BIOTROPICA 46(6): 643–646 2014 10.1111/btp.

12153
INSIGHTS

Fire Damage in Seasonally Flooded and Upland Forests of the Central Amazon

lica F. de Resende1,3, Bruce W. Nelson1, Bernardo M. Flores2, and Danilo R. de Almeida1

Ange
1

Araujo 2936, 69067-375, Manaus, AM, Brazil
INPA – National Institute for Amazon Research, Av. Andre
2
Department of Aquatic Ecology and Water Quality Management, Wageningen University, P.O. Box 47, NL-6700 AA, Wageningen,
The Netherlands

ABSTRACT
Neighboring upland and nutrient-poor seasonally flooded Amazon forests were penetrated by a fire in 2009, providing a natural com-
parative experiment of fire damage for these widespread forest types. In upland, only 16
10% (
2 SEM) of stems and 21
8%
of basal area were lost to fire, while seasonally flooded forest lost 59
13% of stems and 57
13% of basal area. Drier understory
contributes to greater flammability. Much of the area occupied by seasonally flooded woody vegetation (>11.5 percent of the Amazon
region) is vulnerable to fire due to high flammability and slow recovery.

Abstract in Portuguese is available in the online version of this article.

Key words: flammability; forest structure; igap
o; litter layer; relative humidity; tropical forest.

