for cattle, to eliminate undesirable species (both plants and animal) from
pastures and fields, and as a means of clearing areas for planting. He also found that
most burning took place in September. However, further detailed data were not obtained
within the scope of his undergraduate project, especially because actual farmers were not
interviewed.
Farmer knowledge is now being investigated by the present author, taking a decisionanalysis approach (Mistry, 1998a). It is hoped this will throw light on the social,
economic, ecological and political factors influencing fire usage, the understanding of
which could aid the development of fire management policies.
4
Management
Burning is the oldest, cheapest and most widely used management tool in the cerrado
(Coutinho, 1990). It is a natural force and has many beneficial effects including the
stimulation of germination, resprouting, flowering and fruiting, as well as accelerated
nutrient recycling (Coutinho, 1990), all of which are important for animal and human
populations. Prescribed burning also prevents fuel buildup and the occurrence of
uncontrollable wildfires, and is an important determinant of the many fire-adapted
species present in the cerrado (Dias, 1997). However, its misuse, both intentionally and
440
Table 4
Fire regimes used by cerrado farmers
Fire regime
Sequence of actions
Cited by
Rotation of native `green' a) ®eld ! early annual burn
pastures
! `green' regrowth for cattle ! ®eld
b) cerrado ! late annual burn
! `green' regrowth for cattle ! cerrado
Saint-Hilaire, 1827
Rotation of clearings
(`coivaras')
forest ! cut ! burn ! clearing
! brushwood for collecting, and
hunting facilitated ! leave fallow
for more than 40 years ! forest
Posey, 1986
Rotation of charcoalmaking
cerrado ! cut ! remove
wood ! burn ! leave fallow for
10±15 years ! cerrado
Silva and Fel®li, 1992
Rotation of charcoalmaking and natural
pasture
wood ! burn ! regrowth for cattle
! leave fallow for 10±15 years ! cerrado
Saturnino et al., 1977
Opening of natural
pasture
cerrado ! cut ! remove wood `clean'
! burn ! ®eld for cattle
Opening for African
planted pasture
wood ! burn ! plant African grass seed
! African planted pasture for
cattle ! burn
Valverde, 1985
Opening for farming
wood ! rake ! burn ! add fertilizer and
other agricultural inputs ! plant land
Moraes, 1993
Opening for planted
pasture
cerrado ! exploration ! correction of soil
! 1±3 years of farming ! harvest
! burn ! plant grass seed
! planted pasture for cattle
Rotation of cultivation/
post-harvest burn
farming ! harvest ! burn to clear
! plant ! farming
Moraes, 1993: applied to
the cultivation of cotton
Rotation of cultivation/
pre-harvest burn
farming ! burn to clear ! harvest
! regrowth ! farming
Moraes, 1993: applied to
the cultivation of
sugarcane
Renovation of ®rebreaks
cerrado ! clear road with tractor/rake
! burn section of cerrado to
increase ®rebreak ! cerrado
IBDF, 1991
Source: Modi®ed after Dias (1997).
unintentionally, has led to a strong feeling against its usage, and has delayed the
development and implementation of practical fire policies and management strategies
(Pivello and Coutinho, 1996; Pivello and Norton, 1996). This has also been reflected
in practical fire management research for the cerrado, which has been scarce to date.
However, prescribed burns are now being recognized as a suitable tool for cerrado
management and, increasingly, cerrado fire research is being undertaken with a
J. Mistry 441
management aim/perspective. Recent studies with regard to fire history and expert
systems are described below.
a Fire history: The history of recent fires is an important aspect of fire ecology to
understand and determine if correct and effective prescribed burning regimes are to be
applied in order to meet particular management objectives (Whelan, 1995). For example,
proving that an area of cerrado has had a history of fire protection would indicate the
presence of species that are not adapted to fire. If the area was then excessively burnt,
many of these species would become locally extinct.
Fire history studies have been virtually nonexistent in savanna areas, including the
cerrado. Recently, however, a pioneering work by the present author showed how
corticolous lichens could potentially be fire history bioindicators in the cerrado (Mistry,
1996; 1998b; 1998c; 1998d). Fires in the cerrado are generally patchy in behaviour at a
small scale, due to variations in fuel and local climate (see `Fire behaviour' section
above). For example, a fire may scorch only one side of a tree trunk, leave another
completely unscathed, whilst burn a third trunk entire up to two metres, all within a
space of three metres. This small-scale heterogeneity of fire allows lichens living on tree
bark to have a differential distribution over space and time, depending on their fire
survival and recolonization strategies, thus allowing us to build up a detailed picture of
the small-scale fire history of an area (Mistry, 1996). Having this information would be
particularly important for managing a specific resource, or rare species living in certain
areas.
