Forest Ecology and Management 187 (2004) 85–103
The effects of wildfires on wood-eating beetles in deciduous
forests on the southern slope of the Swiss Alps
Marco Morettia,*, Sylvie Barbalatb
a
WSL Swiss Federal Research Institute, Sottostazione Sud delle Alpi, PO Box 57, CH-6504 Bellinzona, Switzerland
b
Rue des Brévards 2, CH-2000 Neuchâtel, Switzerland
Received 15 August 2002; received in revised form 5 February 2003; accepted 12 June 2003
Abstract
The effect of fires on Cerambycidae, Buprestidae and Lucanidae were studied at 23 sites within a chestnut forest in southern
Switzerland. We compared six unburnt sites, two freshly burnt sites, eight sites which burned once at different times in the last 30
years, and seven sites where fires occurred repeatedly in the last 30 years. The diversity and the species composition of the three
xylobiont families were related to various ecological variables at two levels of spatial scale, a small scale of 0.25 ha and a large
scale of 6.25 ha. These variables were: fire frequency, time since the last fire, clear cutting after the fire, forest structure, amount
of dead wood, and habitat mosaic. The fire does not have a direct effect on the xylobiont beetles community at small scale;
however, fire has an indirect effect by maintaining a relatively open forest structure. The mosaic of forest areas burnt with
different frequencies and at different times was an important factor influencing species richness and species composition at the
large spatial scale.
Data presented here supports the strategy to conserve the diversity and includes species composition of xylobiont fauna in
deciduous forests: (i) at small spatial scale, to maintain highly structured and relatively open stands with large amounts of dead
wood and big oak trees; (ii) at large spatial scale, to favour a mosaic of different forest habitats and successional stages. A forest
offering a good structural diversity is important for maintaining landscape complexity and thus a high species richness of
xylophagous beetles.
# 2003 Elsevier B.V. All rights reserved.
Keywords: Forest fires; Post-fire succession; Landscape ecology; Coleoptera; Cerambycidae; Buprestidae; Lucanidae; Biodiversity; Species
richness; Species composition; Conservation; Switzerland
1. Introduction
Fire is one of the most important disturbance factors
in natural ecosystems throughout the world. Recently,
the role of fire was reinterpreted from the viewpoint of
disturbance ecology and biodiversity conservation
*
Corresponding author. Tel.: þ41-91-8215236;
fax: þ41-91-8215239.
E-mail address: [email protected] (M. Moretti).
(e.g. Granström, 1996; Goldammer et al., 1997; White
and Jentsch, 2001). Most studies have been carried out
in fire-prone regions or in specific fire-climax ecosystems such as Mediterranean shrub-land, Savanna,
Chapparal, Boreal forests (see DeBano et al. (1998)
for a review). We know much less about the role of fire
in temperate forests where winter fires predominate
(e.g. Dajoz, 1998; Brawn et al., 2001). This is the case,
for example, in some parts of the southern slopes of the
Alps, where most of the wildfires occur during the
0378-1127/$ – see front matter # 2003 Elsevier B.V. All rights reserved.
doi:10.1016/S0378-1127(03)00314-1
86
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
dormant period between December and April (e.g.
Conedera et al., 1996).
Concerning the effects of fires on fauna, most works
focused on ground and surface dwelling invertebrates
over a time span of not more than 10–15 years (see
Nunes et al. (2000) and Wikars (2001) for a review).
Very little is known about the long-term effect of fire
(e.g. Siemann et al., 1997; York, 2000) often because
precise information is not available about the fire
history at the sites studied. A few studies have considered the response of saproxylic fauna to fire, especially in boreal forests (e.g. Schauermann, 1980;
Muona and Rutanen, 1994; Wikars, 1997). Little is
known about the effect of wildfires on wood-eating
invertebrates in the Mediterranean area, though fire is
an important ecological factor in Mediterranean forests (Trabaud, 2000) and saproxylic invertebrates are
very important in these forests in economic, ecologic
and conservation terms (Speight, 1989; GreatorexDavis and Marrs, 1992; Dajoz, 2000).
Many authors have shown a strong relationship
between xylobiont invertebrates and the three following factors: age of timber, quantity of deadwood and
deadwood connectivity (e.g. Hartmann and Sprecher,
1990; Irmler et al., 1996; Okland et al., 1996; Nilsson
and Baranowski, 1997; Schiegg, 2000; Ranius, 2002).
The influence of fire on these factors is known, but has
not yet been discussed from a functional point of view.
From the forest ecology point of view, it is hence
important to investigate the response of saproxylic
invertebrates to fire as an important functional group
in the food chain of living and dead wood.
Cerambycid, buprestid and lucanid beetles (Coleoptera: Buprestidae, Cerambycidae, Lucanidae) are
xylophagous, being wood feeders mostly during their
larval stages; depending on the species, these may
colonise living trees, dead wood or rotten stumps. The
adults feed on flowers, leaves and stems or do not
feed at all (Hellrigl, 1978; Bense, 1995; Dajoz, 2000).
Among saproxylics these three beetle families are
good forest bioindicators because their ecology is well
known, they are relatively stable taxonomically, and
are known to be sensitive to changes in forest habitat
(Starzyk and Witkowski, 1981; Gutowski, 1995; Barbalat, 1998). Many buprestid and cerambycid beetles
are known to be pyrophilous (Wikars, 1997).
