Forest Ecology and Management 347 (2015) 170179

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Forest Ecology and Management

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Post-re live residuals of maritime pine plantations in Portugal:

Structure, burn severity, and re recurrence
Paulo M. Fernandes a,b,, Manuel M. Fernandes b,1, Carlos Loureiro a
a
b

Centro de Investigao e de Tecnologias Agro-Ambientais e Tecnolgicas (CITAB), Universidade de Trs-os-Montes e Alto Douro, Quinta de Prados, 5000-801, Portugal
Departamento de Cincias Florestais e Arquitetura Paisagista, Universidade de Trs-os-Montes e Alto Douro, Quinta de Prados, 5000-801, Portugal

a r t i c l e

i n f o

Article history:
Received 4 January 2015
Received in revised form 13 March 2015
Accepted 15 March 2015
Available online 30 March 2015
Keywords:
Tree mortality
Fire frequency
Fire severity
Surface re regime
Fuel treatments

a b s t r a c t
The drivers, characteristics and impacts of surface re regimes in the Mediterranean Basin are poorly
understood. We describe the post-re structure of residual maritime pine (Pinus pinaster Ait.) patches
in frequent-re mountain landscapes of northern Portugal and relate it with burn severity and re
history. Live trees within each plot were measured and cumulative burn severity metrics were assessed
at the tree and plot levels. Plot re recurrence and re intervals (mean, MFI; minimum, MinFI; and
maximum, MaxFI) were calculated from a digitized re atlas. Fire thinned from below, with a 15.5-cm
diameter at breast height corresponding to 50% survival probability. Bole char and crown kill (or live
crown base) heights (CKH) were correlated with live tree height; crown ratio averaged 0.38. Each patch
burned 19 times since 1975, with a MFI range of 1.716 years in multiple-re plots. We found wide
variation in stand height (7.822.5 m), basal area (1.447.9 m2 ha1) and tree density (142199 ha1)
but single-storied structures prevailed. Higher re recurrence was associated with lower stand density
and higher percentage of re-scarred trees, indicating cumulative thinning and cambium damage effects.
Bole char height and CKH increased with longer re intervals, i.e. with fuel accumulation and potentially
higher re intensity. MFI (accounting for 70% of the explanation), terrain aspect, and MaxFI explained 55%
of plot-level variation in CKH using regression tree analysis. MFI <4.2 years generated the lowest CKH. For
stands with MFI P4.2 years, CKH increased on slopes facing east or south (drier and warmer) and was
highest when MaxFI P4.8 years. Active crown re was less likely in forest patches stocked at
<20 m2 ha1 and <200 ha1, which should decrease crown re hazard and limit high-severity re if
combined with adequate pruning and surface fuel treatments every 35 years. The study indicates that
patches surviving a rst-entry wildre are likely to persist under a frequent re regime, adding to the
understanding of Mediterranean pines resilience to re and providing quantitative empirical evidence
useful to guide the management of re-prone forests.
2015 Elsevier B.V. All rights reserved.

1. Introduction
Residual live trees are an important biological legacy of disturbance regimes that create heterogeneity in structure, composition,
and function (Franklin et al., 2007), with benets to carbon recovery and biodiversity (Seidl et al., 2014). Differential tree survival to
wildre is dependent on re behaviour variability and the
Corresponding author at: Centro de Investigao e de Tecnologias AgroAmbientais e Tecnolgicas (CITAB), Universidade de Trs-os-Montes e Alto Douro,
Quinta de Prados, 5000-801, Portugal. Tel.: +351 259 350861; fax: +351 259
350480.
E-mail addresses: pfern@utad.pt (P.M. Fernandes), mmfernand@gmail.com (M.
M. Fernandes), carlos.a.r.loureiro@gmail.com (C. Loureiro).
1
Present address: CEGOT Centro de Estudos de Geograa e Ordenamento do
Territrio, Faculdade de Letras da Universidade do Porto, Via Panormica, s/n, 4150564 Porto, Portugal.
http://dx.doi.org/10.1016/j.foreco.2015.03.023
0378-1127/ 2015 Elsevier B.V. All rights reserved.

associated variation in the severity of re effects. Even large and

severe res are characterized by variable re behaviour (Oliveras
et al., 2009; Cruz et al., 2012), resulting in a gradient of re effects
across the landscape, including unburned islands (Boer et al., 2008;
Lee et al., 2009; Romn-Cuesta et al., 2009; Fernandes et al., 2010;
Madoui et al., 2010; Kolden et al., 2012).
The environmental factors that drive re spread and fuel consumption wind, fuel moisture, vegetation type, fuel accumulation
and structure, and topography interact to determine re severity
(Broncano and Retana, 2004; Oliveras et al., 2009; Fernandes et al.,
2010; Clarke et al., 2014; Lecina-Diaz et al., 2014; Viedma et al.,
2014). All these factors are spatially and temporally heterogeneous, so there is ample room for burn severity variation within
any given re. As an example, low-, moderate-, and high-severity
re were attributed to 6.8%, 33% and 60.2% of the 12,697 ha burned
in a maritime pine (Pinus pinaster Ait.) forest in central Spain

P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

(Viedma et al., 2014). On-the-ground assessment and classication

of re severity typically individualizes the existing strata (soil,
understorey, overstorey) through specic indicators (Morgan
et al., 2014). Although correlation is often assumed, substantial
variation in re severity can occur among fuel strata (Jain and
Graham, 2007; Halofsky and Hibbs, 2009; Fernandes et al., 2010),
depending on the interaction between re, fuels, biota, and the
biophysical setting (Sikkink and Keane, 2012).
Disturbance by re in conifer forests ranges from non-lethal,
non-scorching surface res to high-severity crown res,
corresponding to a gradient in re effects and tree mortality
(Franklin et al., 2007) where increasingly larger trees are killed
as burn severity rises (Peterson and Ryan, 1986; Fernandes et al.,
2008). The contribution of crown scorch to tree death depends
essentially on surface re intensity, tree height, and forest type
(Van Wagner, 1973; Peterson and Ryan, 1986). Fire-thinned forests
are typically open and heterogeneous at small spatial scales, forming mosaics of isolated trees, canopy gaps and tree clusters variable
in size and shape (Larson and Churchill, 2012; Kane et al., 2013a;
Lydersen et al., 2013; Fry et al., 2014). Frequent surface re in
USA conifer forests is associated to re resiliency, even when re
occurs under severe weather and drought conditions (e.g.
Stephens et al., 2008), in part because fuel loadings are kept low
and constrain re size and severity (Collins et al., 2009; Malone
et al., 2011; van Wagtendonk et al., 2012).
Research on the re ecology of Mediterranean Basin pines has
been focused on the dynamics of post-re regeneration, probably
because of the contemporary prevalence of high-intensity re,
and much less is known about forest persistence under surface re
regimes (Fernandes et al., 2008). Fire history reconstructions at
centennial or multi-centennial time scales from the analysis of
re-scarred trees in Mediterranean conifer forests support lowor mixed-severity re regimes in maritime pine (Vega, 2000),
Pinus nigra (Ful et al., 2008; Touchan et al., 2012; Christopoulou
et al., 2013), Pinus halepensis (Fournier et al., 2013) and Cedrus
atlantica (Slimani et al., 2014). However, no study has explicitly
examined the linkages between re history, burn severity, and
stand structure in frequently-burned Mediterranean forests.
Crown re development and behaviour is dependent on surface
re behaviour, the existence of vertical fuel continuity, and canopy
bulk density (Van Wagner, 1977; Cruz et al., 2004, 2005, 2006), and
this supports the basic principles for creating re-resistant conifer
forests through fuel treatments such as thinning and prescribed
burning (Agee and Skinner, 2005). Simulation modelling suggests
that stand density management is crucial to decrease the likelihood of severe re (e.g., Fernndez-Alonso et al., 2013; GmezVzquez et al., 2014), but empirical evidence of re-resistant stand
structures in Europe is still scarce and in debate (Oliveras et al.,
2009; Romn-Cuesta et al., 2009; Fernandes et al., 2010; Alvarez
et al., 2012, 2013; Viedma et al., 2014). Ongoing efforts to decrease
burned area and protect forest resources through local and
landscape-scale fuel treatments (e.g., Loureiro et al., 2006) would
benet from a sounder evidence-based approach.
Large-scale afforestation with maritime pine in the western
Mediterranean Basin proceeded throughout the 20th century,
especially in the mountains of Portugal, where most forest cover
disappeared during the mid-late Holocene (Connor et al., 2012).
The resulting dense, even-aged, and ammable plantations have
been implicated in the 1970s re regime shift towards substantially greater re extent (Fernandes et al., 2014). Wildre is the
major factor in the decline of maritime pine occupation in
Portugal, which has diminished 27% between the forest inventories
of 1995 and 2010 (ICNF, 2013). Extensive tracts of the Portuguese
mountain landscapes planted with maritime pine are now shrubdominated, but res commonly leave behind fragments of the former afforestation. While the ability of maritime pine to survive

