JPE575.

fm Page 49 Monday, March 5, 2001 11:43 AM

Journal of Applied
Fire and biodiversity: responses of grass-layer beetles to
Blackwell Science, Ltd

Ecology 2001
38, 49 – 62 experimental fire regimes in an Australian tropical savanna
JÉRÔME ORGEAS* and ALAN N. ANDERSEN
Tropical Ecosystems Research Centre, CSIRO Wildlife and Ecology, PMB 44 Winnellie, NT 0822 Darwin, Australia

Summary
1. Up to 50% or more of the savanna landscapes of northern Australia are burnt each
year, but the effects of these fires on savanna faunas are poorly known, especially for
arthropods.
2. We investigated the responses of grass-layer beetles to three fire regimes (unburnt – fires
excluded; early – annual fires lit early during the dry season; late – annual fires lit late
during the dry season) as part of a landscape-scale fire experiment at Kapalga in
Australia’s Northern Territory, where replicated fire regimes were applied to 15–20-km2
landscape units over a 5-year period. We also sought to identify beetle species that might
act as indicators of particular fire regimes, and examined the extent to which analysis at
the family level is sufficient to reveal patterns evident from species-level analysis.
3. Beetles were sampled using sweep nets during February (mid-wet season) and May
(end of wet season) each year, from 1989 (18 months prior to the imposition of fire treat-
ments) to 1995 (after 5 years of treatments).
4. A total of 3865 beetles from 233 species was recorded, with the dominant families
Chrysomelidae (leaf beetles) and Curculionidae (weevils) contributing 91% of all
beetles and 57% of total species.
5. Total beetle abundance, species richness, family richness and the abundances of one
of the 10 most common species were significantly affected by fire treatment according to
. In all cases, beetle abundance and richness were similar in the two burning treat-
ments, but greater than in the unburnt treatment. These differences were only apparent
during the second half of the experiment. The responses were mirrored by changes in
composition at species, family and guild levels, according to multivariate analysis.
6. According to indicator analysis, five beetle species were significant indicators of late
fires, and one of early fires. All these species were infrequently recorded and therefore of
limited use for management. However, two common species were indicative of burnt
habitats if early and late treatments are considered together.
7. Family richness was highly correlated (r 2 = 0·615) with species richness, and
multivariate analysis of family-level abundances revealed the compositional changes in
relation to fire that were evident in species-level analysis. Family was therefore a reliable
surrogate of species for detecting beetle responses.
8. Although we found significant effects of our experimental treatments, in the broader
context of overall beetle dynamics the assemblages appeared to be remarkably resilient
to fire. This reflects a long history of association with frequent fire, and is consistent with
many other components of the northern Australian fauna, where fire appears to be of
secondary importance to variation in rainfall and soil type.
9. The lack of effect of fire timing on grass-layer beetles calls into question a management
paradigm that focuses largely on fire intensity, and suggests that fire management needs
to be more mindful of fire frequency.
Key-words: beetle assemblages, bioindicators, fire impacts, landscape-scale fire experiment,
northern Australia.
Journal of Applied Ecology (2001) 38, 49–62

© 2001 British *Present address and correspondence: Institut Méditerranéen d’Ecologie et de Paléoécologie, FST St Jérôme, case 461, 13397
Ecological Society Marseille cedex 20, France (fax 33 4 91 28 80 51; e-mail jerome.orgeas@botmed.u-3mrs.fr).
JPE575.fm Page 50 Monday, March 5, 2001 11:43 AM
50
Introduction
J. Orgeas &
A.N. Andersen Fire plays a key role in ecosystem dynamics in many
biomes throughout the world (Whelan 1995; Bond &
van Wilgen 1996) but particularly in tropical savannas
where frequent fire is an inevitable consequence of
the coupling of grassy vegetation with highly seasonal
rainfall and consistently high temperatures (Gillon
1983a; Andersen 1996). Up to 50% or more of the
extensive savanna landscapes of northern Australia are
burnt each year, with most fires being lit by people
(Russell-Smith, Ryan & Durieu 1997). The dominant
fire management paradigm in the region is one of
extensive burning early in the dry season, when grassy
fuels are still relatively moist, in order to reduce the
extent of higher intensity, unmanaged fires that will
inevitably occur later in the dry season (Russell-Smith
1995).
Despite fire being the dominant tool for ecosystem
management in northern Australia, ecological re-
sponses to different fire regimes are poorly known.
Savanna vegetation is resilient to frequent fire to the
extent that most component plant species regenerate
vigorously following fire (Lacey, Walker & Noble
1982), such that ground-layer vegetation completely
recovers (at least superficially) during the first wet sea-
son following fire. However, little is known about the
effects of fire on savanna faunas, or on key processes
that drive ecosystem dynamics over the longer term
(Andersen 1996). In particular, there is a paucity of
information on the effects of fire on insects and other
arthropods, which contribute most to faunal biodiver-
sity and play key roles in ecosystem dynamics. This
is a poor basis for effective ecosystem management
(Andersen 1999).
Here we describe the responses of grass-layer beetles
to different fire regimes in a tropical savanna of north-
ern Australia. Beetles are by far the most diverse of all
insect groups, with more than 400 000 described spe-
cies world-wide (Hammond 1992). Beetles occupy all
consumer trophic levels and, as herbivores, predators
and prey, play key roles in nutrient cycling and energy
flow. Beetles have also proven to be useful bioindicators
for environmental monitoring and assessment (Stork
1990; Luff 1996; Desender & Bosmans 1998; Petit &
Usher 1998; Rodríguez, Pearson & Barrera 1998), with
their high diversity and sensitivity to environmental
conditions providing a fine-grained view of ecological
change (Erwin 1997). Given their variety of functional
roles, the responses of grass-layer beetles to fire is likely
to reflect the responses of a variety of other grass-layer
invertebrates.
A sound understanding of ecological responses to
different fire regimes requires a rigorous experimental
approach, incorporating adequate replication and the
© 2001 British
collection of pretreatment (baseline) data (Whelan
Ecological Society,
Journal of Applied 1995). Although fire ecology has a strong experimental
Ecology, 38, tradition, the reliability of results is often compromised
49–62 by small plot size. Our study was part of a landscape-
scale fire experiment, where replicated fire regimes were
applied to 15–20-km2 landscape units over a 5-year
period (Andersen et al. 1998). An ordinal-level ana-
lysis of grass-layer arthropods during the wet season
has revealed that fire had only a marginal effect on
overall beetle abundance, reflecting a relatively minor
effect of fire on grass-layer vegetation (Andersen &
Müller 2000). However, ordinal-level abundance does
not necessarily reflect responses at the species level.
Here we describe the species-level responses.
There has been considerable recent interest in sim-
plifying insect surveys by restricting analysis to the
highest taxonomic level that is sufficient to reveal the
response in question, rather than necessarily sorting all
specimens to species (Ellis 1985; Gaston & Williams
1993; Vane-Wright, Smith & Kitching 1994). This
would make insects a far more tractable option in the
land assessment and monitoring process. We therefore
also tested the extent to which responses of beetles
obtained from species-level analyses were matched by
those at the family level.
Our specific aims were to: (i) document the effects of
different fire regimes on the richness and composition
of assemblages of grass-layer beetles; (ii) identify beetle
species that might act as indicators of particular fire
regimes; and (iii) examine the extent to which analysis
at the family level is sufficient to reveal patterns evident
from species-level analysis.
Methods
 