The aims of this study are to examine the response
of buprestid, cerambycid and lucanid beetles to fire by
relating their presence to the fire history (fire frequency and time elapsed since last fire), and to assess
how they are influenced by the spatial structure of their
habitat. The main questions are: (1) Do single or
repeated fires affect the diversity of xylophagous
beetles? (2) Which species suffer from fire and which
profit from it? (3) How do the communities respond to
fire frequency and post-fire conditions? (4) Which
environmental factors influence the xylophagous beetles at two spatial scales?
2. Methods
2.1. Study area
This research is part of a multi-disciplinary study
on the effects of sporadic and regular wildfires on
European chestnut forests (Castanea sativa Mill.)
in southern Switzerland (e.g. Delarze et al., 1992;
Conedera et al., 1996; Tinner et al., 1999; Moretti
et al., 2002). The study area is located along a small
geographically homogeneous, south-facing slope
(450–850 m a.s.l.) near Locarno (088440 E; 468090 N),
Canton of Ticino (southern Switzerland) (Fig. 1). The
study area has a moist, warm temperate climate. Unlike
the Mediterranean climate of more southerly regions
the rainfall is higher in summer (June–September:
800 mm) than in winter. For this reason the area is
prone to fast-spreading surface fires during the period
of vegetation dormancy (December–April). Tinner
et al. (1999) suggest that southern Switzerland, and
probably all of the southern slopes of the Alps, are a
fire-prone area and that fire has to be considered as a
natural environmental factor. More details about the
study area are given in Moretti et al. (2002). The
vegetation is dominated by European chestnut (mainly
on acidic soil) that were introduced in the area during
the Roman period, about 2000 years ago (Tinner et al.,
2000; Conedera and Tinner, 2000).
The sampling design was based on a ‘space-fortime substitution’ (Pickett, 1989). For this purpose the
slope was divided into six sectors (A–F, see Fig. 1), in
which a total of 23 study sites were selected according
to their stage of succession following burning. The
Wildfire Database of southern Switzerland (Conedera
et al., 1996) and dendrochronological methods were
the principal sources of information on the fire history
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
87
Fig. 1. Location of the six sectors (A–F) in the study area (dark stippled) on the southern slope of the Alps (Ticino, Switzerland).
of the various sites over the past 30 years (1968–1997).
For the classification of the study sites to the different
categories of fire regime two parameters were used:
frequency as ‘number of fires’ and age as ‘time since
last fire’. The other principal components of the fire
regime (type, season, intensity) differ very little in the
area, i.e. fast spreading surface fires of low to medium
intensity that occur in wintertime. Most of the sites
were left unmanaged during the last 30 years. The time
since the last intervention (such as clear cutting or
thinning) was taken in account. The total area burnt
each year is subject to great annual variation according
to meteorological factors and it ranges from a minimum of 15 ha to a maximum of 70,000 ha (mean value
in 30 years is 730 ha; Conedera et al., 1996).
The sites were classified as follows: sites without
fires in the last 30 years, used as control sites
(unburnt), sites where sampling started 1–2 weeks
after a fire (freshly burnt), sites burnt only once (single
fire), sites burnt three to four times (repeated fires).
Because each sector had a different fire history, it was
not possible to have replicate sites with the same fire
regime in each sector. However, one unburnt (control)
site was defined in each sector (Table 1).
The vegetation structure was different in the 23
study sites, depending on the fire frequency and time
since the last fire (Moretti et al., 2002). Table 2
summarises the most important characteristics.
2.2. Sampling methods
2.2.1. Faunistic data
Cerambycids, buprestids and lucanids were sampled
using ‘‘combi-traps’’ (combination of yellow water pan
and window trap) made of two vertically and crosswise
installed transparent sheets of plexiglass (50 cm 40 cm) mounted over a 45 cm wide, bright yellow
plastic funnel and placed at a height of 1.5 m above
ground (Duelli et al., 1999). The funnels were filled
with water to which soap was added to decrease surface
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M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
Table 1
Description of the study sites (1–23) with regard to fire frequency and time since last firea
Sectors
of the
study
area
Altitude
(m a.s.l.)
A
B
460
560
SE
SE
1C
3C
C
460
S
8C
D
E
F
Asp.
Unburnt
sites (C)
>30 years
(n ¼ 6)
660
590
730
S
SE
S
Burnt sites and time since the last fire
Freshly burnt (F)
<1 year
(n ¼ 2)
Single fire (S)
1–2 years
(n ¼ 2)
7–14 years
(n ¼ 3)
Repeated fires (R)
22–24 years
(n ¼ 3)
2–3 years
(n ¼ 2)
6–17 years
(n ¼ 5)
2S
22 F
23 F
16 C
18 C
21 C
4S
5R
9S
10 Sb
13 S
12 Rc
15 S
14 S
6R
7R
11 Rd
17 S
19 R
20 R
a
C: none, control; F: freshly burnt; S: single fire; R: repeated fires; Asp.: aspect; n: number of study sites.
Thinned 4 years before our study.
c
Thinned 9 years before the last fire.
d
Thinned 10 years before our study.
b
tension, and a bactericide to prevent decomposition of
the specimens. Yellow traps were used in several
studies, as well at the ground level (Hartmann and
Sprecher, 1990) as in the trees (Gutowski, 1995). It
proved to be efficient for the capture of anthophilous
Buprestids, mainly the genus Anthaxia (Barbalat,
1995). The efficiency, selectivity and spatial range
of environmental influence of three methods of trapping saproxylic beetles have been discussed by Okland
(1996). According to this author, window traps are
more suitable for comparing different forest environments; the captures using this method are influenced
by ecological conditions over wide areas, but are
almost unaffected by substrate conditions in the near
surroundings of the traps. We expected our yellow
window traps (combi-traps) to be mainly efficient in
catching flying as well as flower visiting species,
usually attracted by yellow surfaces. Apterous and less
mobile species that live on the ground were sampled
using pitfall traps (plastic funnels, 15 cm diameter)
according to the method of Obrist and Duelli (1996).