171

surface re is well documented (Fernandes and Rigolot, 2007;

Fernandes et al., 2008), scarce attention has been given to tree survival after wildre in relation to re severity (Pimont et al., 2011;
Vega et al., 2011; Catry et al., 2013).
This study addressed maritime pine patches that had survived
one or more wildres in northern Portugal. The general goal was
to examine the links between re history, cumulative burn severity,
and stand structure. We identify size thresholds for tree survival
and describe burn severity at the tree- and plot-level, using eld
data from 50 plots. Correlations between burn severity metrics
are examined. Stand structure and burn severity variables are
related with re history variables, and the determinants of crown
re hazard are inspected as a function of stand density and re history. Our hypothesis is that the structure of post-re pine remnants
should reect some degree of re-induced thinning and pruning,
increasing the likelihood of tree survival to subsequent reburning.
2. Methods
2.1. Study location and design
The study area (41030 41270 N, 7260 7500 W) encompasses
the mountain ranges of Alvo, Maro, and Montemuro in the
northeast of Portugal. Soils are shallow and derived from granite
or schist. Climate is Mediterranean with an oceanic inuence
(Csb sub-type). Mean temperatures in January and July are 79
and 2426 C, respectively, and precipitation occurs mostly from
October to April and varies between 800 and 1400 mm year1
(Ribeiro et al., 1988).
On average, 2.5% of the region burns annually and the median
re return interval is 24 years (Oliveira et al., 2012), but in the
southern part of the study area decreases to 13 years where re
has occurred twice or more since 1975 (Fernandes et al., 2012).
We identied potential sampling areas by overlaying the
Landsat-based digitized re atlas of Portugal with the location of
maritime pine remnants identied in 2005 aerial photography.
The atlas spatial resolution and minimum mapping units are
80 m and 35 ha, respectively (19751983), and 30 m and 5 ha
(1984-present) (Oliveira et al., 2012). In areas where forest remnants were conspicuous we selected nine 25-km2 squares for eld
sampling, each including a range in re recurrences.
We carried out the eld work in 2007 and 2008, excluding
recently (<3 years) burned patches where pine mortality caused
by Scolytidae insects could occur in the future (e.g., Vega et al.,
2011). Upon eld inspection, conditional on accessibility, we chose
a total of 50 sampling plots representative of variation in stand
structure within the selected 25-km2 areas. Distance between plots
was variable but always >50 m. Variability in the size and structure
of forest patches guided the selection of plot size from three
options: (1) plot area equal to patch area in isolated clumps
(<0.05 ha) of trees; (2) 0.05-ha circular plots within relatively
dense stands; and (3) 0.10.5 ha circular plots within relatively
open stands.
2.2. Data collection and processing
2.2.1. Site characteristics and re history
Understory on each plot was qualied as shrub- or grass-dominated and the dominant species were registered. We assessed plot
elevation (m), aspect (), slope (%), and slope position, categorized
as valley or lower slope, upper slope, ridge, or plateau. Plot aspect
(h, 0360) was transformed as a continuous index (Wu et al., 2014):

Aspect index  cosh  2  p=360

which ranges from 1 to 1, with higher values implying higher

exposure to solar radiation and hence warmer and drier conditions;
level terrain was equalled to south aspect.

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P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

We determined re recurrence (the number of times each plot

burned) from the overlapped 19752007 re perimeters. Burn
severity can vary with location within the burned area (e.g.,
Perera et al., 2009), thus for each plot we measured the shortest
linear distances to past res boundaries in a GIS and averaged
the distances. For multiple-re plots we calculated the minimum,
mean, and maximum number of years between consecutive res:
minimum re interval (MinFI), mean re interval (MFI), and
maximum re interval (MaxFI), respectively. At three multiple-re
locations (Pegarinhos, Pinho-Cel, Melces) we acquired supplementary data on re history. Basal cross sections (at 1520 cm
from ground surface) were cut with a chainsaw from four dominant re-scarred live trees. The resulting 12 samples were sanded
and re scars identied and counted. Stand age was estimated
from dominant height (Santos and Almeida, 2003), time since re
for stands resulting from post-re regeneration, or the number of
tree rings counted on the basal cross-sections.
2.2.2. Tree measurements and burn severity
We measured the height and diameter at breast (137 cm)
height (dbh) of all live trees within each plot; we measured dbh
only for standing or downed dead trees. We assessed the following
burn severity indicators for each live tree: bark char depth at 0.5-m
from the ground (Ryan, 1982), presence of re scars, maximum
height of bole char (BCH), evidence of crown combustion, and
height of crown kill (CKH), equivalent to height to live crown base
(Peterson and Ryan, 1986). CKH is the combined outcome of crown
scorch height and bud kill height; the latter is a more relevant
cause of tree mortality than foliage death (Peterson and Ryan,
1986). Heights were measured with a Vertex hypsometer to the
nearest 0.1-m. We use burn severity instead of re severity
because our eld assessments integrated the cumulative effects
of previous res and some degree of ecological response rather
than immediate re effects (Morgan et al., 2014).
We described stand (or patch) structure by calculating basal
area (G, m2 ha1), tree density (no. ha1), the percentage of dominant and co-dominant trees, and stand height (the mean height of
the dominant and co-dominant trees). We expressed BCH and live
crown length as tree height fractions, relative bole char height
(RBCH) and crown ratio (CR) respectively. Burn severity metrics
were averaged for the plot, except re scarring and crown combustion, expressed as plot-level percentages.
BCH is an acceptable proxy for ame size and Byrams reline
intensity IB (McNab, 1977; Windisch and Good, 1991; Finney and
Martin, 1993). We used data from a set of experimental res in
maritime pine stands (Fernandes et al., 2009; n = 74) to establish
a IB-BCH relationship for the head of the re through non-linear
least squares regression:

IB 768:1 BCH0:7723

with an R2 of 0.26 and standard errors of 118.1 and 0.1829, respectively. IB (kW m1) was calculated with total ne fuel consumption
and assuming a heat of combustion of 18,000 kJ kg1. The amount
of variability explained by Eq. (2) is modest, at least in part because
of the effects of varying tree diameter and wind speed (Gutsell and
Johnson, 1996).
Plot-level CKH will be referred to as canopy base height (CBH)
from the point of view of crown re potential. The likelihood of
crowning is determined by CBH, and the passive or active nature
of crowning is determined by canopy bulk density (CBD) (Van
Wagner, 1977; Cruz et al., 2005). Plot-level CBD (in kg m3) was
estimated with an equation developed for the study region
(lvarez, 2013):

CBD 0:0048 G0:8075 H0:1875

N0:1521
0

where H0 is the dominant height (m) and N is tree density (no.

ha1).