The study was part of the Kapalga fire experiment
located in Kakadu National Park, approximately
200 km east of Darwin, in the seasonal tropics of
Australia’s Northern Territory (Andersen et al. 1998).
Annual rainfall is about 1500 mm, with 90% falling
between November and March. Temperatures are high
throughout the year, with the mean monthly temper-
ature ranging from 25 °C in July to 30 °C in November.
Each experimental landscape unit (subsequently
referred to as compartment) was a 15–20-km2 catch-
ment of a minor seasonal stream. Sampling within
compartments occurred along transects running from
the more poorly drained shallow sands of the stream
margins, dominated by various savanna woodlands,
up-slope to the better drained loams supporting savanna
open forest. These two habitats will subsequently be
referred to as woodland and forest, respectively.
We assessed three fire regimes, each replicated three
times, for their impact on grass-layer beetles: (i) early,
compartments burnt annually early in the dry season
(May/June); (ii) late, compartments burnt annually
towards the end of the dry season (September/October);
and (iii) unburnt, fire excluded from compartments.
Apart from some topographic and security constraints,
fire treatments were assigned randomly to compart-
ments (Andersen et al. 1998). To minimize potential
JPE575.fm Page 51 Monday, March 5, 2001 11:43 AM
51 effects of variation due to differences in fire history
Fire and beetle between compartments, fires were excluded from
biodiversity Kapalga for 18 months (1988–89) prior to the applica-
tion of the fire treatments. Treatments were then
applied for 5 years from 1990 to 1994. Given that the
long-term fire history of Kapalga, as in the rest of the
region, is one of frequent burning, the absence of fire is
not an experimental ‘control’. The question ‘How do
beetles respond to fire exclusion?’ is equally as valid as
‘How do beetles respond to annual fires?’

A woodland and forest plot (each 40 × 20 m) was
sampled in each compartment with 45-cm diameter
sweep-nets. Sweep-netting was conducted during
February (mid-wet season) and May (end of wet
season) each year, from 1989 (prior to the imposition
of fire treatments) to 1995 (after 5 years of treatments).
A sweep sample from each plot comprised the pooled
catches of 100 sweeps, collected along five parallel
transects (5 m spacing) of 20 sweeps (Andersen &
Müller 2000). Beetles were sorted to species, and
identified using keys in Lawrence & Britton (1994),
Matthews (1980, 1982, 1984, 1985, 1987, 1992, 1997)
and Zimmerman (1991, 1992, 1993, 1994a, 1994b).
Our identifications were checked and modified by
appropriate specialists (see the Acknowledgements).

Data were first subject to three-way  to test for
effects of habitat, season and fire on the abundance of
all beetles and common families and species, and on
species and family richness. Sampling did not signi-
ficantly deplete the fauna, and we considered that the
fauna at different sampling times was sufficiently dif-
ferent for samples to be considered independent for
statistical purposes. Samples were divided into three
time periods for these analyses: (i) pre, before the imposi-
tion of burning treatments, from February 1989 to May
1990 (n = 4); (ii) post 1, earlier treatment samples, from
February 1991 to May 1993 (n = 6); and (iii) post 2,
later treatment samples, from February 1994 to May
1995 (n = 4). A fire effect was indicated by the fire–time
period interaction term, which takes into account the
pretreatment baseline data, rather than the overall fire
term, which includes pretreatment data. Abundance and
richness data were log- and square root-transformed,
respectively, to satisfy requirements of normality.
Secondly, multivariate analysis was used to identify
compositional patterns in beetle assemblages in rela-
tion to habitat, season and fire. Data were pooled
across the three replicates of each treatment for
each sampling time × habitat combination, giving 84
© 2001 British
sampling units for analysis (14 sampling times × 3
Ecological Society,
Journal of Applied treatments × 2 habitats). The 84 sampling units were
Ecology, 38, ordinated using semi-strong hybrid multidimensional
49–62 ordination (SSH), based on Bray–Curtis dissimilarity
matrices. SSH is a modification of the hybrid Multi-
dimensional Scaling algorithm that uses a single associ-
ation matrix and Guttman rank imaging (Belbin 1994).
It is considered superior to, and more reliable than,
other multidimensional scaling techniques because it
reduces the weighting of large input distances (Belbin
1994). Ordinations were performed at species, family
and feeding-guild levels. Two guild analyses were
performed, one based on larval diet and the other on
adult diet, using information from Lawrence & Britton
(1994; see the Appendix). Fire-related differences in
ordination space were tested for each of the pre, post 1
and post 2 periods using  (Belbin 1994).
Finally, two analyses were performed in relation to
indicator taxa. First, Indicator Value Analysis (IVA;
Dufrêne & Legendre 1997) was performed on all beetle
species and families to identify taxa that reliably
indicate a particular fire treatment. For each species,
IVA provides a coefficient (IndVal) and a statistical test
describing its indicator power. IVA (499 iterations,
random number generator of 5, significance level set at
0·01) was performed on the 84 sampling units described
above for habitat and season, and for the 60 post-
fire sampling units for fire treatment (i.e. excluding
pretreatment samples). Secondly, total beetle species
richness was correlated with family richness and the
richness of dominant families to test how reliable these
might be as surrogates of total species richness.
The different fire treatments had a negligible impact
on the structure of the grass-layer in the plots
(Andersen & Müller 2000), so that any treatment
effects on grass-layer beetles would not be confounded
by fire-induced changes in sampling efficiency.
Results
 
A total of 3865 beetles from 233 species and 26 families
was collected (see the Appendix). The fauna was dom-
inated by two families, Chrysomelidae (leaf beetles)
and Curculionidae (weevils), which together represented
91% (60% and 31%, respectively) of total abundance
and 57% (39% and 18%, respectively) of total species
(Table 1). In turn, the numerical dominance of these
families was in each case due to three exceptionally
abundant species, with these six species representing
51% of all beetles collected. At the other extreme, 101
(43%) species were represented by a single individual.
Analysis of species accumulation (Fig. 1) indicated
that common species were adequately sampled but,
given that the curve continued to have a shallow slope,
it is likely that numerous rare species were not
recorded.
Abundance was substantially higher in forest (2289
beetles) than woodland (1576), but species richness was
almost identical (154 and 155, respectively). Only 76
(32%) species were recorded in both habitats, but this at
least partly reflects inadequate sampling of uncommon
JPE575.fm Page 52 Monday, March 5, 2001 11:43 AM

52 Table 1. Summary of beetle fauna collected by sweep-net at

J. Orgeas & the Kapalga experiment. Data are number of species
A.N. Andersen
Family Forest Woodland Total

Anobiidae 1 0 1
Anthribidae 2 1 3
Apionidae 4 5 7
Attelabidae 1 0 1
Bostrichidae 0 1 1
Buprestidae 1 0 1
Carabidae 0 2 2
Cerambycidae 3 3 4
Chrysomelidae 71 69 91
Cleridae 1 0 1
Coccinellidae 6 6 10
Curculionidae 28 24 41
Dermestidae 0 1 1
Elateridae 9 8 16
Histeridae 1 0 1
Lycidae 2 1 3
Melyridae 4 4 7
Mordellidae 1 1 2
Phalacridae 2 2 3
Pselaphidae 0 1 1
Rhipiphoridae 5 9 12
Scarabaeidae 5 5 9
Scirtidae 2 4 4
Scraptiidae 0 1 1
Staphylinidae 1 3 3
Tenebrionidae 4 4 7
Total 154 155 233

Fig. 2. Changes in (a) mean abundance, (b) species richness

and (c) family richness over the three time periods. Data for
forest and woodland habitats are combined because of a lack
of treatment–period–habitat interaction.

the post 2 period. In each case there was no significant

Fig. 1. Species accumulation across all 252 samples. The fine
line represents the fitted curve.
species. Both abundance and richness were substan-
tially higher in February (3218 beetles from 185 species)
than May (647 beetles from 114 species).
   