For both trap types, the probability of an animal being
caught is a function both of the number of individuals
present and of their activity and ecology. For this
reason, when we use the expression ‘number of individuals’, we include both abundance and activity.
Table 2
Description of some environmental variables sampled at sites with different fire regime (fire frequency and time elapsed since the last fire)
Tree cover (%)
Bush cover (%)
Grass cover (%)
Diameter of dominant trees (cm)
Number of plant species
Number of stools per shoot
Dead stools per shoot
Unburnt sites
>30 years
(n ¼ 6)
Freshly burnt
<1 year
(n ¼ 2)
Single fire
Repeated fires
1–2 years
(n ¼ 2)
7–14 years
(n ¼ 2)
22–24 years
(n ¼ 3)
2–3 years
(n ¼ 2)
6–17 years
(n ¼ 5)
90
5
15
20–30
9
9
2
20
5
5
10–20
9
76
16
40
15
40
20–30
20
28
12
84
15
15
20–30
19
11
6
90
5
25
20–30
11
18
8
50
25
60
10–20
7
27
14
90
15
30
10–20
11
22
13
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
Three combi-traps and three pitfall traps (two triplet
sets of traps) were installed randomly on each of the
23 study sites to achieve a variation of the local habitat
heterogeneity. Combi-traps were close to each pitfalltrap, while the distance between the three trap sites was
always at least 10 m. The traps were emptied weekly
from the beginning of March to the end of September
1997, resulting in a total of 28 sampling periods. All
adult individuals were later identified to species level
using keys of Freude et al. (1964–1983) and Bense
(1995). Nomenclature followed Bense (1995) and
Lohse and Lucht (1992). Reference collections of
labelled and identified beetle specimens are stored at
the Natural History Museum of Lugano, Switzerland.
2.2.2. Environmental variables
Most of the Buprestid and Cerambycid species are
flying insects and some of the most mobile antho- and
heliophilous species exploit food resources over distances of a few hundred meters. With this in mind,
and to gain a better understanding of the dynamics of
wood eating groups in our study region, we considered
two groups of environmental variables at two different
spatial scales: a small scale of 0.25 ha (50 m 50 m)
and a large scale of 6.25 ha (250 m 250 m) where 24
and 17 variables, respectively, were assessed (Tables 3
and 4).
2.3. Data analysis
The samples of the three sets of traps at each study
site (three combi-traps and three pitfall traps) were
pooled in order to avoid autocorrelation; this resulted
in one sample per study site. Mean values (S.E.) of
species richness and number of individuals were
calculated with regard to both ‘number of fire’ and
‘time since last fire’. As homogeneity of variances was
not achieved even by data transformation, non-parametric Kruskal–Wallis ANOVA was applied to the
data (Legendre and Legendre, 1998).
Species diversity was assessed using two measures
with complementary properties
(Magurran, 1991).
P
Shannon index (Hs ¼ pi ln pi , where pi relative
abundance of the ith species), and evenness (Es ¼ Hs /
ln S, where S is the number of species).
Before testing the relationship between environmental variables and faunistic parameters, a principal
component analysis (PCA) was performed to find the
89
correlation among the environmental variables, and to
find which variables are most strongly associated with
the various axes (Tables 3 and 4).
The influence of the environmental variables on the
number of species and individuals was tested by a
stepwise multiple regression at both local and large
spatial scale (Legendre and Legendre, 1998). Environmental effects on the beetle communities was tested by
canonical correspondence analysis (CCA) (ter Braak,
1986); this is a multivariate technique for relating
species composition and abundance to underlying
environmental gradients (direct gradient analysis).
Environmental variables were selected using the
manual stepwise procedure implemented in CANOCO
(ter Braak, 1986), while ‘geographical coordinates’
were allocated as co-variables, in order to control
the geographical location of the sites (Legendre and
Legendre, 1998). This procedure of forward selection
adds environmental variables one at a time, using a
Monte Carlo permutation test (P < 0:05; 500 randomisations), until no other variables significantly explain
residual variation in species composition. For the analysis, we considered only the 32 species for which at
least three individuals were sampled. The number of
individuals of the selected species was log-transformed
(logðxi þ 1Þ), where xi corresponds to the number of
individuals, in order to reduce the weighting of the very
abundant species.
To evaluate the data from a conservation point of
view, we compiled lists of the endangered species
in Switzerland and in Europe by consulting different
sources (Collins and Wells, 1987; Speight, 1989;
Gepp, 1994; Binot et al., 1998).
The assumption of cause–effect relationships
between faunistic and environmental variables was
based on biological interpretation and the convergence
of different statistical relationships. Ecological requirements of the species were found in the abovementioned works as well as in Hellrigl (1978) and in
Bense (1995).
3. Results
3.1. Species richness and diversity after the fire
A total of 1152 individuals representing 56 species
were recorded (40 Cerambycidae, 14 Buprestidae and
90
Table 3
PCA of the site environmental variables (50 m surrounding each site) divided into seven principal axesa
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
a
Variables with values higher than 0.500 were considered to be correlated. In bold type are the variables related to wildfire. The PCA divided the local environmental variables
into seven principal axes explaining a total of 86.1% of the variance. The first two axes explain more than half of the variance. The first one (27.7% of variance) is associated most
strongly with time elapsed since the last fire and therefore also with variables related mainly to both cover and structure of the forest stand (e.g. tree cover, number of shoots per
stool). A group of variables associated with the second axis (18.0% of variance) describes the age and the growth stage of the forest stand (e.g. tree height, diameter of the dominant
trees). This axis is also related to fire frequency.