2.3. Statistical analysis

Data analysis used as experimental units individual trees, to
examine post re survival in relation to tree size and burn severity,
and plots, to identify patterns in the relationships between re
recurrence, burn severity and stand structure. We tested data
variables for normality with the ShapiroWilk test and normal
quantile plots. Non-normal variables were transformed to produce
normal distributions and allow parametric statistical tests, or were
subjected to equivalent nonparametric tests.
We summarized the distributions of live and dead trees by
dbh class, assessed differences in the distributions with a
KolmogorovSmirnov test, and compared the dbh of live and dead
trees with the Wilcoxon rank-sum test. For the sub-set of plots
with live and dead trees, a mixed logistic model was tted with
the xtmelogit command of Stata (StataCorp., 2007) to estimate
the probability of individual tree survival from dbh (the xed
effect), with site and plot as random levels. We summarized live
trees data for each burn severity metric and correlated BCH and
CKH with tree height.
Mean, median and range were reported for the plot-level
descriptors of re recurrence, burn severity, and stand structure.
We assessed whether re history differed by sampling plot size
using Wilcoxon multiple comparison tests. To measure the
strength of association between plot-level burn severity metrics
we used Spearman rank correlation analysis. Linear mixed models
with site as the random effects variable were tted to test whether
forest structure and cumulative burn severity metrics differed
with re recurrence and re intervals; the dependent variables
were log or arcsine square-root transformed.
CKH was selected as the preferred metric of overstorey burn
severity because of its signicance to post-re tree survival and
vigour (Peterson and Ryan, 1986). To understand whether cumulative burn severity was related to re history we modelled CKH for
the multiple-re data subset. We used regression tree analysis
(CART) and included the topographic descriptors and distance to
the re edge along with re return intervals as potential explanatory variables of CKH variation. Regression tree analysis is a recursive partitioning method that repeatedly splits the data and
through sum-of-squares minimization gradually generates more
uniform sub-groups (Death and Fabricius, 2000). Regression tree
analysis overcomes a number of difculties (non-parametric or
unbalanced data; discontinuous or non-linear relationships) and
its results are presented as easily interpreted rules. The minimum
AIC dened the number of splits.
Guidelines useful to prescribe fuel treatments to decrease
crown re hazard were inferred by plotting CBH versus CBD in
relation to stand density and re history variables.
JMP (SAS Institute Inc., 2010) was used for the statistical
analyses, with the above-mentioned exception. Statistical signicance was assumed at the 95% condence level (a = 0.05).
3. Results
Elevation, topographic position, and aspect varied substantially
across the study sites and plots, but moderately steep terrain
prevailed in general (Table 1). Maritime pine comprised the
overstorey in all study plots, with a scattered midstory of
Quercus pyrenaica or Quercus suber at two locations. Shrubs
dominated the understorey in 72% of the plots, mainly Cytisus
multiorus, Cytisus striatus, Ulex europaeus, Ulex minor, Pterospartium
tridentatum, Erica umbellata, Calluna vulgaris and Cistus psilosepalus;
non-woody species (Pteridium aquilinum, Holcus spp., Agrostis spp.,
Rubus spp.) dominated the understorey in the remaining plots.

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P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179
Table 1
Fire history and physiographic information for study sites.

Site (no. plots)

Fire recurrence (Min. Max.)

Elevation (m)

Slope positiona

Aspect

Slope (%)

Teles (5)
S. Tom do Castelo (6)
Alto do Ppulo (6)
Cabea Grande (3)
Pegarinhos (7)
Ribalonga (6)
Pinho-Cel (6)
Fontes (2)
Melces (9)

1
19
13
23
15
12
47
34
35

710740
8201050
805925
500520
450530
650730
700750
690710
850880

2,
2,
2,
2
1,
2,
1,
2
2,

N, S
S, W
N, E, S
N
N, S, W
N, E, S
N, E, S, W
S
N, S, W

1754
016
59
1025
530
015
025
514
015

4
3, 4
3, 4
2
4
2, 3
4

1 Valley or lower slope; 2 Upper slope; 3 Ridge; 4 Plateau.

over 38% of the length of the tree is typically required for the tree
to survive.

0.25
0.20
Dead

0.15

Fraction

0.10

3.2. Stand structure, re history, and burn severity

0.05
0.00
0.25
0.20
Live

0.15
0.10
0.05
0.00
5

10 15 20 25 30 35 40 45 50 55 60

Dbh (cm)
Fig. 1. Dead and live tree diameter distribution by size class in the study plots.

3.1. Surviving trees characteristics

The overall distribution of live trees by size class approached
normality, with most trees measuring 2035 cm in dbh, while
most dead trees had dbh <20 cm (Fig. 1). Dead and live trees differed (p < 0.0001) in their dbh distribution and mean (std. error),
13.6 0.9 cm and 28.0 0.5 cm respectively. The mixed-effects
logistic model with dbh as the independent variable (p < 0.0001)
correctly classied the status (live or dead) of 77.1% of the individual trees, with the 0.50 probability of survival corresponding to
dbh = 15.5 cm.
Burn severity metrics varied widely among trees (Fig. 2),
except bark char depth (not shown) that rated moderate for
84.2% of the trees. Interquartile ranges for CKH and CR were
5.610.6 m and 0.300.48, respectively. Crown combustion and
re scars in the trunk were visible in 39% and 16% of the trees,
respectively. The association of BCH and especially of CKH with
tree height was strong (Fig. 3). If the regression line in Fig. 3 (right
panel) is forced through the origin then its slope changes from
0.72 0.04 to 0.62 0.01. This indicates that live crown retention

BCH (m)

12 0.0

0.2

0.4

RBCH

0.6

Table 2 contains the re history, stand structure and burn severity statistics for the sampled plots. Stand age from tree ring counts
varied between 54 and 67 years and all re scars at those locations
post-dated 1975, with the rst scar formed when trees were 13- to
47-years old (mean = 31 years). Three stands resulted from postre recruitment and were aged between 16 and 28 years. Age estimates for the remaining plots varied from 30 to 60 years. Number
of res and re intervals did not differ between the three plot sizes
(p > 0.05). Sites that burned more than once (72% of the total)
experienced one re every 5 years on average, with 3 and 10 years
as the average MinFI and MaxFI, respectively. Cumulative burn
severity metrics are mostly indicative of surface res of moderately
high intensity (2000 kW m1) with signicant crown combustion. The resulting stand was extremely variable in tree density
and G but was structurally simple as it was essentially composed
of dominant and co-dominant individuals. This was not the general
rule, as shown for the Pegarinhos site where variable re recurrence induced structural variation and created an open stand with
different age classes (Fig. 4).
Plot-level burn severity descriptors were generally correlated,
especially those describing bole char heights and canopy loss
(Table 3). The percentage of re-scarred pines was however
independent of other severity metrics, and crown combustion
was associated with bole char but not with the extent of
re-removed foliage. Tree density, G and CBD decreased with the
number of times burned, and both MFI and MaxFI were positively
associated with stand height and burn severity (Table 4).
Additionally, frequent re increased the proportion of re-scarred
trees, and tree density decreased with higher MaxFI. Bole char
depth and crown combustion were unrelated with re recurrence
or re intervals.
Using the regression tree analysis, three predictors (MFI, MaxFI,
Aspect index) explained 55% of the observed CKH variability in the
plots burned by multiple res (Fig. 5). The rst split was dened by
MFI: re cycles below 4.2 years corresponded to the lowest canopy

12 16

CKH (m)

0.2 0.4 0.6 0.8

CR

Fig. 2. Box-plots with outliers for tree-level burn severity metrics. Whiskers are drawn from the box to the furthest point within 1.5 times the interquartile range (the 3rd
quartile minus the 1st quartile). BCH = average bole char height; RBCH = relative bole char height; CKH = canopy kill height; CR = crown ratio.

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P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

Fig. 3. Relationships of BCH and CKH with tree height, with linear regressions tted (n = 454).