Abundance and richness
Beetle abundance, species richness and family richness
© 2001 British
were all significantly affected by fire treatment (Fig. 2).
Ecological Society,
Journal of Applied Beetle abundance and richness were always greater
Ecology, 38, in the two burning treatments than the unburnt treat-
49–62 ment, but these differences were significant only during
difference between the two burning treatments. Although
the effect of fire was significant, results of 
indicated that fire (maximum F-value of 2·6; P > 0·01)
made a relatively minor contribution to overall vari-
ation in beetle abundance and richness, compared with
the effects of season (F-values ranging from 72·6 to
159·8; P < 0·0005) and habitat (11·8–52·7; P < 0·001).
In most cases the abundance of the most common
beetle species and families varied significantly between
habitats and seasons, but only one of the 10 most com-
mon species and one of the five most common families
were significantly affected by fire (Table 2). Changes in
abundance in relation to fire varied markedly between
the dominant beetle families. Curculionid abundance
increased markedly under both fire treatments, but
declined in the absence of fire, and this was evident even
in the post 1 period; in contrast, chysomelid abundance
was not significantly affected by fire at all (Fig. 3).
However, the curculionid trend was in large part driven
by a single, highly abundant, species.
JPE575.fm Page 53 Monday, March 5, 2001 11:43 AM

53 Table 2. P-values from  of abundance of the 10 most common beetle species (data square root-transformed) and five most
Fire and beetle common families (log-transformed). The significance level has been set at 0·01 to accommodate multiple comparisons
biodiversity
Fire
Season Habitat (treatment × period)

Species
Essolithna sp. 5 (Curculionidae: Entimini) nr. E. mediofusca < 0·0005 < 0·0005 < 0·0005
Scelodonta sp. 8 (Chrysomelidae: Eumolpinae) 0·111 0·002 0·804
Eboo sp. 9 (Chrysomelidae: Eumolpinae) 0·008 < 0·0005 0·323
Ditropidus sp. 11 (Chrysomelidae: Cryptocephalinae) 0·101 0·006 0·024
Hispellinus sp. 17 (Chrysomelidae: Hispinae) 0·065 0·015 0·116
Rhyparida sp. 18 (Chrysomelidae: Eumolpinae) < 0·0005 < 0·0005 0·647
Rhyparida sp. 26 (Chrysomelidae: Eumolpinae) < 0·0005 0·860 0·733
Ditropidus sp. 34 (Chrysomelidae: Cryptocephalinae) < 0·0005 0·004 0·063
Rhyparida sp. 57 (Chrysomelidae: Eumolpinae) < 0·0005 0·317 0·800
Unidentified sp. 58 (Chrysomelidae: Galerucinae) < 0·0005 0·183 0·548
Families
Curculionidae < 0·0005 0·006 0·001
Elateridae < 0·0005 0·014 0·034
Chrysomelidae < 0·0005 < 0·0005 0·548
Cerambycidae < 0·0005 0·060 0·510
Phalacridae 0·008 < 0·0005 0·327

Fig. 3. Changes in mean abundance of (a) Chrysomelidae

and (b) Curculionidae over the three time periods.

Assemblage composition
Fig. 4. SSH ordination (stress = 0·13) based on family-level
Overall beetle composition showed considerable over- abundances. Replicates were combined in 84 sample units and
lap between habitats and seasons, but there was greater plotted according to (a) seasons and (b) habitats. Similar
variability in woodland than forest, and in May com- patterns occurred in SSH ordinations based on species.
pared with February (Fig. 4). This was consistent with
the greater structural diversity of woodland compared
with forest habitats, and the greater variability of
© 2001 British
rainfall in May (end of wet season) compared with no apparent differences in clusters according to fire
Ecological Society,
Journal of Applied February (middle of wet season). Patterns of assem- treatment during the pre period, but the two burning
Ecology, 38, blage composition in relation to fire were similar at treatments had diverged from the unburnt treatment
49–62 species, family and guild levels. In each case, there were by post 2 (Fig. 5), as demonstrated by  (Table 3).
JPE575.fm Page 54 Monday, March 5, 2001 11:43 AM

54 Table 3. P-values from  analyses of SSH clusters,

J. Orgeas & comparing beetle assemblage composition at burnt (early and
late combined) and unburnt sites. Beetle composition was
A.N. Andersen
considered at the level of species (data untransformed, stress
= 0·23), family (untransformed, 0·13) and feeding guilds
(larvae, log-transformed, 0·13; adults, log-transformed, 0·14).
Significant P-values indicate that clusters are significantly
different. In each case, clusters were very similar during pre
and post 1 periods but significantly (or nearly so) different
during the post 2 period (bold text), indicating that repeated
burning significantly influenced beetle composition

Pre Post 1 Post 2

Species 0·404 0·268 0·005

Families 0·877 0·752 0·043
Feeding-guild (larvae) 0·808 0·901 0·008
Feeding-guild (adults) 0·852 0·918 0·053

units (three replicates combined), which seriously

Fig. 5. SSH ordination (stress = 0·13) based on abundances
of nine larval feeding groups, with sample units plotted
separately for the (a) pre burnt and (b) post 2 periods. Similar
patterns occurred in SSH ordinations based on species,
families and adult-feeding guilds (Table 3).
 
Six species were significant indicators of a particular
burning treatment according to indicator analysis,
with five of the six species being indicative of the late
treatment (Table 4). All the indicator species were
infrequently recorded, occurring in < 10% of sample
limited their utility. Pooling early and late treatments as
a single ‘burnt’ treatment for indicator analysis gave
four species as indicators of burned habitats, with two
of the four being frequently recorded (46% and 69% of
sample units, respectively). Additionally, one family
(Curculionidae) was identified as an indicator taxon.
Species and family richness were highly correlated
(r 2 = 0·615, n = 214, P < 0·0005; samples without
beetles excluded from analysis), such that family
richness was a reasonable surrogate of species richness.
Species richness of Chrysomelidae (representing 39%
of total beetle species) was even more highly cor-
related with total species richness (r 2 = 0·839, n = 210,
P < 0·0005), such that chrysomelid richness would
be a more reliable surrogate of species richness.
Curculionid richness (18% of total beetle species) was
not so strongly correlated with total species richness
(r2 = 0·308, n = 142, P < 0·0005).