OVER 11.5 PERCENT OF THE AMAZON BASIN IS OCCUPIED BY SEASON-
ALLY INUNDATED WOODY VEGETATION (Melack & Hess 2010).
Seasonally flooded Amazon forests are home to a high diversity
of trees (Wittmann et al. 2010), endemic invertebrates (Adis et al.
2010) and frugivorous fish (Goulding et al. 1988, Correa et al.
2007). Although inundated for much of the year, the litter layer
of a low open-canopy floodplain forest of the upper Rio Negro
can ignite when relative humidity of the understory drops below
65 percent (Uhl et al. 1988).
Fire also penetrates closed-canopy floodplain forests of the
middle Rio Negro. Using field inventories and high-resolution
images Flores et al. (2014) found average tree mortality of 90 per-
cent from fires associated with droughts—much higher than the
mortality or percent of biomass loss reported after a first burn of
upland forests (Barbosa & Fearnside 1999, Cochrane et al. 1999,
Barlow & Peres 2004). Amazon forests seasonally flooded by
nutrient-poor waters have an aerated root mat and accumulation
of leaf litter which together comprise a stock of fine, quick-drying
fuel two to three times larger than in nearby upland forest
(Kauffmann et al. 1988, Dos Santos & Nelson 2013).
These prior results suggest that flooded forest is less resistant
to fire damage than upland forest. However, no experiment has yet
been performed comparing fire resistance of these two widespread
forest types. The Rio Negro provides few situations to compare
flammability; river channels act as fire breaks between the two for-
est types and ignition sources differ (e.g., fishermen’s campfires are
more abundant in the floodplain). A report by Nelson (2001) com-
pared fire damage between upland and floodplain forests in close
proximity and penetrated by the same fire, but relied only on visual
inspection of satellite images and air photography.
Received 1 February 2014; revision accepted 16 July 2014.
3
Corresponding author; e-mail: gel.florestal@gmail.com
ª 2014 The Association for Tropical Biology and Conservation
Here, we use a natural experiment to address the hypothesis
that given equal ignition opportunity and identical recent rainfall
history, fire will cause greater damage to undisturbed nutrient-
poor flooded forest than undisturbed upland forest (H1). Our
second hypothesis examines cause: The unburned flooded forest
understory has more prolonged periods of flammability than the
unburned upland forest understory (H2).
South of the central sector of the Amazon River are exten-
sive terraces 15–20 m above the present local high water level.
Irion et al. (2010) interpret these as paleo-floodplains deposited
during Pleistocene interglacial periods, such as 110,000 years ago,
when the Amazon River was slightly above its present relative
level. During the final Pleistocene low sea stand the paleo-flood-
plains became upland and were weakly incised by erosion, form-
ing dense dendritic networks of shallow valleys. Post-Pleistocene
sea level rise back-flooded the Amazon main stem and these
shallow incised valleys. Currently, they are seasonally inundated
by a 10 m amplitude annual flood pulse and are colonized by
trees tolerant to periodic inundation. Between the seasonally
flooded valleys, the paleo-floodplain terrace forms flat narrow
interfluves covered by flood intolerant upland tree species.
For three reasons, fires on this landscape provide the ideal
natural experiment to address H1. First, tree cover is continuous
between the two forest types, with no intervening open water or
erosional scarp to impede passage of fire. Second, the two forest
types are spatially intercalated because of the dense network of
seasonally flooded valleys, allowing placement of all plots in close
proximity. As a result, sites can have identical pre-burn rainfall
and equal ignition opportunity when fires course through the sea-
sonally flooded valley forest (Fig. S1). Finally, because ignition
sources for both forest types (swidden fields and pastures) were
absent over much of the area until recent decades, it is easy to
find forest that has burned only once.
643
644 Resende, Nelson, Flores, and Almeida
METHODS
We used a time series of satellite images to identify a region
100 km south of Manaus, where both seasonally flooded valley
forest and upland interfluve forest were exposed to a first burn
in November of 2009 during an El Ni~ no drought (Fig. S1). The
valley forests here are inundated by back-flooded nutrient-poor
local runoff with minor influence of nutrient-rich water from the
Amazon River.
We set up 40 elongate (250 m 9 20 m) inventory plots in
2012/2013. In each plot, we measured basal area (m2/ha) and den-
sity (stems/ha) for all trees >10 cm diameter at breast height (dbh).
Ten of the forty plots were placed in seasonally flooded forest and
ten in upland forest, both of which burned once. The upland plots
were surrounded by burned flooded forests and were no more than
200 m from them, guaranteeing equal ignition opportunity and
identical pre-burn rainfall history. All forest inventories were con-
ducted 3–4 years after the fire to allow completion of post-burn
mortality (Barlow et al. 2003). Pioneer regrowth from seed did not
reach the 10 cm minimum dbh in this time.
We also inventoried ten plots in unburned seasonally flooded
valley forest and ten plots in unburned upland forest. Serving as
proxies for pre-burn structure of the burned plots, these 20 refer-
ence plots were in a large parcel of preserved forest isolated from
ignition sources but within 5 km of all burned plots (Fig. S1).
For the ten burned plots in each forest type, fire-induced struc-
tural changes were expressed as percent loss of basal area and
percent loss of stems, relative to the average basal area and the
average stem density, respectively, of the ten reference plots for
the respective forest type. This provided ten observations of fire
damage per structure attribute per forest type. To test H1, we
compared the means of percent basal area loss and of percent
stem loss between the upland and seasonally flooded burned
sites. To detect differences in fire vulnerability by tree size, we
compared percent loss of stems across five size classes within
each forest type. The 40 inventory plots were spread across five
km2, centered at 3°430 S and 60°140 W, near the southwest end of
A B
FIGURE 1. (A) Stem loss in neighboring seasonally flooded and upland forests exposed to the same fire; (B) basal area loss after same fire; (C) driest one percentile
of relative humidity records for each sensor in understory of unburned flooded forest and unburned upland forest sites over 17 d in the 2013 dry season.
Lago Mamori, a narrow back-flooded lake. All plots were located
at least 500 m inland from the shore to avoid old secondary
forests. No plot had suffered prior mechanized logging.
To address our second hypothesis, we used relative humidity
(RH) and temperature (T) of the understory near the litter layer as
indicators of litter layer flammability. We distributed sensors of RH
and T (Lascar brand, model EL USB 2+) in five unburned plots in
each forest type. Sensors were secured 50 cm above the litter in
meteorological shelters, each made from three nested inverted Sty-
rofoam bowls, providing free air flow and protection from rain and
direct light of understory sunflecks. Each sensor recorded RH and
T in 5 min intervals from 12 to 28 November of 2013, starting
shortly after the annual minimum of the flood cycle. For each sen-
sor, we determined the cutoff for the driest one percent and the
hottest one percent of all measurements. We compared the means
of these cutoffs grouped by undisturbed forest type (N = 5 for
each treatment, upland and flooded forests).
The Lascar RH sensor is protected by a water resistant
membrane that causes an asymptotic response lag. To measure
this, two groups of sensors that started from an RH difference
of 10 percent were placed in the same stable environment. They
took 2 h to converge to an average difference of 5 percent, and
9 h to narrow the gap to 3 percent. So the true minimum RH
near noon of each day will be slightly lower than the recorded
minimum.
RESULTS
FIRE DAMAGE.—Both forest types had reduced stem density in
the burned plots compared to the reference plots (P = 0.001 for
flooded forest, P = 0.037 for upland, Mann–Whitney U-tests).
Confirming hypothesis 1, percent stem loss was much higher in
the burned floodplain plots (P = 0.001, Fig. 1A). These lost
59
13% of their trees (mean
2 SEM), while burned upland
plots lost only 16
10%.
Compared to the reference plots, basal area losses were also
significant in both forest types after a first burn (P = 0.001 for
C