Table 2
Plot-level re history, forest structure and cumulative burn severity metrics for all
plots combined (n = 50, except re intervals, n = 36).
Variable

Range

Mean

Median

Fire recurrence
MFI (yrs.)
MinFI (yrs.)
MaxFI (yrs.)
IB (kW m1)
G (m2 ha1)
No. trees ha1
Dominant and co-dominant trees (%)
Stand height (m)
CBD (kg m3)
Bark char depth
Fire-scarred trees (%)
BCH (m)
RBCH
CKH (m)
CR
Crown combustion (% of trees)

19
1.716
116
322
7994925
1.447.9
142199
14100
7.822.5
0.0050.181
2.43.2
0100
1.211.7
0.070.62
3.614.4
0.230.64
0100

3.1
4.9
3.3
9.9
2126
20.2
368
84
15.5
0.077
2.9
15
4.3
0.29
8.4
0.40
40

3
4.2
2
10
1988
15.5
286
88
16.3
0.071
3.0
12
3.9
0.28
8.5
0.39
32

damage. Shorter MFI was further split by MaxFI, showing higher

CKH when MaxFI P6 years. For plots with MFI P4.2, moister
terrain (Aspect index < 0.71) generated the lowest CKH; in level
terrain and east- and south-facing slopes (Aspect index P 0.71),
plots where MFI P4.8 years experienced higher CKH than those
where MFI <4.8 years. MFI, Aspect index and MaxFI accounted for
78.6%, 18.7%, and 2.7% of the CART explanation, respectively.

where tree death was quite dependent on tree size, as indicated

by the size distribution of dead and live pines (Fig. 1), the 15.5cm dbh threshold for tree death, and high correlation between
BCH and CKH and live trees height (Fig. 3) suggesting that variation
in tree size was more relevant to tree survival than variation in
surface re behaviour. In two of the four maritime pine Spanish
ecotypes studied by Vega et al. (2011), dbh also differed between
dead and live trees. Selective logging of larger dead trees may have
occurred at some sites, in which case the logistic regression
overestimated the likelihood of survival as a function of tree size.
The dbh threshold for tree survival exceeded the 510 cm range
reported for low-intensity burning (Fernandes et al., 2008) and
nearly equalled the ndings of Pimont et al. (2011) for
RBCH = 0.40 and moderate bark char depth.
Previous studies of maritime pine mortality after wildre
indicate virtually certain tree death once crown volume damage
exceeds 68% (Vega et al., 2011), or a 50% tree-death probability
for crown volume damage between 55% and 75%, depending on
bark thickness (Catry et al., 2013). These results reect crown
scorch rather than crown kill, thus are not comparable with ours.
Crown kill volume is lower than crown volume damage when
foliage buds survive below the crown scorch line (Peterson and
Ryan, 1986). The median CR of 0.4 in this study suggests a substantial difference between crown scorch height and CKH in maritime
pine, owing to its large terminal buds protected by foliage
(Fernandes et al., 2008).
4.2. Fire history and burn severity

3.3. Crown re hazard in relation to stand structure and re history

metrics
The combinations of canopy fuel descriptors less likely to support crown re (Fig. 6), i.e. higher CBH and lower CBD, coincide
with G < 20 m2 ha1 and tree densities <200 ha1. The opposite
situation is less well represented and matches G P 20 m2 ha1
and density P400 trees ha1. Active crowning is possible in just
38% of the plots, i.e., when CBD P 0.10 kg m3 ( Agee, 1996; Cruz
et al., 2005). CBD correlates well with tree density (r2 = 0.50) and
especially with G (r2 = 0.94), but CBH is independent (p > 0.05)
from the two stand structure metrics. In relation to re history,
the high CBH low CBD combination requires MFI P4 years and
most plots in that situation burned at least three times.
4. Discussion
4.1. Tree survival
As in other wildre studies (e.g., Ful and Laughlin, 2007), survival of individual pines clearly resulted from a thinning process

The average MFI in this study was 5 years, coinciding with the
lower limit of the MFI range (527 years) reported for
Mediterranean Basin conifer forests (Vega, 2000; Ful et al.,
2008; Touchan et al., 2012; Christopoulou et al., 2013; Fournier
et al., 2013; Slimani et al., 2014). Burned area in northern
Portugal mountains increased by a factor of six from 19531973
to 19742011 (Fernandes et al., 2014). Most pine stands in the
study region were established after the 1940s. Hence, post-1974
re mapping should be representative of the plantations re history, which was supported by the results of our limited re-scar
sampling. A more relevant source of re history error is the occurrence of res smaller than the minimum size detectable from
Landsat imagery. But the possibility of re recurrence overestimation, rather than underestimation, cannot be ruled out. Undetected
unburned islands can occur within a mapped re perimeter, especially when the preceding re is recent and in landscapes
characterized by frequent low-intensity re (Morgan et al., 2001;
Kolden et al., 2012).
Similarly to Lydersen and North (2012), and except for single
re events, the estimated IB express the maximum IB in the plot

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P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

0.2

Fraction

0.15

0.1

0.05

0
5

10

15

20

25

30

35

40

45

50

Dbh (cm)
Fig. 4. Live tree diameter distribution at the Pegarinhos study site. Tree density, G and MFI for the aggregate of sampled plots are 107 ha1 (of which 69% are dominant or
codominant), 5.4 m2 ha1, and 8.1 years, respectively. Pine regeneration between res is evident in the photo.

Table 3
Signicance of Spearman correlation coefcient between plot-level burn severity
metrics (n = 50). (+) and () indicate positive and negative signicant association
( p < 0.05; p < 0.01; p < 0.001), respectively.

Bark char
depth
Fire-scarred
trees
BCH
RBCH
CKH
CR

Fire-scarred
trees

BCH

RBCH

CKH

CR

Crown
combustion

n.s.

(+)

(+)

n.s.

n.s.

(+)

n.s.

n.s.

n.s.

n.s.

n.s.

(+)

(+)
(+)

()
()
()

(+)
(+)
n.s.
n.s.

as recorded by trees, and cannot be ascribed to specic res or

assumed to represent the typical IB to which the trees have been
exposed. Considering the propensity of maritime pine to highintensity re and crowning (Fernandes and Rigolot, 2007), the IB
estimates (7994925 kW m1) are relatively low for summer res
and consistent with partial tree survival. Comparable IB (with
15 m ame lengths) were observed in 2-, 3-, and 13-year old fuels
in the study region during the surface re phases of one experimental re in a maritime pine stand under Very High re danger
rating (Fernandes et al., 2004). This suggests that tree survival
resulted from re-behaviour moderation due to the effects of

Table 4
Effects of re history variables on plot-level forest structure and cumulative burn
severity, with site as a random effect. Dependent variables are log- or arcsinetransformed. (+) and () indicate positive and negative signicant effects ( p < 0.05;

p < 0.01; p < 0.001), respectively. Missing variables (see Table 2) were not
signicantly affected by re history.
Variable

Stand height
Trees density
G
CBD
CKH
BCH
RBCH
Fire-scarred trees

Fire recurrence (n = 50)

n.s.
()
()
()
n.s.
n.s.
n.s.
(+)

single or combined environmental factors: low fuel accumulation,

fuel discontinuity (e.g. impeding torching and crowning), mild re
weather (e.g., low wind speed, high fuel moisture), favourable
topography or location at the least active (rear or ank) sections
of the re front. Thick back and lack of crown injury do not exclude
re-caused tree death, which may arise from smouldering combustion and the associated girdling, as observed in maritime pine
(Burrows et al., 2000). Hence, tree girdling under frequent re is
minimized due to limited accumulation of duff and woody
dead fuels. The occurrence of multiple res at short intervals
presumably also limited IB in the surrounding shrubland, reducing
the gradual tendency to kill trees at the patch-shrubland edge
(e.g., Fernandes et al., 2010). Note however that IB estimates are
solely based in the surviving trees BCH and thus are likely to be
conservative.
The large variation in burn severity metrics among individual
trees and plots (Table 2) reveals variation in re behaviour characteristics between and within res including the response to local
small-scale changes in fuel, weather, and terrain (Fernandes et al.,
2004; McRae et al., 2005; Loudermilk et al., 2012) and its interaction with tree size (Ryan, 2002). While re-behaviour variability
is common to all res, its consequences to tree injury are expected
to be more discernible across the IB range of surface res than in
higher intensity crown res.
Plot-level BCH and RBCH were associated with CKH and CR
(Table 3), as expected in surface res when overstorey foliage is
within reach of lethal temperatures and IB is the major determinant of crown scorch height (Van Wagner, 1973). Crown combustion was less strongly related to the IB proxy (BCH), suggesting a
more relevant role of vertical discontinuity. This is in contrast with

FI (n = 36)
MFI

MinFI

MaxFI

(+)
n.s.
n.s.
n.s.
(+)
(+)
(+)
n.s.

n.s.
n.s.
n.s.
n.s.
n.s.
n.s.
n.s.
n.s.

(+)
()
n.s.
n.s.
(+)
(+)
(+)
n.s.