Table 4. Results of IVA (after Dufrêne & Legendre 1997), computing IndVal coefficient and t-test, and showing species that
indicate a particular fire treatment. In (a), the three treatments are considered separately, whereas in (b) early and late have been
combined to give a single burning treatment. Total abundances/frequency of occurrence (n = 84) in each fire treatment are given
for each species

IndVal index Treatment Unburnt Early Late t

(a)
Chrysomelidae Agetinus sp. 66 24·07 Late 9/3 0/0 106/5 3·154
Melyridae Dicranolaius sp. 3 20·18 Late 2/2 1/1 8/5 3·623
Curculionidae Entimini Myllocerus sp. 256 20 Late 0/0 0/0 17/4 4·457
Curculionidae Entimini unidentified sp. 62a 19·86 Late 4/2 0/0 250/4 2·834
Curculionidae Ceutorhynchini Indohypurus sp. 251 21·71 Late 3/3 0/0 9/5 3·68
Phalacridae Phalacrus sp. 95 26·05 Early 2/2 6/6 0/0 4·933

IndVal index Treatment Unburnt Burnt t

(b)
© 2001 British Chrysomelidae Hispinae Hispellinus sp. 17 45·25 Burnt 23/15 48/24 0·899
Ecological Society, Curculionidae Entimini Essolithna sp. 5 nr. E. mediofusca 69·67 Burnt 131/23 336/35 5·903
Journal of Applied Chrysomelidae Eumolpinae Geloptera sp. 112 22·32 Burnt 2/2 27/9 3·708
Ecology, 38, Chrysomelidae Eumolpinae Rhyparida sp. 114 19·34 Burnt 3/1 28/8 3·361
49–62
JPE575.fm Page 55 Monday, March 5, 2001 11:43 AM
55
Discussion
Fire and beetle
biodiversity
 
The grass-layer at Kapalga supports a diverse beetle
assemblage, with numerous uncommon species likely
to occur in addition to the 233 species recorded.
Unfortunately no comparable data are available from
elsewhere in northern Australia; indeed we are unaware
of any inventory of grass-layer beetle assemblages from
anywhere that can be used for comparative purposes.
However, the total number of species recorded is
similar to the 200 ground-foraging species collected in
pitfall traps during the Kapalga experiment (Blanche,
Andersen & Ludwig 2000). Remarkably, only four
(< 1% total species) species occurred in both grass- and
ground-layer faunas. The dominant families in the grass-
layer were Chrysomelidae and Curculionidae, which is
typical of Australian savannas (J. Orgeas, unpublished
data) and other grassy ecosystems (Cooter 1991;
Siemann, Haarstad & Tilman 1997). The higher
abundance and richness of beetles during the middle of
the wet season (February) compared with end of wet
season (May) can be explained by the erratic nature of
rain during the wet–dry transition (Taylor & Tulloch
1985) and is consistent with general seasonality
patterns of insect activity in the seasonal tropics
(Wolda 1988).
   
Beetle abundance and richness were promoted by fire,
with the two fire treatments having similar effects.
Likewise, species composition was affected by fire, but
not by season of burn. The compositional changes
were also reflected at the family and feeding-guild
levels. These changes were evident only after repeated
application of fire treatments, which underscores the
importance of considering cumulative effects of fire
regimes, rather than individual effects of any particular
fire. We know nothing of the immediate effects of fire
on grass-layer beetles at Kapalga, but other studies of
grass-layer insects demonstrate that a high proportion
of volant taxa escape the flames and rapidly recolonize
(Gillon 1972; Gillon 1983a). Nevertheless, fire is likely
to cause substantial direct mortality of beetles (Gillon
1983a; Siemann, Haarstad & Tilman 1997), such that
the longer term increases in beetle abundance and
richness in burnt compartments must reflect superior
ecological conditions for beetles following repeated
fire.
The specific ecological conditions favouring beetles
in repeatedly burnt habitats are open to conjecture, but
there are several possibilities. First, predation pressure
might be reduced. For example, spiders are likely to be
© 2001 British
major predators of beetles (Wise 1993) and are likely
Ecological Society,
Journal of Applied to suffer greater mortality during fires because they
Ecology, 38, cannot fly. Alternatively, burning might enhance
49–62 habitat structure or food quality for beetles. In particular,
the dominant beetle families (Chrysomelidae and
Curculionidae) are phytophagous (Lawrence & Britton
1994; Zimmerman 1994a), and fire commonly pro-
motes the nutritional status of plant regrowth (Bond &
van Wilgen 1996). Fire had no demonstrable effect on
either plant species richness or floristic composition
at the study sites, and the cover of only one of the
four dominant grass species was affected (Andersen
& Müller 2000), such that changes in the structure
and composition of grass-layer vegetation appeared
minimal. The effect of fire on nutritional quality of
vegetation at Kapalga is unknown.
Grass-layer beetle assemblages at Kapalga were
affected by repeated fires, but not by season of burn.
In other words, the higher intensity typical of late
compared with early fires had no significant effect on
grass-layer beetles. This contrasts with the situation
for ground-foraging beetles at Kapalga, where assem-
blages repeatedly burnt by early fires were indistin-
guishable from those not burnt, but late fires significantly
reduced beetle abundance and richness during the
early wet season (Blanche, Andersen & Ludwig 2000).
Season of fire also had a significant effect on other
ground-active taxa at Kapalga (Andersen & Müller
2000) but not on grasshoppers (A.N. Andersen &
L.M. Low, unpublished data). This suggests the
existence of distinct grass- and ground-layer syndromes
in terms of resilience to fire.
 
We examined the extent to which individual beetle taxa
might be indicative of particular fire regimes. It needs
to be stressed that the indicators we have identified
in this context are not necessarily indicative of other
ecological responses to these treatments: they are environ-
mental rather than ecological indicators (McGeoch
1998). Indicator analysis identified five species that
were indicative of the late treatment and one indicative
of the early one, but they were all infrequent and
therefore of limited use for management. By pooling
early and late treatments we were able to identify two
common species that were indicative of burnt habitats.
Interestingly, no species was indicative of the unburnt
treatment, suggesting an absence of species character-
istic of unburnt habitats. However, this would almost
certainly change in the continued absence of fire if
the understorey vegetation became increasingly
shrubby, as often occurs after long-term fire exclusion
(Andersen 1996).
We also examined the extent to which analyses at the
family level could reveal responses detected by species-
level analysis. Family richness was highly correlated
with species richness, such that the response of species
richness to fire was mirrored by family richness. The
large proportion (10 out of 26, 38%) of families that
were represented by a single species obviously contrib-
uted to the high correlation between family and species
richness, but this was counterbalanced by the fact that
JPE575.fm Page 56 Monday, March 5, 2001 11:43 AM
56 the two richest families contributed more than half of
J. Orgeas & the total species. Similarly, multivariate analysis of
A.N. Andersen family-level abundances revealed the compositional
changes in relation to fire that were evident in species-
level analysis. Family was therefore a reliable surrogate
of species for detecting the responses of grass-layer
beetles to experimental fire regimes.
   