Table 4
PCA of the landscape environmental variables (250 m surrounding each site) divided into six principal axesa
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
a
Variables with values higher than 0.500 were considered to be correlated. The PCA divided the landscape environmental variables into six principal axes explaining a total of
84.6% of the variance. The first two axes explained more than half of the total variance. The first one (25.4% of variance) is associated mainly with the repeatedly burnt areas (three
to five fires) and the managed areas (cut areas). A group of variables associated with the second axis (19.7% of variance) describes principally freshly burnt areas and sporadically
burnt areas (two fires). This axis is also related to unburnt areas.
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M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
Species
richness
92
20
15
10
5
0
a
a
Number of
individuals
Shannon
a
a
a
a
a
a
a
a
30
20
10
0
a
ab
b
0
300
200
100
0
3.0
3.0
2.0
2.0
1.0
1.0
100
50
a
a
> 30
freshly
burnt
a
a
a
2-3
6-17
0.0
0.0
1.5
1.0
Evenness
a
a
1.0
0.5
0.5
0.0
0.0
unburnt
1 fire
3-4 fires
unburnt
Number of fires
1-2
7-14
22-24
single fires
repeated
fires
Time since last fire (years)
Fig. 2. Mean number of species and of individuals (S.E.) and diversity (mean variance) in each study site grouped in classes of number of
fires and time elapsed since the last fire. Bars with different letters are significantly different. The number of species does not differ
significantly. Significantly more individuals were collected in repeated burnt sites (three to four fires) (Mann–Whitney U-test; n ¼ 13;
P < 0:05) and particularly in freshly burnt sites (Mann–Whitney U-test; n ¼ 8; P < 0:05).
2 Lucanidae). Thirty-two species (58%) were sampled
with three or more individuals, while 12 species (22%)
were observed exclusively in one study site.
Species richness at burnt sites was similar to that at
unburnt sites with regard to both fire frequency and to
time elapsed since the last fire (Kruskal–Wallis test;
n ¼ 22; n.s.) (Fig. 2). On the other hand, the number of
individuals varied significantly according to fire frequency (Kruskal–Wallis test; n ¼ 22; P < 0:05), especially at repeatedly burnt sites (Mann–Whitney U-test
with Bonferroni correction; n ¼ 13; P < 0:02),
and according to time elapsed since the last fire
(Kruskal–Wallis test; n ¼ 22; P < 0:05). Large numbers of individuals were trapped on recently burnt
sites where fire occurred repeatedly, but the high variation among the study sites limited the significance
level. Shannon index and evenness show negligible
differences between sites affected by different fire
regimes.
3.2. Environmental factors affecting faunistic
diversity
Table 5 shows that at a small spatial scale (0.25 ha
surrounding the study sites) species number is influenced by three factors: ‘time since the last cut’
(TIME_CUT), ‘tree cover’ (TREECOV) and ‘number
of plant species’ (NSPVEG).
The number of individuals also appears to be
affected by ‘time since the last cut’ and ‘tree cover’,
as well as by ‘number of fires’ (N_FIRE) and ‘distance
between stools’ (DIST_STO). In both cases the
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
93
Table 5
Influence of the environmental variables on the number of species (A) and on the log-number of individuals (B)a
Environmental variables
Coefficient
S.E.
t
P
Time elapsed since the last clear cut
Tree cover in percentage classes
Number of the plant species
0.020
0.006
0.011
0.004
0.002
0.006
4.519
4.010
1.677
0.000
0.001
0.110
(B) log-number of individualsc
N_FIRE
Number of fires
TIME_CUT
Time elapsed since the last clear cut
TREECOV
Tree cover in percentage classes
DIST_STO
Distance between stools
0.072
0.021
0.006
0.152
0.026
0.005
0.001
0.046
2.764
4.650
4.294
3.324
0.013
0.000
0.000
0.004
b
(A) Number of species
TIME_CUT
TREECOV
NSPVEG
a
The variables were selected by stepwise multiple regression analysis among 24 environmental variables collected 0.25 ha surrounding
each study site (small spatial scale).
b 2
R ¼ 0:743, F-ratio ¼ 18:281 and P ¼ 0:000.
c 2
R ¼ 0:828, F-ratio ¼ 21:602 and P ¼ 0:000.
selected environmental variables have high explanatory power (R2 ¼ 0:743, P < 0:001 and R2 ¼ 0:828,
P < 0:001, respectively).
The environmental factors influencing the number
of species at a larger spatial scale (6.25 ha surrounding
the study sites) are ‘unburnt area’ (UNBURNT),
‘rocky area’ (ROCKS), ‘open land area’ and total
burnt area in the last 30 years (from the more recent
one BURNT <1 y to the oldest one BURNT 18–30 y)
(Table 6). At a large spatial scale the number of
individuals seems to be affected by the presence of
half-open habitat like ‘rocky area’ (ROCKS) and ‘path
length’ (PATH), as well as by recently burnt stands
(BURNT 1–3 y; BURNT 7–17 y) and ‘environmental
diversity’ (H_ENV). Both groups of selected variables
are significant (R2 ¼ 0:591, P < 0:05 and R2 ¼ 0:588,
P < 0:05, respectively) but have a lower explanatory
power at a large than at a small spatial scale.