MFI

< 4.2

<6

5.7 1.2

MaxFI

6.9 2.4

< -0.71

8.9 2.6

4.2

Aspect index
< 4.8

10.7 2.1

-0.71

MFI

4.8

12.2 2.0

Fig. 5. Regression tree analysis for canopy kill height (mean std. dev., m) in
multiple-re plots (n = 36, R2 = 0.55, 4 splits). MFI = mean re interval (yrs.);
MaxFI = maximum re interval (yrs.).

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P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

Fig. 6. Scatterplots of canopy base height versus canopy bulk density categorized by stand structure and re history (n = 50, except MFI, n = 36).

empirical crown-re likelihood models (Cruz et al., 2004), developed with data dominated by low CBH, and may also be related
with narrow variation in IB. Bark char depth and crown combustion
were tightly associated, as the former is expected to parallel the
duration of aming combustion: heavier and drier fuels produce
a longer-lasting upward heat ux that increases the odds of igniting the lower canopy (Cruz et al., 2006). Conversely, bark char
depth previously found independent of crown scorch height
(Fernandes et al., 2010) was unrelated to CKH. The fact that on
average only 15% of the live trees in a plot had visible re scars
is in accordance with the thick bark of maritime pine (Fernandes
and Rigolot, 2007) but should also be an outcome of variability
in both tree diameter and re behaviour. The amount of rescarred trees was independent of other burn severity metrics,
namely CKH (Table 3), agreeing with Collins and Stephens
(2007), but increased with re recurrence (Table 4), probably
reecting cumulative trunk injury (Falk et al., 2011). Thick back
and lack of crown injury do not exclude re-caused tree death,
which may arise from smouldering combustion and the associated
girdling, as observed in maritime pine (Burrows et al., 2000).
Hence, frequent re can minimize tree girdling by limiting the
accumulation of duff and woody dead fuels.
4.3. Stand structure in relation to re history
We found a broad range of variation in stand structure metrics
(Table 2), a presumable outcome of variable burn severity levels
(Kane et al., 2013a,b). The negative correlation of stand density variables with re recurrence (Table 4) indicates a cumulative re-thinning effect. Frequent-re pine forests in Spain (MFI  15 years) are
uneven-aged and characterized by higher stand density than typically found in this study, 4801630 trees ha1 (maritime pine;
Vega, 2000) and 483766 trees ha1 with 1931 m2 ha1 (P. nigra;
Ful et al., 2008), respectively. Our lower tree stocking (medians
of 286 trees ha1 and 15.5 m2 ha1) stems from more frequent

burning but also from the fact that most stands are remnants of
high-severity res. Repeated res at short intervals tend to shift tree
distribution to larger size classes, e.g. Holden et al. (2007). The
observed prevalence of single-storied stands (88% of the trees were
dominant or co-dominant, median value) was however unrelated
with re incidence, possibly because most were even-aged and
structurally simple from the beginning. Plots where MaxFI was 14
and 11 years respectively, recorded the highest percentages of
small and intermediate individuals, 86% and 78%. This suggests that
in these plots the amount of time between consecutive res was at
some point sufcient to allow the survival of post-re regeneration
to later res. Spatial heterogeneity in pine forest structure (as in
Fig. 4) is an attribute of low- to mixed-severity re regimes
(Larson and Churchill, 2012) and is expected to confer resilience
to re and other disturbances (Stephens et al., 2008).

4.4. Determinants of burn severity

Higher MFI and MaxFI were both associated with higher burn
severity (Table 4), probably because longer re-free periods
increase fuel accumulation and potential IB. The relationships
between re behaviour and burn severity and time since re
have been experimentally assessed in maritime pine stands and
shown to be produced by changes in surface fuel structure and
loading (Fernandes et al., 2004; Fernandes, 2009). Several NorthAmerican studies in conifer-dominated landscapes denoted
increases in burn severity with time since the previous re, for
variable periods of time (Collins et al., 2007; Malone et al., 2011;
van Wagtendonk et al., 2012; Parks et al., 2014). The regression
tree analysis for CKH in multiple-re plots (Fig. 5) further elucidated how burn severity was affected by re intervals (as fuel
accumulation proxies), plus topography. CKH was mostly determined by MFI, and the successive fuel-age thresholds for increasingly higher CKH were low and varied narrowly (46 years),

P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

owing to fast post-re fuel recovery (Fernandes and Rigolot, 2007;

Rosa et al., 2011).
Aspect was a relevant inuence on CKH when MFI P4 years,
with south aspects (warmer and drier) being prone to more severe
re as found by other studies in Mediterranean pine forest
(Oliveras et al., 2009; Fernandes et al., 2010; Viedma et al.,
2014). The CART analysis did not differentiate between east- and
south-facing slopes, possibly because the prevailing orientation
of wildres in the region (Barros et al., 2012) implies potentially
lower re intensity on west-facing (leeward) slopes. Weather conditions and re spread orientation in relation to slope have an
important role in re behaviour, which also varies markedly along
the re perimeter with consequences to burn severity (Oliveras
et al., 2009; Alvarez et al., 2013; Lecina-Diaz et al., 2014).
Because of the expected masking or overriding effect of these
sources of variability it is remarkable that the local environment
accounted for more than half of CKH variation, indicating relevance
of bottom-up controls of burn severity (Yocom et al., 2014).

4.5. Fire hazard mitigation and management implications

The thinning and pruning effects of re, and especially of recurring re, decreased crown re hazard, namely by cancelling the
potential development of active crowning in almost two thirds of
the plots (Fig. 6). Higher severity res that killed more trees and
raised CBH of the surviving individuals promoted resistance to
subsequent res. The increase in CKH and corresponding decrease
in crowning likelihood was greater for longer re intervals, but
maintenance of the residual stand requires a short re-return
interval.
Open and tall maritime pine plantations have proven to be
resistant to crown re, especially when the build-up of surface
fuels is controlled (Fernandes and Rigolot, 2007). Higher stand density has been linked to higher burn severity, in maritime pine
stands (Viedma et al., 2014) and in North-American pines (Cram
et al., 2006; Amato et al., 2013). High G can however result in
low burn severity, provided that most trees are large, as found in
Spain (Gonzlez et al., 2007; Alvarez et al., 2012). In maritime pine
stands, a dense overstorey tends to limit the shrub layer cover and
height (Castedo-Dorado et al., 2012), which is an important contributor to re intensity and tree mortality (Stephens et al., 2008;
Thompson and Spies, 2009). Whether it results from wildre or
from thinning, canopy opening favours shrub development, and
this has been implicated in subsequent high-severity res
(Holden et al., 2010; Malone et al., 2011; Lydersen and North,
2012; van Wagtendonk et al., 2012; Parks et al., 2014). Wildres
driven by extreme weather in maritime pine forest were especially
severe for G < 7 m2 ha1 (Fernandes et al., 2010), probably owing to
the combination of shrub dominance with the drier and windier
environment that characterizes an open stand. This further highlights the need for combined surface and canopy fuels treatments
(e.g., prescribed burning and thinning), the approach that best
reduces re severity (Martinson and Omi, 2013).
We infer from the results that a silvicultural prescription to
minimize crown re activity (and re hazard and effects in general) in maritime pine stands should target G and tree densities
below 20 m2 ha1 and 200 ha1, respectively, combined with
low-intensity prescribed burning at intervals of 35 years, depending on stand height. Results do not warrant a direct recommendation for pruning height, which depends of tree height. A CBH of
5.4 m was sufcient to prevent crowning of an experimental summer re with 2.5-m ame length in 3-year old fuels (Fernandes
et al., 2004). Our data suggests a 6-m CBH as the minimum
requirement for pruning height, as higher vertical discontinuity
was present in 75% of the study plots.