Although we found significant effects of experimental
fire regimes on grass-layer beetle assemblages, in the
broader context of overall beetle dynamics the assem-
blages appear to be remarkably resilient to fire. The
results need to be viewed in the context that experi-
mental treatments at Kapalga represented the most
extreme fire regimes possible, with complete fire exclu-
sion on one hand and annual fires under the most
extreme fire conditions possible on the other. Fire
exclusion only began affecting grass-layer beetle
assemblages after 3 years, and fire season/intensity had
no significant effect. Only one of the 10 most common
beetle species was significantly affected by any of the
experimental treatments. Moreover,  revealed
that variation in beetle abundance and richness in rela-
tion to fire was small in comparison with variation due
to habitat and, in particular, season. A relatively sub-
dued effect of fire in the context of overall variation
appears to be common for arthropods (Friend & Wil-
liams 1996).
The overall resilience of grass-layer beetles to fire is
consistent with the apparent resilience of grass-layer
arthropods in general at Kapalga, with most ordinal-
level taxa remaining unaffected by experimental fire
regimes (Andersen & Müller 2000). Such resilience
would appear to reflect a long history of association of
grass-layer arthropods with frequent fire in northern
Australia. On the other hand, grass-layer arthropods
appear to be far less resilient to fire at Lamto in west
Africa, where savannas have been relatively recently
derived from forest due to extensive clearing and burn-
ing (Gillon 1983b).
In conclusion, fire at Kapalga appears to be of
secondary importance to grass-layer arthropods,
serving to modify patterns caused primarily by rainfall
and soil type (Walker 1987). This is consistent with the
dynamics of many other components of the northern
Australian fauna, whose populations can fluctuate
enormously from year to year due to variation in the
timing, duration and intensity of rainfall (Ridpath
1985; Taylor & Tulloch 1985). Our results indicate that
grass-layer beetles at Kapalga are not influenced by
when fire occurs during the dry season, but rather
respond to whether or not fire occurs at all. This calls
into question a management paradigm that focuses
© 2001 British
largely on fire intensity, and suggests that fire manage-
Ecological Society,
Journal of Applied ment needs to be more mindful of fire frequency. If
Ecology, 38, the extensive use of prescriptive fires early in the dry
49–62 season results in higher fire frequency, then it might not
be the most effective strategy for the conservation of
biodiversity. These findings highlight the importance
of large-scale replicated experiments for effective
ecosystem management (Carpenter 1996; Ormerod,
Pienkowski & Watkinson 1999).
Acknowledgements
We are most grateful to Lyn Lowe for conducting most
of the field sampling, to Tony Hertog for laboratory
assistance, and to John Ludwig and Tony Griffiths
for advice on data analysis. We also thank Elwood
Zimmerman, Rolf Oberpreiler, Andrew Calder, Tom
Weir and Chris Reid for their assistance with beetle
identifications, and Michael Samways, Jonathan Majer
and Nigel Stork for their comments on the draft
manuscript.
References
Andersen, A.N. (1996) Fire ecology and management.
Landscape and Vegetation Ecology of the Kakadu Region,
Northern Australia (eds C.M. Finlayson & I. Von Oertzen),
pp. 179–196. Kluwer Academic, Holland, the Netherlands.
Andersen, A.N. (1999) My bioindicator or yours? Making the
selection. Journal of Insect Conservation, 3, 1–4.
Andersen, A.N. & Müller, W.J. (2000) Arthropod response to
experimental fire regimes in an Australian tropical savanna:
ordinal-level analysis. Austral Ecology, 25, 199–209.
Andersen, A.N., Braithwaite, R.W., Cook, G.D., Corbett, L.K.,
Williams, R.J., Douglas, M.M., Gill, M.A., Setterfield, S.A.
& Muller, W.J. (1998) Fire research for conservation
management in tropical savannas: introducing the Kapalga
fire experiment. Australian Journal of Ecology, 23, 95–110.
Belbin, L. (1994) Pattern Analysis Package, Technical Reference.
CSIRO, Canberra, Australia.
Blanche, K.R., Andersen, A.N. & Ludwig, J.A. (2000)
Detection of ground-active beetle responses to experimental
fire regimes in an Australian tropical savanna. Ecological
Applications (in press).
Bond, W.J. & van Wilgen, B.W. (1996) Fire and Plants.
Chapman & Hall, London, UK.
Carpenter, S.R. (1996) Microcosm experiments have limited
relevance for community and ecosystem ecology. Ecology,
77, 677– 680.
Cooter, J. (1991) A Coleopterist’s Handbook. The Amateur
Entomologists’ Society, Portsmouth, UK.
Desender, K. & Bosmans, R. (1998) Ground beetles (Cole-
optera, Carabidae) on set-aside fields in the Campine region
and their importance for nature conservation in Flanders
(Belgium). Biodiversity and Conservation, 7, 1485–1493.
Dufrêne, M. & Legendre, P. (1997) Species assemblages
and indicator species: the need for a flexible asymmetrical
approach. Ecological Monographs, 67, 345–366.
Ellis, D. (1985) Taxonomic sufficiency in pollution assessment.
Marine Pollution Bulletin, 16, 459.
Erwin, T.L. (1997) Biodiversity at its utmost: tropical forest
beetles. Biodiversity II. Understanding and Protecting Our
Biological Resources (eds M.L. Reaka-Kudla, D.E. Wilson
& E.O. Wilson), pp. 27–40. Joseph Henry Press, Washington,
DC.
Friend, G.R. & Williams, M.R. (1996) Impact of fire on inver-
tebrate communities in mallee-heath shrublands of south-
western Australia. Pacific Conservation Biology, 2, 244–267.
Gaston, K.J. & Williams, P.H. (1993) Mapping the world’s
species – the higher taxon approach. Biodiversity Letters, 1,
2 – 8.
JPE575.fm Page 57 Monday, March 5, 2001 11:43 AM
57 Gillon, Y. (1972) The effect of bush fire on the principal
Fire and beetle Acridid species of an Ivory Coast savanna. Proceedings of
the Tall Timbers Fire Ecology Conference, 11, 419 – 471.
biodiversity
Gillon, D. (1983a) The fire problem in tropical savannas. Eco-
systems of the World 13: Tropical Savannas (ed. F. Bourlière),
pp. 617– 641. Elsevier, Amsterdam, the Netherlands.
Gillon, Y. (1983b) The invertebrates of the grass-layer. Eco-
systems of the World 13: Tropical Savannas (ed. F. Bourlière),
pp. 289 – 311. Elsevier, Amsterdam, the Netherlands.
Hammond, P. (1992) Species inventory. Global Biodiversity:
Status of the Earth’s Living Resources (ed. B. Groombridge),
pp. 17– 39. Chapman & Hall, London, UK.
Lacey, C.I., Walker, J. & Noble, I.R. (1982) Fire in Australian
tropical savannas. Ecology of Tropical Savannas (eds
B.J. Huntley & B.H. Walker), pp. 246 –272. Springer-
Verlag, Berlin, Germany.
Lawrence, J.F. & Britton, E.B. (1994) Australian Beetles.
Melbourne University Press, Melbourne, Australia.
Luff, M.L. (1996) Use of carabids as environmental indicators
in grasslands and cereals. Annales Zoologici Fennici, 33,
185 –195.
McGeoch, M.A. (1998) The selection, testing and application
of terrestrial insects as bioindicators. Biological Review, 73,
181–201.
Matthews, E.G. (1980) A Guide to the Genera of Beetles of
South Australia Part 1. South Australian Museum,
Adelaide, Australia.
Matthews, E.G. (1982) A Guide to the Genera of Beetles of
South Australia Part 2. South Australian Museum,
Adelaide, Australia.
Matthews, E.G. (1984) A Guide to the Genera of Beetles of
South Australia Part 3. South Australian Museum,
Adelaide, Australia.
Matthews, E.G. (1985) A Guide to the Genera of Beetles of
South Australia Part 4. South Australian Museum,
Adelaide, Australia.
Matthews, E.G. (1987) A Guide to the Genera of Beetles of
South Australia Part 5. South Australian Museum,
Adelaide, Australia.
Matthews, E.G. (1992) A Guide to the Genera of Beetles of
South Australia Part 6. South Australian Museum,
Adelaide, Australia.
Matthews, E.G. (1997) A Guide to the Genera of Beetles of
South Australia Part 7. South Australian Museum,
Adelaide, Australia.
Ormerod, S.J., Pienkowski, M.W. & Watkinson, A.R. (1999)
Communicating the value of ecology. Journal of Applied
Ecology, 36, 847– 855.
Petit, S. & Usher, M.B. (1998) Biodiversity in agricultural
landscapes: the ground beetle communities of woody
uncultivated habitats. Biodiversity and Conservation, 7,
1549 –1561.
© 2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
49–62
Ridpath, M.G. (1985) Ecology in the wet–dry tropics: how
different? Proceedings of the Ecological Society of Australia,
13, 3 – 20.
Rodríguez, J.P., Pearson, D. & Barrera, R.R. (1998) A test
for the adequacy of bioindicator taxa: are tiger beetles
(Coleoptera: Cicindellidae) appropriate indicators for
monitoring the degradation of tropical forests in Venezuela?
Biological Conservation, 83, 69–76.
Russell-Smith, J. (1995) Fire management. Kakadu: Natural
and Cultural Heritage and Management (eds T. Press,
D. Lea, A. Webb & A. Graham), pp. 217–237. Australian
National University Press, Canberra, Australia.
Russell-Smith, J., Ryan, P.G. & Durieu, R. (1997) A LAND-
SATT MSS-derived fire history of Kakadu National
Park, monsoonal northern Australia. 1980–94: seasonal
extent, frequency and patchiness. Journal of Applied Eco-
logy, 34, 748 –766.
Siemann, E., Haarstad, J. & Tilman, D. (1997) Short-term and
long-term effects of burning on oak savanna arthropods.
American Midland Naturalist, 137, 349–361.
Stork, N.E. (1990) The Role of Ground Beetles in Ecological
and Environmental Studies. Intercept Ltd, Andover, the
Netherlands.
Taylor, J.A. & Tulloch, D. (1985) Rainfall in the wet–dry
tropics: extreme events at Darwin and similarities between
years during the period 1870 –1983 inclusive. Australian
Journal of Ecology, 10, 281–295.
Vane-Wright, R.I., Smith, C.R. & Kitching, I.J. (1994)
Systematic assessment of taxic diversity by summation.
Systematics and Conservation Evaluation (eds P.L. Forey,
C.J. Humphries & R.I. Vane-Wright), pp. 309 – 326.
Clarendon Press, Oxford, UK.
Walker, B.H. (1987) Determinants of Tropical Savannas.
IRL Press, Oxford, UK.
Whelan, R.J. (1995) The Ecology of Fire. Cambridge University
Press, Cambridge, UK.
Wise, D.H. (1993) Spiders in Ecological Webs. Cambridge
University Press, Cambridge, UK.
Wolda, H. (1988) Insect seasonality: why? Annual Review of
Ecology and Systematics, 19, 1–18.
Zimmerman, E.C. (1994a) Australian Weevils I. CSIRO,
Melbourne, Australia.
Zimmerman, E.C. (1994b) Australian Weevils II. CSIRO,
Melbourne, Australia.
Zimmerman, E.C. (1991) Australian Weevils V. CSIRO,
Melbourne, Australia.
Zimmerman, E.C. (1992) Australian Weevils VI. CSIRO,
Melbourne, Australia.
Zimmerman, E.C. (1993) Australian Weevils III. CSIRO,
Melbourne, Australia.
Received 9 March 2000; revision received 11 May 2000
JPE575.fm Page 58 Monday, March 5, 2001 11:43 AM