Table 6
Influence of the environmental variables on the number of species (A) and on the log-number of individuals (B)a
Environmental variables
Coefficient
S.E.
t
P
0.002
0.002
0.001
0.002
0.002
0.002
0.002
0.001
0.001
0.001
0.001
0.001
0.001
0.001
2.496
3.382
2.523
2.477
2.853
2.459
2.370
0.025
0.004
0.023
0.026
0.012
0.027
0.032
0.000
0.000
0.000
0.001
0.450
0.000
0.000
0.000
0.000
0.147
2.155
3.334
2.459
2.361
3.070
0.046
0.004
0.025
0.030
0.007
b
(A) Number of species
UNBURNT
ROCKS
OPEN_AREA
BURNT <1 y
BURNT 1–3 y
BURNT 7–17 y
BURNT 18–30 y
Unburnt area since 30 years
Rocky area
Open environments area
Burnt area 0–1 year before the study
Burnt area 1–3 years before the study
Burnt area 7–17 years before the study
Burnt area 18–30 years before the study
(B) log-number of individualsc
ROCKS
Rocky area
BURNT 1–3 y
Burnt area 1–3 years before the study
BURNT 7–17 y
Burnt area 7–17 years before the study
PATH
Path length
H_ENV
Environmental diversity
a
The variables were selected by stepwise multiple regression analysis among 17 variables collected 6.25 ha surrounding each study site
(large spatial scale).
b 2
R ¼ 0:591, F-ratio ¼ 3:097 and P ¼ 0:031.
c 2
R ¼ 0:588, F-ratio ¼ 4:845 and P ¼ 0:006.
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M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
Table 7
Number of species and individuals (in brackets) of the exclusive, scarce, endangered cerambycid, buprestid and lucanid beetles sampled at
sites with different fire regime and clearinga
Class of study site
Exclusive species
Scarce species
Endangered species
Unburnt sites
(n ¼ 6)
Recent fires and
clearing (n ¼ 6)
Single fires
(n ¼ 5)
Repeated
fires (n ¼ 6)
4 (5)
4 (5)
9 (35)
13 (19)
10 (12)
18 (157)
1 (1)
1 (1)
9 (41)
2 (2)
2 (2)
10 (68)
a
Exclusive species: species sampled only in one class of study site; scarce species: species of which three or less individuals were sampled
during the study; endangered species (see Table 8 for the definitions). The class ‘‘recent fires and clearing’’ includes: freshly burnt sites (sites
22 F, 23 F), sites burnt 1–2 years before our study (sites 2 S, 12 R, 17 S) and sites recently managed (site 10 S). For the other classes, see
Table 1.
3.3. Exclusive, rare and endangered species
after the fire
Chlophorus figuratus, Pachtodes cerambyciformis and
Stenopterus rufus).
Among the species collected, only four (each represented by only one specimen) were found exclusively
in unburnt stands; 16 species were observed only at
burnt and clear-cut sites, while 17 species were scarce
in our study sites (Table 7).
Moreover, we found 23 species that are endangered
in different European countries or are indicators of
forests of international conservation importance (Collins and Wells, 1987; Speight, 1989; Binot et al., 1998;
Gepp, 1994) (Table 8); 14 of these were found exclusively at burnt and clear cut sites, and only 2 in unburnt
sites.
3.5. Environmental factors affecting community
succession
3.4. Post-fire succession of the dominant
species
Table 9 shows the succession of the dominant
species after single and repeated fires with regard to
the time elapsed since the last fire. At unburnt sites,
Parmena unifasciata and Leiopus nebulosus were
dominant together with nine co-dominant shade-tolerant species. After the fire the community structure
and the species composition changed, with helio- and
anthophilous species becoming dominant and characterising the burnt sites in the different successional
stages. Some of them were already present in the intact
stands (Agrilus laticornis, A. angustulus, Leptura
maculata, Grammoptera ruficornis and Clytus arietis), while others appeared for the first time, especially
at freshly and recently burnt sites (Callimus angulatus,
At a small spatial scale (0.25 ha surrounding each
study site) canonical correspondence analysis selected
four environmental variables which explained 32.4%
of the variation of species composition of buprestids
and cerambycids (Fig. 3 and Table 10). The first axis,
and all canonical axes together, are significant (Monte
Carlo test, P ¼ 0:005). The variance of the first axis
was mainly due to ‘tree cover’ (TREECOV), which
shows the difference between open and shady sites. On
the one end of axis 1, sites 12, 22 and 23 had been
recently burnt while site 10 was clear cut 4 years before
our study. These sites host a community of heliophilous
species including the cerambycids Stenurella bifasciata, Anastrangalia sanginolenta, Chlorophorus sartor, C. figuratus, Pachytodes cerambyciformis, Leptura
maculata and Stenopterus rufus (Table 11). The correlation between the time elapsed since the last fire and
flowering plant cover was negative (Pearson correlation 0.650; Bonferroni test, n ¼ 22, P ¼ 0:006). On
the other hand, the community of species found in the
unburnt forests and in stands that have not been burnt
for more than 20 years prefer or tolerate shade. This is
the case of the cerambycids Alosterna tabacicolor,
Prionus coriarius, Pogonocherus hispidulus, Parmena
unifasciata and Leiopus nebulosus.
The second axis is related to the ‘occurrence of
oaks’ (QUERCUS) and also partially the ‘amount of
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
95
Table 8
Endangered cerambycid, buprestid and lucanid species sampled at sites with different fire regime and clearinga,b
a
The class ‘‘recent fires and clearing’’ includes: freshly burnt sites (sites 22 F, 23 F), sites burnt 1–2 years before our study (sites 2 S, 12 R,
17 S) and sites recently managed (site 10 S). For the other classes, see Table 1.
b
BC: species included in Appendix II of the Bern Convention (Collins and Wells, 1987); EU: species useful to identify forests of
international importance to natural conservation (Speight, 1989); CH: species of potential national conservation concern (Barbalat, personal
communication); D: red list of Germany (Binot et al., 1998); A: red list of Austria (Gepp, 1994).