177

5. Conclusion
Forest structure variables such as height and density were
highly variable among the patches sampled in this study. This
resulted from preburn structure and the interaction between re
behaviour and tree size, which ultimately determines whether
trees survive and the degree to which the stand is thinned by re.
We found that post-re residual patches of maritime pine are able
to persist under a frequent re regime. An increasingly open forest
with high CBH will tend to develop as more res occur, originating
stands that are less prone to canopy damage and crown re. Fire
recurrence at short intervals tends to progressively thin the patch
and simplify its structure but is also characterized by relatively low
crown re hazard. Burn severity metrics were regulated by re
frequency to some extent, indicating that the length of the re-free
period (via accumulated fuel) is critical to re resistance and patch
persistence.
Our ndings add to the understanding of Mediterranean pines
resilience to re disturbance, adding empirical evidence to better
inform silvicultural and fuel treatment prescriptions for rehazard mitigation. We have described open and vertically
discontinuous stand structures that can be used as models for
fuel-modied areas, either as shaded fuel breaks or area-wide fuel
treatments. Results highlight the importance of both canopy and
surface fuel treatments, including prescribed burning and thinning,
to diminish the likelihood of high-severity re.
Acknowledgments
This study was funded by the European Commission 6th
Framework project FIRE PARADOX (FP6-018505). We acknowledge
the assistance provided by RIBAFLOR - Associao Florestal das
Terras de RibaDouro, Gabinete Tcnico Florestal (Cmara Municipal
de Alij) and Conselho Directivo dos Baldios de Souto de Escaro, as
well as by the land owners involved. We thank Maria Emlia
Silva and Jos Louzada, and Csar Gomes, Antnio Rodrigues and
Armindo Teixeira, for supervising and conducting the eld and
laboratory work related with destructive tree sampling and
processing. The paper beneted from the thoughtful comments
of two anonymous reviewers.
References
Agee, J.K., 1996. The inuence of forest structure on re behavior. In: Proceedings of
the 17th annual forest vegetation management conference, pp. 5268.
Agee, J.K., Skinner, C.N., 2005. Basic principles of forest fuel reduction treatments.
For. Ecol. Manage. 211, 8396. http://dx.doi.org/10.1016/j.foreco.2005.01.034.
Alvarez, A., Gracia, M., Castellnou, M., Retana, J., 2013. Variables that inuence
changes in re severity and their relationship with changes between surface
and crown res in a wind-driven wildre. For. Sci. 59, 139150.
Alvarez, A., Gracia, M., Retana, J., 2012. Fuel types and crown re potencial in Pinus
halepensis forests. Eur. J. For. Res. 131, 463474.
lvarez, J.F., 2013. Caracterizacin de la estructura vertical y dinmica del estrato de
copa en masas de pino maritimo en Portugal. Dissertao de Mestrado em Eng
Florestal, UTAD, Vila Real.
Amato, V.J.W., Lightfoot, D., Stropki, C., Pease, M., 2013. Relationships between tree
stand density and burn severity as measured by the Composite Burn Index
following a ponderosa pine forest wildre in the American Southwest. For. Ecol.
Manage. 302, 7184.
Barros, A.M.G., Pereira, J.M.C., Lund, U.J., 2012. Identifying geographical patterns of
wildre orientation: a watershed-based analysis. For. Ecol. Manage. 264, 98
107.
Boer, M.M., Macfarlane, C., Norris, J., Sadler, R.J., Wallace, J., Grierson, P.F., 2008.
Mapping burned areas and burn severity patterns in SW Australian eucalypt
forest using remotely-sensed changes in leaf area index. Remote Sens. Environ.
112, 43584369. http://dx.doi.org/10.1016/j.rse.2008.08.005.
Broncano, M.J., Retana, J., 2004. Topography and forest composition affecting the
variability in re severity and post-re regeneration occurring after a large re
in the Mediterranean basin. Int. J. Wildland Fire 13, 209216.
Burrows, N., Ward, B., Robinson, A., 2000. Behaviour and some impacts of a large
wildre in the Gnangara maritime pine (Pinus pinaster) plantation, Western
Australia. CALM Sci. 3, 251260.
Castedo-Dorado, F., Gmez-Vzquez, I., Fernandes, P.M., Crecente-Campo, F., 2012.
Shrub fuel characteristics estimated from overstory variables in NW Spain pine

178

P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179

stands. For. Ecol. Manage. 275, 130141. http://dx.doi.org/10.1016/

j.foreco.2012.03.002.
Catry, F.X., Pausas, J.G., Moreira, F., Fernandes, P.M., Rego, F., 2013. Post-re
response variability in Mediterranean Basin tree species in Portugal. Int. J.
Wildland Fire 22, 919932.
Christopoulou, A., Ful, P.Z., Andriopoulos, P., Sarris, D., Arianoutsou, M., 2013.
Dendrochronology-based re history of Pinus nigra forests in Mount Taygetos,
Southern Greece. For. Ecol. Manage. 293, 132139. http://dx.doi.org/10.1016/
j.foreco.2012.12.048.
Clarke, P.J., Knox, K.J.E., Bradstock, R.A., Munoz-Robles, C., Kumar, L., 2014.
Vegetation, terrain and re history shape the impact of extreme weather on
re severity and ecosystem response. J. Veg. Sci. 25, 10331044. http://
dx.doi.org/10.1111/jvs.12166.
Collins, B.M., Kelly, M., van Wagtendonk, J.W., Stephens, S.L., 2007. Spatial patterns
of large natural res in Sierra Nevada wilderness areas. Landsc. Ecol. 22, 545
557. http://dx.doi.org/10.1007/s10980-006-9047-5.
Collins, B.M., Miller, J.D., Thode, A.E., Kelly, M., Van Wagtendonk, J.W., Stephens, S.L.,
2009. Interactions among wildland res in a long-established Sierra Nevada
natural re area. Ecosystems 12, 114128.
Collins, B.M., Stephens, S.L., 2007. Fire scarring patterns in Sierra Nevada wilderness
areas burned by multiple wildland re use res. Fire Ecol. 3, 5367.
Connor, S.E., Arajo, J., van der Knaap, W.O., van Leeuwen, J.F.N., 2012. A long-term
perspective on biomass burning in the Serra da Estrela, Portugal. Quat. Sci. Rev.
55, 114124.
Cram, D., Baker, T., Boren, J., 2006. Wildland re effects in silviculturally treated vs.
untreated stands of New Mexico and Arizona. USDA For. Serv. Rocky Mountain
Research Station Research Paper RMRS-RP-55, Fort Collins, CO.
Cruz, M.G., Alexander, M.E., Wakimoto, R.H., 2004. Modeling the likelihood of crown
re occurrence in conifer forest stands. For. Sci. 50, 640658.
Cruz, M.G., Alexander, M.E., Wakimoto, R.H., 2005. Development and testing of
models for predicting crown re rate of spread in conifer forest stands. Can. J.
For. Res. 35, 16261639. http://dx.doi.org/10.1139/X05-085.
Cruz, M.G., Butler, B.W., Alexander, M.E., 2006. Predicting the ignition of crown fuels
above a spreading surface re. Part II: Model evaluation. Int. J. Wildland Fire 15,
6172.
Cruz, M.G., Sullivan, A.L., Gould, J.S., Sims, N.C., Bannister, A.J., Hollis, J.J., Hurley, R.J.,
2012. Anatomy of a catastrophic wildre: the Black Saturday Kilmore East re
in Victoria, Australia. For. Ecol. Manage. 284, 269285. http://dx.doi.org/
10.1016/j.foreco.2012.02.035.
Death, G., Fabricius, K.E., 2000. Classication and regression trees: a powerful yet
simple technique for ecological data analysis. Ecology 81, 31783192.
Falk, D.A., Heyerdahl, E.K., Brown, P.M., Farris, C., Ful, P.Z., McKenzie, D., Swetnam,
T.W., Taylor, A.H., Van Horne, M.L., 2011. Multi-scale controls of historical
forest-re regimes: new insights from re-scar networks. Front. Ecol. Environ.
9, 446454. http://dx.doi.org/10.1890/100052.
Fernandes, P.A.M., Loureiro, C.A., Botelho, H.N.S., 2004. Fire behaviour and severity
in a maritime pine stand under differing fuel conditions. Ann. For. Sci. 61, 537
544. http://dx.doi.org/10.1051/forest:2004048.
Fernandes, P.M., 2009. Examining fuel treatment longevity through experimental
and simulated surface re behaviour: a maritime pine case study. Can. J. For.
Res. 39, 25292535. http://dx.doi.org/10.1139/x09-145.
Fernandes, P.M., Botelho, H.S., Rego, F.C., Loureiro, C., 2009. Empirical modelling of
surface re behaviour in maritime pine stands. Int. J. Wildland Fire 18, 698710.
http://dx.doi.org/10.1071/wf08023.
Fernandes, P.M., Loureiro, C., Guiomar, N., Pezzatti, G.B., Manso, F.T., Lopes, L., 2014.
The dynamics and drivers of fuel and re in the Portuguese public forest. J.
Environ. Manage. 146, 373382. http://dx.doi.org/10.1016/j.jenvman.2014.
07.049.
Fernandes, P.M., Loureiro, C., Magalhes, M., Ferreira, P., Fernandes, M., 2012. Fuel
age, weather and burn probability in Portugal. Int. J. Wildland Fire 21, 380384.
Fernandes, P.M., Luz, A., Loureiro, C., 2010. Changes in wildre severity from
maritime pine woodland to contiguous forest types in the mountains of
northwestern Portugal. For. Ecol. Manage. 260, 883892. http://dx.doi.org/
10.1016/j.foreco.2010.06.008.
Fernandes, P.M., Rigolot, E., 2007. Fire ecology and management of maritime pine
(Pinus pinaster Ait.). For. Ecol. Manage. 241, 113.
Fernandes, P.M., Vega, J.A., Jimnez, E., Rigolot, E., 2008. Fire resistance of European
pines. For. Ecol. Manage. 256, 246255.
Fernndez-Alonso, J.M., Alberdi, I., lvarez-Gonzlez, J.G., Vega, J.A., Caellas, I.,
Ruiz-Gonzlez, A.D., 2013. Canopy fuel characteristics in relation to crown re
potential in pine stands: analysis, modelling and classication. Eur. J. For. Res.
132, 363377.
Finney, M.A., Martin, R.E., 1993. Modeling effects of prescribed re on younggrowth coast redwood trees. Can. J. For. Res. 23, 11251135.
Fournier, T.P., Battipaglia, G., Brossier, B., Carcaillet, C., 2013. Fire-scars and
polymodal age-structure provide evidence of re-events in an Aleppo pine
population in southern France. Dendrochronologia 31, 159164. http://
dx.doi.org/10.1016/j.dendro.2013.05.001.
Franklin, J.F., Mitchell, R.J., Palik, B.J., 2007. Natural disturbance and stand
development principles for ecological forestry. USDA Forest Service Gen. Tech.
Rep. NRS-GTR-19. Newtown Square, PA.
Fry, D.L., Stephens, S.L., Collins, B.M., North, M.P., Franco-Vizcano, E., Gill, S.J., 2014.
Contrasting spatial patterns in active-re and re-suppressed Mediterranean
climate old-growth mixed conifer forests. PLoS ONE 9, e88985.
Ful, P.Z., Laughlin, D.C., 2007. Wildland re effects on forest stricture over an
altitudinal gradient, Grand Canyon National Park, USA. J. Appl. Ecol. 44, 136146.