58 Appendix
J. Orgeas &
A.N. Andersen List of beetle species collected during the experiment. Total abundance and relative abundance by habitat and season are given
for each species. The species number in each family and subfamily is given in parentheses.

Guilds*
Family/subfamily Total
(number of species) Taxa abundance % forest % February Adults Larvae

ANOBIIDAE (1)
Lasioderma sp. 116 1 100 100 W W
ANTHRIBIDAE (3)
Anthribinae (2) Exillis sp. 323 2 100 100 F F
Atropideres sp. 324 2 0 0 F F
Choraginae (1) Genus H sp. 210 1 100 100 F F
APIONIDAE (7)
Apioninae (7) Apion sp. 44 6 50 17 Ph Ph
Apion sp. 134 4 50 0 Ph Ph
Apion sp. 205 2 0 50 Ph Ph
Apion sp. 258 1 100 100 Ph Ph
Apion sp. 259 1 0 0 Ph Ph
Apion sp. 260 1 100 0 Ph Ph
Apion sp. 261 1 0 100 Ph Ph
ATTELABIDAE (1)
Attelabinae (1) Euops sp. 267 1 100 100 Ph Ph
BOSTRICHIDAE (1)
Bostrichinae (1) Bostrychopsis sp. 276 1 0 100 W W
BUPRESTIDAE (1)
Agrilinae (1) sp. 101 2 100 100 Pol Roo
CARABIDAE (2)
Cicindelinae (2) Cicindela leai (sp. 329) 4 0 100 Pre Pre
Cicindela eparsimpilosa (sp. 330) 1 0 100 Pre Pre
CERAMBYCIDAE (4)
Lychrosis sp. 45 10 80 100 Ph W
Lamiinae (3) Hathliodes sp. 56 43 2 100 Ph W
Hathliodes sp. 277 1 100 100 Ph W
Unknown sp. 340 3 0 100 Ph W
CHRYSOMELIDAE (91)
Bruchinae (3) Callosobruchus sp. 13 36 81 47 See See
Callosobruchus sp. 14 4 100 50 See See
Callosobruchus sp. 97 38 63 53 See See
Chrysomelinae (5) Chalcomela sp. 82 1 100 0 Ph Ph
Phyllocharis hieroglyphica (sp. 88) 1 0 0 Ph Ph
Phyllocharis sp. 302 2 0 100 Ph Ph
Phyllocharis cyanipes (sp. 303) 8 100 88 Ph Ph
sp. 73 1 0 100 Ph Ph
sp. 80 8 50 75 Ph Ph
sp. 304 1 0 100 Ph Ph
Cryptocephalinae (9) Aporocera sp. 152 6 50 100 Ph Sv
Ditropidus sp. 11 71 89 72 Ph Sv
Ditropidus sp. 34 50 84 92 Ph Sv
sp. 115 2 100 100 Ph Sv
sp. 193 4 50 100 Ph Sv
sp. 207 1 100 100 Ph Sv
sp. 211 15 60 87 Ph Sv
sp. 311 4 75 100 Ph Sv
sp. 343 1 0 100 Ph Sv
Eumolpinae (39) Agetinus sp. 64 77 45 91 Ph Roo
Agetinus sp. 66 115 0 100 Ph Roo
Allitus sp. 198 13 62 85 Ph Roo
Deretrichia sp. 29 35 89 91 Ph Roo
Eboo sp. 9 518 91 66 Ph Roo
Eboo sp. 133 45 80 87 Ph Roo
© 2001 British Eboo sp. 208 7 57 100 Ph Roo
Ecological Society, Geloptera sp. 112 29 79 100 Ph Roo
Journal of Applied Pagna sp. 87 24 75 33 Ph Roo
Ecology, 38,
49–62
JPE575.fm Page 59 Monday, March 5, 2001 11:43 AM