Species protected by the Swiss law.
the dead wood’ (DEADWOOD) (Fig. 3 and Table 10).
The oak-depending species at sites 12, 13, 21, 22 and
23 (Table 11) were the buprestids Agrilus sulcicollis,
A. angustulus, A. laticornis, A. graminis and Nalanda
fulgidicollis and the cerambycids Phymatodes alni,
Plagionotus arcuatus and Xylotrechus antilope. The
presence of oak was positively correlated with fire
frequency (Pearson correlation 0.593; Bonferroni test,
n ¼ 12, P ¼ 0:04).
At a large spatial scale (6.25 ha), the CCA selected
four environmental variables, explaining 30.7% of the
variation of the same sites and communities (Table 10).
The variance of the first axis was mainly due to the
‘number of the different environments’ (N_ENV), and
the ‘unburnt area’ (UNBURNT), while the second one
was most closely related to the presence or absence of
recent fire (BURNT 1–3 y). The ‘cut area’ (CUT)
correlated only with the fourth axis. The first axis
96
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
Table 9
The dominant species of the unburnt and burnt sites belonging to different classes of fire-regimesa
Dominant species in
unburnt and burnt sites
Unburnt sites
(control)
>30 years
(n ¼ 6)b
Parmena unifasciata
Leiopus nebulosus
Agrilus laticornis
Leptura maculata
Grammoptera ruficornis
Clytus arietis
Tetrops praeusta
Anaesthetis testacea
Exocentrus adspersus
Molorchus umbellatarum
Phymatodes alni
Agrilus angustulus
Alosterna tabacicolor
Deilus fugax
Prionus coriarius
Pyrrhidium sanguineum
Xylotrechus antilope
Pogonocherus hispidulus
Callimus angulatus
Chlorophorus figuratus
Pachytodes cerambyciformis
Stenopterus rufus
No. of dominant species (*)
No. of co-dominant species (*)
Burnt sites
Freshly burnt Single fires
<1 year
1–2 years
(n ¼ 2)b
(n ¼ 2)b
þ
*
*
*
*
*
*
*
*
*
*
*
þ
þ
þ
þ
þ
2
9
*
*
*
*
*
*
*
þ
*
*
*
þ
þ
þ
þ
*
þ
þ
þ
þ
*
Repeated fires
7–14 years
(n ¼ 2)b
22–24 years
(n ¼ 3)b
2–3 years
(n ¼ 2)b
6–17 years
(n ¼ 5)b
*
þ
þ
*
*
*
þ
*
*
*
*
*
þ
*
*
*
*
*
*
*
*
*
þ
*
*
þ
þ
*
þ
*
þ
*
*
*
*
þ
*
*
*
þ
*
þ
þ
*
*
*
þ
*
*
þ
*
þ
*
*
*
*
*
þ
3
6
2
6
2
11
1
11
*
*
*
3
4
þ
*
*
3
5
a
The species follow the dominance gradient of the unburnt sites. The number of years refers to the time elapsed since the last fire. Mean of
the dominance between study sites: (*) >10%; (*) 3.2–9.9%; (þ) 1–3.1%; () <1%; empty cell means that the species was not sampled.
Study site 10 is omitted.
b
Time since last fire; n represents no. of study sites.
and all canonical axes together are significant (Monte
Carlo test, P ¼ 0:005).
4. Discussion
4.1. Do fast spreading winter fires affect the
diversity of xylophagous beetles?
The numbers of cerambycid, buprestid and lucanid
species recorded in this study (40, 14 and 2, respectively) are consistent with numbers found during
previous research in Switzerland using similar methods (e.g. Hartmann and Sprecher, 1990; Barbalat,
1998; Barbalat and Gétaz, 1999).
Our results show that in the case of fast spreading
winter fires (typical fire regime of the whole southern
slope of the Alps) the fire does not directly affect the
species richness of the studied xylophagous groups. In
fact we found that at a small spatial scale (0.25 ha)
species richness was higher in open sites, regardless
of the occurrence of recent fires (1–3 years since the
last fire) and in recently thinned sites.
At a larger spatial scale (6.25 ha) species richness
and number of individuals seem to be positively
affected by the diversity of sites of different ages
(within the last 30 years), the presence of relatively
open habitats, and the presence of intact forest sites
providing refuge for the most shade-tolerant species.
This result could reflect the presence of a higher
+1.0
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
97
13S
QUERCUS
Axis 2 (5.9% variance explained)
Forest stands with
Oak (Quercus sp.)
21C
8C
15S
11R
12R
TIME_LF
23F
14S
2S
22F
1C
16C
18C
3C
19R
17S
BHD_TREE
TREECOV
5R
9S
10S
4S
7R
20R
-0.8
Open forest
stands
6R
DEADWOOD
-0.8
+0.9
Axis 1 (14.9% variance exlained)
Fig. 3. Ordination diagram based on a canonical correspondence analysis of the spider communities at 23 study sites constrained by four
environmental variables (continuous arrows): QUERCUS ¼ Quercus cover; TREECOV ¼ tree cover; DEADWOOD ¼ dead wood cover;
BHD TREE ¼ breast height diameter of dominant trees; geographical coordinates were allocated as co-variables (F-value first axis: 3.33,
P < 0:05; F-value all axes: 2.16, P < 0:05). The dashed arrow (TIME LF ¼ time since last fire) recalls that the tree cover (TREECOV) is
correlated with the time since the last fire. Study sites are represented by symbols ((&) unburnt site; () freshly burnt site; (*) single burnt
site; (*) repeated burnt site). The variables are defined in Tables 3 and 4.