Ful, P.Z., Ribas, M., Gutirrez, E., Vallejo, R., Kaye, M.W., 2008. Forest structure and
re history in an old Pinus nigra forest, eastern Spain. For. Ecol. Manage. 255,
12341242. http://dx.doi.org/10.1016/j.foreco.2007.10.046.
Gmez-Vzquez, I., Fernandes, P.M., Arias-Rodil, M., Barrio-Anta, M., CastedoDorado, F., 2014. Using density management diagrams to assess crown re
potential in Pinus pinaster Ait. stands. Ann. For. Sci. 71, 473484. http://
dx.doi.org/10.1007/s13595-013-0350-4.
Gonzlez, J.R., Trasobares, A., Palahi, M., Pukkala, T., 2007. Predicting stand damage
and tree survival in burned forests in Catalonia (North-East Spain). Ann. For. Sci.
64, 733742.
Gutsell, S.L., Johnson, E.A., 1996. How re scars are formed: coupling a disturbance
process to its ecological effect. Can. J. For. Res. 26, 166174.
Halofsky, J.E., Hibbs, D.E., 2009. Relationships among indices of re severity in
riparian zones. Int. J. Wildland Fire 18, 584593.
Holden, Z.A., Morgan, P., Hudak, A.T., 2010. Burn severity of areas reburned by
wildres in the Gila National Forest, New Mexico, USA. Fire Ecol. 6, 7785.
http://dx.doi.org/10.4996/reecology.0603077.
Holden, Z.A., Morgan, P., Rollins, M.G., Kavanagh, K., 2007. Effects of multiple
wildland res on ponderosa pine stand structure in two southwestern
wilderness areas, USA. Fire Ecol. 3, 1833.
ICNF, 2013. IFN6 reas dos usos do solo e das espcies orestais de Portugal
continental. Resultados preliminares. Instituto da Conservao da Natureza e
das Florestas. Lisboa.
Jain, T.B., Graham, R.T., 2007. The relation between tree burn severity and forest
structure in the Rocky Mountains. In: Powers, R. (Ed.), Restoring Fire-adapted
Forested Ecosystems: Proceedings of the 2005 National Silviculture Work-shop.
USDA Forest Service, Pacic Southwest Research Station, Albany, CA, pp. 213
250 (General Technical Report PSW-GTR-203).
Kane, V.R., North, M.P., Lutz, J.A., Churchill, D.J., Roberts, S.L., Smith, D.F.,
McGaughey, R.J., Jane, J.T., Brooks, M.L., 2013a. Assessing re effects on forest
spatial structure using a fusion of Landsat and airborne LiDAR data in Yosemite
National Park. Remote Sens. Environ. 151, 89101. http://dx.doi.org/10.1016/
j.rse.2013.07.041.
Kane, V.R., Lutz, J.A., Roberts, S.L., Smith, D.F., McGaughey, R.J., Povak, N.A., Brooks,
M.L., 2013b. Landscape-scale effects of re severity on mixed-conifer and red r
forest structure in Yosemite National Park. For. Ecol. Manage. 287, 1731.
http://dx.doi.org/10.1016/j.foreco.2012.08.044.
Kolden, C.A., Lutz, J.A., Key, C.H., Kane, J.T., van Wagtendonk, J.W., 2012. Mapped
versus actual burned area within wildre perimeters: characterizing the
unburned. For. Ecol. Manage. 286, 3847. http://dx.doi.org/10.1016/
j.foreco.2012.08.020.
Larson, A.J., Churchill, D., 2012. Tree spatial patterns in re-frequent forests of
western North America, including mechanisms of pattern formation and
implications for designing fuel reduction and restoration treatments. For.
Ecol. Manage. 267, 7492.
Lecina-Diaz, J., Alvarez, A., Retana, J., 2014. Extreme re severity patterns in
topographic, convective and wind-driven historical wildres of Mediterranean
pine forests. PLoS ONE 9, e85127.
Lee, S.-W., Lee, M.-B., Lee, Y.-G., Won, M.-S., Kim, J.-J., Hong, S., 2009. Relationship
between landscape structure and burn severity at the landscape and class levels
in Samchuck, South Korea. For. Ecol. Manage. 258, 15941604. http://
dx.doi.org/10.1016/j.foreco.2009.07.017.
Loudermilk, E.L., OBrien, J.J., Mitchell, R.J., Cropper, W.P., Hiers, J.K., Grunwald, S.,
Grego, J., Fernandez-Diaz, J.C., 2012. Linking complex forest fuel structure and
re behaviour at ne scales. Int. J. Wildland Fire 21, 882893.
Loureiro, C., Fernandes, P., Botelho, H., Mateus, P., 2006. A simulation-based test of a
landscape fuel management project in the Maro range of northern Portugal.
For. Ecol. Manage. 234 (S1).
Lydersen, J., North, M., 2012. Topographic Variation in Structure of Mixed-Conifer
Forests Under an Active-Fire Regime. Ecosystems 15, 11341146. http://
dx.doi.org/10.1007/s10021-012-9573-8.
Lydersen, J.M., North, M.P., Knapp, E.E., Collins, B.M., 2013. Quantifying spatial
patterns of tree groups and gaps in mixed-conifer forests: reference conditions
and long-term changes following re suppression and logging. For. Ecol.
Manage. 304, 370382.
Madoui, A., Leduc, A., Gauthier, S., Bergeron, Y., 2010. Spatial pattern analyses of
post-re residual stands in the black spruce boreal forest of western Quebec.
Int. J. Wildland Fire 19, 11101126.
Malone, S.L., Kobziar, L.N., Staudhammer, C.L., Abd-Elrahman, A., 2011. Modeling
relationships among 217 res using remote sensing of burn severity in
Southern pine forests. Remote Sens. 3, 20052028. http://dx.doi.org/10.3390/
rs3092005.
Martinson, E.J., Omi, P.N., 2013. Fuel treatments and re severity: a meta-analysis.
Res. Pap. RMRS-RP-103WWW. U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fort Collins, CO.
McNab, W.H., 1977. An overcrowded loblolly pine stand thinned with re. South. J.
Appl. For. 1, 2426.
McRae, D.J., Jin, J.-Z., Conard, S.G., Sukhinin, A.I., Ivanova, G.A., Blake, T.W., 2005.
Infrared characterization of ne-scale variability in behavior of boreal forest
res. Can. J. For. Res. 35, 21942206. http://dx.doi.org/10.1139/x05-096.
Morgan, P., Hardy, C.C., Swetnam, T.W., Rollins, M.G., Long, D.G., 2001. Mapping re
regimes across time and space: understanding coarse and ne-scale re
patterns. Int. J. Wildland Fire 10, 329342.
Morgan, P., Keane, R.E., Dillon, G.K., Jain, T.B., Hudak, A.T., Karau, E.C., Sikkink, P.G.,
Holden, Z.A., Strand, E.K., 2014. Challenges of assessing re and burn severity using
eld measures, remote sensing and modelling. Int. J. Wildland Fire 23, 10451060.