59 Appendix continued
Fire and beetle
Guilds*
biodiversity
Family/subfamily Total
(number of species) Taxa abundance % forest % February Adults Larvae

Rhyparida sp. 0 2 50 100 Ph Roo

Rhyparida sp. 4 12 92 8 Ph Roo
Rhyparida sp. 18 205 93 98 Ph Roo
Rhyparida sp. 19 40 80 98 Ph Roo
Rhyparida sp. 23 39 72 100 Ph Roo
Rhyparida sp. 26 79 47 97 Ph Roo
Rhyparida sp. 30 30 57 97 Ph Roo
Rhyparida sp. 39 36 83 94 Ph Roo
Rhyparida sp. 40 2 50 50 Ph Roo
Rhyparida sp. 47 11 36 91 Ph Roo
Rhyparida sp. 52 41 71 100 Ph Roo
Rhyparida sp. 57 94 57 95 Ph Roo
Rhyparida sp. 68 2 50 100 Ph Roo
Rhyparida sp. 69 4 25 100 Ph Roo
Rhyparida sp. 71 2 100 100 Ph Roo
Rhyparida sp. 74 26 23 100 Ph Roo
Rhyparida sp. 114 31 65 100 Ph Roo
Rhyparida sp. 156 15 87 73 Ph Roo
Rhyparida sp. 160 7 43 100 Ph Roo
Rhyparida sp. 171 11 100 91 Ph Roo
Rhyparida sp. 182 4 0 100 Ph Roo
Rhyparida sp. 309 2 0 100 Ph Roo
Rhyparida sp. 310 1 0 0 Ph Roo
Rhyparida sp. 313 1 100 100 Ph Roo
Rhyparida sp. 314 3 100 100 Ph Roo
Rhyparida sp. 349 3 67 33 Ph Roo
Scelodonta sp. 8 48 88 38 Ph Roo
sp. 130 3 100 33 Ph Roo
sp. 199 1 0 100 Ph Roo
sp. 315 2 50 50 Ph Roo
sp. 347 2 50 50 Ph Roo
Galerucinae (27) Monolepta sp. 51 12 25 100 Ph Ph
Monolepta sp. 58 236 64 97 Ph Ph
Oides sp. 78 13 77 85 Ph Ph
sp. 31 5 40 60 Ph Ph
sp. 32 14 14 93 Ph Ph
sp. 36 2 100 100 Ph Ph
sp. 46 1 100 100 Ph Ph
sp. 79 20 25 20 Ph Ph
sp. 92 1 0 0 Ph Ph
sp. 148 2 0 100 Ph Ph
sp. 155 4 100 100 Ph Ph
sp. 162 30 100 97 Ph Ph
sp. 185 3 0 100 Ph Ph
sp. 197 1 0 100 Ph Ph
sp. 209 1 0 100 Ph Ph
sp. 316 1 100 100 Ph Ph
sp. 317 3 33 67 Ph Ph
sp. 318 2 100 50 Ph Ph
sp. 319 8 75 88 Ph Ph
sp. 320 6 0 67 Ph Ph
sp. 322 10 30 70 Ph Ph
sp. 325 1 100 100 Ph Ph
sp. 326 5 100 100 Ph Ph
sp. 327 1 0 0 Ph Ph
sp. 328 1 100 0 Ph Ph
Hispinae (7) Eurispa sp. 119 1 0 0 Ph Ph
© 2001 British Eurispa sp. 285 1 0 0 Ph Ph
Ecological Society, Hispellinus sp. 17 71 68 63 Ph Ph
Journal of Applied sp. 28 1 100 100 Ph Ph
Ecology, 38,
49–62
JPE575.fm Page 60 Monday, March 5, 2001 11:43 AM

60 Appendix continued
J. Orgeas &
Guilds*
A.N. Andersen
Family/subfamily Total
(number of species) Taxa abundance % forest % February Adults Larvae

sp. 278 1 0 0 Ph Ph
sp. 279 1 100 100 Ph Ph
sp. 286 1 100 0 Ph Ph

CLERIDAE (1)
Phyllobaeninae (1) Lemidia sp. 204 2 100 100 Pre Pre
COCCINELLIDAE (10)
sp. 174 3 100 67 Pre Pre
Coccinellinae (2) Cleobora sp. 85 1 0 0 Pre Pre
Harmonia sp. 331 1 100 100 Pre Pre
Coccidulinae (5) Rodolia sp. 60 3 33 67 Pre Pre
Rodolia sp. 99 2 50 50 Pre Pre
Rodolia sp. 287 1 0 0 Pre Pre
Rodolia sp. 344 1 100 0 Pre Pre
Rodolia sp. 345 1 0 100 Pre Pre
Chilocorinae (2) Telsimia sp. 164 1 0 100 Pre Pre
Telsimia sp. 281 1 100 0 Pre Pre
CURCULIONIDAE (41)
Adelognatha (Entimini) (26) Essolithna sp. 5 467 80 79 Ph Ph
Myllocerus sp. 76 10 30 70 Ph Ph
Myllocerus sp. 123 8 100 100 Ph Ph
Myllocerus sp. 136 2 50 100 Ph Ph
Myllocerus sp. 151 21 0 95 Ph Ph
Myllocerus sp. 172 10 100 90 Ph Ph
Myllocerus carinatus (sp. 256) 17 0 100 Ph Ph
Myllocerus sp. 274 1 0 0 Ph Ph
sp. 7 9 78 89 Ph Ph
sp. 27 2 100 100 Ph Ph
sp. 42 2 50 100 Ph Ph
sp. 48 1 100 100 Ph Ph
sp. 61 15 100 7 Ph Ph
sp. 62 290 1 96 Ph Ph
sp. 62a 254 1 99 Ph Ph
sp. 77 2 100 50 Ph Ph
sp. 91 1 100 0 Ph Ph
sp. 153 1 100 100 Ph Ph
sp. 176 1 100 100 Ph Ph
sp. 262 1 0 0 Ph Ph
sp. 269 3 0 100 Ph Ph
sp. 270 1 0 100 Ph Ph
sp. 271 1 0 100 Ph Ph
sp. 272 1 100 100 Ph Ph
sp. 273 1 100 100 Ph Ph
sp. 348 1 0 100 Ph Ph
Baridinae (2) sp. 265 1 100 0 Ph Ph
sp. 266 1 100 100 Ph Ph
Ceutorhynchinae (1) ? Indohypurus sp. 251 12 100 58 Ph Ph
Cryptorhynchinae (3) sp. 35 8 88 100 Ph Ph
sp. 35b 23 52 100 Ph Ph
sp. 255 1 0 0 Ph Ph
Curculioninae (6) sp. 22 13 69 23 Ph Ph
sp. 252 1 100 0 Ph Ph
sp. 253 1 100 0 Ph Ph
sp. 254 1 0 0 Ph Ph
sp. 263 1 0 0 Ph Ph
sp. 264 1 0 0 Ph Ph
Gonipterinae (1) Oxyops sp. 257 3 33 100 Ph Ph
© 2001 British Indet. Phanerognatha (1) sp. 86 1 0 0 Ph Ph
Ecological Society, Molytinae (1) sp. 250 1 100 0 Ph Ph
Journal of Applied
Ecology, 38,
49–62
JPE575.fm Page 61 Monday, March 5, 2001 11:43 AM