floodplain, P = 0.007 for upland). Basal area loss was also greater
in the flooded forest (P = 0.001), further supporting hypothesis
1. Upland forests lost only 21
8% of their basal area while
flooded forests lost 57
13% (Fig. 1B).
Higher percent losses from fire in flooded forest are aggra-
vated by starting from pre-burn basal area of 19.4 m2/ha and
tree density of 329 stems/ha, compared to 26.3 m2/ha and 546
stems/ha in upland forest (Table S1; Fig. S2A). Fire-induced
mortality was not selective by tree size within either forest type
for the four dbh classes between 10 and 50 cm (Fig. S2B).
MICROCLIMATE.—The unburned flooded forest understory
reached higher extremes of temperature (P = 0.008) and lower
extremes of relative humidity (P = 0.009) than unburned upland
forest. Among the five sensors in floodplain forest, the lowest
one percentile of humidity was in the range 71–77 percent RH,
compared to 81–87 percent RH for the five sensors in the
upland forest (Fig. 1C). Figure S3 shows a 24 h cycle for all sen-
sors on the driest day of the 17-d period, when one flooded for-
est site dropped briefly below the critical litter flammability level
of 65 percent RH in the understory.
DISCUSSION AND CONCLUSIONS
It is clear from satellite images (Fig. S1) that fire spreads much
more easily through and does greater damage in seasonally flooded
forest than in neighboring upland forest. At least three factors con-
tribute to this higher susceptibility: drier microclimate, higher fine
fuel load and a flammable root mat in the flooded forest.
The lowest one percentile RH cutoff for flooded forest
understory was about 8 percent of RH lower than for upland
forest. Assuming this difference also occurs in years drier than
2013, the flooded forest litter layer will be more flammable and
flammable for longer periods than the upland litter, as postulated
in our second hypothesis. Tree basal area 26 percent lower than
in upland and tree stem density 40 percent lower (Table S1; Fig.
S2A) suggest more canopy gaps and less total leaf area in the
flooded forest. The upland forest has abundant understory leaves,
while flooded forest understory is very open. Forest types with
greater light penetration, lower height and fewer leaves attain the
critical flammability level of 65 percent RH more days of the year
(Uhl et al. 1988, Holdsworth & Uhl 1997).
Regarding comparative fine fuel loads, Dos Santos and Nel-
son (2013) found ~26 Mg/ha of fine fuel as litter and root mat
in nutrient-poor flooded forests of the middle Rio Negro, about
twice the amount in nearby upland forests. More fine fuel
combined with longer periods of flammability will lead to more
frequent and more intense burns during drought years in the
seasonally inundated forest than in the upland. Finally, the flam-
mable root mat makes flooded forest more vulnerable to both
surface fires and ground fires, killing many more trees than in
upland by burning their roots (Flores et al. 2014; Fig. S4).
As on the nutrient-poor Rio Negro (Flores et al. 2014), the
flooded forest at our site showed low post-fire resilience. Woody
regrowth was nearly absent 3–4 yr after a first fire. The burned
Amazon Flooded Forest Fire 645
upland forest had a continuous canopy of surviving pre-burn
trees or of dense regrowth (<10 cm dbh) where the fire had
formed gaps. Near-complete canopy removal combined with slow
post-burn recovery (Flores et al. 2014) leaves the flooded envi-
ronment dominated by low shrubs, forbs and graminoids for
many years, increasing the likelihood of reburning in subsequent
dry seasons (Fig. S4).
Upland forest fires are already widespread during drought
years in the densely occupied portions of Amaz^ onia (Brando et al.
2014). As ignition sources spread with advancing human occupa-
tion, fire will also become frequent and widespread in flooded
forests. Future directions for study include understanding the
underlying causes and consequences of low resistance to fire
damage and of low post-fire resilience in nutrient-poor flood-
plains; the role of fire in forest degradation of nutrient-rich flood-
plains; and predicting and managing fire risk.
ACKNOWLEDGMENTS
We thank Francisco Cavalcante (in memoriam), Pedro Gama,
Livia Resende, Jo~ao Rocha, Vilma Soares, Maria Fernanda da
Silva, Igor do Vale, Aline Lopes and Sebasti~ao Salvino for field
assistance; the Correa family and Tiango Ribeiro for permission
to work on their properties; Milena Holmgren, Adrian Barnett,
Maria Teresa Piedade, Jochen Sch€ ongart and three anonymous
reviewers for revisions; FAPEAM (Amazonas Research Support
Foundation) and CNPq (Brazilian National Research Council) for
financial support.
SUPPORTING INFORMATION
Additional Supporting Information may be found in the online
version of this article:
FIGURE S1. Landsat Thematic Mapper image, 1 yr after fire
showing high damage in dendritic flooded forests and low dam-
age in intercalated upland interfluves.
FIGURE S2. Comparison of stem density by size class
between unburned upland and floodplain, and comparison of
post-fire percent stem loss by size class.
FIGURE S3. Diurnal cycle of relative humidity 50 cm above
the litter layer of unburned floodplain and upland forests on the
driest day recorded.
FIGURE S4. Twelve years after penetration by a fire during
the 1997 drought, floodplain forest on the lower Cuini River, a
tributary of the middle Rio Negro, shows 100 percent tree mor-
tality and almost no woody regrowth.
TABLE S1. Tree basal area and density for 40 inventory plots.
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SUPPORTING INFORMATION