Table 10
Selected environmental variables (manual forward selection; Monte Carlo test, P < 0:05) and correlation values with the first two canonical
axes at both spatial scales: small scale (A) and large scale (B)a
(A) At small spatial scale (0.25 ha)
(B) At large spatial scale (6.25 ha)
Axis 1
Axis 2
Axis 1
Eigenvalues
%_var
%_var (all axes)
0.277
15.1
32.4
0.148
8.0
Environmental variables
Coefficient 1
TREECOV
QUERCUS
DEADWOOD
BHD_TREE
P (axis 1)
P (all axes)
0.806
0.431
0.012
0.137
0.005
0.005
a
Axis 2
Eigenvalues
%_var
%_var (all axes)
0.253
13.8
29.5
Coefficient 2
Environmental variables
Coefficient 1
Coefficient 2
0.130
0.712
0.654
0.108
N_ENV
UNBURNT
BURNT 1–3 y
CUT
P (axis 1)
P (all axes)
0.617
0.594
0.390
0.388
0.010
0.005
0.335
0.075
0.536
0.316
‘Geographical coordinates’ were allocated as co-variables. The variables are defined in Tables 3 and 4.
0.134
7.2
98
Table 11
List of the species and number of individuals diagonalised according to the ‘tree cover’ and ‘time elapsed since the last fire’
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
# Endangered species as explained in Table 8; underline ¼ dominant species (10%) showed in Table 9; N sp ¼ number of species; N ind ¼ number of individuals; % ¼ relative
abundance (dominance); ¼ thinned 4 years before our study; ¼ thinned 10 years before our study; 8 ¼ thinned 9 years before the last fire.
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
number of beetle species but could also be due to a
higher activity of helio- and anthophilous species
coming from the surrounding intact stands. In these
small and highly structured open sites saproxylic
insects can find secondary trophic and breeding conditions as observed by Gutowski (1986, 1995) and
Starzyk and Witkowski (1981) in clear-cut stands of
mature forest in Bialowieza Forest (northeastern
Poland) and in Niepołomice Forest (southern Poland).
Many authors have shown that saproxylic species
require mature forests with either large trees and
dead wood of larger dimensions with a partial canopy
cover, sun-exposed trees and high shrub and herb
cover (e.g. Hartmann and Sprecher, 1990; Hölling,
2000; Niklasson and Drakenberg, 2001; Ranius,
2002), as well as dead wood of smaller dimensions
(e.g. Schiegg, 2001). Our burnt sites are characterised
by a high number of dead young shoots (see Table 2)
as well as by weak or bruised larger shoots (Hofmann
et al., 1998), which are significantly more affected by
the chestnut blight fungus Cryphonectria parasitica
(Murrill) Barr (Prospero et al., 1998) and therefore
more exposed to saproxylic invertebrates.
4.2. How do xylophagous species and
communities respond to fire?
4.2.1. At unburnt sites
Many of the species sampled at unburnt sites were
shade-tolerant, indicating a tendency towards mature
and stable stands. One of the two dominant species,
Parmena unifasciata, is apterous and has therefore
low mobility and low dispersal capacity. Other species
found in unburnt sites are anthophilous and visit
flowers growing in the undergrowth in small sunny
patches in the forests (e.g. Starzyk and Witkowski,
1981; Barbalat, 1998; Bense, 1995).
4.2.2. At freshly burnt sites
During the first year after the fire we noticed a
change in species composition. Parmena unifasciata
and Leiopus nebulosus, which were dominant in
unburnt sites, were no longer found during the first
year after the fire. Parmena unifasciata was found
mainly in control sites and in sites that had not burnt
for at least 6 years. It can be considered as a climax
species which is threatened by fires and which tends
to avoid recently burnt places. The high number of
99
specimens trapped at freshly burnt sites was mainly
due to the abundance of helio- and thermophilous
species. The absence of pyrophilous species indicates
that the species composition depends on the type of
impact that caused an opening, which is also shown in
Fig. 3 and Table 10.
4.2.3. At single fire sites
During the first 1–2 years after a single fire, cerambycid and buprestid species composition still seems
affected by the new post-fire conditions, and it shows
many differences compared with those of intact
stands. The dominant and co-dominant species indicated that the more open and xeric post-fire conditions
could remain until about 7–14 years after the last fire.
Species start recolonising stands after this time.
Nevertheless, species that were dominant in intact
stands were still lacking at this stage, particularly
Parmena unifasciata. The beetle communities at sites
burnt 22–24 years before our study were similar to
those of intact stands. The complete recovery takes
place within 15–20 years.
4.2.4. At repeated fire sites
The species composition of the sites that burned
repeatedly included many anthophilous species. This
is the case of three of the most abundant species of
burnt sites: Leptura maculata, Grammoptera ruficornis and Clytus arietis (Bense, 1995). These are dominant at both singly and repeatedly burnt sites. On the
other hand, the dominant species of intact stands,
Parmena unifasciata and Leiopus nebulosus, remain
sporadic, confirming that they are sensitive to fire.
4.3. Which environmental factors and fire-related
variables influence the xylophagous beetles?
4.3.1. At small spatial scale
In our study, ‘tree cover’ and ‘time elapsed since the
last fire or cut’ influenced the xylophagous beetles at a
small spatial scale. These sites host heliophilous species which require open forest stands or forest clearings for feeding and egg laying. We found a negative
correlation between the time elapsed since the last fire
and flowering plant cover.