The study above culminated in the development of a Lichen Fire History (LFH) key,
which uses lichen community responses on tree bark to detect local fire history, and
comprises a simple, illustrated user guide and checksheets, and can be employed by both
scientific and lay people (Mistry, 1998b; 1998c). The key can estimate the frequency and
behaviour of past fires up to 20 years ago, and would be used in conjunction with other
fire knowledge, such as current fuel loads. Still, the LFH key was developed for a small
area in central Brazil, and further testing and validation need to be undertaken in order
to prescribe correct burning regimes to areas of cerrado.
Other fire history studies in the cerrado have been operating at a higher technological
level than the LFH key, through the use of satellite remote sensing. Various satellite
systems are able to construct recent fire histories by mapping successive active fires, as
well as burn scars, and discerning the fire intensity through interpretation of the spectral
reflectance from the resulting ash layer (Minnich, 1983; Chuvieco and Congalton, 1988;
Justice et al., 1993). For the cerrado, Prins and Menzel (1994), for example, mapped fires
from 1983 to 1991, showing successive expansion of burned areas. Although satellite
imagery can give detailed pictures of fires, as yet, the resolution is not great (up to
10 10 m) (Malingreau, 1990), and cannot, therefore, be effectively applied in terms of
the small-scale heterogeneity seen in cerrado fires or in the management of the relatively
small areas of natural cerrado. Nevertheless, as technology improves, remote sensing is
likely to be one of the most important tools for detecting cerrado fire history.
b Expert systems: Expert systems are decision support tools which handle qualitative
information and heuristic knowledge, enabling specific practical problems to be diagnosed and giving advice on how to solve them (Starfield and Bleloch, 1983; Waterman,
1986). Questions an expert would need to ask in order to make a recommendation
are listed, and depending on the answers the managers give to these questions, a
442
recommendation is produced, according to a series of rules that have also been derived
from experts. For fire management, working expert systems have been successfully
developed for decision-making primarily in Australia (e.g., Davis et al., 1986; Ludwig,
1990) and in the USA (e.g., Rauscher, 1987; Andrews and Chase, 1989; Reinhardt et al.,
1989). For the cerrado, it is only recently that a prototype, FIRETOOL, aimed at assisting
managers of Brazilian savanna national parks and reserves, has been developed (Pivello,
1992).
The expertise necessary for the system was provided by an extensive literature survey
of cerrado fire ecology, and interviews with selected Brazilian scientific researchers and
managers of parks and reserves (Pivello, 1992). This knowledge was then used to
construct FIRETOOL, which comprises four subsystems: STARTING presents the
program and gives instructions on how to design a short-term burning plan; FIRERISK
assesses the risk of a wildfire in a site and its likely intensity; FIREUSE estimates whether
a prescribed burn is necessary and suggests the most appropriate fire regime; and
PROCEDURE gives directions on basic techniques for carrying out controlled fires and
making firebreaks (Pivello and Norton, 1996).
The development of an expert system for fire management in the cerrado is a
breakthrough, as few studies have as yet directly addressed this issue. FIRETOOL was
initially tested against the assessments of one cerrado fire expert (Pivello, 1992), but if it is
to be implemented in everyday fire management, its potential needs to be verified
through further intensive field tests.
IV
Concluding remarks
The study of fire ecology in the cerrado is in a healthy state, having been instigated by
various research groups, namely the Department of Ecology at the University of Brasilia,
and Leopoldo Coutinho and his students at the University of SaÄo Paulo. Present work is
diverse, and yet Pivello and Norton (1996) identify many fire knowledge gaps. Amongst
these, many, such as the effects of fire on soil characteristics and plant dynamics, are
being actively researched. Others, however, such as fire management practised by people
and the effects of fire on faunal populations, are scarce, and yet it is these aspects of fire
ecology which need to be urgently addressed if real management plans and policies are
to be developed, for both natural and semi-natural cerrado areas.
The cerrado is still relatively depopulated compared to other parts of Brazil, although
migration into the area has increased rapidly since the establishment of the new capital
Brasilia in 1960 (6.5 million in 1970 to 12.6 million in 1991) (Alho and Margins, 1995).
Migrators are generally poor subsistence farmers and workers, and for many, fire
constitutes the cheapest and fastest management tool. It is therefore vital to direct future
work with the aim of managing fire in ways that are sustainable for plants, animals and
humans.
Acknowledgements
I would like to thank Andrea Berardi and Professor Philip Stott for their comments and
advice.
J. Mistry 443
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