Fire Damage in Seasonally Flooded and Upland Forests of the

Central Amazon

Angélica Faria de Resende1,3, Bruce W. Nelson1, Bernardo M. Flores2, and Danilo R. Alves

de Almeida1

1
INPA – National Institute for Amazon Research, Av. André Araujo 2936, 69067-375, Manaus,

AM, Brazil
2
Department of Aquatic Ecology and Water Quality Management, Wageningen University, P.O.

Box 47, NL-6700 AA, Wageningen, The Netherlands

Received 1 February 2014; revision accepted 16 July 2014.

3
Corresponding author; e-mail: gel.florestal@gmail.com

1
FIGURE S1. Landsat Thematic Mapper image acquired in November of 2010, one year after
fire, showing high damage in dendritic flooded forests, low damage in intercalated upland
interfluves. Dashed line indicates approximate limit of fire front separating burned plots and
reference plots.

1
 FIGURE S2. (A) Comparison of stem density by size class between unburned upland and
floodplain; (B) Comparison of post-fire percent stem loss by size class. Error bars are ± two
standard errors of the mean.

2
 Relative Humidity (percent) 100
95
90
85
80
75
70
• upland forest
65 + floodplain forest
60
23:00 3:00 7:00 11:00 15:00 19:00 23:00
Time of Day

FIGURE S3. Diurnal cycle of relative humidity 50 cm above the litter layer of unburned
floodplain and upland forests on the driest day recorded, 24 November 2013; data from ten sites.

3
Figure S4. Twelve years after penetration by a fire during the 1997 drought, floodplain forest on
the lower Cuini River, a tributary of the middle Rio Negro, shows 100% tree mortality and
almost no woody regrowth.

4
TABLE S1. Tree basal area and density for 40 inventory plots.

Unburned
Burned Flooded Forest Burned Upland Flooded Forest Unburned Upland

% loss % loss % loss % loss

Plot BA/ha BA Ind./ha Ind. BA/ha BA Ind./ha Ind. Ba/ha Ind./ha Ba/ha Ind./ha
1 8.7 55 78 76 25.1 5 574 -5 18.7 252 27.3 620
2 13.1 32 206 37 25.1 5 332 39 24.4 396 26.6 520
3 10.2 48 154 53 21.0 20 528 3 15.7 318 28.9 502
4 3.1 84 54 84 23.5 11 354 35 15.0 236 28.9 564
5 6.3 68 110 67 22.7 14 518 5 26.6 350 22.6 504
6 12.3 37 196 40 21.4 19 524 4 16.9 292 24.6 556
7 11.6 40 234 29 16.2 39 406 26 19.9 310 25.0 556
8 3.9 80 78 76 15.8 40 372 32 15.8 270 27.2 508
9 11.8 39 190 42 18.2 31 518 5 21.8 480 31.5 528
10 2.8 86 48 85 18.4 30 442 19 19.4 384 21.0 602
Mean 8.4 57 135 59 20.7 21 457 16 19.4 329 26.3 546
SD 4.0 21 69 21 3.5 13 86 16 3.9 75 3.2 41
CV 0.5 0.4 0.5 0.4 0.2 0.6 0.2 1.0 0.2 0.2 0.1 0.1

Abbreviations: BA – Basal Area, Ind. – Number of Individuals, SD – Standard Deviation, CV – Coefficient of Variation.
Percent loss of basal area and of density (Ind./ha) calculated using average of unburned reference plots as proxy for pre-burn values.