61 Appendix continued
Fire and beetle
Guilds*
biodiversity
Family/subfamily Total
(number of species) Taxa abundance % forest % February Adults Larvae

DERMESTIDAE (1)
Orphinus sp. 284 1 0 100 Sa Sa

ELATERIDAE (16)
sp. 1 2 100 100 O Pre
sp. 332 1 0 0 O Pre
sp. 338 1 100 100 O Pre
Pyrophorinae (11) Agrypnus sp. 333 1 0 0 O Pre
Conoderus sp. 25 24 92 96 O Pre
Conoderus sp. 41 5 0 100 O Pre
Conoderus sp. 49 5 100 100 O Pre
Conoderus sp. 50 4 100 100 O Pre
Conoderus sp. 65 1 0 100 O Pre
Conoderus sp. 167 2 100 100 O Pre
Conoderus sp. 191 1 0 100 O Pre
Conoderus sp. 192 7 0 100 O Pre
Conoderus sp. 336 1 100 100 O Pre
Conoderus sp. 337 1 100 100 O Pre
Cardiophorinae (2) Genus H. sp. 335 1 100 0 O Pre
Paracardiophorus sp. 334 1 0 0 O Pre
HISTERIDAE (1)
Clamydopsinae (1) Clamydopsis sp. 268 1 100 100 Pre Pre
LYCIDAE (3)
Trichalus sp. 37 1 100 100 Ph Sv
Trichalus sp. 55 2 0 100 Ph Sv
Trichalus sp. 203 1 100 100 Ph Sv
MELYRIDAE (7)
Malachiinae (7) Balanophorus sp. 280 1 0 100 Ph Pre
Dicranolaius sp. 3 11 9 55 Ph Pre
Dicranolaius sp. 89 1 0 0 Ph Pre
Dicranolaius sp. 102 1 100 0 Ph Pre
Dicranolaius sp. 106 4 0 50 Ph Pre
Dicranolaius sp. 127 2 100 100 Ph Pre
Dicranolaius sp. 158 1 100 100 Ph Pre
MORDELLIDAE (2)
Mordellisteni (2) Glipostena sp. 292 2 100 100 Ph Ph
Mordellistena sp. 291 1 0 100 Ph Ph
PHALACRIDAE (3)
Phalacrus sp. 20 37 100 78 F F
Phalacrus sp. 95 8 88 63 F F
Phalacrus sp. 289 1 0 0 F F
PSELAPHIDAE (1)
Pselaphaulax sp. 288 1 0 100 Pre Pre
RHIPIPHORIDAE (12)
Ptilophorinae (4) Evaniocera sp. 12 1 0 0 Pre Pre
Evaniocera sp. 21 4 50 75 Pre Pre
Evaniocera sp. 290 1 100 100 Pre Pre
Evaniocera sp. 346 1 100 100 Pre Pre
Pelecotominae (1) Pelecotomoides sp. 81 1 100 0 Pre Pre
Rhipiphorinae (7) Macrosiagon sp. 63 4 0 100 Pre Pre
Macrosiagon sp. 177 3 0 100 Pre Pre
Macrosiagon sp. 179 1 0 100 Pre Pre
Macrosiagon sp. 180 2 0 100 Pre Pre
Macrosiagon sp. 181 1 0 100 Pre Pre
Macrosiagon sp. 184 1 0 100 Pre Pre
Macrosiagon sp. 196 2 50 100 Pre Pre
SCARABAEIDAE (9)
© 2001 British Melolonthinae (4) Heteronyx sp. 43 2 100 100 Ph Sv
Ecological Society, Heteronyx sp. 75 5 80 100 Ph Sv
Journal of Applied Heteronyx sp. 299 4 100 100 Ph Sv
Ecology, 38, Liparetrus sp. 300 5 0 100 Ph Sv
49–62
JPE575.fm Page 62 Monday, March 5, 2001 11:43 AM

62 Appendix continued
J. Orgeas &
Guilds*
A.N. Andersen
Family/subfamily Total
(number of species) Taxa abundance % forest % February Adults Larvae

Scarabaeinae (5) Onthophagus sp. 24 1 100 100 Cop Cop

Onthophagus sp. 295 1 0 0 Cop Cop
Onthophagus sp. 296 1 100 0 Cop Cop
Onthophagus sp. 297 1 0 0 Cop Cop
Onthophagus sp. 298 1 0 0 Cop Cop
SCIRTIDAE (4)
Scirtes sp. 59 12 8 75 Sv Sv
Scirtes sp. 161 4 50 50 Sv Sv

Scirtes sp. 282 2 0 0 Sv Sv

Scirtes sp. 283 1 0 0 Sv Sv
SCRAPTIIDAE (1)
Scraptiinae (1) Scraptia sp. 341 1 0 0 Ph Sv
STAPHYLINIDAE (3)
sp. 150 1 0 100 F F
sp. 165 1 0 100 F F
Scaphidiinae (1) Sciatrophes sp. 339 2 50 100 F F
TENEBRIONIDAE (7)
Alleculinae (5) Chromomoea sp. 110 3 100 100 Sv Sv
Chromomoea sp. 294 1 100 0 Sv Sv
Euomma sp. 173 17 82 100 Sv Sv
Homotrysis sp. 33 6 0 100 Sv Sv
sp. 159 1 100 100 Sv Sv
Tenebrioninae (2) Pterohelaeus sp. 275 1 0 0 Sv Sv
Tenebrio sp. 293 1 0 100 Sv Sv

*Sv, saprophagous on vegetation (dead matter); Sa, saprophagous on animals (dead or decaying); F, fungivorous;
Ph, phytophagous (stems, grass, leaves externally); W, wood borer (living and dead logs); O, omnivorous; Pre, predator;
Cop, coprophagous; Roo, living or dead roots; See, seed predator; Pol, pollen and nectar feeder.

© 2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
49–62