P.M. Fernandes et al. / Forest Ecology and Management 347 (2015) 170179
Oliveras, I., Gracia, M., Mor, G., Retana, J., 2009. Factors inuencing the pattern of
re severities in a large wildre under extreme meteorological conditions in the
Mediterranean basin. Int. J. Wildland Fire 18, 755764.
Oliveira, S.L., Pereira, J.M., Carreiras, J.M., 2012. Fire frequency analysis in Portugal
(19752005), using Landsat-based burnt area maps. Int. J. Wildland Fire 21,
4860.
Parks, S.A., Miller, C., Nelson, C.R., Holden, Z.A., 2014. Previous res moderate burn
severity of subsequent wildland res in two large western US wilderness areas.
Ecosystems 17, 2942. http://dx.doi.org/10.1007/s10021-013-9704-x.
Perera, A.H., Dalziel, B.D., Buse, L.J., Routledge, R.G., 2009. Spatial variability of
stand-scale residuals in Ontarios boreal forest res. Can. J. For. Res. 39,
945961.
Peterson, D.L., Ryan, K.C., 1986. Modeling postre conifer mortality for long-range
planning. Environ. Manage. 10, 797808.
Pimont, F., Prodon, R., Rigolot, E., 2011. Comparison of postre mortality in endemic
Corsican black pine (Pinus nigra ssp. laricio) and its direct competitor (Pinus
pinaster). Ann. For. Sci. 68, 425432. http://dx.doi.org/10.1007/s13595-0110031-0.
Ribeiro, O., Lautensach, H., Daveau, S., 1988. Geograa de Portugal. II. O Ritmo
Climtico e a Paisagem. Edies Joo S da Costa, Lisboa.
Romn-Cuesta, R.M., Gracia, M., Retana, J., 2009. Factors inuencing the formation
of unburned forest islands within the perimeter of a large forest re. For. Ecol.
Manage. 258, 7180.
Rosa, I.M.D., Pereira, J.M.C., Tarantola, S., 2011. Atmospheric emissions from
vegetation res in Portugal (19902008): estimates, uncertainty analysis, and
sensitivity analysis. Atmos. Chem. Phys. 11, 26252640. http://dx.doi.org/
10.5194/acp-11-2625-2011.
Ryan, K.C., 1982. Techniques for assessing re damage to trees. In: Lotan, J. (Ed.),
Proceedings of the Symposium: Fire its, Field Effects. Intermountain Fire
Council, pp. 111.
Ryan, K., 2002. Dynamic interactions between forest structure and re behavior in
boreal ecosystems. Silva Fenn. 36, 1339. http://dx.doi.org/10.14214/sf.548.
Santos, C., Almeida, J., 2003. Caracterizao especial de um ndice de produtividade
nos povoamentos de pinheiro-bravo em Portugal. Finisterra 38, 5165.
SAS Institute Inc., 2010. JMP Version 9. Cary, NC.
Seidl, R., Rammer, W., Spies, T.A., 2014. Disturbance legacies increase the resilience
of forest ecosystem structure, composition, and functioning. Ecol. Appl. 24,
20632077. http://dx.doi.org/10.1890/14-0255.1.
Sikkink, P.G., Keane, R.E., 2012. Predicting re severity using surface fuels and
moisture. Res. Pap. RMRS-RP-96. U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station. Fort Collins, CO.

179

Slimani, S., Touchan, R., Derridj, A., Kherchouche, D., Gutirrez, E., 2014. Fire history
of Atlas cedar (Cedrus atlantica Manetti) in Mount Chlia, northern Algeria. J.
Arid Environ. 104, 116123. http://dx.doi.org/10.1016/j.jaridenv.2014.02.008.
StataCorp., 2007. Stata Statistical Software: Release 10. College Station, TX.
Stephens, S.L., Fry, D.L., Franco-Vizcano, E., 2008. Wildre and spatial patterns in
forests in Northwestern Mexico: the United States wishes it had similar re
problems. Ecol. Soc. 13, 10, <http://www.ecologyandsociety.org/vol13/iss2/art10>.
Thompson, J.R., Spies, T.A., 2009. Vegetation and weather explain variation in crown
damage within a large mixed-severity wildre. For. Ecol. Manage. 258, 1684
1694. http://dx.doi.org/10.1016/j.foreco.2009.07.031.
Touchan, R., Baisan, C., Mitsopoulos, I.D., Dimitrakopoulos, A.P., 2012. Fire history in
European black pine (Pinus nigra Arn.) forests of the Valia Kalda, Pindus
mountains, Greece. Tree-Ring Res. 68, 4550.
Van Wagner, C., 1973. Height of crown scorch in forest res. Can. J. For. Res. 3, 373
378.
Van Wagner, C.E., 1977. Conditions for the start and spread of crown re. Can. J. For.
Res. 7, 2334.
Van Wagtendonk, J.W., van Wagtendonk, K.A., Thode, A.E., 2012. Factors associated
with the severity of intersecting res in Yosemite National Park, California, USA.
Fire Ecol. 7, 1131. http://dx.doi.org/10.4996/reecology.0801011.
Vega, J., Jimenez, E., Vega, D., Ortiz, L., Prez, J.R., 2011. Pinus pinaster Ait. tree
mortality following wildre in Spain. For. Ecol. Manage. 261, 22322242. http://
dx.doi.org/10.1016/j.foreco.2010.10.019.
Vega, J.A., 2000. Resistencia vegetativa ante el fuego atravs de la historia de los
incendios. In: Vlez, R. (Ed.), La Defensa Contra Incendios Forestales:
Fundamentos y Experiencias. McGraw-Hill, Madrid, p. 4.66.
Viedma, O., Quesada, J., Torres, I., Santis, A.D., Moreno, J.M., 2014. Fire severity in a
large re in a Pinus pinaster forest is highly predictable from burning conditions,
stand structure, and topography. Ecosystems 114. http://dx.doi.org/10.1007/
s10021-014-9824-y.
Windisch, A.G., Good, R.E., 1991. Fire behavior and stem survival in the New Jersey
pine plains. In: High Intensity Fire in Wildlands: Management Challenges and
Options, Proc. 17th Tall Timbers Fire Ecology Conf., Tall Timbers Res. Stn.,
Tallahassee, pp. 273299.
Wu, Z., He, H.S., Yang, J., Liu, Z., Liang, Y., 2014. Relative effects of climatic and local
factors on re occurrence in boreal forest landscapes of northeastern China. Sci.
Tot. Environ. 493, 472480.
Yocom, L.L., Ful, P.Z., Falk, D.A., Garca-Domnguez, C., Cornejo-Oviedo, E., Brown,
P.M., Villanueva-Daz, J., Cerano, J., Montao, C.C., 2014. Fine-scale factors
inuence re regimes in mixed-conifer forests on three high mountains in
Mexico. Int. J. Wildland Fire 23, 959968.