Moreover, the species composition appears to be
influenced by the occurrence of oak, which hosts oakdepending species, observed especially in open sites.
100
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
Ranius and Jansson (2000), in a study on saproxylic
beetles in old oaks, observed that the species richness
was higher in large trees and sun-exposed trees.
Barbalat (1998) and Barbalat and Gétaz (1999) found
the highest number of species in sun-exposed oak
stands. In our study area oaks and the beetles they
host seem to be favoured by fires and by forest management. We observed a positive correlation between fire
frequency and the presence of oak. While chestnut
trees are managed as coppice, oaks are kept for timber.
Hence, oaks are usually bigger and have a thicker bark
than chestnut trees. In case of a surface fire, most
chestnut shoots are killed, while oak trees are more
likely to survive (Delarze et al., 1992; Hofmann et al.,
1998). Beetle larvae also have a better chance of
surviving a fire under a thick oak bark.
4.3.2. At large spatial scale
The environmental variables explaining the most of
the variance of the buprestid and cerambycid composition at a larger spatial scale (6.25 ha) confirm the
importance of a mosaic of different habitats including
intact stands.
The coexistence of intact stands, open stands, clearings, gaps and sunny places seems to be very important
in maintaining a diverse and abundant wood-eating
fauna. This mosaic of forest landscapes hosts sciaphilous and disturbance-sensitive species, as well as helioand thermophilous species. Similar faunistic situations
have been described for well-structured wood margins,
wind-throw areas, as well as for artificial clearings and
thinned stands (Okland et al., 1996; Simila et al., 2002;
Sippola et al., 2002). The reason seems to be due to
similarities in vegetation structures and microclimatic
conditions between these habitats (Barbalat, 1998;
Duelli and Obrist, 1999).
5. Conclusions and recommendations
5.1. Disturbance and xylophagous diversity
Most species in European forests evolved under
natural disturbance regimes that included at least
occasional fires (e.g. see Bengtsson et al. (2000)
and White and Jentsch (2001) for a review). The
southern slope of the Alps is a fire-prone region
(Tinner et al., 1999) characterised, until the 1950s,
by an intensive fire and forest management history that
influenced environmental conditions, as well as flora
and fauna (Delarze et al., 1992; Tinner et al., 1999;
Moretti et al., 2002).
Our study shows that fast spreading surface fires
of low to medium intensity in winter do not threaten
cerambycid, buprestid and lucanid beetles. We
observed that fire helps create a complex mosaic of
forest patches with different structures, tree composition and dead wood amount; it may even enhance
biodiversity, as postulated by the mosaic concept
(Forman and Gordon, 1986; Duelli et al., 1997; Simila
et al., 2002; Sippola et al., 2002). At small spatial scale
fire improves dynamics in species composition setting
back the process of competitive exclusion, and thus
conforming to the intermediate disturbance hypothesis
(Connell, 1978; Huston, 1994).
5.2. Disturbance and species conservation
On the one hand, disturbances can create landscape
heterogeneity and so promote biodiversity (Simila
et al., 2002). Conversely, several authors mentioned
the problem of stenoecious obligate saproxylic species
with low dispersal capacity and often depending on
high dead wood connectivity (e.g. Speight, 1989;
Dajoz, 2000; Schiegg, 2000; Ranius, 2002).
In our study area, fire seems to favour endangered
species, especially those which need a mosaic of
closed and open stands, as well as some rare species
depending on oak for their development. Therefore, a
higher proportion of oaks favoured by fires in chestnut
stands and fire residuals seem to be very important
factors in conserving saproxylic species (Gandhi et al.,
2001). Moreover, since the disappearance of intensive
management forms, chestnut stands have progressively closed and been colonised by other broadleaved trees (Conedera and Tinner, 2000). These
stands already host a few species indicating the transition to a fauna requiring more mature and stable
forests.
5.3. Management implications
Sites can be opened both by natural disturbance
(windfalls, broken trees, fires) as well as by silvicultural practices, but it is important that dead wood is
not removed (Speight, 1989). These clearings have a
M. Moretti, S. Barbalat / Forest Ecology and Management 187 (2004) 85–103
positive effect for saproxylics even in mature forests
(Simila et al., 2002), while large-scale cuttings for
industrial purposes are one of the main factors negatively affecting obligate saproxylic species richness
(Okland et al., 1996; Grove, 2002).
Our study showed that fire indirectly influences
xylophagous beetles acting by opening up small areas
of vegetation; in this way its effect is similar to that of
coppicing. Fires also favour oak trees and their associated beetle communities by killing young chestnut
shoots. Oak is very important to xylophagous insects
because many of these insects are specifically linked
to it (Dajoz, 2000). At one time oak covered a large
part of the European lowlands as well as the hilly
regions on the southern slope of the Alps (e.g. Burga,
1998). This tree deserves interest and protection especially because most of its original habitat is now
densely populated in Switzerland.
At the same time, residual stands should be preserved from fire in order to maintain a mosaic as
refuges and dispersal pools (Gandhi et al., 2001). A
forest offering a good structural diversity may be
essential for increasing landscape complexity and so
maintaining a high xylophagous beetle richness.
Acknowledgements
We would like to thank P. Duelli, P.J. Edwards and
M. Conedera for their comments on the manuscript.
Many thanks are due to the various persons who helped
with fieldwork (P. Hördeggen, P. Wirz, F. Fibbioli and
K. Sigrist) and who identified and checked the species
(O. Monga and C. Pesarini).
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